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+The Project Gutenberg EBook of Evolution in Modern Thought, by 
+Ernst Haeckel and J. Arthur Thomson and August Weismann
+
+This eBook is for the use of anyone anywhere at no cost and with
+almost no restrictions whatsoever.  You may copy it, give it away or
+re-use it under the terms of the Project Gutenberg License included
+with this eBook or online at www.gutenberg.org
+
+
+Title: Evolution in Modern Thought
+
+Author: Ernst Haeckel
+        J. Arthur Thomson
+        August Weismann
+
+Release Date: August 29, 2007 [EBook #22430]
+
+Language: English
+
+Character set encoding: UTF-8
+
+*** START OF THIS PROJECT GUTENBERG EBOOK EVOLUTION IN MODERN THOUGHT ***
+
+
+
+
+Produced by Marilynda Fraser-Cunliffe, Sankar Viswanathan,
+and the Online Distributed Proofreading Team at
+http://www.pgdp.net
+
+
+
+
+
+
+
+
+
+                         EVOLUTION IN MODERN
+
+                               THOUGHT
+
+
+
+                    BY HAECKEL, THOMSON, WEISMANN
+
+                              AND OTHERS
+
+
+
+
+
+                          THE MODERN LIBRARY
+
+                      PUBLISHERS :: :: NEW YORK
+
+       *       *       *       *       *
+
+
+
+
+CONTENTS
+
+
+I    DARWIN'S PREDECESSORS
+
+     J. Arthur Thomson, Professor of Natural History in the University of
+       Aberdeen
+
+
+II   _The Selection Theory_
+
+     August Weismann, Professor of Zoology in the University of
+       Freiburg (Baden)
+
+
+III  HEREDITY AND VARIATION IN MODERN LIGHTS
+
+     W. Bateson, Professor of Biology in the University of Cambridge
+
+
+IV   "THE DESCENT OF MAN"
+
+     G. Schwalbe, Professor of Anatomy in the University of Strassburg
+
+
+V    CHARLES DARWIN AS AN ANTHROPOLOGIST
+
+     Ernst Haeckel, Professor of Zoology in the University of Jena
+
+
+VI   MENTAL FACTORS IN EVOLUTION
+
+     C. Lloyd Morgan, Professor of Psychology at University College,
+       Bristol
+
+
+VII  THE INFLUENCE OF THE CONCEPTION OF EVOLUTION ON MODERN PHILOSOPHY
+
+       H. Höffding, Professor of Philosophy in the University of Copenhagen
+
+
+VIII THE INFLUENCE OF DARWIN UPON RELIGIOUS THOUGHT
+
+     Rev. P. H. Waggett
+
+
+IX   DARWINISM AND HISTORY
+
+     J. B. Bury, Regious Professor of Modern History in the University
+       of Cambridge
+
+
+X    DARWINISM AND SOCIOLOGY
+
+     C. Bouglé, Professor of Social Philosophy in the University of
+       Toulouse, and Deputy-Professor at the Sorbonne, Paris
+
+       *       *       *       *       *
+
+
+
+EVOLUTION IN MODERN THOUGHT
+
+
+I
+
+DARWIN'S PREDECESSORS
+
+BY J. ARTHUR THOMSON
+
+_Professor of Natural History in the University of Aberdeen_
+
+
+In seeking to discover Darwin's relation to his predecessors it is
+useful to distinguish the various services which he rendered to the
+theory of organic evolution.
+
+(I) As everyone knows, the general idea of the Doctrine of Descent is
+that the plants and animals of the present day are the lineal
+descendants of ancestors on the whole somewhat simpler, that these
+again are descended from yet simpler forms, and so on backwards
+towards the literal "Protozoa" and "Protophyta" about which we
+unfortunately know nothing. Now no one supposes that Darwin originated
+this idea, which in rudiment at least is as old as Aristotle. What
+Darwin did was to make it current intellectual coin. He gave it a form
+that commended itself to the scientific and public intelligence of the
+day, and he won widespread conviction by showing with consummate skill
+that it was an effective formula to work with, a key which no lock
+refused. In a scholarly, critical, and pre-eminently fair-minded way,
+admitting difficulties and removing them, foreseeing objections and
+forestalling them, he showed that the doctrine of descent supplied a
+modal interpretation of how our present-day fauna and flora have come
+to be.
+
+(II) In the second place, Darwin applied the evolution-idea to
+particular problems, such as the descent of man, and showed what a
+powerful organon it is, introducing order into masses of uncorrelated
+facts, interpreting enigmas both of structure and function, both
+bodily and mental, and, best of all, stimulating and guiding further
+investigation. But here again it cannot be claimed that Darwin was
+original. The problem of the descent or ascent of man, and other
+particular cases of evolution, had attracted not a few naturalists
+before Darwin's day, though no one [except Herbert Spencer in the
+psychological domain (1855)] had come near him in precision and
+thoroughness of inquiry.
+
+(III) In the third place, Darwin contributed largely to a knowledge of
+the factors in the evolution-process, especially by his analysis of
+what occurs in the case of domestic animals and cultivated plants, and
+by his elaboration of the theory of Natural Selection which Alfred
+Russel Wallace independently stated at the same time, and of which
+there had been a few previous suggestions of a more or less vague
+description. It was here that Darwin's originality was greatest, for
+he revealed to naturalists the many different forms--often very
+subtle--which natural selection takes, and with the insight of a
+disciplined scientific imagination he realised what a mighty engine of
+progress it has been and is.
+
+(IV) As an epoch-marking contribution, not only to Ætiology but to
+Natural History in the widest sense, we rank the picture which Darwin
+gave to the world of the web of life, that is to say, of the
+inter-relations and linkages in Nature. For the Biology of the
+individual--if that be not a contradiction in terms--no idea is more
+fundamental than that of the correlation of organs, but Darwin's most
+characteristic contribution was not less fundamental,--it was the idea
+of the correlation of organisms. This, again, was not novel; we find
+it in the works of naturalists like Christian Conrad Sprengel,
+Gilbert White, and Alexander von Humboldt, but the realisation of its
+full import was distinctly Darwinian.
+
+
+_As Regards the General Idea of Organic Evolution_
+
+While it is true, as Prof. H. F. Osborn puts it, that "'Before and
+after Darwin' will always be the _ante et post urbem conditam_ of
+biological history," it is also true that the general idea of organic
+evolution is very ancient. In his admirable sketch _From the Greeks to
+Darwin_,[1] Prof. Osborn has shown that several of the ancient
+philosophers looked upon Nature as a gradual development and as still
+in process of change. In the suggestions of Empedocles, to take the
+best instance, there were "four sparks of truth,--first, that the
+development of life was a gradual process; second, that plants were
+evolved before animals; third, that imperfect forms were gradually
+replaced (not succeeded) by perfect forms; fourth, that the natural
+cause of the production of perfect forms was the extinction of the
+imperfect."[2] But the fundamental idea of one stage giving origin to
+another was absent. As the blue Ægean teemed with treasures of beauty
+and threw many upon its shores, so did Nature produce like a fertile
+artist what had to be rejected as well as what was able to survive,
+but the idea of one species emerging out of another was not yet
+conceived.
+
+Aristotle's views of Nature[3] seem to have been more definitely
+evolutionist than those of his predecessors, in this sense, at least,
+that he recognised not only an ascending scale, but a genetic series
+from polyp to man and an age-long movement towards perfection. "It is
+due to the resistance of matter to form that Nature can only rise by
+degrees from lower to higher types." "Nature produces those things
+which, being continually moved by a certain principle contained in
+themselves, arrive at a certain end."
+
+To discern the outcrop of evolution-doctrine in the long interval
+between Aristotle and Bacon seems to be very difficult, and some of
+the instances that have been cited strike one as forced. Epicurus and
+Lucretius, often called poets of evolution, both pictured animals as
+arising directly out of the earth, very much as Milton's lion long
+afterwards pawed its way out. Even when we come to Bruno who wrote
+that "to the sound of the harp of the Universal Apollo (the World
+Spirit), the lower organisms are called by stages to higher, and the
+lower stages are connected by intermediate forms with the higher,"
+there is great room, as Prof. Osborn points out,[4] for difference of
+opinion as to how far he was an evolutionist in our sense of the term.
+
+The awakening of natural science in the sixteenth century brought the
+possibility of a concrete evolution theory nearer, and in the early
+seventeenth century we find evidences of a new spirit--in the
+embryology of Harvey and the classifications of Ray. Besides sober
+naturalists there were speculative dreamers in the sixteenth and
+seventeenth centuries who had at least got beyond static formulae,
+but, as Professor Osborn points out,[5] "it is a very striking fact,
+that the basis of our modern methods of studying the Evolution problem
+was established not by the early naturalists nor by the speculative
+writers, but by the Philosophers." He refers to Bacon, Descartes,
+Leibnitz, Hume, Kant, Lessing, Herder, and Schelling. "They alone were
+upon the main track of modern thought. It is evident that they were
+groping in the dark for a working theory of the Evolution of life, and
+it is remarkable that they clearly perceived from the outset that the
+point to which observation should be directed was not the past but the
+present mutability of species, and further, that this mutability was
+simply the variation of individuals on an extended scale."
+
+Bacon seems to have been one of the first to think definitely about
+the mutability of species, and he was far ahead of his age in his
+suggestion of what we now call a Station of Experimental Evolution.
+Leibnitz discusses in so many words how the species of animals may be
+changed and how intermediate species may once have linked those that
+now seem discontinuous. "All natural orders of beings present but a
+single chain".... "All advances by degrees in Nature, and nothing by
+leaps." Similar evolutionist statements are to be found in the works
+of the other "philosophers," to whom Prof. Osborn refers, who were,
+indeed, more scientific than the naturalists of their day. It must be
+borne in mind that the general idea of organic evolution--that the
+present is the child of the past--is in great part just the idea of
+human history projected upon the natural world, differentiated by the
+qualification that the continuous "Becoming" has been wrought out by
+forces inherent in the organisms themselves and in their environment.
+
+A reference to Kant[6] should come in historical order after Buffon,
+with whose writings he was acquainted, but he seems, along with Herder
+and Schelling, to be best regarded as the culmination of the
+evolutionist philosophers--of those at least who interested themselves
+in scientific problems. In a famous passage he speaks of "the
+agreement of so many kinds of animals in a certain common plan of
+structure" ... an "analogy of forms" which "strengthens the
+supposition that they have an actual blood-relationship, due to
+derivation from a common parent." He speaks of "the great Family of
+creatures, for as a Family we must conceive it, if the above-mentioned
+continuous and connected relationship has a real foundation." Prof.
+Osborn alludes to the scientific caution which led Kant, biology being
+what it was, to refuse to entertain the hope "that a Newton may one
+day arise even to make the production of a blade of grass
+comprehensible, according to natural laws ordained by no intention."
+As Prof. Haeckel finely observes, Darwin rose up as Kant's Newton.[7]
+
+The scientific renaissance brought a wealth of fresh impressions and
+some freedom from the tyranny of tradition, and the twofold stimulus
+stirred the speculative activity of a great variety of men from old
+Claude Duret of Moulins, of whose weird transformism (1609) Dr. Henry
+de Varigny[8] gives us a glimpse, to Lorenz Oken (1779-1851) whose
+writings are such mixtures of sense and nonsense that some regard him
+as a far-seeing prophet and others as a fatuous follower of
+intellectual will-o'-the-wisps. Similarly, for De Maillet, Maupertuis,
+Diderot, Bonnet, and others, we must agree with Professor Osborn that
+they were not actually in the main Evolution movement. Some have been
+included in the roll of honour on very slender evidence, Robinet for
+instance, whose evolutionism seems to us extremely dubious.[9]
+
+The first naturalist to give a broad and concrete expression to the
+evolutionist doctrine of descent was Buffon (1707-1788), but it is
+interesting to recall the fact that his contemporary Linnæus
+(1707-1778), protagonist of the counter-doctrine of the fixity of
+species,[10] went the length of admitting (in 1762) that new species
+might arise by inter-crossing. Buffon's position among the pioneers of
+the evolution-doctrine is weakened by his habit of vacillating between
+his own conclusions and the orthodoxy of the Sorbonne, but there is no
+doubt that he had firm grasp of the general idea of "l'enchaînment des
+êtres."
+
+Erasmus Darwin (1731-1802), probably influenced by Buffon, was another
+firm evolutionist, and the outline of his argument in the
+_Zoonomia_[11] might serve in part at least to-day. "When we revolve
+in our minds the metamorphoses of animals, as from the tadpole to the
+frog; secondly, the changes produced by artificial cultivation, as in
+the breeds of horses, dogs, and sheep; thirdly, the changes produced
+by conditions of climate and of season, as in the sheep of warm
+climates being covered with hair instead of wool, and the hares and
+partridges of northern climates becoming white in winter: when,
+further, we observe the changes of structure produced by habit, as
+seen especially in men of different occupations; or the changes
+produced by artificial mutilation and prenatal influences, as in the
+crossing of species and production of monsters; fourth, when we
+observe the essential unity of plan in all warm-blooded animals,--we
+are led to conclude that they have been alike produced from a similar
+living filament".... "From thus meditating upon the minute portion of
+time in which many of the above changes have been produced, would it
+be too bold to imagine, in the great length of time since the earth
+began to exist, perhaps millions of years before the commencement of
+the history of mankind, that all warm-blooded animals have arisen from
+one living filament?"... "This idea of the gradual generation of all
+things seems to have been as familiar to the ancient philosophers as
+to the modern ones, and to have given rise to the beautiful
+hieroglyphic figure of the π�ω̃τον ὠ�ν, or first great egg,
+produced by night, that is, whose origin is involved in obscurity, and
+animated by Ἔ�ωσ, that is, by Divine Love; from whence
+proceeded all things which exist."
+
+Lamarck (1744-1829) seems to have become an evolutionist
+independently of Erasmus Darwin's influence, though the parallelism
+between them is striking. He probably owed something to Buffon, but he
+developed his theory along a different line. Whatever view be held in
+regard to that theory there is no doubt that Lamarck was a
+thorough-going evolutionist. Professor Haeckel speaks of the
+_Philosophie Zoologique_ as "the first connected and thoroughly
+logical exposition of the theory of descent."[12]
+
+Besides the three old masters, as we may call them, Buffon, Erasmus
+Darwin, and Lamarck, there were other quite convinced pre-Darwinian
+evolutionists. The historian of the theory of descent must take
+account of Treviranus whose _Biology or Philosophy of Animate Nature_
+is full of evolutionary suggestions; of Etienne Geoffroy St. Hilaire,
+who in 1830, before the French Academy of Sciences, fought with
+Cuvier, the fellow-worker of his youth, an intellectual duel on the
+question of descent; of Goethe, one of the founders of morphology and
+the greatest poet of Evolution--who, in his eighty-first year, heard
+the tidings of Geoffrey St. Hilaire's defeat with an interest which
+transcended the political anxieties of the time; and of many others
+who had gained with more or less confidence and clearness a new
+outlook on Nature. It will be remembered that Darwin refers to
+thirty-four more or less evolutionist authors in his Historical
+Sketch, and the list might be added to. Especially when we come near
+to 1858 do the numbers increase, and one of the most remarkable, as
+also most independent champions of the evolution-idea before that date
+was Herbert Spencer, who not only marshalled the arguments in a very
+forcible way in 1852, but applied the formula in detail in his
+_Principles of Psychology_ in 1855.[13]
+
+It is right and proper that we should shake ourselves free from all
+creationist appreciations of Darwin, and that we should recognise the
+services of pre-Darwinian evolutionists who helped to make the time
+ripe, yet one cannot help feeling that the citation of them is apt to
+suggest two fallacies. It may suggest that Darwin simply entered into
+the labours of his predecessors, whereas, as a matter of fact, he knew
+very little about them till after he had been for years at work. To
+write, as Samuel Butler did, "Buffon planted, Erasmus Darwin and
+Lamarck watered, but it was Mr. Darwin who said 'That fruit is ripe,'
+and shook it into his lap" ... seems to us a quite misleading version
+of the facts of the case. The second fallacy which the historical
+citation is a little apt to suggest is that the filiation of ideas is
+a simple problem. On the contrary, the history of an idea, like the
+pedigree of an organism, is often very intricate, and the evolution of
+the evolution-idea is bound up with the whole progress of the world.
+Thus in order to interpret Darwin's clear formulation of the idea of
+organic evolution and his convincing presentation of it, we have to do
+more than go back to his immediate predecessors, such as Buffon,
+Erasmus Darwin, and Lamarck; we have to inquire into the acceptance of
+evolutionary conceptions in regard to other orders of facts, such as
+the earth and the solar system;[14] we have to realise how the growing
+success of scientific interpretation along other lines gave confidence
+to those who refused to admit that there was any domain from which
+science could be excluded as a trespasser; we have to take account of
+the development of philosophical thought, and even of theological and
+religious movements; we should also, if we are wise enough, consider
+social changes. In short, we must abandon the idea that we can
+understand the history of any science as such, without reference to
+contemporary evolution in other departments of activity.
+
+While there were many evolutionists before Darwin, few of them were
+expert naturalists and few were known outside a small circle; what was
+of much more importance was that the genetic view of Nature was
+insinuating itself in regard to other than biological orders of facts,
+here a little and there a little, and that the scientific spirit had
+ripened since the days when Cuvier laughed Lamarck out of court. How
+was it that Darwin succeeded where others had failed? Because, in the
+first place, he had clear visions--"pensées de la jeunesse, executées
+par l'âge mûr"--which a University curriculum had not made impossible,
+which the _Beagle voyage_ made vivid, which an unrivalled British
+doggedness made real--visions of the web of life, of the fountain of
+change within the organism, of the struggle for existence and its
+winnowing, and of the spreading genealogical tree. Because, in the
+second place, he put so much grit into the verification of his
+visions, putting them to the proof in an argument which is of its
+kind--direct demonstration being out of the question--quite
+unequalled. Because, in the third place, he broke down the opposition
+which the most scientific had felt to the seductive modal formula of
+evolution by bringing forward a more plausible theory of the process
+than had been previously suggested. Nor can one forget, since
+questions of this magnitude are human and not merely academic, that he
+wrote so that all men could understand.
+
+
+_As Regards the Factors of Evolution_
+
+It is admitted by all who are acquainted with the history of biology
+that the general idea of organic evolution as expressed in the
+Doctrine of Descent was quite familiar to Darwin's grandfather and to
+others before and after him, as we have briefly indicated. It must
+also be admitted that some of these pioneers of evolutionism did more
+than apply the evolution-idea as a modal formula of becoming, they
+began to inquire into the factors in the process. Thus there were
+pre-Darwinian theories of evolution, and to these we must now briefly
+refer.[15]
+
+In all biological thinking we have to work with the categories
+Organism--Function--Environment, and theories of evolution may be
+classified in relation to these. To some it has always seemed that the
+fundamental fact is the living organism,--a creative agent, a striving
+will, a changeful Proteus, selecting its environment, adjusting itself
+to it, self-differentiating and self-adaptive. The necessity of
+recognising the importance of the organism is admitted by all
+Darwinians who start with inborn variations, but it is open to
+question whether the whole truth of what we might call the Goethian
+position is exhausted in the postulate of inherent variability.
+
+To others it has always seemed that the emphasis should be laid on
+Function,--on use and disuse, on doing and not doing. Practice makes
+perfect; _c'est à force de forger qu'on devient forgeron_. This is one
+of the fundamental ideas of Lamarckism; to some extent it met with
+Darwin's approval; and it finds many supporters to-day. One of the
+ablest of these--Mr. Francis Darwin--has recently given strong reasons
+for combining a modernised Lamarckism with what we usually regard as
+sound Darwinism.[16]
+
+To others it has always seemed that the emphasis should be laid on the
+Environment, which wakes the organism to action, prompts it to change,
+makes dints upon it, moulds it, prunes it, and finally, perhaps, kills it.
+It is again impossible to doubt that there is truth in this view, for even
+if environmentally induced "modifications" be not transmissible,
+environmentally induced "variations" are; and even if the direct influence
+of the environment be less important than many enthusiastic supporters of
+this view--may we call them Buffonians--think, there remains the indirect
+influence which Darwinians in part rely on,--the eliminative process. Even
+if the extreme view be held that the only form of discriminate elimination
+that counts is inter-organismal competition, this might be included under
+the rubric of the animate environment.
+
+In many passages Buffon[17] definitely suggested that environmental
+influences--especially of climate and food--were directly productive
+of changes in organisms, but he did not discuss the question of the
+transmissibility of the modifications so induced, and it is difficult
+to gather from his inconsistent writings what extent of transformation
+he really believed in. Prof. Osborn says of Buffon: "The struggle for
+existence, the elimination of the least-perfected species, the contest
+between the fecundity of certain species and their constant
+destruction, are all clearly expressed in various passages." He quotes
+two of these:[18]
+
+"Le cours ordinaire de la nature vivante, est en général toujours
+constant, toujours le même; son mouvement, toujours régulier, roule
+sur deux points inébranlables: l'un, la fécondité sans bornes donnée à
+toutes les espèces; l'autre, les obstacles sans nombre qui réduisent
+cette fécondité à une mesure déterminée et ne laissent en tout temps
+qu'à peu près la même quantité d'individus de chaque espèce" ... "Les
+espèces les moins parfaites, les plus délicates, les plus pesantes,
+les moins agissantes, les moins armées, etc., ont déjà disparu ou
+disparaîtront.".
+
+Erasmus Darwin[19] had a firm grip of the "idea of the gradual
+formation and improvement of the Animal world," and he had his theory
+of the process. No sentence is more characteristic than this: "All
+animals undergo transformations which are in part produced by their
+own exertions, in response to pleasures and pains, and many of these
+acquired forms or propensities are transmitted to their posterity."
+This is Lamarckism before Lamarck, as his grandson pointed out. His
+central idea is that wants stimulate efforts and that these result in
+improvements which subsequent generations make better still. He
+realised something of the struggle for existence and even pointed out
+that this advantageously checks the rapid multiplication. "As Dr.
+Krause points out, Darwin just misses the connection between this
+struggle and the Survival of the Fittest."[20]
+
+Lamarck[21] (1744-1829) seems to have thought out his theory of
+evolution without any knowledge of Erasmus Darwin's which it closely
+resembled. The central idea of his theory was the cumulative
+inheritance of functional modifications. "Changes in environment bring
+about changes in the habits of animals. Changes in their wants
+necessarily bring about parallel changes in their habits. If new wants
+become constant or very lasting, they form new habits, the new habits
+involve the use of new parts, or a different use of old parts, which
+results finally in the production of new organs and the modification
+of old ones." He differed from Buffon in not attaching importance, as
+far as animals are concerned, to the direct influence of the
+environment, "for environment can effect no direct change whatever
+upon the organisation of animals," but in regard to plants he agreed
+with Buffon that external conditions directly moulded them.
+
+Treviranus[22] (1776-1837), whom Huxley ranked beside Lamarck, was on
+the whole Buffonian, attaching chief importance to the influence of a
+changeful environment both in modifying and in eliminating, but he was
+also Goethian, for instance in his idea that species like individuals
+pass through periods of growth, full bloom, and decline. "Thus, it is
+not only the great catastrophes of Nature which have caused
+extinction, but the completion of cycles of existence, out of which
+new cycles have begun." A characteristic sentence is quoted by Prof.
+Osborn: "In every living being there exists a capability of an endless
+variety of form-assumption; each possesses the power to adapt its
+organisation to the changes of the outer world, and it is this power,
+put into action by the change of the universe, that has raised the
+simple zoophytes of the primitive world to continually higher stages
+of organisation, and has introduced a countless variety of species
+into animate Nature."
+
+Goethe[23] (1749-1832), who knew Buffon's work but not Lamarck's, is
+peculiarly interesting as one of the first to use the evolution-idea
+as a guiding hypothesis, e.g. in the interpretation of vestigial
+structures in man, and to realise that organisms express an attempt to
+make a compromise between specific inertia and individual change. He
+gave the finest expression that science has yet known--if it has known
+it--of the kernel-idea of what is called "bathmism," the idea of an
+"inherent growth-force"--and at the same time he held that "the way of
+life powerfully reacts upon all form" and that the orderly growth of
+form "yields to change from externally acting causes."
+
+Besides Buffon, Erasmus Darwin, Lamarck, Treviranus, and Goethe,
+there were other "pioneers of evolution," whose views have been often
+discussed and appraised. Étienne Geoffroy Saint-Hilaire (1772-1884),
+whose work Goethe so much admired, was on the whole Buffonian,
+emphasising the direct action of the changeful _milieu_. "Species vary
+with their environment, and existing species have descended by
+modification from earlier and somewhat simpler species." He had a
+glimpse of the selection idea, and believed in mutations or sudden
+leaps--induced in the embryonic condition by external influences. The
+complete history of evolution-theories will include many instances of
+guesses at truth which were afterwards substantiated, thus the
+geographer von Buch (1773-1853) detected the importance of the
+Isolation factor on which Wagner, Romanes, Gulick and others have laid
+great stress, but we must content ourselves with recalling one other
+pioneer, the author of the _Vestiges of Creation_ (1844), a work which
+passed through ten editions in nine years and certainly helped to
+harrow the soil for Darwin's sowing. As Darwin said, "it did excellent
+service in this country in calling attention to the subject, in
+removing prejudice, and in thus preparing the ground for the reception
+of analogous views."[24] Its author, Robert Chambers (1802-1871) was
+in part a Buffonian--maintaining that environment moulded organisms
+adaptively, and in part a Goethian--believing in an inherent
+progressive impulse which lifted organisms from one grade of
+organisation to another.
+
+
+_As Regards Natural Selection_
+
+The only thinker to whom Darwin was directly indebted, so far as the
+theory of Natural Selection is concerned, was Malthus, and we may once
+more quote the well-known passage in the Autobiography: "In October,
+1838, that is, fifteen months after I had begun my systematic enquiry,
+I happened to read for amusement 'Malthus on Population,' and being
+well prepared to appreciate the struggle for existence which
+everywhere goes on from long-continued observation of the habits of
+animals and plants, it at once struck me that under these
+circumstances favourable variations would tend to be preserved, and
+unfavourable ones to be destroyed. The result of this would be the
+formation of new species."[25]
+
+Although Malthus gives no adumbration of the idea of Natural Selection
+in his exposition of the eliminative processes which go on in mankind,
+the suggestive value of his essay is undeniable, as is strikingly
+borne out by the fact that it gave to Alfred Russel Wallace also "the
+long-sought clue to the effective agent in the evolution of organic
+species."[26] One day in Ternate when he was resting between fits of
+fever, something brought to his recollection the work of Malthus which
+he had read twelve years before. "I thought of his clear exposition of
+'the positive checks to increase'--disease, accidents, war, and
+famine--which keep down the population of savage races to so much
+lower an average than that of more civilized peoples. It then occurred
+to me that these causes or their equivalents are continually acting in
+the case of animals also; and as animals usually breed much more
+rapidly than does mankind, the destruction every year from these
+causes must be enormous in order to keep down the numbers of each
+species, since they evidently do not increase regularly from year to
+year, as otherwise the world would long ago have been densely crowded
+with those that breed most quickly. Vaguely thinking over the enormous
+and constant destruction which this implied, it occurred to me to ask
+the question, Why do some die and some live? And the answer was
+clearly, that on the whole the best fitted live. From the effects of
+disease the most healthy escaped; from enemies the strongest, the
+swiftest, or the most cunning; from famine the best hunters or those
+with the best digestion; and so on. Then it suddenly flashed upon me
+that this self-acting process would necessarily _improve the race_,
+because in every generation the inferior would inevitably be killed
+off and the superior would remain--that is, _the fittest would
+survive_."[27] We need not apologise for this long quotation, it is a
+tribute to Darwin's magnanimous colleague, the Nestor of the
+evolutionist camp,--and it probably indicates the line of thought
+which Darwin himself followed. It is interesting also to recall the
+fact that in 1852, when Herbert Spencer wrote his famous _Leader_
+article on "The Development Hypothesis" in which he argued powerfully
+for the thesis that the whole animate world is the result of an
+age-long process of natural transformation, he wrote for _The
+Westminster Review_ another important essay, "A Theory of Population
+deduced from the General Law of Animal Fertility," towards the close
+of which he came within an ace of recognising that the struggle for
+existence was a factor in organic evolution. At a time when pressure
+of population was practically interesting men's minds, Darwin,
+Wallace, and Spencer were being independently led from a social
+problem to a biological theory. There could be no better illustration,
+as Prof. Patrick Geddes has pointed out, of the Comtian thesis that
+science is a "social phenomenon."
+
+Therefore, as far more important than any further ferreting out of
+vague hints of Natural Selection in books which Darwin never read, we
+would indicate by a quotation the view that the central idea in
+Darwinism is correlated with contemporary social evolution. "The
+substitution of Darwin for Paley as the chief interpreter of the order
+of nature is currently regarded as the displacement of an
+anthropomorphic view by a purely scientific one: a little reflection,
+however, will show that what has actually happened has been merely the
+replacement of the anthropomorphism of the eighteenth century by that
+of the nineteenth. For the place vacated by Paley's theological and
+metaphysical explanation has simply been occupied by that suggested to
+Darwin and Wallace by Malthus in terms of the prevalent severity of
+industrial competition, and those phenomena of the struggle for
+existence which the light of contemporary economic theory has enabled
+us to discern, have thus come to be temporarily exalted into a
+complete explanation of organic progress."[28] It goes without saying
+that the idea suggested by Malthus was developed by Darwin into a
+biological theory which was then painstakingly verified by being used
+as an interpretative formula, and that the validity of a theory so
+established is not affected by what suggested it, but the practical
+question which this line of thought raises in the mind is this: if
+Biology did thus borrow with such splendid results from social theory,
+why should we not more deliberately repeat the experiment?
+
+Darwin was characteristically frank and generous in admitting that the
+principle of Natural Selection had been independently recognised by
+Dr. W. C. Wells in 1813 and by Mr. Patrick Matthew in 1831, but he had
+no knowledge of these anticipations when he published the first
+edition of _The Origin of Species_. Wells, whose "Essay on Dew" is
+still remembered, read in 1813 before the Royal Society a short paper
+entitled "An Account of a White Female, part of whose skin resembles
+that of a Negro" (published in 1818). In this communication, as Darwin
+said, "he observes, firstly, that all animals tend to vary in some
+degree, and, secondly, that agriculturists improve their domesticated
+animals by selection; and then, he adds, but what is done in this
+latter case 'by art, seems to be done with equal efficacy, though more
+slowly, by nature, in the formation of varieties of mankind, fitted
+for the country which they inhabit.'"[29] Thus Wells had the clear
+idea of survival dependent upon a favourable variation, but he makes
+no more use of the idea and applies it only to man. There is not in
+the paper the least hint that the author ever thought of generalising
+the remarkable sentence quoted above.
+
+Of Mr. Patrick Matthew, who buried his treasure in an appendix to a
+work on _Naval Timber and Arboriculture_, Darwin said that "he clearly
+saw the full force of the principle of natural selection." In 1860
+Darwin wrote--very characteristically--about this to Lyell: "Mr.
+Patrick Matthew publishes a long extract from his work on _Naval
+Timber and Arboriculture_, published in 1831, in which he briefly but
+completely anticipates the theory of Natural Selection. I have ordered
+the book, as some passages are rather obscure, but it is certainly, I
+think, a complete but not developed anticipation. Erasmus always said
+that surely this would be shown to be the case some day. Anyhow, one
+may be excused in not having discovered the fact in a work on Naval
+Timber."[30]
+
+De Quatrefages and De Varigny have maintained that the botanist Naudin
+stated the theory of evolution by natural selection in 1852. He
+explains very clearly the process of artificial selection, and says
+that in the garden we are following Nature's method. "We do not think
+that Nature has made her species in a different fashion from that in
+which we proceed ourselves in order to make our variations." But, as
+Darwin said, "he does not show how selection acts under nature."
+Similarly it must be noted in regard to several pre-Darwinian pictures
+of the struggle for existence (such as Herder's, who wrote in 1790
+"All is in struggle ... each one for himself" and so on), that a
+recognition of this is only the first step in Darwinism.
+
+Profs. E. Perrier and H. F. Osborn have called attention to a
+remarkable anticipation of the selection-idea which is to be found in
+the speculations of Étienne Geoffroy Saint-Hilaire (1825-1828) on the
+evolution of modern Crocodilians from the ancient Teleosaurs. Changing
+environment induced changes in the respiratory system and far-reaching
+consequences followed. The atmosphere, acting upon the pulmonary
+cells, brings about "modifications which are favourable or destructive
+('funestes'); these are inherited, and they influence all the rest of
+the organisation of the animal because if these modifications lead to
+injurious effects the animals which exhibit them perish and are
+replaced by others of a somewhat different form, a form changed so as
+to be adapted to (à la convenance) the new environment."
+
+Prof. E. B. Poulton[31] has shown that the anthropologist James Cowles
+Prichard (1786-1848) must be included even in spite of himself among
+the precursors of Darwin. In some passages of the second edition of
+his _Researches into the Physical History of Mankind_ (1826), he
+certainly talks evolution and anticipates Prof. Weismann in denying
+the transmission of acquired characters. He is, however, sadly
+self-contradictory and his evolutionism weakens in subsequent
+editions--the only ones that Darwin saw. Prof. Poulton finds in
+Prichard's work a recognition of the operation of Natural Selection.
+"After inquiring how it is that 'these varieties are developed and
+preserved in connexion with particular climates and differences of
+local situation,' he gives the following very significant answer: 'One
+cause which tends to maintain this relation is obvious. Individuals
+and families, and even whole colonies perish and disappear in climates
+for which they are, by peculiarity of constitution, not adapted. Of
+this fact proofs have been already mentioned.'" Mr. Francis Darwin and
+Prof. A. C. Seward discuss Prichard's "anticipations" in _More Letters
+of Charles Darwin_, Vol. _I._ p. 43, and come to the conclusion that
+the evolutionary passages are entirely neutralised by others of an
+opposite trend. There is the same difficulty with Buffon.
+
+Hints of the idea of Natural Selection have been detected elsewhere.
+James Watt,[32] for instance, has been reported as one of the
+anticipators (1851). But we need not prolong the inquiry further,
+since Darwin did not know of any anticipations until after he had
+published the immortal work of 1859, and since none of those who got
+hold of the idea made any use of it. What Darwin did was to follow the
+clue which Malthus gave him, to realise, first by genius and
+afterwards by patience, how the complex and subtle struggle for
+existence works out a natural selection of those organisms which vary
+in the direction of fitter adaptation to the conditions of their life.
+So much success attended his application of the Selection-formula that
+for a time he regarded Natural Selection as almost the sole factor in
+evolution, variations being pre-supposed; gradually, however, he came
+to recognise that there was some validity in the factors which had
+been emphasised by Lamarck and by Buffon, and in his well known
+summing up in the sixth edition of the _Origin_ he says of the
+transformation of species: "This has been effected chiefly through the
+natural selection of numerous successive, slight, favourable
+variations; aided in an important manner by the inherited effects of
+the use and disuse of parts; and in an unimportant manner, that is, in
+relation to adaptive structures, whether past or present, by the
+direct action of external conditions, and by variations which seem to
+us in our ignorance to arise spontaneously."
+
+To sum up: the idea of organic evolution, older than Aristotle, slowly
+developed from the stage of suggestion to the stage of verification,
+and the first convincing verification was Darwin's; from being an _a
+priori_ anticipation it has become an interpretation of nature, and
+Darwin is still the chief interpreter; from being a modal
+interpretation it has advanced to the rank of a causal theory, the
+most convincing part of which men will never cease to call Darwinism.
+
+FOOTNOTES:
+
+[Footnote 1: _Columbia University Biological Series_, Vol. I. New York
+and London, 1894. We must acknowledge our great indebtedness to this
+fine piece of work.]
+
+[Footnote 2: _op. cit._ p. 41.]
+
+[Footnote 3: See G. J. Romanes, "Aristotle as a Naturalist,"
+_Contemporary Review_, Vol. lix. p. 275, 1891; G. Pouchet, _La
+Biologie Aristotélique_, Paris, 1885; E. Zeller, _A History of Greek
+Philosophy_, London, 1881, and "Ueber die griechischen Vorgänger
+Darwin's," _Abhandl. Berlin Akad._ 1878, pp. 111-124.]
+
+[Footnote 4: _op. cit._ p. 81.]
+
+[Footnote 5: _op. cit._ p. 87.]
+
+[Footnote 6: See Brock, "Die Stellung Kant's zur Deszendenztheorie,"
+_Biol. Centralbl._ viii. 1889, pp. 641-648. Fritz Schultze, _Kant und
+Darwin_, Jena, 1875.]
+
+[Footnote 7: Mr. Alfred Russel Wallace writes: "We claim for Darwin
+that he is the Newton of natural history, and that, just so surely as
+that the discovery and demonstration by Newton of the law of
+gravitation established order in place of chaos and laid a sure
+foundation for all future study of the starry heavens, so surely has
+Darwin, by his discovery of the law of natural selection and his
+demonstration of the great principle of the preservation of useful
+variations in the struggle for life, not only thrown a flood of light
+on the process of development of the whole organic world, but also
+established a firm foundation for all future study of nature"
+(_Darwinism_, London, 1889, p. 9). See also Prof. Karl Pearson's
+_Grammar of Science_ (2nd edit.), London, 1900, p. 32. See Osborn,
+_op. cit._ p. 100.]
+
+[Footnote 8: _Experimental Evolution_. London, 1892. Chap. I. p. 14.]
+
+[Footnote 9: See J. Arthur Thomson, _The Science of Life_. London,
+1899, Chap. XVI. "Evolution of Evolution Theory."]
+
+[Footnote 10: See Carus Sterne (Ernst Krause), _Die allgemeine
+Weltanschauung in ihrer historischen Entwickelung_. Stuttgart, 1889.
+Chapter entitled "Beständigkeit oder Veränderlichkeit der
+Naturwesen."]
+
+[Footnote 11: _Zoonomia, or the Laws of Organic Life_, 2 vols. London,
+1794; Osborn, _op. cit._ p. 145.]
+
+[Footnote 12: See Alpheus S. Packard, _Lamarck, the Founder of
+Evolution, His Life and Work, with Translations of his writings on
+Organic Evolution_. London, 1901.]
+
+[Footnote 13: See Edward Clodd, _Pioneers of Evolution_, London, p.
+161, 1897.]
+
+[Footnote 14: See Chapter ix. "The Genetic View of Nature" in J. T.
+Merz's _History of European Thought in the Nineteenth Century_, Vol.
+2, Edinburgh and London, 1903.]
+
+[Footnote 15: See Prof. W. A. Locy's _Biology and its Makers_. New
+York, 1908. Part II. "The Doctrine of Organic Evolution."]
+
+[Footnote 16: Presidential Address to the British Association meeting
+at Dublin in 1908.]
+
+[Footnote 17: See in particular Samuel Butler, _Evolution Old and
+New_, London, 1879; J. L. de Lanessan, "Buffon et Darwin," _Revue
+Scientifique_, XLIII. pp. 385-391, 425-432, 1889.]
+
+[Footnote 18: _op. cit._ p. 136.]
+
+[Footnote 19: See Ernest Krause and Charles Darwin, _Erasmus Darwin_,
+London, 1879.]
+
+[Footnote 20: Osborn, _op. cit._ p. 142.]
+
+[Footnote 21: See E. Perrier, _La Philosophie Zoologique avant
+Darwin_, Paris, 1884; A. de Quatrefages, _Darwin et ses Précurseurs
+Français_, Paris, 1870; Packard, _op. cit._; also Claus, _Lamarck als
+Begründer der Descendenzlehre_, Wien, 1888; Haeckel, _Natural History
+of Creation_, Eng. transl. London, 1879; Lang, _Zur Charakteristik der
+Forschungswege von Lamarck und Darwin_, Jena, 1889.]
+
+[Footnote 22: See Huxley's article "Evolution in Biology,"
+_Encyclopaedia Britannica_ (9th edit.), 1879, pp. 744-751, and Sully's
+article, "Evolution in Philosophy," _ibid._ pp. 751-772.]
+
+[Footnote 23: See Haeckel, _Die Naturanschauung von Darwin, Goethe und
+Lamarck_, Jena, 1882.]
+
+[Footnote 24: _Origin of Species_ (6th edit.), p. xvii.]
+
+[Footnote 25: _The Life and Letters of Charles Darwin_, Vol. 1. p. 83.
+London, 1887.]
+
+[Footnote 26: A. R. Wallace, _My Life, a Record of Events and
+Opinions_, London, 1905, Vol. 1, p. 232.]
+
+[Footnote 27: _My Life_, Vol. 1. p. 361.]
+
+[Footnote 28: P. Geddes. article "Biology." _Chambers's
+Encyclopaedia._]
+
+[Footnote 29: _Origin of Species_ (6th edit.), p. xv.]
+
+[Footnote 30: _Life and Letters_, II, p. 301.]
+
+[Footnote 31: _Science Progress_, New Series, Vol. 1. 1897. "A
+Remarkable Anticipation of Modern Views on Evolution." See also Chap.
+VI. in _Essays on Evolution_, Oxford, 1908.]
+
+[Footnote 32: See Prof. Patrick Geddes's article "Variation and
+Selection," _Encyclopaedia Britannica_ (9th edit.) 1888.]
+
+
+
+
+II
+
+THE SELECTION THEORY
+
+BY AUGUST WEISMANN
+
+_Professor of Zoology in the University of Freiburg_ (_Baden_)
+
+
+I. THE IDEA OF SELECTION
+
+Many and diverse were the discoveries made by Charles Darwin in the
+course of a long and strenuous life, but none of them has had so
+far-reaching an influence on the science and thought of his time as
+the theory of selection. I do not believe that the theory of evolution
+would have made its way so easily and so quickly after Darwin took up
+the cudgels in favour of it if he had not been able to support it by a
+principle which was capable of solving, in a simple manner, the
+greatest riddle that living nature presents to us,--I mean the
+purposiveness of every living form relative to the conditions of its
+life and its marvellously exact adaptation to these.
+
+Everyone knows that Darwin was not alone in discovering the principle
+of selection, and that the same idea occurred simultaneously and
+independently to Alfred Russel Wallace. At the memorable meeting of
+the Linnean Society on 1st July, 1858, two papers were read
+(communicated by Lyell and Hooker) both setting forth the same idea of
+selection. One was written by Charles Darwin in Kent, the other by
+Alfred Wallace in Ternate, in the Malay Archipelago. It was a splendid
+proof of the magnanimity of these two investigators, that they thus in
+all friendliness and without envy, united in laying their ideas
+before a scientific tribunal: their names will always shine side by
+side as two of the brightest stars in the scientific sky.
+
+The idea of selection set forth by the two naturalists was at the time
+absolutely new, but it was also so simple that Huxley could say of it
+later, "How extremely stupid not to have thought of that." As Darwin
+was led to the general doctrine of descent, not through the labours of
+his predecessors in the early years of the century, but by his own
+observations, so it was in regard to the principle of selection. He
+was struck by the innumerable cases of adaptation, as, for instance,
+that of the woodpeckers and tree-frogs to climbing, or the hooks and
+feather-like appendages of seeds, which aid in the distribution of
+plants, and he said to himself that an explanation of adaptations was
+the first thing to be sought for in attempting to formulate a theory
+of evolution.
+
+But since adaptations point to _changes_ which have been undergone by
+the ancestral forms of existing species, it is necessary, first of
+all, to inquire how far species in general are _variable_. Thus
+Darwin's attention was directed in the first place to the phenomenon
+of variability, and the use man has made of this, from very early
+times, in the breeding of his domesticated animals and cultivated
+plants. He inquired carefully how breeders set to work, when they
+wished to modify the structure and appearance of a species to their
+own ends, and it was soon clear to him that _selection for breeding
+purposes_ played the chief part.
+
+But how was it possible that such processes should occur in free
+nature? Who is here the breeder, making the selection, choosing out
+one individual to bring forth offspring and rejecting others? That was
+the problem that for a long time remained a riddle to him.
+
+Darwin himself relates how illumination suddenly came to him. He had
+been reading, for his own pleasure, Malthus' book on Population, and,
+as he had long known from numerous observations, that every species
+gives rise to many more descendants than ever attain to maturity, and
+that, therefore, the greater number of the descendants of a species
+perish without reproducing, the idea came to him that the decision as
+to which member of a species was to perish and which was to attain to
+maturity and reproduction might not be a matter of chance, but might
+be determined by the constitution of the individuals themselves,
+according as they were more or less fitted for survival. With this
+idea the foundation of the theory of selection was laid.
+
+In _artificial selection_ the breeder chooses out for pairing only
+such individuals as possess the character desired by him in a somewhat
+higher degree than the rest of the race. Some of the descendants
+inherit this character, often in a still higher degree, and if this
+method be pursued throughout several generations, the race is
+transformed in respect of that particular character.
+
+_Natural selection_ depends on the same three factors as _artificial
+selection_: on _variability_, _inheritance_, and _selection for
+breeding_, but this last is here carried out not by a breeder but by
+what Darwin called the "struggle for existence." This last factor is
+one of the special features of the Darwinian conception of nature.
+That there are carnivorous animals which take heavy toll in every
+generation of the progeny of the animals on which they prey, and that
+there are herbivores which decimate the plants in every generation had
+long been known, but it is only since Darwin's time that sufficient
+attention has been paid to the facts that, in addition to this regular
+destruction, there exists between the members of a species a keen
+competition for space and food, which limits multiplication, and that
+numerous individuals of each species perish because of unfavourable
+climatic conditions. The "struggle for existence," which Darwin
+regarded as taking the place of the human breeder in free nature, is
+not a direct struggle between carnivores and their prey, but is the
+assumed competition for survival between individuals _of the same_
+species, of which, on an average, only those survive to reproduce
+which have the greatest power of resistance, while the others, less
+favourably constituted, perish early. This struggle is so keen, that,
+within a limited area, where the conditions of life have long remained
+unchanged, of every species, whatever be the degree of fertility, only
+two, _on an average_, of the descendants of each pair survive; the
+others succumb either to enemies, or to disadvantages of climate, or
+to accident. A high degree of fertility is thus not an indication of
+the special success of a species, but of the numerous dangers that
+have attended its evolution. Of the six young brought forth by a pair
+of elephants in the course of their lives only two survive in a given
+area; similarly, of the millions of eggs which two thread-worms leave
+behind them only two survive. It is thus possible to estimate the
+dangers which threaten a species by its ratio of elimination, or,
+since this cannot be done directly, by its fertility.
+
+Although a great number of the descendants of each generation fall
+victims to accident, among those that remain it is still the greater
+or less fitness of the organism that determines the "selection for
+breeding purposes," and it would be incomprehensible if, in this
+competition, it were not ultimately, that is, on an average, the best
+equipped which survive, in the sense of living long enough to
+reproduce.
+
+Thus the principle of natural selection is _the selection of the best
+for reproduction_, whether the "best" refers to the whole
+constitution, to one or more parts of the organism, or to one or more
+stages of development. Every organ, every part, every character of an
+animal, fertility and intelligence included, must be improved in this
+manner, and be gradually brought up in the course of generations to
+its highest attainable state of perfection. And not only may
+improvement of parts be brought about in this way, but new parts and
+organs may arise, since, through the slow and minute steps of
+individual or "fluctuating" variations, a part may be added here or
+dropped out there, and thus something new is produced.
+
+The principle of selection solved the riddle as to how what was
+purposive could conceivably be brought about without the intervention
+of a directing power, the riddle which animate nature presents to our
+intelligence at every turn, and in face of which the mind of a Kant
+could find no way out, for he regarded a solution of it as not to be
+hoped for. For, even if we were to assume an evolutionary force that
+is continually transforming the most primitive and the simplest forms
+of life into ever higher forms, and the homogeneity of primitive times
+into the infinite variety of the present, we should still be unable to
+infer from this alone how each of the numberless forms adapted to
+particular conditions of life should have appeared _precisely at the
+right moment in the history of the earth_ to which their adaptations
+were appropriate, and precisely at the proper place in which all the
+conditions of life to which they were adapted occurred: the
+humming-birds at the same time as the flowers; the trichina at the
+same time as the pig; the bark-coloured moth at the same time as the
+oak, and the wasp-like moth at the same time as the wasp which
+protects it. Without processes of selection we should be obliged to
+assume a "pre-established harmony" after the famous Leibnitzian model,
+by means of which the clock of the evolution of organisms is so
+regulated as to strike in exact synchronism with that of the history
+of the earth! All forms of life are strictly adapted to the conditions
+of their life, and can persist under these conditions alone.
+
+There must therefore be an intrinsic connection between the conditions
+and the structural adaptations of the organism, and, _since the
+conditions of life cannot be determined by the animal itself, the
+adaptations must be called forth by the conditions_.
+
+The selection theory teaches us how this is conceivable, since it
+enables us to understand that there is a continual production of what
+is non-purposive as well as of what is purposive, but the purposive
+alone survives, while the non-purposive perishes in the very act of
+arising. This is the old wisdom taught long ago by Empedocles.
+
+
+II. THE LAMARCKIAN PRINCIPLE
+
+Lamarck, as is well known, formulated a definite theory of evolution
+at the beginning of the nineteenth century, exactly fifty years before
+the Darwin-Wallace principle of selection was given to the world. This
+brilliant investigator also endeavoured to support his theory by
+demonstrating forces which might have brought about the
+transformations of the organic world in the course of the ages. In
+addition to other factors, he laid special emphasis on the increased
+or diminished use of the parts of the body, assuming that the
+strengthening or weakening which takes place from this cause during
+the individual life, could be handed on to the offspring, and thus
+intensified and raised to the rank of a specific character. Darwin
+also regarded this _Lamarckian principle_, as it is now generally
+called, as a factor in evolution, but he was not fully convinced of
+the transmissibility of acquired characters.
+
+As I have here to deal only with the theory of selection, I need not
+discuss the Lamarckian hypothesis, but I must express my opinion that
+there is room for much doubt as to the coöperation of this principle
+in evolution. Not only is it difficult to imagine how the transmission
+of functional modifications could take place, but, up to the present
+time, notwithstanding the endeavours of many excellent investigators,
+not a single actual proof of such inheritance has been brought
+forward. Semon's experiments on plants are, according to the botanist
+Pfeffer, not to be relied on, and even the recent, beautiful
+experiments made by Dr. Kammerer on salamanders, cannot, as I hope to
+show elsewhere, be regarded as proof, if only because they do not deal
+at all with functional modifications, that is, with modifications
+brought about by use, and it is to these _alone_ that the Lamarckian
+principle refers.
+
+
+III. OBJECTIONS TO THE THEORY OF SELECTION
+
+
+(_a_) _Saltatory evolution_
+
+The Darwinian doctrine of evolution depends essentially on _the
+cumulative augmentation_ of minute variations in the direction of
+utility. But can such minute variations, which are undoubtedly
+continually appearing among the individuals of the same species,
+possess any selection-value; can they determine which individuals are
+to survive, and which are to succumb; can they be increased by natural
+selection till they attain to the highest development of a purposive
+variation?
+
+To many this seems so improbable that they have urged a theory of
+evolution by leaps from species to species. Kölliker, in 1872,
+compared the evolution of species with the processes which we can
+observe in the individual life in cases of alternation of generations.
+But a polyp only gives rise to a medusa because it has itself arisen
+from one, and there can be no question of a medusa ever having arisen
+suddenly and _de novo_ from a polyp-bud, if only because both forms
+are adapted in their structure as a whole, and in every detail to the
+conditions of their life. A sudden origin, in a natural way, of
+numerous adaptations is inconceivable. Even the degeneration of a
+medusoid from a free-swimming animal to a mere brood-sac (gonophore)
+is not sudden and saltatory, but occurs by imperceptible modifications
+throughout hundreds of years, as we can learn from the numerous stages
+of the process of degeneration persisting at the same time in
+different species.
+
+If, then, the degeneration to a simple brood-sac takes place only by
+very slow transitions, each stage of which may last for centuries, how
+could the much more complex _ascending_ evolution possibly have taken
+place by sudden leaps? I regard this argument as capable of further
+extension, for wherever in nature we come upon degeneration, it is
+taking place by minute steps and with a slowness that makes it not
+directly perceptible, and I believe that this in itself justifies us
+in concluding that _the same must be true of ascending_ evolution. But
+in the latter case the goal can seldom be distinctly recognised while
+in cases of degeneration the starting-point of the process can often
+be inferred, because several nearly related species may represent
+different stages.
+
+In recent years Bateson in particular has championed the idea of
+saltatory, or so-called discontinuous evolution, and has collected a
+number of cases in which more or less marked variations have suddenly
+appeared. These are taken for the most part from among domesticated
+animals which have been bred and crossed for a long time, and it is
+hardly to be wondered at that their much mixed and much influenced
+germ-plasm should, under certain conditions, give rise to remarkable
+phenomena, often indeed producing forms which are strongly suggestive
+of monstrosities, and which would undoubtedly not survive in free
+nature, unprotected by man. I should regard such cases as due to an
+intensified germinal selection--though this is to anticipate a
+little--and from this point of view it cannot be denied that they have
+a special interest. But they seem to me to have no significance as far
+as the transformation of species is concerned, if only because of the
+extreme rarity of their occurrence.
+
+There are, however, many variations which have appeared in a sudden
+and saltatory manner, and some of these Darwin pointed out and
+discussed in detail: the copper beech, the weeping trees, the oak with
+"fern-like leaves," certain garden-flowers, etc. But none of them have
+persisted in free nature, or evolved into permanent types.
+
+On the other hand, wherever enduring types have arisen, we find traces
+of a gradual origin by successive stages, even if, at first sight,
+their origin may appear to have been sudden. This is the case with
+_seasonal Dimorphism_, the first known cases of which exhibited marked
+differences between the two generations, the winter and the summer
+brood. Take for instance the much discussed and studied form
+_Vanessa_ (_Araschnia_) _levana-prorsa_. Here the differences between
+the two forms are so great and so apparently disconnected, that one
+might almost believe it to be a sudden mutation, were it not that old
+transition-stages can be called forth by particular temperatures, and
+we know other butterflies, as for instance our Garden Whites, in which
+the differences between the two generations are not nearly so marked;
+indeed, they are so little apparent that they are scarcely likely to
+be noticed except by experts. Thus here again there are small initial
+steps, some of which, indeed, must be regarded as adaptations, such as
+the green-sprinkled or lightly tinted under-surface which gives them a
+deceptive resemblance to parsley or to Cardamine leaves.
+
+Even if saltatory variations do occur, we cannot assume that these
+_have ever led to forms which are capable of survival under the
+conditions of wild life_. Experience has shown that in plants which
+have suddenly varied the power of persistence is diminished.
+Korschinsky attributes to them weaknesses of organisation in general;
+"they bloom late, ripen few of their seeds, and show great
+sensitiveness to cold." These are not the characters which make for
+success in the struggle for existence.
+
+We must briefly refer here to the views--much discussed in the last
+decade--of H. de Vries, who believes that the roots of transformation
+must be sought for in _saltatory variations arising from internal
+causes_, and distinguishes such _mutations_, as he has called them,
+from ordinary individual variations, in that they breed true, that is,
+with strict in-breeding they are handed on pure to the next
+generation. I have elsewhere endeavoured to point out the weaknesses
+of this theory,[33] and I am the less inclined to return to it here
+that it now appears[34] that the far-reaching conclusions drawn by de
+Vries from his observations on the Evening Primrose, _Oenothera
+lamarckiana_, rest upon a very insecure foundation. The plant from
+which de Vries saw numerous "species"--his "mutations"--arise was not,
+as he assumed, a _wild species_ that had been introduced to Europe
+from America, but was probably a hybrid form which was first
+discovered in the Jardin des Plantes in Paris, and which does not
+appear to exist anywhere in America as a wild species.
+
+This gives a severe shock to the "Mutation theory," for the other
+_actually wild_ species with which de Vries experimented showed no
+"mutations" but yielded only negative results.
+
+Thus we come to the conclusion that Darwin[35] was right in regarding
+transformations as taking place by minute steps, which, if useful, are
+augmented in the course of innumerable generations, because their
+possessors more frequently survive in the struggle for existence.
+
+
+(_b_) _Selection-value of the initial steps_
+
+Is it possible that the insignificant deviations which we know as
+"individual variations" can form the beginning of a process of
+selection? Can they decide which is to perish and which to survive? To
+use a phrase of Romanes, can they have _selection-value_?
+
+Darwin himself answered this question, and brought together many
+excellent examples to show that differences, apparently insignificant
+because very small, might be of decisive importance for the life of
+the possessor. But it is by no means enough to bring forward cases of
+this kind, for the question is not merely whether finished adaptations
+have selection-value, but whether the first beginnings of these, and
+whether the small, I might almost say minimal increments, which have
+led up from these beginnings to the perfect adaptation, have also had
+selection-value. To this question even one who, like myself, has been
+for many years a convinced adherent of the theory of selection, can
+only reply: _We must assume so, but we cannot prove it in any case_.
+It is not upon demonstrative evidence that we rely when we champion
+the doctrine of selection as a scientific truth; we base our argument
+on quite other grounds. Undoubtedly there are many apparently
+insignificant features, which can nevertheless be shown to be
+adaptations--for instance, the thickness of the basin-shaped shell of
+the limpets that live among the breakers on the shore. There can be no
+doubt that the thickness of these shells, combined with their flat
+forms, protects the animals from the force of the waves breaking upon
+them,--but how have they become so thick? What proportion of thickness
+was sufficient to decide that of two variants of a limpet one should
+survive, the other be eliminated? We can say nothing more than that we
+infer from the present state of the shell, that it must have varied in
+regard to differences in shell-thickness, and that these differences
+must have had selection-value,--no proof therefore, but an assumption
+which we must show to be convincing.
+
+For a long time the marvellously complex _radiate_ and _lattice-work_
+skeletons of Radiolarians were regarded as a mere outflow of "Nature's
+infinite wealth of form," as an instance of a purely morphological
+character with no biological significance. But recent investigations
+have shown that these, too, have an adaptive significance (Häcker).
+The same thing has been shown by Schütt in regard to the lowly
+unicellular plants, the Peridineae, which abound alike on the surface
+of the ocean and in its depths. It has been shown that the long
+skeletal processes which grow out from these organisms have
+significance not merely as a supporting skeleton, but also as an
+extension of the superficial area, which increases the contact with
+the water-particles, and prevents the floating organisms from sinking.
+It has been established that the processes are considerably shorter in
+the colder layers of the ocean, and that they may be twelve times as
+long[36] in the warmer layers, thus corresponding to the greater or
+smaller amount of friction which takes place in the denser and less
+dense layers of the water.
+
+The Peridineae of the warmer ocean layers have thus become long-rayed,
+those of the colder layers short-rayed, not through the direct effect
+of friction on the protoplasm, but through processes of selection,
+which favoured the longer rays in warm water, since they kept the
+organism afloat, while those with short rays sank and were eliminated.
+If we put the question as to selection-value in this case, and ask how
+great the variations in the length of processes must be in order to
+possess selection-value; what can we answer except that these
+variations must have been minimal, and yet sufficient to prevent too
+rapid sinking and consequent elimination? Yet this very case would
+give the ideal opportunity for a mathematical calculation of the
+minimal selection-value, although of course it is not feasible from
+lack of data to carry out the actual calculation.
+
+But even in organisms of more than microscopic size there must
+frequently be minute, even microscopic differences which set going the
+process of selection, and regulate its progress to the highest
+possible perfection.
+
+Many tropical trees possess thick, leathery leaves, as a protection
+against the force of the tropical raindrops. The _direct_ influence of
+the rain cannot be the cause of this power of resistance, for the
+leaves, while they were still thin, would simply have been torn to
+pieces. Their toughness must therefore be referred to selection, which
+would favour the trees with slightly thicker leaves, though we cannot
+calculate with any exactness how great the first stages of increase in
+thickness must have been. Our hypothesis receives further support from
+the fact that, in many such trees, the leaves are drawn out into a
+beak-like prolongation (Stahl and Haberlandt) which facilitates the
+rapid falling off of the rain water, and also from the fact that the
+leaves, while they are still young, hang limply down in bunches which
+offer the least possible resistance to the rain. Thus there are here
+three adaptations which can only be interpreted as due to selection.
+The initial stages of these adaptations must undoubtedly have had
+selection-value.
+
+But even in regard to this case we are reasoning in a circle, not
+giving "proofs," and no one who does not wish to believe in the
+selection-value of the initial stages can be forced to do so. Among
+the many pieces of presumptive evidence a particularly weighty one
+seems to me to be _the smallness of the steps of progress_ which we
+can observe in certain cases, as for instance in leaf-imitation among
+butterflies, and in mimicry generally. The resemblance to a leaf, for
+instance of a particular Kallima, seems to us so close as to be
+deceptive, and yet we find in another individual, or it may be in many
+others, a spot added which increases the resemblance, and which could
+not have become fixed unless the increased deceptiveness so produced
+had frequently led to the overlooking of its much persecuted
+possessor. But if we take the selection-value of the initial stages
+for granted, we are confronted with the further question which I
+myself formulated many years ago: How does it happen _that the
+necessary beginnings of a useful variation are always present_? How
+could insects which live upon or among green leaves become all green,
+while those that live on bark become brown? How have the desert
+animals become yellow and the Arctic animals white? Why were the
+necessary variations always present? How could the green locust lay
+brown eggs, or the privet caterpillar develop white and lilac-coloured
+lines on its green skin?
+
+It is of no use answering to this that the question is wrongly
+formulated[37] and that it is the converse that is true; that the
+process of selection takes place in accordance with the variations
+that present themselves. This proposition is undeniably true, but so
+also is another, which apparently negatives it: the variation required
+has in the majority of cases actually presented itself. Selection
+cannot solve this contradiction; it does not call forth the useful
+variation, but simply works upon it. The ultimate reason why one and
+the same insect should occur in green and in brown, as often happens
+in caterpillars and locusts, lies in the fact that variations towards
+brown presented themselves, and so also did variations towards green:
+_the kernel of the riddle lies in the varying_, and for the present we
+can only say, that small variations in different directions present
+themselves in every species. Otherwise so many different kinds of
+variations could not have arisen. I have endeavoured to explain this
+remarkable fact by means of the intimate processes that must take
+place within the germ-plasm, and I shall return to the problem when
+dealing with "germinal selection."
+
+We have, however, to make still greater demands on variation, for it
+is not enough that the necessary variation should occur in isolated
+individuals, because in that case there would be small prospect of its
+being preserved, notwithstanding its utility. Darwin at first
+believed, that even single variations might lead to transformation of
+the species, but later he became convinced that this was impossible,
+at least without the coöperation of other factors, such as isolation
+and sexual selection.
+
+In the case of the _green caterpillars with bright longitudinal
+stripes_, numerous individuals exhibiting this useful variation must
+have been produced to start with. In all higher, that is,
+multicellular organisms, the germ-substance is the source of all
+transmissible variations, and this germ-plasm is not a simple
+substance but is made up of many primary constituents. The question
+can therefore be more precisely stated thus: How does it come about
+that in so many cases the useful variations present themselves in
+numbers just where they are required, the white oblique lines in the
+leaf-caterpillar on the under surface of the body, the accompanying
+coloured stripes just above them? And, further, how has it come about
+that in grass caterpillars, not oblique but longitudinal stripes,
+which are more effective for concealment among grass and plants, have
+been evolved? And finally, how is it that the same Hawk-moth
+caterpillars, which to-day show oblique stripes, possessed
+longitudinal stripes in Tertiary times? We can read this fact from the
+history of their development, and I have before attempted to show the
+biological significance of this change of colour.[38]
+
+For the present I need only draw the conclusion that one and the same
+caterpillar may exhibit the initial stages of both, and that it
+depends on the manner in which these marking elements are
+_intensified_ and _combined_ by natural selection whether whitish
+longitudinal or oblique stripes should result. In this case then the
+"useful variations" were actually "always there," and we see that in
+the same group of Lepidoptera, e.g. species of Sphingidae, evolution
+has occurred in both directions according to whether the form lived
+among grass or on broad leaves with oblique lateral veins, and we can
+observe even now that the species with oblique stripes have
+longitudinal stripes when young, that is to say, while the stripes
+have no biological significance. The white places in the skin which
+gave rise, probably first as small spots, to this protective marking
+could be combined in one way or another according to the requirements
+of the species. They must therefore either have possessed
+selection-value from the first, or, if this was not the case at their
+earliest occurrence, there must have been _some other factors_ which
+raised them to the point of selection-value. I shall return to this in
+discussing germinal selection. But the case may be followed still
+farther, and leads us to the same alternative on a still more secure
+basis.
+
+Many years ago I observed in caterpillars of _Smerinthus populi_ (the
+poplar hawk-moth), which also possess white oblique stripes, that
+certain individuals showed _red spots_ above these stripes; these
+spots occurred only on certain segments, and never flowed together to
+form continuous stripes. In another species (_Smerinthus tiliae_)
+similar blood-red spots unite to form a line-like coloured seam in the
+last stage of larval life, while in _S. ocellata_ rust-red spots
+appear in individual caterpillars, but more rarely than in _S.
+populi_, and they show no tendency to flow together.
+
+Thus we have here the origin of a new character, arising from small
+beginnings, at least in _S. tiliae_, in which species the coloured
+stripes are a normal specific character. In the other species, _S.
+populi_ and _S. ocellata_, we find the beginnings of the same
+variation, in one more rarely than in the other, and we can imagine
+that, in the course of time, in these two species, coloured lines over
+the oblique stripes will arise. In any case these spots are the
+elements of variation, out of which coloured lines _may_ be evolved,
+if they are combined in this direction through the agency of natural
+selection. In _S. populi_ the spots are often small, but sometimes it
+seems as though several had united to form large spots. Whether a
+process of selection in this direction will arise in _S. populi_ and
+_S. ocellata_, or whether it is now going on cannot be determined,
+since we cannot tell in advance what biological value the marking
+might have for these two species. It is conceivable that the spots may
+have no selection-value as far as these species are concerned, and may
+therefore disappear again in the course of phylogeny, or, on the other
+hand, that they may be changed in another direction, for instance
+towards imitation of the rust-red fungoid patches on poplar and willow
+leaves. In any case we may regard the smallest spots as the initial
+stages of variation, the larger as a cumulative summation of these.
+Therefore either these initial stages must already possess
+selection-value, or, as I said before: _There must be some other
+reason for their cumulative summation_. I should like to give one more
+example, in which we can infer, though we cannot directly observe, the
+initial stages.
+
+All the Holothurians or sea-cucumbers have in the skin calcereous
+bodies of different forms, usually thick and irregular, which make the
+skin tough and resistant. In a small group of them--the species of
+Synapta--the calcareous bodies occur in the form of delicate anchors
+of microscopic size. Up till 1897 these anchors, like many other
+delicate microscopic structures, were regarded as curiosities, as
+natural marvels. But a Swedish observer, Oestergren, has recently
+shown that they have a biological significance: they serve the
+footless Synapta as auxiliary organs of locomotion, since, when the
+body swells up in the act of creeping, they press firmly with their
+tips, which are embedded in the skin, against the substratum on which
+the animal creeps, and thus prevent slipping backwards. In other
+Holothurians this slipping is made impossible by the fixing of the
+tube-feet. The anchors act automatically, sinking their tips towards
+the ground when the corresponding part of the body thickens, and
+returning to the original position at an angle of 45 degrees to the
+upper surface when the part becomes thin again. The arms of the anchor
+do not lie in the same plane as the shaft, and thus the curve of the
+arms forms the outermost part of the anchor, and offers no further
+resistance to the gliding of the animal. Every detail of the anchor,
+the curved portion, the little teeth at the head, the arms, etc., can
+be interpreted in the most beautiful way, above all the form of the
+anchor itself, for the two arms prevent it from swaying round to the
+side. The position of the anchors, too, is definite and significant;
+they lie obliquely to the longitudinal axis of the animal, and
+therefore they act alike whether the animal is creeping backwards or
+forwards. Moreover, the tips would pierce through the skin if the
+anchors lay in the longitudinal direction. Synapta burrows in the
+sand; it first pushes in the thin anterior end, and thickens this
+again, thus enlarging the hole, then the anterior tentacles displace
+more sand, the body is worked in a little farther, and the process
+begins anew. In the first act the anchors are passive, but they begin
+to take an active share in the forward movement when the body is
+contracted again. Frequently the animal retains only the posterior end
+buried in the sand, and then the anchors keep it in position, and make
+rapid withdrawal possible.
+
+Thus we have in these apparently random forms of the calcereous
+bodies, complex adaptations in which every little detail as to
+direction, curve, and pointing is exactly determined. That they have
+selection-value in their present perfected form is beyond all doubt,
+since the animals are enabled by means of them to bore rapidly into
+the ground and so to escape from enemies. We do not know what the
+initial stages were, but we cannot doubt that the little improvements,
+which occurred as variations of the originally simple slimy bodies of
+the Holothurians, were preserved because they already possessed
+selection-value for the Synaptidae. For such minute microscopic
+structures whose form is so delicately adapted to the rôle they have
+to play in the life of the animal, cannot have arisen suddenly and as
+a whole, and every new variation of the anchor, that is, in the
+direction of the development of the two arms, and every curving of the
+shaft which prevented the tips from projecting at the wrong time, in
+short, every little adaptation in the modelling of the anchor must
+have possessed selection-value. And that such minute changes of form
+fall within the sphere of fluctuating variations, that is to say,
+_that they occur_ is beyond all doubt.
+
+In many of the Synaptidae the anchors are replaced by calcareous rods
+bent in the form of an S, which are said to act in the same way.
+Others, such as those of the genus Ankyroderma, have anchors which
+project considerably beyond the skin, and, according to Oestergren,
+serve "to catch plant-particles and other substances" and so mask the
+animal. Thus we see that in the Synaptidae the thick and irregular
+calcareous bodies of the Holothurians have been modified and
+transformed in various ways in adaptation to the footlessness of these
+animals, and to the peculiar conditions of their life, and we must
+conclude that the earlier stages of these changes presented themselves
+to the processes of selection in the form of microscopic variations.
+For it is as impossible to think of any origin other than through
+selection in this case as in the case of the toughness, and the
+"drip-tips" of tropical leaves. And as these last could not have been
+produced directly by the beating of the heavy raindrops upon them, so
+the calcareous anchors of Synapta cannot have been produced directly
+by the friction of the sand and mud at the bottom of the sea, and,
+since they are parts whose function is _passive_ the Lamarckian factor
+of use and disuse does not come into question. The conclusion is
+unavoidable, that the microscopically small variations of the
+calcareous bodies in the ancestral forms have been intensified and
+accumulated in a particular direction, till they have led to the
+formation of the anchor. Whether this has taken place by the action of
+natural selection alone, or whether the laws of variation and the
+intimate processes within the germ-plasm have coöperated will become
+clear in the discussion of germinal selection. This whole process of
+adaptation has obviously taken place within the time that has elapsed
+since this group of sea-cucumbers lost their tube-feet, those
+characteristic organs of locomotion which occur in no group except the
+Echinoderms, and yet have totally disappeared in the Synaptidae. And
+after all what would animals that live in sand and mud do with
+tube-feet?
+
+
+(_c_) _Coadaptation_
+
+Darwin pointed out that one of the essential differences between
+artificial and natural selection lies in the fact that the former can
+modify only a few characters, usually only one at a time, while Nature
+preserves in the struggle for existence all the variations of a
+species, at the same time and in a purely mechanical way, if they
+possess selection-value.
+
+Herbert Spencer, though himself an adherent of the theory of selection,
+declared in the beginning of the nineties that in his opinion the range of
+this principle was greatly over-estimated, if the great changes which have
+taken place in so many organisms in the course of ages are to be
+interpreted as due to this process of selection alone, since no
+transformation of any importance can be evolved by itself; it is always
+accompanied by a host of secondary changes. He gives the familiar example
+of the Giant Stag of the Irish peat, the enormous antlers of which required
+not only a much stronger skull cap, but also greater strength of the
+sinews, muscles, nerves and bones of the whole anterior half of the animal,
+if their mass was not to weigh down the animal altogether. It is
+inconceivable, he says, that so many processes of selection should take
+place _simultaneously_, and we are therefore forced to fall back on the
+Lamarckian factor of the use and disuse of functional parts. And how, he
+asks, could natural selection follow two opposite directions of evolution
+in different parts of the body at the same time, as for instance in the
+case of the kangaroo, in which the forelegs must have become shorter, while
+the hind legs and the tail were becoming longer and stronger?
+
+Spencer's main object was to substantiate the validity of the
+Lamarckian principle, the coöperation of which with selection had been
+doubted by many. And it does seem as though this principle, if it
+operates in nature at all, offers a ready and simple explanation of
+all such secondary variations. Not only muscles, but nerves, bones,
+sinews, in short all tissues which function actively, increase in
+strength in proportion as they are used, and conversely they decrease
+when the claims on them diminish. All the parts, therefore, which
+depend on the part that varied first, as for instance the enlarged
+antlers of the Irish Elk, must have been increased or decreased in
+strength, in exact proportion to the claims made upon them,--just as
+is actually the case.
+
+But beautiful as this explanation would be, I regard it as untenable,
+because it assumes the _transmissibility of functional modifications_
+(so-called "acquired" characters), and this is not only
+undemonstrable, but is scarcely theoretically conceivable, for the
+secondary variations which accompany or follow the first as
+correlative variations, occur also in cases in which the animals
+concerned are sterile and _therefore cannot transmit anything to their
+descendants_. This is true of _worker bees_, and particularly of
+_ants_, and I shall here give a brief survey of the present state of
+the problem as it appears to me.
+
+Much has been written on both sides of this question since the
+published controversy on the subject in the nineties between Herbert
+Spencer and myself. I should like to return to the matter in detail,
+if the space at my disposal permitted, because it seems to me that the
+arguments I advanced at that time are equally cogent to-day,
+notwithstanding all the objections that have since been urged against
+them. Moreover, the matter is by no means one of subordinate interest;
+it is the very kernel of the whole question of the reality and value
+of the principle of selection. For if selection alone does not suffice
+to explain "_harmonious adaptation_" as I have called Spencer's
+_Coadaptation_, and if we require to call in the aid of the Lamarckian
+factor it would be questionable whether selection would explain any
+adaptations whatever. In this particular case--of worker bees--the
+Lamarckian factor may be excluded altogether, for it can be
+demonstrated that here at any rate the effects of use and disuse
+cannot be transmitted.
+
+But if it be asked why we are unwilling to admit the coöperation of
+the Darwinian factor of selection and the Lamarckian factor, since
+this would afford us an easy and satisfactory explanation of the
+phenomena, I answer: _Because the Lamarckian principle is fallacious,
+and because by accepting it we close the way towards deeper insight_.
+It is not a spirit of combativeness or a desire for self-vindication
+that induces me to take the field once more against the Lamarckian
+principle, it is the conviction that the progress of our knowledge is
+being obstructed by the acceptance of this fallacious principle, since
+the facile explanation it apparently affords prevents our seeking
+after a truer explanation and a deeper analysis.
+
+The workers in the various species of ants are sterile, that is to
+say, they take no regular part in the reproduction of the species,
+although individuals among them may occasionally lay eggs. In addition
+to this they have lost the wings, and the _receptaculum seminis_, and
+their compound eyes have degenerated to a few facets. How could this
+last change have come about through disuse, since the eyes of workers
+are exposed to light in the same way as are those of the sexual
+insects and thus in this particular case are not liable to "disuse" at
+all? The same is true of the _receptaculum seminis_, which can only
+have been disused as far as its glandular portion and its stalk are
+concerned, and also of the wings, the nerves tracheae and epidermal
+cells of which could not cease to function until the whole wing had
+degenerated, for the chitinous skeleton of the wing does not function
+at all in the active sense.
+
+But, on the other hand, the workers in all species have undergone
+modifications in a positive direction, as, for instance, the greater
+development of brain. In many species large workers have evolved,--the
+so-called _soldiers_, with enormous jaws and teeth, which defend the
+colony,--and in others there are _small_ workers which have taken over
+other special functions, such as the rearing of the young Aphides.
+This kind of division of the workers into two castes occurs among
+several tropical species of ants, but it is also present in the
+Italian species, _Colobopsis truncata_. Beautifully as the size of the
+jaws could be explained as due to the increased use made of them by
+the "soldiers," or the enlarged brain as due to the mental activities
+of the workers, the fact of the infertility of these forms is an
+insurmountable obstacle to accepting such an explanation. Neither jaws
+nor brain can have been evolved on the Lamarckian principle.
+
+The problem of coadaptation is no easier in the case of the ant than
+in the case of the Giant Stag. Darwin himself gave a pretty
+illustration to show how imposing the difference between the two kinds
+of workers in one species would seem if we translated it into human
+terms. In regard to the Driver ants (Anomma) we must picture to
+ourselves a piece of work, "for instance the building of a house,
+being carried on by two kinds of workers, of which one group was five
+feet four inches high, the other sixteen feet high."[39]
+
+Although the ant is a small animal as compared with man or with the
+Irish Elk, the "soldier" with its relatively enormous jaws is hardly
+less heavily burdened than the Elk with its antlers, and in the ant's
+case, too, a strengthening of the skeleton, of the muscles, the nerves
+of the head, and of the legs must have taken place parallel with the
+enlargement of the jaws. _Harmonious adaptation_ (coadaptation) has
+here been active in a high degree, and yet these "soldiers" are
+sterile! There thus remains nothing for it but to refer all their
+adaptations, positive and negative alike, to processes of selection
+which have taken place in the rudiments of the workers within the egg
+and sperm-cells of their parents. There is no way out of the
+difficulty except the one Darwin pointed out. He himself did not find
+the solution of the riddle at once. At first he believed that the case
+of the workers among social insects presented "the most serious
+special difficulty" in the way of his theory of natural selection; and
+it was only after it had become clear to him that it was not the
+sterile insects themselves but their parents that were selected,
+according as they produced more or less well adapted workers, that he
+was able to refer to this very case of the conditions among ants "_in
+order to show the power of natural selection_."[40] He explains his
+view by a simple but interesting illustration. Gardeners have
+produced, by means of long continued artificial selection, a variety
+of Stock, which bears entirely double, and therefore infertile
+flowers.[41] Nevertheless the variety continues to be reproduced from
+seed, because, in addition to the double and infertile flowers, the
+seeds always produce a certain number of single, fertile blossoms, and
+these are used to reproduce the double variety. These single and
+fertile plants correspond "to the males and females of an ant-colony,
+the infertile plants, which are regularly produced in large numbers,
+to the neuter workers of the colony."
+
+This illustration is entirely apt, the only difference between the
+two cases consisting in the fact that the variation in the flower is
+not a useful, but a disadvantageous one, which can only be preserved
+by artificial selection on the part of the gardener, while the
+transformations that have taken place parallel with the sterility of
+the ants are useful, since they procure for the colony an advantage in
+the struggle for existence, and they are therefore preserved by
+natural selection. Even the sterility itself in this case is not
+disadvantageous, since the fertility of the true females has at the
+same time considerably increased. We may therefore regard the sterile
+forms of ants, which have gradually been adapted in several directions
+to varying functions, _as a certain proof_ that selection really takes
+place in the germ-cells of the fathers and mothers of the workers, and
+that _special complexes of primordia_ (_ids_) are present in the
+workers and in the males and females, and these complexes contain the
+primordia of the individual parts (_determinants_). But since all
+living entities vary, the determinants must also vary, now in a
+favourable, now in an unfavourable direction. If a female produces
+eggs, which contain favourably varying determinants in the worker-ids,
+then these eggs will give rise to workers modified in the favourable
+direction, and if this happens with many females, the colony concerned
+will contain a better kind of worker than other colonies.
+
+I digress here in order to give an account of the intimate processes,
+which, according to my view, take place within the germ-plasm, and
+which I have called "_germinal selection_." These processes are of
+importance since they form the roots of variation, which in its turn
+is the root of natural selection. I cannot here do more than give a
+brief outline of the theory in order to show how the Darwin-Wallace
+theory of selection has gained support from it.
+
+With others, I regard the minimal amount of substance which is
+contained within the nucleus of the germ-cells, in the form of rods,
+bands, or granules, as the _germ-substance_ or _germ-plasm_, and I
+call the individual granules _ids_. There is always a multiplicity of
+such ids present in the nucleus, either occurring individually, or
+united in the form of rods or bands (chromosomes). Each id contains
+the primary constituents of a _whole_ individual, so that several ids
+are concerned in the development of a new individual.
+
+In every being of complex structure thousands of primary constituents
+must go to make up a single id; these I call _determinants_, and I
+mean by this name very small individual particles, far below the
+limits of microscopic visibility, vital units which feed, grow, and
+multiply by division. These determinants control the parts of the
+developing embryo,--in what manner need not here concern us. The
+determinants differ among themselves, those of a muscle are
+differently constituted from those of a nerve-cell or a glandular
+cell, etc., and every determinant is in its turn made up of minute
+vital units, which I call _biophores_, or the bearers of life.
+According to my view, these determinants not only assimilate, like
+every other living unit, but they _vary_ in the course of their
+growth, as every living unit does; they may vary qualitatively if the
+elements of which they are composed vary, they may grow and divide
+more or less rapidly, and their variations give rise to
+_corresponding_ variations of the organ, cell, or cell-group which
+they determine. That they are undergoing ceaseless fluctuations in
+regard to size and quality seems to me the inevitable consequence of
+their unequal nutrition; for although the germ-cell as a whole usually
+receives sufficient nutriment, minute fluctuations in the amount
+carried to different parts within the germ-plasm cannot fail to occur.
+
+Now, if a determinant, for instance of a sensory cell, receives for a
+considerable time more abundant nutriment than before, it will grow
+more rapidly--become bigger, and divide more quickly, and, later, when
+the id concerned develops into an embryo, this sensory cell will
+become stronger than in the parents, possibly even twice as strong.
+This is an instance of a _hereditary individual variation_, arising
+from the germ.
+
+The nutritive stream which, according to our hypothesis, favours the
+determinant _N_ by chance, that is, for reasons unknown to us, may
+remain strong for a considerable time, or may decrease again; but even
+in the latter case it is conceivable that the ascending movement of
+the determinant may continue, because the strengthened determinant now
+_actively_ nourishes itself more abundantly,--that is to say, it
+attracts the nutriment to itself, and to a certain extent withdraws it
+from its fellow-determinants. In this way, it may--as it seems to
+me--get into _permanent upward movement, and attain a degree of
+strength from which there is no falling back_. Then positive or
+negative selection sets in, favouring the variations which are
+advantageous, setting aside those which are disadvantageous.
+
+In a similar manner a _downward_ variation of the determinants may
+take place, if its progress be started by a diminished flow of
+nutriment. The determinants which are weakened by this diminished flow
+will have less affinity for attracting nutriment because of their
+diminished strength, and they will assimilate more feebly and grow
+more slowly, unless chance streams of nutriment help them to recover
+themselves. But, as will presently be shown, a change of direction
+cannot take place at _every_ stage of the degenerative process. If a
+certain critical stage of downward progress be passed, even favourable
+conditions of food-supply will no longer suffice permanently to change
+the direction of the variation. Only two cases are conceivable; if the
+determinant corresponds to a _useful_ organ, only its removal can
+bring back the germ-plasm to its former level; therefore personal
+selection removes the id in question, with its determinants, from the
+germ-plasm, by causing the elimination of the individual in the
+struggle for existence. But there is another conceivable case; the
+determinants concerned may be those of an organ which has become
+_useless_, and they will then continue unobstructed, but with
+exceeding slowness, along the downward path, until the organ becomes
+vestigial, and finally disappears altogether.
+
+The fluctuations of the determinants hither and thither may thus be
+transformed into a lasting ascending or descending movement; and _this
+is the crucial point of these germinal processes_.
+
+This is not a fantastic assumption; we can read it in the fact of the
+degeneration of disused parts. _Useless organs are the only ones which
+are not helped to ascend again by personal selection, and therefore in
+their case alone can we form any idea of how the primary constituents
+behave, when they are subject solely to intra-germinal forces_.
+
+The whole determinant system of an id, as I conceive it, is in a state
+of continual fluctuation upwards and downwards. In most cases the
+fluctuations will counteract one another, because the passive streams
+of nutriment soon change, but in many cases the limit from which a
+return is possible will be passed, and then the determinants concerned
+will continue to vary in the same direction, till they attain positive
+or negative selection-value. At this stage personal selection
+intervenes and sets aside the variation if it is disadvantageous, or
+favours--that is to say, preserves--it if it is advantageous. Only
+_the determinant of a useless organ is uninfluenced by personal
+selection_, and, as experience shows, it sinks downwards; that is, the
+organ that corresponds to it degenerates very slowly but
+uninterruptedly till, after what must obviously be an immense stretch
+of time, it disappears from the germ-plasm altogether.
+
+Thus we find in the fact of the degeneration of disused parts the
+proof that not all the fluctuations of a determinant return to
+equilibrium again, but that, when the movement has attained to a
+certain strength, it continues _in the same direction_. We have entire
+certainty in regard to this as far as the downward progress is
+concerned, and we must assume it also in regard to ascending
+variations, as the phenomena of artificial selection certainly justify
+us in doing. If the Japanese breeders were able to lengthen the
+tail-feathers of the cock to six feet, it can only have been because
+the determinants of the tail-feathers in the germ-plasm had already
+struck out a path of ascending variation, and this movement was taken
+advantage of by the breeder, who continually selected for reproduction
+the individuals in which the ascending variation was most marked. For
+all breeding depends upon the unconscious selection of germinal
+variations.
+
+Of course these germinal processes cannot be proved mathematically,
+since we cannot actually see the play of forces of the passive
+fluctuations and their causes. We cannot say how great these
+fluctuations are, and how quickly or slowly, how regularly or
+irregularly they change. Nor do we know how far a determinant must be
+strengthened by the passive flow of the nutritive stream if it is to
+be beyond the danger of unfavourable variations, or how far it must be
+weakened passively before it loses the power of recovering itself by
+its own strength. It is no more possible to bring forward actual
+proofs in this case than it was in regard to the selection-value of
+the initial stages of an adaptation. But if we consider that all
+heritable variations must have their roots in the germ-plasm, and
+further, that when personal selection does not intervene, that is to
+say, in the case of parts which have become useless, a degeneration of
+the part, and therefore also of its determinant must inevitably take
+place; then we must conclude that processes such as I have assumed are
+running their course within the germ-plasm, and we can do this with as
+much certainty as we were able to infer, from the phenomena of
+adaptation, the selection-value of their initial stages. The fact of
+the degeneration of disused parts seems to me to afford irrefutable
+proof that the fluctuations within the germ-plasm _are the real root
+of all hereditary variation_, and the preliminary condition for the
+occurrence of the Darwin-Wallace factor of selection. Germinal
+selection supplies the stones out of which personal selection builds
+her temples and palaces: _adaptations_. The importance for the theory
+of the process of degeneration of disused parts cannot be
+over-estimated, especially when it occurs in sterile animal forms,
+where we are free from the doubt as to the alleged _Lamarckian factor_
+which is apt to confuse our ideas in regard to other cases.
+
+If we regard the variation of the many determinants concerned in the
+transformation of the female into the sterile worker as having come
+about through the gradual transformation of the ids into worker-ids,
+we shall see that the germ-plasm of the sexual ants must contain three
+kinds of ids, male, female, and worker ids, or if the workers have
+diverged into soldiers and nest-builders, then four kinds. We
+understand that the worker-ids arose because their determinants struck
+out a useful path of variation, whether upward or downward, and that
+they continued in this path until the highest attainable degree of
+utility of the parts determined was reached. But in addition to the
+organs of positive or negative selection-value, there were some which
+were indifferent as far as the success and especially the functional
+capacity of the workers was concerned: wings, ovarian tubes,
+_receptaculum seminis_, a number of the facets of the eye, perhaps
+even the whole eye. As to the ovarian tubes it is is possible that
+their degeneration was an advantage for the workers, in saving energy,
+and if so selection would favour the degeneration; but how could the
+presence of eyes diminish the usefulness of the workers to the colony?
+or the minute _receptaculum seminis_, or even the wings? These parts
+have therefore degenerated _because they were of no further value to
+the insect_. But if selection did not influence the setting aside of
+these parts because they were neither of advantage nor of disadvantage
+to the species, then the Darwinian factor of selection is here
+confronted with a puzzle which it cannot solve alone, but which at
+once becomes clear when germinal selection is added. For the
+determinants of organs that have no further value for the organism,
+must, as we have already explained, embark on a gradual course of
+retrograde development.
+
+In ants the degeneration has gone so far that there are no
+wing-rudiments present in _any_ species, as is the case with so many
+butterflies, flies, and locusts, but in the larvae the imaginable
+discs of the wings are still laid down. With regard to the ovaries,
+degeneration has reached different levels in different species of
+ants, as has been shown by the researches of my former pupil,
+Elizabeth Bickford. In many species there are twelve ovarian tubes,
+and they decrease from that number to one; indeed, in one species no
+ovarian tube at all is present. So much at least is certain from what
+has been said, that in this case _everything_ depends on the
+fluctuations of the elements of the germ-plasm. Germinal selection,
+here as elsewhere, presents the variations of the determinants, and
+personal selection favours or rejects these, or,--if it be a question
+of organs which have become useless,--it does not come into play at
+all, and allows the descending variation free course.
+
+It is obvious that even the problem of _coadaptation in sterile
+animals_ can thus be satisfactorily explained. If the determinants are
+oscillating upwards and downwards in continual fluctuation, and
+varying more pronouncedly now in one direction now in the other,
+useful variations of every determinant will continually present
+themselves anew, and may, in the course of generations, be combined
+with one another in various ways. But there is one character of the
+determinants that greatly facilitates this complex process of
+selection, that, after a certain limit has been reached, they go on
+varying in the same direction. From this it follows that development
+along a path once struck out may proceed without the continual
+intervention of personal selection. This factor only operates, so to
+speak, at the beginning, when it selects the determinants which are
+varying in the right direction, and again at the end, when it is
+necessary to put a check upon further variation. In addition to this,
+enormously long periods have been available for all these adaptations,
+as the very gradual transition stages between females and workers in
+many species plainly show, and thus this process of transformation
+loses the marvellous and mysterious character that seemed at the first
+glance to invest it, and takes rank, without any straining, among the
+other processes of selection. It seems to me that, from the facts that
+sterile animal forms can adapt themselves to new vital functions,
+their superfluous parts degenerate, and the parts more used adapt
+themselves in an ascending direction, those less used in a descending
+direction, we must draw the conclusion that harmonious adaptation here
+comes about _without the coöperation of the Lamarckian principle_.
+This conclusion once established, however, we have no reason to refer
+the thousands of cases of harmonious adaptation, which occur in
+exactly the same way among other animals or plants, to a principle,
+the _active intervention of which in the transformation of species is
+nowhere proved. We do not require it to explain the facts, and
+therefore we must not assume it._
+
+The fact of coadaptation, which was supposed to furnish the strongest
+argument against the principle of selection, in reality yields the
+clearest evidence in favour of it. We _must_ assume it, _because no
+other possibility of explanation is open to us, and because these
+adaptations actually exist, that is to say, have really taken place_.
+With this conviction I attempted, as far back as 1894, when the idea
+of germinal selection had not yet occurred to me, to make "harmonious
+adaptation" (coadaptation) more easily intelligible in some way or
+other, and so I was led to the idea, which was subsequently expounded
+in detail by Baldwin, and Lloyd Morgan, and also by Osborn, and Gulick
+as _Organic Selection_. It seemed to me that it was not necessary that
+all the germinal variations required for secondary variations should
+have occurred _simultaneously_, since, for instance, in the case of
+the stag, the bones, muscles, sinews, and nerves would be incited by
+the increasing heaviness of the antlers to greater activity in _the
+individual life_, and so would be strengthened. The antlers can only
+have increased in size by very slow degrees, so that the muscles and
+bones may have been able to keep pace with their growth in the
+individual life, until the requisite germinal variations presented
+themselves. In this way a disharmony between the increasing weight of
+the antlers and the parts which support and move them would be
+avoided, since time would be given for the appropriate germinal
+variations to occur, and so to set agoing the _hereditary_ variation
+of the muscles, sinews and bones.[42]
+
+I still regard this idea as correct, but I attribute less importance
+to "organic selection" than I did at that time, in so far that I do
+not believe that it _alone_ could effect complex harmonious
+adaptations. Germinal selection now seems to me to play the chief part
+in bringing about such adaptations. Something the same is true of the
+principle I have called _Panmixia_. As I became more and more
+convinced, in the course of years, that the _Lamarckian principle_
+ought not to be called in to explain the dwindling of disused parts, I
+believed that this process might be simply explained as due to the
+cessation of the conservative effect of natural selection. I said to
+myself that, from the moment in which a part ceases to be of use,
+natural selection withdraws its hand from it, and then it must
+inevitably fall from the height of its adaptiveness, because inferior
+variants would have as good a chance of persisting as better ones,
+since all grades of fitness of the part in question would be mingled
+with one another indiscriminately. This is undoubtedly true, as
+Romanes pointed out ten years before I did, and this mingling of the
+bad with the good probably does bring about a deterioration of the
+part concerned. But it cannot account for the steady diminution, which
+always occurs when a part is in process of becoming rudimentary, and
+which goes on until it ultimately disappears altogether. The process
+of dwindling cannot therefore be explained as due to panmixia alone:
+we can only find a sufficient explanation in germinal selection.
+
+
+IV. DERIVATIVES OF THE THEORY OF SELECTION
+
+The impetus in all directions given by Darwin through his theory of
+selection has been an immeasurable one, and its influence is still
+felt. It falls within the province of the historian of science to
+enumerate all the ideas which, in the last quarter of the nineteenth
+century, grew out of Darwin's theories, in the endeavour to penetrate
+more deeply into the problem of the evolution of the organic world.
+Within the narrow limits to which this paper is restricted, I cannot
+attempt to discuss any of these.
+
+
+V. ARGUMENTS FOR THE REALITY OF THE PROCESSES OF SELECTION
+
+
+(_a_) _Sexual Selection_
+
+Sexual selection goes hand in hand with natural selection. From the
+very first I have regarded sexual selection as affording an extremely
+important and interesting corroboration of natural selection, but,
+singularly enough, it is precisely against this theory that an adverse
+judgment has been pronounced in so many quarters, and it is only quite
+recently, and probably in proportion as the wealth of facts in proof
+of it penetrates into a wider circle, that we seem to be approaching a
+more general recognition of this side of the problem of adaptations.
+Thus Darwin's words in his preface to the second edition (1874) of his
+book, _The Descent of Man and Sexual Selection_, are being justified:
+"My conviction as to the operation of natural selection remains
+unshaken," and further, "If naturalists were to become more familiar
+with the idea of sexual selection, it would, I think, be accepted to a
+much greater extent, and already it is fully and favourably accepted
+by many competent judges." Darwin was able to speak thus because he
+was already acquainted with an immense mass of facts, which, taken
+together, yield overwhelming evidence of the validity of the principle
+of sexual selection.
+
+_Natural selection_ chooses out for reproduction the individuals that
+are best equipped for the struggle for existence, and it does so at
+every stage of development; it thus improves the species in all its
+stages and forms. _Sexual selection_ operates only on individuals
+that are already capable of reproduction, and does so only in relation
+to the attainment of reproduction. It arises from the rivalry of one
+sex, usually the male, for the possession of the other, usually the
+female. Its influence can therefore only _directly_ affect one sex, in
+that it equips it better for attaining possession of the other. But
+the effect may extend indirectly to the female sex, and thus the whole
+species may be modified, without, however, becoming any more capable
+of resistance in the struggle for existence, for sexual selection only
+gives rise to adaptations which are likely to give their possessor the
+victory over rivals in the struggle for possession of the female, and
+which are therefore peculiar to the wooing sex: the manifold
+"secondary sexual characters." The diversity of these characters is so
+great that I cannot here attempt to give anything approaching a
+complete treatment of them, but I should like to give a sufficient
+number of examples to make the principle itself, in its various modes
+of expression, quite clear.
+
+One of the chief preliminary postulates of sexual selection is the
+unequal number of individuals in the two sexes, for if every male
+immediately finds his mate there can be no competition for the
+possession of the female. Darwin has shown that, for the most part,
+the inequality between the sexes is due simply to the fact that there
+are more males than females, and therefore the males must take some
+pains to secure a mate. But the inequality does not always depend on
+the numerical preponderance of the males, it is often due to polygamy;
+for, if one male claims several females, the number of females in
+proportion to the rest of the males will be reduced. Since it is
+almost always the males that are the wooers, we must expect to find
+the occurrence of secondary sexual characters chiefly among them, and
+to find it especially frequent in polygamous species. And this is
+actually the case.
+
+If we were to try to guess--without knowing the facts--what means the
+male animals make use of to overcome their rivals in the struggle for
+the possession of the female, we might name many kinds of means, but
+it would be difficult to suggest any which is not actually employed in
+some animal group of other. I begin with the mere difference in
+strength, through which the male of many animals is so sharply
+distinguished from the female, as, for instance, the lion, walrus,
+"sea-elephant," and others. Among these the males fight violently for
+the possession of the female, who falls to the victor in the combat.
+In this simple case no one can doubt the operation of selection, and
+there is just as little room for doubt as to the selection-value of
+the initial stages of the variation. Differences in bodily strength
+are apparent even among human beings, although in their case the
+struggle for the possession of the female is no longer decided by
+bodily strength alone.
+
+Combats between male animals are often violent and obstinate, and the
+employment of the natural weapons of the species in this way has led
+to perfecting of these, e.g. the tusks of the boar, the antlers of the
+stag, and the enormous, antler-like jaws of the stag-beetle. Here
+again it is impossible to doubt that variations in these organs
+presented themselves, and that these were considerable enough to be
+decisive in combat, and so to lead to the improvement of the weapon.
+
+Among many animals, however, the females at first withdraw from the
+males; they are coy, and have to be sought out, and sometimes held by
+force. This tracking and grasping of the females by the males has
+given rise to many different characters in the latter, as, for
+instance, the larger eyes of the male bee, and especially of the males
+of the Ephemerids (May-flies), some species of which show, in addition
+to the usual compound eyes, large, so-called turban-eyes, so that the
+whole head is covered with seeing surfaces. In these species the
+females are very greatly in the minority (1-100), and it is easy to
+understand that a keen competition for them must take place, and that,
+when the insects of both sexes are floating freely in the air, an
+unusually wide range of vision will carry with it a decided
+advantage. Here again the actual adaptations are in accordance with
+the preliminary postulates of the theory. We do not know the stages
+through which the eye has passed to its present perfected state, but,
+since the number of simple eyes (facets) has become very much greater
+in the male than in the female, we may assume that their increase is
+due to a gradual duplication of the determinants of the ommatidium in
+the germ-plasm, as I have already indicated in regard to sense-organs
+in general. In this case, again, the selection-value of the initial
+stages hardly admits of doubt; better vision _directly_ secures
+reproduction.
+
+In many cases _the organ of smell_ shows a similar improvement. Many
+lower Crustaceans (Daphnidae) have better developed organs of smell in
+the male sex. The difference is often slight and amounts only to one
+or two olfactory filaments, but certain species show a difference of
+nearly a hundred of these filaments (Leptodora). The same thing occurs
+among insects.
+
+We must briefly consider the clasping or grasping organs which have
+developed in the males among many lower Crustaceans, but here natural
+selection plays its part along with sexual selection, for the union of
+the sexes is an indispensable condition for the maintenance of the
+species, and as Darwin himself pointed out, in many cases the two
+forms of selection merge into each other. This fact has always seemed
+to me to be a proof of natural selection, for, in regard to sexual
+selection, it is quite obvious that the victory of the best-equipped
+could have brought about the improvement only of the organs concerned,
+the factors in the struggle, such as the eye and the olfactory organ.
+
+We come now to the _excitants_; that is, to the group of sexual
+characters whose origin through processes of selection has been most
+frequently called in question. We may cite the _love-calls_ produced
+by many male insects, such as crickets and cicadas. These could only
+have arisen in animal groups in which the female did not rapidly flee
+from the male, but was inclined to accept his wooing from the first.
+Thus, notes like the chirping of the male cricket serve to entice the
+females. At first they were merely the signal which showed the
+presence of a male in the neighbourhood, and the female was gradually
+enticed nearer and nearer by the continued chirping. The male that
+could make himself heard to the greatest distance would obtain the
+largest following, and would transmit the beginnings, and, later, the
+improvement of his voice to the greatest number of descendants. But
+sexual excitement in the female became associated with the hearing of
+the love-call, and then the sound-producing organ of the male began to
+improve, until it attained to the emission of the long-drawn-out soft
+notes of the mole-cricket or the maenad-like cry of the cicadas. I
+cannot here follow the process of development in detail, but will call
+attention to the fact that the original purpose of the voice, the
+announcing of the male's presence, became subsidiary, and the exciting
+of the female became the chief goal to be aimed at. The loudest
+singers awakened the strongest excitement, and the improvement
+resulted as a matter of course. I conceive of the origin of bird-song
+in a somewhat similar manner, first as a means of enticing, then of
+exciting the female.
+
+One more kind of secondary sexual character must here be mentioned:
+the odour which emanates from so many animals at the breeding season.
+It is possible that this odour also served at first merely to give
+notice of the presence of individuals of the other sex, but it soon
+became an excitant, and as the individuals which caused the greatest
+degree of excitement were preferred, it reached as high a pitch of
+perfection as was possible to it. I shall confine myself here to the
+comparatively recently discovered fragrance of butterflies. Since
+Fritz Müller found out that certain Brazilian butterflies gave off
+fragrance "like a flower," we have become acquainted with many such
+cases, and we now know that in all lands, not only many diurnal
+Lepidoptera but nocturnal ones also give off a delicate odour, which
+is agreeable even to man. The ethereal oil to which this fragrance is
+due is secreted by the skin-cells, usually of the wing, as I showed
+soon after the discovery of the _scent-scales_. This is the case in
+the males; the females have no _special_ scent-scales recognisable as
+such by their form, but they must, nevertheless, give off an extremely
+delicate fragrance, although our imperfect organ of smell cannot
+perceive it, for the males become aware of the presence of a female,
+even at night, from a long distance off, and gather round her. We may
+therefore conclude, that both sexes have long given forth a very
+delicate perfume, which announced their presence to others of the same
+species, and that in many species (_not in all_) these small
+beginnings become, in the males, particularly strong scent-scales of
+characteristic form (lute, brush, or lyre-shaped). At first these
+scales were scattered over the surface of the wing, but gradually they
+concentrated themselves, and formed broad, velvety bands, or strong,
+prominent brushes, and they attained their highest pitch of evolution
+when they became enclosed within pits or folds of the skin, which
+could be opened to let the delicious fragrance stream forth suddenly
+towards the female. Thus in this case also we see that characters, the
+original use of which was to bring the sexes together, and so to
+maintain the species, have been evolved in the males into means for
+exciting the female. And we can hardly doubt, that the females are
+most readily enticed to yield to the butterfly that sends out the
+strongest fragrance,--that is to say, that excites them to the highest
+degree. It is a pity that our organs of smell are not fine enough to
+examine the fragrance of male Lepidoptera in general, and to compare
+it with other perfumes which attract these insects.[43] As far as we
+can perceive them they resemble the fragrance of flowers, but there
+are Lepidoptera whose scent suggests musk. A smell of musk is also
+given off by several plants: it is a sexual excitant in the
+musk-deer, the musk-sheep, and the crocodile.
+
+As far as we know, then, it is perfumes similar to those of flowers
+that the male Lepidoptera give off in order to entice their mates and
+this is a further indication that animals, like plants, can to a large
+extent meet the claims made upon them by life, and produce the
+adaptations which are most purposive,--a further proof, too, of my
+proposition that the useful variations, so to speak, are _always
+there_. The flowers developed the perfumes which entice their
+visitors, and the male Lepidoptera developed the perfumes which entice
+and excite their mates.
+
+There are many pretty little problems to be solved in this connection,
+for there are insects, such as some flies, that are attracted by
+smells which are unpleasant to us, like those from decaying flesh and
+carrion. But there are also certain flowers, some orchids for
+instance, which give forth no very agreeable odour, but one which is
+to us repulsive and disgusting; and we should therefore expect that
+the males of such insects would give off a smell unpleasant to us, but
+there is no case known to me in which this has been demonstrated.
+
+In cases such as we have discussed, it is obvious that there is no
+possible explanation except through selection. This brings us to the
+last kind of secondary sexual characters, and the one in regard to
+which doubt has been most frequently expressed,--decorative colours
+and decorative forms, the brilliant plumage of the male pheasant, the
+humming-birds, and the bird of Paradise, as well as the bright colours
+of many species of butterfly, from the beautiful blue of our little
+Lycaenidae to the magnificent azure of the large Morphinae of Brazil.
+In a great many cases, though not by any means in all, the male
+butterflies are "more beautiful" than the females, and in the Tropics
+in particular they shine and glow in the most superb colours. I really
+see no reason why we should doubt the power of sexual selection, and I
+myself stand wholly on Darwin's side. Even though we certainly cannot
+assume that the females exercise a conscious choice of the
+"handsomest" mate, and deliberate like the judges in a court of
+justice over the perfections of their wooers, we have no reason to
+doubt that distinctive forms (decorative feathers), and colours have a
+particularly exciting effect upon the female, just as certain odours
+have among animals of so many different groups, including the
+butterflies. The doubts which existed for a considerable time, as a
+result of fallacious experiments, as to whether the colours of flowers
+really had any influence in attracting butterflies have now been set
+at rest through a series of more careful investigations; we now know
+that the colours of flowers are there on account of the butterflies,
+as Sprengel first showed, and that the blossoms of Phanerogams are
+selected in relation to them, as Darwin pointed out.
+
+Certainly it is not possible to bring forward any convincing proof of
+the origin of decorative colours through sexual selection, but there
+are many weighty arguments in favour of it, and these form a body of
+presumptive evidence so strong that it almost amounts to certainty.
+
+In the first place, there is the analogy with other secondary sexual
+characters. If the song of birds and the chirping of the cricket have
+been evolved through sexual selection, if the penetrating odours of
+male animals,--the crocodile, the musk-deer, the beaver, the
+carnivores, and, finally, the flower-like fragrances of the
+butterflies have been evolved to their present pitch in this way, why
+should decorative colours have arisen in some other way? Why should
+the eye be less sensitive to _specifically male_ colours and other
+_visible_ signs _enticing to the female_, than the olfactory sense to
+specifically male odours, or the sense of hearing to specifically male
+sounds? Moreover, the decorative feathers of birds are almost always
+spread out and displayed before the female during courtship. I have
+elsewhere[44] pointed out that decorative colouring and
+sweet-scentedness may replace one another in Lepidoptera as well as in
+flowers, for just as some modestly coloured flowers (mignonette and
+violet) have often a strong perfume, while strikingly coloured ones
+are sometimes quite devoid of fragrance, so we find that the most
+beautiful and gaily-coloured of our native Lepidoptera, the species of
+Vanessa, have no scent-scales, while these are often markedly
+developed in grey nocturnal Lepidoptera. Both attractions may,
+however, be combined in butterflies, just as in flowers. Of course, we
+cannot explain why both means of attraction should exist in one genus,
+and only one of them in another, since we do not know the minutest
+details of the conditions of life of the genera concerned. But from
+the sporadic distribution of scent-scales in Lepidoptera, and from
+their occurrence or absence in nearly related species, we may conclude
+that fragrance is a relatively _modern_ acquirement, more recent than
+brilliant colouring.
+
+One thing in particular that stamps decorative colouring as a product
+of selection is _its gradual intensification_ by the addition of new
+spots, which we can quite well observe, because in many cases the
+colours have been first acquired by the males, and later transmitted
+to the females by inheritance. The scent-scales are never thus
+transmitted, probably for the same reason that the decorative colours
+of many birds are often not transmitted to the females: because with
+these they would be exposed to too great elimination by enemies.
+Wallace was the first to point out that in species with concealed
+nests the beautiful feathers of the male occurred in the female also,
+as in the parrots, for instance, but this is not the case in species
+which brood on an exposed nest. In the parrots one can often observe
+that the general brilliant colouring of the male is found in the
+female, but that certain spots of colour are absent, and these have
+probably been acquired comparatively recently by the male and have not
+yet been transmitted to the female.
+
+Isolation of the group of individuals which is in process of varying
+is undoubtedly of great value in sexual selection, for even a solitary
+conspicuous variation will become dominant much sooner in a small
+isolated colony, than among a large number of members of a species.
+
+Any one who agrees with me in deriving variations from germinal
+selection will regard that process as an essential aid towards
+explaining the selection of distinctive courtship-characters, such as
+coloured spots, decorative feathers, horny outgrowths in birds and
+reptiles, combs, feather-tufts, and the like, since the beginnings of
+these would be presented with relative frequency in the struggle
+between the determinants within the germ-plasm. The process of
+transmission of decorative feathers to the female results, as Darwin
+pointed out and illustrated by interesting examples, in the
+_colour-transformation of a whole species_, and this process, as the
+phyletically older colouring of young birds shows, must, in the course
+of thousands of years, have repeated itself several times in a line of
+descent.
+
+If we survey the wealth of phenomena presented to us by secondary
+sexual characters, we can hardly fail to be convinced of the truth of
+the principle of sexual selection. And certainly no one who has
+accepted natural selection should reject sexual selection, for, not
+only do the two processes rest upon the same basis, but they merge
+into one another, so that it is often impossible to say how much of a
+particular character depends on one and how much on the other form of
+selection.
+
+
+(_b_) _Natural Selection_
+
+An actual proof of the theory of sexual selection is out of the
+question, if only because we cannot tell when a variation attains to
+selection-value. It is certain that a delicate sense of smell is of
+value to the male moth in his search for the female, but whether the
+possession of one additional olfactory hair, or of ten, or of twenty
+additional hairs leads to the success of its possessor we are unable
+to tell. And we are groping even more in the dark when we discuss the
+excitement caused in the female by agreeable perfumes, or by striking
+and beautiful colours. That these do make an impression is beyond
+doubt; but we can only assume that slight intensifications of them
+give any advantage, and we _must_ assume this _since otherwise
+secondary sexual characters remain inexplicable_.
+
+The same thing is true in regard to natural selection. It is not
+possible to bring forward any actual proof of the selection-value of
+the initial stages, and the stages in the increase of variations, as
+has been already shown. But the selection-value of a finished
+adaptation can in many cases be statistically determined. Cesnola and
+Poulton have made valuable experiments in this direction. The former
+attached forty-five individuals of the green, and sixty-five of the
+brown variety of the praying mantis (_Mantis religiosa_), by a silk
+thread to plants, and watched them for seven days. The insects which
+were on a surface of a colour Similar to their own remained uneaten,
+while twenty-five green insects on brown parts of plants had all
+disappeared in eleven days.
+
+The experiments of Poulton and Sanders[45] were made with 600 pupae of
+_Vanessa urticae_, the "tortoise-shell butterfly." The pupae were
+artificially attached to nettles, tree-trunks, fences, walls, and to
+the ground, some at Oxford, some at St. Helens in the Isle of Wight.
+In the course of a month 93% of the pupae at Oxford were killed,
+chiefly by small birds, while at St. Helens 68% perished. The
+experiments showed very clearly that the colour and character of the
+surface on which the pupa rests--and thus its own conspicuousness--are
+of the greatest importance. At Oxford only the four pupae which were
+fastened to nettles emerged; all the rest--on bark, stones and the
+like--perished. At St. Helens the elimination was as follows: on
+fences where the pupae were conspicuous, 92%; on bark, 66%; on walls,
+54%; and among nettles, 57%. These interesting experiments confirm our
+views as to protective coloration, and show further, _that the ratio
+of elimination in the species is a very high one, and that therefore
+selection must be very keen_.
+
+We may say that the process of selection follows as a logical
+necessity from the fulfilment of the three preliminary postulates of
+the theory: variability, heredity, and the struggle for existence,
+with its enormous ratio of elimination in all species. To this we must
+add a fourth factor, the _intensification_ of variations which Darwin
+established as a fact, and which we are now able to account for
+theoretically on the basis of germinal selection. It may be objected
+that there is considerable uncertainty about this _logical_ proof,
+because of our inability to demonstrate the selection-value of the
+initial stages and the individual stages of increase. We have
+therefore to fall back on _presumptive evidence_. This is to be found
+in _the interpretative value of the theory_. Let us consider this
+point in greater detail.
+
+In the first place it is necessary to emphasize what is often
+overlooked, namely, that the theory not only explains the
+_transformations_ of species, it also explains _their remaining the
+same_; in addition to the principle of varying, it contains within
+itself that of _persisting_. It is part of the essence of selection,
+that it not only causes a part to _vary_ till it has reached its
+highest pitch of adaptation, but that it _maintains it at this pitch.
+This conserving influence of natural selection_ is of great
+importance, and was early recognised by Darwin; it follows naturally
+from the principle of the survival of the fittest.
+
+We understand from this how it is that a species which has become
+fully adapted to certain conditions of life ceases to vary, but
+remains "constant," as long as the conditions of life _for_ it remain
+unchanged, whether this be for thousands of years, or for whole
+geological epochs. But the most convincing proof of the power of the
+principle of selection lies in the innumerable multitude of phenomena
+which cannot be explained in any other way. To this category belong
+all structures which are only _passively_ of advantage to the
+organism, because none of these can have arisen by the alleged
+_Lamarckian principle_. These have been so often discussed that we
+need do no more than indicate them here. Until quite recently the
+sympathetic coloration of animals--for instance, the whiteness of
+Arctic animals--was referred, at least in part, to the _direct_
+influence of external factors, but the facts can best be explained by
+referring them to the processes of selection, for then it is
+unnecessary to make the gratuitous assumption that many species are
+sensitive to the stimulus of cold and that others are not. The great
+majority of Arctic land-animals, mammals and birds, are white, and
+this proves that they were all able to present the variation which was
+most useful for them. The sable is brown, but it lives in trees, where
+the brown colouring protects and conceals it more effectively. The
+musk-sheep (_Ovibos moschatus_) is also brown, and contrasts sharply
+with the ice and snow, but it is protected from beasts of prey by its
+gregarious habit, and therefore it is of advantage to be visible from
+as great a distance as possible. That so many species have been able
+to give rise to white varieties does not depend on a special
+sensitiveness of the skin to the influence of cold, but to the fact
+that Mammals and Birds have a general tendency to vary towards white.
+Even with us, many birds--starlings, blackbirds, swallows,
+etc.--occasionally produce white individuals, but the white variety
+does not persist, because it readily falls a victim to the carnivores.
+This is true of white fawns, foxes, deer, etc. The whiteness,
+therefore, arises from internal causes, and only persists when it is
+useful. A great many animals living in a _green environment_ have
+become clothed in green, especially insects, caterpillars, and
+Mantidae, both persecuted and persecutors.
+
+That it is not the direct effect of the environment which calls forth
+the green colour is shown by the many kinds of caterpillar which rest
+on leaves and feed on them, but are nevertheless brown. These feed by
+night and betake themselves through the day to the trunk of the tree,
+and hide in the furrows of the bark. We cannot, however, conclude
+from this that they were _unable_ to vary towards green, for there are
+Arctic animals which are white only in winter and brown in summer
+(Alpine hare, and the ptarmigan of the Alps), and there are also green
+leaf-insects which remain green only while they are young and
+difficult to see on the leaf, but which become brown again in the last
+stage of larval life, when they have outgrown the leaf. They then
+conceal themselves by day, sometimes only among withered leaves on the
+ground, sometimes in the earth itself. It is interesting that in one
+genus, Chaerocampa, one species is brown in the last stage of larval
+life, another becomes brown earlier, and in many species the last
+stage is not wholly brown, a part remaining green. Whether this is a
+case of a double adaptation, or whether the green is being gradually
+crowded out by the brown, the fact remains that the same species, even
+the same individual, can exhibit both variations. The case is the same
+with many of the leaf-like Orthoptera, as, for instance, the praying
+mantis (_Mantis religiosa_) which we have already mentioned.
+
+But the best proofs are furnished by those of ten-cited cases in which
+the insect bears a deceptive resemblance to another object. We now
+know many such cases, such as the numerous imitations of green or
+withered leaves, which are brought about in the most diverse ways,
+sometimes by mere variations in the form of the insect and in its
+colour, sometimes by an elaborate marking, like that which occurs in
+the Indian leaf-butterflies, _Kallima inachis_. In the single
+butterfly-genus Anaea, in the woods of South America, there are about
+a hundred species which are all gaily coloured on the upper surface,
+and on the reverse side exhibit the most delicate imitation of the
+colouring and pattern of a leaf, generally without any indication of
+the leaf-ribs, but extremely deceptive nevertheless. Anyone who has
+seen only one such butterfly may doubt whether many of the
+insignificant details of the marking can really be of advantage to the
+insect. Such details are for instance the apparent holes and splits
+in the apparently dry or half-rotten leaf, which are usually due to
+the fact that the scales are absent on a circular or oval patch so
+that the colourless wing-membrane lies bare, and one can look through
+the spot as through a window. Whether the bird which is seeking or
+pursuing the butterflies takes these holes for dewdrops, or for the
+work of a devouring insect, does not affect the question; the
+mirror-like spot undoubtedly increases the general deceptiveness, for
+the same thing occurs in many leaf-butterflies, though not in all, and
+in some cases it is replaced in quite a peculiar manner. In one
+species of Anaea (_A. divina_), the resting butterfly looks exactly
+like a leaf out of the outer edge of which a large semi-circular piece
+has been eaten, possibly by a caterpillar; but if we look more closely
+it is obvious that there is no part of the wing absent, and that the
+semi-circular piece is of a clear, pale yellow colour, while the rest
+of the wing is of a strongly contrasted dark brown.
+
+But the deceptive resemblance may be caused in quite a different
+manner. I have often speculated as to what advantage the brilliant
+white C could give to the otherwise dusky-coloured "Comma butterfly"
+(_Grapta C. album_). Poulton's recent observations[46] have shown that
+this represents the imitation of a crack such as is often seen in dry
+leaves, and is very conspicuous because the light shines through it.
+
+The utility obviously lies in presenting to the bird the very familiar
+picture of a broken leaf with a clear shining slit, and we may
+conclude, from the imitation of such small details, that the birds are
+very sharp observers and that the smallest deviation from the usual
+arrests their attention and incites them to closer investigation. It
+is obvious that such detailed--we might almost say such
+subtle--deceptive resemblances could only have come about in the
+course of long ages through the acquirement from time to time of
+something new which heightened the already existing resemblance.
+
+In face of facts like these there can be no question of chance and no
+one has succeeded so far in finding any other explanation to replace
+that by selection. For the rest, the apparent leaves are by no means
+perfect copies of a leaf; many of them only represent the torn or
+broken piece, or the half or two-thirds of a leaf, but then the leaves
+themselves frequently do not present themselves to the eye as a whole,
+but partially concealed among other leaves. Even those butterflies
+which, like the species of Kallima and Anaea, represent the whole of a
+leaf with stalk, ribs, apex, and the whole breadth, are not actual
+copies which would satisfy a botanist; there is often much wanting. In
+Kallima the lateral ribs of the leaf are never all included in the
+markings; there are only two or three on the left side and at more
+four or five on the right, and in many individuals these are rather
+obscure, while in others they are comparatively distinct. This
+furnishes us with fresh evidence in favour of their origin through
+processes of selection, for a botanically perfect picture could not
+arise in this way; there could only be a fixing of such details as
+heightened the deceptive resemblance.
+
+Our postulate of origin through selection also enables us to
+understand why the leaf-imitation is on the lower surface of the wing
+in the diurnal Lepidoptera, and on the upper surface in the nocturnal
+forms, corresponding to the attitude of the wings in the resting
+position of the two groups.
+
+The strongest of all proofs of the theory, however, is afforded by
+cases of true "mimicry," those adaptations discovered by Bates in
+1861, consisting in the imitation of one species by another, which
+becomes more and more like its model. The model is always a species
+that enjoys some special protection from enemies, whether because it
+is unpleasant to taste, or because it is in some way dangerous.
+
+It is chiefly among insects and especially among butterflies that we
+find the greatest number of such cases. Several of these have been
+minutely studied and every detail has been investigated so that it is
+difficult to understand how there can still be disbelief in regard to
+them. If the many and exact observations which have been carefully
+collected and critically discussed for instance by Poulton[47] were
+thoroughly studied the arguments which are still frequently urged
+against mimicry would be found untenable; we can hardly hope to find
+more convincing proof of the actuality of the processes of selection
+than these cases put into our hands. The preliminary postulates of the
+theory of mimicry have been disputed, for instance, that diurnal
+butterflies are persecuted and eaten by birds, but observations
+specially directed towards this point in India, Africa, America and
+Europe have placed it beyond all doubt. If it were necessary I could
+myself furnish an account of my own observations on this point.
+
+In the same way it has been established by experiment and observation
+in the field that in all the great regions of distribution there are
+butterflies which are rejected by birds and lizards, their chief
+enemies, on account of their unpleasant smell or taste. These
+butterflies are usually gaily and conspicuously coloured and thus--as
+Wallace first interpreted it--are furnished with an easily
+recognisable sign: a sign of unpalatableness or _warning colours_. If
+they were not thus recognisable easily and from a distance, they would
+frequently be pecked at by birds, and then rejected because of their
+unpleasant taste; but as it is, the insect-eaters recognise them at
+once as unpalatable booty and ignore them. Such _immune_[48] species,
+wherever they occur, are imitated by other palatable species, which
+thus acquire a certain degree of protection.
+
+It is true that this explanation of the bright, conspicuous colours
+is only a hypothesis, but its foundations--unpalatableness, and the
+liability of other butterflies to be eaten,--are certain, and its
+consequences--the existence of mimetic palatable forms--conform it in
+the most convincing manner. Of the many cases now known I select one,
+which is especially remarkable, and which has been thoroughly
+investigated, _Papilla dardanus_ (_merope_), a large, beautiful,
+diurnal butterfly which ranges from Abyssinia throughout the whole of
+Africa to the south coast of Cape Colony.
+
+The males of this form are everywhere _almost_ the same in colour and
+in form of wings, save for a few variations in the sparse black
+markings on the pale yellow ground. But the females occur in several
+quite different forms and colourings, and one of these only, the
+Abyssinian form, is like the male, while the other three or four are
+_mimetic_, that is to say, they copy a butterfly of quite a different
+family the Danaids, which are among the _immune_ forms. In each region
+the females have thus copied two or three different immune species.
+There is much that is interesting to be said in regard to these
+species, but it would be out of keeping with the general tenor of this
+paper to give details of this very complicated case of polymorphism in
+_P. Dardanus_. Anyone who is interested in the matter will find a full
+and exact statement of the case in as far as we know it, in Poulton's
+_Essays on Evolution_ (pp. 373-375[49]). I need only add that three
+different mimetic female forms have been reared from the eggs of a
+single female in South Africa. The resemblance of the forms to their
+immune models goes so far that even the details of the _local_ forms
+of the models are copied by the mimetic species.
+
+It remains to be said that in Madagascar a butterfly,
+
+_Papilio meriones_, occurs, of which both sexes are very similar in
+form and markings to the non-mimetic male of _P. dardanus_, so that it
+probably represents the ancestor of this latter species.
+
+In face of such facts as these every attempt at another explanation
+must fail. Similarly all the other details of the case fulfil the
+preliminary postulates of selection, and leave no room for any other
+interpretation. That the males do not take on the protective colouring
+is easily explained, because they are in general more numerous, and
+the females are more important for the preservation of the species,
+and must also live longer in order to deposit their eggs. We find the
+same state of things in many other species, and in one case (_Elymnias
+undularis_) in which the male is also mimetically coloured, it copies
+quite a differently coloured immune species from the model followed by
+the female. This is quite intelligible when we consider that if there
+were _too many_ false immune types, the birds would soon discover that
+there were palatable individuals among those with unpalatable warning
+colours. Hence the imitation of different immune species by _Papilio
+dardanus_!
+
+I regret that lack of space prevents my bringing forward more examples
+of mimicry and discussing them fully. But from the case of _Papilio
+dardanus_ alone there is much to be learnt which is of the highest
+importance for our understanding of transformations. It shows us
+chiefly what I once called, somewhat strongly perhaps, _the
+omnipotence of natural selection_ in answer to an opponent who had
+spoken of its "inadequacy." We here see that one and the same species
+is capable of producing four or five different patterns of colouring
+and marking; thus the colouring and marking are not, as has often been
+supposed, a necessary outcome of the specific nature of the species,
+but a true adaptation, which cannot arise as a direct effect of
+climatic conditions, but solely through what I may call the sorting
+out of the variations produced by the species, according to their
+utility. That caterpillars may be either green or brown is already
+something more than could have been expected according to the old
+conception of species, but that one and the same butterfly should be
+now pale yellow, with black; now red with black and pure white; now
+deep black with large, pure white spots; and again black with a large
+ocheous-yellow spot, and many small white and yellow spots; that in
+one sub-species it may be tailed like the ancestral form, and in
+another tailless like its Danaid model,--all this shows a far-reaching
+capacity for variation and adaptation that we could never have
+expected if we did not see the facts before us. How it is possible
+that the primary colour-variations should thus be intensified and
+combined remains a puzzle even now; we are reminded of the modern
+three-colour printing,--perhaps similar combinations of the primary
+colours take place in this case; in any case the direction of these
+primary variations is determined by the artist whom we know as natural
+selection, for there is no other conceivable way in which the model
+could affect the butterfly that is becoming more and more like it. The
+same climate surrounds all four forms of female; they are subject to
+the same conditions of nutrition. Moreover, _Papilio dardanus_ is by
+no means the only species of butterfly which exhibits different kinds
+of colour-pattern on its wings. Many species of the Asiatic genus
+Elymnias have on the upper surface a very good imitation of an immune
+Euploeine (Danainae), often with a steel-blue ground-colour, while the
+under surface is well concealed when the butterfly is at rest,--thus
+there are two kinds of protective coloration each with a different
+meaning! The same thing may be observed in many non-mimetic
+butterflies, for instance in all our species of Vanessa, in which the
+under side shows a grey-brown or brownish-black protective coloration,
+but we do not yet know with certainty what may be the biological
+significance of the gaily coloured upper surface.
+
+In general it may be said that mimetic butterflies are comparatively
+rare species, but there are exceptions, for instance _Limenitis
+archippus_ in North America, of which the immune model (_Danaida
+plexippus_) also occurs in enormous numbers.
+
+In another mimicry-category the imitators are often more numerous than
+the models, namely in the case of the imitation of _dangerous insects_
+by harmless species. Bees and wasps are dreaded for their sting, and
+they are copied by harmless flies of the genera Eristalis and Syrphus,
+and these mimics often occur in swarms about flowering plants without
+damage to themselves or to their models; they are feared and are
+therefore left unmolested.
+
+In regard also to the _faithfulness of the copy_ the facts are quite
+in harmony with the theory, according to which the resemblance must
+have arisen and increased _by degrees_. We can recognise this in many
+cases, for even now the mimetic species show very _varying degrees of
+resemblance_ to their immune model. If we compare, for instance, the
+many different imitators of _Danaida chrysippus_ we find that, with
+their brownish-yellow ground-colour, and the position and size, and
+more or less sharp limitation of their clear marginal spots, they have
+reached very different degrees of nearness to their model. Or compare
+the female of _Elymnias undularis_ with its model _Danaida genutia_;
+there is a general resemblance, but the marking of the Danaida is very
+roughly imitated in Elymnias.
+
+Another fact that bears out the theory of mimicry is, that even when
+the resemblance in colour-pattern is very great, the _wing-venation_,
+which is so constant, and so important in determining the systematic
+position of butterflies, is never affected by the variation. The
+pursuers of the butterfly have no time to trouble about entomological
+intricacies.
+
+I must not pass over a discovery of Poulton's which is of great
+theoretical importance--that mimetic butterflies may reach the same
+effect by very different means.[50] Thus the glass-like transparency
+of the wing of a certain Ithomiine (Methona) and its Pierine mimic
+(_Dismorphia orise_) depends on a diminution in the size of the
+scales; in the Danaine genus Itune it is due to the fewness of the
+scales and in a third imitator, a moth (_Castnia linus var.
+heliconoides_) the glass-like appearance of the wing is due neither to
+diminution nor to absence of scales, but to their absolute
+colourlessness and transparency, and to the fact that they stand
+upright. In another moth mimic (Anthomyza) the arrangement of the
+transparent scales is normal. Thus it is not some unknown external
+influence that has brought about the transparency of the wing in these
+five forms, as has sometimes been supposed. Nor is it a hypothetical
+_internal_ evolutionary tendency, for all three vary in a different
+manner. The cause of this agreement can only lie in selection, which
+preserves and intensifies in each species the favourable variations
+that present themselves. The great faithfulness of the copy is
+astonishing in these cases, for it is not _the whole_ wing which is
+transparent; certain markings are black in colour, and these contrast
+sharply with the glass-like ground. It is obvious that the pursuers of
+these butterflies must be very sharp-sighted, for otherwise the
+agreement between the species could never have been pushed so far. The
+less the enemies see and observe, the more defective must the
+imitation be, and if they had been blind, no visible resemblance
+between the species which required protection could ever have arisen.
+
+A seemingly irreconcilable contradiction to the mimicry theory is
+presented in the following cases, which were known to Bates, who,
+however, never succeeded in bringing them into line with the principle
+of mimicry.
+
+In South America there are, as we have already said, many mimics of
+the immune Ithomiinae (or as Bates called them Heliconidae). Among
+these there occur not merely species which are edible, and thus
+require the protection of a disguise, but others which are rejected on
+account of their unpalatableness. How could the Ithomiine dress have
+developed in their case, and of what use is it, since the species
+would in any case be immune? In Eastern Brazil, for instance, there
+are four butterflies, which bear a most confusing resemblance to one
+another in colour, marking, and form of wing, and all four are
+unpalatable to birds. They belong to four different genera and three
+sub-families, and we have to inquire: Whence came this resemblance and
+what end does it serve? For a long time no satisfactory answer could
+be found, but Fritz Müller,[51] seventeen years after Bates, offered a
+solution to the riddle, when he pointed out that young birds could not
+have an instinctive knowledge of the unpalatableness of the
+Ithomiines, but must learn by experience which species were edible and
+which inedible. Thus each young bird must have tasted at least one
+individual of each inedible species and discovered its unpalatability,
+before it learnt to avoid, and thus to spare the species. But if the
+four species resemble each other very closely the bird will regard
+them all as of the same kind, and avoid them all. Thus there developed
+a process of selection which resulted in the survival of the
+Ithomiine-like individuals, and in so great an increase of resemblance
+between the four species, that they are difficult to distinguish one
+from another even in a collection. The advantage for the four species,
+living side by side as they do e.g. in Bahia, lies in the fact that
+only one individual from the _mimicry-ring_ ("inedible association")
+need be tasted by a young bird, instead of at least four individuals,
+as would otherwise be the case. As the number of young birds is great,
+this makes a considerable difference in the ratio of elimination. The
+four Brazilian species are _Lycorea halia_ (Danainae), _Heliconius
+narcaea_ (_eucrate_) (Heliconinae), _Melinaea ethra_, and _Mechanitis
+lysimnia_ (Ithomiinae).
+
+These interesting mimicry-rings (trusts), which have much significance
+for the theory, have been the subject of numerous and careful
+investigations, and at least their essential features are now fully
+established. Müller took for granted, without making any
+investigations, that young birds only learn by experience to
+distinguish between different kinds of victims. But Lloyd Morgan's[52]
+experiments with young birds proved that this is really the case, and
+at the same time furnished an additional argument against the
+_Lamarckian principle_.
+
+In addition to the mimicry-rings first observed in South America,
+others have been described from Tropical India by Moore, and by
+Poulton and Dixey from Africa, and we may expect to learn many more
+interesting facts in this connection. Here again the preliminary
+postulates of the theory are satisfied. And how much more that would
+lead to the same conclusion might be added!
+
+As in the case of mimicry many species have come to resemble one
+another through processes of selection, so we know whole classes of
+phenomena in which plants and animals have become adapted to one
+another, and have thus been modified to a considerable degree. I refer
+particularly to the relation between flowers and insects. Darwin has
+shown that the originally inconspicuous blossoms of the phanerogams
+were transformed into flowers through the visits of insects, and that,
+conversely, several large orders of insects have been gradually
+modified by their association with flowers, especially as regards the
+parts of their body actively concerned. Bees and butterflies in
+particular have become what they are through their relation to
+flowers. In this case again all that is apparently contradictory to
+the theory can, on closer investigation, be beautifully interpreted in
+corroboration of it. Selection can give rise only to what is of use to
+the organism actually concerned, never to what is of use to some other
+organism, and we must therefore expect to find that in flowers only
+characters of use to _themselves_ have arisen, never characters which
+are of use to insects only, and conversely that in the insects
+characters useful to them and not merely to the plants would have
+originated. For a long time it seemed as if an exception to this rule
+existed in the case of the fertilisation of the yucca blossoms by a
+little moth, _Pronuba yuccasella_. This little moth has a
+sickle-shaped appendage to its mouth-parts which occurs hi no other
+Lepidopteron, and which is used for pushing the yellow pollen into the
+opening of the pistil, thus fertilising the flower. Thus it appears as
+if a new structure, which is useful only to the plant, has arisen in
+the insect. But the difficulty is solved as soon as we learn that the
+moth lays its eggs in the fruit-buds of the Yucca, and that the
+larvae, when they emerge, feed on the developing seeds. In effecting
+the fertilisation of the flower the moth is at the same time making
+provision for its own offspring, since it is only after fertilisation
+that the seeds begin to develop. There is thus nothing to prevent our
+referring this structural adaptation in _Pronuba yuccasella_ to
+processes of selection, which have gradually transformed the maxillary
+palps of the female into the sickle-shaped instrument for collecting
+the pollen, and which have at the same time developed in the insect
+the instinct to press the pollen into the pistil.
+
+In this domain, then, the theory of selection finds nothing but
+corroboration, and it would be impossible to substitute for it any
+other explanation, which now that the facts are so well known, could
+be regarded as a serious rival to it. That selection is a factor, and
+a very powerful factor in the evolution of organisms, can no longer be
+doubted. Even although we cannot bring forward formal proofs of it _in
+detail_, cannot calculate definitely the size of the variations which
+present themselves, and their selection-value, cannot, in short,
+reduce the whole process to a mathematical formula, yet we must assume
+selection, because it is the only possible explanation applicable to
+whole classes of phenomena, and because, on the other hand, it is made
+up of factors which we know can be proved actually to exist, and
+which, _if_ they exist, must of logical necessity coöperate in the
+manner required by the theory. _We must accept it because the
+phenomena of evolution and adaptation must have a natural basis, and
+because it is the only possible explanation of them._[53]
+
+Many people are willing to admit that selection explains adaptations,
+but they maintain that only a part of the phenomena are thus
+explained, because everything does not depend upon adaptation. They
+regard adaptation as, so to speak, a special effort on the part of
+Nature, which she keeps in readiness to meet particularly difficult
+claims of the external world on organisms. But if we look at the
+matter more carefully we shall find that adaptations are by no means
+exceptional, but that they are present everywhere in such enormous
+numbers, that it would be difficult in regard to any structure
+whatever, to prove that adaptation had _not_ played a part in its
+evolution.
+
+How often has the senseless objection been urged against selection
+that it can create nothing, it can only reject. It is true that it
+cannot create either the living substance or the variations of it;
+both must be given. But in rejecting one thing it preserves another,
+intensifies it, combines it, and in this way _creates_ what is new.
+_Everything_ in organisms depends on adaptation; that is to say,
+everything must be admitted through the narrow door of selection,
+otherwise it can take no part in the building up of the whole. But, it
+is asked, what of the direct effect of external conditions,
+temperature, nutrition, climate and the like? Undoubtedly these can
+give rise to variations, but they too must pass through the door of
+selection, and if they cannot do this they are rejected, eliminated
+from the constitution of the species.
+
+It may, perhaps, be objected that such external influences are often
+of a compelling power, and that every animal must submit to them, and
+that thus selection has no choice and can neither select nor reject.
+There may be such cases; let us assume for instance that the effect
+of the cold of the Arctic regions was to make all the mammals become
+black; the result would be that they would all be eliminated by
+selection, and that no mammals would be able to live there at all. But
+in most cases a certain percentage of animals resists these strong
+influences, and thus selection secures a foothold on which to work,
+eliminating the unfavourable variation, and establishing a useful
+colouring, consistent with what is required for the maintenance of the
+species.
+
+Everything depends upon adaptation! We have spoken much of adaptation
+in colouring, in connection with the examples brought into prominence
+by Darwin, because these are conspicuous, easily verified, and at the
+same time convincing for the theory of selection. But is it only
+desert and polar animals whose colouring is determined through
+adaptation? Or the leaf-butterflies, and the mimetic species, or the
+terrifying markings, and "warning-colours" and a thousand other kinds
+of sympathetic colouring? It is, indeed, never the colouring alone
+which makes up the adaptation; the structure of the animal plays a
+part, often a very essential part, in the protective disguise, and
+thus _many_ variations may cooperate towards _one_ common end. And it
+is to be noted that it is by no means only external parts that are
+changed; internal parts are _always_ modified at the same time--for
+instance, the delicate elements of the nervous system on which depend
+the _instinct_ of the insect to hold its wings, when at rest, in a
+perfectly definite position, which, in the leaf-butterfly, has the
+effect of bringing the two pieces on which the marking occurs on the
+anterior and posterior wing into the same direction, and thus
+displaying as a whole the fine curve of the midrib on the seeming
+leaf. But the wing-holding instinct is not regulated in the same way
+in all leaf-butterflies; even our indigenous species of Vanessa, with
+their protective ground-colouring, have quite a distinctive way of
+holding their wings so that the greater part of the anterior wing is
+covered by the posterior when the butterfly is at rest. But the
+protective colouring appears on the posterior wing and on the tip of
+the anterior, _to precisely the distance to which it is left
+uncovered_. This occurs, as Standfuss has shown, in different degrees
+in our two most nearly allied species, the uncovered portion being
+smaller in _V. urticae_ than in _V. polychloros_. In this case, as in
+most leaf-butterflies, the holding of the wing was probably the
+primary character; only after that was thoroughly established did the
+protective marking develop. In any case, the instinctive manner of
+holding the wings is associated with the protective colouring, and
+must remain as it is if the latter is to be effective. How greatly
+instincts may change, that is to say, may be adapted, is shown by the
+case of the Noctuid "shark" moth, _Xylina vetusta_. This form bears a
+most deceptive resemblance to a piece of rotten wood, and the
+appearance is greatly increased by the modification of the innate
+impulse to flight common to so many animals, which has here been
+transformed into an almost contrary instinct. This moth does not fly
+away from danger, but "feigns death," that is, it draws antennae, legs
+and wings close to the body, and remains perfectly motionless. It may
+be touched, picked up, and thrown down again, and still it does not
+move. This remarkable instinct must surely have developed
+simultaneously with the wood-colouring; at all events, both
+coöperating variations are now present, and prove that both the
+external and the most minute internal structure have undergone a
+process of adaptation.
+
+The case is the same with all structural variations of animal parts,
+which are not absolutely insignificant. When the insects acquired
+wings they must also have acquired the mechanism with which to move
+them--the musculature, and the nervous apparatus necessary for its
+automatic regulation. All instincts depend upon compound reflex
+mechanisms and are just as indispensable as the parts they have to set
+in motion, and all may have arisen through processes of selection if
+the reasons which I have elsewhere given for this view are
+correct.[54]
+
+Thus there is no lack of adaptations within the organism, and
+particularly in its most important and complicated parts, so that we
+may say that there is no actively functional organ that has not
+undergone a process of adaptation relative to its function and the
+requirements of the organism. Not only is every gland structurally
+adapted, down to the very minutest histological details, to its
+function, but the function is equally minutely adapted to the needs of
+the body. Every cell in the mucous lining of the intestine is exactly
+regulated in its relation to the different nutritive substances, and
+behaves in quite a different way towards the fats, and towards
+nitrogenous substances, or peptones.
+
+I have elsewhere called attention to the many adaptations of the whale
+to the surrounding medium, and have pointed out--what has long been
+known, but is not universally admitted, even now--that in it a great
+number of important organs have been transformed in adaptation to the
+peculiar conditions of aquatic life, although the ancestors of the
+whale must have lived, like other hair-covered mammals, on land. I
+cited a number of these transformations--the fish-like form of the
+body, the hairlessness of the skin, the transformation of the
+fore-limbs to fins, the disappearance of the hind-limbs and the
+development of a tail fin, the layer of blubber under the skin, which
+affords the protection from cold necessary to a warm-blooded animal,
+the disappearance of the ear-muscles and the auditory passages, the
+displacement of the external nares to the forehead for the greater
+security of the breathing-hole during the brief appearance at the
+surface, and certain remarkable changes in the respiratory and
+circulatory organs which enable the animal to remain for a long time
+under water. I might have added many more, for the list of adaptations
+in the whale to aquatic life is by no means exhausted; they are found
+in the histological structure and in the minutest combinations in the
+nervous system. For it is obvious that a tail-fin must be used in
+quite a different way from a tail, which serves as a fly-brush in
+hoofed animals, or as an aid to springing in the kangaroo or as a
+climbing organ; it will require quite different reflex-mechanisms and
+nerve combinations in the motor centres.
+
+I used this example in order to show how unnecessary it is to assume a
+special internal evolutionary power for the phylogenesis of species,
+for this whole order of whales is, so to speak, _made up of
+adaptations_; it deviates in many essential respects from the usual
+mammalian type, and all the deviations are adaptations to aquatic
+life. But if precisely the most essential features of the organisation
+thus depend upon adaptation, what is left for a phyletic force to do,
+since it is these essential features of the structure it would have to
+determine? There are few people now who believe in a phyletic
+evolutionary power, which is not made up of the forces known to
+us--adaptation and heredity--but the conviction that _every_ part of
+an organism depends upon adaptation has not yet gained a firm footing.
+Nevertheless, I must continue to regard this conception as the correct
+one, as I have long done.
+
+I may be permitted one more example. The feather of a bird is a
+marvellous structure, and no one will deny that as a whole it depends
+upon adaptation. But what part of it _does not_ depend upon
+adaptation? The hollow quill, the shaft with its hard, thin, light
+cortex, and the spongy substance within it, its square section
+compared with the round section of the quill, the flat barbs, their
+short, hooked barbules which, in the flight-feathers, hook into one
+another with just sufficient firmness to resist the pressure of the
+air at each wing-beat, the lightness and firmness of the whole
+apparatus, the elasticity of the vane, and so on. And yet all this
+belongs to an organ which is only passively functional, and therefore
+can have nothing to do with the _Lamarckian principle_. Nor can the
+feather have arisen through some magical effect of temperature,
+moisture, electricity, or specific nutrition, and thus selection is
+again our only anchor of safety.
+
+But--it will be objected--the substance of which the feather consists,
+this peculiar kind of horny substance, did not first arise through
+selection in the course of the evolution of the birds, for it formed
+the covering of the scales of their reptilian ancestors. It is quite
+true that a similar substance covered the scales of the Reptiles, but
+why should it not have arisen among them through selection? Or in what
+other way could it have arisen, since scales are also passively useful
+parts? It is true that if we are only to call adaptation what has been
+acquired by the species we happen to be considering, there would
+remain a great deal that could not be referred to selection; but we
+are postulating an evolution which has stretched back through aeons,
+and in the course of which innumerable adaptations took place, which
+had not merely ephemeral persistence in a genus, a family or a class,
+but which was continued into whole Phyla of animals, with continual
+fresh adaptations to the special conditions of each species, family,
+or class, yet with persistence of the fundamental elements. Thus the
+feather, once acquired, persisted in all birds, and the vertebral
+column, once gained by adaptation in the lowest forms, has persisted
+in all the Vertebrates from Amphioxus upwards, although with constant
+readaptation to the conditions of each particular group. Thus
+everything we can see in animals is adaptation, whether of to-day, or
+of yesterday, or of ages long gone by; every kind of cell, whether
+glandular, muscular, nervous, epidermic, or skeletal, is adapted to
+absolutely definite and specific functions, and every organ which is
+composed of these different kinds of cells contains them in the proper
+proportions, and in the particular arrangement which best serves the
+function of the organ; it is thus adapted to its function.
+
+All parts of the organism are tuned to one another, that is, _they are
+adapted to one another_, and in the same way _the organism as a whole
+is adapted to the conditions of its life, and it is so at every stage
+of its evolution._
+
+But all adaptations _can_ be referred to selection; the only point
+that remains doubtful is whether they all _must_ be referred to it.
+
+However that may be, whether the _Lamarckian principle_ is a factor
+that has coöperated with selection in evolution, or whether it is
+altogether fallacious, the fact remains, that selection is the cause
+of a great part of the phyletic evolution of organisms on our earth.
+Those who agree with me in rejecting the _Lamarckian principle_ will
+regard selection as the only _guiding_ factor in evolution, which
+creates what is new out of the transmissible variations, by ordering
+and arranging these, selecting them in relation to their number and
+size, as the architect does his building-stones so that a particular
+style must result.[55] But the building-stones themselves, the
+variations, have their basis in the influences which cause variation
+in those vital units which are handed on from one generation to
+another, whether, taken together they form the _whole_ organism, as in
+Bacteria and other low forms of life, or only a germ-substance, as in
+unicellular and multicellular organisms.
+
+FOOTNOTES:
+
+[Footnote 33: _Vorträge über Descendenztheorie_, Jena, 1904, II. 269.
+Eng. Transl. London, 1904, II. p. 317.]
+
+[Footnote 34: See Poulton, _Essays on Evolution_, Oxford, 1908. pp.
+xix-xxii.]
+
+[Footnote 35: _Origin of Species_ (6th edit), pp. 176 _et seq._]
+
+[Footnote 36: Chun, _Reise der Valdivia_, Leipzig, 1904.]
+
+[Footnote 37: Plate, _Selektionsprinzip u. Probleme der Artbildung_
+(3rd edit.), Leipzig, 1908.]
+
+[Footnote 38: _Studien zur Descendenz-Theorie_ II., "Die Enstehung der
+Zeichnung bei den Schmetterlings-raupen," Leipzig, 1876.]
+
+[Footnote 39: _Origin of Species_ (6th edit.), p. 232.]
+
+[Footnote 40: _Origin of Species_, p. 233; see also edit. 1, p. 242.]
+
+[Footnote 41: _Ibid._ p. 230.]
+
+[Footnote 42: _The Effect of External Influences upon Development_,
+Romanes Lecture, Oxford, 1894.]
+
+[Footnote 43: See Poulton, _Essays on Evolution_, 1908, pp. 316, 317.]
+
+[Footnote 44: _The Evolution Theory_, London, 1904, I. p. 219.]
+
+[Footnote 45: _Report of the British Association_ (Bristol, 1898),
+London, 1899, pp. 906-909.]
+
+[Footnote 46: _Proc. Ent. Soc._, London, May 6, 1903.]
+
+[Footnote 47: _Essays on Evolution_, 1889-1907, Oxford, 1908,
+_passim_, e.g. p. 269.]
+
+[Footnote 48: The expression does not refer to all the enemies of this
+butterfly; against ichneumon-flies, for instance, their unpleasant
+smell usually gives no protection.]
+
+[Footnote 49: Professor Poulton has corrected some wrong descriptions
+which I had unfortunately overlooked in the Plates of my book
+_Vorträge über Descendenztheorie_, and which refer to _Papilio
+dardanus_ (_merope_). These mistakes are of no importance as far as an
+understanding of the mimicry-theory is concerned, but I hope shortly
+to be able to correct them in a later edition.]
+
+[Footnote 50: _Journ. Linn. Soc. London_ (_Zool._), Vol. xxvi. 1898,
+pp. 598-602.]
+
+[Footnote 51: In _Kosmos_, 1879, p. 100.]
+
+[Footnote 52: _Habit and Instinct_, London. 1896.]
+
+[Footnote 53: This has been discussed in many of my earlier works. See
+for instance _The All-Sufficiency of Natural Selection, a reply to
+Herbert Spencer_, London, 1893.]
+
+[Footnote 54: _The Evolution Theory_, London, 1904, p. 144.]
+
+[Footnote 55: _Variation under Domestication_, 1875, II. pp. 426,
+427.]
+
+
+
+
+III
+
+HEREDITY AND VARIATION IN MODERN LIGHTS
+
+BY W. BATESON, M.A., F.R.S.
+
+_Professor of Biology in the University of Cambridge_
+
+
+Darwin's work has the property of greatness in that it may be admired
+from more aspects than one. For some the perception of the principle
+of Natural Selection stands out as his most wonderful achievement to
+which all the rest is subordinate. Others, among whom I would range
+myself, look up to him rather as the first who plainly distinguished,
+collected, and comprehensively studied that new class of evidence from
+which hereafter a true understanding of the process of Evolution may
+be developed. We each prefer our own standpoint of admiration; but I
+think that it will be in their wider aspect that his labours will most
+command the veneration of posterity.
+
+A treatise written to advance knowledge may be read in two moods. The
+reader may keep his mind passive, willing merely to receive the
+impress of the writer's thought; or he may read with his attention
+strained and alert, asking at every instant how the new knowledge can
+be used in a further advance, watching continually for fresh footholds
+by which to climb higher still. Of Shelley it has been said that he
+was a poet for poets: so Darwin was a naturalist for naturalists. It
+is when his writings are used in the critical and more exacting spirit
+with which we test the outfit for our own enterprise that we learn
+their full value and strength. Whether we glance back and compare his
+performance with the efforts of his predecessors, or look forward
+along the course which modern research is disclosing, we shall honour
+most in him not the rounded merit of finite accomplishment, but the
+creative power by which he inaugurated a line of discovery endless in
+variety and extension. Let us attempt thus to see his work in true
+perspective between the past from which it grew, and the present which
+is its consequence. Darwin attacked the problem of Evolution by
+reference to facts of three classes: Variation; Heredity; Natural
+Selection. His work was not as the laity suppose, a sudden and
+unheralded revelation, but the first fruit of a long and hitherto
+barren controversy. The occurrence of variation from type, and the
+hereditary transmission of such variation had of course been long
+familiar to practical men, and inferences as to the possible bearing
+of those phenomena on the nature of specific difference had been from
+time to time drawn by naturalists. Maupertuis, for example, wrote: "Ce
+qui nous reste à examiner, c'est comment d'un seul individu, il a pu
+naître tant d'espèces si différentes." And again: "La Nature contient
+le fonds de toutes ces variétés: mais le hasard ou l'art les mettent
+en œuvre. C'est ainsi que ceux dont l'industrie s'applique à
+satisfaire le goût des curieux, sont, pour ainsi dire, créateurs
+d'espèces nouvelles."[56]
+
+Such passages, of which many (though few so emphatic) can be found in
+eighteenth century writers, indicate a true perception of the mode of
+Evolution. The speculations hinted at by Buffon,[57] developed by
+Erasmus Darwin, and independently proclaimed above all by Lamarck,
+gave to the doctrine of descent a wide renown. The uniformitarian
+teaching which Lyell deduced from geological observation had gained
+acceptance. The facts of geographical distribution[58] had been shown
+to be obviously inconsistent with the Mosaic legend. Prichard, and
+Lawrence, following the example of Blumenbach, had successfully
+demonstrated that the races of Man could be regarded as different
+forms of one species, contrary to the opinion up till then received.
+These treatises all begin, it is true, with a profound obeisance to
+the sons of Noah, but that performed, they continue on strictly modern
+lines. The question of the mutability of species was thus prominently
+raised.
+
+Those who rate Lamarck no higher than did Huxley in his contemptuous
+phrase "_buccinator tantum_," will scarcely deny that the sound of the
+trumpet had carried far, or that its note was clear. If then there
+were few who had already turned to evolution with positive conviction,
+all scientific men must at least have known that such views had been
+promulgated; and many must, as Huxley says, have taken up his own
+position of "critical expectancy."[59]
+
+Why, then, was it, that Darwin succeeded where the rest had failed?
+The cause of that success was twofold. First, and obviously, in the
+principle of Natural Selection he had a suggestion which would work.
+It might not go the whole way, but it was true as far as it went.
+Evolution could thus in great measure be fairly represented as a
+consequence of demonstrable processes. Darwin seldom endangers the
+mechanism he devised by putting on it strains much greater than it can
+bear. He at least was under no illusion as to the omnipotence of
+Selection; and he introduces none of the forced pleading which in
+recent years has threatened to discredit that principle.
+
+
+
+For example, in the latest text of the _Origin_[60] we find him
+saying:
+
+     "But as my conclusions have lately been much misrepresented,
+     and it has been stated that I attribute the modification of
+     species exclusively to natural selection, I may be permitted
+     to remark that in the first edition of this work, and
+     subsequently, I placed in a most conspicuous
+     position--namely, at the close of the Introduction--the
+     following words: 'I am convinced that natural selection has
+     been the main but not the exclusive means of modification.'"
+
+But apart from the invention of this reasonable hypothesis, which may
+well, as Huxley estimated, "be the guide of biological and
+psychological speculation for the next three or four generations,"
+Darwin made a more significant and imperishable contribution. Not for
+a few generations, but through all ages he should be remembered as the
+first who showed clearly that the problems of Heredity and Variation
+are soluble by observation, and laid down the course by which we must
+proceed to their solution.[61] The moment of inspiration did not come
+with the reading of Malthus, but with the opening of the "first
+note-book on Transmutation of Species."[62] Evolution is a process of
+Variation and Heredity. The older writers, though they had some vague
+idea that it must be so, did not study Variation and Heredity. Darwin
+did, and so begat not a theory, but a science.
+
+The extent to which this is true, the scientific world is only
+beginning to realise. So little was the fact appreciated in Darwin's
+own time that the success of his writings was followed by an almost
+total cessation of work in that special field. Of the causes which led
+to these remarkable consequences I have spoken elsewhere. They
+proceeded from circumstances peculiar to the time; but whatever the
+causes there is no doubt that this statement of the result is
+historically exact, and those who make it their business to collect
+facts elucidating the physiology of Heredity and Variation are well
+aware that they will find little to reward their quest in the leading
+scientific Journals of the Darwinian epoch.
+
+In those thirty years the original stock of evidence current and in
+circulation even underwent a process of attrition. As in the story of
+the Eastern sage who first wrote the collected learning of the
+universe for his sons in a thousand volumes and by successive
+compression and burning reduced them to one and from this by further
+burning distilled the single ejaculation of the Faith "There is no god
+but God and Mohammed is the Prophet of God," which was all his maturer
+wisdom deemed essential:--so in the books of that period do we find
+the _corpus_ of genetic knowledge dwindle to a few prerogative
+instances and these at last to the brief formula of an unquestioned
+creed.
+
+And yet in all else that concerns biological science this period was,
+in very truth, our Golden Age, when the natural history of the earth
+was explored as never before; morphology and embryology were
+exhaustively ransacked; the physiology of plants and animals began to
+rival chemistry and physics in precision of method and in the rapidity
+of its advances; and the foundations of pathology were laid.
+
+In contrast with this immense activity elsewhere the neglect which
+befel the special physiology of Descent, or Genetics as we now call
+it, is astonishing. This may of course be interpreted as meaning that
+the favoured studies seemed to promise a quicker return for effort,
+but it would be more true to say that those who chose these other
+pursuits did so without making any such comparison; for the idea that
+the physiology of Heredity and Variation was a coherent science,
+offering possibilities of extraordinary discovery, was not present to
+their minds at all. In a word, the existence of such a science was
+well nigh forgotten. It is true that in ancillary periodicals, as for
+example those that treat of entomology or horticulture, or in the
+writings of the already isolated systematists,[63] observations with
+this special bearing were from time to time related, but the class of
+fact on which Darwin built his conceptions of Heredity and Variation
+was not seen in the highways of biology. It formed no part of the
+official curriculum of biological students, and found no place among
+the subjects which their teachers were investigating.
+
+During this period nevertheless one distinct advance was made, that
+with which Weismann's name is prominently connected. In Darwin's
+genetic scheme the hereditary transmission of parental experience and
+its consequences played a considerable role. Exactly how great that
+role was supposed to be, he with his habitual caution refrained from
+specifying, for the sufficient reason that he did not know.
+Nevertheless much of the process of Evolution, especially that by
+which organs have become degenerate and rudimentary, was certainly
+attributed by Darwin to such inheritance, though since belief in the
+inheritance of acquired characters fell into dispute, the fact has
+been a good deal overlooked. The _Origin_ without "use and disuse"
+would be a materially different book. A certain vacillation is
+discernible in Darwin's utterances on this question, and the fact gave
+to the astute Butler an opportunity for his most telling attack. The
+discussion which best illustrates the genetic views of the period
+arose in regard to the production of the rudimentary condition of the
+wings of many beetles in the Madeira group of islands, and by
+comparing passages from the _Origin_[64] Butler convicts Darwin of
+saying first that this condition was in the main the result of
+Selection, with disuse aiding, and in another place that the main
+cause of degeneration was disuse, but that Selection had aided. To
+Darwin however I think the point would have seemed one of dialetics
+merely. To him the one paramount purpose was to show that somehow an
+Evolution by means of Variation and Heredity might have brought about
+the facts observed, and whether they had come to pass in the one way
+or the other was a matter of subordinate concern.
+
+To us moderns the question at issue has a diminished significance. For
+over all such debates a change has been brought by Weismann's
+challenge for evidence that use and disuse have any transmitted
+effects at all. Hitherto the transmission of many acquired
+characteristics had seemed to most naturalists so obvious as not to
+call for demonstration.[65] Weismann's demand for facts in support of
+the main proposition revealed at once that none having real cogency
+could be produced. The time-honoured examples were easily shown to be
+capable of different explanations. A few certainly remain which cannot
+be so summarily dismissed, but--though it is manifestly impossible
+here to do justice to such a subject--I think no one will dispute that
+these residual and doubtful phenomena, whatever be their true nature,
+are not of a kind to help us much in the interpretation of any of
+those complex cases of adaptation which on the hypothesis of unguided
+Natural Selection are especially difficult to understand. Use and
+disuse were invoked expressly to help us over these hard places; but
+whatever changes can be induced in offspring by direct treatment of
+the parents, they are not of a kind to encourage hope of real
+assistance from that quarter. It is not to be denied that through the
+collapse of this second line of argument the Selection hypothesis has
+had to take an increased and perilous burden. Various ways of meeting
+the difficulty have been proposed, but these mostly resolve themselves
+into improbable attempts to expand or magnify the powers of Natural
+Selection.
+
+Weismann's interpellation, though negative in purpose, has had a
+lasting and beneficial effect, for through his thorough demolition of
+the old loose and distracting notions of inherited experience, the
+ground has been cleared for the construction of a true knowledge of
+heredity based on experimental fact.
+
+In another way he made a contribution of a more positive character,
+for his elaborate speculations as to the genetic meaning of
+cytological appearances have led to a minute investigation of the
+visible phenomena occurring in those cell-divisions by which
+germ-cells arise. Though the particular views he advocated have very
+largely proved incompatible with the observed facts of heredity, yet
+we must acknowledge that it was chiefly through the stimulus of
+Weismann's ideas that those advances in cytology were made; and though
+the doctrine of the continuity of germ-plasm cannot be maintained in
+the form originally propounded, it is in the main true and
+illuminating.[66] Nevertheless in the present state of knowledge we
+are still as a rule quite unable to connect cytological appearances
+with any genetic consequences and save in one respect (obviously of
+extreme importance--to be spoken of later) the two sets of phenomena
+might, for all we can see, be entirely distinct.
+
+I cannot avoid attaching importance to this want of connection between
+the nuclear phenomena and the features of bodily organisation. All
+attempts to investigate Heredity by cytological means lie under the
+disadvantage that it is the nuclear changes which can alone be
+effectively observed. Important as they must surely be, I have never
+been persuaded that the rest of the cell counts for nothing. What we
+know of the behaviour and variability of chromosomes seems in my
+opinion quite incompatible with the belief that they alone govern
+form, and are the sole agents responsible in heredity.[67]
+
+If, then, progress was to be made in Genetics, work of a different
+kind was required. To learn the laws of Heredity and Variation there
+is no other way than that which Darwin himself followed, the direct
+examination of the phenomena. A beginning could be made by collecting
+fortuitous observations of this class, which have often thrown a
+suggestive light, but such evidence can be at best but superficial and
+some more penetrating instrument of research is required. This can
+only be provided by actual experiments in breeding.
+
+The truth of these general considerations was becoming gradually clear
+to many of us when in 1900 Mendel's work was rediscovered.
+Segregation, a phenomenon of the utmost novelty, was thus revealed.
+From that moment not only in the problem of the origin of species, but
+in all the great problems of biology a new era began. So unexpected
+was the discovery that many naturalists were convinced it was untrue,
+and at once proclaimed Mendel's conclusions as either altogether
+mistaken, or if true, of very limited application. Many fantastic
+notions about the workings of Heredity had been asserted as general
+principles before: this was probably only another fancy of the same
+class.
+
+Nevertheless those who had a preliminary acquaintance with the facts
+of Variation were not wholly unprepared for some such revelation. The
+essential deduction from the discovery of segregation was that the
+characters of living things are dependent on the presence of definite
+elements or factors, which are treated as units in the processes of
+Heredity. These factors can thus be recombined in various ways. They
+act sometimes separately, and sometimes they interact in conduction
+with each other, producing their various effects. All this indicates a
+definiteness and specific order in heredity, and therefore in
+variation. This order cannot by the nature of the case be dependent on
+Natural Selection for its existence, but must be a consequence of the
+fundamental chemical and physical nature of living things. The study
+of Variation had from the first shown that an orderliness of this kind
+was present. The bodies and the properties of livings things are
+cosmic, not chaotic. No matter how low in the scale we go, never do we
+find the slightest hint of a diminution in that all-pervading
+orderliness, nor can we conceive an organism existing for a moment in
+any other state. Moreover not only does this order prevail in normal
+forms, but again and again it is to be seen in newly-sprung varieties,
+which by general consent cannot have been subjected to a prolonged
+Selection. The discovery of Mendelian elements admirably coincided
+with and at once gave a rationale of these facts. Genetic Variation is
+then primarily the consequence of additions to, or omissions from, the
+stock of elements which the species contains. The further
+investigation of the species-problem must thus proceed by the
+analytical method which breeding experiments provide.
+
+In the nine years which have elapsed since Mendel's clue became
+generally known, progress has been rapid. We now understand the
+process by which a polymorphic race maintains its polymorphism. When a
+family consists of dissimilar members, given the numerical proportions
+in which these members are occurring, we can represent their
+composition symbolically and state what types can be transmitted by
+the various members. The difficulty of the "swamping effects of
+inter-crossing" is practically at an end. Even the famous puzzle of
+sex-limited inheritance is solved, at all events in its more regular
+manifestations, and we know now how it is brought about that the
+normal sisters of a colour-blind man can transmit the colour-blindness
+while his normal brothers cannot transmit it.
+
+We are still only on the fringe of the inquiry. It can be seen
+extending and ramifying in many directions. To enumerate these here
+would be impossible. A whole new range of possibilities is being
+brought into view by study of the inter-relations between the simple
+factors. By following up the evidence as to segregation, indications
+have been obtained which can only be interpreted as meaning that when
+many factors are being simultaneously redistributed among the
+germ-cells, certain of them exert what must be described as a
+repulsion upon other factors. We cannot surmise whither this discovery
+may lead.
+
+In the new light all the old problems wear a fresh aspect. Upon the
+question of the nature of Sex, for example, the bearing of Mendelian
+evidence is close. Elsewhere I have shown that from several sets of
+parallel experiments the conclusion is almost forced upon us that, in
+the types investigated, of the two sexes the female is to be regarded
+as heterozygous in sex, containing one unpaired dominant element,
+while the male is similarly homozygous in the absence of that
+element.[68] It is not a little remarkable that on this point--which
+is the only one where observations of the nuclear processes of
+gameto-genesis have yet been brought into relation with the visible
+characteristics of the organisms themselves--there should be
+diametrical opposition between the results of breeding experiments and
+those derived from cytology.
+
+Those who have followed the researches of the American school will be
+aware that, after it had been found in certain insects that the
+spermatozoa were of two kinds according as they contained or did not
+contain the accessory chromosome, E. B. Wilson succeeded in proving
+that the sperms possessing this accessory body were destined to form
+_females_ on fertilisation, while sperms without it form males, the
+eggs being apparently indifferent. Perhaps the most striking of all
+this series of observations is that lately made by T. H. Morgan,[69]
+since confirmed by von Baehr, that in a Phylloxeran two kinds of
+spermatids are formed, respectively with and without an accessory (in
+this case, _double_) chromosome. Of these, only those possessing the
+accessory body become functional spermatozoa, the others degenerating.
+We have thus an elucidation of the puzzling fact that in these forms
+fertilisation results in the formation of _females_ only. How the
+males are formed--for of course males are eventually produced by the
+parthenogenetic females--we do not know.
+
+If the accessory body is really to be regarded as bearing the factor
+for femaleness, then in Mendelian terms female is DD and male is DR.
+The eggs are indifferent and the spermatozoa are each male, _or_
+female. But according to the evidence derived from a study of the
+sex-limited descent of certain features in other animals the
+conclusion seems equally clear that in them female must be regarded as
+DR and male as RR. The eggs are thus each either male or female and
+the spermatozoa are indifferent. How this contradictory evidence is to
+be reconciled we do not yet know. The breeding work concerns fowls,
+canaries, and the Currant moth (_Abraxas grossulariata_). The
+accessory chromosome has been now observed in most of the great
+divisions of insects,[70] except, as it happens, Lepidoptera. At first
+sight it seems difficult to suppose that a feature apparently so
+fundamental as sex should be differently constituted in different
+animals, but that seems at present the least improbable inference. I
+mention these two groups of facts as illustrating the nature and
+methods of modern genetic work. We must proceed by minute and specific
+analytical investigation. Wherever we look we find traces of the
+operation of precise and specific rules.
+
+In the light of present knowledge it is evident that before we can
+attack the Species-problem with any hope of success there are vast
+arrears to be made up. He would be a bold man who would now assert
+that there was no sense in which the term Species might not have a
+strict and concrete meaning in contradistinction to the term Variety.
+We have been taught to regard the difference between species and
+variety as one of degree. I think it unlikely that this conclusion
+will bear the test of further research. To Darwin the question, What
+is a variation? presented no difficulties. Any difference between
+parent and offspring was a variation. Now we have to be more precise.
+First we must, as de Vries has shown, distinguish real, genetic,
+variation from _fluctuational_ variations, due to environmental and
+other accidents, which cannot be transmitted. Having excluded these
+sources of error the variations observed must be expressed in terms of
+the factors to which they are due before their significance can be
+understood. For example, numbers of the variations seen under
+domestication, and not a few witnessed in nature, are simply the
+consequence of some ingredient being in an unknown way omitted from
+the composition of the varying individual. The variation may on the
+contrary be due to the addition of some new element, but to prove that
+it is so is by no means an easy matter. Casual observation is useless,
+for though these latter variations will always be dominants, yet many
+dominant characteristics may arise from another cause, namely the
+meeting of complementary factors, and special study of each case in
+two generations at least is needed before these two phenomena can be
+distinguished.
+
+When such considerations are fully appreciated it will be realised
+that medleys of most dissimilar occurrences are all confused together
+under the term Variation. One of the first objects of genetic analysis
+is to disentangle this mass of confusion.
+
+To those who have made no study of heredity it sometimes appears that
+the question of the effect of conditions in causing variation is one
+which we should immediately investigate, but a little thought will
+show that before any critical inquiry into such possibilities can be
+attempted, a knowledge of the working of heredity under conditions as
+far as possible uniform must be obtained. At the time when Darwin was
+writing, if a plant brought into cultivation gave off an albino
+variety, such an event was without hesitation ascribed to the change
+of life. Now we see that albino _gametes_, germs, that is to say,
+which are destitute of the pigment-forming factor, may have been
+originally produced by individuals standing an indefinite number of
+generations back in the ancestry of the actual albino, and it is
+indeed almost certain that the variation to which the appearance of
+the albino is due cannot have taken place in a generation later than
+that of the grandparents. It is true that when a new _dominant_
+appears we should feel greater confidence that we were witnessing the
+original variation, but such events are of extreme rarity, and no such
+case has come under the notice of an experimenter in modern times, as
+far as I am aware. That they must have appeared is clear enough.
+Nothing corresponding to the Brown-breasted Game fowl is known wild,
+yet that colour is a most definite dominant, and at some moment since
+_Gallus bankiva_ was domesticated, the element on which that special
+colour depends must have at least once been formed in the germ-cell of
+a fowl; but we need harder evidence than any which has yet been
+produced before we can declare that this novelty came through
+over-feeding, or change of climate, or any other disturbance
+consequent on domestication. When we reflect on the intricacies of
+genetic problems as we must now conceive them there come moments when
+we feel almost thankful that the Mendelian principles were unknown to
+Darwin. The time called for a bold pronouncement, and he made it, to
+our lasting profit and delight. With fuller knowledge we pass once
+more into a period of cautious expectation and reserve.
+
+In every arduous enterprise it is pleasanter to look back at
+difficulties overcome than forward to those which still seem
+insurmountable, but in the next stage there is nothing to be stained
+by disguising the fact that the attributes of living things are not
+what we used to suppose. If they are more complex in the sense that
+the properties they display are throughout so regular[71] that the
+Selection of minute random variations is an unacceptable account of
+the origin of their diversity, yet by virtue of that very regularity
+the problem is limited in scope and thus simplified.
+
+To begin with, we must relegate Selection to its proper place.
+Selection permits the viable to continue and decides that the
+non-viable shall perish; just as the temperature of our atmosphere
+decides that no liquid carbon shall be found on the face of the earth:
+but we do not suppose that the form of the diamond has been gradually
+achieved by a process of Selection. So again, as the course of descent
+branches in the successive generations, Selection determines along
+which branch Evolution shall proceed, but it does not decide what
+novelties that branch shall bring forth. "_La Nature contient le fonds
+de toutes ces variétés, mais le hazard ou l'art les mettent en
+Å“uvre_," as Maupertuis most truly said.
+
+Not till knowledge of the genetic properties of organisms has attained
+to far greater completeness can evolutionary speculations have more
+than a suggestive value. By genetic experiment, cytology and
+physiological chemistry aiding, we may hope to acquire such knowledge.
+In 1872 Nathusius wrote:[72] "Das Gesetz der Vererbung ist noch nicht
+erkannt; der Apfel ist noch nicht vom Baum der Erkenntniss gefallen,
+welcher, der Sage nach, Newton auf den rechten Weg zur Ergründung der
+Gravitationsgesetze führte." We cannot pretend that the words are not
+still true, but in Mendelian analysis the seeds of that apple-tree at
+last are sown.
+
+If we were asked what discovery would do most to forward our inquiry,
+what one bit of knowledge would more than any other illuminate the
+problem, I think we may give the answer without hesitation. The
+greatest advance that we can foresee will be made when it is found
+possible to connect the geometrical phenomena of development with the
+chemical. The geometrical symmetry of living things is the key to a
+knowledge of their regularity, and the forces which cause it. In the
+symmetry of the dividing cell the basis of that resemblance we call
+Heredity is contained. To imitate the morphological phenomena of life
+we have to devise a system which can divide. It must be able to
+divide, and to segment as--grossly--a vibrating plate or rod does, or
+as an icicle can do as it becomes ribbed in a continuous stream of
+water; but with this distinction, that the distribution of chemical
+differences and properties must simultaneously be decided and disposed
+in orderly relation to the pattern of the segmentation. Even if a
+model which would do this could be constructed it might prove to be a
+useful beginning.
+
+This may be looking too far ahead. If we had to choose some one piece
+of more proximate knowledge which we would more especially like to
+acquire, I suppose we should ask for the secret of interracial
+sterility. Nothing has yet been discovered to remove the grave
+difficulty, by which Huxley in particular was so much oppressed, that
+among the many varieties produced under domestication--which we all
+regard as analogous to the species seen in nature--no clear case of
+interracial sterility has been demonstrated. The phenomenon is
+probably the only one to which the domesticated products seem to
+afford no parallel. No solution of the difficulty can be offered which
+has positive value, but it is perhaps worth considering the facts in
+the light of modern ideas. It should be observed that we are not
+discussing incompatibility of two species to produce offspring (a
+totally distinct phenomenon), but the sterility of the offspring which
+many of them do produce.
+
+When two species, both perfectly fertile severally, produce on crossing a
+sterile progeny, there is a presumption that the sterility is due to the
+development in the hybrid of some substance which can only be formed by the
+meeting of two complementary factors. That some such account is correct in
+essence may be inferred from the well-known observation that if the hybrid
+is not totally sterile but only partially so, and thus is able to form some
+good germ-cells which develop into new individuals, the sterility of these
+daughter-individuals is sensibly reduced or may be entirely absent. The
+fertility once re-established, the sterility does not return in the later
+progeny, a fact strongly suggestive of segregation. Now if the sterility of
+the cross-bred be really the consequence of the meeting of two
+complementary factors, we see that the phenomenon could only be produced
+among the divergent offspring of one species by the acquisition of at least
+_two_ new factors; for if the acquisition of a single factor caused
+sterility the line would then end. Moreover each factor must be separately
+acquired by distinct individuals, for if both were present together, the
+possessors would by hypothesis be sterile. And in order to imitate the case
+of species each of these factors must be acquired by distinct breeds. The
+factors need not, and probably would not, produce any other perceptible
+effects; they might, like the colour-factors present in white flowers, make
+no difference in the form or other characters. Not till the cross was
+actually made between the two complementary individuals would either factor
+come into play, and the effects even then might be unobserved until an
+attempt was made to breed from the cross-bred.
+
+Next, if the factors responsible for sterility were acquired, they
+would in all probability be peculiar to certain individuals and would
+not readily be distributed to the whole breed. Any member of the breed
+also into which _both_ the factors were introduced would drop out of
+the pedigree by virtue of its sterility. Hence the evidence that the
+various domesticated breeds say of dogs or fowls can when mated
+together produce fertile offspring, is beside the mark. The real
+question is, Do they ever produce sterile offspring? I think the
+evidence is clearly that sometimes they do, oftener perhaps than is
+commonly supposed. These suggestions are quite amenable to
+experimental tests. The most obvious way to begin is to get a pair of
+parents which are known to have had any sterile offspring, and to find
+the proportions in which these steriles were produced. If, as I
+anticipate, these proportions are found to be definite, the rest is
+simple.
+
+In passing, certain other considerations may be referred to. First,
+that there are observations favouring the view that the production of
+totally sterile cross-breds is seldom a universal property of two
+species, and that it may be a matter of individuals, which is just
+what on the view here proposed would be expected. Moreover, as we all
+know now, though incompatibility may be dependent to some extent on
+the degree to which the species are dissimilar, no such principle can
+be demonstrated to determine sterility or fertility in general. For
+example, though all our Finches can breed together, the hybrids are
+all sterile. Of Ducks some species can breed together without
+producing the slightest sterility; others have totally sterile
+offspring, and so on. The hybrids between several _genera_ of Orchids
+are perfectly fertile on the female side, and some on the male side
+also, but the hybrids produced between the Turnip (_Brassica napus_)
+and the Swede (_Brassica campestris_), which, according to our
+estimates of affinity, should be nearly allied forms, are totally
+sterile.[73] Lastly, it may be recalled that in sterility we are
+almost certainly considering a meristic phenomenon. _Failure to
+divide_ is, we may feel fairly sure, the immediate "cause" of the
+sterility. Now, though we know very little about the heredity of
+meristic differences, all that we do know points to the conclusion
+that the less-divided is dominant to the more-divided, and we are thus
+justified in supposing that there are factors which can arrest or
+prevent cell-division. My conjecture therefore is that in the case of
+sterility of cross-breds we see the effect produced by a complementary
+pair of such factors. This and many similar problems are now open to
+our analysis.
+
+The question is sometimes asked, Do the new lights on Variation and
+Heredity make the process of Evolution easier to understand? On the
+whole the answer may be given that they do. There is some appearance
+of loss of simplicity, but the gain is real. As was said above, the
+time is not ripe for the discussion of the origin of species. With
+faith in Evolution unshaken--if indeed the word faith can be used in
+application to that which is certain--we look on the manner and
+causation of adapted differentiation as still wholly mysterious. As
+Samuel Butler so truly said: "To me it seems that the 'Origin of
+Variation,' whatever it is, is the only true 'Origin of Species,'"[74]
+and of that Origin not one of us knows anything. But given
+Variation--and it is given: assuming further that the variations are
+not guided into paths of adaptation--and both to the Darwinian and to
+the modern school this hypothesis appears to be sound if unproven--an
+evolution of species proceeding by definite steps is more, rather than
+less, easy to imagine than an evolution proceeding by the accumulation
+of indefinite and insensible steps. Those who have lost themselves in
+contemplating the miracles of Adaptation (whether real or spurious)
+have not unnaturally fixed their hopes rather on the indefinite than
+on the definite changes. The reasons are obvious. By suggesting that
+the steps through which an adaptative mechanism arose were indefinite
+and insensible, all further trouble is spared. While it could be said
+that species arise by an insensible and imperceptible process of
+variation, there was clearly no use in tiring ourselves by trying to
+perceive that process. This labour-saving counsel found great favour.
+All that had to be done to develop evolution-theory was to discover
+the good in everything, a task which, in the complete absence of any
+control or test whereby to check the truth of the discovery, is not
+very onerous. The doctrine "_que tout est au mieux_" was therefore
+preached with fresh vigour, and examples of that illuminating
+principle were discovered with a facility that Pangloss himself might
+have envied, till at last even the spectators wearied of such dazzling
+performances.
+
+But in all seriousness, why should indefinite and unlimited variation
+have been regarded as a more probable account of the origin of
+Adaptation? Only, I think, because the obstacle was shifted one plane
+back, and so looked rather less prominent. The abundance of
+Adaptation, we all grant, is an immense, almost an unsurpassable
+difficulty in all non-Lamarckian views of Evolution; but if the steps
+by which that adaptation arose were fortuitious, to imagine them
+insensible is assuredly no help. In one most important respect indeed,
+as has often been observed, it is a multiplication of troubles. For
+the smaller the steps, the less could Natural Selection act upon them.
+Definite variations--and of the occurrence of definite variations in
+abundance we have now the most convincing proof--have at least the
+obvious merit that they can make and often do make a real difference
+in the chances of life.
+
+There is another aspect of the Adaptation problem to which I can
+allude very briefly. May not our present ideas of the universality and
+precision of Adaptation be greatly exaggerated? The fit of organism to
+its environment is not after all so very close--a proposition
+unwelcome perhaps, but one which could be illustrated by very copious
+evidence. Natural Selection is stern, but she has her tolerant moods.
+
+We have now most certain and irrefragable proof that much definiteness
+exists in living things apart from Selection, and also much that may
+very well have been preserved and so in a sense constituted by
+Selection. Here the matter is likely to rest. There is a passage in
+the sixth edition of the _Origin_ which has I think been overlooked.
+On page 70 Darwin says, "The tuft of hair on the breast of the wild
+turkey-cock cannot be of any use, and it is doubtful whether it can be
+ornamental in the eyes of the female bird." This tuft of hair is a
+most definite and unusual structure, and I am afraid that the remark
+that it "cannot be of any use" may have been made inadvertently; but
+it may have been intended, for in the first edition the usual
+qualification was given and must therefore have been deliberately
+excised. Anyhow I should like to think that Darwin did throw over that
+tuft of hair, and that he felt relief when he had done so. Whether
+however we have his great authority for such a course or not, I feel
+quite sure that we shall be rightly interpreting the facts of nature
+if we cease to expect to find purposefulness wherever we meet with
+definite structures or patterns. Such things are, as often as not, I
+suspect rather of the nature of tool-marks, mere incidents of
+manufacture, benefiting their possessor not more than the wire-marks
+in a sheet of paper, or the ribbing on the bottom of an oriental plate
+renders those objects more attractive in our eyes.
+
+If Variation may be in any way definite, the question once more
+arises, may it not be definite in direction? The belief that it is has
+had many supporters, from Lamarck onwards, who held that it was guided
+by need, and others who, like Nägeli, while laying no emphasis on
+need, yet were convinced that there was guidance of some kind. The
+latter view under the name of "Orthogenesis," devised I believe by
+Eimer, at the present day commends itself to some naturalists. The
+objection to such a suggestion is of course that no fragment of real
+evidence can be produced in its support. On the other hand, with the
+experimental proof that variation consists largely in the unpacking
+and repacking of an original complexity, it is not so certain as we
+might like to think that the order of these events is not
+predetermined.
+
+For instance the original "pack" may have been made in such a way that
+at the _n_th division of the germ-cells of a Sweet Pea a colour-factor
+might be dropped, and that at the _n_+_n_th division the hooded
+variety be given off, and so on. I see no ground whatever for holding
+such a view, but in fairness the possibility should not be forgotten,
+and in the light of modern research it scarcely looks so absurdly
+improbable as before.
+
+No one can survey the work of recent years without perceiving that
+evolutionary orthodoxy developed too fast, and that a great deal has
+got to come down; but this satisfaction at least remains, that in the
+experimental methods which Mendel inaugurated, we have means of
+reaching certainty in regard to the physiology of Heredity and
+Variation upon which a more lasting structure may be built.
+
+FOOTNOTES:
+
+[Footnote 56: _Vénus Physique, contenant deux Dissertations, l'une sur
+l'origine des Hommes et des Animaux_; _Et l'autre sur l'origine des
+Noirs_, La Haye, 1746, pp. 124 and 129. For an introduction to the
+writings of Maupertuis I am indebted to an article by Professor
+Lovejoy in _Popular Sci. Monthly_, 1902.]
+
+[Footnote 57: For the fullest account of the views of these pioneers
+of Evolution, see the works of Samuel Butler, especially _Evolution,
+Old and New_ (2nd edit.) 1882. Butler's claims on behalf of Buffon
+have met with some acceptance; but after reading what Butler has said,
+and a considerable part of Buffon's own works, the word "hinted" seems
+to me a sufficiently correct description of the part he played. It is
+interesting to note that in the chapter on the Ass, which contains
+some of his evolutionary passages, there is a reference to "_plusieurs
+idées très-élevées sur la génération_" contained in the Letters of
+Maupertuis.]
+
+[Footnote 58: See especially W. Lawrence, _Lectures on Physiology_,
+London, 1823, pp. 213 f.]
+
+[Footnote 59: See the chapter contributed to the _Life and Letters of
+Charles Darwin_, II. p. 195. I do not clearly understand the sense in
+which Darwin wrote (Autobiography, _ibid._ I. p. 87): "It has
+sometimes been said that the success of the _Origin_ proved 'that the
+subject was in the air,' or 'that men's minds were prepared for it.' I
+do not think that this is strictly true, for I occasionally sounded
+not a few naturalists, and never happened to come across a single one
+who seemed to doubt about the permanence of species." This experience
+may perhaps have been an accident due to Darwin's isolation. The
+literature of the period abounds with indications of "critical
+expectancy." A most interesting expression of that feeling is given in
+the charming account of the "Early Days of Darwinism" by Alfred
+Newton, _Macmillan's Magazine_, LVII. 1888, p. 241. He tells how in
+1858 when spending a dreary summer in Iceland, he and his friend, the
+ornithologist John Wolley, in default of active occupation, spent
+their days in discussion. "Both of us taking a keen interest in
+Natural History, it was but reasonable that a question, which in those
+days was always coming up wherever two or more naturalists were
+gathered together, should be continually recurring. That question was,
+'What is a species?' and connected therewith was the other question,
+'How did a species begin?'... Now we were of course fairly well
+acquainted with what had been published on these subjects." He then
+enumerates some of these publications, mentioning among others T.
+Vernon Wollaston's _Variation of Species_--a work which has in my
+opinion never been adequately appreciated. He proceeds: "Of course we
+never arrived at anything like a solution of these problems, general
+or special, but we felt very strongly that a solution ought to be
+found, and that quickly, if the study of Botany and Zoology was to
+make any great advance." He then describes how on his return home he
+received the famous number of the _Linnean Journal_ on a certain
+evening. "I sat up late that night to read it; and never shall I
+forget the impression it made upon me. Herein was contained a
+perfectly simple solution of all the difficulties which had been
+troubling me for months past.... I went to bed satisfied that a
+solution had been found."]
+
+[Footnote 60: _Origin_, 6th edit. (1882), p. 421.]
+
+[Footnote 61: Whatever be our estimate of the importance of Natural
+Selection, in this we all agree. Samuel Butler, the most brilliant,
+and by far the most interesting of Darwin's opponents--whose works are
+at length emerging from oblivion--in his Preface (1882) to the 2nd
+edition of _Evolution, Old and New_, repeats his earlier expression of
+homage to one whom he had come to regard as an enemy: "To the end of
+time, if the question be asked, 'Who taught people to believe in
+Evolution?' the answer must be that it was Mr. Darwin. This is true,
+and it is hard to see what palm of higher praise can be awarded to any
+philosopher."]
+
+[Footnote 62: _Life and Letters_, I. pp. 276 and 83.]
+
+[Footnote 63: This isolation of the systematists is the one most
+melancholy sequela of Darwinism. It seems an irony that we should read
+in the peroration to the _Origin_ that when the Darwinian view is
+accepted "Systematists will be able to pursue their labours as at
+present; but they will not be incessantly haunted by the shadowy doubt
+whether this or that form be a true species. This, I feel sure, and I
+speak after experience, will be no slight relief. The endless disputes
+whether or not some fifty species of British brambles are good species
+will cease." _Origin_, 6th edit. (1882), p. 425. True they have ceased
+to attract the attention of those who lead opinion, but anyone who
+will turn to the literature of systematics will find that they have
+not ceased in any other sense. Should there not be something
+disquieting in the fact that among the workers who come most into
+contact with specific differences, are to be found the only men who
+have failed to be persuaded of the unreality of those differences?]
+
+[Footnote 64: 6th edit. pp. 109 and 401. See Butler, _Essays on Life,
+Art, and Science_, p. 265, reprinted 1908, and _Evolution, Old and
+New_, chap. XXII. (2nd edit.), 1882.]
+
+[Footnote 65: W. Lawrence was one of the few who consistently
+maintained the contrary opinion. Prichard, who previously had
+expressed himself in the same sense, does not, I believe, repeat these
+views in his later writings, and there are signs that he came to
+believe in the transmission of acquired habits. See Lawrence, _Lect.
+Physiol._ 1823, pp. 436-437, 447. Prichard, Edin. Inaug. Disp. 1808
+[not seen by me], quoted _ibid._ and _Nat. Hist. Man_, 1843, pp. 34
+f.]
+
+[Footnote 66: It is interesting to see how nearly Butler was led by
+natural penetration, and from absolutely opposite conclusions, back to
+this underlying truth: "So that each ovum when impregnate should be
+considered not as descended from its ancestors, but as being a
+continuation of the personality of every ovum in the chain of its
+ancestry, which every ovum _it actually is_ quite as truly as the
+octogenarian _is_ the same identity with the ovum from which he has
+been developed. This process cannot stop short of the primordial cell,
+which again will probably turn out to be but a brief resting-place. We
+therefore prove each one of us to _be actually_ the primordial cell
+which never died nor dies, but has differentiated itself into the life
+of the world, all living beings whatever, being one with it and
+members one of another," _Life and Habit_, 1878, p. 86.]
+
+[Footnote 67: This view is no doubt contrary to the received opinion.
+I am however interested to see it lately maintained by Driesch
+(_Science and Philosophy of the Organism_, London, 1907, p. 233), and
+from the recent observations of Godlewski it has received distinct
+experimental support.]
+
+[Footnote 68: In other words, the ova are each _either_ female, _or
+male_ (i.e. non-female), but the sperms are all non-female.]
+
+[Footnote 69: Morgan, _Proc. Soc. Exp. Biol. Med._ V. 1908, and von
+Baehr, _Zool. Anz._ XXXII. p. 507, 1908.]
+
+[Footnote 70: As Wilson has proved, the unpaired body is not a
+universal feature even in those orders in which it has been observed.
+Nearly allied types may differ. In some it is altogether unpaired. In
+others it is paired with a body of much smaller size, and by selection
+of various types all gradations can be demonstrated ranging to the
+condition in which the members of the pair are indistinguishable from
+each other.]
+
+[Footnote 71: I have in view, for example, the marvellous and specific
+phenomena of regeneration, and those discovered by the students of
+"_Entwicklungsmechanik_." The circumstances of its occurrence here
+preclude any suggestion that this regularity has been brought about by
+the workings of Selection. The attempts thus to represent the
+phenomena have resulted in mere parodies of scientific reasoning.]
+
+[Footnote 72: _Vorträge über Viehzucht und Rassenerkenntniss_, p. 120,
+Berlin, 1872.]
+
+[Footnote 73: See Sutton, A. W., _Journ. Linn. Soc._ XXXVIII. p. 341,
+1908.]
+
+[Footnote 74: _Life and Habit_, London, p. 263, 1878]
+
+
+
+
+IV
+
+"THE DESCENT OF MAN"
+
+BY G. SCHWALBE
+
+_Professor of Anatomy in the University of Strassburg_
+
+
+The problem of the origin of the human race, of the descent of man, is
+ranked by Huxley in his epoch-making book _Man's Place in Nature_, as
+the deepest with which biology has to concern itself, "the question of
+questions,"--the problem which underlies all others. In the same
+brilliant and lucid exposition, which appeared in 1863, soon after the
+publication of Darwin's _Origin of Species_, Huxley stated his own
+views in regard to this great problem. He tells us how the idea of a
+natural descent of man gradually grew up in his mind. It was
+especially the assertions of Owen in regard to the total difference
+between the human and the simian brain that called forth strong
+dissent from the great anatomist Huxley, and he easily succeeded in
+showing that Owen's supposed differences had no real existence; he
+even established, on the basis of his own anatomical investigations,
+the proposition that the anatomical differences between the Marmoset
+and the Chimpanzee are much greater than those between the Chimpanzee
+and Man.
+
+But why do we thus introduce the study of Darwin's _Descent of Man_,
+which is to occupy us here, by insisting on the fact that Huxley had
+taken the field in defence of the descent of man in 1863, while
+Darwin's book on the subject did not appear till 1871? It is in order
+that we may clearly understand how it happened that from this time
+onwards Darwin and Huxley followed the same great aim in the most
+intimate association.
+
+Huxley and Darwin working at the same _Problema maximum_! Huxley
+fiery, impetuous, eager for battle, contemptuous of the resistance of
+a dull world, or energetically triumphing over it. Darwin calm,
+weighing every problem slowly, letting it mature thoroughly,--not a
+fighter, yet having the greater and more lasting influence by virtue
+of his immense mass of critically sifted proofs. Darwin's friend,
+Huxley, was the first to do him justice, to understand his nature, and
+to find in it the reason why the detailed and carefully considered
+book on the descent of man made its appearance so late. Huxley, always
+generous, never thought of claiming priority for himself. In
+enthusiastic language he tells how Darwin's immortal work, _The Origin
+of Species_, first shed light for him on the problem of the descent of
+man; the recognition of a _vera causa_ in the transformation of
+species illuminated his thoughts as with a flash. He was now content
+to leave what perplexed him, what he could not yet solve, as he says
+himself, "in the mighty hands of Darwin." Happy in the bustle of
+strife against old and deep-rooted prejudices, against intolerance and
+superstition, he wielded his sharp weapons on Darwin's behalf; wearing
+Darwin's armour he joyously overthrew adversary after adversary.
+Darwin spoke of Huxley as his "general agent."[75] Huxley says of
+himself "I am Darwin's bulldog."[76]
+
+Thus Huxley openly acknowledged that it was Darwin's _Origin of
+Species_ that first set the problem of the descent of man in its true
+light, that made the question of the origin of the human race a
+pressing one. That this was the logical consequence of his book Darwin
+himself had long felt. He had been reproached with intentionally
+shirking the application of his theory to Man. Let us hear what he
+says on this point in his autobiography: "As soon as I had become, in
+the year 1837 or 1838, convinced that species were mutable
+productions, I could not avoid the belief that man must come under the
+same law. Accordingly I collected notes on the subject for my own
+satisfaction, and not for a long time with any intention of
+publishing. Although in the 'Origin of Species' the derivation of any
+particular species is never discussed, yet I thought it best, in order
+_that no honourable man should accuse me of concealing my views_,[77]
+to add that by the work 'light would be thrown on the origin of man
+and his history.' It would have been useless and injurious to the
+success of the book to have paraded, without giving any evidence, my
+conviction with respect to his origin."[78]
+
+In a letter written in January, 1860, to the Rev. L. Blomefield,
+Darwin expresses himself in similar terms. "With respect to man, I am
+very far from wishing to obtrude my belief; but I thought it dishonest
+to quite conceal my opinion."[79]
+
+The brief allusion in the _Origin of Species_ is so far from prominent
+and so incidental that it was excusable to assume that Darwin had not
+touched upon the descent of man in this work. It was solely the desire
+to have his mass of evidence sufficiently complete, solely Darwin's
+great characteristic of never publishing till he had carefully weighed
+all aspects of his subject for years, solely, in short, his most
+fastidious scientific conscience that restrained him from challenging
+the world in 1859 with a book in which the theory of the descent of
+man was fully set forth. Three years, frequently interrupted by
+ill-health, were needed for the actual writing of the book:[80] the
+first edition, which appeared in 1871, was followed in 1874 by a much
+improved second edition, the preparation of which he very reluctantly
+undertook.[81]
+
+This, briefly, is the history of the work, which, with the _Origin of
+Species_, marks an epoch in the history of biological sciences--the
+work with which the cautious, peace-loving investigator ventured forth
+from his contemplative life into the arena of strife and unrest, and
+laid himself open to all the annoyances that deep-rooted belief and
+prejudice, and the prevailing tendency of scientific thought at the
+time could devise.
+
+Darwin did not take this step lightly. Of great interest in this
+connection is a letter written to Wallace on Dec. 22, 1857,[82] in
+which he says, "You ask me whether I shall discuss 'man.' I think I
+shall avoid the whole subject, as so surrounded with prejudices;
+though I fully admit that it is the highest and most interesting
+problem for the naturalist." But his conscientiousness compelled him
+to state briefly his opinion on the subject in the _Origin of Species_
+in 1859. Nevertheless he did not escape reproaches for having been so
+reticent. This is unmistakably apparent from a letter to Fritz Müller
+dated Feb. 22 [1869?], in which he says: "I am thinking of writing a
+little essay on the Origin of Mankind, as I have been taunted with
+concealing my opinions."[83]
+
+It might be thought that Darwin behaved thus hesitatingly, and was so
+slow in deciding on the full publication of his collected material in
+regard to the descent of man, because he had religious difficulties to
+overcome.
+
+But this was not the case, as we can see from his admirable confession
+of faith, the publication of which we owe to his son Francis.[84]
+Whoever wishes really to understand the lofty character of this great
+man should read these immortal lines in which he unfolds to us in
+simple and straightforward words the development of his conception of
+the universe. He describes how, though he was still quite orthodox
+during his voyage round the world on board the _Beagle_, he came
+gradually to see, shortly afterwards (1836-1839) that the Old
+Testament was no more to be trusted than the Sacred Books of the
+Hindoos; the miracles by which Christianity is supported, the
+discrepancies between the accounts in the different Gospels, gradually
+led him to disbelieve in Christianity as a divine revelation. "Thus,"
+he writes,[85] "disbelief crept over me at a very slow rate, but was
+at last complete. The rate was so slow that I felt no distress." But
+Darwin was too modest to presume to go beyond the limits laid down by
+science. He wanted nothing more than to be able to go, freely and
+unhampered by belief in authority or in the Bible, as far as human
+knowledge could lead him. We learn this from the concluding words of
+his chapter on religion "The mystery of the beginning of all things is
+insoluble by us; and I for one must be content to remain an
+Agnostic."[86]
+
+Darwin was always very unwilling to give publicity to his views in
+regard to religion. In a letter to Asa Gray on May 22, 1860,[87] he
+declares that it is always painful to him to have to enter into
+discussion of religious problems. He had, he said, no intention of
+writing atheistically.
+
+Finally, let us cite one characteristic sentence from a letter from
+Darwin to C. Ridley[88] (Nov. 28, 1878). A clergyman, Dr. Pusey, had
+asserted that Darwin had written the _Origin of Species_ with some
+relation to theology. Darwin writes emphatically, "Many years ago when
+I was collecting facts for the 'Origin,' my belief in what is called a
+personal God was as firm as that of Dr. Pusey himself, and as to the
+eternity of matter I never troubled myself about such insoluble
+questions." The expression "many years ago" refers to the time of his
+voyage round the world, as has already been pointed out. Darwin means
+by this utterance that the views which had gradually developed in his
+mind in regard to the origin of species were quite compatible with the
+faith of the Church.
+
+If we consider all these utterances of Darwin in regard to religion
+and to his outlook on life (Weltanschauung), we shall see at least so
+much, that religious reflection could in no way have influenced him in
+regard to the writing and publishing of his book on _The Descent of
+Man_. Darwin had early won for himself freedom of thought, and to this
+freedom he remained true to the end of his life, uninfluenced by the
+customs and opinions of the world around him.
+
+Darwin was thus inwardly fortified and armed against the host of
+calumnies, accusations, and attacks called forth by the publication of
+the _Origin of Species_, and to an even greater extent by the
+appearance of the _Descent of Man_. But in his defence he could rely
+on the aid of a band of distinguished auxiliaries of the rarest
+ability. His faithful confederate, Huxley, was joined by the botanist
+Hooker, and, after longer resistance, by the famous geologist Lyell,
+whose "conversion" afforded Darwin peculiar satisfaction. All three
+took the field with enthusiasm in defence of the natural descent of
+man. From Wallace, on the other hand, though he shared with him the
+idea of natural selection, Darwin got no support in this matter.
+Wallace expressed himself in a strange manner. He admitted everything
+in regard to the morphological descent of man, but maintained, in a
+mystic way, that something else, something of a spiritual nature must
+have been added to what man inherited from his animal ancestors.
+Darwin, whose esteem for Wallace was extraordinarily high, could not
+understand how he could give utterance to such a mystical view in
+regard to man; the idea seemed to him so "incredibly strange" that he
+thought some one else must have added these sentences to Wallace's
+paper.
+
+Even now there are thinkers who, like Wallace, shrink from applying to
+man the ultimate consequences of the theory of descent. The idea that
+man is derived from ape-like forms is to them unpleasant and
+humiliating.
+
+So far I have been depicting the development of Darwin's work on the
+descent of man. In what follows I shall endeavour to give a condensed
+survey of the contents of the book.
+
+It must at once be said that the contents of Darwin's work fall into
+two parts, dealing with entirely different subjects. _The Descent of
+Man_ includes a very detailed investigation in regard to secondary
+sexual characters in the animal series, and on this investigation
+Darwin founded a new theory, that of sexual selection. With
+astonishing patience he gathered together an immense mass of material,
+and showed, in regard to Arthropods and Vertebrates, the wide
+distribution of secondary characters, which develop almost exclusively
+in the male, and which enable him, on the one hand, to get the better
+of his rivals in the struggle for the female by the greater perfection
+of his weapons, and, on the other hand, to offer greater allurements
+to the female through the higher development of decorative characters,
+of song, or of scent-producing glands. The best equipped males will
+thus crowd out the less well-equipped in the matter of reproduction,
+and thus the relevant characters will be increased and perfected
+through sexual selection. It is, of course, a necessary assumption
+that these secondary sexual characters may be transmitted to the
+female, although perhaps in rudimentary form.
+
+As we have said, this story of sexual selection takes up a great deal
+of space in Darwin's book, and it need only be considered here in so
+far as Darwin applied it to the descent of man. To this latter problem
+the whole of Part I is devoted, while Part III contains a discussion
+of sexual selection in relation to man, and a general summary. Part
+II treats of sexual selection in general, and may be disregarded in
+our present study. Moreover, many interesting details must necessarily
+be passed over in what follows, for want of space.
+
+The first part of the _Descent of Man_ begins with an enumeration of
+the proofs of the animal descent of man taken from the structure of
+the human body. Darwin chiefly emphasises the fact that the human body
+consists of the same organs and of the same tissues as those of the
+other mammals; he shows also that man is subject to the same diseases
+and tormented by the same parasites as the apes. He further dwells on
+the general agreement exhibited by young embryonic forms, and he
+illustrates this by two figures placed one above the other, one
+representing a human embryo, after Ecker, the other a dog embryo,
+after Bischoff.[89]
+
+Darwin finds further proofs of the animal origin of man in the reduced
+structures, in themselves extremely variable, which are either
+absolutely useless to their possessors, or of so little use that they
+could never have developed under existing conditions. Of such vestiges
+he enumerates: the defective development of the _panniculus carnosus_
+(muscle of the skin) so widely distributed among mammals, the
+ear-muscles, the occasional persistence of the animal ear-point in
+man, the rudimentary nictitating membrane (_plica semilunaris_) in the
+human eye, the slight development of the organ of smell, the general
+hairiness of the human body, the frequently defective development or
+entire absence of the third molar (the wisdom tooth), the vermiform
+appendix, the occasional reappearance of a bony canal (_foramen
+supracondyloideum_) at the lower end of the humerus, the rudimentary
+tail of man (the so-called taillessness), and so on. Of these
+rudimentary structures the occasional occurrence of the animal
+ear-point in man is most fully discussed. Darwin's attention was
+called to this interesting structure by the sculptor Woolner. He
+figures such a case observed in man, and also the head of an alleged
+orang-foetus, the photograph of which he received from Nitsche.
+
+Darwin's interpretation of Woolner's case as having arisen through a
+folding over of the free edge of a pointed ear has been fully borne
+out by my investigations on the external ear.[90] In particular, it
+was established by these investigations that the human foetus, about
+the middle of its embryonic life, possesses a pointed ear somewhat
+similar to that of the monkey genus Macacus. One of Darwin's
+statements in regard to the head of the orang-foetus must be
+corrected. A _large_ ear with a point is shown in the photograph,[91]
+but it can easily be demonstrated--and Deniker has already pointed
+this out--that the figure is not that of an orang foetus at all, for
+that form has much smaller ears with no point; nor can it be a
+gibbon-foetus, as Deniker supposes, for the gibbon ear is also without
+a point. I myself regard it as that of a Macacus-embryo. But this
+mistake, which is due to Nitsche, in no way affects the fact
+recognised by Darwin, that ear-forms showing the point characteristic
+of the animal ear occur in man with extraordinary frequency.
+
+Finally, there is a discussion of those rudimentary structures which
+occur only in _one_ sex, such as the rudimentary mammary glands in the
+male, the vesicula prostatica, which corresponds to the uterus of the
+female, and others. All these facts tell in favour of the common
+descent of man and all other vertebrates. The conclusion of this
+section is characteristic: "_It is only our natural prejudice, and
+that arrogance which made our forefathers declare that they were
+descended from demi-gods, which leads us to demur to this conclusion.
+But the time will before long come, when it will be thought wonderful
+that naturalists, who were well acquainted with the comparative
+structure and development of man, and other mammals, should have
+believed that each was the work of a separate act of creation._"[92]
+
+In the second chapter there is a more detailed discussion, again based
+upon an extraordinary wealth of facts, of the problem as to the manner
+in which, and the causes through which, man evolved from a lower form.
+Precisely the same causes are here suggested for the origin of man, as
+for the origin of species in general. Variability, which is a
+necessary assumption in regard to all transformations, occurs in man
+to a high degree. Moreover, the rapid multiplication of the human race
+creates conditions which necessitate an energetic struggle for
+existence, and thus afford scope for the intervention of natural
+selection. Of the exercise of _artificial_ selection in the human
+race, there is nothing to be said, unless we cite such cases as the
+grenadiers of Frederick William I, or the population of ancient
+Sparta. In the passages already referred to and in those which follow,
+the transmission of acquired characters, upon which Darwin does not
+dwell, is taken for granted. In man, direct effects of changed
+conditions can be demonstrated (for instance in regard to bodily
+size), and there are also proofs of the influence exerted on his
+physical constitution by increased use or disuse. Reference is here
+made to the fact, established by Forbes, that the Quechua Indians of
+the high plateaus of Peru show a striking development of lungs and
+thorax, as a result of living constantly at high altitudes.
+
+Such special forms of variation as arrests of development
+(microcephalism) and reversion to lower forms are next discussed.
+Darwin himself felt[93] that these subjects are so nearly related to
+the cases mentioned in the first chapter, that many of them might as
+well have been dealt with there. It seems to me that it would have
+been better so, for the citation of additional instances of reversion
+at this place rather disturbs the logical sequence of his ideas as to
+the conditions which have brought about the evolution of man from
+lower forms. The instances of reversion here discussed are
+microcephalism, which Darwin wrongly interpreted as atavistic,
+supernumerary mammae, supernumerary digits, bicornuate uterus, the
+development of abnormal muscles, and so on. Brief mention is also made
+of correlative variations observed in man.
+
+Darwin next discusses the question as to the manner in which man
+attained to the erect position from the state of a climbing quadruped.
+Here again he puts the influence of Natural Selection in the first
+rank. The immediate progenitors of man had to maintain a struggle for
+existence in which success was to the more intelligent, and to those
+with social instincts. The hand of these climbing ancestors, which had
+little skill and served mainly for locomotion, could only undergo
+further development when some early member of the Primate series came
+to live more on the ground and less among trees.
+
+A bipedal existence thus became possible, and with it the liberation
+of the hand from locomotion, and the one-sided development of the
+human foot. The upright position brought about correlated variations
+in the bodily structure; with the free use of the hand it became
+possible to manufacture weapons and to use them; and this again
+resulted in a degeneration of the powerful canine teeth and the jaws,
+which were then no longer necessary for defence. Above all, however,
+the intelligence immediately increased, and with it skull and brain.
+The nakedness of man, and the absence of a tail (rudimentariness of
+the tail vertebrae) are next discussed. Darwin is inclined to
+attribute the nakedness of man, not to the action of natural selection
+on ancestors who originally inhabited a tropical land, but to sexual
+selection, which, for aesthetic reasons, brought about the loss of the
+hairy covering in man, or primarily in woman. An interesting
+discussion of the loss of the tail, which, however, man shares with
+the anthropoid apes, some other monkeys and lemurs, forms the
+conclusion of the almost superabundant material which Darwin worked
+up in the second chapter. His object was to show that some of the most
+distinctive human characters are in all probability directly or
+indirectly due to natural selection. With characteristic modesty he
+adds:[94] "Hence, if I have erred in giving to natural selection great
+power, which I am very far from admitting, or in having exaggerated
+its power, which is in itself probable, I have at least, as I hope,
+done good service in aiding to overthrow the dogma of separate
+creations." At the end of the chapter he touches upon the objection as
+to man's helpless and defenceless condition. Against this he urges his
+intelligence and social instincts.
+
+The two following chapters contain a detailed discussion of the
+objections drawn from the supposed great differences between the
+mental powers of men and animals. Darwin at once admits that the
+differences are enormous, but not that any fundamental difference
+between the two can be found. Very characteristic of him is the
+following passage: "In what manner the mental powers were first
+developed in the lowest organisms, is as hopeless an enquiry as how
+life itself first originated. These are problems for the distant
+future, if they are ever to be solved by man."[95]
+
+After some brief observations on instinct and intelligence, Darwin
+brings forward evidence to show that the greater number of the
+emotional states, such as pleasure and pain, happiness and misery,
+love and hate are common to man and the higher animals. He goes on to
+give various examples showing that wonder and curiosity, imitation,
+attention, memory and imagination (dreams of animals), can also be
+observed in the higher mammals, especially in apes. In regard even to
+reason there are no sharply defined limits. A certain faculty of
+deliberation is characteristic of some animals, and the more
+thoroughly we know an animal the more intelligence we are inclined to
+credit it with. Examples are brought forward of the intelligent and
+deliberate actions of apes, dogs and elephants. But although no
+sharply defined differences exist between man and animals, there is,
+nevertheless, a series of other mental powers which are
+characteristics usually regarded as absolutely peculiar to man. Some
+of these characteristics are examined in detail, and it is shown that
+the arguments drawn from them are not conclusive. Man alone is said to
+be capable of progressive improvement; but against this must be placed
+as something analogous in animals, the fact that they learn cunning
+and caution through long continued persecution. Even the use of tools
+is not in itself peculiar to man (monkeys use sticks, stones and
+twigs), but man alone fashions and uses implements _designed for a
+special purpose_. In this connection the remarks taken from Lubbock in
+regard to the origin and gradual development of the earliest flint
+implements will be read with interest; these are similar to the
+observations on modern eoliths, and their bearing on the development
+of the stone industry. It is interesting to learn from a letter to
+Hooker,[96] that Darwin himself at first doubted whether the stone
+implements discovered by Boucher de Perthes were really of the nature
+of tools. With the relentless candour as to himself which
+characterised him, he writes four years later in a letter to Lyell in
+regard to this view of Boucher de Perthes' discoveries: "I know
+something about his errors, and looked at his book many years ago, and
+am ashamed to think that I concluded the whole was rubbish! Yet he has
+done for man something like what Agassiz did for glaciers."[97]
+
+To return to Darwin's further comparisons between the higher mental powers
+of man and animals; He takes much of the force from the argument that man
+alone is capable of abstraction and self-consciousness by his own
+observations on dogs. One of the main differences between man and animals,
+speech, receives detailed treatment. He points out that various animals
+(birds, monkeys, dogs) have a large number of different sounds for
+different emotions, that, further, man produces in common with animals a
+whole series of inarticulate cries combined with gestures, and that dogs
+learn to understand whole sentences of human speech. In regard to human
+language, Darwin expresses a view contrary to that held by Max Müller:[98]
+"I cannot doubt that language owes its origin to the imitation and
+modification of various natural sounds, the voices of other animals, and
+man's own instinctive cries, aided by signs and gestures." The development
+of actual language presupposes a higher degree of intelligence than is
+found in any kind of ape. Darwin remarks on this point:[99] "The fact of
+the higher apes not using their vocal organs for speech no doubt depends on
+their intelligence not having been sufficiently advanced."
+
+The sense of beauty, too, has been alleged to be peculiar to man. In
+refutation of this assertion Darwin points to the decorative colours
+of birds, which are used for display. And to the last objection, that
+man alone has religion, that he alone has a belief in God, it is
+answered "that numerous races have existed, and still exist, who have
+no idea of one or more gods, and who have no words in their languages
+to express such an idea."[100]
+
+The result of the investigations recorded in this chapter is to show
+that, great as the difference in mental powers between man and the
+higher animals may be, it is undoubtedly only a difference "of degree
+and not of kind."[101]
+
+In the fourth chapter Darwin deals with the _moral sense_ or
+_conscience_, which is the most important of all differences between
+man and animals. It is a result of social instincts, which lead to
+sympathy for other members of the same society, to non-egoistic
+actions for the good of others. Darwin shows that social tendencies
+are found among many animals, and that among these love and
+kin-sympathy exist, and he gives examples of animals (especially dogs)
+which may exhibit characters that we should call moral in man (e.g.
+disinterested self-sacrifice for the sake of others). The early
+ape-like progenitors of the human race were undoubtedly social. With
+the increase of intelligence the moral sense develops farther; with
+the acquisition of speech public opinion arises, and finally, moral
+sense becomes habit. The rest of Darwin's detailed discussions on
+moral philosophy may be passed over.
+
+The fifth chapter may be very briefly summarised. In it Darwin shows
+that the intellectual and moral faculties are perfected through
+natural selection. He inquires how it can come about that a tribe at a
+low level of evolution attains to a higher, although the best and
+bravest among them often pay for their fidelity and courage with their
+lives without leaving any descendants. In this case it is the
+sentiment of glory, praise and blame, the admiration of others, which
+bring about the increase of the better members of the tribe. Property,
+fixed dwellings, and the association of families into a community are
+also indispensable requirements for civilisation. In the longer second
+section of the fifth chapter Darwin acts mainly as recorder. On the
+basis of numerous investigations, especially those of Greg, Wallace,
+and Galton, he inquires how far the influence of natural selection can
+be demonstrated in regard to civilised nations. In the final section,
+which deals with the proofs that all civilised nations were once
+barbarians, Darwin again uses the results gained by other
+investigators, such as Lubbock and Tylor. There are two sets of facts
+which prove the proposition in question. In the first place, we find
+traces of a former lower state in the customs and beliefs of all
+civilised nations, and in the second place, there are proofs to show
+that savage races are independently able to raise themselves a few
+steps in the scale of civilisation, and that they have thus raised
+themselves.
+
+In the sixth chapter of the work, Morphology comes into the foreground
+once more. Darwin first goes back, however, to the argument based on
+the great difference between the mental powers of the highest animals
+and those of man. That this is only quantitative, not qualitative, he
+has already shown. Very instructive in this connection is the
+reference to the enormous difference in mental powers in another
+class. No one would draw from the fact that the cochineal insect
+(Coccus) and the ant exhibit enormous differences in their mental
+powers, the conclusion that the ant should therefore be regarded as
+something quite distinct, and withdrawn from the class of insects
+altogether.
+
+Darwin next attempts to establish the _specific_ genealogical tree of
+man, and carefully weighs the differences and resemblances between the
+different families of the Primates. The erect position of man is an
+adaptive character, just as are the various characters referable to
+aquatic life in the seals, which, notwithstanding these, are ranked as
+a mere family of the carnivores. The following utterance is very
+characteristic of Darwin:[102] "If man had not been his own
+classifier, he would never have thought of founding a separate order
+for his own reception." In numerous characters not mentioned in
+systematic works, in the features of the face, in the form of the
+nose, in the structure of the external ear, man resembles the apes.
+The arrangement of the hair in man has also much in common with the
+apes; as also the occurrence of hair on the forehead of the human
+embryo, the beard, the convergence of the hair of the upper and under
+arm towards the elbow, which occurs not only in the anthropoid apes,
+but also in some American monkeys. Darwin here adopts Wallace's
+explanation of the origin of the ascending direction of the hair in
+the forearm of the orang,--that it has arisen through the habit of
+holding the hands over the head in rain. But this explanation cannot
+be maintained when we consider that this disposition of the hair is
+widely distributed among the most different mammals, being found in
+the dog, in the sloth, and in many of the lower monkeys.
+
+After further careful analysis of the anatomical characters Darwin
+reaches the conclusion that the New World monkeys (Platyrrhine) may be
+excluded from the genealogical tree altogether, but that man is an
+offshoot from the Old World monkeys (Catarrhine) whose progenitors
+existed as far back as the Miocene period. Among these Old World
+monkeys the forms to which man shows the greatest resemblance are the
+anthropoid apes, which, like him, possess neither tail nor ischial
+callosities. The platyrrhine and catarrhine monkeys have their
+primitive ancestor among extinct forms of the Lemuridae. Darwin also
+touches on the question of the original home of the human race and
+supposes that it may have been in Africa, because it is there that
+man's nearest relatives, the gorilla and the chimpanzee, are found.
+But he regards speculation on this point as useless. It is remarkable
+that, in this connection, Darwin regards the loss of the hair-covering
+in man as having some relation to a warm climate, while elsewhere he
+is inclined to make sexual selection responsible for it. Darwin
+recognises the great gap between man and his nearest relatives, but
+similar gaps exist at other parts of the mammalian genealogical tree:
+the allied forms have become extinct. After the extermination of the
+lower races of mankind, on the one hand, and of the anthropoid apes on
+the other, which will undoubtedly take place, the gulf will be greater
+than ever, since the baboons will then bound it on the one side, and
+the white races on the other. Little weight need be attached to the
+lack of fossil remains to fill up this gap, since the discovery of
+these depends upon chance. The last part of the chapter is devoted to
+a discussion of the earlier stages in the genealogy of man. Here
+Darwin accepts in the main the genealogical tree, which had meantime
+been published by Haeckel, who traces the pedigree back through
+Monotrems, Reptiles, Amphibians, and Fishes, to Amphioxus.
+
+Then follows an attempt to reconstruct, from the atavistic characters,
+a picture of our primitive ancestor who was undoubtedly an arboreal
+animal. The occurrence of rudiments of parts in one sex which only
+come to full development in the other is next discussed. This state of
+things Darwin regards as derived from an original hermaphroditism. In
+regard to the mammary glands of the male he does not accept the theory
+that they are vestigial, but considers them rather as not fully
+developed.
+
+The last chapter of Part I deals with the question whether the
+different races of man are to be regarded as different species, or as
+sub-species of a race of monophyletic origin. The striking differences
+between the races are first emphasised, and the question of the
+fertility or infertility of hybrids is discussed. That fertility is
+the more usual is shown by the excessive fertility of the hybrid
+population of Brazil. This, and the great variability of the
+distinguishing characters of the different races, as well as the fact
+that all grades of transition stages are found between these, while
+considerable general agreement exists, tell in favour of the unity of
+the races and lead to the conclusion that they all had a common
+primitive ancestor.
+
+Darwin therefore classifies all the different races as sub-species of
+_one and the same species_. Then follows an interesting inquiry into
+the reasons for the extinction of human races. He recognises as the
+ultimate reason the injurious effects of a change of the conditions of
+life, which may bring about an increase in infantile mortality, and a
+diminished fertility. It is precisely the reproductive system, among
+animals also, which is most susceptible to changes in the environment.
+
+The final section of this chapter deals with the formation of the
+races of mankind. Darwin discusses the question how far the direct
+effect of different conditions of life, or the inherited effects of
+increased use or disuse may have brought about the characteristic
+differences between the different races. Even in regard to the origin
+of the colour of the skin he rejects the transmitted effects of an
+original difference of climate as an explanation. In so doing he is
+following his tendency to exclude Lamarckian explanations as far as
+possible. But here he makes gratuitous difficulties from which, since
+natural selection fails, there is no escape except by bringing in the
+principle of sexual selection, to which, he regarded it as possible,
+skin-colouring, arrangement of hair, and form of features might be
+traced. But with his characteristic conscientiousness he guards
+himself thus: "I do not intend to assert that sexual selection will
+account for all the differences between the races."[103]
+
+I may be permitted a remark as to Darwin's attitude towards Lamarck.
+While, at an earlier stage, when he was engaged in the preliminary
+labours for his immortal work, _The Origin of Species_, Darwin
+expresses himself very forcibly against the views of Lamarck, speaking
+of Lamarckian "nonsense,"[104] and of Lamarck's "absurd, though clever
+work"[105] and expressly declaring, "I attribute very little to the
+direct action of climate, etc."[106] yet in later life he became more
+and more convinced of the influence of external conditions. In 1876,
+that is, two years after the appearance of the second edition of _The
+Descent of Man_, he writes with his usual candid honesty: "In my
+opinion the greatest error which I have committed, has been not
+allowing sufficient weight to the direct action of the environment,
+i.e. food, climate, etc. independently of a natural selection."[107]
+It is certain from this change of opinion that, if he had been able to
+make up his mind to issue a third edition of _The Descent of Man_, he
+would have ascribed a much greater influence to the effect of
+external conditions in explaining the different characters of the
+races of man than he did in the second edition. He would also
+undoubtedly have attributed less influence to sexual selection as a
+factor in the origin of the different bodily characteristics, if
+indeed he would not have excluded it altogether.
+
+In Part III of the _Descent_ two additional chapters are devoted to
+the discussion of sexual selection in relation to man. These may be
+very briefly referred to. Darwin here seeks to show that sexual
+selection has been operative on man and his primitive progenitor.
+Space fails me to follow out his interesting arguments. I can only
+mention that he is inclined to trace back hairlessness, the
+development of the beard in man, and the characteristic colour of the
+different human races to sexual selection. Since bareness of the skin
+could be no advantage, but rather a disadvantage, this character
+cannot have been brought about by natural selection. Darwin also
+rejected a direct influence of climate as a cause of the origin of the
+skin-colour. I have already expressed the opinion, based on the
+development of his views as shown in his letters, that in a third
+edition Darwin would probably have laid more stress on the influence
+of external environment. He himself feels that there are gaps in his
+proofs here, and says in self-criticism: "The views here advanced, on
+the part which sexual selection has played in the history of man, want
+scientific precision."[108] I need here only point out that it is
+impossible to explain the graduated stages of skin-colour by sexual
+selection, since it would have produced races sharply defined by their
+colour and not united to other races by transition stages, and this,
+it is well known, is not the case. Moreover, the fact established by
+me,[109] that in all races the ventral side of the trunk is paler than
+the dorsal side, and the inner surface of the extremities paler than
+the outer side, cannot be explained by sexual selection in the
+Darwinian sense.
+
+With this I conclude my brief survey of the rich contents of Darwin's
+book. I may be permitted to conclude by quoting the magnificent final
+words of _The Descent of Man_: "We must, however, acknowledge, as it
+seems to me, that man, with all his noble qualities, with sympathy
+which feels for the most debased, with benevolence which extends not
+only to other men but to the humblest living creature, with his
+god-like intellect which has penetrated into the movements and
+constitution of the solar system--with all these exalted powers--Man
+still bears in his bodily frame the indelible stamp of his lowly
+origin."[110]
+
+What has been the fate of Darwin's doctrines since his great
+achievement? How have they been received and followed up by the
+scientific and lay world? And what do the successors of the mighty
+hero and genius think now in regard to the origin of the human race?
+
+At the present time we are incomparably more favourably placed than
+Darwin was for answering this question of all questions. We have at
+our command an incomparably greater wealth of material than he had at
+his disposal. And we are more fortunate than he in this respect, that
+we now know transition-forms which help to fill up the gap, still
+great, between the lowest human races and the highest apes. Let us
+consider for a little the more essential additions to our knowledge
+since the publication of _The Descent of Man_.
+
+Since that time our knowledge of animal embryos has increased
+enormously. While Darwin was obliged to content himself with comparing
+a human embryo with that of a dog, there are now available the
+youngest embryos of monkeys of all possible groups (Orang, Gibbon,
+Semnopithecus, Macacus), thanks to Selenka's most successful tour in
+the East Indies in search of such material. We can now compare
+corresponding stages of the lower monkeys and of the Anthropoid apes
+with human embryos, and convince ourselves of their great resemblance
+to one another, thus strengthening enormously the armour prepared by
+Darwin in defence of his view on man's nearest relatives. It may be
+said that Selenka's material fills up the blanks in Darwin's array of
+proofs in the most satisfactory manner.
+
+The deepening of our knowledge of comparative anatomy also gives us much
+surer foundations than those on which Darwin was obliged to build. Just of
+late there have been many workers in the domain of the anatomy of apes and
+lemurs, and their investigations extend to the most different organs. Our
+knowledge of fossil apes and lemurs has also become much wider and more
+exact since Darwin's time: the fossil lemurs have been especially worked up
+by Cope, Forsyth Major, Ameghino, and others. Darwin knew very little about
+fossil monkeys. He mentions two or three anthropoid apes as occurring in
+the Miocene of Europe,[111] but only names _Dryopithecus_, the largest form
+from the Miocene of France. It was erroneously supposed that this form was
+related to _Hylobates_. We now know not only a form that actually stands
+near to the gibbon (_Pliopithecus_), and remains of other anthropoids
+(_Pliohylobates_ and the fossil chimpanzee, _Palaeopithecus_), but also
+several lower catarrhine monkeys, of which _Mesopithecus_, a form nearly
+related to the modern Sacred Monkeys (a species of _Semnopithecus_) and
+found in strata of the Miocene period in Greece, is the most important.
+Quite recently, too, Ameghino's investigations have made us acquainted with
+fossil monkeys from South America (_Anthropops_, _Homunculus_), which,
+according to their discoverer, are to be regarded as in the line of human
+descent.
+
+What Darwin missed most of all--intermediate forms between apes and
+man--has been recently furnished. E. Dubois, as is well known,
+discovered in 1893, near Trinil in Java, in the alluvial deposits of
+the river Bengawan, an important form represented by a skull-cap, some
+molars, and a femur. His opinion--much disputed as it has been--that
+in this form, which he named _Pithecanthropus_, he has found a
+long-desired transition-form is shared by the present writer. And
+although the geological age of these fossils, which, according to
+Dubois, belong to the uppermost Tertiary series, the Pliocene has
+recently been fixed at a later date (the older Diluvium), the
+_morphological value_ of these interesting remains, that is, the
+intermediate position of _Pithecanthropus_, still holds good. Volz
+says with justice,[112] that even if _Pithecanthropus_ is not _the_
+missing link, it is undoubtedly _a_ missing link.
+
+As on the one hand there has been found in _Pithecanthropus_ a form
+which, though intermediate between apes and man, is nevertheless more
+closely allied to the apes, so on the other hand, much progress has
+been made since Darwin's day in the discovery and description of the
+oldest human remains. Since the famous roof of a skull and the bones
+of the extremities belonging to it were found in 1856 in the
+Neandertal near Düsseldorf, the most varied judgments have been
+expressed in regard to the significance of the remains and of the
+skull in particular. In Darwin's _Descent of Man_ there is only a
+passing allusion to them[113] in connection with the discussion of the
+skull-capacity, although the investigations of Schaaffhausen, King,
+and Huxley were then known. I believe I have shown, in a series of
+papers, that the skull in question belongs to a form different from
+any of the races of man now living, and, with King and Cope, I regard
+it as at least a different species from living man, and have therefore
+designated it _Homo primigenius_. The form unquestionably belongs to
+the older Diluvium, and in the later Diluvium human forms already
+appear, which agree in all essential points with existing human races.
+
+As far back as 1886 the value of the Neandertal skull was greatly
+enhanced by Fraipont's discovery of two skulls and skeletons from Spy
+in Belgium. These are excellently described by their discoverer,[114]
+and are regarded as belonging to the same group of forms as the
+Neandertal remains. In 1899 and the following years came the discovery
+by Gorjanovi�-Kramberger of different skeletal parts of at least
+ten individuals in a cave near Krapina in Croatia.[115] It is in
+particular the form of the lower jaw which is different from that of
+all recent races of man, and which clearly indicates the lowly
+position of _Homo primigenius_, while, on the other hand, the
+long-known skull from Gibraltar, which I[116] have referred to _Homo
+primigenius_, and which has lately been examined in detail by
+Sollas,[117] has made us acquainted with the surprising shape of the
+eye-orbit, of the nose, and of the whole upper part of the face.
+Isolated lower jaws found at La Naulette in Belgium, and at Malarnaud
+in France, increase our material which is now as abundant as could be
+desired. The most recent discovery of all is that of a skull dug up in
+August of this year [1908] by Klaatsch and Hauser in the lower grotto
+of the Le Moustier in Southern France, but this skull has not yet been
+fully described. Thus _Homo primigenius_ must also be regarded as
+occupying a position in the gap existing between the highest apes and
+the lowest human races, _Pithecanthropus_, standing in the lower part
+of it, and _Homo primigenius_ in the higher, near man. In order to
+prevent misunderstanding, I should like here to emphasise that in
+arranging this structural series--anthropoid apes, _Pithecanthropus_,
+_Homo primigenius_, _Homo sapiens_--I have no intention of
+establishing it as a direct genealogical series. I shall have
+something to say in regard to the genetic relations of these forms,
+one to another, when discussing the different theories of descent
+current at the present day.[118]
+
+In quite a different domain from that of morphological relationship,
+namely in the physiological study of the blood, results have recently
+been gained which are of the highest importance to the doctrine of
+descent. Uhlenhuth, Nuttall, and others have established the fact that
+the blood-serum of a rabbit which has previously had human blood
+injected into it, forms a precipitate with human blood. This
+biological reaction was tried with a great variety of mammalian
+species, and it was found that those far removed from man gave no
+precipitate under these conditions. But as in other cases among
+mammals all nearly related forms yield an almost equally marked
+precipitate, so the serum of a rabbit treated with human blood and
+then added to the blood of an anthropoid ape gives _almost_ as marked
+a precipitate as in human blood; the reaction to the blood of the
+lower Eastern monkeys is weaker, that to the Western monkeys weaker
+still; indeed in this last case there is only a slight clouding after
+a considerable time and no actual precipitate. The blood of the
+Lemuridae (Nuttall) gives no reaction or an extremely weak one, that
+of the other mammals none whatever. We have in this not only a proof
+of the literal blood relationship between man and apes, but the degree
+of relationship with the different main groups of apes can be
+determined beyond possibility of mistake.
+
+Finally, it must be briefly mentioned that in regard to remains of
+human handicraft also, the material at our disposal has greatly
+increased of late years, that, as a result of this, the opinions of
+archaeologists have undergone many changes, and that, in particular,
+their views in regard to the age of the human race have been greatly
+influenced. There is a tendency at the present time to refer the
+origin of man back to Tertiary times. It is true that no remains of
+Tertiary man have been found, but flints have been discovered which,
+according to the opinion of most investigators, bear traces either of
+use, or of very primitive workmanship. Since Rutot's time, following
+Mortillet's example, investigators have called these "eoliths," and
+they have been traced back by Verworn to the Miocene of the Auvergne,
+and by Rutot even to the upper Oligocene. Although these eoliths are
+even nowadays the subject of many different views, the preoccupation
+with them has kept the problem of the age of the human race
+continually before us.
+
+Geology, too, has made great progress since the days of Darwin and
+Lyell, and has endeavoured with satisfactory results to arrange the
+human remains of the Diluvial period in chronological order (Penck). I
+do not intend to enter upon the question of the primitive home of the
+human race; since the space at my disposal will not allow of my
+touching even very briefly upon all the departments of science which
+are concerned in the problem of the descent of man. How Darwin would
+have rejoiced over each of the discoveries here briefly outlined! What
+use he would have made of the new and precious material, which would
+have prevented the discouragement from which he suffered when
+preparing the second edition of _The Descent of Man_! But it was not
+granted to him to see this progress towards filling up the gaps in his
+edifice of which he was so painfully conscious.
+
+He did, however, have the satisfaction of seeing his ideas steadily
+gaining ground, notwithstanding much hostility and deep-rooted
+prejudice. Even in the years between the appearance of _The Origin of
+Species_ and of the first edition of the _Descent_, the idea of a
+natural descent of man, which was only briefly indicated in the work
+of 1859, had been eagerly welcomed in some quarters. It has been
+already pointed out how brilliantly Huxley contributed to the defence
+and diffusion of Darwin's doctrines, and how in _Man's Place in
+Nature_ he has given us a classic work as a foundation for the
+doctrine of the descent of man. As Huxley was Darwin's champion in
+England, so in Germany Carl Vogt, in particular, made himself master
+of the Darwinian ideas. But above all it was Haeckel who, in energy,
+eagerness for battle, and knowledge may be placed side by side with
+Huxley, who took over the leadership in the controversy over the new
+conception of the universe. As far back as 1866, in his _Generelle
+Morphologie_, he had inquired minutely into the question of the
+descent of man, and not content with urging merely the general theory
+of descent from lower animal forms, he drew up for the first time
+genealogical trees showing the close structural relationships of the
+different animal groups; the last of these illustrated the
+relationships of Mammals, and among them of all groups of the
+Primates, including man. It was Haeckel's genealogical trees that
+formed the basis of the special discussion of the relationships of
+man, in the sixth chapter of Darwin's _Descent of Man_.
+
+In the last section of this essay I shall return to Haeckel's
+conception of the special descent of man, the main features of which
+he still upholds, and rightly so. Haeckel has contributed more than
+any one else to the spread of the Darwinian doctrine.
+
+I can only allow myself a few words as to the spread of the theory of
+the natural descent of man in other countries. The Parisian
+anthropological school, founded and guided by the genius of Broca,
+took up the idea of the descent of man, and made many notable
+contributions to it (Broca, Manouvrier, Mahoudeau, Deniker and
+others). In England itself Darwin's work did not die. Huxley took care
+of that, for he, with his lofty and unprejudiced mind, dominated and
+inspired English biology until his death on June 29, 1895. He had the
+satisfaction shortly before his death of learning of Dubois'
+discovery, which he illustrated by a humourous sketch.[119] But there
+are still many followers in Darwin's footsteps in England. Keane has
+worked at the special genealogical tree of the Primates; Keith has
+inquired which of the anthropoid apes has the greatest number of
+characters in common with man; Morris concerns himself with the
+evolution of man in general, especially with his acquisition of the
+erect position. The recent discoveries of _Pithecanthropus_ and _Homo
+primigenius_ are being vigorously discussed; but the present writer is
+not in a position to form an opinion of the extent to which the idea
+of descent has penetrated throughout England generally.
+
+In Italy independent work in the domain of the descent of man is being
+produced, especially by Morselli; with him are associated, in the
+investigation of related problems, Sergi and Giuffrida-Ruggeri. From
+the ranks of American investigators we may single out in particular
+the eminent geologist Cope, who championed with much decision the idea
+of the specific difference of _Homo neandertalensis_ (_primigenius_)
+and maintained a more direct descent of man from the fossil Lemuridae.
+In South America too, in Argentina, new life is stirring in this
+department of science. Ameghino in Buenos Ayres has awakened the
+fossil primates of the Pampas formation to new life; he even believes
+that in his _Tetraprothomo_, represented by a femur, he has discovered
+a direct ancestor of man. Lehmann-Nitsche is working at the other side
+of the gulf between apes and man, and he describes a remarkable first
+cervical vertebra (atlas) from Monte Hermoso as belonging to a form
+which may bear the same relation to _Homo sapiens_ in South America as
+_Homo primigenius_ does in the Old World. After a minute investigation
+he establishes a human species _Homo neogaeus_, while Ameghino
+ascribes this atlas vertebra to his _Tetraprothomo_.
+
+Thus throughout the whole scientific world there is arising a new
+life, an eager endeavour to get nearer to Huxley's _problema maximum_,
+to penetrate more deeply into the origin of the human race. There are
+to-day very few experts in anatomy and zoology who deny the animal
+descent of man in general. Religious considerations, old prejudices,
+the reluctance to accept man, who so far surpasses mentally all other
+creatures, as descended from "soulless" animals, prevent a few
+investigators from giving full adherence to the doctrine. But there
+are very few of these who still postulate a special act of creation
+for man. Although the majority of experts in anatomy and zoology
+accept unconditionally the descent of man from lower forms, there is
+much diversity of opinion among them in regard to the special line of
+descent.
+
+In trying to establish any special hypothesis of descent, whether by
+the graphic method of drawing up genealogical trees or otherwise, let
+us always bear in mind Darwin's words[120] and use them as a critical
+guiding line: "As we have no record of the lines of descent, the
+pedigree can be discovered only by observing the degrees of
+resemblance between the beings which are to be classed." Darwin
+carries this further by stating "that resemblances in several
+unimportant structures, in useless and rudimentary organs, or not now
+functionally active, or in an embryological condition, are by far the
+most serviceable for classification."[121] It has also to be
+remembered that _numerous_ separate points of agreement are of much
+greater importance than the amount of similarity or dissimilarity in a
+few points.
+
+The hypotheses as to descent current at the present day may be divided
+into two main groups. The first group seeks for the roots of the human
+race not among any of the families of the apes--the anatomically
+nearest forms--nor among their very similar but less specialised
+ancestral forms, the fossil representatives of which we can know only
+in part, but, setting the monkeys on one side, it seeks for them lower
+down among the fossil Eocene Pseudo-lemuridae or Lemuridae (Cope), or
+even among the primitive pentadactylous Eocene forms, which may
+either have led directly to the evolution of man (Adloff), or have
+given rise to an ancestral form common to apes and men (Klaatsch,[122]
+Giuffrida-Ruggeri). The common ancestral form, from which man and apes
+are thus supposed to have arisen independently, may explain the
+numerous resemblances which actually exist between them. That is to
+say, all the characters upon which the great structural resemblance
+between apes and man depends must have been present in their common
+ancestor. Let us take an example of such a common character. The bony
+external ear-passage is in general as highly developed in the lower
+Eastern monkeys and the anthropoid apes as in man. This character
+must, therefore, have already been present in the common primitive
+form. In that case it is not easy to understand why the Western
+monkeys have not also inherited the character, instead of possessing
+only a tympanic ring. But it becomes more intelligible if we assume
+that forms with a primitive tympanic ring were the original type, and
+that from these were evolved, on the one hand, the existing New World
+monkeys with persistent tympanic ring, and on the other an ancestral
+form common to the lower Old World monkeys, the anthropoid apes and
+man. For man shares with these the character in question, and it is
+also one of the "unimportant" characters required by Darwin. Thus we
+have two divergent lines arising from the ancestral form, the Western
+monkeys (Platyrrhine) on the one hand, and an ancestral form common to
+the lower Eastern monkeys, the anthropoid apes, and man, on the other.
+But considerations similar to those which showed it to be impossible
+that man should have developed from an ancestor common to him and the
+monkeys, yet outside of and parallel with these, may be urged also
+against the likelihood of a parallel evolution of the lower Eastern
+monkeys, the anthropoid apes, and man. The anthropoid apes have in
+common with man many characters which are not present in the lower
+Old World monkeys. These characters must therefore have been present
+in the ancestral form common to the three groups. But here, again, it
+is difficult to understand why the lower Eastern monkeys should not
+also have inherited these characters. As this is not the case, there
+remains no alternative but to assume divergent evolution from an
+indifferent form. The lower Eastern monkeys are carrying on the
+evolution in one direction--I might almost say towards a blind
+alley--while anthropoids and men have struck out a progressive path,
+at first in common, which explains the many points of resemblance
+between them, without regarding man as derived directly from the
+anthropoids. Their many striking points of agreement indicate a common
+descent, and cannot be explained as phenomena of convergence.
+
+I believe I have shown in the above sketch that a theory which derives
+man directly from lower forms without regarding apes as
+transition-types leads _ad absurdum_. The close structural
+relationship between man and monkeys can only be understood if both
+are brought into the same line of evolution. To trace man's line of
+descent directly back to the old Eocene mammals, alongside of, but
+with no relation to these very similar forms, is to abandon the method
+of exact comparison, which, as Darwin rightly recognised, alone
+justifies us in drawing up genealogical trees on the basis of
+resemblances and differences. The farther down we go the more does the
+ground slip from beneath our feet. Even the Lemuridae show very
+numerous divergent conditions, much more so the Eocene mammals
+(Creodonta, Condylarthra), the chief resemblance of which to man
+consists in the possession of pentadactylous hands and feet! Thus the
+farther course of the line of descent disappears in the darkness of
+the ancestry of the mammals. With just as much reason we might pass by
+the Vertebrates altogether, and go back to the lower Invertebrates,
+but in that case it would be much easier to say that man has arisen
+independently, and has evolved, without relation to any animals, from
+the lowest primitive form to his present isolated and dominant
+position. But this would be to deny all value to classification, which
+must after all be the ultimate basis of a genealogical tree. We can,
+as Darwin rightly observed, only infer the line of descent from the
+degree of resemblance between single forms. If we regard man as
+directly derived from primitive forms very far back, we have no way of
+explaining the many points of agreement between him and the monkeys in
+general, and the anthropoid apes in particular. These must remain an
+inexplicable marvel.
+
+I have thus, I trust, shown that the first class of special theories
+of descent, which assumes that man has developed, parallel with the
+monkeys, but without relation to them, from very low primitive forms
+cannot be upheld, because it fails to take into account the close
+structural affinity of man and monkeys. I cannot but regard this
+hypothesis as lamentably retrograde, for it makes impossible any
+application of the facts that have been discovered in the course of
+the anatomical and embryological study of man and monkeys, and indeed
+prejudges investigations of that class as pointless. The whole method
+is perverted; an unjustifiable theory of descent is first formulated
+with the aid of the imagination, and then we are asked to declare that
+all structural relations between man and monkeys, and between the
+different groups of the latter, are valueless,--the fact being that
+they are the only true basis on which a genealogical tree can be
+constructed.
+
+So much for this most modern method of classification, which has
+probably found adherents because it would deliver us from the
+relationship to apes which many people so much dislike. In contrast to
+it we have the second class of special hypotheses of descent, which
+keeps strictly to the nearest structural relationship. This is the
+only basis that justifies the drawing up of a special hypothesis of
+descent. If this fundamental proposition be recognised, it will be
+admitted that the doctrine of special descent upheld by Haeckel, and
+set forth in Darwin's _Descent of Man_, is still valid to-day. In the
+genealogical tree, man's place is quite close to the anthropoid apes;
+these again have as their nearest relatives the lower Old World
+monkeys, and their progenitors must be sought among the less
+differentiated Platyrrhine monkeys, whose most important characters
+have been handed on to the present day New World monkeys. How the
+different genera are to be arranged within the general scheme
+indicated depends in the main on the classificatory value attributed
+to individual characters. This is particularly true in regard to
+_Pithecanthropus_, which I consider as the root of a branch which has
+sprung from the anthropoid ape root and has led up to man; the latter
+I have designated the family of the Hominidae.
+
+For the rest, there are, as we have said, various possible ways of
+constructing the narrower genealogy within the limits of this branch
+including men and apes, and these methods will probably continue to
+change with the accumulation of new facts. Haeckel himself has
+modified his genealogical tree of the Primates in certain details
+since the publication of his _Generelle Morphologie_ in 1866, but its
+general basis remains the same.[123] All the special genealogical
+trees drawn up on the lines laid down by Haeckel and Darwin--and that
+of Dubois may be specially mentioned--are based, in general, on the
+close relationship of monkeys and men, although they may vary in
+detail. Various hypotheses have been formulated on these lines, with
+special reference to the evolution of man. _Pithecanthropus_ is
+regarded by some authorities as the direct ancestor of man, by others
+as a side-track failure in the attempt at the evolution of man. The
+problem of the monophyletic or polyphyletic origin of the human race
+has also been much discussed. Sergi[124] inclines towards the
+assumption of a polyphyletic origin of the three main races of man,
+the African primitive form of which has given rise also to the
+gorilla and chimpanzee, the Asiatic to the Orang, the Gibbon, and
+_Pithecanthropus_. Kollmann regards existing human races as derived
+from small primitive races (pigmies), and considers that _Homo
+primigenius_ must have arisen in a secondary and degenerative manner.
+
+But this is not the place, nor have I the space to criticise the
+various special theories of descent. One, however, must receive
+particular notice. According to Ameghino, the South American monkeys
+(_Pitheculites_) from the oldest Tertiary of the Pampas are the forms
+from which have arisen the existing American monkeys on the one hand,
+and on the other, the extinct South American Homunculidae, which are
+also small forms. From these last, anthropoid apes and man have, he
+believes, been evolved. Among the progenitors of man, Ameghino reckons
+the form discovered by him (_Tetraprothomo_), from which a South
+American primitive man, _Homo pampaeus_, might be directly evolved,
+while on the other hand all the lower Old World monkeys may have
+arisen from older fossil South American forms (Clenialitidae), the
+distribution of which may be explained by the bridge formerly existing
+between South America and Africa, as may be the derivation of all
+existing human races from _Homo pampaeus_.[125] The fossil forms
+discovered by Ameghino deserve the most minute investigation, as does
+also the fossil man from South America of which Lehmann-Nitsche[126]
+has made a thorough study.
+
+It is obvious that, notwithstanding the necessity for fitting man's
+line of descent into the genealogical tree of the Primates, especially
+the apes, opinions in regard to it differ greatly in detail. This
+could not be otherwise, since the different Primate forms, especially
+the fossile forms, are still far from being exhaustively known. But
+one thing remains certain,--the idea of the close relationship between
+man and monkeys set forth in Darwin's _Descent of Man_. Only those who
+deny the many points of agreement, the sole basis of classification,
+and thus of a natural genealogical tree, can look upon the position of
+Darwin and Haeckel as antiquated, or as standing on an insufficient
+foundation. For such a genealogical tree is nothing more than a
+summarised representation of what is known in regard to the degree of
+resemblance between the different forms.
+
+Darwin's work in regard to the descent of man has not been surpassed;
+the more we immerse ourselves in the study of the structural
+relationships between apes and man, the more is our path illumined by
+the clear light radiating from him, and through his calm and
+deliberate investigation, based on a mass of material in the
+accumulation of which he has never had an equal. Darwin's fame will be
+bound up for all time with the unprejudiced investigation of the
+question of all questions, the descent of the human race.
+
+FOOTNOTES:
+
+[Footnote 75: _Life and Letters of Thomas Henry Huxley_, Vol. I. p.
+171, London, 1900.]
+
+[Footnote 76: _Ibid._, p. 363.]
+
+[Footnote 77: No italics in original.]
+
+[Footnote 78: _Life and Letters of Charles Darwin_, Vol. I. p. 93.]
+
+[Footnote 79: _Ibid._ Vol. II. p. 263.]
+
+[Footnote 80: _Ibid._ Vol. I. p. 94.]
+
+[Footnote 81: _Life and Letters_, Vol. III. p. 175.]
+
+[Footnote 82: _Ibid._ Vol. II. p. 109.]
+
+[Footnote 83: _Ibid._ Vol. III. p. 112.]
+
+[Footnote 84: _Ibid._ Vol. I. pp. 304-317.]
+
+[Footnote 85: _Life and Letters_, Vol. I. p. 309.]
+
+[Footnote 86: _Loc. cit._ p. 313.]
+
+[Footnote 87: _Ibid._ Vol. II. p. 310.]
+
+[Footnote 88: _Ibid._ Vol. III. p. 236. ["C. Ridley," Mr. Francis
+Darwin points out to me, should be H. N. Ridley. A.C.S.]]
+
+[Footnote 89: _Descent of Man_ (Popular Edit., 1901), fig. 1, p. 14.]
+
+[Footnote 90: G. Schwalbe, "Das Darwin'sche Spitzohr beim menschlichen
+Embryo," _Anatom. Anzeiger_, 1889, pp. 176-189, and other papers.]
+
+[Footnote 91: _Descent of Man_, fig. 3, p. 24.]
+
+[Footnote 92: _Descent of man_, p. 6.]
+
+[Footnote 93: _Ibid._ p. 54.]
+
+[Footnote 94: _Descent of Man_, p. 92.]
+
+[Footnote 95: _Ibid._ p. 100.]
+
+[Footnote 96: _Life and letters_, Vol. II. p. 161, June 22, 1859.]
+
+[Footnote 97: _Ibid._ Vol. III. p. 15, March 17, 1863.]
+
+[Footnote 98: _Descent of Man_, p. 132.]
+
+[Footnote 99: _Ibid._ pp. 136, 137.]
+
+[Footnote 100: _Ibid._ p. 143.]
+
+[Footnote 101: _Ibid._ p. 193.]
+
+[Footnote 102: _Descent of Man_, p. 231.]
+
+[Footnote 103: _Descent of Man_, p. 308.]
+
+[Footnote 104: _Life and Letters_, Vol. II. p. 23.]
+
+[Footnote 105: _Loc. cit._ p. 39.]
+
+[Footnote 106: _Loc. cit._ (1856), p. 82.]
+
+[Footnote 107: _Ibid._ Vol. III p. 159.]
+
+[Footnote 108: _Descent of Man_, p. 924.]
+
+[Footnote 109: "Die Hautfarbe des Menschen," _Mitteilungen der
+Anthropologischen Gessellschaft in Wien_, Vol. XXXIV. pp. 331-352.]
+
+[Footnote 110: _Ibid._ p. 947.]
+
+[Footnote 111: _Descent of Man_, p. 240.]
+
+[Footnote 112: "Das geologische Alter der Pithecanthropus-Schichten
+bei Trinil, Ost-Java." _Neues Jahrb. f. Mineralogie_. Festband, 1907.]
+
+[Footnote 113: _Descent of Man_, p. 82.]
+
+[Footnote 114: "La race humaine de Néanderthal ou de Canstatt en
+Belgique." _Arch. de Biologie_, VII. 1887.]
+
+[Footnote 115: Gorjanovi�-Kramberger. _Der diluviale Mensch van
+Krapina in Kroatien_, 1906.]
+
+[Footnote 116: _Studien zur Vorgeschichte des Menschen_, 1906, pp. 154
+ff.]
+
+[Footnote 117: "On the cranial and facial characters of the Neandertal
+Race." _Trans. R. Soc._ London, vol. 199, 1908, p. 281.]
+
+[Footnote 118: Since this essay was written Schoetensack has
+discovered near Heidelberg and briefly described an exceedingly
+interesting lower jaw from rocks between the Pliocene and Diluvial
+beds. This exhibits interesting differences from the forms of lower
+jaw of _Homo primigenius_. (Schoetensack, _Der Unterkiefer des Homo
+heidelbergensis_, Leipzig, 1908.) G. S.]
+
+[Footnote 119: _Life and Letters of Thomas Henry Huxley_, Vol. II. p.
+394.]
+
+[Footnote 120: _Descent of Man_, p. 229.]
+
+[Footnote 121: _Loc. cit._]
+
+[Footnote 122: Klaatsch in his last publications speaks in the main
+only of an ancestral form common to men and anthropoid apes.]
+
+[Footnote 123: Haeckels latest genealogical tree is to be found in his
+most recent work, _Unsere Ahnenreihe_. Jena, 1908.]
+
+[Footnote 124: Sergi, G. _Europa_, 1908.]
+
+[Footnote 125: _See_ Ameghino's latest paper, "_Notas preliminaries
+sobre el Tetraprothomo argentinus_," etc. _Anales del Museo nacional
+de Buenos Aires_, XVI. pp. 107-242, 1907.]
+
+[Footnote 126: "Nouvelles recherches sur la formation pampéenne et
+l'homme fossile de la République Argentine." _Rivista del Museo de la
+Plata_, T. XIV. pp. 193-488.]
+
+
+
+
+V
+
+CHARLES DARWIN AS AN ANTHROPOLOGIST
+
+BY ERNST HAECKEL
+
+_Professor of Zoology in the University of Jena_
+
+
+The great advance that anthropology has made in the second half of the
+nineteenth century is due, in the first place, to Darwin's discovery
+of the origin of man. No other problem in the whole field of research
+is so momentous as that of "Man's place in nature," which was justly
+described by Huxley (1863) as the most fundamental of all questions.
+Yet the scientific solution of this problem was impossible until the
+theory of descent had been established.
+
+It is now a hundred years since the great French biologist Jean
+Lamarck published his _Philosophie Zoologique_. By a remarkable
+coincidence the year in which that work was issued, 1809, was the year
+of the birth of his most distinguished successor, Charles Darwin.
+Lamarck had already recognised that the descent of man from a series
+of other Vertebrates--that is, from a series of Ape-like Primates--was
+essentially involved in the general theory of transformation which he
+had erected on a broad inductive basis; and he had sufficient
+penetration to detect the agencies that had been at work in the
+evolution of the erect bimanous man from the arboreal and quadrumanous
+ape. He had, however, few empirical arguments to advance in support of
+his hypothesis, and it could not be established until the further
+development of the biological sciences--the founding of comparative
+embryology by Baer (1828) and of the cell-theory by Schleiden and
+Schwann (1838), the advance of physiology under Johannes Müller
+(1833), and the enormous progress of palaeontology and comparative
+anatomy between 1820 and 1860--provided this necessary foundation.
+Darwin was the first to coordinate the ample results of these lines of
+research. With no less comprehensiveness than discrimination he
+consolidated them as a basis of a modified theory of descent, and
+associated with them his own theory of natural selection, which we
+take to be distinctive of "Darwinism" in the stricter sense. The
+illuminating truth of these cumulative arguments was so great in every
+branch of biology that, in spite of the most vehement opposition, the
+battle was won within a single decade, and Darwin secured the general
+admiration and recognition that had been denied to his forerunner,
+Lamarck, up to the hour of his death (1829).
+
+Before, however, we consider the momentous influence that Darwinism
+has had in anthropology, we shall find it useful to glance at its
+history in the course of the last half century, and notice the various
+theories that have contributed to its advance. The first attempt to
+give extensive expression to the reform of biology by Darwin's work
+will be found in my _Generelle Morphologie_ (1866)[127] which was
+followed by a more popular treatment of the subject in my _Natürliche
+Schöpfungsgeschichte_ (1868),[128] a compilation from the earlier
+work. In the first volume of the _Generelle Morphologie_ I endeavoured
+to show the great importance of evolution in settling the fundamental
+questions of biological philosophy, especially in regard to
+comparative anatomy. In the second volume I dealt broadly with the
+principle of evolution, distinguishing ontogeny and phylogeny as its
+two coordinate main branches, and associating the two in the
+Biogenetic Law. The Law may be formulated thus: "Ontogeny (embryology
+or the development of the individual) is a concise and compressed
+recapitulation of phylogeny (the palaeontological or genealogical
+series) conditioned by laws of heredity and adaptation." The
+"Systematic introduction to general evolution," with which the second
+volume of the _Generelle Morphologie_ opens, was the first attempt to
+draw up a natural system of organisms (in harmony with the principles
+of Lamarck and Darwin) in the form of a hypothetical pedigree, and was
+provisionally set forth in eight genealogical tables.
+
+In the nineteenth chapter of the _Generelle Morphologie_--a part of
+which has been republished, without any alteration, after a lapse of
+forty years--I made a critical study of Lamarck's theory of descent
+and of Darwin's theory of selection, and endeavoured to bring the
+complex phenomena of heredity and adaptation under definite laws for
+the first time. Heredity I divided into conservative and progressive:
+adaptation into indirect (or potential) and direct (or actual). I then
+found it possible to give some explanation of the correlation of the
+two physiological functions in the struggle for life (selection), and
+to indicate the important laws of divergence (or differentiation) and
+complexity (or division of labor), which are the direct and inevitable
+outcome of selection. Finally, I marked off dysteleology as the
+science of the aimless (vestigial, abortive, atrophied, and useless)
+organs and parts of the body. In all this I worked from a strictly
+monistic standpoint, and sought to explain all biological phenomena on
+the mechanical and naturalistic lines that had long been recognised in
+the study of inorganic nature. Then (1866), as now, being convinced of
+the unity of nature, the fundamental identity of the agencies at work
+in the inorganic and the organic worlds, I discarded vitalism,
+teleology, and all hypotheses of a mystic character.
+
+It was clear from the first that it was essential, in the monistic
+conception of evolution, to distinguish between the laws of
+conservative and progressive heredity. Conservative heredity maintains
+from generation to generation the enduring characters of the species.
+Each organism transmits to its descendants a part of the morphological
+and physiological qualities that it has received from its parents and
+ancestors. On the other hand, progressive heredity brings new
+characters to the species--characters that were not found in preceding
+generations. Each organism may transmit to its offspring a part of the
+morphological and physiological features that it has itself acquired,
+by adaptation, in the course of its individual career, through the use
+or disuse of particular organs, the influence of environment, climate,
+nutrition, etc. At that time I gave the name of "progressive heredity"
+to this inheritance of acquired characters, as a short and convenient
+expression, but have since changed the term to "transformative
+heredity" (as distinguished from conservative). This term is
+preferable, as inherited regressive modifications (degeneration,
+retrograde metamorphosis, etc.) come under the same head.
+
+Transformative heredity--or the transmission of acquired
+characters--is one of the most important principles in evolutionary
+science. Unless we admit it most of the facts of comparative anatomy
+and physiology are inexplicable. That was the conviction of Darwin no
+less than of Lamarck, of Spencer as well as Virchow, of Huxley as well
+as Gegenbaur, indeed of the great majority of speculative biologists.
+This fundamental principle was for the first time called in question
+and assailed in 1885 by August Weismann of Freiburg, the eminent
+zoologist to whom the theory of evolution owes a great deal of
+valuable support, and who has attained distinction by his extension of
+the theory of selection. In explanation of the phenomena of heredity
+he introduced a new theory, the "theory of the continuity of the
+germ-plasm." According to him the living substance in all organisms
+consists of two quite distinct kinds of plasm, somatic and germinal.
+The permanent germ-plasm, or the active substance of the two
+germ-cells (egg-cell and sperm-cell), passes unchanged through a
+series of generations, and is not affected by environmental
+influences. The environment modifies only the soma-plasm, the organs
+and tissues of the body. The modifications that these parts undergo
+through the influence of the environment or their own activity (use
+and habit), do not affect the germ-plasm, and cannot therefore be
+transmitted.
+
+This theory of the continuity of the germ-plasm has been expounded by
+Weismann during the last twenty-four years in a number of able
+volumes, and is regarded by many biologists, such as Mr. Francis
+Galton, Sir E. Ray Lankester, and Professor J. Arthur Thomson (who has
+recently made a thorough-going defence of it in his important work
+_Heredity_),[129] as the most striking advance in evolutionary
+science. On the other hand, the theory has been rejected by Herbert
+Spencer, Sir W. Turner, Gegenbaur, Kölliker, Hertwig, and many others.
+For my part I have, with all respect for the distinguished Darwinian,
+contested the theory from the first, because its whole foundation
+seems to me erroneous, and its deductions do not seem to be in accord
+with the main facts of comparative morphology and physiology.
+Weismann's theory in its entirety is a finely conceived molecular
+hypothesis, but it is devoid of empirical basis. The notion of the
+absolute and permanent independence of the germ-plasm, as
+distinguished from the soma-plasm, is purely speculative; as is also
+the theory of germinal selection. The determinants, ids, and idants,
+are purely hypothetical elements. The experiments that have been
+devised to demonstrate their existence really prove nothing.
+
+It seems to me quite improper to describe this hypothetical structure
+as "Neodarwinism." Darwin was just as convinced as Lamarck of the
+transmission of acquired characters and its great importance in the
+scheme of evolution. I had the good fortune to visit Darwin at Down
+three times and discuss with him the main principles of his system,
+and on each occasion we were fully agreed as to the incalculable
+importance of what I may call transformative inheritance. It is only
+proper to point out that Weismann's theory of the germ-plasm is in
+express contradiction to the fundamental principles of Darwin and
+Lamarck. Nor is it more acceptable in what one may call its
+"ultradarwinism"--the idea that the theory of selection explains
+everything in the evolution of the organic world. This belief in the
+"omnipotence of natural selection" was not shared by Darwin himself.
+Assuredly, I regard it as of the utmost value, as the process of
+natural selection through the struggle for life affords an explanation
+of the mechanical origin of the adapted organisation. It solves the
+great problem: how could the finely adapted structure of the animal or
+plant body be formed unless it was built on a preconceived plan? It
+thus enables us to dispense with the teleology of the metaphysician
+and the dualist, and to set aside the old mythological and poetic
+legends of creation. The idea had occurred in vague form to the great
+Empedocles 2000 years before the time of Darwin, but it was reserved
+for modern research to give it ample expression. Nevertheless, natural
+selection does not of itself give the solution of all our evolutionary
+problems. It has to be taken in conjunction with the transformism of
+Lamarck, with which it is in complete harmony.
+
+The monumental greatness of Charles Darwin, who surpasses every other
+student of science in the nineteenth century by the loftiness of his
+monistic conception of nature and the progressive influence of his
+ideas, is perhaps best seen in the fact that not one of his many
+successors has succeeded in modifying his theory of descent in any
+essential point or in discovering an entirely new standpoint in the
+interpretation of the organic world. Neither Nägeli nor Weismann,
+neither De Vries nor Roux, has done this. Nägeli, in his
+_Mechanisch-Physiologische Theorie der Abstammungslehre_[130] which is
+to a great extent in agreement with Weismann, constructed a theory of
+the idioplasm, that represents it (like the germ-plasm) as developing
+continuously in a definite direction from internal causes. But his
+internal "principle of progress" is at the bottom just as teleological
+as the vital force of the Vitalists, and the micella structure of the
+idioplasm is just as hypothetical as the "dominant" structure of the
+germ-plasm. In 1889 Moritz Wagner sought to explain the origin of
+species by migration and isolation, and on that basis constructed a
+special "migration-theory." This, however, is not out of harmony with
+the theory of selection. It merely elevates one single factor in the
+theory to a predominant position. Isolation is only a special case of
+selection, as I had pointed out in the fifteenth chapter of my
+_Natural history of creation_. The "mutation-theory" of De Vries,[131]
+that would explain the origin of species by sudden and saltatory
+variations rather than by gradual modification, is regarded by many
+botanists as a great step in advance, but it is generally rejected by
+zoologists. It affords no explanation of the facts of adaptation, and
+has no causal value.
+
+Much more important than these theories is that of Wilhelm Roux[132]
+of "the struggle of parts within the organism, a supplementation of
+the theory of mechanical adaptation." He explains the functional
+autoformation of the purposive structure by a combination of Darwin's
+principle of selection with Lamarck's idea of transformative heredity,
+and applies the two in conjunction to the facts of histology. He lays
+stress on the significance of functional adaptation, which I had
+described in 1866, under the head of cumulative adaptation, as the
+most important factor in evolution. Pointing out its influence in the
+cell-life of the tissues, he puts "cellular selection" above "personal
+selection," and shows how the finest conceivable adaptations in the
+structure of the tissue may be brought about quite mechanically,
+without preconceived plan. This "mechanical teleology" is a valuable
+extension of Darwin's monistic principle of selection to the whole
+field of cellular physiology and histology, and is wholly destructive
+of dualistic vitalism.
+
+The most important advance that evolution has made since Darwin and
+the most valuable amplification of his theory of selection is, in my
+opinion, the work of Richard Semon: _Die Mneme als erhaltendes Prinzip
+im Wechsel des organischen Geschehens_.[133] He offers a psychological
+explanation of the facts of heredity by reducing them to a process of
+(unconscious) memory. The physiologist Ewald Hering had shown in 1870
+that memory must be regarded as a general function of organic matter,
+and that we are quite unable to explain the chief vital phenomena,
+especially those of reproduction and inheritance, unless we admit this
+unconscious memory. In my essay _Die Perigenesis der Plastidule_[134]
+I elaborated this far-reaching idea, and applied the physical
+principle of transmitted motion to the plastidules, or active
+molecules of plasm. I concluded that "heredity is the memory of the
+plastidules, and variability their power of comprehension." This
+"provisional attempt to give a mechanical explanation of the
+elementary processes of evolution" I afterwards extended by showing
+that sensitiveness is (as Carl Nägeli, Ernst Mach, and Albrecht Rau
+express it) a general quality of matter. This form of panpsychism
+finds its simplest expression in the "trinity of substance."
+
+To the two fundamental attributes that Spinoza ascribed to
+substance--Extension (matter as occupying space) and Cogitation
+(energy, force)--we now add the third fundamental quality of Psychoma
+(sensitiveness, soul). I further elaborated this trinitarian
+conception of substance in the nineteenth chapter of my _Die
+Lebenswunder_ (1904),[135] and it seems to me well calculated to
+afford a monistic solution of many of the antitheses of philosophy.
+
+This important Mneme-theory of Semon and the luminous physiological
+experiments and observations associated with it not only throw
+considerable light on transformative inheritance, but provide a sound
+physiological foundation for the biogenetic law. I had endeavoured to
+show in 1874, in the first chapter of my _Anthropogenie_,[136] that
+this fundamental law of organic evolution holds good generally, and
+that there is everywhere a direct causal connection between ontogeny
+and phylogeny. "Phylogenesis is the mechanical cause of ontogenesis;"
+in other words, "The evolution of the stem or race is--in accordance
+with the laws of heredity and adaptation--the real cause of all the
+changes that appear, in a condensed form, in the development of the
+individual organism from the ovum, in either the embryo or the larva."
+
+It is now fifty years since Charles Darwin pointed out, in the
+thirteenth chapter of his epoch-making _Origin of Species_, the
+fundamental importance of embryology in connection with his theory of
+descent:
+
+"The leading facts in embryology, which are second to none in
+importance, are explained on the principle of variations in the many
+descendants from some one ancient progenitor, having appeared at a not
+very early period of life, and having been inherited at a
+corresponding period."[137]
+
+He then shows that the striking resemblance of the embryos and larvae
+of closely related animals, which in the mature stage belong to widely
+different species and genera, can only be explained by their descent
+from a common progenitor. Fritz Müller made a closer study of these
+important phenomena in the instructive instance of the Crustacean
+larva, as given in his able work _Für Darwin_[138] (1864). I then, in
+1872, extended the range so as to include all animals (with the
+exception of the unicellular Protozoa) and showed, by means of the
+theory of the Gastraea, that all multicellular, tissue-forming
+animals--all the Metazoa--develop in essentially the same way from the
+primary germ-layers.
+
+I conceived the embryonic form, in which the whole structure consists of
+only two layers of cells, and is known as the gastrula, to be the
+ontogenetic recapitulation, maintained by tenacious heredity, of a
+primitive common progenitor of all the Metazoa, the Gastraea. At a later
+date (1895) Monticelli discovered that this conjectural ancestral form is
+still preserved in certain primitive Coelenterata--Pemmatodiscus,
+Kunstleria, and the nearly-related Orthonectida.
+
+The general application of the biogenetic law to all classes of
+animals and plants has been proved in my _Systematische
+Phylogenie_.[139] It has, however, been frequently challenged, both by
+botanists and zoologists, chiefly owing to the fact that many have
+failed to distinguish its two essential elements, palingenesis and
+cenogenesis. As early as 1874 I had emphasised, in the first chapter
+of my _Evolution of Man_, the importance of discriminating carefully
+between these two sets of phenomena:
+
+"In the evolutionary appreciation of the facts of embryology we must
+take particular care to distinguish sharply and clearly between the
+primary, palingenetic evolutionary processes and the secondary,
+cenogenetic processes. The palingenetic phenomena, or embryonic
+_recapitulations_, are due to heredity, to the transmission of
+characters from one generation to another. They enable us to draw
+direct inferences in regard to corresponding structures in the
+development of the species (e.g. the chorda or the branchial arches in
+all vertebrate embryos). The cenogenetic phenomena, on the other hand,
+or the embryonic _variations_, cannot be traced to inheritance from a
+mature ancestor, but are due to the adaption of the embryo or the
+larva to certain conditions of its individual development (e.g. the
+amnion, the allantois, and the vitelline arteries in the embryos of
+the higher vertebrates). These cenogenetic phenomena are later
+additions; we must not infer from them that there were corresponding
+processes in the ancestral history, and hence they are apt to
+mislead."
+
+The fundamental importance of these facts of comparative anatomy,
+atavism, and the rudimentary organs, was pointed out by Darwin in the
+first part of his classic work, _The Descent of Man and Selection in
+Relation to Sex_ (1871).[140] In the "General summary and conclusion"
+(chap. xxi.) he was able to say, with perfect justice: "He who is not
+content to look, like a savage, at the phenomena of nature as
+disconnected, cannot any longer believe that man is the work of a
+separate act of creation. He will be forced to admit that the close
+resemblance of the embryo of man to that, for instance, of a dog--the
+construction of his skull, limbs, and whole frame on the same plan
+with that of other mammals, independently of the uses to which the
+parts may be put--the occasional reappearance of various structures,
+for instance of several muscles, which man does not normally possess,
+but which are common to the Quadrumana--and a crowd of analogous
+facts--all point in the plainest manner to the conclusion that man is
+the co-descendant with other mammals of a common progenitor."
+
+These few lines of Darwin's have a greater scientific value than
+hundreds of those so-called "anthropological treatises," which give
+detailed descriptions of single organs, or mathematical tables with
+series of numbers and what are claimed to be "exact analyses," but are
+devoid of synoptic conclusions and a philosophical spirit.
+
+Charles Darwin is not generally recognised as a great anthropologist,
+nor does the school of modern anthropologists regard him as a leading
+authority. In Germany, especially, the great majority of the members
+of the anthropological societies took up an attitude of hostility to
+him from the very beginning of the controversy in 1860. _The Descent
+of Man_ was not merely rejected, but even the discussion of it was
+forbidden on the ground that it was "unscientific."
+
+The centre of this inveterate hostility for thirty years--especially
+after 1877--was Rudolph Virchow of Berlin, the leading investigator
+in pathological anatomy, who did so much for the reform of medicine by
+his establishment of cellular pathology in 1858. As a prominent
+representative of "exact" or "descriptive" anthropology, and lacking a
+broad equipment in comparative anatomy and ontogeny, he was unable to
+accept the theory of descent. In earlier years, and especially during
+his splendid period of activity at Würzburg (1848-1856), he had been a
+consistent free-thinker, and had in a number of able articles
+(collected in his _Gesammelte Abhandlungen_)[141] upheld the unity of
+human nature, the inseparability of body and spirit. In later years at
+Berlin, where he was more occupied with political work and sociology
+(especially after 1866), he abandoned the positive monistic position
+for one of agnosticism and scepticism, and made concessions to the
+dualistic dogma of a spiritual world apart from the material frame.
+
+In the course of a Scientific Congress at Munich in 1877 the conflict
+of these antithetic views of nature came into sharp relief. At this
+memorable Congress I had undertaken to deliver the first address
+(September 18th) on the subject of "Modern evolution in relation to
+the whole of science." I maintained that Darwin's theory not only
+solved the great problem of the origin of species, but that its
+implications, especially in regard to the nature of man, threw
+considerable light on the whole of science, and on anthropology in
+particular. The discovery of the real origin of man by evolution from
+a long series of mammal ancestors threw light on his place in nature
+in every respect, as Huxley had already shown in his excellent
+lectures of 1863. Just as all the organs and tissues of the human body
+had originated from those of the nearest related mammals, certain
+ape-like forms, so we were bound to conclude that his mental qualities
+also had been derived from those of his extinct primate ancestor.
+
+This monistic view of the origin and nature of man, which is now admitted
+by nearly all who have the requisite acquaintance with biology, and
+approach the subject without prejudice, encountered a sharp opposition at
+that time. The opposition found its strongest expression in an address that
+Virchow delivered at Munich four days afterwards (September 22nd), on "The
+freedom of science in the modern State." He spoke of the theory of
+evolution as an unproved hypothesis, and declared that it ought not to be
+taught in the schools, because it was dangerous to the State. "We must
+not," he said, "teach that man has descended from the ape or any other
+animal." When Darwin, usually so lenient in his judgment, read the English
+translation of Virchow's speech, he expressed his disapproval in strong
+terms. But the great authority that Virchow had--an authority well founded
+in pathology and sociology--and his prestige as president of the German
+Anthropological Society, had the effect of preventing any member of the
+Society from raising serious opposition to him for thirty years. Numbers of
+journals and treatises repeated his dogmatic statement: "It is quite
+certain that man has descended neither from the ape nor from any other
+animal." In this he persisted till his death in 1902. Since that time the
+whole position of German anthropology has changed. The question is no
+longer whether man was created by a distinct supernatural act or evolved
+from other mammals, but to which line of the animal hierarchy we must look
+for the actual series of ancestors. The interested reader will find an
+account of this "battle of Munich" (1877) in my three Berlin lectures
+(April, 1905), _Der Kampf um die Entwickelungs-Gedanken_.[142]
+
+The main points in our genealogical tree were clearly recognised by
+Darwin in the sixth chapter of the _Descent of Man_. Lowly organised
+fishes, like the lancelot (Amphioxus), are descended from lower
+invertebrates resembling the larvae of an existing Tunicate
+(Appendicularia). From these primitive fishes were evolved higher
+fishes of the ganoid type and others of the type of Lepidosiren
+(Dipneusta). It is a very small step from these to the Amphibia:
+
+"In the class of animals the steps are not difficult to conceive which
+led from the ancient Monotremata to the ancient Marsupials; and from
+these to the early progenitors of the placental mammals. We may thus
+ascend to the Lemuridae; and the interval is not very wide from these
+to the Simiadae. The Simiadae then branched off into two great stems,
+the New World and Old World monkeys; and from the latter, at a remote
+period, Man, the wonder and glory of the Universe, proceeded."[143]
+
+In these few lines Darwin clearly indicated the way in which we were
+to conceive our ancestral series within the vertebrates. It is fully
+confirmed by all the arguments of comparative anatomy and embryology,
+of palaeontology and physiology; and all the research of the
+subsequent forty years have gone to establish it. The deep interest in
+geology which Darwin maintained throughout his life and his complete
+knowledge of palaeontology enabled him to grasp the fundamental
+importance of the palaeontological record more clearly than
+anthropologists and zoologists usually do.
+
+There has been much debate in subsequent decades whether Darwin
+himself maintained that man was descended from the ape, and many
+writers have sought to deny it. But the lines I have quoted _verbatim_
+from the conclusion of the sixth chapter of the _Descent of Man_
+(1871) leave no doubt that he was as firmly convinced of it as was his
+great precursor Jean Lamarck in 1809. Moreover, Darwin adds, with
+particular explicitness, in the "general summary and conclusion"
+(chap. xxi.) of that standard work:[144]
+
+"By considering the embryological structure of man--the homologies
+which he presents with the lower animals,--the rudiments which he
+retains,--and the reversions to which he is liable, we can partly
+recall in imagination the former condition of our early progenitors;
+and can approximately place them in their proper place in the
+zoological series. We thus learn that man is descended from a hairy,
+tailed quadruped, probably arboreal in its habits, and an inhabitant
+of the Old World. This creature, if its whole structure had been
+examined by a naturalist, would have been classed amongst the
+Quadrumana, as surely as the still more ancient progenitor of the Old
+and New World monkeys."
+
+These clear and definite lines leave no doubt that Darwin--so critical
+and cautious in regard to important conclusions--was quite as firmly
+convinced of the descent of man from the apes (the Catarrhinae, in
+particular) as Lamarck was in 1809 and Huxley in 1863.
+
+It is to be noted particularly that, in these and other observations
+on the subject, Darwin decidedly assumes the monophyletic origin of
+the mammals, including man. It is my own conviction that this is of
+the greatest importance. A number of difficult questions in regard to
+the development of man, in respect of anatomy, physiology, psychology,
+and embryology, are easily settled if we do not merely extend our
+_progonotaxis_ to our nearest relatives, the anthropoid apes and the
+tailed monkeys from which these have descended, but go further back
+and find an ancestor in the group of the Lemuridae, and still further
+back to the Marsupials and Monotremata. The essential identity of all
+the Mammals in point of anatomical structure and embryonic
+development--in spite of their astonishing differences in external
+appearance and habits of life--is so palpably significant that modern
+zoologists are agreed in the hypothesis that they have all sprung from
+a common root, and that this root may be sought in the earlier
+Palaeozoic Amphibia.
+
+The fundamental importance of this comparative morphology of the
+Mammals, as a sound basis of scientific anthropology, was recognised
+just before the beginning of the nineteenth century, when Lamarck
+first emphasised (1794) the division of the animal kingdom into
+Vertebrates and Invertebrates. Even thirteen years earlier (1781),
+when Goethe made a close study of the mammal skeleton in the
+Anatomical Institute at Jena, he was intensely interested to find that
+the composition of the skull was the same in man as in the other
+mammals. His discovery of the _os inter-maxillare_ in man (1784),
+which was contradicted by most of the anatomists of the time, and his
+ingenious "vertebral theory of the skull," were the splendid fruit of
+his morphological studies. They remind us how Germany's greatest
+philosopher and poet was for many years ardently absorbed in the
+comparative anatomy of man and the mammals, and how he divined that
+their wonderful identity in structure was no mere superficial
+resemblance, but pointed to a deep internal connection. In my
+_Generelle Morphologie_ (1866), in which I published the first
+attempts to construct phylogenetic trees, I have given a number of
+remarkable theses of Goethe, which may be called "phyletic
+prophecies." They justify us in regarding him as a precursor of
+Darwin.
+
+In the ensuing forty years I have made many conscientious efforts to
+penetrate further along that line of anthropological research that was
+opened up by Goethe, Lamarck, and Darwin. I have brought together the many
+valuable results that have constantly been reached in comparative anatomy,
+physiology, ontogeny, and palaeontology, and maintained the effort to
+reform the classification of animals and plants in an evolutionary sense.
+The first rough drafts of pedigrees that were published in the _Generelle
+Morphologie_ have been improved time after time in the ten editions of my
+_Natürlich Schöpfungsgeschichte_ (1868-1902).[145] A sounded basis for my
+phyletic hypotheses, derived from a discriminating combination of the three
+great records--morphology, ontogeny, and palaeontology--was provided in the
+three volumes of my _Systematische Phylogenie_[146] (1894 Protists and
+Plants, 1895 Vertebrates, 1896 Invertebrates).
+
+In my _Anthropogenie_[147] I endeavoured to employ all the known
+facts of comparative ontogeny (embryology) for the purpose of
+completing my scheme of human phylogeny (evolution). I attempted to
+sketch the historical development of each organ of the body, beginning
+with the most elementary structures in the germ-layers of the
+Gastraea. At the same time I drew up a corrected statement of the most
+important steps in the line of our ancestral series.
+
+At the fourth International Congress of Zoology at Cambridge (August
+26th, 1898) I delivered an address on "Our present knowledge of the
+Descent of Man." It was translated into English, enriched with many
+valuable notes and additions, by my friend and pupil in earlier days
+Dr. Hans Gadow (Cambridge), and published under the title: _The Last
+Link: our present knowledge of the Descent of Man_[148] The
+determination of the chief animal forms that occur in the line of our
+ancestry is there restricted to thirty types, and these are
+distributed in six main groups.
+
+The first half of this "Progonotaxis hominis," which has no support
+from fossil evidence, comprises three groups: (i) Protista
+(unicellular organisms, 1-5): (ii) Invertebrate Metazoa (Coelenteria
+6-8, Vermalia 9-11): (iii) Monorrhine Vertebrates (Acrania 12-13,
+Cyclostoma 14-15). The second half, which is based on fossil records,
+also comprises three groups: (iv) Palaeozoic cold-blooded Craniota
+(Fishes 16-18, Amphibia 19, Reptiles 20): (v) Mesozoic Mammals
+(Monotrema 21, Marsupialia 22, Mallotheria 23): (vi) Cenozoic Primates
+(Lemuridae 24-25, Tailed Apes 26-27, Anthropomorpha 28-30). An
+improved and enlarged edition of this hypothetic "Progonotaxis
+hominis" was published in 1908, in my essay _Unsere Ahnenreihe_.[149]
+
+If I have succeeded in furthering, in some degree, by these
+anthropological works, the solution of the great problem of Man's
+place in nature, and particularly in helping to trace the definite
+stages in our ancestral series, I owe the success, not merely to the
+vast progress that biology has made in the last half century, but
+largely to the luminous example of the great investigators who have
+applied themselves to the problem, with so much assiduity and genius,
+for a century and a quarter--I mean Goethe and Lamarck, Gegenbaur and
+Huxley, but, above all, Charles Darwin. It was the great genius of
+Darwin that first brought together that symmetrical temple of
+scientific knowledge, the theory of descent. It was Darwin who put the
+crown on the edifice by his theory of natural selection. Not until
+this broad inductive law was firmly established was it possible to
+vindicate the special conclusion, the descent of man from a series of
+other Vertebrates. By his illuminating discovery Darwin did more for
+anthropology than thousands of those writers, who are more
+specifically titled anthropologists, have done by their technical
+treatises. We may, indeed, say that it is not merely as an exact
+observer and ingenious experimenter, but as a distinguished
+anthropologist and far-seeing thinker, that Darwin takes his place
+among the greatest men of science of the nineteenth century.
+
+To appreciate fully the immortal merit of Darwin in connection with
+anthropology, we must remember that not only did his chief work, _The
+Origin of Species_, which opened up a new era in natural history in
+1859, sustain the most virulent and widespread opposition for a
+lengthy period, but even thirty years later, when its principles were
+generally recognised and adopted, the application of them to man was
+energetically contested by many high scientific authorities. Even
+Alfred Russel Wallace, who discovered the principle of natural
+selection independently in 1858, did not concede that it was
+applicable to the higher mental and moral qualities of man. Dr.
+Wallace still holds a spiritualist and dualist view of the nature of
+man, contending that he is composed of a material frame (descended
+from the apes) and an immortal immaterial soul (infused by a higher
+power). This dual conception, moreover, is still predominant in the
+wide circles of modern theology and metaphysics, and has the general
+and influential adherence of the more conservative classes of society.
+
+In strict contradiction to this mystical dualism, which is generally
+connected with teleology and vitalism, Darwin always maintained the
+complete unity of human nature, and showed convincingly that the
+psychological side of man was developed, in the same way as the body,
+from the less advanced soul of the anthropoid ape, and, at a still
+more remote period, from the cerebral functions of the older
+vertebrates. The eighth chapter of the _Origin of Species_, which is
+devoted to instinct, contains weighty evidence that the instincts of
+animals are subject, like all other vital processes, to the general
+laws of historic development. The special instincts of particular
+species were formed by adaptation, and the modifications thus acquired
+were handed on to posterity by heredity; in their formation and
+preservation natural selection plays the same part as in the
+transformation of every other physiological function. The higher moral
+qualities of civilised man have been derived from the lower mental
+functions of the uncultivated barbarians and savages, and these in
+turn from the social instincts of the mammals. This natural and
+monistic psychology of Darwin's was afterwards more fully developed by
+his friend George Romanes in his excellent works _Mental Evolution in
+Animals_ and _Mental Evolution in Man_.[150]
+
+Many valuable and most interesting contributions to this monistic
+psychology of man were made by Darwin in his fine work on _The Descent
+of Man and Selection in Relation to Sex_, and again in his
+supplementary work, _The Expression of the Emotions in Man and
+Animals_. To understand the historical development of Darwin's
+anthropology one must read his life and the introduction to _The
+Descent of Man_. From the moment that he was convinced of the truth
+of the principle of descent--that is to say, from his thirtieth year,
+in 1838--he recognised clearly that man could not be excluded from its
+range. He recognised as a logical necessity the important conclusion
+that "man is the co-descendant with other species of some ancient,
+lower, and extinct form." For many years he gathered notes and
+arguments in support of this thesis, and for the purpose of showing
+the probable line of man's ancestry. But in the first edition of _The
+Origin of Species_ (1859) he restricted himself to the single line,
+that by this work "light would be thrown on the origin of man and his
+history." In the fifty years that have elapsed since that time the
+science of the origin and nature of man has made astonishing progress,
+and we are now fairly agreed in a monistic conception of nature that
+regards the whole universe, including man, as a wonderful unity,
+governed by unalterable and eternal laws. In my philosophical book
+_Die Welträtsel_ (1899)[151] and in the supplementary volume _Die
+Lebenswunder_ (1904)[152] I have endeavoured to show that this pure
+monism is securely established, and that the admission of the
+all-powerful rule of the same principle of evolution throughout the
+universe compels us to formulate a single supreme law--the
+all-embracing "Law of Substance," or the united laws of the constancy
+of matter and the conservation of energy. We should never have reached
+this supreme general conception if Charles Darwin--a "monistic
+philosopher" in the true sense of the word--had not prepared the way
+by his theory of descent by natural selection, and crowned the great
+work of his life by the association of this theory with a naturalistic
+anthropology.
+
+FOOTNOTES:
+
+[Footnote 127: _Generelle Morphologie der Organismen_, 2 vols.,
+Berlin, 1866.]
+
+[Footnote 128: Eng. transl.; _The History of Creation_, London, 1876.]
+
+[Footnote 129: London, 1908.]
+
+[Footnote 130: Munich, 1884.]
+
+[Footnote 131: _Die Mutationstheorie_, Leipzig, 1903.]
+
+[Footnote 132: _Der Kampf der Theile im Organismus_, Leipzig, 1881.]
+
+[Footnote 133: Leipzig, 1904.]
+
+[Footnote 134: Berlin, 1876.]
+
+[Footnote 135: _Wonders of Life_, London and New York, 1904.]
+
+[Footnote 136: Eng. transl.; _The Evolution of Man_, 2 vols., London,
+1879 and 1905.]
+
+[Footnote 137: _Origin of Species_ (6th edit.), p. 396.]
+
+[Footnote 138: Eng. transl.; _Facts and Arguments for Darwin_, London,
+1869.]
+
+[Footnote 139: 3 vols., Berlin, 1894-96.]
+
+[Footnote 140: _Descent of Man_ (Popular Edit.), p. 927.]
+
+[Footnote 141: _Gesammelte Abhandlungen zur wissenschaftlichen
+Medizin_, Berlin, 1856.]
+
+[Footnote 142: Eng. transl.; _Last Words on Evolution_, London, 1906.]
+
+[Footnote 143: _Descent of Man_, (Popular Edit.), p. 255.]
+
+[Footnote 144: _Descent of Man_, p. 930.]
+
+[Footnote 145: Eng. transl.; _The History of Creation_, London, 1876.]
+
+[Footnote 146: Berlin, 1894-96.]
+
+[Footnote 147: Leipzig, 1874, 5th edit. 1905. Eng. transl.; _The
+Evolution of Man_, London, 1905.]
+
+[Footnote 148: London, 1898.]
+
+[Footnote 149: _Festschrift zur 350-jährigen Jubelfeier der Thüringer
+Universität Jena_. Jena. 1908.]
+
+[Footnote 150: London, 1885; 1888.]
+
+[Footnote 151: _The Riddle of the Universe_, London and New York,
+1900.]
+
+[Footnote 152: _The Wonders of Life_, London and New York, 1904.]
+
+
+
+
+VI
+
+MENTAL FACTORS IN EVOLUTION
+
+BY C. LLOYD MORGAN, LLD., F.R.S
+
+
+In developing his conception of organic evolution Charles Darwin was
+of necessity brought into contact with some of the problems of mental
+evolution. In _The Origin of Species_ he devoted a chapter to "the
+diversities of instinct and of the other mental faculties in animals
+of the same class."[153] When he passed to the detailed consideration
+of _The Descent of Man_, it was part of his object to show "that there
+is no fundamental difference between man and the higher mammals in
+their mental faculties."[154] "If no organic being excepting man," he
+said, "had possessed any mental power, or if his powers had been of a
+wholly different nature, from those of the lower animals, then we
+should never have been able to convince ourselves that our high
+faculties had been gradually developed."[155] In his discussion of
+_The Expression of the Emotions_ it was important for his purpose
+"fully to recognise that actions readily become associated with other
+actions and with various states of the mind."[156] His hypothesis of
+sexual selection is largely dependent upon the exercise of choice on
+the part of the female and her preference for "not only the more
+attractive but at the same time the more vigourous and vicious
+males."[157] Mental processes and physiological processes were for
+Darwin closely correlated; and he accepted the conclusion "that the
+nervous system not only regulates most of the existing functions of
+the body, but has indirectly influenced the progressive development of
+various bodily structures and of certain mental qualities."[158]
+
+Throughout his treatment, mental evolution was for Darwin incidental
+to and contributory to organic evolution. For specialised research in
+comparative and genetic psychology, as an independent field of
+investigation, he had neither the time nor the requisite training.
+None the less his writings and the spirit of his work have exercised a
+profound influence on this department of evolutionary thought. And,
+for those who follow Darwin's lead, mental evolution is still in a
+measure subservient to organic evolution. Mental processes are the
+accompaniments or concomitants of the functional activity of specially
+differentiated parts of the organism. They are in some way dependent
+on physiological and physical conditions. But though they are not
+physical in their nature, and though it is difficult or impossible to
+conceive that they are physical in their origin, they are, for Darwin
+and his followers, factors in the evolutionary process in its physical
+or organic aspect. By the physiologist within his special and
+well-defined universe of discourse they may be properly regarded as
+epiphenomena; but by the naturalist in his more catholic survey of
+nature they cannot be so regarded, and were not so regarded by Darwin.
+Intelligence has contributed to evolution of which it is in a sense a
+product.
+
+The facts of observation or of inference which Darwin accepted are
+these: Conscious experience accompanies some of the modes of animal
+behaviour; it is concomitant with certain physiological processes;
+these processes are the outcome of development in the individual and
+evolution in the race; the accompanying mental processes undergo a
+like development. Into the subtle philosophical questions which arise
+out of the naïve acceptance of such a creed it was not Darwin's
+province to enter; "I have nothing to do," he said,[159] "with the
+origin of the mental powers, any more than I have with that of life
+itself." He dealt with the natural history of organisms, including not
+only their structure but their modes of behaviour; with the natural
+history of the states of consciousness which accompany some of their
+actions; and with the relation of behaviour to experience. We will
+endeavour to follow Darwin in his modesty and candour in making no
+pretence to give ultimate explanations. But we must note one of the
+implications of this self-denying ordinance of science. Development
+and evolution imply continuity. For Darwin and his followers the
+continuity is organic through physical heredity. Apart from
+speculative hypothesis, legitimate enough in its proper place but here
+out of court, we know nothing of continuity of mental evolution as
+such: consciousness appears afresh in each succeeding generation.
+Hence it is that for those who follow Darwin's lead, mental evolution
+is and must ever be, within his universe of discourse, subservient to
+organic evolution. Only in so far as conscious experience, or its
+neural correlate, effects some changes in organic structure can it
+influence the course of heredity; and conversely only in so far as
+changes in organic structure are transmitted through heredity, is
+mental evolution rendered possible. Such is the logical outcome of
+Darwin's teaching.
+
+Those who abide by the cardinal results of this teaching are bound to
+regard all behaviour as the expression of the functional activities of
+the living tissues of the organism, and all conscious experience as
+correlated with such activities. For the purposes of scientific
+treatment, mental processes are one mode of expression of the same
+changes of which the physiological processes accompanying behaviour
+are another mode of expression. This is simply accepted as a fact
+which others may seek to explain. The behaviour itself is the adaptive
+application of the energies of the organism; it is called forth by
+some form of presentation or stimulation brought to bear on the
+organism by the environment. This presentation is always an individual
+or personal matter. But in order that the organism may be fitted to
+respond to the presentation of the environment it must have undergone
+in some way a suitable preparation. According to the theory of
+evolution this preparation is primarily racial and is transmitted
+through heredity. Darwin's main thesis was that the method of
+preparation is predominantly by natural selection. Subordinate to
+racial preparation, and always dependent thereon, is individual or
+personal preparation through some kind of acquisition; of which the
+guidance of behaviour through individually won experience is a typical
+example. We here introduce the mental factor because the facts seem to
+justify the inference. Thus there are some modes of behaviour which
+are wholly and solely dependent upon inherited racial preparation;
+there are other modes of behaviour which are also dependent, in part
+at least, on individual preparation. In the former case the behaviour
+is adaptive on the first occurrence of the appropriate presentation;
+in the latter case accommodation to circumstances is only reached
+after a greater or less amount of acquired organic modification of
+structure, often accompanied (as we assume) in the higher animals by
+acquired experience. Logically and biologically the two classes of
+behaviour are clearly distinguishable: but the analysis of complex
+cases of behaviour where the two factors coöperate, is difficult and
+requires careful and critical study of life-history.
+
+The foundations of the mental life are laid in the conscious
+experience that accompanies those modes of behaviour, dependent
+entirely on racial preparation, which may broadly be described as
+instinctive. In the eighth chapter of _The Origin of Species_ Darwin
+says,[160] "I will not attempt any definition of instinct.... Every
+one understands what is meant, when it is said that instinct impels
+the cuckoo to migrate and to lay her eggs in other birds' nests. An
+action, which we ourselves require experience to enable us to perform,
+when performed by an animal, more especially by a very young one,
+without experience, and when performed by many individuals in the same
+way, without their knowing for what purpose it is performed, is
+usually said to be instinctive." And in the summary at the close of
+the chapter he says,[161] "I have endeavoured briefly to show that the
+mental qualities of our domestic animals vary, and that the variations
+are inherited. Still more briefly I have attempted to show that
+instincts vary slightly in a state of nature. No one will dispute that
+instincts are of the highest importance to each animal. Therefore
+there is no real difficulty, under changing conditions of life, in
+natural selection accumulating to any extent slight modifications of
+instinct which are in any way useful. In many cases habit or use and
+disuse have probably come into play."
+
+Into the details of Darwin's treatment there is neither space nor need
+to enter. There are some ambiguous passages; but it may be said that
+for him, as for his followers to-day, instinctive behaviour is wholly
+the result of racial preparation transmitted through organic heredity.
+For the performance of the instinctive act no individual preparation
+under the guidance of personal experience is necessary. It is true
+that Darwin quotes with approval Huber's saying that "a little dose of
+judgment or reason often comes into play, even with animals low in the
+scale of nature."[162] But we may fairly interpret his meaning to be
+that in behaviour, which is commonly called instinctive, some element
+of intelligent guidance is often combined. If this be conceded the
+strictly instinctive performance (or part of the performance) is the
+outcome of heredity and due to the direct transmission of parental or
+ancestral aptitudes. Hence the instinctive response as such depends
+entirely on how the nervous mechanism has been built up through
+heredity; while intelligent behaviour, or the intelligent factor in
+behaviour, depends also on how the nervous mechanism has been modified
+and moulded by use during its development and concurrently with the
+growth of individual experience in the customary situations of daily
+life. Of course it is essential to the Darwinian thesis that what Sir
+E. Ray Lankester has termed "educability," not less than instinct, is
+hereditary. But it is also essential to the understanding of this
+thesis that the differentiae of the hereditary factor should be
+clearly grasped.
+
+For Darwin there were two modes of racial preparation, (_1_) natural
+selection, and (_2_) the establishment of individually acquired habit.
+He showed that instincts are subject to hereditary variation; he saw
+that instincts are also subject to modification through acquisition in
+the course of individual life. He believed that not only the
+variations but also, to some extent, the modifications are inherited.
+He therefore held that some instincts (the greater number) are due to
+natural selection but that others (less numerous) are due, or partly
+due, to the inheritance of acquired habits. The latter involve
+Lamarckian inheritance, which of late years has been the centre of so
+much controversy. It is noteworthy however that Darwin laid especial
+emphasis on the fact that many of the most typical and also the most
+complex instincts--those of neuter insects--do not admit of such an
+interpretation. "I am surprised," he says,[163] "that no one has
+hitherto advanced this demonstrative case of neuter insects, against
+the well-known doctrine of inherited habit, as advanced by Lamarck."
+None the less Darwin admitted this doctrine as supplementary to that
+which was more distinctively his own--for example in the case of the
+instincts of domesticated animals. Still, even in such cases, "it may
+be doubted," he says,[164] "whether any one would have thought of
+training a dog to point, had not some one dog naturally shown a
+tendency in this line ... so that habit and some degree of selection
+have probably concurred in civilising by inheritance our dogs." But in
+the interpretation of the instincts of domesticated animals, a more
+recently suggested hypothesis, that of organic selection,[165] may be
+helpful. According to this hypothesis any intelligent modification of
+behaviour which is subject to selection is probably coincident in
+direction with an inherited tendency to behave in this fashion. Hence
+in such behaviour there are two factors: (1) an incipient variation in
+the line of such behaviour, and (2) an acquired modification by which
+the behaviour is carried further along the same line. Under natural
+selection those organisms in which the two factors coöperate are
+likely to survive. Under artificial selection they are deliberately
+chosen out from among the rest.
+
+Organic selection has been termed a compromise between the more
+strictly Darwinian and the Lamarckian principles of interpretation.
+But it is not in any sense a compromise. The principle of
+interpretation of that which is instinctive and hereditary is wholly
+Darwinian. It is true that some of the facts of observation relied
+upon by Lamarckians are introduced. For Lamarckians however the
+modifications which are admittedly factors in survival, are regarded
+as the parents of inherited variations; for believers in organic
+selection they are only the foster-parents or nurses. It is because
+organic selection is the direct outcome of and a natural extension of
+Darwin's cardinal thesis that some reference to it here is
+justifiable. The matter may be put with the utmost brevity as follows:
+(1) Variations (V) occur, some of which are in the direction of
+increased adaptation (+), others in the direction of decreased
+adaptation (-).
+
+(2) Acquired modifications (M) also occur. Some of these are in the
+direction of increased accommodation to circumstances (+), while
+others are in the direction of diminished accommodation (-). Four
+major combinations are
+
+   (_b_) + V with - M,      (_c_) - V with + M,
+
+   (_a_) + V with + M,      (_d_) - V with - M.
+
+Of these (_d_) must inevitably be eliminated while (_a_) are selected.
+The predominant survival of (_a_) entails the survival of the adaptive
+variations which are inherited. The contributory acquisitions (+ M)
+are not inherited; but there are none the less factors in determining
+the survival of the coincident variations. It is surely abundantly
+clear that this is Darwinism and has no tincture of Lamarck's
+essential principle, the inheritance of acquired characters.
+
+Whether Darwin himself would have accepted this interpretation of some
+at least of the evidence put forward by Lamarckians is unfortunately a
+matter of conjecture. The fact remains that in his interpretation of
+instinct and in allied questions he accepted the inheritance of
+individually acquired modifications of behaviour and structure.
+
+Darwin was chiefly concerned with instinct from the biological rather
+than from the psychological point of view. Indeed it must be confessed
+that, from the latter standpoint, his conception of instinct as a
+"mental faculty" which "impels" an animal to the performance of
+certain actions, scarcely affords a satisfactory basis for genetic
+treatment. To carry out the spirit of Darwin's teaching it is
+necessary to link more closely biological and psychological evolution.
+The first step towards this is to interpret the phenomena of
+instinctive behaviour in terms of stimulation and response. It may be
+well to take a particular case. Swimming on the part of a duckling is,
+from the biological point of view, a typical example of instinctive
+behaviour. Gently lower a recently hatched bird into water:
+coordinated movements of the limbs follow in rhythmical sequence. The
+behaviour is new to the individual though it is no doubt closely
+related to that of walking, which is no less instinctive. There is a
+group of stimuli afforded by the "presentation" which results from
+partial immersion: upon this there follows as a complex response an
+application of the functional activities in swimming; the sequence of
+adaptive application on the appropriate presentation is determined by
+racial preparation. We know, it is true, but little of the
+physiological details of what takes place in the central nervous
+system; but in broad outline the nature of the organic mechanism and
+the manner of its functioning may at least be provisionally
+conjectured in the present state of physiological knowledge. Similarly
+in the case of the pecking of newly-hatched chicks; there is a visual
+presentation, there is probably a coöperating group of stimuli from
+the alimentary tract in need of food, there is an adaptive application
+of the activities in a definite mode of behaviour. Like data are
+afforded in a great number of cases of instinctive procedure,
+sometimes occurring very early in life, not infrequently deferred
+until the organism is more fully developed, but all of them dependent
+upon racial preparation. No doubt there is some range of variation in
+the behaviour, just such variation as the theory of natural selection
+demands. But there can be no question that the higher animals inherit
+a bodily organisation and a nervous system, the functional working of
+which gives rise to those inherited modes of behaviour which are
+termed instinctive.
+
+It is to be noted that the term "instinctive" is here employed in the
+adjectival form as a descriptive heading under which may be grouped
+many and various modes of behaviour due to racial preparation. We
+speak of these as inherited; but in strictness what is transmitted
+through heredity is the complex of anatomical and physiological
+conditions under which, in appropriate circumstances, the organism so
+behaves. So far the term "instinctive" has a restricted biological
+connotation in terms of behaviour. But the connecting link between
+biological evolution and psychological evolution is to be sought,--as
+Darwin fully realised,--in the phenomena of instinct, broadly
+considered. The term "instinctive" has also a psychological
+connotation. What is that connotation?
+
+Let us take the case of the swimming duckling or the pecking chick,
+and fix our attention on the first instinctive performance. Grant that
+just as there is, strictly speaking, no inherited behaviour, but only
+the conditions which render such behaviour under appropriate
+circumstances possible; so too there is no inherited experience, but
+only the conditions which render such experience possible; then the
+cerebral conditions in both cases are the same. The biological
+behaviour-complex, including the total stimulation and the total
+response with the intervening or resultant processes in the sensorium,
+is accompanied by an experience-complex including the initial
+stimulation-consciousness and resulting response-consciousness. In the
+experience-complex are comprised data which in psychological analysis
+are grouped under the headings of cognition, affective tone and
+conation. But the complex is probably experienced as an unanalysed
+whole. If then we use the term "instinctive" so as to comprise all
+congenital modes of behaviour which contribute to experience, we are
+in a position to grasp the view that the net result in consciousness
+constitutes what we may term the primary tissue of experience. To the
+development of this experience each instinctive act contributes. The
+nature and manner of organisation of this primary tissue of experience
+are dependent on inherited biological aptitudes; but they are from the
+outset onwards subject to secondary development dependent on acquired
+aptitudes. Biological values are supplemented by psychological values
+in terms of satisfaction or the reverse.
+
+In our study of instinct we have to select some particular phase of
+animal behaviour and isolate it so far as is possible from the life of
+which it is a part. But the animal is a going concern, restlessly
+active in many ways. Many instinctive performances, as Darwin pointed
+out,[166] are serial in their nature. But the whole of active life is
+a serial and coordinated business. The particular instinctive
+performance is only an episode in a life-history, and every mode of
+behaviour is more or less closely correlated with other modes. This
+coordination of behaviour is accompanied by a correlation of the modes
+of primary experience. We may classify the instinctive modes of
+behaviour and their accompanying modes of instinctive experience under
+as many heads as may be convenient for our purposes of interpretation,
+and label them instincts of self-preservation, of pugnacity, of
+acquisition, the reproductive instincts, the parental instincts, and
+so forth. An instinct, in this sense of the term (for example the
+parental instinct), may be described as a specialised part of the
+primary tissue of experience differentiated in relation to some
+definite biological end. Under such an instinct will fall a large
+number of particular and often well-defined modes of behaviour, each
+with its own peculiar mode of experience.
+
+It is no doubt exceedingly difficult as a matter of observation and of
+inference securely based thereon to distinguish what is primary from
+what is in part due to secondary acquisition--a fact which Darwin
+fully appreciated. Animals are educable in different degrees; but
+where they are educable they begin to profit by experience from the
+first. Only, therefore, on the occasion of the first instinctive act
+of a given type can the experience gained be regarded as _wholly_
+primary; all subsequent performance is liable to be in some degree,
+sometimes more, sometimes less, modified by the acquired disposition
+which the initial behaviour engenders. But the early stages of
+acquisition are always along the lines predetermined by instinctive
+differentiation. It is the task of comparative psychology to
+distinguish the primary tissue of experience from its secondary and
+acquired modifications. We cannot follow up the matter in further
+detail. It must here suffice to suggest that this conception of
+instinct as a primary form of experience lends itself better to
+natural history treatment than Darwin's conception of an impelling
+force, and that it is in line with the main trend of Darwin's thought.
+
+In a characteristic work,--characteristic in wealth of detail, in
+closeness and fidelity of observation, in breadth of outlook, in
+candour and modesty,--Darwin dealt with _The Expression of the
+Emotions in Man and Animals_. Sir Charles Bell in his _Anatomy of
+Expression_ had contended that many of man's facial muscles had been
+specially created for the sole purpose of being instrumental in the
+expression of his emotions. Darwin claimed that a natural explanation,
+consistent with the doctrine of evolution, could in many cases be
+given and would in other cases be afforded by an extension of the
+principles he advocated. "No doubt," he said,[167] "as long as man and
+all other animals are viewed as independent creations, an effectual
+stop is put to our natural desire to investigate as far as possible
+the causes of Expression. By this doctrine, anything and everything
+can be equally well explained.... With mankind, some expressions ...
+can hardly be understood, except on the belief that man once existed
+in a much lower and animal-like condition. The community of certain
+expressions in distinct though allied species ... is rendered somewhat
+more intelligible, if we believe in their descent from a common
+progenitor. He who admits on general grounds that the structure and
+habits of all animals have been gradually evolved, will look at the
+whole subject of Expression in a new and interesting light."
+
+Darwin relied on three principles of explanation. "The first of these
+principles is, that movements which are serviceable in gratifying some
+desire, or in relieving some sensation, if often repeated, become so
+habitual that they are performed, whether or not of any service,
+whenever the same desire or sensation is felt, even in a very weak
+degree."[168] The modes of expression which fall under this head have
+become instinctive through the hereditary transmission of acquired
+habit. "As far as we can judge, only a few expressive movements are
+learnt by each individual; that is, were consciously and voluntarily
+performed during the early years of life for some definite object, or
+in imitation of others, and then became habitual. The far greater
+number of the movements of expression, and all the more important
+ones, are innate or inherited; and such cannot be said to depend on
+the will of the individual. Nevertheless, all those included under our
+first principle were at first voluntarily performed for a definite
+object,--namely, to escape some danger, to relieve some distress, or
+to gratify some desire."[169]
+
+"Our second principle is that of antithesis. The habit of voluntarily
+performing opposite movements under opposite impulses has become
+firmly established in us by the practice of our whole lives. Hence, if
+certain actions have been regularly performed, in accordance with our
+first principle, under a certain frame of mind, there will be a strong
+and involuntary tendency to the performance of directly opposite
+actions, whether or not these are of any use, under the excitement of
+an opposite frame of mind."[170] This principle of antithesis has not
+been widely accepted. Nor is Darwin's own position easy to grasp.
+
+"Our third principle," he says,[171] "is the direct action of the
+excited nervous system on the body, independently of the will, and
+independently, in large part, of habit. Experience shows that
+nerve-force is generated and set free whenever the cerebro-spinal
+system is excited. The direction which this nerve-force follows is
+necessarily determined by the lines of connection between the
+nerve-cells, with each other and with various parts of the body."
+
+Lack of space prevents our following up the details of Darwin's
+treatment of expression. Whether we accept or do not accept his three
+principles of explanation we must regard his work as a masterpiece of
+descriptive analysis, packed full of observations possessing lasting
+value. For a further development of the subject it is essential that
+the instinctive factors in expression should be more fully
+distinguished from those which are individually acquired--a difficult
+task--and that the instinctive factors should be rediscussed in the
+light of modern doctrines of heredity, with a view to determining
+whether Lamarckian inheritance, on which Darwin so largely relied, is
+necessary for an interpretation of the facts.
+
+The whole subject as Darwin realised is very complex. Even the term
+"expression" has a certain amount of ambiguity. When the emotion is in
+full flood, the animal fights, flees, or faints. Is this full-tide
+effect to be regarded as expression; or are we to restrict the term to
+the premonitory or residual effects--the bared canine when the
+fighting mood is being roused, the ruffled fur when reminiscent
+representations of the object inducing anger cross the mind? Broadly
+considered both should be included. The activity of premonitory
+expression as a means of communication was recognised by Darwin; he
+might, perhaps, have emphasised it more strongly in dealing with the
+lower animals. Man so largely relies on a special means of
+communication, that of language, that he sometimes fails to realise
+that for animals with their keen powers of perception, and dependent
+as they are on such means of communication, the more strictly
+biological means of expression are full of subtle suggestiveness. Many
+modes of expression, otherwise useless, are signs of behaviour that
+may be anticipated,--signs which stimulate the appropriate attitude of
+response. This would not, however, serve to account for the utility of
+the organic accompaniments--heart-affection, respiratory changes,
+vaso-motor effects and so forth, together with heightened muscular
+tone,--on all of which Darwin lays stress[172] under his third
+principle. The biological value of all this is, however, of great
+importance, though Darwin was hardly in a position to take it fully
+into account.
+
+Having regard to the instinctive and hereditary factors of emotional
+expression we may ask whether Darwin's third principle does not alone
+suffice as an explanation. Whether we admit or reject Lamarckian
+inheritance it would appear that all hereditary expression must be due
+to pre-established connections within the central nervous system and
+to a transmitted provision for coordinated response under the
+appropriate stimulation. If this be so, Darwin's first and second
+principles are subordinate and ancillary to the third, an expression,
+so far as it is instinctive or heredity, being "the direct result of
+the constitution of the nervous system."
+
+Darwin accepted the emotions themselves as hereditary or acquired
+states of mind and devoted his attention to their expression. But
+these emotions themselves are genetic products and as such dependent
+on organic conditions. It remained, therefore, for psychologists who
+accepted evolution and sought to build on biological foundations to
+trace the genesis of these modes of animal and human experience. The
+subject has been independently developed by Professors Lange and
+James;[173] and some modification of their view is regarded by many
+evolutionists as affording the best explanation of the facts. We must
+fix our attention on the lower emotions, such as anger or fear, and on
+their first occurrence in the life of the individual organism. It is a
+matter of observation that if a group of young birds which have been
+hatched in an incubator are frightened by an appropriate presentation,
+auditory or visual, they instinctively respond in special ways. If we
+speak of this response as the expression, we find that there are many
+factors. There are certain visible modes of behaviour, crouching at
+once, scattering and then crouching, remaining motionless, the braced
+muscles sustaining an attitude of arrest, and so forth, There are also
+certain visceral or organic effects, such as affections of the heart
+and respiration. These can be readily observed by taking the young
+bird in the hand. Other effects cannot be readily observed; vaso-motor
+changes, affections of the alimentary canal, the skin and so forth.
+Now the essence of the James-Lange view, as applied to these
+congenital effects, is that though we are justified in speaking of
+them as effects of the stimulation, we are not justified, without
+further evidence, in speaking of them as effects of the emotional
+state. May it not rather be that the emotion as a primary mode of
+experience is the concomitant of the net result of the organic
+situation--the initial presentation, the instinctive mode of
+behaviour, the visceral disturbances?
+
+According to this interpretation the primary tissue of experience of
+the emotional order, felt as an unanalysed complex, is generated by
+the stimulation of the sensorium by afferent or incoming physiological
+impulses from the special senses, from the organs concerned in the
+responsive behaviour, from the viscere and vaso-motor system.
+
+Some psychologists, however, contend that the emotional experience is
+generated in the sensorium prior to, and not subsequent to, the
+behaviour-response and the visceral disturbances. It is a direct and
+not an indirect outcome of the presentation to the special senses. Be
+this as it may, there is a growing tendency to bring into the closest
+possible relation, or even to identify, instinct and emotion in their
+primary genesis. The central core of all such interpretations is that
+instinctive behaviour and experience, its emotional accompaniments,
+and its expression, are but different aspects of the outcome of the
+same organic occurrences. Such emotions are, therefore, only a
+distinguishable aspect of the primary tissue of experience and exhibit
+a like differentiation. Here again a biological foundation is laid for
+a psychological doctrine of the mental development of the individual.
+
+The intimate relation between emotion as a psychological mode of
+experience and expression as a group of organic conditions has an
+important bearing on biological interpretation. The emotion, as the
+psychological accompaniment of orderly disturbances in the central
+nervous system, profoundly influences behaviour and often renders it
+more vigourous and more effective. The utility of the emotions in the
+struggle for existence can, therefore, scarcely be over-estimated.
+Just as keenness of perception has survival-value; just as it is
+obviously subject to variation; just as it must be enhanced under
+natural selection, whether individually acquired increments are
+inherited or not; and just as its value lies not only in this or that
+special perceptive act but in its importance for life as a whole; so
+the vigourous effectiveness of activity has survival-value; it is
+subject to variation; it must be enhanced under natural selection; and
+its importance lies not only in particular modes of behaviour but in
+its value for life as a whole. If emotion and its expression as a
+congenital endowment are but different aspects of the same biological
+occurrence; and if this is a powerful supplement to vigour
+effectiveness and persistency of behaviour, it must on Darwin's
+principles be subject to natural selection.
+
+If we include under the expression of the emotions not only the
+premonitory symptoms of the initial phases of the organic and mental
+state, not only the signs or conditions of half-tide emotion, but the
+full-tide manifestation of an emotion which dominates the situation,
+we are naturally led on to the consideration of many of the phenomena
+which are discussed under the head of sexual selection. The subject is
+difficult and complex, and it was treated by Darwin with all the
+strength he could summon to the task. It can only be dealt with here
+from a special point of view--that which may serve to illustrate the
+influence of certain mental factors on the course of evolution. From
+this point of view too much stress can scarcely be laid on the
+dominance of emotion during the period of courtship and pairing in the
+more highly organised animals. It is a period of maximum vigour,
+maximum activity, and, correlated with special modes of behaviour and
+special organic and visceral accompaniments, a period also of maximum
+emotional excitement. The combats of males, their dances and aerial
+evolutions, their elaborate behaviour and display, or the flood of
+song in birds, are emotional expressions which are at any rate
+coincident in time with sexual periodicity. From the combat of the
+males there follows on Darwin's principles the elimination of those
+which are deficient in bodily vigour, deficient in special structures,
+offensive or protective, which contribute to success, deficient in the
+emotional supplement of which persistent and whole-hearted fighting is
+the expression, and deficient in alertness and skill which are the
+outcome of the psychological development of the powers of perception.
+Few biologists question that we have here a mode of selection of much
+importance, though its influence on psychological evolution often
+fails to receive its due emphasis. Mr. Wallace[174] regards it as "a
+form of natural selection"; "to it," he says, "we must impute the
+development of the exceptional strength, size, and activity of the
+male, together with the possession of special offensive and defensive
+weapons, and of all other characters which arise from the development
+of these or are correlated with them." So far there is little
+disagreement among the followers of Darwin--for Mr. Wallace, with fine
+magnanimity, has always preferred to be ranked as such,
+notwithstanding his right, on which a smaller man would have
+constantly insisted, to the claim of independent originator of the
+doctrine of natural selection. So far with regard to sexual selection
+Darwin and Mr. Wallace are agreed; so far and no farther. For Darwin,
+says Mr. Wallace,[175] "has extended the principle into a totally
+different field of action, which has none of that character of
+constancy and of inevitable result that attaches to natural selection,
+including male rivalry; for by far the larger portion of the
+phenomena, which he endeavours to explain by the direct action of
+sexual selection, can only be so explained on the hypothesis that the
+immediate agency is female choice or preference. It is to this that he
+imputes the origin of all secondary sexual characters other than
+weapons of offence and defence.... In this extension of sexual
+selection to include the action of female choice or preference, and in
+the attempt to give to that choice such wide-reaching effects, I am
+unable to follow him more than a very little way."
+
+Into the details of Mr. Wallace's criticisms it is impossible to enter
+here. We cannot discuss either the mode of origin of the variations in
+structure which have rendered secondary sexual characters possible or
+the modes of selection other than sexual which have rendered them,
+within narrow limits, specifically constant. Mendelism and mutation
+theories may have something to say on the subject when these theories
+have been more fully correlated with the basal principles of
+selection. It is noteworthy that Mr. Wallace says:[176] "Besides the
+acquisition of weapons by the male for the purpose of fighting with
+other males, there are some other sexual characters which may have
+been produced by natural selection. Such are the various sounds and
+odours which are peculiar to the male, and which serve as a call to
+the female or as an indication of his presence. These are evidently a
+valuable addition to the means of recognition of the two sexes, and
+are a further indication, that the pairing season has arrived; and the
+production, intensification, and differentiation of these sounds and
+odours are clearly within the power of natural selection. The same
+remark will apply to the peculiar calls of birds, and even to the
+singing of the males." Why the same remark should not apply to their
+colours and adornments is not obvious. What is obvious is that "means
+of recognition" and "indication that the pairing season has arrived"
+are dependent on the perceptive powers of the female who recognises
+and for whom the indication has meaning. The hypothesis of female
+preference, stripped of the aesthetic surplusage which is
+psychologically both unnecessary and unproven, is really only
+different in degree from that which Mr. Wallace admits in principle
+when he says that it is probable that the female is pleased or excited
+by the display.
+
+Let us for our present purpose leave on one side and regard as _sub
+judice_ the question whether the specific details of secondary sexual
+characters are the outcome of female choice. For us the question is
+whether certain psychological accompaniments of the pairing situation
+have influenced the course of evolution and whether these
+psychological accompaniments are themselves the outcome of evolution.
+As a matter of observation, specially differentiated modes of
+behaviour, often very elaborate, frequently requiring highly developed
+skill, and apparently highly charged with emotional tone, are the
+precursors of pairing. They are generally confined to the males, whose
+fierce combats during the period of sexual activity are part of the
+emotional manifestation. It is inconceivable that they have no
+biological meaning; and it is difficult to conceive that they have any
+other biological end than to evoke in the generally more passive
+female the pairing impulse. They, are based on instinctive foundations
+ingrained in the nervous constitution through natural (or may we not
+say sexual?) selection in virtue of their profound utility. They are
+called into play by a specialised presentation such as the sight or
+the scent of the female at, or a little in advance of, a critical
+period of the physiological rhythm. There is no necessity that the
+male should have any knowledge of the end to which his strenuous
+activity leads up. In presence of the female there is an elaborate
+application of all the energies of behaviour, just because ages of
+racial preparation have made him biologically and emotionally what he
+is--a functionally sexual male that must dance or sing or go through
+hereditary movements of display, when the appropriate stimulation
+comes. Of course after the first successful courtship his future
+behaviour will be in some degree modified by his previous experience.
+No doubt during his first courtship he is gaining the primary data of
+a peculiarly rich experience, instinctive and emotional. But the
+biological foundations of the behaviour of courtship are laid in the
+hereditary coordinations. It would seem that in some cases, not indeed
+in all, perhaps especially in those cases in which secondary sexual
+behaviour is most highly evolved,--correlative with the ardour of the
+male is a certain amount of reluctance in the female. The pairing act
+on her part only takes place after prolonged stimulation, for
+affording which the behaviour of male courtship is the requisite
+presentation. The most vigourous, defiant and mettlesome male is
+preferred just because he alone affords a contributory stimulation
+adequate to evoke the pairing impulse with its attendant emotional
+tone.
+
+It is true that this places female preference or choice on a much
+lower psychological plane than Darwin in some passages seems to
+contemplate where, for example, he says that the female appreciates
+the display of the male and places to her credit a taste for the
+beautiful. But Darwin himself distinctly states[177] that "it is not
+probable that she consciously deliberates; but she is most excited or
+attracted by the most beautiful, or melodious, or gallant males." The
+view here put forward, which has been developed by Prof. Groos,[178]
+therefore seems to have Darwin's own sanction. The phenomena are not
+only biological; there are psychological elements as well. One can
+hardly suppose that the female is unconscious of the male's presence;
+the final yielding must surely be accompanied by heightened emotional
+tone. Whether we call it choice or not is merely a matter of
+definition of terms. The behaviour is in part determined by
+supplementary psychological values. Prof. Groos regards the coyness of
+females as "a most efficient means of preventing the too early and too
+frequent yielding to the sexual impulse."[179] Be that as it may, it
+is, in any case, if we grant the facts, a means through which male
+sexual behaviour with all its biological and psychological
+implications, is raised to a level otherwise perhaps unattainable by
+natural means, while in the female it affords opportunities for the
+development in the individual and evolution in the race of what we may
+follow Darwin in calling appreciation, if we empty this word of the
+aesthetic implications which have gathered round it in the mental life
+of man.
+
+Regarded from this standpoint of sexual selection, broadly considered,
+has probably been of great importance. The psychological
+accompaniments of the pairing situation have profoundly influenced the
+course of biological evolution and are themselves the outcome of that
+evolution.
+
+Darwin makes only passing reference to those modes of behaviour in
+animals which go by the name of play. "Nothing," he says,[180] "is
+more common than for animals to take pleasure in practising whatever
+instinct they follow at other times for some real good." This is one
+of the very numerous cases in which a hint of the master has served to
+stimulate research in his disciples. It was left to Prof. Groos to
+develop this subject on evolutionary lines and to elaborate in a
+masterly manner Darwin's suggestion. "The utility of play," he
+says,[181] "is incalculable. This utility consists in the practice and
+exercise it affords for some of the more important duties of
+life,"--that is to say, for the performance of activities which will
+in adult life be essential to survival. He urges[182] that "the play
+of young animals has its origin in the fact that certain very
+important instincts appear at a time when the animal does not
+seriously need them." It is, however, questionable whether any
+instincts appear at a time when they are not needed. And it is
+questionable whether the instinctive and emotional attitude of the
+play-fight, to take one example, can be identified with those which
+accompany fighting in earnest, though no doubt they are closely
+related and have some common factors. It is probable that play, as
+preparatory behaviour, differs in biological detail (as it almost
+certainly does in emotional attributes) from the earnest of after-life
+and that it has been evolved through differentiation and integration
+of the primary tissue of experience, as a preparation through which
+certain essential modes of skill may be acquired--those animals in
+which the preparatory play-propensity was not inherited in due force
+and requisite amount being subsequently eliminated in the struggle for
+existence. In any case there is little question that Prof. Groos is
+right in basing the play-propensity on instinctive foundations.[183]
+None the less, as he contends, the essential biological value of play
+is that it is a means of training the educable nerve-tissue, of
+developing that part of the brain which is modified by experience and
+which thus acquires new characters, of elaborating the secondary
+tissue of experience on the predetermined lines of instinctive
+differentiation and thus furthering the psychological activities which
+are included under the comprehensive term "intelligent."
+
+In _The Descent of Man_ Darwin dealt at some length with intelligence
+and the higher mental faculties.[184] His object, he says, is to show
+that there is no fundamental difference between man and the higher
+mammals in their mental faculties; that these faculties are variable
+and the variations tend to be inherited; and that under natural
+selection beneficial variations of all kinds will have been preserved
+and injurious ones eliminated.
+
+Darwin was too good an observer and too honest a man to minimise the
+"enormous difference" between the level of mental attainment of
+civilised man and that reached by any animal. His contention was that
+the difference, great as it is, is one of degree and not of kind. He
+realised that, in the development of the mental faculties of man, new
+factors in evolution have supervened--factors which play but a
+subordinate and subsidiary part in animal intelligence.
+Intercommunication by means of language, approbation and blame, and
+all that arises out of reflective thought, are but foreshadowed in the
+mental life of animals. Still he contends that these may be explained
+on the doctrine of evolution. He urges[185] "that man is variable in
+body and mind; and that the variations are induced, either directly or
+indirectly, by the same general causes, and obey the same general
+laws, as with the lower animals." He correlates mental development
+with the evolution of the brain.[186] "As the various mental faculties
+gradually developed themselves, the brain would almost certainly
+become larger. No one, I presume, doubts that the large proportion
+which the size of man's brain bears to his body, compared to the same
+proportion in the gorilla or orang, is closely connected with his
+higher mental powers." "With respect to the lower animals," he
+says,[187] "M. E. Lartet,[188] by comparing the crania of tertiary and
+recent mammals belonging to the same groups, has come to the
+remarkable conclusion that the brain is generally larger and the
+convolutions are more complex in the more recent form."
+
+Sir E. Ray Lankester has sought to express in the simplest terms the
+implications of the increase in size of the cerebrum. "In what," he
+asks, "does the advantage of a larger cerebral mass consist?" "Man,"
+he replies, "is born with fewer ready-made tricks of the
+nerve-centres--these performances of an inherited nervous mechanism so
+often called by the ill-defined term 'instincts'--than are the monkeys
+or any other animal. Correlated with the absence of inherited
+ready-made mechanism, man has a greater capacity of developing in the
+course of his individual growth similar nervous mechanisms (similar
+to but not identical with those of 'instinct') than any other
+animal.... The power of being educated--'educability' as we may term
+it--is what man possesses in excess as compared with the apes. I think
+we are justified in forming the hypothesis that it is this
+'educability' which is the correlative of the increased size of the
+cerebrum." There has been natural selection of the more educable
+animals, for "the character which we describe as 'educability' can be
+transmitted, it is a congenital character. But the _results_ of
+education can _not_ be transmitted. In each generation they have to be
+acquired afresh, and with increased 'educability' they are more
+readily acquired and a larger variety of them.... The fact is that
+there is no community between the mechanisms of instinct and the
+mechanisms of intelligence, and that the latter are later in the
+history of the evolution of the brain than the former and can only
+develop in proportion as the former become feeble and defective."[189]
+
+In this statement we have a good example of the further development of
+views which Darwin foreshadowed but did not thoroughly work out. It
+states the biological case clearly and tersely. Plasticity of
+behaviour in special accommodation to special circumstances is of
+survival value; it depends upon acquired characters; it is correlated
+with increase in size and complexity of the cerebrum; under natural
+selection therefore the larger and more complex cerebrum as the organ
+of plastic behaviour has been the outcome of natural selection. We
+have thus the biological foundations for a further development of
+genetic psychology.
+
+There are diversities of opinion, as Darwin showed, with regard to the
+range of instinct in man and the higher animals as contrasted with
+lower types. Darwin himself said[190] that "Man, perhaps, has somewhat
+fewer instincts than those possessed by the animals which come next to
+him in the series." On the other hand, Prof. Wm. James says[191] that
+man is probably the animal with most instincts. The true position is
+that man and the higher animals have fewer complete and self-sufficing
+instincts than those which stand lower in the scale of mental
+evolution, but that they have an equally large or perhaps larger mass
+of instinctive raw material which may furnish the stuff to be
+elaborated by intelligent processes. There is, perhaps, a greater
+abundance of the primary tissue of experience to be refashioned and
+integrated by secondary modification; there is probably the same
+differentiation in relation to the determining biological ends, but
+there is at the outset less differentiation of the particular and
+specific modes of behaviour. The specialised instinctive performances
+and their concomitant experience-complexes are at the outset more
+indefinite. Only through acquired connections, correlated with
+experience, do they become definitely organised.
+
+The full working-out of the delicate and subtle relationship of
+instinct and educability--that is, of the hereditary and the acquired
+factors in the mental life--is the task which lies before genetic and
+comparative psychology. They interact throughout the whole of life,
+and their interactions are very complex. No one can read the chapters
+of _The Descent of Man_ which Darwin devotes to a consideration of the
+mental characters of man and animals without noticing, on the one
+hand, how sedulous he is in his search for hereditary foundations,
+and, on the other hand, how fully he realises the importance of
+acquired habits of mind. The fact that educability itself has innate
+tendencies--is in fact a partially differentiated educability--renders
+the unravelling of the factors of mental progress all the more
+difficult.
+
+In his comparison of the mental powers of men and animals it was
+essential that Darwin should lay stress on points of similarity rather
+than on points of difference. Seeking to establish a doctrine of
+evolution, with its basal concept of continuity of process and
+community of character, he was bound to render clear and to emphasise
+the contention that the difference in mind between man and the higher
+animals, great as it is, is one of degree and not of kind. To this end
+Darwin not only recorded a large number of valuable observations of
+his own, and collected a considerable body of information from
+reliable sources, he presented the whole subject in a new light and
+showed that a natural history of mind might be written and that this
+method of study offered a wide and rich field for investigation. Of
+course those who regarded the study of mind only as a branch of
+metaphysics smiled at the philosophical ineptitude of the mere man of
+science. But the investigation, on natural history lines, has been
+prosecuted with a large measure of success. Much indeed still remains
+to be done; for special training is required, and the workers are
+still few. Promise for the future is however afforded by the fact that
+investigation is prosecuted on experimental lines and that something
+like organised methods of research are taking form. There is now but
+little reliance on casual observations recorded by those who have not
+undergone the necessary discipline in these methods. There is also
+some change of emphasis in formulating conclusions. Now that the
+general evolutionary thesis is fully and freely accepted by those who
+carry on such researches, more stress is laid on the differentiation
+of the stages of evolutionary advance than on the fact of their
+underlying community of nature. The conceptual intelligence which is
+especially characteristic of the higher mental procedure of man is
+more firmly distinguished from the perceptual intelligence which he
+shares with the lower animals--distinguished now as a higher product
+of evolution, no longer as differing in origin or different in kind.
+Some progress has been made, on the one hand in rendering an account
+of intelligent profiting by experience under the guidance of pleasure
+and pain in the perceptual field, on lines predetermined by
+instinctive differentiation for biological ends, and on the other hand
+in elucidating the method of conceptual thought employed, for
+example, by the investigator himself in interpreting the perceptual
+experience of the lower animals.
+
+Thus there is a growing tendency to realise more fully that there are
+two orders of educability--first an educability of the perceptual
+intelligence based on the biological foundation of instinct, and
+secondly an educability of the conceptual intelligence which
+refashions and rearranges the data afforded by previous inheritance
+and acquisition. It is in relation to this second and higher order of
+educability that the cerebrum of man shows so large an increase of
+mass and a yet larger increase of effective surface through its rich
+convolutions. It is through educability of this order that the human
+child is brought intellectually and affectively into touch with the
+ideal constructions by means of which man has endeavoured, with more
+or less success, to reach an interpretation of nature, and to guide
+the course of the further evolution of his race--ideal constructions
+which form part of man's environment.
+
+It formed no part of Darwin's purpose to consider, save in broad
+outline, the methods, or to discuss in any fulness of detail the
+results of the process by which a differentiation of the mental
+faculties of man from those of the lower animals has been brought
+about--a differentiation the existence of which he again and again
+acknowledges. His purpose was rather to show that, notwithstanding
+this differentiation, there is basal community in kind. This must be
+remembered in considering his treatment of the biological foundations
+on which man's systems of ethics are built. He definitely stated that
+he approached the subject "exclusively from the side of natural
+history."[192] His general conclusion is that the moral sense is
+fundamentally identical with the social instincts, which have been
+developed for the good of the community; and he suggests that the
+concept which thus enables us to interpret the biological ground-plan
+of morals also enables us to frame a rational ideal of the moral end.
+"As the social instincts," he says,[193] "both of man and the lower
+animals have no doubt been developed by nearly the same steps, it
+would be advisable, if found practicable, to use the same definition
+in both cases, and to take as the standard of morality, the general
+good or welfare of the community, rather than the general happiness."
+But the kind of community for the good of which the social instincts
+of animals and primitive men were biologically developed may be
+different from that which is the product of civilisation, as Darwin no
+doubt realised. Darwin's contention was that conscience is a social
+instinct and has been evolved because it is useful to the tribe in the
+struggle for existence against other tribes. One the other hand J. S.
+Mill urged that the moral feelings are not innate but acquired, and
+Bain held the same view, believing that the moral sense is acquired by
+each individual during his life-time. Darwin, who notes[194] their
+opinion with his usual candour, adds that "on the general theory of
+evolution this is at least extremely improbable." It is impossible to
+enter into the question here: much turns on the exact connotation of
+the terms "conscience" and "moral sense," and on the meaning we attach
+to the statement that the moral sense is fundamentally identical with
+the social instincts.
+
+Presumably the majority of those who approach the subjects discussed
+in the third, fourth, and fifth chapters of _The Descent of Man_ in
+the full conviction that mental phenomena, not less than organic
+phenomena, have a natural genesis, would, without hesitation, admit
+that the intellectual and moral systems of civilised man are ideal
+constructions, the products of conceptual thought, and that as such
+they are, in their developed form, acquired. The moral sentiments are
+the emotional analogues of highly developed concepts. This does not
+however imply that they are outside the range of natural history
+treatment. Even though it may be desirable to differentiate the moral
+conduct of men from the social behaviour of animals (to which some
+such term as "pre-moral" or "quasi-moral" may be applied), still the
+fact remains that, as Darwin showed, there is abundant evidence of the
+occurrence of such social behaviour--social behaviour which, even
+granted that it is in large part intelligently acquired, and is itself
+so far a product of educability, is of survival value. It makes for
+that integration without which no social group could hold together and
+escape elimination. Furthermore, even if we grant that such behaviour
+is intelligently acquired, that is to say arises through the
+modification of hereditary instincts and emotions, the fact remains
+that only through these instinctive and emotional data is afforded the
+primary tissue of the experience which is susceptible of such
+modification.
+
+Darwin sought to show, and succeeded in showing, that for the
+intellectual and moral life there are instinctive foundations which a
+biological treatment alone can disclose. It is true that he did not in
+all cases analytically distinguish the foundations from the
+superstructure. Even to-day we are scarcely in a position to do so
+adequately. But his treatment was of great value in giving an impetus
+to further research. This value indeed can scarcely be over-estimated.
+And when the natural history of the mental operations shall have been
+written, the cardinal fact will stand forth, that the instinctive and
+emotional foundations are the outcome of biological evolution and have
+been ingrained in the race through natural selection. We shall more
+clearly realise that educability itself is a product of natural
+selection, though the specific results acquired through cerebral
+modifications are not transmitted through heredity. It will, perhaps,
+also be realised that the instinctive foundations of social behaviour
+are, for us, somewhat out of date and have undergone but little change
+throughout the progress of civilisation, because natural selection has
+long since ceased to be the dominant factor in human progress. The
+history of human progress has been mainly the history of man's higher
+educability, the products of which he has projected on to his
+environment. This educability remains on the average what it was a
+dozen generations ago; but the thought-woven tapestry of his
+surroundings is refashioned and improved by each succeeding
+generation. Few men have in greater measure enriched the
+thought-environment with which it is the aim of education to bring
+educable human beings into vital contact, than has Charles Darwin. His
+special field of work was the wide province of biology; but he did
+much to help us to realise that mental factors have contributed to
+organic evolution and that in man, the highest product of Evolution,
+they have reached a position of unquestioned supremacy.
+
+FOOTNOTES:
+
+[Footnote 153: _Origin of Species_ (6th edit.), p. 205.]
+
+[Footnote 154: _Descent of man_ (2nd edit. 1888), Vol. I. p. 99;
+Popular edit. p. 99.]
+
+[Footnote 155: _Ibid._ p. 99.]
+
+[Footnote 156: _The Expression of the Emotions_ (2nd edit.), p. 32.]
+
+[Footnote 157: _Descent of Man_, Vol. II. p. 435.]
+
+[Footnote 158: _Ibid._ 437, 438.]
+
+[Footnote 159: _Origin of Species_ (6th edit.), p. 205.]
+
+[Footnote 160: _Origin of Species_ (6th edit.), p. 205.]
+
+[Footnote 161: _Ibid._ p. 233.]
+
+[Footnote 162: _Ibid._ p. 205.]
+
+[Footnote 163: _Origin of Species_ (6th edit.), p. 233.]
+
+[Footnote 164: _Origin of Species_, pp. 210, 211.]
+
+[Footnote 165: Independently suggested, on somewhat different lines,
+by Profs. J. Mark Baldwin, Henry F. Osborn and the writer.]
+
+[Footnote 166: _Origin of Species_ (6th edit.), p. 206.]
+
+[Footnote 167: _Expression of the Emotions_, p. 13. The passage is
+here somewhat condensed.]
+
+[Footnote 168: _Ibid._ p. 368.]
+
+[Footnote 169: _Expression of the Emotions_, pp. 373, 374.]
+
+[Footnote 170: _Ibid._ p. 368.]
+
+[Footnote 171: _Ibid._ p. 369.]
+
+[Footnote 172: _Expression of the Emotions_, pp. 65 ff.]
+
+[Footnote 173: Cf. William James, _Principles of Psychology_, Vol. II.
+Chap. XXV, New York, 1890.]
+
+[Footnote 174: _Darwinism_, pp. 282, 283, London, 1889.]
+
+[Footnote 175: _Ibid._ p. 283.]
+
+[Footnote 176: _Darwinism_, pp. 283, 284.]
+
+[Footnote 177: _Descent of Man_ (2nd edit.), Vol. II. pp. 136, 137;
+(Popular edit.), pp. 642, 643.]
+
+[Footnote 178: _The Play of Animals_, p. 244, London, 1898.]
+
+[Footnote 179: _Ibid._ p. 283.]
+
+[Footnote 180: _Descent of Man_, Vol. II. p. 60; (Popular edit.), p.
+566.]
+
+[Footnote 181: _The Play of Animals_, p. 76.]
+
+[Footnote 182: _Ibid._ p. 75.]
+
+[Footnote 183: _The Play of Animals_ p. 24.]
+
+[Footnote 184: _Descent of Man_ (1st edit.), Chaps. II, III, V; (2nd
+edit.), Chaps. III, IV, V.]
+
+[Footnote 185: _Descent of Man_, Vol. I. pp. 70, 71; (Popular edit.),
+pp. 70, 71.]
+
+[Footnote 186: _Ibid._ p. 81.]
+
+[Footnote 187: _Ibid._ (Popular edit.), p. 82.]
+
+[Footnote 188: _Comptes Rendus des Sciences_, June 1, 1868.]
+
+[Footnote 189: _Nature_, Vol. LXI. pp. 624, 625 (1900).]
+
+[Footnote 190: _Descent of Man_, Vol. I. p. 100.]
+
+[Footnote 191: _Principles of Psychology_, Vol. II. p. 289.]
+
+[Footnote 192: _Descent of Man_, Vol. I. p. 149.]
+
+[Footnote 193: _Descent of Man_, p. 185.]
+
+[Footnote 194: _Ibid._ p. 150 (footnote).]
+
+
+
+
+VII
+
+THE INFLUENCE OF THE CONCEPTION OF EVOLUTION ON MODERN PHILOSOPHY
+
+BY H. HÖFFDING
+
+_Professor of Philosophy in the University of Copenhagen_
+
+
+I
+
+It is difficult to draw a sharp line between philosophy and natural
+science. The naturalist who introduces a new principle, or
+demonstrates a fact which throws a new light on existence, not only
+renders an important service to philosophy but is himself a
+philosopher in the broader sense of the word. The aim of philosophy in
+the stricter sense is to attain points of view from which the
+fundamental phenomena and the principles of the special sciences can
+be seen in their relative importance and connection. But philosophy in
+this stricter sense has always been influenced by philosophy in the
+broader sense. Greek philosophy came under the influence of logic and
+mathematics, modern philosophy under the influence of natural science.
+The name of Charles Darwin stands with those of Galileo, Newton, and
+Robert Mayer--names which denote new problems and great alterations in
+our conception of the universe.
+
+First of all we must lay stress on Darwin's own personality. His deep love
+of truth, his indefatigable inquiry, his wide horizon, and his steady
+self-criticism make him a scientific model, even if his results and
+theories should eventually come to possess mainly an historical interest.
+In the intellectual domain the primary object is to reach high summits
+from which wide surveys are possible, to reach them toiling honestly
+upwards by the way of experience, and then not to turn dizzy when a summit
+is gained. Darwinians have sometimes turned dizzy, but Darwin never. He saw
+from the first the great importance of his hypothesis, not only because of
+its solution of the old problem as to the value of the concept of species,
+not only because of the grand picture of natural evolution which it
+unrolls, but also because of the life and inspiration its method would
+impart to the study of comparative anatomy, of instinct and of heredity,
+and finally because of the influence it would exert on the whole conception
+of existence. He wrote in his note-book in the year 1837: "My theory would
+give zest to recent and fossil comparative anatomy; it would lead to the
+study of instinct, heredity, and mind-heredity, whole [of]
+metaphysics."[195]
+
+We can distinguish four main points in which Darwin's investigations
+possess philosophical importance.
+
+The evolution hypothesis is much older than Darwin; it is, indeed, one
+of the oldest guessings of human thought. In the eighteenth century is
+was put forward by Diderot and Lamettrie and suggested by Kant (1786).
+As we shall see later, it was held also by several philosophers in the
+first half of the nineteenth century. In his preface to _The Origin of
+Species_, Darwin mentions the naturalists who were his forerunners.
+But he has set forth the hypothesis of evolution in so energetic and
+thorough a manner that it perforce attracts the attention of all
+thoughtful men in a much higher degree than it did before the
+publication of the _Origin_.
+
+And further, the importance of his teaching rests on the fact that he,
+much more than his predecessors, even than Lamarck, sought a
+foundation for his hypothesis in definite facts. Modern science began
+by demanding--with Kepler and Newton--evidence of _varae causae_; this
+demand Darwin industriously set himself to satisfy--hence the wealth
+of material which he collected by his observations and his
+experiments. He not only revived an old hypothesis, but he saw the
+necessity of verifying it by facts. Whether the special cause on which
+he founded the explanation of the origin of species--Natural
+Selection--is sufficient, is now a subject of discussion. He himself
+had some doubt in regard to this question, and the criticisms which
+are directed against his hypothesis hit Darwinism rather than Darwin.
+In his indefatigable search for empirical evidence he is a model even
+for his antagonists: he has compelled them to approach the problems of
+life along other lines than those which were formerly followed.
+
+Whether the special cause to which Darwin appealed is sufficient or not, at
+least to it is probably due the greater part of the influence which he has
+exerted on the general trend of thought. "Struggle for existence" and
+"natural selection" are principles which have been applied, more or less,
+in every department of thought. Recent research, it is true, has discovered
+greater empirical discontinuity--leaps, "mutations"--whereas Darwin
+believed in the importance of small variations slowly accumulated. It has
+also been shown by the experimental method, which in recent biological work
+has succeeded Darwin's more historical method, that types once constituted
+possess great permanence, the fluctuations being restricted within clearly
+defined boundaries. The problem has become more precise, both as to
+variation and as to heredity. The inner conditions of life have in both
+respects shown a greater independence than Darwin had supposed in his
+theory, though he always admitted that the cause of variation was to him a
+great enigma, "a most perplexing problem," and that the struggle for life
+could only occur where variation existed. But, at any rate, it was of the
+greatest importance that Darwin gave a living impression of the struggle
+for life which is everywhere going on, and to which even the highest forms
+of existence must be amenable. The philosophical importance of these ideas
+does not stand or fall with the answer to the question, whether natural
+selection is a sufficient explanation of the origin of species or not it
+has an independent, positive value for everyone who will observe life and
+reality with an unbiased mind.
+
+In accentuating the struggle for life Darwin stands as a
+characteristically English thinker: he continues a train of ideas
+which Hobbes and Malthus had already begun. Moreover in his critical
+views as to the conception of species he had English forerunners; in
+the middle ages Occam and Duns Scotus, in the eighteenth century
+Berkeley and Hume. In his moral philosophy, as we shall see later, he
+is an adherent of the school which is represented by Hutcheson, Home
+and Adam Smith. Because he is no philosopher in the stricter sense of
+the term, it is of great interest to see that his attitude of mind is
+that of the great thinkers of his nation.
+
+In considering Darwin's influence on philosophy we will begin with an
+examination of the attitude of philosophy to the conception of
+evolution at the time when _The Origin of Species_ appeared. We will
+then examine the effects which the theory of evolution, and especially
+the idea of the struggle for life, has had, and naturally must have,
+on the discussion of philosophical problems.
+
+
+II
+
+When _The Origin of Species_ appeared fifty years ago Romantic
+speculation, Schelling's and Hegel's philosophy, still reigned on the
+continent, while in England Positivism, the philosophy of Comte and
+Stuart Mill, represented the most important trend of thought. German
+speculation had much to say on evolution, it even pretended to be a
+philosophy of evolution. But then the word "evolution" was to be taken
+in an ideal, not in a real, sense. To speculative thought the forms
+and types of nature formed a system of ideas, within which any form
+could lead us by continuous transitions to any other. It was a
+classificatory system which was regarded as a divine world of thought
+or images, within which metamorphoses could go on--a condition
+comparable with that in the mind of the poet when one image follows
+another with imperceptible changes. Goethe's ideas of evolution, as
+expressed in his _Metamorphosen der Pflanzen und der Thiere_, belong
+to this category; it is, therefore, incorrect to call him a forerunner
+of Darwin. Schelling and Hegel held the same idea; Hegel expressly
+rejected the conception of a real evolution in time as coarse and
+materialistic. "Nature," he says, "is to be considered as a _system of
+stages_, the one necessarily arising from the other, and being the
+nearest truth of that from which it proceeds; but not in such a way
+that the one is _naturally_ generated by the other; on the contrary
+[their connection lies] in the inner idea which is the ground of
+nature. The _metamorphosis_ can be ascribed only to the notion as
+such, because it alone is evolution.... It has been a clumsy idea in
+the older as well as in the newer philosophy of nature, to regard the
+transformation and the transition from one natural form and sphere to
+a higher as an outward and actual production."[196]
+
+The only one of the philosophers of Romanticism who believed in a
+real, historical evolution, a real production of new species, was
+Oken.[197] Danish philosophers, such as Treschow (1812) and Sibbern
+(1846), have also broached the idea of an historical evolution of all
+living beings from the lowest to the highest. Schopenhauer's
+philosophy has a more realistic character than that of Schelling's and
+Hegel's, his diametrical opposites, although he also belongs to the
+romantic school of thought. His philosophical and psychological views
+were greatly influenced by French naturalists and philosophers,
+especially by Cabanis and Lamarck. He praises the "ever memorable
+Lamarck," because he laid so much stress on the "will to live." But he
+repudiates as a "wonderful error" the idea that the organs of animals
+should have reached their present perfection through a development in
+time, during the course of innumerable generations. It was, he said, a
+consequence of the low standard of contemporary French philosophy,
+that Lamarck came to the idea of the construction of living beings in
+time through succession![198]
+
+The positivistic stream of thought was not more in favour of a real
+evolution than was the Romantic school. Its aim was to adhere to
+positive facts: it looked with suspicion on far-reaching speculation.
+Comte laid great stress on the discontinuity found between the
+different kingdoms of nature, as well as within each single kingdom.
+As he regarded as unscientific every attempt to reduce the number of
+physical forces, so he rejected entirely the hypothesis of Lamarck
+concerning the evolution of species; the idea of species would in his
+eyes absolutely lose its importance if a transition from species to
+species under the influence of conditions of life were admitted. His
+disciples (Littré, Robin) continued to direct against Darwin the
+polemics which their master had employed against Lamarck. Stuart Mill,
+who, in the theory of knowledge, represented the empirical or
+positivistic movement in philosophy--like his English forerunners from
+Locke to Hume--founded his theory of knowledge and morals on the
+experience of the single individual. He sympathised with the theory of
+the original likeness of all individuals and derived their
+differences, on which he practically and theoretically laid much
+stress, from the influence both of experience and education, and,
+generally, of physical and social causes. He admitted an individual
+evolution, and, in the human species, an evolution based on social
+progress; but no physiological evolution of species. He was afraid
+that the hypothesis of heredity would carry us back to the old theory
+of "innate" ideas.
+
+Darwin was more empirical than Comte and Mill; experience disclosed to
+him a deeper continuity than they could find; closer than before the
+nature and fate of the single individual were shown to be interwoven
+in the great web binding the life of the species with nature as a
+whole. And the continuity which so many idealistic philosophers could
+find only in the world of thought, he showed to be present in the
+world of reality.
+
+
+III
+
+Darwin's energetic renewal of the old idea of evolution has its chief
+importance in strengthening the conviction of this real continuity in
+the world, of continuity in the series of form and events. It was a
+great support for all those who were prepared to base their conception
+of life on scientific grounds. Together with the recently discovered
+law of the conservation of energy, it helped to produce the great
+realistic movement which characterises the last third of the
+nineteenth century. After the decline of the Romantic movement people
+wished to have firmer ground under their feet and reality now asserted
+itself in a more emphatic manner than in the period of Romanticism. It
+was easy for Hegel to proclaim that "the real" was "the rational," and
+that "the rational" was "the real": reality itself existed for him
+only in the interpretation of ideal reason, and if there was anything
+which could not be merged in the higher unity of thought, then it was
+only an example of the "impotence of nature to hold to the idea." But
+now concepts are to be founded on nature and not on any system of
+categories too confidently deduced _à priori_. The new devotion to
+nature had its recompense in itself, because the new points of view
+made us see that nature could indeed "hold to ideas," though perhaps
+not to those which we had cogitated beforehand.
+
+A most important question for philosophers to answer was whether the
+new views were compatible with an idealistic conception of life and
+existence. Some proclaimed that we have now no need of any philosophy
+beyond the principles of the conservation of matter and energy and the
+principle of natural evolution: existence should and could be
+definitely and completely explained by the laws of material nature.
+But abler thinkers saw that the thing was not so simple. They were
+prepared to give the new views their just place and to examine what
+alterations the old views must undergo in order to be brought into
+harmony with the new data.
+
+The realistic character of Darwin's theory was shown not only in the
+idea of natural continuity, but also, and not least, in the idea of
+the cause whereby organic life advances step by step. This idea--the
+idea of the struggle for life--implied that nothing could persist, if
+it had no power to maintain itself under the given conditions. Inner
+value alone does not decide. Idealism was here put to its hardest
+trial. In continuous evolution it could perhaps still find an analogy
+to the inner evolution of ideas in the mind; but in the demand for
+power in order to struggle with outward conditions Realism seemed to
+announce itself in its most brutal form. Every form of Idealism had to
+ask itself seriously how it was going to "struggle for life" with this
+new Realism.
+
+We will now give a short account of the position which leading
+thinkers in different countries have taken up in regard to this
+question.
+
+I. Herbert Spencer was the philosopher whose mind was best prepared by his
+own previous thinking to admit the theory of Darwin to a place in his
+conception of the world. His criticism of the arguments which had been put
+forward against the hypothesis of Lamarck, showed that Spencer, as a young
+man, was an adherent to the evolution idea. In his _Social Statics_ (1850)
+he applied this idea to human life and moral civilisation. In 1852 he wrote
+an essay on _The Development Hypothesis_, in which he definitely stated his
+belief that the differentiation of species, like the differentiation within
+a single organism, was the result of development. In the first edition of
+his _Psychology_ (1855) he took a step which put him in opposition to the
+older English school (from Locke to Mill): he acknowledged "innate ideas"
+so far as to admit the tendency of acquired habits to be inherited in the
+course of generations, so that the nature and functions of the individual
+are only to be understood through its connection with the life of the
+species. In 1857, in his essay on _Progress_, he propounded the law of
+differentiation as a general law of evolution, verified by examples from
+all regions of experience, the evolution of species being only one of these
+examples. On the effect which the appearance of _The Origin of Species_ had
+on his mind he writes in his _Autobiography_: "Up to that time ... I held
+that the sole cause of organic evolution is the inheritance of
+functionally-produced modifications. The _Origin of Species_ made it clear
+to me that I was wrong, and that the larger part of the facts cannot be due
+to any such cause.... To have the theory of organic evolution justified was
+of course to get further support for that theory of evolution at large with
+which ... all my conceptions were bound up."[199] Instead of the
+metaphorical expression "natural selection," Spencer introduced the term
+"survival of the fittest," which found favour with Darwin as well as with
+Wallace.
+
+In working out his ideas of evolution, Spencer found that
+differentiation was not the only form of evolution. In its simplest
+form evolution is mainly a concentration, previously scattered
+elements being integrated and losing independent movement.
+Differentiation is only forthcoming when minor wholes arise within a
+greater whole. And the highest form of evolution is reached when there
+is a harmony between concentration and differentiation, a harmony
+which Spencer calls equilibration and which he defines as a moving
+equilibrium. At the same time this definition enables him to
+illustrate the expression "survival of the fittest." "Every living
+organism exhibits such a moving equilibrium--a balanced set of
+functions constituting its life; and the overthrow of this balanced
+set of functions or moving equilibrium is what we call death. Some
+individuals in a species are so constituted that their moving
+equilibria are less easily overthrown than those of other
+individuals; and these are the fittest which survive, or, in Mr.
+Darwin's language, they are the select which nature preserves."[200]
+Not only in the domain of organic life, but in all domains, the summit
+of evolution is, according to Spencer, characterised by such a
+harmony--by a moving equilibrium.
+
+Spencer's analysis of the concept of evolution, based on a great
+variety of examples, has made this concept clearer and more definite
+than before. It contains the three elements; integration,
+differentiation and equilibration. It is true that a concept which is
+to be valid for all domains of experience must have an abstract
+character, and between the several domains there is, strictly
+speaking, only a relation of analogy. So there is only analogy between
+psychical and physical evolution. But this is no serious objection,
+because general concepts do not express more than analogies between
+the phenomena which they represent. Spencer takes his leading forms
+from the material world in defining evolution (in the simplest form)
+as integration of matter and dissipation of movement; but as he--not
+always quite consistently[201]--assumed a correspondence of mind and
+matter, he could very well give these terms an indirect importance for
+psychical evolution. Spencer has always, in my opinion with full
+right, repudiated the ascription of materialism. He is no more a
+materialist than Spinoza. In his _Principles of Psychology_ (§ 63) he
+expressed himself very clearly: "Though it seems easier to translate
+so-called matter into so-called spirit, than to translate so-called
+spirit into so-called matter--which latter is indeed wholly
+impossible--yet no translation can carry us beyond our symbols." These
+words lead us naturally to a group of thinkers whose starting-point
+was psychical evolution. But we have still one aspect of Spencer's
+philosophy to mention.
+
+Spencer founded his "laws of evolution" on an inductive basis, but he
+was convinced that they could be deduced from the law of the
+conservation of energy. Such a deduction is, perhaps, possible for the
+more elementary forms of evolution, integration and differentiation;
+but it is not possible for the highest form, the equilibration, which
+is a harmony of integration and differentiation. Spencer can no more
+deduce the necessity for the eventual appearance of "moving
+equilibria" of harmonious totalities than Hegel could guarantee the
+"higher unities" in which all contradictions should be reconciled. In
+Spencer's hands the theory of evolution acquired a more decidedly
+optimistic character than in Darwin's; but I shall deal later with the
+relation of Darwin's hypothesis to the opposition of optimism and
+pessimism.
+
+II. While the starting-point of Spencer was biological or
+cosmological, psychical evolution being conceived as in analogy with
+physical, a group of eminent thinkers--in Germany Wundt, in France
+Fouillée, in Italy Ardigò--took, each in his own manner, their
+starting-point in psychical evolution as an original fact and as a
+type of all evolution, the hypothesis of Darwin coming in as a
+corroboration and as a special example. They maintain the continuity
+of evolution; they find this character most prominent in psychical
+evolution, and this is for them a motive to demand a corresponding
+continuity in the material, especially in the organic domain.
+
+To Wundt and Fouillée the concept of will is prominent. They see the
+type of all evolution in the transformation of the life of will from
+blind impulse to conscious choice; the theories of Lamarck and Darwin
+are used to support the view that there is in nature a tendency to
+evolution in steady reciprocity with external conditions. The struggle
+for life is here only a secondary fact. Its apparent prominence is
+explained by the circumstance that the influence of external
+conditions is easily made out, while inner conditions can be verified
+only through their effects. For Ardigò the evolution of thought was
+the starting-point and the type: in the evolution of a scientific
+hypothesis we see a progress from the indefinite (_indistinto_) to the
+definite (_distinto_), and this is a characteristic of all evolution,
+as Ardigò has pointed out in a series of works. The opposition between
+_indistinto_ and _distinto_ corresponds to Spencer's opposition
+between homogeneity and heterogeneity. The hypothesis of the origin of
+differences of species from more simple forms is a special example of
+the general law of evolution.
+
+In the views of Wundt and Fouillée we find the fundamental idea of
+idealism psychical phenomena as expressions of the innermost nature of
+existence. They differ from the older Idealism in the great stress
+which they lay on evolution as a real, historical process which is
+going on through steady conflict with external conditions. The
+Romantic dread of reality is broken. It is beyond doubt that Darwin's
+emphasis on the struggle for life as a necessary condition of
+evolution has been a very important factor in carrying philosophy back
+to reality from the heaven of pure ideas. The philosophy of Ardigò, on
+the other side, appears more as a continuation and deepening of
+positivism, though the Italian thinker arrived at his point of view
+independently of French-English positivism. The idea of continuous
+evolution is here maintained in opposition to Comte's and Mill's
+philosophy of discontinuity. From Wundt and Fouillée Ardigò differs in
+conceiving psychical evolution not as an immediate revelation of the
+innermost nature of existence, but only as a single, though the most
+accessible example, of evolution.
+
+III. To the French philosophers Boutroux and Bergson, evolution proper
+is continuous and qualitative, while outer experience and physical
+science give us fragments only, sporadic processes and mechanical
+combinations. To Bergson, in his recent work _L'Evolution Créatrice_,
+evolution consists in an _élan de vie_ which to our fragmentary
+observation and analytic reflexion appears as broken into a manifold
+of elements and processes. The concept of matter in its scientific
+form is the result of this breaking asunder, essential for all
+scientific reflexion. In these conceptions the strongest opposition
+between inner and outer conditions of evolution is expressed: in the
+domain of internal conditions spontaneous development of qualitative
+forms--in the domain of external conditions discontinuity and
+mechanical combination.
+
+We see, then, that the theory of evolution has influenced philosophy
+in a variety of forms. It has made idealistic thinkers revise their
+relation to the real world; it has led positivistic thinkers to find a
+closer connection between the facts on which they based their views;
+it has made us all open our eyes for new possibilities to arise
+through the _prima facie_ inexplicable "spontaneous" variations which
+are the condition of all evolution. This last point is one of peculiar
+interest. Deeper than speculative philosophy and mechanical science
+saw in the days of their triumph, we catch sight of new streams, whose
+sources and laws we have still to discover. Most sharply does this
+appear in the theory of mutation, which is only a stronger
+accentuation of a main point in Darwinism. It is interesting to see
+that an analogous problem comes into the foreground in physics through
+the discovery of radioactive phenomena, and in psychology through the
+assumption of psychical new formations (as held by Boutroux, William
+James and Bergson). From this side, Darwin's ideas, as well as the
+analogous ideas in other domains, incite us to renewed examination of
+our first principles, their rationality and their value. On the other
+hand, his theory of the struggle for existence challenges us to
+examine the conditions and discuss the outlook as to the persistence
+of human life and society and of the values that belong to them. It is
+not enough to hope (or fear?) the rising of new forms; we have also to
+investigate the possibility of upholding the forms and ideals which
+have hitherto been the bases of human life. Darwin has here given his
+age the most earnest and most impressive lesson. This side of Darwin's
+theory is of peculiar interest to some special philosophical problems
+to which I now pass.
+
+
+IV
+
+Among philosophical problems the problem of knowledge has in the last
+century occupied a foremost place. It is natural, then, to ask how
+Darwin and the hypothesis whose most eminent representative he is,
+stand to this problem.
+
+Darwin started an hypothesis. But every hypothesis is won by inference
+from certain presuppositions, and every inference is based on the
+general principles of human thought. The evolution hypothesis
+presupposes, then, human thought and its principles. And not only the
+abstract logical principles are thus pre-supposed. The evolution
+hypothesis purports to be not only a formal arrangement of phenomena,
+but to express also the law of a real process. It supposes, then, that
+the real data--all that in our knowledge which we do not produce
+ourselves, but which we in the main simply receive--are subject to
+laws which are at least analogous to the logical relations of our
+thoughts; in other words, it assumes the validity of the principle of
+causality. If organic species could arise without cause there would be
+no use in framing hypotheses. Only if we assume the principle of
+causality, is there a problem to solve.
+
+Though Darwinism has had a great influence on philosophy considered as
+a striving after a scientific view of the world, yet here is a point
+of view--the epistemological--where philosophy is not only independent
+but reaches beyond any result of natural science. Perhaps it will be
+said: the powers and functions of organic beings only persist (perhaps
+also only arise) when they correspond sufficiently to the conditions
+under which the struggle of life is to go on. Human thought itself is,
+then, a variation (or a mutation) which has been able to persist and
+to survive. Is not, then, the problem of knowledge solved by the
+evolution hypothesis? Spencer had given an affirmative answer to this
+question before the appearance of _The Origin of Species_. For the
+individual, he said, there is an _à priori_, original, basis (or
+_Anlage_) for all mental life; but in the species all powers have
+developed in reciprocity with extendal conditions. Knowledge is here
+considered from the practical point of view, as a weapon in the
+struggle for life, as an "organon" which has been continuously in use
+for generations. In recent years the economic or pragmatic
+epistemology, as developed by Avenarius and Mach in Germany, and by
+James in America, points in the same direction. Science, it is said,
+only maintains those principles and presuppositions which are
+necessary to the simplest and clearest orientation be applied to
+experience and to practical work, will successively be eliminated.
+
+In these views a striking and important application is made of the
+idea of struggle for life to the development of human thought. Thought
+must, as all other things in the world, struggle for life. But this
+whole consideration belongs to psychology, not to the theory of
+knowledge (epistemology), which is concerned only with the validity of
+knowledge, not with its historical origin. Every hypothesis to explain
+the origin of knowledge must submit to cross-examination by the theory
+of knowledge, because it works with the fundamental forms and
+principles of human thought. We cannot go further back than these
+forms and principles, which it is the aim of epistemology to ascertain
+and for which no further reason can be given.[202]
+
+But there is another side of the problem which is, perhaps, of more
+importance and which epistemology generally overlooks. If new
+variations can arise, not only in organic but perhaps also in
+inorganic nature, new tasks are placed before the human mind. The
+question is, then, if it has forms in which there is room for the new
+matter? We are here touching a possibility which the great master of
+epistemology did not bring to light. Kant supposed confidently that no
+other matter of knowledge could stream forth from the dark source
+which he called "the thing-in-itself," than such as could be
+synthesised in our existing forms of knowledge. He mentions the
+possibility of other forms than the human, and warns us against the
+dogmatic assumption that the human conception of existence should be
+absolutely adequate. But he seems to be quite sure that the
+thing-in-itself works constantly, and consequently always gives us
+only what our powers can master. This assumption was a consequence of
+Kant's rationalistic tendency, but one for which no warrant can be
+given. Evolutionism and systematism are opposing tendencies which can
+never be absolutely harmonised one with the other. Evolution may at
+any time break some form which the system-monger regards as finally
+established. Darwin himself felt a great difference in looking at
+variation as an evolutionist and as a systematist. When he was working
+at his evolution theory, he was very glad to find variations; but they
+were a hindrance to him when he worked as a systematist, in preparing
+his work on Cirripedia. He says in a letter: "I had thought the same
+parts of the same species more resemble (than they do anyhow in
+Cirripedia) objects cast in the same mould. Systematic work would be
+easy were it not for this confounded variation, which, however, is
+pleasant to me as a speculatist, though odious to me as a
+systematist."[203] He could indeed be angry with variations even as an
+evolutionist; but then only because he could not explain them, not
+because he could not classify them. "If, as I must think, external
+conditions produce little _direct_ effect, what the devil determines
+each particular variation?"[204] What Darwin experienced in this
+particular domain holds good of all knowledge. All knowledge is
+systematic, in so far as it strives to put phenomena in quite definite
+relations, one to another. But the systematisation can never be
+complete. And here Darwin has contributed much to widen the world, for
+us. He has shown us forces and tendencies in nature which make
+absolute systems impossible, at the same time that they give us new
+objects and problems. There is still a place for what Lessing called
+"the unceasing striving after truth," while "absolute truth" (in the
+sense of a closed system) is unattainable so long as life and
+experience are going on.
+
+There is here a special remark to be made. As we have seen above,
+recent research has shown that natural selection or struggle for life
+is no explanation of variations. Hugo de Vries distinguishes between
+partial and embryonal variations, or between variations and mutations,
+only the last-named being heritable, and therefore of importance for
+the origin of new species. But the existence of variations is not only
+of interest for the problem of the origin of species; it has also a
+more general interest. An individual does not lose its importance for
+knowledge, because its qualities are not heritable. On the contrary,
+in higher beings at least, individual peculiarities will become more
+and more independent objects of interest. Knowledge takes account of
+the biographies not only of species, but also of individuals: it seeks
+to find the law of development of the single individual.[205] As
+Leibnitz said long ago, individuality consists in the law of the
+changes of a being: "La loi du changement fait l'individualité de
+chaque substance." Here is a world which is almost new for science,
+which till now has mainly occupied itself with general laws and forms.
+But these are ultimately only means to understand the individual
+phenomena, in whose nature and history a manifold of laws and forms
+always coöperate. The importance of this remark will appear in the
+sequel.
+
+
+V
+
+To many people the Darwinian theory of natural selection or struggle
+for existence seemed to change the whole conception of life, and
+particularly all the conditions on which the validity of ethical ideas
+depends. If only that has persistence which can be adapted to a given
+condition, what will then be the fate of our ideals, of our standards
+of good and evil? Blind force seems to reign, and the only thing that
+counts seems to be the most heedless use of power. Darwinism, it was
+said, has proclaimed brutality. No other difference seems permanent
+save that between the sound, powerful and happy on the one side, the
+sick, feeble and unhappy on the other; and every attempt to alleviate
+this difference seems to lead to general enervation. Some of those who
+interpreted Darwinism in this manner felt an aesthetic delight in
+contemplating the heedlessness and energy of the great struggle for
+existence and anticipated the realisation of a higher human type as
+the outcome of it: so Nietzsche and his followers. Others recognising
+the same consequences in Darwinism regarded these as one of the
+strongest objections against it; so Dühring and Kropotkin (in his
+earlier works).
+
+This interpretation of Darwinism was frequent in the interval between
+the two main works of Darwin--_The Origin of Species_ and _The Descent
+of Man_. But even during this interval it was evident to an attentive
+reader that Darwin himself did not found his standard of good and evil
+on the features of the life of nature he had emphasised so strongly.
+He did not justify the ways along which nature reached its ends; he
+only pointed them out. The "real" was not to him, as to Hegel, one
+with the "rational." Darwin has, indeed, by his whole conception of
+nature, rendered a great service to ethics in making the difference
+between the life of nature and the ethical life appear in so strong a
+light. The ethical problem could now be stated in a sharper form than
+before. But this was not the first time that the idea of the struggle
+for life was put in relation to the ethical problem. In the
+seventeenth century Thomas Hobbes gave the first impulse to the whole
+modern discussion of ethical principles in his theory of _bellum
+omnium contra omnes_. Men, he taught, are in the state of nature
+enemies one of another, and they live either in fright or in the glory
+of power. But it was not the opinion of Hobbes that this made ethics
+impossible. On the contrary, he found a standard for virtue and vice
+in the fact that some qualities and actions have a tendency to bring
+us out of the state of war and to secure peace, while other qualities
+have a contrary tendency. In the eighteenth century even Immanuel
+Kant's ideal ethics had--so far as can be seen--a similar origin.
+Shortly before the foundation of his definitive ethics, Kant wrote his
+_Idee zu einer allgemeinen Weltgeschichte_ (1784), where--in a way
+which reminds us of Hobbes, and is prophetic of Darwin--he describes
+the forward-driving power of struggle in the human world. It is here
+as with the struggle of the trees for light and air, through which
+they compete with one another in height. Anxiety about war can only be
+allayed by an ordinance which gives everyone his full liberty under
+acknowledgment of the equal liberty of others. And such ordinance and
+acknowledgment are also attributes of the content of the moral law, as
+Kant proclaimed it in the year after the publication of his essay
+(1785).[206] Kant really came to his ethics by the way of evolution,
+though he afterwards disavowed it. Similarly the same line of thought
+may be traced in Hegel though it has been disguised in the form of
+speculative dialectics.[207] And in Schopenhauer's theory of the blind
+will to live and its abrogation by the ethical feeling, which is
+founded on universal sympathy, we have a more individualistic form of
+the same idea.
+
+It was, then, not entirely a foreign point of view which Darwin
+introduced into ethical thought, even if we take no account of the
+poetical character of the word "struggle" and of the more direct
+adaptation, through the use and non-use of power, which Darwin also
+emphasised. In _The Descent of Man_ he has devoted a special
+chapter[208] to a discussion of the origin of the ethical
+consciousness. The characteristic expression of this consciousness he
+found, just as Kant did, in the idea of "ought"; it was the origin of
+this new idea which should be explained. His hypothesis was that the
+ethical "ought" has its origin in the social and parental instincts,
+which, as well as other instincts (e.g. the instinct of
+self-preservation), lie deeper than pleasure and pain. In many
+species, not least in the human species, these instincts are fostered
+by natural selection; and when the powers of memory and comparison are
+developed, so that single acts can be valued according to the claims
+of the deep social instinct, then consciousness of duty and remorse
+are possible. Blind instinct has developed to conscious ethical will.
+
+As already stated, Darwin, as a moral philosopher belongs to the
+school that was founded by Shaftesbury, and was afterwards represented
+by Hutcheson, Hume, Adam Smith, Comte and Spencer. His merit is,
+first, that he has given this tendency of thought a biological
+foundation, and that he has stamped on it a doughty character in
+showing that ethical ideas and sentiments, rightly conceived, are
+forces which are at work in the struggle for life.
+
+There are still many questions to solve. Not only does the ethical
+development within the human species contain features still
+unexplained;[209] but we are confronted by the great problem whether
+after all a genetic historical theory can be of decisive importance
+here. To every consequent ethical consciousness there is a standard of
+value, a primordial value which determines the single ethical
+judgments as their last presupposition, and the "rightness" of this
+basis, the "value" of this value can as little be discussed as the
+"rationality" of our logical principles. There is here revealed a
+possibility of ethical scepticism which evolutionistic ethics (as well
+as intuitive or rationalistic ethics) has overlooked. No demonstration
+can show that the results of the ethical development are definitive
+and universal. We meet here again with the important opposition of
+systematisation and evolution. There will, I think, always be an open
+question here, though comparative ethics, of which we have so far only
+the first attempts, can do much to throw light on it.
+
+It would carry us too far to discuss all the philosophical works on
+ethics, which have been influenced directly or indirectly by
+evolutionism. I may, however, here refer to the book of C. M.
+Williams, _A Review of the Systems of Ethics founded on the Theory of
+Evolution_,[210] in which, besides Darwin, the following authors are
+reviewed: Wallace, Haeckel, Spencer, Fiske, Rolph, Barratt, Stephen,
+Carneri, Höffding, Gizycki, Alexander, Rée. As works which criticise
+evolutionistic ethics from an intuitive point of view and in an
+instructive way, may be cited: Guyau, _La morale anglaise
+contemporaine_,[211] and Sorley, _Ethics of Naturalism_. I will only
+mention some interesting contributions to ethical discussion which can
+be found in Darwinism besides the idea of struggle for life.
+
+The attention which Darwin has directed to variations has opened our
+eyes to the differences in human nature as well as in nature
+generally. There is here a fact of great importance for ethical
+thought, no matter from what ultimate premiss it starts. Only from a
+very abstract point of view can different individuals be treated in
+the same manner. The most eminent ethical thinkers, men such as Jeremy
+Bentham and Immanuel Kant, who discussed ethical questions from very
+opposite standpoints, agreed in regarding all men as equal in respect
+of ethical endowment. In regard to Bentham, Leslie Stephen remarks:
+"He is determined to be thoroughly empirical, to take men as he found
+them. But his utilitarianism supposed that men's views of happiness
+and utility were uniform and clear, and that all that was wanted was
+to show them the means by which their ends could be reached."[212] And
+Kant supposed that every man would find the "categorical imperative"
+in his consciousness, when he came to sober reflexion, and that all
+would have the same qualifications to follow it. But if continual
+variations, great or small, are going on in human nature, it is the
+duty of ethics to make allowance for them, both in making claims, and
+in valuing what is done. A new set of ethical problems have their
+origin here.[213] It is an interesting fact that Stuart Mill's book
+_On Liberty_ appeared in the same year as _The Origin of Species_.
+Though Mill agreed with Bentham about the original equality of all
+men's endowments, he regarded individual differences as a necessary
+result of physical and social influences, and he claimed that free
+play shall be allowed to differences of character so far as is
+possible without injury to other men. It is a condition of individual
+and social progress that a man's mode of action should be determined
+by his own character and not by tradition and custom, nor by abstract
+rules. This view was to be corroborated by the theory of Darwin.
+
+But here we have reached a point of view from which the criticism,
+which in recent years has often been directed against Darwin--that
+small variations are of no importance in the struggle for life--is of
+no weight. From an ethical standpoint, and particularly from the
+ethical standpoint of Darwin himself, it is a duty to foster
+individual differences that can be valuable, even though they can
+neither be of service for physical preservation nor be physically
+inherited. The distinction between variation and mutation is here
+without importance. It is quite natural that biologists should be
+particularly interested in such variations as can be inherited and
+produce new species. But in the human world there is not only a
+physical, but also a mental and social heredity. When an ideal human
+character has taken form, then there is shaped a type, which through
+imitation and influence can become an important factor in subsequent
+development, even if it cannot form a species in the biological sense
+of the word. Spiritually strong men often succumb in the physical
+struggle for life; but they can nevertheless be victorious through the
+typical influence they exert, perhaps on very distant generations, if
+the remembrance of them is kept alive, be it in legendary or in
+historical form. Their very failure can show that a type has taken
+form which is maintained at all risks, a standard of life which is
+adhered to in spite of the strongest opposition. The question "to be
+or not to be" can be put from very different levels of being: it has
+too often been considered a consequence of Darwinism that this
+question is only to be put from the lowest level. When a stage is
+reached, where ideal (ethical, intellectual, aesthetic) interests are
+concerned, the struggle for life is a struggle for the preservation of
+this stage. The giving up of a higher standard of life is a sort of
+death; for there is not only a physical, there is also a spiritual,
+death.
+
+
+VI
+
+The Socratic character of Darwin's mind appears in his wariness in
+drawing the last consequences of his doctrine, in contrast both with
+the audacious theories of so many of his followers and with the
+consequences which his antagonists were busy in drawing. Though he, as
+we have seen, saw from the beginning that his hypothesis would
+occasion "a whole of metaphysics," he was himself very reserved as to
+the ultimate questions, and his answers to such questions were
+extorted from him.
+
+As to the question of optimism and pessimism, Darwin held that though
+pain and suffering were very often the ways by which animals were led
+to pursue that course of action which is most beneficial to the
+species, yet pleasurable feelings were the most habitual guides. "We
+see this in the pleasure from exertion, even occasionally from great
+exertion of the body or mind, in the pleasure of our daily meals, and
+especially in the pleasure derived from sociability, and from loving
+our families." But there was to him so much suffering in the world
+that it was a strong argument against the existence of an intelligent
+First Cause.[214]
+
+It seems to me that Darwin was not so clear on another question, that
+of the relation between improvement and adaptation. He wrote to Lyell:
+"When you contrast natural selection and 'improvement,' you seem
+always to overlook ... that every step in the natural selection of
+each species implies improvement in that species _in relation to its
+condition of life_.... Improvement implies, I suppose, _each form
+obtaining many parts or organs_, all excellently adapted for their
+functions." "All this," he adds, "seems to me quite compatible with
+certain forms fitted for simple conditions, remaining unaltered, or
+being, degraded."[215] But the great question is, if the conditions of
+life will in the long run favour "improvement" in the sense of
+differentiation (or harmony of differentiation and integration). Many
+beings are best adapted to their conditions of life if they have few
+organs and few necessities. Pessimism would not only be the
+consequence, if suffering outweighed happiness, but also if the most
+elementary forms of happiness were predominant, or if there were a
+tendency to reduce the standard of life to the simplest possible, the
+contentment of inertia or stable equilibrium. There are animals which
+are very highly differentiated and active in their young state, but
+later lose their complex organisation and concentrate themselves on
+the one function of nutrition. In the human world analogies to this
+sort of adaptation are not wanting. Young "idealists" very often end
+as old "Philistines." Adaptation and progress are not the same.
+
+Another question of great importance in respect to human evolution is,
+whether there will be always a possibility for the existence of an
+impulse to progress, an impulse to make great claims on life, to be
+active and to alter the conditions of life instead of adapting to them
+in a passive manner. Many people do not develop because they have too
+few necessities, and because they have no power to imagine other
+conditions of life than those under which they live. In his remarks on
+"the pleasure from exertion" Darwin has a point of contact with the
+practical idealism of former times--with the ideas of Lessing and
+Goethe, of Condorcet and Fichte. The continual striving which was the
+condition of salvation to Faust's soul, is also the condition of
+salvation to mankind. There is a holy fire which we ought to keep
+burning, if adaptation is really to be improvement. If, as I have
+tried to show in my _Philosophy of Religion_, the innermost core of
+all religion is faith in the persistence of value in the world, and if
+the highest values express themselves in the cry "Excelsior!" then the
+capital point is, that this cry should always be heard and followed.
+We have here a corollary of the theory of evolution in its application
+to human life.
+
+Darwin declared himself an agnostic, not only because he could not
+harmonise the large amount of suffering in the world with the idea of
+a God as its first cause, but also because he "was aware that if we
+admit a first cause, the mind still craves to know whence it came and
+how it arose."[216] He saw, as Kant had seen before him and expressed
+in his _Kritik der Urtheilskraft_, that we cannot accept either of the
+only two possibilities which we are able to conceive: chance (or brute
+force) and design. Neither mechanism nor teleology can give an
+absolute answer to ultimate questions. The universe, and especially
+the organic life in it, can neither be explained as a mere
+combination of absolute elements nor as the effect of a constructing
+thought. Darwin concluded, as Kant, and before him Spinoza, that the
+oppositions and distinctions which our experience presents, cannot
+safely be regarded as valid for existence in itself. And, with Kant
+and Fichte, he found his stronghold in the conviction that man has
+something to do, even if he cannot solve all enigmas. "The safest
+conclusion seems to me that the whole subject is beyond the scope of
+man's intellect; but man can do his duty."[217]
+
+Is this the last word of human thought? Does not the possibility, that
+man can do his duty, suppose that the conditions of life allow of
+continuous ethical striving, so that there is a certain harmony
+between cosmic order and human ideals? Darwin himself has shown how
+the consciousness of duty can arise as a natural result of evolution.
+Moreover there are lines of evolution which have their end in ethical
+idealism, in a kingdom of values, which must struggle for life as all
+things in the world must do, but a kingdom which has its firm
+foundation in reality.
+
+FOOTNOTES:
+
+[Footnote 195: _Life and Letters of Charles Darwin_, Vol. I. p. 8.]
+
+[Footnote 196: _Encyclopädie der philosophischen Wissenschaften_ (4th
+edit.), Berlin, 1845, § 249.]
+
+[Footnote 197: _Lehrbuch der Naturphilosophie_, Jena, 1809.]
+
+[Footnote 198: _Ueber den Willen in der Natur_ (2nd edit.), Frankfurt
+a. M., 1854, pp. 41-43.]
+
+[Footnote 199: Spencer, _Autobiography_, Vol. II. p. 50, London and
+New York, 1904.]
+
+[Footnote 200: _Autobiography_, Vol. II. p. 100.]
+
+[Footnote 201: Cf. my letter to him 1876, now printed in Duncan's
+_Life and Letters of Herbert Spencer_, p. 178. London, 1908.]
+
+[Footnote 202: The present writer, many years ago, in his _Psychology_
+(Copenhagen, 1882; Eng. transl. London, 1891), criticised the
+evolutionistic treatment of the problem of knowledge from the Kantian
+point of view.]
+
+[Footnote 203: _Life and Letters_, Vol. II. p. 37.]
+
+[Footnote 204: _Ibid._ p. 232.]
+
+[Footnote 205: The new science of Ecology occupies an intermediate
+position between the biography of species and the biography of
+individuals. Compare _Congress of Arts and Science_, St. Louis, Vol.
+V. 1906 (The Reports of Drude and Robinson) and the work of my
+colleague, E. Warming.]
+
+[Footnote 206: Cf. my _History of Modern Philosophy_ (Eng. transl.
+London, 1900), I. pp. 76-79.]
+
+[Footnote 207: "Herrschaft und Knechtschaft," _Phönomenologie des
+Geistes_, IV. A., Leiden, 1907.]
+
+[Footnote 208: _The Descent of Man_, Vol. I. Ch. iii.]
+
+[Footnote 209: The works of Westermarck and Hobhouse throw new light
+on many of these features.]
+
+[Footnote 210: New York and London, 1893.]
+
+[Footnote 211: Paris, 1879.]
+
+[Footnote 212: _English literature and society in the eighteenth
+century_, London, 1904, p. 187.]
+
+[Footnote 213: Cf. my paper, "The law of relativity in Ethics,"
+_International Journal of Ethics_, Vol. I. 1891, pp. 37-62.]
+
+[Footnote 214: _Life and Letters_, Vol. I. p. 310.]
+
+[Footnote 215: _Ibid._ Vol. II. p. 177.]
+
+[Footnote 216: _Life and Letters_, Vol. 1. p. 306.]
+
+[Footnote 217: _Life and Letters_, p. 307.]
+
+
+
+
+VIII
+
+THE INFLUENCE OF DARWIN UPON RELIGIOUS THOUGHT
+
+BY P. N. WAGGETT, M.A., S.S.J.E.
+
+
+I
+
+The object of this paper is first to point out certain elements of the
+Darwinian influence upon Religious thought, and then to show reason
+for the conclusion that it has been, from a Christian point of view,
+satisfactory. I shall not proceed further to urge that the Christian
+apologetic in relation to biology has been successful. A variety of
+opinions may be held on this question, without disturbing the
+conclusion that the movements of readjustment have been beneficial to
+those who remain Christians, and this by making them more Christian
+and not only more liberal. The theologians may sometimes have
+retreated, but there has been an advance of theology. I know that this
+account incurs the charge of optimism. It is not the worst that could
+be made. The influence has been limited in personal range, unequal,
+even divergent, in operation, and accompanied by the appearance of
+waste and mischievous products. The estimate which follows requires
+for due balance a full development of many qualifying considerations.
+For this I lack space, but I must at least distinguish my view from
+the popular one that our difficulties about religion and natural
+science have come to an end.
+
+Concerning the older questions about origins--the origin of the
+world, of species, of man, of reason, conscience, religion--a large
+measure of understanding has been reached by some thoughtful men. But
+meanwhile new questions have arisen, questions about conduct,
+regarding both the reality of morals and the rule of right action for
+individuals and societies. And these problems, still far from
+solution, may also be traced to the influence of Darwin. For they
+arise from the renewed attention to heredity, brought about by the
+search for the causes of variation, without which the study of the
+selection of variations has no sufficient basis.
+
+Even the existing understanding about origins is very far from
+universal. On these points there were always thoughtful men who denied
+the necessity of conflict, and there are still thoughtful men who deny
+the possibility of a truce.
+
+It must further be remembered that the earlier discussion now, as I
+hope to show, producing favourable results, created also for a time
+grave damage, not only in the disturbance of faith and the loss of
+men--a loss not repaired by a change in the currents of debate--but in
+what I believe to be a still more serious respect. I mean the
+introduction of a habit of facile and untested hypothesis in religious
+as in other departments of thought.
+
+Darwin is not responsible for this, but he is in part the cause of it.
+Great ideas are dangerous guests in narrow minds; and thus it has
+happened that Darwin--the most patient of scientific workers, in whom
+hypothesis waited upon research, or if it provisionally outstepped it
+did so only with the most scrupulously careful acknowledgment--has led
+smaller and less conscientious men in natural science, in history, and
+in theology to an over-eager confidence in probable conjecture and a
+loose grip upon the facts of experience. It is not too much to say
+that in many quarters the age of materialism was the least
+matter-of-fact age conceivable, and the age of science the age which
+showed least of the patient temper of inquiry.
+
+I have indicated, as shortly as I could, some losses and dangers
+which in a balanced account of Darwin's influence would be discussed
+at length.
+
+One other loss must be mentioned. It is a defect in our thought which,
+in some quarters, has by itself almost cancelled all the advantages
+secured. I mean the exaggerated emphasis on uniformity or continuity;
+the unwillingness to rest any part of faith or of our practical
+expectation upon anything that from any point of view can be called
+exceptional. The high degree of success reached by naturalists in
+tracing, or reasonably conjecturing, the small beginnings of great
+differences, has led the inconsiderate to believe that anything may in
+time become anything else.
+
+It is true that this exaggeration of the belief in uniformity has
+produced in turn its own perilous reaction. From refusing to believe
+whatever can be called exceptional, some have come to believe whatever
+can be called wonderful.
+
+But, on the whole, the discontinuous or highly various character of
+experience received for many years too little deliberate attention.
+The conception of uniformity which is a necessity of scientific
+description has been taken for the substance of history. We have
+accepted a postulate of scientific method as if it were a conclusion
+of scientific demonstration. In the name of a generalisation which,
+however just on the lines of a particular method, is the prize of a
+difficult exploit of reflexion, we have discarded the direct
+impressions of experience; or, perhaps it is more true to say, we have
+used for the criticism of alleged experiences a doctrine of uniformity
+which is only valid in the region of abstract science. For every
+science depends for its advance upon limitation of attention, upon the
+selection out of the whole content of consciousness of that part or
+aspect which is measurable by the method of the science. Accordingly
+there is a science of life which rightly displays the unity underlying
+all its manifestations. But there is another view of life, equally
+valid, and practically sometimes more important, which recognises the
+immediate and lasting effect of crisis, difference, and revolution.
+Our ardour for the demonstration of uniformity of process and of
+minute continuous change needs to be balanced by a recognition of the
+catastrophic element in experience, and also by a recognition of the
+exceptional significance for us of events which may be perfectly
+regular from an impersonal point of view.
+
+An exorbitant jealousy of miracle, revelation, and ultimate moral
+distinctions has been imported from evolutionary science into
+religious thought. And it has been a damaging influence, because it
+has taken men's attention from facts, and fixed them upon theories.
+
+
+II
+
+With this acknowledgment of important drawbacks, requiring many words
+for their proper description, I proceed to indicate certain results of
+Darwin's doctrine which I believe to be in the long run wholly
+beneficial to Christian thought. These are:
+
+The encouragement in theology of that evolutionary method of
+observation and study, which has shaped all modern research:
+
+The recoil of Christian apologetics towards the ground of religious
+experience, a recoil produced by the pressure of scientific criticism
+upon other supports of faith:
+
+The restatement, or the recovery of ancient forms of statement, of the
+doctrines of Creation and of divine Design in Nature, consequent upon
+the discussion of evolution and of natural selection as its guiding
+factor.
+
+(1) The first of these is quite possibly the most important of all. It
+was well defined in a notable paper read by Dr. Gore, now Bishop of
+Birmingham, to the Church Congress at Shrewsbury in 1896. We have
+learnt a new caution both in ascribing and in denying significance to
+items of evidence, in utterance or in event. There has been, as in
+art, a study of values, which secures perspective and solidity in our
+representation of facts. On the one hand, a given utterance or event
+cannot be drawn into evidence as if all items were of equal
+consequence, like sovereigns in a bag. The question whence and whither
+must be asked, and the particular thing measured as part of a series.
+Thus measured it is not less truly important, but it may be important
+in a lower degree. On the other hand, and for exactly the same reason,
+nothing that is real is unimportant. The "failures" are not mere
+mistakes. We see them, in St. Augustine's words, as "scholar's faults
+which men praise in hope of fruit."
+
+We cannot safely trace the origin of the evolutionistic method to the
+influence of natural science. The view is tenable that theology led
+the way. Probably this is a case of alternate and reciprocal debt.
+Quite certainly the evolutionist method in theology, in Christian
+history and in the estimate of scripture, has received vast
+reinforcement from biology, in which evolution has been the ever
+present and ever victorious conception.
+
+(2) The second effect named is the new willingness of Christian
+thinkers to take definite account of religious experience. This is
+related to Darwin through the general pressure upon religious faith of
+scientific criticism. The great advance of our knowledge of organisms
+has been an important element in the general advance of science. It
+has acted, by the varied requirements of the theory of organisms, upon
+all other branches of natural inquiry, and it held for a long time
+that leading place in public attention which is now occupied by
+speculative physics. Consequently it contributed largely to our
+present estimation of science as the supreme judge in all matters of
+inquiry,[218] to the supposed destruction of mystery and the
+disparagement of metaphysics which marked the last age, as well as to
+the just recommendation of scientific method in branches of learning
+where the direct acquisitions of natural science had no place.
+
+Besides this, the new application of the idea of law and mechanical
+regularity to the organic world seemed to rob faith of a kind of
+refuge. The romantics had, as Berthelot[219] shows, appealed to life
+to redress the judgments drawn from mechanism. Now, in Spencer,
+evolution gave us a vitalist mechanic or mechanical vitalism, and the
+appeal seemed cut off. We may return to this point later when we
+consider evolution; at present I only endeavour to indicate that
+general pressure of scientific criticism which drove men of faith to
+seek the grounds of reassurance in a science of their own; in a method
+of experiment, of observation, of hypothesis checked by known facts.
+It is impossible for me to do more than glance across the threshold of
+this subject. But it is necessary to say that the method is in an
+elementary stage of revival. The imposing success that belongs to
+natural science is absent: we fall short of the unchallengeable
+unanimity of the Biologists on fundamentals. The experimental method
+with its sure repetitions cannot be applied to our subject-matter. But
+we have something like the observational method of palaeontology and
+geographical distribution; and in biology there are still men who
+think that the large examination of varieties by way of geography and
+the search of strata is as truly scientific, uses as genuinely the
+logical method of difference, and is as fruitful in sure conclusions
+as the quasi-chemical analysis of Mendelian laboratory work, of which
+last I desire to express my humble admiration. Religion also has its
+observational work in the larger and possibly more arduous manner.
+
+But the scientific work in religion makes its way through difficulties
+and dangers. We are far from having found the formula of its
+combination with the historical elements of our apologetic. It is
+exposed, therefore, to a damaging fire not only from unspiritualist
+psychology and pathology but also from the side of scholastic dogma.
+It is hard to admit on equal terms a partner to the old undivided rule
+of books and learning. With Charles Lamb, we cry in some distress,
+"must knowledge come to me, if it come at all, by some awkward
+experiment of intuition, and no longer by this familiar process of
+reading?"[220] and we are answered that the old process has an
+imperishable value, only we have not yet made clear its connection
+with other contributions. And all the work is young, liable to be
+drawn into unprofitable excursions, side-tracked by self-deceit and
+pretence; and it fatally attracts, like the older mysticism, the
+curiosity and the expository powers of those least in sympathy with
+it, ready writers who, with all the air of extended research, have
+been content with narrow grounds for induction. There is a danger,
+besides, which accompanies even the most genuine work of this science
+and must be provided against by all its serious students. I mean the
+danger of unbalanced introspection both for individuals and for
+societies; of a preoccupation comparable to our modern social
+preoccupation with bodily health; of reflexion upon mental states not
+accompanied by exercise and growth of the mental powers; the danger of
+contemplating will and neglecting work, of analysing conviction and
+not criticising evidence.
+
+Still, in spite of dangers and mistakes, the work remains full of
+hopeful indications, and, in the best examples,[221] it is truly
+scientific in its determination to know the very truth, to tell what
+we think, not what we think we ought to think,[222] truly scientific
+in its employment of hypothesis and verification, and in growing
+conviction of the reality of its subject-matter through the repeated
+victories of a mastery which advances, like science, in the Baconian
+road of obedience. It is reasonable to hope that progress in this
+respect will be more rapid and sure when religious study enlists more
+men affected by scientific desire and endowed with scientific
+capacity.
+
+The class of investigating minds is a small one, possibly even smaller
+than that of reflecting minds. Very few persons at any period are able
+to find out anything whatever. There are few observers, few
+discoverers, few who even wish to discover truth. In how many
+societies the problems of philology which face every person who speaks
+English are left unattempted! And if the inquiring or the successfully
+inquiring class of minds is small, much smaller, of course, is the
+class of those possessing the scientific aptitude in an eminent
+degree. During the last age this most distinguished class was to a
+very great extent absorbed in the study of phenomena, a study which
+had fallen into arrears. For we stood possessed, in rudiment, of means
+of observation, means for travelling and acquisition, qualifying men
+for a larger knowledge than had yet been attempted. These were now to
+be directed with new accuracy and ardour upon the fabric and behaviour
+of the world of sense. Our debt to the great masters in physical
+science who overtook and almost outstripped the task cannot be
+measured; and, under the honourable leadership of Ruskin, we may all
+well do penance if we have failed "in the respect due to their great
+powers of thought, or in the admiration due to the far scope of their
+discovery."[223] With what miraculous mental energy and divine good
+fortune--as Romans said of their soldiers--did our men of curiosity
+face the apparently impenetrable mysteries of nature! And how natural
+it was that immense accessions of knowledge, unrelated to the
+spiritual facts of life, should discredit Christian faith, by the
+apparent superiority of the new work to the feeble and unprogressive
+knowledge of Christian believers! The day is coming when men of this
+mental character and rank, of this curiosity, this energy and this
+good fortune in investigation, will be employed in opening mysteries
+of a spiritual nature. They will silence with masterful witness the
+over-confident denials of naturalism. They will be in danger of the
+widespread recognition which thirty years ago accompanied every
+utterance of Huxley, Tyndall, Spencer. They will contribute, in spite
+of adulation, to the advance of sober religious and moral science.
+
+And this result will be due to Darwin, first because by raising the
+dignity of natural science, he encouraged the development of the
+scientific mind; secondly because he gave to religious students the
+example of patient and ardent investigation; and thirdly because by
+the pressure of naturalistic criticism the religious have been driven
+to ascertain the causes of their own convictions, a work in which they
+were not without the sympathy of men of science.[224]
+
+In leaving the subject of scientific religious inquiry, I will only
+add that I do not believe it receives any important help--and
+certainly it suffers incidentally much damaging interruption--from the
+study of abnormal manifestations or abnormal conditions of
+personality.
+
+(3) Both of the above effects seem to me of high, perhaps the very
+highest, importance to faith and to thought. But, under the third
+head, I name two which are more directly traceable to the personal
+work of Darwin, and more definitely characteristic of the age in which
+his influence was paramount: viz. the influence of the two conceptions
+of evolution and natural selection upon the doctrine of creation and
+of design respectively.
+
+It is impossible here, though it is necessary for a complete sketch of
+the matter, to distinguish the different elements and channels of this
+Darwinian influence; in Darwin's own writings, in the vigourous
+polemic of Huxley, and strangely enough, but very actually for popular
+thought, in the teaching of the definitely anti-Darwinian evolutionist
+Spencer.
+
+Under the head of the directly and purely Darwinian elements I should
+class as preeminent the work of Wallace and of Bates; for no two sets
+of facts have done more to fix in ordinary intelligent minds a belief
+in organic evolution and in natural selection as its guiding factor
+than the facts of geographical distribution and of protective colour
+and mimicry. The facts of geology were difficult to grasp and the
+public and theologians heard more often of the imperfection than of
+the extent of the geological record. The witness of embryology,
+depending to a great extent upon microscopic work, was and is beyond
+the appreciation of persons occupied in fields of work other than
+biology.
+
+
+III
+
+From the influence in religion of scientific modes of thought we pass
+to the influence of particular biological conceptions. The former
+effect comes by way of analogy, example, encouragement and challenge;
+inspiring or provoking kindred or similar modes of thought in the
+field of theology; the latter by a collision of opinions upon matters
+of fact or conjecture which seem to concern both science and religion.
+
+In the case of Darwinism the story of this collision is familiar, and
+falls under the heads of evolution and natural selection, the doctrine
+of descent with modification, and the doctrine of its guidance or
+determination by the struggle for existence between related varieties.
+These doctrines, though associated and interdependent, and in popular
+thought not only combined but confused, must be considered separately.
+It is true that the ancient doctrine of Evolution, in spite of the
+ingenuity and ardour of Lamarck, remained a dream tantalising the
+intellectual ambition of naturalists, until the day when Darwin made
+it conceivable by suggesting the machinery of its guidance. And,
+further, the idea of natural selection has so effectively opened the
+door of research and stimulated observation in a score of principal
+directions that, even if the Darwinian explanation became one day much
+less convincing than, in spite of recent criticism, it now is, yet its
+passing, supposing it to pass, would leave the doctrine of Evolution
+immeasurably and permanently strengthened. For in the interests of the
+theory of selection, "Für Darwin," as Müller wrote, facts have been
+collected which remain in any case evidence of the reality of descent
+with modification.
+
+But still, though thus united in the modern history of convictions,
+though united and confused in the collision of biological and
+traditional opinion, yet evolution and natural selection must be
+separated in theological no less than in biological estimation.
+Evolution seemed inconsistent with Creation; natural selection with
+Providence and Divine design.
+
+Discussion was maintained about these points for many years and with
+much dark heat. It ranged over many particular topics and engaged
+minds different in tone, in quality, and in accomplishment. There was
+at most times a degree of misconception. Some naturalists attributed
+to theologians in general a poverty of thought which belonged really
+to men of a particular temper or training. The "timid theism"
+discerned in Darwin by so cautious a theologian as Liddon[225] was
+supposed by many biologists to be the necessary foundation of an
+honest Christianity. It was really more characteristic of devout
+_naturalists_ like Philip Henry Gosse, than of religious believers as
+such.[226] The study of theologians more considerable and even more
+typically conservative than Liddon does not confirm the description of
+religious intolerance given in good faith, but in serious ignorance,
+by a disputant so acute, so observant and so candid as Huxley.
+Something hid from each other's knowledge the devoted pilgrims in two
+great ways of thought. The truth may be, that naturalists took their
+view of what creation was from Christian men of science who naturally
+looked in their own special studies for the supports and illustrations
+of their religious belief. Of almost every labourious student it may
+be said: "_Hic ab arte sua non recessit_." And both the believing and
+the denying naturalists, confining habitual attention to a part of
+experience, are apt to affirm and deny with trenchant vigour and
+something of a narrow clearness "_Qui respiciunt ad pauca, de facili
+pronunciant_."[227]
+
+Newman says of some secular teachers that "they persuade the world of
+what is false by urging upon it what is true." Of some early opponents
+of Darwin it might be said by a candid friend that, in all sincerity
+of devotion to truth, they tried to persuade the world of what is true
+by urging upon it what is false. If naturalists took their version of
+orthodoxy from amateurs in theology, some conservative Christians,
+instead of learning what evolution meant to its regular exponents,
+took their view of it from celebrated persons, not of the front rank
+in theology or in thought, but eager to take account of public
+movements and able to arrest public attention.
+
+Cleverness and eloquence on both sides certainly had their share in
+producing the very great and general disturbance of men's minds in the
+early days of Darwinian teaching. But by far the greater part of that
+disturbance was due to the practical novelty and the profound
+importance of the teaching itself, and to the fact that the
+controversy about evolution quickly became much more public than any
+controversy of equal seriousness had been for many generations.
+
+We must not think lightly of that great disturbance because it has, in
+some real sense, done its work, and because it is impossible in days
+of more coolness and light, to recover a full sense of its very real
+difficulties.
+
+Those who would know them better should add to the calm records of
+Darwin[228] and to the story of Huxley's impassioned championship, all
+that they can learn of George Romanes.[229] For his life was absorbed
+in this very struggle and reproduced its stages. It began in a certain
+assured simplicity of biblical interpretation; it went on, through the
+glories and adventures of a paladin in Darwin's train, to the darkness
+and dismay of a man who saw all his most cherished beliefs rendered,
+as he thought, incredible.[230] He lived to find the freer faith for
+which process and purpose are not irreconcilable, but necessary to one
+another. His development, scientific, intellectual and moral, was
+itself of high significance; and its record is of unique value to our
+own generation, so near the age of that doubt and yet so far from it;
+certainly still much in need of the caution and courage by which past
+endurance prepares men for new emergencies. We have little enough
+reason to be sure that in the discussions awaiting us we shall do as
+well as our predecessors in theirs. Remembering their endurance of
+mental pain, their ardour in mental labour, the heroic temper and the
+high sincerity of controversialists on either side, we may well speak
+of our fathers in such words of modesty and self-judgment as Drayton
+used when he sang the victors of Agincourt. The progress of biblical
+study, in the departments of Introduction and Exegesis, resulting in
+the recovery of a point of view anciently tolerated if not prevalent,
+has altered some of the conditions of that discussion. In the years
+near 1858, the witness of Scripture was adduced both by Christian
+advocates and their critics as if unmistakably irreconcilable with
+Evolution.
+
+Huxley[231] found the path of the blameless naturalist everywhere
+blocked by "Moses": the believer in revelation was generally held to
+be forced to a choice between revealed cosmogony and the scientific
+account of origins. It is not clear how far the change in Biblical
+interpretation is due to natural science, and how far to the vital
+movements of theological study which have been quite independent of
+the controversy about species. It belongs to a general renewal of
+Christian movement, the recovery of a heritage. "Special
+Creation"--really a biological rather than a theological
+conception,--seems in its rigid form to have been a recent element
+even in English biblical orthodoxy.
+
+The Middle Ages had no suspicion that religious faith forbad inquiry
+into the natural origination of the different forms of life.
+Bartholomaeus Anglicus, an English Franciscan of the thirteenth
+century, was a mutationist in his way, as Aristotle, "the Philosopher"
+of the Christian Schoolmen, had been in his. So late as the
+seventeenth century, as we learn not only from early proceedings of
+the Royal Society, but from a writer so homely and so regularly pious
+as Walton, the variation of species and "spontaneous" generations had
+no theological bearing, except as instances of that various wonder of
+the world which in devout minds is food for devotion.
+
+It was in the eighteenth century that the harder statement took shape.
+Something in the preciseness of that age, its exaltation of law, its
+cold passion for a stable and measured universe, its cold denial, its
+cold affirmation of the power of God, a God of ice, is the occasion of
+that rigidity of religious thought about the living world which Darwin
+by accident challenged, or rather by one of those movements of genius
+which, Goethe[232] declares, are "elevated above all earthly control."
+
+If religious thought in the eighteenth century was aimed at a fixed
+and nearly finite world of spirit, it followed in all these respects
+the secular and critical lead. "La philosophie réformatrice du
+XVIII^{e} siècle[233] ramenait la nature et la société à des
+mécanismes que la pensée réfléchie peut concevoir et récomposer." In
+fact, religion in a mechanical age is condemned if it takes any but a
+mechanical tone. Butler's thought was too moving, too vital, too
+evolutionary, for the sceptics of his time. In a rationalist,
+encyclopaedic period, religion also must give hard outline to its
+facts, it must be able to display its secret to any sensible man in
+the language used by all sensible men. Milton's prophetic genius
+furnished the eighteenth century, out of the depth of the passionate
+age before it, with the theological tone it was to need. In spite of
+the austere magnificence of his devotion, he gives to smaller souls a
+dangerous lead. The rigidity of Scripture exegesis belonged to this
+stately but imperfectly sensitive mode of thought. It passed away with
+the influence of the older rationalists whose precise denials matched
+the precise and limited affirmations of the static orthodoxy.
+
+I shall, then, leave the specially biblical aspect of the
+debate--interesting as it is and even useful, as in Huxley's
+correspondence with the Duke of Argyll and others in 1892[234]--in
+order to consider without complication the permanent elements of
+Christian thought brought into question by the teaching of evolution.
+
+Such permanent elements are the doctrine of God as Creator of the
+universe, and the doctrine of man as spiritual and unique. Upon both
+the doctrine of evolution seemed to fall with crushing force.
+
+With regard to Man I leave out, acknowledging a grave omission, the
+doctrine of the Fall and of Sin. And I do so because these have not
+yet, as I believe, been adequately treated: here the fruitful reaction
+to the stimulus of evolution is yet to come. The doctrine of sin,
+indeed, falls principally within the scope of that discussion which
+has followed or displaced the Darwinian; and without it the Fall
+cannot be usefully considered. For the question about the Fall is a
+question not merely of origins, but of the interpretation of moral
+facts whose moral reality must first be established.
+
+I confine myself therefore to Creation and the dignity of man.
+
+The meaning of evolution, in the most general terms, is that the
+differentiation of forms is not essentially separate from their
+behaviour and use; that if these are within the scope of study, that
+is also; that the world has taken the form we see by movements not
+unlike those we now see in progress; that what may be called proximate
+origins are continuous in the way of force and matter, continuous in
+the way of life, with actual occurrences and actual characteristics.
+All this has no revolutionary bearing upon the question of ultimate
+origins. The whole is a statement about process. It says nothing to
+metaphysicians about cause. It simply brings within the scope of
+observation or conjecture that series of changes which has given their
+special characters to the different parts of the world we see. In
+particular, evolutionary science aspires to the discovery of the
+process or order of the appearance of life itself: if it were to
+achieve its aim it could say nothing of the cause of this or indeed of
+the most familiar occurrences. We should have become spectators or
+convinced historians of an event which, in respect of its cause and
+ultimate meaning, would be still impenetrable.
+
+With regard to the origin of species, supposing life already
+established, biological science has the well founded hopes and the
+measure of success with which we are all familiar. All this has, it
+would seem, little chance of collision with a consistent theism, a
+doctrine which has its own difficulties unconnected with any
+particular view of order or process. But when it was stated that
+species had arisen by processes through which new species were still
+being made, evolutionism came into collision with a statement,
+traditionally religious, that species were formed and fixed once for
+all and long ago.
+
+What is the theological import of such a statement when it is regarded
+as essential to belief in God? Simply that God's activity, with
+respect to the formation of living creatures, ceased at some point in
+past time.
+
+"God rested" is made the touchstone of orthodoxy. And when, under the
+pressure of the evidences, we found ourselves obliged to acknowledge
+and assert the present and persistent power of God, in the maintenance
+and in the continued formation of "types," what happened was the
+abolition of a time-limit. We were forced only to a bolder claim, to
+a theistic language less halting, more consistent, more thorough in
+its own line, as well as better qualified to assimilate and modify
+such schemes as Von Hartmann's philosophy of the Unconscious--a
+philosophy, by the way, quite intolerant of a merely mechanical
+evolution.[235]
+
+Here was not the retrenchment of an extravagant assertion, but the
+expansion of one which was faltering and inadequate. The traditional
+statement did not need paring down so as to pass the meshes of a new
+and exacting criticism. It was itself a net meant to surround and
+enclose experience; and we must increase its size and close its mesh
+to hold newly disclosed facts of life. The world, which had seemed a
+fixed picture or model, gained first perspective and then solidity and
+movement. We had a glimpse of organic _history_; and Christian thought
+became more living and more assured as it met the larger view of life.
+
+However unsatisfactory the new attitude might be to our critics, to
+Christians the reform was positive. What was discarded was a
+limitation, a negation. The movement was essentially conservative,
+even actually reconstructive. For the language disused was a language
+inconsistent with the definitions of orthodoxy; it set bounds to the
+infinite, and by implication withdrew from the creative rule all such
+processes as could be brought within the descriptions of research. It
+ascribed fixity and finality to that "creature" in which an apostle
+taught us to recognise the birth-struggles of an unexhausted progress.
+It tended to banish mystery from the world we see, and to confine it
+to a remote first age.
+
+In the reformed, the restored, language of religion, Creation became
+again not a link in a rational series to complete a circle of the
+sciences, but the mysterious and permanent relation between the
+infinite and the finite, between the moving changes we know in part,
+and the Power, after the fashion of that observation, unknown, which
+is itself "unmoved all motion's source."[236]
+
+With regard to man it is hardly necessary, even were it possible, to
+illustrate the application of this bolder faith. When the record of
+his high extraction fell under dispute, we were driven to a
+contemplation of the whole of his life, rather than of a part and that
+part out of sight. We remembered again, out of Aristotle, that the
+result of a process interprets its beginnings. We were obliged to read
+the title of such dignity as we may claim, in results and still more
+in aspirations.
+
+Some men still measure the value of great present facts in
+life--reason and virtue and sacrifice--by what a self-disparaged
+reason can collect of the meaner rudiments of these noble gifts. Mr.
+Balfour has admirably displayed the discrepancy, in this view, between
+the alleged origin and the alleged authority of reason. Such an
+argument ought to be used not to discredit the confident reason, but
+to illuminate and dignify its dark beginnings, and to show that at
+every step in the long course of growth a Power was at work which is
+not included in any term or in all the terms of the series.
+
+I submit that the more men know of actual Christian teaching, its
+fidelity to the past, and its sincerity in face of discovery, the more
+certainly they will judge that the stimulus of the doctrine of
+evolution has produced in the long run vigour as well as flexibility
+in the doctrine of Creation and of man.
+
+I pass from Evolution in general to Natural Selection.
+
+The character in religious language which I have for short called
+mechanical was not absent in the argument from design as stated before
+Darwin. It seemed to have reference to a world conceived as fixed. It
+pointed, not to the plastic capacity and energy of living matter, but
+to the fixed adaptation of this and that organ to an unchanging place
+or function.
+
+Mr. Hobhouse has given us the valuable phrase "a niche of organic
+opportunity." Such a phrase would have borne a different sense in
+non-evolutionary thought. In that thought, the opportunity was an
+opportunity for the Creative Power, and Design appeared in the
+preparation of the organism to fit the niche. The idea of the niche
+and its occupant growing together from simpler to more complex mutual
+adjustment was unwelcome to this teleology. If the adaptation was
+traced to the influence, through competition, of the environment, the
+old teleology lost an illustration and a proof. For the cogency of the
+proof in every instance depended upon the absence of explanation.
+Where the process of adaptation was discerned, the evidence of Purpose
+or Design was weak. It was strong only when the natural antecedents
+were not discovered, strongest when they could be declared
+undiscoverable.
+
+Paley's favourite word is "Contrivance"; and for him contrivance is
+most certain where production is most obscure. He points out the
+physiological advantage of the _valvulae conniventes_ to man, and the
+advantage for teleology of the fact that they cannot have been formed
+by "action and pressure." What is not due to pressure may be
+attributed to design, and when a "mechanical" process more subtle than
+pressure was suggested, the case for design was so far weakened. The
+cumulative proof from the multitude of instances began to disappear
+when, in selection, a natural sequence was suggested in which all the
+adaptations might be reached by the motive power of life, and
+especially when, as in Darwin's teaching, there was full recognition
+of the reactions of life to the stimulus of circumstance. "The
+organism fits the niche," said the teleologist, "because the Creator
+formed it so as to fit." "The organism fits the niche," said the
+naturalist, "because unless it fitted it could not exist." "It was
+fitted to survive," said the theologian. "It survives because it
+fits," said the selectionist. The two forms of statement are not
+incompatible; but the new statement, by provision of an ideally
+universal explanation of process, was hostile to a doctrine of purpose
+which relied upon evidences always exceptional however numerous.
+Science persistently presses on to find the universal machinery of
+adaptation in this planet; and whether this be found in selection, or
+in direct-effect, or in vital reactions resulting in large changes, or
+in a combination of these and other factors, it must always be opposed
+to the conception of a Divine Power here and there but not everywhere
+active.
+
+For science, the Divine must be constant, operative everywhere and in
+every quality and power, in environment and in organism, in stimulus
+and in reaction, in variation and in struggle, in hereditary
+equilibrium, and in "the unstable state of species"; equally present
+on both sides of every strain, in all pressures and in all
+resistances, in short in the general wonder of life and the world. And
+this is exactly what the Divine Power must be for religious faith.
+
+The point I wish once more to make is that the necessary readjustment
+of teleology, so as to make it depend upon the contemplation of the
+whole instead of a part, is advantageous quite as much to theology as
+to science. For the older view failed in courage. Here again our
+theism was not sufficiently theistic.
+
+Where results seemed inevitable, it dared not claim them as God-given.
+In the argument from Design it spoke not of God in the sense of
+theology, but of a Contriver, immensely, not infinitely wise and good,
+working within a world, the scene, rather than the ever dependent
+outcome, of His Wisdom; working in such emergencies and opportunities
+as occurred, by forces not altogether within His control, towards an
+end beyond Himself. It gave us, instead of the awful reverence due to
+the Cause of all substance and form, all love and wisdom, a
+dangerously detached appreciation of an ingenuity and benevolence
+meritorious in aim and often surprisingly successful in contrivance.
+
+The old teleology was more useful to science than to religion, and
+the design-naturalists ought to be gratefully remembered by
+Biologists. Their search for evidences led them to an eager study of
+adaptations and of minute forms, a study such as we have now an
+incentive to in the theory of Natural Selection. One hardly meets with
+the same ardour in microscopical research until we come to modern
+workers. But the argument from Design was never of great importance to
+faith. Still, to rid it of this character was worth all the stress and
+anxiety of the gallant old war. If Darwin had done nothing else for
+us, we are to-day deeply in his debt for this. The world is not less
+venerable to us now, not less eloquent of the causing mind, rather
+much more eloquent and sacred. But our wonder is not that "the
+underjaw of the swine works under the ground" or in any or all of
+those particular adaptations which Paley collected with so much skill,
+but that a purpose transcending, though resembling, our own purposes,
+is everywhere manifest; that what we live in is a whole, mutually
+sustaining, eventful and beautiful, where the "dead" forces feed the
+energies of life, and life sustains a stranger existence, able in some
+real measure to contemplate the whole, of which, mechanically
+considered, it is a minor product and a rare ingredient. Here, again,
+the change was altogether positive. It was not the escape of a vessel
+in a storm with loss of spars and rigging, not a shortening of sail to
+save the masts and make a port of refuge. It was rather the emergence
+from narrow channels to an open sea. We had propelled the great ship,
+finding purchase here and there for slow and uncertain movement. Now,
+in deep water, we spread large canvas to a favouring breeze.
+
+The scattered traces of design might be forgotten or obliterated. But
+the broad impression of Order became plainer when seen at due distance
+and in sufficient range of effect, and the evidence of love and wisdom
+in the universe could be trusted more securely for the loss of the
+particular calculation of their machinery.
+
+Many other topics of faith are affected by modern biology. In some of
+these we have learnt at present only a wise caution, a wise
+uncertainty. We stand before the newly unfolded spectacle of
+suffering, silenced; with faith not scientifically reassured but still
+holding fast certain other clues of conviction. In many important
+topics we are at a loss. But in others, and among them those I have
+mentioned, we have passed beyond this negative state and find faith
+positively strengthened and more fully expressed.
+
+We have gained also a language and a habit of thought more fit for the
+great and dark problems that remain, less liable to damaging
+conflicts, equipped for more rapid assimilation of knowledge. And by
+this change biology itself is a gainer. For, relieved of fruitless
+encounters with popular religion, it may advance with surer aim along
+the path of really scientific life-study which was reopened for modern
+men by the publication of _The Origin of Species_.
+
+Charles Darwin regretted that, in following science, he had not done
+"more direct good"[237] to his fellow-creatures. He has, in fact,
+rendered substantial service to interests bound up with the daily
+conduct and hopes of common men; for his work has led to improvements
+in the preaching of the Christian faith.
+
+FOOTNOTES:
+
+[Footnote 218: F. R. Tennant: "The Being of God in the light of
+Physical Science," in _Essays on some theological questions of the
+day_. London, 1905.]
+
+[Footnote 219: _Evolutionisme et Platonisme_, pp. 45, 46, 47. Paris,
+1908.]
+
+[Footnote 220: _Essays of Elia_, "New Year's Eve," p. 41; Ainger's
+edition. London, 1899.]
+
+[Footnote 221: Such an example is given in Baron F. von Hügel's
+recently finished book, the result of thirty years' research: _The
+Mystical Element of Religion, as studied in Saint Catherine of Genoa
+and her Friends_. London, 1908.]
+
+[Footnote 222: G. Tyrrell, in _Mediaevalism_, has a chapter which is
+full of the important _moral_ element in a scientific attitude. "The
+only infallible guardian of truth is the spirit of truthfulness."
+_Mediaevalism_, p. 182, London, 1908.]
+
+[Footnote 223: _Queen of the Air_, Preface, p. vii. London, 1906.]
+
+[Footnote 224: The scientific rank of its writer justifies the
+insertion of the following letter from the late Sir John
+Burdon-Sanderson to me. In the lecture referred to I had described the
+methods of Professor Moseley in teaching Biology as affording a
+suggestion of the scientific treatment of religion.
+
+OXFORD,
+
+_April 30, 1902_.
+
+DEAR SIR:
+
+     I feel that I must express to you my thanks for the
+     discourse which I had the pleasure of listening to yesterday
+     afternoon.
+
+     I do not mean to say that I was able to follow all that you
+     said as to the identity of Method in the two fields of
+     Science and Religion, but I recognise that the "mysticism"
+     of which you spoke gives us the only way by which the two
+     fields can be brought into relation.
+
+     Among much that was memorable, nothing interested me more
+     than what you said of Moseley.
+
+     No one, I am sure, knew better than you the value of his
+     teaching and in what that value consisted.
+
+Yours faithfully,
+
+J. BURDON-SANDERSON.
+
+]
+
+[Footnote 225: H. P. Liddon, _The Recovery of S. Thomas_; a sermon
+preached in St. Paul's, London, on April 23rd, 1882 (the Sunday after
+Darwin's death).]
+
+[Footnote 226: Dr. Pusey (_Unscience not Science adverse to Faith_,
+1878) writes: "The questions as to 'species,' of what variations the
+animal world is capable, whether the species be more or fewer, whether
+accidental variations may become hereditary ... and the like,
+naturally fall under the province of science. In all these questions
+Mr. Darwin's careful observations gained for him a deserved
+approbation and confidence."]
+
+[Footnote 227: Aristotle, in Bacon, quoted by Newman in his _Idea of a
+University_, p. 78. London, 1873.]
+
+[Footnote 228: _Life and Letters_ and _More Letters of Charles
+Darwin._]
+
+[Footnote 229: _Life and Letters_, London, 1896. _Thoughts on
+Religion_, London, 1895. _Candid Examination of Theism_, London,
+1878.]
+
+[Footnote 230: "Never in the history of man has so terrific a calamity
+befallen the race as that which all who look may now (viz. in
+consequence of the scientific victory of Darwin) behold advancing as a
+deluge black with destruction, resistless in might, uprooting our most
+cherished hopes, engulphing our most precious creed, and burying our
+highest life in mindless destruction."--_A Candid Examination of
+Theism_, p. 51.]
+
+[Footnote 231: _Science and Christian Tradition._ London, 1904.]
+
+[Footnote 232: "No productiveness of the highest kind ... is in the
+power of anyone."--_Conversations of Goethe with Eckermann and Soret_.
+London, 1850.]
+
+[Footnote 233: Berthelot, _Evolutionisme et Platonisme_, Paris, 1908,
+p. 45.]
+
+[Footnote 234: _Times_, 1892, _passim._]
+
+[Footnote 235: See Von Hartmann's _Wahrheit und Irrthum in
+Darwinismus_. Berlin, 1875.]
+
+[Footnote 236: Hymn of the Church--
+
+    Rerum Deus tenax vigor,
+    Immotus in te permanens.
+
+]
+
+[Footnote 237: _Life and Letters_, Vol. III. p. 359.]
+
+
+
+
+IX
+
+DARWINISM AND HISTORY
+
+BY J. B. BURY, LITT.D., LL.D.
+
+_Regius Professor of Modern History in the University of Cambridge_
+
+
+1. Evolution, and the principles associated with the Darwinian theory,
+could not fail to exert a considerable influence on the studies
+connected with the history of civilised man. The speculations which
+are known as "philosophy of history," as well as the sciences of
+anthropology, ethnography, and sociology (sciences which though they
+stand on their own feet are for the historian auxiliary), have been
+deeply affected by these principles. Historiographers, indeed, have
+with few exceptions made little attempt to apply them; but the growth
+of historical study in the nineteenth century has been determined and
+characterised by the same general principle which has underlain the
+simultaneous developments of the study of nature, namely the _genetic
+idea_. The "historical" conception of nature, which has produced the
+history of the solar system, the story of the earth, the genealogies
+of telluric organisms, and has revolutionised natural science, belongs
+to the same order of thought as the conception of human history as a
+continuous, genetic, causal process--a conception which has
+revolutionised historical research and made it scientific. Before
+proceeding to consider the application of evolutional principles, it
+will be pertinent to notice the rise of this new view.
+
+2. With the Greeks and Romans history had been either a descriptive
+record or had been written in practical interests. The most eminent
+of the ancient historians were pragmatical; that is, they regarded
+history as an instructress in statesmanship, or in the art of war, or
+in morals. Their records reached back such a short way, their
+experience was so brief, that they never attained to the conception of
+continuous process, or realised the significance of time; and they
+never viewed the history of human societies as a phenomenon to be
+investigated for its own sake. In the middle ages there was still less
+chance of the emergence of the ideas of progress and development. Such
+notions were excluded by the fundamental doctrines of the dominant
+religion which bounded and bound men's minds. As the course of history
+was held to be determined from hour to hour by the arbitrary will of
+an extra cosmic person, there could be no self-contained causal
+development, only a dispensation imposed from without. And as it was
+believed that the world was within no great distance from the end of
+this dispensation, there was no motive to take much interest in
+understanding the temporal, which was to be only temporary.
+
+The intellectual movements of the fifteenth and sixteenth centuries
+prepared the way for a new conception, but it did not emerge
+immediately. The historians of the Renaissance period simply reverted
+to the ancient pragmatical view. For Machiavelli, exactly as for
+Thucydides and Polybius, the use of studying history was instruction
+in the art of politics. The Renaissance itself was the appearance of a
+new culture, different from anything that had gone before; but at the
+time men were not conscious of this; they saw clearly that the
+traditions of classical antiquity had been lost for a long period, and
+they were seeking to revive them, but otherwise they did not perceive
+that the world had moved, and that their own spirit, culture, and
+conditions were entirely unlike those of the thirteenth century. It
+was hardly till the seventeenth century that the presence of a new
+age, as different from the middle ages as from the ages of Greece and
+Rome, was fully realised. It was then that the triple division of
+ancient, medieval, and modern was first applied to the history of
+western civilisation. Whatever objections may be urged against this
+division, which has now become almost a category of thought, it marks
+a most significant advance in man's view of his own past. He has
+become conscious of the immense changes in civilisation which have
+come about slowly in the course of time, and history confronts him
+with a new aspect. He has to explain how those changes have been
+produced, how the transformations were effected. The appearance of
+this problem was almost simultaneous with the rise of rationalism, and
+the great historians and thinkers of the eighteenth century, such as
+Montesquieu, Voltaire, Gibbon, attempted to explain the movement of
+civilisation by purely natural causes. These brilliant writers
+prepared the way for the genetic history of the following century. But
+in the spirit of the _Aufklärung_, that eighteenth-century
+Enlightenment to which they belonged, they were concerned to judge all
+phenomena before the tribunal of reason; and the apotheosis of
+"reason" tended to foster a certain superior _a priori_ attitude,
+which was not favourable to objective treatment and was incompatible
+with a "historical sense." Moreover the traditions of pragmatical
+historiography had by no means disappeared.
+
+3. In the first quarter of the nineteenth century the meaning of
+genetic history was fully realised. "Genetic" perhaps is as good a
+word as can be found for the conception which in this century was
+applied to so many branches of knowledge in the spheres both of nature
+and of mind. It does not commit us to the doctrine proper of
+evolution, nor yet to any teleological hypothesis such as is implied
+in "progress." For history it meant that the present condition of the
+human race is simply and strictly the result of a causal series (or
+set of causal series)--a continuous succession of changes, where each
+state arises causally out of the preceding; and that the business of
+historians is to trace this genetic process, to explain each change,
+and ultimately to grasp' the complete development of the life of
+humanity. Three influential writers, who appeared at this stage and
+helped to initiate a new period of research, may specially be
+mentioned. Ranke in 1824 definitely repudiated the pragmatical view
+which ascribes to history the duties of an instructress, and with no
+less decision renounced the function, assumed by the historians of the
+_Aufklärung_, to judge the past; it was his business, he said, merely
+to show how things really happened. Niebuhr was already working in the
+same spirit and did more than any other writer to establish the
+principle that historical transactions must be related to the ideas
+and conditions of their age. Savigny about the same time founded the
+"historical school" of law. He sought to show that law was not the
+creation of an enlightened will, but grew out of custom and was
+developed by a series of adaptations and rejections, thus applying the
+conception of evolution. He helped to diffuse the notion that all the
+institutions of a society or a nation are as closely interconnected as
+the parts of a living organism.
+
+4. The conception of the history of man as a causal development meant
+the elevation of historical inquiry to the dignity of a science. Just
+as the study of bees cannot become scientific so long as the student's
+interest in them is only to procure honey or to derive moral lessons
+from the labours of "the little busy bee," so the history of human
+societies cannot become the object of pure scientific investigation so
+long as man estimates its value in pragmatical scales. Nor can it
+become a science until it is conceived as lying entirely within a
+sphere in which the law of cause and effect has unreserved and
+unrestricted dominion. On the other hand, once history is envisaged as
+a causal process, which contains within itself the explanation of the
+development of man from his primitive state to the point which he has
+reached, such a process necessarily becomes the object of scientific
+investigation and the interest in it is scientific curiosity.
+
+At the same time, the instruments were sharpened and refined. Here
+Wolf, a philologist with historical instinct, was a pioneer. His
+_Prolegomena_ to Homer (1795) announced new modes of attack.
+Historical investigation was soon transformed by the elaboration of
+new methods.
+
+5. "Progress" involves a judgment of value, which is not involved in
+the conception of history as a genetic process. It is also an idea
+distinct from that of evolution. Nevertheless it is closely related to
+the ideas which revolutionised history at the beginning of the last
+century; it swam into men's ken simultaneously; and it helped
+effectively to establish the notion of history as a continuous process
+and to emphasise the significance of time. Passing over earlier
+anticipations, I may point to a _Discours_ of Turgot (1750), where
+history is presented as a process in which "the total mass of the
+human race" "marches continually though sometimes slowly to an ever
+increasing perfection." That is a clear statement of the conception
+which Turgot's friend Condorcet elaborated in the famous work,
+published in 1795, _Esquisse d'un tableau historique des progrès de
+l'esprit humain_. This work first treated with explicit fulness the
+idea to which a leading role was to fall in the ideology of the
+nineteenth century. Condorcet's book reflects the triumphs of the
+_Tiers état_, whose growing importance had also inspired Turgot; it
+was the political changes in the eighteenth century which led to the
+doctrine, emphatically formulated by Condorcet, that the masses are
+the most important element in the historical process. I dwell on this
+because, though Condorcet had no idea of evolution, the predominant
+importance of the masses was the assumption which made it possible to
+apply evolutional principles to history. And it enabled Condorcet
+himself to maintain that the history of civilisation, a progress still
+far from being complete, was a development conditioned by general
+laws.
+
+6. The assimilation of society to an organism, which was a governing
+notion in the school of Savigny, and the conception of progress,
+combined to produce the idea of an organic development, in which the
+historian has to determine the central principle or leading character.
+This is illustrated by the apotheosis of democracy in Tocqueville's
+_Démocratie en Amérique_, where the theory is maintained that "the
+gradual and progressive development of equality is at once the past
+and the future of the history of men." The same two principles are
+combined in the doctrine of Spencer (who held that society is an
+organism, though he also contemplated its being what he calls a
+"super-organic aggregate"),[238] that social evolution is a
+progressive change from militarism to industrialism.
+
+7. The idea of development assumed another form in the speculations of
+German idealism. Hegel conceived the successive periods of history as
+corresponding to the ascending phases or ideas in the self-evolution
+of his Absolute Being. His _Lectures on the Philosophy of History_
+were published in 1837 after his death. His philosophy had a
+considerable effect, direct and indirect, on the treatment of history
+by historians, and although he was superficial and unscientific
+himself in dealing with historical phenomena, he contributed much
+towards making the idea of historical development familiar. Ranke was
+influenced, if not by Hegel himself, at least by the Idealistic
+philosophies of which Hegel's was the greatest. He was inclined to
+conceive the stages in the process of history as marked by
+incarnations, as it were, of ideas, and sometimes speaks as if the
+ideas were independent forces, with hands and feet. But while Hegel
+determined his ideas by _a priori_ logic, Ranke obtained his by
+induction--by a strict investigation of the phenomena; so that he was
+scientific in his method and work, and was influenced by Hegelian
+prepossessions only in the kind of significance which he was disposed
+to ascribe to his results. It is to be noted that the theory of Hegel
+implied a judgment of value; the movement was a progress towards
+perfection.
+
+8. In France, Comte approached the subject from a different side, and
+exercised, outside Germany, a far wider influence than Hegel. The 4th
+volume of his _Cours de philosophie positive_, which appeared in 1839,
+created sociology and treated history as a part of this new science,
+namely as "social dynamics." Comte sought the key for unfolding
+historical development, in what he called the social-psychological
+point of view, and he worked out the two ideas which had been
+enunciated by Condorcet: that the historian's attention should be
+directed not, as hitherto, principally to eminent individuals, but to
+the collective behaviour of the masses, as being the most important
+element in the process; and that, as in nature, so in history, there
+are general laws, necessary and constant, which condition the
+development. The two points are intimately connected, for it is only
+when the masses are moved into the foreground that regularity,
+uniformity, and law can be conceived as applicable. To determine the
+social-psychological laws which have controlled the development is,
+according to Comte, the task of sociologists and historians.
+
+9. The hypothesis of general laws operative in history was carried
+further in a book which appeared in England twenty years later and
+exercised an influence in Europe far beyond its intrinsic merit,
+Buckle's _History of Civilisation in England_ (1857-61). Buckle owed
+much to Comte, and followed him, or rather outdid him, in regarding
+intellect as the most important factor conditioning the upward
+development of man, so that progress, according to him, consisted in
+the victory of the intellectual over the moral laws.
+
+10. The tendency of Comte and Buckle to assimilate history to the
+sciences of nature by reducing it to general "laws," derived stimulus
+and plausibility from the vista offered by the study of statistics,
+in which the Belgian Quetelet, whose book _Sur l'homme_ appeared in
+1835, discerned endless possibilities. The astonishing uniformities
+which statistical inquiry disclosed led to the belief that it was only
+a question of collecting a sufficient amount of statistical material,
+to enable us to predict how a given social group will act in a
+particular case. Bourdeau, a disciple of this school, looks forward to
+the time when historical science will become entirely quantitative.
+The actions of prominent individuals, which are generally considered
+to have altered or determined the course of things, are obviously not
+amenable to statistical computation or explicable by general laws.
+Thinkers like Buckle sought to minimise their importance or explain
+them away.
+
+11. These indications may suffice to show that the new efforts to
+interpret history which marked the first half of the nineteenth
+century were governed by conceptions closely related to those which
+were current in the field of natural science and which resulted in the
+doctrine of evolution. The genetic principle, progressive development,
+general laws, the significance of time, the conception of society as
+an organic aggregate, the metaphysical theory of history as the
+self-evolution of spirit,--all these ideas show that historical
+inquiry had been advancing independently on somewhat parallel lines to
+the sciences of nature. It was necessary to bring this out in order to
+appreciate the influence of Darwinism.
+
+12. In the course of the dozen years which elapsed between the
+appearances of _The Origin of Species_ (observe that the first volume
+of Buckle's work was published just two years before) and of _The
+Descent of Man_ (1871), the hypothesis of Lamarck that man is the
+co-descendant with other species of some lower extinct form was
+admitted to have been raised to the rank of an established fact by
+most thinkers whose brains were not working under the constraint of
+theological authority.
+
+One important effect of the discovery of this fact (I am not speaking
+now of the Darwinian explanation) was to assign to history a definite
+place in the coordinated whole of knowledge, and relate it more
+closely to other sciences. It had indeed a defined logical place in
+systems such as Hegel's and Comte's; but Darwinism certified its
+standing convincingly and without more ado. The prevailing doctrine
+that man was created _ex abrupto_ had placed history in an isolated
+position, disconnected with the sciences of nature. Anthropology,
+which deals with the animal _anthropos_, now comes into line with
+zoology, and brings it into relation with history.[239] Man's
+condition at the present day is the result of a series of
+transformations, going back to the most primitive phase of society,
+which is the ideal (unattainable) beginning of history. But that
+beginning had emerged without any breach of continuity from a
+development which carries us back to a quadrimane ancestor, still
+further back (according to Darwin's conjecture) to a marine animal of
+the ascidian type, and then through remoter periods to the lowest form
+of organism. It is essential in this theory that though links have
+been lost there was no break in the gradual development; and this
+conception of a continuous progress in the evolution of life,
+resulting in the appearance of uncivilised Anthropos, helped to
+reinforce, and increase a belief in, the conception of the history of
+civilised Anthropos as itself also a continuous progressive
+development.
+
+13. Thus the diffusion of the Darwinian theory of the origin of man,
+by emphasising the idea of continuity and breaking down the barriers
+between the human and animal kingdoms, has had an important effect in
+establishing the position of history among the sciences which deal
+with telluric development. The perspective of history is merged in a
+larger perspective of development. As one of the objects of biology is
+to find the exact steps in the genealogy of man from the lowest
+organic form, so the scope of history is to determine the stages in
+the unique causal series from the most rudimentary to the present
+state of human civilisation.
+
+It is to be observed that the interest in historical research implied
+by this conception need not be that of Comte. In the Positive
+Philosophy history is part of sociology; the interest in it is to
+discover the sociological laws. In the view of which I have just
+spoken, history is permitted to be an end in itself; the
+reconstruction of the genetic process is an independent interest. For
+the purpose of the reconstruction, sociology, as well as physical
+geography, biology, psychology, is necessary; the sociologist and the
+historian play into each other's hands; but the object of the former
+is to establish generalisations; the aim of the latter is to trace in
+detail a singular causal sequence.
+
+14. The success of the evolutional theory helped to discredit the
+assumption or at least the invocation of transcendent causes.
+Philosophically of course it is compatible with theism, but historians
+have for the most part desisted from invoking the naive conception of
+a "god in history" to explain historical movements. A historian may be
+a theist; but, so far as his work is concerned, this particular belief
+is otiose. Otherwise indeed (as was remarked above) history could not
+be a science; for with a _deus ex machina_ who can be brought on the
+stage to solve difficulties scientific treatment is a farce. The
+transcendent element had appeared in a more subtle form through the
+influence of German philosophy. I noticed how Ranke is prone to refer
+to ideas as if they were transcendent existences manifesting
+themselves in the successive movements of history. It is intelligible
+to speak of certain ideas as controlling, in a given period,--for
+instance, the idea of nationality; but from the scientific point of
+view, such ideas have no existence outside the minds of individuals
+and are purely psychical forces; and a historical "idea," if it does
+not exist in this form, is merely a way of expressing a synthesis of
+the historian himself.
+
+15. From the more general influence of Darwinism on the place of
+history in the system of human knowledge, we may turn to the influence
+of the principles and methods by which Darwin explained development.
+It had been recognised even by ancient writers (such as Aristotle and
+Polybius) that physical circumstances (geography, climate) were
+factors conditioning the character and history of a race or society.
+In the sixteenth century Bodin emphasised these factors, and many
+subsequent writers took them into account. The investigations of
+Darwin, which brought them into the foreground, naturally promoted
+attempts to discover in them the chief key to the growth of
+civilisation. Comte had expressly denounced the notion that the
+biological methods of Lamarck could be applied to social man. Buckle
+had taken account of natural influences, but had relegated them to a
+secondary plane, compared with psychological factors. But the
+Darwinian theory made it tempting to explain the development of
+civilisation in terms of "adaptation to environment," "struggle for
+existence," "natural selection," "survival of the fittest," etc.[240]
+
+The operation of these principles cannot be denied. Man is still an
+animal, subject to zoological as well as mechanical laws. The dark
+influence of heredity continues to be effective; and psychical
+development had begun in lower organic forms,--perhaps with life
+itself. The organic and the social struggles for existence are
+manifestations of the same principle. Environment and climatic
+influence must be called in to explain not only the differentiation of
+the great racial sections of humanity, but also the varieties within
+these sub-species and, it may be, the assimilation of distinct
+varieties. Ritter's _Anthropogeography_ has opened a useful line of
+research. But on the other hand, it is urged that, in explaining the
+course of history, these principles do not take us very far, and that
+it is chiefly for the primitive ultra-prehistoric period that they can
+account for human development. It may be said that, so far as concerns
+the actions and movements of men which are the subject of recorded
+history, physical environment has ceased to act mechanically, and in
+order to affect their actions must affect their wills first; and that
+this psychical character of the causal relations substantially alters
+the problem. The development of human societies, it may be argued,
+derives a completely new character from the dominance of the conscious
+psychical element, creating as it does new conditions (inventions,
+social institutions, etc.) which limit and counteract the operation of
+natural selection, and control and modify the influence of physical
+environment. Most thinkers agree now that the chief clews to the
+growth of civilisation must be sought in the psychological sphere.
+Imitation, for instance, is a principle which is probably more
+significant for the explanation of human development than natural
+selection. Darwin himself was conscious that his principles had only a
+very restricted application in this sphere, as is evident from his
+cautious and tentative remarks in the 5th chapter of his _Descent of
+Man_. He applied natural selection to the growth of the intellectual
+faculties and of the fundamental social instincts, and also to the
+differentiation of the great races or "sub-species" (Caucasian,
+African, etc.) which differ in anthropological character.[241]
+
+16. But if it is admitted that the governing factors which concern the
+student of social development are of the psychical order, the
+preliminary success of natural science in explaining organic evolution
+by general principles encouraged sociologists to hope that social
+evolution could be explained on general principles also. The idea of
+Condorcet, Buckle, and others, that history could be assimilated to
+the natural sciences was powerfully reinforced, and the notion that
+the actual historical process, and every social movement involved in
+it, can be accounted for by sociological generalisations, so-called
+"laws," is still entertained by many, in one form or another.
+Dissentients from this view do not deny that the generalisations at
+which the sociologist arrives by the comparative method, by the
+analysis of social factors, and by psychological deduction may be an
+aid to the historian; but they deny that such uniformities are laws or
+contain an explanation of the phenomena. They can point to the element
+of chance coincidence. This element must have played a part in the
+events of organic evolution, but it has probably in a larger measure
+helped to determine events in social evolution. The collision of two
+unconnected sequences may be fraught with great results. The sudden
+death of a leader or a marriage without issue, to take simple cases,
+has again and again led to permanent political consequences. More
+emphasis is laid on the decisive actions of individuals, which cannot
+be reduced under generalisations and which deflect the course of
+events. If the significance of the individual will had been
+exaggerated to the neglect of the collective activity of the social
+aggregate before Condorcet, his doctrine tended to eliminate as
+unimportant the roles of prominent men, and by means of this
+elimination it was possible to found sociology. But it may be urged
+that it is patent on the face of history that its course has
+constantly been shaped and modified by the wills of individuals,[242]
+which are by no means always the expression of the collective will;
+and that the appearance of such personalities at the given moments is
+not a necessary outcome of the conditions and cannot be deduced. Nor
+is there any proof that, if such and such an individual had not been
+born, some one else would have arisen to do what he did. In some cases
+there is no reason to think that what happened need ever have come to
+pass. In other cases, it seems evident that the actual change was
+inevitable, but in default of the man who initiated and guided it, it
+might have been postponed, and, postponed or not, might have borne a
+different cachet. I may illustrate by an instance which has just come
+under my notice. Modern painting was founded by Giotto, and the
+Italian expedition of Charles VIII, near the close of the sixteenth
+century, introduced into France the fashion of imitating Italian
+painters. But for Giotto and Charles VIII, French painting might have
+been very different. It may be said that "if Giotto had not appeared,
+some other great imitator would have played a role analogous to his,
+and that without Charles VIII there would have been the commerce with
+Italy, which in the long run would have sufficed to place France in
+relation with Italian artists. But the equivalent of Giotto might have
+been deferred for a century and probably would have been different;
+and commercial relations would have required ages to produce the
+_rayonnement imitatif_ of Italian art in France, which the expedition
+of the royal adventurer provoked in a few years."[243] Instances
+furnished by political history are simply endless. Can we conjecture
+how events would have moved if the son of Philip of Macedon had been
+an incompetent? The aggressive action of Prussia which astonished
+Europe in 1740 determined the subsequent history of Germany; but that
+action was anything but inevitable; it depended entirely on the
+personality of Frederick the Great.
+
+Hence it may be argued that the action of individual wills is a
+determining and disturbing factor, too significant and effective to
+allow history to be grasped by sociological formulae. The types and
+general forms of development which the sociologist attempts to
+disengage can only assist the historian in understanding the actual
+course of events. It is in the special domains of economic history and
+_Culturgeschichte_ which have come to the front in modern times that
+generalisation is most fruitful, but even in these it may be contended
+that it furnishes only partial explanations.
+
+17. The truth is that Darwinism itself offers the best illustration of
+the insufficiency of general laws to account for historical
+development. The part played by coincidence, and the part played by
+individuals--limited by, and related to, general social
+conditions--render it impossible to deduce the course of the past
+history of man or to predict the future. But it is just the same with
+organic development. Darwin (or any other zoologist) could not deduce
+the actual course of evolution from general principles. Given an
+organism and its environment, he could not show that it must evolve
+into a more complex organism of a definite predetermined type; knowing
+what it has evolved into, he could attempt to discover and assign the
+determining causes. General principles do not account for a particular
+sequence; they embody necessary conditions; but there is a chapter of
+accidents too. It is the same in the case of history.
+
+18. Among the evolutional attempts to subsume the course of history under
+general syntheses, perhaps the most important is that of Lamprecht, whose
+"kulturhistorische" attempt to discover and assign the determining causes.
+German history, exhibits the (indirect) influence of the Comtist school. It
+is based upon psychology, which, in his views, holds among the sciences of
+mind (_Geisteswissenschaften_) the same place (that of a
+_Grundwissenschaft_) which mechanics holds among the sciences of nature.
+History, by the same comparison, corresponds to biology, and, according to
+him, it can only become scientific if it is reduced to general concepts
+(_Begriffe_). Historical movements and events are of a psychical character,
+and Lamprecht conceives a given phase of civilisation as "a collective
+psychical condition (_seelischer Gesamtzustand_)" controlling the period,
+"a diapason which penetrates all psychical phenomena and thereby all
+historical events of the time."[244] He has worked out a series of such
+phases, "ages of changing psychical diapason," in his _Deutsche
+Geschichte_, with the aim of showing that all the feelings and actions of
+each age can be explained by the diapason; and has attempted to prove that
+these diapasons are exhibited in other social developments, and are
+consequently not singular but typical. He maintains further that these ages
+succeed each other in a definite order; the principle being that the
+collective psychical development begins with the homogeneity of all the
+individual members of a society and, through heightened psychical activity,
+advances in the form of a continually increasing differentiation of the
+individuals (this is akin to the Spencerian formula). This process,
+evolving psychical freedom from psychical constraint, exhibits a series of
+psychical phenomena which define successive periods of civilisation. The
+process depends on two simple principles, that no idea can disappear
+without leaving behind it an effect or influence, and that all psychical
+life, whether in a person or a society, means change, the acquisition of
+new mental contents. It follows that the new have to come to terms with the
+old, and this leads to a synthesis which determines the character of a new
+age. Hence the ages of civilisation are defined as the "highest concepts
+for subsuming without exception all psychical phenomena of the development
+of human societies, that is, of all historical events."[245] Lamprecht
+deduces the idea of a special historical science, which might be called
+"historical ethnology," dealing with the ages of civilisation, and bearing
+the same relation to (descriptive or narrative) history as ethnology to
+ethnography. Such a science obviously corresponds to Comte's social
+dynamics, and the comparative method, on which Comte laid so much emphasis,
+is the principal instrument of Lamprecht.
+
+19. I have dwelt on the fundamental ideas of Lamprecht, because they
+are not yet widely known in England, and because his system is the
+ablest product of the sociological school of historians. It carries
+the more weight as its author himself is a historical specialist, and
+his historical syntheses deserve the most careful consideration. But
+there is much in the process of development which on such assumptions
+is not explained, especially the initiative of individuals. Historical
+development does not proceed in a right line, without the choice of
+diverging. Again and again, several roads are open to it, of which it
+chooses one--why? On Lamprecht's method, we may be able to assign the
+conditions which limit the psychical activity of men at a particular
+stage of evolution, but within those limits the individual has so many
+options, such a wide room for moving, that the definition of those
+conditions, the "psychical diapasons," is only part of the explanation
+of the particular development. The heel of Achilles in all historical
+speculations of this class has been the role of the individual.
+
+The increasing prominence of economic history has tended to encourage
+the view that history can be explained in terms of general concepts or
+types. Marx and his school based their theory of human development on
+the conditions of production, by which, according to them, all social
+movements and historical changes are entirely controlled. The leading
+part which economic factors play in Lamprecht's system is significant,
+illustrating the fact that economic changes admit most readily this
+kind of treatment, because they have been less subject to direction or
+interference by individual pioneers.
+
+Perhaps it may be thought that the conception of _social environment_
+(essentially psychical), on which Lamprecht's "psychical diapasons"
+depend, is the most valuable and fertile conception that the historian
+owes to the suggestion of the science of biology--the conception of
+all particular historical actions and movements as (1) related to and
+conditioned by the social environment, and (2) gradually bringing
+about a transformation of that environment. But no given
+transformation can be proved to be necessary (predetermined). And
+types of development do not represent laws; their meaning and value
+lie in the help they may give to the historian, in investigating a
+certain period of civilisation, to enable him to discover the
+inter-relations among the diverse features which it presents. They
+are, as some one has said, an instrument of heuretic method.
+
+20. The man engaged in special historical researches--which have been
+pursued unremittingly for a century past, according to scientific
+methods of investigating evidence (initiated by Wolf, Niebuhr,
+Ranke)--have for the most part worked on the assumptions of genetic
+history or at least followed in the footsteps of those who fully
+grasped the genetic point of view. But their aim has been to collect
+and sift evidence, and determine particular facts; comparatively few
+have given serious thought to the lines of research and the
+speculations which have been considered in this paper. They have been
+reasonably shy of compromising their work by applying theories which
+are still much debated and immature. But historiography cannot
+permanently evade the questions raised by these theories. One may
+venture to say that no historical change or transformation will be
+fully understood until it is explained how social environment acted on
+the individual components of the society (both immediately and by
+heredity), and how the individuals reacted upon their environment. The
+problem is psychical, but it is analogous to the main problem of the
+biologist.
+
+FOOTNOTES:
+
+[Footnote 238: A society presents suggestive analogies with an
+organism, but it certainly is not an organism, and sociologists who
+draw inferences from the assumption of its organic nature must fall
+into error. A vital organism and a society are radically distinguished
+by the fact that the individual components of the former, namely the
+cells, are morphologically as well as functionally differentiated,
+whereas the individuals which compose a society are morphologically
+homogeneous and only functionally differentiated. The resemblances and
+the differences are worked out in E. de Majewski's striking book, _La
+Science de la Civilisation_. Paris. 1908.]
+
+[Footnote 239: It is to be observed that history is (not only
+different in scope but) not co-extensive with anthropology _in time_.
+For it deals only with the development of man in societies, whereas
+anthropology includes in its definition the proto-anthropic period
+when _anthropos_ was still non-social, whether he lived in herds like
+the chimpanzee, or alone like the male ourang-outang. (It has been
+well shown by Majewski that congregations--herds, flocks, packs,
+&c.--of animals are not _societies_; the characteristic of a society
+is differentiation of function. Bee hives, ant hills, may be called
+quasi-societies; but in their case the classes which perform distinct
+functions are morphologically different.)]
+
+[Footnote 240: Recently O. Seeck has applied these principles to the
+decline of Graeco-Roman civilisation in his _Untergang der antiken
+Welt_, 2 vols., Berlin, 1895, 1901.]
+
+[Footnote 241: Darwinian formulae may be suggestive by way of analogy.
+For instance, it is characteristic of social advance that a multitude
+of inventions, schemes and plans are framed which are never carried
+out, similar to, or designed for the same end as, an invention or plan
+which is actually adopted because it has chanced to suit better the
+particular conditions of the hour (just as the works accomplished by
+an individual statesman, artist or savant are usually only a residue
+of the numerous projects conceived by his brain). This process in
+which so much abortive production occurs is analogous to elimination
+by natural selection.]
+
+[Footnote 242: We can ignore here the metaphysical question of
+freewill and determinism. For the character of the individual's brain
+depends in any case on ante-natal accidents and coincidences, and so
+it may be said that the role of individuals ultimately depends on
+chance,--the accidental coincidence of independent sequences.]
+
+[Footnote 243: I have taken this example from G. Tarde's _La logique
+sociale_ (p. 403), Paris, 1904, where it is used for quite a different
+purpose.]
+
+[Footnote 244: _Die kulturhistorische Methode_, Berlin, 1900, p. 26.]
+
+[Footnote 245: _Ibid._ pp. 28, 29.]
+
+
+
+
+X
+
+DARWINISM AND SOCIOLOGY
+
+BY C. BOUGLÉ
+
+_Professor of Social Philosophy in the University of Toulouse and
+Deputy-Professor at the Sorbonne, Paris_
+
+
+How has our conception of social phenomena, and of their history, been
+affected by Darwin's conception of Nature and the laws of its
+transformation? To what extent and in what particular respects have
+the discoveries and hypotheses of the author of _The Origin of
+Species_ aided the efforts of those who have sought to construct a
+science of society?
+
+To such a question it is certainly not easy to give any brief or
+precise answer. We find traces of Darwinism almost everywhere.
+Sociological systems differing widely from each other have laid claim
+to its authority; while, on the other hand, its influence has often
+made itself felt only in combination with other influences. The
+Darwinian thread is worked into a hundred patterns along with other
+threads.
+
+To deal with the problem, we must, it seems, first of all distinguish
+the more general conclusions in regard to the evolution of living
+beings, which are the outcome of Darwinism, from the particular
+explanations it offers of the ways and means by which that evolution
+is effected. That is to say, we must, as far as possible, estimate
+separately the influence of Darwin as an evolutionist and Darwin as a
+selectionist.
+
+The nineteenth century, said Cournot, has witnessed a mighty effort to
+"réintégrer l'homme dans la nature." From divers quarters there has
+been a methodical reaction against the persistent dualism of the
+Cartesian tradition, which was itself the unconscious heir of the
+Christian tradition. Even the philosophy of the eighteenth century,
+materialistic as were for the most part the tendencies of its leaders,
+seemed to revere man as a being apart, concerning whom laws might be
+formulated _à priori_. To bring him down from his pedestal there was
+needed the marked predominance of positive researches wherein no
+account was taken of the "pride of man." There can be no doubt that
+Darwin has done much to familiarise us with this attitude. Take for
+instance the first part of _The Descent of Man_: it is an accumulation
+of typical facts, all tending to diminish the distance between us and
+our brothers, the lower animals. One might say that the naturalist had
+here taken as his motto, "Whosoever shall exalt himself shall be
+abased; and he that shall humble himself shall be exalted." Homologous
+structures, the survival in man of certain organs of animals, the
+rudiments in the animal of certain human faculties, a multitude of
+facts of this sort, led Darwin to the conclusion that there is no
+ground for supposing that the "king of the universe" is exempt from
+universal laws. Thus belief in the _imperium in imperio_ has been, as
+it were, whittled away by the progress of the naturalistic spirit,
+itself continually strengthened by the conquests of the natural
+sciences. The tendency may, indeed, drag the social sciences into
+overstrained analogies, such, for instance, as the assimilation of
+societies to organisms. But it will, at least, have had the merit of
+helping sociology to shake off the pre-conception that the groups
+formed by men are artificial, and that history is completely at the
+mercy of chance. Some years before the appearance of _The Origin of
+Species_, August Comte had pointed out the importance, as regards the
+unification of positive knowledge, of the conviction that the social
+world, the last refuge of spiritualism, is itself subject to
+determinism. It cannot be doubted that the movement of thought which
+Darwin's discoveries promoted contributed to the spread of this
+conviction, by breaking down the traditional barrier which cut man off
+from Nature.
+
+But Nature, according to modern naturalists, is no immutable thing: it
+is rather perpetual movement, continual progression. Their discoveries
+batter a breach directly into the Aristotelian notion of species; they
+refuse to see in the animal world a collection of immutable types,
+distinct from all eternity, and corresponding, as Cuvier said, to so
+many particular thoughts of the Creator. Darwin especially
+congratulated himself upon having been able to deal this doctrine the
+_coup de grâce_: immutability is, he says, his chief enemy; and he is
+concerned to show--therein following up Lyell's work--that everything
+in the organic world, as in the inorganic, is explained by insensible
+but incessant transformations. "Nature makes no leaps"--"Nature knows
+no gaps": these two _dicta_ form, as it were, the two landmarks
+between which Darwin's idea of transformation is worked out. That is
+to say, the development of Darwinism is calculated to further the
+application of the philosophy of Becoming to the study of human
+institutions.
+
+The progress of the natural sciences thus brings unexpected
+reinforcements to the revolution which the progress of historical
+discipline had begun. The first attempt to constitute an actual
+science of social phenomena--that, namely, of the economists--had
+resulted in laws which were called natural, and which were believed to
+be eternal and universal, valid for all times and all places. But this
+perpetuality, brother, as Knies said, of the immutability of the old
+zoology, did not long hold out against the ever-swelling tide of the
+historical movement. Knowledge of the transformations that had taken
+place in language, of the early phases of the family, of religion, of
+property, had all favoured the revival of the Heraclitean view:
+π�ντα �ει̃. As to the categories of political economy, it was
+soon to be recognised, as by Lasalle, that they too are only
+historical. The philosophy of history, moreover, gave expression
+under various forms to the same tendency. Hegel declares that "all
+that is real is rational," but at the same time he shows that all that
+is real is ephemeral, and that for history there is nothing fixed
+beneath the sun. It is this sense of universal evolution that Darwin
+came with fresh authority to enlarge. It was in the name of biological
+facts themselves that he taught us to see only slow metamorphoses in
+the history of institutions, and to be always on the outlook for
+survivals side by side with rudimentary forms. Anyone who reads
+_Primitive Culture_, by Tylor,--a writer closely connected with
+Darwin--will be able to estimate the services which these cardinal
+ideas were to render to the social sciences when the age of
+comparative research had succeeded to that of _à priori_ construction.
+
+Let us note, moreover, that the philosophy of Becoming in passing through
+the Darwinian biology became, as it were, filtered; it got rid of those
+traces of finalism, which, under different forms, it had preserved through
+all the systems of German Romanticism. Even in Herbert Spencer, it has been
+plausibly argued, one can detect something of that sort of mystic
+confidence in forces spontaneously directing life, which forms the very
+essence of those systems. But Darwin's observations were precisely
+calculated to render such an hypothesis futile. At first people may have
+failed to see this; and we call to mind the ponderous sarcasms of Flourens
+when he objected to the theory of Natural Selection that it attributed to
+nature a power of free choice. "Nature endowed with will! That was the
+final error of last century; but the nineteenth no longer deals in
+personifications."[246] In fact Darwin himself put his readers on their
+guard against the metaphors he was obliged to use. The processes by which
+he explains the survival of the fittest are far from affording any
+indication of the design of some transcendent breeder. Nor, if we look
+closely, do they even imply immanent effort in the animal; the sorting out
+can be brought about mechanically, simply by the action of the environment.
+In this connection Huxley could with good reason maintain that Darwin's
+originality consisted in showing how harmonies which hitherto had been
+taken to imply the agency of intelligence and will could be explained
+without any such intervention. So, when later on, objective sociology
+declares that, even when social phenomena are in question, all finalist
+preconceptions must be distrusted if a science is to be constituted, it is
+to Darwin that its thanks are due; he had long been clearing paths for it
+which lay well away from the old familiar road trodden by so many theories
+of evolution.
+
+This anti-finalist doctrine, when fully worked out, was, moreover,
+calculated to aid in the needful dissociation of two notions: that of
+evolution and that of progress. In application to society these had
+long been confounded; and, as a consequence, the general idea seemed
+to be that only one type of evolution was here possible. Do we not
+detect such a view in Comte's sociology, and perhaps even in Herbert
+Spencer's? Whoever, indeed, assumes an end for evolution is naturally
+inclined to think that only one road leads to that end. But those
+whose minds the Darwinian theory has enlightened are aware that the
+transformations of living beings depend primarily upon their
+conditions, and that it is these conditions which are the agents of
+selection from among individual variations. Hence, it immediately
+follows that transformations are not necessarily improvements. Here,
+Darwin's thought hesitated. Logically his theory proves, as Ray
+Lankester pointed out, that the struggle for existence may have as its
+outcome degeneration as well as amelioration: evolution may be
+regressive as well as progressive. Then, too--and this is especially
+to be borne in mind--each species takes its good where it finds it,
+seeks its own path and survives as best it can. Apply this notion to
+society and you arrive at the theory of multilinear evolution.
+Divergencies will no longer surprise you. You will be forewarned not
+to apply to all civilisations the same measure of progress, and you
+will recognise that types of evolution may differ just as social
+species themselves differ. Have we not here one of the conceptions
+which mark off sociology proper from the old philosophy of history?
+
+       *       *       *       *       *
+
+But if we are to estimate the influence of Darwinism upon sociological
+conceptions, we must not dwell only upon the way in which Darwin
+impressed the general notion of evolution upon the minds of thinkers.
+We must go into details. We must consider the influence of the
+particular theories by which he explained the mechanism of this
+evolution. The name of the author of _The Origin of Species_ has been
+especially attached, as everyone knows, to the doctrines of "natural
+selection" and of "struggle for existence," completed by the notion of
+"individual variation." These doctrines were turned to account by very
+different schools of social philosophy. Pessimistic and optimistic,
+aristocratic and democratic, individualistic and socialistic systems
+were to war with each other for years by casting scraps of Darwinism
+at each other's heads.
+
+It was the spectacle of human contrivance that suggested to Darwin his
+conception of natural selection. It was in studying the methods of
+pigeon breeders that he divined the processes by which nature, in the
+absence of design, obtains analogous results in the differentiation of
+types. As soon as the importance of artificial selection in the
+transformation of species of animals was understood, reflection
+naturally turned to the human species, and the question arose, How far
+do men observe, in connection with themselves, those laws of which
+they make practical application in the case of animals? Here we come
+upon one of the ideas which guided the researches of Gallon, Darwin's
+cousin. The author of _Inquiries into Human Faculty and its
+Development_,[247] has often expressed his surprise that, considering
+all the precautions taken, for example, in the breeding of horses,
+none whatever are taken in the breeding of the human species. It seems
+to be forgotten that the species suffers when the "fittest" are not
+able to perpetuate their type. Ritchie, in his _Darwinism and
+Politics_[248] reminds us of Darwin's remark that the institution of
+the peerage might be defended on the ground that peers, owing to the
+prestige they enjoy, are enabled to select as wives "the most
+beautiful and charming women out of the lower ranks."[249] But, says
+Galton, it is as often as not "heiresses" that they pick out, and
+birth statistics seem to show that these are either less robust or
+less fecund than others. The truth is that considerations continue to
+preside over marriage which are entirely foreign to the improvement of
+type, much as this is a condition of general progress. Hence the
+importance of completing Odin's and De Candolle's statistics which are
+designed to show how characters are incorporated in organisms, how
+they are transmitted, how lost, and according to what law eugenic,
+elements depart from the mean or return to it.
+
+But thinkers do not always content themselves with undertaking merely
+the minute researches which the idea of Selection suggests. They are
+eager to defend this or that thesis. In the name of this idea certain
+social anthropologists have recast the conception of the process of
+civilisation, and have affirmed that Social Selection generally works
+against the trend of Natural Selection. Vacher de Lapouge--following
+up an observation by Broca on the point--enumerates the various
+institutions, or customs, such as the celibacy of priests and military
+conscription, which cause elimination or sterilisation of the bearers
+of certain superior qualities, intellectual or physical. In a more
+general way he attacks the democratic movement, a movement, as P.
+Bourget says, which is "anti-physical" and contrary to the natural
+laws of progress; though it has been inspired "by the dreams of that
+most visionary of all centuries, the eighteenth."[250] The "Equality"
+which levels down and mixes (justly condemned, he holds, by the Comte
+de Gobineau), prevents the aristocracy of the blond dolichocephales
+from holding the position and playing the part which, in the interests
+of all, should belong to them. Otto Ammon, in his _Natural Selection
+in Man_, and in _The Social Order and its Natural Bases_,[251]
+defended analogous doctrines in Germany; setting the curve
+representing frequency of talent over against that of income, he
+attempted to show that all democratic measures which aim at promoting
+the rise in the social scale of the talented are useless, if not
+dangerous; that they only increase the panmixia, to the great
+detriment of the species and of society.
+
+Among the aristocratic theories which Darwinism has thus inspired we
+must reckon that of Nietzsche. It is well known that in order to
+complete his philosophy he added biological studies to his
+philological; and more than once in his remarks upon the _Wille zur
+Macht_ he definitely alludes to Darwin; though it must be confessed
+that it is generally in order to proclaim the insufficiency of the
+processes by which Darwin seeks to explain the genesis of species.
+Nevertheless, Nietzsche's mind is completely possessed by an ideal of
+Selection. He, too, has a horror of panmixia. The naturalists'
+conception of "the fittest" is joined by him to that of the "hero" of
+romance to furnish a basis for his doctrine of the Superman. Let us
+hasten to add, moreover, that at the very moment when support was
+being sought in the theory of Selection for the various forms of the
+aristocratic doctrine, those same forms were being battered down on
+another side by means of that very theory. Attention was drawn to the
+fact that by virtue of the laws which Darwin himself had discovered
+isolation leads to etiolation. There is a risk that the privilege
+which withdraws the privileged elements of Society from competition
+will cause them to degenerate. In fact, Jacoby in his _Studies in
+Selection, in connexion with Heredity in Man_,[252] concludes that
+"sterility, mental debility, premature death and, finally, the
+extinction of the stock were not specially and exclusively the fate of
+sovereign dynasties; all privileged classes, all families in
+exclusively elevated positions share the fate of reigning families,
+although in a minor degree and in direct proportion to the loftiness
+of their social standing. From the mass of human beings spring
+individuals, families, races, which tend to raise themselves above the
+common level; painfully they climb the rugged heights, attain the
+summits of power, of wealth, of intelligence, of talent, and then, no
+sooner are they there than they topple down and disappear in gulfs of
+mental and physical degeneracy." The demographical researches of
+Hansen[253] (following up and completing Dumont's) tended, indeed, to
+show that urban as well as feudal aristocracies, burgher classes as
+well as noble castes, were liable to become effete. Hence it might
+well be concluded that the democratic movement, operating as it does
+to break down class barriers, was promoting instead of impeding human
+selection.
+
+       *       *       *       *       *
+
+So we see that, according to the point of view, very different
+conclusions have been drawn from the application of the Darwinian idea
+of Selection to human society. Darwin's other central idea, closely
+bound up with this, that, namely, of the "struggle for existence" also
+has been diversely utilised. But discussion has chiefly centered upon
+its signification. And while some endeavour to extend its application
+to everything, we find others trying to limit its range. The
+conception of a "struggle for existence" has in the present day been
+taken up into the social sciences from natural science, and adopted.
+But originally it descended from social science to natural. Darwin's
+law is, as he himself said, only Malthus' law generalised and extended
+to the animal world: a growing disproportion between the supply of
+food and the number of the living is the fatal order whence arises the
+necessity of universal struggle, a struggle which, to the great
+advantage of the species, allows only the best equipped individuals to
+survive. Nature is regarded by Huxley as an immense arena where all
+living beings are gladiators.[254]
+
+Such a generalisation was well adapted to feed the stream of
+pessimistic thought; and it furnished to the apologists of war, in
+particular, new arguments, weighted with all the authority which in
+these days attaches to scientific deliverances. If people no longer
+say, as Bonald did, and Moltke after him, that war is a providential
+fact, they yet lay stress on the point that it is a natural fact. To
+the peace party Dragomirov's objection is urged that its attempts are
+contrary to the fundamental laws of nature, and that no sea wall can
+hold against breakers that come with such gathered force.
+
+But in yet another quarter Darwinism was represented as opposed to
+philanthropic intervention. The defenders of the orthodox political
+economy found in it support for their tenets. Since in the organic
+world universal struggle is the condition of progress, it seemed
+obvious that free competition must be allowed to reign unchecked in
+the economic world. Attempts to curb it were in the highest degree
+imprudent. The spirit of Liberalism here seemed in conformity with the
+trend of nature: in this respect, at least, contemporary naturalism,
+offspring of the discoveries of the nineteenth century, brought
+reinforcements to the individualist doctrine, begotten of the
+speculations of the eighteenth: but only, it appeared, to turn mankind
+away for ever from humanitarian dreams. Would those whom such
+conclusions repelled be content to oppose to nature's imperatives
+only the protests of the heart? There were some who declared, like
+Brunetière, that the laws in question, valid though they might be for
+the animal kingdom, were not applicable to the human. And so a return
+was made to the classic dualism. This indeed seems to be the line that
+Huxley took, when, for instance, he opposed to the cosmic process an
+ethical process which was its reverse.
+
+But the number of thinkers whom this antithesis does not satisfy grows
+daily. Although the pessimism which claims authorisation from Darwin's
+doctrines is repugnant to them, they still are unable to accept the
+dualism which leaves a gulf between man and nature. And their
+endeavour is to link the two by showing that while Darwin's laws
+obtain in both kingdoms, the conditions of their application are not
+the same: their forms, and, consequently, their results, vary with the
+varying mediums in which the struggle of living beings takes place,
+with the means these beings have at disposal, with the ends even which
+they propose to themselves.
+
+Here we have the explanation of the fact that among determined
+opponents of war partisans of the "struggle for existence" can be
+found: there are disciples of Darwin in the peace party. Novicow, for
+example, admits the "_combat universel_" of which Le Dantec[255]
+speaks; but he remarks that at different stages of evolution, at
+different stages of life the same weapons are not necessarily
+employed. Struggles of brute force, armed hand to hand conflicts, may
+have been a necessity in the early phases of human societies.
+Nowadays, although competition may remain inevitable and
+indispensable, it can assume milder forms. Economic rivalries,
+struggles between intellectual influences, suffice to stimulate
+progress: the processes which these admit are, in the actual state of
+civilisation, the only ones which attain their end without waste, the
+only ones logical. From one end to the other of the ladder of life,
+struggle is the order of the day; but more and more as the higher
+rungs are reached, it takes on characters which are proportionately
+more "humane."
+
+Reflections of this kind permit the introduction into the economic
+order of limitations to the doctrine of "laisser faire, laisser
+passer." This appeals, it is said, to the example of nature where
+creatures, left to themselves, struggle without truce and without
+mercy; but the fact is forgotten that upon industrial battlefields the
+conditions are different. The competitors here are not left simply to
+their natural energies: they are variously handicapped. A rich store
+of artificial resources exists in which some participate and others do
+not. The sides then are unequal; and as a consequence the result of
+the struggle is falsified. "In the animal world," said De
+Laveleye,[256] criticising Spencer, "the fate of each creature is
+determined by its individual qualities; whereas in civilised societies
+a man may obtain the highest position and the most beautiful wife
+because he is rich and well-born, although he may be ugly, idle or
+improvident; and then it is he who will perpetuate the species. The
+wealthy man, ill constituted, incapable, sickly, enjoys his riches and
+establishes his stock under the protection of the laws." Haycraft in
+England and Jentsch in Germany have strongly emphasised these
+"anomalies," which nevertheless are the rule. That is to say that even
+from a Darwinian point of view all social reforms can readily be
+justified which aim at diminishing, as Wallace said, inequalities at
+the start.
+
+But we can go further still. Whence comes the idea that all measures
+inspired by the sentiment of solidarity are contrary to Nature's
+trend? Observe her carefully, and she will not give lessons only in
+individualism. Side by side with the struggle for existence do we not
+find in operation what Lanessan calls "association for existence."
+Long ago, Espinas had drawn attention to "societies of animals,"
+temporary or permanent, and to the kind of morality that arose in
+them. Since then, naturalists have often insisted upon the importance
+of various forms of symbiosis. Kropotkin in _Mutual Aid_ has chosen
+to enumerate many examples of altruism furnished by animals to
+mankind. Geddes and Thomson went so far as to maintain that "Each of
+the greater steps of progress is in fact associated with an increased
+measure of subordination of individual competition to reproductive or
+social ends, and of interspecific competition to co-operative,
+association."[257] Experience shows, according to Geddes, that the
+types which are fittest to surmount great obstacles are not so much
+those who engage in the fiercest competitive struggle for existence,
+as those who contrive to temper it. From all these observations there
+resulted, along with a limitation of Darwinian pessimism, some
+encouragement for the aspirations of the collectivists.
+
+And Darwin himself would, doubtless, have subscribed to these
+rectifications. He never insisted, like his rival, Wallace, upon the
+necessity of the solitary struggle of creatures in a state of nature,
+each for himself and against all. On the contrary, in _The Descent of
+Man_, he pointed out the serviceableness of the social instincts, and
+corroborated Bagehot's statements when the latter, applying laws of
+physics to politics, showed the great advantage societies derived from
+intercourse and communion. Again, the theory of sexual evolution which
+makes the evolution of types depend increasingly upon preferences,
+judgments, mental factors, surely offers something to qualify what
+seems hard and brutal in the theory of natural selection.
+
+But, as often happens with disciples, the Darwinians had out-Darwined
+Darwin. The extravagances of social Darwinism provoked a useful
+reaction; and thus people were led to seek, even in the animal
+kingdom, for facts of solidarity which would serve to justify humane
+effort.
+
+       *       *       *       *       *
+
+On quite another line, however, an attempt has been made to connect
+socialist tendencies with Darwinian principles. Marx and Darwin have
+been confronted; and writers have undertaken to show that the work of
+the German philosopher fell readily into line with that of the English
+naturalist and was a development of it. Such has been the endeavour of
+Ferri in Italy and of Woltmann in Germany, not to mention others. The
+founders of "scientific socialism" had, moreover, themselves thought
+of this reconciliation. They make more than one allusion to Darwin in
+works which appeared after 1859. And sometimes they use his theory to
+define by contrast their own ideal. They remark that the capitalist
+system, by giving free course to individual competition, ends indeed
+in a _bellum omnium contra omnes_; and they make it clear that
+Darwinism, thus understood, is as repugnant to them as to Dühring.
+
+But it is at the scientific and not at the moral point of view that
+they place themselves when they connect their economic history with
+Darwin's work. Thanks to this unifying hypothesis, they claim to have
+constructed--as Marx does in his preface to _Das Kapital_--a veritable
+natural history of social evolution. Engels speaks in praise of his
+friend Marx as having discovered the true mainspring of history hidden
+under the veil of idealism and sentimentalism, and as having
+proclaimed in the _primum vivere_ the inevitableness of the struggle
+for existence. Marx himself, in _Das Kapital_, indicated another
+analogy when he dwelt upon the importance of a general technology for
+the explanation of this psychology:--a history of tools which would be
+to social organs what Darwinism is to the organs of animal species.
+And the very importance they attach to tools, to apparatus, to
+machines, abundantly proves that neither Marx nor Engels were likely
+to forget the special characters which mark off the human world from
+the animal. The former always remains to a great extent an artificial
+world. Inventions change the face of its institutions. New modes of
+production revolutionise not only modes of government, but modes even
+of collective thought. Therefore it is that the evolution of society
+is controlled by laws special to it, of which the spectacle of nature
+offers no suggestion.
+
+If, however, even in this special sphere, it can still be urged that
+the evolution of the material conditions of society is in accord with
+Darwin's theory, it is because the influence of the methods of
+production is itself to be explained by the incessant strife of the
+various classes with each other. So that in the end Marx, like Darwin,
+finds the source of all progress is in struggle. Both are grandsons of
+Heraclitus:--π�λεμος πατἠ� π�ντων. It sometimes happens, in
+these days, that the doctrine of revolutionary socialism is contrasted
+as rude and healthy with what may seem to be the enervating tendency
+of "solidarist" philanthropy: the apologists of the doctrine then
+pride themselves above all upon their faithfulness to Darwinian
+principles.
+
+       *       *       *       *       *
+
+So far we have been mainly concerned to show the use that social
+philosophies have made of the Darwinian laws for practical purposes:
+in order to orientate society towards their ideals each school tries
+to show that the authority of natural science is on its side. But even
+in the most objective of theories, those which systematically make
+abstraction of all political tendencies in order to study the social
+reality in itself, traces of Darwinism are readily to be found.
+
+Let us take for example Durkheim's theory of Division of Labour.[258]
+The conclusions he derives from it are that whenever professional
+specialisation causes multiplication of distinct branches of activity,
+we get organic solidarity--implying differences--substituted for
+mechanical solidarity, based upon likenesses. The umbilical cord, as
+Marx said, which connects the individual consciousness with the
+collective consciousness is cut. The personality becomes more and more
+emancipated. But on what does this phenomenon, so big with
+consequences, itself depend? The author goes to social morphology for
+the answer: it is, he says, the growing density of population which
+brings with it this increasing differentiation of activities. But,
+again, why? Because the greater density, in thrusting men up against
+each other, augments the intensity of their competition for the means
+of existence; and for the problems which society thus has to face
+differentiation of functions presents itself as the gentlest solution.
+
+Here one sees that the writer borrows directly from Darwin.
+Competition is at its maximum between similars, Darwin had declared;
+different species, not laying claim to the same food, could more
+easily coexist. Here lay the explanation of the fact that upon the
+same oak hundreds of different insects might be found. Other things
+being equal, the same applies to society. He who finds some unadopted
+specialty possesses a means of his own for getting a living. It is by
+this division of their manifold tasks that men contrive not to crush
+each other. Here we obviously have a Darwinian law serving as
+intermediary in the explanation of that progress of division of labour
+which itself explains so much in the social evolution.
+
+And we might take another example, at the other end of the series of
+sociological systems. G. Tarde is a sociologist with the most
+pronounced anti-naturalistic views. He has attempted to show that all
+application of the laws of natural science to society is misleading.
+In his _Opposition Universelle_ he has directly combatted all forms of
+sociological Darwinism. According to him the idea that the evolution
+of society can be traced on the same plan as the evolution of species
+is chimerical. Social evolution is at the mercy of all kinds of
+inventions, which by virtue of the laws of imitation modify, through
+individual to individual, through neighbourhood to neighbourhood, the
+general state of those beliefs and desires which are the only
+"quantities" whose variation matters to the sociologist. But, it may
+be rejoined, that however psychical the forces may be, they are none
+the less subject to Darwinian laws. They compete with each other; they
+struggle for the mastery of minds. Between types of ideas, as between
+organic forms, selection operates. And though it may be that these
+types are ushered into the arena by unexpected discoveries, we yet
+recognise in the psychological accidents, which Tarde places at the
+base of everything, near relatives of those small accidental
+variations upon which Darwin builds. Thus, accepting Tarde's own
+representations, it is quite possible to express in Darwinian terms,
+with the necessary transpositions, one of the most idealistic
+sociologies that have ever been constructed.
+
+These few examples suffice. They enable us to estimate the extent of
+the field of influence of Darwinism. It affects sociology not only
+through the agency of its advocates but through that of its opponents.
+The questions to which it has given rise have proved no less fruitful
+than the solutions it has suggested. In short, few doctrines, in the
+history of social philosophy, will have produced on their passage a
+finer crop of ideas.
+
+FOOTNOTES:
+
+[Footnote 246: P. Flourens, _Examen du Livre de M. Darwin sur
+l'Origine des Espèces_, p. 53, Paris, 1864. See also Huxley,
+"Criticisms on the _Origin of Species," Collected Essays_, Vol. II, p.
+102, London, 1902.]
+
+[Footnote 247: _Inquiries into Human Faculty_, pp. 1, 2, 3 sq.,
+London, 1883.]
+
+[Footnote 248: _Darwinism and Politics_, pp. 9, 22, London, 1889.]
+
+[Footnote 249: _Life and Letters of Charles Darwin_, II. p. 385.]
+
+[Footnote 250: V. de Lapouge, _Les Sélections sociales_, p. 259,
+Paris, 1896.]
+
+[Footnote 251: _Die natärliche Auslese beim Menschen_, Jena, 1893; _Du
+Gesellschaftsordnung und ihre natürlichen Grundlagen. Entwurf einer
+Sozialanthropologie_, Jena, 1896.]
+
+[Footnote 252: _Etudes sur la Sélection dans ses rapports avec
+l'hérédité chez l'homme_, Paris, p. 481, 1881.]
+
+[Footnote 253: _Die drei Bevölkerungsstufen_, Munich, 1889.]
+
+[Footnote 254: _Evolution and Ethics_, p. 200; _Collected Essays_,
+Vol. IX, London, 1894.]
+
+[Footnote 255: _Les Luttes entre Sociétés humaines et leurs phases
+successives_, Paris, 1893.]
+
+[Footnote 256: _Le socialisme contemporain_, p. 384 (6th edit.),
+Paris, 1891.]
+
+[Footnote 257: Geddes and Thomson, _The Evolution of Sex_, p. 311,
+London, 1889.]
+
+[Footnote 258: _De la Division du Travail social_, Paris. 1893.]
+
+
+
+
+INDEX
+
+
+_Abraxas grossulariata_, 100
+
+Acquired characters, transmission of, 20, 28, 42, 94, 120, 149, 171, 173
+
+_Acraea johnstoni_, 290
+[Transcriber's Note: No such page number or reference seen]
+
+
+Adaptation, 24, 27, 34, 39, 42-45, 50, 58, 79-86, 106, 107
+
+Adloff, 140
+
+Alexander, 217
+
+Ameghino, 132, 138
+
+Ammon, O., Works of, 271
+
+_Anaea divina_, 69
+
+Anglicus, Bartholomaeus, 237
+
+Ankyroderma, 40
+
+Anomma, 44
+
+Anthropops, 132
+
+Ants, modifications of, 43-46, 51
+
+Ardigò, 207, 208
+
+Argyll, Huxley and the Duke of, 238
+
+Aristotle, 3, 237, 240
+
+Avenarius, 211
+
+
+Bacon, on mutability of species, 4, 5
+
+Baehr, von, on Cytology, 99
+
+Bain, 194
+
+Baldwin, J. M., 53, Foot Note 165
+
+Balfour, A. J., 241
+
+Barratt, 217
+
+Bates, H. W., on Mimicry, 70, 76
+  --232
+
+BATESON, W., on _Heredity and
+Variation in Modern Lights_, 87-110
+  --on discontinuous evolution, 30
+
+Bathmism, 14
+
+Bells (Sir Charles) _Anatomy of Expression_, 177
+
+Bentham, Jeremy, 217, 218
+
+Bergson, H., 208
+
+Berkeley, 200
+
+Berthelot, 228
+
+Bickford, E., experiments on degeneration by, 52
+
+Biophores, 47
+
+Blumenbach, 89
+
+Bodin, 256
+
+Bonald, on war, 273
+
+Bonnet, 6
+
+BOUGLÉ, C., on _Darwinism and Sociology_, 264-280
+
+Bourdeau, 253
+
+Bourget, P., 270
+
+Boutroux, 208
+
+Brassica, hybrids of, 106
+
+_Brassica Napus_, 106
+
+Broca, 137, 270
+
+Brock, on Kant, Foot Note 6
+
+Brunetière, 274
+
+Bruno, on Evolution, 4
+
+Buch, von, 15
+
+Buckle, 252, 253, 256, 258
+
+Buffon, 6-15, 21, 88
+
+Burdon-Sanderson, J., letter from, Foot Note 224
+
+BURY, J. B., on _Darwinism and History_, 246-263
+
+Butler, Samuel, 9, Foot Note 17, Foot Note 57, Foot Note 61, 94, Foot Note 66, 107
+
+Butterflies, mimicry in, 65-83
+  --sexual characters in, 59-63
+
+
+Cabanis, 201
+
+Candolle, de, 270
+
+Carneri, 217
+
+_Castnia linus_, 76
+
+Caterpillars, variation in, 36, 37
+
+Cesnola, experiments on Mantis by, 65
+
+Chaerocampa, colouring of, 68
+
+Chambers, R., _The Vestiges of Creation_ by, 15
+
+Chromosomes and Chromomeres, 47, 96-100
+
+Chun, Foot Note 36
+
+Claus, Foot Note 21
+
+Clodd, E., Foot Note 13
+
+Coadaptation, 41-54
+
+_Colobopsis truncata_, 44
+
+Colour, E. B. Poulton, in relation to Sexual Selection, 61-65
+
+Comte, A., 200-203, 252-255, 262, 265
+
+Condorcet, 221, 250, 252, 258
+
+Cope, 138
+
+Correlation of organisms, Darwin's idea of the, 2
+
+Cournot, 265
+
+Cuvier, 9, 10, 266, 268
+
+Cytology and heredity, 95, 96, 99, 100
+
+
+_Danaida chrysippus_, 75
+
+_Danaida genutia_, 75
+
+_D. Plexippus_, 75
+
+Dantec, Le, 274
+
+Darwin, Charles, as an Anthropologist, 146-165
+  --on ants, 44
+  --and S. Butler, Foot Note 61, 94
+  --on Cirripedia, 212
+  --on the Descent of Man, 111-145
+  --evolutionist authors referred to in the _Origin_ by, 9
+
+Darwin, Charles, and Haeckel, 137
+  --and History, 246-263
+  --and Huxley, 112
+  --on Lamarck, 28, 129
+  --on Language, 124
+  --and Malthus, 16, 24, 91
+  --on Patrick Matthew, 19
+  --on mental evolution, 166-196
+  --on Natural Selection, 21, 41, 54, 55, 122
+  --a "Naturalist for Naturalists," 87
+  --his personality, 187
+  --his influence on Philosophy, 197-222
+  --predecessors of, 1-22
+  --his views on religion, etc., 115, 116, 219-222
+  --his influence on religious thought, 223-245
+  --causes of his success, 10, 90
+
+Darwin, Charles, on the _Vestiges of Creation_, 15
+  --and Wallace, 23, 183
+  --on evolution, 7-15, 88
+  --on Lamarckism, 11
+
+Darwin, F., on Prichard's "Anticipations," 21
+
+Darwinism, Sociology, Evolution and, 17-18
+
+Degeneration, 49-51, 93
+
+Deniker, 137
+
+Descartes, 4
+
+Descent, history of doctrine of, 1
+
+_Descent of Man_, G. Schwalbe on _The_, 111-145
+  --rejection in Germany of _The_, 156
+
+Diderot, 6, 198
+
+Dimorphism, seasonal, 30
+
+_Dismorphia orise_, 75
+
+Dragomirov, 273
+
+Driesch, Foot Note 67
+
+Dryopithecus, 132
+
+Dubois, E., on Pithecanthropus, 132, 137
+
+Dühring, 214, 277
+
+Duns Scotus, 200
+
+Duret, C., 6
+
+Durkheim, on division of labour, 278
+
+
+Ecology, Foot Note 205
+
+Eimer, 109
+
+_Elymnias undularis_, 73, 75
+
+Embryology, the Origin of Species and, 154, 155
+
+Empedocles, 3, 27, 151
+
+Engels, 277
+
+Environment, action of, 12, 13, 15
+
+Epicurus, a poet of Evolution, 4
+
+Eristalis, 75
+
+Espinas, 275
+
+Evolution, and creation, 233
+  --conception of, 3-5, 9, 148, 151, 198
+  --discontinuous, 30
+  --experimental, 5, 7
+  --factors of, 11-15
+  --mental, 194
+  --Lloyd Morgan on mental factors in, 166-196
+  --Darwinism and Social, 18
+  --Saltatory, 29-32
+  --Herbert Spencer on, 204-207
+  --Philosophers and modern methods of studying, 4
+
+Expression of the Emotions, 177-184
+
+
+Ferri, 277
+
+Ferrier, his work on the brain, 523
+[Transcriber's note: No such page number or reference seen]
+
+Fichte, 222
+
+Flourens, 267
+
+Flowers and Insects, 61, 78
+
+Fouillée, 207, 208
+
+Fraipont, on skulls from Spy, 134
+
+
+GADOW, 162
+
+_Gallus bankiva_, 102
+
+Gallon, F., 125, 150, 269
+
+Geddes, P., 17, Foot Note 32
+
+Geddes, P. and A. W. Thomson, 276
+
+Gegenbaur, 150, 163
+
+Genetics, 93, 96
+
+_Germ-plasm_, continuity of, 95
+--Weismann on, 46-51
+
+Germinal Selection, 36, 37, 46-51, 64
+
+Gibbon, 248
+
+Giuffrida-Ruggeri, 138, 140
+
+Giotto, 259
+
+Gizycki, 217
+
+Goethe and Evolution, 8, 14, 15, 201
+--on the relation between Man and Mammals, 161, 163
+--221
+
+Gore, Dr., 226
+
+Gorjanovi�-Kramberger, 134
+
+Gosse, P. H., 234
+
+_Grapta C. album_, 69
+
+Groos, 187, 188
+
+Gulick, 15, 53
+
+Guyau, 217
+
+
+Haberlandt, G., 34
+
+HAECKEL, E., on _Charles Darwin as an Anthropologist_, 146-165
+  --and Darwin, 135-151, 137, 146-165
+  --on the Descent of Man, 137, 143
+  --on Lamarck, 8, Foot Note 21
+  --a leader in the Darwinian controversy, 137
+  --217
+
+Häcker, 33
+
+Hansen, 272
+
+Hartmann, von, 240
+
+Harvey, 4
+
+Haycraft, 275
+
+Hegel, 201, 203, 215, 251, 252, 255
+
+Heraclitus, 278
+
+Herder, 4, 5, 20
+
+Heredity and Cytology, 95, 96
+  --Haeckel on, 147, 148, 149, 153
+  --and Variation, 87-110
+  --219, 224
+
+Hering, E., on Memory, 153
+
+Hertwig, O., 150
+
+History, Darwin and, 246-263
+
+Hobbes, T., 200, 215
+
+Hobhouse, 242
+
+HÖFFDING, H., on _The Influence of the Conception of Evolution
+                             on Modern Philosophy_, 197-222
+
+Holothurians, calcareous bodies in skin of, 37-41
+
+_Homo heidelbergensis_, Foot Note 118
+
+_H. neandertalensis_, 138
+
+_H. pampaeus_, 144
+
+_H. primigenius_, 133, 134, 138, 144
+
+_Homunculus_, 132
+
+Hooker, Sir J. D., and Darwin, 23, 116
+
+Huber, 170
+
+Hügel, F. von, Foot Note 221
+
+Hume, 200
+
+Hutcheson, 216
+
+Huxley, T. H., and Darwin, 112, 116, 268
+  --and the Duke of Argyll, 238
+  --on Lamarck, 89
+  --on Man, 111, 112, 137, 146, 156, 160, 163
+  --on Selection, 24, 91
+  --on transmission of acquired characters, 149
+  --14, 24, 104, 231-236, 273, 274
+
+Hybrids, Sterility of, 104, 105, 106
+
+
+Inheritance of acquired characters, 93, 94
+
+Insects and Flowers, 60, 61, 78, 79
+
+Instinct, 122, 172-175
+
+Irish Elk, an example of coadaptation, 41, 42, 45
+
+
+Jacoby, _Studies in Selection_ by, 272
+
+James, W., 180, 191, 211
+
+Jentsch, 275
+
+
+Kallima, protective colouring of, 35, 68, 70
+
+_K. inachis_, 68
+
+Kammerer's experiments on Salamanders, 28
+
+Kant, I., 4, 5, 6, 27, 198, 211, 212, 217, 221, 222
+
+Keane, on the Primates, 138
+
+Keith, on Anthropoid Apes, 138
+
+Kepler, 198
+
+Klaatsch, on Ancestry of Man, 140
+
+Klaatsch and Hauser, 134
+
+Knies, 266
+
+Kölliker, his views on Evolution, 29, 150
+
+Kollmann, on origin of human races, 144
+
+Korschinsky, 31
+
+Krause, E., Foot Note 10, 13
+
+Kropotkin, 214, 275
+
+
+Lamarck, his division of the Animal Kingdom, 160, 161
+  --Darwin's opinion of, 129
+  --on Evolution, 9-14, 21, 25, 171, 172, 173, 179, 180, 201, 202, 253
+  --on Man, 146, 148, 160, 163
+  --89, 109, 201, 202, 233
+
+Lamarckian principle, 28, 41-44, 50-54, 67, 84, 86
+
+Lamb, C., 229
+
+Lamettrie, 198
+
+Lamprecht, 260-263
+
+Lanessan, J. L. de, Foot Note 17, 275
+
+Lang, Foot Note 21
+
+Lange, 180
+
+Language, Darwin on, 123, 124
+  --Evolution and the Science of, 178, 179, 188
+
+Lankester, Sir E. Ray, on degeneration, 268
+  --on educability, 170, 189
+
+Lankester, Sir E. Ray, on the germ-plasm theory, 150
+
+Lapouge, Vacher de, 270
+
+Lartet, M. E., 189
+
+Lasalle, 266
+
+Laveleye, de, 275
+
+Lawrence, W., 89, Foot Note 65
+
+Lehmann-Nitsche, 138, 144
+
+Leibnitz, 4, 5, 213
+
+Lepidoptera, variation in, 37, 60-63
+
+Lessing, 4, 221
+
+Liddon, H. P., 234
+
+_Limenitis archippus_, 74
+
+Linnaeus, 6
+
+Locy, W. A., Foot Note 15
+
+Lovejoy, Foot Note 56
+
+Lubbock, 125
+
+Lucretius, a poet of Evolution, 4
+
+Lyell, Sir Charles, and Darwin, 23, 116
+  --the uniformitarian teaching of, 89
+
+
+Macacus, ear of, 119
+
+Mach, E., 153, 211
+
+Mahoudeau, 137
+
+Maillet, de, 6
+
+Majewski, Foot Note 238, Foot Note 239
+
+Malthus, his influence on Darwin, 16-18, 21, 24, 91
+  --200, 273
+
+Man, Descent of, 126, 127, 128, 131-145, 156-165, 189, 254, 265
+  --mental and moral qualities of animals and, 122-126, 164, 188-192
+  --pre-Darwinian views on the Descent of, 1
+
+Man, Tertiary flints worked by, 136
+
+_Man_, G. Schwalbe on Darwin's _Descent of_, 111-145
+
+Manouvrier, 137
+
+_Mantis religiosa_, colour experiments on, 65, 68
+
+Marx, 262, 276-278
+
+Matthew, P., and Natural Selection, 18, 19
+
+Maupertuis, 6, 88, 103
+
+Mayer, R., 197
+
+_Mechanitis lysimnia_, 77
+
+_Melinaea ethra_, 77
+
+Mendel, 97-100, 184, 228
+
+Merz, J. T., Foot Note 14
+
+Mesopithecus, 132
+
+Mill, J. S., 193, 200, 202, 218
+
+Mimicry, 70-82
+
+Moltke, on war, 273
+
+Monkeys, fossil, 132
+
+Montesquieu, 248
+
+Monticelli, 155
+
+MORGAN, C. LLOYD, on _Mental Factors in Evolution_, 166-196
+  --on Organic Selection, 53
+
+Morgan, T. H., 99
+
+Morselli, 138
+
+Mortillet, 136
+
+Moseley, Foot Note 224
+
+Muller, Fritz, _Für Darwin_ by, 154
+  --on Mimicry, 233
+  --59, 77
+
+Muller, J., 147
+
+Müller, Max, on language, 124
+
+Mutation, 15, 31, 184, 199, 209
+
+
+Nägeli, 109, 151, 153
+
+Nathusius, 103
+
+Natural Selection, Darwin's views on, 90, 91, 122, 149
+  --Darwin and Wallace on, 2, 163, 183
+  --and design, 241, 242
+  --and educability, 195
+  --and human development, 125, 256, 257
+  --16-20, 25, 26, 41, 55-58, 64-86, 87-96, 199, 233
+
+Neandertal skulls, 133, 134
+
+Neodarwinism, 150
+
+Newton, A., Foot Note 59
+
+Newton, I., 197, 198
+
+Niebuhr, 249, 263
+
+Nietzsche, 214, 271
+
+Nitsche, 119
+
+Novicow, 274
+
+Nuttall, G. H. F., 135
+
+
+Occam, 200
+
+Odin, 270
+
+Oecology, see Ecology
+
+_Oenothera lamarckiana_, 32
+
+Oestergren, on Holothurians, 37-39
+
+Oken, L., 7, 201
+
+Organic Selection, 53, 54, 172, 173
+
+Orthogenesis, 109
+
+Osborn, H. F., 53, Foot Note 165
+  --_From the Greeks to Darwin_ by, 3-5, 12, 14, 20
+
+_Ovibos moschatus_, 67
+
+Owen, Sir Richard, 111
+
+
+Packard, A. S., Foot Note 12, Foot Note 18
+
+Palaeopithecus, 132
+
+Paley, 18, 242, 244
+
+Panmixia, Weismann's principle of, 54
+
+_Papilio dardanus_, 72, 73, 74
+
+_P. meriones_, 73
+
+_P. merope_, 72
+
+Pearson, K., Foot Note 7
+
+Penck, 136
+
+Peridineae, 33
+
+Perrier, E., Foot Note 21, 20
+
+Perthes, B. de, 123
+
+Pfeffer, W., 28
+
+Philosophy, influence of the conception of evolution on modern, 197-222
+
+Pithecanthropus, 133, 134, 138, 143
+
+Pitheculites, 144
+
+Plate, Foot Note 37
+
+Pliopithecus, 132
+
+Pouchet, G., Foot Note 3
+
+POULTON, E. B., experiments on Butterflies by, 65
+  --on J. C. Prichard, 20
+  --on Mimicry, 69, 71, 75, 78
+  --Foot Note 34, Foot Note 43, Foot Note 49, Foot Note 55
+
+Prichard, J. C., 20, 21, 89, Foot Note 65
+
+_Pronuba yuccasella_, 79
+
+Protective resemblance, 65-70
+
+Pusey, 115
+
+
+Quatrefages, A. de, Foot Note 21, 19
+
+
+Radiolarians, 33
+
+Ranke, 249, 251, 255, 263
+
+Rau, A., 153
+
+Ray, J., 4
+
+Regeneration, Foot Note 71
+
+Religious thought, Darwin's influence on, 223-245
+
+Reversion, 120, 121
+
+Ridley, H. N., Foot Note 88
+
+Ritchie, 270
+
+Robinet, 6
+
+Rolph, 217
+
+Romanes, G. J., Foot Note 3, 15, 32, 54, 164, 234
+
+Roux, 151, 152
+
+Ruskin, 230
+
+Rutot, 136
+
+Saint-Hilaire, E. G. de, 8, 15, 20
+
+Saltatory Evolution, 29-32 (see also Mutations)
+
+Sanders, experiments on Vanessa by, 65
+
+Savigny, 249
+
+Schelling, 4, 5, 200, 201
+
+Schleiden and Schwann, Cell-theory of, 147
+
+Schoetensack, on _Homo heidelbergensis_, Foot Note 118
+
+Schütt, 23
+
+SCHWALBE, G., on _The Descent of Man_, 111-145
+
+Seeck, O., Foot Note 240
+
+Segregation, 97, 98
+
+Selection, artificial, 24, 25, 26, 41, 45, 120, 269-272
+  --germinal, 35, 36, 46-52, 64
+
+Selection, natural (see Natural Selection)
+  --organic, 53, 171, 172
+  --sexual, 55-64, 117, 118
+  --social and natural, 271
+  --23-86, 103, 129, 130
+
+Selenka, 131
+
+Semnopithecus, 132
+
+Semon, R., 28, 153
+
+Sergi, 138, 143
+
+Sex, recent investigations on, 99, 100
+
+Sibbern, 201
+
+_Smerinthus ocellata_, 38
+
+_Smerinthus populi_, 38
+
+_S. tiliae_, 38
+
+Smith, A., 200
+
+Sociology,  Darwinism and, 264-280
+  --History and, 255
+
+Sollas, W. J., 134
+
+Sorley, W. R., 217
+
+Species and varieties, 100
+
+Spencer, H., on evolution, 204-209
+  --on the theory of Selection, 41
+
+Spencer, H., on Sociology, 268
+  --on the transmission of acquired characters, 149
+  --on Weismann, 41, 150
+  --2, 17, 217, 231, 268
+
+Sphingidae, variation in, 37
+
+Spinoza, 153, 206
+
+Standfuss, 82
+
+Stephen, L., 217
+
+Sterility in hybrids, 104-106
+
+Sterne, C, Foot Note 10
+
+Struggle for existence, 25, 26, 272-274
+
+Sutton, A. W., Foot Note 73
+
+Synapta, calcareous bodies in skin of, 38-41
+
+Syrphus, 75
+
+
+Tarde, G., 279
+
+Tennant, F. R., Foot Note 218
+
+Tetraprothomo, 138, 144
+
+THOMSON, J. A., on _Darwin's Predecessors_, 1-22
+  --150
+  --and P. Geddes, 276
+
+Treschow, 201
+
+Treviranus, 8, 14, 15
+
+Turgot, 249
+
+Turner, Sir W., 150
+
+Tylor, 267
+
+Tyndall, W., 267
+
+Tyrrell, G, Foot Note 222
+
+
+Uhlenhuth, on blood reactions, 135
+
+Use and disuse, 28, 41-43, 48-54, 94, 95, 119, 149
+
+
+Vanessa, 63
+
+_V. levana_, 31
+
+_V. polychloros_, 82
+
+_V. urticae_, 65, 82
+
+Variability, Darwin's attention directed to, 24
+  --W. Bateson on, 87-110
+  --causes of, 200
+
+Variation, Darwin's views as an evolutionist, and as a systematist, on, 212
+  --and heredity, 87-110
+  --minute, 28-32
+  --in relation to species, 100, 101
+
+Varigny, H. de, 6, 19
+
+Verworn, 136
+
+_Vestiges of Creation_, Darwin on _The_, 15
+
+Virchow, his opposition to Darwin, 157, 158
+  --on the transmission of acquired characters, 149
+
+Vogt, 137
+
+Voltaire, 248
+
+VRIES, H. de, the Mutation theory of, 31, 101, 151, 213
+
+
+WAGGETT, Rev. P. N., on _The Influence of Darwin upon Religious
+                                    Thought_, 223-245
+
+Wallace, A. R., on Colour, 63, 71
+  --and Darwin, Foot Note 7, 23, 183
+  --on the Descent of Man, 116
+  --on Malthus, 17
+  --on Natural Selection, 2, 16, 163, 232
+
+Wallace, A. R., on social reforms, 275, 276
+  --on Sexual Selection, 183, 184
+
+Walton, 237
+
+Watt, J., and Natural Selection, 21
+
+WEISMANN, A., on _The Selection Theory_, 23-86
+  --his germ-plasm theory, 46-51, 149, 150
+  --and Prichard, 20
+  --and Spencer, 42
+
+Weismann, A., on the transmission of acquired characters, 93-95
+  --156
+
+Wells, W. C, and Natural Selection, 18
+
+White, G., 3
+
+Williams, C. M., 217
+
+Wilson, E. B., on cytology, 99
+
+Wolf, 249
+
+Wollaston's, T. V., _Variation of Species_, Foot Note 59
+
+Woltmann, 277
+
+Woolner, 118
+
+Wundt, on language, 207, 208
+
+
+_Xylina vetusta_, 82
+
+
+Yucca, fertilisation of, 78, 79
+
+
+Zeller, E., Foot Note 3
+
+_Zoonomia_, Erasmus Darwin's, 7
+
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+The Project Gutenberg EBook of Evolution in Modern Thought, by 
+Ernst Haeckel and J. Arthur Thomson and August Weismann
+
+This eBook is for the use of anyone anywhere at no cost and with
+almost no restrictions whatsoever.  You may copy it, give it away or
+re-use it under the terms of the Project Gutenberg License included
+with this eBook or online at www.gutenberg.org
+
+
+Title: Evolution in Modern Thought
+
+Author: Ernst Haeckel
+        J. Arthur Thomson
+        August Weismann
+
+Release Date: August 29, 2007 [EBook #22430]
+
+Language: English
+
+Character set encoding: ISO-8859-1
+
+*** START OF THIS PROJECT GUTENBERG EBOOK EVOLUTION IN MODERN THOUGHT ***
+
+
+
+
+Produced by Marilynda Fraser-Cunliffe, Sankar Viswanathan,
+and the Online Distributed Proofreading Team at
+http://www.pgdp.net
+
+
+
+
+
+
+
+
+
+                         EVOLUTION IN MODERN
+
+                               THOUGHT
+
+
+
+                    BY HAECKEL, THOMSON, WEISMANN
+
+                              AND OTHERS
+
+
+
+
+
+                          THE MODERN LIBRARY
+
+                      PUBLISHERS :: :: NEW YORK
+
+       *       *       *       *       *
+
+
+
+
+CONTENTS
+
+
+I    DARWIN'S PREDECESSORS
+
+     J. Arthur Thomson, Professor of Natural History in the University of
+       Aberdeen
+
+
+II   _The Selection Theory_
+
+     August Weismann, Professor of Zoology in the University of
+       Freiburg (Baden)
+
+
+III  HEREDITY AND VARIATION IN MODERN LIGHTS
+
+     W. Bateson, Professor of Biology in the University of Cambridge
+
+
+IV   "THE DESCENT OF MAN"
+
+     G. Schwalbe, Professor of Anatomy in the University of Strassburg
+
+
+V    CHARLES DARWIN AS AN ANTHROPOLOGIST
+
+     Ernst Haeckel, Professor of Zoology in the University of Jena
+
+
+VI   MENTAL FACTORS IN EVOLUTION
+
+     C. Lloyd Morgan, Professor of Psychology at University College,
+       Bristol
+
+
+VII  THE INFLUENCE OF THE CONCEPTION OF EVOLUTION ON MODERN PHILOSOPHY
+
+       H. Höffding, Professor of Philosophy in the University of Copenhagen
+
+
+VIII THE INFLUENCE OF DARWIN UPON RELIGIOUS THOUGHT
+
+     Rev. P. H. Waggett
+
+
+IX   DARWINISM AND HISTORY
+
+     J. B. Bury, Regious Professor of Modern History in the University
+       of Cambridge
+
+
+X    DARWINISM AND SOCIOLOGY
+
+     C. Bouglé, Professor of Social Philosophy in the University of
+       Toulouse, and Deputy-Professor at the Sorbonne, Paris
+
+       *       *       *       *       *
+
+
+
+EVOLUTION IN MODERN THOUGHT
+
+
+I
+
+DARWIN'S PREDECESSORS
+
+BY J. ARTHUR THOMSON
+
+_Professor of Natural History in the University of Aberdeen_
+
+
+In seeking to discover Darwin's relation to his predecessors it is
+useful to distinguish the various services which he rendered to the
+theory of organic evolution.
+
+(I) As everyone knows, the general idea of the Doctrine of Descent is
+that the plants and animals of the present day are the lineal
+descendants of ancestors on the whole somewhat simpler, that these
+again are descended from yet simpler forms, and so on backwards
+towards the literal "Protozoa" and "Protophyta" about which we
+unfortunately know nothing. Now no one supposes that Darwin originated
+this idea, which in rudiment at least is as old as Aristotle. What
+Darwin did was to make it current intellectual coin. He gave it a form
+that commended itself to the scientific and public intelligence of the
+day, and he won widespread conviction by showing with consummate skill
+that it was an effective formula to work with, a key which no lock
+refused. In a scholarly, critical, and pre-eminently fair-minded way,
+admitting difficulties and removing them, foreseeing objections and
+forestalling them, he showed that the doctrine of descent supplied a
+modal interpretation of how our present-day fauna and flora have come
+to be.
+
+(II) In the second place, Darwin applied the evolution-idea to
+particular problems, such as the descent of man, and showed what a
+powerful organon it is, introducing order into masses of uncorrelated
+facts, interpreting enigmas both of structure and function, both
+bodily and mental, and, best of all, stimulating and guiding further
+investigation. But here again it cannot be claimed that Darwin was
+original. The problem of the descent or ascent of man, and other
+particular cases of evolution, had attracted not a few naturalists
+before Darwin's day, though no one [except Herbert Spencer in the
+psychological domain (1855)] had come near him in precision and
+thoroughness of inquiry.
+
+(III) In the third place, Darwin contributed largely to a knowledge of
+the factors in the evolution-process, especially by his analysis of
+what occurs in the case of domestic animals and cultivated plants, and
+by his elaboration of the theory of Natural Selection which Alfred
+Russel Wallace independently stated at the same time, and of which
+there had been a few previous suggestions of a more or less vague
+description. It was here that Darwin's originality was greatest, for
+he revealed to naturalists the many different forms--often very
+subtle--which natural selection takes, and with the insight of a
+disciplined scientific imagination he realised what a mighty engine of
+progress it has been and is.
+
+(IV) As an epoch-marking contribution, not only to Ætiology but to
+Natural History in the widest sense, we rank the picture which Darwin
+gave to the world of the web of life, that is to say, of the
+inter-relations and linkages in Nature. For the Biology of the
+individual--if that be not a contradiction in terms--no idea is more
+fundamental than that of the correlation of organs, but Darwin's most
+characteristic contribution was not less fundamental,--it was the idea
+of the correlation of organisms. This, again, was not novel; we find
+it in the works of naturalists like Christian Conrad Sprengel,
+Gilbert White, and Alexander von Humboldt, but the realisation of its
+full import was distinctly Darwinian.
+
+
+_As Regards the General Idea of Organic Evolution_
+
+While it is true, as Prof. H. F. Osborn puts it, that "'Before and
+after Darwin' will always be the _ante et post urbem conditam_ of
+biological history," it is also true that the general idea of organic
+evolution is very ancient. In his admirable sketch _From the Greeks to
+Darwin_,[1] Prof. Osborn has shown that several of the ancient
+philosophers looked upon Nature as a gradual development and as still
+in process of change. In the suggestions of Empedocles, to take the
+best instance, there were "four sparks of truth,--first, that the
+development of life was a gradual process; second, that plants were
+evolved before animals; third, that imperfect forms were gradually
+replaced (not succeeded) by perfect forms; fourth, that the natural
+cause of the production of perfect forms was the extinction of the
+imperfect."[2] But the fundamental idea of one stage giving origin to
+another was absent. As the blue Ægean teemed with treasures of beauty
+and threw many upon its shores, so did Nature produce like a fertile
+artist what had to be rejected as well as what was able to survive,
+but the idea of one species emerging out of another was not yet
+conceived.
+
+Aristotle's views of Nature[3] seem to have been more definitely
+evolutionist than those of his predecessors, in this sense, at least,
+that he recognised not only an ascending scale, but a genetic series
+from polyp to man and an age-long movement towards perfection. "It is
+due to the resistance of matter to form that Nature can only rise by
+degrees from lower to higher types." "Nature produces those things
+which, being continually moved by a certain principle contained in
+themselves, arrive at a certain end."
+
+To discern the outcrop of evolution-doctrine in the long interval
+between Aristotle and Bacon seems to be very difficult, and some of
+the instances that have been cited strike one as forced. Epicurus and
+Lucretius, often called poets of evolution, both pictured animals as
+arising directly out of the earth, very much as Milton's lion long
+afterwards pawed its way out. Even when we come to Bruno who wrote
+that "to the sound of the harp of the Universal Apollo (the World
+Spirit), the lower organisms are called by stages to higher, and the
+lower stages are connected by intermediate forms with the higher,"
+there is great room, as Prof. Osborn points out,[4] for difference of
+opinion as to how far he was an evolutionist in our sense of the term.
+
+The awakening of natural science in the sixteenth century brought the
+possibility of a concrete evolution theory nearer, and in the early
+seventeenth century we find evidences of a new spirit--in the
+embryology of Harvey and the classifications of Ray. Besides sober
+naturalists there were speculative dreamers in the sixteenth and
+seventeenth centuries who had at least got beyond static formulae,
+but, as Professor Osborn points out,[5] "it is a very striking fact,
+that the basis of our modern methods of studying the Evolution problem
+was established not by the early naturalists nor by the speculative
+writers, but by the Philosophers." He refers to Bacon, Descartes,
+Leibnitz, Hume, Kant, Lessing, Herder, and Schelling. "They alone were
+upon the main track of modern thought. It is evident that they were
+groping in the dark for a working theory of the Evolution of life, and
+it is remarkable that they clearly perceived from the outset that the
+point to which observation should be directed was not the past but the
+present mutability of species, and further, that this mutability was
+simply the variation of individuals on an extended scale."
+
+Bacon seems to have been one of the first to think definitely about
+the mutability of species, and he was far ahead of his age in his
+suggestion of what we now call a Station of Experimental Evolution.
+Leibnitz discusses in so many words how the species of animals may be
+changed and how intermediate species may once have linked those that
+now seem discontinuous. "All natural orders of beings present but a
+single chain".... "All advances by degrees in Nature, and nothing by
+leaps." Similar evolutionist statements are to be found in the works
+of the other "philosophers," to whom Prof. Osborn refers, who were,
+indeed, more scientific than the naturalists of their day. It must be
+borne in mind that the general idea of organic evolution--that the
+present is the child of the past--is in great part just the idea of
+human history projected upon the natural world, differentiated by the
+qualification that the continuous "Becoming" has been wrought out by
+forces inherent in the organisms themselves and in their environment.
+
+A reference to Kant[6] should come in historical order after Buffon,
+with whose writings he was acquainted, but he seems, along with Herder
+and Schelling, to be best regarded as the culmination of the
+evolutionist philosophers--of those at least who interested themselves
+in scientific problems. In a famous passage he speaks of "the
+agreement of so many kinds of animals in a certain common plan of
+structure" ... an "analogy of forms" which "strengthens the
+supposition that they have an actual blood-relationship, due to
+derivation from a common parent." He speaks of "the great Family of
+creatures, for as a Family we must conceive it, if the above-mentioned
+continuous and connected relationship has a real foundation." Prof.
+Osborn alludes to the scientific caution which led Kant, biology being
+what it was, to refuse to entertain the hope "that a Newton may one
+day arise even to make the production of a blade of grass
+comprehensible, according to natural laws ordained by no intention."
+As Prof. Haeckel finely observes, Darwin rose up as Kant's Newton.[7]
+
+The scientific renaissance brought a wealth of fresh impressions and
+some freedom from the tyranny of tradition, and the twofold stimulus
+stirred the speculative activity of a great variety of men from old
+Claude Duret of Moulins, of whose weird transformism (1609) Dr. Henry
+de Varigny[8] gives us a glimpse, to Lorenz Oken (1779-1851) whose
+writings are such mixtures of sense and nonsense that some regard him
+as a far-seeing prophet and others as a fatuous follower of
+intellectual will-o'-the-wisps. Similarly, for De Maillet, Maupertuis,
+Diderot, Bonnet, and others, we must agree with Professor Osborn that
+they were not actually in the main Evolution movement. Some have been
+included in the roll of honour on very slender evidence, Robinet for
+instance, whose evolutionism seems to us extremely dubious.[9]
+
+The first naturalist to give a broad and concrete expression to the
+evolutionist doctrine of descent was Buffon (1707-1788), but it is
+interesting to recall the fact that his contemporary Linnæus
+(1707-1778), protagonist of the counter-doctrine of the fixity of
+species,[10] went the length of admitting (in 1762) that new species
+might arise by inter-crossing. Buffon's position among the pioneers of
+the evolution-doctrine is weakened by his habit of vacillating between
+his own conclusions and the orthodoxy of the Sorbonne, but there is no
+doubt that he had firm grasp of the general idea of "l'enchaînment des
+êtres."
+
+Erasmus Darwin (1731-1802), probably influenced by Buffon, was another
+firm evolutionist, and the outline of his argument in the
+_Zoonomia_[11] might serve in part at least to-day. "When we revolve
+in our minds the metamorphoses of animals, as from the tadpole to the
+frog; secondly, the changes produced by artificial cultivation, as in
+the breeds of horses, dogs, and sheep; thirdly, the changes produced
+by conditions of climate and of season, as in the sheep of warm
+climates being covered with hair instead of wool, and the hares and
+partridges of northern climates becoming white in winter: when,
+further, we observe the changes of structure produced by habit, as
+seen especially in men of different occupations; or the changes
+produced by artificial mutilation and prenatal influences, as in the
+crossing of species and production of monsters; fourth, when we
+observe the essential unity of plan in all warm-blooded animals,--we
+are led to conclude that they have been alike produced from a similar
+living filament".... "From thus meditating upon the minute portion of
+time in which many of the above changes have been produced, would it
+be too bold to imagine, in the great length of time since the earth
+began to exist, perhaps millions of years before the commencement of
+the history of mankind, that all warm-blooded animals have arisen from
+one living filament?"... "This idea of the gradual generation of all
+things seems to have been as familiar to the ancient philosophers as
+to the modern ones, and to have given rise to the beautiful
+hieroglyphic figure of the [Greek: prôton ôon], or first great egg,
+produced by night, that is, whose origin is involved in obscurity, and
+animated by [Greek: Erôs], that is, by Divine Love; from whence
+proceeded all things which exist."
+
+Lamarck (1744-1829) seems to have become an evolutionist
+independently of Erasmus Darwin's influence, though the parallelism
+between them is striking. He probably owed something to Buffon, but he
+developed his theory along a different line. Whatever view be held in
+regard to that theory there is no doubt that Lamarck was a
+thorough-going evolutionist. Professor Haeckel speaks of the
+_Philosophie Zoologique_ as "the first connected and thoroughly
+logical exposition of the theory of descent."[12]
+
+Besides the three old masters, as we may call them, Buffon, Erasmus
+Darwin, and Lamarck, there were other quite convinced pre-Darwinian
+evolutionists. The historian of the theory of descent must take
+account of Treviranus whose _Biology or Philosophy of Animate Nature_
+is full of evolutionary suggestions; of Etienne Geoffroy St. Hilaire,
+who in 1830, before the French Academy of Sciences, fought with
+Cuvier, the fellow-worker of his youth, an intellectual duel on the
+question of descent; of Goethe, one of the founders of morphology and
+the greatest poet of Evolution--who, in his eighty-first year, heard
+the tidings of Geoffrey St. Hilaire's defeat with an interest which
+transcended the political anxieties of the time; and of many others
+who had gained with more or less confidence and clearness a new
+outlook on Nature. It will be remembered that Darwin refers to
+thirty-four more or less evolutionist authors in his Historical
+Sketch, and the list might be added to. Especially when we come near
+to 1858 do the numbers increase, and one of the most remarkable, as
+also most independent champions of the evolution-idea before that date
+was Herbert Spencer, who not only marshalled the arguments in a very
+forcible way in 1852, but applied the formula in detail in his
+_Principles of Psychology_ in 1855.[13]
+
+It is right and proper that we should shake ourselves free from all
+creationist appreciations of Darwin, and that we should recognise the
+services of pre-Darwinian evolutionists who helped to make the time
+ripe, yet one cannot help feeling that the citation of them is apt to
+suggest two fallacies. It may suggest that Darwin simply entered into
+the labours of his predecessors, whereas, as a matter of fact, he knew
+very little about them till after he had been for years at work. To
+write, as Samuel Butler did, "Buffon planted, Erasmus Darwin and
+Lamarck watered, but it was Mr. Darwin who said 'That fruit is ripe,'
+and shook it into his lap" ... seems to us a quite misleading version
+of the facts of the case. The second fallacy which the historical
+citation is a little apt to suggest is that the filiation of ideas is
+a simple problem. On the contrary, the history of an idea, like the
+pedigree of an organism, is often very intricate, and the evolution of
+the evolution-idea is bound up with the whole progress of the world.
+Thus in order to interpret Darwin's clear formulation of the idea of
+organic evolution and his convincing presentation of it, we have to do
+more than go back to his immediate predecessors, such as Buffon,
+Erasmus Darwin, and Lamarck; we have to inquire into the acceptance of
+evolutionary conceptions in regard to other orders of facts, such as
+the earth and the solar system;[14] we have to realise how the growing
+success of scientific interpretation along other lines gave confidence
+to those who refused to admit that there was any domain from which
+science could be excluded as a trespasser; we have to take account of
+the development of philosophical thought, and even of theological and
+religious movements; we should also, if we are wise enough, consider
+social changes. In short, we must abandon the idea that we can
+understand the history of any science as such, without reference to
+contemporary evolution in other departments of activity.
+
+While there were many evolutionists before Darwin, few of them were
+expert naturalists and few were known outside a small circle; what was
+of much more importance was that the genetic view of Nature was
+insinuating itself in regard to other than biological orders of facts,
+here a little and there a little, and that the scientific spirit had
+ripened since the days when Cuvier laughed Lamarck out of court. How
+was it that Darwin succeeded where others had failed? Because, in the
+first place, he had clear visions--"pensées de la jeunesse, executées
+par l'âge mûr"--which a University curriculum had not made impossible,
+which the _Beagle voyage_ made vivid, which an unrivalled British
+doggedness made real--visions of the web of life, of the fountain of
+change within the organism, of the struggle for existence and its
+winnowing, and of the spreading genealogical tree. Because, in the
+second place, he put so much grit into the verification of his
+visions, putting them to the proof in an argument which is of its
+kind--direct demonstration being out of the question--quite
+unequalled. Because, in the third place, he broke down the opposition
+which the most scientific had felt to the seductive modal formula of
+evolution by bringing forward a more plausible theory of the process
+than had been previously suggested. Nor can one forget, since
+questions of this magnitude are human and not merely academic, that he
+wrote so that all men could understand.
+
+
+_As Regards the Factors of Evolution_
+
+It is admitted by all who are acquainted with the history of biology
+that the general idea of organic evolution as expressed in the
+Doctrine of Descent was quite familiar to Darwin's grandfather and to
+others before and after him, as we have briefly indicated. It must
+also be admitted that some of these pioneers of evolutionism did more
+than apply the evolution-idea as a modal formula of becoming, they
+began to inquire into the factors in the process. Thus there were
+pre-Darwinian theories of evolution, and to these we must now briefly
+refer.[15]
+
+In all biological thinking we have to work with the categories
+Organism--Function--Environment, and theories of evolution may be
+classified in relation to these. To some it has always seemed that the
+fundamental fact is the living organism,--a creative agent, a striving
+will, a changeful Proteus, selecting its environment, adjusting itself
+to it, self-differentiating and self-adaptive. The necessity of
+recognising the importance of the organism is admitted by all
+Darwinians who start with inborn variations, but it is open to
+question whether the whole truth of what we might call the Goethian
+position is exhausted in the postulate of inherent variability.
+
+To others it has always seemed that the emphasis should be laid on
+Function,--on use and disuse, on doing and not doing. Practice makes
+perfect; _c'est à force de forger qu'on devient forgeron_. This is one
+of the fundamental ideas of Lamarckism; to some extent it met with
+Darwin's approval; and it finds many supporters to-day. One of the
+ablest of these--Mr. Francis Darwin--has recently given strong reasons
+for combining a modernised Lamarckism with what we usually regard as
+sound Darwinism.[16]
+
+To others it has always seemed that the emphasis should be laid on the
+Environment, which wakes the organism to action, prompts it to change,
+makes dints upon it, moulds it, prunes it, and finally, perhaps, kills it.
+It is again impossible to doubt that there is truth in this view, for even
+if environmentally induced "modifications" be not transmissible,
+environmentally induced "variations" are; and even if the direct influence
+of the environment be less important than many enthusiastic supporters of
+this view--may we call them Buffonians--think, there remains the indirect
+influence which Darwinians in part rely on,--the eliminative process. Even
+if the extreme view be held that the only form of discriminate elimination
+that counts is inter-organismal competition, this might be included under
+the rubric of the animate environment.
+
+In many passages Buffon[17] definitely suggested that environmental
+influences--especially of climate and food--were directly productive
+of changes in organisms, but he did not discuss the question of the
+transmissibility of the modifications so induced, and it is difficult
+to gather from his inconsistent writings what extent of transformation
+he really believed in. Prof. Osborn says of Buffon: "The struggle for
+existence, the elimination of the least-perfected species, the contest
+between the fecundity of certain species and their constant
+destruction, are all clearly expressed in various passages." He quotes
+two of these:[18]
+
+"Le cours ordinaire de la nature vivante, est en général toujours
+constant, toujours le même; son mouvement, toujours régulier, roule
+sur deux points inébranlables: l'un, la fécondité sans bornes donnée à
+toutes les espèces; l'autre, les obstacles sans nombre qui réduisent
+cette fécondité à une mesure déterminée et ne laissent en tout temps
+qu'à peu près la même quantité d'individus de chaque espèce" ... "Les
+espèces les moins parfaites, les plus délicates, les plus pesantes,
+les moins agissantes, les moins armées, etc., ont déjà disparu ou
+disparaîtront.".
+
+Erasmus Darwin[19] had a firm grip of the "idea of the gradual
+formation and improvement of the Animal world," and he had his theory
+of the process. No sentence is more characteristic than this: "All
+animals undergo transformations which are in part produced by their
+own exertions, in response to pleasures and pains, and many of these
+acquired forms or propensities are transmitted to their posterity."
+This is Lamarckism before Lamarck, as his grandson pointed out. His
+central idea is that wants stimulate efforts and that these result in
+improvements which subsequent generations make better still. He
+realised something of the struggle for existence and even pointed out
+that this advantageously checks the rapid multiplication. "As Dr.
+Krause points out, Darwin just misses the connection between this
+struggle and the Survival of the Fittest."[20]
+
+Lamarck[21] (1744-1829) seems to have thought out his theory of
+evolution without any knowledge of Erasmus Darwin's which it closely
+resembled. The central idea of his theory was the cumulative
+inheritance of functional modifications. "Changes in environment bring
+about changes in the habits of animals. Changes in their wants
+necessarily bring about parallel changes in their habits. If new wants
+become constant or very lasting, they form new habits, the new habits
+involve the use of new parts, or a different use of old parts, which
+results finally in the production of new organs and the modification
+of old ones." He differed from Buffon in not attaching importance, as
+far as animals are concerned, to the direct influence of the
+environment, "for environment can effect no direct change whatever
+upon the organisation of animals," but in regard to plants he agreed
+with Buffon that external conditions directly moulded them.
+
+Treviranus[22] (1776-1837), whom Huxley ranked beside Lamarck, was on
+the whole Buffonian, attaching chief importance to the influence of a
+changeful environment both in modifying and in eliminating, but he was
+also Goethian, for instance in his idea that species like individuals
+pass through periods of growth, full bloom, and decline. "Thus, it is
+not only the great catastrophes of Nature which have caused
+extinction, but the completion of cycles of existence, out of which
+new cycles have begun." A characteristic sentence is quoted by Prof.
+Osborn: "In every living being there exists a capability of an endless
+variety of form-assumption; each possesses the power to adapt its
+organisation to the changes of the outer world, and it is this power,
+put into action by the change of the universe, that has raised the
+simple zoophytes of the primitive world to continually higher stages
+of organisation, and has introduced a countless variety of species
+into animate Nature."
+
+Goethe[23] (1749-1832), who knew Buffon's work but not Lamarck's, is
+peculiarly interesting as one of the first to use the evolution-idea
+as a guiding hypothesis, e.g. in the interpretation of vestigial
+structures in man, and to realise that organisms express an attempt to
+make a compromise between specific inertia and individual change. He
+gave the finest expression that science has yet known--if it has known
+it--of the kernel-idea of what is called "bathmism," the idea of an
+"inherent growth-force"--and at the same time he held that "the way of
+life powerfully reacts upon all form" and that the orderly growth of
+form "yields to change from externally acting causes."
+
+Besides Buffon, Erasmus Darwin, Lamarck, Treviranus, and Goethe,
+there were other "pioneers of evolution," whose views have been often
+discussed and appraised. Étienne Geoffroy Saint-Hilaire (1772-1884),
+whose work Goethe so much admired, was on the whole Buffonian,
+emphasising the direct action of the changeful _milieu_. "Species vary
+with their environment, and existing species have descended by
+modification from earlier and somewhat simpler species." He had a
+glimpse of the selection idea, and believed in mutations or sudden
+leaps--induced in the embryonic condition by external influences. The
+complete history of evolution-theories will include many instances of
+guesses at truth which were afterwards substantiated, thus the
+geographer von Buch (1773-1853) detected the importance of the
+Isolation factor on which Wagner, Romanes, Gulick and others have laid
+great stress, but we must content ourselves with recalling one other
+pioneer, the author of the _Vestiges of Creation_ (1844), a work which
+passed through ten editions in nine years and certainly helped to
+harrow the soil for Darwin's sowing. As Darwin said, "it did excellent
+service in this country in calling attention to the subject, in
+removing prejudice, and in thus preparing the ground for the reception
+of analogous views."[24] Its author, Robert Chambers (1802-1871) was
+in part a Buffonian--maintaining that environment moulded organisms
+adaptively, and in part a Goethian--believing in an inherent
+progressive impulse which lifted organisms from one grade of
+organisation to another.
+
+
+_As Regards Natural Selection_
+
+The only thinker to whom Darwin was directly indebted, so far as the
+theory of Natural Selection is concerned, was Malthus, and we may once
+more quote the well-known passage in the Autobiography: "In October,
+1838, that is, fifteen months after I had begun my systematic enquiry,
+I happened to read for amusement 'Malthus on Population,' and being
+well prepared to appreciate the struggle for existence which
+everywhere goes on from long-continued observation of the habits of
+animals and plants, it at once struck me that under these
+circumstances favourable variations would tend to be preserved, and
+unfavourable ones to be destroyed. The result of this would be the
+formation of new species."[25]
+
+Although Malthus gives no adumbration of the idea of Natural Selection
+in his exposition of the eliminative processes which go on in mankind,
+the suggestive value of his essay is undeniable, as is strikingly
+borne out by the fact that it gave to Alfred Russel Wallace also "the
+long-sought clue to the effective agent in the evolution of organic
+species."[26] One day in Ternate when he was resting between fits of
+fever, something brought to his recollection the work of Malthus which
+he had read twelve years before. "I thought of his clear exposition of
+'the positive checks to increase'--disease, accidents, war, and
+famine--which keep down the population of savage races to so much
+lower an average than that of more civilized peoples. It then occurred
+to me that these causes or their equivalents are continually acting in
+the case of animals also; and as animals usually breed much more
+rapidly than does mankind, the destruction every year from these
+causes must be enormous in order to keep down the numbers of each
+species, since they evidently do not increase regularly from year to
+year, as otherwise the world would long ago have been densely crowded
+with those that breed most quickly. Vaguely thinking over the enormous
+and constant destruction which this implied, it occurred to me to ask
+the question, Why do some die and some live? And the answer was
+clearly, that on the whole the best fitted live. From the effects of
+disease the most healthy escaped; from enemies the strongest, the
+swiftest, or the most cunning; from famine the best hunters or those
+with the best digestion; and so on. Then it suddenly flashed upon me
+that this self-acting process would necessarily _improve the race_,
+because in every generation the inferior would inevitably be killed
+off and the superior would remain--that is, _the fittest would
+survive_."[27] We need not apologise for this long quotation, it is a
+tribute to Darwin's magnanimous colleague, the Nestor of the
+evolutionist camp,--and it probably indicates the line of thought
+which Darwin himself followed. It is interesting also to recall the
+fact that in 1852, when Herbert Spencer wrote his famous _Leader_
+article on "The Development Hypothesis" in which he argued powerfully
+for the thesis that the whole animate world is the result of an
+age-long process of natural transformation, he wrote for _The
+Westminster Review_ another important essay, "A Theory of Population
+deduced from the General Law of Animal Fertility," towards the close
+of which he came within an ace of recognising that the struggle for
+existence was a factor in organic evolution. At a time when pressure
+of population was practically interesting men's minds, Darwin,
+Wallace, and Spencer were being independently led from a social
+problem to a biological theory. There could be no better illustration,
+as Prof. Patrick Geddes has pointed out, of the Comtian thesis that
+science is a "social phenomenon."
+
+Therefore, as far more important than any further ferreting out of
+vague hints of Natural Selection in books which Darwin never read, we
+would indicate by a quotation the view that the central idea in
+Darwinism is correlated with contemporary social evolution. "The
+substitution of Darwin for Paley as the chief interpreter of the order
+of nature is currently regarded as the displacement of an
+anthropomorphic view by a purely scientific one: a little reflection,
+however, will show that what has actually happened has been merely the
+replacement of the anthropomorphism of the eighteenth century by that
+of the nineteenth. For the place vacated by Paley's theological and
+metaphysical explanation has simply been occupied by that suggested to
+Darwin and Wallace by Malthus in terms of the prevalent severity of
+industrial competition, and those phenomena of the struggle for
+existence which the light of contemporary economic theory has enabled
+us to discern, have thus come to be temporarily exalted into a
+complete explanation of organic progress."[28] It goes without saying
+that the idea suggested by Malthus was developed by Darwin into a
+biological theory which was then painstakingly verified by being used
+as an interpretative formula, and that the validity of a theory so
+established is not affected by what suggested it, but the practical
+question which this line of thought raises in the mind is this: if
+Biology did thus borrow with such splendid results from social theory,
+why should we not more deliberately repeat the experiment?
+
+Darwin was characteristically frank and generous in admitting that the
+principle of Natural Selection had been independently recognised by
+Dr. W. C. Wells in 1813 and by Mr. Patrick Matthew in 1831, but he had
+no knowledge of these anticipations when he published the first
+edition of _The Origin of Species_. Wells, whose "Essay on Dew" is
+still remembered, read in 1813 before the Royal Society a short paper
+entitled "An Account of a White Female, part of whose skin resembles
+that of a Negro" (published in 1818). In this communication, as Darwin
+said, "he observes, firstly, that all animals tend to vary in some
+degree, and, secondly, that agriculturists improve their domesticated
+animals by selection; and then, he adds, but what is done in this
+latter case 'by art, seems to be done with equal efficacy, though more
+slowly, by nature, in the formation of varieties of mankind, fitted
+for the country which they inhabit.'"[29] Thus Wells had the clear
+idea of survival dependent upon a favourable variation, but he makes
+no more use of the idea and applies it only to man. There is not in
+the paper the least hint that the author ever thought of generalising
+the remarkable sentence quoted above.
+
+Of Mr. Patrick Matthew, who buried his treasure in an appendix to a
+work on _Naval Timber and Arboriculture_, Darwin said that "he clearly
+saw the full force of the principle of natural selection." In 1860
+Darwin wrote--very characteristically--about this to Lyell: "Mr.
+Patrick Matthew publishes a long extract from his work on _Naval
+Timber and Arboriculture_, published in 1831, in which he briefly but
+completely anticipates the theory of Natural Selection. I have ordered
+the book, as some passages are rather obscure, but it is certainly, I
+think, a complete but not developed anticipation. Erasmus always said
+that surely this would be shown to be the case some day. Anyhow, one
+may be excused in not having discovered the fact in a work on Naval
+Timber."[30]
+
+De Quatrefages and De Varigny have maintained that the botanist Naudin
+stated the theory of evolution by natural selection in 1852. He
+explains very clearly the process of artificial selection, and says
+that in the garden we are following Nature's method. "We do not think
+that Nature has made her species in a different fashion from that in
+which we proceed ourselves in order to make our variations." But, as
+Darwin said, "he does not show how selection acts under nature."
+Similarly it must be noted in regard to several pre-Darwinian pictures
+of the struggle for existence (such as Herder's, who wrote in 1790
+"All is in struggle ... each one for himself" and so on), that a
+recognition of this is only the first step in Darwinism.
+
+Profs. E. Perrier and H. F. Osborn have called attention to a
+remarkable anticipation of the selection-idea which is to be found in
+the speculations of Étienne Geoffroy Saint-Hilaire (1825-1828) on the
+evolution of modern Crocodilians from the ancient Teleosaurs. Changing
+environment induced changes in the respiratory system and far-reaching
+consequences followed. The atmosphere, acting upon the pulmonary
+cells, brings about "modifications which are favourable or destructive
+('funestes'); these are inherited, and they influence all the rest of
+the organisation of the animal because if these modifications lead to
+injurious effects the animals which exhibit them perish and are
+replaced by others of a somewhat different form, a form changed so as
+to be adapted to (à la convenance) the new environment."
+
+Prof. E. B. Poulton[31] has shown that the anthropologist James Cowles
+Prichard (1786-1848) must be included even in spite of himself among
+the precursors of Darwin. In some passages of the second edition of
+his _Researches into the Physical History of Mankind_ (1826), he
+certainly talks evolution and anticipates Prof. Weismann in denying
+the transmission of acquired characters. He is, however, sadly
+self-contradictory and his evolutionism weakens in subsequent
+editions--the only ones that Darwin saw. Prof. Poulton finds in
+Prichard's work a recognition of the operation of Natural Selection.
+"After inquiring how it is that 'these varieties are developed and
+preserved in connexion with particular climates and differences of
+local situation,' he gives the following very significant answer: 'One
+cause which tends to maintain this relation is obvious. Individuals
+and families, and even whole colonies perish and disappear in climates
+for which they are, by peculiarity of constitution, not adapted. Of
+this fact proofs have been already mentioned.'" Mr. Francis Darwin and
+Prof. A. C. Seward discuss Prichard's "anticipations" in _More Letters
+of Charles Darwin_, Vol. _I._ p. 43, and come to the conclusion that
+the evolutionary passages are entirely neutralised by others of an
+opposite trend. There is the same difficulty with Buffon.
+
+Hints of the idea of Natural Selection have been detected elsewhere.
+James Watt,[32] for instance, has been reported as one of the
+anticipators (1851). But we need not prolong the inquiry further,
+since Darwin did not know of any anticipations until after he had
+published the immortal work of 1859, and since none of those who got
+hold of the idea made any use of it. What Darwin did was to follow the
+clue which Malthus gave him, to realise, first by genius and
+afterwards by patience, how the complex and subtle struggle for
+existence works out a natural selection of those organisms which vary
+in the direction of fitter adaptation to the conditions of their life.
+So much success attended his application of the Selection-formula that
+for a time he regarded Natural Selection as almost the sole factor in
+evolution, variations being pre-supposed; gradually, however, he came
+to recognise that there was some validity in the factors which had
+been emphasised by Lamarck and by Buffon, and in his well known
+summing up in the sixth edition of the _Origin_ he says of the
+transformation of species: "This has been effected chiefly through the
+natural selection of numerous successive, slight, favourable
+variations; aided in an important manner by the inherited effects of
+the use and disuse of parts; and in an unimportant manner, that is, in
+relation to adaptive structures, whether past or present, by the
+direct action of external conditions, and by variations which seem to
+us in our ignorance to arise spontaneously."
+
+To sum up: the idea of organic evolution, older than Aristotle, slowly
+developed from the stage of suggestion to the stage of verification,
+and the first convincing verification was Darwin's; from being an _a
+priori_ anticipation it has become an interpretation of nature, and
+Darwin is still the chief interpreter; from being a modal
+interpretation it has advanced to the rank of a causal theory, the
+most convincing part of which men will never cease to call Darwinism.
+
+FOOTNOTES:
+
+[Footnote 1: _Columbia University Biological Series_, Vol. I. New York
+and London, 1894. We must acknowledge our great indebtedness to this
+fine piece of work.]
+
+[Footnote 2: _op. cit._ p. 41.]
+
+[Footnote 3: See G. J. Romanes, "Aristotle as a Naturalist,"
+_Contemporary Review_, Vol. lix. p. 275, 1891; G. Pouchet, _La
+Biologie Aristotélique_, Paris, 1885; E. Zeller, _A History of Greek
+Philosophy_, London, 1881, and "Ueber die griechischen Vorgänger
+Darwin's," _Abhandl. Berlin Akad._ 1878, pp. 111-124.]
+
+[Footnote 4: _op. cit._ p. 81.]
+
+[Footnote 5: _op. cit._ p. 87.]
+
+[Footnote 6: See Brock, "Die Stellung Kant's zur Deszendenztheorie,"
+_Biol. Centralbl._ viii. 1889, pp. 641-648. Fritz Schultze, _Kant und
+Darwin_, Jena, 1875.]
+
+[Footnote 7: Mr. Alfred Russel Wallace writes: "We claim for Darwin
+that he is the Newton of natural history, and that, just so surely as
+that the discovery and demonstration by Newton of the law of
+gravitation established order in place of chaos and laid a sure
+foundation for all future study of the starry heavens, so surely has
+Darwin, by his discovery of the law of natural selection and his
+demonstration of the great principle of the preservation of useful
+variations in the struggle for life, not only thrown a flood of light
+on the process of development of the whole organic world, but also
+established a firm foundation for all future study of nature"
+(_Darwinism_, London, 1889, p. 9). See also Prof. Karl Pearson's
+_Grammar of Science_ (2nd edit.), London, 1900, p. 32. See Osborn,
+_op. cit._ p. 100.]
+
+[Footnote 8: _Experimental Evolution_. London, 1892. Chap. I. p. 14.]
+
+[Footnote 9: See J. Arthur Thomson, _The Science of Life_. London,
+1899, Chap. XVI. "Evolution of Evolution Theory."]
+
+[Footnote 10: See Carus Sterne (Ernst Krause), _Die allgemeine
+Weltanschauung in ihrer historischen Entwickelung_. Stuttgart, 1889.
+Chapter entitled "Beständigkeit oder Veränderlichkeit der
+Naturwesen."]
+
+[Footnote 11: _Zoonomia, or the Laws of Organic Life_, 2 vols. London,
+1794; Osborn, _op. cit._ p. 145.]
+
+[Footnote 12: See Alpheus S. Packard, _Lamarck, the Founder of
+Evolution, His Life and Work, with Translations of his writings on
+Organic Evolution_. London, 1901.]
+
+[Footnote 13: See Edward Clodd, _Pioneers of Evolution_, London, p.
+161, 1897.]
+
+[Footnote 14: See Chapter ix. "The Genetic View of Nature" in J. T.
+Merz's _History of European Thought in the Nineteenth Century_, Vol.
+2, Edinburgh and London, 1903.]
+
+[Footnote 15: See Prof. W. A. Locy's _Biology and its Makers_. New
+York, 1908. Part II. "The Doctrine of Organic Evolution."]
+
+[Footnote 16: Presidential Address to the British Association meeting
+at Dublin in 1908.]
+
+[Footnote 17: See in particular Samuel Butler, _Evolution Old and
+New_, London, 1879; J. L. de Lanessan, "Buffon et Darwin," _Revue
+Scientifique_, XLIII. pp. 385-391, 425-432, 1889.]
+
+[Footnote 18: _op. cit._ p. 136.]
+
+[Footnote 19: See Ernest Krause and Charles Darwin, _Erasmus Darwin_,
+London, 1879.]
+
+[Footnote 20: Osborn, _op. cit._ p. 142.]
+
+[Footnote 21: See E. Perrier, _La Philosophie Zoologique avant
+Darwin_, Paris, 1884; A. de Quatrefages, _Darwin et ses Précurseurs
+Français_, Paris, 1870; Packard, _op. cit._; also Claus, _Lamarck als
+Begründer der Descendenzlehre_, Wien, 1888; Haeckel, _Natural History
+of Creation_, Eng. transl. London, 1879; Lang, _Zur Charakteristik der
+Forschungswege von Lamarck und Darwin_, Jena, 1889.]
+
+[Footnote 22: See Huxley's article "Evolution in Biology,"
+_Encyclopaedia Britannica_ (9th edit.), 1879, pp. 744-751, and Sully's
+article, "Evolution in Philosophy," _ibid._ pp. 751-772.]
+
+[Footnote 23: See Haeckel, _Die Naturanschauung von Darwin, Goethe und
+Lamarck_, Jena, 1882.]
+
+[Footnote 24: _Origin of Species_ (6th edit.), p. xvii.]
+
+[Footnote 25: _The Life and Letters of Charles Darwin_, Vol. 1. p. 83.
+London, 1887.]
+
+[Footnote 26: A. R. Wallace, _My Life, a Record of Events and
+Opinions_, London, 1905, Vol. 1, p. 232.]
+
+[Footnote 27: _My Life_, Vol. 1. p. 361.]
+
+[Footnote 28: P. Geddes. article "Biology." _Chambers's
+Encyclopaedia._]
+
+[Footnote 29: _Origin of Species_ (6th edit.), p. xv.]
+
+[Footnote 30: _Life and Letters_, II, p. 301.]
+
+[Footnote 31: _Science Progress_, New Series, Vol. 1. 1897. "A
+Remarkable Anticipation of Modern Views on Evolution." See also Chap.
+VI. in _Essays on Evolution_, Oxford, 1908.]
+
+[Footnote 32: See Prof. Patrick Geddes's article "Variation and
+Selection," _Encyclopaedia Britannica_ (9th edit.) 1888.]
+
+
+
+
+II
+
+THE SELECTION THEORY
+
+BY AUGUST WEISMANN
+
+_Professor of Zoology in the University of Freiburg_ (_Baden_)
+
+
+I. THE IDEA OF SELECTION
+
+Many and diverse were the discoveries made by Charles Darwin in the
+course of a long and strenuous life, but none of them has had so
+far-reaching an influence on the science and thought of his time as
+the theory of selection. I do not believe that the theory of evolution
+would have made its way so easily and so quickly after Darwin took up
+the cudgels in favour of it if he had not been able to support it by a
+principle which was capable of solving, in a simple manner, the
+greatest riddle that living nature presents to us,--I mean the
+purposiveness of every living form relative to the conditions of its
+life and its marvellously exact adaptation to these.
+
+Everyone knows that Darwin was not alone in discovering the principle
+of selection, and that the same idea occurred simultaneously and
+independently to Alfred Russel Wallace. At the memorable meeting of
+the Linnean Society on 1st July, 1858, two papers were read
+(communicated by Lyell and Hooker) both setting forth the same idea of
+selection. One was written by Charles Darwin in Kent, the other by
+Alfred Wallace in Ternate, in the Malay Archipelago. It was a splendid
+proof of the magnanimity of these two investigators, that they thus in
+all friendliness and without envy, united in laying their ideas
+before a scientific tribunal: their names will always shine side by
+side as two of the brightest stars in the scientific sky.
+
+The idea of selection set forth by the two naturalists was at the time
+absolutely new, but it was also so simple that Huxley could say of it
+later, "How extremely stupid not to have thought of that." As Darwin
+was led to the general doctrine of descent, not through the labours of
+his predecessors in the early years of the century, but by his own
+observations, so it was in regard to the principle of selection. He
+was struck by the innumerable cases of adaptation, as, for instance,
+that of the woodpeckers and tree-frogs to climbing, or the hooks and
+feather-like appendages of seeds, which aid in the distribution of
+plants, and he said to himself that an explanation of adaptations was
+the first thing to be sought for in attempting to formulate a theory
+of evolution.
+
+But since adaptations point to _changes_ which have been undergone by
+the ancestral forms of existing species, it is necessary, first of
+all, to inquire how far species in general are _variable_. Thus
+Darwin's attention was directed in the first place to the phenomenon
+of variability, and the use man has made of this, from very early
+times, in the breeding of his domesticated animals and cultivated
+plants. He inquired carefully how breeders set to work, when they
+wished to modify the structure and appearance of a species to their
+own ends, and it was soon clear to him that _selection for breeding
+purposes_ played the chief part.
+
+But how was it possible that such processes should occur in free
+nature? Who is here the breeder, making the selection, choosing out
+one individual to bring forth offspring and rejecting others? That was
+the problem that for a long time remained a riddle to him.
+
+Darwin himself relates how illumination suddenly came to him. He had
+been reading, for his own pleasure, Malthus' book on Population, and,
+as he had long known from numerous observations, that every species
+gives rise to many more descendants than ever attain to maturity, and
+that, therefore, the greater number of the descendants of a species
+perish without reproducing, the idea came to him that the decision as
+to which member of a species was to perish and which was to attain to
+maturity and reproduction might not be a matter of chance, but might
+be determined by the constitution of the individuals themselves,
+according as they were more or less fitted for survival. With this
+idea the foundation of the theory of selection was laid.
+
+In _artificial selection_ the breeder chooses out for pairing only
+such individuals as possess the character desired by him in a somewhat
+higher degree than the rest of the race. Some of the descendants
+inherit this character, often in a still higher degree, and if this
+method be pursued throughout several generations, the race is
+transformed in respect of that particular character.
+
+_Natural selection_ depends on the same three factors as _artificial
+selection_: on _variability_, _inheritance_, and _selection for
+breeding_, but this last is here carried out not by a breeder but by
+what Darwin called the "struggle for existence." This last factor is
+one of the special features of the Darwinian conception of nature.
+That there are carnivorous animals which take heavy toll in every
+generation of the progeny of the animals on which they prey, and that
+there are herbivores which decimate the plants in every generation had
+long been known, but it is only since Darwin's time that sufficient
+attention has been paid to the facts that, in addition to this regular
+destruction, there exists between the members of a species a keen
+competition for space and food, which limits multiplication, and that
+numerous individuals of each species perish because of unfavourable
+climatic conditions. The "struggle for existence," which Darwin
+regarded as taking the place of the human breeder in free nature, is
+not a direct struggle between carnivores and their prey, but is the
+assumed competition for survival between individuals _of the same_
+species, of which, on an average, only those survive to reproduce
+which have the greatest power of resistance, while the others, less
+favourably constituted, perish early. This struggle is so keen, that,
+within a limited area, where the conditions of life have long remained
+unchanged, of every species, whatever be the degree of fertility, only
+two, _on an average_, of the descendants of each pair survive; the
+others succumb either to enemies, or to disadvantages of climate, or
+to accident. A high degree of fertility is thus not an indication of
+the special success of a species, but of the numerous dangers that
+have attended its evolution. Of the six young brought forth by a pair
+of elephants in the course of their lives only two survive in a given
+area; similarly, of the millions of eggs which two thread-worms leave
+behind them only two survive. It is thus possible to estimate the
+dangers which threaten a species by its ratio of elimination, or,
+since this cannot be done directly, by its fertility.
+
+Although a great number of the descendants of each generation fall
+victims to accident, among those that remain it is still the greater
+or less fitness of the organism that determines the "selection for
+breeding purposes," and it would be incomprehensible if, in this
+competition, it were not ultimately, that is, on an average, the best
+equipped which survive, in the sense of living long enough to
+reproduce.
+
+Thus the principle of natural selection is _the selection of the best
+for reproduction_, whether the "best" refers to the whole
+constitution, to one or more parts of the organism, or to one or more
+stages of development. Every organ, every part, every character of an
+animal, fertility and intelligence included, must be improved in this
+manner, and be gradually brought up in the course of generations to
+its highest attainable state of perfection. And not only may
+improvement of parts be brought about in this way, but new parts and
+organs may arise, since, through the slow and minute steps of
+individual or "fluctuating" variations, a part may be added here or
+dropped out there, and thus something new is produced.
+
+The principle of selection solved the riddle as to how what was
+purposive could conceivably be brought about without the intervention
+of a directing power, the riddle which animate nature presents to our
+intelligence at every turn, and in face of which the mind of a Kant
+could find no way out, for he regarded a solution of it as not to be
+hoped for. For, even if we were to assume an evolutionary force that
+is continually transforming the most primitive and the simplest forms
+of life into ever higher forms, and the homogeneity of primitive times
+into the infinite variety of the present, we should still be unable to
+infer from this alone how each of the numberless forms adapted to
+particular conditions of life should have appeared _precisely at the
+right moment in the history of the earth_ to which their adaptations
+were appropriate, and precisely at the proper place in which all the
+conditions of life to which they were adapted occurred: the
+humming-birds at the same time as the flowers; the trichina at the
+same time as the pig; the bark-coloured moth at the same time as the
+oak, and the wasp-like moth at the same time as the wasp which
+protects it. Without processes of selection we should be obliged to
+assume a "pre-established harmony" after the famous Leibnitzian model,
+by means of which the clock of the evolution of organisms is so
+regulated as to strike in exact synchronism with that of the history
+of the earth! All forms of life are strictly adapted to the conditions
+of their life, and can persist under these conditions alone.
+
+There must therefore be an intrinsic connection between the conditions
+and the structural adaptations of the organism, and, _since the
+conditions of life cannot be determined by the animal itself, the
+adaptations must be called forth by the conditions_.
+
+The selection theory teaches us how this is conceivable, since it
+enables us to understand that there is a continual production of what
+is non-purposive as well as of what is purposive, but the purposive
+alone survives, while the non-purposive perishes in the very act of
+arising. This is the old wisdom taught long ago by Empedocles.
+
+
+II. THE LAMARCKIAN PRINCIPLE
+
+Lamarck, as is well known, formulated a definite theory of evolution
+at the beginning of the nineteenth century, exactly fifty years before
+the Darwin-Wallace principle of selection was given to the world. This
+brilliant investigator also endeavoured to support his theory by
+demonstrating forces which might have brought about the
+transformations of the organic world in the course of the ages. In
+addition to other factors, he laid special emphasis on the increased
+or diminished use of the parts of the body, assuming that the
+strengthening or weakening which takes place from this cause during
+the individual life, could be handed on to the offspring, and thus
+intensified and raised to the rank of a specific character. Darwin
+also regarded this _Lamarckian principle_, as it is now generally
+called, as a factor in evolution, but he was not fully convinced of
+the transmissibility of acquired characters.
+
+As I have here to deal only with the theory of selection, I need not
+discuss the Lamarckian hypothesis, but I must express my opinion that
+there is room for much doubt as to the coöperation of this principle
+in evolution. Not only is it difficult to imagine how the transmission
+of functional modifications could take place, but, up to the present
+time, notwithstanding the endeavours of many excellent investigators,
+not a single actual proof of such inheritance has been brought
+forward. Semon's experiments on plants are, according to the botanist
+Pfeffer, not to be relied on, and even the recent, beautiful
+experiments made by Dr. Kammerer on salamanders, cannot, as I hope to
+show elsewhere, be regarded as proof, if only because they do not deal
+at all with functional modifications, that is, with modifications
+brought about by use, and it is to these _alone_ that the Lamarckian
+principle refers.
+
+
+III. OBJECTIONS TO THE THEORY OF SELECTION
+
+
+(_a_) _Saltatory evolution_
+
+The Darwinian doctrine of evolution depends essentially on _the
+cumulative augmentation_ of minute variations in the direction of
+utility. But can such minute variations, which are undoubtedly
+continually appearing among the individuals of the same species,
+possess any selection-value; can they determine which individuals are
+to survive, and which are to succumb; can they be increased by natural
+selection till they attain to the highest development of a purposive
+variation?
+
+To many this seems so improbable that they have urged a theory of
+evolution by leaps from species to species. Kölliker, in 1872,
+compared the evolution of species with the processes which we can
+observe in the individual life in cases of alternation of generations.
+But a polyp only gives rise to a medusa because it has itself arisen
+from one, and there can be no question of a medusa ever having arisen
+suddenly and _de novo_ from a polyp-bud, if only because both forms
+are adapted in their structure as a whole, and in every detail to the
+conditions of their life. A sudden origin, in a natural way, of
+numerous adaptations is inconceivable. Even the degeneration of a
+medusoid from a free-swimming animal to a mere brood-sac (gonophore)
+is not sudden and saltatory, but occurs by imperceptible modifications
+throughout hundreds of years, as we can learn from the numerous stages
+of the process of degeneration persisting at the same time in
+different species.
+
+If, then, the degeneration to a simple brood-sac takes place only by
+very slow transitions, each stage of which may last for centuries, how
+could the much more complex _ascending_ evolution possibly have taken
+place by sudden leaps? I regard this argument as capable of further
+extension, for wherever in nature we come upon degeneration, it is
+taking place by minute steps and with a slowness that makes it not
+directly perceptible, and I believe that this in itself justifies us
+in concluding that _the same must be true of ascending_ evolution. But
+in the latter case the goal can seldom be distinctly recognised while
+in cases of degeneration the starting-point of the process can often
+be inferred, because several nearly related species may represent
+different stages.
+
+In recent years Bateson in particular has championed the idea of
+saltatory, or so-called discontinuous evolution, and has collected a
+number of cases in which more or less marked variations have suddenly
+appeared. These are taken for the most part from among domesticated
+animals which have been bred and crossed for a long time, and it is
+hardly to be wondered at that their much mixed and much influenced
+germ-plasm should, under certain conditions, give rise to remarkable
+phenomena, often indeed producing forms which are strongly suggestive
+of monstrosities, and which would undoubtedly not survive in free
+nature, unprotected by man. I should regard such cases as due to an
+intensified germinal selection--though this is to anticipate a
+little--and from this point of view it cannot be denied that they have
+a special interest. But they seem to me to have no significance as far
+as the transformation of species is concerned, if only because of the
+extreme rarity of their occurrence.
+
+There are, however, many variations which have appeared in a sudden
+and saltatory manner, and some of these Darwin pointed out and
+discussed in detail: the copper beech, the weeping trees, the oak with
+"fern-like leaves," certain garden-flowers, etc. But none of them have
+persisted in free nature, or evolved into permanent types.
+
+On the other hand, wherever enduring types have arisen, we find traces
+of a gradual origin by successive stages, even if, at first sight,
+their origin may appear to have been sudden. This is the case with
+_seasonal Dimorphism_, the first known cases of which exhibited marked
+differences between the two generations, the winter and the summer
+brood. Take for instance the much discussed and studied form
+_Vanessa_ (_Araschnia_) _levana-prorsa_. Here the differences between
+the two forms are so great and so apparently disconnected, that one
+might almost believe it to be a sudden mutation, were it not that old
+transition-stages can be called forth by particular temperatures, and
+we know other butterflies, as for instance our Garden Whites, in which
+the differences between the two generations are not nearly so marked;
+indeed, they are so little apparent that they are scarcely likely to
+be noticed except by experts. Thus here again there are small initial
+steps, some of which, indeed, must be regarded as adaptations, such as
+the green-sprinkled or lightly tinted under-surface which gives them a
+deceptive resemblance to parsley or to Cardamine leaves.
+
+Even if saltatory variations do occur, we cannot assume that these
+_have ever led to forms which are capable of survival under the
+conditions of wild life_. Experience has shown that in plants which
+have suddenly varied the power of persistence is diminished.
+Korschinsky attributes to them weaknesses of organisation in general;
+"they bloom late, ripen few of their seeds, and show great
+sensitiveness to cold." These are not the characters which make for
+success in the struggle for existence.
+
+We must briefly refer here to the views--much discussed in the last
+decade--of H. de Vries, who believes that the roots of transformation
+must be sought for in _saltatory variations arising from internal
+causes_, and distinguishes such _mutations_, as he has called them,
+from ordinary individual variations, in that they breed true, that is,
+with strict in-breeding they are handed on pure to the next
+generation. I have elsewhere endeavoured to point out the weaknesses
+of this theory,[33] and I am the less inclined to return to it here
+that it now appears[34] that the far-reaching conclusions drawn by de
+Vries from his observations on the Evening Primrose, _Oenothera
+lamarckiana_, rest upon a very insecure foundation. The plant from
+which de Vries saw numerous "species"--his "mutations"--arise was not,
+as he assumed, a _wild species_ that had been introduced to Europe
+from America, but was probably a hybrid form which was first
+discovered in the Jardin des Plantes in Paris, and which does not
+appear to exist anywhere in America as a wild species.
+
+This gives a severe shock to the "Mutation theory," for the other
+_actually wild_ species with which de Vries experimented showed no
+"mutations" but yielded only negative results.
+
+Thus we come to the conclusion that Darwin[35] was right in regarding
+transformations as taking place by minute steps, which, if useful, are
+augmented in the course of innumerable generations, because their
+possessors more frequently survive in the struggle for existence.
+
+
+(_b_) _Selection-value of the initial steps_
+
+Is it possible that the insignificant deviations which we know as
+"individual variations" can form the beginning of a process of
+selection? Can they decide which is to perish and which to survive? To
+use a phrase of Romanes, can they have _selection-value_?
+
+Darwin himself answered this question, and brought together many
+excellent examples to show that differences, apparently insignificant
+because very small, might be of decisive importance for the life of
+the possessor. But it is by no means enough to bring forward cases of
+this kind, for the question is not merely whether finished adaptations
+have selection-value, but whether the first beginnings of these, and
+whether the small, I might almost say minimal increments, which have
+led up from these beginnings to the perfect adaptation, have also had
+selection-value. To this question even one who, like myself, has been
+for many years a convinced adherent of the theory of selection, can
+only reply: _We must assume so, but we cannot prove it in any case_.
+It is not upon demonstrative evidence that we rely when we champion
+the doctrine of selection as a scientific truth; we base our argument
+on quite other grounds. Undoubtedly there are many apparently
+insignificant features, which can nevertheless be shown to be
+adaptations--for instance, the thickness of the basin-shaped shell of
+the limpets that live among the breakers on the shore. There can be no
+doubt that the thickness of these shells, combined with their flat
+forms, protects the animals from the force of the waves breaking upon
+them,--but how have they become so thick? What proportion of thickness
+was sufficient to decide that of two variants of a limpet one should
+survive, the other be eliminated? We can say nothing more than that we
+infer from the present state of the shell, that it must have varied in
+regard to differences in shell-thickness, and that these differences
+must have had selection-value,--no proof therefore, but an assumption
+which we must show to be convincing.
+
+For a long time the marvellously complex _radiate_ and _lattice-work_
+skeletons of Radiolarians were regarded as a mere outflow of "Nature's
+infinite wealth of form," as an instance of a purely morphological
+character with no biological significance. But recent investigations
+have shown that these, too, have an adaptive significance (Häcker).
+The same thing has been shown by Schütt in regard to the lowly
+unicellular plants, the Peridineae, which abound alike on the surface
+of the ocean and in its depths. It has been shown that the long
+skeletal processes which grow out from these organisms have
+significance not merely as a supporting skeleton, but also as an
+extension of the superficial area, which increases the contact with
+the water-particles, and prevents the floating organisms from sinking.
+It has been established that the processes are considerably shorter in
+the colder layers of the ocean, and that they may be twelve times as
+long[36] in the warmer layers, thus corresponding to the greater or
+smaller amount of friction which takes place in the denser and less
+dense layers of the water.
+
+The Peridineae of the warmer ocean layers have thus become long-rayed,
+those of the colder layers short-rayed, not through the direct effect
+of friction on the protoplasm, but through processes of selection,
+which favoured the longer rays in warm water, since they kept the
+organism afloat, while those with short rays sank and were eliminated.
+If we put the question as to selection-value in this case, and ask how
+great the variations in the length of processes must be in order to
+possess selection-value; what can we answer except that these
+variations must have been minimal, and yet sufficient to prevent too
+rapid sinking and consequent elimination? Yet this very case would
+give the ideal opportunity for a mathematical calculation of the
+minimal selection-value, although of course it is not feasible from
+lack of data to carry out the actual calculation.
+
+But even in organisms of more than microscopic size there must
+frequently be minute, even microscopic differences which set going the
+process of selection, and regulate its progress to the highest
+possible perfection.
+
+Many tropical trees possess thick, leathery leaves, as a protection
+against the force of the tropical raindrops. The _direct_ influence of
+the rain cannot be the cause of this power of resistance, for the
+leaves, while they were still thin, would simply have been torn to
+pieces. Their toughness must therefore be referred to selection, which
+would favour the trees with slightly thicker leaves, though we cannot
+calculate with any exactness how great the first stages of increase in
+thickness must have been. Our hypothesis receives further support from
+the fact that, in many such trees, the leaves are drawn out into a
+beak-like prolongation (Stahl and Haberlandt) which facilitates the
+rapid falling off of the rain water, and also from the fact that the
+leaves, while they are still young, hang limply down in bunches which
+offer the least possible resistance to the rain. Thus there are here
+three adaptations which can only be interpreted as due to selection.
+The initial stages of these adaptations must undoubtedly have had
+selection-value.
+
+But even in regard to this case we are reasoning in a circle, not
+giving "proofs," and no one who does not wish to believe in the
+selection-value of the initial stages can be forced to do so. Among
+the many pieces of presumptive evidence a particularly weighty one
+seems to me to be _the smallness of the steps of progress_ which we
+can observe in certain cases, as for instance in leaf-imitation among
+butterflies, and in mimicry generally. The resemblance to a leaf, for
+instance of a particular Kallima, seems to us so close as to be
+deceptive, and yet we find in another individual, or it may be in many
+others, a spot added which increases the resemblance, and which could
+not have become fixed unless the increased deceptiveness so produced
+had frequently led to the overlooking of its much persecuted
+possessor. But if we take the selection-value of the initial stages
+for granted, we are confronted with the further question which I
+myself formulated many years ago: How does it happen _that the
+necessary beginnings of a useful variation are always present_? How
+could insects which live upon or among green leaves become all green,
+while those that live on bark become brown? How have the desert
+animals become yellow and the Arctic animals white? Why were the
+necessary variations always present? How could the green locust lay
+brown eggs, or the privet caterpillar develop white and lilac-coloured
+lines on its green skin?
+
+It is of no use answering to this that the question is wrongly
+formulated[37] and that it is the converse that is true; that the
+process of selection takes place in accordance with the variations
+that present themselves. This proposition is undeniably true, but so
+also is another, which apparently negatives it: the variation required
+has in the majority of cases actually presented itself. Selection
+cannot solve this contradiction; it does not call forth the useful
+variation, but simply works upon it. The ultimate reason why one and
+the same insect should occur in green and in brown, as often happens
+in caterpillars and locusts, lies in the fact that variations towards
+brown presented themselves, and so also did variations towards green:
+_the kernel of the riddle lies in the varying_, and for the present we
+can only say, that small variations in different directions present
+themselves in every species. Otherwise so many different kinds of
+variations could not have arisen. I have endeavoured to explain this
+remarkable fact by means of the intimate processes that must take
+place within the germ-plasm, and I shall return to the problem when
+dealing with "germinal selection."
+
+We have, however, to make still greater demands on variation, for it
+is not enough that the necessary variation should occur in isolated
+individuals, because in that case there would be small prospect of its
+being preserved, notwithstanding its utility. Darwin at first
+believed, that even single variations might lead to transformation of
+the species, but later he became convinced that this was impossible,
+at least without the coöperation of other factors, such as isolation
+and sexual selection.
+
+In the case of the _green caterpillars with bright longitudinal
+stripes_, numerous individuals exhibiting this useful variation must
+have been produced to start with. In all higher, that is,
+multicellular organisms, the germ-substance is the source of all
+transmissible variations, and this germ-plasm is not a simple
+substance but is made up of many primary constituents. The question
+can therefore be more precisely stated thus: How does it come about
+that in so many cases the useful variations present themselves in
+numbers just where they are required, the white oblique lines in the
+leaf-caterpillar on the under surface of the body, the accompanying
+coloured stripes just above them? And, further, how has it come about
+that in grass caterpillars, not oblique but longitudinal stripes,
+which are more effective for concealment among grass and plants, have
+been evolved? And finally, how is it that the same Hawk-moth
+caterpillars, which to-day show oblique stripes, possessed
+longitudinal stripes in Tertiary times? We can read this fact from the
+history of their development, and I have before attempted to show the
+biological significance of this change of colour.[38]
+
+For the present I need only draw the conclusion that one and the same
+caterpillar may exhibit the initial stages of both, and that it
+depends on the manner in which these marking elements are
+_intensified_ and _combined_ by natural selection whether whitish
+longitudinal or oblique stripes should result. In this case then the
+"useful variations" were actually "always there," and we see that in
+the same group of Lepidoptera, e.g. species of Sphingidae, evolution
+has occurred in both directions according to whether the form lived
+among grass or on broad leaves with oblique lateral veins, and we can
+observe even now that the species with oblique stripes have
+longitudinal stripes when young, that is to say, while the stripes
+have no biological significance. The white places in the skin which
+gave rise, probably first as small spots, to this protective marking
+could be combined in one way or another according to the requirements
+of the species. They must therefore either have possessed
+selection-value from the first, or, if this was not the case at their
+earliest occurrence, there must have been _some other factors_ which
+raised them to the point of selection-value. I shall return to this in
+discussing germinal selection. But the case may be followed still
+farther, and leads us to the same alternative on a still more secure
+basis.
+
+Many years ago I observed in caterpillars of _Smerinthus populi_ (the
+poplar hawk-moth), which also possess white oblique stripes, that
+certain individuals showed _red spots_ above these stripes; these
+spots occurred only on certain segments, and never flowed together to
+form continuous stripes. In another species (_Smerinthus tiliae_)
+similar blood-red spots unite to form a line-like coloured seam in the
+last stage of larval life, while in _S. ocellata_ rust-red spots
+appear in individual caterpillars, but more rarely than in _S.
+populi_, and they show no tendency to flow together.
+
+Thus we have here the origin of a new character, arising from small
+beginnings, at least in _S. tiliae_, in which species the coloured
+stripes are a normal specific character. In the other species, _S.
+populi_ and _S. ocellata_, we find the beginnings of the same
+variation, in one more rarely than in the other, and we can imagine
+that, in the course of time, in these two species, coloured lines over
+the oblique stripes will arise. In any case these spots are the
+elements of variation, out of which coloured lines _may_ be evolved,
+if they are combined in this direction through the agency of natural
+selection. In _S. populi_ the spots are often small, but sometimes it
+seems as though several had united to form large spots. Whether a
+process of selection in this direction will arise in _S. populi_ and
+_S. ocellata_, or whether it is now going on cannot be determined,
+since we cannot tell in advance what biological value the marking
+might have for these two species. It is conceivable that the spots may
+have no selection-value as far as these species are concerned, and may
+therefore disappear again in the course of phylogeny, or, on the other
+hand, that they may be changed in another direction, for instance
+towards imitation of the rust-red fungoid patches on poplar and willow
+leaves. In any case we may regard the smallest spots as the initial
+stages of variation, the larger as a cumulative summation of these.
+Therefore either these initial stages must already possess
+selection-value, or, as I said before: _There must be some other
+reason for their cumulative summation_. I should like to give one more
+example, in which we can infer, though we cannot directly observe, the
+initial stages.
+
+All the Holothurians or sea-cucumbers have in the skin calcereous
+bodies of different forms, usually thick and irregular, which make the
+skin tough and resistant. In a small group of them--the species of
+Synapta--the calcareous bodies occur in the form of delicate anchors
+of microscopic size. Up till 1897 these anchors, like many other
+delicate microscopic structures, were regarded as curiosities, as
+natural marvels. But a Swedish observer, Oestergren, has recently
+shown that they have a biological significance: they serve the
+footless Synapta as auxiliary organs of locomotion, since, when the
+body swells up in the act of creeping, they press firmly with their
+tips, which are embedded in the skin, against the substratum on which
+the animal creeps, and thus prevent slipping backwards. In other
+Holothurians this slipping is made impossible by the fixing of the
+tube-feet. The anchors act automatically, sinking their tips towards
+the ground when the corresponding part of the body thickens, and
+returning to the original position at an angle of 45 degrees to the
+upper surface when the part becomes thin again. The arms of the anchor
+do not lie in the same plane as the shaft, and thus the curve of the
+arms forms the outermost part of the anchor, and offers no further
+resistance to the gliding of the animal. Every detail of the anchor,
+the curved portion, the little teeth at the head, the arms, etc., can
+be interpreted in the most beautiful way, above all the form of the
+anchor itself, for the two arms prevent it from swaying round to the
+side. The position of the anchors, too, is definite and significant;
+they lie obliquely to the longitudinal axis of the animal, and
+therefore they act alike whether the animal is creeping backwards or
+forwards. Moreover, the tips would pierce through the skin if the
+anchors lay in the longitudinal direction. Synapta burrows in the
+sand; it first pushes in the thin anterior end, and thickens this
+again, thus enlarging the hole, then the anterior tentacles displace
+more sand, the body is worked in a little farther, and the process
+begins anew. In the first act the anchors are passive, but they begin
+to take an active share in the forward movement when the body is
+contracted again. Frequently the animal retains only the posterior end
+buried in the sand, and then the anchors keep it in position, and make
+rapid withdrawal possible.
+
+Thus we have in these apparently random forms of the calcereous
+bodies, complex adaptations in which every little detail as to
+direction, curve, and pointing is exactly determined. That they have
+selection-value in their present perfected form is beyond all doubt,
+since the animals are enabled by means of them to bore rapidly into
+the ground and so to escape from enemies. We do not know what the
+initial stages were, but we cannot doubt that the little improvements,
+which occurred as variations of the originally simple slimy bodies of
+the Holothurians, were preserved because they already possessed
+selection-value for the Synaptidae. For such minute microscopic
+structures whose form is so delicately adapted to the rôle they have
+to play in the life of the animal, cannot have arisen suddenly and as
+a whole, and every new variation of the anchor, that is, in the
+direction of the development of the two arms, and every curving of the
+shaft which prevented the tips from projecting at the wrong time, in
+short, every little adaptation in the modelling of the anchor must
+have possessed selection-value. And that such minute changes of form
+fall within the sphere of fluctuating variations, that is to say,
+_that they occur_ is beyond all doubt.
+
+In many of the Synaptidae the anchors are replaced by calcareous rods
+bent in the form of an S, which are said to act in the same way.
+Others, such as those of the genus Ankyroderma, have anchors which
+project considerably beyond the skin, and, according to Oestergren,
+serve "to catch plant-particles and other substances" and so mask the
+animal. Thus we see that in the Synaptidae the thick and irregular
+calcareous bodies of the Holothurians have been modified and
+transformed in various ways in adaptation to the footlessness of these
+animals, and to the peculiar conditions of their life, and we must
+conclude that the earlier stages of these changes presented themselves
+to the processes of selection in the form of microscopic variations.
+For it is as impossible to think of any origin other than through
+selection in this case as in the case of the toughness, and the
+"drip-tips" of tropical leaves. And as these last could not have been
+produced directly by the beating of the heavy raindrops upon them, so
+the calcareous anchors of Synapta cannot have been produced directly
+by the friction of the sand and mud at the bottom of the sea, and,
+since they are parts whose function is _passive_ the Lamarckian factor
+of use and disuse does not come into question. The conclusion is
+unavoidable, that the microscopically small variations of the
+calcareous bodies in the ancestral forms have been intensified and
+accumulated in a particular direction, till they have led to the
+formation of the anchor. Whether this has taken place by the action of
+natural selection alone, or whether the laws of variation and the
+intimate processes within the germ-plasm have coöperated will become
+clear in the discussion of germinal selection. This whole process of
+adaptation has obviously taken place within the time that has elapsed
+since this group of sea-cucumbers lost their tube-feet, those
+characteristic organs of locomotion which occur in no group except the
+Echinoderms, and yet have totally disappeared in the Synaptidae. And
+after all what would animals that live in sand and mud do with
+tube-feet?
+
+
+(_c_) _Coadaptation_
+
+Darwin pointed out that one of the essential differences between
+artificial and natural selection lies in the fact that the former can
+modify only a few characters, usually only one at a time, while Nature
+preserves in the struggle for existence all the variations of a
+species, at the same time and in a purely mechanical way, if they
+possess selection-value.
+
+Herbert Spencer, though himself an adherent of the theory of selection,
+declared in the beginning of the nineties that in his opinion the range of
+this principle was greatly over-estimated, if the great changes which have
+taken place in so many organisms in the course of ages are to be
+interpreted as due to this process of selection alone, since no
+transformation of any importance can be evolved by itself; it is always
+accompanied by a host of secondary changes. He gives the familiar example
+of the Giant Stag of the Irish peat, the enormous antlers of which required
+not only a much stronger skull cap, but also greater strength of the
+sinews, muscles, nerves and bones of the whole anterior half of the animal,
+if their mass was not to weigh down the animal altogether. It is
+inconceivable, he says, that so many processes of selection should take
+place _simultaneously_, and we are therefore forced to fall back on the
+Lamarckian factor of the use and disuse of functional parts. And how, he
+asks, could natural selection follow two opposite directions of evolution
+in different parts of the body at the same time, as for instance in the
+case of the kangaroo, in which the forelegs must have become shorter, while
+the hind legs and the tail were becoming longer and stronger?
+
+Spencer's main object was to substantiate the validity of the
+Lamarckian principle, the coöperation of which with selection had been
+doubted by many. And it does seem as though this principle, if it
+operates in nature at all, offers a ready and simple explanation of
+all such secondary variations. Not only muscles, but nerves, bones,
+sinews, in short all tissues which function actively, increase in
+strength in proportion as they are used, and conversely they decrease
+when the claims on them diminish. All the parts, therefore, which
+depend on the part that varied first, as for instance the enlarged
+antlers of the Irish Elk, must have been increased or decreased in
+strength, in exact proportion to the claims made upon them,--just as
+is actually the case.
+
+But beautiful as this explanation would be, I regard it as untenable,
+because it assumes the _transmissibility of functional modifications_
+(so-called "acquired" characters), and this is not only
+undemonstrable, but is scarcely theoretically conceivable, for the
+secondary variations which accompany or follow the first as
+correlative variations, occur also in cases in which the animals
+concerned are sterile and _therefore cannot transmit anything to their
+descendants_. This is true of _worker bees_, and particularly of
+_ants_, and I shall here give a brief survey of the present state of
+the problem as it appears to me.
+
+Much has been written on both sides of this question since the
+published controversy on the subject in the nineties between Herbert
+Spencer and myself. I should like to return to the matter in detail,
+if the space at my disposal permitted, because it seems to me that the
+arguments I advanced at that time are equally cogent to-day,
+notwithstanding all the objections that have since been urged against
+them. Moreover, the matter is by no means one of subordinate interest;
+it is the very kernel of the whole question of the reality and value
+of the principle of selection. For if selection alone does not suffice
+to explain "_harmonious adaptation_" as I have called Spencer's
+_Coadaptation_, and if we require to call in the aid of the Lamarckian
+factor it would be questionable whether selection would explain any
+adaptations whatever. In this particular case--of worker bees--the
+Lamarckian factor may be excluded altogether, for it can be
+demonstrated that here at any rate the effects of use and disuse
+cannot be transmitted.
+
+But if it be asked why we are unwilling to admit the coöperation of
+the Darwinian factor of selection and the Lamarckian factor, since
+this would afford us an easy and satisfactory explanation of the
+phenomena, I answer: _Because the Lamarckian principle is fallacious,
+and because by accepting it we close the way towards deeper insight_.
+It is not a spirit of combativeness or a desire for self-vindication
+that induces me to take the field once more against the Lamarckian
+principle, it is the conviction that the progress of our knowledge is
+being obstructed by the acceptance of this fallacious principle, since
+the facile explanation it apparently affords prevents our seeking
+after a truer explanation and a deeper analysis.
+
+The workers in the various species of ants are sterile, that is to
+say, they take no regular part in the reproduction of the species,
+although individuals among them may occasionally lay eggs. In addition
+to this they have lost the wings, and the _receptaculum seminis_, and
+their compound eyes have degenerated to a few facets. How could this
+last change have come about through disuse, since the eyes of workers
+are exposed to light in the same way as are those of the sexual
+insects and thus in this particular case are not liable to "disuse" at
+all? The same is true of the _receptaculum seminis_, which can only
+have been disused as far as its glandular portion and its stalk are
+concerned, and also of the wings, the nerves tracheae and epidermal
+cells of which could not cease to function until the whole wing had
+degenerated, for the chitinous skeleton of the wing does not function
+at all in the active sense.
+
+But, on the other hand, the workers in all species have undergone
+modifications in a positive direction, as, for instance, the greater
+development of brain. In many species large workers have evolved,--the
+so-called _soldiers_, with enormous jaws and teeth, which defend the
+colony,--and in others there are _small_ workers which have taken over
+other special functions, such as the rearing of the young Aphides.
+This kind of division of the workers into two castes occurs among
+several tropical species of ants, but it is also present in the
+Italian species, _Colobopsis truncata_. Beautifully as the size of the
+jaws could be explained as due to the increased use made of them by
+the "soldiers," or the enlarged brain as due to the mental activities
+of the workers, the fact of the infertility of these forms is an
+insurmountable obstacle to accepting such an explanation. Neither jaws
+nor brain can have been evolved on the Lamarckian principle.
+
+The problem of coadaptation is no easier in the case of the ant than
+in the case of the Giant Stag. Darwin himself gave a pretty
+illustration to show how imposing the difference between the two kinds
+of workers in one species would seem if we translated it into human
+terms. In regard to the Driver ants (Anomma) we must picture to
+ourselves a piece of work, "for instance the building of a house,
+being carried on by two kinds of workers, of which one group was five
+feet four inches high, the other sixteen feet high."[39]
+
+Although the ant is a small animal as compared with man or with the
+Irish Elk, the "soldier" with its relatively enormous jaws is hardly
+less heavily burdened than the Elk with its antlers, and in the ant's
+case, too, a strengthening of the skeleton, of the muscles, the nerves
+of the head, and of the legs must have taken place parallel with the
+enlargement of the jaws. _Harmonious adaptation_ (coadaptation) has
+here been active in a high degree, and yet these "soldiers" are
+sterile! There thus remains nothing for it but to refer all their
+adaptations, positive and negative alike, to processes of selection
+which have taken place in the rudiments of the workers within the egg
+and sperm-cells of their parents. There is no way out of the
+difficulty except the one Darwin pointed out. He himself did not find
+the solution of the riddle at once. At first he believed that the case
+of the workers among social insects presented "the most serious
+special difficulty" in the way of his theory of natural selection; and
+it was only after it had become clear to him that it was not the
+sterile insects themselves but their parents that were selected,
+according as they produced more or less well adapted workers, that he
+was able to refer to this very case of the conditions among ants "_in
+order to show the power of natural selection_."[40] He explains his
+view by a simple but interesting illustration. Gardeners have
+produced, by means of long continued artificial selection, a variety
+of Stock, which bears entirely double, and therefore infertile
+flowers.[41] Nevertheless the variety continues to be reproduced from
+seed, because, in addition to the double and infertile flowers, the
+seeds always produce a certain number of single, fertile blossoms, and
+these are used to reproduce the double variety. These single and
+fertile plants correspond "to the males and females of an ant-colony,
+the infertile plants, which are regularly produced in large numbers,
+to the neuter workers of the colony."
+
+This illustration is entirely apt, the only difference between the
+two cases consisting in the fact that the variation in the flower is
+not a useful, but a disadvantageous one, which can only be preserved
+by artificial selection on the part of the gardener, while the
+transformations that have taken place parallel with the sterility of
+the ants are useful, since they procure for the colony an advantage in
+the struggle for existence, and they are therefore preserved by
+natural selection. Even the sterility itself in this case is not
+disadvantageous, since the fertility of the true females has at the
+same time considerably increased. We may therefore regard the sterile
+forms of ants, which have gradually been adapted in several directions
+to varying functions, _as a certain proof_ that selection really takes
+place in the germ-cells of the fathers and mothers of the workers, and
+that _special complexes of primordia_ (_ids_) are present in the
+workers and in the males and females, and these complexes contain the
+primordia of the individual parts (_determinants_). But since all
+living entities vary, the determinants must also vary, now in a
+favourable, now in an unfavourable direction. If a female produces
+eggs, which contain favourably varying determinants in the worker-ids,
+then these eggs will give rise to workers modified in the favourable
+direction, and if this happens with many females, the colony concerned
+will contain a better kind of worker than other colonies.
+
+I digress here in order to give an account of the intimate processes,
+which, according to my view, take place within the germ-plasm, and
+which I have called "_germinal selection_." These processes are of
+importance since they form the roots of variation, which in its turn
+is the root of natural selection. I cannot here do more than give a
+brief outline of the theory in order to show how the Darwin-Wallace
+theory of selection has gained support from it.
+
+With others, I regard the minimal amount of substance which is
+contained within the nucleus of the germ-cells, in the form of rods,
+bands, or granules, as the _germ-substance_ or _germ-plasm_, and I
+call the individual granules _ids_. There is always a multiplicity of
+such ids present in the nucleus, either occurring individually, or
+united in the form of rods or bands (chromosomes). Each id contains
+the primary constituents of a _whole_ individual, so that several ids
+are concerned in the development of a new individual.
+
+In every being of complex structure thousands of primary constituents
+must go to make up a single id; these I call _determinants_, and I
+mean by this name very small individual particles, far below the
+limits of microscopic visibility, vital units which feed, grow, and
+multiply by division. These determinants control the parts of the
+developing embryo,--in what manner need not here concern us. The
+determinants differ among themselves, those of a muscle are
+differently constituted from those of a nerve-cell or a glandular
+cell, etc., and every determinant is in its turn made up of minute
+vital units, which I call _biophores_, or the bearers of life.
+According to my view, these determinants not only assimilate, like
+every other living unit, but they _vary_ in the course of their
+growth, as every living unit does; they may vary qualitatively if the
+elements of which they are composed vary, they may grow and divide
+more or less rapidly, and their variations give rise to
+_corresponding_ variations of the organ, cell, or cell-group which
+they determine. That they are undergoing ceaseless fluctuations in
+regard to size and quality seems to me the inevitable consequence of
+their unequal nutrition; for although the germ-cell as a whole usually
+receives sufficient nutriment, minute fluctuations in the amount
+carried to different parts within the germ-plasm cannot fail to occur.
+
+Now, if a determinant, for instance of a sensory cell, receives for a
+considerable time more abundant nutriment than before, it will grow
+more rapidly--become bigger, and divide more quickly, and, later, when
+the id concerned develops into an embryo, this sensory cell will
+become stronger than in the parents, possibly even twice as strong.
+This is an instance of a _hereditary individual variation_, arising
+from the germ.
+
+The nutritive stream which, according to our hypothesis, favours the
+determinant _N_ by chance, that is, for reasons unknown to us, may
+remain strong for a considerable time, or may decrease again; but even
+in the latter case it is conceivable that the ascending movement of
+the determinant may continue, because the strengthened determinant now
+_actively_ nourishes itself more abundantly,--that is to say, it
+attracts the nutriment to itself, and to a certain extent withdraws it
+from its fellow-determinants. In this way, it may--as it seems to
+me--get into _permanent upward movement, and attain a degree of
+strength from which there is no falling back_. Then positive or
+negative selection sets in, favouring the variations which are
+advantageous, setting aside those which are disadvantageous.
+
+In a similar manner a _downward_ variation of the determinants may
+take place, if its progress be started by a diminished flow of
+nutriment. The determinants which are weakened by this diminished flow
+will have less affinity for attracting nutriment because of their
+diminished strength, and they will assimilate more feebly and grow
+more slowly, unless chance streams of nutriment help them to recover
+themselves. But, as will presently be shown, a change of direction
+cannot take place at _every_ stage of the degenerative process. If a
+certain critical stage of downward progress be passed, even favourable
+conditions of food-supply will no longer suffice permanently to change
+the direction of the variation. Only two cases are conceivable; if the
+determinant corresponds to a _useful_ organ, only its removal can
+bring back the germ-plasm to its former level; therefore personal
+selection removes the id in question, with its determinants, from the
+germ-plasm, by causing the elimination of the individual in the
+struggle for existence. But there is another conceivable case; the
+determinants concerned may be those of an organ which has become
+_useless_, and they will then continue unobstructed, but with
+exceeding slowness, along the downward path, until the organ becomes
+vestigial, and finally disappears altogether.
+
+The fluctuations of the determinants hither and thither may thus be
+transformed into a lasting ascending or descending movement; and _this
+is the crucial point of these germinal processes_.
+
+This is not a fantastic assumption; we can read it in the fact of the
+degeneration of disused parts. _Useless organs are the only ones which
+are not helped to ascend again by personal selection, and therefore in
+their case alone can we form any idea of how the primary constituents
+behave, when they are subject solely to intra-germinal forces_.
+
+The whole determinant system of an id, as I conceive it, is in a state
+of continual fluctuation upwards and downwards. In most cases the
+fluctuations will counteract one another, because the passive streams
+of nutriment soon change, but in many cases the limit from which a
+return is possible will be passed, and then the determinants concerned
+will continue to vary in the same direction, till they attain positive
+or negative selection-value. At this stage personal selection
+intervenes and sets aside the variation if it is disadvantageous, or
+favours--that is to say, preserves--it if it is advantageous. Only
+_the determinant of a useless organ is uninfluenced by personal
+selection_, and, as experience shows, it sinks downwards; that is, the
+organ that corresponds to it degenerates very slowly but
+uninterruptedly till, after what must obviously be an immense stretch
+of time, it disappears from the germ-plasm altogether.
+
+Thus we find in the fact of the degeneration of disused parts the
+proof that not all the fluctuations of a determinant return to
+equilibrium again, but that, when the movement has attained to a
+certain strength, it continues _in the same direction_. We have entire
+certainty in regard to this as far as the downward progress is
+concerned, and we must assume it also in regard to ascending
+variations, as the phenomena of artificial selection certainly justify
+us in doing. If the Japanese breeders were able to lengthen the
+tail-feathers of the cock to six feet, it can only have been because
+the determinants of the tail-feathers in the germ-plasm had already
+struck out a path of ascending variation, and this movement was taken
+advantage of by the breeder, who continually selected for reproduction
+the individuals in which the ascending variation was most marked. For
+all breeding depends upon the unconscious selection of germinal
+variations.
+
+Of course these germinal processes cannot be proved mathematically,
+since we cannot actually see the play of forces of the passive
+fluctuations and their causes. We cannot say how great these
+fluctuations are, and how quickly or slowly, how regularly or
+irregularly they change. Nor do we know how far a determinant must be
+strengthened by the passive flow of the nutritive stream if it is to
+be beyond the danger of unfavourable variations, or how far it must be
+weakened passively before it loses the power of recovering itself by
+its own strength. It is no more possible to bring forward actual
+proofs in this case than it was in regard to the selection-value of
+the initial stages of an adaptation. But if we consider that all
+heritable variations must have their roots in the germ-plasm, and
+further, that when personal selection does not intervene, that is to
+say, in the case of parts which have become useless, a degeneration of
+the part, and therefore also of its determinant must inevitably take
+place; then we must conclude that processes such as I have assumed are
+running their course within the germ-plasm, and we can do this with as
+much certainty as we were able to infer, from the phenomena of
+adaptation, the selection-value of their initial stages. The fact of
+the degeneration of disused parts seems to me to afford irrefutable
+proof that the fluctuations within the germ-plasm _are the real root
+of all hereditary variation_, and the preliminary condition for the
+occurrence of the Darwin-Wallace factor of selection. Germinal
+selection supplies the stones out of which personal selection builds
+her temples and palaces: _adaptations_. The importance for the theory
+of the process of degeneration of disused parts cannot be
+over-estimated, especially when it occurs in sterile animal forms,
+where we are free from the doubt as to the alleged _Lamarckian factor_
+which is apt to confuse our ideas in regard to other cases.
+
+If we regard the variation of the many determinants concerned in the
+transformation of the female into the sterile worker as having come
+about through the gradual transformation of the ids into worker-ids,
+we shall see that the germ-plasm of the sexual ants must contain three
+kinds of ids, male, female, and worker ids, or if the workers have
+diverged into soldiers and nest-builders, then four kinds. We
+understand that the worker-ids arose because their determinants struck
+out a useful path of variation, whether upward or downward, and that
+they continued in this path until the highest attainable degree of
+utility of the parts determined was reached. But in addition to the
+organs of positive or negative selection-value, there were some which
+were indifferent as far as the success and especially the functional
+capacity of the workers was concerned: wings, ovarian tubes,
+_receptaculum seminis_, a number of the facets of the eye, perhaps
+even the whole eye. As to the ovarian tubes it is is possible that
+their degeneration was an advantage for the workers, in saving energy,
+and if so selection would favour the degeneration; but how could the
+presence of eyes diminish the usefulness of the workers to the colony?
+or the minute _receptaculum seminis_, or even the wings? These parts
+have therefore degenerated _because they were of no further value to
+the insect_. But if selection did not influence the setting aside of
+these parts because they were neither of advantage nor of disadvantage
+to the species, then the Darwinian factor of selection is here
+confronted with a puzzle which it cannot solve alone, but which at
+once becomes clear when germinal selection is added. For the
+determinants of organs that have no further value for the organism,
+must, as we have already explained, embark on a gradual course of
+retrograde development.
+
+In ants the degeneration has gone so far that there are no
+wing-rudiments present in _any_ species, as is the case with so many
+butterflies, flies, and locusts, but in the larvae the imaginable
+discs of the wings are still laid down. With regard to the ovaries,
+degeneration has reached different levels in different species of
+ants, as has been shown by the researches of my former pupil,
+Elizabeth Bickford. In many species there are twelve ovarian tubes,
+and they decrease from that number to one; indeed, in one species no
+ovarian tube at all is present. So much at least is certain from what
+has been said, that in this case _everything_ depends on the
+fluctuations of the elements of the germ-plasm. Germinal selection,
+here as elsewhere, presents the variations of the determinants, and
+personal selection favours or rejects these, or,--if it be a question
+of organs which have become useless,--it does not come into play at
+all, and allows the descending variation free course.
+
+It is obvious that even the problem of _coadaptation in sterile
+animals_ can thus be satisfactorily explained. If the determinants are
+oscillating upwards and downwards in continual fluctuation, and
+varying more pronouncedly now in one direction now in the other,
+useful variations of every determinant will continually present
+themselves anew, and may, in the course of generations, be combined
+with one another in various ways. But there is one character of the
+determinants that greatly facilitates this complex process of
+selection, that, after a certain limit has been reached, they go on
+varying in the same direction. From this it follows that development
+along a path once struck out may proceed without the continual
+intervention of personal selection. This factor only operates, so to
+speak, at the beginning, when it selects the determinants which are
+varying in the right direction, and again at the end, when it is
+necessary to put a check upon further variation. In addition to this,
+enormously long periods have been available for all these adaptations,
+as the very gradual transition stages between females and workers in
+many species plainly show, and thus this process of transformation
+loses the marvellous and mysterious character that seemed at the first
+glance to invest it, and takes rank, without any straining, among the
+other processes of selection. It seems to me that, from the facts that
+sterile animal forms can adapt themselves to new vital functions,
+their superfluous parts degenerate, and the parts more used adapt
+themselves in an ascending direction, those less used in a descending
+direction, we must draw the conclusion that harmonious adaptation here
+comes about _without the coöperation of the Lamarckian principle_.
+This conclusion once established, however, we have no reason to refer
+the thousands of cases of harmonious adaptation, which occur in
+exactly the same way among other animals or plants, to a principle,
+the _active intervention of which in the transformation of species is
+nowhere proved. We do not require it to explain the facts, and
+therefore we must not assume it._
+
+The fact of coadaptation, which was supposed to furnish the strongest
+argument against the principle of selection, in reality yields the
+clearest evidence in favour of it. We _must_ assume it, _because no
+other possibility of explanation is open to us, and because these
+adaptations actually exist, that is to say, have really taken place_.
+With this conviction I attempted, as far back as 1894, when the idea
+of germinal selection had not yet occurred to me, to make "harmonious
+adaptation" (coadaptation) more easily intelligible in some way or
+other, and so I was led to the idea, which was subsequently expounded
+in detail by Baldwin, and Lloyd Morgan, and also by Osborn, and Gulick
+as _Organic Selection_. It seemed to me that it was not necessary that
+all the germinal variations required for secondary variations should
+have occurred _simultaneously_, since, for instance, in the case of
+the stag, the bones, muscles, sinews, and nerves would be incited by
+the increasing heaviness of the antlers to greater activity in _the
+individual life_, and so would be strengthened. The antlers can only
+have increased in size by very slow degrees, so that the muscles and
+bones may have been able to keep pace with their growth in the
+individual life, until the requisite germinal variations presented
+themselves. In this way a disharmony between the increasing weight of
+the antlers and the parts which support and move them would be
+avoided, since time would be given for the appropriate germinal
+variations to occur, and so to set agoing the _hereditary_ variation
+of the muscles, sinews and bones.[42]
+
+I still regard this idea as correct, but I attribute less importance
+to "organic selection" than I did at that time, in so far that I do
+not believe that it _alone_ could effect complex harmonious
+adaptations. Germinal selection now seems to me to play the chief part
+in bringing about such adaptations. Something the same is true of the
+principle I have called _Panmixia_. As I became more and more
+convinced, in the course of years, that the _Lamarckian principle_
+ought not to be called in to explain the dwindling of disused parts, I
+believed that this process might be simply explained as due to the
+cessation of the conservative effect of natural selection. I said to
+myself that, from the moment in which a part ceases to be of use,
+natural selection withdraws its hand from it, and then it must
+inevitably fall from the height of its adaptiveness, because inferior
+variants would have as good a chance of persisting as better ones,
+since all grades of fitness of the part in question would be mingled
+with one another indiscriminately. This is undoubtedly true, as
+Romanes pointed out ten years before I did, and this mingling of the
+bad with the good probably does bring about a deterioration of the
+part concerned. But it cannot account for the steady diminution, which
+always occurs when a part is in process of becoming rudimentary, and
+which goes on until it ultimately disappears altogether. The process
+of dwindling cannot therefore be explained as due to panmixia alone:
+we can only find a sufficient explanation in germinal selection.
+
+
+IV. DERIVATIVES OF THE THEORY OF SELECTION
+
+The impetus in all directions given by Darwin through his theory of
+selection has been an immeasurable one, and its influence is still
+felt. It falls within the province of the historian of science to
+enumerate all the ideas which, in the last quarter of the nineteenth
+century, grew out of Darwin's theories, in the endeavour to penetrate
+more deeply into the problem of the evolution of the organic world.
+Within the narrow limits to which this paper is restricted, I cannot
+attempt to discuss any of these.
+
+
+V. ARGUMENTS FOR THE REALITY OF THE PROCESSES OF SELECTION
+
+
+(_a_) _Sexual Selection_
+
+Sexual selection goes hand in hand with natural selection. From the
+very first I have regarded sexual selection as affording an extremely
+important and interesting corroboration of natural selection, but,
+singularly enough, it is precisely against this theory that an adverse
+judgment has been pronounced in so many quarters, and it is only quite
+recently, and probably in proportion as the wealth of facts in proof
+of it penetrates into a wider circle, that we seem to be approaching a
+more general recognition of this side of the problem of adaptations.
+Thus Darwin's words in his preface to the second edition (1874) of his
+book, _The Descent of Man and Sexual Selection_, are being justified:
+"My conviction as to the operation of natural selection remains
+unshaken," and further, "If naturalists were to become more familiar
+with the idea of sexual selection, it would, I think, be accepted to a
+much greater extent, and already it is fully and favourably accepted
+by many competent judges." Darwin was able to speak thus because he
+was already acquainted with an immense mass of facts, which, taken
+together, yield overwhelming evidence of the validity of the principle
+of sexual selection.
+
+_Natural selection_ chooses out for reproduction the individuals that
+are best equipped for the struggle for existence, and it does so at
+every stage of development; it thus improves the species in all its
+stages and forms. _Sexual selection_ operates only on individuals
+that are already capable of reproduction, and does so only in relation
+to the attainment of reproduction. It arises from the rivalry of one
+sex, usually the male, for the possession of the other, usually the
+female. Its influence can therefore only _directly_ affect one sex, in
+that it equips it better for attaining possession of the other. But
+the effect may extend indirectly to the female sex, and thus the whole
+species may be modified, without, however, becoming any more capable
+of resistance in the struggle for existence, for sexual selection only
+gives rise to adaptations which are likely to give their possessor the
+victory over rivals in the struggle for possession of the female, and
+which are therefore peculiar to the wooing sex: the manifold
+"secondary sexual characters." The diversity of these characters is so
+great that I cannot here attempt to give anything approaching a
+complete treatment of them, but I should like to give a sufficient
+number of examples to make the principle itself, in its various modes
+of expression, quite clear.
+
+One of the chief preliminary postulates of sexual selection is the
+unequal number of individuals in the two sexes, for if every male
+immediately finds his mate there can be no competition for the
+possession of the female. Darwin has shown that, for the most part,
+the inequality between the sexes is due simply to the fact that there
+are more males than females, and therefore the males must take some
+pains to secure a mate. But the inequality does not always depend on
+the numerical preponderance of the males, it is often due to polygamy;
+for, if one male claims several females, the number of females in
+proportion to the rest of the males will be reduced. Since it is
+almost always the males that are the wooers, we must expect to find
+the occurrence of secondary sexual characters chiefly among them, and
+to find it especially frequent in polygamous species. And this is
+actually the case.
+
+If we were to try to guess--without knowing the facts--what means the
+male animals make use of to overcome their rivals in the struggle for
+the possession of the female, we might name many kinds of means, but
+it would be difficult to suggest any which is not actually employed in
+some animal group of other. I begin with the mere difference in
+strength, through which the male of many animals is so sharply
+distinguished from the female, as, for instance, the lion, walrus,
+"sea-elephant," and others. Among these the males fight violently for
+the possession of the female, who falls to the victor in the combat.
+In this simple case no one can doubt the operation of selection, and
+there is just as little room for doubt as to the selection-value of
+the initial stages of the variation. Differences in bodily strength
+are apparent even among human beings, although in their case the
+struggle for the possession of the female is no longer decided by
+bodily strength alone.
+
+Combats between male animals are often violent and obstinate, and the
+employment of the natural weapons of the species in this way has led
+to perfecting of these, e.g. the tusks of the boar, the antlers of the
+stag, and the enormous, antler-like jaws of the stag-beetle. Here
+again it is impossible to doubt that variations in these organs
+presented themselves, and that these were considerable enough to be
+decisive in combat, and so to lead to the improvement of the weapon.
+
+Among many animals, however, the females at first withdraw from the
+males; they are coy, and have to be sought out, and sometimes held by
+force. This tracking and grasping of the females by the males has
+given rise to many different characters in the latter, as, for
+instance, the larger eyes of the male bee, and especially of the males
+of the Ephemerids (May-flies), some species of which show, in addition
+to the usual compound eyes, large, so-called turban-eyes, so that the
+whole head is covered with seeing surfaces. In these species the
+females are very greatly in the minority (1-100), and it is easy to
+understand that a keen competition for them must take place, and that,
+when the insects of both sexes are floating freely in the air, an
+unusually wide range of vision will carry with it a decided
+advantage. Here again the actual adaptations are in accordance with
+the preliminary postulates of the theory. We do not know the stages
+through which the eye has passed to its present perfected state, but,
+since the number of simple eyes (facets) has become very much greater
+in the male than in the female, we may assume that their increase is
+due to a gradual duplication of the determinants of the ommatidium in
+the germ-plasm, as I have already indicated in regard to sense-organs
+in general. In this case, again, the selection-value of the initial
+stages hardly admits of doubt; better vision _directly_ secures
+reproduction.
+
+In many cases _the organ of smell_ shows a similar improvement. Many
+lower Crustaceans (Daphnidae) have better developed organs of smell in
+the male sex. The difference is often slight and amounts only to one
+or two olfactory filaments, but certain species show a difference of
+nearly a hundred of these filaments (Leptodora). The same thing occurs
+among insects.
+
+We must briefly consider the clasping or grasping organs which have
+developed in the males among many lower Crustaceans, but here natural
+selection plays its part along with sexual selection, for the union of
+the sexes is an indispensable condition for the maintenance of the
+species, and as Darwin himself pointed out, in many cases the two
+forms of selection merge into each other. This fact has always seemed
+to me to be a proof of natural selection, for, in regard to sexual
+selection, it is quite obvious that the victory of the best-equipped
+could have brought about the improvement only of the organs concerned,
+the factors in the struggle, such as the eye and the olfactory organ.
+
+We come now to the _excitants_; that is, to the group of sexual
+characters whose origin through processes of selection has been most
+frequently called in question. We may cite the _love-calls_ produced
+by many male insects, such as crickets and cicadas. These could only
+have arisen in animal groups in which the female did not rapidly flee
+from the male, but was inclined to accept his wooing from the first.
+Thus, notes like the chirping of the male cricket serve to entice the
+females. At first they were merely the signal which showed the
+presence of a male in the neighbourhood, and the female was gradually
+enticed nearer and nearer by the continued chirping. The male that
+could make himself heard to the greatest distance would obtain the
+largest following, and would transmit the beginnings, and, later, the
+improvement of his voice to the greatest number of descendants. But
+sexual excitement in the female became associated with the hearing of
+the love-call, and then the sound-producing organ of the male began to
+improve, until it attained to the emission of the long-drawn-out soft
+notes of the mole-cricket or the maenad-like cry of the cicadas. I
+cannot here follow the process of development in detail, but will call
+attention to the fact that the original purpose of the voice, the
+announcing of the male's presence, became subsidiary, and the exciting
+of the female became the chief goal to be aimed at. The loudest
+singers awakened the strongest excitement, and the improvement
+resulted as a matter of course. I conceive of the origin of bird-song
+in a somewhat similar manner, first as a means of enticing, then of
+exciting the female.
+
+One more kind of secondary sexual character must here be mentioned:
+the odour which emanates from so many animals at the breeding season.
+It is possible that this odour also served at first merely to give
+notice of the presence of individuals of the other sex, but it soon
+became an excitant, and as the individuals which caused the greatest
+degree of excitement were preferred, it reached as high a pitch of
+perfection as was possible to it. I shall confine myself here to the
+comparatively recently discovered fragrance of butterflies. Since
+Fritz Müller found out that certain Brazilian butterflies gave off
+fragrance "like a flower," we have become acquainted with many such
+cases, and we now know that in all lands, not only many diurnal
+Lepidoptera but nocturnal ones also give off a delicate odour, which
+is agreeable even to man. The ethereal oil to which this fragrance is
+due is secreted by the skin-cells, usually of the wing, as I showed
+soon after the discovery of the _scent-scales_. This is the case in
+the males; the females have no _special_ scent-scales recognisable as
+such by their form, but they must, nevertheless, give off an extremely
+delicate fragrance, although our imperfect organ of smell cannot
+perceive it, for the males become aware of the presence of a female,
+even at night, from a long distance off, and gather round her. We may
+therefore conclude, that both sexes have long given forth a very
+delicate perfume, which announced their presence to others of the same
+species, and that in many species (_not in all_) these small
+beginnings become, in the males, particularly strong scent-scales of
+characteristic form (lute, brush, or lyre-shaped). At first these
+scales were scattered over the surface of the wing, but gradually they
+concentrated themselves, and formed broad, velvety bands, or strong,
+prominent brushes, and they attained their highest pitch of evolution
+when they became enclosed within pits or folds of the skin, which
+could be opened to let the delicious fragrance stream forth suddenly
+towards the female. Thus in this case also we see that characters, the
+original use of which was to bring the sexes together, and so to
+maintain the species, have been evolved in the males into means for
+exciting the female. And we can hardly doubt, that the females are
+most readily enticed to yield to the butterfly that sends out the
+strongest fragrance,--that is to say, that excites them to the highest
+degree. It is a pity that our organs of smell are not fine enough to
+examine the fragrance of male Lepidoptera in general, and to compare
+it with other perfumes which attract these insects.[43] As far as we
+can perceive them they resemble the fragrance of flowers, but there
+are Lepidoptera whose scent suggests musk. A smell of musk is also
+given off by several plants: it is a sexual excitant in the
+musk-deer, the musk-sheep, and the crocodile.
+
+As far as we know, then, it is perfumes similar to those of flowers
+that the male Lepidoptera give off in order to entice their mates and
+this is a further indication that animals, like plants, can to a large
+extent meet the claims made upon them by life, and produce the
+adaptations which are most purposive,--a further proof, too, of my
+proposition that the useful variations, so to speak, are _always
+there_. The flowers developed the perfumes which entice their
+visitors, and the male Lepidoptera developed the perfumes which entice
+and excite their mates.
+
+There are many pretty little problems to be solved in this connection,
+for there are insects, such as some flies, that are attracted by
+smells which are unpleasant to us, like those from decaying flesh and
+carrion. But there are also certain flowers, some orchids for
+instance, which give forth no very agreeable odour, but one which is
+to us repulsive and disgusting; and we should therefore expect that
+the males of such insects would give off a smell unpleasant to us, but
+there is no case known to me in which this has been demonstrated.
+
+In cases such as we have discussed, it is obvious that there is no
+possible explanation except through selection. This brings us to the
+last kind of secondary sexual characters, and the one in regard to
+which doubt has been most frequently expressed,--decorative colours
+and decorative forms, the brilliant plumage of the male pheasant, the
+humming-birds, and the bird of Paradise, as well as the bright colours
+of many species of butterfly, from the beautiful blue of our little
+Lycaenidae to the magnificent azure of the large Morphinae of Brazil.
+In a great many cases, though not by any means in all, the male
+butterflies are "more beautiful" than the females, and in the Tropics
+in particular they shine and glow in the most superb colours. I really
+see no reason why we should doubt the power of sexual selection, and I
+myself stand wholly on Darwin's side. Even though we certainly cannot
+assume that the females exercise a conscious choice of the
+"handsomest" mate, and deliberate like the judges in a court of
+justice over the perfections of their wooers, we have no reason to
+doubt that distinctive forms (decorative feathers), and colours have a
+particularly exciting effect upon the female, just as certain odours
+have among animals of so many different groups, including the
+butterflies. The doubts which existed for a considerable time, as a
+result of fallacious experiments, as to whether the colours of flowers
+really had any influence in attracting butterflies have now been set
+at rest through a series of more careful investigations; we now know
+that the colours of flowers are there on account of the butterflies,
+as Sprengel first showed, and that the blossoms of Phanerogams are
+selected in relation to them, as Darwin pointed out.
+
+Certainly it is not possible to bring forward any convincing proof of
+the origin of decorative colours through sexual selection, but there
+are many weighty arguments in favour of it, and these form a body of
+presumptive evidence so strong that it almost amounts to certainty.
+
+In the first place, there is the analogy with other secondary sexual
+characters. If the song of birds and the chirping of the cricket have
+been evolved through sexual selection, if the penetrating odours of
+male animals,--the crocodile, the musk-deer, the beaver, the
+carnivores, and, finally, the flower-like fragrances of the
+butterflies have been evolved to their present pitch in this way, why
+should decorative colours have arisen in some other way? Why should
+the eye be less sensitive to _specifically male_ colours and other
+_visible_ signs _enticing to the female_, than the olfactory sense to
+specifically male odours, or the sense of hearing to specifically male
+sounds? Moreover, the decorative feathers of birds are almost always
+spread out and displayed before the female during courtship. I have
+elsewhere[44] pointed out that decorative colouring and
+sweet-scentedness may replace one another in Lepidoptera as well as in
+flowers, for just as some modestly coloured flowers (mignonette and
+violet) have often a strong perfume, while strikingly coloured ones
+are sometimes quite devoid of fragrance, so we find that the most
+beautiful and gaily-coloured of our native Lepidoptera, the species of
+Vanessa, have no scent-scales, while these are often markedly
+developed in grey nocturnal Lepidoptera. Both attractions may,
+however, be combined in butterflies, just as in flowers. Of course, we
+cannot explain why both means of attraction should exist in one genus,
+and only one of them in another, since we do not know the minutest
+details of the conditions of life of the genera concerned. But from
+the sporadic distribution of scent-scales in Lepidoptera, and from
+their occurrence or absence in nearly related species, we may conclude
+that fragrance is a relatively _modern_ acquirement, more recent than
+brilliant colouring.
+
+One thing in particular that stamps decorative colouring as a product
+of selection is _its gradual intensification_ by the addition of new
+spots, which we can quite well observe, because in many cases the
+colours have been first acquired by the males, and later transmitted
+to the females by inheritance. The scent-scales are never thus
+transmitted, probably for the same reason that the decorative colours
+of many birds are often not transmitted to the females: because with
+these they would be exposed to too great elimination by enemies.
+Wallace was the first to point out that in species with concealed
+nests the beautiful feathers of the male occurred in the female also,
+as in the parrots, for instance, but this is not the case in species
+which brood on an exposed nest. In the parrots one can often observe
+that the general brilliant colouring of the male is found in the
+female, but that certain spots of colour are absent, and these have
+probably been acquired comparatively recently by the male and have not
+yet been transmitted to the female.
+
+Isolation of the group of individuals which is in process of varying
+is undoubtedly of great value in sexual selection, for even a solitary
+conspicuous variation will become dominant much sooner in a small
+isolated colony, than among a large number of members of a species.
+
+Any one who agrees with me in deriving variations from germinal
+selection will regard that process as an essential aid towards
+explaining the selection of distinctive courtship-characters, such as
+coloured spots, decorative feathers, horny outgrowths in birds and
+reptiles, combs, feather-tufts, and the like, since the beginnings of
+these would be presented with relative frequency in the struggle
+between the determinants within the germ-plasm. The process of
+transmission of decorative feathers to the female results, as Darwin
+pointed out and illustrated by interesting examples, in the
+_colour-transformation of a whole species_, and this process, as the
+phyletically older colouring of young birds shows, must, in the course
+of thousands of years, have repeated itself several times in a line of
+descent.
+
+If we survey the wealth of phenomena presented to us by secondary
+sexual characters, we can hardly fail to be convinced of the truth of
+the principle of sexual selection. And certainly no one who has
+accepted natural selection should reject sexual selection, for, not
+only do the two processes rest upon the same basis, but they merge
+into one another, so that it is often impossible to say how much of a
+particular character depends on one and how much on the other form of
+selection.
+
+
+(_b_) _Natural Selection_
+
+An actual proof of the theory of sexual selection is out of the
+question, if only because we cannot tell when a variation attains to
+selection-value. It is certain that a delicate sense of smell is of
+value to the male moth in his search for the female, but whether the
+possession of one additional olfactory hair, or of ten, or of twenty
+additional hairs leads to the success of its possessor we are unable
+to tell. And we are groping even more in the dark when we discuss the
+excitement caused in the female by agreeable perfumes, or by striking
+and beautiful colours. That these do make an impression is beyond
+doubt; but we can only assume that slight intensifications of them
+give any advantage, and we _must_ assume this _since otherwise
+secondary sexual characters remain inexplicable_.
+
+The same thing is true in regard to natural selection. It is not
+possible to bring forward any actual proof of the selection-value of
+the initial stages, and the stages in the increase of variations, as
+has been already shown. But the selection-value of a finished
+adaptation can in many cases be statistically determined. Cesnola and
+Poulton have made valuable experiments in this direction. The former
+attached forty-five individuals of the green, and sixty-five of the
+brown variety of the praying mantis (_Mantis religiosa_), by a silk
+thread to plants, and watched them for seven days. The insects which
+were on a surface of a colour Similar to their own remained uneaten,
+while twenty-five green insects on brown parts of plants had all
+disappeared in eleven days.
+
+The experiments of Poulton and Sanders[45] were made with 600 pupae of
+_Vanessa urticae_, the "tortoise-shell butterfly." The pupae were
+artificially attached to nettles, tree-trunks, fences, walls, and to
+the ground, some at Oxford, some at St. Helens in the Isle of Wight.
+In the course of a month 93% of the pupae at Oxford were killed,
+chiefly by small birds, while at St. Helens 68% perished. The
+experiments showed very clearly that the colour and character of the
+surface on which the pupa rests--and thus its own conspicuousness--are
+of the greatest importance. At Oxford only the four pupae which were
+fastened to nettles emerged; all the rest--on bark, stones and the
+like--perished. At St. Helens the elimination was as follows: on
+fences where the pupae were conspicuous, 92%; on bark, 66%; on walls,
+54%; and among nettles, 57%. These interesting experiments confirm our
+views as to protective coloration, and show further, _that the ratio
+of elimination in the species is a very high one, and that therefore
+selection must be very keen_.
+
+We may say that the process of selection follows as a logical
+necessity from the fulfilment of the three preliminary postulates of
+the theory: variability, heredity, and the struggle for existence,
+with its enormous ratio of elimination in all species. To this we must
+add a fourth factor, the _intensification_ of variations which Darwin
+established as a fact, and which we are now able to account for
+theoretically on the basis of germinal selection. It may be objected
+that there is considerable uncertainty about this _logical_ proof,
+because of our inability to demonstrate the selection-value of the
+initial stages and the individual stages of increase. We have
+therefore to fall back on _presumptive evidence_. This is to be found
+in _the interpretative value of the theory_. Let us consider this
+point in greater detail.
+
+In the first place it is necessary to emphasize what is often
+overlooked, namely, that the theory not only explains the
+_transformations_ of species, it also explains _their remaining the
+same_; in addition to the principle of varying, it contains within
+itself that of _persisting_. It is part of the essence of selection,
+that it not only causes a part to _vary_ till it has reached its
+highest pitch of adaptation, but that it _maintains it at this pitch.
+This conserving influence of natural selection_ is of great
+importance, and was early recognised by Darwin; it follows naturally
+from the principle of the survival of the fittest.
+
+We understand from this how it is that a species which has become
+fully adapted to certain conditions of life ceases to vary, but
+remains "constant," as long as the conditions of life _for_ it remain
+unchanged, whether this be for thousands of years, or for whole
+geological epochs. But the most convincing proof of the power of the
+principle of selection lies in the innumerable multitude of phenomena
+which cannot be explained in any other way. To this category belong
+all structures which are only _passively_ of advantage to the
+organism, because none of these can have arisen by the alleged
+_Lamarckian principle_. These have been so often discussed that we
+need do no more than indicate them here. Until quite recently the
+sympathetic coloration of animals--for instance, the whiteness of
+Arctic animals--was referred, at least in part, to the _direct_
+influence of external factors, but the facts can best be explained by
+referring them to the processes of selection, for then it is
+unnecessary to make the gratuitous assumption that many species are
+sensitive to the stimulus of cold and that others are not. The great
+majority of Arctic land-animals, mammals and birds, are white, and
+this proves that they were all able to present the variation which was
+most useful for them. The sable is brown, but it lives in trees, where
+the brown colouring protects and conceals it more effectively. The
+musk-sheep (_Ovibos moschatus_) is also brown, and contrasts sharply
+with the ice and snow, but it is protected from beasts of prey by its
+gregarious habit, and therefore it is of advantage to be visible from
+as great a distance as possible. That so many species have been able
+to give rise to white varieties does not depend on a special
+sensitiveness of the skin to the influence of cold, but to the fact
+that Mammals and Birds have a general tendency to vary towards white.
+Even with us, many birds--starlings, blackbirds, swallows,
+etc.--occasionally produce white individuals, but the white variety
+does not persist, because it readily falls a victim to the carnivores.
+This is true of white fawns, foxes, deer, etc. The whiteness,
+therefore, arises from internal causes, and only persists when it is
+useful. A great many animals living in a _green environment_ have
+become clothed in green, especially insects, caterpillars, and
+Mantidae, both persecuted and persecutors.
+
+That it is not the direct effect of the environment which calls forth
+the green colour is shown by the many kinds of caterpillar which rest
+on leaves and feed on them, but are nevertheless brown. These feed by
+night and betake themselves through the day to the trunk of the tree,
+and hide in the furrows of the bark. We cannot, however, conclude
+from this that they were _unable_ to vary towards green, for there are
+Arctic animals which are white only in winter and brown in summer
+(Alpine hare, and the ptarmigan of the Alps), and there are also green
+leaf-insects which remain green only while they are young and
+difficult to see on the leaf, but which become brown again in the last
+stage of larval life, when they have outgrown the leaf. They then
+conceal themselves by day, sometimes only among withered leaves on the
+ground, sometimes in the earth itself. It is interesting that in one
+genus, Chaerocampa, one species is brown in the last stage of larval
+life, another becomes brown earlier, and in many species the last
+stage is not wholly brown, a part remaining green. Whether this is a
+case of a double adaptation, or whether the green is being gradually
+crowded out by the brown, the fact remains that the same species, even
+the same individual, can exhibit both variations. The case is the same
+with many of the leaf-like Orthoptera, as, for instance, the praying
+mantis (_Mantis religiosa_) which we have already mentioned.
+
+But the best proofs are furnished by those of ten-cited cases in which
+the insect bears a deceptive resemblance to another object. We now
+know many such cases, such as the numerous imitations of green or
+withered leaves, which are brought about in the most diverse ways,
+sometimes by mere variations in the form of the insect and in its
+colour, sometimes by an elaborate marking, like that which occurs in
+the Indian leaf-butterflies, _Kallima inachis_. In the single
+butterfly-genus Anaea, in the woods of South America, there are about
+a hundred species which are all gaily coloured on the upper surface,
+and on the reverse side exhibit the most delicate imitation of the
+colouring and pattern of a leaf, generally without any indication of
+the leaf-ribs, but extremely deceptive nevertheless. Anyone who has
+seen only one such butterfly may doubt whether many of the
+insignificant details of the marking can really be of advantage to the
+insect. Such details are for instance the apparent holes and splits
+in the apparently dry or half-rotten leaf, which are usually due to
+the fact that the scales are absent on a circular or oval patch so
+that the colourless wing-membrane lies bare, and one can look through
+the spot as through a window. Whether the bird which is seeking or
+pursuing the butterflies takes these holes for dewdrops, or for the
+work of a devouring insect, does not affect the question; the
+mirror-like spot undoubtedly increases the general deceptiveness, for
+the same thing occurs in many leaf-butterflies, though not in all, and
+in some cases it is replaced in quite a peculiar manner. In one
+species of Anaea (_A. divina_), the resting butterfly looks exactly
+like a leaf out of the outer edge of which a large semi-circular piece
+has been eaten, possibly by a caterpillar; but if we look more closely
+it is obvious that there is no part of the wing absent, and that the
+semi-circular piece is of a clear, pale yellow colour, while the rest
+of the wing is of a strongly contrasted dark brown.
+
+But the deceptive resemblance may be caused in quite a different
+manner. I have often speculated as to what advantage the brilliant
+white C could give to the otherwise dusky-coloured "Comma butterfly"
+(_Grapta C. album_). Poulton's recent observations[46] have shown that
+this represents the imitation of a crack such as is often seen in dry
+leaves, and is very conspicuous because the light shines through it.
+
+The utility obviously lies in presenting to the bird the very familiar
+picture of a broken leaf with a clear shining slit, and we may
+conclude, from the imitation of such small details, that the birds are
+very sharp observers and that the smallest deviation from the usual
+arrests their attention and incites them to closer investigation. It
+is obvious that such detailed--we might almost say such
+subtle--deceptive resemblances could only have come about in the
+course of long ages through the acquirement from time to time of
+something new which heightened the already existing resemblance.
+
+In face of facts like these there can be no question of chance and no
+one has succeeded so far in finding any other explanation to replace
+that by selection. For the rest, the apparent leaves are by no means
+perfect copies of a leaf; many of them only represent the torn or
+broken piece, or the half or two-thirds of a leaf, but then the leaves
+themselves frequently do not present themselves to the eye as a whole,
+but partially concealed among other leaves. Even those butterflies
+which, like the species of Kallima and Anaea, represent the whole of a
+leaf with stalk, ribs, apex, and the whole breadth, are not actual
+copies which would satisfy a botanist; there is often much wanting. In
+Kallima the lateral ribs of the leaf are never all included in the
+markings; there are only two or three on the left side and at more
+four or five on the right, and in many individuals these are rather
+obscure, while in others they are comparatively distinct. This
+furnishes us with fresh evidence in favour of their origin through
+processes of selection, for a botanically perfect picture could not
+arise in this way; there could only be a fixing of such details as
+heightened the deceptive resemblance.
+
+Our postulate of origin through selection also enables us to
+understand why the leaf-imitation is on the lower surface of the wing
+in the diurnal Lepidoptera, and on the upper surface in the nocturnal
+forms, corresponding to the attitude of the wings in the resting
+position of the two groups.
+
+The strongest of all proofs of the theory, however, is afforded by
+cases of true "mimicry," those adaptations discovered by Bates in
+1861, consisting in the imitation of one species by another, which
+becomes more and more like its model. The model is always a species
+that enjoys some special protection from enemies, whether because it
+is unpleasant to taste, or because it is in some way dangerous.
+
+It is chiefly among insects and especially among butterflies that we
+find the greatest number of such cases. Several of these have been
+minutely studied and every detail has been investigated so that it is
+difficult to understand how there can still be disbelief in regard to
+them. If the many and exact observations which have been carefully
+collected and critically discussed for instance by Poulton[47] were
+thoroughly studied the arguments which are still frequently urged
+against mimicry would be found untenable; we can hardly hope to find
+more convincing proof of the actuality of the processes of selection
+than these cases put into our hands. The preliminary postulates of the
+theory of mimicry have been disputed, for instance, that diurnal
+butterflies are persecuted and eaten by birds, but observations
+specially directed towards this point in India, Africa, America and
+Europe have placed it beyond all doubt. If it were necessary I could
+myself furnish an account of my own observations on this point.
+
+In the same way it has been established by experiment and observation
+in the field that in all the great regions of distribution there are
+butterflies which are rejected by birds and lizards, their chief
+enemies, on account of their unpleasant smell or taste. These
+butterflies are usually gaily and conspicuously coloured and thus--as
+Wallace first interpreted it--are furnished with an easily
+recognisable sign: a sign of unpalatableness or _warning colours_. If
+they were not thus recognisable easily and from a distance, they would
+frequently be pecked at by birds, and then rejected because of their
+unpleasant taste; but as it is, the insect-eaters recognise them at
+once as unpalatable booty and ignore them. Such _immune_[48] species,
+wherever they occur, are imitated by other palatable species, which
+thus acquire a certain degree of protection.
+
+It is true that this explanation of the bright, conspicuous colours
+is only a hypothesis, but its foundations--unpalatableness, and the
+liability of other butterflies to be eaten,--are certain, and its
+consequences--the existence of mimetic palatable forms--conform it in
+the most convincing manner. Of the many cases now known I select one,
+which is especially remarkable, and which has been thoroughly
+investigated, _Papilla dardanus_ (_merope_), a large, beautiful,
+diurnal butterfly which ranges from Abyssinia throughout the whole of
+Africa to the south coast of Cape Colony.
+
+The males of this form are everywhere _almost_ the same in colour and
+in form of wings, save for a few variations in the sparse black
+markings on the pale yellow ground. But the females occur in several
+quite different forms and colourings, and one of these only, the
+Abyssinian form, is like the male, while the other three or four are
+_mimetic_, that is to say, they copy a butterfly of quite a different
+family the Danaids, which are among the _immune_ forms. In each region
+the females have thus copied two or three different immune species.
+There is much that is interesting to be said in regard to these
+species, but it would be out of keeping with the general tenor of this
+paper to give details of this very complicated case of polymorphism in
+_P. Dardanus_. Anyone who is interested in the matter will find a full
+and exact statement of the case in as far as we know it, in Poulton's
+_Essays on Evolution_ (pp. 373-375[49]). I need only add that three
+different mimetic female forms have been reared from the eggs of a
+single female in South Africa. The resemblance of the forms to their
+immune models goes so far that even the details of the _local_ forms
+of the models are copied by the mimetic species.
+
+It remains to be said that in Madagascar a butterfly,
+
+_Papilio meriones_, occurs, of which both sexes are very similar in
+form and markings to the non-mimetic male of _P. dardanus_, so that it
+probably represents the ancestor of this latter species.
+
+In face of such facts as these every attempt at another explanation
+must fail. Similarly all the other details of the case fulfil the
+preliminary postulates of selection, and leave no room for any other
+interpretation. That the males do not take on the protective colouring
+is easily explained, because they are in general more numerous, and
+the females are more important for the preservation of the species,
+and must also live longer in order to deposit their eggs. We find the
+same state of things in many other species, and in one case (_Elymnias
+undularis_) in which the male is also mimetically coloured, it copies
+quite a differently coloured immune species from the model followed by
+the female. This is quite intelligible when we consider that if there
+were _too many_ false immune types, the birds would soon discover that
+there were palatable individuals among those with unpalatable warning
+colours. Hence the imitation of different immune species by _Papilio
+dardanus_!
+
+I regret that lack of space prevents my bringing forward more examples
+of mimicry and discussing them fully. But from the case of _Papilio
+dardanus_ alone there is much to be learnt which is of the highest
+importance for our understanding of transformations. It shows us
+chiefly what I once called, somewhat strongly perhaps, _the
+omnipotence of natural selection_ in answer to an opponent who had
+spoken of its "inadequacy." We here see that one and the same species
+is capable of producing four or five different patterns of colouring
+and marking; thus the colouring and marking are not, as has often been
+supposed, a necessary outcome of the specific nature of the species,
+but a true adaptation, which cannot arise as a direct effect of
+climatic conditions, but solely through what I may call the sorting
+out of the variations produced by the species, according to their
+utility. That caterpillars may be either green or brown is already
+something more than could have been expected according to the old
+conception of species, but that one and the same butterfly should be
+now pale yellow, with black; now red with black and pure white; now
+deep black with large, pure white spots; and again black with a large
+ocheous-yellow spot, and many small white and yellow spots; that in
+one sub-species it may be tailed like the ancestral form, and in
+another tailless like its Danaid model,--all this shows a far-reaching
+capacity for variation and adaptation that we could never have
+expected if we did not see the facts before us. How it is possible
+that the primary colour-variations should thus be intensified and
+combined remains a puzzle even now; we are reminded of the modern
+three-colour printing,--perhaps similar combinations of the primary
+colours take place in this case; in any case the direction of these
+primary variations is determined by the artist whom we know as natural
+selection, for there is no other conceivable way in which the model
+could affect the butterfly that is becoming more and more like it. The
+same climate surrounds all four forms of female; they are subject to
+the same conditions of nutrition. Moreover, _Papilio dardanus_ is by
+no means the only species of butterfly which exhibits different kinds
+of colour-pattern on its wings. Many species of the Asiatic genus
+Elymnias have on the upper surface a very good imitation of an immune
+Euploeine (Danainae), often with a steel-blue ground-colour, while the
+under surface is well concealed when the butterfly is at rest,--thus
+there are two kinds of protective coloration each with a different
+meaning! The same thing may be observed in many non-mimetic
+butterflies, for instance in all our species of Vanessa, in which the
+under side shows a grey-brown or brownish-black protective coloration,
+but we do not yet know with certainty what may be the biological
+significance of the gaily coloured upper surface.
+
+In general it may be said that mimetic butterflies are comparatively
+rare species, but there are exceptions, for instance _Limenitis
+archippus_ in North America, of which the immune model (_Danaida
+plexippus_) also occurs in enormous numbers.
+
+In another mimicry-category the imitators are often more numerous than
+the models, namely in the case of the imitation of _dangerous insects_
+by harmless species. Bees and wasps are dreaded for their sting, and
+they are copied by harmless flies of the genera Eristalis and Syrphus,
+and these mimics often occur in swarms about flowering plants without
+damage to themselves or to their models; they are feared and are
+therefore left unmolested.
+
+In regard also to the _faithfulness of the copy_ the facts are quite
+in harmony with the theory, according to which the resemblance must
+have arisen and increased _by degrees_. We can recognise this in many
+cases, for even now the mimetic species show very _varying degrees of
+resemblance_ to their immune model. If we compare, for instance, the
+many different imitators of _Danaida chrysippus_ we find that, with
+their brownish-yellow ground-colour, and the position and size, and
+more or less sharp limitation of their clear marginal spots, they have
+reached very different degrees of nearness to their model. Or compare
+the female of _Elymnias undularis_ with its model _Danaida genutia_;
+there is a general resemblance, but the marking of the Danaida is very
+roughly imitated in Elymnias.
+
+Another fact that bears out the theory of mimicry is, that even when
+the resemblance in colour-pattern is very great, the _wing-venation_,
+which is so constant, and so important in determining the systematic
+position of butterflies, is never affected by the variation. The
+pursuers of the butterfly have no time to trouble about entomological
+intricacies.
+
+I must not pass over a discovery of Poulton's which is of great
+theoretical importance--that mimetic butterflies may reach the same
+effect by very different means.[50] Thus the glass-like transparency
+of the wing of a certain Ithomiine (Methona) and its Pierine mimic
+(_Dismorphia orise_) depends on a diminution in the size of the
+scales; in the Danaine genus Itune it is due to the fewness of the
+scales and in a third imitator, a moth (_Castnia linus var.
+heliconoides_) the glass-like appearance of the wing is due neither to
+diminution nor to absence of scales, but to their absolute
+colourlessness and transparency, and to the fact that they stand
+upright. In another moth mimic (Anthomyza) the arrangement of the
+transparent scales is normal. Thus it is not some unknown external
+influence that has brought about the transparency of the wing in these
+five forms, as has sometimes been supposed. Nor is it a hypothetical
+_internal_ evolutionary tendency, for all three vary in a different
+manner. The cause of this agreement can only lie in selection, which
+preserves and intensifies in each species the favourable variations
+that present themselves. The great faithfulness of the copy is
+astonishing in these cases, for it is not _the whole_ wing which is
+transparent; certain markings are black in colour, and these contrast
+sharply with the glass-like ground. It is obvious that the pursuers of
+these butterflies must be very sharp-sighted, for otherwise the
+agreement between the species could never have been pushed so far. The
+less the enemies see and observe, the more defective must the
+imitation be, and if they had been blind, no visible resemblance
+between the species which required protection could ever have arisen.
+
+A seemingly irreconcilable contradiction to the mimicry theory is
+presented in the following cases, which were known to Bates, who,
+however, never succeeded in bringing them into line with the principle
+of mimicry.
+
+In South America there are, as we have already said, many mimics of
+the immune Ithomiinae (or as Bates called them Heliconidae). Among
+these there occur not merely species which are edible, and thus
+require the protection of a disguise, but others which are rejected on
+account of their unpalatableness. How could the Ithomiine dress have
+developed in their case, and of what use is it, since the species
+would in any case be immune? In Eastern Brazil, for instance, there
+are four butterflies, which bear a most confusing resemblance to one
+another in colour, marking, and form of wing, and all four are
+unpalatable to birds. They belong to four different genera and three
+sub-families, and we have to inquire: Whence came this resemblance and
+what end does it serve? For a long time no satisfactory answer could
+be found, but Fritz Müller,[51] seventeen years after Bates, offered a
+solution to the riddle, when he pointed out that young birds could not
+have an instinctive knowledge of the unpalatableness of the
+Ithomiines, but must learn by experience which species were edible and
+which inedible. Thus each young bird must have tasted at least one
+individual of each inedible species and discovered its unpalatability,
+before it learnt to avoid, and thus to spare the species. But if the
+four species resemble each other very closely the bird will regard
+them all as of the same kind, and avoid them all. Thus there developed
+a process of selection which resulted in the survival of the
+Ithomiine-like individuals, and in so great an increase of resemblance
+between the four species, that they are difficult to distinguish one
+from another even in a collection. The advantage for the four species,
+living side by side as they do e.g. in Bahia, lies in the fact that
+only one individual from the _mimicry-ring_ ("inedible association")
+need be tasted by a young bird, instead of at least four individuals,
+as would otherwise be the case. As the number of young birds is great,
+this makes a considerable difference in the ratio of elimination. The
+four Brazilian species are _Lycorea halia_ (Danainae), _Heliconius
+narcaea_ (_eucrate_) (Heliconinae), _Melinaea ethra_, and _Mechanitis
+lysimnia_ (Ithomiinae).
+
+These interesting mimicry-rings (trusts), which have much significance
+for the theory, have been the subject of numerous and careful
+investigations, and at least their essential features are now fully
+established. Müller took for granted, without making any
+investigations, that young birds only learn by experience to
+distinguish between different kinds of victims. But Lloyd Morgan's[52]
+experiments with young birds proved that this is really the case, and
+at the same time furnished an additional argument against the
+_Lamarckian principle_.
+
+In addition to the mimicry-rings first observed in South America,
+others have been described from Tropical India by Moore, and by
+Poulton and Dixey from Africa, and we may expect to learn many more
+interesting facts in this connection. Here again the preliminary
+postulates of the theory are satisfied. And how much more that would
+lead to the same conclusion might be added!
+
+As in the case of mimicry many species have come to resemble one
+another through processes of selection, so we know whole classes of
+phenomena in which plants and animals have become adapted to one
+another, and have thus been modified to a considerable degree. I refer
+particularly to the relation between flowers and insects. Darwin has
+shown that the originally inconspicuous blossoms of the phanerogams
+were transformed into flowers through the visits of insects, and that,
+conversely, several large orders of insects have been gradually
+modified by their association with flowers, especially as regards the
+parts of their body actively concerned. Bees and butterflies in
+particular have become what they are through their relation to
+flowers. In this case again all that is apparently contradictory to
+the theory can, on closer investigation, be beautifully interpreted in
+corroboration of it. Selection can give rise only to what is of use to
+the organism actually concerned, never to what is of use to some other
+organism, and we must therefore expect to find that in flowers only
+characters of use to _themselves_ have arisen, never characters which
+are of use to insects only, and conversely that in the insects
+characters useful to them and not merely to the plants would have
+originated. For a long time it seemed as if an exception to this rule
+existed in the case of the fertilisation of the yucca blossoms by a
+little moth, _Pronuba yuccasella_. This little moth has a
+sickle-shaped appendage to its mouth-parts which occurs hi no other
+Lepidopteron, and which is used for pushing the yellow pollen into the
+opening of the pistil, thus fertilising the flower. Thus it appears as
+if a new structure, which is useful only to the plant, has arisen in
+the insect. But the difficulty is solved as soon as we learn that the
+moth lays its eggs in the fruit-buds of the Yucca, and that the
+larvae, when they emerge, feed on the developing seeds. In effecting
+the fertilisation of the flower the moth is at the same time making
+provision for its own offspring, since it is only after fertilisation
+that the seeds begin to develop. There is thus nothing to prevent our
+referring this structural adaptation in _Pronuba yuccasella_ to
+processes of selection, which have gradually transformed the maxillary
+palps of the female into the sickle-shaped instrument for collecting
+the pollen, and which have at the same time developed in the insect
+the instinct to press the pollen into the pistil.
+
+In this domain, then, the theory of selection finds nothing but
+corroboration, and it would be impossible to substitute for it any
+other explanation, which now that the facts are so well known, could
+be regarded as a serious rival to it. That selection is a factor, and
+a very powerful factor in the evolution of organisms, can no longer be
+doubted. Even although we cannot bring forward formal proofs of it _in
+detail_, cannot calculate definitely the size of the variations which
+present themselves, and their selection-value, cannot, in short,
+reduce the whole process to a mathematical formula, yet we must assume
+selection, because it is the only possible explanation applicable to
+whole classes of phenomena, and because, on the other hand, it is made
+up of factors which we know can be proved actually to exist, and
+which, _if_ they exist, must of logical necessity coöperate in the
+manner required by the theory. _We must accept it because the
+phenomena of evolution and adaptation must have a natural basis, and
+because it is the only possible explanation of them._[53]
+
+Many people are willing to admit that selection explains adaptations,
+but they maintain that only a part of the phenomena are thus
+explained, because everything does not depend upon adaptation. They
+regard adaptation as, so to speak, a special effort on the part of
+Nature, which she keeps in readiness to meet particularly difficult
+claims of the external world on organisms. But if we look at the
+matter more carefully we shall find that adaptations are by no means
+exceptional, but that they are present everywhere in such enormous
+numbers, that it would be difficult in regard to any structure
+whatever, to prove that adaptation had _not_ played a part in its
+evolution.
+
+How often has the senseless objection been urged against selection
+that it can create nothing, it can only reject. It is true that it
+cannot create either the living substance or the variations of it;
+both must be given. But in rejecting one thing it preserves another,
+intensifies it, combines it, and in this way _creates_ what is new.
+_Everything_ in organisms depends on adaptation; that is to say,
+everything must be admitted through the narrow door of selection,
+otherwise it can take no part in the building up of the whole. But, it
+is asked, what of the direct effect of external conditions,
+temperature, nutrition, climate and the like? Undoubtedly these can
+give rise to variations, but they too must pass through the door of
+selection, and if they cannot do this they are rejected, eliminated
+from the constitution of the species.
+
+It may, perhaps, be objected that such external influences are often
+of a compelling power, and that every animal must submit to them, and
+that thus selection has no choice and can neither select nor reject.
+There may be such cases; let us assume for instance that the effect
+of the cold of the Arctic regions was to make all the mammals become
+black; the result would be that they would all be eliminated by
+selection, and that no mammals would be able to live there at all. But
+in most cases a certain percentage of animals resists these strong
+influences, and thus selection secures a foothold on which to work,
+eliminating the unfavourable variation, and establishing a useful
+colouring, consistent with what is required for the maintenance of the
+species.
+
+Everything depends upon adaptation! We have spoken much of adaptation
+in colouring, in connection with the examples brought into prominence
+by Darwin, because these are conspicuous, easily verified, and at the
+same time convincing for the theory of selection. But is it only
+desert and polar animals whose colouring is determined through
+adaptation? Or the leaf-butterflies, and the mimetic species, or the
+terrifying markings, and "warning-colours" and a thousand other kinds
+of sympathetic colouring? It is, indeed, never the colouring alone
+which makes up the adaptation; the structure of the animal plays a
+part, often a very essential part, in the protective disguise, and
+thus _many_ variations may cooperate towards _one_ common end. And it
+is to be noted that it is by no means only external parts that are
+changed; internal parts are _always_ modified at the same time--for
+instance, the delicate elements of the nervous system on which depend
+the _instinct_ of the insect to hold its wings, when at rest, in a
+perfectly definite position, which, in the leaf-butterfly, has the
+effect of bringing the two pieces on which the marking occurs on the
+anterior and posterior wing into the same direction, and thus
+displaying as a whole the fine curve of the midrib on the seeming
+leaf. But the wing-holding instinct is not regulated in the same way
+in all leaf-butterflies; even our indigenous species of Vanessa, with
+their protective ground-colouring, have quite a distinctive way of
+holding their wings so that the greater part of the anterior wing is
+covered by the posterior when the butterfly is at rest. But the
+protective colouring appears on the posterior wing and on the tip of
+the anterior, _to precisely the distance to which it is left
+uncovered_. This occurs, as Standfuss has shown, in different degrees
+in our two most nearly allied species, the uncovered portion being
+smaller in _V. urticae_ than in _V. polychloros_. In this case, as in
+most leaf-butterflies, the holding of the wing was probably the
+primary character; only after that was thoroughly established did the
+protective marking develop. In any case, the instinctive manner of
+holding the wings is associated with the protective colouring, and
+must remain as it is if the latter is to be effective. How greatly
+instincts may change, that is to say, may be adapted, is shown by the
+case of the Noctuid "shark" moth, _Xylina vetusta_. This form bears a
+most deceptive resemblance to a piece of rotten wood, and the
+appearance is greatly increased by the modification of the innate
+impulse to flight common to so many animals, which has here been
+transformed into an almost contrary instinct. This moth does not fly
+away from danger, but "feigns death," that is, it draws antennae, legs
+and wings close to the body, and remains perfectly motionless. It may
+be touched, picked up, and thrown down again, and still it does not
+move. This remarkable instinct must surely have developed
+simultaneously with the wood-colouring; at all events, both
+coöperating variations are now present, and prove that both the
+external and the most minute internal structure have undergone a
+process of adaptation.
+
+The case is the same with all structural variations of animal parts,
+which are not absolutely insignificant. When the insects acquired
+wings they must also have acquired the mechanism with which to move
+them--the musculature, and the nervous apparatus necessary for its
+automatic regulation. All instincts depend upon compound reflex
+mechanisms and are just as indispensable as the parts they have to set
+in motion, and all may have arisen through processes of selection if
+the reasons which I have elsewhere given for this view are
+correct.[54]
+
+Thus there is no lack of adaptations within the organism, and
+particularly in its most important and complicated parts, so that we
+may say that there is no actively functional organ that has not
+undergone a process of adaptation relative to its function and the
+requirements of the organism. Not only is every gland structurally
+adapted, down to the very minutest histological details, to its
+function, but the function is equally minutely adapted to the needs of
+the body. Every cell in the mucous lining of the intestine is exactly
+regulated in its relation to the different nutritive substances, and
+behaves in quite a different way towards the fats, and towards
+nitrogenous substances, or peptones.
+
+I have elsewhere called attention to the many adaptations of the whale
+to the surrounding medium, and have pointed out--what has long been
+known, but is not universally admitted, even now--that in it a great
+number of important organs have been transformed in adaptation to the
+peculiar conditions of aquatic life, although the ancestors of the
+whale must have lived, like other hair-covered mammals, on land. I
+cited a number of these transformations--the fish-like form of the
+body, the hairlessness of the skin, the transformation of the
+fore-limbs to fins, the disappearance of the hind-limbs and the
+development of a tail fin, the layer of blubber under the skin, which
+affords the protection from cold necessary to a warm-blooded animal,
+the disappearance of the ear-muscles and the auditory passages, the
+displacement of the external nares to the forehead for the greater
+security of the breathing-hole during the brief appearance at the
+surface, and certain remarkable changes in the respiratory and
+circulatory organs which enable the animal to remain for a long time
+under water. I might have added many more, for the list of adaptations
+in the whale to aquatic life is by no means exhausted; they are found
+in the histological structure and in the minutest combinations in the
+nervous system. For it is obvious that a tail-fin must be used in
+quite a different way from a tail, which serves as a fly-brush in
+hoofed animals, or as an aid to springing in the kangaroo or as a
+climbing organ; it will require quite different reflex-mechanisms and
+nerve combinations in the motor centres.
+
+I used this example in order to show how unnecessary it is to assume a
+special internal evolutionary power for the phylogenesis of species,
+for this whole order of whales is, so to speak, _made up of
+adaptations_; it deviates in many essential respects from the usual
+mammalian type, and all the deviations are adaptations to aquatic
+life. But if precisely the most essential features of the organisation
+thus depend upon adaptation, what is left for a phyletic force to do,
+since it is these essential features of the structure it would have to
+determine? There are few people now who believe in a phyletic
+evolutionary power, which is not made up of the forces known to
+us--adaptation and heredity--but the conviction that _every_ part of
+an organism depends upon adaptation has not yet gained a firm footing.
+Nevertheless, I must continue to regard this conception as the correct
+one, as I have long done.
+
+I may be permitted one more example. The feather of a bird is a
+marvellous structure, and no one will deny that as a whole it depends
+upon adaptation. But what part of it _does not_ depend upon
+adaptation? The hollow quill, the shaft with its hard, thin, light
+cortex, and the spongy substance within it, its square section
+compared with the round section of the quill, the flat barbs, their
+short, hooked barbules which, in the flight-feathers, hook into one
+another with just sufficient firmness to resist the pressure of the
+air at each wing-beat, the lightness and firmness of the whole
+apparatus, the elasticity of the vane, and so on. And yet all this
+belongs to an organ which is only passively functional, and therefore
+can have nothing to do with the _Lamarckian principle_. Nor can the
+feather have arisen through some magical effect of temperature,
+moisture, electricity, or specific nutrition, and thus selection is
+again our only anchor of safety.
+
+But--it will be objected--the substance of which the feather consists,
+this peculiar kind of horny substance, did not first arise through
+selection in the course of the evolution of the birds, for it formed
+the covering of the scales of their reptilian ancestors. It is quite
+true that a similar substance covered the scales of the Reptiles, but
+why should it not have arisen among them through selection? Or in what
+other way could it have arisen, since scales are also passively useful
+parts? It is true that if we are only to call adaptation what has been
+acquired by the species we happen to be considering, there would
+remain a great deal that could not be referred to selection; but we
+are postulating an evolution which has stretched back through aeons,
+and in the course of which innumerable adaptations took place, which
+had not merely ephemeral persistence in a genus, a family or a class,
+but which was continued into whole Phyla of animals, with continual
+fresh adaptations to the special conditions of each species, family,
+or class, yet with persistence of the fundamental elements. Thus the
+feather, once acquired, persisted in all birds, and the vertebral
+column, once gained by adaptation in the lowest forms, has persisted
+in all the Vertebrates from Amphioxus upwards, although with constant
+readaptation to the conditions of each particular group. Thus
+everything we can see in animals is adaptation, whether of to-day, or
+of yesterday, or of ages long gone by; every kind of cell, whether
+glandular, muscular, nervous, epidermic, or skeletal, is adapted to
+absolutely definite and specific functions, and every organ which is
+composed of these different kinds of cells contains them in the proper
+proportions, and in the particular arrangement which best serves the
+function of the organ; it is thus adapted to its function.
+
+All parts of the organism are tuned to one another, that is, _they are
+adapted to one another_, and in the same way _the organism as a whole
+is adapted to the conditions of its life, and it is so at every stage
+of its evolution._
+
+But all adaptations _can_ be referred to selection; the only point
+that remains doubtful is whether they all _must_ be referred to it.
+
+However that may be, whether the _Lamarckian principle_ is a factor
+that has coöperated with selection in evolution, or whether it is
+altogether fallacious, the fact remains, that selection is the cause
+of a great part of the phyletic evolution of organisms on our earth.
+Those who agree with me in rejecting the _Lamarckian principle_ will
+regard selection as the only _guiding_ factor in evolution, which
+creates what is new out of the transmissible variations, by ordering
+and arranging these, selecting them in relation to their number and
+size, as the architect does his building-stones so that a particular
+style must result.[55] But the building-stones themselves, the
+variations, have their basis in the influences which cause variation
+in those vital units which are handed on from one generation to
+another, whether, taken together they form the _whole_ organism, as in
+Bacteria and other low forms of life, or only a germ-substance, as in
+unicellular and multicellular organisms.
+
+FOOTNOTES:
+
+[Footnote 33: _Vorträge über Descendenztheorie_, Jena, 1904, II. 269.
+Eng. Transl. London, 1904, II. p. 317.]
+
+[Footnote 34: See Poulton, _Essays on Evolution_, Oxford, 1908. pp.
+xix-xxii.]
+
+[Footnote 35: _Origin of Species_ (6th edit), pp. 176 _et seq._]
+
+[Footnote 36: Chun, _Reise der Valdivia_, Leipzig, 1904.]
+
+[Footnote 37: Plate, _Selektionsprinzip u. Probleme der Artbildung_
+(3rd edit.), Leipzig, 1908.]
+
+[Footnote 38: _Studien zur Descendenz-Theorie_ II., "Die Enstehung der
+Zeichnung bei den Schmetterlings-raupen," Leipzig, 1876.]
+
+[Footnote 39: _Origin of Species_ (6th edit.), p. 232.]
+
+[Footnote 40: _Origin of Species_, p. 233; see also edit. 1, p. 242.]
+
+[Footnote 41: _Ibid._ p. 230.]
+
+[Footnote 42: _The Effect of External Influences upon Development_,
+Romanes Lecture, Oxford, 1894.]
+
+[Footnote 43: See Poulton, _Essays on Evolution_, 1908, pp. 316, 317.]
+
+[Footnote 44: _The Evolution Theory_, London, 1904, I. p. 219.]
+
+[Footnote 45: _Report of the British Association_ (Bristol, 1898),
+London, 1899, pp. 906-909.]
+
+[Footnote 46: _Proc. Ent. Soc._, London, May 6, 1903.]
+
+[Footnote 47: _Essays on Evolution_, 1889-1907, Oxford, 1908,
+_passim_, e.g. p. 269.]
+
+[Footnote 48: The expression does not refer to all the enemies of this
+butterfly; against ichneumon-flies, for instance, their unpleasant
+smell usually gives no protection.]
+
+[Footnote 49: Professor Poulton has corrected some wrong descriptions
+which I had unfortunately overlooked in the Plates of my book
+_Vorträge über Descendenztheorie_, and which refer to _Papilio
+dardanus_ (_merope_). These mistakes are of no importance as far as an
+understanding of the mimicry-theory is concerned, but I hope shortly
+to be able to correct them in a later edition.]
+
+[Footnote 50: _Journ. Linn. Soc. London_ (_Zool._), Vol. xxvi. 1898,
+pp. 598-602.]
+
+[Footnote 51: In _Kosmos_, 1879, p. 100.]
+
+[Footnote 52: _Habit and Instinct_, London. 1896.]
+
+[Footnote 53: This has been discussed in many of my earlier works. See
+for instance _The All-Sufficiency of Natural Selection, a reply to
+Herbert Spencer_, London, 1893.]
+
+[Footnote 54: _The Evolution Theory_, London, 1904, p. 144.]
+
+[Footnote 55: _Variation under Domestication_, 1875, II. pp. 426,
+427.]
+
+
+
+
+III
+
+HEREDITY AND VARIATION IN MODERN LIGHTS
+
+BY W. BATESON, M.A., F.R.S.
+
+_Professor of Biology in the University of Cambridge_
+
+
+Darwin's work has the property of greatness in that it may be admired
+from more aspects than one. For some the perception of the principle
+of Natural Selection stands out as his most wonderful achievement to
+which all the rest is subordinate. Others, among whom I would range
+myself, look up to him rather as the first who plainly distinguished,
+collected, and comprehensively studied that new class of evidence from
+which hereafter a true understanding of the process of Evolution may
+be developed. We each prefer our own standpoint of admiration; but I
+think that it will be in their wider aspect that his labours will most
+command the veneration of posterity.
+
+A treatise written to advance knowledge may be read in two moods. The
+reader may keep his mind passive, willing merely to receive the
+impress of the writer's thought; or he may read with his attention
+strained and alert, asking at every instant how the new knowledge can
+be used in a further advance, watching continually for fresh footholds
+by which to climb higher still. Of Shelley it has been said that he
+was a poet for poets: so Darwin was a naturalist for naturalists. It
+is when his writings are used in the critical and more exacting spirit
+with which we test the outfit for our own enterprise that we learn
+their full value and strength. Whether we glance back and compare his
+performance with the efforts of his predecessors, or look forward
+along the course which modern research is disclosing, we shall honour
+most in him not the rounded merit of finite accomplishment, but the
+creative power by which he inaugurated a line of discovery endless in
+variety and extension. Let us attempt thus to see his work in true
+perspective between the past from which it grew, and the present which
+is its consequence. Darwin attacked the problem of Evolution by
+reference to facts of three classes: Variation; Heredity; Natural
+Selection. His work was not as the laity suppose, a sudden and
+unheralded revelation, but the first fruit of a long and hitherto
+barren controversy. The occurrence of variation from type, and the
+hereditary transmission of such variation had of course been long
+familiar to practical men, and inferences as to the possible bearing
+of those phenomena on the nature of specific difference had been from
+time to time drawn by naturalists. Maupertuis, for example, wrote: "Ce
+qui nous reste à examiner, c'est comment d'un seul individu, il a pu
+naître tant d'espèces si différentes." And again: "La Nature contient
+le fonds de toutes ces variétés: mais le hasard ou l'art les mettent
+en oeuvre. C'est ainsi que ceux dont l'industrie s'applique à
+satisfaire le goût des curieux, sont, pour ainsi dire, créateurs
+d'espèces nouvelles."[56]
+
+Such passages, of which many (though few so emphatic) can be found in
+eighteenth century writers, indicate a true perception of the mode of
+Evolution. The speculations hinted at by Buffon,[57] developed by
+Erasmus Darwin, and independently proclaimed above all by Lamarck,
+gave to the doctrine of descent a wide renown. The uniformitarian
+teaching which Lyell deduced from geological observation had gained
+acceptance. The facts of geographical distribution[58] had been shown
+to be obviously inconsistent with the Mosaic legend. Prichard, and
+Lawrence, following the example of Blumenbach, had successfully
+demonstrated that the races of Man could be regarded as different
+forms of one species, contrary to the opinion up till then received.
+These treatises all begin, it is true, with a profound obeisance to
+the sons of Noah, but that performed, they continue on strictly modern
+lines. The question of the mutability of species was thus prominently
+raised.
+
+Those who rate Lamarck no higher than did Huxley in his contemptuous
+phrase "_buccinator tantum_," will scarcely deny that the sound of the
+trumpet had carried far, or that its note was clear. If then there
+were few who had already turned to evolution with positive conviction,
+all scientific men must at least have known that such views had been
+promulgated; and many must, as Huxley says, have taken up his own
+position of "critical expectancy."[59]
+
+Why, then, was it, that Darwin succeeded where the rest had failed?
+The cause of that success was twofold. First, and obviously, in the
+principle of Natural Selection he had a suggestion which would work.
+It might not go the whole way, but it was true as far as it went.
+Evolution could thus in great measure be fairly represented as a
+consequence of demonstrable processes. Darwin seldom endangers the
+mechanism he devised by putting on it strains much greater than it can
+bear. He at least was under no illusion as to the omnipotence of
+Selection; and he introduces none of the forced pleading which in
+recent years has threatened to discredit that principle.
+
+
+
+For example, in the latest text of the _Origin_[60] we find him
+saying:
+
+     "But as my conclusions have lately been much misrepresented,
+     and it has been stated that I attribute the modification of
+     species exclusively to natural selection, I may be permitted
+     to remark that in the first edition of this work, and
+     subsequently, I placed in a most conspicuous
+     position--namely, at the close of the Introduction--the
+     following words: 'I am convinced that natural selection has
+     been the main but not the exclusive means of modification.'"
+
+But apart from the invention of this reasonable hypothesis, which may
+well, as Huxley estimated, "be the guide of biological and
+psychological speculation for the next three or four generations,"
+Darwin made a more significant and imperishable contribution. Not for
+a few generations, but through all ages he should be remembered as the
+first who showed clearly that the problems of Heredity and Variation
+are soluble by observation, and laid down the course by which we must
+proceed to their solution.[61] The moment of inspiration did not come
+with the reading of Malthus, but with the opening of the "first
+note-book on Transmutation of Species."[62] Evolution is a process of
+Variation and Heredity. The older writers, though they had some vague
+idea that it must be so, did not study Variation and Heredity. Darwin
+did, and so begat not a theory, but a science.
+
+The extent to which this is true, the scientific world is only
+beginning to realise. So little was the fact appreciated in Darwin's
+own time that the success of his writings was followed by an almost
+total cessation of work in that special field. Of the causes which led
+to these remarkable consequences I have spoken elsewhere. They
+proceeded from circumstances peculiar to the time; but whatever the
+causes there is no doubt that this statement of the result is
+historically exact, and those who make it their business to collect
+facts elucidating the physiology of Heredity and Variation are well
+aware that they will find little to reward their quest in the leading
+scientific Journals of the Darwinian epoch.
+
+In those thirty years the original stock of evidence current and in
+circulation even underwent a process of attrition. As in the story of
+the Eastern sage who first wrote the collected learning of the
+universe for his sons in a thousand volumes and by successive
+compression and burning reduced them to one and from this by further
+burning distilled the single ejaculation of the Faith "There is no god
+but God and Mohammed is the Prophet of God," which was all his maturer
+wisdom deemed essential:--so in the books of that period do we find
+the _corpus_ of genetic knowledge dwindle to a few prerogative
+instances and these at last to the brief formula of an unquestioned
+creed.
+
+And yet in all else that concerns biological science this period was,
+in very truth, our Golden Age, when the natural history of the earth
+was explored as never before; morphology and embryology were
+exhaustively ransacked; the physiology of plants and animals began to
+rival chemistry and physics in precision of method and in the rapidity
+of its advances; and the foundations of pathology were laid.
+
+In contrast with this immense activity elsewhere the neglect which
+befel the special physiology of Descent, or Genetics as we now call
+it, is astonishing. This may of course be interpreted as meaning that
+the favoured studies seemed to promise a quicker return for effort,
+but it would be more true to say that those who chose these other
+pursuits did so without making any such comparison; for the idea that
+the physiology of Heredity and Variation was a coherent science,
+offering possibilities of extraordinary discovery, was not present to
+their minds at all. In a word, the existence of such a science was
+well nigh forgotten. It is true that in ancillary periodicals, as for
+example those that treat of entomology or horticulture, or in the
+writings of the already isolated systematists,[63] observations with
+this special bearing were from time to time related, but the class of
+fact on which Darwin built his conceptions of Heredity and Variation
+was not seen in the highways of biology. It formed no part of the
+official curriculum of biological students, and found no place among
+the subjects which their teachers were investigating.
+
+During this period nevertheless one distinct advance was made, that
+with which Weismann's name is prominently connected. In Darwin's
+genetic scheme the hereditary transmission of parental experience and
+its consequences played a considerable role. Exactly how great that
+role was supposed to be, he with his habitual caution refrained from
+specifying, for the sufficient reason that he did not know.
+Nevertheless much of the process of Evolution, especially that by
+which organs have become degenerate and rudimentary, was certainly
+attributed by Darwin to such inheritance, though since belief in the
+inheritance of acquired characters fell into dispute, the fact has
+been a good deal overlooked. The _Origin_ without "use and disuse"
+would be a materially different book. A certain vacillation is
+discernible in Darwin's utterances on this question, and the fact gave
+to the astute Butler an opportunity for his most telling attack. The
+discussion which best illustrates the genetic views of the period
+arose in regard to the production of the rudimentary condition of the
+wings of many beetles in the Madeira group of islands, and by
+comparing passages from the _Origin_[64] Butler convicts Darwin of
+saying first that this condition was in the main the result of
+Selection, with disuse aiding, and in another place that the main
+cause of degeneration was disuse, but that Selection had aided. To
+Darwin however I think the point would have seemed one of dialetics
+merely. To him the one paramount purpose was to show that somehow an
+Evolution by means of Variation and Heredity might have brought about
+the facts observed, and whether they had come to pass in the one way
+or the other was a matter of subordinate concern.
+
+To us moderns the question at issue has a diminished significance. For
+over all such debates a change has been brought by Weismann's
+challenge for evidence that use and disuse have any transmitted
+effects at all. Hitherto the transmission of many acquired
+characteristics had seemed to most naturalists so obvious as not to
+call for demonstration.[65] Weismann's demand for facts in support of
+the main proposition revealed at once that none having real cogency
+could be produced. The time-honoured examples were easily shown to be
+capable of different explanations. A few certainly remain which cannot
+be so summarily dismissed, but--though it is manifestly impossible
+here to do justice to such a subject--I think no one will dispute that
+these residual and doubtful phenomena, whatever be their true nature,
+are not of a kind to help us much in the interpretation of any of
+those complex cases of adaptation which on the hypothesis of unguided
+Natural Selection are especially difficult to understand. Use and
+disuse were invoked expressly to help us over these hard places; but
+whatever changes can be induced in offspring by direct treatment of
+the parents, they are not of a kind to encourage hope of real
+assistance from that quarter. It is not to be denied that through the
+collapse of this second line of argument the Selection hypothesis has
+had to take an increased and perilous burden. Various ways of meeting
+the difficulty have been proposed, but these mostly resolve themselves
+into improbable attempts to expand or magnify the powers of Natural
+Selection.
+
+Weismann's interpellation, though negative in purpose, has had a
+lasting and beneficial effect, for through his thorough demolition of
+the old loose and distracting notions of inherited experience, the
+ground has been cleared for the construction of a true knowledge of
+heredity based on experimental fact.
+
+In another way he made a contribution of a more positive character,
+for his elaborate speculations as to the genetic meaning of
+cytological appearances have led to a minute investigation of the
+visible phenomena occurring in those cell-divisions by which
+germ-cells arise. Though the particular views he advocated have very
+largely proved incompatible with the observed facts of heredity, yet
+we must acknowledge that it was chiefly through the stimulus of
+Weismann's ideas that those advances in cytology were made; and though
+the doctrine of the continuity of germ-plasm cannot be maintained in
+the form originally propounded, it is in the main true and
+illuminating.[66] Nevertheless in the present state of knowledge we
+are still as a rule quite unable to connect cytological appearances
+with any genetic consequences and save in one respect (obviously of
+extreme importance--to be spoken of later) the two sets of phenomena
+might, for all we can see, be entirely distinct.
+
+I cannot avoid attaching importance to this want of connection between
+the nuclear phenomena and the features of bodily organisation. All
+attempts to investigate Heredity by cytological means lie under the
+disadvantage that it is the nuclear changes which can alone be
+effectively observed. Important as they must surely be, I have never
+been persuaded that the rest of the cell counts for nothing. What we
+know of the behaviour and variability of chromosomes seems in my
+opinion quite incompatible with the belief that they alone govern
+form, and are the sole agents responsible in heredity.[67]
+
+If, then, progress was to be made in Genetics, work of a different
+kind was required. To learn the laws of Heredity and Variation there
+is no other way than that which Darwin himself followed, the direct
+examination of the phenomena. A beginning could be made by collecting
+fortuitous observations of this class, which have often thrown a
+suggestive light, but such evidence can be at best but superficial and
+some more penetrating instrument of research is required. This can
+only be provided by actual experiments in breeding.
+
+The truth of these general considerations was becoming gradually clear
+to many of us when in 1900 Mendel's work was rediscovered.
+Segregation, a phenomenon of the utmost novelty, was thus revealed.
+From that moment not only in the problem of the origin of species, but
+in all the great problems of biology a new era began. So unexpected
+was the discovery that many naturalists were convinced it was untrue,
+and at once proclaimed Mendel's conclusions as either altogether
+mistaken, or if true, of very limited application. Many fantastic
+notions about the workings of Heredity had been asserted as general
+principles before: this was probably only another fancy of the same
+class.
+
+Nevertheless those who had a preliminary acquaintance with the facts
+of Variation were not wholly unprepared for some such revelation. The
+essential deduction from the discovery of segregation was that the
+characters of living things are dependent on the presence of definite
+elements or factors, which are treated as units in the processes of
+Heredity. These factors can thus be recombined in various ways. They
+act sometimes separately, and sometimes they interact in conduction
+with each other, producing their various effects. All this indicates a
+definiteness and specific order in heredity, and therefore in
+variation. This order cannot by the nature of the case be dependent on
+Natural Selection for its existence, but must be a consequence of the
+fundamental chemical and physical nature of living things. The study
+of Variation had from the first shown that an orderliness of this kind
+was present. The bodies and the properties of livings things are
+cosmic, not chaotic. No matter how low in the scale we go, never do we
+find the slightest hint of a diminution in that all-pervading
+orderliness, nor can we conceive an organism existing for a moment in
+any other state. Moreover not only does this order prevail in normal
+forms, but again and again it is to be seen in newly-sprung varieties,
+which by general consent cannot have been subjected to a prolonged
+Selection. The discovery of Mendelian elements admirably coincided
+with and at once gave a rationale of these facts. Genetic Variation is
+then primarily the consequence of additions to, or omissions from, the
+stock of elements which the species contains. The further
+investigation of the species-problem must thus proceed by the
+analytical method which breeding experiments provide.
+
+In the nine years which have elapsed since Mendel's clue became
+generally known, progress has been rapid. We now understand the
+process by which a polymorphic race maintains its polymorphism. When a
+family consists of dissimilar members, given the numerical proportions
+in which these members are occurring, we can represent their
+composition symbolically and state what types can be transmitted by
+the various members. The difficulty of the "swamping effects of
+inter-crossing" is practically at an end. Even the famous puzzle of
+sex-limited inheritance is solved, at all events in its more regular
+manifestations, and we know now how it is brought about that the
+normal sisters of a colour-blind man can transmit the colour-blindness
+while his normal brothers cannot transmit it.
+
+We are still only on the fringe of the inquiry. It can be seen
+extending and ramifying in many directions. To enumerate these here
+would be impossible. A whole new range of possibilities is being
+brought into view by study of the inter-relations between the simple
+factors. By following up the evidence as to segregation, indications
+have been obtained which can only be interpreted as meaning that when
+many factors are being simultaneously redistributed among the
+germ-cells, certain of them exert what must be described as a
+repulsion upon other factors. We cannot surmise whither this discovery
+may lead.
+
+In the new light all the old problems wear a fresh aspect. Upon the
+question of the nature of Sex, for example, the bearing of Mendelian
+evidence is close. Elsewhere I have shown that from several sets of
+parallel experiments the conclusion is almost forced upon us that, in
+the types investigated, of the two sexes the female is to be regarded
+as heterozygous in sex, containing one unpaired dominant element,
+while the male is similarly homozygous in the absence of that
+element.[68] It is not a little remarkable that on this point--which
+is the only one where observations of the nuclear processes of
+gameto-genesis have yet been brought into relation with the visible
+characteristics of the organisms themselves--there should be
+diametrical opposition between the results of breeding experiments and
+those derived from cytology.
+
+Those who have followed the researches of the American school will be
+aware that, after it had been found in certain insects that the
+spermatozoa were of two kinds according as they contained or did not
+contain the accessory chromosome, E. B. Wilson succeeded in proving
+that the sperms possessing this accessory body were destined to form
+_females_ on fertilisation, while sperms without it form males, the
+eggs being apparently indifferent. Perhaps the most striking of all
+this series of observations is that lately made by T. H. Morgan,[69]
+since confirmed by von Baehr, that in a Phylloxeran two kinds of
+spermatids are formed, respectively with and without an accessory (in
+this case, _double_) chromosome. Of these, only those possessing the
+accessory body become functional spermatozoa, the others degenerating.
+We have thus an elucidation of the puzzling fact that in these forms
+fertilisation results in the formation of _females_ only. How the
+males are formed--for of course males are eventually produced by the
+parthenogenetic females--we do not know.
+
+If the accessory body is really to be regarded as bearing the factor
+for femaleness, then in Mendelian terms female is DD and male is DR.
+The eggs are indifferent and the spermatozoa are each male, _or_
+female. But according to the evidence derived from a study of the
+sex-limited descent of certain features in other animals the
+conclusion seems equally clear that in them female must be regarded as
+DR and male as RR. The eggs are thus each either male or female and
+the spermatozoa are indifferent. How this contradictory evidence is to
+be reconciled we do not yet know. The breeding work concerns fowls,
+canaries, and the Currant moth (_Abraxas grossulariata_). The
+accessory chromosome has been now observed in most of the great
+divisions of insects,[70] except, as it happens, Lepidoptera. At first
+sight it seems difficult to suppose that a feature apparently so
+fundamental as sex should be differently constituted in different
+animals, but that seems at present the least improbable inference. I
+mention these two groups of facts as illustrating the nature and
+methods of modern genetic work. We must proceed by minute and specific
+analytical investigation. Wherever we look we find traces of the
+operation of precise and specific rules.
+
+In the light of present knowledge it is evident that before we can
+attack the Species-problem with any hope of success there are vast
+arrears to be made up. He would be a bold man who would now assert
+that there was no sense in which the term Species might not have a
+strict and concrete meaning in contradistinction to the term Variety.
+We have been taught to regard the difference between species and
+variety as one of degree. I think it unlikely that this conclusion
+will bear the test of further research. To Darwin the question, What
+is a variation? presented no difficulties. Any difference between
+parent and offspring was a variation. Now we have to be more precise.
+First we must, as de Vries has shown, distinguish real, genetic,
+variation from _fluctuational_ variations, due to environmental and
+other accidents, which cannot be transmitted. Having excluded these
+sources of error the variations observed must be expressed in terms of
+the factors to which they are due before their significance can be
+understood. For example, numbers of the variations seen under
+domestication, and not a few witnessed in nature, are simply the
+consequence of some ingredient being in an unknown way omitted from
+the composition of the varying individual. The variation may on the
+contrary be due to the addition of some new element, but to prove that
+it is so is by no means an easy matter. Casual observation is useless,
+for though these latter variations will always be dominants, yet many
+dominant characteristics may arise from another cause, namely the
+meeting of complementary factors, and special study of each case in
+two generations at least is needed before these two phenomena can be
+distinguished.
+
+When such considerations are fully appreciated it will be realised
+that medleys of most dissimilar occurrences are all confused together
+under the term Variation. One of the first objects of genetic analysis
+is to disentangle this mass of confusion.
+
+To those who have made no study of heredity it sometimes appears that
+the question of the effect of conditions in causing variation is one
+which we should immediately investigate, but a little thought will
+show that before any critical inquiry into such possibilities can be
+attempted, a knowledge of the working of heredity under conditions as
+far as possible uniform must be obtained. At the time when Darwin was
+writing, if a plant brought into cultivation gave off an albino
+variety, such an event was without hesitation ascribed to the change
+of life. Now we see that albino _gametes_, germs, that is to say,
+which are destitute of the pigment-forming factor, may have been
+originally produced by individuals standing an indefinite number of
+generations back in the ancestry of the actual albino, and it is
+indeed almost certain that the variation to which the appearance of
+the albino is due cannot have taken place in a generation later than
+that of the grandparents. It is true that when a new _dominant_
+appears we should feel greater confidence that we were witnessing the
+original variation, but such events are of extreme rarity, and no such
+case has come under the notice of an experimenter in modern times, as
+far as I am aware. That they must have appeared is clear enough.
+Nothing corresponding to the Brown-breasted Game fowl is known wild,
+yet that colour is a most definite dominant, and at some moment since
+_Gallus bankiva_ was domesticated, the element on which that special
+colour depends must have at least once been formed in the germ-cell of
+a fowl; but we need harder evidence than any which has yet been
+produced before we can declare that this novelty came through
+over-feeding, or change of climate, or any other disturbance
+consequent on domestication. When we reflect on the intricacies of
+genetic problems as we must now conceive them there come moments when
+we feel almost thankful that the Mendelian principles were unknown to
+Darwin. The time called for a bold pronouncement, and he made it, to
+our lasting profit and delight. With fuller knowledge we pass once
+more into a period of cautious expectation and reserve.
+
+In every arduous enterprise it is pleasanter to look back at
+difficulties overcome than forward to those which still seem
+insurmountable, but in the next stage there is nothing to be stained
+by disguising the fact that the attributes of living things are not
+what we used to suppose. If they are more complex in the sense that
+the properties they display are throughout so regular[71] that the
+Selection of minute random variations is an unacceptable account of
+the origin of their diversity, yet by virtue of that very regularity
+the problem is limited in scope and thus simplified.
+
+To begin with, we must relegate Selection to its proper place.
+Selection permits the viable to continue and decides that the
+non-viable shall perish; just as the temperature of our atmosphere
+decides that no liquid carbon shall be found on the face of the earth:
+but we do not suppose that the form of the diamond has been gradually
+achieved by a process of Selection. So again, as the course of descent
+branches in the successive generations, Selection determines along
+which branch Evolution shall proceed, but it does not decide what
+novelties that branch shall bring forth. "_La Nature contient le fonds
+de toutes ces variétés, mais le hazard ou l'art les mettent en
+oeuvre_," as Maupertuis most truly said.
+
+Not till knowledge of the genetic properties of organisms has attained
+to far greater completeness can evolutionary speculations have more
+than a suggestive value. By genetic experiment, cytology and
+physiological chemistry aiding, we may hope to acquire such knowledge.
+In 1872 Nathusius wrote:[72] "Das Gesetz der Vererbung ist noch nicht
+erkannt; der Apfel ist noch nicht vom Baum der Erkenntniss gefallen,
+welcher, der Sage nach, Newton auf den rechten Weg zur Ergründung der
+Gravitationsgesetze führte." We cannot pretend that the words are not
+still true, but in Mendelian analysis the seeds of that apple-tree at
+last are sown.
+
+If we were asked what discovery would do most to forward our inquiry,
+what one bit of knowledge would more than any other illuminate the
+problem, I think we may give the answer without hesitation. The
+greatest advance that we can foresee will be made when it is found
+possible to connect the geometrical phenomena of development with the
+chemical. The geometrical symmetry of living things is the key to a
+knowledge of their regularity, and the forces which cause it. In the
+symmetry of the dividing cell the basis of that resemblance we call
+Heredity is contained. To imitate the morphological phenomena of life
+we have to devise a system which can divide. It must be able to
+divide, and to segment as--grossly--a vibrating plate or rod does, or
+as an icicle can do as it becomes ribbed in a continuous stream of
+water; but with this distinction, that the distribution of chemical
+differences and properties must simultaneously be decided and disposed
+in orderly relation to the pattern of the segmentation. Even if a
+model which would do this could be constructed it might prove to be a
+useful beginning.
+
+This may be looking too far ahead. If we had to choose some one piece
+of more proximate knowledge which we would more especially like to
+acquire, I suppose we should ask for the secret of interracial
+sterility. Nothing has yet been discovered to remove the grave
+difficulty, by which Huxley in particular was so much oppressed, that
+among the many varieties produced under domestication--which we all
+regard as analogous to the species seen in nature--no clear case of
+interracial sterility has been demonstrated. The phenomenon is
+probably the only one to which the domesticated products seem to
+afford no parallel. No solution of the difficulty can be offered which
+has positive value, but it is perhaps worth considering the facts in
+the light of modern ideas. It should be observed that we are not
+discussing incompatibility of two species to produce offspring (a
+totally distinct phenomenon), but the sterility of the offspring which
+many of them do produce.
+
+When two species, both perfectly fertile severally, produce on crossing a
+sterile progeny, there is a presumption that the sterility is due to the
+development in the hybrid of some substance which can only be formed by the
+meeting of two complementary factors. That some such account is correct in
+essence may be inferred from the well-known observation that if the hybrid
+is not totally sterile but only partially so, and thus is able to form some
+good germ-cells which develop into new individuals, the sterility of these
+daughter-individuals is sensibly reduced or may be entirely absent. The
+fertility once re-established, the sterility does not return in the later
+progeny, a fact strongly suggestive of segregation. Now if the sterility of
+the cross-bred be really the consequence of the meeting of two
+complementary factors, we see that the phenomenon could only be produced
+among the divergent offspring of one species by the acquisition of at least
+_two_ new factors; for if the acquisition of a single factor caused
+sterility the line would then end. Moreover each factor must be separately
+acquired by distinct individuals, for if both were present together, the
+possessors would by hypothesis be sterile. And in order to imitate the case
+of species each of these factors must be acquired by distinct breeds. The
+factors need not, and probably would not, produce any other perceptible
+effects; they might, like the colour-factors present in white flowers, make
+no difference in the form or other characters. Not till the cross was
+actually made between the two complementary individuals would either factor
+come into play, and the effects even then might be unobserved until an
+attempt was made to breed from the cross-bred.
+
+Next, if the factors responsible for sterility were acquired, they
+would in all probability be peculiar to certain individuals and would
+not readily be distributed to the whole breed. Any member of the breed
+also into which _both_ the factors were introduced would drop out of
+the pedigree by virtue of its sterility. Hence the evidence that the
+various domesticated breeds say of dogs or fowls can when mated
+together produce fertile offspring, is beside the mark. The real
+question is, Do they ever produce sterile offspring? I think the
+evidence is clearly that sometimes they do, oftener perhaps than is
+commonly supposed. These suggestions are quite amenable to
+experimental tests. The most obvious way to begin is to get a pair of
+parents which are known to have had any sterile offspring, and to find
+the proportions in which these steriles were produced. If, as I
+anticipate, these proportions are found to be definite, the rest is
+simple.
+
+In passing, certain other considerations may be referred to. First,
+that there are observations favouring the view that the production of
+totally sterile cross-breds is seldom a universal property of two
+species, and that it may be a matter of individuals, which is just
+what on the view here proposed would be expected. Moreover, as we all
+know now, though incompatibility may be dependent to some extent on
+the degree to which the species are dissimilar, no such principle can
+be demonstrated to determine sterility or fertility in general. For
+example, though all our Finches can breed together, the hybrids are
+all sterile. Of Ducks some species can breed together without
+producing the slightest sterility; others have totally sterile
+offspring, and so on. The hybrids between several _genera_ of Orchids
+are perfectly fertile on the female side, and some on the male side
+also, but the hybrids produced between the Turnip (_Brassica napus_)
+and the Swede (_Brassica campestris_), which, according to our
+estimates of affinity, should be nearly allied forms, are totally
+sterile.[73] Lastly, it may be recalled that in sterility we are
+almost certainly considering a meristic phenomenon. _Failure to
+divide_ is, we may feel fairly sure, the immediate "cause" of the
+sterility. Now, though we know very little about the heredity of
+meristic differences, all that we do know points to the conclusion
+that the less-divided is dominant to the more-divided, and we are thus
+justified in supposing that there are factors which can arrest or
+prevent cell-division. My conjecture therefore is that in the case of
+sterility of cross-breds we see the effect produced by a complementary
+pair of such factors. This and many similar problems are now open to
+our analysis.
+
+The question is sometimes asked, Do the new lights on Variation and
+Heredity make the process of Evolution easier to understand? On the
+whole the answer may be given that they do. There is some appearance
+of loss of simplicity, but the gain is real. As was said above, the
+time is not ripe for the discussion of the origin of species. With
+faith in Evolution unshaken--if indeed the word faith can be used in
+application to that which is certain--we look on the manner and
+causation of adapted differentiation as still wholly mysterious. As
+Samuel Butler so truly said: "To me it seems that the 'Origin of
+Variation,' whatever it is, is the only true 'Origin of Species,'"[74]
+and of that Origin not one of us knows anything. But given
+Variation--and it is given: assuming further that the variations are
+not guided into paths of adaptation--and both to the Darwinian and to
+the modern school this hypothesis appears to be sound if unproven--an
+evolution of species proceeding by definite steps is more, rather than
+less, easy to imagine than an evolution proceeding by the accumulation
+of indefinite and insensible steps. Those who have lost themselves in
+contemplating the miracles of Adaptation (whether real or spurious)
+have not unnaturally fixed their hopes rather on the indefinite than
+on the definite changes. The reasons are obvious. By suggesting that
+the steps through which an adaptative mechanism arose were indefinite
+and insensible, all further trouble is spared. While it could be said
+that species arise by an insensible and imperceptible process of
+variation, there was clearly no use in tiring ourselves by trying to
+perceive that process. This labour-saving counsel found great favour.
+All that had to be done to develop evolution-theory was to discover
+the good in everything, a task which, in the complete absence of any
+control or test whereby to check the truth of the discovery, is not
+very onerous. The doctrine "_que tout est au mieux_" was therefore
+preached with fresh vigour, and examples of that illuminating
+principle were discovered with a facility that Pangloss himself might
+have envied, till at last even the spectators wearied of such dazzling
+performances.
+
+But in all seriousness, why should indefinite and unlimited variation
+have been regarded as a more probable account of the origin of
+Adaptation? Only, I think, because the obstacle was shifted one plane
+back, and so looked rather less prominent. The abundance of
+Adaptation, we all grant, is an immense, almost an unsurpassable
+difficulty in all non-Lamarckian views of Evolution; but if the steps
+by which that adaptation arose were fortuitious, to imagine them
+insensible is assuredly no help. In one most important respect indeed,
+as has often been observed, it is a multiplication of troubles. For
+the smaller the steps, the less could Natural Selection act upon them.
+Definite variations--and of the occurrence of definite variations in
+abundance we have now the most convincing proof--have at least the
+obvious merit that they can make and often do make a real difference
+in the chances of life.
+
+There is another aspect of the Adaptation problem to which I can
+allude very briefly. May not our present ideas of the universality and
+precision of Adaptation be greatly exaggerated? The fit of organism to
+its environment is not after all so very close--a proposition
+unwelcome perhaps, but one which could be illustrated by very copious
+evidence. Natural Selection is stern, but she has her tolerant moods.
+
+We have now most certain and irrefragable proof that much definiteness
+exists in living things apart from Selection, and also much that may
+very well have been preserved and so in a sense constituted by
+Selection. Here the matter is likely to rest. There is a passage in
+the sixth edition of the _Origin_ which has I think been overlooked.
+On page 70 Darwin says, "The tuft of hair on the breast of the wild
+turkey-cock cannot be of any use, and it is doubtful whether it can be
+ornamental in the eyes of the female bird." This tuft of hair is a
+most definite and unusual structure, and I am afraid that the remark
+that it "cannot be of any use" may have been made inadvertently; but
+it may have been intended, for in the first edition the usual
+qualification was given and must therefore have been deliberately
+excised. Anyhow I should like to think that Darwin did throw over that
+tuft of hair, and that he felt relief when he had done so. Whether
+however we have his great authority for such a course or not, I feel
+quite sure that we shall be rightly interpreting the facts of nature
+if we cease to expect to find purposefulness wherever we meet with
+definite structures or patterns. Such things are, as often as not, I
+suspect rather of the nature of tool-marks, mere incidents of
+manufacture, benefiting their possessor not more than the wire-marks
+in a sheet of paper, or the ribbing on the bottom of an oriental plate
+renders those objects more attractive in our eyes.
+
+If Variation may be in any way definite, the question once more
+arises, may it not be definite in direction? The belief that it is has
+had many supporters, from Lamarck onwards, who held that it was guided
+by need, and others who, like Nägeli, while laying no emphasis on
+need, yet were convinced that there was guidance of some kind. The
+latter view under the name of "Orthogenesis," devised I believe by
+Eimer, at the present day commends itself to some naturalists. The
+objection to such a suggestion is of course that no fragment of real
+evidence can be produced in its support. On the other hand, with the
+experimental proof that variation consists largely in the unpacking
+and repacking of an original complexity, it is not so certain as we
+might like to think that the order of these events is not
+predetermined.
+
+For instance the original "pack" may have been made in such a way that
+at the _n_th division of the germ-cells of a Sweet Pea a colour-factor
+might be dropped, and that at the _n_+_n_th division the hooded
+variety be given off, and so on. I see no ground whatever for holding
+such a view, but in fairness the possibility should not be forgotten,
+and in the light of modern research it scarcely looks so absurdly
+improbable as before.
+
+No one can survey the work of recent years without perceiving that
+evolutionary orthodoxy developed too fast, and that a great deal has
+got to come down; but this satisfaction at least remains, that in the
+experimental methods which Mendel inaugurated, we have means of
+reaching certainty in regard to the physiology of Heredity and
+Variation upon which a more lasting structure may be built.
+
+FOOTNOTES:
+
+[Footnote 56: _Vénus Physique, contenant deux Dissertations, l'une sur
+l'origine des Hommes et des Animaux_; _Et l'autre sur l'origine des
+Noirs_, La Haye, 1746, pp. 124 and 129. For an introduction to the
+writings of Maupertuis I am indebted to an article by Professor
+Lovejoy in _Popular Sci. Monthly_, 1902.]
+
+[Footnote 57: For the fullest account of the views of these pioneers
+of Evolution, see the works of Samuel Butler, especially _Evolution,
+Old and New_ (2nd edit.) 1882. Butler's claims on behalf of Buffon
+have met with some acceptance; but after reading what Butler has said,
+and a considerable part of Buffon's own works, the word "hinted" seems
+to me a sufficiently correct description of the part he played. It is
+interesting to note that in the chapter on the Ass, which contains
+some of his evolutionary passages, there is a reference to "_plusieurs
+idées très-élevées sur la génération_" contained in the Letters of
+Maupertuis.]
+
+[Footnote 58: See especially W. Lawrence, _Lectures on Physiology_,
+London, 1823, pp. 213 f.]
+
+[Footnote 59: See the chapter contributed to the _Life and Letters of
+Charles Darwin_, II. p. 195. I do not clearly understand the sense in
+which Darwin wrote (Autobiography, _ibid._ I. p. 87): "It has
+sometimes been said that the success of the _Origin_ proved 'that the
+subject was in the air,' or 'that men's minds were prepared for it.' I
+do not think that this is strictly true, for I occasionally sounded
+not a few naturalists, and never happened to come across a single one
+who seemed to doubt about the permanence of species." This experience
+may perhaps have been an accident due to Darwin's isolation. The
+literature of the period abounds with indications of "critical
+expectancy." A most interesting expression of that feeling is given in
+the charming account of the "Early Days of Darwinism" by Alfred
+Newton, _Macmillan's Magazine_, LVII. 1888, p. 241. He tells how in
+1858 when spending a dreary summer in Iceland, he and his friend, the
+ornithologist John Wolley, in default of active occupation, spent
+their days in discussion. "Both of us taking a keen interest in
+Natural History, it was but reasonable that a question, which in those
+days was always coming up wherever two or more naturalists were
+gathered together, should be continually recurring. That question was,
+'What is a species?' and connected therewith was the other question,
+'How did a species begin?'... Now we were of course fairly well
+acquainted with what had been published on these subjects." He then
+enumerates some of these publications, mentioning among others T.
+Vernon Wollaston's _Variation of Species_--a work which has in my
+opinion never been adequately appreciated. He proceeds: "Of course we
+never arrived at anything like a solution of these problems, general
+or special, but we felt very strongly that a solution ought to be
+found, and that quickly, if the study of Botany and Zoology was to
+make any great advance." He then describes how on his return home he
+received the famous number of the _Linnean Journal_ on a certain
+evening. "I sat up late that night to read it; and never shall I
+forget the impression it made upon me. Herein was contained a
+perfectly simple solution of all the difficulties which had been
+troubling me for months past.... I went to bed satisfied that a
+solution had been found."]
+
+[Footnote 60: _Origin_, 6th edit. (1882), p. 421.]
+
+[Footnote 61: Whatever be our estimate of the importance of Natural
+Selection, in this we all agree. Samuel Butler, the most brilliant,
+and by far the most interesting of Darwin's opponents--whose works are
+at length emerging from oblivion--in his Preface (1882) to the 2nd
+edition of _Evolution, Old and New_, repeats his earlier expression of
+homage to one whom he had come to regard as an enemy: "To the end of
+time, if the question be asked, 'Who taught people to believe in
+Evolution?' the answer must be that it was Mr. Darwin. This is true,
+and it is hard to see what palm of higher praise can be awarded to any
+philosopher."]
+
+[Footnote 62: _Life and Letters_, I. pp. 276 and 83.]
+
+[Footnote 63: This isolation of the systematists is the one most
+melancholy sequela of Darwinism. It seems an irony that we should read
+in the peroration to the _Origin_ that when the Darwinian view is
+accepted "Systematists will be able to pursue their labours as at
+present; but they will not be incessantly haunted by the shadowy doubt
+whether this or that form be a true species. This, I feel sure, and I
+speak after experience, will be no slight relief. The endless disputes
+whether or not some fifty species of British brambles are good species
+will cease." _Origin_, 6th edit. (1882), p. 425. True they have ceased
+to attract the attention of those who lead opinion, but anyone who
+will turn to the literature of systematics will find that they have
+not ceased in any other sense. Should there not be something
+disquieting in the fact that among the workers who come most into
+contact with specific differences, are to be found the only men who
+have failed to be persuaded of the unreality of those differences?]
+
+[Footnote 64: 6th edit. pp. 109 and 401. See Butler, _Essays on Life,
+Art, and Science_, p. 265, reprinted 1908, and _Evolution, Old and
+New_, chap. XXII. (2nd edit.), 1882.]
+
+[Footnote 65: W. Lawrence was one of the few who consistently
+maintained the contrary opinion. Prichard, who previously had
+expressed himself in the same sense, does not, I believe, repeat these
+views in his later writings, and there are signs that he came to
+believe in the transmission of acquired habits. See Lawrence, _Lect.
+Physiol._ 1823, pp. 436-437, 447. Prichard, Edin. Inaug. Disp. 1808
+[not seen by me], quoted _ibid._ and _Nat. Hist. Man_, 1843, pp. 34
+f.]
+
+[Footnote 66: It is interesting to see how nearly Butler was led by
+natural penetration, and from absolutely opposite conclusions, back to
+this underlying truth: "So that each ovum when impregnate should be
+considered not as descended from its ancestors, but as being a
+continuation of the personality of every ovum in the chain of its
+ancestry, which every ovum _it actually is_ quite as truly as the
+octogenarian _is_ the same identity with the ovum from which he has
+been developed. This process cannot stop short of the primordial cell,
+which again will probably turn out to be but a brief resting-place. We
+therefore prove each one of us to _be actually_ the primordial cell
+which never died nor dies, but has differentiated itself into the life
+of the world, all living beings whatever, being one with it and
+members one of another," _Life and Habit_, 1878, p. 86.]
+
+[Footnote 67: This view is no doubt contrary to the received opinion.
+I am however interested to see it lately maintained by Driesch
+(_Science and Philosophy of the Organism_, London, 1907, p. 233), and
+from the recent observations of Godlewski it has received distinct
+experimental support.]
+
+[Footnote 68: In other words, the ova are each _either_ female, _or
+male_ (i.e. non-female), but the sperms are all non-female.]
+
+[Footnote 69: Morgan, _Proc. Soc. Exp. Biol. Med._ V. 1908, and von
+Baehr, _Zool. Anz._ XXXII. p. 507, 1908.]
+
+[Footnote 70: As Wilson has proved, the unpaired body is not a
+universal feature even in those orders in which it has been observed.
+Nearly allied types may differ. In some it is altogether unpaired. In
+others it is paired with a body of much smaller size, and by selection
+of various types all gradations can be demonstrated ranging to the
+condition in which the members of the pair are indistinguishable from
+each other.]
+
+[Footnote 71: I have in view, for example, the marvellous and specific
+phenomena of regeneration, and those discovered by the students of
+"_Entwicklungsmechanik_." The circumstances of its occurrence here
+preclude any suggestion that this regularity has been brought about by
+the workings of Selection. The attempts thus to represent the
+phenomena have resulted in mere parodies of scientific reasoning.]
+
+[Footnote 72: _Vorträge über Viehzucht und Rassenerkenntniss_, p. 120,
+Berlin, 1872.]
+
+[Footnote 73: See Sutton, A. W., _Journ. Linn. Soc._ XXXVIII. p. 341,
+1908.]
+
+[Footnote 74: _Life and Habit_, London, p. 263, 1878]
+
+
+
+
+IV
+
+"THE DESCENT OF MAN"
+
+BY G. SCHWALBE
+
+_Professor of Anatomy in the University of Strassburg_
+
+
+The problem of the origin of the human race, of the descent of man, is
+ranked by Huxley in his epoch-making book _Man's Place in Nature_, as
+the deepest with which biology has to concern itself, "the question of
+questions,"--the problem which underlies all others. In the same
+brilliant and lucid exposition, which appeared in 1863, soon after the
+publication of Darwin's _Origin of Species_, Huxley stated his own
+views in regard to this great problem. He tells us how the idea of a
+natural descent of man gradually grew up in his mind. It was
+especially the assertions of Owen in regard to the total difference
+between the human and the simian brain that called forth strong
+dissent from the great anatomist Huxley, and he easily succeeded in
+showing that Owen's supposed differences had no real existence; he
+even established, on the basis of his own anatomical investigations,
+the proposition that the anatomical differences between the Marmoset
+and the Chimpanzee are much greater than those between the Chimpanzee
+and Man.
+
+But why do we thus introduce the study of Darwin's _Descent of Man_,
+which is to occupy us here, by insisting on the fact that Huxley had
+taken the field in defence of the descent of man in 1863, while
+Darwin's book on the subject did not appear till 1871? It is in order
+that we may clearly understand how it happened that from this time
+onwards Darwin and Huxley followed the same great aim in the most
+intimate association.
+
+Huxley and Darwin working at the same _Problema maximum_! Huxley
+fiery, impetuous, eager for battle, contemptuous of the resistance of
+a dull world, or energetically triumphing over it. Darwin calm,
+weighing every problem slowly, letting it mature thoroughly,--not a
+fighter, yet having the greater and more lasting influence by virtue
+of his immense mass of critically sifted proofs. Darwin's friend,
+Huxley, was the first to do him justice, to understand his nature, and
+to find in it the reason why the detailed and carefully considered
+book on the descent of man made its appearance so late. Huxley, always
+generous, never thought of claiming priority for himself. In
+enthusiastic language he tells how Darwin's immortal work, _The Origin
+of Species_, first shed light for him on the problem of the descent of
+man; the recognition of a _vera causa_ in the transformation of
+species illuminated his thoughts as with a flash. He was now content
+to leave what perplexed him, what he could not yet solve, as he says
+himself, "in the mighty hands of Darwin." Happy in the bustle of
+strife against old and deep-rooted prejudices, against intolerance and
+superstition, he wielded his sharp weapons on Darwin's behalf; wearing
+Darwin's armour he joyously overthrew adversary after adversary.
+Darwin spoke of Huxley as his "general agent."[75] Huxley says of
+himself "I am Darwin's bulldog."[76]
+
+Thus Huxley openly acknowledged that it was Darwin's _Origin of
+Species_ that first set the problem of the descent of man in its true
+light, that made the question of the origin of the human race a
+pressing one. That this was the logical consequence of his book Darwin
+himself had long felt. He had been reproached with intentionally
+shirking the application of his theory to Man. Let us hear what he
+says on this point in his autobiography: "As soon as I had become, in
+the year 1837 or 1838, convinced that species were mutable
+productions, I could not avoid the belief that man must come under the
+same law. Accordingly I collected notes on the subject for my own
+satisfaction, and not for a long time with any intention of
+publishing. Although in the 'Origin of Species' the derivation of any
+particular species is never discussed, yet I thought it best, in order
+_that no honourable man should accuse me of concealing my views_,[77]
+to add that by the work 'light would be thrown on the origin of man
+and his history.' It would have been useless and injurious to the
+success of the book to have paraded, without giving any evidence, my
+conviction with respect to his origin."[78]
+
+In a letter written in January, 1860, to the Rev. L. Blomefield,
+Darwin expresses himself in similar terms. "With respect to man, I am
+very far from wishing to obtrude my belief; but I thought it dishonest
+to quite conceal my opinion."[79]
+
+The brief allusion in the _Origin of Species_ is so far from prominent
+and so incidental that it was excusable to assume that Darwin had not
+touched upon the descent of man in this work. It was solely the desire
+to have his mass of evidence sufficiently complete, solely Darwin's
+great characteristic of never publishing till he had carefully weighed
+all aspects of his subject for years, solely, in short, his most
+fastidious scientific conscience that restrained him from challenging
+the world in 1859 with a book in which the theory of the descent of
+man was fully set forth. Three years, frequently interrupted by
+ill-health, were needed for the actual writing of the book:[80] the
+first edition, which appeared in 1871, was followed in 1874 by a much
+improved second edition, the preparation of which he very reluctantly
+undertook.[81]
+
+This, briefly, is the history of the work, which, with the _Origin of
+Species_, marks an epoch in the history of biological sciences--the
+work with which the cautious, peace-loving investigator ventured forth
+from his contemplative life into the arena of strife and unrest, and
+laid himself open to all the annoyances that deep-rooted belief and
+prejudice, and the prevailing tendency of scientific thought at the
+time could devise.
+
+Darwin did not take this step lightly. Of great interest in this
+connection is a letter written to Wallace on Dec. 22, 1857,[82] in
+which he says, "You ask me whether I shall discuss 'man.' I think I
+shall avoid the whole subject, as so surrounded with prejudices;
+though I fully admit that it is the highest and most interesting
+problem for the naturalist." But his conscientiousness compelled him
+to state briefly his opinion on the subject in the _Origin of Species_
+in 1859. Nevertheless he did not escape reproaches for having been so
+reticent. This is unmistakably apparent from a letter to Fritz Müller
+dated Feb. 22 [1869?], in which he says: "I am thinking of writing a
+little essay on the Origin of Mankind, as I have been taunted with
+concealing my opinions."[83]
+
+It might be thought that Darwin behaved thus hesitatingly, and was so
+slow in deciding on the full publication of his collected material in
+regard to the descent of man, because he had religious difficulties to
+overcome.
+
+But this was not the case, as we can see from his admirable confession
+of faith, the publication of which we owe to his son Francis.[84]
+Whoever wishes really to understand the lofty character of this great
+man should read these immortal lines in which he unfolds to us in
+simple and straightforward words the development of his conception of
+the universe. He describes how, though he was still quite orthodox
+during his voyage round the world on board the _Beagle_, he came
+gradually to see, shortly afterwards (1836-1839) that the Old
+Testament was no more to be trusted than the Sacred Books of the
+Hindoos; the miracles by which Christianity is supported, the
+discrepancies between the accounts in the different Gospels, gradually
+led him to disbelieve in Christianity as a divine revelation. "Thus,"
+he writes,[85] "disbelief crept over me at a very slow rate, but was
+at last complete. The rate was so slow that I felt no distress." But
+Darwin was too modest to presume to go beyond the limits laid down by
+science. He wanted nothing more than to be able to go, freely and
+unhampered by belief in authority or in the Bible, as far as human
+knowledge could lead him. We learn this from the concluding words of
+his chapter on religion "The mystery of the beginning of all things is
+insoluble by us; and I for one must be content to remain an
+Agnostic."[86]
+
+Darwin was always very unwilling to give publicity to his views in
+regard to religion. In a letter to Asa Gray on May 22, 1860,[87] he
+declares that it is always painful to him to have to enter into
+discussion of religious problems. He had, he said, no intention of
+writing atheistically.
+
+Finally, let us cite one characteristic sentence from a letter from
+Darwin to C. Ridley[88] (Nov. 28, 1878). A clergyman, Dr. Pusey, had
+asserted that Darwin had written the _Origin of Species_ with some
+relation to theology. Darwin writes emphatically, "Many years ago when
+I was collecting facts for the 'Origin,' my belief in what is called a
+personal God was as firm as that of Dr. Pusey himself, and as to the
+eternity of matter I never troubled myself about such insoluble
+questions." The expression "many years ago" refers to the time of his
+voyage round the world, as has already been pointed out. Darwin means
+by this utterance that the views which had gradually developed in his
+mind in regard to the origin of species were quite compatible with the
+faith of the Church.
+
+If we consider all these utterances of Darwin in regard to religion
+and to his outlook on life (Weltanschauung), we shall see at least so
+much, that religious reflection could in no way have influenced him in
+regard to the writing and publishing of his book on _The Descent of
+Man_. Darwin had early won for himself freedom of thought, and to this
+freedom he remained true to the end of his life, uninfluenced by the
+customs and opinions of the world around him.
+
+Darwin was thus inwardly fortified and armed against the host of
+calumnies, accusations, and attacks called forth by the publication of
+the _Origin of Species_, and to an even greater extent by the
+appearance of the _Descent of Man_. But in his defence he could rely
+on the aid of a band of distinguished auxiliaries of the rarest
+ability. His faithful confederate, Huxley, was joined by the botanist
+Hooker, and, after longer resistance, by the famous geologist Lyell,
+whose "conversion" afforded Darwin peculiar satisfaction. All three
+took the field with enthusiasm in defence of the natural descent of
+man. From Wallace, on the other hand, though he shared with him the
+idea of natural selection, Darwin got no support in this matter.
+Wallace expressed himself in a strange manner. He admitted everything
+in regard to the morphological descent of man, but maintained, in a
+mystic way, that something else, something of a spiritual nature must
+have been added to what man inherited from his animal ancestors.
+Darwin, whose esteem for Wallace was extraordinarily high, could not
+understand how he could give utterance to such a mystical view in
+regard to man; the idea seemed to him so "incredibly strange" that he
+thought some one else must have added these sentences to Wallace's
+paper.
+
+Even now there are thinkers who, like Wallace, shrink from applying to
+man the ultimate consequences of the theory of descent. The idea that
+man is derived from ape-like forms is to them unpleasant and
+humiliating.
+
+So far I have been depicting the development of Darwin's work on the
+descent of man. In what follows I shall endeavour to give a condensed
+survey of the contents of the book.
+
+It must at once be said that the contents of Darwin's work fall into
+two parts, dealing with entirely different subjects. _The Descent of
+Man_ includes a very detailed investigation in regard to secondary
+sexual characters in the animal series, and on this investigation
+Darwin founded a new theory, that of sexual selection. With
+astonishing patience he gathered together an immense mass of material,
+and showed, in regard to Arthropods and Vertebrates, the wide
+distribution of secondary characters, which develop almost exclusively
+in the male, and which enable him, on the one hand, to get the better
+of his rivals in the struggle for the female by the greater perfection
+of his weapons, and, on the other hand, to offer greater allurements
+to the female through the higher development of decorative characters,
+of song, or of scent-producing glands. The best equipped males will
+thus crowd out the less well-equipped in the matter of reproduction,
+and thus the relevant characters will be increased and perfected
+through sexual selection. It is, of course, a necessary assumption
+that these secondary sexual characters may be transmitted to the
+female, although perhaps in rudimentary form.
+
+As we have said, this story of sexual selection takes up a great deal
+of space in Darwin's book, and it need only be considered here in so
+far as Darwin applied it to the descent of man. To this latter problem
+the whole of Part I is devoted, while Part III contains a discussion
+of sexual selection in relation to man, and a general summary. Part
+II treats of sexual selection in general, and may be disregarded in
+our present study. Moreover, many interesting details must necessarily
+be passed over in what follows, for want of space.
+
+The first part of the _Descent of Man_ begins with an enumeration of
+the proofs of the animal descent of man taken from the structure of
+the human body. Darwin chiefly emphasises the fact that the human body
+consists of the same organs and of the same tissues as those of the
+other mammals; he shows also that man is subject to the same diseases
+and tormented by the same parasites as the apes. He further dwells on
+the general agreement exhibited by young embryonic forms, and he
+illustrates this by two figures placed one above the other, one
+representing a human embryo, after Ecker, the other a dog embryo,
+after Bischoff.[89]
+
+Darwin finds further proofs of the animal origin of man in the reduced
+structures, in themselves extremely variable, which are either
+absolutely useless to their possessors, or of so little use that they
+could never have developed under existing conditions. Of such vestiges
+he enumerates: the defective development of the _panniculus carnosus_
+(muscle of the skin) so widely distributed among mammals, the
+ear-muscles, the occasional persistence of the animal ear-point in
+man, the rudimentary nictitating membrane (_plica semilunaris_) in the
+human eye, the slight development of the organ of smell, the general
+hairiness of the human body, the frequently defective development or
+entire absence of the third molar (the wisdom tooth), the vermiform
+appendix, the occasional reappearance of a bony canal (_foramen
+supracondyloideum_) at the lower end of the humerus, the rudimentary
+tail of man (the so-called taillessness), and so on. Of these
+rudimentary structures the occasional occurrence of the animal
+ear-point in man is most fully discussed. Darwin's attention was
+called to this interesting structure by the sculptor Woolner. He
+figures such a case observed in man, and also the head of an alleged
+orang-foetus, the photograph of which he received from Nitsche.
+
+Darwin's interpretation of Woolner's case as having arisen through a
+folding over of the free edge of a pointed ear has been fully borne
+out by my investigations on the external ear.[90] In particular, it
+was established by these investigations that the human foetus, about
+the middle of its embryonic life, possesses a pointed ear somewhat
+similar to that of the monkey genus Macacus. One of Darwin's
+statements in regard to the head of the orang-foetus must be
+corrected. A _large_ ear with a point is shown in the photograph,[91]
+but it can easily be demonstrated--and Deniker has already pointed
+this out--that the figure is not that of an orang foetus at all, for
+that form has much smaller ears with no point; nor can it be a
+gibbon-foetus, as Deniker supposes, for the gibbon ear is also without
+a point. I myself regard it as that of a Macacus-embryo. But this
+mistake, which is due to Nitsche, in no way affects the fact
+recognised by Darwin, that ear-forms showing the point characteristic
+of the animal ear occur in man with extraordinary frequency.
+
+Finally, there is a discussion of those rudimentary structures which
+occur only in _one_ sex, such as the rudimentary mammary glands in the
+male, the vesicula prostatica, which corresponds to the uterus of the
+female, and others. All these facts tell in favour of the common
+descent of man and all other vertebrates. The conclusion of this
+section is characteristic: "_It is only our natural prejudice, and
+that arrogance which made our forefathers declare that they were
+descended from demi-gods, which leads us to demur to this conclusion.
+But the time will before long come, when it will be thought wonderful
+that naturalists, who were well acquainted with the comparative
+structure and development of man, and other mammals, should have
+believed that each was the work of a separate act of creation._"[92]
+
+In the second chapter there is a more detailed discussion, again based
+upon an extraordinary wealth of facts, of the problem as to the manner
+in which, and the causes through which, man evolved from a lower form.
+Precisely the same causes are here suggested for the origin of man, as
+for the origin of species in general. Variability, which is a
+necessary assumption in regard to all transformations, occurs in man
+to a high degree. Moreover, the rapid multiplication of the human race
+creates conditions which necessitate an energetic struggle for
+existence, and thus afford scope for the intervention of natural
+selection. Of the exercise of _artificial_ selection in the human
+race, there is nothing to be said, unless we cite such cases as the
+grenadiers of Frederick William I, or the population of ancient
+Sparta. In the passages already referred to and in those which follow,
+the transmission of acquired characters, upon which Darwin does not
+dwell, is taken for granted. In man, direct effects of changed
+conditions can be demonstrated (for instance in regard to bodily
+size), and there are also proofs of the influence exerted on his
+physical constitution by increased use or disuse. Reference is here
+made to the fact, established by Forbes, that the Quechua Indians of
+the high plateaus of Peru show a striking development of lungs and
+thorax, as a result of living constantly at high altitudes.
+
+Such special forms of variation as arrests of development
+(microcephalism) and reversion to lower forms are next discussed.
+Darwin himself felt[93] that these subjects are so nearly related to
+the cases mentioned in the first chapter, that many of them might as
+well have been dealt with there. It seems to me that it would have
+been better so, for the citation of additional instances of reversion
+at this place rather disturbs the logical sequence of his ideas as to
+the conditions which have brought about the evolution of man from
+lower forms. The instances of reversion here discussed are
+microcephalism, which Darwin wrongly interpreted as atavistic,
+supernumerary mammae, supernumerary digits, bicornuate uterus, the
+development of abnormal muscles, and so on. Brief mention is also made
+of correlative variations observed in man.
+
+Darwin next discusses the question as to the manner in which man
+attained to the erect position from the state of a climbing quadruped.
+Here again he puts the influence of Natural Selection in the first
+rank. The immediate progenitors of man had to maintain a struggle for
+existence in which success was to the more intelligent, and to those
+with social instincts. The hand of these climbing ancestors, which had
+little skill and served mainly for locomotion, could only undergo
+further development when some early member of the Primate series came
+to live more on the ground and less among trees.
+
+A bipedal existence thus became possible, and with it the liberation
+of the hand from locomotion, and the one-sided development of the
+human foot. The upright position brought about correlated variations
+in the bodily structure; with the free use of the hand it became
+possible to manufacture weapons and to use them; and this again
+resulted in a degeneration of the powerful canine teeth and the jaws,
+which were then no longer necessary for defence. Above all, however,
+the intelligence immediately increased, and with it skull and brain.
+The nakedness of man, and the absence of a tail (rudimentariness of
+the tail vertebrae) are next discussed. Darwin is inclined to
+attribute the nakedness of man, not to the action of natural selection
+on ancestors who originally inhabited a tropical land, but to sexual
+selection, which, for aesthetic reasons, brought about the loss of the
+hairy covering in man, or primarily in woman. An interesting
+discussion of the loss of the tail, which, however, man shares with
+the anthropoid apes, some other monkeys and lemurs, forms the
+conclusion of the almost superabundant material which Darwin worked
+up in the second chapter. His object was to show that some of the most
+distinctive human characters are in all probability directly or
+indirectly due to natural selection. With characteristic modesty he
+adds:[94] "Hence, if I have erred in giving to natural selection great
+power, which I am very far from admitting, or in having exaggerated
+its power, which is in itself probable, I have at least, as I hope,
+done good service in aiding to overthrow the dogma of separate
+creations." At the end of the chapter he touches upon the objection as
+to man's helpless and defenceless condition. Against this he urges his
+intelligence and social instincts.
+
+The two following chapters contain a detailed discussion of the
+objections drawn from the supposed great differences between the
+mental powers of men and animals. Darwin at once admits that the
+differences are enormous, but not that any fundamental difference
+between the two can be found. Very characteristic of him is the
+following passage: "In what manner the mental powers were first
+developed in the lowest organisms, is as hopeless an enquiry as how
+life itself first originated. These are problems for the distant
+future, if they are ever to be solved by man."[95]
+
+After some brief observations on instinct and intelligence, Darwin
+brings forward evidence to show that the greater number of the
+emotional states, such as pleasure and pain, happiness and misery,
+love and hate are common to man and the higher animals. He goes on to
+give various examples showing that wonder and curiosity, imitation,
+attention, memory and imagination (dreams of animals), can also be
+observed in the higher mammals, especially in apes. In regard even to
+reason there are no sharply defined limits. A certain faculty of
+deliberation is characteristic of some animals, and the more
+thoroughly we know an animal the more intelligence we are inclined to
+credit it with. Examples are brought forward of the intelligent and
+deliberate actions of apes, dogs and elephants. But although no
+sharply defined differences exist between man and animals, there is,
+nevertheless, a series of other mental powers which are
+characteristics usually regarded as absolutely peculiar to man. Some
+of these characteristics are examined in detail, and it is shown that
+the arguments drawn from them are not conclusive. Man alone is said to
+be capable of progressive improvement; but against this must be placed
+as something analogous in animals, the fact that they learn cunning
+and caution through long continued persecution. Even the use of tools
+is not in itself peculiar to man (monkeys use sticks, stones and
+twigs), but man alone fashions and uses implements _designed for a
+special purpose_. In this connection the remarks taken from Lubbock in
+regard to the origin and gradual development of the earliest flint
+implements will be read with interest; these are similar to the
+observations on modern eoliths, and their bearing on the development
+of the stone industry. It is interesting to learn from a letter to
+Hooker,[96] that Darwin himself at first doubted whether the stone
+implements discovered by Boucher de Perthes were really of the nature
+of tools. With the relentless candour as to himself which
+characterised him, he writes four years later in a letter to Lyell in
+regard to this view of Boucher de Perthes' discoveries: "I know
+something about his errors, and looked at his book many years ago, and
+am ashamed to think that I concluded the whole was rubbish! Yet he has
+done for man something like what Agassiz did for glaciers."[97]
+
+To return to Darwin's further comparisons between the higher mental powers
+of man and animals; He takes much of the force from the argument that man
+alone is capable of abstraction and self-consciousness by his own
+observations on dogs. One of the main differences between man and animals,
+speech, receives detailed treatment. He points out that various animals
+(birds, monkeys, dogs) have a large number of different sounds for
+different emotions, that, further, man produces in common with animals a
+whole series of inarticulate cries combined with gestures, and that dogs
+learn to understand whole sentences of human speech. In regard to human
+language, Darwin expresses a view contrary to that held by Max Müller:[98]
+"I cannot doubt that language owes its origin to the imitation and
+modification of various natural sounds, the voices of other animals, and
+man's own instinctive cries, aided by signs and gestures." The development
+of actual language presupposes a higher degree of intelligence than is
+found in any kind of ape. Darwin remarks on this point:[99] "The fact of
+the higher apes not using their vocal organs for speech no doubt depends on
+their intelligence not having been sufficiently advanced."
+
+The sense of beauty, too, has been alleged to be peculiar to man. In
+refutation of this assertion Darwin points to the decorative colours
+of birds, which are used for display. And to the last objection, that
+man alone has religion, that he alone has a belief in God, it is
+answered "that numerous races have existed, and still exist, who have
+no idea of one or more gods, and who have no words in their languages
+to express such an idea."[100]
+
+The result of the investigations recorded in this chapter is to show
+that, great as the difference in mental powers between man and the
+higher animals may be, it is undoubtedly only a difference "of degree
+and not of kind."[101]
+
+In the fourth chapter Darwin deals with the _moral sense_ or
+_conscience_, which is the most important of all differences between
+man and animals. It is a result of social instincts, which lead to
+sympathy for other members of the same society, to non-egoistic
+actions for the good of others. Darwin shows that social tendencies
+are found among many animals, and that among these love and
+kin-sympathy exist, and he gives examples of animals (especially dogs)
+which may exhibit characters that we should call moral in man (e.g.
+disinterested self-sacrifice for the sake of others). The early
+ape-like progenitors of the human race were undoubtedly social. With
+the increase of intelligence the moral sense develops farther; with
+the acquisition of speech public opinion arises, and finally, moral
+sense becomes habit. The rest of Darwin's detailed discussions on
+moral philosophy may be passed over.
+
+The fifth chapter may be very briefly summarised. In it Darwin shows
+that the intellectual and moral faculties are perfected through
+natural selection. He inquires how it can come about that a tribe at a
+low level of evolution attains to a higher, although the best and
+bravest among them often pay for their fidelity and courage with their
+lives without leaving any descendants. In this case it is the
+sentiment of glory, praise and blame, the admiration of others, which
+bring about the increase of the better members of the tribe. Property,
+fixed dwellings, and the association of families into a community are
+also indispensable requirements for civilisation. In the longer second
+section of the fifth chapter Darwin acts mainly as recorder. On the
+basis of numerous investigations, especially those of Greg, Wallace,
+and Galton, he inquires how far the influence of natural selection can
+be demonstrated in regard to civilised nations. In the final section,
+which deals with the proofs that all civilised nations were once
+barbarians, Darwin again uses the results gained by other
+investigators, such as Lubbock and Tylor. There are two sets of facts
+which prove the proposition in question. In the first place, we find
+traces of a former lower state in the customs and beliefs of all
+civilised nations, and in the second place, there are proofs to show
+that savage races are independently able to raise themselves a few
+steps in the scale of civilisation, and that they have thus raised
+themselves.
+
+In the sixth chapter of the work, Morphology comes into the foreground
+once more. Darwin first goes back, however, to the argument based on
+the great difference between the mental powers of the highest animals
+and those of man. That this is only quantitative, not qualitative, he
+has already shown. Very instructive in this connection is the
+reference to the enormous difference in mental powers in another
+class. No one would draw from the fact that the cochineal insect
+(Coccus) and the ant exhibit enormous differences in their mental
+powers, the conclusion that the ant should therefore be regarded as
+something quite distinct, and withdrawn from the class of insects
+altogether.
+
+Darwin next attempts to establish the _specific_ genealogical tree of
+man, and carefully weighs the differences and resemblances between the
+different families of the Primates. The erect position of man is an
+adaptive character, just as are the various characters referable to
+aquatic life in the seals, which, notwithstanding these, are ranked as
+a mere family of the carnivores. The following utterance is very
+characteristic of Darwin:[102] "If man had not been his own
+classifier, he would never have thought of founding a separate order
+for his own reception." In numerous characters not mentioned in
+systematic works, in the features of the face, in the form of the
+nose, in the structure of the external ear, man resembles the apes.
+The arrangement of the hair in man has also much in common with the
+apes; as also the occurrence of hair on the forehead of the human
+embryo, the beard, the convergence of the hair of the upper and under
+arm towards the elbow, which occurs not only in the anthropoid apes,
+but also in some American monkeys. Darwin here adopts Wallace's
+explanation of the origin of the ascending direction of the hair in
+the forearm of the orang,--that it has arisen through the habit of
+holding the hands over the head in rain. But this explanation cannot
+be maintained when we consider that this disposition of the hair is
+widely distributed among the most different mammals, being found in
+the dog, in the sloth, and in many of the lower monkeys.
+
+After further careful analysis of the anatomical characters Darwin
+reaches the conclusion that the New World monkeys (Platyrrhine) may be
+excluded from the genealogical tree altogether, but that man is an
+offshoot from the Old World monkeys (Catarrhine) whose progenitors
+existed as far back as the Miocene period. Among these Old World
+monkeys the forms to which man shows the greatest resemblance are the
+anthropoid apes, which, like him, possess neither tail nor ischial
+callosities. The platyrrhine and catarrhine monkeys have their
+primitive ancestor among extinct forms of the Lemuridae. Darwin also
+touches on the question of the original home of the human race and
+supposes that it may have been in Africa, because it is there that
+man's nearest relatives, the gorilla and the chimpanzee, are found.
+But he regards speculation on this point as useless. It is remarkable
+that, in this connection, Darwin regards the loss of the hair-covering
+in man as having some relation to a warm climate, while elsewhere he
+is inclined to make sexual selection responsible for it. Darwin
+recognises the great gap between man and his nearest relatives, but
+similar gaps exist at other parts of the mammalian genealogical tree:
+the allied forms have become extinct. After the extermination of the
+lower races of mankind, on the one hand, and of the anthropoid apes on
+the other, which will undoubtedly take place, the gulf will be greater
+than ever, since the baboons will then bound it on the one side, and
+the white races on the other. Little weight need be attached to the
+lack of fossil remains to fill up this gap, since the discovery of
+these depends upon chance. The last part of the chapter is devoted to
+a discussion of the earlier stages in the genealogy of man. Here
+Darwin accepts in the main the genealogical tree, which had meantime
+been published by Haeckel, who traces the pedigree back through
+Monotrems, Reptiles, Amphibians, and Fishes, to Amphioxus.
+
+Then follows an attempt to reconstruct, from the atavistic characters,
+a picture of our primitive ancestor who was undoubtedly an arboreal
+animal. The occurrence of rudiments of parts in one sex which only
+come to full development in the other is next discussed. This state of
+things Darwin regards as derived from an original hermaphroditism. In
+regard to the mammary glands of the male he does not accept the theory
+that they are vestigial, but considers them rather as not fully
+developed.
+
+The last chapter of Part I deals with the question whether the
+different races of man are to be regarded as different species, or as
+sub-species of a race of monophyletic origin. The striking differences
+between the races are first emphasised, and the question of the
+fertility or infertility of hybrids is discussed. That fertility is
+the more usual is shown by the excessive fertility of the hybrid
+population of Brazil. This, and the great variability of the
+distinguishing characters of the different races, as well as the fact
+that all grades of transition stages are found between these, while
+considerable general agreement exists, tell in favour of the unity of
+the races and lead to the conclusion that they all had a common
+primitive ancestor.
+
+Darwin therefore classifies all the different races as sub-species of
+_one and the same species_. Then follows an interesting inquiry into
+the reasons for the extinction of human races. He recognises as the
+ultimate reason the injurious effects of a change of the conditions of
+life, which may bring about an increase in infantile mortality, and a
+diminished fertility. It is precisely the reproductive system, among
+animals also, which is most susceptible to changes in the environment.
+
+The final section of this chapter deals with the formation of the
+races of mankind. Darwin discusses the question how far the direct
+effect of different conditions of life, or the inherited effects of
+increased use or disuse may have brought about the characteristic
+differences between the different races. Even in regard to the origin
+of the colour of the skin he rejects the transmitted effects of an
+original difference of climate as an explanation. In so doing he is
+following his tendency to exclude Lamarckian explanations as far as
+possible. But here he makes gratuitous difficulties from which, since
+natural selection fails, there is no escape except by bringing in the
+principle of sexual selection, to which, he regarded it as possible,
+skin-colouring, arrangement of hair, and form of features might be
+traced. But with his characteristic conscientiousness he guards
+himself thus: "I do not intend to assert that sexual selection will
+account for all the differences between the races."[103]
+
+I may be permitted a remark as to Darwin's attitude towards Lamarck.
+While, at an earlier stage, when he was engaged in the preliminary
+labours for his immortal work, _The Origin of Species_, Darwin
+expresses himself very forcibly against the views of Lamarck, speaking
+of Lamarckian "nonsense,"[104] and of Lamarck's "absurd, though clever
+work"[105] and expressly declaring, "I attribute very little to the
+direct action of climate, etc."[106] yet in later life he became more
+and more convinced of the influence of external conditions. In 1876,
+that is, two years after the appearance of the second edition of _The
+Descent of Man_, he writes with his usual candid honesty: "In my
+opinion the greatest error which I have committed, has been not
+allowing sufficient weight to the direct action of the environment,
+i.e. food, climate, etc. independently of a natural selection."[107]
+It is certain from this change of opinion that, if he had been able to
+make up his mind to issue a third edition of _The Descent of Man_, he
+would have ascribed a much greater influence to the effect of
+external conditions in explaining the different characters of the
+races of man than he did in the second edition. He would also
+undoubtedly have attributed less influence to sexual selection as a
+factor in the origin of the different bodily characteristics, if
+indeed he would not have excluded it altogether.
+
+In Part III of the _Descent_ two additional chapters are devoted to
+the discussion of sexual selection in relation to man. These may be
+very briefly referred to. Darwin here seeks to show that sexual
+selection has been operative on man and his primitive progenitor.
+Space fails me to follow out his interesting arguments. I can only
+mention that he is inclined to trace back hairlessness, the
+development of the beard in man, and the characteristic colour of the
+different human races to sexual selection. Since bareness of the skin
+could be no advantage, but rather a disadvantage, this character
+cannot have been brought about by natural selection. Darwin also
+rejected a direct influence of climate as a cause of the origin of the
+skin-colour. I have already expressed the opinion, based on the
+development of his views as shown in his letters, that in a third
+edition Darwin would probably have laid more stress on the influence
+of external environment. He himself feels that there are gaps in his
+proofs here, and says in self-criticism: "The views here advanced, on
+the part which sexual selection has played in the history of man, want
+scientific precision."[108] I need here only point out that it is
+impossible to explain the graduated stages of skin-colour by sexual
+selection, since it would have produced races sharply defined by their
+colour and not united to other races by transition stages, and this,
+it is well known, is not the case. Moreover, the fact established by
+me,[109] that in all races the ventral side of the trunk is paler than
+the dorsal side, and the inner surface of the extremities paler than
+the outer side, cannot be explained by sexual selection in the
+Darwinian sense.
+
+With this I conclude my brief survey of the rich contents of Darwin's
+book. I may be permitted to conclude by quoting the magnificent final
+words of _The Descent of Man_: "We must, however, acknowledge, as it
+seems to me, that man, with all his noble qualities, with sympathy
+which feels for the most debased, with benevolence which extends not
+only to other men but to the humblest living creature, with his
+god-like intellect which has penetrated into the movements and
+constitution of the solar system--with all these exalted powers--Man
+still bears in his bodily frame the indelible stamp of his lowly
+origin."[110]
+
+What has been the fate of Darwin's doctrines since his great
+achievement? How have they been received and followed up by the
+scientific and lay world? And what do the successors of the mighty
+hero and genius think now in regard to the origin of the human race?
+
+At the present time we are incomparably more favourably placed than
+Darwin was for answering this question of all questions. We have at
+our command an incomparably greater wealth of material than he had at
+his disposal. And we are more fortunate than he in this respect, that
+we now know transition-forms which help to fill up the gap, still
+great, between the lowest human races and the highest apes. Let us
+consider for a little the more essential additions to our knowledge
+since the publication of _The Descent of Man_.
+
+Since that time our knowledge of animal embryos has increased
+enormously. While Darwin was obliged to content himself with comparing
+a human embryo with that of a dog, there are now available the
+youngest embryos of monkeys of all possible groups (Orang, Gibbon,
+Semnopithecus, Macacus), thanks to Selenka's most successful tour in
+the East Indies in search of such material. We can now compare
+corresponding stages of the lower monkeys and of the Anthropoid apes
+with human embryos, and convince ourselves of their great resemblance
+to one another, thus strengthening enormously the armour prepared by
+Darwin in defence of his view on man's nearest relatives. It may be
+said that Selenka's material fills up the blanks in Darwin's array of
+proofs in the most satisfactory manner.
+
+The deepening of our knowledge of comparative anatomy also gives us much
+surer foundations than those on which Darwin was obliged to build. Just of
+late there have been many workers in the domain of the anatomy of apes and
+lemurs, and their investigations extend to the most different organs. Our
+knowledge of fossil apes and lemurs has also become much wider and more
+exact since Darwin's time: the fossil lemurs have been especially worked up
+by Cope, Forsyth Major, Ameghino, and others. Darwin knew very little about
+fossil monkeys. He mentions two or three anthropoid apes as occurring in
+the Miocene of Europe,[111] but only names _Dryopithecus_, the largest form
+from the Miocene of France. It was erroneously supposed that this form was
+related to _Hylobates_. We now know not only a form that actually stands
+near to the gibbon (_Pliopithecus_), and remains of other anthropoids
+(_Pliohylobates_ and the fossil chimpanzee, _Palaeopithecus_), but also
+several lower catarrhine monkeys, of which _Mesopithecus_, a form nearly
+related to the modern Sacred Monkeys (a species of _Semnopithecus_) and
+found in strata of the Miocene period in Greece, is the most important.
+Quite recently, too, Ameghino's investigations have made us acquainted with
+fossil monkeys from South America (_Anthropops_, _Homunculus_), which,
+according to their discoverer, are to be regarded as in the line of human
+descent.
+
+What Darwin missed most of all--intermediate forms between apes and
+man--has been recently furnished. E. Dubois, as is well known,
+discovered in 1893, near Trinil in Java, in the alluvial deposits of
+the river Bengawan, an important form represented by a skull-cap, some
+molars, and a femur. His opinion--much disputed as it has been--that
+in this form, which he named _Pithecanthropus_, he has found a
+long-desired transition-form is shared by the present writer. And
+although the geological age of these fossils, which, according to
+Dubois, belong to the uppermost Tertiary series, the Pliocene has
+recently been fixed at a later date (the older Diluvium), the
+_morphological value_ of these interesting remains, that is, the
+intermediate position of _Pithecanthropus_, still holds good. Volz
+says with justice,[112] that even if _Pithecanthropus_ is not _the_
+missing link, it is undoubtedly _a_ missing link.
+
+As on the one hand there has been found in _Pithecanthropus_ a form
+which, though intermediate between apes and man, is nevertheless more
+closely allied to the apes, so on the other hand, much progress has
+been made since Darwin's day in the discovery and description of the
+oldest human remains. Since the famous roof of a skull and the bones
+of the extremities belonging to it were found in 1856 in the
+Neandertal near Düsseldorf, the most varied judgments have been
+expressed in regard to the significance of the remains and of the
+skull in particular. In Darwin's _Descent of Man_ there is only a
+passing allusion to them[113] in connection with the discussion of the
+skull-capacity, although the investigations of Schaaffhausen, King,
+and Huxley were then known. I believe I have shown, in a series of
+papers, that the skull in question belongs to a form different from
+any of the races of man now living, and, with King and Cope, I regard
+it as at least a different species from living man, and have therefore
+designated it _Homo primigenius_. The form unquestionably belongs to
+the older Diluvium, and in the later Diluvium human forms already
+appear, which agree in all essential points with existing human races.
+
+As far back as 1886 the value of the Neandertal skull was greatly
+enhanced by Fraipont's discovery of two skulls and skeletons from Spy
+in Belgium. These are excellently described by their discoverer,[114]
+and are regarded as belonging to the same group of forms as the
+Neandertal remains. In 1899 and the following years came the discovery
+by Gorjanovic-Kramberger of different skeletal parts of at least
+ten individuals in a cave near Krapina in Croatia.[115] It is in
+particular the form of the lower jaw which is different from that of
+all recent races of man, and which clearly indicates the lowly
+position of _Homo primigenius_, while, on the other hand, the
+long-known skull from Gibraltar, which I[116] have referred to _Homo
+primigenius_, and which has lately been examined in detail by
+Sollas,[117] has made us acquainted with the surprising shape of the
+eye-orbit, of the nose, and of the whole upper part of the face.
+Isolated lower jaws found at La Naulette in Belgium, and at Malarnaud
+in France, increase our material which is now as abundant as could be
+desired. The most recent discovery of all is that of a skull dug up in
+August of this year [1908] by Klaatsch and Hauser in the lower grotto
+of the Le Moustier in Southern France, but this skull has not yet been
+fully described. Thus _Homo primigenius_ must also be regarded as
+occupying a position in the gap existing between the highest apes and
+the lowest human races, _Pithecanthropus_, standing in the lower part
+of it, and _Homo primigenius_ in the higher, near man. In order to
+prevent misunderstanding, I should like here to emphasise that in
+arranging this structural series--anthropoid apes, _Pithecanthropus_,
+_Homo primigenius_, _Homo sapiens_--I have no intention of
+establishing it as a direct genealogical series. I shall have
+something to say in regard to the genetic relations of these forms,
+one to another, when discussing the different theories of descent
+current at the present day.[118]
+
+In quite a different domain from that of morphological relationship,
+namely in the physiological study of the blood, results have recently
+been gained which are of the highest importance to the doctrine of
+descent. Uhlenhuth, Nuttall, and others have established the fact that
+the blood-serum of a rabbit which has previously had human blood
+injected into it, forms a precipitate with human blood. This
+biological reaction was tried with a great variety of mammalian
+species, and it was found that those far removed from man gave no
+precipitate under these conditions. But as in other cases among
+mammals all nearly related forms yield an almost equally marked
+precipitate, so the serum of a rabbit treated with human blood and
+then added to the blood of an anthropoid ape gives _almost_ as marked
+a precipitate as in human blood; the reaction to the blood of the
+lower Eastern monkeys is weaker, that to the Western monkeys weaker
+still; indeed in this last case there is only a slight clouding after
+a considerable time and no actual precipitate. The blood of the
+Lemuridae (Nuttall) gives no reaction or an extremely weak one, that
+of the other mammals none whatever. We have in this not only a proof
+of the literal blood relationship between man and apes, but the degree
+of relationship with the different main groups of apes can be
+determined beyond possibility of mistake.
+
+Finally, it must be briefly mentioned that in regard to remains of
+human handicraft also, the material at our disposal has greatly
+increased of late years, that, as a result of this, the opinions of
+archaeologists have undergone many changes, and that, in particular,
+their views in regard to the age of the human race have been greatly
+influenced. There is a tendency at the present time to refer the
+origin of man back to Tertiary times. It is true that no remains of
+Tertiary man have been found, but flints have been discovered which,
+according to the opinion of most investigators, bear traces either of
+use, or of very primitive workmanship. Since Rutot's time, following
+Mortillet's example, investigators have called these "eoliths," and
+they have been traced back by Verworn to the Miocene of the Auvergne,
+and by Rutot even to the upper Oligocene. Although these eoliths are
+even nowadays the subject of many different views, the preoccupation
+with them has kept the problem of the age of the human race
+continually before us.
+
+Geology, too, has made great progress since the days of Darwin and
+Lyell, and has endeavoured with satisfactory results to arrange the
+human remains of the Diluvial period in chronological order (Penck). I
+do not intend to enter upon the question of the primitive home of the
+human race; since the space at my disposal will not allow of my
+touching even very briefly upon all the departments of science which
+are concerned in the problem of the descent of man. How Darwin would
+have rejoiced over each of the discoveries here briefly outlined! What
+use he would have made of the new and precious material, which would
+have prevented the discouragement from which he suffered when
+preparing the second edition of _The Descent of Man_! But it was not
+granted to him to see this progress towards filling up the gaps in his
+edifice of which he was so painfully conscious.
+
+He did, however, have the satisfaction of seeing his ideas steadily
+gaining ground, notwithstanding much hostility and deep-rooted
+prejudice. Even in the years between the appearance of _The Origin of
+Species_ and of the first edition of the _Descent_, the idea of a
+natural descent of man, which was only briefly indicated in the work
+of 1859, had been eagerly welcomed in some quarters. It has been
+already pointed out how brilliantly Huxley contributed to the defence
+and diffusion of Darwin's doctrines, and how in _Man's Place in
+Nature_ he has given us a classic work as a foundation for the
+doctrine of the descent of man. As Huxley was Darwin's champion in
+England, so in Germany Carl Vogt, in particular, made himself master
+of the Darwinian ideas. But above all it was Haeckel who, in energy,
+eagerness for battle, and knowledge may be placed side by side with
+Huxley, who took over the leadership in the controversy over the new
+conception of the universe. As far back as 1866, in his _Generelle
+Morphologie_, he had inquired minutely into the question of the
+descent of man, and not content with urging merely the general theory
+of descent from lower animal forms, he drew up for the first time
+genealogical trees showing the close structural relationships of the
+different animal groups; the last of these illustrated the
+relationships of Mammals, and among them of all groups of the
+Primates, including man. It was Haeckel's genealogical trees that
+formed the basis of the special discussion of the relationships of
+man, in the sixth chapter of Darwin's _Descent of Man_.
+
+In the last section of this essay I shall return to Haeckel's
+conception of the special descent of man, the main features of which
+he still upholds, and rightly so. Haeckel has contributed more than
+any one else to the spread of the Darwinian doctrine.
+
+I can only allow myself a few words as to the spread of the theory of
+the natural descent of man in other countries. The Parisian
+anthropological school, founded and guided by the genius of Broca,
+took up the idea of the descent of man, and made many notable
+contributions to it (Broca, Manouvrier, Mahoudeau, Deniker and
+others). In England itself Darwin's work did not die. Huxley took care
+of that, for he, with his lofty and unprejudiced mind, dominated and
+inspired English biology until his death on June 29, 1895. He had the
+satisfaction shortly before his death of learning of Dubois'
+discovery, which he illustrated by a humourous sketch.[119] But there
+are still many followers in Darwin's footsteps in England. Keane has
+worked at the special genealogical tree of the Primates; Keith has
+inquired which of the anthropoid apes has the greatest number of
+characters in common with man; Morris concerns himself with the
+evolution of man in general, especially with his acquisition of the
+erect position. The recent discoveries of _Pithecanthropus_ and _Homo
+primigenius_ are being vigorously discussed; but the present writer is
+not in a position to form an opinion of the extent to which the idea
+of descent has penetrated throughout England generally.
+
+In Italy independent work in the domain of the descent of man is being
+produced, especially by Morselli; with him are associated, in the
+investigation of related problems, Sergi and Giuffrida-Ruggeri. From
+the ranks of American investigators we may single out in particular
+the eminent geologist Cope, who championed with much decision the idea
+of the specific difference of _Homo neandertalensis_ (_primigenius_)
+and maintained a more direct descent of man from the fossil Lemuridae.
+In South America too, in Argentina, new life is stirring in this
+department of science. Ameghino in Buenos Ayres has awakened the
+fossil primates of the Pampas formation to new life; he even believes
+that in his _Tetraprothomo_, represented by a femur, he has discovered
+a direct ancestor of man. Lehmann-Nitsche is working at the other side
+of the gulf between apes and man, and he describes a remarkable first
+cervical vertebra (atlas) from Monte Hermoso as belonging to a form
+which may bear the same relation to _Homo sapiens_ in South America as
+_Homo primigenius_ does in the Old World. After a minute investigation
+he establishes a human species _Homo neogaeus_, while Ameghino
+ascribes this atlas vertebra to his _Tetraprothomo_.
+
+Thus throughout the whole scientific world there is arising a new
+life, an eager endeavour to get nearer to Huxley's _problema maximum_,
+to penetrate more deeply into the origin of the human race. There are
+to-day very few experts in anatomy and zoology who deny the animal
+descent of man in general. Religious considerations, old prejudices,
+the reluctance to accept man, who so far surpasses mentally all other
+creatures, as descended from "soulless" animals, prevent a few
+investigators from giving full adherence to the doctrine. But there
+are very few of these who still postulate a special act of creation
+for man. Although the majority of experts in anatomy and zoology
+accept unconditionally the descent of man from lower forms, there is
+much diversity of opinion among them in regard to the special line of
+descent.
+
+In trying to establish any special hypothesis of descent, whether by
+the graphic method of drawing up genealogical trees or otherwise, let
+us always bear in mind Darwin's words[120] and use them as a critical
+guiding line: "As we have no record of the lines of descent, the
+pedigree can be discovered only by observing the degrees of
+resemblance between the beings which are to be classed." Darwin
+carries this further by stating "that resemblances in several
+unimportant structures, in useless and rudimentary organs, or not now
+functionally active, or in an embryological condition, are by far the
+most serviceable for classification."[121] It has also to be
+remembered that _numerous_ separate points of agreement are of much
+greater importance than the amount of similarity or dissimilarity in a
+few points.
+
+The hypotheses as to descent current at the present day may be divided
+into two main groups. The first group seeks for the roots of the human
+race not among any of the families of the apes--the anatomically
+nearest forms--nor among their very similar but less specialised
+ancestral forms, the fossil representatives of which we can know only
+in part, but, setting the monkeys on one side, it seeks for them lower
+down among the fossil Eocene Pseudo-lemuridae or Lemuridae (Cope), or
+even among the primitive pentadactylous Eocene forms, which may
+either have led directly to the evolution of man (Adloff), or have
+given rise to an ancestral form common to apes and men (Klaatsch,[122]
+Giuffrida-Ruggeri). The common ancestral form, from which man and apes
+are thus supposed to have arisen independently, may explain the
+numerous resemblances which actually exist between them. That is to
+say, all the characters upon which the great structural resemblance
+between apes and man depends must have been present in their common
+ancestor. Let us take an example of such a common character. The bony
+external ear-passage is in general as highly developed in the lower
+Eastern monkeys and the anthropoid apes as in man. This character
+must, therefore, have already been present in the common primitive
+form. In that case it is not easy to understand why the Western
+monkeys have not also inherited the character, instead of possessing
+only a tympanic ring. But it becomes more intelligible if we assume
+that forms with a primitive tympanic ring were the original type, and
+that from these were evolved, on the one hand, the existing New World
+monkeys with persistent tympanic ring, and on the other an ancestral
+form common to the lower Old World monkeys, the anthropoid apes and
+man. For man shares with these the character in question, and it is
+also one of the "unimportant" characters required by Darwin. Thus we
+have two divergent lines arising from the ancestral form, the Western
+monkeys (Platyrrhine) on the one hand, and an ancestral form common to
+the lower Eastern monkeys, the anthropoid apes, and man, on the other.
+But considerations similar to those which showed it to be impossible
+that man should have developed from an ancestor common to him and the
+monkeys, yet outside of and parallel with these, may be urged also
+against the likelihood of a parallel evolution of the lower Eastern
+monkeys, the anthropoid apes, and man. The anthropoid apes have in
+common with man many characters which are not present in the lower
+Old World monkeys. These characters must therefore have been present
+in the ancestral form common to the three groups. But here, again, it
+is difficult to understand why the lower Eastern monkeys should not
+also have inherited these characters. As this is not the case, there
+remains no alternative but to assume divergent evolution from an
+indifferent form. The lower Eastern monkeys are carrying on the
+evolution in one direction--I might almost say towards a blind
+alley--while anthropoids and men have struck out a progressive path,
+at first in common, which explains the many points of resemblance
+between them, without regarding man as derived directly from the
+anthropoids. Their many striking points of agreement indicate a common
+descent, and cannot be explained as phenomena of convergence.
+
+I believe I have shown in the above sketch that a theory which derives
+man directly from lower forms without regarding apes as
+transition-types leads _ad absurdum_. The close structural
+relationship between man and monkeys can only be understood if both
+are brought into the same line of evolution. To trace man's line of
+descent directly back to the old Eocene mammals, alongside of, but
+with no relation to these very similar forms, is to abandon the method
+of exact comparison, which, as Darwin rightly recognised, alone
+justifies us in drawing up genealogical trees on the basis of
+resemblances and differences. The farther down we go the more does the
+ground slip from beneath our feet. Even the Lemuridae show very
+numerous divergent conditions, much more so the Eocene mammals
+(Creodonta, Condylarthra), the chief resemblance of which to man
+consists in the possession of pentadactylous hands and feet! Thus the
+farther course of the line of descent disappears in the darkness of
+the ancestry of the mammals. With just as much reason we might pass by
+the Vertebrates altogether, and go back to the lower Invertebrates,
+but in that case it would be much easier to say that man has arisen
+independently, and has evolved, without relation to any animals, from
+the lowest primitive form to his present isolated and dominant
+position. But this would be to deny all value to classification, which
+must after all be the ultimate basis of a genealogical tree. We can,
+as Darwin rightly observed, only infer the line of descent from the
+degree of resemblance between single forms. If we regard man as
+directly derived from primitive forms very far back, we have no way of
+explaining the many points of agreement between him and the monkeys in
+general, and the anthropoid apes in particular. These must remain an
+inexplicable marvel.
+
+I have thus, I trust, shown that the first class of special theories
+of descent, which assumes that man has developed, parallel with the
+monkeys, but without relation to them, from very low primitive forms
+cannot be upheld, because it fails to take into account the close
+structural affinity of man and monkeys. I cannot but regard this
+hypothesis as lamentably retrograde, for it makes impossible any
+application of the facts that have been discovered in the course of
+the anatomical and embryological study of man and monkeys, and indeed
+prejudges investigations of that class as pointless. The whole method
+is perverted; an unjustifiable theory of descent is first formulated
+with the aid of the imagination, and then we are asked to declare that
+all structural relations between man and monkeys, and between the
+different groups of the latter, are valueless,--the fact being that
+they are the only true basis on which a genealogical tree can be
+constructed.
+
+So much for this most modern method of classification, which has
+probably found adherents because it would deliver us from the
+relationship to apes which many people so much dislike. In contrast to
+it we have the second class of special hypotheses of descent, which
+keeps strictly to the nearest structural relationship. This is the
+only basis that justifies the drawing up of a special hypothesis of
+descent. If this fundamental proposition be recognised, it will be
+admitted that the doctrine of special descent upheld by Haeckel, and
+set forth in Darwin's _Descent of Man_, is still valid to-day. In the
+genealogical tree, man's place is quite close to the anthropoid apes;
+these again have as their nearest relatives the lower Old World
+monkeys, and their progenitors must be sought among the less
+differentiated Platyrrhine monkeys, whose most important characters
+have been handed on to the present day New World monkeys. How the
+different genera are to be arranged within the general scheme
+indicated depends in the main on the classificatory value attributed
+to individual characters. This is particularly true in regard to
+_Pithecanthropus_, which I consider as the root of a branch which has
+sprung from the anthropoid ape root and has led up to man; the latter
+I have designated the family of the Hominidae.
+
+For the rest, there are, as we have said, various possible ways of
+constructing the narrower genealogy within the limits of this branch
+including men and apes, and these methods will probably continue to
+change with the accumulation of new facts. Haeckel himself has
+modified his genealogical tree of the Primates in certain details
+since the publication of his _Generelle Morphologie_ in 1866, but its
+general basis remains the same.[123] All the special genealogical
+trees drawn up on the lines laid down by Haeckel and Darwin--and that
+of Dubois may be specially mentioned--are based, in general, on the
+close relationship of monkeys and men, although they may vary in
+detail. Various hypotheses have been formulated on these lines, with
+special reference to the evolution of man. _Pithecanthropus_ is
+regarded by some authorities as the direct ancestor of man, by others
+as a side-track failure in the attempt at the evolution of man. The
+problem of the monophyletic or polyphyletic origin of the human race
+has also been much discussed. Sergi[124] inclines towards the
+assumption of a polyphyletic origin of the three main races of man,
+the African primitive form of which has given rise also to the
+gorilla and chimpanzee, the Asiatic to the Orang, the Gibbon, and
+_Pithecanthropus_. Kollmann regards existing human races as derived
+from small primitive races (pigmies), and considers that _Homo
+primigenius_ must have arisen in a secondary and degenerative manner.
+
+But this is not the place, nor have I the space to criticise the
+various special theories of descent. One, however, must receive
+particular notice. According to Ameghino, the South American monkeys
+(_Pitheculites_) from the oldest Tertiary of the Pampas are the forms
+from which have arisen the existing American monkeys on the one hand,
+and on the other, the extinct South American Homunculidae, which are
+also small forms. From these last, anthropoid apes and man have, he
+believes, been evolved. Among the progenitors of man, Ameghino reckons
+the form discovered by him (_Tetraprothomo_), from which a South
+American primitive man, _Homo pampaeus_, might be directly evolved,
+while on the other hand all the lower Old World monkeys may have
+arisen from older fossil South American forms (Clenialitidae), the
+distribution of which may be explained by the bridge formerly existing
+between South America and Africa, as may be the derivation of all
+existing human races from _Homo pampaeus_.[125] The fossil forms
+discovered by Ameghino deserve the most minute investigation, as does
+also the fossil man from South America of which Lehmann-Nitsche[126]
+has made a thorough study.
+
+It is obvious that, notwithstanding the necessity for fitting man's
+line of descent into the genealogical tree of the Primates, especially
+the apes, opinions in regard to it differ greatly in detail. This
+could not be otherwise, since the different Primate forms, especially
+the fossile forms, are still far from being exhaustively known. But
+one thing remains certain,--the idea of the close relationship between
+man and monkeys set forth in Darwin's _Descent of Man_. Only those who
+deny the many points of agreement, the sole basis of classification,
+and thus of a natural genealogical tree, can look upon the position of
+Darwin and Haeckel as antiquated, or as standing on an insufficient
+foundation. For such a genealogical tree is nothing more than a
+summarised representation of what is known in regard to the degree of
+resemblance between the different forms.
+
+Darwin's work in regard to the descent of man has not been surpassed;
+the more we immerse ourselves in the study of the structural
+relationships between apes and man, the more is our path illumined by
+the clear light radiating from him, and through his calm and
+deliberate investigation, based on a mass of material in the
+accumulation of which he has never had an equal. Darwin's fame will be
+bound up for all time with the unprejudiced investigation of the
+question of all questions, the descent of the human race.
+
+FOOTNOTES:
+
+[Footnote 75: _Life and Letters of Thomas Henry Huxley_, Vol. I. p.
+171, London, 1900.]
+
+[Footnote 76: _Ibid._, p. 363.]
+
+[Footnote 77: No italics in original.]
+
+[Footnote 78: _Life and Letters of Charles Darwin_, Vol. I. p. 93.]
+
+[Footnote 79: _Ibid._ Vol. II. p. 263.]
+
+[Footnote 80: _Ibid._ Vol. I. p. 94.]
+
+[Footnote 81: _Life and Letters_, Vol. III. p. 175.]
+
+[Footnote 82: _Ibid._ Vol. II. p. 109.]
+
+[Footnote 83: _Ibid._ Vol. III. p. 112.]
+
+[Footnote 84: _Ibid._ Vol. I. pp. 304-317.]
+
+[Footnote 85: _Life and Letters_, Vol. I. p. 309.]
+
+[Footnote 86: _Loc. cit._ p. 313.]
+
+[Footnote 87: _Ibid._ Vol. II. p. 310.]
+
+[Footnote 88: _Ibid._ Vol. III. p. 236. ["C. Ridley," Mr. Francis
+Darwin points out to me, should be H. N. Ridley. A.C.S.]]
+
+[Footnote 89: _Descent of Man_ (Popular Edit., 1901), fig. 1, p. 14.]
+
+[Footnote 90: G. Schwalbe, "Das Darwin'sche Spitzohr beim menschlichen
+Embryo," _Anatom. Anzeiger_, 1889, pp. 176-189, and other papers.]
+
+[Footnote 91: _Descent of Man_, fig. 3, p. 24.]
+
+[Footnote 92: _Descent of man_, p. 6.]
+
+[Footnote 93: _Ibid._ p. 54.]
+
+[Footnote 94: _Descent of Man_, p. 92.]
+
+[Footnote 95: _Ibid._ p. 100.]
+
+[Footnote 96: _Life and letters_, Vol. II. p. 161, June 22, 1859.]
+
+[Footnote 97: _Ibid._ Vol. III. p. 15, March 17, 1863.]
+
+[Footnote 98: _Descent of Man_, p. 132.]
+
+[Footnote 99: _Ibid._ pp. 136, 137.]
+
+[Footnote 100: _Ibid._ p. 143.]
+
+[Footnote 101: _Ibid._ p. 193.]
+
+[Footnote 102: _Descent of Man_, p. 231.]
+
+[Footnote 103: _Descent of Man_, p. 308.]
+
+[Footnote 104: _Life and Letters_, Vol. II. p. 23.]
+
+[Footnote 105: _Loc. cit._ p. 39.]
+
+[Footnote 106: _Loc. cit._ (1856), p. 82.]
+
+[Footnote 107: _Ibid._ Vol. III p. 159.]
+
+[Footnote 108: _Descent of Man_, p. 924.]
+
+[Footnote 109: "Die Hautfarbe des Menschen," _Mitteilungen der
+Anthropologischen Gessellschaft in Wien_, Vol. XXXIV. pp. 331-352.]
+
+[Footnote 110: _Ibid._ p. 947.]
+
+[Footnote 111: _Descent of Man_, p. 240.]
+
+[Footnote 112: "Das geologische Alter der Pithecanthropus-Schichten
+bei Trinil, Ost-Java." _Neues Jahrb. f. Mineralogie_. Festband, 1907.]
+
+[Footnote 113: _Descent of Man_, p. 82.]
+
+[Footnote 114: "La race humaine de Néanderthal ou de Canstatt en
+Belgique." _Arch. de Biologie_, VII. 1887.]
+
+[Footnote 115: Gorjanovic-Kramberger. _Der diluviale Mensch van
+Krapina in Kroatien_, 1906.]
+
+[Footnote 116: _Studien zur Vorgeschichte des Menschen_, 1906, pp. 154
+ff.]
+
+[Footnote 117: "On the cranial and facial characters of the Neandertal
+Race." _Trans. R. Soc._ London, vol. 199, 1908, p. 281.]
+
+[Footnote 118: Since this essay was written Schoetensack has
+discovered near Heidelberg and briefly described an exceedingly
+interesting lower jaw from rocks between the Pliocene and Diluvial
+beds. This exhibits interesting differences from the forms of lower
+jaw of _Homo primigenius_. (Schoetensack, _Der Unterkiefer des Homo
+heidelbergensis_, Leipzig, 1908.) G. S.]
+
+[Footnote 119: _Life and Letters of Thomas Henry Huxley_, Vol. II. p.
+394.]
+
+[Footnote 120: _Descent of Man_, p. 229.]
+
+[Footnote 121: _Loc. cit._]
+
+[Footnote 122: Klaatsch in his last publications speaks in the main
+only of an ancestral form common to men and anthropoid apes.]
+
+[Footnote 123: Haeckels latest genealogical tree is to be found in his
+most recent work, _Unsere Ahnenreihe_. Jena, 1908.]
+
+[Footnote 124: Sergi, G. _Europa_, 1908.]
+
+[Footnote 125: _See_ Ameghino's latest paper, "_Notas preliminaries
+sobre el Tetraprothomo argentinus_," etc. _Anales del Museo nacional
+de Buenos Aires_, XVI. pp. 107-242, 1907.]
+
+[Footnote 126: "Nouvelles recherches sur la formation pampéenne et
+l'homme fossile de la République Argentine." _Rivista del Museo de la
+Plata_, T. XIV. pp. 193-488.]
+
+
+
+
+V
+
+CHARLES DARWIN AS AN ANTHROPOLOGIST
+
+BY ERNST HAECKEL
+
+_Professor of Zoology in the University of Jena_
+
+
+The great advance that anthropology has made in the second half of the
+nineteenth century is due, in the first place, to Darwin's discovery
+of the origin of man. No other problem in the whole field of research
+is so momentous as that of "Man's place in nature," which was justly
+described by Huxley (1863) as the most fundamental of all questions.
+Yet the scientific solution of this problem was impossible until the
+theory of descent had been established.
+
+It is now a hundred years since the great French biologist Jean
+Lamarck published his _Philosophie Zoologique_. By a remarkable
+coincidence the year in which that work was issued, 1809, was the year
+of the birth of his most distinguished successor, Charles Darwin.
+Lamarck had already recognised that the descent of man from a series
+of other Vertebrates--that is, from a series of Ape-like Primates--was
+essentially involved in the general theory of transformation which he
+had erected on a broad inductive basis; and he had sufficient
+penetration to detect the agencies that had been at work in the
+evolution of the erect bimanous man from the arboreal and quadrumanous
+ape. He had, however, few empirical arguments to advance in support of
+his hypothesis, and it could not be established until the further
+development of the biological sciences--the founding of comparative
+embryology by Baer (1828) and of the cell-theory by Schleiden and
+Schwann (1838), the advance of physiology under Johannes Müller
+(1833), and the enormous progress of palaeontology and comparative
+anatomy between 1820 and 1860--provided this necessary foundation.
+Darwin was the first to coordinate the ample results of these lines of
+research. With no less comprehensiveness than discrimination he
+consolidated them as a basis of a modified theory of descent, and
+associated with them his own theory of natural selection, which we
+take to be distinctive of "Darwinism" in the stricter sense. The
+illuminating truth of these cumulative arguments was so great in every
+branch of biology that, in spite of the most vehement opposition, the
+battle was won within a single decade, and Darwin secured the general
+admiration and recognition that had been denied to his forerunner,
+Lamarck, up to the hour of his death (1829).
+
+Before, however, we consider the momentous influence that Darwinism
+has had in anthropology, we shall find it useful to glance at its
+history in the course of the last half century, and notice the various
+theories that have contributed to its advance. The first attempt to
+give extensive expression to the reform of biology by Darwin's work
+will be found in my _Generelle Morphologie_ (1866)[127] which was
+followed by a more popular treatment of the subject in my _Natürliche
+Schöpfungsgeschichte_ (1868),[128] a compilation from the earlier
+work. In the first volume of the _Generelle Morphologie_ I endeavoured
+to show the great importance of evolution in settling the fundamental
+questions of biological philosophy, especially in regard to
+comparative anatomy. In the second volume I dealt broadly with the
+principle of evolution, distinguishing ontogeny and phylogeny as its
+two coordinate main branches, and associating the two in the
+Biogenetic Law. The Law may be formulated thus: "Ontogeny (embryology
+or the development of the individual) is a concise and compressed
+recapitulation of phylogeny (the palaeontological or genealogical
+series) conditioned by laws of heredity and adaptation." The
+"Systematic introduction to general evolution," with which the second
+volume of the _Generelle Morphologie_ opens, was the first attempt to
+draw up a natural system of organisms (in harmony with the principles
+of Lamarck and Darwin) in the form of a hypothetical pedigree, and was
+provisionally set forth in eight genealogical tables.
+
+In the nineteenth chapter of the _Generelle Morphologie_--a part of
+which has been republished, without any alteration, after a lapse of
+forty years--I made a critical study of Lamarck's theory of descent
+and of Darwin's theory of selection, and endeavoured to bring the
+complex phenomena of heredity and adaptation under definite laws for
+the first time. Heredity I divided into conservative and progressive:
+adaptation into indirect (or potential) and direct (or actual). I then
+found it possible to give some explanation of the correlation of the
+two physiological functions in the struggle for life (selection), and
+to indicate the important laws of divergence (or differentiation) and
+complexity (or division of labor), which are the direct and inevitable
+outcome of selection. Finally, I marked off dysteleology as the
+science of the aimless (vestigial, abortive, atrophied, and useless)
+organs and parts of the body. In all this I worked from a strictly
+monistic standpoint, and sought to explain all biological phenomena on
+the mechanical and naturalistic lines that had long been recognised in
+the study of inorganic nature. Then (1866), as now, being convinced of
+the unity of nature, the fundamental identity of the agencies at work
+in the inorganic and the organic worlds, I discarded vitalism,
+teleology, and all hypotheses of a mystic character.
+
+It was clear from the first that it was essential, in the monistic
+conception of evolution, to distinguish between the laws of
+conservative and progressive heredity. Conservative heredity maintains
+from generation to generation the enduring characters of the species.
+Each organism transmits to its descendants a part of the morphological
+and physiological qualities that it has received from its parents and
+ancestors. On the other hand, progressive heredity brings new
+characters to the species--characters that were not found in preceding
+generations. Each organism may transmit to its offspring a part of the
+morphological and physiological features that it has itself acquired,
+by adaptation, in the course of its individual career, through the use
+or disuse of particular organs, the influence of environment, climate,
+nutrition, etc. At that time I gave the name of "progressive heredity"
+to this inheritance of acquired characters, as a short and convenient
+expression, but have since changed the term to "transformative
+heredity" (as distinguished from conservative). This term is
+preferable, as inherited regressive modifications (degeneration,
+retrograde metamorphosis, etc.) come under the same head.
+
+Transformative heredity--or the transmission of acquired
+characters--is one of the most important principles in evolutionary
+science. Unless we admit it most of the facts of comparative anatomy
+and physiology are inexplicable. That was the conviction of Darwin no
+less than of Lamarck, of Spencer as well as Virchow, of Huxley as well
+as Gegenbaur, indeed of the great majority of speculative biologists.
+This fundamental principle was for the first time called in question
+and assailed in 1885 by August Weismann of Freiburg, the eminent
+zoologist to whom the theory of evolution owes a great deal of
+valuable support, and who has attained distinction by his extension of
+the theory of selection. In explanation of the phenomena of heredity
+he introduced a new theory, the "theory of the continuity of the
+germ-plasm." According to him the living substance in all organisms
+consists of two quite distinct kinds of plasm, somatic and germinal.
+The permanent germ-plasm, or the active substance of the two
+germ-cells (egg-cell and sperm-cell), passes unchanged through a
+series of generations, and is not affected by environmental
+influences. The environment modifies only the soma-plasm, the organs
+and tissues of the body. The modifications that these parts undergo
+through the influence of the environment or their own activity (use
+and habit), do not affect the germ-plasm, and cannot therefore be
+transmitted.
+
+This theory of the continuity of the germ-plasm has been expounded by
+Weismann during the last twenty-four years in a number of able
+volumes, and is regarded by many biologists, such as Mr. Francis
+Galton, Sir E. Ray Lankester, and Professor J. Arthur Thomson (who has
+recently made a thorough-going defence of it in his important work
+_Heredity_),[129] as the most striking advance in evolutionary
+science. On the other hand, the theory has been rejected by Herbert
+Spencer, Sir W. Turner, Gegenbaur, Kölliker, Hertwig, and many others.
+For my part I have, with all respect for the distinguished Darwinian,
+contested the theory from the first, because its whole foundation
+seems to me erroneous, and its deductions do not seem to be in accord
+with the main facts of comparative morphology and physiology.
+Weismann's theory in its entirety is a finely conceived molecular
+hypothesis, but it is devoid of empirical basis. The notion of the
+absolute and permanent independence of the germ-plasm, as
+distinguished from the soma-plasm, is purely speculative; as is also
+the theory of germinal selection. The determinants, ids, and idants,
+are purely hypothetical elements. The experiments that have been
+devised to demonstrate their existence really prove nothing.
+
+It seems to me quite improper to describe this hypothetical structure
+as "Neodarwinism." Darwin was just as convinced as Lamarck of the
+transmission of acquired characters and its great importance in the
+scheme of evolution. I had the good fortune to visit Darwin at Down
+three times and discuss with him the main principles of his system,
+and on each occasion we were fully agreed as to the incalculable
+importance of what I may call transformative inheritance. It is only
+proper to point out that Weismann's theory of the germ-plasm is in
+express contradiction to the fundamental principles of Darwin and
+Lamarck. Nor is it more acceptable in what one may call its
+"ultradarwinism"--the idea that the theory of selection explains
+everything in the evolution of the organic world. This belief in the
+"omnipotence of natural selection" was not shared by Darwin himself.
+Assuredly, I regard it as of the utmost value, as the process of
+natural selection through the struggle for life affords an explanation
+of the mechanical origin of the adapted organisation. It solves the
+great problem: how could the finely adapted structure of the animal or
+plant body be formed unless it was built on a preconceived plan? It
+thus enables us to dispense with the teleology of the metaphysician
+and the dualist, and to set aside the old mythological and poetic
+legends of creation. The idea had occurred in vague form to the great
+Empedocles 2000 years before the time of Darwin, but it was reserved
+for modern research to give it ample expression. Nevertheless, natural
+selection does not of itself give the solution of all our evolutionary
+problems. It has to be taken in conjunction with the transformism of
+Lamarck, with which it is in complete harmony.
+
+The monumental greatness of Charles Darwin, who surpasses every other
+student of science in the nineteenth century by the loftiness of his
+monistic conception of nature and the progressive influence of his
+ideas, is perhaps best seen in the fact that not one of his many
+successors has succeeded in modifying his theory of descent in any
+essential point or in discovering an entirely new standpoint in the
+interpretation of the organic world. Neither Nägeli nor Weismann,
+neither De Vries nor Roux, has done this. Nägeli, in his
+_Mechanisch-Physiologische Theorie der Abstammungslehre_[130] which is
+to a great extent in agreement with Weismann, constructed a theory of
+the idioplasm, that represents it (like the germ-plasm) as developing
+continuously in a definite direction from internal causes. But his
+internal "principle of progress" is at the bottom just as teleological
+as the vital force of the Vitalists, and the micella structure of the
+idioplasm is just as hypothetical as the "dominant" structure of the
+germ-plasm. In 1889 Moritz Wagner sought to explain the origin of
+species by migration and isolation, and on that basis constructed a
+special "migration-theory." This, however, is not out of harmony with
+the theory of selection. It merely elevates one single factor in the
+theory to a predominant position. Isolation is only a special case of
+selection, as I had pointed out in the fifteenth chapter of my
+_Natural history of creation_. The "mutation-theory" of De Vries,[131]
+that would explain the origin of species by sudden and saltatory
+variations rather than by gradual modification, is regarded by many
+botanists as a great step in advance, but it is generally rejected by
+zoologists. It affords no explanation of the facts of adaptation, and
+has no causal value.
+
+Much more important than these theories is that of Wilhelm Roux[132]
+of "the struggle of parts within the organism, a supplementation of
+the theory of mechanical adaptation." He explains the functional
+autoformation of the purposive structure by a combination of Darwin's
+principle of selection with Lamarck's idea of transformative heredity,
+and applies the two in conjunction to the facts of histology. He lays
+stress on the significance of functional adaptation, which I had
+described in 1866, under the head of cumulative adaptation, as the
+most important factor in evolution. Pointing out its influence in the
+cell-life of the tissues, he puts "cellular selection" above "personal
+selection," and shows how the finest conceivable adaptations in the
+structure of the tissue may be brought about quite mechanically,
+without preconceived plan. This "mechanical teleology" is a valuable
+extension of Darwin's monistic principle of selection to the whole
+field of cellular physiology and histology, and is wholly destructive
+of dualistic vitalism.
+
+The most important advance that evolution has made since Darwin and
+the most valuable amplification of his theory of selection is, in my
+opinion, the work of Richard Semon: _Die Mneme als erhaltendes Prinzip
+im Wechsel des organischen Geschehens_.[133] He offers a psychological
+explanation of the facts of heredity by reducing them to a process of
+(unconscious) memory. The physiologist Ewald Hering had shown in 1870
+that memory must be regarded as a general function of organic matter,
+and that we are quite unable to explain the chief vital phenomena,
+especially those of reproduction and inheritance, unless we admit this
+unconscious memory. In my essay _Die Perigenesis der Plastidule_[134]
+I elaborated this far-reaching idea, and applied the physical
+principle of transmitted motion to the plastidules, or active
+molecules of plasm. I concluded that "heredity is the memory of the
+plastidules, and variability their power of comprehension." This
+"provisional attempt to give a mechanical explanation of the
+elementary processes of evolution" I afterwards extended by showing
+that sensitiveness is (as Carl Nägeli, Ernst Mach, and Albrecht Rau
+express it) a general quality of matter. This form of panpsychism
+finds its simplest expression in the "trinity of substance."
+
+To the two fundamental attributes that Spinoza ascribed to
+substance--Extension (matter as occupying space) and Cogitation
+(energy, force)--we now add the third fundamental quality of Psychoma
+(sensitiveness, soul). I further elaborated this trinitarian
+conception of substance in the nineteenth chapter of my _Die
+Lebenswunder_ (1904),[135] and it seems to me well calculated to
+afford a monistic solution of many of the antitheses of philosophy.
+
+This important Mneme-theory of Semon and the luminous physiological
+experiments and observations associated with it not only throw
+considerable light on transformative inheritance, but provide a sound
+physiological foundation for the biogenetic law. I had endeavoured to
+show in 1874, in the first chapter of my _Anthropogenie_,[136] that
+this fundamental law of organic evolution holds good generally, and
+that there is everywhere a direct causal connection between ontogeny
+and phylogeny. "Phylogenesis is the mechanical cause of ontogenesis;"
+in other words, "The evolution of the stem or race is--in accordance
+with the laws of heredity and adaptation--the real cause of all the
+changes that appear, in a condensed form, in the development of the
+individual organism from the ovum, in either the embryo or the larva."
+
+It is now fifty years since Charles Darwin pointed out, in the
+thirteenth chapter of his epoch-making _Origin of Species_, the
+fundamental importance of embryology in connection with his theory of
+descent:
+
+"The leading facts in embryology, which are second to none in
+importance, are explained on the principle of variations in the many
+descendants from some one ancient progenitor, having appeared at a not
+very early period of life, and having been inherited at a
+corresponding period."[137]
+
+He then shows that the striking resemblance of the embryos and larvae
+of closely related animals, which in the mature stage belong to widely
+different species and genera, can only be explained by their descent
+from a common progenitor. Fritz Müller made a closer study of these
+important phenomena in the instructive instance of the Crustacean
+larva, as given in his able work _Für Darwin_[138] (1864). I then, in
+1872, extended the range so as to include all animals (with the
+exception of the unicellular Protozoa) and showed, by means of the
+theory of the Gastraea, that all multicellular, tissue-forming
+animals--all the Metazoa--develop in essentially the same way from the
+primary germ-layers.
+
+I conceived the embryonic form, in which the whole structure consists of
+only two layers of cells, and is known as the gastrula, to be the
+ontogenetic recapitulation, maintained by tenacious heredity, of a
+primitive common progenitor of all the Metazoa, the Gastraea. At a later
+date (1895) Monticelli discovered that this conjectural ancestral form is
+still preserved in certain primitive Coelenterata--Pemmatodiscus,
+Kunstleria, and the nearly-related Orthonectida.
+
+The general application of the biogenetic law to all classes of
+animals and plants has been proved in my _Systematische
+Phylogenie_.[139] It has, however, been frequently challenged, both by
+botanists and zoologists, chiefly owing to the fact that many have
+failed to distinguish its two essential elements, palingenesis and
+cenogenesis. As early as 1874 I had emphasised, in the first chapter
+of my _Evolution of Man_, the importance of discriminating carefully
+between these two sets of phenomena:
+
+"In the evolutionary appreciation of the facts of embryology we must
+take particular care to distinguish sharply and clearly between the
+primary, palingenetic evolutionary processes and the secondary,
+cenogenetic processes. The palingenetic phenomena, or embryonic
+_recapitulations_, are due to heredity, to the transmission of
+characters from one generation to another. They enable us to draw
+direct inferences in regard to corresponding structures in the
+development of the species (e.g. the chorda or the branchial arches in
+all vertebrate embryos). The cenogenetic phenomena, on the other hand,
+or the embryonic _variations_, cannot be traced to inheritance from a
+mature ancestor, but are due to the adaption of the embryo or the
+larva to certain conditions of its individual development (e.g. the
+amnion, the allantois, and the vitelline arteries in the embryos of
+the higher vertebrates). These cenogenetic phenomena are later
+additions; we must not infer from them that there were corresponding
+processes in the ancestral history, and hence they are apt to
+mislead."
+
+The fundamental importance of these facts of comparative anatomy,
+atavism, and the rudimentary organs, was pointed out by Darwin in the
+first part of his classic work, _The Descent of Man and Selection in
+Relation to Sex_ (1871).[140] In the "General summary and conclusion"
+(chap. xxi.) he was able to say, with perfect justice: "He who is not
+content to look, like a savage, at the phenomena of nature as
+disconnected, cannot any longer believe that man is the work of a
+separate act of creation. He will be forced to admit that the close
+resemblance of the embryo of man to that, for instance, of a dog--the
+construction of his skull, limbs, and whole frame on the same plan
+with that of other mammals, independently of the uses to which the
+parts may be put--the occasional reappearance of various structures,
+for instance of several muscles, which man does not normally possess,
+but which are common to the Quadrumana--and a crowd of analogous
+facts--all point in the plainest manner to the conclusion that man is
+the co-descendant with other mammals of a common progenitor."
+
+These few lines of Darwin's have a greater scientific value than
+hundreds of those so-called "anthropological treatises," which give
+detailed descriptions of single organs, or mathematical tables with
+series of numbers and what are claimed to be "exact analyses," but are
+devoid of synoptic conclusions and a philosophical spirit.
+
+Charles Darwin is not generally recognised as a great anthropologist,
+nor does the school of modern anthropologists regard him as a leading
+authority. In Germany, especially, the great majority of the members
+of the anthropological societies took up an attitude of hostility to
+him from the very beginning of the controversy in 1860. _The Descent
+of Man_ was not merely rejected, but even the discussion of it was
+forbidden on the ground that it was "unscientific."
+
+The centre of this inveterate hostility for thirty years--especially
+after 1877--was Rudolph Virchow of Berlin, the leading investigator
+in pathological anatomy, who did so much for the reform of medicine by
+his establishment of cellular pathology in 1858. As a prominent
+representative of "exact" or "descriptive" anthropology, and lacking a
+broad equipment in comparative anatomy and ontogeny, he was unable to
+accept the theory of descent. In earlier years, and especially during
+his splendid period of activity at Würzburg (1848-1856), he had been a
+consistent free-thinker, and had in a number of able articles
+(collected in his _Gesammelte Abhandlungen_)[141] upheld the unity of
+human nature, the inseparability of body and spirit. In later years at
+Berlin, where he was more occupied with political work and sociology
+(especially after 1866), he abandoned the positive monistic position
+for one of agnosticism and scepticism, and made concessions to the
+dualistic dogma of a spiritual world apart from the material frame.
+
+In the course of a Scientific Congress at Munich in 1877 the conflict
+of these antithetic views of nature came into sharp relief. At this
+memorable Congress I had undertaken to deliver the first address
+(September 18th) on the subject of "Modern evolution in relation to
+the whole of science." I maintained that Darwin's theory not only
+solved the great problem of the origin of species, but that its
+implications, especially in regard to the nature of man, threw
+considerable light on the whole of science, and on anthropology in
+particular. The discovery of the real origin of man by evolution from
+a long series of mammal ancestors threw light on his place in nature
+in every respect, as Huxley had already shown in his excellent
+lectures of 1863. Just as all the organs and tissues of the human body
+had originated from those of the nearest related mammals, certain
+ape-like forms, so we were bound to conclude that his mental qualities
+also had been derived from those of his extinct primate ancestor.
+
+This monistic view of the origin and nature of man, which is now admitted
+by nearly all who have the requisite acquaintance with biology, and
+approach the subject without prejudice, encountered a sharp opposition at
+that time. The opposition found its strongest expression in an address that
+Virchow delivered at Munich four days afterwards (September 22nd), on "The
+freedom of science in the modern State." He spoke of the theory of
+evolution as an unproved hypothesis, and declared that it ought not to be
+taught in the schools, because it was dangerous to the State. "We must
+not," he said, "teach that man has descended from the ape or any other
+animal." When Darwin, usually so lenient in his judgment, read the English
+translation of Virchow's speech, he expressed his disapproval in strong
+terms. But the great authority that Virchow had--an authority well founded
+in pathology and sociology--and his prestige as president of the German
+Anthropological Society, had the effect of preventing any member of the
+Society from raising serious opposition to him for thirty years. Numbers of
+journals and treatises repeated his dogmatic statement: "It is quite
+certain that man has descended neither from the ape nor from any other
+animal." In this he persisted till his death in 1902. Since that time the
+whole position of German anthropology has changed. The question is no
+longer whether man was created by a distinct supernatural act or evolved
+from other mammals, but to which line of the animal hierarchy we must look
+for the actual series of ancestors. The interested reader will find an
+account of this "battle of Munich" (1877) in my three Berlin lectures
+(April, 1905), _Der Kampf um die Entwickelungs-Gedanken_.[142]
+
+The main points in our genealogical tree were clearly recognised by
+Darwin in the sixth chapter of the _Descent of Man_. Lowly organised
+fishes, like the lancelot (Amphioxus), are descended from lower
+invertebrates resembling the larvae of an existing Tunicate
+(Appendicularia). From these primitive fishes were evolved higher
+fishes of the ganoid type and others of the type of Lepidosiren
+(Dipneusta). It is a very small step from these to the Amphibia:
+
+"In the class of animals the steps are not difficult to conceive which
+led from the ancient Monotremata to the ancient Marsupials; and from
+these to the early progenitors of the placental mammals. We may thus
+ascend to the Lemuridae; and the interval is not very wide from these
+to the Simiadae. The Simiadae then branched off into two great stems,
+the New World and Old World monkeys; and from the latter, at a remote
+period, Man, the wonder and glory of the Universe, proceeded."[143]
+
+In these few lines Darwin clearly indicated the way in which we were
+to conceive our ancestral series within the vertebrates. It is fully
+confirmed by all the arguments of comparative anatomy and embryology,
+of palaeontology and physiology; and all the research of the
+subsequent forty years have gone to establish it. The deep interest in
+geology which Darwin maintained throughout his life and his complete
+knowledge of palaeontology enabled him to grasp the fundamental
+importance of the palaeontological record more clearly than
+anthropologists and zoologists usually do.
+
+There has been much debate in subsequent decades whether Darwin
+himself maintained that man was descended from the ape, and many
+writers have sought to deny it. But the lines I have quoted _verbatim_
+from the conclusion of the sixth chapter of the _Descent of Man_
+(1871) leave no doubt that he was as firmly convinced of it as was his
+great precursor Jean Lamarck in 1809. Moreover, Darwin adds, with
+particular explicitness, in the "general summary and conclusion"
+(chap. xxi.) of that standard work:[144]
+
+"By considering the embryological structure of man--the homologies
+which he presents with the lower animals,--the rudiments which he
+retains,--and the reversions to which he is liable, we can partly
+recall in imagination the former condition of our early progenitors;
+and can approximately place them in their proper place in the
+zoological series. We thus learn that man is descended from a hairy,
+tailed quadruped, probably arboreal in its habits, and an inhabitant
+of the Old World. This creature, if its whole structure had been
+examined by a naturalist, would have been classed amongst the
+Quadrumana, as surely as the still more ancient progenitor of the Old
+and New World monkeys."
+
+These clear and definite lines leave no doubt that Darwin--so critical
+and cautious in regard to important conclusions--was quite as firmly
+convinced of the descent of man from the apes (the Catarrhinae, in
+particular) as Lamarck was in 1809 and Huxley in 1863.
+
+It is to be noted particularly that, in these and other observations
+on the subject, Darwin decidedly assumes the monophyletic origin of
+the mammals, including man. It is my own conviction that this is of
+the greatest importance. A number of difficult questions in regard to
+the development of man, in respect of anatomy, physiology, psychology,
+and embryology, are easily settled if we do not merely extend our
+_progonotaxis_ to our nearest relatives, the anthropoid apes and the
+tailed monkeys from which these have descended, but go further back
+and find an ancestor in the group of the Lemuridae, and still further
+back to the Marsupials and Monotremata. The essential identity of all
+the Mammals in point of anatomical structure and embryonic
+development--in spite of their astonishing differences in external
+appearance and habits of life--is so palpably significant that modern
+zoologists are agreed in the hypothesis that they have all sprung from
+a common root, and that this root may be sought in the earlier
+Palaeozoic Amphibia.
+
+The fundamental importance of this comparative morphology of the
+Mammals, as a sound basis of scientific anthropology, was recognised
+just before the beginning of the nineteenth century, when Lamarck
+first emphasised (1794) the division of the animal kingdom into
+Vertebrates and Invertebrates. Even thirteen years earlier (1781),
+when Goethe made a close study of the mammal skeleton in the
+Anatomical Institute at Jena, he was intensely interested to find that
+the composition of the skull was the same in man as in the other
+mammals. His discovery of the _os inter-maxillare_ in man (1784),
+which was contradicted by most of the anatomists of the time, and his
+ingenious "vertebral theory of the skull," were the splendid fruit of
+his morphological studies. They remind us how Germany's greatest
+philosopher and poet was for many years ardently absorbed in the
+comparative anatomy of man and the mammals, and how he divined that
+their wonderful identity in structure was no mere superficial
+resemblance, but pointed to a deep internal connection. In my
+_Generelle Morphologie_ (1866), in which I published the first
+attempts to construct phylogenetic trees, I have given a number of
+remarkable theses of Goethe, which may be called "phyletic
+prophecies." They justify us in regarding him as a precursor of
+Darwin.
+
+In the ensuing forty years I have made many conscientious efforts to
+penetrate further along that line of anthropological research that was
+opened up by Goethe, Lamarck, and Darwin. I have brought together the many
+valuable results that have constantly been reached in comparative anatomy,
+physiology, ontogeny, and palaeontology, and maintained the effort to
+reform the classification of animals and plants in an evolutionary sense.
+The first rough drafts of pedigrees that were published in the _Generelle
+Morphologie_ have been improved time after time in the ten editions of my
+_Natürlich Schöpfungsgeschichte_ (1868-1902).[145] A sounded basis for my
+phyletic hypotheses, derived from a discriminating combination of the three
+great records--morphology, ontogeny, and palaeontology--was provided in the
+three volumes of my _Systematische Phylogenie_[146] (1894 Protists and
+Plants, 1895 Vertebrates, 1896 Invertebrates).
+
+In my _Anthropogenie_[147] I endeavoured to employ all the known
+facts of comparative ontogeny (embryology) for the purpose of
+completing my scheme of human phylogeny (evolution). I attempted to
+sketch the historical development of each organ of the body, beginning
+with the most elementary structures in the germ-layers of the
+Gastraea. At the same time I drew up a corrected statement of the most
+important steps in the line of our ancestral series.
+
+At the fourth International Congress of Zoology at Cambridge (August
+26th, 1898) I delivered an address on "Our present knowledge of the
+Descent of Man." It was translated into English, enriched with many
+valuable notes and additions, by my friend and pupil in earlier days
+Dr. Hans Gadow (Cambridge), and published under the title: _The Last
+Link: our present knowledge of the Descent of Man_[148] The
+determination of the chief animal forms that occur in the line of our
+ancestry is there restricted to thirty types, and these are
+distributed in six main groups.
+
+The first half of this "Progonotaxis hominis," which has no support
+from fossil evidence, comprises three groups: (i) Protista
+(unicellular organisms, 1-5): (ii) Invertebrate Metazoa (Coelenteria
+6-8, Vermalia 9-11): (iii) Monorrhine Vertebrates (Acrania 12-13,
+Cyclostoma 14-15). The second half, which is based on fossil records,
+also comprises three groups: (iv) Palaeozoic cold-blooded Craniota
+(Fishes 16-18, Amphibia 19, Reptiles 20): (v) Mesozoic Mammals
+(Monotrema 21, Marsupialia 22, Mallotheria 23): (vi) Cenozoic Primates
+(Lemuridae 24-25, Tailed Apes 26-27, Anthropomorpha 28-30). An
+improved and enlarged edition of this hypothetic "Progonotaxis
+hominis" was published in 1908, in my essay _Unsere Ahnenreihe_.[149]
+
+If I have succeeded in furthering, in some degree, by these
+anthropological works, the solution of the great problem of Man's
+place in nature, and particularly in helping to trace the definite
+stages in our ancestral series, I owe the success, not merely to the
+vast progress that biology has made in the last half century, but
+largely to the luminous example of the great investigators who have
+applied themselves to the problem, with so much assiduity and genius,
+for a century and a quarter--I mean Goethe and Lamarck, Gegenbaur and
+Huxley, but, above all, Charles Darwin. It was the great genius of
+Darwin that first brought together that symmetrical temple of
+scientific knowledge, the theory of descent. It was Darwin who put the
+crown on the edifice by his theory of natural selection. Not until
+this broad inductive law was firmly established was it possible to
+vindicate the special conclusion, the descent of man from a series of
+other Vertebrates. By his illuminating discovery Darwin did more for
+anthropology than thousands of those writers, who are more
+specifically titled anthropologists, have done by their technical
+treatises. We may, indeed, say that it is not merely as an exact
+observer and ingenious experimenter, but as a distinguished
+anthropologist and far-seeing thinker, that Darwin takes his place
+among the greatest men of science of the nineteenth century.
+
+To appreciate fully the immortal merit of Darwin in connection with
+anthropology, we must remember that not only did his chief work, _The
+Origin of Species_, which opened up a new era in natural history in
+1859, sustain the most virulent and widespread opposition for a
+lengthy period, but even thirty years later, when its principles were
+generally recognised and adopted, the application of them to man was
+energetically contested by many high scientific authorities. Even
+Alfred Russel Wallace, who discovered the principle of natural
+selection independently in 1858, did not concede that it was
+applicable to the higher mental and moral qualities of man. Dr.
+Wallace still holds a spiritualist and dualist view of the nature of
+man, contending that he is composed of a material frame (descended
+from the apes) and an immortal immaterial soul (infused by a higher
+power). This dual conception, moreover, is still predominant in the
+wide circles of modern theology and metaphysics, and has the general
+and influential adherence of the more conservative classes of society.
+
+In strict contradiction to this mystical dualism, which is generally
+connected with teleology and vitalism, Darwin always maintained the
+complete unity of human nature, and showed convincingly that the
+psychological side of man was developed, in the same way as the body,
+from the less advanced soul of the anthropoid ape, and, at a still
+more remote period, from the cerebral functions of the older
+vertebrates. The eighth chapter of the _Origin of Species_, which is
+devoted to instinct, contains weighty evidence that the instincts of
+animals are subject, like all other vital processes, to the general
+laws of historic development. The special instincts of particular
+species were formed by adaptation, and the modifications thus acquired
+were handed on to posterity by heredity; in their formation and
+preservation natural selection plays the same part as in the
+transformation of every other physiological function. The higher moral
+qualities of civilised man have been derived from the lower mental
+functions of the uncultivated barbarians and savages, and these in
+turn from the social instincts of the mammals. This natural and
+monistic psychology of Darwin's was afterwards more fully developed by
+his friend George Romanes in his excellent works _Mental Evolution in
+Animals_ and _Mental Evolution in Man_.[150]
+
+Many valuable and most interesting contributions to this monistic
+psychology of man were made by Darwin in his fine work on _The Descent
+of Man and Selection in Relation to Sex_, and again in his
+supplementary work, _The Expression of the Emotions in Man and
+Animals_. To understand the historical development of Darwin's
+anthropology one must read his life and the introduction to _The
+Descent of Man_. From the moment that he was convinced of the truth
+of the principle of descent--that is to say, from his thirtieth year,
+in 1838--he recognised clearly that man could not be excluded from its
+range. He recognised as a logical necessity the important conclusion
+that "man is the co-descendant with other species of some ancient,
+lower, and extinct form." For many years he gathered notes and
+arguments in support of this thesis, and for the purpose of showing
+the probable line of man's ancestry. But in the first edition of _The
+Origin of Species_ (1859) he restricted himself to the single line,
+that by this work "light would be thrown on the origin of man and his
+history." In the fifty years that have elapsed since that time the
+science of the origin and nature of man has made astonishing progress,
+and we are now fairly agreed in a monistic conception of nature that
+regards the whole universe, including man, as a wonderful unity,
+governed by unalterable and eternal laws. In my philosophical book
+_Die Welträtsel_ (1899)[151] and in the supplementary volume _Die
+Lebenswunder_ (1904)[152] I have endeavoured to show that this pure
+monism is securely established, and that the admission of the
+all-powerful rule of the same principle of evolution throughout the
+universe compels us to formulate a single supreme law--the
+all-embracing "Law of Substance," or the united laws of the constancy
+of matter and the conservation of energy. We should never have reached
+this supreme general conception if Charles Darwin--a "monistic
+philosopher" in the true sense of the word--had not prepared the way
+by his theory of descent by natural selection, and crowned the great
+work of his life by the association of this theory with a naturalistic
+anthropology.
+
+FOOTNOTES:
+
+[Footnote 127: _Generelle Morphologie der Organismen_, 2 vols.,
+Berlin, 1866.]
+
+[Footnote 128: Eng. transl.; _The History of Creation_, London, 1876.]
+
+[Footnote 129: London, 1908.]
+
+[Footnote 130: Munich, 1884.]
+
+[Footnote 131: _Die Mutationstheorie_, Leipzig, 1903.]
+
+[Footnote 132: _Der Kampf der Theile im Organismus_, Leipzig, 1881.]
+
+[Footnote 133: Leipzig, 1904.]
+
+[Footnote 134: Berlin, 1876.]
+
+[Footnote 135: _Wonders of Life_, London and New York, 1904.]
+
+[Footnote 136: Eng. transl.; _The Evolution of Man_, 2 vols., London,
+1879 and 1905.]
+
+[Footnote 137: _Origin of Species_ (6th edit.), p. 396.]
+
+[Footnote 138: Eng. transl.; _Facts and Arguments for Darwin_, London,
+1869.]
+
+[Footnote 139: 3 vols., Berlin, 1894-96.]
+
+[Footnote 140: _Descent of Man_ (Popular Edit.), p. 927.]
+
+[Footnote 141: _Gesammelte Abhandlungen zur wissenschaftlichen
+Medizin_, Berlin, 1856.]
+
+[Footnote 142: Eng. transl.; _Last Words on Evolution_, London, 1906.]
+
+[Footnote 143: _Descent of Man_, (Popular Edit.), p. 255.]
+
+[Footnote 144: _Descent of Man_, p. 930.]
+
+[Footnote 145: Eng. transl.; _The History of Creation_, London, 1876.]
+
+[Footnote 146: Berlin, 1894-96.]
+
+[Footnote 147: Leipzig, 1874, 5th edit. 1905. Eng. transl.; _The
+Evolution of Man_, London, 1905.]
+
+[Footnote 148: London, 1898.]
+
+[Footnote 149: _Festschrift zur 350-jährigen Jubelfeier der Thüringer
+Universität Jena_. Jena. 1908.]
+
+[Footnote 150: London, 1885; 1888.]
+
+[Footnote 151: _The Riddle of the Universe_, London and New York,
+1900.]
+
+[Footnote 152: _The Wonders of Life_, London and New York, 1904.]
+
+
+
+
+VI
+
+MENTAL FACTORS IN EVOLUTION
+
+BY C. LLOYD MORGAN, LLD., F.R.S
+
+
+In developing his conception of organic evolution Charles Darwin was
+of necessity brought into contact with some of the problems of mental
+evolution. In _The Origin of Species_ he devoted a chapter to "the
+diversities of instinct and of the other mental faculties in animals
+of the same class."[153] When he passed to the detailed consideration
+of _The Descent of Man_, it was part of his object to show "that there
+is no fundamental difference between man and the higher mammals in
+their mental faculties."[154] "If no organic being excepting man," he
+said, "had possessed any mental power, or if his powers had been of a
+wholly different nature, from those of the lower animals, then we
+should never have been able to convince ourselves that our high
+faculties had been gradually developed."[155] In his discussion of
+_The Expression of the Emotions_ it was important for his purpose
+"fully to recognise that actions readily become associated with other
+actions and with various states of the mind."[156] His hypothesis of
+sexual selection is largely dependent upon the exercise of choice on
+the part of the female and her preference for "not only the more
+attractive but at the same time the more vigourous and vicious
+males."[157] Mental processes and physiological processes were for
+Darwin closely correlated; and he accepted the conclusion "that the
+nervous system not only regulates most of the existing functions of
+the body, but has indirectly influenced the progressive development of
+various bodily structures and of certain mental qualities."[158]
+
+Throughout his treatment, mental evolution was for Darwin incidental
+to and contributory to organic evolution. For specialised research in
+comparative and genetic psychology, as an independent field of
+investigation, he had neither the time nor the requisite training.
+None the less his writings and the spirit of his work have exercised a
+profound influence on this department of evolutionary thought. And,
+for those who follow Darwin's lead, mental evolution is still in a
+measure subservient to organic evolution. Mental processes are the
+accompaniments or concomitants of the functional activity of specially
+differentiated parts of the organism. They are in some way dependent
+on physiological and physical conditions. But though they are not
+physical in their nature, and though it is difficult or impossible to
+conceive that they are physical in their origin, they are, for Darwin
+and his followers, factors in the evolutionary process in its physical
+or organic aspect. By the physiologist within his special and
+well-defined universe of discourse they may be properly regarded as
+epiphenomena; but by the naturalist in his more catholic survey of
+nature they cannot be so regarded, and were not so regarded by Darwin.
+Intelligence has contributed to evolution of which it is in a sense a
+product.
+
+The facts of observation or of inference which Darwin accepted are
+these: Conscious experience accompanies some of the modes of animal
+behaviour; it is concomitant with certain physiological processes;
+these processes are the outcome of development in the individual and
+evolution in the race; the accompanying mental processes undergo a
+like development. Into the subtle philosophical questions which arise
+out of the naïve acceptance of such a creed it was not Darwin's
+province to enter; "I have nothing to do," he said,[159] "with the
+origin of the mental powers, any more than I have with that of life
+itself." He dealt with the natural history of organisms, including not
+only their structure but their modes of behaviour; with the natural
+history of the states of consciousness which accompany some of their
+actions; and with the relation of behaviour to experience. We will
+endeavour to follow Darwin in his modesty and candour in making no
+pretence to give ultimate explanations. But we must note one of the
+implications of this self-denying ordinance of science. Development
+and evolution imply continuity. For Darwin and his followers the
+continuity is organic through physical heredity. Apart from
+speculative hypothesis, legitimate enough in its proper place but here
+out of court, we know nothing of continuity of mental evolution as
+such: consciousness appears afresh in each succeeding generation.
+Hence it is that for those who follow Darwin's lead, mental evolution
+is and must ever be, within his universe of discourse, subservient to
+organic evolution. Only in so far as conscious experience, or its
+neural correlate, effects some changes in organic structure can it
+influence the course of heredity; and conversely only in so far as
+changes in organic structure are transmitted through heredity, is
+mental evolution rendered possible. Such is the logical outcome of
+Darwin's teaching.
+
+Those who abide by the cardinal results of this teaching are bound to
+regard all behaviour as the expression of the functional activities of
+the living tissues of the organism, and all conscious experience as
+correlated with such activities. For the purposes of scientific
+treatment, mental processes are one mode of expression of the same
+changes of which the physiological processes accompanying behaviour
+are another mode of expression. This is simply accepted as a fact
+which others may seek to explain. The behaviour itself is the adaptive
+application of the energies of the organism; it is called forth by
+some form of presentation or stimulation brought to bear on the
+organism by the environment. This presentation is always an individual
+or personal matter. But in order that the organism may be fitted to
+respond to the presentation of the environment it must have undergone
+in some way a suitable preparation. According to the theory of
+evolution this preparation is primarily racial and is transmitted
+through heredity. Darwin's main thesis was that the method of
+preparation is predominantly by natural selection. Subordinate to
+racial preparation, and always dependent thereon, is individual or
+personal preparation through some kind of acquisition; of which the
+guidance of behaviour through individually won experience is a typical
+example. We here introduce the mental factor because the facts seem to
+justify the inference. Thus there are some modes of behaviour which
+are wholly and solely dependent upon inherited racial preparation;
+there are other modes of behaviour which are also dependent, in part
+at least, on individual preparation. In the former case the behaviour
+is adaptive on the first occurrence of the appropriate presentation;
+in the latter case accommodation to circumstances is only reached
+after a greater or less amount of acquired organic modification of
+structure, often accompanied (as we assume) in the higher animals by
+acquired experience. Logically and biologically the two classes of
+behaviour are clearly distinguishable: but the analysis of complex
+cases of behaviour where the two factors coöperate, is difficult and
+requires careful and critical study of life-history.
+
+The foundations of the mental life are laid in the conscious
+experience that accompanies those modes of behaviour, dependent
+entirely on racial preparation, which may broadly be described as
+instinctive. In the eighth chapter of _The Origin of Species_ Darwin
+says,[160] "I will not attempt any definition of instinct.... Every
+one understands what is meant, when it is said that instinct impels
+the cuckoo to migrate and to lay her eggs in other birds' nests. An
+action, which we ourselves require experience to enable us to perform,
+when performed by an animal, more especially by a very young one,
+without experience, and when performed by many individuals in the same
+way, without their knowing for what purpose it is performed, is
+usually said to be instinctive." And in the summary at the close of
+the chapter he says,[161] "I have endeavoured briefly to show that the
+mental qualities of our domestic animals vary, and that the variations
+are inherited. Still more briefly I have attempted to show that
+instincts vary slightly in a state of nature. No one will dispute that
+instincts are of the highest importance to each animal. Therefore
+there is no real difficulty, under changing conditions of life, in
+natural selection accumulating to any extent slight modifications of
+instinct which are in any way useful. In many cases habit or use and
+disuse have probably come into play."
+
+Into the details of Darwin's treatment there is neither space nor need
+to enter. There are some ambiguous passages; but it may be said that
+for him, as for his followers to-day, instinctive behaviour is wholly
+the result of racial preparation transmitted through organic heredity.
+For the performance of the instinctive act no individual preparation
+under the guidance of personal experience is necessary. It is true
+that Darwin quotes with approval Huber's saying that "a little dose of
+judgment or reason often comes into play, even with animals low in the
+scale of nature."[162] But we may fairly interpret his meaning to be
+that in behaviour, which is commonly called instinctive, some element
+of intelligent guidance is often combined. If this be conceded the
+strictly instinctive performance (or part of the performance) is the
+outcome of heredity and due to the direct transmission of parental or
+ancestral aptitudes. Hence the instinctive response as such depends
+entirely on how the nervous mechanism has been built up through
+heredity; while intelligent behaviour, or the intelligent factor in
+behaviour, depends also on how the nervous mechanism has been modified
+and moulded by use during its development and concurrently with the
+growth of individual experience in the customary situations of daily
+life. Of course it is essential to the Darwinian thesis that what Sir
+E. Ray Lankester has termed "educability," not less than instinct, is
+hereditary. But it is also essential to the understanding of this
+thesis that the differentiae of the hereditary factor should be
+clearly grasped.
+
+For Darwin there were two modes of racial preparation, (_1_) natural
+selection, and (_2_) the establishment of individually acquired habit.
+He showed that instincts are subject to hereditary variation; he saw
+that instincts are also subject to modification through acquisition in
+the course of individual life. He believed that not only the
+variations but also, to some extent, the modifications are inherited.
+He therefore held that some instincts (the greater number) are due to
+natural selection but that others (less numerous) are due, or partly
+due, to the inheritance of acquired habits. The latter involve
+Lamarckian inheritance, which of late years has been the centre of so
+much controversy. It is noteworthy however that Darwin laid especial
+emphasis on the fact that many of the most typical and also the most
+complex instincts--those of neuter insects--do not admit of such an
+interpretation. "I am surprised," he says,[163] "that no one has
+hitherto advanced this demonstrative case of neuter insects, against
+the well-known doctrine of inherited habit, as advanced by Lamarck."
+None the less Darwin admitted this doctrine as supplementary to that
+which was more distinctively his own--for example in the case of the
+instincts of domesticated animals. Still, even in such cases, "it may
+be doubted," he says,[164] "whether any one would have thought of
+training a dog to point, had not some one dog naturally shown a
+tendency in this line ... so that habit and some degree of selection
+have probably concurred in civilising by inheritance our dogs." But in
+the interpretation of the instincts of domesticated animals, a more
+recently suggested hypothesis, that of organic selection,[165] may be
+helpful. According to this hypothesis any intelligent modification of
+behaviour which is subject to selection is probably coincident in
+direction with an inherited tendency to behave in this fashion. Hence
+in such behaviour there are two factors: (1) an incipient variation in
+the line of such behaviour, and (2) an acquired modification by which
+the behaviour is carried further along the same line. Under natural
+selection those organisms in which the two factors coöperate are
+likely to survive. Under artificial selection they are deliberately
+chosen out from among the rest.
+
+Organic selection has been termed a compromise between the more
+strictly Darwinian and the Lamarckian principles of interpretation.
+But it is not in any sense a compromise. The principle of
+interpretation of that which is instinctive and hereditary is wholly
+Darwinian. It is true that some of the facts of observation relied
+upon by Lamarckians are introduced. For Lamarckians however the
+modifications which are admittedly factors in survival, are regarded
+as the parents of inherited variations; for believers in organic
+selection they are only the foster-parents or nurses. It is because
+organic selection is the direct outcome of and a natural extension of
+Darwin's cardinal thesis that some reference to it here is
+justifiable. The matter may be put with the utmost brevity as follows:
+(1) Variations (V) occur, some of which are in the direction of
+increased adaptation (+), others in the direction of decreased
+adaptation (-).
+
+(2) Acquired modifications (M) also occur. Some of these are in the
+direction of increased accommodation to circumstances (+), while
+others are in the direction of diminished accommodation (-). Four
+major combinations are
+
+   (_b_) + V with - M,      (_c_) - V with + M,
+
+   (_a_) + V with + M,      (_d_) - V with - M.
+
+Of these (_d_) must inevitably be eliminated while (_a_) are selected.
+The predominant survival of (_a_) entails the survival of the adaptive
+variations which are inherited. The contributory acquisitions (+ M)
+are not inherited; but there are none the less factors in determining
+the survival of the coincident variations. It is surely abundantly
+clear that this is Darwinism and has no tincture of Lamarck's
+essential principle, the inheritance of acquired characters.
+
+Whether Darwin himself would have accepted this interpretation of some
+at least of the evidence put forward by Lamarckians is unfortunately a
+matter of conjecture. The fact remains that in his interpretation of
+instinct and in allied questions he accepted the inheritance of
+individually acquired modifications of behaviour and structure.
+
+Darwin was chiefly concerned with instinct from the biological rather
+than from the psychological point of view. Indeed it must be confessed
+that, from the latter standpoint, his conception of instinct as a
+"mental faculty" which "impels" an animal to the performance of
+certain actions, scarcely affords a satisfactory basis for genetic
+treatment. To carry out the spirit of Darwin's teaching it is
+necessary to link more closely biological and psychological evolution.
+The first step towards this is to interpret the phenomena of
+instinctive behaviour in terms of stimulation and response. It may be
+well to take a particular case. Swimming on the part of a duckling is,
+from the biological point of view, a typical example of instinctive
+behaviour. Gently lower a recently hatched bird into water:
+coordinated movements of the limbs follow in rhythmical sequence. The
+behaviour is new to the individual though it is no doubt closely
+related to that of walking, which is no less instinctive. There is a
+group of stimuli afforded by the "presentation" which results from
+partial immersion: upon this there follows as a complex response an
+application of the functional activities in swimming; the sequence of
+adaptive application on the appropriate presentation is determined by
+racial preparation. We know, it is true, but little of the
+physiological details of what takes place in the central nervous
+system; but in broad outline the nature of the organic mechanism and
+the manner of its functioning may at least be provisionally
+conjectured in the present state of physiological knowledge. Similarly
+in the case of the pecking of newly-hatched chicks; there is a visual
+presentation, there is probably a coöperating group of stimuli from
+the alimentary tract in need of food, there is an adaptive application
+of the activities in a definite mode of behaviour. Like data are
+afforded in a great number of cases of instinctive procedure,
+sometimes occurring very early in life, not infrequently deferred
+until the organism is more fully developed, but all of them dependent
+upon racial preparation. No doubt there is some range of variation in
+the behaviour, just such variation as the theory of natural selection
+demands. But there can be no question that the higher animals inherit
+a bodily organisation and a nervous system, the functional working of
+which gives rise to those inherited modes of behaviour which are
+termed instinctive.
+
+It is to be noted that the term "instinctive" is here employed in the
+adjectival form as a descriptive heading under which may be grouped
+many and various modes of behaviour due to racial preparation. We
+speak of these as inherited; but in strictness what is transmitted
+through heredity is the complex of anatomical and physiological
+conditions under which, in appropriate circumstances, the organism so
+behaves. So far the term "instinctive" has a restricted biological
+connotation in terms of behaviour. But the connecting link between
+biological evolution and psychological evolution is to be sought,--as
+Darwin fully realised,--in the phenomena of instinct, broadly
+considered. The term "instinctive" has also a psychological
+connotation. What is that connotation?
+
+Let us take the case of the swimming duckling or the pecking chick,
+and fix our attention on the first instinctive performance. Grant that
+just as there is, strictly speaking, no inherited behaviour, but only
+the conditions which render such behaviour under appropriate
+circumstances possible; so too there is no inherited experience, but
+only the conditions which render such experience possible; then the
+cerebral conditions in both cases are the same. The biological
+behaviour-complex, including the total stimulation and the total
+response with the intervening or resultant processes in the sensorium,
+is accompanied by an experience-complex including the initial
+stimulation-consciousness and resulting response-consciousness. In the
+experience-complex are comprised data which in psychological analysis
+are grouped under the headings of cognition, affective tone and
+conation. But the complex is probably experienced as an unanalysed
+whole. If then we use the term "instinctive" so as to comprise all
+congenital modes of behaviour which contribute to experience, we are
+in a position to grasp the view that the net result in consciousness
+constitutes what we may term the primary tissue of experience. To the
+development of this experience each instinctive act contributes. The
+nature and manner of organisation of this primary tissue of experience
+are dependent on inherited biological aptitudes; but they are from the
+outset onwards subject to secondary development dependent on acquired
+aptitudes. Biological values are supplemented by psychological values
+in terms of satisfaction or the reverse.
+
+In our study of instinct we have to select some particular phase of
+animal behaviour and isolate it so far as is possible from the life of
+which it is a part. But the animal is a going concern, restlessly
+active in many ways. Many instinctive performances, as Darwin pointed
+out,[166] are serial in their nature. But the whole of active life is
+a serial and coordinated business. The particular instinctive
+performance is only an episode in a life-history, and every mode of
+behaviour is more or less closely correlated with other modes. This
+coordination of behaviour is accompanied by a correlation of the modes
+of primary experience. We may classify the instinctive modes of
+behaviour and their accompanying modes of instinctive experience under
+as many heads as may be convenient for our purposes of interpretation,
+and label them instincts of self-preservation, of pugnacity, of
+acquisition, the reproductive instincts, the parental instincts, and
+so forth. An instinct, in this sense of the term (for example the
+parental instinct), may be described as a specialised part of the
+primary tissue of experience differentiated in relation to some
+definite biological end. Under such an instinct will fall a large
+number of particular and often well-defined modes of behaviour, each
+with its own peculiar mode of experience.
+
+It is no doubt exceedingly difficult as a matter of observation and of
+inference securely based thereon to distinguish what is primary from
+what is in part due to secondary acquisition--a fact which Darwin
+fully appreciated. Animals are educable in different degrees; but
+where they are educable they begin to profit by experience from the
+first. Only, therefore, on the occasion of the first instinctive act
+of a given type can the experience gained be regarded as _wholly_
+primary; all subsequent performance is liable to be in some degree,
+sometimes more, sometimes less, modified by the acquired disposition
+which the initial behaviour engenders. But the early stages of
+acquisition are always along the lines predetermined by instinctive
+differentiation. It is the task of comparative psychology to
+distinguish the primary tissue of experience from its secondary and
+acquired modifications. We cannot follow up the matter in further
+detail. It must here suffice to suggest that this conception of
+instinct as a primary form of experience lends itself better to
+natural history treatment than Darwin's conception of an impelling
+force, and that it is in line with the main trend of Darwin's thought.
+
+In a characteristic work,--characteristic in wealth of detail, in
+closeness and fidelity of observation, in breadth of outlook, in
+candour and modesty,--Darwin dealt with _The Expression of the
+Emotions in Man and Animals_. Sir Charles Bell in his _Anatomy of
+Expression_ had contended that many of man's facial muscles had been
+specially created for the sole purpose of being instrumental in the
+expression of his emotions. Darwin claimed that a natural explanation,
+consistent with the doctrine of evolution, could in many cases be
+given and would in other cases be afforded by an extension of the
+principles he advocated. "No doubt," he said,[167] "as long as man and
+all other animals are viewed as independent creations, an effectual
+stop is put to our natural desire to investigate as far as possible
+the causes of Expression. By this doctrine, anything and everything
+can be equally well explained.... With mankind, some expressions ...
+can hardly be understood, except on the belief that man once existed
+in a much lower and animal-like condition. The community of certain
+expressions in distinct though allied species ... is rendered somewhat
+more intelligible, if we believe in their descent from a common
+progenitor. He who admits on general grounds that the structure and
+habits of all animals have been gradually evolved, will look at the
+whole subject of Expression in a new and interesting light."
+
+Darwin relied on three principles of explanation. "The first of these
+principles is, that movements which are serviceable in gratifying some
+desire, or in relieving some sensation, if often repeated, become so
+habitual that they are performed, whether or not of any service,
+whenever the same desire or sensation is felt, even in a very weak
+degree."[168] The modes of expression which fall under this head have
+become instinctive through the hereditary transmission of acquired
+habit. "As far as we can judge, only a few expressive movements are
+learnt by each individual; that is, were consciously and voluntarily
+performed during the early years of life for some definite object, or
+in imitation of others, and then became habitual. The far greater
+number of the movements of expression, and all the more important
+ones, are innate or inherited; and such cannot be said to depend on
+the will of the individual. Nevertheless, all those included under our
+first principle were at first voluntarily performed for a definite
+object,--namely, to escape some danger, to relieve some distress, or
+to gratify some desire."[169]
+
+"Our second principle is that of antithesis. The habit of voluntarily
+performing opposite movements under opposite impulses has become
+firmly established in us by the practice of our whole lives. Hence, if
+certain actions have been regularly performed, in accordance with our
+first principle, under a certain frame of mind, there will be a strong
+and involuntary tendency to the performance of directly opposite
+actions, whether or not these are of any use, under the excitement of
+an opposite frame of mind."[170] This principle of antithesis has not
+been widely accepted. Nor is Darwin's own position easy to grasp.
+
+"Our third principle," he says,[171] "is the direct action of the
+excited nervous system on the body, independently of the will, and
+independently, in large part, of habit. Experience shows that
+nerve-force is generated and set free whenever the cerebro-spinal
+system is excited. The direction which this nerve-force follows is
+necessarily determined by the lines of connection between the
+nerve-cells, with each other and with various parts of the body."
+
+Lack of space prevents our following up the details of Darwin's
+treatment of expression. Whether we accept or do not accept his three
+principles of explanation we must regard his work as a masterpiece of
+descriptive analysis, packed full of observations possessing lasting
+value. For a further development of the subject it is essential that
+the instinctive factors in expression should be more fully
+distinguished from those which are individually acquired--a difficult
+task--and that the instinctive factors should be rediscussed in the
+light of modern doctrines of heredity, with a view to determining
+whether Lamarckian inheritance, on which Darwin so largely relied, is
+necessary for an interpretation of the facts.
+
+The whole subject as Darwin realised is very complex. Even the term
+"expression" has a certain amount of ambiguity. When the emotion is in
+full flood, the animal fights, flees, or faints. Is this full-tide
+effect to be regarded as expression; or are we to restrict the term to
+the premonitory or residual effects--the bared canine when the
+fighting mood is being roused, the ruffled fur when reminiscent
+representations of the object inducing anger cross the mind? Broadly
+considered both should be included. The activity of premonitory
+expression as a means of communication was recognised by Darwin; he
+might, perhaps, have emphasised it more strongly in dealing with the
+lower animals. Man so largely relies on a special means of
+communication, that of language, that he sometimes fails to realise
+that for animals with their keen powers of perception, and dependent
+as they are on such means of communication, the more strictly
+biological means of expression are full of subtle suggestiveness. Many
+modes of expression, otherwise useless, are signs of behaviour that
+may be anticipated,--signs which stimulate the appropriate attitude of
+response. This would not, however, serve to account for the utility of
+the organic accompaniments--heart-affection, respiratory changes,
+vaso-motor effects and so forth, together with heightened muscular
+tone,--on all of which Darwin lays stress[172] under his third
+principle. The biological value of all this is, however, of great
+importance, though Darwin was hardly in a position to take it fully
+into account.
+
+Having regard to the instinctive and hereditary factors of emotional
+expression we may ask whether Darwin's third principle does not alone
+suffice as an explanation. Whether we admit or reject Lamarckian
+inheritance it would appear that all hereditary expression must be due
+to pre-established connections within the central nervous system and
+to a transmitted provision for coordinated response under the
+appropriate stimulation. If this be so, Darwin's first and second
+principles are subordinate and ancillary to the third, an expression,
+so far as it is instinctive or heredity, being "the direct result of
+the constitution of the nervous system."
+
+Darwin accepted the emotions themselves as hereditary or acquired
+states of mind and devoted his attention to their expression. But
+these emotions themselves are genetic products and as such dependent
+on organic conditions. It remained, therefore, for psychologists who
+accepted evolution and sought to build on biological foundations to
+trace the genesis of these modes of animal and human experience. The
+subject has been independently developed by Professors Lange and
+James;[173] and some modification of their view is regarded by many
+evolutionists as affording the best explanation of the facts. We must
+fix our attention on the lower emotions, such as anger or fear, and on
+their first occurrence in the life of the individual organism. It is a
+matter of observation that if a group of young birds which have been
+hatched in an incubator are frightened by an appropriate presentation,
+auditory or visual, they instinctively respond in special ways. If we
+speak of this response as the expression, we find that there are many
+factors. There are certain visible modes of behaviour, crouching at
+once, scattering and then crouching, remaining motionless, the braced
+muscles sustaining an attitude of arrest, and so forth, There are also
+certain visceral or organic effects, such as affections of the heart
+and respiration. These can be readily observed by taking the young
+bird in the hand. Other effects cannot be readily observed; vaso-motor
+changes, affections of the alimentary canal, the skin and so forth.
+Now the essence of the James-Lange view, as applied to these
+congenital effects, is that though we are justified in speaking of
+them as effects of the stimulation, we are not justified, without
+further evidence, in speaking of them as effects of the emotional
+state. May it not rather be that the emotion as a primary mode of
+experience is the concomitant of the net result of the organic
+situation--the initial presentation, the instinctive mode of
+behaviour, the visceral disturbances?
+
+According to this interpretation the primary tissue of experience of
+the emotional order, felt as an unanalysed complex, is generated by
+the stimulation of the sensorium by afferent or incoming physiological
+impulses from the special senses, from the organs concerned in the
+responsive behaviour, from the viscere and vaso-motor system.
+
+Some psychologists, however, contend that the emotional experience is
+generated in the sensorium prior to, and not subsequent to, the
+behaviour-response and the visceral disturbances. It is a direct and
+not an indirect outcome of the presentation to the special senses. Be
+this as it may, there is a growing tendency to bring into the closest
+possible relation, or even to identify, instinct and emotion in their
+primary genesis. The central core of all such interpretations is that
+instinctive behaviour and experience, its emotional accompaniments,
+and its expression, are but different aspects of the outcome of the
+same organic occurrences. Such emotions are, therefore, only a
+distinguishable aspect of the primary tissue of experience and exhibit
+a like differentiation. Here again a biological foundation is laid for
+a psychological doctrine of the mental development of the individual.
+
+The intimate relation between emotion as a psychological mode of
+experience and expression as a group of organic conditions has an
+important bearing on biological interpretation. The emotion, as the
+psychological accompaniment of orderly disturbances in the central
+nervous system, profoundly influences behaviour and often renders it
+more vigourous and more effective. The utility of the emotions in the
+struggle for existence can, therefore, scarcely be over-estimated.
+Just as keenness of perception has survival-value; just as it is
+obviously subject to variation; just as it must be enhanced under
+natural selection, whether individually acquired increments are
+inherited or not; and just as its value lies not only in this or that
+special perceptive act but in its importance for life as a whole; so
+the vigourous effectiveness of activity has survival-value; it is
+subject to variation; it must be enhanced under natural selection; and
+its importance lies not only in particular modes of behaviour but in
+its value for life as a whole. If emotion and its expression as a
+congenital endowment are but different aspects of the same biological
+occurrence; and if this is a powerful supplement to vigour
+effectiveness and persistency of behaviour, it must on Darwin's
+principles be subject to natural selection.
+
+If we include under the expression of the emotions not only the
+premonitory symptoms of the initial phases of the organic and mental
+state, not only the signs or conditions of half-tide emotion, but the
+full-tide manifestation of an emotion which dominates the situation,
+we are naturally led on to the consideration of many of the phenomena
+which are discussed under the head of sexual selection. The subject is
+difficult and complex, and it was treated by Darwin with all the
+strength he could summon to the task. It can only be dealt with here
+from a special point of view--that which may serve to illustrate the
+influence of certain mental factors on the course of evolution. From
+this point of view too much stress can scarcely be laid on the
+dominance of emotion during the period of courtship and pairing in the
+more highly organised animals. It is a period of maximum vigour,
+maximum activity, and, correlated with special modes of behaviour and
+special organic and visceral accompaniments, a period also of maximum
+emotional excitement. The combats of males, their dances and aerial
+evolutions, their elaborate behaviour and display, or the flood of
+song in birds, are emotional expressions which are at any rate
+coincident in time with sexual periodicity. From the combat of the
+males there follows on Darwin's principles the elimination of those
+which are deficient in bodily vigour, deficient in special structures,
+offensive or protective, which contribute to success, deficient in the
+emotional supplement of which persistent and whole-hearted fighting is
+the expression, and deficient in alertness and skill which are the
+outcome of the psychological development of the powers of perception.
+Few biologists question that we have here a mode of selection of much
+importance, though its influence on psychological evolution often
+fails to receive its due emphasis. Mr. Wallace[174] regards it as "a
+form of natural selection"; "to it," he says, "we must impute the
+development of the exceptional strength, size, and activity of the
+male, together with the possession of special offensive and defensive
+weapons, and of all other characters which arise from the development
+of these or are correlated with them." So far there is little
+disagreement among the followers of Darwin--for Mr. Wallace, with fine
+magnanimity, has always preferred to be ranked as such,
+notwithstanding his right, on which a smaller man would have
+constantly insisted, to the claim of independent originator of the
+doctrine of natural selection. So far with regard to sexual selection
+Darwin and Mr. Wallace are agreed; so far and no farther. For Darwin,
+says Mr. Wallace,[175] "has extended the principle into a totally
+different field of action, which has none of that character of
+constancy and of inevitable result that attaches to natural selection,
+including male rivalry; for by far the larger portion of the
+phenomena, which he endeavours to explain by the direct action of
+sexual selection, can only be so explained on the hypothesis that the
+immediate agency is female choice or preference. It is to this that he
+imputes the origin of all secondary sexual characters other than
+weapons of offence and defence.... In this extension of sexual
+selection to include the action of female choice or preference, and in
+the attempt to give to that choice such wide-reaching effects, I am
+unable to follow him more than a very little way."
+
+Into the details of Mr. Wallace's criticisms it is impossible to enter
+here. We cannot discuss either the mode of origin of the variations in
+structure which have rendered secondary sexual characters possible or
+the modes of selection other than sexual which have rendered them,
+within narrow limits, specifically constant. Mendelism and mutation
+theories may have something to say on the subject when these theories
+have been more fully correlated with the basal principles of
+selection. It is noteworthy that Mr. Wallace says:[176] "Besides the
+acquisition of weapons by the male for the purpose of fighting with
+other males, there are some other sexual characters which may have
+been produced by natural selection. Such are the various sounds and
+odours which are peculiar to the male, and which serve as a call to
+the female or as an indication of his presence. These are evidently a
+valuable addition to the means of recognition of the two sexes, and
+are a further indication, that the pairing season has arrived; and the
+production, intensification, and differentiation of these sounds and
+odours are clearly within the power of natural selection. The same
+remark will apply to the peculiar calls of birds, and even to the
+singing of the males." Why the same remark should not apply to their
+colours and adornments is not obvious. What is obvious is that "means
+of recognition" and "indication that the pairing season has arrived"
+are dependent on the perceptive powers of the female who recognises
+and for whom the indication has meaning. The hypothesis of female
+preference, stripped of the aesthetic surplusage which is
+psychologically both unnecessary and unproven, is really only
+different in degree from that which Mr. Wallace admits in principle
+when he says that it is probable that the female is pleased or excited
+by the display.
+
+Let us for our present purpose leave on one side and regard as _sub
+judice_ the question whether the specific details of secondary sexual
+characters are the outcome of female choice. For us the question is
+whether certain psychological accompaniments of the pairing situation
+have influenced the course of evolution and whether these
+psychological accompaniments are themselves the outcome of evolution.
+As a matter of observation, specially differentiated modes of
+behaviour, often very elaborate, frequently requiring highly developed
+skill, and apparently highly charged with emotional tone, are the
+precursors of pairing. They are generally confined to the males, whose
+fierce combats during the period of sexual activity are part of the
+emotional manifestation. It is inconceivable that they have no
+biological meaning; and it is difficult to conceive that they have any
+other biological end than to evoke in the generally more passive
+female the pairing impulse. They, are based on instinctive foundations
+ingrained in the nervous constitution through natural (or may we not
+say sexual?) selection in virtue of their profound utility. They are
+called into play by a specialised presentation such as the sight or
+the scent of the female at, or a little in advance of, a critical
+period of the physiological rhythm. There is no necessity that the
+male should have any knowledge of the end to which his strenuous
+activity leads up. In presence of the female there is an elaborate
+application of all the energies of behaviour, just because ages of
+racial preparation have made him biologically and emotionally what he
+is--a functionally sexual male that must dance or sing or go through
+hereditary movements of display, when the appropriate stimulation
+comes. Of course after the first successful courtship his future
+behaviour will be in some degree modified by his previous experience.
+No doubt during his first courtship he is gaining the primary data of
+a peculiarly rich experience, instinctive and emotional. But the
+biological foundations of the behaviour of courtship are laid in the
+hereditary coordinations. It would seem that in some cases, not indeed
+in all, perhaps especially in those cases in which secondary sexual
+behaviour is most highly evolved,--correlative with the ardour of the
+male is a certain amount of reluctance in the female. The pairing act
+on her part only takes place after prolonged stimulation, for
+affording which the behaviour of male courtship is the requisite
+presentation. The most vigourous, defiant and mettlesome male is
+preferred just because he alone affords a contributory stimulation
+adequate to evoke the pairing impulse with its attendant emotional
+tone.
+
+It is true that this places female preference or choice on a much
+lower psychological plane than Darwin in some passages seems to
+contemplate where, for example, he says that the female appreciates
+the display of the male and places to her credit a taste for the
+beautiful. But Darwin himself distinctly states[177] that "it is not
+probable that she consciously deliberates; but she is most excited or
+attracted by the most beautiful, or melodious, or gallant males." The
+view here put forward, which has been developed by Prof. Groos,[178]
+therefore seems to have Darwin's own sanction. The phenomena are not
+only biological; there are psychological elements as well. One can
+hardly suppose that the female is unconscious of the male's presence;
+the final yielding must surely be accompanied by heightened emotional
+tone. Whether we call it choice or not is merely a matter of
+definition of terms. The behaviour is in part determined by
+supplementary psychological values. Prof. Groos regards the coyness of
+females as "a most efficient means of preventing the too early and too
+frequent yielding to the sexual impulse."[179] Be that as it may, it
+is, in any case, if we grant the facts, a means through which male
+sexual behaviour with all its biological and psychological
+implications, is raised to a level otherwise perhaps unattainable by
+natural means, while in the female it affords opportunities for the
+development in the individual and evolution in the race of what we may
+follow Darwin in calling appreciation, if we empty this word of the
+aesthetic implications which have gathered round it in the mental life
+of man.
+
+Regarded from this standpoint of sexual selection, broadly considered,
+has probably been of great importance. The psychological
+accompaniments of the pairing situation have profoundly influenced the
+course of biological evolution and are themselves the outcome of that
+evolution.
+
+Darwin makes only passing reference to those modes of behaviour in
+animals which go by the name of play. "Nothing," he says,[180] "is
+more common than for animals to take pleasure in practising whatever
+instinct they follow at other times for some real good." This is one
+of the very numerous cases in which a hint of the master has served to
+stimulate research in his disciples. It was left to Prof. Groos to
+develop this subject on evolutionary lines and to elaborate in a
+masterly manner Darwin's suggestion. "The utility of play," he
+says,[181] "is incalculable. This utility consists in the practice and
+exercise it affords for some of the more important duties of
+life,"--that is to say, for the performance of activities which will
+in adult life be essential to survival. He urges[182] that "the play
+of young animals has its origin in the fact that certain very
+important instincts appear at a time when the animal does not
+seriously need them." It is, however, questionable whether any
+instincts appear at a time when they are not needed. And it is
+questionable whether the instinctive and emotional attitude of the
+play-fight, to take one example, can be identified with those which
+accompany fighting in earnest, though no doubt they are closely
+related and have some common factors. It is probable that play, as
+preparatory behaviour, differs in biological detail (as it almost
+certainly does in emotional attributes) from the earnest of after-life
+and that it has been evolved through differentiation and integration
+of the primary tissue of experience, as a preparation through which
+certain essential modes of skill may be acquired--those animals in
+which the preparatory play-propensity was not inherited in due force
+and requisite amount being subsequently eliminated in the struggle for
+existence. In any case there is little question that Prof. Groos is
+right in basing the play-propensity on instinctive foundations.[183]
+None the less, as he contends, the essential biological value of play
+is that it is a means of training the educable nerve-tissue, of
+developing that part of the brain which is modified by experience and
+which thus acquires new characters, of elaborating the secondary
+tissue of experience on the predetermined lines of instinctive
+differentiation and thus furthering the psychological activities which
+are included under the comprehensive term "intelligent."
+
+In _The Descent of Man_ Darwin dealt at some length with intelligence
+and the higher mental faculties.[184] His object, he says, is to show
+that there is no fundamental difference between man and the higher
+mammals in their mental faculties; that these faculties are variable
+and the variations tend to be inherited; and that under natural
+selection beneficial variations of all kinds will have been preserved
+and injurious ones eliminated.
+
+Darwin was too good an observer and too honest a man to minimise the
+"enormous difference" between the level of mental attainment of
+civilised man and that reached by any animal. His contention was that
+the difference, great as it is, is one of degree and not of kind. He
+realised that, in the development of the mental faculties of man, new
+factors in evolution have supervened--factors which play but a
+subordinate and subsidiary part in animal intelligence.
+Intercommunication by means of language, approbation and blame, and
+all that arises out of reflective thought, are but foreshadowed in the
+mental life of animals. Still he contends that these may be explained
+on the doctrine of evolution. He urges[185] "that man is variable in
+body and mind; and that the variations are induced, either directly or
+indirectly, by the same general causes, and obey the same general
+laws, as with the lower animals." He correlates mental development
+with the evolution of the brain.[186] "As the various mental faculties
+gradually developed themselves, the brain would almost certainly
+become larger. No one, I presume, doubts that the large proportion
+which the size of man's brain bears to his body, compared to the same
+proportion in the gorilla or orang, is closely connected with his
+higher mental powers." "With respect to the lower animals," he
+says,[187] "M. E. Lartet,[188] by comparing the crania of tertiary and
+recent mammals belonging to the same groups, has come to the
+remarkable conclusion that the brain is generally larger and the
+convolutions are more complex in the more recent form."
+
+Sir E. Ray Lankester has sought to express in the simplest terms the
+implications of the increase in size of the cerebrum. "In what," he
+asks, "does the advantage of a larger cerebral mass consist?" "Man,"
+he replies, "is born with fewer ready-made tricks of the
+nerve-centres--these performances of an inherited nervous mechanism so
+often called by the ill-defined term 'instincts'--than are the monkeys
+or any other animal. Correlated with the absence of inherited
+ready-made mechanism, man has a greater capacity of developing in the
+course of his individual growth similar nervous mechanisms (similar
+to but not identical with those of 'instinct') than any other
+animal.... The power of being educated--'educability' as we may term
+it--is what man possesses in excess as compared with the apes. I think
+we are justified in forming the hypothesis that it is this
+'educability' which is the correlative of the increased size of the
+cerebrum." There has been natural selection of the more educable
+animals, for "the character which we describe as 'educability' can be
+transmitted, it is a congenital character. But the _results_ of
+education can _not_ be transmitted. In each generation they have to be
+acquired afresh, and with increased 'educability' they are more
+readily acquired and a larger variety of them.... The fact is that
+there is no community between the mechanisms of instinct and the
+mechanisms of intelligence, and that the latter are later in the
+history of the evolution of the brain than the former and can only
+develop in proportion as the former become feeble and defective."[189]
+
+In this statement we have a good example of the further development of
+views which Darwin foreshadowed but did not thoroughly work out. It
+states the biological case clearly and tersely. Plasticity of
+behaviour in special accommodation to special circumstances is of
+survival value; it depends upon acquired characters; it is correlated
+with increase in size and complexity of the cerebrum; under natural
+selection therefore the larger and more complex cerebrum as the organ
+of plastic behaviour has been the outcome of natural selection. We
+have thus the biological foundations for a further development of
+genetic psychology.
+
+There are diversities of opinion, as Darwin showed, with regard to the
+range of instinct in man and the higher animals as contrasted with
+lower types. Darwin himself said[190] that "Man, perhaps, has somewhat
+fewer instincts than those possessed by the animals which come next to
+him in the series." On the other hand, Prof. Wm. James says[191] that
+man is probably the animal with most instincts. The true position is
+that man and the higher animals have fewer complete and self-sufficing
+instincts than those which stand lower in the scale of mental
+evolution, but that they have an equally large or perhaps larger mass
+of instinctive raw material which may furnish the stuff to be
+elaborated by intelligent processes. There is, perhaps, a greater
+abundance of the primary tissue of experience to be refashioned and
+integrated by secondary modification; there is probably the same
+differentiation in relation to the determining biological ends, but
+there is at the outset less differentiation of the particular and
+specific modes of behaviour. The specialised instinctive performances
+and their concomitant experience-complexes are at the outset more
+indefinite. Only through acquired connections, correlated with
+experience, do they become definitely organised.
+
+The full working-out of the delicate and subtle relationship of
+instinct and educability--that is, of the hereditary and the acquired
+factors in the mental life--is the task which lies before genetic and
+comparative psychology. They interact throughout the whole of life,
+and their interactions are very complex. No one can read the chapters
+of _The Descent of Man_ which Darwin devotes to a consideration of the
+mental characters of man and animals without noticing, on the one
+hand, how sedulous he is in his search for hereditary foundations,
+and, on the other hand, how fully he realises the importance of
+acquired habits of mind. The fact that educability itself has innate
+tendencies--is in fact a partially differentiated educability--renders
+the unravelling of the factors of mental progress all the more
+difficult.
+
+In his comparison of the mental powers of men and animals it was
+essential that Darwin should lay stress on points of similarity rather
+than on points of difference. Seeking to establish a doctrine of
+evolution, with its basal concept of continuity of process and
+community of character, he was bound to render clear and to emphasise
+the contention that the difference in mind between man and the higher
+animals, great as it is, is one of degree and not of kind. To this end
+Darwin not only recorded a large number of valuable observations of
+his own, and collected a considerable body of information from
+reliable sources, he presented the whole subject in a new light and
+showed that a natural history of mind might be written and that this
+method of study offered a wide and rich field for investigation. Of
+course those who regarded the study of mind only as a branch of
+metaphysics smiled at the philosophical ineptitude of the mere man of
+science. But the investigation, on natural history lines, has been
+prosecuted with a large measure of success. Much indeed still remains
+to be done; for special training is required, and the workers are
+still few. Promise for the future is however afforded by the fact that
+investigation is prosecuted on experimental lines and that something
+like organised methods of research are taking form. There is now but
+little reliance on casual observations recorded by those who have not
+undergone the necessary discipline in these methods. There is also
+some change of emphasis in formulating conclusions. Now that the
+general evolutionary thesis is fully and freely accepted by those who
+carry on such researches, more stress is laid on the differentiation
+of the stages of evolutionary advance than on the fact of their
+underlying community of nature. The conceptual intelligence which is
+especially characteristic of the higher mental procedure of man is
+more firmly distinguished from the perceptual intelligence which he
+shares with the lower animals--distinguished now as a higher product
+of evolution, no longer as differing in origin or different in kind.
+Some progress has been made, on the one hand in rendering an account
+of intelligent profiting by experience under the guidance of pleasure
+and pain in the perceptual field, on lines predetermined by
+instinctive differentiation for biological ends, and on the other hand
+in elucidating the method of conceptual thought employed, for
+example, by the investigator himself in interpreting the perceptual
+experience of the lower animals.
+
+Thus there is a growing tendency to realise more fully that there are
+two orders of educability--first an educability of the perceptual
+intelligence based on the biological foundation of instinct, and
+secondly an educability of the conceptual intelligence which
+refashions and rearranges the data afforded by previous inheritance
+and acquisition. It is in relation to this second and higher order of
+educability that the cerebrum of man shows so large an increase of
+mass and a yet larger increase of effective surface through its rich
+convolutions. It is through educability of this order that the human
+child is brought intellectually and affectively into touch with the
+ideal constructions by means of which man has endeavoured, with more
+or less success, to reach an interpretation of nature, and to guide
+the course of the further evolution of his race--ideal constructions
+which form part of man's environment.
+
+It formed no part of Darwin's purpose to consider, save in broad
+outline, the methods, or to discuss in any fulness of detail the
+results of the process by which a differentiation of the mental
+faculties of man from those of the lower animals has been brought
+about--a differentiation the existence of which he again and again
+acknowledges. His purpose was rather to show that, notwithstanding
+this differentiation, there is basal community in kind. This must be
+remembered in considering his treatment of the biological foundations
+on which man's systems of ethics are built. He definitely stated that
+he approached the subject "exclusively from the side of natural
+history."[192] His general conclusion is that the moral sense is
+fundamentally identical with the social instincts, which have been
+developed for the good of the community; and he suggests that the
+concept which thus enables us to interpret the biological ground-plan
+of morals also enables us to frame a rational ideal of the moral end.
+"As the social instincts," he says,[193] "both of man and the lower
+animals have no doubt been developed by nearly the same steps, it
+would be advisable, if found practicable, to use the same definition
+in both cases, and to take as the standard of morality, the general
+good or welfare of the community, rather than the general happiness."
+But the kind of community for the good of which the social instincts
+of animals and primitive men were biologically developed may be
+different from that which is the product of civilisation, as Darwin no
+doubt realised. Darwin's contention was that conscience is a social
+instinct and has been evolved because it is useful to the tribe in the
+struggle for existence against other tribes. One the other hand J. S.
+Mill urged that the moral feelings are not innate but acquired, and
+Bain held the same view, believing that the moral sense is acquired by
+each individual during his life-time. Darwin, who notes[194] their
+opinion with his usual candour, adds that "on the general theory of
+evolution this is at least extremely improbable." It is impossible to
+enter into the question here: much turns on the exact connotation of
+the terms "conscience" and "moral sense," and on the meaning we attach
+to the statement that the moral sense is fundamentally identical with
+the social instincts.
+
+Presumably the majority of those who approach the subjects discussed
+in the third, fourth, and fifth chapters of _The Descent of Man_ in
+the full conviction that mental phenomena, not less than organic
+phenomena, have a natural genesis, would, without hesitation, admit
+that the intellectual and moral systems of civilised man are ideal
+constructions, the products of conceptual thought, and that as such
+they are, in their developed form, acquired. The moral sentiments are
+the emotional analogues of highly developed concepts. This does not
+however imply that they are outside the range of natural history
+treatment. Even though it may be desirable to differentiate the moral
+conduct of men from the social behaviour of animals (to which some
+such term as "pre-moral" or "quasi-moral" may be applied), still the
+fact remains that, as Darwin showed, there is abundant evidence of the
+occurrence of such social behaviour--social behaviour which, even
+granted that it is in large part intelligently acquired, and is itself
+so far a product of educability, is of survival value. It makes for
+that integration without which no social group could hold together and
+escape elimination. Furthermore, even if we grant that such behaviour
+is intelligently acquired, that is to say arises through the
+modification of hereditary instincts and emotions, the fact remains
+that only through these instinctive and emotional data is afforded the
+primary tissue of the experience which is susceptible of such
+modification.
+
+Darwin sought to show, and succeeded in showing, that for the
+intellectual and moral life there are instinctive foundations which a
+biological treatment alone can disclose. It is true that he did not in
+all cases analytically distinguish the foundations from the
+superstructure. Even to-day we are scarcely in a position to do so
+adequately. But his treatment was of great value in giving an impetus
+to further research. This value indeed can scarcely be over-estimated.
+And when the natural history of the mental operations shall have been
+written, the cardinal fact will stand forth, that the instinctive and
+emotional foundations are the outcome of biological evolution and have
+been ingrained in the race through natural selection. We shall more
+clearly realise that educability itself is a product of natural
+selection, though the specific results acquired through cerebral
+modifications are not transmitted through heredity. It will, perhaps,
+also be realised that the instinctive foundations of social behaviour
+are, for us, somewhat out of date and have undergone but little change
+throughout the progress of civilisation, because natural selection has
+long since ceased to be the dominant factor in human progress. The
+history of human progress has been mainly the history of man's higher
+educability, the products of which he has projected on to his
+environment. This educability remains on the average what it was a
+dozen generations ago; but the thought-woven tapestry of his
+surroundings is refashioned and improved by each succeeding
+generation. Few men have in greater measure enriched the
+thought-environment with which it is the aim of education to bring
+educable human beings into vital contact, than has Charles Darwin. His
+special field of work was the wide province of biology; but he did
+much to help us to realise that mental factors have contributed to
+organic evolution and that in man, the highest product of Evolution,
+they have reached a position of unquestioned supremacy.
+
+FOOTNOTES:
+
+[Footnote 153: _Origin of Species_ (6th edit.), p. 205.]
+
+[Footnote 154: _Descent of man_ (2nd edit. 1888), Vol. I. p. 99;
+Popular edit. p. 99.]
+
+[Footnote 155: _Ibid._ p. 99.]
+
+[Footnote 156: _The Expression of the Emotions_ (2nd edit.), p. 32.]
+
+[Footnote 157: _Descent of Man_, Vol. II. p. 435.]
+
+[Footnote 158: _Ibid._ 437, 438.]
+
+[Footnote 159: _Origin of Species_ (6th edit.), p. 205.]
+
+[Footnote 160: _Origin of Species_ (6th edit.), p. 205.]
+
+[Footnote 161: _Ibid._ p. 233.]
+
+[Footnote 162: _Ibid._ p. 205.]
+
+[Footnote 163: _Origin of Species_ (6th edit.), p. 233.]
+
+[Footnote 164: _Origin of Species_, pp. 210, 211.]
+
+[Footnote 165: Independently suggested, on somewhat different lines,
+by Profs. J. Mark Baldwin, Henry F. Osborn and the writer.]
+
+[Footnote 166: _Origin of Species_ (6th edit.), p. 206.]
+
+[Footnote 167: _Expression of the Emotions_, p. 13. The passage is
+here somewhat condensed.]
+
+[Footnote 168: _Ibid._ p. 368.]
+
+[Footnote 169: _Expression of the Emotions_, pp. 373, 374.]
+
+[Footnote 170: _Ibid._ p. 368.]
+
+[Footnote 171: _Ibid._ p. 369.]
+
+[Footnote 172: _Expression of the Emotions_, pp. 65 ff.]
+
+[Footnote 173: Cf. William James, _Principles of Psychology_, Vol. II.
+Chap. XXV, New York, 1890.]
+
+[Footnote 174: _Darwinism_, pp. 282, 283, London, 1889.]
+
+[Footnote 175: _Ibid._ p. 283.]
+
+[Footnote 176: _Darwinism_, pp. 283, 284.]
+
+[Footnote 177: _Descent of Man_ (2nd edit.), Vol. II. pp. 136, 137;
+(Popular edit.), pp. 642, 643.]
+
+[Footnote 178: _The Play of Animals_, p. 244, London, 1898.]
+
+[Footnote 179: _Ibid._ p. 283.]
+
+[Footnote 180: _Descent of Man_, Vol. II. p. 60; (Popular edit.), p.
+566.]
+
+[Footnote 181: _The Play of Animals_, p. 76.]
+
+[Footnote 182: _Ibid._ p. 75.]
+
+[Footnote 183: _The Play of Animals_ p. 24.]
+
+[Footnote 184: _Descent of Man_ (1st edit.), Chaps. II, III, V; (2nd
+edit.), Chaps. III, IV, V.]
+
+[Footnote 185: _Descent of Man_, Vol. I. pp. 70, 71; (Popular edit.),
+pp. 70, 71.]
+
+[Footnote 186: _Ibid._ p. 81.]
+
+[Footnote 187: _Ibid._ (Popular edit.), p. 82.]
+
+[Footnote 188: _Comptes Rendus des Sciences_, June 1, 1868.]
+
+[Footnote 189: _Nature_, Vol. LXI. pp. 624, 625 (1900).]
+
+[Footnote 190: _Descent of Man_, Vol. I. p. 100.]
+
+[Footnote 191: _Principles of Psychology_, Vol. II. p. 289.]
+
+[Footnote 192: _Descent of Man_, Vol. I. p. 149.]
+
+[Footnote 193: _Descent of Man_, p. 185.]
+
+[Footnote 194: _Ibid._ p. 150 (footnote).]
+
+
+
+
+VII
+
+THE INFLUENCE OF THE CONCEPTION OF EVOLUTION ON MODERN PHILOSOPHY
+
+BY H. HÖFFDING
+
+_Professor of Philosophy in the University of Copenhagen_
+
+
+I
+
+It is difficult to draw a sharp line between philosophy and natural
+science. The naturalist who introduces a new principle, or
+demonstrates a fact which throws a new light on existence, not only
+renders an important service to philosophy but is himself a
+philosopher in the broader sense of the word. The aim of philosophy in
+the stricter sense is to attain points of view from which the
+fundamental phenomena and the principles of the special sciences can
+be seen in their relative importance and connection. But philosophy in
+this stricter sense has always been influenced by philosophy in the
+broader sense. Greek philosophy came under the influence of logic and
+mathematics, modern philosophy under the influence of natural science.
+The name of Charles Darwin stands with those of Galileo, Newton, and
+Robert Mayer--names which denote new problems and great alterations in
+our conception of the universe.
+
+First of all we must lay stress on Darwin's own personality. His deep love
+of truth, his indefatigable inquiry, his wide horizon, and his steady
+self-criticism make him a scientific model, even if his results and
+theories should eventually come to possess mainly an historical interest.
+In the intellectual domain the primary object is to reach high summits
+from which wide surveys are possible, to reach them toiling honestly
+upwards by the way of experience, and then not to turn dizzy when a summit
+is gained. Darwinians have sometimes turned dizzy, but Darwin never. He saw
+from the first the great importance of his hypothesis, not only because of
+its solution of the old problem as to the value of the concept of species,
+not only because of the grand picture of natural evolution which it
+unrolls, but also because of the life and inspiration its method would
+impart to the study of comparative anatomy, of instinct and of heredity,
+and finally because of the influence it would exert on the whole conception
+of existence. He wrote in his note-book in the year 1837: "My theory would
+give zest to recent and fossil comparative anatomy; it would lead to the
+study of instinct, heredity, and mind-heredity, whole [of]
+metaphysics."[195]
+
+We can distinguish four main points in which Darwin's investigations
+possess philosophical importance.
+
+The evolution hypothesis is much older than Darwin; it is, indeed, one
+of the oldest guessings of human thought. In the eighteenth century is
+was put forward by Diderot and Lamettrie and suggested by Kant (1786).
+As we shall see later, it was held also by several philosophers in the
+first half of the nineteenth century. In his preface to _The Origin of
+Species_, Darwin mentions the naturalists who were his forerunners.
+But he has set forth the hypothesis of evolution in so energetic and
+thorough a manner that it perforce attracts the attention of all
+thoughtful men in a much higher degree than it did before the
+publication of the _Origin_.
+
+And further, the importance of his teaching rests on the fact that he,
+much more than his predecessors, even than Lamarck, sought a
+foundation for his hypothesis in definite facts. Modern science began
+by demanding--with Kepler and Newton--evidence of _varae causae_; this
+demand Darwin industriously set himself to satisfy--hence the wealth
+of material which he collected by his observations and his
+experiments. He not only revived an old hypothesis, but he saw the
+necessity of verifying it by facts. Whether the special cause on which
+he founded the explanation of the origin of species--Natural
+Selection--is sufficient, is now a subject of discussion. He himself
+had some doubt in regard to this question, and the criticisms which
+are directed against his hypothesis hit Darwinism rather than Darwin.
+In his indefatigable search for empirical evidence he is a model even
+for his antagonists: he has compelled them to approach the problems of
+life along other lines than those which were formerly followed.
+
+Whether the special cause to which Darwin appealed is sufficient or not, at
+least to it is probably due the greater part of the influence which he has
+exerted on the general trend of thought. "Struggle for existence" and
+"natural selection" are principles which have been applied, more or less,
+in every department of thought. Recent research, it is true, has discovered
+greater empirical discontinuity--leaps, "mutations"--whereas Darwin
+believed in the importance of small variations slowly accumulated. It has
+also been shown by the experimental method, which in recent biological work
+has succeeded Darwin's more historical method, that types once constituted
+possess great permanence, the fluctuations being restricted within clearly
+defined boundaries. The problem has become more precise, both as to
+variation and as to heredity. The inner conditions of life have in both
+respects shown a greater independence than Darwin had supposed in his
+theory, though he always admitted that the cause of variation was to him a
+great enigma, "a most perplexing problem," and that the struggle for life
+could only occur where variation existed. But, at any rate, it was of the
+greatest importance that Darwin gave a living impression of the struggle
+for life which is everywhere going on, and to which even the highest forms
+of existence must be amenable. The philosophical importance of these ideas
+does not stand or fall with the answer to the question, whether natural
+selection is a sufficient explanation of the origin of species or not it
+has an independent, positive value for everyone who will observe life and
+reality with an unbiased mind.
+
+In accentuating the struggle for life Darwin stands as a
+characteristically English thinker: he continues a train of ideas
+which Hobbes and Malthus had already begun. Moreover in his critical
+views as to the conception of species he had English forerunners; in
+the middle ages Occam and Duns Scotus, in the eighteenth century
+Berkeley and Hume. In his moral philosophy, as we shall see later, he
+is an adherent of the school which is represented by Hutcheson, Home
+and Adam Smith. Because he is no philosopher in the stricter sense of
+the term, it is of great interest to see that his attitude of mind is
+that of the great thinkers of his nation.
+
+In considering Darwin's influence on philosophy we will begin with an
+examination of the attitude of philosophy to the conception of
+evolution at the time when _The Origin of Species_ appeared. We will
+then examine the effects which the theory of evolution, and especially
+the idea of the struggle for life, has had, and naturally must have,
+on the discussion of philosophical problems.
+
+
+II
+
+When _The Origin of Species_ appeared fifty years ago Romantic
+speculation, Schelling's and Hegel's philosophy, still reigned on the
+continent, while in England Positivism, the philosophy of Comte and
+Stuart Mill, represented the most important trend of thought. German
+speculation had much to say on evolution, it even pretended to be a
+philosophy of evolution. But then the word "evolution" was to be taken
+in an ideal, not in a real, sense. To speculative thought the forms
+and types of nature formed a system of ideas, within which any form
+could lead us by continuous transitions to any other. It was a
+classificatory system which was regarded as a divine world of thought
+or images, within which metamorphoses could go on--a condition
+comparable with that in the mind of the poet when one image follows
+another with imperceptible changes. Goethe's ideas of evolution, as
+expressed in his _Metamorphosen der Pflanzen und der Thiere_, belong
+to this category; it is, therefore, incorrect to call him a forerunner
+of Darwin. Schelling and Hegel held the same idea; Hegel expressly
+rejected the conception of a real evolution in time as coarse and
+materialistic. "Nature," he says, "is to be considered as a _system of
+stages_, the one necessarily arising from the other, and being the
+nearest truth of that from which it proceeds; but not in such a way
+that the one is _naturally_ generated by the other; on the contrary
+[their connection lies] in the inner idea which is the ground of
+nature. The _metamorphosis_ can be ascribed only to the notion as
+such, because it alone is evolution.... It has been a clumsy idea in
+the older as well as in the newer philosophy of nature, to regard the
+transformation and the transition from one natural form and sphere to
+a higher as an outward and actual production."[196]
+
+The only one of the philosophers of Romanticism who believed in a
+real, historical evolution, a real production of new species, was
+Oken.[197] Danish philosophers, such as Treschow (1812) and Sibbern
+(1846), have also broached the idea of an historical evolution of all
+living beings from the lowest to the highest. Schopenhauer's
+philosophy has a more realistic character than that of Schelling's and
+Hegel's, his diametrical opposites, although he also belongs to the
+romantic school of thought. His philosophical and psychological views
+were greatly influenced by French naturalists and philosophers,
+especially by Cabanis and Lamarck. He praises the "ever memorable
+Lamarck," because he laid so much stress on the "will to live." But he
+repudiates as a "wonderful error" the idea that the organs of animals
+should have reached their present perfection through a development in
+time, during the course of innumerable generations. It was, he said, a
+consequence of the low standard of contemporary French philosophy,
+that Lamarck came to the idea of the construction of living beings in
+time through succession![198]
+
+The positivistic stream of thought was not more in favour of a real
+evolution than was the Romantic school. Its aim was to adhere to
+positive facts: it looked with suspicion on far-reaching speculation.
+Comte laid great stress on the discontinuity found between the
+different kingdoms of nature, as well as within each single kingdom.
+As he regarded as unscientific every attempt to reduce the number of
+physical forces, so he rejected entirely the hypothesis of Lamarck
+concerning the evolution of species; the idea of species would in his
+eyes absolutely lose its importance if a transition from species to
+species under the influence of conditions of life were admitted. His
+disciples (Littré, Robin) continued to direct against Darwin the
+polemics which their master had employed against Lamarck. Stuart Mill,
+who, in the theory of knowledge, represented the empirical or
+positivistic movement in philosophy--like his English forerunners from
+Locke to Hume--founded his theory of knowledge and morals on the
+experience of the single individual. He sympathised with the theory of
+the original likeness of all individuals and derived their
+differences, on which he practically and theoretically laid much
+stress, from the influence both of experience and education, and,
+generally, of physical and social causes. He admitted an individual
+evolution, and, in the human species, an evolution based on social
+progress; but no physiological evolution of species. He was afraid
+that the hypothesis of heredity would carry us back to the old theory
+of "innate" ideas.
+
+Darwin was more empirical than Comte and Mill; experience disclosed to
+him a deeper continuity than they could find; closer than before the
+nature and fate of the single individual were shown to be interwoven
+in the great web binding the life of the species with nature as a
+whole. And the continuity which so many idealistic philosophers could
+find only in the world of thought, he showed to be present in the
+world of reality.
+
+
+III
+
+Darwin's energetic renewal of the old idea of evolution has its chief
+importance in strengthening the conviction of this real continuity in
+the world, of continuity in the series of form and events. It was a
+great support for all those who were prepared to base their conception
+of life on scientific grounds. Together with the recently discovered
+law of the conservation of energy, it helped to produce the great
+realistic movement which characterises the last third of the
+nineteenth century. After the decline of the Romantic movement people
+wished to have firmer ground under their feet and reality now asserted
+itself in a more emphatic manner than in the period of Romanticism. It
+was easy for Hegel to proclaim that "the real" was "the rational," and
+that "the rational" was "the real": reality itself existed for him
+only in the interpretation of ideal reason, and if there was anything
+which could not be merged in the higher unity of thought, then it was
+only an example of the "impotence of nature to hold to the idea." But
+now concepts are to be founded on nature and not on any system of
+categories too confidently deduced _à priori_. The new devotion to
+nature had its recompense in itself, because the new points of view
+made us see that nature could indeed "hold to ideas," though perhaps
+not to those which we had cogitated beforehand.
+
+A most important question for philosophers to answer was whether the
+new views were compatible with an idealistic conception of life and
+existence. Some proclaimed that we have now no need of any philosophy
+beyond the principles of the conservation of matter and energy and the
+principle of natural evolution: existence should and could be
+definitely and completely explained by the laws of material nature.
+But abler thinkers saw that the thing was not so simple. They were
+prepared to give the new views their just place and to examine what
+alterations the old views must undergo in order to be brought into
+harmony with the new data.
+
+The realistic character of Darwin's theory was shown not only in the
+idea of natural continuity, but also, and not least, in the idea of
+the cause whereby organic life advances step by step. This idea--the
+idea of the struggle for life--implied that nothing could persist, if
+it had no power to maintain itself under the given conditions. Inner
+value alone does not decide. Idealism was here put to its hardest
+trial. In continuous evolution it could perhaps still find an analogy
+to the inner evolution of ideas in the mind; but in the demand for
+power in order to struggle with outward conditions Realism seemed to
+announce itself in its most brutal form. Every form of Idealism had to
+ask itself seriously how it was going to "struggle for life" with this
+new Realism.
+
+We will now give a short account of the position which leading
+thinkers in different countries have taken up in regard to this
+question.
+
+I. Herbert Spencer was the philosopher whose mind was best prepared by his
+own previous thinking to admit the theory of Darwin to a place in his
+conception of the world. His criticism of the arguments which had been put
+forward against the hypothesis of Lamarck, showed that Spencer, as a young
+man, was an adherent to the evolution idea. In his _Social Statics_ (1850)
+he applied this idea to human life and moral civilisation. In 1852 he wrote
+an essay on _The Development Hypothesis_, in which he definitely stated his
+belief that the differentiation of species, like the differentiation within
+a single organism, was the result of development. In the first edition of
+his _Psychology_ (1855) he took a step which put him in opposition to the
+older English school (from Locke to Mill): he acknowledged "innate ideas"
+so far as to admit the tendency of acquired habits to be inherited in the
+course of generations, so that the nature and functions of the individual
+are only to be understood through its connection with the life of the
+species. In 1857, in his essay on _Progress_, he propounded the law of
+differentiation as a general law of evolution, verified by examples from
+all regions of experience, the evolution of species being only one of these
+examples. On the effect which the appearance of _The Origin of Species_ had
+on his mind he writes in his _Autobiography_: "Up to that time ... I held
+that the sole cause of organic evolution is the inheritance of
+functionally-produced modifications. The _Origin of Species_ made it clear
+to me that I was wrong, and that the larger part of the facts cannot be due
+to any such cause.... To have the theory of organic evolution justified was
+of course to get further support for that theory of evolution at large with
+which ... all my conceptions were bound up."[199] Instead of the
+metaphorical expression "natural selection," Spencer introduced the term
+"survival of the fittest," which found favour with Darwin as well as with
+Wallace.
+
+In working out his ideas of evolution, Spencer found that
+differentiation was not the only form of evolution. In its simplest
+form evolution is mainly a concentration, previously scattered
+elements being integrated and losing independent movement.
+Differentiation is only forthcoming when minor wholes arise within a
+greater whole. And the highest form of evolution is reached when there
+is a harmony between concentration and differentiation, a harmony
+which Spencer calls equilibration and which he defines as a moving
+equilibrium. At the same time this definition enables him to
+illustrate the expression "survival of the fittest." "Every living
+organism exhibits such a moving equilibrium--a balanced set of
+functions constituting its life; and the overthrow of this balanced
+set of functions or moving equilibrium is what we call death. Some
+individuals in a species are so constituted that their moving
+equilibria are less easily overthrown than those of other
+individuals; and these are the fittest which survive, or, in Mr.
+Darwin's language, they are the select which nature preserves."[200]
+Not only in the domain of organic life, but in all domains, the summit
+of evolution is, according to Spencer, characterised by such a
+harmony--by a moving equilibrium.
+
+Spencer's analysis of the concept of evolution, based on a great
+variety of examples, has made this concept clearer and more definite
+than before. It contains the three elements; integration,
+differentiation and equilibration. It is true that a concept which is
+to be valid for all domains of experience must have an abstract
+character, and between the several domains there is, strictly
+speaking, only a relation of analogy. So there is only analogy between
+psychical and physical evolution. But this is no serious objection,
+because general concepts do not express more than analogies between
+the phenomena which they represent. Spencer takes his leading forms
+from the material world in defining evolution (in the simplest form)
+as integration of matter and dissipation of movement; but as he--not
+always quite consistently[201]--assumed a correspondence of mind and
+matter, he could very well give these terms an indirect importance for
+psychical evolution. Spencer has always, in my opinion with full
+right, repudiated the ascription of materialism. He is no more a
+materialist than Spinoza. In his _Principles of Psychology_ (§ 63) he
+expressed himself very clearly: "Though it seems easier to translate
+so-called matter into so-called spirit, than to translate so-called
+spirit into so-called matter--which latter is indeed wholly
+impossible--yet no translation can carry us beyond our symbols." These
+words lead us naturally to a group of thinkers whose starting-point
+was psychical evolution. But we have still one aspect of Spencer's
+philosophy to mention.
+
+Spencer founded his "laws of evolution" on an inductive basis, but he
+was convinced that they could be deduced from the law of the
+conservation of energy. Such a deduction is, perhaps, possible for the
+more elementary forms of evolution, integration and differentiation;
+but it is not possible for the highest form, the equilibration, which
+is a harmony of integration and differentiation. Spencer can no more
+deduce the necessity for the eventual appearance of "moving
+equilibria" of harmonious totalities than Hegel could guarantee the
+"higher unities" in which all contradictions should be reconciled. In
+Spencer's hands the theory of evolution acquired a more decidedly
+optimistic character than in Darwin's; but I shall deal later with the
+relation of Darwin's hypothesis to the opposition of optimism and
+pessimism.
+
+II. While the starting-point of Spencer was biological or
+cosmological, psychical evolution being conceived as in analogy with
+physical, a group of eminent thinkers--in Germany Wundt, in France
+Fouillée, in Italy Ardigò--took, each in his own manner, their
+starting-point in psychical evolution as an original fact and as a
+type of all evolution, the hypothesis of Darwin coming in as a
+corroboration and as a special example. They maintain the continuity
+of evolution; they find this character most prominent in psychical
+evolution, and this is for them a motive to demand a corresponding
+continuity in the material, especially in the organic domain.
+
+To Wundt and Fouillée the concept of will is prominent. They see the
+type of all evolution in the transformation of the life of will from
+blind impulse to conscious choice; the theories of Lamarck and Darwin
+are used to support the view that there is in nature a tendency to
+evolution in steady reciprocity with external conditions. The struggle
+for life is here only a secondary fact. Its apparent prominence is
+explained by the circumstance that the influence of external
+conditions is easily made out, while inner conditions can be verified
+only through their effects. For Ardigò the evolution of thought was
+the starting-point and the type: in the evolution of a scientific
+hypothesis we see a progress from the indefinite (_indistinto_) to the
+definite (_distinto_), and this is a characteristic of all evolution,
+as Ardigò has pointed out in a series of works. The opposition between
+_indistinto_ and _distinto_ corresponds to Spencer's opposition
+between homogeneity and heterogeneity. The hypothesis of the origin of
+differences of species from more simple forms is a special example of
+the general law of evolution.
+
+In the views of Wundt and Fouillée we find the fundamental idea of
+idealism psychical phenomena as expressions of the innermost nature of
+existence. They differ from the older Idealism in the great stress
+which they lay on evolution as a real, historical process which is
+going on through steady conflict with external conditions. The
+Romantic dread of reality is broken. It is beyond doubt that Darwin's
+emphasis on the struggle for life as a necessary condition of
+evolution has been a very important factor in carrying philosophy back
+to reality from the heaven of pure ideas. The philosophy of Ardigò, on
+the other side, appears more as a continuation and deepening of
+positivism, though the Italian thinker arrived at his point of view
+independently of French-English positivism. The idea of continuous
+evolution is here maintained in opposition to Comte's and Mill's
+philosophy of discontinuity. From Wundt and Fouillée Ardigò differs in
+conceiving psychical evolution not as an immediate revelation of the
+innermost nature of existence, but only as a single, though the most
+accessible example, of evolution.
+
+III. To the French philosophers Boutroux and Bergson, evolution proper
+is continuous and qualitative, while outer experience and physical
+science give us fragments only, sporadic processes and mechanical
+combinations. To Bergson, in his recent work _L'Evolution Créatrice_,
+evolution consists in an _élan de vie_ which to our fragmentary
+observation and analytic reflexion appears as broken into a manifold
+of elements and processes. The concept of matter in its scientific
+form is the result of this breaking asunder, essential for all
+scientific reflexion. In these conceptions the strongest opposition
+between inner and outer conditions of evolution is expressed: in the
+domain of internal conditions spontaneous development of qualitative
+forms--in the domain of external conditions discontinuity and
+mechanical combination.
+
+We see, then, that the theory of evolution has influenced philosophy
+in a variety of forms. It has made idealistic thinkers revise their
+relation to the real world; it has led positivistic thinkers to find a
+closer connection between the facts on which they based their views;
+it has made us all open our eyes for new possibilities to arise
+through the _prima facie_ inexplicable "spontaneous" variations which
+are the condition of all evolution. This last point is one of peculiar
+interest. Deeper than speculative philosophy and mechanical science
+saw in the days of their triumph, we catch sight of new streams, whose
+sources and laws we have still to discover. Most sharply does this
+appear in the theory of mutation, which is only a stronger
+accentuation of a main point in Darwinism. It is interesting to see
+that an analogous problem comes into the foreground in physics through
+the discovery of radioactive phenomena, and in psychology through the
+assumption of psychical new formations (as held by Boutroux, William
+James and Bergson). From this side, Darwin's ideas, as well as the
+analogous ideas in other domains, incite us to renewed examination of
+our first principles, their rationality and their value. On the other
+hand, his theory of the struggle for existence challenges us to
+examine the conditions and discuss the outlook as to the persistence
+of human life and society and of the values that belong to them. It is
+not enough to hope (or fear?) the rising of new forms; we have also to
+investigate the possibility of upholding the forms and ideals which
+have hitherto been the bases of human life. Darwin has here given his
+age the most earnest and most impressive lesson. This side of Darwin's
+theory is of peculiar interest to some special philosophical problems
+to which I now pass.
+
+
+IV
+
+Among philosophical problems the problem of knowledge has in the last
+century occupied a foremost place. It is natural, then, to ask how
+Darwin and the hypothesis whose most eminent representative he is,
+stand to this problem.
+
+Darwin started an hypothesis. But every hypothesis is won by inference
+from certain presuppositions, and every inference is based on the
+general principles of human thought. The evolution hypothesis
+presupposes, then, human thought and its principles. And not only the
+abstract logical principles are thus pre-supposed. The evolution
+hypothesis purports to be not only a formal arrangement of phenomena,
+but to express also the law of a real process. It supposes, then, that
+the real data--all that in our knowledge which we do not produce
+ourselves, but which we in the main simply receive--are subject to
+laws which are at least analogous to the logical relations of our
+thoughts; in other words, it assumes the validity of the principle of
+causality. If organic species could arise without cause there would be
+no use in framing hypotheses. Only if we assume the principle of
+causality, is there a problem to solve.
+
+Though Darwinism has had a great influence on philosophy considered as
+a striving after a scientific view of the world, yet here is a point
+of view--the epistemological--where philosophy is not only independent
+but reaches beyond any result of natural science. Perhaps it will be
+said: the powers and functions of organic beings only persist (perhaps
+also only arise) when they correspond sufficiently to the conditions
+under which the struggle of life is to go on. Human thought itself is,
+then, a variation (or a mutation) which has been able to persist and
+to survive. Is not, then, the problem of knowledge solved by the
+evolution hypothesis? Spencer had given an affirmative answer to this
+question before the appearance of _The Origin of Species_. For the
+individual, he said, there is an _à priori_, original, basis (or
+_Anlage_) for all mental life; but in the species all powers have
+developed in reciprocity with extendal conditions. Knowledge is here
+considered from the practical point of view, as a weapon in the
+struggle for life, as an "organon" which has been continuously in use
+for generations. In recent years the economic or pragmatic
+epistemology, as developed by Avenarius and Mach in Germany, and by
+James in America, points in the same direction. Science, it is said,
+only maintains those principles and presuppositions which are
+necessary to the simplest and clearest orientation be applied to
+experience and to practical work, will successively be eliminated.
+
+In these views a striking and important application is made of the
+idea of struggle for life to the development of human thought. Thought
+must, as all other things in the world, struggle for life. But this
+whole consideration belongs to psychology, not to the theory of
+knowledge (epistemology), which is concerned only with the validity of
+knowledge, not with its historical origin. Every hypothesis to explain
+the origin of knowledge must submit to cross-examination by the theory
+of knowledge, because it works with the fundamental forms and
+principles of human thought. We cannot go further back than these
+forms and principles, which it is the aim of epistemology to ascertain
+and for which no further reason can be given.[202]
+
+But there is another side of the problem which is, perhaps, of more
+importance and which epistemology generally overlooks. If new
+variations can arise, not only in organic but perhaps also in
+inorganic nature, new tasks are placed before the human mind. The
+question is, then, if it has forms in which there is room for the new
+matter? We are here touching a possibility which the great master of
+epistemology did not bring to light. Kant supposed confidently that no
+other matter of knowledge could stream forth from the dark source
+which he called "the thing-in-itself," than such as could be
+synthesised in our existing forms of knowledge. He mentions the
+possibility of other forms than the human, and warns us against the
+dogmatic assumption that the human conception of existence should be
+absolutely adequate. But he seems to be quite sure that the
+thing-in-itself works constantly, and consequently always gives us
+only what our powers can master. This assumption was a consequence of
+Kant's rationalistic tendency, but one for which no warrant can be
+given. Evolutionism and systematism are opposing tendencies which can
+never be absolutely harmonised one with the other. Evolution may at
+any time break some form which the system-monger regards as finally
+established. Darwin himself felt a great difference in looking at
+variation as an evolutionist and as a systematist. When he was working
+at his evolution theory, he was very glad to find variations; but they
+were a hindrance to him when he worked as a systematist, in preparing
+his work on Cirripedia. He says in a letter: "I had thought the same
+parts of the same species more resemble (than they do anyhow in
+Cirripedia) objects cast in the same mould. Systematic work would be
+easy were it not for this confounded variation, which, however, is
+pleasant to me as a speculatist, though odious to me as a
+systematist."[203] He could indeed be angry with variations even as an
+evolutionist; but then only because he could not explain them, not
+because he could not classify them. "If, as I must think, external
+conditions produce little _direct_ effect, what the devil determines
+each particular variation?"[204] What Darwin experienced in this
+particular domain holds good of all knowledge. All knowledge is
+systematic, in so far as it strives to put phenomena in quite definite
+relations, one to another. But the systematisation can never be
+complete. And here Darwin has contributed much to widen the world, for
+us. He has shown us forces and tendencies in nature which make
+absolute systems impossible, at the same time that they give us new
+objects and problems. There is still a place for what Lessing called
+"the unceasing striving after truth," while "absolute truth" (in the
+sense of a closed system) is unattainable so long as life and
+experience are going on.
+
+There is here a special remark to be made. As we have seen above,
+recent research has shown that natural selection or struggle for life
+is no explanation of variations. Hugo de Vries distinguishes between
+partial and embryonal variations, or between variations and mutations,
+only the last-named being heritable, and therefore of importance for
+the origin of new species. But the existence of variations is not only
+of interest for the problem of the origin of species; it has also a
+more general interest. An individual does not lose its importance for
+knowledge, because its qualities are not heritable. On the contrary,
+in higher beings at least, individual peculiarities will become more
+and more independent objects of interest. Knowledge takes account of
+the biographies not only of species, but also of individuals: it seeks
+to find the law of development of the single individual.[205] As
+Leibnitz said long ago, individuality consists in the law of the
+changes of a being: "La loi du changement fait l'individualité de
+chaque substance." Here is a world which is almost new for science,
+which till now has mainly occupied itself with general laws and forms.
+But these are ultimately only means to understand the individual
+phenomena, in whose nature and history a manifold of laws and forms
+always coöperate. The importance of this remark will appear in the
+sequel.
+
+
+V
+
+To many people the Darwinian theory of natural selection or struggle
+for existence seemed to change the whole conception of life, and
+particularly all the conditions on which the validity of ethical ideas
+depends. If only that has persistence which can be adapted to a given
+condition, what will then be the fate of our ideals, of our standards
+of good and evil? Blind force seems to reign, and the only thing that
+counts seems to be the most heedless use of power. Darwinism, it was
+said, has proclaimed brutality. No other difference seems permanent
+save that between the sound, powerful and happy on the one side, the
+sick, feeble and unhappy on the other; and every attempt to alleviate
+this difference seems to lead to general enervation. Some of those who
+interpreted Darwinism in this manner felt an aesthetic delight in
+contemplating the heedlessness and energy of the great struggle for
+existence and anticipated the realisation of a higher human type as
+the outcome of it: so Nietzsche and his followers. Others recognising
+the same consequences in Darwinism regarded these as one of the
+strongest objections against it; so Dühring and Kropotkin (in his
+earlier works).
+
+This interpretation of Darwinism was frequent in the interval between
+the two main works of Darwin--_The Origin of Species_ and _The Descent
+of Man_. But even during this interval it was evident to an attentive
+reader that Darwin himself did not found his standard of good and evil
+on the features of the life of nature he had emphasised so strongly.
+He did not justify the ways along which nature reached its ends; he
+only pointed them out. The "real" was not to him, as to Hegel, one
+with the "rational." Darwin has, indeed, by his whole conception of
+nature, rendered a great service to ethics in making the difference
+between the life of nature and the ethical life appear in so strong a
+light. The ethical problem could now be stated in a sharper form than
+before. But this was not the first time that the idea of the struggle
+for life was put in relation to the ethical problem. In the
+seventeenth century Thomas Hobbes gave the first impulse to the whole
+modern discussion of ethical principles in his theory of _bellum
+omnium contra omnes_. Men, he taught, are in the state of nature
+enemies one of another, and they live either in fright or in the glory
+of power. But it was not the opinion of Hobbes that this made ethics
+impossible. On the contrary, he found a standard for virtue and vice
+in the fact that some qualities and actions have a tendency to bring
+us out of the state of war and to secure peace, while other qualities
+have a contrary tendency. In the eighteenth century even Immanuel
+Kant's ideal ethics had--so far as can be seen--a similar origin.
+Shortly before the foundation of his definitive ethics, Kant wrote his
+_Idee zu einer allgemeinen Weltgeschichte_ (1784), where--in a way
+which reminds us of Hobbes, and is prophetic of Darwin--he describes
+the forward-driving power of struggle in the human world. It is here
+as with the struggle of the trees for light and air, through which
+they compete with one another in height. Anxiety about war can only be
+allayed by an ordinance which gives everyone his full liberty under
+acknowledgment of the equal liberty of others. And such ordinance and
+acknowledgment are also attributes of the content of the moral law, as
+Kant proclaimed it in the year after the publication of his essay
+(1785).[206] Kant really came to his ethics by the way of evolution,
+though he afterwards disavowed it. Similarly the same line of thought
+may be traced in Hegel though it has been disguised in the form of
+speculative dialectics.[207] And in Schopenhauer's theory of the blind
+will to live and its abrogation by the ethical feeling, which is
+founded on universal sympathy, we have a more individualistic form of
+the same idea.
+
+It was, then, not entirely a foreign point of view which Darwin
+introduced into ethical thought, even if we take no account of the
+poetical character of the word "struggle" and of the more direct
+adaptation, through the use and non-use of power, which Darwin also
+emphasised. In _The Descent of Man_ he has devoted a special
+chapter[208] to a discussion of the origin of the ethical
+consciousness. The characteristic expression of this consciousness he
+found, just as Kant did, in the idea of "ought"; it was the origin of
+this new idea which should be explained. His hypothesis was that the
+ethical "ought" has its origin in the social and parental instincts,
+which, as well as other instincts (e.g. the instinct of
+self-preservation), lie deeper than pleasure and pain. In many
+species, not least in the human species, these instincts are fostered
+by natural selection; and when the powers of memory and comparison are
+developed, so that single acts can be valued according to the claims
+of the deep social instinct, then consciousness of duty and remorse
+are possible. Blind instinct has developed to conscious ethical will.
+
+As already stated, Darwin, as a moral philosopher belongs to the
+school that was founded by Shaftesbury, and was afterwards represented
+by Hutcheson, Hume, Adam Smith, Comte and Spencer. His merit is,
+first, that he has given this tendency of thought a biological
+foundation, and that he has stamped on it a doughty character in
+showing that ethical ideas and sentiments, rightly conceived, are
+forces which are at work in the struggle for life.
+
+There are still many questions to solve. Not only does the ethical
+development within the human species contain features still
+unexplained;[209] but we are confronted by the great problem whether
+after all a genetic historical theory can be of decisive importance
+here. To every consequent ethical consciousness there is a standard of
+value, a primordial value which determines the single ethical
+judgments as their last presupposition, and the "rightness" of this
+basis, the "value" of this value can as little be discussed as the
+"rationality" of our logical principles. There is here revealed a
+possibility of ethical scepticism which evolutionistic ethics (as well
+as intuitive or rationalistic ethics) has overlooked. No demonstration
+can show that the results of the ethical development are definitive
+and universal. We meet here again with the important opposition of
+systematisation and evolution. There will, I think, always be an open
+question here, though comparative ethics, of which we have so far only
+the first attempts, can do much to throw light on it.
+
+It would carry us too far to discuss all the philosophical works on
+ethics, which have been influenced directly or indirectly by
+evolutionism. I may, however, here refer to the book of C. M.
+Williams, _A Review of the Systems of Ethics founded on the Theory of
+Evolution_,[210] in which, besides Darwin, the following authors are
+reviewed: Wallace, Haeckel, Spencer, Fiske, Rolph, Barratt, Stephen,
+Carneri, Höffding, Gizycki, Alexander, Rée. As works which criticise
+evolutionistic ethics from an intuitive point of view and in an
+instructive way, may be cited: Guyau, _La morale anglaise
+contemporaine_,[211] and Sorley, _Ethics of Naturalism_. I will only
+mention some interesting contributions to ethical discussion which can
+be found in Darwinism besides the idea of struggle for life.
+
+The attention which Darwin has directed to variations has opened our
+eyes to the differences in human nature as well as in nature
+generally. There is here a fact of great importance for ethical
+thought, no matter from what ultimate premiss it starts. Only from a
+very abstract point of view can different individuals be treated in
+the same manner. The most eminent ethical thinkers, men such as Jeremy
+Bentham and Immanuel Kant, who discussed ethical questions from very
+opposite standpoints, agreed in regarding all men as equal in respect
+of ethical endowment. In regard to Bentham, Leslie Stephen remarks:
+"He is determined to be thoroughly empirical, to take men as he found
+them. But his utilitarianism supposed that men's views of happiness
+and utility were uniform and clear, and that all that was wanted was
+to show them the means by which their ends could be reached."[212] And
+Kant supposed that every man would find the "categorical imperative"
+in his consciousness, when he came to sober reflexion, and that all
+would have the same qualifications to follow it. But if continual
+variations, great or small, are going on in human nature, it is the
+duty of ethics to make allowance for them, both in making claims, and
+in valuing what is done. A new set of ethical problems have their
+origin here.[213] It is an interesting fact that Stuart Mill's book
+_On Liberty_ appeared in the same year as _The Origin of Species_.
+Though Mill agreed with Bentham about the original equality of all
+men's endowments, he regarded individual differences as a necessary
+result of physical and social influences, and he claimed that free
+play shall be allowed to differences of character so far as is
+possible without injury to other men. It is a condition of individual
+and social progress that a man's mode of action should be determined
+by his own character and not by tradition and custom, nor by abstract
+rules. This view was to be corroborated by the theory of Darwin.
+
+But here we have reached a point of view from which the criticism,
+which in recent years has often been directed against Darwin--that
+small variations are of no importance in the struggle for life--is of
+no weight. From an ethical standpoint, and particularly from the
+ethical standpoint of Darwin himself, it is a duty to foster
+individual differences that can be valuable, even though they can
+neither be of service for physical preservation nor be physically
+inherited. The distinction between variation and mutation is here
+without importance. It is quite natural that biologists should be
+particularly interested in such variations as can be inherited and
+produce new species. But in the human world there is not only a
+physical, but also a mental and social heredity. When an ideal human
+character has taken form, then there is shaped a type, which through
+imitation and influence can become an important factor in subsequent
+development, even if it cannot form a species in the biological sense
+of the word. Spiritually strong men often succumb in the physical
+struggle for life; but they can nevertheless be victorious through the
+typical influence they exert, perhaps on very distant generations, if
+the remembrance of them is kept alive, be it in legendary or in
+historical form. Their very failure can show that a type has taken
+form which is maintained at all risks, a standard of life which is
+adhered to in spite of the strongest opposition. The question "to be
+or not to be" can be put from very different levels of being: it has
+too often been considered a consequence of Darwinism that this
+question is only to be put from the lowest level. When a stage is
+reached, where ideal (ethical, intellectual, aesthetic) interests are
+concerned, the struggle for life is a struggle for the preservation of
+this stage. The giving up of a higher standard of life is a sort of
+death; for there is not only a physical, there is also a spiritual,
+death.
+
+
+VI
+
+The Socratic character of Darwin's mind appears in his wariness in
+drawing the last consequences of his doctrine, in contrast both with
+the audacious theories of so many of his followers and with the
+consequences which his antagonists were busy in drawing. Though he, as
+we have seen, saw from the beginning that his hypothesis would
+occasion "a whole of metaphysics," he was himself very reserved as to
+the ultimate questions, and his answers to such questions were
+extorted from him.
+
+As to the question of optimism and pessimism, Darwin held that though
+pain and suffering were very often the ways by which animals were led
+to pursue that course of action which is most beneficial to the
+species, yet pleasurable feelings were the most habitual guides. "We
+see this in the pleasure from exertion, even occasionally from great
+exertion of the body or mind, in the pleasure of our daily meals, and
+especially in the pleasure derived from sociability, and from loving
+our families." But there was to him so much suffering in the world
+that it was a strong argument against the existence of an intelligent
+First Cause.[214]
+
+It seems to me that Darwin was not so clear on another question, that
+of the relation between improvement and adaptation. He wrote to Lyell:
+"When you contrast natural selection and 'improvement,' you seem
+always to overlook ... that every step in the natural selection of
+each species implies improvement in that species _in relation to its
+condition of life_.... Improvement implies, I suppose, _each form
+obtaining many parts or organs_, all excellently adapted for their
+functions." "All this," he adds, "seems to me quite compatible with
+certain forms fitted for simple conditions, remaining unaltered, or
+being, degraded."[215] But the great question is, if the conditions of
+life will in the long run favour "improvement" in the sense of
+differentiation (or harmony of differentiation and integration). Many
+beings are best adapted to their conditions of life if they have few
+organs and few necessities. Pessimism would not only be the
+consequence, if suffering outweighed happiness, but also if the most
+elementary forms of happiness were predominant, or if there were a
+tendency to reduce the standard of life to the simplest possible, the
+contentment of inertia or stable equilibrium. There are animals which
+are very highly differentiated and active in their young state, but
+later lose their complex organisation and concentrate themselves on
+the one function of nutrition. In the human world analogies to this
+sort of adaptation are not wanting. Young "idealists" very often end
+as old "Philistines." Adaptation and progress are not the same.
+
+Another question of great importance in respect to human evolution is,
+whether there will be always a possibility for the existence of an
+impulse to progress, an impulse to make great claims on life, to be
+active and to alter the conditions of life instead of adapting to them
+in a passive manner. Many people do not develop because they have too
+few necessities, and because they have no power to imagine other
+conditions of life than those under which they live. In his remarks on
+"the pleasure from exertion" Darwin has a point of contact with the
+practical idealism of former times--with the ideas of Lessing and
+Goethe, of Condorcet and Fichte. The continual striving which was the
+condition of salvation to Faust's soul, is also the condition of
+salvation to mankind. There is a holy fire which we ought to keep
+burning, if adaptation is really to be improvement. If, as I have
+tried to show in my _Philosophy of Religion_, the innermost core of
+all religion is faith in the persistence of value in the world, and if
+the highest values express themselves in the cry "Excelsior!" then the
+capital point is, that this cry should always be heard and followed.
+We have here a corollary of the theory of evolution in its application
+to human life.
+
+Darwin declared himself an agnostic, not only because he could not
+harmonise the large amount of suffering in the world with the idea of
+a God as its first cause, but also because he "was aware that if we
+admit a first cause, the mind still craves to know whence it came and
+how it arose."[216] He saw, as Kant had seen before him and expressed
+in his _Kritik der Urtheilskraft_, that we cannot accept either of the
+only two possibilities which we are able to conceive: chance (or brute
+force) and design. Neither mechanism nor teleology can give an
+absolute answer to ultimate questions. The universe, and especially
+the organic life in it, can neither be explained as a mere
+combination of absolute elements nor as the effect of a constructing
+thought. Darwin concluded, as Kant, and before him Spinoza, that the
+oppositions and distinctions which our experience presents, cannot
+safely be regarded as valid for existence in itself. And, with Kant
+and Fichte, he found his stronghold in the conviction that man has
+something to do, even if he cannot solve all enigmas. "The safest
+conclusion seems to me that the whole subject is beyond the scope of
+man's intellect; but man can do his duty."[217]
+
+Is this the last word of human thought? Does not the possibility, that
+man can do his duty, suppose that the conditions of life allow of
+continuous ethical striving, so that there is a certain harmony
+between cosmic order and human ideals? Darwin himself has shown how
+the consciousness of duty can arise as a natural result of evolution.
+Moreover there are lines of evolution which have their end in ethical
+idealism, in a kingdom of values, which must struggle for life as all
+things in the world must do, but a kingdom which has its firm
+foundation in reality.
+
+FOOTNOTES:
+
+[Footnote 195: _Life and Letters of Charles Darwin_, Vol. I. p. 8.]
+
+[Footnote 196: _Encyclopädie der philosophischen Wissenschaften_ (4th
+edit.), Berlin, 1845, § 249.]
+
+[Footnote 197: _Lehrbuch der Naturphilosophie_, Jena, 1809.]
+
+[Footnote 198: _Ueber den Willen in der Natur_ (2nd edit.), Frankfurt
+a. M., 1854, pp. 41-43.]
+
+[Footnote 199: Spencer, _Autobiography_, Vol. II. p. 50, London and
+New York, 1904.]
+
+[Footnote 200: _Autobiography_, Vol. II. p. 100.]
+
+[Footnote 201: Cf. my letter to him 1876, now printed in Duncan's
+_Life and Letters of Herbert Spencer_, p. 178. London, 1908.]
+
+[Footnote 202: The present writer, many years ago, in his _Psychology_
+(Copenhagen, 1882; Eng. transl. London, 1891), criticised the
+evolutionistic treatment of the problem of knowledge from the Kantian
+point of view.]
+
+[Footnote 203: _Life and Letters_, Vol. II. p. 37.]
+
+[Footnote 204: _Ibid._ p. 232.]
+
+[Footnote 205: The new science of Ecology occupies an intermediate
+position between the biography of species and the biography of
+individuals. Compare _Congress of Arts and Science_, St. Louis, Vol.
+V. 1906 (The Reports of Drude and Robinson) and the work of my
+colleague, E. Warming.]
+
+[Footnote 206: Cf. my _History of Modern Philosophy_ (Eng. transl.
+London, 1900), I. pp. 76-79.]
+
+[Footnote 207: "Herrschaft und Knechtschaft," _Phönomenologie des
+Geistes_, IV. A., Leiden, 1907.]
+
+[Footnote 208: _The Descent of Man_, Vol. I. Ch. iii.]
+
+[Footnote 209: The works of Westermarck and Hobhouse throw new light
+on many of these features.]
+
+[Footnote 210: New York and London, 1893.]
+
+[Footnote 211: Paris, 1879.]
+
+[Footnote 212: _English literature and society in the eighteenth
+century_, London, 1904, p. 187.]
+
+[Footnote 213: Cf. my paper, "The law of relativity in Ethics,"
+_International Journal of Ethics_, Vol. I. 1891, pp. 37-62.]
+
+[Footnote 214: _Life and Letters_, Vol. I. p. 310.]
+
+[Footnote 215: _Ibid._ Vol. II. p. 177.]
+
+[Footnote 216: _Life and Letters_, Vol. 1. p. 306.]
+
+[Footnote 217: _Life and Letters_, p. 307.]
+
+
+
+
+VIII
+
+THE INFLUENCE OF DARWIN UPON RELIGIOUS THOUGHT
+
+BY P. N. WAGGETT, M.A., S.S.J.E.
+
+
+I
+
+The object of this paper is first to point out certain elements of the
+Darwinian influence upon Religious thought, and then to show reason
+for the conclusion that it has been, from a Christian point of view,
+satisfactory. I shall not proceed further to urge that the Christian
+apologetic in relation to biology has been successful. A variety of
+opinions may be held on this question, without disturbing the
+conclusion that the movements of readjustment have been beneficial to
+those who remain Christians, and this by making them more Christian
+and not only more liberal. The theologians may sometimes have
+retreated, but there has been an advance of theology. I know that this
+account incurs the charge of optimism. It is not the worst that could
+be made. The influence has been limited in personal range, unequal,
+even divergent, in operation, and accompanied by the appearance of
+waste and mischievous products. The estimate which follows requires
+for due balance a full development of many qualifying considerations.
+For this I lack space, but I must at least distinguish my view from
+the popular one that our difficulties about religion and natural
+science have come to an end.
+
+Concerning the older questions about origins--the origin of the
+world, of species, of man, of reason, conscience, religion--a large
+measure of understanding has been reached by some thoughtful men. But
+meanwhile new questions have arisen, questions about conduct,
+regarding both the reality of morals and the rule of right action for
+individuals and societies. And these problems, still far from
+solution, may also be traced to the influence of Darwin. For they
+arise from the renewed attention to heredity, brought about by the
+search for the causes of variation, without which the study of the
+selection of variations has no sufficient basis.
+
+Even the existing understanding about origins is very far from
+universal. On these points there were always thoughtful men who denied
+the necessity of conflict, and there are still thoughtful men who deny
+the possibility of a truce.
+
+It must further be remembered that the earlier discussion now, as I
+hope to show, producing favourable results, created also for a time
+grave damage, not only in the disturbance of faith and the loss of
+men--a loss not repaired by a change in the currents of debate--but in
+what I believe to be a still more serious respect. I mean the
+introduction of a habit of facile and untested hypothesis in religious
+as in other departments of thought.
+
+Darwin is not responsible for this, but he is in part the cause of it.
+Great ideas are dangerous guests in narrow minds; and thus it has
+happened that Darwin--the most patient of scientific workers, in whom
+hypothesis waited upon research, or if it provisionally outstepped it
+did so only with the most scrupulously careful acknowledgment--has led
+smaller and less conscientious men in natural science, in history, and
+in theology to an over-eager confidence in probable conjecture and a
+loose grip upon the facts of experience. It is not too much to say
+that in many quarters the age of materialism was the least
+matter-of-fact age conceivable, and the age of science the age which
+showed least of the patient temper of inquiry.
+
+I have indicated, as shortly as I could, some losses and dangers
+which in a balanced account of Darwin's influence would be discussed
+at length.
+
+One other loss must be mentioned. It is a defect in our thought which,
+in some quarters, has by itself almost cancelled all the advantages
+secured. I mean the exaggerated emphasis on uniformity or continuity;
+the unwillingness to rest any part of faith or of our practical
+expectation upon anything that from any point of view can be called
+exceptional. The high degree of success reached by naturalists in
+tracing, or reasonably conjecturing, the small beginnings of great
+differences, has led the inconsiderate to believe that anything may in
+time become anything else.
+
+It is true that this exaggeration of the belief in uniformity has
+produced in turn its own perilous reaction. From refusing to believe
+whatever can be called exceptional, some have come to believe whatever
+can be called wonderful.
+
+But, on the whole, the discontinuous or highly various character of
+experience received for many years too little deliberate attention.
+The conception of uniformity which is a necessity of scientific
+description has been taken for the substance of history. We have
+accepted a postulate of scientific method as if it were a conclusion
+of scientific demonstration. In the name of a generalisation which,
+however just on the lines of a particular method, is the prize of a
+difficult exploit of reflexion, we have discarded the direct
+impressions of experience; or, perhaps it is more true to say, we have
+used for the criticism of alleged experiences a doctrine of uniformity
+which is only valid in the region of abstract science. For every
+science depends for its advance upon limitation of attention, upon the
+selection out of the whole content of consciousness of that part or
+aspect which is measurable by the method of the science. Accordingly
+there is a science of life which rightly displays the unity underlying
+all its manifestations. But there is another view of life, equally
+valid, and practically sometimes more important, which recognises the
+immediate and lasting effect of crisis, difference, and revolution.
+Our ardour for the demonstration of uniformity of process and of
+minute continuous change needs to be balanced by a recognition of the
+catastrophic element in experience, and also by a recognition of the
+exceptional significance for us of events which may be perfectly
+regular from an impersonal point of view.
+
+An exorbitant jealousy of miracle, revelation, and ultimate moral
+distinctions has been imported from evolutionary science into
+religious thought. And it has been a damaging influence, because it
+has taken men's attention from facts, and fixed them upon theories.
+
+
+II
+
+With this acknowledgment of important drawbacks, requiring many words
+for their proper description, I proceed to indicate certain results of
+Darwin's doctrine which I believe to be in the long run wholly
+beneficial to Christian thought. These are:
+
+The encouragement in theology of that evolutionary method of
+observation and study, which has shaped all modern research:
+
+The recoil of Christian apologetics towards the ground of religious
+experience, a recoil produced by the pressure of scientific criticism
+upon other supports of faith:
+
+The restatement, or the recovery of ancient forms of statement, of the
+doctrines of Creation and of divine Design in Nature, consequent upon
+the discussion of evolution and of natural selection as its guiding
+factor.
+
+(1) The first of these is quite possibly the most important of all. It
+was well defined in a notable paper read by Dr. Gore, now Bishop of
+Birmingham, to the Church Congress at Shrewsbury in 1896. We have
+learnt a new caution both in ascribing and in denying significance to
+items of evidence, in utterance or in event. There has been, as in
+art, a study of values, which secures perspective and solidity in our
+representation of facts. On the one hand, a given utterance or event
+cannot be drawn into evidence as if all items were of equal
+consequence, like sovereigns in a bag. The question whence and whither
+must be asked, and the particular thing measured as part of a series.
+Thus measured it is not less truly important, but it may be important
+in a lower degree. On the other hand, and for exactly the same reason,
+nothing that is real is unimportant. The "failures" are not mere
+mistakes. We see them, in St. Augustine's words, as "scholar's faults
+which men praise in hope of fruit."
+
+We cannot safely trace the origin of the evolutionistic method to the
+influence of natural science. The view is tenable that theology led
+the way. Probably this is a case of alternate and reciprocal debt.
+Quite certainly the evolutionist method in theology, in Christian
+history and in the estimate of scripture, has received vast
+reinforcement from biology, in which evolution has been the ever
+present and ever victorious conception.
+
+(2) The second effect named is the new willingness of Christian
+thinkers to take definite account of religious experience. This is
+related to Darwin through the general pressure upon religious faith of
+scientific criticism. The great advance of our knowledge of organisms
+has been an important element in the general advance of science. It
+has acted, by the varied requirements of the theory of organisms, upon
+all other branches of natural inquiry, and it held for a long time
+that leading place in public attention which is now occupied by
+speculative physics. Consequently it contributed largely to our
+present estimation of science as the supreme judge in all matters of
+inquiry,[218] to the supposed destruction of mystery and the
+disparagement of metaphysics which marked the last age, as well as to
+the just recommendation of scientific method in branches of learning
+where the direct acquisitions of natural science had no place.
+
+Besides this, the new application of the idea of law and mechanical
+regularity to the organic world seemed to rob faith of a kind of
+refuge. The romantics had, as Berthelot[219] shows, appealed to life
+to redress the judgments drawn from mechanism. Now, in Spencer,
+evolution gave us a vitalist mechanic or mechanical vitalism, and the
+appeal seemed cut off. We may return to this point later when we
+consider evolution; at present I only endeavour to indicate that
+general pressure of scientific criticism which drove men of faith to
+seek the grounds of reassurance in a science of their own; in a method
+of experiment, of observation, of hypothesis checked by known facts.
+It is impossible for me to do more than glance across the threshold of
+this subject. But it is necessary to say that the method is in an
+elementary stage of revival. The imposing success that belongs to
+natural science is absent: we fall short of the unchallengeable
+unanimity of the Biologists on fundamentals. The experimental method
+with its sure repetitions cannot be applied to our subject-matter. But
+we have something like the observational method of palaeontology and
+geographical distribution; and in biology there are still men who
+think that the large examination of varieties by way of geography and
+the search of strata is as truly scientific, uses as genuinely the
+logical method of difference, and is as fruitful in sure conclusions
+as the quasi-chemical analysis of Mendelian laboratory work, of which
+last I desire to express my humble admiration. Religion also has its
+observational work in the larger and possibly more arduous manner.
+
+But the scientific work in religion makes its way through difficulties
+and dangers. We are far from having found the formula of its
+combination with the historical elements of our apologetic. It is
+exposed, therefore, to a damaging fire not only from unspiritualist
+psychology and pathology but also from the side of scholastic dogma.
+It is hard to admit on equal terms a partner to the old undivided rule
+of books and learning. With Charles Lamb, we cry in some distress,
+"must knowledge come to me, if it come at all, by some awkward
+experiment of intuition, and no longer by this familiar process of
+reading?"[220] and we are answered that the old process has an
+imperishable value, only we have not yet made clear its connection
+with other contributions. And all the work is young, liable to be
+drawn into unprofitable excursions, side-tracked by self-deceit and
+pretence; and it fatally attracts, like the older mysticism, the
+curiosity and the expository powers of those least in sympathy with
+it, ready writers who, with all the air of extended research, have
+been content with narrow grounds for induction. There is a danger,
+besides, which accompanies even the most genuine work of this science
+and must be provided against by all its serious students. I mean the
+danger of unbalanced introspection both for individuals and for
+societies; of a preoccupation comparable to our modern social
+preoccupation with bodily health; of reflexion upon mental states not
+accompanied by exercise and growth of the mental powers; the danger of
+contemplating will and neglecting work, of analysing conviction and
+not criticising evidence.
+
+Still, in spite of dangers and mistakes, the work remains full of
+hopeful indications, and, in the best examples,[221] it is truly
+scientific in its determination to know the very truth, to tell what
+we think, not what we think we ought to think,[222] truly scientific
+in its employment of hypothesis and verification, and in growing
+conviction of the reality of its subject-matter through the repeated
+victories of a mastery which advances, like science, in the Baconian
+road of obedience. It is reasonable to hope that progress in this
+respect will be more rapid and sure when religious study enlists more
+men affected by scientific desire and endowed with scientific
+capacity.
+
+The class of investigating minds is a small one, possibly even smaller
+than that of reflecting minds. Very few persons at any period are able
+to find out anything whatever. There are few observers, few
+discoverers, few who even wish to discover truth. In how many
+societies the problems of philology which face every person who speaks
+English are left unattempted! And if the inquiring or the successfully
+inquiring class of minds is small, much smaller, of course, is the
+class of those possessing the scientific aptitude in an eminent
+degree. During the last age this most distinguished class was to a
+very great extent absorbed in the study of phenomena, a study which
+had fallen into arrears. For we stood possessed, in rudiment, of means
+of observation, means for travelling and acquisition, qualifying men
+for a larger knowledge than had yet been attempted. These were now to
+be directed with new accuracy and ardour upon the fabric and behaviour
+of the world of sense. Our debt to the great masters in physical
+science who overtook and almost outstripped the task cannot be
+measured; and, under the honourable leadership of Ruskin, we may all
+well do penance if we have failed "in the respect due to their great
+powers of thought, or in the admiration due to the far scope of their
+discovery."[223] With what miraculous mental energy and divine good
+fortune--as Romans said of their soldiers--did our men of curiosity
+face the apparently impenetrable mysteries of nature! And how natural
+it was that immense accessions of knowledge, unrelated to the
+spiritual facts of life, should discredit Christian faith, by the
+apparent superiority of the new work to the feeble and unprogressive
+knowledge of Christian believers! The day is coming when men of this
+mental character and rank, of this curiosity, this energy and this
+good fortune in investigation, will be employed in opening mysteries
+of a spiritual nature. They will silence with masterful witness the
+over-confident denials of naturalism. They will be in danger of the
+widespread recognition which thirty years ago accompanied every
+utterance of Huxley, Tyndall, Spencer. They will contribute, in spite
+of adulation, to the advance of sober religious and moral science.
+
+And this result will be due to Darwin, first because by raising the
+dignity of natural science, he encouraged the development of the
+scientific mind; secondly because he gave to religious students the
+example of patient and ardent investigation; and thirdly because by
+the pressure of naturalistic criticism the religious have been driven
+to ascertain the causes of their own convictions, a work in which they
+were not without the sympathy of men of science.[224]
+
+In leaving the subject of scientific religious inquiry, I will only
+add that I do not believe it receives any important help--and
+certainly it suffers incidentally much damaging interruption--from the
+study of abnormal manifestations or abnormal conditions of
+personality.
+
+(3) Both of the above effects seem to me of high, perhaps the very
+highest, importance to faith and to thought. But, under the third
+head, I name two which are more directly traceable to the personal
+work of Darwin, and more definitely characteristic of the age in which
+his influence was paramount: viz. the influence of the two conceptions
+of evolution and natural selection upon the doctrine of creation and
+of design respectively.
+
+It is impossible here, though it is necessary for a complete sketch of
+the matter, to distinguish the different elements and channels of this
+Darwinian influence; in Darwin's own writings, in the vigourous
+polemic of Huxley, and strangely enough, but very actually for popular
+thought, in the teaching of the definitely anti-Darwinian evolutionist
+Spencer.
+
+Under the head of the directly and purely Darwinian elements I should
+class as preeminent the work of Wallace and of Bates; for no two sets
+of facts have done more to fix in ordinary intelligent minds a belief
+in organic evolution and in natural selection as its guiding factor
+than the facts of geographical distribution and of protective colour
+and mimicry. The facts of geology were difficult to grasp and the
+public and theologians heard more often of the imperfection than of
+the extent of the geological record. The witness of embryology,
+depending to a great extent upon microscopic work, was and is beyond
+the appreciation of persons occupied in fields of work other than
+biology.
+
+
+III
+
+From the influence in religion of scientific modes of thought we pass
+to the influence of particular biological conceptions. The former
+effect comes by way of analogy, example, encouragement and challenge;
+inspiring or provoking kindred or similar modes of thought in the
+field of theology; the latter by a collision of opinions upon matters
+of fact or conjecture which seem to concern both science and religion.
+
+In the case of Darwinism the story of this collision is familiar, and
+falls under the heads of evolution and natural selection, the doctrine
+of descent with modification, and the doctrine of its guidance or
+determination by the struggle for existence between related varieties.
+These doctrines, though associated and interdependent, and in popular
+thought not only combined but confused, must be considered separately.
+It is true that the ancient doctrine of Evolution, in spite of the
+ingenuity and ardour of Lamarck, remained a dream tantalising the
+intellectual ambition of naturalists, until the day when Darwin made
+it conceivable by suggesting the machinery of its guidance. And,
+further, the idea of natural selection has so effectively opened the
+door of research and stimulated observation in a score of principal
+directions that, even if the Darwinian explanation became one day much
+less convincing than, in spite of recent criticism, it now is, yet its
+passing, supposing it to pass, would leave the doctrine of Evolution
+immeasurably and permanently strengthened. For in the interests of the
+theory of selection, "Für Darwin," as Müller wrote, facts have been
+collected which remain in any case evidence of the reality of descent
+with modification.
+
+But still, though thus united in the modern history of convictions,
+though united and confused in the collision of biological and
+traditional opinion, yet evolution and natural selection must be
+separated in theological no less than in biological estimation.
+Evolution seemed inconsistent with Creation; natural selection with
+Providence and Divine design.
+
+Discussion was maintained about these points for many years and with
+much dark heat. It ranged over many particular topics and engaged
+minds different in tone, in quality, and in accomplishment. There was
+at most times a degree of misconception. Some naturalists attributed
+to theologians in general a poverty of thought which belonged really
+to men of a particular temper or training. The "timid theism"
+discerned in Darwin by so cautious a theologian as Liddon[225] was
+supposed by many biologists to be the necessary foundation of an
+honest Christianity. It was really more characteristic of devout
+_naturalists_ like Philip Henry Gosse, than of religious believers as
+such.[226] The study of theologians more considerable and even more
+typically conservative than Liddon does not confirm the description of
+religious intolerance given in good faith, but in serious ignorance,
+by a disputant so acute, so observant and so candid as Huxley.
+Something hid from each other's knowledge the devoted pilgrims in two
+great ways of thought. The truth may be, that naturalists took their
+view of what creation was from Christian men of science who naturally
+looked in their own special studies for the supports and illustrations
+of their religious belief. Of almost every labourious student it may
+be said: "_Hic ab arte sua non recessit_." And both the believing and
+the denying naturalists, confining habitual attention to a part of
+experience, are apt to affirm and deny with trenchant vigour and
+something of a narrow clearness "_Qui respiciunt ad pauca, de facili
+pronunciant_."[227]
+
+Newman says of some secular teachers that "they persuade the world of
+what is false by urging upon it what is true." Of some early opponents
+of Darwin it might be said by a candid friend that, in all sincerity
+of devotion to truth, they tried to persuade the world of what is true
+by urging upon it what is false. If naturalists took their version of
+orthodoxy from amateurs in theology, some conservative Christians,
+instead of learning what evolution meant to its regular exponents,
+took their view of it from celebrated persons, not of the front rank
+in theology or in thought, but eager to take account of public
+movements and able to arrest public attention.
+
+Cleverness and eloquence on both sides certainly had their share in
+producing the very great and general disturbance of men's minds in the
+early days of Darwinian teaching. But by far the greater part of that
+disturbance was due to the practical novelty and the profound
+importance of the teaching itself, and to the fact that the
+controversy about evolution quickly became much more public than any
+controversy of equal seriousness had been for many generations.
+
+We must not think lightly of that great disturbance because it has, in
+some real sense, done its work, and because it is impossible in days
+of more coolness and light, to recover a full sense of its very real
+difficulties.
+
+Those who would know them better should add to the calm records of
+Darwin[228] and to the story of Huxley's impassioned championship, all
+that they can learn of George Romanes.[229] For his life was absorbed
+in this very struggle and reproduced its stages. It began in a certain
+assured simplicity of biblical interpretation; it went on, through the
+glories and adventures of a paladin in Darwin's train, to the darkness
+and dismay of a man who saw all his most cherished beliefs rendered,
+as he thought, incredible.[230] He lived to find the freer faith for
+which process and purpose are not irreconcilable, but necessary to one
+another. His development, scientific, intellectual and moral, was
+itself of high significance; and its record is of unique value to our
+own generation, so near the age of that doubt and yet so far from it;
+certainly still much in need of the caution and courage by which past
+endurance prepares men for new emergencies. We have little enough
+reason to be sure that in the discussions awaiting us we shall do as
+well as our predecessors in theirs. Remembering their endurance of
+mental pain, their ardour in mental labour, the heroic temper and the
+high sincerity of controversialists on either side, we may well speak
+of our fathers in such words of modesty and self-judgment as Drayton
+used when he sang the victors of Agincourt. The progress of biblical
+study, in the departments of Introduction and Exegesis, resulting in
+the recovery of a point of view anciently tolerated if not prevalent,
+has altered some of the conditions of that discussion. In the years
+near 1858, the witness of Scripture was adduced both by Christian
+advocates and their critics as if unmistakably irreconcilable with
+Evolution.
+
+Huxley[231] found the path of the blameless naturalist everywhere
+blocked by "Moses": the believer in revelation was generally held to
+be forced to a choice between revealed cosmogony and the scientific
+account of origins. It is not clear how far the change in Biblical
+interpretation is due to natural science, and how far to the vital
+movements of theological study which have been quite independent of
+the controversy about species. It belongs to a general renewal of
+Christian movement, the recovery of a heritage. "Special
+Creation"--really a biological rather than a theological
+conception,--seems in its rigid form to have been a recent element
+even in English biblical orthodoxy.
+
+The Middle Ages had no suspicion that religious faith forbad inquiry
+into the natural origination of the different forms of life.
+Bartholomaeus Anglicus, an English Franciscan of the thirteenth
+century, was a mutationist in his way, as Aristotle, "the Philosopher"
+of the Christian Schoolmen, had been in his. So late as the
+seventeenth century, as we learn not only from early proceedings of
+the Royal Society, but from a writer so homely and so regularly pious
+as Walton, the variation of species and "spontaneous" generations had
+no theological bearing, except as instances of that various wonder of
+the world which in devout minds is food for devotion.
+
+It was in the eighteenth century that the harder statement took shape.
+Something in the preciseness of that age, its exaltation of law, its
+cold passion for a stable and measured universe, its cold denial, its
+cold affirmation of the power of God, a God of ice, is the occasion of
+that rigidity of religious thought about the living world which Darwin
+by accident challenged, or rather by one of those movements of genius
+which, Goethe[232] declares, are "elevated above all earthly control."
+
+If religious thought in the eighteenth century was aimed at a fixed
+and nearly finite world of spirit, it followed in all these respects
+the secular and critical lead. "La philosophie réformatrice du
+XVIII^{e} siècle[233] ramenait la nature et la société à des
+mécanismes que la pensée réfléchie peut concevoir et récomposer." In
+fact, religion in a mechanical age is condemned if it takes any but a
+mechanical tone. Butler's thought was too moving, too vital, too
+evolutionary, for the sceptics of his time. In a rationalist,
+encyclopaedic period, religion also must give hard outline to its
+facts, it must be able to display its secret to any sensible man in
+the language used by all sensible men. Milton's prophetic genius
+furnished the eighteenth century, out of the depth of the passionate
+age before it, with the theological tone it was to need. In spite of
+the austere magnificence of his devotion, he gives to smaller souls a
+dangerous lead. The rigidity of Scripture exegesis belonged to this
+stately but imperfectly sensitive mode of thought. It passed away with
+the influence of the older rationalists whose precise denials matched
+the precise and limited affirmations of the static orthodoxy.
+
+I shall, then, leave the specially biblical aspect of the
+debate--interesting as it is and even useful, as in Huxley's
+correspondence with the Duke of Argyll and others in 1892[234]--in
+order to consider without complication the permanent elements of
+Christian thought brought into question by the teaching of evolution.
+
+Such permanent elements are the doctrine of God as Creator of the
+universe, and the doctrine of man as spiritual and unique. Upon both
+the doctrine of evolution seemed to fall with crushing force.
+
+With regard to Man I leave out, acknowledging a grave omission, the
+doctrine of the Fall and of Sin. And I do so because these have not
+yet, as I believe, been adequately treated: here the fruitful reaction
+to the stimulus of evolution is yet to come. The doctrine of sin,
+indeed, falls principally within the scope of that discussion which
+has followed or displaced the Darwinian; and without it the Fall
+cannot be usefully considered. For the question about the Fall is a
+question not merely of origins, but of the interpretation of moral
+facts whose moral reality must first be established.
+
+I confine myself therefore to Creation and the dignity of man.
+
+The meaning of evolution, in the most general terms, is that the
+differentiation of forms is not essentially separate from their
+behaviour and use; that if these are within the scope of study, that
+is also; that the world has taken the form we see by movements not
+unlike those we now see in progress; that what may be called proximate
+origins are continuous in the way of force and matter, continuous in
+the way of life, with actual occurrences and actual characteristics.
+All this has no revolutionary bearing upon the question of ultimate
+origins. The whole is a statement about process. It says nothing to
+metaphysicians about cause. It simply brings within the scope of
+observation or conjecture that series of changes which has given their
+special characters to the different parts of the world we see. In
+particular, evolutionary science aspires to the discovery of the
+process or order of the appearance of life itself: if it were to
+achieve its aim it could say nothing of the cause of this or indeed of
+the most familiar occurrences. We should have become spectators or
+convinced historians of an event which, in respect of its cause and
+ultimate meaning, would be still impenetrable.
+
+With regard to the origin of species, supposing life already
+established, biological science has the well founded hopes and the
+measure of success with which we are all familiar. All this has, it
+would seem, little chance of collision with a consistent theism, a
+doctrine which has its own difficulties unconnected with any
+particular view of order or process. But when it was stated that
+species had arisen by processes through which new species were still
+being made, evolutionism came into collision with a statement,
+traditionally religious, that species were formed and fixed once for
+all and long ago.
+
+What is the theological import of such a statement when it is regarded
+as essential to belief in God? Simply that God's activity, with
+respect to the formation of living creatures, ceased at some point in
+past time.
+
+"God rested" is made the touchstone of orthodoxy. And when, under the
+pressure of the evidences, we found ourselves obliged to acknowledge
+and assert the present and persistent power of God, in the maintenance
+and in the continued formation of "types," what happened was the
+abolition of a time-limit. We were forced only to a bolder claim, to
+a theistic language less halting, more consistent, more thorough in
+its own line, as well as better qualified to assimilate and modify
+such schemes as Von Hartmann's philosophy of the Unconscious--a
+philosophy, by the way, quite intolerant of a merely mechanical
+evolution.[235]
+
+Here was not the retrenchment of an extravagant assertion, but the
+expansion of one which was faltering and inadequate. The traditional
+statement did not need paring down so as to pass the meshes of a new
+and exacting criticism. It was itself a net meant to surround and
+enclose experience; and we must increase its size and close its mesh
+to hold newly disclosed facts of life. The world, which had seemed a
+fixed picture or model, gained first perspective and then solidity and
+movement. We had a glimpse of organic _history_; and Christian thought
+became more living and more assured as it met the larger view of life.
+
+However unsatisfactory the new attitude might be to our critics, to
+Christians the reform was positive. What was discarded was a
+limitation, a negation. The movement was essentially conservative,
+even actually reconstructive. For the language disused was a language
+inconsistent with the definitions of orthodoxy; it set bounds to the
+infinite, and by implication withdrew from the creative rule all such
+processes as could be brought within the descriptions of research. It
+ascribed fixity and finality to that "creature" in which an apostle
+taught us to recognise the birth-struggles of an unexhausted progress.
+It tended to banish mystery from the world we see, and to confine it
+to a remote first age.
+
+In the reformed, the restored, language of religion, Creation became
+again not a link in a rational series to complete a circle of the
+sciences, but the mysterious and permanent relation between the
+infinite and the finite, between the moving changes we know in part,
+and the Power, after the fashion of that observation, unknown, which
+is itself "unmoved all motion's source."[236]
+
+With regard to man it is hardly necessary, even were it possible, to
+illustrate the application of this bolder faith. When the record of
+his high extraction fell under dispute, we were driven to a
+contemplation of the whole of his life, rather than of a part and that
+part out of sight. We remembered again, out of Aristotle, that the
+result of a process interprets its beginnings. We were obliged to read
+the title of such dignity as we may claim, in results and still more
+in aspirations.
+
+Some men still measure the value of great present facts in
+life--reason and virtue and sacrifice--by what a self-disparaged
+reason can collect of the meaner rudiments of these noble gifts. Mr.
+Balfour has admirably displayed the discrepancy, in this view, between
+the alleged origin and the alleged authority of reason. Such an
+argument ought to be used not to discredit the confident reason, but
+to illuminate and dignify its dark beginnings, and to show that at
+every step in the long course of growth a Power was at work which is
+not included in any term or in all the terms of the series.
+
+I submit that the more men know of actual Christian teaching, its
+fidelity to the past, and its sincerity in face of discovery, the more
+certainly they will judge that the stimulus of the doctrine of
+evolution has produced in the long run vigour as well as flexibility
+in the doctrine of Creation and of man.
+
+I pass from Evolution in general to Natural Selection.
+
+The character in religious language which I have for short called
+mechanical was not absent in the argument from design as stated before
+Darwin. It seemed to have reference to a world conceived as fixed. It
+pointed, not to the plastic capacity and energy of living matter, but
+to the fixed adaptation of this and that organ to an unchanging place
+or function.
+
+Mr. Hobhouse has given us the valuable phrase "a niche of organic
+opportunity." Such a phrase would have borne a different sense in
+non-evolutionary thought. In that thought, the opportunity was an
+opportunity for the Creative Power, and Design appeared in the
+preparation of the organism to fit the niche. The idea of the niche
+and its occupant growing together from simpler to more complex mutual
+adjustment was unwelcome to this teleology. If the adaptation was
+traced to the influence, through competition, of the environment, the
+old teleology lost an illustration and a proof. For the cogency of the
+proof in every instance depended upon the absence of explanation.
+Where the process of adaptation was discerned, the evidence of Purpose
+or Design was weak. It was strong only when the natural antecedents
+were not discovered, strongest when they could be declared
+undiscoverable.
+
+Paley's favourite word is "Contrivance"; and for him contrivance is
+most certain where production is most obscure. He points out the
+physiological advantage of the _valvulae conniventes_ to man, and the
+advantage for teleology of the fact that they cannot have been formed
+by "action and pressure." What is not due to pressure may be
+attributed to design, and when a "mechanical" process more subtle than
+pressure was suggested, the case for design was so far weakened. The
+cumulative proof from the multitude of instances began to disappear
+when, in selection, a natural sequence was suggested in which all the
+adaptations might be reached by the motive power of life, and
+especially when, as in Darwin's teaching, there was full recognition
+of the reactions of life to the stimulus of circumstance. "The
+organism fits the niche," said the teleologist, "because the Creator
+formed it so as to fit." "The organism fits the niche," said the
+naturalist, "because unless it fitted it could not exist." "It was
+fitted to survive," said the theologian. "It survives because it
+fits," said the selectionist. The two forms of statement are not
+incompatible; but the new statement, by provision of an ideally
+universal explanation of process, was hostile to a doctrine of purpose
+which relied upon evidences always exceptional however numerous.
+Science persistently presses on to find the universal machinery of
+adaptation in this planet; and whether this be found in selection, or
+in direct-effect, or in vital reactions resulting in large changes, or
+in a combination of these and other factors, it must always be opposed
+to the conception of a Divine Power here and there but not everywhere
+active.
+
+For science, the Divine must be constant, operative everywhere and in
+every quality and power, in environment and in organism, in stimulus
+and in reaction, in variation and in struggle, in hereditary
+equilibrium, and in "the unstable state of species"; equally present
+on both sides of every strain, in all pressures and in all
+resistances, in short in the general wonder of life and the world. And
+this is exactly what the Divine Power must be for religious faith.
+
+The point I wish once more to make is that the necessary readjustment
+of teleology, so as to make it depend upon the contemplation of the
+whole instead of a part, is advantageous quite as much to theology as
+to science. For the older view failed in courage. Here again our
+theism was not sufficiently theistic.
+
+Where results seemed inevitable, it dared not claim them as God-given.
+In the argument from Design it spoke not of God in the sense of
+theology, but of a Contriver, immensely, not infinitely wise and good,
+working within a world, the scene, rather than the ever dependent
+outcome, of His Wisdom; working in such emergencies and opportunities
+as occurred, by forces not altogether within His control, towards an
+end beyond Himself. It gave us, instead of the awful reverence due to
+the Cause of all substance and form, all love and wisdom, a
+dangerously detached appreciation of an ingenuity and benevolence
+meritorious in aim and often surprisingly successful in contrivance.
+
+The old teleology was more useful to science than to religion, and
+the design-naturalists ought to be gratefully remembered by
+Biologists. Their search for evidences led them to an eager study of
+adaptations and of minute forms, a study such as we have now an
+incentive to in the theory of Natural Selection. One hardly meets with
+the same ardour in microscopical research until we come to modern
+workers. But the argument from Design was never of great importance to
+faith. Still, to rid it of this character was worth all the stress and
+anxiety of the gallant old war. If Darwin had done nothing else for
+us, we are to-day deeply in his debt for this. The world is not less
+venerable to us now, not less eloquent of the causing mind, rather
+much more eloquent and sacred. But our wonder is not that "the
+underjaw of the swine works under the ground" or in any or all of
+those particular adaptations which Paley collected with so much skill,
+but that a purpose transcending, though resembling, our own purposes,
+is everywhere manifest; that what we live in is a whole, mutually
+sustaining, eventful and beautiful, where the "dead" forces feed the
+energies of life, and life sustains a stranger existence, able in some
+real measure to contemplate the whole, of which, mechanically
+considered, it is a minor product and a rare ingredient. Here, again,
+the change was altogether positive. It was not the escape of a vessel
+in a storm with loss of spars and rigging, not a shortening of sail to
+save the masts and make a port of refuge. It was rather the emergence
+from narrow channels to an open sea. We had propelled the great ship,
+finding purchase here and there for slow and uncertain movement. Now,
+in deep water, we spread large canvas to a favouring breeze.
+
+The scattered traces of design might be forgotten or obliterated. But
+the broad impression of Order became plainer when seen at due distance
+and in sufficient range of effect, and the evidence of love and wisdom
+in the universe could be trusted more securely for the loss of the
+particular calculation of their machinery.
+
+Many other topics of faith are affected by modern biology. In some of
+these we have learnt at present only a wise caution, a wise
+uncertainty. We stand before the newly unfolded spectacle of
+suffering, silenced; with faith not scientifically reassured but still
+holding fast certain other clues of conviction. In many important
+topics we are at a loss. But in others, and among them those I have
+mentioned, we have passed beyond this negative state and find faith
+positively strengthened and more fully expressed.
+
+We have gained also a language and a habit of thought more fit for the
+great and dark problems that remain, less liable to damaging
+conflicts, equipped for more rapid assimilation of knowledge. And by
+this change biology itself is a gainer. For, relieved of fruitless
+encounters with popular religion, it may advance with surer aim along
+the path of really scientific life-study which was reopened for modern
+men by the publication of _The Origin of Species_.
+
+Charles Darwin regretted that, in following science, he had not done
+"more direct good"[237] to his fellow-creatures. He has, in fact,
+rendered substantial service to interests bound up with the daily
+conduct and hopes of common men; for his work has led to improvements
+in the preaching of the Christian faith.
+
+FOOTNOTES:
+
+[Footnote 218: F. R. Tennant: "The Being of God in the light of
+Physical Science," in _Essays on some theological questions of the
+day_. London, 1905.]
+
+[Footnote 219: _Evolutionisme et Platonisme_, pp. 45, 46, 47. Paris,
+1908.]
+
+[Footnote 220: _Essays of Elia_, "New Year's Eve," p. 41; Ainger's
+edition. London, 1899.]
+
+[Footnote 221: Such an example is given in Baron F. von Hügel's
+recently finished book, the result of thirty years' research: _The
+Mystical Element of Religion, as studied in Saint Catherine of Genoa
+and her Friends_. London, 1908.]
+
+[Footnote 222: G. Tyrrell, in _Mediaevalism_, has a chapter which is
+full of the important _moral_ element in a scientific attitude. "The
+only infallible guardian of truth is the spirit of truthfulness."
+_Mediaevalism_, p. 182, London, 1908.]
+
+[Footnote 223: _Queen of the Air_, Preface, p. vii. London, 1906.]
+
+[Footnote 224: The scientific rank of its writer justifies the
+insertion of the following letter from the late Sir John
+Burdon-Sanderson to me. In the lecture referred to I had described the
+methods of Professor Moseley in teaching Biology as affording a
+suggestion of the scientific treatment of religion.
+
+OXFORD,
+
+_April 30, 1902_.
+
+DEAR SIR:
+
+     I feel that I must express to you my thanks for the
+     discourse which I had the pleasure of listening to yesterday
+     afternoon.
+
+     I do not mean to say that I was able to follow all that you
+     said as to the identity of Method in the two fields of
+     Science and Religion, but I recognise that the "mysticism"
+     of which you spoke gives us the only way by which the two
+     fields can be brought into relation.
+
+     Among much that was memorable, nothing interested me more
+     than what you said of Moseley.
+
+     No one, I am sure, knew better than you the value of his
+     teaching and in what that value consisted.
+
+Yours faithfully,
+
+J. BURDON-SANDERSON.
+
+]
+
+[Footnote 225: H. P. Liddon, _The Recovery of S. Thomas_; a sermon
+preached in St. Paul's, London, on April 23rd, 1882 (the Sunday after
+Darwin's death).]
+
+[Footnote 226: Dr. Pusey (_Unscience not Science adverse to Faith_,
+1878) writes: "The questions as to 'species,' of what variations the
+animal world is capable, whether the species be more or fewer, whether
+accidental variations may become hereditary ... and the like,
+naturally fall under the province of science. In all these questions
+Mr. Darwin's careful observations gained for him a deserved
+approbation and confidence."]
+
+[Footnote 227: Aristotle, in Bacon, quoted by Newman in his _Idea of a
+University_, p. 78. London, 1873.]
+
+[Footnote 228: _Life and Letters_ and _More Letters of Charles
+Darwin._]
+
+[Footnote 229: _Life and Letters_, London, 1896. _Thoughts on
+Religion_, London, 1895. _Candid Examination of Theism_, London,
+1878.]
+
+[Footnote 230: "Never in the history of man has so terrific a calamity
+befallen the race as that which all who look may now (viz. in
+consequence of the scientific victory of Darwin) behold advancing as a
+deluge black with destruction, resistless in might, uprooting our most
+cherished hopes, engulphing our most precious creed, and burying our
+highest life in mindless destruction."--_A Candid Examination of
+Theism_, p. 51.]
+
+[Footnote 231: _Science and Christian Tradition._ London, 1904.]
+
+[Footnote 232: "No productiveness of the highest kind ... is in the
+power of anyone."--_Conversations of Goethe with Eckermann and Soret_.
+London, 1850.]
+
+[Footnote 233: Berthelot, _Evolutionisme et Platonisme_, Paris, 1908,
+p. 45.]
+
+[Footnote 234: _Times_, 1892, _passim._]
+
+[Footnote 235: See Von Hartmann's _Wahrheit und Irrthum in
+Darwinismus_. Berlin, 1875.]
+
+[Footnote 236: Hymn of the Church--
+
+    Rerum Deus tenax vigor,
+    Immotus in te permanens.
+
+]
+
+[Footnote 237: _Life and Letters_, Vol. III. p. 359.]
+
+
+
+
+IX
+
+DARWINISM AND HISTORY
+
+BY J. B. BURY, LITT.D., LL.D.
+
+_Regius Professor of Modern History in the University of Cambridge_
+
+
+1. Evolution, and the principles associated with the Darwinian theory,
+could not fail to exert a considerable influence on the studies
+connected with the history of civilised man. The speculations which
+are known as "philosophy of history," as well as the sciences of
+anthropology, ethnography, and sociology (sciences which though they
+stand on their own feet are for the historian auxiliary), have been
+deeply affected by these principles. Historiographers, indeed, have
+with few exceptions made little attempt to apply them; but the growth
+of historical study in the nineteenth century has been determined and
+characterised by the same general principle which has underlain the
+simultaneous developments of the study of nature, namely the _genetic
+idea_. The "historical" conception of nature, which has produced the
+history of the solar system, the story of the earth, the genealogies
+of telluric organisms, and has revolutionised natural science, belongs
+to the same order of thought as the conception of human history as a
+continuous, genetic, causal process--a conception which has
+revolutionised historical research and made it scientific. Before
+proceeding to consider the application of evolutional principles, it
+will be pertinent to notice the rise of this new view.
+
+2. With the Greeks and Romans history had been either a descriptive
+record or had been written in practical interests. The most eminent
+of the ancient historians were pragmatical; that is, they regarded
+history as an instructress in statesmanship, or in the art of war, or
+in morals. Their records reached back such a short way, their
+experience was so brief, that they never attained to the conception of
+continuous process, or realised the significance of time; and they
+never viewed the history of human societies as a phenomenon to be
+investigated for its own sake. In the middle ages there was still less
+chance of the emergence of the ideas of progress and development. Such
+notions were excluded by the fundamental doctrines of the dominant
+religion which bounded and bound men's minds. As the course of history
+was held to be determined from hour to hour by the arbitrary will of
+an extra cosmic person, there could be no self-contained causal
+development, only a dispensation imposed from without. And as it was
+believed that the world was within no great distance from the end of
+this dispensation, there was no motive to take much interest in
+understanding the temporal, which was to be only temporary.
+
+The intellectual movements of the fifteenth and sixteenth centuries
+prepared the way for a new conception, but it did not emerge
+immediately. The historians of the Renaissance period simply reverted
+to the ancient pragmatical view. For Machiavelli, exactly as for
+Thucydides and Polybius, the use of studying history was instruction
+in the art of politics. The Renaissance itself was the appearance of a
+new culture, different from anything that had gone before; but at the
+time men were not conscious of this; they saw clearly that the
+traditions of classical antiquity had been lost for a long period, and
+they were seeking to revive them, but otherwise they did not perceive
+that the world had moved, and that their own spirit, culture, and
+conditions were entirely unlike those of the thirteenth century. It
+was hardly till the seventeenth century that the presence of a new
+age, as different from the middle ages as from the ages of Greece and
+Rome, was fully realised. It was then that the triple division of
+ancient, medieval, and modern was first applied to the history of
+western civilisation. Whatever objections may be urged against this
+division, which has now become almost a category of thought, it marks
+a most significant advance in man's view of his own past. He has
+become conscious of the immense changes in civilisation which have
+come about slowly in the course of time, and history confronts him
+with a new aspect. He has to explain how those changes have been
+produced, how the transformations were effected. The appearance of
+this problem was almost simultaneous with the rise of rationalism, and
+the great historians and thinkers of the eighteenth century, such as
+Montesquieu, Voltaire, Gibbon, attempted to explain the movement of
+civilisation by purely natural causes. These brilliant writers
+prepared the way for the genetic history of the following century. But
+in the spirit of the _Aufklärung_, that eighteenth-century
+Enlightenment to which they belonged, they were concerned to judge all
+phenomena before the tribunal of reason; and the apotheosis of
+"reason" tended to foster a certain superior _a priori_ attitude,
+which was not favourable to objective treatment and was incompatible
+with a "historical sense." Moreover the traditions of pragmatical
+historiography had by no means disappeared.
+
+3. In the first quarter of the nineteenth century the meaning of
+genetic history was fully realised. "Genetic" perhaps is as good a
+word as can be found for the conception which in this century was
+applied to so many branches of knowledge in the spheres both of nature
+and of mind. It does not commit us to the doctrine proper of
+evolution, nor yet to any teleological hypothesis such as is implied
+in "progress." For history it meant that the present condition of the
+human race is simply and strictly the result of a causal series (or
+set of causal series)--a continuous succession of changes, where each
+state arises causally out of the preceding; and that the business of
+historians is to trace this genetic process, to explain each change,
+and ultimately to grasp' the complete development of the life of
+humanity. Three influential writers, who appeared at this stage and
+helped to initiate a new period of research, may specially be
+mentioned. Ranke in 1824 definitely repudiated the pragmatical view
+which ascribes to history the duties of an instructress, and with no
+less decision renounced the function, assumed by the historians of the
+_Aufklärung_, to judge the past; it was his business, he said, merely
+to show how things really happened. Niebuhr was already working in the
+same spirit and did more than any other writer to establish the
+principle that historical transactions must be related to the ideas
+and conditions of their age. Savigny about the same time founded the
+"historical school" of law. He sought to show that law was not the
+creation of an enlightened will, but grew out of custom and was
+developed by a series of adaptations and rejections, thus applying the
+conception of evolution. He helped to diffuse the notion that all the
+institutions of a society or a nation are as closely interconnected as
+the parts of a living organism.
+
+4. The conception of the history of man as a causal development meant
+the elevation of historical inquiry to the dignity of a science. Just
+as the study of bees cannot become scientific so long as the student's
+interest in them is only to procure honey or to derive moral lessons
+from the labours of "the little busy bee," so the history of human
+societies cannot become the object of pure scientific investigation so
+long as man estimates its value in pragmatical scales. Nor can it
+become a science until it is conceived as lying entirely within a
+sphere in which the law of cause and effect has unreserved and
+unrestricted dominion. On the other hand, once history is envisaged as
+a causal process, which contains within itself the explanation of the
+development of man from his primitive state to the point which he has
+reached, such a process necessarily becomes the object of scientific
+investigation and the interest in it is scientific curiosity.
+
+At the same time, the instruments were sharpened and refined. Here
+Wolf, a philologist with historical instinct, was a pioneer. His
+_Prolegomena_ to Homer (1795) announced new modes of attack.
+Historical investigation was soon transformed by the elaboration of
+new methods.
+
+5. "Progress" involves a judgment of value, which is not involved in
+the conception of history as a genetic process. It is also an idea
+distinct from that of evolution. Nevertheless it is closely related to
+the ideas which revolutionised history at the beginning of the last
+century; it swam into men's ken simultaneously; and it helped
+effectively to establish the notion of history as a continuous process
+and to emphasise the significance of time. Passing over earlier
+anticipations, I may point to a _Discours_ of Turgot (1750), where
+history is presented as a process in which "the total mass of the
+human race" "marches continually though sometimes slowly to an ever
+increasing perfection." That is a clear statement of the conception
+which Turgot's friend Condorcet elaborated in the famous work,
+published in 1795, _Esquisse d'un tableau historique des progrès de
+l'esprit humain_. This work first treated with explicit fulness the
+idea to which a leading role was to fall in the ideology of the
+nineteenth century. Condorcet's book reflects the triumphs of the
+_Tiers état_, whose growing importance had also inspired Turgot; it
+was the political changes in the eighteenth century which led to the
+doctrine, emphatically formulated by Condorcet, that the masses are
+the most important element in the historical process. I dwell on this
+because, though Condorcet had no idea of evolution, the predominant
+importance of the masses was the assumption which made it possible to
+apply evolutional principles to history. And it enabled Condorcet
+himself to maintain that the history of civilisation, a progress still
+far from being complete, was a development conditioned by general
+laws.
+
+6. The assimilation of society to an organism, which was a governing
+notion in the school of Savigny, and the conception of progress,
+combined to produce the idea of an organic development, in which the
+historian has to determine the central principle or leading character.
+This is illustrated by the apotheosis of democracy in Tocqueville's
+_Démocratie en Amérique_, where the theory is maintained that "the
+gradual and progressive development of equality is at once the past
+and the future of the history of men." The same two principles are
+combined in the doctrine of Spencer (who held that society is an
+organism, though he also contemplated its being what he calls a
+"super-organic aggregate"),[238] that social evolution is a
+progressive change from militarism to industrialism.
+
+7. The idea of development assumed another form in the speculations of
+German idealism. Hegel conceived the successive periods of history as
+corresponding to the ascending phases or ideas in the self-evolution
+of his Absolute Being. His _Lectures on the Philosophy of History_
+were published in 1837 after his death. His philosophy had a
+considerable effect, direct and indirect, on the treatment of history
+by historians, and although he was superficial and unscientific
+himself in dealing with historical phenomena, he contributed much
+towards making the idea of historical development familiar. Ranke was
+influenced, if not by Hegel himself, at least by the Idealistic
+philosophies of which Hegel's was the greatest. He was inclined to
+conceive the stages in the process of history as marked by
+incarnations, as it were, of ideas, and sometimes speaks as if the
+ideas were independent forces, with hands and feet. But while Hegel
+determined his ideas by _a priori_ logic, Ranke obtained his by
+induction--by a strict investigation of the phenomena; so that he was
+scientific in his method and work, and was influenced by Hegelian
+prepossessions only in the kind of significance which he was disposed
+to ascribe to his results. It is to be noted that the theory of Hegel
+implied a judgment of value; the movement was a progress towards
+perfection.
+
+8. In France, Comte approached the subject from a different side, and
+exercised, outside Germany, a far wider influence than Hegel. The 4th
+volume of his _Cours de philosophie positive_, which appeared in 1839,
+created sociology and treated history as a part of this new science,
+namely as "social dynamics." Comte sought the key for unfolding
+historical development, in what he called the social-psychological
+point of view, and he worked out the two ideas which had been
+enunciated by Condorcet: that the historian's attention should be
+directed not, as hitherto, principally to eminent individuals, but to
+the collective behaviour of the masses, as being the most important
+element in the process; and that, as in nature, so in history, there
+are general laws, necessary and constant, which condition the
+development. The two points are intimately connected, for it is only
+when the masses are moved into the foreground that regularity,
+uniformity, and law can be conceived as applicable. To determine the
+social-psychological laws which have controlled the development is,
+according to Comte, the task of sociologists and historians.
+
+9. The hypothesis of general laws operative in history was carried
+further in a book which appeared in England twenty years later and
+exercised an influence in Europe far beyond its intrinsic merit,
+Buckle's _History of Civilisation in England_ (1857-61). Buckle owed
+much to Comte, and followed him, or rather outdid him, in regarding
+intellect as the most important factor conditioning the upward
+development of man, so that progress, according to him, consisted in
+the victory of the intellectual over the moral laws.
+
+10. The tendency of Comte and Buckle to assimilate history to the
+sciences of nature by reducing it to general "laws," derived stimulus
+and plausibility from the vista offered by the study of statistics,
+in which the Belgian Quetelet, whose book _Sur l'homme_ appeared in
+1835, discerned endless possibilities. The astonishing uniformities
+which statistical inquiry disclosed led to the belief that it was only
+a question of collecting a sufficient amount of statistical material,
+to enable us to predict how a given social group will act in a
+particular case. Bourdeau, a disciple of this school, looks forward to
+the time when historical science will become entirely quantitative.
+The actions of prominent individuals, which are generally considered
+to have altered or determined the course of things, are obviously not
+amenable to statistical computation or explicable by general laws.
+Thinkers like Buckle sought to minimise their importance or explain
+them away.
+
+11. These indications may suffice to show that the new efforts to
+interpret history which marked the first half of the nineteenth
+century were governed by conceptions closely related to those which
+were current in the field of natural science and which resulted in the
+doctrine of evolution. The genetic principle, progressive development,
+general laws, the significance of time, the conception of society as
+an organic aggregate, the metaphysical theory of history as the
+self-evolution of spirit,--all these ideas show that historical
+inquiry had been advancing independently on somewhat parallel lines to
+the sciences of nature. It was necessary to bring this out in order to
+appreciate the influence of Darwinism.
+
+12. In the course of the dozen years which elapsed between the
+appearances of _The Origin of Species_ (observe that the first volume
+of Buckle's work was published just two years before) and of _The
+Descent of Man_ (1871), the hypothesis of Lamarck that man is the
+co-descendant with other species of some lower extinct form was
+admitted to have been raised to the rank of an established fact by
+most thinkers whose brains were not working under the constraint of
+theological authority.
+
+One important effect of the discovery of this fact (I am not speaking
+now of the Darwinian explanation) was to assign to history a definite
+place in the coordinated whole of knowledge, and relate it more
+closely to other sciences. It had indeed a defined logical place in
+systems such as Hegel's and Comte's; but Darwinism certified its
+standing convincingly and without more ado. The prevailing doctrine
+that man was created _ex abrupto_ had placed history in an isolated
+position, disconnected with the sciences of nature. Anthropology,
+which deals with the animal _anthropos_, now comes into line with
+zoology, and brings it into relation with history.[239] Man's
+condition at the present day is the result of a series of
+transformations, going back to the most primitive phase of society,
+which is the ideal (unattainable) beginning of history. But that
+beginning had emerged without any breach of continuity from a
+development which carries us back to a quadrimane ancestor, still
+further back (according to Darwin's conjecture) to a marine animal of
+the ascidian type, and then through remoter periods to the lowest form
+of organism. It is essential in this theory that though links have
+been lost there was no break in the gradual development; and this
+conception of a continuous progress in the evolution of life,
+resulting in the appearance of uncivilised Anthropos, helped to
+reinforce, and increase a belief in, the conception of the history of
+civilised Anthropos as itself also a continuous progressive
+development.
+
+13. Thus the diffusion of the Darwinian theory of the origin of man,
+by emphasising the idea of continuity and breaking down the barriers
+between the human and animal kingdoms, has had an important effect in
+establishing the position of history among the sciences which deal
+with telluric development. The perspective of history is merged in a
+larger perspective of development. As one of the objects of biology is
+to find the exact steps in the genealogy of man from the lowest
+organic form, so the scope of history is to determine the stages in
+the unique causal series from the most rudimentary to the present
+state of human civilisation.
+
+It is to be observed that the interest in historical research implied
+by this conception need not be that of Comte. In the Positive
+Philosophy history is part of sociology; the interest in it is to
+discover the sociological laws. In the view of which I have just
+spoken, history is permitted to be an end in itself; the
+reconstruction of the genetic process is an independent interest. For
+the purpose of the reconstruction, sociology, as well as physical
+geography, biology, psychology, is necessary; the sociologist and the
+historian play into each other's hands; but the object of the former
+is to establish generalisations; the aim of the latter is to trace in
+detail a singular causal sequence.
+
+14. The success of the evolutional theory helped to discredit the
+assumption or at least the invocation of transcendent causes.
+Philosophically of course it is compatible with theism, but historians
+have for the most part desisted from invoking the naive conception of
+a "god in history" to explain historical movements. A historian may be
+a theist; but, so far as his work is concerned, this particular belief
+is otiose. Otherwise indeed (as was remarked above) history could not
+be a science; for with a _deus ex machina_ who can be brought on the
+stage to solve difficulties scientific treatment is a farce. The
+transcendent element had appeared in a more subtle form through the
+influence of German philosophy. I noticed how Ranke is prone to refer
+to ideas as if they were transcendent existences manifesting
+themselves in the successive movements of history. It is intelligible
+to speak of certain ideas as controlling, in a given period,--for
+instance, the idea of nationality; but from the scientific point of
+view, such ideas have no existence outside the minds of individuals
+and are purely psychical forces; and a historical "idea," if it does
+not exist in this form, is merely a way of expressing a synthesis of
+the historian himself.
+
+15. From the more general influence of Darwinism on the place of
+history in the system of human knowledge, we may turn to the influence
+of the principles and methods by which Darwin explained development.
+It had been recognised even by ancient writers (such as Aristotle and
+Polybius) that physical circumstances (geography, climate) were
+factors conditioning the character and history of a race or society.
+In the sixteenth century Bodin emphasised these factors, and many
+subsequent writers took them into account. The investigations of
+Darwin, which brought them into the foreground, naturally promoted
+attempts to discover in them the chief key to the growth of
+civilisation. Comte had expressly denounced the notion that the
+biological methods of Lamarck could be applied to social man. Buckle
+had taken account of natural influences, but had relegated them to a
+secondary plane, compared with psychological factors. But the
+Darwinian theory made it tempting to explain the development of
+civilisation in terms of "adaptation to environment," "struggle for
+existence," "natural selection," "survival of the fittest," etc.[240]
+
+The operation of these principles cannot be denied. Man is still an
+animal, subject to zoological as well as mechanical laws. The dark
+influence of heredity continues to be effective; and psychical
+development had begun in lower organic forms,--perhaps with life
+itself. The organic and the social struggles for existence are
+manifestations of the same principle. Environment and climatic
+influence must be called in to explain not only the differentiation of
+the great racial sections of humanity, but also the varieties within
+these sub-species and, it may be, the assimilation of distinct
+varieties. Ritter's _Anthropogeography_ has opened a useful line of
+research. But on the other hand, it is urged that, in explaining the
+course of history, these principles do not take us very far, and that
+it is chiefly for the primitive ultra-prehistoric period that they can
+account for human development. It may be said that, so far as concerns
+the actions and movements of men which are the subject of recorded
+history, physical environment has ceased to act mechanically, and in
+order to affect their actions must affect their wills first; and that
+this psychical character of the causal relations substantially alters
+the problem. The development of human societies, it may be argued,
+derives a completely new character from the dominance of the conscious
+psychical element, creating as it does new conditions (inventions,
+social institutions, etc.) which limit and counteract the operation of
+natural selection, and control and modify the influence of physical
+environment. Most thinkers agree now that the chief clews to the
+growth of civilisation must be sought in the psychological sphere.
+Imitation, for instance, is a principle which is probably more
+significant for the explanation of human development than natural
+selection. Darwin himself was conscious that his principles had only a
+very restricted application in this sphere, as is evident from his
+cautious and tentative remarks in the 5th chapter of his _Descent of
+Man_. He applied natural selection to the growth of the intellectual
+faculties and of the fundamental social instincts, and also to the
+differentiation of the great races or "sub-species" (Caucasian,
+African, etc.) which differ in anthropological character.[241]
+
+16. But if it is admitted that the governing factors which concern the
+student of social development are of the psychical order, the
+preliminary success of natural science in explaining organic evolution
+by general principles encouraged sociologists to hope that social
+evolution could be explained on general principles also. The idea of
+Condorcet, Buckle, and others, that history could be assimilated to
+the natural sciences was powerfully reinforced, and the notion that
+the actual historical process, and every social movement involved in
+it, can be accounted for by sociological generalisations, so-called
+"laws," is still entertained by many, in one form or another.
+Dissentients from this view do not deny that the generalisations at
+which the sociologist arrives by the comparative method, by the
+analysis of social factors, and by psychological deduction may be an
+aid to the historian; but they deny that such uniformities are laws or
+contain an explanation of the phenomena. They can point to the element
+of chance coincidence. This element must have played a part in the
+events of organic evolution, but it has probably in a larger measure
+helped to determine events in social evolution. The collision of two
+unconnected sequences may be fraught with great results. The sudden
+death of a leader or a marriage without issue, to take simple cases,
+has again and again led to permanent political consequences. More
+emphasis is laid on the decisive actions of individuals, which cannot
+be reduced under generalisations and which deflect the course of
+events. If the significance of the individual will had been
+exaggerated to the neglect of the collective activity of the social
+aggregate before Condorcet, his doctrine tended to eliminate as
+unimportant the roles of prominent men, and by means of this
+elimination it was possible to found sociology. But it may be urged
+that it is patent on the face of history that its course has
+constantly been shaped and modified by the wills of individuals,[242]
+which are by no means always the expression of the collective will;
+and that the appearance of such personalities at the given moments is
+not a necessary outcome of the conditions and cannot be deduced. Nor
+is there any proof that, if such and such an individual had not been
+born, some one else would have arisen to do what he did. In some cases
+there is no reason to think that what happened need ever have come to
+pass. In other cases, it seems evident that the actual change was
+inevitable, but in default of the man who initiated and guided it, it
+might have been postponed, and, postponed or not, might have borne a
+different cachet. I may illustrate by an instance which has just come
+under my notice. Modern painting was founded by Giotto, and the
+Italian expedition of Charles VIII, near the close of the sixteenth
+century, introduced into France the fashion of imitating Italian
+painters. But for Giotto and Charles VIII, French painting might have
+been very different. It may be said that "if Giotto had not appeared,
+some other great imitator would have played a role analogous to his,
+and that without Charles VIII there would have been the commerce with
+Italy, which in the long run would have sufficed to place France in
+relation with Italian artists. But the equivalent of Giotto might have
+been deferred for a century and probably would have been different;
+and commercial relations would have required ages to produce the
+_rayonnement imitatif_ of Italian art in France, which the expedition
+of the royal adventurer provoked in a few years."[243] Instances
+furnished by political history are simply endless. Can we conjecture
+how events would have moved if the son of Philip of Macedon had been
+an incompetent? The aggressive action of Prussia which astonished
+Europe in 1740 determined the subsequent history of Germany; but that
+action was anything but inevitable; it depended entirely on the
+personality of Frederick the Great.
+
+Hence it may be argued that the action of individual wills is a
+determining and disturbing factor, too significant and effective to
+allow history to be grasped by sociological formulae. The types and
+general forms of development which the sociologist attempts to
+disengage can only assist the historian in understanding the actual
+course of events. It is in the special domains of economic history and
+_Culturgeschichte_ which have come to the front in modern times that
+generalisation is most fruitful, but even in these it may be contended
+that it furnishes only partial explanations.
+
+17. The truth is that Darwinism itself offers the best illustration of
+the insufficiency of general laws to account for historical
+development. The part played by coincidence, and the part played by
+individuals--limited by, and related to, general social
+conditions--render it impossible to deduce the course of the past
+history of man or to predict the future. But it is just the same with
+organic development. Darwin (or any other zoologist) could not deduce
+the actual course of evolution from general principles. Given an
+organism and its environment, he could not show that it must evolve
+into a more complex organism of a definite predetermined type; knowing
+what it has evolved into, he could attempt to discover and assign the
+determining causes. General principles do not account for a particular
+sequence; they embody necessary conditions; but there is a chapter of
+accidents too. It is the same in the case of history.
+
+18. Among the evolutional attempts to subsume the course of history under
+general syntheses, perhaps the most important is that of Lamprecht, whose
+"kulturhistorische" attempt to discover and assign the determining causes.
+German history, exhibits the (indirect) influence of the Comtist school. It
+is based upon psychology, which, in his views, holds among the sciences of
+mind (_Geisteswissenschaften_) the same place (that of a
+_Grundwissenschaft_) which mechanics holds among the sciences of nature.
+History, by the same comparison, corresponds to biology, and, according to
+him, it can only become scientific if it is reduced to general concepts
+(_Begriffe_). Historical movements and events are of a psychical character,
+and Lamprecht conceives a given phase of civilisation as "a collective
+psychical condition (_seelischer Gesamtzustand_)" controlling the period,
+"a diapason which penetrates all psychical phenomena and thereby all
+historical events of the time."[244] He has worked out a series of such
+phases, "ages of changing psychical diapason," in his _Deutsche
+Geschichte_, with the aim of showing that all the feelings and actions of
+each age can be explained by the diapason; and has attempted to prove that
+these diapasons are exhibited in other social developments, and are
+consequently not singular but typical. He maintains further that these ages
+succeed each other in a definite order; the principle being that the
+collective psychical development begins with the homogeneity of all the
+individual members of a society and, through heightened psychical activity,
+advances in the form of a continually increasing differentiation of the
+individuals (this is akin to the Spencerian formula). This process,
+evolving psychical freedom from psychical constraint, exhibits a series of
+psychical phenomena which define successive periods of civilisation. The
+process depends on two simple principles, that no idea can disappear
+without leaving behind it an effect or influence, and that all psychical
+life, whether in a person or a society, means change, the acquisition of
+new mental contents. It follows that the new have to come to terms with the
+old, and this leads to a synthesis which determines the character of a new
+age. Hence the ages of civilisation are defined as the "highest concepts
+for subsuming without exception all psychical phenomena of the development
+of human societies, that is, of all historical events."[245] Lamprecht
+deduces the idea of a special historical science, which might be called
+"historical ethnology," dealing with the ages of civilisation, and bearing
+the same relation to (descriptive or narrative) history as ethnology to
+ethnography. Such a science obviously corresponds to Comte's social
+dynamics, and the comparative method, on which Comte laid so much emphasis,
+is the principal instrument of Lamprecht.
+
+19. I have dwelt on the fundamental ideas of Lamprecht, because they
+are not yet widely known in England, and because his system is the
+ablest product of the sociological school of historians. It carries
+the more weight as its author himself is a historical specialist, and
+his historical syntheses deserve the most careful consideration. But
+there is much in the process of development which on such assumptions
+is not explained, especially the initiative of individuals. Historical
+development does not proceed in a right line, without the choice of
+diverging. Again and again, several roads are open to it, of which it
+chooses one--why? On Lamprecht's method, we may be able to assign the
+conditions which limit the psychical activity of men at a particular
+stage of evolution, but within those limits the individual has so many
+options, such a wide room for moving, that the definition of those
+conditions, the "psychical diapasons," is only part of the explanation
+of the particular development. The heel of Achilles in all historical
+speculations of this class has been the role of the individual.
+
+The increasing prominence of economic history has tended to encourage
+the view that history can be explained in terms of general concepts or
+types. Marx and his school based their theory of human development on
+the conditions of production, by which, according to them, all social
+movements and historical changes are entirely controlled. The leading
+part which economic factors play in Lamprecht's system is significant,
+illustrating the fact that economic changes admit most readily this
+kind of treatment, because they have been less subject to direction or
+interference by individual pioneers.
+
+Perhaps it may be thought that the conception of _social environment_
+(essentially psychical), on which Lamprecht's "psychical diapasons"
+depend, is the most valuable and fertile conception that the historian
+owes to the suggestion of the science of biology--the conception of
+all particular historical actions and movements as (1) related to and
+conditioned by the social environment, and (2) gradually bringing
+about a transformation of that environment. But no given
+transformation can be proved to be necessary (predetermined). And
+types of development do not represent laws; their meaning and value
+lie in the help they may give to the historian, in investigating a
+certain period of civilisation, to enable him to discover the
+inter-relations among the diverse features which it presents. They
+are, as some one has said, an instrument of heuretic method.
+
+20. The man engaged in special historical researches--which have been
+pursued unremittingly for a century past, according to scientific
+methods of investigating evidence (initiated by Wolf, Niebuhr,
+Ranke)--have for the most part worked on the assumptions of genetic
+history or at least followed in the footsteps of those who fully
+grasped the genetic point of view. But their aim has been to collect
+and sift evidence, and determine particular facts; comparatively few
+have given serious thought to the lines of research and the
+speculations which have been considered in this paper. They have been
+reasonably shy of compromising their work by applying theories which
+are still much debated and immature. But historiography cannot
+permanently evade the questions raised by these theories. One may
+venture to say that no historical change or transformation will be
+fully understood until it is explained how social environment acted on
+the individual components of the society (both immediately and by
+heredity), and how the individuals reacted upon their environment. The
+problem is psychical, but it is analogous to the main problem of the
+biologist.
+
+FOOTNOTES:
+
+[Footnote 238: A society presents suggestive analogies with an
+organism, but it certainly is not an organism, and sociologists who
+draw inferences from the assumption of its organic nature must fall
+into error. A vital organism and a society are radically distinguished
+by the fact that the individual components of the former, namely the
+cells, are morphologically as well as functionally differentiated,
+whereas the individuals which compose a society are morphologically
+homogeneous and only functionally differentiated. The resemblances and
+the differences are worked out in E. de Majewski's striking book, _La
+Science de la Civilisation_. Paris. 1908.]
+
+[Footnote 239: It is to be observed that history is (not only
+different in scope but) not co-extensive with anthropology _in time_.
+For it deals only with the development of man in societies, whereas
+anthropology includes in its definition the proto-anthropic period
+when _anthropos_ was still non-social, whether he lived in herds like
+the chimpanzee, or alone like the male ourang-outang. (It has been
+well shown by Majewski that congregations--herds, flocks, packs,
+&c.--of animals are not _societies_; the characteristic of a society
+is differentiation of function. Bee hives, ant hills, may be called
+quasi-societies; but in their case the classes which perform distinct
+functions are morphologically different.)]
+
+[Footnote 240: Recently O. Seeck has applied these principles to the
+decline of Graeco-Roman civilisation in his _Untergang der antiken
+Welt_, 2 vols., Berlin, 1895, 1901.]
+
+[Footnote 241: Darwinian formulae may be suggestive by way of analogy.
+For instance, it is characteristic of social advance that a multitude
+of inventions, schemes and plans are framed which are never carried
+out, similar to, or designed for the same end as, an invention or plan
+which is actually adopted because it has chanced to suit better the
+particular conditions of the hour (just as the works accomplished by
+an individual statesman, artist or savant are usually only a residue
+of the numerous projects conceived by his brain). This process in
+which so much abortive production occurs is analogous to elimination
+by natural selection.]
+
+[Footnote 242: We can ignore here the metaphysical question of
+freewill and determinism. For the character of the individual's brain
+depends in any case on ante-natal accidents and coincidences, and so
+it may be said that the role of individuals ultimately depends on
+chance,--the accidental coincidence of independent sequences.]
+
+[Footnote 243: I have taken this example from G. Tarde's _La logique
+sociale_ (p. 403), Paris, 1904, where it is used for quite a different
+purpose.]
+
+[Footnote 244: _Die kulturhistorische Methode_, Berlin, 1900, p. 26.]
+
+[Footnote 245: _Ibid._ pp. 28, 29.]
+
+
+
+
+X
+
+DARWINISM AND SOCIOLOGY
+
+BY C. BOUGLÉ
+
+_Professor of Social Philosophy in the University of Toulouse and
+Deputy-Professor at the Sorbonne, Paris_
+
+
+How has our conception of social phenomena, and of their history, been
+affected by Darwin's conception of Nature and the laws of its
+transformation? To what extent and in what particular respects have
+the discoveries and hypotheses of the author of _The Origin of
+Species_ aided the efforts of those who have sought to construct a
+science of society?
+
+To such a question it is certainly not easy to give any brief or
+precise answer. We find traces of Darwinism almost everywhere.
+Sociological systems differing widely from each other have laid claim
+to its authority; while, on the other hand, its influence has often
+made itself felt only in combination with other influences. The
+Darwinian thread is worked into a hundred patterns along with other
+threads.
+
+To deal with the problem, we must, it seems, first of all distinguish
+the more general conclusions in regard to the evolution of living
+beings, which are the outcome of Darwinism, from the particular
+explanations it offers of the ways and means by which that evolution
+is effected. That is to say, we must, as far as possible, estimate
+separately the influence of Darwin as an evolutionist and Darwin as a
+selectionist.
+
+The nineteenth century, said Cournot, has witnessed a mighty effort to
+"réintégrer l'homme dans la nature." From divers quarters there has
+been a methodical reaction against the persistent dualism of the
+Cartesian tradition, which was itself the unconscious heir of the
+Christian tradition. Even the philosophy of the eighteenth century,
+materialistic as were for the most part the tendencies of its leaders,
+seemed to revere man as a being apart, concerning whom laws might be
+formulated _à priori_. To bring him down from his pedestal there was
+needed the marked predominance of positive researches wherein no
+account was taken of the "pride of man." There can be no doubt that
+Darwin has done much to familiarise us with this attitude. Take for
+instance the first part of _The Descent of Man_: it is an accumulation
+of typical facts, all tending to diminish the distance between us and
+our brothers, the lower animals. One might say that the naturalist had
+here taken as his motto, "Whosoever shall exalt himself shall be
+abased; and he that shall humble himself shall be exalted." Homologous
+structures, the survival in man of certain organs of animals, the
+rudiments in the animal of certain human faculties, a multitude of
+facts of this sort, led Darwin to the conclusion that there is no
+ground for supposing that the "king of the universe" is exempt from
+universal laws. Thus belief in the _imperium in imperio_ has been, as
+it were, whittled away by the progress of the naturalistic spirit,
+itself continually strengthened by the conquests of the natural
+sciences. The tendency may, indeed, drag the social sciences into
+overstrained analogies, such, for instance, as the assimilation of
+societies to organisms. But it will, at least, have had the merit of
+helping sociology to shake off the pre-conception that the groups
+formed by men are artificial, and that history is completely at the
+mercy of chance. Some years before the appearance of _The Origin of
+Species_, August Comte had pointed out the importance, as regards the
+unification of positive knowledge, of the conviction that the social
+world, the last refuge of spiritualism, is itself subject to
+determinism. It cannot be doubted that the movement of thought which
+Darwin's discoveries promoted contributed to the spread of this
+conviction, by breaking down the traditional barrier which cut man off
+from Nature.
+
+But Nature, according to modern naturalists, is no immutable thing: it
+is rather perpetual movement, continual progression. Their discoveries
+batter a breach directly into the Aristotelian notion of species; they
+refuse to see in the animal world a collection of immutable types,
+distinct from all eternity, and corresponding, as Cuvier said, to so
+many particular thoughts of the Creator. Darwin especially
+congratulated himself upon having been able to deal this doctrine the
+_coup de grâce_: immutability is, he says, his chief enemy; and he is
+concerned to show--therein following up Lyell's work--that everything
+in the organic world, as in the inorganic, is explained by insensible
+but incessant transformations. "Nature makes no leaps"--"Nature knows
+no gaps": these two _dicta_ form, as it were, the two landmarks
+between which Darwin's idea of transformation is worked out. That is
+to say, the development of Darwinism is calculated to further the
+application of the philosophy of Becoming to the study of human
+institutions.
+
+The progress of the natural sciences thus brings unexpected
+reinforcements to the revolution which the progress of historical
+discipline had begun. The first attempt to constitute an actual
+science of social phenomena--that, namely, of the economists--had
+resulted in laws which were called natural, and which were believed to
+be eternal and universal, valid for all times and all places. But this
+perpetuality, brother, as Knies said, of the immutability of the old
+zoology, did not long hold out against the ever-swelling tide of the
+historical movement. Knowledge of the transformations that had taken
+place in language, of the early phases of the family, of religion, of
+property, had all favoured the revival of the Heraclitean view:
+[Greek: panta rei]. As to the categories of political economy, it was
+soon to be recognised, as by Lasalle, that they too are only
+historical. The philosophy of history, moreover, gave expression
+under various forms to the same tendency. Hegel declares that "all
+that is real is rational," but at the same time he shows that all that
+is real is ephemeral, and that for history there is nothing fixed
+beneath the sun. It is this sense of universal evolution that Darwin
+came with fresh authority to enlarge. It was in the name of biological
+facts themselves that he taught us to see only slow metamorphoses in
+the history of institutions, and to be always on the outlook for
+survivals side by side with rudimentary forms. Anyone who reads
+_Primitive Culture_, by Tylor,--a writer closely connected with
+Darwin--will be able to estimate the services which these cardinal
+ideas were to render to the social sciences when the age of
+comparative research had succeeded to that of _à priori_ construction.
+
+Let us note, moreover, that the philosophy of Becoming in passing through
+the Darwinian biology became, as it were, filtered; it got rid of those
+traces of finalism, which, under different forms, it had preserved through
+all the systems of German Romanticism. Even in Herbert Spencer, it has been
+plausibly argued, one can detect something of that sort of mystic
+confidence in forces spontaneously directing life, which forms the very
+essence of those systems. But Darwin's observations were precisely
+calculated to render such an hypothesis futile. At first people may have
+failed to see this; and we call to mind the ponderous sarcasms of Flourens
+when he objected to the theory of Natural Selection that it attributed to
+nature a power of free choice. "Nature endowed with will! That was the
+final error of last century; but the nineteenth no longer deals in
+personifications."[246] In fact Darwin himself put his readers on their
+guard against the metaphors he was obliged to use. The processes by which
+he explains the survival of the fittest are far from affording any
+indication of the design of some transcendent breeder. Nor, if we look
+closely, do they even imply immanent effort in the animal; the sorting out
+can be brought about mechanically, simply by the action of the environment.
+In this connection Huxley could with good reason maintain that Darwin's
+originality consisted in showing how harmonies which hitherto had been
+taken to imply the agency of intelligence and will could be explained
+without any such intervention. So, when later on, objective sociology
+declares that, even when social phenomena are in question, all finalist
+preconceptions must be distrusted if a science is to be constituted, it is
+to Darwin that its thanks are due; he had long been clearing paths for it
+which lay well away from the old familiar road trodden by so many theories
+of evolution.
+
+This anti-finalist doctrine, when fully worked out, was, moreover,
+calculated to aid in the needful dissociation of two notions: that of
+evolution and that of progress. In application to society these had
+long been confounded; and, as a consequence, the general idea seemed
+to be that only one type of evolution was here possible. Do we not
+detect such a view in Comte's sociology, and perhaps even in Herbert
+Spencer's? Whoever, indeed, assumes an end for evolution is naturally
+inclined to think that only one road leads to that end. But those
+whose minds the Darwinian theory has enlightened are aware that the
+transformations of living beings depend primarily upon their
+conditions, and that it is these conditions which are the agents of
+selection from among individual variations. Hence, it immediately
+follows that transformations are not necessarily improvements. Here,
+Darwin's thought hesitated. Logically his theory proves, as Ray
+Lankester pointed out, that the struggle for existence may have as its
+outcome degeneration as well as amelioration: evolution may be
+regressive as well as progressive. Then, too--and this is especially
+to be borne in mind--each species takes its good where it finds it,
+seeks its own path and survives as best it can. Apply this notion to
+society and you arrive at the theory of multilinear evolution.
+Divergencies will no longer surprise you. You will be forewarned not
+to apply to all civilisations the same measure of progress, and you
+will recognise that types of evolution may differ just as social
+species themselves differ. Have we not here one of the conceptions
+which mark off sociology proper from the old philosophy of history?
+
+       *       *       *       *       *
+
+But if we are to estimate the influence of Darwinism upon sociological
+conceptions, we must not dwell only upon the way in which Darwin
+impressed the general notion of evolution upon the minds of thinkers.
+We must go into details. We must consider the influence of the
+particular theories by which he explained the mechanism of this
+evolution. The name of the author of _The Origin of Species_ has been
+especially attached, as everyone knows, to the doctrines of "natural
+selection" and of "struggle for existence," completed by the notion of
+"individual variation." These doctrines were turned to account by very
+different schools of social philosophy. Pessimistic and optimistic,
+aristocratic and democratic, individualistic and socialistic systems
+were to war with each other for years by casting scraps of Darwinism
+at each other's heads.
+
+It was the spectacle of human contrivance that suggested to Darwin his
+conception of natural selection. It was in studying the methods of
+pigeon breeders that he divined the processes by which nature, in the
+absence of design, obtains analogous results in the differentiation of
+types. As soon as the importance of artificial selection in the
+transformation of species of animals was understood, reflection
+naturally turned to the human species, and the question arose, How far
+do men observe, in connection with themselves, those laws of which
+they make practical application in the case of animals? Here we come
+upon one of the ideas which guided the researches of Gallon, Darwin's
+cousin. The author of _Inquiries into Human Faculty and its
+Development_,[247] has often expressed his surprise that, considering
+all the precautions taken, for example, in the breeding of horses,
+none whatever are taken in the breeding of the human species. It seems
+to be forgotten that the species suffers when the "fittest" are not
+able to perpetuate their type. Ritchie, in his _Darwinism and
+Politics_[248] reminds us of Darwin's remark that the institution of
+the peerage might be defended on the ground that peers, owing to the
+prestige they enjoy, are enabled to select as wives "the most
+beautiful and charming women out of the lower ranks."[249] But, says
+Galton, it is as often as not "heiresses" that they pick out, and
+birth statistics seem to show that these are either less robust or
+less fecund than others. The truth is that considerations continue to
+preside over marriage which are entirely foreign to the improvement of
+type, much as this is a condition of general progress. Hence the
+importance of completing Odin's and De Candolle's statistics which are
+designed to show how characters are incorporated in organisms, how
+they are transmitted, how lost, and according to what law eugenic,
+elements depart from the mean or return to it.
+
+But thinkers do not always content themselves with undertaking merely
+the minute researches which the idea of Selection suggests. They are
+eager to defend this or that thesis. In the name of this idea certain
+social anthropologists have recast the conception of the process of
+civilisation, and have affirmed that Social Selection generally works
+against the trend of Natural Selection. Vacher de Lapouge--following
+up an observation by Broca on the point--enumerates the various
+institutions, or customs, such as the celibacy of priests and military
+conscription, which cause elimination or sterilisation of the bearers
+of certain superior qualities, intellectual or physical. In a more
+general way he attacks the democratic movement, a movement, as P.
+Bourget says, which is "anti-physical" and contrary to the natural
+laws of progress; though it has been inspired "by the dreams of that
+most visionary of all centuries, the eighteenth."[250] The "Equality"
+which levels down and mixes (justly condemned, he holds, by the Comte
+de Gobineau), prevents the aristocracy of the blond dolichocephales
+from holding the position and playing the part which, in the interests
+of all, should belong to them. Otto Ammon, in his _Natural Selection
+in Man_, and in _The Social Order and its Natural Bases_,[251]
+defended analogous doctrines in Germany; setting the curve
+representing frequency of talent over against that of income, he
+attempted to show that all democratic measures which aim at promoting
+the rise in the social scale of the talented are useless, if not
+dangerous; that they only increase the panmixia, to the great
+detriment of the species and of society.
+
+Among the aristocratic theories which Darwinism has thus inspired we
+must reckon that of Nietzsche. It is well known that in order to
+complete his philosophy he added biological studies to his
+philological; and more than once in his remarks upon the _Wille zur
+Macht_ he definitely alludes to Darwin; though it must be confessed
+that it is generally in order to proclaim the insufficiency of the
+processes by which Darwin seeks to explain the genesis of species.
+Nevertheless, Nietzsche's mind is completely possessed by an ideal of
+Selection. He, too, has a horror of panmixia. The naturalists'
+conception of "the fittest" is joined by him to that of the "hero" of
+romance to furnish a basis for his doctrine of the Superman. Let us
+hasten to add, moreover, that at the very moment when support was
+being sought in the theory of Selection for the various forms of the
+aristocratic doctrine, those same forms were being battered down on
+another side by means of that very theory. Attention was drawn to the
+fact that by virtue of the laws which Darwin himself had discovered
+isolation leads to etiolation. There is a risk that the privilege
+which withdraws the privileged elements of Society from competition
+will cause them to degenerate. In fact, Jacoby in his _Studies in
+Selection, in connexion with Heredity in Man_,[252] concludes that
+"sterility, mental debility, premature death and, finally, the
+extinction of the stock were not specially and exclusively the fate of
+sovereign dynasties; all privileged classes, all families in
+exclusively elevated positions share the fate of reigning families,
+although in a minor degree and in direct proportion to the loftiness
+of their social standing. From the mass of human beings spring
+individuals, families, races, which tend to raise themselves above the
+common level; painfully they climb the rugged heights, attain the
+summits of power, of wealth, of intelligence, of talent, and then, no
+sooner are they there than they topple down and disappear in gulfs of
+mental and physical degeneracy." The demographical researches of
+Hansen[253] (following up and completing Dumont's) tended, indeed, to
+show that urban as well as feudal aristocracies, burgher classes as
+well as noble castes, were liable to become effete. Hence it might
+well be concluded that the democratic movement, operating as it does
+to break down class barriers, was promoting instead of impeding human
+selection.
+
+       *       *       *       *       *
+
+So we see that, according to the point of view, very different
+conclusions have been drawn from the application of the Darwinian idea
+of Selection to human society. Darwin's other central idea, closely
+bound up with this, that, namely, of the "struggle for existence" also
+has been diversely utilised. But discussion has chiefly centered upon
+its signification. And while some endeavour to extend its application
+to everything, we find others trying to limit its range. The
+conception of a "struggle for existence" has in the present day been
+taken up into the social sciences from natural science, and adopted.
+But originally it descended from social science to natural. Darwin's
+law is, as he himself said, only Malthus' law generalised and extended
+to the animal world: a growing disproportion between the supply of
+food and the number of the living is the fatal order whence arises the
+necessity of universal struggle, a struggle which, to the great
+advantage of the species, allows only the best equipped individuals to
+survive. Nature is regarded by Huxley as an immense arena where all
+living beings are gladiators.[254]
+
+Such a generalisation was well adapted to feed the stream of
+pessimistic thought; and it furnished to the apologists of war, in
+particular, new arguments, weighted with all the authority which in
+these days attaches to scientific deliverances. If people no longer
+say, as Bonald did, and Moltke after him, that war is a providential
+fact, they yet lay stress on the point that it is a natural fact. To
+the peace party Dragomirov's objection is urged that its attempts are
+contrary to the fundamental laws of nature, and that no sea wall can
+hold against breakers that come with such gathered force.
+
+But in yet another quarter Darwinism was represented as opposed to
+philanthropic intervention. The defenders of the orthodox political
+economy found in it support for their tenets. Since in the organic
+world universal struggle is the condition of progress, it seemed
+obvious that free competition must be allowed to reign unchecked in
+the economic world. Attempts to curb it were in the highest degree
+imprudent. The spirit of Liberalism here seemed in conformity with the
+trend of nature: in this respect, at least, contemporary naturalism,
+offspring of the discoveries of the nineteenth century, brought
+reinforcements to the individualist doctrine, begotten of the
+speculations of the eighteenth: but only, it appeared, to turn mankind
+away for ever from humanitarian dreams. Would those whom such
+conclusions repelled be content to oppose to nature's imperatives
+only the protests of the heart? There were some who declared, like
+Brunetière, that the laws in question, valid though they might be for
+the animal kingdom, were not applicable to the human. And so a return
+was made to the classic dualism. This indeed seems to be the line that
+Huxley took, when, for instance, he opposed to the cosmic process an
+ethical process which was its reverse.
+
+But the number of thinkers whom this antithesis does not satisfy grows
+daily. Although the pessimism which claims authorisation from Darwin's
+doctrines is repugnant to them, they still are unable to accept the
+dualism which leaves a gulf between man and nature. And their
+endeavour is to link the two by showing that while Darwin's laws
+obtain in both kingdoms, the conditions of their application are not
+the same: their forms, and, consequently, their results, vary with the
+varying mediums in which the struggle of living beings takes place,
+with the means these beings have at disposal, with the ends even which
+they propose to themselves.
+
+Here we have the explanation of the fact that among determined
+opponents of war partisans of the "struggle for existence" can be
+found: there are disciples of Darwin in the peace party. Novicow, for
+example, admits the "_combat universel_" of which Le Dantec[255]
+speaks; but he remarks that at different stages of evolution, at
+different stages of life the same weapons are not necessarily
+employed. Struggles of brute force, armed hand to hand conflicts, may
+have been a necessity in the early phases of human societies.
+Nowadays, although competition may remain inevitable and
+indispensable, it can assume milder forms. Economic rivalries,
+struggles between intellectual influences, suffice to stimulate
+progress: the processes which these admit are, in the actual state of
+civilisation, the only ones which attain their end without waste, the
+only ones logical. From one end to the other of the ladder of life,
+struggle is the order of the day; but more and more as the higher
+rungs are reached, it takes on characters which are proportionately
+more "humane."
+
+Reflections of this kind permit the introduction into the economic
+order of limitations to the doctrine of "laisser faire, laisser
+passer." This appeals, it is said, to the example of nature where
+creatures, left to themselves, struggle without truce and without
+mercy; but the fact is forgotten that upon industrial battlefields the
+conditions are different. The competitors here are not left simply to
+their natural energies: they are variously handicapped. A rich store
+of artificial resources exists in which some participate and others do
+not. The sides then are unequal; and as a consequence the result of
+the struggle is falsified. "In the animal world," said De
+Laveleye,[256] criticising Spencer, "the fate of each creature is
+determined by its individual qualities; whereas in civilised societies
+a man may obtain the highest position and the most beautiful wife
+because he is rich and well-born, although he may be ugly, idle or
+improvident; and then it is he who will perpetuate the species. The
+wealthy man, ill constituted, incapable, sickly, enjoys his riches and
+establishes his stock under the protection of the laws." Haycraft in
+England and Jentsch in Germany have strongly emphasised these
+"anomalies," which nevertheless are the rule. That is to say that even
+from a Darwinian point of view all social reforms can readily be
+justified which aim at diminishing, as Wallace said, inequalities at
+the start.
+
+But we can go further still. Whence comes the idea that all measures
+inspired by the sentiment of solidarity are contrary to Nature's
+trend? Observe her carefully, and she will not give lessons only in
+individualism. Side by side with the struggle for existence do we not
+find in operation what Lanessan calls "association for existence."
+Long ago, Espinas had drawn attention to "societies of animals,"
+temporary or permanent, and to the kind of morality that arose in
+them. Since then, naturalists have often insisted upon the importance
+of various forms of symbiosis. Kropotkin in _Mutual Aid_ has chosen
+to enumerate many examples of altruism furnished by animals to
+mankind. Geddes and Thomson went so far as to maintain that "Each of
+the greater steps of progress is in fact associated with an increased
+measure of subordination of individual competition to reproductive or
+social ends, and of interspecific competition to co-operative,
+association."[257] Experience shows, according to Geddes, that the
+types which are fittest to surmount great obstacles are not so much
+those who engage in the fiercest competitive struggle for existence,
+as those who contrive to temper it. From all these observations there
+resulted, along with a limitation of Darwinian pessimism, some
+encouragement for the aspirations of the collectivists.
+
+And Darwin himself would, doubtless, have subscribed to these
+rectifications. He never insisted, like his rival, Wallace, upon the
+necessity of the solitary struggle of creatures in a state of nature,
+each for himself and against all. On the contrary, in _The Descent of
+Man_, he pointed out the serviceableness of the social instincts, and
+corroborated Bagehot's statements when the latter, applying laws of
+physics to politics, showed the great advantage societies derived from
+intercourse and communion. Again, the theory of sexual evolution which
+makes the evolution of types depend increasingly upon preferences,
+judgments, mental factors, surely offers something to qualify what
+seems hard and brutal in the theory of natural selection.
+
+But, as often happens with disciples, the Darwinians had out-Darwined
+Darwin. The extravagances of social Darwinism provoked a useful
+reaction; and thus people were led to seek, even in the animal
+kingdom, for facts of solidarity which would serve to justify humane
+effort.
+
+       *       *       *       *       *
+
+On quite another line, however, an attempt has been made to connect
+socialist tendencies with Darwinian principles. Marx and Darwin have
+been confronted; and writers have undertaken to show that the work of
+the German philosopher fell readily into line with that of the English
+naturalist and was a development of it. Such has been the endeavour of
+Ferri in Italy and of Woltmann in Germany, not to mention others. The
+founders of "scientific socialism" had, moreover, themselves thought
+of this reconciliation. They make more than one allusion to Darwin in
+works which appeared after 1859. And sometimes they use his theory to
+define by contrast their own ideal. They remark that the capitalist
+system, by giving free course to individual competition, ends indeed
+in a _bellum omnium contra omnes_; and they make it clear that
+Darwinism, thus understood, is as repugnant to them as to Dühring.
+
+But it is at the scientific and not at the moral point of view that
+they place themselves when they connect their economic history with
+Darwin's work. Thanks to this unifying hypothesis, they claim to have
+constructed--as Marx does in his preface to _Das Kapital_--a veritable
+natural history of social evolution. Engels speaks in praise of his
+friend Marx as having discovered the true mainspring of history hidden
+under the veil of idealism and sentimentalism, and as having
+proclaimed in the _primum vivere_ the inevitableness of the struggle
+for existence. Marx himself, in _Das Kapital_, indicated another
+analogy when he dwelt upon the importance of a general technology for
+the explanation of this psychology:--a history of tools which would be
+to social organs what Darwinism is to the organs of animal species.
+And the very importance they attach to tools, to apparatus, to
+machines, abundantly proves that neither Marx nor Engels were likely
+to forget the special characters which mark off the human world from
+the animal. The former always remains to a great extent an artificial
+world. Inventions change the face of its institutions. New modes of
+production revolutionise not only modes of government, but modes even
+of collective thought. Therefore it is that the evolution of society
+is controlled by laws special to it, of which the spectacle of nature
+offers no suggestion.
+
+If, however, even in this special sphere, it can still be urged that
+the evolution of the material conditions of society is in accord with
+Darwin's theory, it is because the influence of the methods of
+production is itself to be explained by the incessant strife of the
+various classes with each other. So that in the end Marx, like Darwin,
+finds the source of all progress is in struggle. Both are grandsons of
+Heraclitus:--[Greek: polemos patêr pantôn]. It sometimes happens, in
+these days, that the doctrine of revolutionary socialism is contrasted
+as rude and healthy with what may seem to be the enervating tendency
+of "solidarist" philanthropy: the apologists of the doctrine then
+pride themselves above all upon their faithfulness to Darwinian
+principles.
+
+       *       *       *       *       *
+
+So far we have been mainly concerned to show the use that social
+philosophies have made of the Darwinian laws for practical purposes:
+in order to orientate society towards their ideals each school tries
+to show that the authority of natural science is on its side. But even
+in the most objective of theories, those which systematically make
+abstraction of all political tendencies in order to study the social
+reality in itself, traces of Darwinism are readily to be found.
+
+Let us take for example Durkheim's theory of Division of Labour.[258]
+The conclusions he derives from it are that whenever professional
+specialisation causes multiplication of distinct branches of activity,
+we get organic solidarity--implying differences--substituted for
+mechanical solidarity, based upon likenesses. The umbilical cord, as
+Marx said, which connects the individual consciousness with the
+collective consciousness is cut. The personality becomes more and more
+emancipated. But on what does this phenomenon, so big with
+consequences, itself depend? The author goes to social morphology for
+the answer: it is, he says, the growing density of population which
+brings with it this increasing differentiation of activities. But,
+again, why? Because the greater density, in thrusting men up against
+each other, augments the intensity of their competition for the means
+of existence; and for the problems which society thus has to face
+differentiation of functions presents itself as the gentlest solution.
+
+Here one sees that the writer borrows directly from Darwin.
+Competition is at its maximum between similars, Darwin had declared;
+different species, not laying claim to the same food, could more
+easily coexist. Here lay the explanation of the fact that upon the
+same oak hundreds of different insects might be found. Other things
+being equal, the same applies to society. He who finds some unadopted
+specialty possesses a means of his own for getting a living. It is by
+this division of their manifold tasks that men contrive not to crush
+each other. Here we obviously have a Darwinian law serving as
+intermediary in the explanation of that progress of division of labour
+which itself explains so much in the social evolution.
+
+And we might take another example, at the other end of the series of
+sociological systems. G. Tarde is a sociologist with the most
+pronounced anti-naturalistic views. He has attempted to show that all
+application of the laws of natural science to society is misleading.
+In his _Opposition Universelle_ he has directly combatted all forms of
+sociological Darwinism. According to him the idea that the evolution
+of society can be traced on the same plan as the evolution of species
+is chimerical. Social evolution is at the mercy of all kinds of
+inventions, which by virtue of the laws of imitation modify, through
+individual to individual, through neighbourhood to neighbourhood, the
+general state of those beliefs and desires which are the only
+"quantities" whose variation matters to the sociologist. But, it may
+be rejoined, that however psychical the forces may be, they are none
+the less subject to Darwinian laws. They compete with each other; they
+struggle for the mastery of minds. Between types of ideas, as between
+organic forms, selection operates. And though it may be that these
+types are ushered into the arena by unexpected discoveries, we yet
+recognise in the psychological accidents, which Tarde places at the
+base of everything, near relatives of those small accidental
+variations upon which Darwin builds. Thus, accepting Tarde's own
+representations, it is quite possible to express in Darwinian terms,
+with the necessary transpositions, one of the most idealistic
+sociologies that have ever been constructed.
+
+These few examples suffice. They enable us to estimate the extent of
+the field of influence of Darwinism. It affects sociology not only
+through the agency of its advocates but through that of its opponents.
+The questions to which it has given rise have proved no less fruitful
+than the solutions it has suggested. In short, few doctrines, in the
+history of social philosophy, will have produced on their passage a
+finer crop of ideas.
+
+FOOTNOTES:
+
+[Footnote 246: P. Flourens, _Examen du Livre de M. Darwin sur
+l'Origine des Espèces_, p. 53, Paris, 1864. See also Huxley,
+"Criticisms on the _Origin of Species," Collected Essays_, Vol. II, p.
+102, London, 1902.]
+
+[Footnote 247: _Inquiries into Human Faculty_, pp. 1, 2, 3 sq.,
+London, 1883.]
+
+[Footnote 248: _Darwinism and Politics_, pp. 9, 22, London, 1889.]
+
+[Footnote 249: _Life and Letters of Charles Darwin_, II. p. 385.]
+
+[Footnote 250: V. de Lapouge, _Les Sélections sociales_, p. 259,
+Paris, 1896.]
+
+[Footnote 251: _Die natärliche Auslese beim Menschen_, Jena, 1893; _Du
+Gesellschaftsordnung und ihre natürlichen Grundlagen. Entwurf einer
+Sozialanthropologie_, Jena, 1896.]
+
+[Footnote 252: _Etudes sur la Sélection dans ses rapports avec
+l'hérédité chez l'homme_, Paris, p. 481, 1881.]
+
+[Footnote 253: _Die drei Bevölkerungsstufen_, Munich, 1889.]
+
+[Footnote 254: _Evolution and Ethics_, p. 200; _Collected Essays_,
+Vol. IX, London, 1894.]
+
+[Footnote 255: _Les Luttes entre Sociétés humaines et leurs phases
+successives_, Paris, 1893.]
+
+[Footnote 256: _Le socialisme contemporain_, p. 384 (6th edit.),
+Paris, 1891.]
+
+[Footnote 257: Geddes and Thomson, _The Evolution of Sex_, p. 311,
+London, 1889.]
+
+[Footnote 258: _De la Division du Travail social_, Paris. 1893.]
+
+
+
+
+INDEX
+
+
+_Abraxas grossulariata_, 100
+
+Acquired characters, transmission of, 20, 28, 42, 94, 120, 149, 171, 173
+
+_Acraea johnstoni_, 290
+[Transcriber's Note: No such page number or reference seen.]
+
+Adaptation, 24, 27, 34, 39, 42-45, 50, 58, 79-86, 106, 107
+
+Adloff, 140
+
+Alexander, 217
+
+Ameghino, 132, 138
+
+Ammon, O., Works of, 271
+
+_Anaea divina_, 69
+
+Anglicus, Bartholomaeus, 237
+
+Ankyroderma, 40
+
+Anomma, 44
+
+Anthropops, 132
+
+Ants, modifications of, 43-46, 51
+
+Ardigò, 207, 208
+
+Argyll, Huxley and the Duke of, 238
+
+Aristotle, 3, 237, 240
+
+Avenarius, 211
+
+
+Bacon, on mutability of species, 4, 5
+
+Baehr, von, on Cytology, 99
+
+Bain, 194
+
+Baldwin, J. M., 53, Foot Note 165
+
+Balfour, A. J., 241
+
+Barratt, 217
+
+Bates, H. W., on Mimicry, 70, 76
+  --232
+
+BATESON, W., on _Heredity and
+Variation in Modern Lights_, 87-110
+  --on discontinuous evolution, 30
+
+Bathmism, 14
+
+Bells (Sir Charles) _Anatomy of Expression_, 177
+
+Bentham, Jeremy, 217, 218
+
+Bergson, H., 208
+
+Berkeley, 200
+
+Berthelot, 228
+
+Bickford, E., experiments on degeneration by, 52
+
+Biophores, 47
+
+Blumenbach, 89
+
+Bodin, 256
+
+Bonald, on war, 273
+
+Bonnet, 6
+
+BOUGLÉ, C., on _Darwinism and Sociology_, 264-280
+
+Bourdeau, 253
+
+Bourget, P., 270
+
+Boutroux, 208
+
+Brassica, hybrids of, 106
+
+_Brassica Napus_, 106
+
+Broca, 137, 270
+
+Brock, on Kant, Foot Note 6
+
+Brunetière, 274
+
+Bruno, on Evolution, 4
+
+Buch, von, 15
+
+Buckle, 252, 253, 256, 258
+
+Buffon, 6-15, 21, 88
+
+Burdon-Sanderson, J., letter from, Foot Note 224
+
+BURY, J. B., on _Darwinism and History_, 246-263
+
+Butler, Samuel, 9, Foot Note 17, Foot Note 57, Foot Note 61, 94, Foot Note 66, 107
+
+Butterflies, mimicry in, 65-83
+  --sexual characters in, 59-63
+
+
+Cabanis, 201
+
+Candolle, de, 270
+
+Carneri, 217
+
+_Castnia linus_, 76
+
+Caterpillars, variation in, 36, 37
+
+Cesnola, experiments on Mantis by, 65
+
+Chaerocampa, colouring of, 68
+
+Chambers, R., _The Vestiges of Creation_ by, 15
+
+Chromosomes and Chromomeres, 47, 96-100
+
+Chun, Foot Note 36
+
+Claus, Foot Note 21
+
+Clodd, E., Foot Note 13
+
+Coadaptation, 41-54
+
+_Colobopsis truncata_, 44
+
+Colour, E. B. Poulton, in relation to Sexual Selection, 61-65
+
+Comte, A., 200-203, 252-255, 262, 265
+
+Condorcet, 221, 250, 252, 258
+
+Cope, 138
+
+Correlation of organisms, Darwin's idea of the, 2
+
+Cournot, 265
+
+Cuvier, 9, 10, 266, 268
+
+Cytology and heredity, 95, 96, 99, 100
+
+
+_Danaida chrysippus_, 75
+
+_Danaida genutia_, 75
+
+_D. Plexippus_, 75
+
+Dantec, Le, 274
+
+Darwin, Charles, as an Anthropologist, 146-165
+  --on ants, 44
+  --and S. Butler, Foot Note 61, 94
+  --on Cirripedia, 212
+  --on the Descent of Man, 111-145
+  --evolutionist authors referred to in the _Origin_ by, 9
+
+Darwin, Charles, and Haeckel, 137
+  --and History, 246-263
+  --and Huxley, 112
+  --on Lamarck, 28, 129
+  --on Language, 124
+  --and Malthus, 16, 24, 91
+  --on Patrick Matthew, 19
+  --on mental evolution, 166-196
+  --on Natural Selection, 21, 41, 54, 55, 122
+  --a "Naturalist for Naturalists," 87
+  --his personality, 187
+  --his influence on Philosophy, 197-222
+  --predecessors of, 1-22
+  --his views on religion, etc., 115, 116, 219-222
+  --his influence on religious thought, 223-245
+  --causes of his success, 10, 90
+
+Darwin, Charles, on the _Vestiges of Creation_, 15
+  --and Wallace, 23, 183
+  --on evolution, 7-15, 88
+  --on Lamarckism, 11
+
+Darwin, F., on Prichard's "Anticipations," 21
+
+Darwinism, Sociology, Evolution and, 17-18
+
+Degeneration, 49-51, 93
+
+Deniker, 137
+
+Descartes, 4
+
+Descent, history of doctrine of, 1
+
+_Descent of Man_, G. Schwalbe on _The_, 111-145
+  --rejection in Germany of _The_, 156
+
+Diderot, 6, 198
+
+Dimorphism, seasonal, 30
+
+_Dismorphia orise_, 75
+
+Dragomirov, 273
+
+Driesch, Foot Note 67
+
+Dryopithecus, 132
+
+Dubois, E., on Pithecanthropus, 132, 137
+
+Dühring, 214, 277
+
+Duns Scotus, 200
+
+Duret, C., 6
+
+Durkheim, on division of labour, 278
+
+
+Ecology, Foot Note 205
+
+Eimer, 109
+
+_Elymnias undularis_, 73, 75
+
+Embryology, the Origin of Species and, 154, 155
+
+Empedocles, 3, 27, 151
+
+Engels, 277
+
+Environment, action of, 12, 13, 15
+
+Epicurus, a poet of Evolution, 4
+
+Eristalis, 75
+
+Espinas, 275
+
+Evolution, and creation, 233
+  --conception of, 3-5, 9, 148, 151, 198
+  --discontinuous, 30
+  --experimental, 5, 7
+  --factors of, 11-15
+  --mental, 194
+  --Lloyd Morgan on mental factors in, 166-196
+  --Darwinism and Social, 18
+  --Saltatory, 29-32
+  --Herbert Spencer on, 204-207
+  --Philosophers and modern methods of studying, 4
+
+Expression of the Emotions, 177-184
+
+
+Ferri, 277
+
+Ferrier, his work on the brain, 523
+[Transcriber's note: No such page number or reference seen]
+
+Fichte, 222
+
+Flourens, 267
+
+Flowers and Insects, 61, 78
+
+Fouillée, 207, 208
+
+Fraipont, on skulls from Spy, 134
+
+
+GADOW, 162
+
+_Gallus bankiva_, 102
+
+Gallon, F., 125, 150, 269
+
+Geddes, P., 17, Foot Note 32
+
+Geddes, P. and A. W. Thomson, 276
+
+Gegenbaur, 150, 163
+
+Genetics, 93, 96
+
+_Germ-plasm_, continuity of, 95
+--Weismann on, 46-51
+
+Germinal Selection, 36, 37, 46-51, 64
+
+Gibbon, 248
+
+Giuffrida-Ruggeri, 138, 140
+
+Giotto, 259
+
+Gizycki, 217
+
+Goethe and Evolution, 8, 14, 15, 201
+--on the relation between Man and Mammals, 161, 163
+--221
+
+Gore, Dr., 226
+
+Gorjanovic-Kramberger, 134
+
+Gosse, P. H., 234
+
+_Grapta C. album_, 69
+
+Groos, 187, 188
+
+Gulick, 15, 53
+
+Guyau, 217
+
+
+Haberlandt, G., 34
+
+HAECKEL, E., on _Charles Darwin as an Anthropologist_, 146-165
+  --and Darwin, 135-151, 137, 146-165
+  --on the Descent of Man, 137, 143
+  --on Lamarck, 8, Foot Note 21
+  --a leader in the Darwinian controversy, 137
+  --217
+
+Häcker, 33
+
+Hansen, 272
+
+Hartmann, von, 240
+
+Harvey, 4
+
+Haycraft, 275
+
+Hegel, 201, 203, 215, 251, 252, 255
+
+Heraclitus, 278
+
+Herder, 4, 5, 20
+
+Heredity and Cytology, 95, 96
+  --Haeckel on, 147, 148, 149, 153
+  --and Variation, 87-110
+  --219, 224
+
+Hering, E., on Memory, 153
+
+Hertwig, O., 150
+
+History, Darwin and, 246-263
+
+Hobbes, T., 200, 215
+
+Hobhouse, 242
+
+HÖFFDING, H., on _The Influence of the Conception of Evolution
+                             on Modern Philosophy_, 197-222
+
+Holothurians, calcareous bodies in skin of, 37-41
+
+_Homo heidelbergensis_, Foot Note 118
+
+_H. neandertalensis_, 138
+
+_H. pampaeus_, 144
+
+_H. primigenius_, 133, 134, 138, 144
+
+_Homunculus_, 132
+
+Hooker, Sir J. D., and Darwin, 23, 116
+
+Huber, 170
+
+Hügel, F. von, Foot Note 221
+
+Hume, 200
+
+Hutcheson, 216
+
+Huxley, T. H., and Darwin, 112, 116, 268
+  --and the Duke of Argyll, 238
+  --on Lamarck, 89
+  --on Man, 111, 112, 137, 146, 156, 160, 163
+  --on Selection, 24, 91
+  --on transmission of acquired characters, 149
+  --14, 24, 104, 231-236, 273, 274
+
+Hybrids, Sterility of, 104, 105, 106
+
+
+Inheritance of acquired characters, 93, 94
+
+Insects and Flowers, 60, 61, 78, 79
+
+Instinct, 122, 172-175
+
+Irish Elk, an example of coadaptation, 41, 42, 45
+
+
+Jacoby, _Studies in Selection_ by, 272
+
+James, W., 180, 191, 211
+
+Jentsch, 275
+
+
+Kallima, protective colouring of, 35, 68, 70
+
+_K. inachis_, 68
+
+Kammerer's experiments on Salamanders, 28
+
+Kant, I., 4, 5, 6, 27, 198, 211, 212, 217, 221, 222
+
+Keane, on the Primates, 138
+
+Keith, on Anthropoid Apes, 138
+
+Kepler, 198
+
+Klaatsch, on Ancestry of Man, 140
+
+Klaatsch and Hauser, 134
+
+Knies, 266
+
+Kölliker, his views on Evolution, 29, 150
+
+Kollmann, on origin of human races, 144
+
+Korschinsky, 31
+
+Krause, E., Foot Note 10, 13
+
+Kropotkin, 214, 275
+
+
+Lamarck, his division of the Animal Kingdom, 160, 161
+  --Darwin's opinion of, 129
+  --on Evolution, 9-14, 21, 25, 171, 172, 173, 179, 180, 201, 202, 253
+  --on Man, 146, 148, 160, 163
+  --89, 109, 201, 202, 233
+
+Lamarckian principle, 28, 41-44, 50-54, 67, 84, 86
+
+Lamb, C., 229
+
+Lamettrie, 198
+
+Lamprecht, 260-263
+
+Lanessan, J. L. de, Foot Note 17, 275
+
+Lang, Foot Note 21
+
+Lange, 180
+
+Language, Darwin on, 123, 124
+  --Evolution and the Science of, 178, 179, 188
+
+Lankester, Sir E. Ray, on degeneration, 268
+  --on educability, 170, 189
+
+Lankester, Sir E. Ray, on the germ-plasm theory, 150
+
+Lapouge, Vacher de, 270
+
+Lartet, M. E., 189
+
+Lasalle, 266
+
+Laveleye, de, 275
+
+Lawrence, W., 89, Foot Note 65
+
+Lehmann-Nitsche, 138, 144
+
+Leibnitz, 4, 5, 213
+
+Lepidoptera, variation in, 37, 60-63
+
+Lessing, 4, 221
+
+Liddon, H. P., 234
+
+_Limenitis archippus_, 74
+
+Linnaeus, 6
+
+Locy, W. A., Foot Note 15
+
+Lovejoy, Foot Note 56
+
+Lubbock, 125
+
+Lucretius, a poet of Evolution, 4
+
+Lyell, Sir Charles, and Darwin, 23, 116
+  --the uniformitarian teaching of, 89
+
+
+Macacus, ear of, 119
+
+Mach, E., 153, 211
+
+Mahoudeau, 137
+
+Maillet, de, 6
+
+Majewski, Foot Note 238, Foot Note 239
+
+Malthus, his influence on Darwin, 16-18, 21, 24, 91
+  --200, 273
+
+Man, Descent of, 126, 127, 128, 131-145, 156-165, 189, 254, 265
+  --mental and moral qualities of animals and, 122-126, 164, 188-192
+  --pre-Darwinian views on the Descent of, 1
+
+Man, Tertiary flints worked by, 136
+
+_Man_, G. Schwalbe on Darwin's _Descent of_, 111-145
+
+Manouvrier, 137
+
+_Mantis religiosa_, colour experiments on, 65, 68
+
+Marx, 262, 276-278
+
+Matthew, P., and Natural Selection, 18, 19
+
+Maupertuis, 6, 88, 103
+
+Mayer, R., 197
+
+_Mechanitis lysimnia_, 77
+
+_Melinaea ethra_, 77
+
+Mendel, 97-100, 184, 228
+
+Merz, J. T., Foot Note 14
+
+Mesopithecus, 132
+
+Mill, J. S., 193, 200, 202, 218
+
+Mimicry, 70-82
+
+Moltke, on war, 273
+
+Monkeys, fossil, 132
+
+Montesquieu, 248
+
+Monticelli, 155
+
+MORGAN, C. LLOYD, on _Mental Factors in Evolution_, 166-196
+  --on Organic Selection, 53
+
+Morgan, T. H., 99
+
+Morselli, 138
+
+Mortillet, 136
+
+Moseley, Foot Note 224
+
+Muller, Fritz, _Für Darwin_ by, 154
+  --on Mimicry, 233
+  --59, 77
+
+Muller, J., 147
+
+Müller, Max, on language, 124
+
+Mutation, 15, 31, 184, 199, 209
+
+
+Nägeli, 109, 151, 153
+
+Nathusius, 103
+
+Natural Selection, Darwin's views on, 90, 91, 122, 149
+  --Darwin and Wallace on, 2, 163, 183
+  --and design, 241, 242
+  --and educability, 195
+  --and human development, 125, 256, 257
+  --16-20, 25, 26, 41, 55-58, 64-86, 87-96, 199, 233
+
+Neandertal skulls, 133, 134
+
+Neodarwinism, 150
+
+Newton, A., Foot Note 59
+
+Newton, I., 197, 198
+
+Niebuhr, 249, 263
+
+Nietzsche, 214, 271
+
+Nitsche, 119
+
+Novicow, 274
+
+Nuttall, G. H. F., 135
+
+
+Occam, 200
+
+Odin, 270
+
+Oecology, see Ecology
+
+_Oenothera lamarckiana_, 32
+
+Oestergren, on Holothurians, 37-39
+
+Oken, L., 7, 201
+
+Organic Selection, 53, 54, 172, 173
+
+Orthogenesis, 109
+
+Osborn, H. F., 53, Foot Note 165
+  --_From the Greeks to Darwin_ by, 3-5, 12, 14, 20
+
+_Ovibos moschatus_, 67
+
+Owen, Sir Richard, 111
+
+
+Packard, A. S., Foot Note 12, Foot Note 18
+
+Palaeopithecus, 132
+
+Paley, 18, 242, 244
+
+Panmixia, Weismann's principle of, 54
+
+_Papilio dardanus_, 72, 73, 74
+
+_P. meriones_, 73
+
+_P. merope_, 72
+
+Pearson, K., Foot Note 7
+
+Penck, 136
+
+Peridineae, 33
+
+Perrier, E., Foot Note 21, 20
+
+Perthes, B. de, 123
+
+Pfeffer, W., 28
+
+Philosophy, influence of the conception of evolution on modern, 197-222
+
+Pithecanthropus, 133, 134, 138, 143
+
+Pitheculites, 144
+
+Plate, Foot Note 37
+
+Pliopithecus, 132
+
+Pouchet, G., Foot Note 3
+
+POULTON, E. B., experiments on Butterflies by, 65
+  --on J. C. Prichard, 20
+  --on Mimicry, 69, 71, 75, 78
+  --Foot Note 34, Foot Note 43, Foot Note 49, Foot Note 55
+
+Prichard, J. C., 20, 21, 89, Foot Note 65
+
+_Pronuba yuccasella_, 79
+
+Protective resemblance, 65-70
+
+Pusey, 115
+
+
+Quatrefages, A. de, Foot Note 21, 19
+
+
+Radiolarians, 33
+
+Ranke, 249, 251, 255, 263
+
+Rau, A., 153
+
+Ray, J., 4
+
+Regeneration, Foot Note 71
+
+Religious thought, Darwin's influence on, 223-245
+
+Reversion, 120, 121
+
+Ridley, H. N., Foot Note 88
+
+Ritchie, 270
+
+Robinet, 6
+
+Rolph, 217
+
+Romanes, G. J., Foot Note 3, 15, 32, 54, 164, 234
+
+Roux, 151, 152
+
+Ruskin, 230
+
+Rutot, 136
+
+Saint-Hilaire, E. G. de, 8, 15, 20
+
+Saltatory Evolution, 29-32 (see also Mutations)
+
+Sanders, experiments on Vanessa by, 65
+
+Savigny, 249
+
+Schelling, 4, 5, 200, 201
+
+Schleiden and Schwann, Cell-theory of, 147
+
+Schoetensack, on _Homo heidelbergensis_, Foot Note 118
+
+Schütt, 23
+
+SCHWALBE, G., on _The Descent of Man_, 111-145
+
+Seeck, O., Foot Note 240
+
+Segregation, 97, 98
+
+Selection, artificial, 24, 25, 26, 41, 45, 120, 269-272
+  --germinal, 35, 36, 46-52, 64
+
+Selection, natural (see Natural Selection)
+  --organic, 53, 171, 172
+  --sexual, 55-64, 117, 118
+  --social and natural, 271
+  --23-86, 103, 129, 130
+
+Selenka, 131
+
+Semnopithecus, 132
+
+Semon, R., 28, 153
+
+Sergi, 138, 143
+
+Sex, recent investigations on, 99, 100
+
+Sibbern, 201
+
+_Smerinthus ocellata_, 38
+
+_Smerinthus populi_, 38
+
+_S. tiliae_, 38
+
+Smith, A., 200
+
+Sociology,  Darwinism and, 264-280
+  --History and, 255
+
+Sollas, W. J., 134
+
+Sorley, W. R., 217
+
+Species and varieties, 100
+
+Spencer, H., on evolution, 204-209
+  --on the theory of Selection, 41
+
+Spencer, H., on Sociology, 268
+  --on the transmission of acquired characters, 149
+  --on Weismann, 41, 150
+  --2, 17, 217, 231, 268
+
+Sphingidae, variation in, 37
+
+Spinoza, 153, 206
+
+Standfuss, 82
+
+Stephen, L., 217
+
+Sterility in hybrids, 104-106
+
+Sterne, C, Foot Note 10
+
+Struggle for existence, 25, 26, 272-274
+
+Sutton, A. W., Foot Note 73
+
+Synapta, calcareous bodies in skin of, 38-41
+
+Syrphus, 75
+
+
+Tarde, G., 279
+
+Tennant, F. R., Foot Note 218
+
+Tetraprothomo, 138, 144
+
+THOMSON, J. A., on _Darwin's Predecessors_, 1-22
+  --150
+  --and P. Geddes, 276
+
+Treschow, 201
+
+Treviranus, 8, 14, 15
+
+Turgot, 249
+
+Turner, Sir W., 150
+
+Tylor, 267
+
+Tyndall, W., 267
+
+Tyrrell, G, Foot Note 222
+
+
+Uhlenhuth, on blood reactions, 135
+
+Use and disuse, 28, 41-43, 48-54, 94, 95, 119, 149
+
+
+Vanessa, 63
+
+_V. levana_, 31
+
+_V. polychloros_, 82
+
+_V. urticae_, 65, 82
+
+Variability, Darwin's attention directed to, 24
+  --W. Bateson on, 87-110
+  --causes of, 200
+
+Variation, Darwin's views as an evolutionist, and as a systematist, on, 212
+  --and heredity, 87-110
+  --minute, 28-32
+  --in relation to species, 100, 101
+
+Varigny, H. de, 6, 19
+
+Verworn, 136
+
+_Vestiges of Creation_, Darwin on _The_, 15
+
+Virchow, his opposition to Darwin, 157, 158
+  --on the transmission of acquired characters, 149
+
+Vogt, 137
+
+Voltaire, 248
+
+VRIES, H. de, the Mutation theory of, 31, 101, 151, 213
+
+
+WAGGETT, Rev. P. N., on _The Influence of Darwin upon Religious
+                                    Thought_, 223-245
+
+Wallace, A. R., on Colour, 63, 71
+  --and Darwin, Foot Note 7, 23, 183
+  --on the Descent of Man, 116
+  --on Malthus, 17
+  --on Natural Selection, 2, 16, 163, 232
+
+Wallace, A. R., on social reforms, 275, 276
+  --on Sexual Selection, 183, 184
+
+Walton, 237
+
+Watt, J., and Natural Selection, 21
+
+WEISMANN, A., on _The Selection Theory_, 23-86
+  --his germ-plasm theory, 46-51, 149, 150
+  --and Prichard, 20
+  --and Spencer, 42
+
+Weismann, A., on the transmission of acquired characters, 93-95
+  --156
+
+Wells, W. C, and Natural Selection, 18
+
+White, G., 3
+
+Williams, C. M., 217
+
+Wilson, E. B., on cytology, 99
+
+Wolf, 249
+
+Wollaston's, T. V., _Variation of Species_, Foot Note 59
+
+Woltmann, 277
+
+Woolner, 118
+
+Wundt, on language, 207, 208
+
+
+_Xylina vetusta_, 82
+
+
+Yucca, fertilisation of, 78, 79
+
+
+Zeller, E., Foot Note 3
+
+_Zoonomia_, Erasmus Darwin's, 7
+
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+
+The Project Gutenberg EBook of Evolution in Modern Thought, by 
+Ernst Haeckel and J. Arthur Thomson and August Weismann
+
+This eBook is for the use of anyone anywhere at no cost and with
+almost no restrictions whatsoever.  You may copy it, give it away or
+re-use it under the terms of the Project Gutenberg License included
+with this eBook or online at www.gutenberg.org
+
+
+Title: Evolution in Modern Thought
+
+Author: Ernst Haeckel
+        J. Arthur Thomson
+        August Weismann
+
+Release Date: August 29, 2007 [EBook #22430]
+
+Language: English
+
+Character set encoding: ISO-8859-1
+
+*** START OF THIS PROJECT GUTENBERG EBOOK EVOLUTION IN MODERN THOUGHT ***
+
+
+
+
+Produced by Marilynda Fraser-Cunliffe, Sankar Viswanathan,
+and the Online Distributed Proofreading Team at
+http://www.pgdp.net
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+
+
+
+
+
+</pre>
+
+<div class="center"><img src="images/image_001.jpg" alt="Cover Page" width="500" height="804" /></div>
+<p>&nbsp;</p>
+<h1>EVOLUTION IN MODERN<br />
+THOUGHT</h1>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+<h2>BY HAECKEL, THOMSON, WEISMANN<br />
+AND OTHERS</h2>
+<p>&nbsp;</p>
+<p class="center"><img src="images/image_002.jpg" alt="Seal" width="75" height="99" /></p>
+<p>&nbsp;</p>
+<p>&nbsp;</p>
+<h3>THE MODERN LIBRARY</h3>
+
+<h4>PUBLISHERS&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;:: ::&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;NEW YORK</h4>
+
+
+
+
+<hr style="width: 65%;" />
+<h2><a name="CONTENTS" id="CONTENTS"></a>CONTENTS</h2>
+
+<table summary="Contents">
+<tr><td></td>
+  <td>&nbsp;</td>
+  <td>&nbsp;</td>
+  <td class="tocpg f1">PAGE</td></tr>
+<tr><td class="tocch">I</td>
+  <td>&nbsp;</td>
+  <td><span class="smcap"><a href="#I">Darwin's Predecessors</a></span></td>
+<td class="tocpg"><a href="#Page_1">1</a></td>
+</tr>
+<tr><td></td>
+  <td>&nbsp;</td>
+  <td>J. Arthur Thomson, Professor of Natural
+History in the University of Aberdeen</td></tr>
+<tr><td class="tocch">II</td>
+  <td>&nbsp;</td>
+  <td><i><a href="#II">The Selection Theory</a></i></td>
+<td class="tocpg"><a href="#Page_23">23</a></td>
+</tr>
+<tr><td></td>
+  <td>&nbsp;</td>
+  <td>August Weismann, Professor of Zoology
+in the University of Freiburg (Baden)</td></tr>
+<tr><td class="tocch">III</td>
+  <td>&nbsp;</td>
+  <td><span class="smcap"><a href="#III">Heredity and Variation in Modern Lights</a></span></td>
+<td class="tocpg"><a href="#Page_87">87</a></td>
+</tr>
+<tr><td></td>
+  <td>&nbsp;</td>
+  <td>W. Bateson, Professor of Biology in the
+University of Cambridge</td></tr>
+<tr><td class="tocch">IV</td>
+  <td>&nbsp;</td>
+  <td><a href="#IV">"<span class="smcap">The Descent of Man</span>"</a></td>
+<td class="tocpg"><a href="#Page_111">111</a></td>
+</tr>
+<tr><td></td>
+  <td>&nbsp;</td>
+  <td>G. Schwalbe, Professor of Anatomy in
+the University of Strassburg</td></tr>
+<tr><td class="tocch">V</td>
+  <td>&nbsp;</td>
+  <td><span class="smcap"><a href="#V">Charles Darwin as an Anthropologist</a></span></td>
+<td class="tocpg"><a href="#Page_146">146</a></td>
+</tr>
+<tr><td></td>
+  <td>&nbsp;</td>
+  <td>Ernst Haeckel, Professor of Zoology in
+the University of Jena</td></tr>
+<tr><td class="tocch">VI</td>
+  <td>&nbsp;</td>
+  <td><span class="smcap"><a href="#VI">Mental Factors in Evolution</a></span></td>
+<td class="tocpg"><a href="#Page_166">166</a></td>
+</tr>
+<tr><td></td>
+  <td>&nbsp;</td>
+  <td>C. Lloyd Morgan, Professor of Psychology
+at University College, Bristol</td></tr>
+<tr><td class="tocch">VII</td>
+  <td>&nbsp;</td>
+  <td><span class="smcap"><a href="#VII">The Influence of the Conception of
+Evolution on Modern Philosophy</a></span></td>
+<td class="tocpg"><a href="#Page_197">197</a></td>
+</tr>
+<tr><td></td>
+  <td>&nbsp;</td>
+  <td>H. H&ouml;ffding, Professor of Philosophy in
+the University of Copenhagen</td></tr>
+<tr><td class="tocch">VIII</td>
+  <td>&nbsp;</td>
+  <td><span class="smcap"><a href="#VIII">The Influence of Darwin Upon Religious
+Thought</a></span></td>
+<td class="tocpg"><a href="#Page_223">223</a></td>
+</tr>
+<tr><td></td>
+  <td>&nbsp;</td>
+  <td>Rev. P. H. Waggett</td></tr>
+<tr><td class="tocch">IX</td>
+  <td>&nbsp;</td>
+  <td><span class="smcap"><a href="#IX">Darwinism and History</a></span></td>
+<td class="tocpg"><a href="#Page_246">246</a></td>
+</tr>
+<tr><td></td>
+  <td>&nbsp;</td>
+  <td>J. B. Bury, Regious Professor of Modern
+History in the University of Cambridge</td></tr>
+<tr><td class="tocch">X</td>
+  <td>&nbsp;</td>
+  <td><span class="smcap"><a href="#X">Darwinism and Sociology</a></span></td>
+<td class="tocpg"><a href="#Page_264">264</a></td>
+</tr>
+<tr><td></td>
+  <td>&nbsp;</td>
+  <td>C. Bougl&eacute;, Professor of Social Philosophy
+in the University of Toulouse, and Deputy-Professor
+at the Sorbonne, Paris</td></tr>
+</table>
+
+
+
+<hr style="width: 65%;" />
+<p><span class='pagenum'><a name="Page_1" id="Page_1">[1]</a></span></p>
+<h2>EVOLUTION IN MODERN THOUGHT</h2>
+
+
+
+<hr style="width: 65%;" />
+<h2><a name="I" id="I"></a>I</h2>
+
+<h2>DARWIN'S PREDECESSORS</h2>
+<p class="center"><span class="smcap1">By J. Arthur Thomson</span></p>
+<h4><i>Professor of Natural History in the University of Aberdeen</i></h4>
+<p>In seeking to discover Darwin's relation to his predecessors it is
+useful to distinguish the various services which he rendered to the
+theory of organic evolution.</p>
+
+<p>(I) As everyone knows, the general idea of the Doctrine of Descent is
+that the plants and animals of the present day are the lineal
+descendants of ancestors on the whole somewhat simpler, that these
+again are descended from yet simpler forms, and so on backwards
+towards the literal "Protozoa" and "Protophyta" about which we
+unfortunately know nothing. Now no one supposes that Darwin originated
+this idea, which in rudiment at least is as old as Aristotle. What
+Darwin did was to make it current intellectual coin. He gave it a form
+that commended itself to the scientific and public intelligence of the
+day, and he won widespread conviction by showing with consummate skill
+that it was an effective formula to work with, a key which no lock
+refused. In a scholarly, critical, and pre-eminently fair-minded way,
+admitting difficulties and removing them, foreseeing objections and
+forestalling them, he showed that the doctrine<span class='pagenum'><a name="Page_2" id="Page_2">[2]</a></span> of descent supplied a
+modal interpretation of how our present-day fauna and flora have come
+to be.</p>
+
+<p>(II) In the second place, Darwin applied the evolution-idea to
+particular problems, such as the descent of man, and showed what a
+powerful organon it is, introducing order into masses of uncorrelated
+facts, interpreting enigmas both of structure and function, both
+bodily and mental, and, best of all, stimulating and guiding further
+investigation. But here again it cannot be claimed that Darwin was
+original. The problem of the descent or ascent of man, and other
+particular cases of evolution, had attracted not a few naturalists
+before Darwin's day, though no one [except Herbert Spencer in the
+psychological domain (1855)] had come near him in precision and
+thoroughness of inquiry.</p>
+
+<p>(III) In the third place, Darwin contributed largely to a knowledge of
+the factors in the evolution-process, especially by his analysis of
+what occurs in the case of domestic animals and cultivated plants, and
+by his elaboration of the theory of Natural Selection which Alfred
+Russel Wallace independently stated at the same time, and of which
+there had been a few previous suggestions of a more or less vague
+description. It was here that Darwin's originality was greatest, for
+he revealed to naturalists the many different forms&mdash;often very
+subtle&mdash;which natural selection takes, and with the insight of a
+disciplined scientific imagination he realised what a mighty engine of
+progress it has been and is.</p>
+
+<p>(IV) As an epoch-marking contribution, not only to &AElig;tiology but to
+Natural History in the widest sense, we rank the picture which Darwin
+gave to the world of the web of life, that is to say, of the
+inter-relations and linkages in Nature. For the Biology of the
+individual&mdash;if that be not a contradiction in terms&mdash;no idea is more
+fundamental than that of the correlation of organs, but Darwin's most
+characteristic contribution was not less fundamental,&mdash;it was the idea
+of the correlation of organisms. This, again, was not novel; we find
+it in the works of naturalists like Christian<span class='pagenum'><a name="Page_3" id="Page_3">[3]</a></span> Conrad Sprengel,
+Gilbert White, and Alexander von Humboldt, but the realisation of its
+full import was distinctly Darwinian.</p>
+
+
+<p><i>As Regards the General Idea of Organic Evolution</i></p>
+
+<p>While it is true, as Prof. H. F. Osborn puts it, that "'Before and
+after Darwin' will always be the <i>ante et post urbem conditam</i> of
+biological history," it is also true that the general idea of organic
+evolution is very ancient. In his admirable sketch <i>From the Greeks to
+Darwin</i>,<a name="FNanchor_1_1" id="FNanchor_1_1"></a><a href="#Footnote_1_1" class="fnanchor">[1]</a> Prof. Osborn has shown that several of the ancient
+philosophers looked upon Nature as a gradual development and as still
+in process of change. In the suggestions of Empedocles, to take the
+best instance, there were "four sparks of truth,&mdash;first, that the
+development of life was a gradual process; second, that plants were
+evolved before animals; third, that imperfect forms were gradually
+replaced (not succeeded) by perfect forms; fourth, that the natural
+cause of the production of perfect forms was the extinction of the
+imperfect."<a name="FNanchor_2_2" id="FNanchor_2_2"></a><a href="#Footnote_2_2" class="fnanchor">[2]</a> But the fundamental idea of one stage giving origin to
+another was absent. As the blue &AElig;gean teemed with treasures of beauty
+and threw many upon its shores, so did Nature produce like a fertile
+artist what had to be rejected as well as what was able to survive,
+but the idea of one species emerging out of another was not yet
+conceived.</p>
+
+<p><span class='pagenum'><a name="Page_4" id="Page_4">[4]</a></span></p><p>Aristotle's views of Nature<a name="FNanchor_3_3" id="FNanchor_3_3"></a><a href="#Footnote_3_3" class="fnanchor">[3]</a> seem to have been more definitely
+evolutionist than those of his predecessors, in this sense, at least,
+that he recognised not only an ascending scale, but a genetic series
+from polyp to man and an age-long movement towards perfection. "It is
+due to the resistance of matter to form that Nature can only rise by
+degrees from lower to higher types." "Nature produces those things
+which, being continually moved by a certain principle contained in
+themselves, arrive at a certain end."</p>
+
+<p>To discern the outcrop of evolution-doctrine in the long interval
+between Aristotle and Bacon seems to be very difficult, and some of
+the instances that have been cited strike one as forced. Epicurus and
+Lucretius, often called poets of evolution, both pictured animals as
+arising directly out of the earth, very much as Milton's lion long
+afterwards pawed its way out. Even when we come to Bruno who wrote
+that "to the sound of the harp of the Universal Apollo (the World
+Spirit), the lower organisms are called by stages to higher, and the
+lower stages are connected by intermediate forms with the higher,"
+there is great room, as Prof. Osborn points out,<a name="FNanchor_4_4" id="FNanchor_4_4"></a><a href="#Footnote_4_4" class="fnanchor">[4]</a> for difference of
+opinion as to how far he was an evolutionist in our sense of the term.</p>
+
+<p>The awakening of natural science in the sixteenth century brought the
+possibility of a concrete evolution theory nearer, and in the early
+seventeenth century we find evidences of a new spirit&mdash;in the
+embryology of Harvey and the classifications of Ray. Besides sober
+naturalists there were speculative dreamers in the sixteenth and
+seventeenth centuries who had at least got beyond static formulae,
+but, as Professor Osborn points out,<a name="FNanchor_5_5" id="FNanchor_5_5"></a><a href="#Footnote_5_5" class="fnanchor">[5]</a> "it is a very striking fact,
+that the basis of our modern methods of studying the Evolution problem
+was established not by the early naturalists nor by the speculative
+writers, but by the Philosophers." He refers to Bacon, Descartes,
+Leibnitz, Hume, Kant, Lessing, Herder, and Schelling. "They alone were
+upon the main track of modern thought. It is evident that they were
+groping in the dark for a working theory of the Evolution of life, and
+it is remarkable that they clearly perceived from the outset that the
+point to which observation should be directed was not the past but the
+present mutability of species, and further, that this mutability was
+simply the variation of individuals on an extended scale."</p>
+
+<p><span class='pagenum'><a name="Page_5" id="Page_5">[5]</a></span></p><p>Bacon seems to have been one of the first to think definitely about
+the mutability of species, and he was far ahead of his age in his
+suggestion of what we now call a Station of Experimental Evolution.
+Leibnitz discusses in so many words how the species of animals may be
+changed and how intermediate species may once have linked those that
+now seem discontinuous. "All natural orders of beings present but a
+single chain".... "All advances by degrees in Nature, and nothing by
+leaps." Similar evolutionist statements are to be found in the works
+of the other "philosophers," to whom Prof. Osborn refers, who were,
+indeed, more scientific than the naturalists of their day. It must be
+borne in mind that the general idea of organic evolution&mdash;that the
+present is the child of the past&mdash;is in great part just the idea of
+human history projected upon the natural world, differentiated by the
+qualification that the continuous "Becoming" has been wrought out by
+forces inherent in the organisms themselves and in their environment.</p>
+
+<p>A reference to Kant<a name="FNanchor_6_6" id="FNanchor_6_6"></a><a href="#Footnote_6_6" class="fnanchor">[6]</a> should come in historical order after Buffon,
+with whose writings he was acquainted, but he seems, along with Herder
+and Schelling, to be best regarded as the culmination of the
+evolutionist philosophers&mdash;of those at least who interested themselves
+in scientific problems. In a famous passage he speaks of "the
+agreement of so many kinds of animals in a certain common plan of
+structure" ... an "analogy of forms" which "strengthens the
+supposition that they have an actual blood-relationship, due to
+derivation from a common parent." He speaks of "the great Family of
+creatures, for as a Family we must conceive it, if the above-mentioned
+continuous and connected relationship has a real foundation." Prof.
+Osborn alludes to the scientific caution which led Kant, biology being
+what it was, to refuse to entertain the hope "that a Newton may one
+day arise even to make the production of a blade of grass
+comprehensible, according to natural laws ordained by no intention."
+As Prof. Haeckel finely observes, Darwin rose up as Kant's Newton.<a name="FNanchor_7_7" id="FNanchor_7_7"></a><a href="#Footnote_7_7" class="fnanchor">[7]</a></p>
+
+<p><span class='pagenum'><a name="Page_6" id="Page_6">[6]</a></span></p><p>The scientific renaissance brought a wealth of fresh impressions and
+some freedom from the tyranny of tradition, and the twofold stimulus
+stirred the speculative activity of a great variety of men from old
+Claude Duret of Moulins, of whose weird transformism (1609) Dr. Henry
+de Varigny<a name="FNanchor_8_8" id="FNanchor_8_8"></a><a href="#Footnote_8_8" class="fnanchor">[8]</a> gives us a glimpse, to Lorenz Oken (1779-1851) whose
+writings are such mixtures of sense and nonsense that some regard him
+as a far-seeing prophet and others as a fatuous follower of
+intellectual will-o'-the-wisps. Similarly, for De Maillet, Maupertuis,
+Diderot, Bonnet, and others, we must agree with Professor Osborn that
+they were not actually in the main Evolution movement. Some have been
+included in the roll of honour on very slender evidence, Robinet for
+instance, whose evolutionism seems to us extremely dubious.<a name="FNanchor_9_9" id="FNanchor_9_9"></a><a href="#Footnote_9_9" class="fnanchor">[9]</a></p>
+
+<p><span class='pagenum'><a name="Page_7" id="Page_7">[7]</a></span></p><p>The first naturalist to give a broad and concrete expression to the
+evolutionist doctrine of descent was Buffon (1707-1788), but it is
+interesting to recall the fact that his contemporary Linn&aelig;us
+(1707-1778), protagonist of the counter-doctrine of the fixity of
+species,<a name="FNanchor_10_10" id="FNanchor_10_10"></a><a href="#Footnote_10_10" class="fnanchor">[10]</a> went the length of admitting (in 1762) that new species
+might arise by inter-crossing. Buffon's position among the pioneers of
+the evolution-doctrine is weakened by his habit of vacillating between
+his own conclusions and the orthodoxy of the Sorbonne, but there is no
+doubt that he had firm grasp of the general idea of "l'encha&icirc;nment des
+&ecirc;tres."</p>
+
+<p>Erasmus Darwin (1731-1802), probably influenced by Buffon, was another
+firm evolutionist, and the outline of his argument in the
+<i>Zoonomia</i><a name="FNanchor_11_11" id="FNanchor_11_11"></a><a href="#Footnote_11_11" class="fnanchor">[11]</a> might serve in part at least to-day. "When we revolve
+in our minds the metamorphoses of animals, as from the tadpole to the
+frog; secondly, the changes produced by artificial cultivation, as in
+the breeds of horses, dogs, and sheep; thirdly, the changes produced
+by conditions of climate and of season, as in the sheep of warm
+climates being covered with hair instead of wool, and the hares and
+partridges of northern climates becoming white in winter: when,
+further, we observe the changes of structure produced by habit, as
+seen especially in men of different occupations; or the changes
+produced by artificial mutilation and prenatal influences, as in the
+crossing of species and production of monsters; fourth, when we
+observe the essential unity of plan in all warm-blooded animals,&mdash;we
+are led to conclude that they have been alike produced from a similar
+living filament".... "From thus meditating upon the minute portion of
+time in which many of the above changes have been produced, would it
+be too bold to imagine, in the great length of time since the earth
+began to exist, perhaps millions of years before the commencement of
+the history of mankind, that all warm-blooded animals have arisen from
+one living filament?"... "This idea of the gradual generation of all
+things seems to have been as familiar to the ancient philosophers as
+to the modern ones, and to have given rise to the beautiful
+hieroglyphic figure of the &#960;&#961;&#969;&#771;&#964;&#959;&#957; &#8032;&#8001;&#957;, or first great egg,
+produced by night, that is, whose origin is involved in obscurity, and
+animated by &#7964;&#961;&#969;&#962;, that is, by Divine Love; from whence
+proceeded all things which exist."</p>
+
+<p><span class='pagenum'><a name="Page_8" id="Page_8">[8]</a></span></p><p>Lamarck (1744-1829) seems to have become an evolutionist
+independently of Erasmus Darwin's influence, though the parallelism
+between them is striking. He probably owed something to Buffon, but he
+developed his theory along a different line. Whatever view be held in
+regard to that theory there is no doubt that Lamarck was a
+thorough-going evolutionist. Professor Haeckel speaks of the
+<i>Philosophie Zoologique</i> as "the first connected and thoroughly
+logical exposition of the theory of descent."<a name="FNanchor_12_12" id="FNanchor_12_12"></a><a href="#Footnote_12_12" class="fnanchor">[12]</a></p>
+
+<p>Besides the three old masters, as we may call them, Buffon, Erasmus
+Darwin, and Lamarck, there were other quite convinced pre-Darwinian
+evolutionists. The historian of the theory of descent must take
+account of Treviranus whose <i>Biology or Philosophy of Animate Nature</i>
+is full of evolutionary suggestions; of Etienne Geoffroy St. Hilaire,
+who in 1830, before the French Academy of Sciences, fought with
+Cuvier, the fellow-worker of his youth, an intellectual duel on the
+question of descent; of Goethe, one of the founders of morphology and
+the greatest poet of Evolution&mdash;who, in his eighty-first year, heard
+the tidings of Geoffrey St. Hilaire's defeat with an interest which
+transcended the political anxieties of the time; and of many others
+who had gained with more or less confidence and clearness a new
+outlook on Nature. It will be remembered that Darwin refers to
+thirty-four more or less evolutionist authors in his Historical
+Sketch, and the list might be added to. Especially when we come near
+to 1858 do the numbers increase, and one of the most remarkable, as
+also most independent champions of the evolution-idea before that date
+was Herbert Spencer, who not only marshalled the arguments in a very
+forcible way in 1852, but applied the formula in detail in his
+<i>Principles of Psychology</i> in 1855.<a name="FNanchor_13_13" id="FNanchor_13_13"></a><a href="#Footnote_13_13" class="fnanchor">[13]</a></p>
+
+<p><span class='pagenum'><a name="Page_9" id="Page_9">[9]</a></span></p><p>It is right and proper that we should shake ourselves free from all
+creationist appreciations of Darwin, and that we should recognise the
+services of pre-Darwinian evolutionists who helped to make the time
+ripe, yet one cannot help feeling that the citation of them is apt to
+suggest two fallacies. It may suggest that Darwin simply entered into
+the labours of his predecessors, whereas, as a matter of fact, he knew
+very little about them till after he had been for years at work. To
+write, as Samuel Butler did, "Buffon planted, Erasmus Darwin and
+Lamarck watered, but it was Mr. Darwin who said 'That fruit is ripe,'
+and shook it into his lap" ... seems to us a quite misleading version
+of the facts of the case. The second fallacy which the historical
+citation is a little apt to suggest is that the filiation of ideas is
+a simple problem. On the contrary, the history of an idea, like the
+pedigree of an organism, is often very intricate, and the evolution of
+the evolution-idea is bound up with the whole progress of the world.
+Thus in order to interpret Darwin's clear formulation of the idea of
+organic evolution and his convincing presentation of it, we have to do
+more than go back to his immediate predecessors, such as Buffon,
+Erasmus Darwin, and Lamarck; we have to inquire into the acceptance of
+evolutionary conceptions in regard to other orders of facts, such as
+the earth and the solar system;<a name="FNanchor_14_14" id="FNanchor_14_14"></a><a href="#Footnote_14_14" class="fnanchor">[14]</a> we have to realise how the growing
+success of scientific interpretation along other lines gave confidence
+to those who refused to admit that there was any domain from which
+science could be excluded as a trespasser; we have to take account of
+the development of philosophical thought, and even of theological and
+religious movements; we should also, if we are wise enough, consider
+social changes. In short, we must abandon the idea that we can
+understand the history of any science as such, without reference to
+contemporary evolution in other departments of activity.</p>
+
+<p><span class='pagenum'><a name="Page_10" id="Page_10">[10]</a></span></p><p>While there were many evolutionists before Darwin, few of them were
+expert naturalists and few were known outside a small circle; what was
+of much more importance was that the genetic view of Nature was
+insinuating itself in regard to other than biological orders of facts,
+here a little and there a little, and that the scientific spirit had
+ripened since the days when Cuvier laughed Lamarck out of court. How
+was it that Darwin succeeded where others had failed? Because, in the
+first place, he had clear visions&mdash;"pens&eacute;es de la jeunesse, execut&eacute;es
+par l'&acirc;ge m&ucirc;r"&mdash;which a University curriculum had not made impossible,
+which the <i>Beagle voyage</i> made vivid, which an unrivalled British
+doggedness made real&mdash;visions of the web of life, of the fountain of
+change within the organism, of the struggle for existence and its
+winnowing, and of the spreading genealogical tree. Because, in the
+second place, he put so much grit into the verification of his
+visions, putting them to the proof in an argument which is of its
+kind&mdash;direct demonstration being out of the question&mdash;quite
+unequalled. Because, in the third place, he broke down the opposition
+which the most scientific had felt to the seductive modal formula of
+evolution by bringing forward a more plausible theory of the process
+than had been previously suggested. Nor can one forget, since
+questions of this magnitude are human and not merely academic, that he
+wrote so that all men could understand.<span class='pagenum'><a name="Page_11" id="Page_11">[11]</a></span></p>
+
+
+<p><i>As Regards the Factors of Evolution</i></p>
+
+<p>It is admitted by all who are acquainted with the history of biology
+that the general idea of organic evolution as expressed in the
+Doctrine of Descent was quite familiar to Darwin's grandfather and to
+others before and after him, as we have briefly indicated. It must
+also be admitted that some of these pioneers of evolutionism did more
+than apply the evolution-idea as a modal formula of becoming, they
+began to inquire into the factors in the process. Thus there were
+pre-Darwinian theories of evolution, and to these we must now briefly
+refer.<a name="FNanchor_15_15" id="FNanchor_15_15"></a><a href="#Footnote_15_15" class="fnanchor">[15]</a></p>
+
+<p>In all biological thinking we have to work with the categories
+Organism&mdash;Function&mdash;Environment, and theories of evolution may be
+classified in relation to these. To some it has always seemed that the
+fundamental fact is the living organism,&mdash;a creative agent, a striving
+will, a changeful Proteus, selecting its environment, adjusting itself
+to it, self-differentiating and self-adaptive. The necessity of
+recognising the importance of the organism is admitted by all
+Darwinians who start with inborn variations, but it is open to
+question whether the whole truth of what we might call the Goethian
+position is exhausted in the postulate of inherent variability.</p>
+
+<p>To others it has always seemed that the emphasis should be laid on
+Function,&mdash;on use and disuse, on doing and not doing. Practice makes
+perfect; <i>c'est &agrave; force de forger qu'on devient forgeron</i>. This is one
+of the fundamental ideas of Lamarckism; to some extent it met with
+Darwin's approval; and it finds many supporters to-day. One of the
+ablest of these&mdash;Mr. Francis Darwin&mdash;has recently given strong reasons
+for combining a modernised Lamarckism with what we usually regard as
+sound Darwinism.<a name="FNanchor_16_16" id="FNanchor_16_16"></a><a href="#Footnote_16_16" class="fnanchor">[16]</a></p>
+
+<p><span class='pagenum'><a name="Page_12" id="Page_12">[12]</a></span></p><p>To others it has always seemed that the emphasis should be laid on
+the Environment, which wakes the organism to action, prompts it to
+change, makes dints upon it, moulds it, prunes it, and finally,
+perhaps, kills it. It is again impossible to doubt that there is truth
+in this view, for even if environmentally induced "modifications" be
+not transmissible, environmentally induced "variations" are; and even
+if the direct influence of the environment be less important than many
+enthusiastic supporters of this view&mdash;may we call them
+Buffonians&mdash;think, there remains the indirect influence which
+Darwinians in part rely on,&mdash;the eliminative process. Even if the
+extreme view be held that the only form of discriminate elimination
+that counts is inter-organismal competition, this might be included
+under the rubric of the animate environment.</p>
+
+<p>In many passages Buffon<a name="FNanchor_17_17" id="FNanchor_17_17"></a><a href="#Footnote_17_17" class="fnanchor">[17]</a> definitely suggested that environmental
+influences&mdash;especially of climate and food&mdash;were directly productive
+of changes in organisms, but he did not discuss the question of the
+transmissibility of the modifications so induced, and it is difficult
+to gather from his inconsistent writings what extent of transformation
+he really believed in. Prof. Osborn says of Buffon: "The struggle for
+existence, the elimination of the least-perfected species, the contest
+between the fecundity of certain species and their constant
+destruction, are all clearly expressed in various passages." He quotes
+two of these:<a name="FNanchor_18_18" id="FNanchor_18_18"></a><a href="#Footnote_18_18" class="fnanchor">[18]</a></p>
+
+<p>"Le cours ordinaire de la nature vivante, est en g&eacute;n&eacute;ral toujours
+constant, toujours le m&ecirc;me; son mouvement, toujours r&eacute;gulier, roule
+sur deux points in&eacute;branlables: l'un, la f&eacute;condit&eacute; sans bornes donn&eacute;e &agrave;
+toutes les esp&egrave;ces; l'autre, les obstacles sans nombre qui r&eacute;duisent
+cette f&eacute;condit&eacute; &agrave; une mesure d&eacute;termin&eacute;e et ne laissent en tout temps
+qu'&agrave; peu pr&egrave;s la m&ecirc;me quantit&eacute; d'individus de chaque esp&egrave;ce" ... "Les
+esp&egrave;ces les moins parfaites, les plus d&eacute;licates, les plus pesantes,
+les moins agissantes, les moins arm&eacute;es, etc., ont d&eacute;j&agrave; disparu ou
+dispara&icirc;tront.".</p>
+
+<p><span class='pagenum'><a name="Page_13" id="Page_13">[13]</a></span></p><p>Erasmus Darwin<a name="FNanchor_19_19" id="FNanchor_19_19"></a><a href="#Footnote_19_19" class="fnanchor">[19]</a> had a firm grip of the "idea of the gradual
+formation and improvement of the Animal world," and he had his theory
+of the process. No sentence is more characteristic than this: "All
+animals undergo transformations which are in part produced by their
+own exertions, in response to pleasures and pains, and many of these
+acquired forms or propensities are transmitted to their posterity."
+This is Lamarckism before Lamarck, as his grandson pointed out. His
+central idea is that wants stimulate efforts and that these result in
+improvements which subsequent generations make better still. He
+realised something of the struggle for existence and even pointed out
+that this advantageously checks the rapid multiplication. "As Dr.
+Krause points out, Darwin just misses the connection between this
+struggle and the Survival of the Fittest."<a name="FNanchor_20_20" id="FNanchor_20_20"></a><a href="#Footnote_20_20" class="fnanchor">[20]</a></p>
+
+<p>Lamarck<a name="FNanchor_21_21" id="FNanchor_21_21"></a><a href="#Footnote_21_21" class="fnanchor">[21]</a> (1744-1829) seems to have thought out his theory of
+evolution without any knowledge of Erasmus Darwin's which it closely
+resembled. The central idea of his theory was the cumulative
+inheritance of functional modifications. "Changes in environment bring
+about changes in the habits of animals. Changes in their wants
+necessarily bring about parallel changes in their habits. If new wants
+become constant or very lasting, they form new habits, the new habits
+involve the use of new parts, or a different use of old parts, which
+results finally in the production of new organs and the modification
+of old ones." He differed from Buffon in not attaching importance, as
+far as animals are concerned, to the direct influence of the
+environment, "for environment can effect no direct change whatever
+upon the organisation of animals," but in regard to plants he agreed
+with Buffon that external conditions directly moulded them.</p>
+
+<p><span class='pagenum'><a name="Page_14" id="Page_14">[14]</a></span></p><p>Treviranus<a name="FNanchor_22_22" id="FNanchor_22_22"></a><a href="#Footnote_22_22" class="fnanchor">[22]</a> (1776-1837), whom Huxley ranked beside Lamarck, was on
+the whole Buffonian, attaching chief importance to the influence of a
+changeful environment both in modifying and in eliminating, but he was
+also Goethian, for instance in his idea that species like individuals
+pass through periods of growth, full bloom, and decline. "Thus, it is
+not only the great catastrophes of Nature which have caused
+extinction, but the completion of cycles of existence, out of which
+new cycles have begun." A characteristic sentence is quoted by Prof.
+Osborn: "In every living being there exists a capability of an endless
+variety of form-assumption; each possesses the power to adapt its
+organisation to the changes of the outer world, and it is this power,
+put into action by the change of the universe, that has raised the
+simple zoophytes of the primitive world to continually higher stages
+of organisation, and has introduced a countless variety of species
+into animate Nature."</p>
+
+<p>Goethe<a name="FNanchor_23_23" id="FNanchor_23_23"></a><a href="#Footnote_23_23" class="fnanchor">[23]</a> (1749-1832), who knew Buffon's work but not Lamarck's, is
+peculiarly interesting as one of the first to use the evolution-idea
+as a guiding hypothesis, e.g. in the interpretation of vestigial
+structures in man, and to realise that organisms express an attempt to
+make a compromise between specific inertia and individual change. He
+gave the finest expression that science has yet known&mdash;if it has known
+it&mdash;of the kernel-idea of what is called "bathmism," the idea of an
+"inherent growth-force"&mdash;and at the same time he held that "the way of
+life powerfully reacts upon all form" and that the orderly growth of
+form "yields to change from externally acting causes."</p>
+
+<p><span class='pagenum'><a name="Page_15" id="Page_15">[15]</a></span></p><p>Besides Buffon, Erasmus Darwin, Lamarck, Treviranus, and Goethe,
+there were other "pioneers of evolution," whose views have been often
+discussed and appraised. &Eacute;tienne Geoffroy Saint-Hilaire (1772-1884),
+whose work Goethe so much admired, was on the whole Buffonian,
+emphasising the direct action of the changeful <i>milieu</i>. "Species vary
+with their environment, and existing species have descended by
+modification from earlier and somewhat simpler species." He had a
+glimpse of the selection idea, and believed in mutations or sudden
+leaps&mdash;induced in the embryonic condition by external influences. The
+complete history of evolution-theories will include many instances of
+guesses at truth which were afterwards substantiated, thus the
+geographer von Buch (1773-1853) detected the importance of the
+Isolation factor on which Wagner, Romanes, Gulick and others have laid
+great stress, but we must content ourselves with recalling one other
+pioneer, the author of the <i>Vestiges of Creation</i> (1844), a work which
+passed through ten editions in nine years and certainly helped to
+harrow the soil for Darwin's sowing. As Darwin said, "it did excellent
+service in this country in calling attention to the subject, in
+removing prejudice, and in thus preparing the ground for the reception
+of analogous views."<a name="FNanchor_24_24" id="FNanchor_24_24"></a><a href="#Footnote_24_24" class="fnanchor">[24]</a> Its author, Robert Chambers (1802-1871) was
+in part a Buffonian&mdash;maintaining that environment moulded organisms
+adaptively, and in part a Goethian&mdash;believing in an inherent
+progressive impulse which lifted organisms from one grade of
+organisation to another. <span class='pagenum'><a name="Page_16" id="Page_16">[16]</a></span></p>
+
+
+<p><i>As Regards Natural Selection</i></p>
+
+<p>The only thinker to whom Darwin was directly indebted, so far as the
+theory of Natural Selection is concerned, was Malthus, and we may once
+more quote the well-known passage in the Autobiography: "In October,
+1838, that is, fifteen months after I had begun my systematic enquiry,
+I happened to read for amusement 'Malthus on Population,' and being
+well prepared to appreciate the struggle for existence which
+everywhere goes on from long-continued observation of the habits of
+animals and plants, it at once struck me that under these
+circumstances favourable variations would tend to be preserved, and
+unfavourable ones to be destroyed. The result of this would be the
+formation of new species."<a name="FNanchor_25_25" id="FNanchor_25_25"></a><a href="#Footnote_25_25" class="fnanchor">[25]</a></p>
+
+<p>Although Malthus gives no adumbration of the idea of Natural Selection
+in his exposition of the eliminative processes which go on in mankind,
+the suggestive value of his essay is undeniable, as is strikingly
+borne out by the fact that it gave to Alfred Russel Wallace also "the
+long-sought clue to the effective agent in the evolution of organic
+species."<a name="FNanchor_26_26" id="FNanchor_26_26"></a><a href="#Footnote_26_26" class="fnanchor">[26]</a> One day in Ternate when he was resting between fits of
+fever, something brought to his recollection the work of Malthus which
+he had read twelve years before. "I thought of his clear exposition of
+'the positive checks to increase'&mdash;disease, accidents, war, and
+famine&mdash;which keep down the population of savage races to so much
+lower an average than that of more civilized peoples. It then occurred
+to me that these causes or their equivalents are continually acting in
+the case of animals also; and as animals usually breed much more
+rapidly than does mankind, the destruction every year from these
+causes must be enormous in order to keep down the numbers of each
+species, since they <span class='pagenum'><a name="Page_17" id="Page_17">[17]</a></span>evidently do not increase regularly from year to
+year, as otherwise the world would long ago have been densely crowded
+with those that breed most quickly. Vaguely thinking over the enormous
+and constant destruction which this implied, it occurred to me to ask
+the question, Why do some die and some live? And the answer was
+clearly, that on the whole the best fitted live. From the effects of
+disease the most healthy escaped; from enemies the strongest, the
+swiftest, or the most cunning; from famine the best hunters or those
+with the best digestion; and so on. Then it suddenly flashed upon me
+that this self-acting process would necessarily <i>improve the race</i>,
+because in every generation the inferior would inevitably be killed
+off and the superior would remain&mdash;that is, <i>the fittest would
+survive</i>."<a name="FNanchor_27_27" id="FNanchor_27_27"></a><a href="#Footnote_27_27" class="fnanchor">[27]</a> We need not apologise for this long quotation, it is a
+tribute to Darwin's magnanimous colleague, the Nestor of the
+evolutionist camp,&mdash;and it probably indicates the line of thought
+which Darwin himself followed. It is interesting also to recall the
+fact that in 1852, when Herbert Spencer wrote his famous <i>Leader</i>
+article on "The Development Hypothesis" in which he argued powerfully
+for the thesis that the whole animate world is the result of an
+age-long process of natural transformation, he wrote for <i>The
+Westminster Review</i> another important essay, "A Theory of Population
+deduced from the General Law of Animal Fertility," towards the close
+of which he came within an ace of recognising that the struggle for
+existence was a factor in organic evolution. At a time when pressure
+of population was practically interesting men's minds, Darwin,
+Wallace, and Spencer were being independently led from a social
+problem to a biological theory. There could be no better illustration,
+as Prof. Patrick Geddes has pointed out, of the Comtian thesis that
+science is a "social phenomenon."</p>
+
+<p>Therefore, as far more important than any further ferreting out of
+vague hints of Natural Selection in books which Darwin never read, we
+would indicate by a quotation the <span class='pagenum'><a name="Page_18" id="Page_18">[18]</a></span>view that the central idea in
+Darwinism is correlated with contemporary social evolution. "The
+substitution of Darwin for Paley as the chief interpreter of the order
+of nature is currently regarded as the displacement of an
+anthropomorphic view by a purely scientific one: a little reflection,
+however, will show that what has actually happened has been merely the
+replacement of the anthropomorphism of the eighteenth century by that
+of the nineteenth. For the place vacated by Paley's theological and
+metaphysical explanation has simply been occupied by that suggested to
+Darwin and Wallace by Malthus in terms of the prevalent severity of
+industrial competition, and those phenomena of the struggle for
+existence which the light of contemporary economic theory has enabled
+us to discern, have thus come to be temporarily exalted into a
+complete explanation of organic progress."<a name="FNanchor_28_28" id="FNanchor_28_28"></a><a href="#Footnote_28_28" class="fnanchor">[28]</a> It goes without saying
+that the idea suggested by Malthus was developed by Darwin into a
+biological theory which was then painstakingly verified by being used
+as an interpretative formula, and that the validity of a theory so
+established is not affected by what suggested it, but the practical
+question which this line of thought raises in the mind is this: if
+Biology did thus borrow with such splendid results from social theory,
+why should we not more deliberately repeat the experiment?</p>
+
+<p>Darwin was characteristically frank and generous in admitting that the
+principle of Natural Selection had been independently recognised by
+Dr. W. C. Wells in 1813 and by Mr. Patrick Matthew in 1831, but he had
+no knowledge of these anticipations when he published the first
+edition of <i>The Origin of Species</i>. Wells, whose "Essay on Dew" is
+still remembered, read in 1813 before the Royal Society a short paper
+entitled "An Account of a White Female, part of whose skin resembles
+that of a Negro" (published in 1818). In this communication, as Darwin
+said, "he observes, firstly, that all animals tend to vary in some
+degree, and, secondly, that agriculturists improve their domesticated
+<span class='pagenum'><a name="Page_19" id="Page_19">[19]</a></span>animals by selection; and then, he adds, but what is done in this
+latter case 'by art, seems to be done with equal efficacy, though more
+slowly, by nature, in the formation of varieties of mankind, fitted
+for the country which they inhabit.'"<a name="FNanchor_29_29" id="FNanchor_29_29"></a><a href="#Footnote_29_29" class="fnanchor">[29]</a> Thus Wells had the clear
+idea of survival dependent upon a favourable variation, but he makes
+no more use of the idea and applies it only to man. There is not in
+the paper the least hint that the author ever thought of generalising
+the remarkable sentence quoted above.</p>
+
+<p>Of Mr. Patrick Matthew, who buried his treasure in an appendix to a
+work on <i>Naval Timber and Arboriculture</i>, Darwin said that "he clearly
+saw the full force of the principle of natural selection." In 1860
+Darwin wrote&mdash;very characteristically&mdash;about this to Lyell: "Mr.
+Patrick Matthew publishes a long extract from his work on <i>Naval
+Timber and Arboriculture</i>, published in 1831, in which he briefly but
+completely anticipates the theory of Natural Selection. I have ordered
+the book, as some passages are rather obscure, but it is certainly, I
+think, a complete but not developed anticipation. Erasmus always said
+that surely this would be shown to be the case some day. Anyhow, one
+may be excused in not having discovered the fact in a work on Naval
+Timber."<a name="FNanchor_30_30" id="FNanchor_30_30"></a><a href="#Footnote_30_30" class="fnanchor">[30]</a></p>
+
+<p>De Quatrefages and De Varigny have maintained that the botanist Naudin
+stated the theory of evolution by natural selection in 1852. He
+explains very clearly the process of artificial selection, and says
+that in the garden we are following Nature's method. "We do not think
+that Nature has made her species in a different fashion from that in
+which we proceed ourselves in order to make our variations." But, as
+Darwin said, "he does not show how selection acts under nature."
+Similarly it must be noted in regard to several pre-Darwinian pictures
+of the struggle for existence (such as <span class='pagenum'><a name="Page_20" id="Page_20">[20]</a></span>Herder's, who wrote in 1790
+"All is in struggle ... each one for himself" and so on), that a
+recognition of this is only the first step in Darwinism.</p>
+
+<p>Profs. E. Perrier and H. F. Osborn have called attention to a
+remarkable anticipation of the selection-idea which is to be found in
+the speculations of &Eacute;tienne Geoffroy Saint-Hilaire (1825-1828) on the
+evolution of modern Crocodilians from the ancient Teleosaurs. Changing
+environment induced changes in the respiratory system and far-reaching
+consequences followed. The atmosphere, acting upon the pulmonary
+cells, brings about "modifications which are favourable or destructive
+('funestes'); these are inherited, and they influence all the rest of
+the organisation of the animal because if these modifications lead to
+injurious effects the animals which exhibit them perish and are
+replaced by others of a somewhat different form, a form changed so as
+to be adapted to (&agrave; la convenance) the new environment."</p>
+
+<p>Prof. E. B. Poulton<a name="FNanchor_31_31" id="FNanchor_31_31"></a><a href="#Footnote_31_31" class="fnanchor">[31]</a> has shown that the anthropologist James Cowles
+Prichard (1786-1848) must be included even in spite of himself among
+the precursors of Darwin. In some passages of the second edition of
+his <i>Researches into the Physical History of Mankind</i> (1826), he
+certainly talks evolution and anticipates Prof. Weismann in denying
+the transmission of acquired characters. He is, however, sadly
+self-contradictory and his evolutionism weakens in subsequent
+editions&mdash;the only ones that Darwin saw. Prof. Poulton finds in
+Prichard's work a recognition of the operation of Natural Selection.
+"After inquiring how it is that 'these varieties are developed and
+preserved in connexion with particular climates and differences of
+local situation,' he gives the following very significant answer: 'One
+cause which tends to maintain this relation is obvious. Individuals
+and families, and even whole colonies perish and disappear in climates
+for which they are, by peculiarity of constitution, <span class='pagenum'><a name="Page_21" id="Page_21">[21]</a></span>not adapted. Of
+this fact proofs have been already mentioned.'" Mr. Francis Darwin and
+Prof. A. C. Seward discuss Prichard's "anticipations" in <i>More Letters
+of Charles Darwin</i>, Vol. <i>I.</i> p. 43, and come to the conclusion that
+the evolutionary passages are entirely neutralised by others of an
+opposite trend. There is the same difficulty with Buffon.</p>
+
+<p>Hints of the idea of Natural Selection have been detected elsewhere.
+James Watt,<a name="FNanchor_32_32" id="FNanchor_32_32"></a><a href="#Footnote_32_32" class="fnanchor">[32]</a> for instance, has been reported as one of the
+anticipators (1851). But we need not prolong the inquiry further,
+since Darwin did not know of any anticipations until after he had
+published the immortal work of 1859, and since none of those who got
+hold of the idea made any use of it. What Darwin did was to follow the
+clue which Malthus gave him, to realise, first by genius and
+afterwards by patience, how the complex and subtle struggle for
+existence works out a natural selection of those organisms which vary
+in the direction of fitter adaptation to the conditions of their life.
+So much success attended his application of the Selection-formula that
+for a time he regarded Natural Selection as almost the sole factor in
+evolution, variations being pre-supposed; gradually, however, he came
+to recognise that there was some validity in the factors which had
+been emphasised by Lamarck and by Buffon, and in his well known
+summing up in the sixth edition of the <i>Origin</i> he says of the
+transformation of species: "This has been effected chiefly through the
+natural selection of numerous successive, slight, favourable
+variations; aided in an important manner by the inherited effects of
+the use and disuse of parts; and in an unimportant manner, that is, in
+relation to adaptive structures, whether past or present, by the
+direct action of external conditions, and by variations which seem to
+us in our ignorance to arise spontaneously."</p>
+
+<p>To sum up: the idea of organic evolution, older than Aristotle, slowly
+developed from the stage of suggestion to the stage of verification,
+and the first convincing verification <span class='pagenum'><a name="Page_22" id="Page_22">[22]</a></span>was Darwin's; from being an <i>a
+priori</i> anticipation it has become an interpretation of nature, and
+Darwin is still the chief interpreter; from being a modal
+interpretation it has advanced to the rank of a causal theory, the
+most convincing part of which men will never cease to call Darwinism.</p>
+
+<div class="footnotes"><h3>FOOTNOTES:</h3>
+
+<div class="footnote"><p><a name="Footnote_1_1" id="Footnote_1_1"></a><a href="#FNanchor_1_1"><span class="label">[1]</span></a> <i>Columbia University Biological Series</i>, Vol. I. New York
+and London, 1894. We must acknowledge our great indebtedness to this
+fine piece of work.</p></div>
+
+<div class="footnote"><p><a name="Footnote_2_2" id="Footnote_2_2"></a><a href="#FNanchor_2_2"><span class="label">[2]</span></a> <i>op. cit.</i> p. 41.</p></div>
+
+<div class="footnote"><p><a name="Footnote_3_3" id="Footnote_3_3"></a><a href="#FNanchor_3_3"><span class="label">[3]</span></a> See G. J. Romanes, "Aristotle as a Naturalist,"
+<i>Contemporary Review</i>, Vol. lix. p. 275, 1891; G. Pouchet, <i>La
+Biologie Aristot&eacute;lique</i>, Paris, 1885; E. Zeller, <i>A History of Greek
+Philosophy</i>, London, 1881, and "Ueber die griechischen Vorg&auml;nger
+Darwin's," <i>Abhandl. Berlin Akad.</i> 1878, pp. 111-124.</p></div>
+
+<div class="footnote"><p><a name="Footnote_4_4" id="Footnote_4_4"></a><a href="#FNanchor_4_4"><span class="label">[4]</span></a> <i>op. cit.</i> p. 81.</p></div>
+
+<div class="footnote"><p><a name="Footnote_5_5" id="Footnote_5_5"></a><a href="#FNanchor_5_5"><span class="label">[5]</span></a> <i>op. cit.</i> p. 87.</p></div>
+
+<div class="footnote"><p><a name="Footnote_6_6" id="Footnote_6_6"></a><a href="#FNanchor_6_6"><span class="label">[6]</span></a> See Brock, "Die Stellung Kant's zur Deszendenztheorie,"
+<i>Biol. Centralbl.</i> viii. 1889, pp. 641-648. Fritz Schultze, <i>Kant und
+Darwin</i>, Jena, 1875.</p></div>
+
+<div class="footnote"><p><a name="Footnote_7_7" id="Footnote_7_7"></a><a href="#FNanchor_7_7"><span class="label">[7]</span></a> Mr. Alfred Russel Wallace writes: "We claim for Darwin
+that he is the Newton of natural history, and that, just so surely as
+that the discovery and demonstration by Newton of the law of
+gravitation established order in place of chaos and laid a sure
+foundation for all future study of the starry heavens, so surely has
+Darwin, by his discovery of the law of natural selection and his
+demonstration of the great principle of the preservation of useful
+variations in the struggle for life, not only thrown a flood of light
+on the process of development of the whole organic world, but also
+established a firm foundation for all future study of nature"
+(<i>Darwinism</i>, London, 1889, p. 9). See also Prof. Karl Pearson's
+<i>Grammar of Science</i> (2nd edit.), London, 1900, p. 32. See Osborn,
+<i>op. cit.</i> p. 100.</p></div>
+
+<div class="footnote"><p><a name="Footnote_8_8" id="Footnote_8_8"></a><a href="#FNanchor_8_8"><span class="label">[8]</span></a> <i>Experimental Evolution</i>. London, 1892. Chap. I. p. 14.</p></div>
+
+<div class="footnote"><p><a name="Footnote_9_9" id="Footnote_9_9"></a><a href="#FNanchor_9_9"><span class="label">[9]</span></a> See J. Arthur Thomson, <i>The Science of Life</i>. London,
+1899, Chap. <span class="smcap">XVI</span>. "Evolution of Evolution Theory."</p></div>
+
+<div class="footnote"><p><a name="Footnote_10_10" id="Footnote_10_10"></a><a href="#FNanchor_10_10"><span class="label">[10]</span></a> See Carus Sterne (Ernst Krause), <i>Die allgemeine
+Weltanschauung in ihrer historischen Entwickelung</i>. Stuttgart, 1889.
+Chapter entitled "Best&auml;ndigkeit oder Ver&auml;nderlichkeit der
+Naturwesen."</p></div>
+
+<div class="footnote"><p><a name="Footnote_11_11" id="Footnote_11_11"></a><a href="#FNanchor_11_11"><span class="label">[11]</span></a> <i>Zoonomia, or the Laws of Organic Life</i>, 2 vols. London,
+1794; Osborn, <i>op. cit.</i> p. 145.</p></div>
+
+<div class="footnote"><p><a name="Footnote_12_12" id="Footnote_12_12"></a><a href="#FNanchor_12_12"><span class="label">[12]</span></a> See Alpheus S. Packard, <i>Lamarck, the Founder of
+Evolution, His Life and Work, with Translations of his writings on
+Organic Evolution</i>. London, 1901.</p></div>
+
+<div class="footnote"><p><a name="Footnote_13_13" id="Footnote_13_13"></a><a href="#FNanchor_13_13"><span class="label">[13]</span></a> See Edward Clodd, <i>Pioneers of Evolution</i>, London, p.
+161, 1897.</p></div>
+
+<div class="footnote"><p><a name="Footnote_14_14" id="Footnote_14_14"></a><a href="#FNanchor_14_14"><span class="label">[14]</span></a> See Chapter ix. "The Genetic View of Nature" in J. T.
+Merz's <i>History of European Thought in the Nineteenth Century</i>, Vol.
+2, Edinburgh and London, 1903.</p></div>
+
+<div class="footnote"><p><a name="Footnote_15_15" id="Footnote_15_15"></a><a href="#FNanchor_15_15"><span class="label">[15]</span></a> See Prof. W. A. Locy's <i>Biology and its Makers</i>. New
+York, 1908. Part II. "The Doctrine of Organic Evolution."</p></div>
+
+<div class="footnote"><p><a name="Footnote_16_16" id="Footnote_16_16"></a><a href="#FNanchor_16_16"><span class="label">[16]</span></a> Presidential Address to the British Association meeting
+at Dublin in 1908.</p></div>
+
+<div class="footnote"><p><a name="Footnote_17_17" id="Footnote_17_17"></a><a href="#FNanchor_17_17"><span class="label">[17]</span></a> See in particular Samuel Butler, <i>Evolution Old and
+New</i>, London, 1879; J. L. de Lanessan, "Buffon et Darwin," <i>Revue
+Scientifique</i>, <span class="smcap">XLIII</span>. pp. 385-391, 425-432, 1889.</p></div>
+
+<div class="footnote"><p><a name="Footnote_18_18" id="Footnote_18_18"></a><a href="#FNanchor_18_18"><span class="label">[18]</span></a> <i>op. cit.</i> p. 136.</p></div>
+
+<div class="footnote"><p><a name="Footnote_19_19" id="Footnote_19_19"></a><a href="#FNanchor_19_19"><span class="label">[19]</span></a> See Ernest Krause and Charles Darwin, <i>Erasmus Darwin</i>,
+London, 1879.</p></div>
+
+<div class="footnote"><p><a name="Footnote_20_20" id="Footnote_20_20"></a><a href="#FNanchor_20_20"><span class="label">[20]</span></a> Osborn, <i>op. cit.</i> p. 142.</p></div>
+
+<div class="footnote"><p><a name="Footnote_21_21" id="Footnote_21_21"></a><a href="#FNanchor_21_21"><span class="label">[21]</span></a> See E. Perrier, <i>La Philosophie Zoologique avant
+Darwin</i>, Paris, 1884; A. de Quatrefages, <i>Darwin et ses Pr&eacute;curseurs
+Fran&ccedil;ais</i>, Paris, 1870; Packard, <i>op. cit.</i>; also Claus, <i>Lamarck als
+Begr&uuml;nder der Descendenzlehre</i>, Wien, 1888; Haeckel, <i>Natural History
+of Creation</i>, Eng. transl. London, 1879; Lang, <i>Zur Charakteristik der
+Forschungswege von Lamarck und Darwin</i>, Jena, 1889.</p></div>
+
+<div class="footnote"><p><a name="Footnote_22_22" id="Footnote_22_22"></a><a href="#FNanchor_22_22"><span class="label">[22]</span></a> See Huxley's article "Evolution in Biology,"
+<i>Encyclopaedia Britannica</i> (9th edit.), 1879, pp. 744-751, and Sully's
+article, "Evolution in Philosophy," <i>ibid.</i> pp. 751-772.</p></div>
+
+<div class="footnote"><p><a name="Footnote_23_23" id="Footnote_23_23"></a><a href="#FNanchor_23_23"><span class="label">[23]</span></a> See Haeckel, <i>Die Naturanschauung von Darwin, Goethe und
+Lamarck</i>, Jena, 1882.</p></div>
+
+<div class="footnote"><p><a name="Footnote_24_24" id="Footnote_24_24"></a><a href="#FNanchor_24_24"><span class="label">[24]</span></a> <i>Origin of Species</i> (6th edit.), p. xvii.</p></div>
+
+<div class="footnote"><p><a name="Footnote_25_25" id="Footnote_25_25"></a><a href="#FNanchor_25_25"><span class="label">[25]</span></a> <i>The Life and Letters of Charles Darwin</i>, Vol. 1. p. 83.
+London, 1887.</p></div>
+
+<div class="footnote"><p><a name="Footnote_26_26" id="Footnote_26_26"></a><a href="#FNanchor_26_26"><span class="label">[26]</span></a> A. R. Wallace, <i>My Life, a Record of Events and
+Opinions</i>, London, 1905, Vol. 1, p. 232.</p></div>
+
+<div class="footnote"><p><a name="Footnote_27_27" id="Footnote_27_27"></a><a href="#FNanchor_27_27"><span class="label">[27]</span></a> <i>My Life</i>, Vol. 1. p. 361.</p></div>
+
+<div class="footnote"><p><a name="Footnote_28_28" id="Footnote_28_28"></a><a href="#FNanchor_28_28"><span class="label">[28]</span></a> P. Geddes. article "Biology." <i>Chambers's
+Encyclopaedia.</i></p></div>
+
+<div class="footnote"><p><a name="Footnote_29_29" id="Footnote_29_29"></a><a href="#FNanchor_29_29"><span class="label">[29]</span></a> <i>Origin of Species</i> (6th edit.), p. xv.</p></div>
+
+<div class="footnote"><p><a name="Footnote_30_30" id="Footnote_30_30"></a><a href="#FNanchor_30_30"><span class="label">[30]</span></a> <i>Life and Letters</i>, <span class="smcap">ii</span>, p. 301.</p></div>
+
+<div class="footnote"><p><a name="Footnote_31_31" id="Footnote_31_31"></a><a href="#FNanchor_31_31"><span class="label">[31]</span></a> <i>Science Progress</i>, New Series, Vol. 1. 1897. "A
+Remarkable Anticipation of Modern Views on Evolution." See also Chap.
+<span class="smcap">vi.</span> in <i>Essays on Evolution</i>, Oxford, 1908.</p></div>
+
+<div class="footnote"><p><a name="Footnote_32_32" id="Footnote_32_32"></a><a href="#FNanchor_32_32"><span class="label">[32]</span></a> See Prof. Patrick Geddes's article "Variation and
+Selection," <i>Encyclopaedia Britannica</i> (9th edit.) 1888.</p></div>
+</div>
+
+
+<hr style="width: 65%;" />
+<p><span class='pagenum'><a name="Page_23" id="Page_23">[23]</a></span></p>
+<h2><a name="II" id="II"></a>II</h2>
+
+<h2>THE SELECTION THEORY</h2>
+<p class="center"><span class="smcap1">By August Weismann</span></p>
+<h4><i>Professor of Zoology in the University of Freiburg</i> (<i>Baden</i>)</h4>
+<h3>I. <span class="smcap">The Idea of Selection</span></h3>
+<p>Many and diverse were the discoveries made by Charles Darwin in the
+course of a long and strenuous life, but none of them has had so
+far-reaching an influence on the science and thought of his time as
+the theory of selection. I do not believe that the theory of evolution
+would have made its way so easily and so quickly after Darwin took up
+the cudgels in favour of it if he had not been able to support it by a
+principle which was capable of solving, in a simple manner, the
+greatest riddle that living nature presents to us,&mdash;I mean the
+purposiveness of every living form relative to the conditions of its
+life and its marvellously exact adaptation to these.</p>
+
+<p>Everyone knows that Darwin was not alone in discovering the principle
+of selection, and that the same idea occurred simultaneously and
+independently to Alfred Russel Wallace. At the memorable meeting of
+the Linnean Society on 1st July, 1858, two papers were read
+(communicated by Lyell and Hooker) both setting forth the same idea of
+selection. One was written by Charles Darwin in Kent, the other by
+Alfred Wallace in Ternate, in the Malay Archipelago. It was a splendid
+proof of the magnanimity of these two investigators, that they thus in
+all friendliness and without envy,<span class='pagenum'><a name="Page_24" id="Page_24">[24]</a></span> united in laying their ideas
+before a scientific tribunal: their names will always shine side by
+side as two of the brightest stars in the scientific sky.</p>
+
+<p>The idea of selection set forth by the two naturalists was at the time
+absolutely new, but it was also so simple that Huxley could say of it
+later, "How extremely stupid not to have thought of that." As Darwin
+was led to the general doctrine of descent, not through the labours of
+his predecessors in the early years of the century, but by his own
+observations, so it was in regard to the principle of selection. He
+was struck by the innumerable cases of adaptation, as, for instance,
+that of the woodpeckers and tree-frogs to climbing, or the hooks and
+feather-like appendages of seeds, which aid in the distribution of
+plants, and he said to himself that an explanation of adaptations was
+the first thing to be sought for in attempting to formulate a theory
+of evolution.</p>
+
+<p>But since adaptations point to <i>changes</i> which have been undergone by
+the ancestral forms of existing species, it is necessary, first of
+all, to inquire how far species in general are <i>variable</i>. Thus
+Darwin's attention was directed in the first place to the phenomenon
+of variability, and the use man has made of this, from very early
+times, in the breeding of his domesticated animals and cultivated
+plants. He inquired carefully how breeders set to work, when they
+wished to modify the structure and appearance of a species to their
+own ends, and it was soon clear to him that <i>selection for breeding
+purposes</i> played the chief part.</p>
+
+<p>But how was it possible that such processes should occur in free
+nature? Who is here the breeder, making the selection, choosing out
+one individual to bring forth offspring and rejecting others? That was
+the problem that for a long time remained a riddle to him.</p>
+
+<p>Darwin himself relates how illumination suddenly came to him. He had
+been reading, for his own pleasure, Malthus' book on Population, and,
+as he had long known from numerous observations, that every species
+gives rise to many<span class='pagenum'><a name="Page_25" id="Page_25">[25]</a></span> more descendants than ever attain to maturity, and
+that, therefore, the greater number of the descendants of a species
+perish without reproducing, the idea came to him that the decision as
+to which member of a species was to perish and which was to attain to
+maturity and reproduction might not be a matter of chance, but might
+be determined by the constitution of the individuals themselves,
+according as they were more or less fitted for survival. With this
+idea the foundation of the theory of selection was laid.</p>
+
+<p>In <i>artificial selection</i> the breeder chooses out for pairing only
+such individuals as possess the character desired by him in a somewhat
+higher degree than the rest of the race. Some of the descendants
+inherit this character, often in a still higher degree, and if this
+method be pursued throughout several generations, the race is
+transformed in respect of that particular character.</p>
+
+<p><i>Natural selection</i> depends on the same three factors as <i>artificial
+selection</i>: on <i>variability</i>, <i>inheritance</i>, and <i>selection for
+breeding</i>, but this last is here carried out not by a breeder but by
+what Darwin called the "struggle for existence." This last factor is
+one of the special features of the Darwinian conception of nature.
+That there are carnivorous animals which take heavy toll in every
+generation of the progeny of the animals on which they prey, and that
+there are herbivores which decimate the plants in every generation had
+long been known, but it is only since Darwin's time that sufficient
+attention has been paid to the facts that, in addition to this regular
+destruction, there exists between the members of a species a keen
+competition for space and food, which limits multiplication, and that
+numerous individuals of each species perish because of unfavourable
+climatic conditions. The "struggle for existence," which Darwin
+regarded as taking the place of the human breeder in free nature, is
+not a direct struggle between carnivores and their prey, but is the
+assumed competition for survival between individuals <i>of the same</i>
+species, of which, on an average, only those survive to reproduce
+which have the greatest power<span class='pagenum'><a name="Page_26" id="Page_26">[26]</a></span> of resistance, while the others, less
+favourably constituted, perish early. This struggle is so keen, that,
+within a limited area, where the conditions of life have long remained
+unchanged, of every species, whatever be the degree of fertility, only
+two, <i>on an average</i>, of the descendants of each pair survive; the
+others succumb either to enemies, or to disadvantages of climate, or
+to accident. A high degree of fertility is thus not an indication of
+the special success of a species, but of the numerous dangers that
+have attended its evolution. Of the six young brought forth by a pair
+of elephants in the course of their lives only two survive in a given
+area; similarly, of the millions of eggs which two thread-worms leave
+behind them only two survive. It is thus possible to estimate the
+dangers which threaten a species by its ratio of elimination, or,
+since this cannot be done directly, by its fertility.</p>
+
+<p>Although a great number of the descendants of each generation fall
+victims to accident, among those that remain it is still the greater
+or less fitness of the organism that determines the "selection for
+breeding purposes," and it would be incomprehensible if, in this
+competition, it were not ultimately, that is, on an average, the best
+equipped which survive, in the sense of living long enough to
+reproduce.</p>
+
+<p>Thus the principle of natural selection is <i>the selection of the best
+for reproduction</i>, whether the "best" refers to the whole
+constitution, to one or more parts of the organism, or to one or more
+stages of development. Every organ, every part, every character of an
+animal, fertility and intelligence included, must be improved in this
+manner, and be gradually brought up in the course of generations to
+its highest attainable state of perfection. And not only may
+improvement of parts be brought about in this way, but new parts and
+organs may arise, since, through the slow and minute steps of
+individual or "fluctuating" variations, a part may be added here or
+dropped out there, and thus something new is produced.</p>
+
+<p>The principle of selection solved the riddle as to how<span class='pagenum'><a name="Page_27" id="Page_27">[27]</a></span> what was
+purposive could conceivably be brought about without the intervention
+of a directing power, the riddle which animate nature presents to our
+intelligence at every turn, and in face of which the mind of a Kant
+could find no way out, for he regarded a solution of it as not to be
+hoped for. For, even if we were to assume an evolutionary force that
+is continually transforming the most primitive and the simplest forms
+of life into ever higher forms, and the homogeneity of primitive times
+into the infinite variety of the present, we should still be unable to
+infer from this alone how each of the numberless forms adapted to
+particular conditions of life should have appeared <i>precisely at the
+right moment in the history of the earth</i> to which their adaptations
+were appropriate, and precisely at the proper place in which all the
+conditions of life to which they were adapted occurred: the
+humming-birds at the same time as the flowers; the trichina at the
+same time as the pig; the bark-coloured moth at the same time as the
+oak, and the wasp-like moth at the same time as the wasp which
+protects it. Without processes of selection we should be obliged to
+assume a "pre-established harmony" after the famous Leibnitzian model,
+by means of which the clock of the evolution of organisms is so
+regulated as to strike in exact synchronism with that of the history
+of the earth! All forms of life are strictly adapted to the conditions
+of their life, and can persist under these conditions alone.</p>
+
+<p>There must therefore be an intrinsic connection between the conditions
+and the structural adaptations of the organism, and, <i>since the
+conditions of life cannot be determined by the animal itself, the
+adaptations must be called forth by the conditions</i>.</p>
+
+<p>The selection theory teaches us how this is conceivable, since it
+enables us to understand that there is a continual production of what
+is non-purposive as well as of what is purposive, but the purposive
+alone survives, while the non-purposive perishes in the very act of
+arising. This is the old wisdom taught long ago by Empedocles.<span class='pagenum'><a name="Page_28" id="Page_28">[28]</a></span></p>
+
+
+<p>II. <span class="smcap">The Lamarckian Principle</span></p>
+
+<p>Lamarck, as is well known, formulated a definite theory of evolution
+at the beginning of the nineteenth century, exactly fifty years before
+the Darwin-Wallace principle of selection was given to the world. This
+brilliant investigator also endeavoured to support his theory by
+demonstrating forces which might have brought about the
+transformations of the organic world in the course of the ages. In
+addition to other factors, he laid special emphasis on the increased
+or diminished use of the parts of the body, assuming that the
+strengthening or weakening which takes place from this cause during
+the individual life, could be handed on to the offspring, and thus
+intensified and raised to the rank of a specific character. Darwin
+also regarded this <i>Lamarckian principle</i>, as it is now generally
+called, as a factor in evolution, but he was not fully convinced of
+the transmissibility of acquired characters.</p>
+
+<p>As I have here to deal only with the theory of selection, I need not
+discuss the Lamarckian hypothesis, but I must express my opinion that
+there is room for much doubt as to the co&ouml;peration of this principle
+in evolution. Not only is it difficult to imagine how the transmission
+of functional modifications could take place, but, up to the present
+time, notwithstanding the endeavours of many excellent investigators,
+not a single actual proof of such inheritance has been brought
+forward. Semon's experiments on plants are, according to the botanist
+Pfeffer, not to be relied on, and even the recent, beautiful
+experiments made by Dr. Kammerer on salamanders, cannot, as I hope to
+show elsewhere, be regarded as proof, if only because they do not deal
+at all with functional modifications, that is, with modifications
+brought about by use, and it is to these <i>alone</i> that the Lamarckian
+principle refers.<span class='pagenum'><a name="Page_29" id="Page_29">[29]</a></span></p>
+
+
+<p>III. <span class="smcap">Objections to the Theory of Selection</span></p>
+
+
+<p>(<i>a</i>) <i>Saltatory evolution</i></p>
+
+<p>The Darwinian doctrine of evolution depends essentially on <i>the
+cumulative augmentation</i> of minute variations in the direction of
+utility. But can such minute variations, which are undoubtedly
+continually appearing among the individuals of the same species,
+possess any selection-value; can they determine which individuals are
+to survive, and which are to succumb; can they be increased by natural
+selection till they attain to the highest development of a purposive
+variation?</p>
+
+<p>To many this seems so improbable that they have urged a theory of
+evolution by leaps from species to species. K&ouml;lliker, in 1872,
+compared the evolution of species with the processes which we can
+observe in the individual life in cases of alternation of generations.
+But a polyp only gives rise to a medusa because it has itself arisen
+from one, and there can be no question of a medusa ever having arisen
+suddenly and <i>de novo</i> from a polyp-bud, if only because both forms
+are adapted in their structure as a whole, and in every detail to the
+conditions of their life. A sudden origin, in a natural way, of
+numerous adaptations is inconceivable. Even the degeneration of a
+medusoid from a free-swimming animal to a mere brood-sac (gonophore)
+is not sudden and saltatory, but occurs by imperceptible modifications
+throughout hundreds of years, as we can learn from the numerous stages
+of the process of degeneration persisting at the same time in
+different species.</p>
+
+<p>If, then, the degeneration to a simple brood-sac takes place only by
+very slow transitions, each stage of which may last for centuries, how
+could the much more complex <i>ascending</i> evolution possibly have taken
+place by sudden leaps? I regard this argument as capable of further
+extension, for wherever in nature we come upon degeneration, it<span class='pagenum'><a name="Page_30" id="Page_30">[30]</a></span> is
+taking place by minute steps and with a slowness that makes it not
+directly perceptible, and I believe that this in itself justifies us
+in concluding that <i>the same must be true of ascending</i> evolution. But
+in the latter case the goal can seldom be distinctly recognised while
+in cases of degeneration the starting-point of the process can often
+be inferred, because several nearly related species may represent
+different stages.</p>
+
+<p>In recent years Bateson in particular has championed the idea of
+saltatory, or so-called discontinuous evolution, and has collected a
+number of cases in which more or less marked variations have suddenly
+appeared. These are taken for the most part from among domesticated
+animals which have been bred and crossed for a long time, and it is
+hardly to be wondered at that their much mixed and much influenced
+germ-plasm should, under certain conditions, give rise to remarkable
+phenomena, often indeed producing forms which are strongly suggestive
+of monstrosities, and which would undoubtedly not survive in free
+nature, unprotected by man. I should regard such cases as due to an
+intensified germinal selection&mdash;though this is to anticipate a
+little&mdash;and from this point of view it cannot be denied that they have
+a special interest. But they seem to me to have no significance as far
+as the transformation of species is concerned, if only because of the
+extreme rarity of their occurrence.</p>
+
+<p>There are, however, many variations which have appeared in a sudden
+and saltatory manner, and some of these Darwin pointed out and
+discussed in detail: the copper beech, the weeping trees, the oak with
+"fern-like leaves," certain garden-flowers, etc. But none of them have
+persisted in free nature, or evolved into permanent types.</p>
+
+<p>On the other hand, wherever enduring types have arisen, we find traces
+of a gradual origin by successive stages, even if, at first sight,
+their origin may appear to have been sudden. This is the case with
+<i>seasonal Dimorphism</i>, the first known cases of which exhibited marked
+differences between the two generations, the winter and the summer
+brood. Take for<span class='pagenum'><a name="Page_31" id="Page_31">[31]</a></span> instance the much discussed and studied form
+<i>Vanessa</i> (<i>Araschnia</i>) <i>levana-prorsa</i>. Here the differences between
+the two forms are so great and so apparently disconnected, that one
+might almost believe it to be a sudden mutation, were it not that old
+transition-stages can be called forth by particular temperatures, and
+we know other butterflies, as for instance our Garden Whites, in which
+the differences between the two generations are not nearly so marked;
+indeed, they are so little apparent that they are scarcely likely to
+be noticed except by experts. Thus here again there are small initial
+steps, some of which, indeed, must be regarded as adaptations, such as
+the green-sprinkled or lightly tinted under-surface which gives them a
+deceptive resemblance to parsley or to Cardamine leaves.</p>
+
+<p>Even if saltatory variations do occur, we cannot assume that these
+<i>have ever led to forms which are capable of survival under the
+conditions of wild life</i>. Experience has shown that in plants which
+have suddenly varied the power of persistence is diminished.
+Korschinsky attributes to them weaknesses of organisation in general;
+"they bloom late, ripen few of their seeds, and show great
+sensitiveness to cold." These are not the characters which make for
+success in the struggle for existence.</p>
+
+<p>We must briefly refer here to the views&mdash;much discussed in the last
+decade&mdash;of H. de Vries, who believes that the roots of transformation
+must be sought for in <i>saltatory variations arising from internal
+causes</i>, and distinguishes such <i>mutations</i>, as he has called them,
+from ordinary individual variations, in that they breed true, that is,
+with strict in-breeding they are handed on pure to the next
+generation. I have elsewhere endeavoured to point out the weaknesses
+of this theory,<a name="FNanchor_33_33" id="FNanchor_33_33"></a><a href="#Footnote_33_33" class="fnanchor">[33]</a> and I am the less inclined to return to it here
+that it now appears<a name="FNanchor_34_34" id="FNanchor_34_34"></a><a href="#Footnote_34_34" class="fnanchor">[34]</a> that the far-reaching conclusions drawn by de
+Vries from his observations on the Evening Primrose, <span class='pagenum'><a name="Page_32" id="Page_32">[32]</a></span><i>Oenothera
+lamarckiana</i>, rest upon a very insecure foundation. The plant from
+which de Vries saw numerous "species"&mdash;his "mutations"&mdash;arise was not,
+as he assumed, a <i>wild species</i> that had been introduced to Europe
+from America, but was probably a hybrid form which was first
+discovered in the Jardin des Plantes in Paris, and which does not
+appear to exist anywhere in America as a wild species.</p>
+
+<p>This gives a severe shock to the "Mutation theory," for the other
+<i>actually wild</i> species with which de Vries experimented showed no
+"mutations" but yielded only negative results.</p>
+
+<p>Thus we come to the conclusion that Darwin<a name="FNanchor_35_35" id="FNanchor_35_35"></a><a href="#Footnote_35_35" class="fnanchor">[35]</a> was right in regarding
+transformations as taking place by minute steps, which, if useful, are
+augmented in the course of innumerable generations, because their
+possessors more frequently survive in the struggle for existence.</p>
+
+
+<p>(<i>b</i>) <i>Selection-value of the initial steps</i></p>
+
+<p>Is it possible that the insignificant deviations which we know as
+"individual variations" can form the beginning of a process of
+selection? Can they decide which is to perish and which to survive? To
+use a phrase of Romanes, can they have <i>selection-value</i>?</p>
+
+<p>Darwin himself answered this question, and brought together many
+excellent examples to show that differences, apparently insignificant
+because very small, might be of decisive importance for the life of
+the possessor. But it is by no means enough to bring forward cases of
+this kind, for the question is not merely whether finished adaptations
+have selection-value, but whether the first beginnings of these, and
+whether the small, I might almost say minimal increments, which have
+led up from these beginnings to the perfect adaptation, have also had
+selection-value. To this question even one who, like myself, has been
+for many years a convinced adherent of the theory of selection, can
+only <span class='pagenum'><a name="Page_33" id="Page_33">[33]</a></span>reply: <i>We must assume so, but we cannot prove it in any case</i>.
+It is not upon demonstrative evidence that we rely when we champion
+the doctrine of selection as a scientific truth; we base our argument
+on quite other grounds. Undoubtedly there are many apparently
+insignificant features, which can nevertheless be shown to be
+adaptations&mdash;for instance, the thickness of the basin-shaped shell of
+the limpets that live among the breakers on the shore. There can be no
+doubt that the thickness of these shells, combined with their flat
+forms, protects the animals from the force of the waves breaking upon
+them,&mdash;but how have they become so thick? What proportion of thickness
+was sufficient to decide that of two variants of a limpet one should
+survive, the other be eliminated? We can say nothing more than that we
+infer from the present state of the shell, that it must have varied in
+regard to differences in shell-thickness, and that these differences
+must have had selection-value,&mdash;no proof therefore, but an assumption
+which we must show to be convincing.</p>
+
+<p>For a long time the marvellously complex <i>radiate</i> and <i>lattice-work</i>
+skeletons of Radiolarians were regarded as a mere outflow of "Nature's
+infinite wealth of form," as an instance of a purely morphological
+character with no biological significance. But recent investigations
+have shown that these, too, have an adaptive significance (H&auml;cker).
+The same thing has been shown by Sch&uuml;tt in regard to the lowly
+unicellular plants, the Peridineae, which abound alike on the surface
+of the ocean and in its depths. It has been shown that the long
+skeletal processes which grow out from these organisms have
+significance not merely as a supporting skeleton, but also as an
+extension of the superficial area, which increases the contact with
+the water-particles, and prevents the floating organisms from sinking.
+It has been established that the processes are considerably shorter in
+the colder layers of the ocean, and that they may be twelve times as
+long<a name="FNanchor_36_36" id="FNanchor_36_36"></a><a href="#Footnote_36_36" class="fnanchor">[36]</a> in the warmer layers, thus corresponding to <span class='pagenum'><a name="Page_34" id="Page_34">[34]</a></span>the greater or
+smaller amount of friction which takes place in the denser and less
+dense layers of the water.</p>
+
+<p>The Peridineae of the warmer ocean layers have thus become long-rayed,
+those of the colder layers short-rayed, not through the direct effect
+of friction on the protoplasm, but through processes of selection,
+which favoured the longer rays in warm water, since they kept the
+organism afloat, while those with short rays sank and were eliminated.
+If we put the question as to selection-value in this case, and ask how
+great the variations in the length of processes must be in order to
+possess selection-value; what can we answer except that these
+variations must have been minimal, and yet sufficient to prevent too
+rapid sinking and consequent elimination? Yet this very case would
+give the ideal opportunity for a mathematical calculation of the
+minimal selection-value, although of course it is not feasible from
+lack of data to carry out the actual calculation.</p>
+
+<p>But even in organisms of more than microscopic size there must
+frequently be minute, even microscopic differences which set going the
+process of selection, and regulate its progress to the highest
+possible perfection.</p>
+
+<p>Many tropical trees possess thick, leathery leaves, as a protection
+against the force of the tropical raindrops. The <i>direct</i> influence of
+the rain cannot be the cause of this power of resistance, for the
+leaves, while they were still thin, would simply have been torn to
+pieces. Their toughness must therefore be referred to selection, which
+would favour the trees with slightly thicker leaves, though we cannot
+calculate with any exactness how great the first stages of increase in
+thickness must have been. Our hypothesis receives further support from
+the fact that, in many such trees, the leaves are drawn out into a
+beak-like prolongation (Stahl and Haberlandt) which facilitates the
+rapid falling off of the rain water, and also from the fact that the
+leaves, while they are still young, hang limply down in bunches which
+offer the least possible resistance to the rain. Thus there are here
+three adaptations which can only be interpreted as<span class='pagenum'><a name="Page_35" id="Page_35">[35]</a></span> due to selection.
+The initial stages of these adaptations must undoubtedly have had
+selection-value.</p>
+
+<p>But even in regard to this case we are reasoning in a circle, not
+giving "proofs," and no one who does not wish to believe in the
+selection-value of the initial stages can be forced to do so. Among
+the many pieces of presumptive evidence a particularly weighty one
+seems to me to be <i>the smallness of the steps of progress</i> which we
+can observe in certain cases, as for instance in leaf-imitation among
+butterflies, and in mimicry generally. The resemblance to a leaf, for
+instance of a particular Kallima, seems to us so close as to be
+deceptive, and yet we find in another individual, or it may be in many
+others, a spot added which increases the resemblance, and which could
+not have become fixed unless the increased deceptiveness so produced
+had frequently led to the overlooking of its much persecuted
+possessor. But if we take the selection-value of the initial stages
+for granted, we are confronted with the further question which I
+myself formulated many years ago: How does it happen <i>that the
+necessary beginnings of a useful variation are always present</i>? How
+could insects which live upon or among green leaves become all green,
+while those that live on bark become brown? How have the desert
+animals become yellow and the Arctic animals white? Why were the
+necessary variations always present? How could the green locust lay
+brown eggs, or the privet caterpillar develop white and lilac-coloured
+lines on its green skin?</p>
+
+<p>It is of no use answering to this that the question is wrongly
+formulated<a name="FNanchor_37_37" id="FNanchor_37_37"></a><a href="#Footnote_37_37" class="fnanchor">[37]</a> and that it is the converse that is true; that the
+process of selection takes place in accordance with the variations
+that present themselves. This proposition is undeniably true, but so
+also is another, which apparently negatives it: the variation required
+has in the majority of cases actually presented itself. Selection
+cannot solve this contradiction; it does not call forth the useful
+variation, but <span class='pagenum'><a name="Page_36" id="Page_36">[36]</a></span>simply works upon it. The ultimate reason why one and
+the same insect should occur in green and in brown, as often happens
+in caterpillars and locusts, lies in the fact that variations towards
+brown presented themselves, and so also did variations towards green:
+<i>the kernel of the riddle lies in the varying</i>, and for the present we
+can only say, that small variations in different directions present
+themselves in every species. Otherwise so many different kinds of
+variations could not have arisen. I have endeavoured to explain this
+remarkable fact by means of the intimate processes that must take
+place within the germ-plasm, and I shall return to the problem when
+dealing with "germinal selection."</p>
+
+<p>We have, however, to make still greater demands on variation, for it
+is not enough that the necessary variation should occur in isolated
+individuals, because in that case there would be small prospect of its
+being preserved, notwithstanding its utility. Darwin at first
+believed, that even single variations might lead to transformation of
+the species, but later he became convinced that this was impossible,
+at least without the co&ouml;peration of other factors, such as isolation
+and sexual selection.</p>
+
+<p>In the case of the <i>green caterpillars with bright longitudinal
+stripes</i>, numerous individuals exhibiting this useful variation must
+have been produced to start with. In all higher, that is,
+multicellular organisms, the germ-substance is the source of all
+transmissible variations, and this germ-plasm is not a simple
+substance but is made up of many primary constituents. The question
+can therefore be more precisely stated thus: How does it come about
+that in so many cases the useful variations present themselves in
+numbers just where they are required, the white oblique lines in the
+leaf-caterpillar on the under surface of the body, the accompanying
+coloured stripes just above them? And, further, how has it come about
+that in grass caterpillars, not oblique but longitudinal stripes,
+which are more effective for concealment among grass and plants, have
+been evolved?<span class='pagenum'><a name="Page_37" id="Page_37">[37]</a></span> And finally, how is it that the same Hawk-moth
+caterpillars, which to-day show oblique stripes, possessed
+longitudinal stripes in Tertiary times? We can read this fact from the
+history of their development, and I have before attempted to show the
+biological significance of this change of colour.<a name="FNanchor_38_38" id="FNanchor_38_38"></a><a href="#Footnote_38_38" class="fnanchor">[38]</a></p>
+
+<p>For the present I need only draw the conclusion that one and the same
+caterpillar may exhibit the initial stages of both, and that it
+depends on the manner in which these marking elements are
+<i>intensified</i> and <i>combined</i> by natural selection whether whitish
+longitudinal or oblique stripes should result. In this case then the
+"useful variations" were actually "always there," and we see that in
+the same group of Lepidoptera, e.g. species of Sphingidae, evolution
+has occurred in both directions according to whether the form lived
+among grass or on broad leaves with oblique lateral veins, and we can
+observe even now that the species with oblique stripes have
+longitudinal stripes when young, that is to say, while the stripes
+have no biological significance. The white places in the skin which
+gave rise, probably first as small spots, to this protective marking
+could be combined in one way or another according to the requirements
+of the species. They must therefore either have possessed
+selection-value from the first, or, if this was not the case at their
+earliest occurrence, there must have been <i>some other factors</i> which
+raised them to the point of selection-value. I shall return to this in
+discussing germinal selection. But the case may be followed still
+farther, and leads us to the same alternative on a still more secure
+basis.</p>
+
+<p>Many years ago I observed in caterpillars of <i>Smerinthus populi</i> (the
+poplar hawk-moth), which also possess white oblique stripes, that
+certain individuals showed <i>red spots</i> above these stripes; these
+spots occurred only on certain segments, and never flowed together to
+form continuous stripes. In another species (<i>Smerinthus tiliae</i>)
+similar blood-red spots unite to form a line-like coloured seam in the
+last <span class='pagenum'><a name="Page_38" id="Page_38">[38]</a></span>stage of larval life, while in <i>S. ocellata</i> rust-red spots
+appear in individual caterpillars, but more rarely than in <i>S.
+populi</i>, and they show no tendency to flow together.</p>
+
+<p>Thus we have here the origin of a new character, arising from small
+beginnings, at least in <i>S. tiliae</i>, in which species the coloured
+stripes are a normal specific character. In the other species, <i>S.
+populi</i> and <i>S. ocellata</i>, we find the beginnings of the same
+variation, in one more rarely than in the other, and we can imagine
+that, in the course of time, in these two species, coloured lines over
+the oblique stripes will arise. In any case these spots are the
+elements of variation, out of which coloured lines <i>may</i> be evolved,
+if they are combined in this direction through the agency of natural
+selection. In <i>S. populi</i> the spots are often small, but sometimes it
+seems as though several had united to form large spots. Whether a
+process of selection in this direction will arise in <i>S. populi</i> and
+<i>S. ocellata</i>, or whether it is now going on cannot be determined,
+since we cannot tell in advance what biological value the marking
+might have for these two species. It is conceivable that the spots may
+have no selection-value as far as these species are concerned, and may
+therefore disappear again in the course of phylogeny, or, on the other
+hand, that they may be changed in another direction, for instance
+towards imitation of the rust-red fungoid patches on poplar and willow
+leaves. In any case we may regard the smallest spots as the initial
+stages of variation, the larger as a cumulative summation of these.
+Therefore either these initial stages must already possess
+selection-value, or, as I said before: <i>There must be some other
+reason for their cumulative summation</i>. I should like to give one more
+example, in which we can infer, though we cannot directly observe, the
+initial stages.</p>
+
+<p>All the Holothurians or sea-cucumbers have in the skin calcereous
+bodies of different forms, usually thick and irregular, which make the
+skin tough and resistant. In a small group of them&mdash;the species of
+Synapta&mdash;the calcareous bodies occur in the form of delicate anchors
+of microscopic<span class='pagenum'><a name="Page_39" id="Page_39">[39]</a></span> size. Up till 1897 these anchors, like many other
+delicate microscopic structures, were regarded as curiosities, as
+natural marvels. But a Swedish observer, Oestergren, has recently
+shown that they have a biological significance: they serve the
+footless Synapta as auxiliary organs of locomotion, since, when the
+body swells up in the act of creeping, they press firmly with their
+tips, which are embedded in the skin, against the substratum on which
+the animal creeps, and thus prevent slipping backwards. In other
+Holothurians this slipping is made impossible by the fixing of the
+tube-feet. The anchors act automatically, sinking their tips towards
+the ground when the corresponding part of the body thickens, and
+returning to the original position at an angle of 45 degrees to the
+upper surface when the part becomes thin again. The arms of the anchor
+do not lie in the same plane as the shaft, and thus the curve of the
+arms forms the outermost part of the anchor, and offers no further
+resistance to the gliding of the animal. Every detail of the anchor,
+the curved portion, the little teeth at the head, the arms, etc., can
+be interpreted in the most beautiful way, above all the form of the
+anchor itself, for the two arms prevent it from swaying round to the
+side. The position of the anchors, too, is definite and significant;
+they lie obliquely to the longitudinal axis of the animal, and
+therefore they act alike whether the animal is creeping backwards or
+forwards. Moreover, the tips would pierce through the skin if the
+anchors lay in the longitudinal direction. Synapta burrows in the
+sand; it first pushes in the thin anterior end, and thickens this
+again, thus enlarging the hole, then the anterior tentacles displace
+more sand, the body is worked in a little farther, and the process
+begins anew. In the first act the anchors are passive, but they begin
+to take an active share in the forward movement when the body is
+contracted again. Frequently the animal retains only the posterior end
+buried in the sand, and then the anchors keep it in position, and make
+rapid withdrawal possible.</p>
+
+<p>Thus we have in these apparently random forms of the<span class='pagenum'><a name="Page_40" id="Page_40">[40]</a></span> calcereous
+bodies, complex adaptations in which every little detail as to
+direction, curve, and pointing is exactly determined. That they have
+selection-value in their present perfected form is beyond all doubt,
+since the animals are enabled by means of them to bore rapidly into
+the ground and so to escape from enemies. We do not know what the
+initial stages were, but we cannot doubt that the little improvements,
+which occurred as variations of the originally simple slimy bodies of
+the Holothurians, were preserved because they already possessed
+selection-value for the Synaptidae. For such minute microscopic
+structures whose form is so delicately adapted to the r&ocirc;le they have
+to play in the life of the animal, cannot have arisen suddenly and as
+a whole, and every new variation of the anchor, that is, in the
+direction of the development of the two arms, and every curving of the
+shaft which prevented the tips from projecting at the wrong time, in
+short, every little adaptation in the modelling of the anchor must
+have possessed selection-value. And that such minute changes of form
+fall within the sphere of fluctuating variations, that is to say,
+<i>that they occur</i> is beyond all doubt.</p>
+
+<p>In many of the Synaptidae the anchors are replaced by calcareous rods
+bent in the form of an S, which are said to act in the same way.
+Others, such as those of the genus Ankyroderma, have anchors which
+project considerably beyond the skin, and, according to Oestergren,
+serve "to catch plant-particles and other substances" and so mask the
+animal. Thus we see that in the Synaptidae the thick and irregular
+calcareous bodies of the Holothurians have been modified and
+transformed in various ways in adaptation to the footlessness of these
+animals, and to the peculiar conditions of their life, and we must
+conclude that the earlier stages of these changes presented themselves
+to the processes of selection in the form of microscopic variations.
+For it is as impossible to think of any origin other than through
+selection in this case as in the case of the toughness, and the
+"drip-tips" of tropical leaves. And as these last could not<span class='pagenum'><a name="Page_41" id="Page_41">[41]</a></span> have been
+produced directly by the beating of the heavy raindrops upon them, so
+the calcareous anchors of Synapta cannot have been produced directly
+by the friction of the sand and mud at the bottom of the sea, and,
+since they are parts whose function is <i>passive</i> the Lamarckian factor
+of use and disuse does not come into question. The conclusion is
+unavoidable, that the microscopically small variations of the
+calcareous bodies in the ancestral forms have been intensified and
+accumulated in a particular direction, till they have led to the
+formation of the anchor. Whether this has taken place by the action of
+natural selection alone, or whether the laws of variation and the
+intimate processes within the germ-plasm have co&ouml;perated will become
+clear in the discussion of germinal selection. This whole process of
+adaptation has obviously taken place within the time that has elapsed
+since this group of sea-cucumbers lost their tube-feet, those
+characteristic organs of locomotion which occur in no group except the
+Echinoderms, and yet have totally disappeared in the Synaptidae. And
+after all what would animals that live in sand and mud do with
+tube-feet?</p>
+
+
+<p>(<i>c</i>) <i>Coadaptation</i></p>
+
+<p>Darwin pointed out that one of the essential differences between
+artificial and natural selection lies in the fact that the former can
+modify only a few characters, usually only one at a time, while Nature
+preserves in the struggle for existence all the variations of a
+species, at the same time and in a purely mechanical way, if they
+possess selection-value.</p>
+
+<p>Herbert Spencer, though himself an adherent of the theory of
+selection, declared in the beginning of the nineties that in his
+opinion the range of this principle was greatly over-estimated, if the
+great changes which have taken place in so many organisms in the
+course of ages are to be interpreted as due to this process of
+selection alone, since no transformation of any importance can be
+evolved by itself; it is always accompanied by a host of secondary
+changes. He gives the<span class='pagenum'><a name="Page_42" id="Page_42">[42]</a></span> familiar example of the Giant Stag of the Irish
+peat, the enormous antlers of which required not only a much stronger
+skull cap, but also greater strength of the sinews, muscles, nerves
+and bones of the whole anterior half of the animal, if their mass was
+not to weigh down the animal altogether. It is inconceivable, he says,
+that so many processes of selection should take place
+<i>simultaneously</i>, and we are therefore forced to fall back on the
+Lamarckian factor of the use and disuse of functional parts. And how,
+he asks, could natural selection follow two opposite directions of
+evolution in different parts of the body at the same time, as for
+instance in the case of the kangaroo, in which the forelegs must have
+become shorter, while the hind legs and the tail were becoming longer
+and stronger?</p>
+
+<p>Spencer's main object was to substantiate the validity of the
+Lamarckian principle, the co&ouml;peration of which with selection had been
+doubted by many. And it does seem as though this principle, if it
+operates in nature at all, offers a ready and simple explanation of
+all such secondary variations. Not only muscles, but nerves, bones,
+sinews, in short all tissues which function actively, increase in
+strength in proportion as they are used, and conversely they decrease
+when the claims on them diminish. All the parts, therefore, which
+depend on the part that varied first, as for instance the enlarged
+antlers of the Irish Elk, must have been increased or decreased in
+strength, in exact proportion to the claims made upon them,&mdash;just as
+is actually the case.</p>
+
+<p>But beautiful as this explanation would be, I regard it as untenable,
+because it assumes the <i>transmissibility of functional modifications</i>
+(so-called "acquired" characters), and this is not only
+undemonstrable, but is scarcely theoretically conceivable, for the
+secondary variations which accompany or follow the first as
+correlative variations, occur also in cases in which the animals
+concerned are sterile and <i>therefore cannot transmit anything to their
+descendants</i>. This is true of <i>worker bees</i>, and particularly of
+<i>ants</i>, and I shall here<span class='pagenum'><a name="Page_43" id="Page_43">[43]</a></span> give a brief survey of the present state of
+the problem as it appears to me.</p>
+
+<p>Much has been written on both sides of this question since the
+published controversy on the subject in the nineties between Herbert
+Spencer and myself. I should like to return to the matter in detail,
+if the space at my disposal permitted, because it seems to me that the
+arguments I advanced at that time are equally cogent to-day,
+notwithstanding all the objections that have since been urged against
+them. Moreover, the matter is by no means one of subordinate interest;
+it is the very kernel of the whole question of the reality and value
+of the principle of selection. For if selection alone does not suffice
+to explain "<i>harmonious adaptation</i>" as I have called Spencer's
+<i>Coadaptation</i>, and if we require to call in the aid of the Lamarckian
+factor it would be questionable whether selection would explain any
+adaptations whatever. In this particular case&mdash;of worker bees&mdash;the
+Lamarckian factor may be excluded altogether, for it can be
+demonstrated that here at any rate the effects of use and disuse
+cannot be transmitted.</p>
+
+<p>But if it be asked why we are unwilling to admit the co&ouml;peration of
+the Darwinian factor of selection and the Lamarckian factor, since
+this would afford us an easy and satisfactory explanation of the
+phenomena, I answer: <i>Because the Lamarckian principle is fallacious,
+and because by accepting it we close the way towards deeper insight</i>.
+It is not a spirit of combativeness or a desire for self-vindication
+that induces me to take the field once more against the Lamarckian
+principle, it is the conviction that the progress of our knowledge is
+being obstructed by the acceptance of this fallacious principle, since
+the facile explanation it apparently affords prevents our seeking
+after a truer explanation and a deeper analysis.</p>
+
+<p>The workers in the various species of ants are sterile, that is to
+say, they take no regular part in the reproduction of the species,
+although individuals among them may occasionally lay eggs. In addition
+to this they have lost the wings, and<span class='pagenum'><a name="Page_44" id="Page_44">[44]</a></span> the <i>receptaculum seminis</i>, and
+their compound eyes have degenerated to a few facets. How could this
+last change have come about through disuse, since the eyes of workers
+are exposed to light in the same way as are those of the sexual
+insects and thus in this particular case are not liable to "disuse" at
+all? The same is true of the <i>receptaculum seminis</i>, which can only
+have been disused as far as its glandular portion and its stalk are
+concerned, and also of the wings, the nerves tracheae and epidermal
+cells of which could not cease to function until the whole wing had
+degenerated, for the chitinous skeleton of the wing does not function
+at all in the active sense.</p>
+
+<p>But, on the other hand, the workers in all species have undergone
+modifications in a positive direction, as, for instance, the greater
+development of brain. In many species large workers have evolved,&mdash;the
+so-called <i>soldiers</i>, with enormous jaws and teeth, which defend the
+colony,&mdash;and in others there are <i>small</i> workers which have taken over
+other special functions, such as the rearing of the young Aphides.
+This kind of division of the workers into two castes occurs among
+several tropical species of ants, but it is also present in the
+Italian species, <i>Colobopsis truncata</i>. Beautifully as the size of the
+jaws could be explained as due to the increased use made of them by
+the "soldiers," or the enlarged brain as due to the mental activities
+of the workers, the fact of the infertility of these forms is an
+insurmountable obstacle to accepting such an explanation. Neither jaws
+nor brain can have been evolved on the Lamarckian principle.</p>
+
+<p>The problem of coadaptation is no easier in the case of the ant than
+in the case of the Giant Stag. Darwin himself gave a pretty
+illustration to show how imposing the difference between the two kinds
+of workers in one species would seem if we translated it into human
+terms. In regard to the Driver ants (Anomma) we must picture to
+ourselves a piece of work, "for instance the building of a house,
+being carried on by two kinds of workers, of which one group was five
+feet four inches high, the other sixteen feet high."<a name="FNanchor_39_39" id="FNanchor_39_39"></a><a href="#Footnote_39_39" class="fnanchor">[39]</a> <span class='pagenum'><a name="Page_45" id="Page_45">[45]</a></span></p>
+
+<p>Although the ant is a small animal as compared with man or with the
+Irish Elk, the "soldier" with its relatively enormous jaws is hardly
+less heavily burdened than the Elk with its antlers, and in the ant's
+case, too, a strengthening of the skeleton, of the muscles, the nerves
+of the head, and of the legs must have taken place parallel with the
+enlargement of the jaws. <i>Harmonious adaptation</i> (coadaptation) has
+here been active in a high degree, and yet these "soldiers" are
+sterile! There thus remains nothing for it but to refer all their
+adaptations, positive and negative alike, to processes of selection
+which have taken place in the rudiments of the workers within the egg
+and sperm-cells of their parents. There is no way out of the
+difficulty except the one Darwin pointed out. He himself did not find
+the solution of the riddle at once. At first he believed that the case
+of the workers among social insects presented "the most serious
+special difficulty" in the way of his theory of natural selection; and
+it was only after it had become clear to him that it was not the
+sterile insects themselves but their parents that were selected,
+according as they produced more or less well adapted workers, that he
+was able to refer to this very case of the conditions among ants "<i>in
+order to show the power of natural selection</i>."<a name="FNanchor_40_40" id="FNanchor_40_40"></a><a href="#Footnote_40_40" class="fnanchor">[40]</a> He explains his
+view by a simple but interesting illustration. Gardeners have
+produced, by means of long continued artificial selection, a variety
+of Stock, which bears entirely double, and therefore infertile
+flowers.<a name="FNanchor_41_41" id="FNanchor_41_41"></a><a href="#Footnote_41_41" class="fnanchor">[41]</a> Nevertheless the variety continues to be reproduced from
+seed, because, in addition to the double and infertile flowers, the
+seeds always produce a certain number of single, fertile blossoms, and
+these are used to reproduce the double variety. These single and
+fertile plants correspond "to the males and females of an ant-colony,
+the infertile plants, which are regularly produced in large numbers,
+to the neuter workers of the colony."</p>
+
+<p>This illustration is entirely apt, the only difference between <span class='pagenum'><a name="Page_46" id="Page_46">[46]</a></span>the
+two cases consisting in the fact that the variation in the flower is
+not a useful, but a disadvantageous one, which can only be preserved
+by artificial selection on the part of the gardener, while the
+transformations that have taken place parallel with the sterility of
+the ants are useful, since they procure for the colony an advantage in
+the struggle for existence, and they are therefore preserved by
+natural selection. Even the sterility itself in this case is not
+disadvantageous, since the fertility of the true females has at the
+same time considerably increased. We may therefore regard the sterile
+forms of ants, which have gradually been adapted in several directions
+to varying functions, <i>as a certain proof</i> that selection really takes
+place in the germ-cells of the fathers and mothers of the workers, and
+that <i>special complexes of primordia</i> (<i>ids</i>) are present in the
+workers and in the males and females, and these complexes contain the
+primordia of the individual parts (<i>determinants</i>). But since all
+living entities vary, the determinants must also vary, now in a
+favourable, now in an unfavourable direction. If a female produces
+eggs, which contain favourably varying determinants in the worker-ids,
+then these eggs will give rise to workers modified in the favourable
+direction, and if this happens with many females, the colony concerned
+will contain a better kind of worker than other colonies.</p>
+
+<p>I digress here in order to give an account of the intimate processes,
+which, according to my view, take place within the germ-plasm, and
+which I have called "<i>germinal selection</i>." These processes are of
+importance since they form the roots of variation, which in its turn
+is the root of natural selection. I cannot here do more than give a
+brief outline of the theory in order to show how the Darwin-Wallace
+theory of selection has gained support from it.</p>
+
+<p>With others, I regard the minimal amount of substance which is
+contained within the nucleus of the germ-cells, in the form of rods,
+bands, or granules, as the <i>germ-substance</i> or <i>germ-plasm</i>, and I
+call the individual granules <i>ids</i>. There is always a multiplicity of
+such ids present in the nucleus<span class='pagenum'><a name="Page_47" id="Page_47">[47]</a></span>, either occurring individually, or
+united in the form of rods or bands (chromosomes). Each id contains
+the primary constituents of a <i>whole</i> individual, so that several ids
+are concerned in the development of a new individual.</p>
+
+<p>In every being of complex structure thousands of primary constituents
+must go to make up a single id; these I call <i>determinants</i>, and I
+mean by this name very small individual particles, far below the
+limits of microscopic visibility, vital units which feed, grow, and
+multiply by division. These determinants control the parts of the
+developing embryo,&mdash;in what manner need not here concern us. The
+determinants differ among themselves, those of a muscle are
+differently constituted from those of a nerve-cell or a glandular
+cell, etc., and every determinant is in its turn made up of minute
+vital units, which I call <i>biophores</i>, or the bearers of life.
+According to my view, these determinants not only assimilate, like
+every other living unit, but they <i>vary</i> in the course of their
+growth, as every living unit does; they may vary qualitatively if the
+elements of which they are composed vary, they may grow and divide
+more or less rapidly, and their variations give rise to
+<i>corresponding</i> variations of the organ, cell, or cell-group which
+they determine. That they are undergoing ceaseless fluctuations in
+regard to size and quality seems to me the inevitable consequence of
+their unequal nutrition; for although the germ-cell as a whole usually
+receives sufficient nutriment, minute fluctuations in the amount
+carried to different parts within the germ-plasm cannot fail to occur.</p>
+
+<p>Now, if a determinant, for instance of a sensory cell, receives for a
+considerable time more abundant nutriment than before, it will grow
+more rapidly&mdash;become bigger, and divide more quickly, and, later, when
+the id concerned develops into an embryo, this sensory cell will
+become stronger than in the parents, possibly even twice as strong.
+This is an instance of a <i>hereditary individual variation</i>, arising
+from the germ.</p>
+
+<p>The nutritive stream which, according to our hypothesis,<span class='pagenum'><a name="Page_48" id="Page_48">[48]</a></span> favours the
+determinant <i>N</i> by chance, that is, for reasons unknown to us, may
+remain strong for a considerable time, or may decrease again; but even
+in the latter case it is conceivable that the ascending movement of
+the determinant may continue, because the strengthened determinant now
+<i>actively</i> nourishes itself more abundantly,&mdash;that is to say, it
+attracts the nutriment to itself, and to a certain extent withdraws it
+from its fellow-determinants. In this way, it may&mdash;as it seems to
+me&mdash;get into <i>permanent upward movement, and attain a degree of
+strength from which there is no falling back</i>. Then positive or
+negative selection sets in, favouring the variations which are
+advantageous, setting aside those which are disadvantageous.</p>
+
+<p>In a similar manner a <i>downward</i> variation of the determinants may
+take place, if its progress be started by a diminished flow of
+nutriment. The determinants which are weakened by this diminished flow
+will have less affinity for attracting nutriment because of their
+diminished strength, and they will assimilate more feebly and grow
+more slowly, unless chance streams of nutriment help them to recover
+themselves. But, as will presently be shown, a change of direction
+cannot take place at <i>every</i> stage of the degenerative process. If a
+certain critical stage of downward progress be passed, even favourable
+conditions of food-supply will no longer suffice permanently to change
+the direction of the variation. Only two cases are conceivable; if the
+determinant corresponds to a <i>useful</i> organ, only its removal can
+bring back the germ-plasm to its former level; therefore personal
+selection removes the id in question, with its determinants, from the
+germ-plasm, by causing the elimination of the individual in the
+struggle for existence. But there is another conceivable case; the
+determinants concerned may be those of an organ which has become
+<i>useless</i>, and they will then continue unobstructed, but with
+exceeding slowness, along the downward path, until the organ becomes
+vestigial, and finally disappears altogether.</p>
+
+<p>The fluctuations of the determinants hither and thither<span class='pagenum'><a name="Page_49" id="Page_49">[49]</a></span> may thus be
+transformed into a lasting ascending or descending movement; and <i>this
+is the crucial point of these germinal processes</i>.</p>
+
+<p>This is not a fantastic assumption; we can read it in the fact of the
+degeneration of disused parts. <i>Useless organs are the only ones which
+are not helped to ascend again by personal selection, and therefore in
+their case alone can we form any idea of how the primary constituents
+behave, when they are subject solely to intra-germinal forces</i>.</p>
+
+<p>The whole determinant system of an id, as I conceive it, is in a state
+of continual fluctuation upwards and downwards. In most cases the
+fluctuations will counteract one another, because the passive streams
+of nutriment soon change, but in many cases the limit from which a
+return is possible will be passed, and then the determinants concerned
+will continue to vary in the same direction, till they attain positive
+or negative selection-value. At this stage personal selection
+intervenes and sets aside the variation if it is disadvantageous, or
+favours&mdash;that is to say, preserves&mdash;it if it is advantageous. Only
+<i>the determinant of a useless organ is uninfluenced by personal
+selection</i>, and, as experience shows, it sinks downwards; that is, the
+organ that corresponds to it degenerates very slowly but
+uninterruptedly till, after what must obviously be an immense stretch
+of time, it disappears from the germ-plasm altogether.</p>
+
+<p>Thus we find in the fact of the degeneration of disused parts the
+proof that not all the fluctuations of a determinant return to
+equilibrium again, but that, when the movement has attained to a
+certain strength, it continues <i>in the same direction</i>. We have entire
+certainty in regard to this as far as the downward progress is
+concerned, and we must assume it also in regard to ascending
+variations, as the phenomena of artificial selection certainly justify
+us in doing. If the Japanese breeders were able to lengthen the
+tail-feathers of the cock to six feet, it can only have been because
+the determinants of the tail-feathers in the germ-plasm had already
+struck out a path of ascending variation, and this movement<span class='pagenum'><a name="Page_50" id="Page_50">[50]</a></span> was taken
+advantage of by the breeder, who continually selected for reproduction
+the individuals in which the ascending variation was most marked. For
+all breeding depends upon the unconscious selection of germinal
+variations.</p>
+
+<p>Of course these germinal processes cannot be proved mathematically,
+since we cannot actually see the play of forces of the passive
+fluctuations and their causes. We cannot say how great these
+fluctuations are, and how quickly or slowly, how regularly or
+irregularly they change. Nor do we know how far a determinant must be
+strengthened by the passive flow of the nutritive stream if it is to
+be beyond the danger of unfavourable variations, or how far it must be
+weakened passively before it loses the power of recovering itself by
+its own strength. It is no more possible to bring forward actual
+proofs in this case than it was in regard to the selection-value of
+the initial stages of an adaptation. But if we consider that all
+heritable variations must have their roots in the germ-plasm, and
+further, that when personal selection does not intervene, that is to
+say, in the case of parts which have become useless, a degeneration of
+the part, and therefore also of its determinant must inevitably take
+place; then we must conclude that processes such as I have assumed are
+running their course within the germ-plasm, and we can do this with as
+much certainty as we were able to infer, from the phenomena of
+adaptation, the selection-value of their initial stages. The fact of
+the degeneration of disused parts seems to me to afford irrefutable
+proof that the fluctuations within the germ-plasm <i>are the real root
+of all hereditary variation</i>, and the preliminary condition for the
+occurrence of the Darwin-Wallace factor of selection. Germinal
+selection supplies the stones out of which personal selection builds
+her temples and palaces: <i>adaptations</i>. The importance for the theory
+of the process of degeneration of disused parts cannot be
+over-estimated, especially when it occurs in sterile animal forms,
+where we are free from the doubt as to the alleged <i>Lamarckian factor</i>
+which is apt to confuse our ideas in regard to other cases.<span class='pagenum'><a name="Page_51" id="Page_51">[51]</a></span></p>
+
+<p>If we regard the variation of the many determinants concerned in the
+transformation of the female into the sterile worker as having come
+about through the gradual transformation of the ids into worker-ids,
+we shall see that the germ-plasm of the sexual ants must contain three
+kinds of ids, male, female, and worker ids, or if the workers have
+diverged into soldiers and nest-builders, then four kinds. We
+understand that the worker-ids arose because their determinants struck
+out a useful path of variation, whether upward or downward, and that
+they continued in this path until the highest attainable degree of
+utility of the parts determined was reached. But in addition to the
+organs of positive or negative selection-value, there were some which
+were indifferent as far as the success and especially the functional
+capacity of the workers was concerned: wings, ovarian tubes,
+<i>receptaculum seminis</i>, a number of the facets of the eye, perhaps
+even the whole eye. As to the ovarian tubes it is is possible that
+their degeneration was an advantage for the workers, in saving energy,
+and if so selection would favour the degeneration; but how could the
+presence of eyes diminish the usefulness of the workers to the colony?
+or the minute <i>receptaculum seminis</i>, or even the wings? These parts
+have therefore degenerated <i>because they were of no further value to
+the insect</i>. But if selection did not influence the setting aside of
+these parts because they were neither of advantage nor of disadvantage
+to the species, then the Darwinian factor of selection is here
+confronted with a puzzle which it cannot solve alone, but which at
+once becomes clear when germinal selection is added. For the
+determinants of organs that have no further value for the organism,
+must, as we have already explained, embark on a gradual course of
+retrograde development.</p>
+
+<p>In ants the degeneration has gone so far that there are no
+wing-rudiments present in <i>any</i> species, as is the case with so many
+butterflies, flies, and locusts, but in the larvae the imaginable
+discs of the wings are still laid down. With regard to the ovaries,
+degeneration has reached different levels<span class='pagenum'><a name="Page_52" id="Page_52">[52]</a></span> in different species of
+ants, as has been shown by the researches of my former pupil,
+Elizabeth Bickford. In many species there are twelve ovarian tubes,
+and they decrease from that number to one; indeed, in one species no
+ovarian tube at all is present. So much at least is certain from what
+has been said, that in this case <i>everything</i> depends on the
+fluctuations of the elements of the germ-plasm. Germinal selection,
+here as elsewhere, presents the variations of the determinants, and
+personal selection favours or rejects these, or,&mdash;if it be a question
+of organs which have become useless,&mdash;it does not come into play at
+all, and allows the descending variation free course.</p>
+
+<p>It is obvious that even the problem of <i>coadaptation in sterile
+animals</i> can thus be satisfactorily explained. If the determinants are
+oscillating upwards and downwards in continual fluctuation, and
+varying more pronouncedly now in one direction now in the other,
+useful variations of every determinant will continually present
+themselves anew, and may, in the course of generations, be combined
+with one another in various ways. But there is one character of the
+determinants that greatly facilitates this complex process of
+selection, that, after a certain limit has been reached, they go on
+varying in the same direction. From this it follows that development
+along a path once struck out may proceed without the continual
+intervention of personal selection. This factor only operates, so to
+speak, at the beginning, when it selects the determinants which are
+varying in the right direction, and again at the end, when it is
+necessary to put a check upon further variation. In addition to this,
+enormously long periods have been available for all these adaptations,
+as the very gradual transition stages between females and workers in
+many species plainly show, and thus this process of transformation
+loses the marvellous and mysterious character that seemed at the first
+glance to invest it, and takes rank, without any straining, among the
+other processes of selection. It seems to me that, from the facts that
+sterile animal forms can adapt themselves to new vital<span class='pagenum'><a name="Page_53" id="Page_53">[53]</a></span> functions,
+their superfluous parts degenerate, and the parts more used adapt
+themselves in an ascending direction, those less used in a descending
+direction, we must draw the conclusion that harmonious adaptation here
+comes about <i>without the co&ouml;peration of the Lamarckian principle</i>.
+This conclusion once established, however, we have no reason to refer
+the thousands of cases of harmonious adaptation, which occur in
+exactly the same way among other animals or plants, to a principle,
+the <i>active intervention of which in the transformation of species is
+nowhere proved. We do not require it to explain the facts, and
+therefore we must not assume it.</i></p>
+
+<p>The fact of coadaptation, which was supposed to furnish the strongest
+argument against the principle of selection, in reality yields the
+clearest evidence in favour of it. We <i>must</i> assume it, <i>because no
+other possibility of explanation is open to us, and because these
+adaptations actually exist, that is to say, have really taken place</i>.
+With this conviction I attempted, as far back as 1894, when the idea
+of germinal selection had not yet occurred to me, to make "harmonious
+adaptation" (coadaptation) more easily intelligible in some way or
+other, and so I was led to the idea, which was subsequently expounded
+in detail by Baldwin, and Lloyd Morgan, and also by Osborn, and Gulick
+as <i>Organic Selection</i>. It seemed to me that it was not necessary that
+all the germinal variations required for secondary variations should
+have occurred <i>simultaneously</i>, since, for instance, in the case of
+the stag, the bones, muscles, sinews, and nerves would be incited by
+the increasing heaviness of the antlers to greater activity in <i>the
+individual life</i>, and so would be strengthened. The antlers can only
+have increased in size by very slow degrees, so that the muscles and
+bones may have been able to keep pace with their growth in the
+individual life, until the requisite germinal variations presented
+themselves. In this way a disharmony between the increasing weight of
+the antlers and the parts which support and move them would be
+avoided, since time would be given for the appropriate germinal
+variations to occur, and<span class='pagenum'><a name="Page_54" id="Page_54">[54]</a></span> so to set agoing the <i>hereditary</i> variation
+of the muscles, sinews and bones.<a name="FNanchor_42_42" id="FNanchor_42_42"></a><a href="#Footnote_42_42" class="fnanchor">[42]</a></p>
+
+<p>I still regard this idea as correct, but I attribute less importance
+to "organic selection" than I did at that time, in so far that I do
+not believe that it <i>alone</i> could effect complex harmonious
+adaptations. Germinal selection now seems to me to play the chief part
+in bringing about such adaptations. Something the same is true of the
+principle I have called <i>Panmixia</i>. As I became more and more
+convinced, in the course of years, that the <i>Lamarckian principle</i>
+ought not to be called in to explain the dwindling of disused parts, I
+believed that this process might be simply explained as due to the
+cessation of the conservative effect of natural selection. I said to
+myself that, from the moment in which a part ceases to be of use,
+natural selection withdraws its hand from it, and then it must
+inevitably fall from the height of its adaptiveness, because inferior
+variants would have as good a chance of persisting as better ones,
+since all grades of fitness of the part in question would be mingled
+with one another indiscriminately. This is undoubtedly true, as
+Romanes pointed out ten years before I did, and this mingling of the
+bad with the good probably does bring about a deterioration of the
+part concerned. But it cannot account for the steady diminution, which
+always occurs when a part is in process of becoming rudimentary, and
+which goes on until it ultimately disappears altogether. The process
+of dwindling cannot therefore be explained as due to panmixia alone:
+we can only find a sufficient explanation in germinal selection.</p>
+
+
+<p>IV. <span class="smcap">Derivatives of the Theory of Selection</span></p>
+
+<p>The impetus in all directions given by Darwin through his theory of
+selection has been an immeasurable one, and its influence is still
+felt. It falls within the province of the <span class='pagenum'><a name="Page_55" id="Page_55">[55]</a></span>historian of science to
+enumerate all the ideas which, in the last quarter of the nineteenth
+century, grew out of Darwin's theories, in the endeavour to penetrate
+more deeply into the problem of the evolution of the organic world.
+Within the narrow limits to which this paper is restricted, I cannot
+attempt to discuss any of these.</p>
+
+
+<p>V. <span class="smcap">Arguments for the Reality of the Processes of Selection</span></p>
+
+
+<p>(<i>a</i>) <i>Sexual Selection</i></p>
+
+<p>Sexual selection goes hand in hand with natural selection. From the
+very first I have regarded sexual selection as affording an extremely
+important and interesting corroboration of natural selection, but,
+singularly enough, it is precisely against this theory that an adverse
+judgment has been pronounced in so many quarters, and it is only quite
+recently, and probably in proportion as the wealth of facts in proof
+of it penetrates into a wider circle, that we seem to be approaching a
+more general recognition of this side of the problem of adaptations.
+Thus Darwin's words in his preface to the second edition (1874) of his
+book, <i>The Descent of Man and Sexual Selection</i>, are being justified:
+"My conviction as to the operation of natural selection remains
+unshaken," and further, "If naturalists were to become more familiar
+with the idea of sexual selection, it would, I think, be accepted to a
+much greater extent, and already it is fully and favourably accepted
+by many competent judges." Darwin was able to speak thus because he
+was already acquainted with an immense mass of facts, which, taken
+together, yield overwhelming evidence of the validity of the principle
+of sexual selection.</p>
+
+<p><i>Natural selection</i> chooses out for reproduction the individuals that
+are best equipped for the struggle for existence, and it does so at
+every stage of development; it thus improves the species in all its
+stages and forms. <i>Sexual selec<span class='pagenum'><a name="Page_56" id="Page_56">[56]</a></span>tion</i> operates only on individuals
+that are already capable of reproduction, and does so only in relation
+to the attainment of reproduction. It arises from the rivalry of one
+sex, usually the male, for the possession of the other, usually the
+female. Its influence can therefore only <i>directly</i> affect one sex, in
+that it equips it better for attaining possession of the other. But
+the effect may extend indirectly to the female sex, and thus the whole
+species may be modified, without, however, becoming any more capable
+of resistance in the struggle for existence, for sexual selection only
+gives rise to adaptations which are likely to give their possessor the
+victory over rivals in the struggle for possession of the female, and
+which are therefore peculiar to the wooing sex: the manifold
+"secondary sexual characters." The diversity of these characters is so
+great that I cannot here attempt to give anything approaching a
+complete treatment of them, but I should like to give a sufficient
+number of examples to make the principle itself, in its various modes
+of expression, quite clear.</p>
+
+<p>One of the chief preliminary postulates of sexual selection is the
+unequal number of individuals in the two sexes, for if every male
+immediately finds his mate there can be no competition for the
+possession of the female. Darwin has shown that, for the most part,
+the inequality between the sexes is due simply to the fact that there
+are more males than females, and therefore the males must take some
+pains to secure a mate. But the inequality does not always depend on
+the numerical preponderance of the males, it is often due to polygamy;
+for, if one male claims several females, the number of females in
+proportion to the rest of the males will be reduced. Since it is
+almost always the males that are the wooers, we must expect to find
+the occurrence of secondary sexual characters chiefly among them, and
+to find it especially frequent in polygamous species. And this is
+actually the case.</p>
+
+<p>If we were to try to guess&mdash;without knowing the facts&mdash;what means the
+male animals make use of to overcome their<span class='pagenum'><a name="Page_57" id="Page_57">[57]</a></span> rivals in the struggle for
+the possession of the female, we might name many kinds of means, but
+it would be difficult to suggest any which is not actually employed in
+some animal group of other. I begin with the mere difference in
+strength, through which the male of many animals is so sharply
+distinguished from the female, as, for instance, the lion, walrus,
+"sea-elephant," and others. Among these the males fight violently for
+the possession of the female, who falls to the victor in the combat.
+In this simple case no one can doubt the operation of selection, and
+there is just as little room for doubt as to the selection-value of
+the initial stages of the variation. Differences in bodily strength
+are apparent even among human beings, although in their case the
+struggle for the possession of the female is no longer decided by
+bodily strength alone.</p>
+
+<p>Combats between male animals are often violent and obstinate, and the
+employment of the natural weapons of the species in this way has led
+to perfecting of these, e.g. the tusks of the boar, the antlers of the
+stag, and the enormous, antler-like jaws of the stag-beetle. Here
+again it is impossible to doubt that variations in these organs
+presented themselves, and that these were considerable enough to be
+decisive in combat, and so to lead to the improvement of the weapon.</p>
+
+<p>Among many animals, however, the females at first withdraw from the
+males; they are coy, and have to be sought out, and sometimes held by
+force. This tracking and grasping of the females by the males has
+given rise to many different characters in the latter, as, for
+instance, the larger eyes of the male bee, and especially of the males
+of the Ephemerids (May-flies), some species of which show, in addition
+to the usual compound eyes, large, so-called turban-eyes, so that the
+whole head is covered with seeing surfaces. In these species the
+females are very greatly in the minority (1-100), and it is easy to
+understand that a keen competition for them must take place, and that,
+when the insects of both sexes are floating freely in the air, an
+unusually wide<span class='pagenum'><a name="Page_58" id="Page_58">[58]</a></span> range of vision will carry with it a decided
+advantage. Here again the actual adaptations are in accordance with
+the preliminary postulates of the theory. We do not know the stages
+through which the eye has passed to its present perfected state, but,
+since the number of simple eyes (facets) has become very much greater
+in the male than in the female, we may assume that their increase is
+due to a gradual duplication of the determinants of the ommatidium in
+the germ-plasm, as I have already indicated in regard to sense-organs
+in general. In this case, again, the selection-value of the initial
+stages hardly admits of doubt; better vision <i>directly</i> secures
+reproduction.</p>
+
+<p>In many cases <i>the organ of smell</i> shows a similar improvement. Many
+lower Crustaceans (Daphnidae) have better developed organs of smell in
+the male sex. The difference is often slight and amounts only to one
+or two olfactory filaments, but certain species show a difference of
+nearly a hundred of these filaments (Leptodora). The same thing occurs
+among insects.</p>
+
+<p>We must briefly consider the clasping or grasping organs which have
+developed in the males among many lower Crustaceans, but here natural
+selection plays its part along with sexual selection, for the union of
+the sexes is an indispensable condition for the maintenance of the
+species, and as Darwin himself pointed out, in many cases the two
+forms of selection merge into each other. This fact has always seemed
+to me to be a proof of natural selection, for, in regard to sexual
+selection, it is quite obvious that the victory of the best-equipped
+could have brought about the improvement only of the organs concerned,
+the factors in the struggle, such as the eye and the olfactory organ.</p>
+
+<p>We come now to the <i>excitants</i>; that is, to the group of sexual
+characters whose origin through processes of selection has been most
+frequently called in question. We may cite the <i>love-calls</i> produced
+by many male insects, such as crickets and cicadas. These could only
+have arisen in animal groups in which the female did not rapidly flee
+from the<span class='pagenum'><a name="Page_59" id="Page_59">[59]</a></span> male, but was inclined to accept his wooing from the first.
+Thus, notes like the chirping of the male cricket serve to entice the
+females. At first they were merely the signal which showed the
+presence of a male in the neighbourhood, and the female was gradually
+enticed nearer and nearer by the continued chirping. The male that
+could make himself heard to the greatest distance would obtain the
+largest following, and would transmit the beginnings, and, later, the
+improvement of his voice to the greatest number of descendants. But
+sexual excitement in the female became associated with the hearing of
+the love-call, and then the sound-producing organ of the male began to
+improve, until it attained to the emission of the long-drawn-out soft
+notes of the mole-cricket or the maenad-like cry of the cicadas. I
+cannot here follow the process of development in detail, but will call
+attention to the fact that the original purpose of the voice, the
+announcing of the male's presence, became subsidiary, and the exciting
+of the female became the chief goal to be aimed at. The loudest
+singers awakened the strongest excitement, and the improvement
+resulted as a matter of course. I conceive of the origin of bird-song
+in a somewhat similar manner, first as a means of enticing, then of
+exciting the female.</p>
+
+<p>One more kind of secondary sexual character must here be mentioned:
+the odour which emanates from so many animals at the breeding season.
+It is possible that this odour also served at first merely to give
+notice of the presence of individuals of the other sex, but it soon
+became an excitant, and as the individuals which caused the greatest
+degree of excitement were preferred, it reached as high a pitch of
+perfection as was possible to it. I shall confine myself here to the
+comparatively recently discovered fragrance of butterflies. Since
+Fritz M&uuml;ller found out that certain Brazilian butterflies gave off
+fragrance "like a flower," we have become acquainted with many such
+cases, and we now know that in all lands, not only many diurnal
+Lepidoptera but nocturnal ones also give off a delicate odour<span class='pagenum'><a name="Page_60" id="Page_60">[60]</a></span>, which
+is agreeable even to man. The ethereal oil to which this fragrance is
+due is secreted by the skin-cells, usually of the wing, as I showed
+soon after the discovery of the <i>scent-scales</i>. This is the case in
+the males; the females have no <i>special</i> scent-scales recognisable as
+such by their form, but they must, nevertheless, give off an extremely
+delicate fragrance, although our imperfect organ of smell cannot
+perceive it, for the males become aware of the presence of a female,
+even at night, from a long distance off, and gather round her. We may
+therefore conclude, that both sexes have long given forth a very
+delicate perfume, which announced their presence to others of the same
+species, and that in many species (<i>not in all</i>) these small
+beginnings become, in the males, particularly strong scent-scales of
+characteristic form (lute, brush, or lyre-shaped). At first these
+scales were scattered over the surface of the wing, but gradually they
+concentrated themselves, and formed broad, velvety bands, or strong,
+prominent brushes, and they attained their highest pitch of evolution
+when they became enclosed within pits or folds of the skin, which
+could be opened to let the delicious fragrance stream forth suddenly
+towards the female. Thus in this case also we see that characters, the
+original use of which was to bring the sexes together, and so to
+maintain the species, have been evolved in the males into means for
+exciting the female. And we can hardly doubt, that the females are
+most readily enticed to yield to the butterfly that sends out the
+strongest fragrance,&mdash;that is to say, that excites them to the highest
+degree. It is a pity that our organs of smell are not fine enough to
+examine the fragrance of male Lepidoptera in general, and to compare
+it with other perfumes which attract these insects.<a name="FNanchor_43_43" id="FNanchor_43_43"></a><a href="#Footnote_43_43" class="fnanchor">[43]</a> As far as we
+can perceive them they resemble the fragrance of flowers, but there
+are Lepidoptera whose scent suggests musk. A smell of musk is also
+given off by several plants: it is a <span class='pagenum'><a name="Page_61" id="Page_61">[61]</a></span>sexual excitant in the
+musk-deer, the musk-sheep, and the crocodile.</p>
+
+<p>As far as we know, then, it is perfumes similar to those of flowers
+that the male Lepidoptera give off in order to entice their mates and
+this is a further indication that animals, like plants, can to a large
+extent meet the claims made upon them by life, and produce the
+adaptations which are most purposive,&mdash;a further proof, too, of my
+proposition that the useful variations, so to speak, are <i>always
+there</i>. The flowers developed the perfumes which entice their
+visitors, and the male Lepidoptera developed the perfumes which entice
+and excite their mates.</p>
+
+<p>There are many pretty little problems to be solved in this connection,
+for there are insects, such as some flies, that are attracted by
+smells which are unpleasant to us, like those from decaying flesh and
+carrion. But there are also certain flowers, some orchids for
+instance, which give forth no very agreeable odour, but one which is
+to us repulsive and disgusting; and we should therefore expect that
+the males of such insects would give off a smell unpleasant to us, but
+there is no case known to me in which this has been demonstrated.</p>
+
+<p>In cases such as we have discussed, it is obvious that there is no
+possible explanation except through selection. This brings us to the
+last kind of secondary sexual characters, and the one in regard to
+which doubt has been most frequently expressed,&mdash;decorative colours
+and decorative forms, the brilliant plumage of the male pheasant, the
+humming-birds, and the bird of Paradise, as well as the bright colours
+of many species of butterfly, from the beautiful blue of our little
+Lycaenidae to the magnificent azure of the large Morphinae of Brazil.
+In a great many cases, though not by any means in all, the male
+butterflies are "more beautiful" than the females, and in the Tropics
+in particular they shine and glow in the most superb colours. I really
+see no reason why we should doubt the power of sexual selection, and I
+myself stand wholly on Darwin's side. Even though we<span class='pagenum'><a name="Page_62" id="Page_62">[62]</a></span> certainly cannot
+assume that the females exercise a conscious choice of the
+"handsomest" mate, and deliberate like the judges in a court of
+justice over the perfections of their wooers, we have no reason to
+doubt that distinctive forms (decorative feathers), and colours have a
+particularly exciting effect upon the female, just as certain odours
+have among animals of so many different groups, including the
+butterflies. The doubts which existed for a considerable time, as a
+result of fallacious experiments, as to whether the colours of flowers
+really had any influence in attracting butterflies have now been set
+at rest through a series of more careful investigations; we now know
+that the colours of flowers are there on account of the butterflies,
+as Sprengel first showed, and that the blossoms of Phanerogams are
+selected in relation to them, as Darwin pointed out.</p>
+
+<p>Certainly it is not possible to bring forward any convincing proof of
+the origin of decorative colours through sexual selection, but there
+are many weighty arguments in favour of it, and these form a body of
+presumptive evidence so strong that it almost amounts to certainty.</p>
+
+<p>In the first place, there is the analogy with other secondary sexual
+characters. If the song of birds and the chirping of the cricket have
+been evolved through sexual selection, if the penetrating odours of
+male animals,&mdash;the crocodile, the musk-deer, the beaver, the
+carnivores, and, finally, the flower-like fragrances of the
+butterflies have been evolved to their present pitch in this way, why
+should decorative colours have arisen in some other way? Why should
+the eye be less sensitive to <i>specifically male</i> colours and other
+<i>visible</i> signs <i>enticing to the female</i>, than the olfactory sense to
+specifically male odours, or the sense of hearing to specifically male
+sounds? Moreover, the decorative feathers of birds are almost always
+spread out and displayed before the female during courtship. I have
+elsewhere<a name="FNanchor_44_44" id="FNanchor_44_44"></a><a href="#Footnote_44_44" class="fnanchor">[44]</a> pointed out that decorative colouring and
+sweet-scentedness may replace one another in Lepidoptera as well as in
+flowers, for <span class='pagenum'><a name="Page_63" id="Page_63">[63]</a></span>just as some modestly coloured flowers (mignonette and
+violet) have often a strong perfume, while strikingly coloured ones
+are sometimes quite devoid of fragrance, so we find that the most
+beautiful and gaily-coloured of our native Lepidoptera, the species of
+Vanessa, have no scent-scales, while these are often markedly
+developed in grey nocturnal Lepidoptera. Both attractions may,
+however, be combined in butterflies, just as in flowers. Of course, we
+cannot explain why both means of attraction should exist in one genus,
+and only one of them in another, since we do not know the minutest
+details of the conditions of life of the genera concerned. But from
+the sporadic distribution of scent-scales in Lepidoptera, and from
+their occurrence or absence in nearly related species, we may conclude
+that fragrance is a relatively <i>modern</i> acquirement, more recent than
+brilliant colouring.</p>
+
+<p>One thing in particular that stamps decorative colouring as a product
+of selection is <i>its gradual intensification</i> by the addition of new
+spots, which we can quite well observe, because in many cases the
+colours have been first acquired by the males, and later transmitted
+to the females by inheritance. The scent-scales are never thus
+transmitted, probably for the same reason that the decorative colours
+of many birds are often not transmitted to the females: because with
+these they would be exposed to too great elimination by enemies.
+Wallace was the first to point out that in species with concealed
+nests the beautiful feathers of the male occurred in the female also,
+as in the parrots, for instance, but this is not the case in species
+which brood on an exposed nest. In the parrots one can often observe
+that the general brilliant colouring of the male is found in the
+female, but that certain spots of colour are absent, and these have
+probably been acquired comparatively recently by the male and have not
+yet been transmitted to the female.</p>
+
+<p>Isolation of the group of individuals which is in process of varying
+is undoubtedly of great value in sexual selection, for even a solitary
+conspicuous variation will become dom<span class='pagenum'><a name="Page_64" id="Page_64">[64]</a></span>inant much sooner in a small
+isolated colony, than among a large number of members of a species.</p>
+
+<p>Any one who agrees with me in deriving variations from germinal
+selection will regard that process as an essential aid towards
+explaining the selection of distinctive courtship-characters, such as
+coloured spots, decorative feathers, horny outgrowths in birds and
+reptiles, combs, feather-tufts, and the like, since the beginnings of
+these would be presented with relative frequency in the struggle
+between the determinants within the germ-plasm. The process of
+transmission of decorative feathers to the female results, as Darwin
+pointed out and illustrated by interesting examples, in the
+<i>colour-transformation of a whole species</i>, and this process, as the
+phyletically older colouring of young birds shows, must, in the course
+of thousands of years, have repeated itself several times in a line of
+descent.</p>
+
+<p>If we survey the wealth of phenomena presented to us by secondary
+sexual characters, we can hardly fail to be convinced of the truth of
+the principle of sexual selection. And certainly no one who has
+accepted natural selection should reject sexual selection, for, not
+only do the two processes rest upon the same basis, but they merge
+into one another, so that it is often impossible to say how much of a
+particular character depends on one and how much on the other form of
+selection.</p>
+
+
+<p>(<i>b</i>) <i>Natural Selection</i></p>
+
+<p>An actual proof of the theory of sexual selection is out of the
+question, if only because we cannot tell when a variation attains to
+selection-value. It is certain that a delicate sense of smell is of
+value to the male moth in his search for the female, but whether the
+possession of one additional olfactory hair, or of ten, or of twenty
+additional hairs leads to the success of its possessor we are unable
+to tell. And we are groping even more in the dark when we discuss the
+excitement caused in the female by agreeable perfumes, or<span class='pagenum'><a name="Page_65" id="Page_65">[65]</a></span> by striking
+and beautiful colours. That these do make an impression is beyond
+doubt; but we can only assume that slight intensifications of them
+give any advantage, and we <i>must</i> assume this <i>since otherwise
+secondary sexual characters remain inexplicable</i>.</p>
+
+<p>The same thing is true in regard to natural selection. It is not
+possible to bring forward any actual proof of the selection-value of
+the initial stages, and the stages in the increase of variations, as
+has been already shown. But the selection-value of a finished
+adaptation can in many cases be statistically determined. Cesnola and
+Poulton have made valuable experiments in this direction. The former
+attached forty-five individuals of the green, and sixty-five of the
+brown variety of the praying mantis (<i>Mantis religiosa</i>), by a silk
+thread to plants, and watched them for seven days. The insects which
+were on a surface of a colour Similar to their own remained uneaten,
+while twenty-five green insects on brown parts of plants had all
+disappeared in eleven days.</p>
+
+<p>The experiments of Poulton and Sanders<a name="FNanchor_45_45" id="FNanchor_45_45"></a><a href="#Footnote_45_45" class="fnanchor">[45]</a> were made with 600 pupae of
+<i>Vanessa urticae</i>, the "tortoise-shell butterfly." The pupae were
+artificially attached to nettles, tree-trunks, fences, walls, and to
+the ground, some at Oxford, some at St. Helens in the Isle of Wight.
+In the course of a month 93% of the pupae at Oxford were killed,
+chiefly by small birds, while at St. Helens 68% perished. The
+experiments showed very clearly that the colour and character of the
+surface on which the pupa rests&mdash;and thus its own conspicuousness&mdash;are
+of the greatest importance. At Oxford only the four pupae which were
+fastened to nettles emerged; all the rest&mdash;on bark, stones and the
+like&mdash;perished. At St. Helens the elimination was as follows: on
+fences where the pupae were conspicuous, 92%; on bark, 66%; on walls,
+54%; and among nettles, 57%. These interesting experiments confirm our
+views as to <span class='pagenum'><a name="Page_66" id="Page_66">[66]</a></span>protective coloration, and show further, <i>that the ratio
+of elimination in the species is a very high one, and that therefore
+selection must be very keen</i>.</p>
+
+<p>We may say that the process of selection follows as a logical
+necessity from the fulfilment of the three preliminary postulates of
+the theory: variability, heredity, and the struggle for existence,
+with its enormous ratio of elimination in all species. To this we must
+add a fourth factor, the <i>intensification</i> of variations which Darwin
+established as a fact, and which we are now able to account for
+theoretically on the basis of germinal selection. It may be objected
+that there is considerable uncertainty about this <i>logical</i> proof,
+because of our inability to demonstrate the selection-value of the
+initial stages and the individual stages of increase. We have
+therefore to fall back on <i>presumptive evidence</i>. This is to be found
+in <i>the interpretative value of the theory</i>. Let us consider this
+point in greater detail.</p>
+
+<p>In the first place it is necessary to emphasize what is often
+overlooked, namely, that the theory not only explains the
+<i>transformations</i> of species, it also explains <i>their remaining the
+same</i>; in addition to the principle of varying, it contains within
+itself that of <i>persisting</i>. It is part of the essence of selection,
+that it not only causes a part to <i>vary</i> till it has reached its
+highest pitch of adaptation, but that it <i>maintains it at this pitch.
+This conserving influence of natural selection</i> is of great
+importance, and was early recognised by Darwin; it follows naturally
+from the principle of the survival of the fittest.</p>
+
+<p>We understand from this how it is that a species which has become
+fully adapted to certain conditions of life ceases to vary, but
+remains "constant," as long as the conditions of life <i>for</i> it remain
+unchanged, whether this be for thousands of years, or for whole
+geological epochs. But the most convincing proof of the power of the
+principle of selection lies in the innumerable multitude of phenomena
+which cannot be explained in any other way. To this category belong
+all structures which are only <i>passively</i> of advantage to the<span class='pagenum'><a name="Page_67" id="Page_67">[67]</a></span>
+organism, because none of these can have arisen by the alleged
+<i>Lamarckian principle</i>. These have been so often discussed that we
+need do no more than indicate them here. Until quite recently the
+sympathetic coloration of animals&mdash;for instance, the whiteness of
+Arctic animals&mdash;was referred, at least in part, to the <i>direct</i>
+influence of external factors, but the facts can best be explained by
+referring them to the processes of selection, for then it is
+unnecessary to make the gratuitous assumption that many species are
+sensitive to the stimulus of cold and that others are not. The great
+majority of Arctic land-animals, mammals and birds, are white, and
+this proves that they were all able to present the variation which was
+most useful for them. The sable is brown, but it lives in trees, where
+the brown colouring protects and conceals it more effectively. The
+musk-sheep (<i>Ovibos moschatus</i>) is also brown, and contrasts sharply
+with the ice and snow, but it is protected from beasts of prey by its
+gregarious habit, and therefore it is of advantage to be visible from
+as great a distance as possible. That so many species have been able
+to give rise to white varieties does not depend on a special
+sensitiveness of the skin to the influence of cold, but to the fact
+that Mammals and Birds have a general tendency to vary towards white.
+Even with us, many birds&mdash;starlings, blackbirds, swallows,
+etc.&mdash;occasionally produce white individuals, but the white variety
+does not persist, because it readily falls a victim to the carnivores.
+This is true of white fawns, foxes, deer, etc. The whiteness,
+therefore, arises from internal causes, and only persists when it is
+useful. A great many animals living in a <i>green environment</i> have
+become clothed in green, especially insects, caterpillars, and
+Mantidae, both persecuted and persecutors.</p>
+
+<p>That it is not the direct effect of the environment which calls forth
+the green colour is shown by the many kinds of caterpillar which rest
+on leaves and feed on them, but are nevertheless brown. These feed by
+night and betake themselves through the day to the trunk of the tree,
+and<span class='pagenum'><a name="Page_68" id="Page_68">[68]</a></span> hide in the furrows of the bark. We cannot, however, conclude
+from this that they were <i>unable</i> to vary towards green, for there are
+Arctic animals which are white only in winter and brown in summer
+(Alpine hare, and the ptarmigan of the Alps), and there are also green
+leaf-insects which remain green only while they are young and
+difficult to see on the leaf, but which become brown again in the last
+stage of larval life, when they have outgrown the leaf. They then
+conceal themselves by day, sometimes only among withered leaves on the
+ground, sometimes in the earth itself. It is interesting that in one
+genus, Chaerocampa, one species is brown in the last stage of larval
+life, another becomes brown earlier, and in many species the last
+stage is not wholly brown, a part remaining green. Whether this is a
+case of a double adaptation, or whether the green is being gradually
+crowded out by the brown, the fact remains that the same species, even
+the same individual, can exhibit both variations. The case is the same
+with many of the leaf-like Orthoptera, as, for instance, the praying
+mantis (<i>Mantis religiosa</i>) which we have already mentioned.</p>
+
+<p>But the best proofs are furnished by those of ten-cited cases in which
+the insect bears a deceptive resemblance to another object. We now
+know many such cases, such as the numerous imitations of green or
+withered leaves, which are brought about in the most diverse ways,
+sometimes by mere variations in the form of the insect and in its
+colour, sometimes by an elaborate marking, like that which occurs in
+the Indian leaf-butterflies, <i>Kallima inachis</i>. In the single
+butterfly-genus Anaea, in the woods of South America, there are about
+a hundred species which are all gaily coloured on the upper surface,
+and on the reverse side exhibit the most delicate imitation of the
+colouring and pattern of a leaf, generally without any indication of
+the leaf-ribs, but extremely deceptive nevertheless. Anyone who has
+seen only one such butterfly may doubt whether many of the
+insignificant details of the marking can really be of advantage to the
+insect. Such details are for instance the apparent holes and<span class='pagenum'><a name="Page_69" id="Page_69">[69]</a></span> splits
+in the apparently dry or half-rotten leaf, which are usually due to
+the fact that the scales are absent on a circular or oval patch so
+that the colourless wing-membrane lies bare, and one can look through
+the spot as through a window. Whether the bird which is seeking or
+pursuing the butterflies takes these holes for dewdrops, or for the
+work of a devouring insect, does not affect the question; the
+mirror-like spot undoubtedly increases the general deceptiveness, for
+the same thing occurs in many leaf-butterflies, though not in all, and
+in some cases it is replaced in quite a peculiar manner. In one
+species of Anaea (<i>A. divina</i>), the resting butterfly looks exactly
+like a leaf out of the outer edge of which a large semi-circular piece
+has been eaten, possibly by a caterpillar; but if we look more closely
+it is obvious that there is no part of the wing absent, and that the
+semi-circular piece is of a clear, pale yellow colour, while the rest
+of the wing is of a strongly contrasted dark brown.</p>
+
+<p>But the deceptive resemblance may be caused in quite a different
+manner. I have often speculated as to what advantage the brilliant
+white C could give to the otherwise dusky-coloured "Comma butterfly"
+(<i>Grapta C. album</i>). Poulton's recent observations<a name="FNanchor_46_46" id="FNanchor_46_46"></a><a href="#Footnote_46_46" class="fnanchor">[46]</a> have shown that
+this represents the imitation of a crack such as is often seen in dry
+leaves, and is very conspicuous because the light shines through it.</p>
+
+<p>The utility obviously lies in presenting to the bird the very familiar
+picture of a broken leaf with a clear shining slit, and we may
+conclude, from the imitation of such small details, that the birds are
+very sharp observers and that the smallest deviation from the usual
+arrests their attention and incites them to closer investigation. It
+is obvious that such detailed&mdash;we might almost say such
+subtle&mdash;deceptive resemblances could only have come about in the
+course of long ages through the acquirement from time to time of
+<span class='pagenum'><a name="Page_70" id="Page_70">[70]</a></span>something new which heightened the already existing resemblance.</p>
+
+<p>In face of facts like these there can be no question of chance and no
+one has succeeded so far in finding any other explanation to replace
+that by selection. For the rest, the apparent leaves are by no means
+perfect copies of a leaf; many of them only represent the torn or
+broken piece, or the half or two-thirds of a leaf, but then the leaves
+themselves frequently do not present themselves to the eye as a whole,
+but partially concealed among other leaves. Even those butterflies
+which, like the species of Kallima and Anaea, represent the whole of a
+leaf with stalk, ribs, apex, and the whole breadth, are not actual
+copies which would satisfy a botanist; there is often much wanting. In
+Kallima the lateral ribs of the leaf are never all included in the
+markings; there are only two or three on the left side and at more
+four or five on the right, and in many individuals these are rather
+obscure, while in others they are comparatively distinct. This
+furnishes us with fresh evidence in favour of their origin through
+processes of selection, for a botanically perfect picture could not
+arise in this way; there could only be a fixing of such details as
+heightened the deceptive resemblance.</p>
+
+<p>Our postulate of origin through selection also enables us to
+understand why the leaf-imitation is on the lower surface of the wing
+in the diurnal Lepidoptera, and on the upper surface in the nocturnal
+forms, corresponding to the attitude of the wings in the resting
+position of the two groups.</p>
+
+<p>The strongest of all proofs of the theory, however, is afforded by
+cases of true "mimicry," those adaptations discovered by Bates in
+1861, consisting in the imitation of one species by another, which
+becomes more and more like its model. The model is always a species
+that enjoys some special protection from enemies, whether because it
+is unpleasant to taste, or because it is in some way dangerous.</p>
+
+<p>It is chiefly among insects and especially among butterflies that we
+find the greatest number of such cases. Several<span class='pagenum'><a name="Page_71" id="Page_71">[71]</a></span> of these have been
+minutely studied and every detail has been investigated so that it is
+difficult to understand how there can still be disbelief in regard to
+them. If the many and exact observations which have been carefully
+collected and critically discussed for instance by Poulton<a name="FNanchor_47_47" id="FNanchor_47_47"></a><a href="#Footnote_47_47" class="fnanchor">[47]</a> were
+thoroughly studied the arguments which are still frequently urged
+against mimicry would be found untenable; we can hardly hope to find
+more convincing proof of the actuality of the processes of selection
+than these cases put into our hands. The preliminary postulates of the
+theory of mimicry have been disputed, for instance, that diurnal
+butterflies are persecuted and eaten by birds, but observations
+specially directed towards this point in India, Africa, America and
+Europe have placed it beyond all doubt. If it were necessary I could
+myself furnish an account of my own observations on this point.</p>
+
+<p>In the same way it has been established by experiment and observation
+in the field that in all the great regions of distribution there are
+butterflies which are rejected by birds and lizards, their chief
+enemies, on account of their unpleasant smell or taste. These
+butterflies are usually gaily and conspicuously coloured and thus&mdash;as
+Wallace first interpreted it&mdash;are furnished with an easily
+recognisable sign: a sign of unpalatableness or <i>warning colours</i>. If
+they were not thus recognisable easily and from a distance, they would
+frequently be pecked at by birds, and then rejected because of their
+unpleasant taste; but as it is, the insect-eaters recognise them at
+once as unpalatable booty and ignore them. Such <i>immune</i><a name="FNanchor_48_48" id="FNanchor_48_48"></a><a href="#Footnote_48_48" class="fnanchor">[48]</a> species,
+wherever they occur, are imitated by other palatable species, which
+thus acquire a certain degree of protection.</p>
+
+<p>It is true that this explanation of the bright, conspicuous <span class='pagenum'><a name="Page_72" id="Page_72">[72]</a></span>colours
+is only a hypothesis, but its foundations&mdash;unpalatableness, and the
+liability of other butterflies to be eaten,&mdash;are certain, and its
+consequences&mdash;the existence of mimetic palatable forms&mdash;conform it in
+the most convincing manner. Of the many cases now known I select one,
+which is especially remarkable, and which has been thoroughly
+investigated, <i>Papilla dardanus</i> (<i>merope</i>), a large, beautiful,
+diurnal butterfly which ranges from Abyssinia throughout the whole of
+Africa to the south coast of Cape Colony.</p>
+
+<p>The males of this form are everywhere <i>almost</i> the same in colour and
+in form of wings, save for a few variations in the sparse black
+markings on the pale yellow ground. But the females occur in several
+quite different forms and colourings, and one of these only, the
+Abyssinian form, is like the male, while the other three or four are
+<i>mimetic</i>, that is to say, they copy a butterfly of quite a different
+family the Danaids, which are among the <i>immune</i> forms. In each region
+the females have thus copied two or three different immune species.
+There is much that is interesting to be said in regard to these
+species, but it would be out of keeping with the general tenor of this
+paper to give details of this very complicated case of polymorphism in
+<i>P. Dardanus</i>. Anyone who is interested in the matter will find a full
+and exact statement of the case in as far as we know it, in Poulton's
+<i>Essays on Evolution</i> (pp. 373-375<a name="FNanchor_49_49" id="FNanchor_49_49"></a><a href="#Footnote_49_49" class="fnanchor">[49]</a>). I need only add that three
+different mimetic female forms have been reared from the eggs of a
+single female in South Africa. The resemblance of the forms to their
+immune models goes so far that even the details of the <i>local</i> forms
+of the models are copied by the mimetic species.</p>
+
+<p>It remains to be said that in Madagascar a butterfly, <span class='pagenum'><a name="Page_73" id="Page_73">[73]</a></span></p>
+
+<p><i>Papilio meriones</i>, occurs, of which both sexes are very similar in
+form and markings to the non-mimetic male of <i>P. dardanus</i>, so that it
+probably represents the ancestor of this latter species.</p>
+
+<p>In face of such facts as these every attempt at another explanation
+must fail. Similarly all the other details of the case fulfil the
+preliminary postulates of selection, and leave no room for any other
+interpretation. That the males do not take on the protective colouring
+is easily explained, because they are in general more numerous, and
+the females are more important for the preservation of the species,
+and must also live longer in order to deposit their eggs. We find the
+same state of things in many other species, and in one case (<i>Elymnias
+undularis</i>) in which the male is also mimetically coloured, it copies
+quite a differently coloured immune species from the model followed by
+the female. This is quite intelligible when we consider that if there
+were <i>too many</i> false immune types, the birds would soon discover that
+there were palatable individuals among those with unpalatable warning
+colours. Hence the imitation of different immune species by <i>Papilio
+dardanus</i>!</p>
+
+<p>I regret that lack of space prevents my bringing forward more examples
+of mimicry and discussing them fully. But from the case of <i>Papilio
+dardanus</i> alone there is much to be learnt which is of the highest
+importance for our understanding of transformations. It shows us
+chiefly what I once called, somewhat strongly perhaps, <i>the
+omnipotence of natural selection</i> in answer to an opponent who had
+spoken of its "inadequacy." We here see that one and the same species
+is capable of producing four or five different patterns of colouring
+and marking; thus the colouring and marking are not, as has often been
+supposed, a necessary outcome of the specific nature of the species,
+but a true adaptation, which cannot arise as a direct effect of
+climatic conditions, but solely through what I may call the sorting
+out of the variations produced by the species, according to their
+utility. That caterpillars may be either green or brown is already<span class='pagenum'><a name="Page_74" id="Page_74">[74]</a></span>
+something more than could have been expected according to the old
+conception of species, but that one and the same butterfly should be
+now pale yellow, with black; now red with black and pure white; now
+deep black with large, pure white spots; and again black with a large
+ocheous-yellow spot, and many small white and yellow spots; that in
+one sub-species it may be tailed like the ancestral form, and in
+another tailless like its Danaid model,&mdash;all this shows a far-reaching
+capacity for variation and adaptation that we could never have
+expected if we did not see the facts before us. How it is possible
+that the primary colour-variations should thus be intensified and
+combined remains a puzzle even now; we are reminded of the modern
+three-colour printing,&mdash;perhaps similar combinations of the primary
+colours take place in this case; in any case the direction of these
+primary variations is determined by the artist whom we know as natural
+selection, for there is no other conceivable way in which the model
+could affect the butterfly that is becoming more and more like it. The
+same climate surrounds all four forms of female; they are subject to
+the same conditions of nutrition. Moreover, <i>Papilio dardanus</i> is by
+no means the only species of butterfly which exhibits different kinds
+of colour-pattern on its wings. Many species of the Asiatic genus
+Elymnias have on the upper surface a very good imitation of an immune
+Euploeine (Danainae), often with a steel-blue ground-colour, while the
+under surface is well concealed when the butterfly is at rest,&mdash;thus
+there are two kinds of protective coloration each with a different
+meaning! The same thing may be observed in many non-mimetic
+butterflies, for instance in all our species of Vanessa, in which the
+under side shows a grey-brown or brownish-black protective coloration,
+but we do not yet know with certainty what may be the biological
+significance of the gaily coloured upper surface.</p>
+
+<p>In general it may be said that mimetic butterflies are comparatively
+rare species, but there are exceptions, for instance <i>Limenitis
+archippus</i> in North America, of which the<span class='pagenum'><a name="Page_75" id="Page_75">[75]</a></span> immune model (<i>Danaida
+plexippus</i>) also occurs in enormous numbers.</p>
+
+<p>In another mimicry-category the imitators are often more numerous than
+the models, namely in the case of the imitation of <i>dangerous insects</i>
+by harmless species. Bees and wasps are dreaded for their sting, and
+they are copied by harmless flies of the genera Eristalis and Syrphus,
+and these mimics often occur in swarms about flowering plants without
+damage to themselves or to their models; they are feared and are
+therefore left unmolested.</p>
+
+<p>In regard also to the <i>faithfulness of the copy</i> the facts are quite
+in harmony with the theory, according to which the resemblance must
+have arisen and increased <i>by degrees</i>. We can recognise this in many
+cases, for even now the mimetic species show very <i>varying degrees of
+resemblance</i> to their immune model. If we compare, for instance, the
+many different imitators of <i>Danaida chrysippus</i> we find that, with
+their brownish-yellow ground-colour, and the position and size, and
+more or less sharp limitation of their clear marginal spots, they have
+reached very different degrees of nearness to their model. Or compare
+the female of <i>Elymnias undularis</i> with its model <i>Danaida genutia</i>;
+there is a general resemblance, but the marking of the Danaida is very
+roughly imitated in Elymnias.</p>
+
+<p>Another fact that bears out the theory of mimicry is, that even when
+the resemblance in colour-pattern is very great, the <i>wing-venation</i>,
+which is so constant, and so important in determining the systematic
+position of butterflies, is never affected by the variation. The
+pursuers of the butterfly have no time to trouble about entomological
+intricacies.</p>
+
+<p>I must not pass over a discovery of Poulton's which is of great
+theoretical importance&mdash;that mimetic butterflies may reach the same
+effect by very different means.<a name="FNanchor_50_50" id="FNanchor_50_50"></a><a href="#Footnote_50_50" class="fnanchor">[50]</a> Thus the glass-like transparency
+of the wing of a certain Ithomiine (Methona) and its Pierine mimic
+(<i>Dismorphia orise</i>) <span class='pagenum'><a name="Page_76" id="Page_76">[76]</a></span>depends on a diminution in the size of the
+scales; in the Danaine genus Itune it is due to the fewness of the
+scales and in a third imitator, a moth (<i>Castnia linus var.
+heliconoides</i>) the glass-like appearance of the wing is due neither to
+diminution nor to absence of scales, but to their absolute
+colourlessness and transparency, and to the fact that they stand
+upright. In another moth mimic (Anthomyza) the arrangement of the
+transparent scales is normal. Thus it is not some unknown external
+influence that has brought about the transparency of the wing in these
+five forms, as has sometimes been supposed. Nor is it a hypothetical
+<i>internal</i> evolutionary tendency, for all three vary in a different
+manner. The cause of this agreement can only lie in selection, which
+preserves and intensifies in each species the favourable variations
+that present themselves. The great faithfulness of the copy is
+astonishing in these cases, for it is not <i>the whole</i> wing which is
+transparent; certain markings are black in colour, and these contrast
+sharply with the glass-like ground. It is obvious that the pursuers of
+these butterflies must be very sharp-sighted, for otherwise the
+agreement between the species could never have been pushed so far. The
+less the enemies see and observe, the more defective must the
+imitation be, and if they had been blind, no visible resemblance
+between the species which required protection could ever have arisen.</p>
+
+<p>A seemingly irreconcilable contradiction to the mimicry theory is
+presented in the following cases, which were known to Bates, who,
+however, never succeeded in bringing them into line with the principle
+of mimicry.</p>
+
+<p>In South America there are, as we have already said, many mimics of
+the immune Ithomiinae (or as Bates called them Heliconidae). Among
+these there occur not merely species which are edible, and thus
+require the protection of a disguise, but others which are rejected on
+account of their unpalatableness. How could the Ithomiine dress have
+developed in their case, and of what use is it, since the species
+would in any case be immune? In Eastern Brazil, for in<span class='pagenum'><a name="Page_77" id="Page_77">[77]</a></span>stance, there
+are four butterflies, which bear a most confusing resemblance to one
+another in colour, marking, and form of wing, and all four are
+unpalatable to birds. They belong to four different genera and three
+sub-families, and we have to inquire: Whence came this resemblance and
+what end does it serve? For a long time no satisfactory answer could
+be found, but Fritz M&uuml;ller,<a name="FNanchor_51_51" id="FNanchor_51_51"></a><a href="#Footnote_51_51" class="fnanchor">[51]</a> seventeen years after Bates, offered a
+solution to the riddle, when he pointed out that young birds could not
+have an instinctive knowledge of the unpalatableness of the
+Ithomiines, but must learn by experience which species were edible and
+which inedible. Thus each young bird must have tasted at least one
+individual of each inedible species and discovered its unpalatability,
+before it learnt to avoid, and thus to spare the species. But if the
+four species resemble each other very closely the bird will regard
+them all as of the same kind, and avoid them all. Thus there developed
+a process of selection which resulted in the survival of the
+Ithomiine-like individuals, and in so great an increase of resemblance
+between the four species, that they are difficult to distinguish one
+from another even in a collection. The advantage for the four species,
+living side by side as they do e.g. in Bahia, lies in the fact that
+only one individual from the <i>mimicry-ring</i> ("inedible association")
+need be tasted by a young bird, instead of at least four individuals,
+as would otherwise be the case. As the number of young birds is great,
+this makes a considerable difference in the ratio of elimination. The
+four Brazilian species are <i>Lycorea halia</i> (Danainae), <i>Heliconius
+narcaea</i> (<i>eucrate</i>) (Heliconinae), <i>Melinaea ethra</i>, and <i>Mechanitis
+lysimnia</i> (Ithomiinae).</p>
+
+<p>These interesting mimicry-rings (trusts), which have much significance
+for the theory, have been the subject of numerous and careful
+investigations, and at least their essential features are now fully
+established. M&uuml;ller took for granted, without making any
+investigations, that young birds only <span class='pagenum'><a name="Page_78" id="Page_78">[78]</a></span>learn by experience to
+distinguish between different kinds of victims. But Lloyd Morgan's<a name="FNanchor_52_52" id="FNanchor_52_52"></a><a href="#Footnote_52_52" class="fnanchor">[52]</a>
+experiments with young birds proved that this is really the case, and
+at the same time furnished an additional argument against the
+<i>Lamarckian principle</i>.</p>
+
+<p>In addition to the mimicry-rings first observed in South America,
+others have been described from Tropical India by Moore, and by
+Poulton and Dixey from Africa, and we may expect to learn many more
+interesting facts in this connection. Here again the preliminary
+postulates of the theory are satisfied. And how much more that would
+lead to the same conclusion might be added!</p>
+
+<p>As in the case of mimicry many species have come to resemble one
+another through processes of selection, so we know whole classes of
+phenomena in which plants and animals have become adapted to one
+another, and have thus been modified to a considerable degree. I refer
+particularly to the relation between flowers and insects. Darwin has
+shown that the originally inconspicuous blossoms of the phanerogams
+were transformed into flowers through the visits of insects, and that,
+conversely, several large orders of insects have been gradually
+modified by their association with flowers, especially as regards the
+parts of their body actively concerned. Bees and butterflies in
+particular have become what they are through their relation to
+flowers. In this case again all that is apparently contradictory to
+the theory can, on closer investigation, be beautifully interpreted in
+corroboration of it. Selection can give rise only to what is of use to
+the organism actually concerned, never to what is of use to some other
+organism, and we must therefore expect to find that in flowers only
+characters of use to <i>themselves</i> have arisen, never characters which
+are of use to insects only, and conversely that in the insects
+characters useful to them and not merely to the plants would have
+originated. For a long time it seemed as if an exception to this rule
+existed in the <span class='pagenum'><a name="Page_79" id="Page_79">[79]</a></span>case of the fertilisation of the yucca blossoms by a
+little moth, <i>Pronuba yuccasella</i>. This little moth has a
+sickle-shaped appendage to its mouth-parts which occurs hi no other
+Lepidopteron, and which is used for pushing the yellow pollen into the
+opening of the pistil, thus fertilising the flower. Thus it appears as
+if a new structure, which is useful only to the plant, has arisen in
+the insect. But the difficulty is solved as soon as we learn that the
+moth lays its eggs in the fruit-buds of the Yucca, and that the
+larvae, when they emerge, feed on the developing seeds. In effecting
+the fertilisation of the flower the moth is at the same time making
+provision for its own offspring, since it is only after fertilisation
+that the seeds begin to develop. There is thus nothing to prevent our
+referring this structural adaptation in <i>Pronuba yuccasella</i> to
+processes of selection, which have gradually transformed the maxillary
+palps of the female into the sickle-shaped instrument for collecting
+the pollen, and which have at the same time developed in the insect
+the instinct to press the pollen into the pistil.</p>
+
+<p>In this domain, then, the theory of selection finds nothing but
+corroboration, and it would be impossible to substitute for it any
+other explanation, which now that the facts are so well known, could
+be regarded as a serious rival to it. That selection is a factor, and
+a very powerful factor in the evolution of organisms, can no longer be
+doubted. Even although we cannot bring forward formal proofs of it <i>in
+detail</i>, cannot calculate definitely the size of the variations which
+present themselves, and their selection-value, cannot, in short,
+reduce the whole process to a mathematical formula, yet we must assume
+selection, because it is the only possible explanation applicable to
+whole classes of phenomena, and because, on the other hand, it is made
+up of factors which we know can be proved actually to exist, and
+which, <i>if</i> they exist, must of logical necessity co&ouml;perate in the
+manner required by the theory. <i>We must accept it because the
+phenomena of evolution and adaptation must have<span class='pagenum'><a name="Page_80" id="Page_80">[80]</a></span> a natural basis, and
+because it is the only possible explanation of them.</i><a name="FNanchor_53_53" id="FNanchor_53_53"></a><a href="#Footnote_53_53" class="fnanchor">[53]</a></p>
+
+<p>Many people are willing to admit that selection explains adaptations,
+but they maintain that only a part of the phenomena are thus
+explained, because everything does not depend upon adaptation. They
+regard adaptation as, so to speak, a special effort on the part of
+Nature, which she keeps in readiness to meet particularly difficult
+claims of the external world on organisms. But if we look at the
+matter more carefully we shall find that adaptations are by no means
+exceptional, but that they are present everywhere in such enormous
+numbers, that it would be difficult in regard to any structure
+whatever, to prove that adaptation had <i>not</i> played a part in its
+evolution.</p>
+
+<p>How often has the senseless objection been urged against selection
+that it can create nothing, it can only reject. It is true that it
+cannot create either the living substance or the variations of it;
+both must be given. But in rejecting one thing it preserves another,
+intensifies it, combines it, and in this way <i>creates</i> what is new.
+<i>Everything</i> in organisms depends on adaptation; that is to say,
+everything must be admitted through the narrow door of selection,
+otherwise it can take no part in the building up of the whole. But, it
+is asked, what of the direct effect of external conditions,
+temperature, nutrition, climate and the like? Undoubtedly these can
+give rise to variations, but they too must pass through the door of
+selection, and if they cannot do this they are rejected, eliminated
+from the constitution of the species.</p>
+
+<p>It may, perhaps, be objected that such external influences are often
+of a compelling power, and that every animal must submit to them, and
+that thus selection has no choice and can neither select nor reject.
+There may be such cases; <span class='pagenum'><a name="Page_81" id="Page_81">[81]</a></span>let us assume for instance that the effect
+of the cold of the Arctic regions was to make all the mammals become
+black; the result would be that they would all be eliminated by
+selection, and that no mammals would be able to live there at all. But
+in most cases a certain percentage of animals resists these strong
+influences, and thus selection secures a foothold on which to work,
+eliminating the unfavourable variation, and establishing a useful
+colouring, consistent with what is required for the maintenance of the
+species.</p>
+
+<p>Everything depends upon adaptation! We have spoken much of adaptation
+in colouring, in connection with the examples brought into prominence
+by Darwin, because these are conspicuous, easily verified, and at the
+same time convincing for the theory of selection. But is it only
+desert and polar animals whose colouring is determined through
+adaptation? Or the leaf-butterflies, and the mimetic species, or the
+terrifying markings, and "warning-colours" and a thousand other kinds
+of sympathetic colouring? It is, indeed, never the colouring alone
+which makes up the adaptation; the structure of the animal plays a
+part, often a very essential part, in the protective disguise, and
+thus <i>many</i> variations may cooperate towards <i>one</i> common end. And it
+is to be noted that it is by no means only external parts that are
+changed; internal parts are <i>always</i> modified at the same time&mdash;for
+instance, the delicate elements of the nervous system on which depend
+the <i>instinct</i> of the insect to hold its wings, when at rest, in a
+perfectly definite position, which, in the leaf-butterfly, has the
+effect of bringing the two pieces on which the marking occurs on the
+anterior and posterior wing into the same direction, and thus
+displaying as a whole the fine curve of the midrib on the seeming
+leaf. But the wing-holding instinct is not regulated in the same way
+in all leaf-butterflies; even our indigenous species of Vanessa, with
+their protective ground-colouring, have quite a distinctive way of
+holding their wings so that the greater part of the anterior wing is
+covered by the posterior when the butterfly is at rest. But<span class='pagenum'><a name="Page_82" id="Page_82">[82]</a></span> the
+protective colouring appears on the posterior wing and on the tip of
+the anterior, <i>to precisely the distance to which it is left
+uncovered</i>. This occurs, as Standfuss has shown, in different degrees
+in our two most nearly allied species, the uncovered portion being
+smaller in <i>V. urticae</i> than in <i>V. polychloros</i>. In this case, as in
+most leaf-butterflies, the holding of the wing was probably the
+primary character; only after that was thoroughly established did the
+protective marking develop. In any case, the instinctive manner of
+holding the wings is associated with the protective colouring, and
+must remain as it is if the latter is to be effective. How greatly
+instincts may change, that is to say, may be adapted, is shown by the
+case of the Noctuid "shark" moth, <i>Xylina vetusta</i>. This form bears a
+most deceptive resemblance to a piece of rotten wood, and the
+appearance is greatly increased by the modification of the innate
+impulse to flight common to so many animals, which has here been
+transformed into an almost contrary instinct. This moth does not fly
+away from danger, but "feigns death," that is, it draws antennae, legs
+and wings close to the body, and remains perfectly motionless. It may
+be touched, picked up, and thrown down again, and still it does not
+move. This remarkable instinct must surely have developed
+simultaneously with the wood-colouring; at all events, both
+co&ouml;perating variations are now present, and prove that both the
+external and the most minute internal structure have undergone a
+process of adaptation.</p>
+
+<p>The case is the same with all structural variations of animal parts,
+which are not absolutely insignificant. When the insects acquired
+wings they must also have acquired the mechanism with which to move
+them&mdash;the musculature, and the nervous apparatus necessary for its
+automatic regulation. All instincts depend upon compound reflex
+mechanisms and are just as indispensable as the parts they have to set
+in motion, and all may have arisen through processes of selection if
+the reasons which I have elsewhere given for this view are
+correct.<a name="FNanchor_54_54" id="FNanchor_54_54"></a><a href="#Footnote_54_54" class="fnanchor">[54]</a> <span class='pagenum'><a name="Page_83" id="Page_83">[83]</a></span></p>
+
+<p>Thus there is no lack of adaptations within the organism, and
+particularly in its most important and complicated parts, so that we
+may say that there is no actively functional organ that has not
+undergone a process of adaptation relative to its function and the
+requirements of the organism. Not only is every gland structurally
+adapted, down to the very minutest histological details, to its
+function, but the function is equally minutely adapted to the needs of
+the body. Every cell in the mucous lining of the intestine is exactly
+regulated in its relation to the different nutritive substances, and
+behaves in quite a different way towards the fats, and towards
+nitrogenous substances, or peptones.</p>
+
+<p>I have elsewhere called attention to the many adaptations of the whale
+to the surrounding medium, and have pointed out&mdash;what has long been
+known, but is not universally admitted, even now&mdash;that in it a great
+number of important organs have been transformed in adaptation to the
+peculiar conditions of aquatic life, although the ancestors of the
+whale must have lived, like other hair-covered mammals, on land. I
+cited a number of these transformations&mdash;the fish-like form of the
+body, the hairlessness of the skin, the transformation of the
+fore-limbs to fins, the disappearance of the hind-limbs and the
+development of a tail fin, the layer of blubber under the skin, which
+affords the protection from cold necessary to a warm-blooded animal,
+the disappearance of the ear-muscles and the auditory passages, the
+displacement of the external nares to the forehead for the greater
+security of the breathing-hole during the brief appearance at the
+surface, and certain remarkable changes in the respiratory and
+circulatory organs which enable the animal to remain for a long time
+under water. I might have added many more, for the list of adaptations
+in the whale to aquatic life is by no means exhausted; they are found
+in the histological structure and in the minutest combinations in the
+nervous system. For it is obvious that a tail-fin must be<span class='pagenum'><a name="Page_84" id="Page_84">[84]</a></span> used in
+quite a different way from a tail, which serves as a fly-brush in
+hoofed animals, or as an aid to springing in the kangaroo or as a
+climbing organ; it will require quite different reflex-mechanisms and
+nerve combinations in the motor centres.</p>
+
+<p>I used this example in order to show how unnecessary it is to assume a
+special internal evolutionary power for the phylogenesis of species,
+for this whole order of whales is, so to speak, <i>made up of
+adaptations</i>; it deviates in many essential respects from the usual
+mammalian type, and all the deviations are adaptations to aquatic
+life. But if precisely the most essential features of the organisation
+thus depend upon adaptation, what is left for a phyletic force to do,
+since it is these essential features of the structure it would have to
+determine? There are few people now who believe in a phyletic
+evolutionary power, which is not made up of the forces known to
+us&mdash;adaptation and heredity&mdash;but the conviction that <i>every</i> part of
+an organism depends upon adaptation has not yet gained a firm footing.
+Nevertheless, I must continue to regard this conception as the correct
+one, as I have long done.</p>
+
+<p>I may be permitted one more example. The feather of a bird is a
+marvellous structure, and no one will deny that as a whole it depends
+upon adaptation. But what part of it <i>does not</i> depend upon
+adaptation? The hollow quill, the shaft with its hard, thin, light
+cortex, and the spongy substance within it, its square section
+compared with the round section of the quill, the flat barbs, their
+short, hooked barbules which, in the flight-feathers, hook into one
+another with just sufficient firmness to resist the pressure of the
+air at each wing-beat, the lightness and firmness of the whole
+apparatus, the elasticity of the vane, and so on. And yet all this
+belongs to an organ which is only passively functional, and therefore
+can have nothing to do with the <i>Lamarckian principle</i>. Nor can the
+feather have arisen through some magical effect of temperature,
+moisture, electricity, or spe<span class='pagenum'><a name="Page_85" id="Page_85">[85]</a></span>cific nutrition, and thus selection is
+again our only anchor of safety.</p>
+
+<p>But&mdash;it will be objected&mdash;the substance of which the feather consists,
+this peculiar kind of horny substance, did not first arise through
+selection in the course of the evolution of the birds, for it formed
+the covering of the scales of their reptilian ancestors. It is quite
+true that a similar substance covered the scales of the Reptiles, but
+why should it not have arisen among them through selection? Or in what
+other way could it have arisen, since scales are also passively useful
+parts? It is true that if we are only to call adaptation what has been
+acquired by the species we happen to be considering, there would
+remain a great deal that could not be referred to selection; but we
+are postulating an evolution which has stretched back through aeons,
+and in the course of which innumerable adaptations took place, which
+had not merely ephemeral persistence in a genus, a family or a class,
+but which was continued into whole Phyla of animals, with continual
+fresh adaptations to the special conditions of each species, family,
+or class, yet with persistence of the fundamental elements. Thus the
+feather, once acquired, persisted in all birds, and the vertebral
+column, once gained by adaptation in the lowest forms, has persisted
+in all the Vertebrates from Amphioxus upwards, although with constant
+readaptation to the conditions of each particular group. Thus
+everything we can see in animals is adaptation, whether of to-day, or
+of yesterday, or of ages long gone by; every kind of cell, whether
+glandular, muscular, nervous, epidermic, or skeletal, is adapted to
+absolutely definite and specific functions, and every organ which is
+composed of these different kinds of cells contains them in the proper
+proportions, and in the particular arrangement which best serves the
+function of the organ; it is thus adapted to its function.</p>
+
+<p>All parts of the organism are tuned to one another, that is, <i>they are
+adapted to one another</i>, and in the same way<span class='pagenum'><a name="Page_86" id="Page_86">[86]</a></span> <i>the organism as a whole
+is adapted to the conditions of its life, and it is so at every stage
+of its evolution.</i></p>
+
+<p>But all adaptations <i>can</i> be referred to selection; the only point
+that remains doubtful is whether they all <i>must</i> be referred to it.</p>
+
+<p>However that may be, whether the <i>Lamarckian principle</i> is a factor
+that has co&ouml;perated with selection in evolution, or whether it is
+altogether fallacious, the fact remains, that selection is the cause
+of a great part of the phyletic evolution of organisms on our earth.
+Those who agree with me in rejecting the <i>Lamarckian principle</i> will
+regard selection as the only <i>guiding</i> factor in evolution, which
+creates what is new out of the transmissible variations, by ordering
+and arranging these, selecting them in relation to their number and
+size, as the architect does his building-stones so that a particular
+style must result.<a name="FNanchor_55_55" id="FNanchor_55_55"></a><a href="#Footnote_55_55" class="fnanchor">[55]</a> But the building-stones themselves, the
+variations, have their basis in the influences which cause variation
+in those vital units which are handed on from one generation to
+another, whether, taken together they form the <i>whole</i> organism, as in
+Bacteria and other low forms of life, or only a germ-substance, as in
+unicellular and multicellular organisms. </p>
+
+<div class="footnotes"><h3>FOOTNOTES:</h3>
+
+<div class="footnote"><p><a name="Footnote_33_33" id="Footnote_33_33"></a><a href="#FNanchor_33_33"><span class="label">[33]</span></a> <i>Vortr&auml;ge &uuml;ber Descendenztheorie</i>, Jena, 1904, <span class="smcap">ii</span>. 269.
+Eng. Transl. London, 1904, <span class="smcap">ii</span>. p. 317.</p></div>
+
+<div class="footnote"><p><a name="Footnote_34_34" id="Footnote_34_34"></a><a href="#FNanchor_34_34"><span class="label">[34]</span></a> See Poulton, <i>Essays on Evolution</i>, Oxford, 1908. pp.
+xix-xxii.</p></div>
+
+<div class="footnote"><p><a name="Footnote_35_35" id="Footnote_35_35"></a><a href="#FNanchor_35_35"><span class="label">[35]</span></a> <i>Origin of Species</i> (6th edit), pp. 176 <i>et seq.</i></p></div>
+
+<div class="footnote"><p><a name="Footnote_36_36" id="Footnote_36_36"></a><a href="#FNanchor_36_36"><span class="label">[36]</span></a> Chun, <i>Reise der Valdivia</i>, Leipzig, 1904.</p></div>
+
+<div class="footnote"><p><a name="Footnote_37_37" id="Footnote_37_37"></a><a href="#FNanchor_37_37"><span class="label">[37]</span></a> Plate, <i>Selektionsprinzip u. Probleme der Artbildung</i>
+(3rd edit.), Leipzig, 1908.</p></div>
+
+<div class="footnote"><p><a name="Footnote_38_38" id="Footnote_38_38"></a><a href="#FNanchor_38_38"><span class="label">[38]</span></a> <i>Studien zur Descendenz-Theorie</i> <span class="smcap">ii</span>., "Die Enstehung der
+Zeichnung bei den Schmetterlings-raupen," Leipzig, 1876.</p></div>
+
+<div class="footnote"><p><a name="Footnote_39_39" id="Footnote_39_39"></a><a href="#FNanchor_39_39"><span class="label">[39]</span></a> <i>Origin of Species</i> (6th edit.), p. 232.</p></div>
+
+<div class="footnote"><p><a name="Footnote_40_40" id="Footnote_40_40"></a><a href="#FNanchor_40_40"><span class="label">[40]</span></a> <i>Origin of Species</i>, p. 233; see also edit. 1, p. 242.</p></div>
+
+<div class="footnote"><p><a name="Footnote_41_41" id="Footnote_41_41"></a><a href="#FNanchor_41_41"><span class="label">[41]</span></a> <i>Ibid.</i> p. 230.</p></div>
+
+<div class="footnote"><p><a name="Footnote_42_42" id="Footnote_42_42"></a><a href="#FNanchor_42_42"><span class="label">[42]</span></a> <i>The Effect of External Influences upon Development</i>,
+Romanes Lecture, Oxford, 1894.</p></div>
+
+<div class="footnote"><p><a name="Footnote_43_43" id="Footnote_43_43"></a><a href="#FNanchor_43_43"><span class="label">[43]</span></a> See Poulton, <i>Essays on Evolution</i>, 1908, pp. 316, 317.</p></div>
+
+<div class="footnote"><p><a name="Footnote_44_44" id="Footnote_44_44"></a><a href="#FNanchor_44_44"><span class="label">[44]</span></a> <i>The Evolution Theory</i>, London, 1904, <span class="smcap">i</span>. p. 219.</p></div>
+
+<div class="footnote"><p><a name="Footnote_45_45" id="Footnote_45_45"></a><a href="#FNanchor_45_45"><span class="label">[45]</span></a> <i>Report of the British Association</i> (Bristol, 1898),
+London, 1899, pp. 906-909.</p></div>
+
+<div class="footnote"><p><a name="Footnote_46_46" id="Footnote_46_46"></a><a href="#FNanchor_46_46"><span class="label">[46]</span></a> <i>Proc. Ent. Soc.</i>, London, May 6, 1903.</p></div>
+
+<div class="footnote"><p><a name="Footnote_47_47" id="Footnote_47_47"></a><a href="#FNanchor_47_47"><span class="label">[47]</span></a> <i>Essays on Evolution</i>, 1889-1907, Oxford, 1908,
+<i>passim</i>, e.g. p. 269.</p></div>
+
+<div class="footnote"><p><a name="Footnote_48_48" id="Footnote_48_48"></a><a href="#FNanchor_48_48"><span class="label">[48]</span></a> The expression does not refer to all the enemies of this
+butterfly; against ichneumon-flies, for instance, their unpleasant
+smell usually gives no protection.</p></div>
+
+<div class="footnote"><p><a name="Footnote_49_49" id="Footnote_49_49"></a><a href="#FNanchor_49_49"><span class="label">[49]</span></a> Professor Poulton has corrected some wrong descriptions
+which I had unfortunately overlooked in the Plates of my book
+<i>Vortr&auml;ge &uuml;ber Descendenztheorie</i>, and which refer to <i>Papilio
+dardanus</i> (<i>merope</i>). These mistakes are of no importance as far as an
+understanding of the mimicry-theory is concerned, but I hope shortly
+to be able to correct them in a later edition.</p></div>
+
+<div class="footnote"><p><a name="Footnote_50_50" id="Footnote_50_50"></a><a href="#FNanchor_50_50"><span class="label">[50]</span></a> <i>Journ. Linn. Soc. London</i> (<i>Zool.</i>), Vol. xxvi. 1898,
+pp. 598-602.</p></div>
+
+<div class="footnote"><p><a name="Footnote_51_51" id="Footnote_51_51"></a><a href="#FNanchor_51_51"><span class="label">[51]</span></a> In <i>Kosmos</i>, 1879, p. 100.</p></div>
+
+<div class="footnote"><p><a name="Footnote_52_52" id="Footnote_52_52"></a><a href="#FNanchor_52_52"><span class="label">[52]</span></a> <i>Habit and Instinct</i>, London. 1896.</p></div>
+
+<div class="footnote"><p><a name="Footnote_53_53" id="Footnote_53_53"></a><a href="#FNanchor_53_53"><span class="label">[53]</span></a> This has been discussed in many of my earlier works. See
+for instance <i>The All-Sufficiency of Natural Selection, a reply to
+Herbert Spencer</i>, London, 1893.</p></div>
+
+<div class="footnote"><p><a name="Footnote_54_54" id="Footnote_54_54"></a><a href="#FNanchor_54_54"><span class="label">[54]</span></a> <i>The Evolution Theory</i>, London, 1904, p. 144.</p></div>
+
+<div class="footnote"><p><a name="Footnote_55_55" id="Footnote_55_55"></a><a href="#FNanchor_55_55"><span class="label">[55]</span></a> <i>Variation under Domestication</i>, 1875, <span class="smcap">ii</span>. pp. 426,
+427.</p></div>
+</div>
+
+
+<hr style="width: 65%;" />
+<p><span class='pagenum'><a name="Page_87" id="Page_87">[87]</a></span></p>
+<h2><a name="III" id="III"></a>III</h2>
+
+<h2>HEREDITY AND VARIATION IN MODERN LIGHTS</h2>
+<p class="center"><span class="smcap1">By W. Bateson, M.A., F.R.S.</span></p>
+<h4><i>Professor of Biology in the University of Cambridge</i></h4>
+
+<p>Darwin's work has the property of greatness in that it may be admired
+from more aspects than one. For some the perception of the principle
+of Natural Selection stands out as his most wonderful achievement to
+which all the rest is subordinate. Others, among whom I would range
+myself, look up to him rather as the first who plainly distinguished,
+collected, and comprehensively studied that new class of evidence from
+which hereafter a true understanding of the process of Evolution may
+be developed. We each prefer our own standpoint of admiration; but I
+think that it will be in their wider aspect that his labours will most
+command the veneration of posterity.</p>
+
+<p>A treatise written to advance knowledge may be read in two moods. The
+reader may keep his mind passive, willing merely to receive the
+impress of the writer's thought; or he may read with his attention
+strained and alert, asking at every instant how the new knowledge can
+be used in a further advance, watching continually for fresh footholds
+by which to climb higher still. Of Shelley it has been said that he
+was a poet for poets: so Darwin was a naturalist for naturalists. It
+is when his writings are used in the critical and more exacting spirit
+with which we test the outfit for our own enterprise that we learn
+their full value and<span class='pagenum'><a name="Page_88" id="Page_88">[88]</a></span> strength. Whether we glance back and compare his
+performance with the efforts of his predecessors, or look forward
+along the course which modern research is disclosing, we shall honour
+most in him not the rounded merit of finite accomplishment, but the
+creative power by which he inaugurated a line of discovery endless in
+variety and extension. Let us attempt thus to see his work in true
+perspective between the past from which it grew, and the present which
+is its consequence. Darwin attacked the problem of Evolution by
+reference to facts of three classes: Variation; Heredity; Natural
+Selection. His work was not as the laity suppose, a sudden and
+unheralded revelation, but the first fruit of a long and hitherto
+barren controversy. The occurrence of variation from type, and the
+hereditary transmission of such variation had of course been long
+familiar to practical men, and inferences as to the possible bearing
+of those phenomena on the nature of specific difference had been from
+time to time drawn by naturalists. Maupertuis, for example, wrote: "Ce
+qui nous reste &agrave; examiner, c'est comment d'un seul individu, il a pu
+na&icirc;tre tant d'esp&egrave;ces si diff&eacute;rentes." And again: "La Nature contient
+le fonds de toutes ces vari&eacute;t&eacute;s: mais le hasard ou l'art les mettent
+en &oelig;uvre. C'est ainsi que ceux dont l'industrie s'applique &agrave;
+satisfaire le go&ucirc;t des curieux, sont, pour ainsi dire, cr&eacute;ateurs
+d'esp&egrave;ces nouvelles."<a name="FNanchor_56_56" id="FNanchor_56_56"></a><a href="#Footnote_56_56" class="fnanchor">[56]</a></p>
+
+<p>Such passages, of which many (though few so emphatic) can be found in
+eighteenth century writers, indicate a true perception of the mode of
+Evolution. The speculations hinted at by Buffon,<a name="FNanchor_57_57" id="FNanchor_57_57"></a><a href="#Footnote_57_57" class="fnanchor">[57]</a> developed by
+Erasmus Darwin, and <span class='pagenum'><a name="Page_89" id="Page_89">[89]</a></span>independently proclaimed above all by Lamarck,
+gave to the doctrine of descent a wide renown. The uniformitarian
+teaching which Lyell deduced from geological observation had gained
+acceptance. The facts of geographical distribution<a name="FNanchor_58_58" id="FNanchor_58_58"></a><a href="#Footnote_58_58" class="fnanchor">[58]</a> had been shown
+to be obviously inconsistent with the Mosaic legend. Prichard, and
+Lawrence, following the example of Blumenbach, had successfully
+demonstrated that the races of Man could be regarded as different
+forms of one species, contrary to the opinion up till then received.
+These treatises all begin, it is true, with a profound obeisance to
+the sons of Noah, but that performed, they continue on strictly modern
+lines. The question of the mutability of species was thus prominently
+raised.</p>
+
+<p>Those who rate Lamarck no higher than did Huxley in his contemptuous
+phrase "<i>buccinator tantum</i>," will scarcely deny that the sound of the
+trumpet had carried far, or that its note was clear. If then there
+were few who had already turned to evolution with positive conviction,
+all scientific men must at least have known that such views had been
+promulgated; and many must, as Huxley says, have taken up his own
+position of "critical expectancy."<a name="FNanchor_59_59" id="FNanchor_59_59"></a><a href="#Footnote_59_59" class="fnanchor">[59]</a> <span class='pagenum'><a name="Page_90" id="Page_90">[90]</a></span></p>
+
+<p>Why, then, was it, that Darwin succeeded where the rest had failed?
+The cause of that success was twofold. First, and obviously, in the
+principle of Natural Selection he had a suggestion which would work.
+It might not go the whole way, but it was true as far as it went.
+Evolution could thus in great measure be fairly represented as a
+consequence of demonstrable processes. Darwin seldom endangers the
+mechanism he devised by putting on it strains much greater than it can
+bear. He at least was under no illusion as to the omnipotence of
+Selection; and he introduces none of the forced pleading which in
+recent years has threatened to discredit that principle.</p>
+
+<p><span class='pagenum'><a name="Page_91" id="Page_91">[91]</a></span></p>
+
+<p>For example, in the latest text of the <i>Origin</i><a name="FNanchor_60_60" id="FNanchor_60_60"></a><a href="#Footnote_60_60" class="fnanchor">[60]</a> we find him
+saying:</p>
+
+<div class="blockquot"><p>"But as my conclusions have lately been much misrepresented,
+and it has been stated that I attribute the modification of
+species exclusively to natural selection, I may be permitted
+to remark that in the first edition of this work, and
+subsequently, I placed in a most conspicuous
+position&mdash;namely, at the close of the Introduction&mdash;the
+following words: 'I am convinced that natural selection has
+been the main but not the exclusive means of modification.'" </p></div>
+
+<p>But apart from the invention of this reasonable hypothesis, which may
+well, as Huxley estimated, "be the guide of biological and
+psychological speculation for the next three or four generations,"
+Darwin made a more significant and imperishable contribution. Not for
+a few generations, but through all ages he should be remembered as the
+first who showed clearly that the problems of Heredity and Variation
+are soluble by observation, and laid down the course by which we must
+proceed to their solution.<a name="FNanchor_61_61" id="FNanchor_61_61"></a><a href="#Footnote_61_61" class="fnanchor">[61]</a> The moment of inspiration did not come
+with the reading of Malthus, but with the opening of the "first
+note-book on Transmutation of Species."<a name="FNanchor_62_62" id="FNanchor_62_62"></a><a href="#Footnote_62_62" class="fnanchor">[62]</a> Evolution is a process of
+Variation and Heredity. The older writers, though they had some vague
+idea that it must be so, did not study Variation and Heredity. Darwin
+did, and so begat not a theory, but a science.</p>
+
+<p><span class='pagenum'><a name="Page_92" id="Page_92">[92]</a></span></p><p>The extent to which this is true, the scientific world is only
+beginning to realise. So little was the fact appreciated in Darwin's
+own time that the success of his writings was followed by an almost
+total cessation of work in that special field. Of the causes which led
+to these remarkable consequences I have spoken elsewhere. They
+proceeded from circumstances peculiar to the time; but whatever the
+causes there is no doubt that this statement of the result is
+historically exact, and those who make it their business to collect
+facts elucidating the physiology of Heredity and Variation are well
+aware that they will find little to reward their quest in the leading
+scientific Journals of the Darwinian epoch.</p>
+
+<p>In those thirty years the original stock of evidence current and in
+circulation even underwent a process of attrition. As in the story of
+the Eastern sage who first wrote the collected learning of the
+universe for his sons in a thousand volumes and by successive
+compression and burning reduced them to one and from this by further
+burning distilled the single ejaculation of the Faith "There is no god
+but God and Mohammed is the Prophet of God," which was all his maturer
+wisdom deemed essential:&mdash;so in the books of that period do we find
+the <i>corpus</i> of genetic knowledge dwindle to a few prerogative
+instances and these at last to the brief formula of an unquestioned
+creed.</p>
+
+<p>And yet in all else that concerns biological science this period was,
+in very truth, our Golden Age, when the natural history of the earth
+was explored as never before; morphology and embryology were
+exhaustively ransacked; the physiology of plants and animals began to
+rival chemistry and physics in precision of method and in the rapidity
+of its advances; and the foundations of pathology were laid.</p>
+
+<p>In contrast with this immense activity elsewhere the neglect which
+befel the special physiology of Descent, or Genetics as we now call
+it, is astonishing. This may of course be interpreted as meaning that
+the favoured studies seemed to promise a quicker return for effort,
+but it would be more true to say that those who chose these other
+pursuits did<span class='pagenum'><a name="Page_93" id="Page_93">[93]</a></span> so without making any such comparison; for the idea that
+the physiology of Heredity and Variation was a coherent science,
+offering possibilities of extraordinary discovery, was not present to
+their minds at all. In a word, the existence of such a science was
+well nigh forgotten. It is true that in ancillary periodicals, as for
+example those that treat of entomology or horticulture, or in the
+writings of the already isolated systematists,<a name="FNanchor_63_63" id="FNanchor_63_63"></a><a href="#Footnote_63_63" class="fnanchor">[63]</a> observations with
+this special bearing were from time to time related, but the class of
+fact on which Darwin built his conceptions of Heredity and Variation
+was not seen in the highways of biology. It formed no part of the
+official curriculum of biological students, and found no place among
+the subjects which their teachers were investigating.</p>
+
+<p>During this period nevertheless one distinct advance was made, that
+with which Weismann's name is prominently connected. In Darwin's
+genetic scheme the hereditary transmission of parental experience and
+its consequences played a considerable role. Exactly how great that
+role was supposed to be, he with his habitual caution refrained from
+specifying, for the sufficient reason that he did not know.
+Nevertheless much of the process of Evolution, especially that by
+which organs have become degenerate and rudimentary, was certainly
+attributed by Darwin to such inheritance, <span class='pagenum'><a name="Page_94" id="Page_94">[94]</a></span>though since belief in the
+inheritance of acquired characters fell into dispute, the fact has
+been a good deal overlooked. The <i>Origin</i> without "use and disuse"
+would be a materially different book. A certain vacillation is
+discernible in Darwin's utterances on this question, and the fact gave
+to the astute Butler an opportunity for his most telling attack. The
+discussion which best illustrates the genetic views of the period
+arose in regard to the production of the rudimentary condition of the
+wings of many beetles in the Madeira group of islands, and by
+comparing passages from the <i>Origin</i><a name="FNanchor_64_64" id="FNanchor_64_64"></a><a href="#Footnote_64_64" class="fnanchor">[64]</a> Butler convicts Darwin of
+saying first that this condition was in the main the result of
+Selection, with disuse aiding, and in another place that the main
+cause of degeneration was disuse, but that Selection had aided. To
+Darwin however I think the point would have seemed one of dialetics
+merely. To him the one paramount purpose was to show that somehow an
+Evolution by means of Variation and Heredity might have brought about
+the facts observed, and whether they had come to pass in the one way
+or the other was a matter of subordinate concern.</p>
+
+<p>To us moderns the question at issue has a diminished significance. For
+over all such debates a change has been brought by Weismann's
+challenge for evidence that use and disuse have any transmitted
+effects at all. Hitherto the transmission of many acquired
+characteristics had seemed to most naturalists so obvious as not to
+call for demonstration.<a name="FNanchor_65_65" id="FNanchor_65_65"></a><a href="#Footnote_65_65" class="fnanchor">[65]</a> Weismann's demand for facts in support of
+the main <span class='pagenum'><a name="Page_95" id="Page_95">[95]</a></span>proposition revealed at once that none having real cogency
+could be produced. The time-honoured examples were easily shown to be
+capable of different explanations. A few certainly remain which cannot
+be so summarily dismissed, but&mdash;though it is manifestly impossible
+here to do justice to such a subject&mdash;I think no one will dispute that
+these residual and doubtful phenomena, whatever be their true nature,
+are not of a kind to help us much in the interpretation of any of
+those complex cases of adaptation which on the hypothesis of unguided
+Natural Selection are especially difficult to understand. Use and
+disuse were invoked expressly to help us over these hard places; but
+whatever changes can be induced in offspring by direct treatment of
+the parents, they are not of a kind to encourage hope of real
+assistance from that quarter. It is not to be denied that through the
+collapse of this second line of argument the Selection hypothesis has
+had to take an increased and perilous burden. Various ways of meeting
+the difficulty have been proposed, but these mostly resolve themselves
+into improbable attempts to expand or magnify the powers of Natural
+Selection.</p>
+
+<p>Weismann's interpellation, though negative in purpose, has had a
+lasting and beneficial effect, for through his thorough demolition of
+the old loose and distracting notions of inherited experience, the
+ground has been cleared for the construction of a true knowledge of
+heredity based on experimental fact.</p>
+
+<p>In another way he made a contribution of a more positive character,
+for his elaborate speculations as to the genetic meaning of
+cytological appearances have led to a minute investigation of the
+visible phenomena occurring in those cell-divisions by which
+germ-cells arise. Though the particular views he advocated have very
+largely proved incompatible with the observed facts of heredity, yet
+we must acknowledge that it was chiefly through the stimulus of
+Weismann's ideas that those advances in cytology were made; and though
+the doctrine of the continuity of germ<span class='pagenum'><a name="Page_96" id="Page_96">[96]</a></span>-plasm cannot be maintained in
+the form originally propounded, it is in the main true and
+illuminating.<a name="FNanchor_66_66" id="FNanchor_66_66"></a><a href="#Footnote_66_66" class="fnanchor">[66]</a> Nevertheless in the present state of knowledge we
+are still as a rule quite unable to connect cytological appearances
+with any genetic consequences and save in one respect (obviously of
+extreme importance&mdash;to be spoken of later) the two sets of phenomena
+might, for all we can see, be entirely distinct.</p>
+
+<p>I cannot avoid attaching importance to this want of connection between
+the nuclear phenomena and the features of bodily organisation. All
+attempts to investigate Heredity by cytological means lie under the
+disadvantage that it is the nuclear changes which can alone be
+effectively observed. Important as they must surely be, I have never
+been persuaded that the rest of the cell counts for nothing. What we
+know of the behaviour and variability of chromosomes seems in my
+opinion quite incompatible with the belief that they alone govern
+form, and are the sole agents responsible in heredity.<a name="FNanchor_67_67" id="FNanchor_67_67"></a><a href="#Footnote_67_67" class="fnanchor">[67]</a></p>
+
+<p>If, then, progress was to be made in Genetics, work of a different
+kind was required. To learn the laws of Heredity and Variation there
+is no other way than that which Darwin <span class='pagenum'><a name="Page_97" id="Page_97">[97]</a></span>himself followed, the direct
+examination of the phenomena. A beginning could be made by collecting
+fortuitous observations of this class, which have often thrown a
+suggestive light, but such evidence can be at best but superficial and
+some more penetrating instrument of research is required. This can
+only be provided by actual experiments in breeding.</p>
+
+<p>The truth of these general considerations was becoming gradually clear
+to many of us when in 1900 Mendel's work was rediscovered.
+Segregation, a phenomenon of the utmost novelty, was thus revealed.
+From that moment not only in the problem of the origin of species, but
+in all the great problems of biology a new era began. So unexpected
+was the discovery that many naturalists were convinced it was untrue,
+and at once proclaimed Mendel's conclusions as either altogether
+mistaken, or if true, of very limited application. Many fantastic
+notions about the workings of Heredity had been asserted as general
+principles before: this was probably only another fancy of the same
+class.</p>
+
+<p>Nevertheless those who had a preliminary acquaintance with the facts
+of Variation were not wholly unprepared for some such revelation. The
+essential deduction from the discovery of segregation was that the
+characters of living things are dependent on the presence of definite
+elements or factors, which are treated as units in the processes of
+Heredity. These factors can thus be recombined in various ways. They
+act sometimes separately, and sometimes they interact in conduction
+with each other, producing their various effects. All this indicates a
+definiteness and specific order in heredity, and therefore in
+variation. This order cannot by the nature of the case be dependent on
+Natural Selection for its existence, but must be a consequence of the
+fundamental chemical and physical nature of living things. The study
+of Variation had from the first shown that an orderliness of this kind
+was present. The bodies and the properties of livings things are
+cosmic, not chaotic. No matter how low in the scale we go, never do we
+find the slightest hint of a diminution in that all-pervading
+orderli<span class='pagenum'><a name="Page_98" id="Page_98">[98]</a></span>ness, nor can we conceive an organism existing for a moment in
+any other state. Moreover not only does this order prevail in normal
+forms, but again and again it is to be seen in newly-sprung varieties,
+which by general consent cannot have been subjected to a prolonged
+Selection. The discovery of Mendelian elements admirably coincided
+with and at once gave a rationale of these facts. Genetic Variation is
+then primarily the consequence of additions to, or omissions from, the
+stock of elements which the species contains. The further
+investigation of the species-problem must thus proceed by the
+analytical method which breeding experiments provide.</p>
+
+<p>In the nine years which have elapsed since Mendel's clue became
+generally known, progress has been rapid. We now understand the
+process by which a polymorphic race maintains its polymorphism. When a
+family consists of dissimilar members, given the numerical proportions
+in which these members are occurring, we can represent their
+composition symbolically and state what types can be transmitted by
+the various members. The difficulty of the "swamping effects of
+inter-crossing" is practically at an end. Even the famous puzzle of
+sex-limited inheritance is solved, at all events in its more regular
+manifestations, and we know now how it is brought about that the
+normal sisters of a colour-blind man can transmit the colour-blindness
+while his normal brothers cannot transmit it.</p>
+
+<p>We are still only on the fringe of the inquiry. It can be seen
+extending and ramifying in many directions. To enumerate these here
+would be impossible. A whole new range of possibilities is being
+brought into view by study of the inter-relations between the simple
+factors. By following up the evidence as to segregation, indications
+have been obtained which can only be interpreted as meaning that when
+many factors are being simultaneously redistributed among the
+germ-cells, certain of them exert what must be described as a
+repulsion upon other factors. We cannot surmise whither this discovery
+may lead.<span class='pagenum'><a name="Page_99" id="Page_99">[99]</a></span></p>
+
+<p>In the new light all the old problems wear a fresh aspect. Upon the
+question of the nature of Sex, for example, the bearing of Mendelian
+evidence is close. Elsewhere I have shown that from several sets of
+parallel experiments the conclusion is almost forced upon us that, in
+the types investigated, of the two sexes the female is to be regarded
+as heterozygous in sex, containing one unpaired dominant element,
+while the male is similarly homozygous in the absence of that
+element.<a name="FNanchor_68_68" id="FNanchor_68_68"></a><a href="#Footnote_68_68" class="fnanchor">[68]</a> It is not a little remarkable that on this point&mdash;which
+is the only one where observations of the nuclear processes of
+gameto-genesis have yet been brought into relation with the visible
+characteristics of the organisms themselves&mdash;there should be
+diametrical opposition between the results of breeding experiments and
+those derived from cytology.</p>
+
+<p>Those who have followed the researches of the American school will be
+aware that, after it had been found in certain insects that the
+spermatozoa were of two kinds according as they contained or did not
+contain the accessory chromosome, E. B. Wilson succeeded in proving
+that the sperms possessing this accessory body were destined to form
+<i>females</i> on fertilisation, while sperms without it form males, the
+eggs being apparently indifferent. Perhaps the most striking of all
+this series of observations is that lately made by T. H. Morgan,<a name="FNanchor_69_69" id="FNanchor_69_69"></a><a href="#Footnote_69_69" class="fnanchor">[69]</a>
+since confirmed by von Baehr, that in a Phylloxeran two kinds of
+spermatids are formed, respectively with and without an accessory (in
+this case, <i>double</i>) chromosome. Of these, only those possessing the
+accessory body become functional spermatozoa, the others degenerating.
+We have thus an elucidation of the puzzling fact that in these forms
+fertilisation results in the formation of <i>females</i> only. How the
+males are formed&mdash;for of course males are eventually produced by the
+parthenogenetic females&mdash;we do not know.</p>
+
+<p><span class='pagenum'><a name="Page_100" id="Page_100">[100]</a></span></p><p>If the accessory body is really to be regarded as bearing the factor
+for femaleness, then in Mendelian terms female is DD and male is DR.
+The eggs are indifferent and the spermatozoa are each male, <i>or</i>
+female. But according to the evidence derived from a study of the
+sex-limited descent of certain features in other animals the
+conclusion seems equally clear that in them female must be regarded as
+DR and male as RR. The eggs are thus each either male or female and
+the spermatozoa are indifferent. How this contradictory evidence is to
+be reconciled we do not yet know. The breeding work concerns fowls,
+canaries, and the Currant moth (<i>Abraxas grossulariata</i>). The
+accessory chromosome has been now observed in most of the great
+divisions of insects,<a name="FNanchor_70_70" id="FNanchor_70_70"></a><a href="#Footnote_70_70" class="fnanchor">[70]</a> except, as it happens, Lepidoptera. At first
+sight it seems difficult to suppose that a feature apparently so
+fundamental as sex should be differently constituted in different
+animals, but that seems at present the least improbable inference. I
+mention these two groups of facts as illustrating the nature and
+methods of modern genetic work. We must proceed by minute and specific
+analytical investigation. Wherever we look we find traces of the
+operation of precise and specific rules.</p>
+
+<p>In the light of present knowledge it is evident that before we can
+attack the Species-problem with any hope of success there are vast
+arrears to be made up. He would be a bold man who would now assert
+that there was no sense in which the term Species might not have a
+strict and concrete meaning in contradistinction to the term Variety.
+We have been taught to regard the difference between species and
+variety as one of degree. I think it unlikely that this conclusion
+will bear the test of further research. To Darwin <span class='pagenum'><a name="Page_101" id="Page_101">[101]</a></span>the question, What
+is a variation? presented no difficulties. Any difference between
+parent and offspring was a variation. Now we have to be more precise.
+First we must, as de Vries has shown, distinguish real, genetic,
+variation from <i>fluctuational</i> variations, due to environmental and
+other accidents, which cannot be transmitted. Having excluded these
+sources of error the variations observed must be expressed in terms of
+the factors to which they are due before their significance can be
+understood. For example, numbers of the variations seen under
+domestication, and not a few witnessed in nature, are simply the
+consequence of some ingredient being in an unknown way omitted from
+the composition of the varying individual. The variation may on the
+contrary be due to the addition of some new element, but to prove that
+it is so is by no means an easy matter. Casual observation is useless,
+for though these latter variations will always be dominants, yet many
+dominant characteristics may arise from another cause, namely the
+meeting of complementary factors, and special study of each case in
+two generations at least is needed before these two phenomena can be
+distinguished.</p>
+
+<p>When such considerations are fully appreciated it will be realised
+that medleys of most dissimilar occurrences are all confused together
+under the term Variation. One of the first objects of genetic analysis
+is to disentangle this mass of confusion.</p>
+
+<p>To those who have made no study of heredity it sometimes appears that
+the question of the effect of conditions in causing variation is one
+which we should immediately investigate, but a little thought will
+show that before any critical inquiry into such possibilities can be
+attempted, a knowledge of the working of heredity under conditions as
+far as possible uniform must be obtained. At the time when Darwin was
+writing, if a plant brought into cultivation gave off an albino
+variety, such an event was without hesitation ascribed to the change
+of life. Now we see that albino <i>gametes</i>, germs, that is to say,
+which are destitute of<span class='pagenum'><a name="Page_102" id="Page_102">[102]</a></span> the pigment-forming factor, may have been
+originally produced by individuals standing an indefinite number of
+generations back in the ancestry of the actual albino, and it is
+indeed almost certain that the variation to which the appearance of
+the albino is due cannot have taken place in a generation later than
+that of the grandparents. It is true that when a new <i>dominant</i>
+appears we should feel greater confidence that we were witnessing the
+original variation, but such events are of extreme rarity, and no such
+case has come under the notice of an experimenter in modern times, as
+far as I am aware. That they must have appeared is clear enough.
+Nothing corresponding to the Brown-breasted Game fowl is known wild,
+yet that colour is a most definite dominant, and at some moment since
+<i>Gallus bankiva</i> was domesticated, the element on which that special
+colour depends must have at least once been formed in the germ-cell of
+a fowl; but we need harder evidence than any which has yet been
+produced before we can declare that this novelty came through
+over-feeding, or change of climate, or any other disturbance
+consequent on domestication. When we reflect on the intricacies of
+genetic problems as we must now conceive them there come moments when
+we feel almost thankful that the Mendelian principles were unknown to
+Darwin. The time called for a bold pronouncement, and he made it, to
+our lasting profit and delight. With fuller knowledge we pass once
+more into a period of cautious expectation and reserve.</p>
+
+<p>In every arduous enterprise it is pleasanter to look back at
+difficulties overcome than forward to those which still seem
+insurmountable, but in the next stage there is nothing to be stained
+by disguising the fact that the attributes of living things are not
+what we used to suppose. If they are more complex in the sense that
+the properties they display are throughout so regular<a name="FNanchor_71_71" id="FNanchor_71_71"></a><a href="#Footnote_71_71" class="fnanchor">[71]</a> that the
+Selection of minute random <span class='pagenum'><a name="Page_103" id="Page_103">[103]</a></span>variations is an unacceptable account of
+the origin of their diversity, yet by virtue of that very regularity
+the problem is limited in scope and thus simplified.</p>
+
+<p>To begin with, we must relegate Selection to its proper place.
+Selection permits the viable to continue and decides that the
+non-viable shall perish; just as the temperature of our atmosphere
+decides that no liquid carbon shall be found on the face of the earth:
+but we do not suppose that the form of the diamond has been gradually
+achieved by a process of Selection. So again, as the course of descent
+branches in the successive generations, Selection determines along
+which branch Evolution shall proceed, but it does not decide what
+novelties that branch shall bring forth. "<i>La Nature contient le fonds
+de toutes ces vari&eacute;t&eacute;s, mais le hazard ou l'art les mettent en
+&oelig;uvre</i>," as Maupertuis most truly said.</p>
+
+<p>Not till knowledge of the genetic properties of organisms has attained
+to far greater completeness can evolutionary speculations have more
+than a suggestive value. By genetic experiment, cytology and
+physiological chemistry aiding, we may hope to acquire such knowledge.
+In 1872 Nathusius wrote:<a name="FNanchor_72_72" id="FNanchor_72_72"></a><a href="#Footnote_72_72" class="fnanchor">[72]</a> "Das Gesetz der Vererbung ist noch nicht
+erkannt; der Apfel ist noch nicht vom Baum der Erkenntniss gefallen,
+welcher, der Sage nach, Newton auf den rechten Weg zur Ergr&uuml;ndung der
+Gravitationsgesetze f&uuml;hrte." We cannot pretend that the words are not
+still true, but in Mendelian analysis the seeds of that apple-tree at
+last are sown.</p>
+
+<p>If we were asked what discovery would do most to forward our inquiry,
+what one bit of knowledge would more than any other illuminate the
+problem, I think we may give the answer without hesitation. The
+greatest advance that we can foresee will be made when it is found
+possible to <span class='pagenum'><a name="Page_104" id="Page_104">[104]</a></span>connect the geometrical phenomena of development with the
+chemical. The geometrical symmetry of living things is the key to a
+knowledge of their regularity, and the forces which cause it. In the
+symmetry of the dividing cell the basis of that resemblance we call
+Heredity is contained. To imitate the morphological phenomena of life
+we have to devise a system which can divide. It must be able to
+divide, and to segment as&mdash;grossly&mdash;a vibrating plate or rod does, or
+as an icicle can do as it becomes ribbed in a continuous stream of
+water; but with this distinction, that the distribution of chemical
+differences and properties must simultaneously be decided and disposed
+in orderly relation to the pattern of the segmentation. Even if a
+model which would do this could be constructed it might prove to be a
+useful beginning.</p>
+
+<p>This may be looking too far ahead. If we had to choose some one piece
+of more proximate knowledge which we would more especially like to
+acquire, I suppose we should ask for the secret of interracial
+sterility. Nothing has yet been discovered to remove the grave
+difficulty, by which Huxley in particular was so much oppressed, that
+among the many varieties produced under domestication&mdash;which we all
+regard as analogous to the species seen in nature&mdash;no clear case of
+interracial sterility has been demonstrated. The phenomenon is
+probably the only one to which the domesticated products seem to
+afford no parallel. No solution of the difficulty can be offered which
+has positive value, but it is perhaps worth considering the facts in
+the light of modern ideas. It should be observed that we are not
+discussing incompatibility of two species to produce offspring (a
+totally distinct phenomenon), but the sterility of the offspring which
+many of them do produce.</p>
+
+<p>When two species, both perfectly fertile severally, produce on
+crossing a sterile progeny, there is a presumption that the sterility
+is due to the development in the hybrid of some substance which can
+only be formed by the meeting of two complementary factors. That some
+such account<span class='pagenum'><a name="Page_105" id="Page_105">[105]</a></span> is correct in essence may be inferred from the
+well-known observation that if the hybrid is not totally sterile but
+only partially so, and thus is able to form some good germ-cells which
+develop into new individuals, the sterility of these
+daughter-individuals is sensibly reduced or may be entirely absent.
+The fertility once re-established, the sterility does not return in
+the later progeny, a fact strongly suggestive of segregation. Now if
+the sterility of the cross-bred be really the consequence of the
+meeting of two complementary factors, we see that the phenomenon could
+only be produced among the divergent offspring of one species by the
+acquisition of at least <i>two</i> new factors; for if the acquisition of a
+single factor caused sterility the line would then end. Moreover each
+factor must be separately acquired by distinct individuals, for if
+both were present together, the possessors would by hypothesis be
+sterile. And in order to imitate the case of species each of these
+factors must be acquired by distinct breeds. The factors need not, and
+probably would not, produce any other perceptible effects; they might,
+like the colour-factors present in white flowers, make no difference
+in the form or other characters. Not till the cross was actually made
+between the two complementary individuals would either factor come
+into play, and the effects even then might be unobserved until an
+attempt was made to breed from the cross-bred.</p>
+
+<p>Next, if the factors responsible for sterility were acquired, they
+would in all probability be peculiar to certain individuals and would
+not readily be distributed to the whole breed. Any member of the breed
+also into which <i>both</i> the factors were introduced would drop out of
+the pedigree by virtue of its sterility. Hence the evidence that the
+various domesticated breeds say of dogs or fowls can when mated
+together produce fertile offspring, is beside the mark. The real
+question is, Do they ever produce sterile offspring? I think the
+evidence is clearly that sometimes they do, oftener perhaps than is
+commonly supposed. These suggestions are quite amenable to
+experimental tests. The most obvious<span class='pagenum'><a name="Page_106" id="Page_106">[106]</a></span> way to begin is to get a pair of
+parents which are known to have had any sterile offspring, and to find
+the proportions in which these steriles were produced. If, as I
+anticipate, these proportions are found to be definite, the rest is
+simple.</p>
+
+<p>In passing, certain other considerations may be referred to. First,
+that there are observations favouring the view that the production of
+totally sterile cross-breds is seldom a universal property of two
+species, and that it may be a matter of individuals, which is just
+what on the view here proposed would be expected. Moreover, as we all
+know now, though incompatibility may be dependent to some extent on
+the degree to which the species are dissimilar, no such principle can
+be demonstrated to determine sterility or fertility in general. For
+example, though all our Finches can breed together, the hybrids are
+all sterile. Of Ducks some species can breed together without
+producing the slightest sterility; others have totally sterile
+offspring, and so on. The hybrids between several <i>genera</i> of Orchids
+are perfectly fertile on the female side, and some on the male side
+also, but the hybrids produced between the Turnip (<i>Brassica napus</i>)
+and the Swede (<i>Brassica campestris</i>), which, according to our
+estimates of affinity, should be nearly allied forms, are totally
+sterile.<a name="FNanchor_73_73" id="FNanchor_73_73"></a><a href="#Footnote_73_73" class="fnanchor">[73]</a> Lastly, it may be recalled that in sterility we are
+almost certainly considering a meristic phenomenon. <i>Failure to
+divide</i> is, we may feel fairly sure, the immediate "cause" of the
+sterility. Now, though we know very little about the heredity of
+meristic differences, all that we do know points to the conclusion
+that the less-divided is dominant to the more-divided, and we are thus
+justified in supposing that there are factors which can arrest or
+prevent cell-division. My conjecture therefore is that in the case of
+sterility of cross-breds we see the effect produced by a complementary
+pair of such factors. This and many similar problems are now open to
+our analysis.</p>
+
+<p>The question is sometimes asked, Do the new lights on Variation and
+Heredity make the process of Evolution <span class='pagenum'><a name="Page_107" id="Page_107">[107]</a></span>easier to understand? On the
+whole the answer may be given that they do. There is some appearance
+of loss of simplicity, but the gain is real. As was said above, the
+time is not ripe for the discussion of the origin of species. With
+faith in Evolution unshaken&mdash;if indeed the word faith can be used in
+application to that which is certain&mdash;we look on the manner and
+causation of adapted differentiation as still wholly mysterious. As
+Samuel Butler so truly said: "To me it seems that the 'Origin of
+Variation,' whatever it is, is the only true 'Origin of Species,'"<a name="FNanchor_74_74" id="FNanchor_74_74"></a><a href="#Footnote_74_74" class="fnanchor">[74]</a>
+and of that Origin not one of us knows anything. But given
+Variation&mdash;and it is given: assuming further that the variations are
+not guided into paths of adaptation&mdash;and both to the Darwinian and to
+the modern school this hypothesis appears to be sound if unproven&mdash;an
+evolution of species proceeding by definite steps is more, rather than
+less, easy to imagine than an evolution proceeding by the accumulation
+of indefinite and insensible steps. Those who have lost themselves in
+contemplating the miracles of Adaptation (whether real or spurious)
+have not unnaturally fixed their hopes rather on the indefinite than
+on the definite changes. The reasons are obvious. By suggesting that
+the steps through which an adaptative mechanism arose were indefinite
+and insensible, all further trouble is spared. While it could be said
+that species arise by an insensible and imperceptible process of
+variation, there was clearly no use in tiring ourselves by trying to
+perceive that process. This labour-saving counsel found great favour.
+All that had to be done to develop evolution-theory was to discover
+the good in everything, a task which, in the complete absence of any
+control or test whereby to check the truth of the discovery, is not
+very onerous. The doctrine "<i>que tout est au mieux</i>" was therefore
+preached with fresh vigour, and examples of that illuminating
+principle were discovered with a facility that Pangloss himself might
+have envied, till at last even the spectators wearied of such dazzling
+performances. <span class='pagenum'><a name="Page_108" id="Page_108">[108]</a></span></p>
+
+<p>But in all seriousness, why should indefinite and unlimited variation
+have been regarded as a more probable account of the origin of
+Adaptation? Only, I think, because the obstacle was shifted one plane
+back, and so looked rather less prominent. The abundance of
+Adaptation, we all grant, is an immense, almost an unsurpassable
+difficulty in all non-Lamarckian views of Evolution; but if the steps
+by which that adaptation arose were fortuitious, to imagine them
+insensible is assuredly no help. In one most important respect indeed,
+as has often been observed, it is a multiplication of troubles. For
+the smaller the steps, the less could Natural Selection act upon them.
+Definite variations&mdash;and of the occurrence of definite variations in
+abundance we have now the most convincing proof&mdash;have at least the
+obvious merit that they can make and often do make a real difference
+in the chances of life.</p>
+
+<p>There is another aspect of the Adaptation problem to which I can
+allude very briefly. May not our present ideas of the universality and
+precision of Adaptation be greatly exaggerated? The fit of organism to
+its environment is not after all so very close&mdash;a proposition
+unwelcome perhaps, but one which could be illustrated by very copious
+evidence. Natural Selection is stern, but she has her tolerant moods.</p>
+
+<p>We have now most certain and irrefragable proof that much definiteness
+exists in living things apart from Selection, and also much that may
+very well have been preserved and so in a sense constituted by
+Selection. Here the matter is likely to rest. There is a passage in
+the sixth edition of the <i>Origin</i> which has I think been overlooked.
+On page 70 Darwin says, "The tuft of hair on the breast of the wild
+turkey-cock cannot be of any use, and it is doubtful whether it can be
+ornamental in the eyes of the female bird." This tuft of hair is a
+most definite and unusual structure, and I am afraid that the remark
+that it "cannot be of any use" may have been made inadvertently; but
+it may have been intended, for in the first edition the usual
+qualification was given and must therefore have been delib<span class='pagenum'><a name="Page_109" id="Page_109">[109]</a></span>erately
+excised. Anyhow I should like to think that Darwin did throw over that
+tuft of hair, and that he felt relief when he had done so. Whether
+however we have his great authority for such a course or not, I feel
+quite sure that we shall be rightly interpreting the facts of nature
+if we cease to expect to find purposefulness wherever we meet with
+definite structures or patterns. Such things are, as often as not, I
+suspect rather of the nature of tool-marks, mere incidents of
+manufacture, benefiting their possessor not more than the wire-marks
+in a sheet of paper, or the ribbing on the bottom of an oriental plate
+renders those objects more attractive in our eyes.</p>
+
+<p>If Variation may be in any way definite, the question once more
+arises, may it not be definite in direction? The belief that it is has
+had many supporters, from Lamarck onwards, who held that it was guided
+by need, and others who, like N&auml;geli, while laying no emphasis on
+need, yet were convinced that there was guidance of some kind. The
+latter view under the name of "Orthogenesis," devised I believe by
+Eimer, at the present day commends itself to some naturalists. The
+objection to such a suggestion is of course that no fragment of real
+evidence can be produced in its support. On the other hand, with the
+experimental proof that variation consists largely in the unpacking
+and repacking of an original complexity, it is not so certain as we
+might like to think that the order of these events is not
+predetermined.</p>
+
+<p>For instance the original "pack" may have been made in such a way that
+at the <i>n</i>th division of the germ-cells of a Sweet Pea a colour-factor
+might be dropped, and that at the <i>n</i>+<i>n</i>th division the hooded
+variety be given off, and so on. I see no ground whatever for holding
+such a view, but in fairness the possibility should not be forgotten,
+and in the light of modern research it scarcely looks so absurdly
+improbable as before.</p>
+
+<p>No one can survey the work of recent years without perceiving that
+evolutionary orthodoxy developed too fast, and that a great deal has
+got to come down; but this satisfaction<span class='pagenum'><a name="Page_110" id="Page_110">[110]</a></span> at least remains, that in the
+experimental methods which Mendel inaugurated, we have means of
+reaching certainty in regard to the physiology of Heredity and
+Variation upon which a more lasting structure may be built.</p>
+
+<div class="footnotes"><h3>FOOTNOTES:</h3>
+
+<div class="footnote"><p><a name="Footnote_56_56" id="Footnote_56_56"></a><a href="#FNanchor_56_56"><span class="label">[56]</span></a> <i>V&eacute;nus Physique, contenant deux Dissertations, l'une sur
+l'origine des Hommes et des Animaux</i>; <i>Et l'autre sur l'origine des
+Noirs</i>, La Haye, 1746, pp. 124 and 129. For an introduction to the
+writings of Maupertuis I am indebted to an article by Professor
+Lovejoy in <i>Popular Sci. Monthly</i>, 1902.</p></div>
+
+<div class="footnote"><p><a name="Footnote_57_57" id="Footnote_57_57"></a><a href="#FNanchor_57_57"><span class="label">[57]</span></a> For the fullest account of the views of these pioneers
+of Evolution, see the works of Samuel Butler, especially <i>Evolution,
+Old and New</i> (2nd edit.) 1882. Butler's claims on behalf of Buffon
+have met with some acceptance; but after reading what Butler has said,
+and a considerable part of Buffon's own works, the word "hinted" seems
+to me a sufficiently correct description of the part he played. It is
+interesting to note that in the chapter on the Ass, which contains
+some of his evolutionary passages, there is a reference to "<i>plusieurs
+id&eacute;es tr&egrave;s-&eacute;lev&eacute;es sur la g&eacute;n&eacute;ration</i>" contained in the Letters of
+Maupertuis.</p></div>
+
+<div class="footnote"><p><a name="Footnote_58_58" id="Footnote_58_58"></a><a href="#FNanchor_58_58"><span class="label">[58]</span></a> See especially W. Lawrence, <i>Lectures on Physiology</i>,
+London, 1823, pp. 213 f.</p></div>
+
+<div class="footnote"><p><a name="Footnote_59_59" id="Footnote_59_59"></a><a href="#FNanchor_59_59"><span class="label">[59]</span></a> See the chapter contributed to the <i>Life and Letters of
+Charles Darwin</i>, <span class="smcap">ii</span>. p. 195. I do not clearly understand the sense in
+which Darwin wrote (Autobiography, <i>ibid.</i> <span class="smcap">i</span>. p. 87): "It has
+sometimes been said that the success of the <i>Origin</i> proved 'that the
+subject was in the air,' or 'that men's minds were prepared for it.' I
+do not think that this is strictly true, for I occasionally sounded
+not a few naturalists, and never happened to come across a single one
+who seemed to doubt about the permanence of species." This experience
+may perhaps have been an accident due to Darwin's isolation. The
+literature of the period abounds with indications of "critical
+expectancy." A most interesting expression of that feeling is given in
+the charming account of the "Early Days of Darwinism" by Alfred
+Newton, <i>Macmillan's Magazine</i>, <span class="smcap">lvii</span>. 1888, p. 241. He tells how in
+1858 when spending a dreary summer in Iceland, he and his friend, the
+ornithologist John Wolley, in default of active occupation, spent
+their days in discussion. "Both of us taking a keen interest in
+Natural History, it was but reasonable that a question, which in those
+days was always coming up wherever two or more naturalists were
+gathered together, should be continually recurring. That question was,
+'What is a species?' and connected therewith was the other question,
+'How did a species begin?'... Now we were of course fairly well
+acquainted with what had been published on these subjects." He then
+enumerates some of these publications, mentioning among others T.
+Vernon Wollaston's <i>Variation of Species</i>&mdash;a work which has in my
+opinion never been adequately appreciated. He proceeds: "Of course we
+never arrived at anything like a solution of these problems, general
+or special, but we felt very strongly that a solution ought to be
+found, and that quickly, if the study of Botany and Zoology was to
+make any great advance." He then describes how on his return home he
+received the famous number of the <i>Linnean Journal</i> on a certain
+evening. "I sat up late that night to read it; and never shall I
+forget the impression it made upon me. Herein was contained a
+perfectly simple solution of all the difficulties which had been
+troubling me for months past.... I went to bed satisfied that a
+solution had been found."</p></div>
+
+<div class="footnote"><p><a name="Footnote_60_60" id="Footnote_60_60"></a><a href="#FNanchor_60_60"><span class="label">[60]</span></a> <i>Origin</i>, 6th edit. (1882), p. 421.</p></div>
+
+<div class="footnote"><p><a name="Footnote_61_61" id="Footnote_61_61"></a><a href="#FNanchor_61_61"><span class="label">[61]</span></a> Whatever be our estimate of the importance of Natural
+Selection, in this we all agree. Samuel Butler, the most brilliant,
+and by far the most interesting of Darwin's opponents&mdash;whose works are
+at length emerging from oblivion&mdash;in his Preface (1882) to the 2nd
+edition of <i>Evolution, Old and New</i>, repeats his earlier expression of
+homage to one whom he had come to regard as an enemy: "To the end of
+time, if the question be asked, 'Who taught people to believe in
+Evolution?' the answer must be that it was Mr. Darwin. This is true,
+and it is hard to see what palm of higher praise can be awarded to any
+philosopher."</p></div>
+
+<div class="footnote"><p><a name="Footnote_62_62" id="Footnote_62_62"></a><a href="#FNanchor_62_62"><span class="label">[62]</span></a> <i>Life and Letters</i>, <span class="smcap">i</span>. pp. 276 and 83.</p></div>
+
+<div class="footnote"><p><a name="Footnote_63_63" id="Footnote_63_63"></a><a href="#FNanchor_63_63"><span class="label">[63]</span></a> This isolation of the systematists is the one most
+melancholy sequela of Darwinism. It seems an irony that we should read
+in the peroration to the <i>Origin</i> that when the Darwinian view is
+accepted "Systematists will be able to pursue their labours as at
+present; but they will not be incessantly haunted by the shadowy doubt
+whether this or that form be a true species. This, I feel sure, and I
+speak after experience, will be no slight relief. The endless disputes
+whether or not some fifty species of British brambles are good species
+will cease." <i>Origin</i>, 6th edit. (1882), p. 425. True they have ceased
+to attract the attention of those who lead opinion, but anyone who
+will turn to the literature of systematics will find that they have
+not ceased in any other sense. Should there not be something
+disquieting in the fact that among the workers who come most into
+contact with specific differences, are to be found the only men who
+have failed to be persuaded of the unreality of those differences?</p></div>
+
+<div class="footnote"><p><a name="Footnote_64_64" id="Footnote_64_64"></a><a href="#FNanchor_64_64"><span class="label">[64]</span></a> 6th edit. pp. 109 and 401. See Butler, <i>Essays on Life,
+Art, and Science</i>, p. 265, reprinted 1908, and <i>Evolution, Old and
+New</i>, chap. <span class="smcap">xxii</span>. (2nd edit.), 1882.</p></div>
+
+<div class="footnote"><p><a name="Footnote_65_65" id="Footnote_65_65"></a><a href="#FNanchor_65_65"><span class="label">[65]</span></a> W. Lawrence was one of the few who consistently
+maintained the contrary opinion. Prichard, who previously had
+expressed himself in the same sense, does not, I believe, repeat these
+views in his later writings, and there are signs that he came to
+believe in the transmission of acquired habits. See Lawrence, <i>Lect.
+Physiol.</i> 1823, pp. 436-437, 447. Prichard, Edin. Inaug. Disp. 1808
+[not seen by me], quoted <i>ibid.</i> and <i>Nat. Hist. Man</i>, 1843, pp. 34
+f.</p></div>
+
+<div class="footnote"><p><a name="Footnote_66_66" id="Footnote_66_66"></a><a href="#FNanchor_66_66"><span class="label">[66]</span></a> It is interesting to see how nearly Butler was led by
+natural penetration, and from absolutely opposite conclusions, back to
+this underlying truth: "So that each ovum when impregnate should be
+considered not as descended from its ancestors, but as being a
+continuation of the personality of every ovum in the chain of its
+ancestry, which every ovum <i>it actually is</i> quite as truly as the
+octogenarian <i>is</i> the same identity with the ovum from which he has
+been developed. This process cannot stop short of the primordial cell,
+which again will probably turn out to be but a brief resting-place. We
+therefore prove each one of us to <i>be actually</i> the primordial cell
+which never died nor dies, but has differentiated itself into the life
+of the world, all living beings whatever, being one with it and
+members one of another," <i>Life and Habit</i>, 1878, p. 86.</p></div>
+
+<div class="footnote"><p><a name="Footnote_67_67" id="Footnote_67_67"></a><a href="#FNanchor_67_67"><span class="label">[67]</span></a> This view is no doubt contrary to the received opinion.
+I am however interested to see it lately maintained by Driesch
+(<i>Science and Philosophy of the Organism</i>, London, 1907, p. 233), and
+from the recent observations of Godlewski it has received distinct
+experimental support.</p></div>
+
+<div class="footnote"><p><a name="Footnote_68_68" id="Footnote_68_68"></a><a href="#FNanchor_68_68"><span class="label">[68]</span></a> In other words, the ova are each <i>either</i> female, <i>or
+male</i> (i.e. non-female), but the sperms are all non-female.</p></div>
+
+<div class="footnote"><p><a name="Footnote_69_69" id="Footnote_69_69"></a><a href="#FNanchor_69_69"><span class="label">[69]</span></a> Morgan, <i>Proc. Soc. Exp. Biol. Med.</i> <span class="smcap">v</span>. 1908, and von
+Baehr, <i>Zool. Anz.</i> <span class="smcap">xxxii</span>. p. 507, 1908.</p></div>
+
+<div class="footnote"><p><a name="Footnote_70_70" id="Footnote_70_70"></a><a href="#FNanchor_70_70"><span class="label">[70]</span></a> As Wilson has proved, the unpaired body is not a
+universal feature even in those orders in which it has been observed.
+Nearly allied types may differ. In some it is altogether unpaired. In
+others it is paired with a body of much smaller size, and by selection
+of various types all gradations can be demonstrated ranging to the
+condition in which the members of the pair are indistinguishable from
+each other.</p></div>
+
+<div class="footnote"><p><a name="Footnote_71_71" id="Footnote_71_71"></a><a href="#FNanchor_71_71"><span class="label">[71]</span></a> I have in view, for example, the marvellous and specific
+phenomena of regeneration, and those discovered by the students of
+"<i>Entwicklungsmechanik</i>." The circumstances of its occurrence here
+preclude any suggestion that this regularity has been brought about by
+the workings of Selection. The attempts thus to represent the
+phenomena have resulted in mere parodies of scientific reasoning.</p></div>
+
+<div class="footnote"><p><a name="Footnote_72_72" id="Footnote_72_72"></a><a href="#FNanchor_72_72"><span class="label">[72]</span></a> <i>Vortr&auml;ge &uuml;ber Viehzucht und Rassenerkenntniss</i>, p. 120,
+Berlin, 1872.</p></div>
+
+<div class="footnote"><p><a name="Footnote_73_73" id="Footnote_73_73"></a><a href="#FNanchor_73_73"><span class="label">[73]</span></a> See Sutton, A. W., <i>Journ. Linn. Soc.</i> <span class="smcap">xxxviii</span>. p. 341,
+1908.</p></div>
+
+<div class="footnote"><p><a name="Footnote_74_74" id="Footnote_74_74"></a><a href="#FNanchor_74_74"><span class="label">[74]</span></a> <i>Life and Habit</i>, London, p. 263, 1878</p></div>
+</div>
+
+
+<hr style="width: 65%;" />
+<p><span class='pagenum'><a name="Page_111" id="Page_111">[111]</a></span></p>
+<h2><a name="IV" id="IV"></a>IV</h2>
+
+<h2>"THE DESCENT OF MAN"</h2>
+<p class="center"><span class="smcap1">By G. Schwalbe</span></p>
+<h4><i>Professor of Anatomy in the University of Strassburg</i></h4>
+<p>The problem of the origin of the human race, of the descent of man, is
+ranked by Huxley in his epoch-making book <i>Man's Place in Nature</i>, as
+the deepest with which biology has to concern itself, "the question of
+questions,"&mdash;the problem which underlies all others. In the same
+brilliant and lucid exposition, which appeared in 1863, soon after the
+publication of Darwin's <i>Origin of Species</i>, Huxley stated his own
+views in regard to this great problem. He tells us how the idea of a
+natural descent of man gradually grew up in his mind. It was
+especially the assertions of Owen in regard to the total difference
+between the human and the simian brain that called forth strong
+dissent from the great anatomist Huxley, and he easily succeeded in
+showing that Owen's supposed differences had no real existence; he
+even established, on the basis of his own anatomical investigations,
+the proposition that the anatomical differences between the Marmoset
+and the Chimpanzee are much greater than those between the Chimpanzee
+and Man.</p>
+
+<p>But why do we thus introduce the study of Darwin's <i>Descent of Man</i>,
+which is to occupy us here, by insisting on the fact that Huxley had
+taken the field in defence of the descent of man in 1863, while
+Darwin's book on the subject did not appear till 1871? It is in order
+that we may<span class='pagenum'><a name="Page_112" id="Page_112">[112]</a></span> clearly understand how it happened that from this time
+onwards Darwin and Huxley followed the same great aim in the most
+intimate association.</p>
+
+<p>Huxley and Darwin working at the same <i>Problema maximum</i>! Huxley
+fiery, impetuous, eager for battle, contemptuous of the resistance of
+a dull world, or energetically triumphing over it. Darwin calm,
+weighing every problem slowly, letting it mature thoroughly,&mdash;not a
+fighter, yet having the greater and more lasting influence by virtue
+of his immense mass of critically sifted proofs. Darwin's friend,
+Huxley, was the first to do him justice, to understand his nature, and
+to find in it the reason why the detailed and carefully considered
+book on the descent of man made its appearance so late. Huxley, always
+generous, never thought of claiming priority for himself. In
+enthusiastic language he tells how Darwin's immortal work, <i>The Origin
+of Species</i>, first shed light for him on the problem of the descent of
+man; the recognition of a <i>vera causa</i> in the transformation of
+species illuminated his thoughts as with a flash. He was now content
+to leave what perplexed him, what he could not yet solve, as he says
+himself, "in the mighty hands of Darwin." Happy in the bustle of
+strife against old and deep-rooted prejudices, against intolerance and
+superstition, he wielded his sharp weapons on Darwin's behalf; wearing
+Darwin's armour he joyously overthrew adversary after adversary.
+Darwin spoke of Huxley as his "general agent."<a name="FNanchor_75_75" id="FNanchor_75_75"></a><a href="#Footnote_75_75" class="fnanchor">[75]</a> Huxley says of
+himself "I am Darwin's bulldog."<a name="FNanchor_76_76" id="FNanchor_76_76"></a><a href="#Footnote_76_76" class="fnanchor">[76]</a></p>
+
+<p>Thus Huxley openly acknowledged that it was Darwin's <i>Origin of
+Species</i> that first set the problem of the descent of man in its true
+light, that made the question of the origin of the human race a
+pressing one. That this was the logical consequence of his book Darwin
+himself had long felt. He had been reproached with intentionally
+shirking <span class='pagenum'><a name="Page_113" id="Page_113">[113]</a></span>the application of his theory to Man. Let us hear what he
+says on this point in his autobiography: "As soon as I had become, in
+the year 1837 or 1838, convinced that species were mutable
+productions, I could not avoid the belief that man must come under the
+same law. Accordingly I collected notes on the subject for my own
+satisfaction, and not for a long time with any intention of
+publishing. Although in the 'Origin of Species' the derivation of any
+particular species is never discussed, yet I thought it best, in order
+<i>that no honourable man should accuse me of concealing my views</i>,<a name="FNanchor_77_77" id="FNanchor_77_77"></a><a href="#Footnote_77_77" class="fnanchor">[77]</a>
+to add that by the work 'light would be thrown on the origin of man
+and his history.' It would have been useless and injurious to the
+success of the book to have paraded, without giving any evidence, my
+conviction with respect to his origin."<a name="FNanchor_78_78" id="FNanchor_78_78"></a><a href="#Footnote_78_78" class="fnanchor">[78]</a></p>
+
+<p>In a letter written in January, 1860, to the Rev. L. Blomefield,
+Darwin expresses himself in similar terms. "With respect to man, I am
+very far from wishing to obtrude my belief; but I thought it dishonest
+to quite conceal my opinion."<a name="FNanchor_79_79" id="FNanchor_79_79"></a><a href="#Footnote_79_79" class="fnanchor">[79]</a></p>
+
+<p>The brief allusion in the <i>Origin of Species</i> is so far from prominent
+and so incidental that it was excusable to assume that Darwin had not
+touched upon the descent of man in this work. It was solely the desire
+to have his mass of evidence sufficiently complete, solely Darwin's
+great characteristic of never publishing till he had carefully weighed
+all aspects of his subject for years, solely, in short, his most
+fastidious scientific conscience that restrained him from challenging
+the world in 1859 with a book in which the theory of the descent of
+man was fully set forth. Three years, frequently interrupted by
+ill-health, were needed for the actual writing of the book:<a name="FNanchor_80_80" id="FNanchor_80_80"></a><a href="#Footnote_80_80" class="fnanchor">[80]</a> the
+first edition, which appeared <span class='pagenum'><a name="Page_114" id="Page_114">[114]</a></span>in 1871, was followed in 1874 by a much
+improved second edition, the preparation of which he very reluctantly
+undertook.<a name="FNanchor_81_81" id="FNanchor_81_81"></a><a href="#Footnote_81_81" class="fnanchor">[81]</a></p>
+
+<p>This, briefly, is the history of the work, which, with the <i>Origin of
+Species</i>, marks an epoch in the history of biological sciences&mdash;the
+work with which the cautious, peace-loving investigator ventured forth
+from his contemplative life into the arena of strife and unrest, and
+laid himself open to all the annoyances that deep-rooted belief and
+prejudice, and the prevailing tendency of scientific thought at the
+time could devise.</p>
+
+<p>Darwin did not take this step lightly. Of great interest in this
+connection is a letter written to Wallace on Dec. 22, 1857,<a name="FNanchor_82_82" id="FNanchor_82_82"></a><a href="#Footnote_82_82" class="fnanchor">[82]</a> in
+which he says, "You ask me whether I shall discuss 'man.' I think I
+shall avoid the whole subject, as so surrounded with prejudices;
+though I fully admit that it is the highest and most interesting
+problem for the naturalist." But his conscientiousness compelled him
+to state briefly his opinion on the subject in the <i>Origin of Species</i>
+in 1859. Nevertheless he did not escape reproaches for having been so
+reticent. This is unmistakably apparent from a letter to Fritz M&uuml;ller
+dated Feb. 22 [1869?], in which he says: "I am thinking of writing a
+little essay on the Origin of Mankind, as I have been taunted with
+concealing my opinions."<a name="FNanchor_83_83" id="FNanchor_83_83"></a><a href="#Footnote_83_83" class="fnanchor">[83]</a></p>
+
+<p>It might be thought that Darwin behaved thus hesitatingly, and was so
+slow in deciding on the full publication of his collected material in
+regard to the descent of man, because he had religious difficulties to
+overcome.</p>
+
+<p>But this was not the case, as we can see from his admirable confession
+of faith, the publication of which we owe to his son Francis.<a name="FNanchor_84_84" id="FNanchor_84_84"></a><a href="#Footnote_84_84" class="fnanchor">[84]</a>
+Whoever wishes really to understand the lofty character of this great
+man should read these <span class='pagenum'><a name="Page_115" id="Page_115">[115]</a></span>immortal lines in which he unfolds to us in
+simple and straightforward words the development of his conception of
+the universe. He describes how, though he was still quite orthodox
+during his voyage round the world on board the <i>Beagle</i>, he came
+gradually to see, shortly afterwards (1836-1839) that the Old
+Testament was no more to be trusted than the Sacred Books of the
+Hindoos; the miracles by which Christianity is supported, the
+discrepancies between the accounts in the different Gospels, gradually
+led him to disbelieve in Christianity as a divine revelation. "Thus,"
+he writes,<a name="FNanchor_85_85" id="FNanchor_85_85"></a><a href="#Footnote_85_85" class="fnanchor">[85]</a> "disbelief crept over me at a very slow rate, but was
+at last complete. The rate was so slow that I felt no distress." But
+Darwin was too modest to presume to go beyond the limits laid down by
+science. He wanted nothing more than to be able to go, freely and
+unhampered by belief in authority or in the Bible, as far as human
+knowledge could lead him. We learn this from the concluding words of
+his chapter on religion "The mystery of the beginning of all things is
+insoluble by us; and I for one must be content to remain an
+Agnostic."<a name="FNanchor_86_86" id="FNanchor_86_86"></a><a href="#Footnote_86_86" class="fnanchor">[86]</a></p>
+
+<p>Darwin was always very unwilling to give publicity to his views in
+regard to religion. In a letter to Asa Gray on May 22, 1860,<a name="FNanchor_87_87" id="FNanchor_87_87"></a><a href="#Footnote_87_87" class="fnanchor">[87]</a> he
+declares that it is always painful to him to have to enter into
+discussion of religious problems. He had, he said, no intention of
+writing atheistically.</p>
+
+<p>Finally, let us cite one characteristic sentence from a letter from
+Darwin to C. Ridley<a name="FNanchor_88_88" id="FNanchor_88_88"></a><a href="#Footnote_88_88" class="fnanchor">[88]</a> (Nov. 28, 1878). A clergyman, Dr. Pusey, had
+asserted that Darwin had written the <i>Origin of Species</i> with some
+relation to theology. Darwin writes emphatically, "Many years ago when
+I was collecting facts for the 'Origin,' my belief in what is called a
+personal God <span class='pagenum'><a name="Page_116" id="Page_116">[116]</a></span>was as firm as that of Dr. Pusey himself, and as to the
+eternity of matter I never troubled myself about such insoluble
+questions." The expression "many years ago" refers to the time of his
+voyage round the world, as has already been pointed out. Darwin means
+by this utterance that the views which had gradually developed in his
+mind in regard to the origin of species were quite compatible with the
+faith of the Church.</p>
+
+<p>If we consider all these utterances of Darwin in regard to religion
+and to his outlook on life (Weltanschauung), we shall see at least so
+much, that religious reflection could in no way have influenced him in
+regard to the writing and publishing of his book on <i>The Descent of
+Man</i>. Darwin had early won for himself freedom of thought, and to this
+freedom he remained true to the end of his life, uninfluenced by the
+customs and opinions of the world around him.</p>
+
+<p>Darwin was thus inwardly fortified and armed against the host of
+calumnies, accusations, and attacks called forth by the publication of
+the <i>Origin of Species</i>, and to an even greater extent by the
+appearance of the <i>Descent of Man</i>. But in his defence he could rely
+on the aid of a band of distinguished auxiliaries of the rarest
+ability. His faithful confederate, Huxley, was joined by the botanist
+Hooker, and, after longer resistance, by the famous geologist Lyell,
+whose "conversion" afforded Darwin peculiar satisfaction. All three
+took the field with enthusiasm in defence of the natural descent of
+man. From Wallace, on the other hand, though he shared with him the
+idea of natural selection, Darwin got no support in this matter.
+Wallace expressed himself in a strange manner. He admitted everything
+in regard to the morphological descent of man, but maintained, in a
+mystic way, that something else, something of a spiritual nature must
+have been added to what man inherited from his animal ancestors.
+Darwin, whose esteem for Wallace was extraordinarily high, could not
+understand how he could give utterance to such a mystical view in
+regard to man; the idea seemed to him so "incredibly strange"<span class='pagenum'><a name="Page_117" id="Page_117">[117]</a></span> that he
+thought some one else must have added these sentences to Wallace's
+paper.</p>
+
+<p>Even now there are thinkers who, like Wallace, shrink from applying to
+man the ultimate consequences of the theory of descent. The idea that
+man is derived from ape-like forms is to them unpleasant and
+humiliating.</p>
+
+<p>So far I have been depicting the development of Darwin's work on the
+descent of man. In what follows I shall endeavour to give a condensed
+survey of the contents of the book.</p>
+
+<p>It must at once be said that the contents of Darwin's work fall into
+two parts, dealing with entirely different subjects. <i>The Descent of
+Man</i> includes a very detailed investigation in regard to secondary
+sexual characters in the animal series, and on this investigation
+Darwin founded a new theory, that of sexual selection. With
+astonishing patience he gathered together an immense mass of material,
+and showed, in regard to Arthropods and Vertebrates, the wide
+distribution of secondary characters, which develop almost exclusively
+in the male, and which enable him, on the one hand, to get the better
+of his rivals in the struggle for the female by the greater perfection
+of his weapons, and, on the other hand, to offer greater allurements
+to the female through the higher development of decorative characters,
+of song, or of scent-producing glands. The best equipped males will
+thus crowd out the less well-equipped in the matter of reproduction,
+and thus the relevant characters will be increased and perfected
+through sexual selection. It is, of course, a necessary assumption
+that these secondary sexual characters may be transmitted to the
+female, although perhaps in rudimentary form.</p>
+
+<p>As we have said, this story of sexual selection takes up a great deal
+of space in Darwin's book, and it need only be considered here in so
+far as Darwin applied it to the descent of man. To this latter problem
+the whole of Part I is devoted, while Part III contains a discussion
+of sexual selection in relation to man, and a general summary. Part<span class='pagenum'><a name="Page_118" id="Page_118">[118]</a></span>
+II treats of sexual selection in general, and may be disregarded in
+our present study. Moreover, many interesting details must necessarily
+be passed over in what follows, for want of space.</p>
+
+<p>The first part of the <i>Descent of Man</i> begins with an enumeration of
+the proofs of the animal descent of man taken from the structure of
+the human body. Darwin chiefly emphasises the fact that the human body
+consists of the same organs and of the same tissues as those of the
+other mammals; he shows also that man is subject to the same diseases
+and tormented by the same parasites as the apes. He further dwells on
+the general agreement exhibited by young embryonic forms, and he
+illustrates this by two figures placed one above the other, one
+representing a human embryo, after Ecker, the other a dog embryo,
+after Bischoff.<a name="FNanchor_89_89" id="FNanchor_89_89"></a><a href="#Footnote_89_89" class="fnanchor">[89]</a></p>
+
+<p>Darwin finds further proofs of the animal origin of man in the reduced
+structures, in themselves extremely variable, which are either
+absolutely useless to their possessors, or of so little use that they
+could never have developed under existing conditions. Of such vestiges
+he enumerates: the defective development of the <i>panniculus carnosus</i>
+(muscle of the skin) so widely distributed among mammals, the
+ear-muscles, the occasional persistence of the animal ear-point in
+man, the rudimentary nictitating membrane (<i>plica semilunaris</i>) in the
+human eye, the slight development of the organ of smell, the general
+hairiness of the human body, the frequently defective development or
+entire absence of the third molar (the wisdom tooth), the vermiform
+appendix, the occasional reappearance of a bony canal (<i>foramen
+supracondyloideum</i>) at the lower end of the humerus, the rudimentary
+tail of man (the so-called taillessness), and so on. Of these
+rudimentary structures the occasional occurrence of the animal
+ear-point in man is most fully discussed. Darwin's attention was
+called to this interesting structure by <span class='pagenum'><a name="Page_119" id="Page_119">[119]</a></span>the sculptor Woolner. He
+figures such a case observed in man, and also the head of an alleged
+orang-foetus, the photograph of which he received from Nitsche.</p>
+
+<p>Darwin's interpretation of Woolner's case as having arisen through a
+folding over of the free edge of a pointed ear has been fully borne
+out by my investigations on the external ear.<a name="FNanchor_90_90" id="FNanchor_90_90"></a><a href="#Footnote_90_90" class="fnanchor">[90]</a> In particular, it
+was established by these investigations that the human foetus, about
+the middle of its embryonic life, possesses a pointed ear somewhat
+similar to that of the monkey genus Macacus. One of Darwin's
+statements in regard to the head of the orang-foetus must be
+corrected. A <i>large</i> ear with a point is shown in the photograph,<a name="FNanchor_91_91" id="FNanchor_91_91"></a><a href="#Footnote_91_91" class="fnanchor">[91]</a>
+but it can easily be demonstrated&mdash;and Deniker has already pointed
+this out&mdash;that the figure is not that of an orang foetus at all, for
+that form has much smaller ears with no point; nor can it be a
+gibbon-foetus, as Deniker supposes, for the gibbon ear is also without
+a point. I myself regard it as that of a Macacus-embryo. But this
+mistake, which is due to Nitsche, in no way affects the fact
+recognised by Darwin, that ear-forms showing the point characteristic
+of the animal ear occur in man with extraordinary frequency.</p>
+
+<p>Finally, there is a discussion of those rudimentary structures which
+occur only in <i>one</i> sex, such as the rudimentary mammary glands in the
+male, the vesicula prostatica, which corresponds to the uterus of the
+female, and others. All these facts tell in favour of the common
+descent of man and all other vertebrates. The conclusion of this
+section is characteristic: "<i>It is only our natural prejudice, and
+that arrogance which made our forefathers declare that they were
+descended from demi-gods, which leads us to demur to this conclusion.
+But the time will before long come, when it will be thought wonderful
+that naturalists, who were well acquainted with the comparative
+structure and development<span class='pagenum'><a name="Page_120" id="Page_120">[120]</a></span> of man, and other mammals, should have
+believed that each was the work of a separate act of creation.</i>"<a name="FNanchor_92_92" id="FNanchor_92_92"></a><a href="#Footnote_92_92" class="fnanchor">[92]</a></p>
+
+<p>In the second chapter there is a more detailed discussion, again based
+upon an extraordinary wealth of facts, of the problem as to the manner
+in which, and the causes through which, man evolved from a lower form.
+Precisely the same causes are here suggested for the origin of man, as
+for the origin of species in general. Variability, which is a
+necessary assumption in regard to all transformations, occurs in man
+to a high degree. Moreover, the rapid multiplication of the human race
+creates conditions which necessitate an energetic struggle for
+existence, and thus afford scope for the intervention of natural
+selection. Of the exercise of <i>artificial</i> selection in the human
+race, there is nothing to be said, unless we cite such cases as the
+grenadiers of Frederick William I, or the population of ancient
+Sparta. In the passages already referred to and in those which follow,
+the transmission of acquired characters, upon which Darwin does not
+dwell, is taken for granted. In man, direct effects of changed
+conditions can be demonstrated (for instance in regard to bodily
+size), and there are also proofs of the influence exerted on his
+physical constitution by increased use or disuse. Reference is here
+made to the fact, established by Forbes, that the Quechua Indians of
+the high plateaus of Peru show a striking development of lungs and
+thorax, as a result of living constantly at high altitudes.</p>
+
+<p>Such special forms of variation as arrests of development
+(microcephalism) and reversion to lower forms are next discussed.
+Darwin himself felt<a name="FNanchor_93_93" id="FNanchor_93_93"></a><a href="#Footnote_93_93" class="fnanchor">[93]</a> that these subjects are so nearly related to
+the cases mentioned in the first chapter, that many of them might as
+well have been dealt with there. It seems to me that it would have
+been better so, for the citation of additional instances of reversion
+at this place rather disturbs the logical sequence of his ideas as to
+the <span class='pagenum'><a name="Page_121" id="Page_121">[121]</a></span>conditions which have brought about the evolution of man from
+lower forms. The instances of reversion here discussed are
+microcephalism, which Darwin wrongly interpreted as atavistic,
+supernumerary mammae, supernumerary digits, bicornuate uterus, the
+development of abnormal muscles, and so on. Brief mention is also made
+of correlative variations observed in man.</p>
+
+<p>Darwin next discusses the question as to the manner in which man
+attained to the erect position from the state of a climbing quadruped.
+Here again he puts the influence of Natural Selection in the first
+rank. The immediate progenitors of man had to maintain a struggle for
+existence in which success was to the more intelligent, and to those
+with social instincts. The hand of these climbing ancestors, which had
+little skill and served mainly for locomotion, could only undergo
+further development when some early member of the Primate series came
+to live more on the ground and less among trees.</p>
+
+<p>A bipedal existence thus became possible, and with it the liberation
+of the hand from locomotion, and the one-sided development of the
+human foot. The upright position brought about correlated variations
+in the bodily structure; with the free use of the hand it became
+possible to manufacture weapons and to use them; and this again
+resulted in a degeneration of the powerful canine teeth and the jaws,
+which were then no longer necessary for defence. Above all, however,
+the intelligence immediately increased, and with it skull and brain.
+The nakedness of man, and the absence of a tail (rudimentariness of
+the tail vertebrae) are next discussed. Darwin is inclined to
+attribute the nakedness of man, not to the action of natural selection
+on ancestors who originally inhabited a tropical land, but to sexual
+selection, which, for aesthetic reasons, brought about the loss of the
+hairy covering in man, or primarily in woman. An interesting
+discussion of the loss of the tail, which, however, man shares with
+the anthropoid apes, some other monkeys and lemurs, forms the
+conclusion of the al<span class='pagenum'><a name="Page_122" id="Page_122">[122]</a></span>most superabundant material which Darwin worked
+up in the second chapter. His object was to show that some of the most
+distinctive human characters are in all probability directly or
+indirectly due to natural selection. With characteristic modesty he
+adds:<a name="FNanchor_94_94" id="FNanchor_94_94"></a><a href="#Footnote_94_94" class="fnanchor">[94]</a> "Hence, if I have erred in giving to natural selection great
+power, which I am very far from admitting, or in having exaggerated
+its power, which is in itself probable, I have at least, as I hope,
+done good service in aiding to overthrow the dogma of separate
+creations." At the end of the chapter he touches upon the objection as
+to man's helpless and defenceless condition. Against this he urges his
+intelligence and social instincts.</p>
+
+<p>The two following chapters contain a detailed discussion of the
+objections drawn from the supposed great differences between the
+mental powers of men and animals. Darwin at once admits that the
+differences are enormous, but not that any fundamental difference
+between the two can be found. Very characteristic of him is the
+following passage: "In what manner the mental powers were first
+developed in the lowest organisms, is as hopeless an enquiry as how
+life itself first originated. These are problems for the distant
+future, if they are ever to be solved by man."<a name="FNanchor_95_95" id="FNanchor_95_95"></a><a href="#Footnote_95_95" class="fnanchor">[95]</a></p>
+
+<p>After some brief observations on instinct and intelligence, Darwin
+brings forward evidence to show that the greater number of the
+emotional states, such as pleasure and pain, happiness and misery,
+love and hate are common to man and the higher animals. He goes on to
+give various examples showing that wonder and curiosity, imitation,
+attention, memory and imagination (dreams of animals), can also be
+observed in the higher mammals, especially in apes. In regard even to
+reason there are no sharply defined limits. A certain faculty of
+deliberation is characteristic of some animals, and the more
+thoroughly we know an animal the more intelligence we are inclined to
+credit it with. Examples <span class='pagenum'><a name="Page_123" id="Page_123">[123]</a></span>are brought forward of the intelligent and
+deliberate actions of apes, dogs and elephants. But although no
+sharply defined differences exist between man and animals, there is,
+nevertheless, a series of other mental powers which are
+characteristics usually regarded as absolutely peculiar to man. Some
+of these characteristics are examined in detail, and it is shown that
+the arguments drawn from them are not conclusive. Man alone is said to
+be capable of progressive improvement; but against this must be placed
+as something analogous in animals, the fact that they learn cunning
+and caution through long continued persecution. Even the use of tools
+is not in itself peculiar to man (monkeys use sticks, stones and
+twigs), but man alone fashions and uses implements <i>designed for a
+special purpose</i>. In this connection the remarks taken from Lubbock in
+regard to the origin and gradual development of the earliest flint
+implements will be read with interest; these are similar to the
+observations on modern eoliths, and their bearing on the development
+of the stone industry. It is interesting to learn from a letter to
+Hooker,<a name="FNanchor_96_96" id="FNanchor_96_96"></a><a href="#Footnote_96_96" class="fnanchor">[96]</a> that Darwin himself at first doubted whether the stone
+implements discovered by Boucher de Perthes were really of the nature
+of tools. With the relentless candour as to himself which
+characterised him, he writes four years later in a letter to Lyell in
+regard to this view of Boucher de Perthes' discoveries: "I know
+something about his errors, and looked at his book many years ago, and
+am ashamed to think that I concluded the whole was rubbish! Yet he has
+done for man something like what Agassiz did for glaciers."<a name="FNanchor_97_97" id="FNanchor_97_97"></a><a href="#Footnote_97_97" class="fnanchor">[97]</a></p>
+
+<p>To return to Darwin's further comparisons between the higher mental
+powers of man and animals; He takes much of the force from the
+argument that man alone is capable of abstraction and
+self-consciousness by his own observations on dogs. One of the main
+differences between man and <span class='pagenum'><a name="Page_124" id="Page_124">[124]</a></span>animals, speech, receives detailed
+treatment. He points out that various animals (birds, monkeys, dogs)
+have a large number of different sounds for different emotions, that,
+further, man produces in common with animals a whole series of
+inarticulate cries combined with gestures, and that dogs learn to
+understand whole sentences of human speech. In regard to human
+language, Darwin expresses a view contrary to that held by Max
+M&uuml;ller:<a name="FNanchor_98_98" id="FNanchor_98_98"></a><a href="#Footnote_98_98" class="fnanchor">[98]</a> "I cannot doubt that language owes its origin to the
+imitation and modification of various natural sounds, the voices of
+other animals, and man's own instinctive cries, aided by signs and
+gestures." The development of actual language presupposes a higher
+degree of intelligence than is found in any kind of ape. Darwin
+remarks on this point:<a name="FNanchor_99_99" id="FNanchor_99_99"></a><a href="#Footnote_99_99" class="fnanchor">[99]</a> "The fact of the higher apes not using
+their vocal organs for speech no doubt depends on their intelligence
+not having been sufficiently advanced."</p>
+
+<p>The sense of beauty, too, has been alleged to be peculiar to man. In
+refutation of this assertion Darwin points to the decorative colours
+of birds, which are used for display. And to the last objection, that
+man alone has religion, that he alone has a belief in God, it is
+answered "that numerous races have existed, and still exist, who have
+no idea of one or more gods, and who have no words in their languages
+to express such an idea."<a name="FNanchor_100_100" id="FNanchor_100_100"></a><a href="#Footnote_100_100" class="fnanchor">[100]</a></p>
+
+<p>The result of the investigations recorded in this chapter is to show
+that, great as the difference in mental powers between man and the
+higher animals may be, it is undoubtedly only a difference "of degree
+and not of kind."<a name="FNanchor_101_101" id="FNanchor_101_101"></a><a href="#Footnote_101_101" class="fnanchor">[101]</a></p>
+
+<p>In the fourth chapter Darwin deals with the <i>moral sense</i> or
+<i>conscience</i>, which is the most important of all differences between
+man and animals. It is a result of social instincts, <span class='pagenum'><a name="Page_125" id="Page_125">[125]</a></span>which lead to
+sympathy for other members of the same society, to non-egoistic
+actions for the good of others. Darwin shows that social tendencies
+are found among many animals, and that among these love and
+kin-sympathy exist, and he gives examples of animals (especially dogs)
+which may exhibit characters that we should call moral in man (e.g.
+disinterested self-sacrifice for the sake of others). The early
+ape-like progenitors of the human race were undoubtedly social. With
+the increase of intelligence the moral sense develops farther; with
+the acquisition of speech public opinion arises, and finally, moral
+sense becomes habit. The rest of Darwin's detailed discussions on
+moral philosophy may be passed over.</p>
+
+<p>The fifth chapter may be very briefly summarised. In it Darwin shows
+that the intellectual and moral faculties are perfected through
+natural selection. He inquires how it can come about that a tribe at a
+low level of evolution attains to a higher, although the best and
+bravest among them often pay for their fidelity and courage with their
+lives without leaving any descendants. In this case it is the
+sentiment of glory, praise and blame, the admiration of others, which
+bring about the increase of the better members of the tribe. Property,
+fixed dwellings, and the association of families into a community are
+also indispensable requirements for civilisation. In the longer second
+section of the fifth chapter Darwin acts mainly as recorder. On the
+basis of numerous investigations, especially those of Greg, Wallace,
+and Galton, he inquires how far the influence of natural selection can
+be demonstrated in regard to civilised nations. In the final section,
+which deals with the proofs that all civilised nations were once
+barbarians, Darwin again uses the results gained by other
+investigators, such as Lubbock and Tylor. There are two sets of facts
+which prove the proposition in question. In the first place, we find
+traces of a former lower state in the customs and beliefs of all
+civilised nations, and in the second place, there are proofs to show
+that savage races are independently<span class='pagenum'><a name="Page_126" id="Page_126">[126]</a></span> able to raise themselves a few
+steps in the scale of civilisation, and that they have thus raised
+themselves.</p>
+
+<p>In the sixth chapter of the work, Morphology comes into the foreground
+once more. Darwin first goes back, however, to the argument based on
+the great difference between the mental powers of the highest animals
+and those of man. That this is only quantitative, not qualitative, he
+has already shown. Very instructive in this connection is the
+reference to the enormous difference in mental powers in another
+class. No one would draw from the fact that the cochineal insect
+(Coccus) and the ant exhibit enormous differences in their mental
+powers, the conclusion that the ant should therefore be regarded as
+something quite distinct, and withdrawn from the class of insects
+altogether.</p>
+
+<p>Darwin next attempts to establish the <i>specific</i> genealogical tree of
+man, and carefully weighs the differences and resemblances between the
+different families of the Primates. The erect position of man is an
+adaptive character, just as are the various characters referable to
+aquatic life in the seals, which, notwithstanding these, are ranked as
+a mere family of the carnivores. The following utterance is very
+characteristic of Darwin:<a name="FNanchor_102_102" id="FNanchor_102_102"></a><a href="#Footnote_102_102" class="fnanchor">[102]</a> "If man had not been his own
+classifier, he would never have thought of founding a separate order
+for his own reception." In numerous characters not mentioned in
+systematic works, in the features of the face, in the form of the
+nose, in the structure of the external ear, man resembles the apes.
+The arrangement of the hair in man has also much in common with the
+apes; as also the occurrence of hair on the forehead of the human
+embryo, the beard, the convergence of the hair of the upper and under
+arm towards the elbow, which occurs not only in the anthropoid apes,
+but also in some American monkeys. Darwin here adopts Wallace's
+explanation of the origin of the ascending direction of the hair in
+the forearm of the orang,&mdash;that it has arisen through the habit of
+holding the hands <span class='pagenum'><a name="Page_127" id="Page_127">[127]</a></span>over the head in rain. But this explanation cannot
+be maintained when we consider that this disposition of the hair is
+widely distributed among the most different mammals, being found in
+the dog, in the sloth, and in many of the lower monkeys.</p>
+
+<p>After further careful analysis of the anatomical characters Darwin
+reaches the conclusion that the New World monkeys (Platyrrhine) may be
+excluded from the genealogical tree altogether, but that man is an
+offshoot from the Old World monkeys (Catarrhine) whose progenitors
+existed as far back as the Miocene period. Among these Old World
+monkeys the forms to which man shows the greatest resemblance are the
+anthropoid apes, which, like him, possess neither tail nor ischial
+callosities. The platyrrhine and catarrhine monkeys have their
+primitive ancestor among extinct forms of the Lemuridae. Darwin also
+touches on the question of the original home of the human race and
+supposes that it may have been in Africa, because it is there that
+man's nearest relatives, the gorilla and the chimpanzee, are found.
+But he regards speculation on this point as useless. It is remarkable
+that, in this connection, Darwin regards the loss of the hair-covering
+in man as having some relation to a warm climate, while elsewhere he
+is inclined to make sexual selection responsible for it. Darwin
+recognises the great gap between man and his nearest relatives, but
+similar gaps exist at other parts of the mammalian genealogical tree:
+the allied forms have become extinct. After the extermination of the
+lower races of mankind, on the one hand, and of the anthropoid apes on
+the other, which will undoubtedly take place, the gulf will be greater
+than ever, since the baboons will then bound it on the one side, and
+the white races on the other. Little weight need be attached to the
+lack of fossil remains to fill up this gap, since the discovery of
+these depends upon chance. The last part of the chapter is devoted to
+a discussion of the earlier stages in the genealogy of man. Here
+Darwin accepts in the main the genealogical tree, which had meantime
+been published by Haeckel, who<span class='pagenum'><a name="Page_128" id="Page_128">[128]</a></span> traces the pedigree back through
+Monotrems, Reptiles, Amphibians, and Fishes, to Amphioxus.</p>
+
+<p>Then follows an attempt to reconstruct, from the atavistic characters,
+a picture of our primitive ancestor who was undoubtedly an arboreal
+animal. The occurrence of rudiments of parts in one sex which only
+come to full development in the other is next discussed. This state of
+things Darwin regards as derived from an original hermaphroditism. In
+regard to the mammary glands of the male he does not accept the theory
+that they are vestigial, but considers them rather as not fully
+developed.</p>
+
+<p>The last chapter of Part I deals with the question whether the
+different races of man are to be regarded as different species, or as
+sub-species of a race of monophyletic origin. The striking differences
+between the races are first emphasised, and the question of the
+fertility or infertility of hybrids is discussed. That fertility is
+the more usual is shown by the excessive fertility of the hybrid
+population of Brazil. This, and the great variability of the
+distinguishing characters of the different races, as well as the fact
+that all grades of transition stages are found between these, while
+considerable general agreement exists, tell in favour of the unity of
+the races and lead to the conclusion that they all had a common
+primitive ancestor.</p>
+
+<p>Darwin therefore classifies all the different races as sub-species of
+<i>one and the same species</i>. Then follows an interesting inquiry into
+the reasons for the extinction of human races. He recognises as the
+ultimate reason the injurious effects of a change of the conditions of
+life, which may bring about an increase in infantile mortality, and a
+diminished fertility. It is precisely the reproductive system, among
+animals also, which is most susceptible to changes in the environment.</p>
+
+<p>The final section of this chapter deals with the formation of the
+races of mankind. Darwin discusses the question how far the direct
+effect of different conditions of life, or the inherited effects of
+increased use or disuse may have<span class='pagenum'><a name="Page_129" id="Page_129">[129]</a></span> brought about the characteristic
+differences between the different races. Even in regard to the origin
+of the colour of the skin he rejects the transmitted effects of an
+original difference of climate as an explanation. In so doing he is
+following his tendency to exclude Lamarckian explanations as far as
+possible. But here he makes gratuitous difficulties from which, since
+natural selection fails, there is no escape except by bringing in the
+principle of sexual selection, to which, he regarded it as possible,
+skin-colouring, arrangement of hair, and form of features might be
+traced. But with his characteristic conscientiousness he guards
+himself thus: "I do not intend to assert that sexual selection will
+account for all the differences between the races."<a name="FNanchor_103_103" id="FNanchor_103_103"></a><a href="#Footnote_103_103" class="fnanchor">[103]</a></p>
+
+<p>I may be permitted a remark as to Darwin's attitude towards Lamarck.
+While, at an earlier stage, when he was engaged in the preliminary
+labours for his immortal work, <i>The Origin of Species</i>, Darwin
+expresses himself very forcibly against the views of Lamarck, speaking
+of Lamarckian "nonsense,"<a name="FNanchor_104_104" id="FNanchor_104_104"></a><a href="#Footnote_104_104" class="fnanchor">[104]</a> and of Lamarck's "absurd, though clever
+work"<a name="FNanchor_105_105" id="FNanchor_105_105"></a><a href="#Footnote_105_105" class="fnanchor">[105]</a> and expressly declaring, "I attribute very little to the
+direct action of climate, etc."<a name="FNanchor_106_106" id="FNanchor_106_106"></a><a href="#Footnote_106_106" class="fnanchor">[106]</a> yet in later life he became more
+and more convinced of the influence of external conditions. In 1876,
+that is, two years after the appearance of the second edition of <i>The
+Descent of Man</i>, he writes with his usual candid honesty: "In my
+opinion the greatest error which I have committed, has been not
+allowing sufficient weight to the direct action of the environment,
+i.e. food, climate, etc. independently of a natural selection."<a name="FNanchor_107_107" id="FNanchor_107_107"></a><a href="#Footnote_107_107" class="fnanchor">[107]</a>
+It is certain from this change of opinion that, if he had been able to
+make up his mind to issue a third edition of <i>The Descent of Man</i>, he
+would have ascribed a much greater influence <span class='pagenum'><a name="Page_130" id="Page_130">[130]</a></span>to the effect of
+external conditions in explaining the different characters of the
+races of man than he did in the second edition. He would also
+undoubtedly have attributed less influence to sexual selection as a
+factor in the origin of the different bodily characteristics, if
+indeed he would not have excluded it altogether.</p>
+
+<p>In Part III of the <i>Descent</i> two additional chapters are devoted to
+the discussion of sexual selection in relation to man. These may be
+very briefly referred to. Darwin here seeks to show that sexual
+selection has been operative on man and his primitive progenitor.
+Space fails me to follow out his interesting arguments. I can only
+mention that he is inclined to trace back hairlessness, the
+development of the beard in man, and the characteristic colour of the
+different human races to sexual selection. Since bareness of the skin
+could be no advantage, but rather a disadvantage, this character
+cannot have been brought about by natural selection. Darwin also
+rejected a direct influence of climate as a cause of the origin of the
+skin-colour. I have already expressed the opinion, based on the
+development of his views as shown in his letters, that in a third
+edition Darwin would probably have laid more stress on the influence
+of external environment. He himself feels that there are gaps in his
+proofs here, and says in self-criticism: "The views here advanced, on
+the part which sexual selection has played in the history of man, want
+scientific precision."<a name="FNanchor_108_108" id="FNanchor_108_108"></a><a href="#Footnote_108_108" class="fnanchor">[108]</a> I need here only point out that it is
+impossible to explain the graduated stages of skin-colour by sexual
+selection, since it would have produced races sharply defined by their
+colour and not united to other races by transition stages, and this,
+it is well known, is not the case. Moreover, the fact established by
+me,<a name="FNanchor_109_109" id="FNanchor_109_109"></a><a href="#Footnote_109_109" class="fnanchor">[109]</a> that in all races the ventral side of the trunk is paler than
+the dorsal side, and the inner surface of <span class='pagenum'><a name="Page_131" id="Page_131">[131]</a></span>the extremities paler than
+the outer side, cannot be explained by sexual selection in the
+Darwinian sense.</p>
+
+<p>With this I conclude my brief survey of the rich contents of Darwin's
+book. I may be permitted to conclude by quoting the magnificent final
+words of <i>The Descent of Man</i>: "We must, however, acknowledge, as it
+seems to me, that man, with all his noble qualities, with sympathy
+which feels for the most debased, with benevolence which extends not
+only to other men but to the humblest living creature, with his
+god-like intellect which has penetrated into the movements and
+constitution of the solar system&mdash;with all these exalted powers&mdash;Man
+still bears in his bodily frame the indelible stamp of his lowly
+origin."<a name="FNanchor_110_110" id="FNanchor_110_110"></a><a href="#Footnote_110_110" class="fnanchor">[110]</a></p>
+
+<p>What has been the fate of Darwin's doctrines since his great
+achievement? How have they been received and followed up by the
+scientific and lay world? And what do the successors of the mighty
+hero and genius think now in regard to the origin of the human race?</p>
+
+<p>At the present time we are incomparably more favourably placed than
+Darwin was for answering this question of all questions. We have at
+our command an incomparably greater wealth of material than he had at
+his disposal. And we are more fortunate than he in this respect, that
+we now know transition-forms which help to fill up the gap, still
+great, between the lowest human races and the highest apes. Let us
+consider for a little the more essential additions to our knowledge
+since the publication of <i>The Descent of Man</i>.</p>
+
+<p>Since that time our knowledge of animal embryos has increased
+enormously. While Darwin was obliged to content himself with comparing
+a human embryo with that of a dog, there are now available the
+youngest embryos of monkeys of all possible groups (Orang, Gibbon,
+Semnopithecus, Macacus), thanks to Selenka's most successful tour in
+the East Indies in search of such material. We can now compare
+corresponding stages of the lower monkeys and of <span class='pagenum'><a name="Page_132" id="Page_132">[132]</a></span>the Anthropoid apes
+with human embryos, and convince ourselves of their great resemblance
+to one another, thus strengthening enormously the armour prepared by
+Darwin in defence of his view on man's nearest relatives. It may be
+said that Selenka's material fills up the blanks in Darwin's array of
+proofs in the most satisfactory manner.</p>
+
+<p>The deepening of our knowledge of comparative anatomy also gives us
+much surer foundations than those on which Darwin was obliged to
+build. Just of late there have been many workers in the domain of the
+anatomy of apes and lemurs, and their investigations extend to the
+most different organs. Our knowledge of fossil apes and lemurs has
+also become much wider and more exact since Darwin's time: the fossil
+lemurs have been especially worked up by Cope, Forsyth Major,
+Ameghino, and others. Darwin knew very little about fossil monkeys. He
+mentions two or three anthropoid apes as occurring in the Miocene of
+Europe,<a name="FNanchor_111_111" id="FNanchor_111_111"></a><a href="#Footnote_111_111" class="fnanchor">[111]</a> but only names <i>Dryopithecus</i>, the largest form from the
+Miocene of France. It was erroneously supposed that this form was
+related to <i>Hylobates</i>. We now know not only a form that actually
+stands near to the gibbon (<i>Pliopithecus</i>), and remains of other
+anthropoids (<i>Pliohylobates</i> and the fossil chimpanzee,
+<i>Palaeopithecus</i>), but also several lower catarrhine monkeys, of which
+<i>Mesopithecus</i>, a form nearly related to the modern Sacred Monkeys (a
+species of <i>Semnopithecus</i>) and found in strata of the Miocene period
+in Greece, is the most important. Quite recently, too, Ameghino's
+investigations have made us acquainted with fossil monkeys from South
+America (<i>Anthropops</i>, <i>Homunculus</i>), which, according to their
+discoverer, are to be regarded as in the line of human descent.</p>
+
+<p>What Darwin missed most of all&mdash;intermediate forms between apes and
+man&mdash;has been recently furnished. E. Dubois, as is well known,
+discovered in 1893, near Trinil in Java, in the alluvial deposits of
+the river Bengawan, an important form represented by a skull-cap, some
+molars, <span class='pagenum'><a name="Page_133" id="Page_133">[133]</a></span>and a femur. His opinion&mdash;much disputed as it has been&mdash;that
+in this form, which he named <i>Pithecanthropus</i>, he has found a
+long-desired transition-form is shared by the present writer. And
+although the geological age of these fossils, which, according to
+Dubois, belong to the uppermost Tertiary series, the Pliocene has
+recently been fixed at a later date (the older Diluvium), the
+<i>morphological value</i> of these interesting remains, that is, the
+intermediate position of <i>Pithecanthropus</i>, still holds good. Volz
+says with justice,<a name="FNanchor_112_112" id="FNanchor_112_112"></a><a href="#Footnote_112_112" class="fnanchor">[112]</a> that even if <i>Pithecanthropus</i> is not <i>the</i>
+missing link, it is undoubtedly <i>a</i> missing link.</p>
+
+<p>As on the one hand there has been found in <i>Pithecanthropus</i> a form
+which, though intermediate between apes and man, is nevertheless more
+closely allied to the apes, so on the other hand, much progress has
+been made since Darwin's day in the discovery and description of the
+oldest human remains. Since the famous roof of a skull and the bones
+of the extremities belonging to it were found in 1856 in the
+Neandertal near D&uuml;sseldorf, the most varied judgments have been
+expressed in regard to the significance of the remains and of the
+skull in particular. In Darwin's <i>Descent of Man</i> there is only a
+passing allusion to them<a name="FNanchor_113_113" id="FNanchor_113_113"></a><a href="#Footnote_113_113" class="fnanchor">[113]</a> in connection with the discussion of the
+skull-capacity, although the investigations of Schaaffhausen, King,
+and Huxley were then known. I believe I have shown, in a series of
+papers, that the skull in question belongs to a form different from
+any of the races of man now living, and, with King and Cope, I regard
+it as at least a different species from living man, and have therefore
+designated it <i>Homo primigenius</i>. The form unquestionably belongs to
+the older Diluvium, and in the later Diluvium human forms already
+appear, which agree in all essential points with existing human races.</p>
+
+<p><span class='pagenum'><a name="Page_134" id="Page_134">[134]</a></span></p>
+<p>As far back as 1886 the value of the Neandertal skull was greatly
+enhanced by Fraipont's discovery of two skulls and skeletons from Spy
+in Belgium. These are excellently described by their discoverer,<a name="FNanchor_114_114" id="FNanchor_114_114"></a><a href="#Footnote_114_114" class="fnanchor">[114]</a>
+and are regarded as belonging to the same group of forms as the
+Neandertal remains. In 1899 and the following years came the discovery
+by Gorjanovi&#269;-Kramberger of different skeletal parts of at least
+ten individuals in a cave near Krapina in Croatia.<a name="FNanchor_115_115" id="FNanchor_115_115"></a><a href="#Footnote_115_115" class="fnanchor">[115]</a> It is in
+particular the form of the lower jaw which is different from that of
+all recent races of man, and which clearly indicates the lowly
+position of <i>Homo primigenius</i>, while, on the other hand, the
+long-known skull from Gibraltar, which I<a name="FNanchor_116_116" id="FNanchor_116_116"></a><a href="#Footnote_116_116" class="fnanchor">[116]</a> have referred to <i>Homo
+primigenius</i>, and which has lately been examined in detail by
+Sollas,<a name="FNanchor_117_117" id="FNanchor_117_117"></a><a href="#Footnote_117_117" class="fnanchor">[117]</a> has made us acquainted with the surprising shape of the
+eye-orbit, of the nose, and of the whole upper part of the face.
+Isolated lower jaws found at La Naulette in Belgium, and at Malarnaud
+in France, increase our material which is now as abundant as could be
+desired. The most recent discovery of all is that of a skull dug up in
+August of this year [1908] by Klaatsch and Hauser in the lower grotto
+of the Le Moustier in Southern France, but this skull has not yet been
+fully described. Thus <i>Homo primigenius</i> must also be regarded as
+occupying a position in the gap existing between the highest apes and
+the lowest human races, <i>Pithecanthropus</i>, standing in the lower part
+of it, and <i>Homo primigenius</i> in the higher, near man. In order to
+prevent misunderstanding, I should like here to emphasise that in
+arranging this structural series&mdash;anthropoid apes, <i>Pithecanthropus</i>,
+<i>Homo primigenius</i>, <i>Homo sapiens</i>&mdash;I have no intention of
+establishing <span class='pagenum'><a name="Page_135" id="Page_135">[135]</a></span>it as a direct genealogical series. I shall have
+something to say in regard to the genetic relations of these forms,
+one to another, when discussing the different theories of descent
+current at the present day.<a name="FNanchor_118_118" id="FNanchor_118_118"></a><a href="#Footnote_118_118" class="fnanchor">[118]</a></p>
+
+<p>In quite a different domain from that of morphological relationship,
+namely in the physiological study of the blood, results have recently
+been gained which are of the highest importance to the doctrine of
+descent. Uhlenhuth, Nuttall, and others have established the fact that
+the blood-serum of a rabbit which has previously had human blood
+injected into it, forms a precipitate with human blood. This
+biological reaction was tried with a great variety of mammalian
+species, and it was found that those far removed from man gave no
+precipitate under these conditions. But as in other cases among
+mammals all nearly related forms yield an almost equally marked
+precipitate, so the serum of a rabbit treated with human blood and
+then added to the blood of an anthropoid ape gives <i>almost</i> as marked
+a precipitate as in human blood; the reaction to the blood of the
+lower Eastern monkeys is weaker, that to the Western monkeys weaker
+still; indeed in this last case there is only a slight clouding after
+a considerable time and no actual precipitate. The blood of the
+Lemuridae (Nuttall) gives no reaction or an extremely weak one, that
+of the other mammals none whatever. We have in this not only a proof
+of the literal blood relationship between man and apes, but the degree
+of relationship with the different main groups of apes can be
+determined beyond possibility of mistake.</p>
+
+<p>Finally, it must be briefly mentioned that in regard to remains of
+human handicraft also, the material at our disposal has greatly
+increased of late years, that, as a result of this, the opinions of
+archaeologists have undergone many <span class='pagenum'><a name="Page_136" id="Page_136">[136]</a></span>changes, and that, in particular,
+their views in regard to the age of the human race have been greatly
+influenced. There is a tendency at the present time to refer the
+origin of man back to Tertiary times. It is true that no remains of
+Tertiary man have been found, but flints have been discovered which,
+according to the opinion of most investigators, bear traces either of
+use, or of very primitive workmanship. Since Rutot's time, following
+Mortillet's example, investigators have called these "eoliths," and
+they have been traced back by Verworn to the Miocene of the Auvergne,
+and by Rutot even to the upper Oligocene. Although these eoliths are
+even nowadays the subject of many different views, the preoccupation
+with them has kept the problem of the age of the human race
+continually before us.</p>
+
+<p>Geology, too, has made great progress since the days of Darwin and
+Lyell, and has endeavoured with satisfactory results to arrange the
+human remains of the Diluvial period in chronological order (Penck). I
+do not intend to enter upon the question of the primitive home of the
+human race; since the space at my disposal will not allow of my
+touching even very briefly upon all the departments of science which
+are concerned in the problem of the descent of man. How Darwin would
+have rejoiced over each of the discoveries here briefly outlined! What
+use he would have made of the new and precious material, which would
+have prevented the discouragement from which he suffered when
+preparing the second edition of <i>The Descent of Man</i>! But it was not
+granted to him to see this progress towards filling up the gaps in his
+edifice of which he was so painfully conscious.</p>
+
+<p>He did, however, have the satisfaction of seeing his ideas steadily
+gaining ground, notwithstanding much hostility and deep-rooted
+prejudice. Even in the years between the appearance of <i>The Origin of
+Species</i> and of the first edition of the <i>Descent</i>, the idea of a
+natural descent of man, which was only briefly indicated in the work
+of 1859, had been eagerly welcomed in some quarters. It has been
+already<span class='pagenum'><a name="Page_137" id="Page_137">[137]</a></span> pointed out how brilliantly Huxley contributed to the defence
+and diffusion of Darwin's doctrines, and how in <i>Man's Place in
+Nature</i> he has given us a classic work as a foundation for the
+doctrine of the descent of man. As Huxley was Darwin's champion in
+England, so in Germany Carl Vogt, in particular, made himself master
+of the Darwinian ideas. But above all it was Haeckel who, in energy,
+eagerness for battle, and knowledge may be placed side by side with
+Huxley, who took over the leadership in the controversy over the new
+conception of the universe. As far back as 1866, in his <i>Generelle
+Morphologie</i>, he had inquired minutely into the question of the
+descent of man, and not content with urging merely the general theory
+of descent from lower animal forms, he drew up for the first time
+genealogical trees showing the close structural relationships of the
+different animal groups; the last of these illustrated the
+relationships of Mammals, and among them of all groups of the
+Primates, including man. It was Haeckel's genealogical trees that
+formed the basis of the special discussion of the relationships of
+man, in the sixth chapter of Darwin's <i>Descent of Man</i>.</p>
+
+<p>In the last section of this essay I shall return to Haeckel's
+conception of the special descent of man, the main features of which
+he still upholds, and rightly so. Haeckel has contributed more than
+any one else to the spread of the Darwinian doctrine.</p>
+
+<p>I can only allow myself a few words as to the spread of the theory of
+the natural descent of man in other countries. The Parisian
+anthropological school, founded and guided by the genius of Broca,
+took up the idea of the descent of man, and made many notable
+contributions to it (Broca, Manouvrier, Mahoudeau, Deniker and
+others). In England itself Darwin's work did not die. Huxley took care
+of that, for he, with his lofty and unprejudiced mind, dominated and
+inspired English biology until his death on June 29, 1895. He had the
+satisfaction shortly before his death of learning of Dubois'
+discovery, which he illustrated by a<span class='pagenum'><a name="Page_138" id="Page_138">[138]</a></span> humourous sketch.<a name="FNanchor_119_119" id="FNanchor_119_119"></a><a href="#Footnote_119_119" class="fnanchor">[119]</a> But there
+are still many followers in Darwin's footsteps in England. Keane has
+worked at the special genealogical tree of the Primates; Keith has
+inquired which of the anthropoid apes has the greatest number of
+characters in common with man; Morris concerns himself with the
+evolution of man in general, especially with his acquisition of the
+erect position. The recent discoveries of <i>Pithecanthropus</i> and <i>Homo
+primigenius</i> are being vigorously discussed; but the present writer is
+not in a position to form an opinion of the extent to which the idea
+of descent has penetrated throughout England generally.</p>
+
+<p>In Italy independent work in the domain of the descent of man is being
+produced, especially by Morselli; with him are associated, in the
+investigation of related problems, Sergi and Giuffrida-Ruggeri. From
+the ranks of American investigators we may single out in particular
+the eminent geologist Cope, who championed with much decision the idea
+of the specific difference of <i>Homo neandertalensis</i> (<i>primigenius</i>)
+and maintained a more direct descent of man from the fossil Lemuridae.
+In South America too, in Argentina, new life is stirring in this
+department of science. Ameghino in Buenos Ayres has awakened the
+fossil primates of the Pampas formation to new life; he even believes
+that in his <i>Tetraprothomo</i>, represented by a femur, he has discovered
+a direct ancestor of man. Lehmann-Nitsche is working at the other side
+of the gulf between apes and man, and he describes a remarkable first
+cervical vertebra (atlas) from Monte Hermoso as belonging to a form
+which may bear the same relation to <i>Homo sapiens</i> in South America as
+<i>Homo primigenius</i> does in the Old World. After a minute investigation
+he establishes a human species <i>Homo neogaeus</i>, while Ameghino
+ascribes this atlas vertebra to his <i>Tetraprothomo</i>.</p>
+
+<p>Thus throughout the whole scientific world there is arising a new
+life, an eager endeavour to get nearer to Huxley's <i>problema maximum</i>,
+to penetrate more deeply into the <span class='pagenum'><a name="Page_139" id="Page_139">[139]</a></span>origin of the human race. There are
+to-day very few experts in anatomy and zoology who deny the animal
+descent of man in general. Religious considerations, old prejudices,
+the reluctance to accept man, who so far surpasses mentally all other
+creatures, as descended from "soulless" animals, prevent a few
+investigators from giving full adherence to the doctrine. But there
+are very few of these who still postulate a special act of creation
+for man. Although the majority of experts in anatomy and zoology
+accept unconditionally the descent of man from lower forms, there is
+much diversity of opinion among them in regard to the special line of
+descent.</p>
+
+<p>In trying to establish any special hypothesis of descent, whether by
+the graphic method of drawing up genealogical trees or otherwise, let
+us always bear in mind Darwin's words<a name="FNanchor_120_120" id="FNanchor_120_120"></a><a href="#Footnote_120_120" class="fnanchor">[120]</a> and use them as a critical
+guiding line: "As we have no record of the lines of descent, the
+pedigree can be discovered only by observing the degrees of
+resemblance between the beings which are to be classed." Darwin
+carries this further by stating "that resemblances in several
+unimportant structures, in useless and rudimentary organs, or not now
+functionally active, or in an embryological condition, are by far the
+most serviceable for classification."<a name="FNanchor_121_121" id="FNanchor_121_121"></a><a href="#Footnote_121_121" class="fnanchor">[121]</a> It has also to be
+remembered that <i>numerous</i> separate points of agreement are of much
+greater importance than the amount of similarity or dissimilarity in a
+few points.</p>
+
+<p>The hypotheses as to descent current at the present day may be divided
+into two main groups. The first group seeks for the roots of the human
+race not among any of the families of the apes&mdash;the anatomically
+nearest forms&mdash;nor among their very similar but less specialised
+ancestral forms, the fossil representatives of which we can know only
+in part, but, setting the monkeys on one side, it seeks for them lower
+down among the fossil Eocene Pseudo-lemuridae or Lemuridae (Cope), or
+even among the primitive pentadactylous <span class='pagenum'><a name="Page_140" id="Page_140">[140]</a></span>Eocene forms, which may
+either have led directly to the evolution of man (Adloff), or have
+given rise to an ancestral form common to apes and men (Klaatsch,<a name="FNanchor_122_122" id="FNanchor_122_122"></a><a href="#Footnote_122_122" class="fnanchor">[122]</a>
+Giuffrida-Ruggeri). The common ancestral form, from which man and apes
+are thus supposed to have arisen independently, may explain the
+numerous resemblances which actually exist between them. That is to
+say, all the characters upon which the great structural resemblance
+between apes and man depends must have been present in their common
+ancestor. Let us take an example of such a common character. The bony
+external ear-passage is in general as highly developed in the lower
+Eastern monkeys and the anthropoid apes as in man. This character
+must, therefore, have already been present in the common primitive
+form. In that case it is not easy to understand why the Western
+monkeys have not also inherited the character, instead of possessing
+only a tympanic ring. But it becomes more intelligible if we assume
+that forms with a primitive tympanic ring were the original type, and
+that from these were evolved, on the one hand, the existing New World
+monkeys with persistent tympanic ring, and on the other an ancestral
+form common to the lower Old World monkeys, the anthropoid apes and
+man. For man shares with these the character in question, and it is
+also one of the "unimportant" characters required by Darwin. Thus we
+have two divergent lines arising from the ancestral form, the Western
+monkeys (Platyrrhine) on the one hand, and an ancestral form common to
+the lower Eastern monkeys, the anthropoid apes, and man, on the other.
+But considerations similar to those which showed it to be impossible
+that man should have developed from an ancestor common to him and the
+monkeys, yet outside of and parallel with these, may be urged also
+against the likelihood of a parallel evolution of the lower Eastern
+monkeys, the anthropoid apes, and man. The anthropoid apes have in
+common with <span class='pagenum'><a name="Page_141" id="Page_141">[141]</a></span>man many characters which are not present in the lower
+Old World monkeys. These characters must therefore have been present
+in the ancestral form common to the three groups. But here, again, it
+is difficult to understand why the lower Eastern monkeys should not
+also have inherited these characters. As this is not the case, there
+remains no alternative but to assume divergent evolution from an
+indifferent form. The lower Eastern monkeys are carrying on the
+evolution in one direction&mdash;I might almost say towards a blind
+alley&mdash;while anthropoids and men have struck out a progressive path,
+at first in common, which explains the many points of resemblance
+between them, without regarding man as derived directly from the
+anthropoids. Their many striking points of agreement indicate a common
+descent, and cannot be explained as phenomena of convergence.</p>
+
+<p>I believe I have shown in the above sketch that a theory which derives
+man directly from lower forms without regarding apes as
+transition-types leads <i>ad absurdum</i>. The close structural
+relationship between man and monkeys can only be understood if both
+are brought into the same line of evolution. To trace man's line of
+descent directly back to the old Eocene mammals, alongside of, but
+with no relation to these very similar forms, is to abandon the method
+of exact comparison, which, as Darwin rightly recognised, alone
+justifies us in drawing up genealogical trees on the basis of
+resemblances and differences. The farther down we go the more does the
+ground slip from beneath our feet. Even the Lemuridae show very
+numerous divergent conditions, much more so the Eocene mammals
+(Creodonta, Condylarthra), the chief resemblance of which to man
+consists in the possession of pentadactylous hands and feet! Thus the
+farther course of the line of descent disappears in the darkness of
+the ancestry of the mammals. With just as much reason we might pass by
+the Vertebrates altogether, and go back to the lower Invertebrates,
+but in that case it would be much easier to say that man has arisen
+indepen<span class='pagenum'><a name="Page_142" id="Page_142">[142]</a></span>dently, and has evolved, without relation to any animals, from
+the lowest primitive form to his present isolated and dominant
+position. But this would be to deny all value to classification, which
+must after all be the ultimate basis of a genealogical tree. We can,
+as Darwin rightly observed, only infer the line of descent from the
+degree of resemblance between single forms. If we regard man as
+directly derived from primitive forms very far back, we have no way of
+explaining the many points of agreement between him and the monkeys in
+general, and the anthropoid apes in particular. These must remain an
+inexplicable marvel.</p>
+
+<p>I have thus, I trust, shown that the first class of special theories
+of descent, which assumes that man has developed, parallel with the
+monkeys, but without relation to them, from very low primitive forms
+cannot be upheld, because it fails to take into account the close
+structural affinity of man and monkeys. I cannot but regard this
+hypothesis as lamentably retrograde, for it makes impossible any
+application of the facts that have been discovered in the course of
+the anatomical and embryological study of man and monkeys, and indeed
+prejudges investigations of that class as pointless. The whole method
+is perverted; an unjustifiable theory of descent is first formulated
+with the aid of the imagination, and then we are asked to declare that
+all structural relations between man and monkeys, and between the
+different groups of the latter, are valueless,&mdash;the fact being that
+they are the only true basis on which a genealogical tree can be
+constructed.</p>
+
+<p>So much for this most modern method of classification, which has
+probably found adherents because it would deliver us from the
+relationship to apes which many people so much dislike. In contrast to
+it we have the second class of special hypotheses of descent, which
+keeps strictly to the nearest structural relationship. This is the
+only basis that justifies the drawing up of a special hypothesis of
+descent. If this fundamental proposition be recognised, it will be
+admitted that the doctrine of special descent upheld<span class='pagenum'><a name="Page_143" id="Page_143">[143]</a></span> by Haeckel, and
+set forth in Darwin's <i>Descent of Man</i>, is still valid to-day. In the
+genealogical tree, man's place is quite close to the anthropoid apes;
+these again have as their nearest relatives the lower Old World
+monkeys, and their progenitors must be sought among the less
+differentiated Platyrrhine monkeys, whose most important characters
+have been handed on to the present day New World monkeys. How the
+different genera are to be arranged within the general scheme
+indicated depends in the main on the classificatory value attributed
+to individual characters. This is particularly true in regard to
+<i>Pithecanthropus</i>, which I consider as the root of a branch which has
+sprung from the anthropoid ape root and has led up to man; the latter
+I have designated the family of the Hominidae.</p>
+
+<p>For the rest, there are, as we have said, various possible ways of
+constructing the narrower genealogy within the limits of this branch
+including men and apes, and these methods will probably continue to
+change with the accumulation of new facts. Haeckel himself has
+modified his genealogical tree of the Primates in certain details
+since the publication of his <i>Generelle Morphologie</i> in 1866, but its
+general basis remains the same.<a name="FNanchor_123_123" id="FNanchor_123_123"></a><a href="#Footnote_123_123" class="fnanchor">[123]</a> All the special genealogical
+trees drawn up on the lines laid down by Haeckel and Darwin&mdash;and that
+of Dubois may be specially mentioned&mdash;are based, in general, on the
+close relationship of monkeys and men, although they may vary in
+detail. Various hypotheses have been formulated on these lines, with
+special reference to the evolution of man. <i>Pithecanthropus</i> is
+regarded by some authorities as the direct ancestor of man, by others
+as a side-track failure in the attempt at the evolution of man. The
+problem of the monophyletic or polyphyletic origin of the human race
+has also been much discussed. Sergi<a name="FNanchor_124_124" id="FNanchor_124_124"></a><a href="#Footnote_124_124" class="fnanchor">[124]</a> inclines towards the
+assumption of a polyphyletic origin of the three main races of man,
+the African primitive form of which <span class='pagenum'><a name="Page_144" id="Page_144">[144]</a></span>has given rise also to the
+gorilla and chimpanzee, the Asiatic to the Orang, the Gibbon, and
+<i>Pithecanthropus</i>. Kollmann regards existing human races as derived
+from small primitive races (pigmies), and considers that <i>Homo
+primigenius</i> must have arisen in a secondary and degenerative manner.</p>
+
+<p>But this is not the place, nor have I the space to criticise the
+various special theories of descent. One, however, must receive
+particular notice. According to Ameghino, the South American monkeys
+(<i>Pitheculites</i>) from the oldest Tertiary of the Pampas are the forms
+from which have arisen the existing American monkeys on the one hand,
+and on the other, the extinct South American Homunculidae, which are
+also small forms. From these last, anthropoid apes and man have, he
+believes, been evolved. Among the progenitors of man, Ameghino reckons
+the form discovered by him (<i>Tetraprothomo</i>), from which a South
+American primitive man, <i>Homo pampaeus</i>, might be directly evolved,
+while on the other hand all the lower Old World monkeys may have
+arisen from older fossil South American forms (Clenialitidae), the
+distribution of which may be explained by the bridge formerly existing
+between South America and Africa, as may be the derivation of all
+existing human races from <i>Homo pampaeus</i>.<a name="FNanchor_125_125" id="FNanchor_125_125"></a><a href="#Footnote_125_125" class="fnanchor">[125]</a> The fossil forms
+discovered by Ameghino deserve the most minute investigation, as does
+also the fossil man from South America of which Lehmann-Nitsche<a name="FNanchor_126_126" id="FNanchor_126_126"></a><a href="#Footnote_126_126" class="fnanchor">[126]</a>
+has made a thorough study.</p>
+
+<p>It is obvious that, notwithstanding the necessity for fitting man's
+line of descent into the genealogical tree of the Primates, especially
+the apes, opinions in regard to it differ greatly in detail. This
+could not be otherwise, since the different Primate forms, especially
+the fossile forms, are still <span class='pagenum'><a name="Page_145" id="Page_145">[145]</a></span>far from being exhaustively known. But
+one thing remains certain,&mdash;the idea of the close relationship between
+man and monkeys set forth in Darwin's <i>Descent of Man</i>. Only those who
+deny the many points of agreement, the sole basis of classification,
+and thus of a natural genealogical tree, can look upon the position of
+Darwin and Haeckel as antiquated, or as standing on an insufficient
+foundation. For such a genealogical tree is nothing more than a
+summarised representation of what is known in regard to the degree of
+resemblance between the different forms.</p>
+
+<p>Darwin's work in regard to the descent of man has not been surpassed;
+the more we immerse ourselves in the study of the structural
+relationships between apes and man, the more is our path illumined by
+the clear light radiating from him, and through his calm and
+deliberate investigation, based on a mass of material in the
+accumulation of which he has never had an equal. Darwin's fame will be
+bound up for all time with the unprejudiced investigation of the
+question of all questions, the descent of the human race.</p>
+
+<div class="footnotes"><h3>FOOTNOTES:</h3>
+
+<div class="footnote"><p><a name="Footnote_75_75" id="Footnote_75_75"></a><a href="#FNanchor_75_75"><span class="label">[75]</span></a> <i>Life and Letters of Thomas Henry Huxley</i>, Vol. I. p.
+171, London, 1900.</p></div>
+
+<div class="footnote"><p><a name="Footnote_76_76" id="Footnote_76_76"></a><a href="#FNanchor_76_76"><span class="label">[76]</span></a> <i>Ibid.</i>, p. 363.</p></div>
+
+<div class="footnote"><p><a name="Footnote_77_77" id="Footnote_77_77"></a><a href="#FNanchor_77_77"><span class="label">[77]</span></a> No italics in original.</p></div>
+
+<div class="footnote"><p><a name="Footnote_78_78" id="Footnote_78_78"></a><a href="#FNanchor_78_78"><span class="label">[78]</span></a> <i>Life and Letters of Charles Darwin</i>, Vol. I. p. 93.</p></div>
+
+<div class="footnote"><p><a name="Footnote_79_79" id="Footnote_79_79"></a><a href="#FNanchor_79_79"><span class="label">[79]</span></a> <i>Ibid.</i> Vol. II. p. 263.</p></div>
+
+<div class="footnote"><p><a name="Footnote_80_80" id="Footnote_80_80"></a><a href="#FNanchor_80_80"><span class="label">[80]</span></a> <i>Ibid.</i> Vol. I. p. 94.</p></div>
+
+<div class="footnote"><p><a name="Footnote_81_81" id="Footnote_81_81"></a><a href="#FNanchor_81_81"><span class="label">[81]</span></a> <i>Life and Letters</i>, Vol. III. p. 175.</p></div>
+
+<div class="footnote"><p><a name="Footnote_82_82" id="Footnote_82_82"></a><a href="#FNanchor_82_82"><span class="label">[82]</span></a> <i>Ibid.</i> Vol. II. p. 109.</p></div>
+
+<div class="footnote"><p><a name="Footnote_83_83" id="Footnote_83_83"></a><a href="#FNanchor_83_83"><span class="label">[83]</span></a> <i>Ibid.</i> Vol. III. p. 112.</p></div>
+
+<div class="footnote"><p><a name="Footnote_84_84" id="Footnote_84_84"></a><a href="#FNanchor_84_84"><span class="label">[84]</span></a> <i>Ibid.</i> Vol. I. pp. 304-317.</p></div>
+
+<div class="footnote"><p><a name="Footnote_85_85" id="Footnote_85_85"></a><a href="#FNanchor_85_85"><span class="label">[85]</span></a> <i>Life and Letters</i>, Vol. I. p. 309.</p></div>
+
+<div class="footnote"><p><a name="Footnote_86_86" id="Footnote_86_86"></a><a href="#FNanchor_86_86"><span class="label">[86]</span></a> <i>Loc. cit.</i> p. 313.</p></div>
+
+<div class="footnote"><p><a name="Footnote_87_87" id="Footnote_87_87"></a><a href="#FNanchor_87_87"><span class="label">[87]</span></a> <i>Ibid.</i> Vol. II. p. 310.</p></div>
+
+<div class="footnote"><p><a name="Footnote_88_88" id="Footnote_88_88"></a><a href="#FNanchor_88_88"><span class="label">[88]</span></a> <i>Ibid.</i> Vol. III. p. 236. ["C. Ridley," Mr. Francis
+Darwin points out to me, should be H. N. Ridley. A.C.S.]</p></div>
+
+<div class="footnote"><p><a name="Footnote_89_89" id="Footnote_89_89"></a><a href="#FNanchor_89_89"><span class="label">[89]</span></a> <i>Descent of Man</i> (Popular Edit., 1901), fig. 1, p. 14.</p></div>
+
+<div class="footnote"><p><a name="Footnote_90_90" id="Footnote_90_90"></a><a href="#FNanchor_90_90"><span class="label">[90]</span></a> G. Schwalbe, "Das Darwin'sche Spitzohr beim menschlichen
+Embryo," <i>Anatom. Anzeiger</i>, 1889, pp. 176-189, and other papers.</p></div>
+
+<div class="footnote"><p><a name="Footnote_91_91" id="Footnote_91_91"></a><a href="#FNanchor_91_91"><span class="label">[91]</span></a> <i>Descent of Man</i>, fig. 3, p. 24.</p></div>
+
+<div class="footnote"><p><a name="Footnote_92_92" id="Footnote_92_92"></a><a href="#FNanchor_92_92"><span class="label">[92]</span></a> <i>Descent of man</i>, p. 6.</p></div>
+
+<div class="footnote"><p><a name="Footnote_93_93" id="Footnote_93_93"></a><a href="#FNanchor_93_93"><span class="label">[93]</span></a> <i>Ibid.</i> p. 54.</p></div>
+
+<div class="footnote"><p><a name="Footnote_94_94" id="Footnote_94_94"></a><a href="#FNanchor_94_94"><span class="label">[94]</span></a> <i>Descent of Man</i>, p. 92.</p></div>
+
+<div class="footnote"><p><a name="Footnote_95_95" id="Footnote_95_95"></a><a href="#FNanchor_95_95"><span class="label">[95]</span></a> <i>Ibid.</i> p. 100.</p></div>
+
+<div class="footnote"><p><a name="Footnote_96_96" id="Footnote_96_96"></a><a href="#FNanchor_96_96"><span class="label">[96]</span></a> <i>Life and letters</i>, Vol. II. p. 161, June 22, 1859.</p></div>
+
+<div class="footnote"><p><a name="Footnote_97_97" id="Footnote_97_97"></a><a href="#FNanchor_97_97"><span class="label">[97]</span></a> <i>Ibid.</i> Vol. III. p. 15, March 17, 1863.</p></div>
+
+<div class="footnote"><p><a name="Footnote_98_98" id="Footnote_98_98"></a><a href="#FNanchor_98_98"><span class="label">[98]</span></a> <i>Descent of Man</i>, p. 132.</p></div>
+
+<div class="footnote"><p><a name="Footnote_99_99" id="Footnote_99_99"></a><a href="#FNanchor_99_99"><span class="label">[99]</span></a> <i>Ibid.</i> pp. 136, 137.</p></div>
+
+<div class="footnote"><p><a name="Footnote_100_100" id="Footnote_100_100"></a><a href="#FNanchor_100_100"><span class="label">[100]</span></a> <i>Ibid.</i> p. 143.</p></div>
+
+<div class="footnote"><p><a name="Footnote_101_101" id="Footnote_101_101"></a><a href="#FNanchor_101_101"><span class="label">[101]</span></a> <i>Ibid.</i> p. 193.</p></div>
+
+<div class="footnote"><p><a name="Footnote_102_102" id="Footnote_102_102"></a><a href="#FNanchor_102_102"><span class="label">[102]</span></a> <i>Descent of Man</i>, p. 231.</p></div>
+
+<div class="footnote"><p><a name="Footnote_103_103" id="Footnote_103_103"></a><a href="#FNanchor_103_103"><span class="label">[103]</span></a> <i>Descent of Man</i>, p. 308.</p></div>
+
+<div class="footnote"><p><a name="Footnote_104_104" id="Footnote_104_104"></a><a href="#FNanchor_104_104"><span class="label">[104]</span></a> <i>Life and Letters</i>, Vol. II. p. 23.</p></div>
+
+<div class="footnote"><p><a name="Footnote_105_105" id="Footnote_105_105"></a><a href="#FNanchor_105_105"><span class="label">[105]</span></a> <i>Loc. cit.</i> p. 39.</p></div>
+
+<div class="footnote"><p><a name="Footnote_106_106" id="Footnote_106_106"></a><a href="#FNanchor_106_106"><span class="label">[106]</span></a> <i>Loc. cit.</i> (1856), p. 82.</p></div>
+
+<div class="footnote"><p><a name="Footnote_107_107" id="Footnote_107_107"></a><a href="#FNanchor_107_107"><span class="label">[107]</span></a> <i>Ibid.</i> Vol. III p. 159.</p></div>
+
+<div class="footnote"><p><a name="Footnote_108_108" id="Footnote_108_108"></a><a href="#FNanchor_108_108"><span class="label">[108]</span></a> <i>Descent of Man</i>, p. 924.</p></div>
+
+<div class="footnote"><p><a name="Footnote_109_109" id="Footnote_109_109"></a><a href="#FNanchor_109_109"><span class="label">[109]</span></a> "Die Hautfarbe des Menschen," <i>Mitteilungen der
+Anthropologischen Gessellschaft in Wien</i>, Vol. <span class="smcap">xxxiv</span>. pp. 331-352.</p></div>
+
+<div class="footnote"><p><a name="Footnote_110_110" id="Footnote_110_110"></a><a href="#FNanchor_110_110"><span class="label">[110]</span></a> <i>Ibid.</i> p. 947.</p></div>
+
+<div class="footnote"><p><a name="Footnote_111_111" id="Footnote_111_111"></a><a href="#FNanchor_111_111"><span class="label">[111]</span></a> <i>Descent of Man</i>, p. 240.</p></div>
+
+<div class="footnote"><p><a name="Footnote_112_112" id="Footnote_112_112"></a><a href="#FNanchor_112_112"><span class="label">[112]</span></a> "Das geologische Alter der Pithecanthropus-Schichten
+bei Trinil, Ost-Java." <i>Neues Jahrb. f. Mineralogie</i>. Festband, 1907.</p></div>
+
+<div class="footnote"><p><a name="Footnote_113_113" id="Footnote_113_113"></a><a href="#FNanchor_113_113"><span class="label">[113]</span></a> <i>Descent of Man</i>, p. 82.</p></div>
+
+<div class="footnote"><p><a name="Footnote_114_114" id="Footnote_114_114"></a><a href="#FNanchor_114_114"><span class="label">[114]</span></a> "La race humaine de N&eacute;anderthal ou de Canstatt en
+Belgique." <i>Arch. de Biologie</i>, VII. 1887.</p></div>
+
+<div class="footnote"><p><a name="Footnote_115_115" id="Footnote_115_115"></a><a href="#FNanchor_115_115"><span class="label">[115]</span></a> Gorjanovi&#269;-Kramberger. <i>Der diluviale Mensch van
+Krapina in Kroatien</i>, 1906.</p></div>
+
+<div class="footnote"><p><a name="Footnote_116_116" id="Footnote_116_116"></a><a href="#FNanchor_116_116"><span class="label">[116]</span></a> <i>Studien zur Vorgeschichte des Menschen</i>, 1906, pp. 154
+ff.</p></div>
+
+<div class="footnote"><p><a name="Footnote_117_117" id="Footnote_117_117"></a><a href="#FNanchor_117_117"><span class="label">[117]</span></a> "On the cranial and facial characters of the Neandertal
+Race." <i>Trans. R. Soc.</i> London, vol. 199, 1908, p. 281.</p></div>
+
+<div class="footnote"><p><a name="Footnote_118_118" id="Footnote_118_118"></a><a href="#FNanchor_118_118"><span class="label">[118]</span></a> Since this essay was written Schoetensack has
+discovered near Heidelberg and briefly described an exceedingly
+interesting lower jaw from rocks between the Pliocene and Diluvial
+beds. This exhibits interesting differences from the forms of lower
+jaw of <i>Homo primigenius</i>. (Schoetensack, <i>Der Unterkiefer des Homo
+heidelbergensis</i>, Leipzig, 1908.) G. S.</p></div>
+
+<div class="footnote"><p><a name="Footnote_119_119" id="Footnote_119_119"></a><a href="#FNanchor_119_119"><span class="label">[119]</span></a> <i>Life and Letters of Thomas Henry Huxley</i>, Vol. II. p.
+394.</p></div>
+
+<div class="footnote"><p><a name="Footnote_120_120" id="Footnote_120_120"></a><a href="#FNanchor_120_120"><span class="label">[120]</span></a> <i>Descent of Man</i>, p. 229.</p></div>
+
+<div class="footnote"><p><a name="Footnote_121_121" id="Footnote_121_121"></a><a href="#FNanchor_121_121"><span class="label">[121]</span></a> <i>Loc. cit.</i></p></div>
+
+<div class="footnote"><p><a name="Footnote_122_122" id="Footnote_122_122"></a><a href="#FNanchor_122_122"><span class="label">[122]</span></a> Klaatsch in his last publications speaks in the main
+only of an ancestral form common to men and anthropoid apes.</p></div>
+
+<div class="footnote"><p><a name="Footnote_123_123" id="Footnote_123_123"></a><a href="#FNanchor_123_123"><span class="label">[123]</span></a> Haeckels latest genealogical tree is to be found in his
+most recent work, <i>Unsere Ahnenreihe</i>. Jena, 1908.</p></div>
+
+<div class="footnote"><p><a name="Footnote_124_124" id="Footnote_124_124"></a><a href="#FNanchor_124_124"><span class="label">[124]</span></a> Sergi, G. <i>Europa</i>, 1908.</p></div>
+
+<div class="footnote"><p><a name="Footnote_125_125" id="Footnote_125_125"></a><a href="#FNanchor_125_125"><span class="label">[125]</span></a> <i>See</i> Ameghino's latest paper, "<i>Notas preliminaries
+sobre el Tetraprothomo argentinus</i>," etc. <i>Anales del Museo nacional
+de Buenos Aires</i>, <span class="smcap">xvi</span>. pp. 107-242, 1907.</p></div>
+
+<div class="footnote"><p><a name="Footnote_126_126" id="Footnote_126_126"></a><a href="#FNanchor_126_126"><span class="label">[126]</span></a> "Nouvelles recherches sur la formation pamp&eacute;enne et
+l'homme fossile de la R&eacute;publique Argentine." <i>Rivista del Museo de la
+Plata</i>, T. <span class="smcap">xiv</span>. pp. 193-488.</p></div>
+</div>
+
+
+<hr style="width: 65%;" />
+<p><span class='pagenum'><a name="Page_146" id="Page_146">[146]</a></span></p>
+<h2><a name="V" id="V"></a>V</h2>
+
+<h2>CHARLES DARWIN AS AN ANTHROPOLOGIST</h2>
+<p class="center"><span class="smcap1">By Ernst Haeckel</span></p>
+<h4><i>Professor of Zoology in the University of Jena</i></h4>
+
+<p>The great advance that anthropology has made in the second half of the
+nineteenth century is due, in the first place, to Darwin's discovery
+of the origin of man. No other problem in the whole field of research
+is so momentous as that of "Man's place in nature," which was justly
+described by Huxley (1863) as the most fundamental of all questions.
+Yet the scientific solution of this problem was impossible until the
+theory of descent had been established.</p>
+
+<p>It is now a hundred years since the great French biologist Jean
+Lamarck published his <i>Philosophie Zoologique</i>. By a remarkable
+coincidence the year in which that work was issued, 1809, was the year
+of the birth of his most distinguished successor, Charles Darwin.
+Lamarck had already recognised that the descent of man from a series
+of other Vertebrates&mdash;that is, from a series of Ape-like Primates&mdash;was
+essentially involved in the general theory of transformation which he
+had erected on a broad inductive basis; and he had sufficient
+penetration to detect the agencies that had been at work in the
+evolution of the erect bimanous man from the arboreal and quadrumanous
+ape. He had, however, few empirical arguments to advance in support of
+his hypothesis, and it could not be established until the further
+development of the biological sciences&mdash;the founding of comparative
+embryology by Baer (1828) and of the cell-<span class='pagenum'><a name="Page_147" id="Page_147">[147]</a></span>theory by Schleiden and
+Schwann (1838), the advance of physiology under Johannes M&uuml;ller
+(1833), and the enormous progress of palaeontology and comparative
+anatomy between 1820 and 1860&mdash;provided this necessary foundation.
+Darwin was the first to coordinate the ample results of these lines of
+research. With no less comprehensiveness than discrimination he
+consolidated them as a basis of a modified theory of descent, and
+associated with them his own theory of natural selection, which we
+take to be distinctive of "Darwinism" in the stricter sense. The
+illuminating truth of these cumulative arguments was so great in every
+branch of biology that, in spite of the most vehement opposition, the
+battle was won within a single decade, and Darwin secured the general
+admiration and recognition that had been denied to his forerunner,
+Lamarck, up to the hour of his death (1829).</p>
+
+<p>Before, however, we consider the momentous influence that Darwinism
+has had in anthropology, we shall find it useful to glance at its
+history in the course of the last half century, and notice the various
+theories that have contributed to its advance. The first attempt to
+give extensive expression to the reform of biology by Darwin's work
+will be found in my <i>Generelle Morphologie</i> (1866)<a name="FNanchor_127_127" id="FNanchor_127_127"></a><a href="#Footnote_127_127" class="fnanchor">[127]</a> which was
+followed by a more popular treatment of the subject in my <i>Nat&uuml;rliche
+Sch&ouml;pfungsgeschichte</i> (1868),<a name="FNanchor_128_128" id="FNanchor_128_128"></a><a href="#Footnote_128_128" class="fnanchor">[128]</a> a compilation from the earlier
+work. In the first volume of the <i>Generelle Morphologie</i> I endeavoured
+to show the great importance of evolution in settling the fundamental
+questions of biological philosophy, especially in regard to
+comparative anatomy. In the second volume I dealt broadly with the
+principle of evolution, distinguishing ontogeny and phylogeny as its
+two coordinate main branches, and associating the two in the
+Biogenetic Law. The Law may be formulated thus: "Ontogeny (embryology
+or the development of the individual) is a concise and compressed
+recapitulation of phylogeny <span class='pagenum'><a name="Page_148" id="Page_148">[148]</a></span>(the palaeontological or genealogical
+series) conditioned by laws of heredity and adaptation." The
+"Systematic introduction to general evolution," with which the second
+volume of the <i>Generelle Morphologie</i> opens, was the first attempt to
+draw up a natural system of organisms (in harmony with the principles
+of Lamarck and Darwin) in the form of a hypothetical pedigree, and was
+provisionally set forth in eight genealogical tables.</p>
+
+<p>In the nineteenth chapter of the <i>Generelle Morphologie</i>&mdash;a part of
+which has been republished, without any alteration, after a lapse of
+forty years&mdash;I made a critical study of Lamarck's theory of descent
+and of Darwin's theory of selection, and endeavoured to bring the
+complex phenomena of heredity and adaptation under definite laws for
+the first time. Heredity I divided into conservative and progressive:
+adaptation into indirect (or potential) and direct (or actual). I then
+found it possible to give some explanation of the correlation of the
+two physiological functions in the struggle for life (selection), and
+to indicate the important laws of divergence (or differentiation) and
+complexity (or division of labor), which are the direct and inevitable
+outcome of selection. Finally, I marked off dysteleology as the
+science of the aimless (vestigial, abortive, atrophied, and useless)
+organs and parts of the body. In all this I worked from a strictly
+monistic standpoint, and sought to explain all biological phenomena on
+the mechanical and naturalistic lines that had long been recognised in
+the study of inorganic nature. Then (1866), as now, being convinced of
+the unity of nature, the fundamental identity of the agencies at work
+in the inorganic and the organic worlds, I discarded vitalism,
+teleology, and all hypotheses of a mystic character.</p>
+
+<p>It was clear from the first that it was essential, in the monistic
+conception of evolution, to distinguish between the laws of
+conservative and progressive heredity. Conservative heredity maintains
+from generation to generation the enduring characters of the species.
+Each organism transmits to its descendants a part of the morphological
+and<span class='pagenum'><a name="Page_149" id="Page_149">[149]</a></span> physiological qualities that it has received from its parents and
+ancestors. On the other hand, progressive heredity brings new
+characters to the species&mdash;characters that were not found in preceding
+generations. Each organism may transmit to its offspring a part of the
+morphological and physiological features that it has itself acquired,
+by adaptation, in the course of its individual career, through the use
+or disuse of particular organs, the influence of environment, climate,
+nutrition, etc. At that time I gave the name of "progressive heredity"
+to this inheritance of acquired characters, as a short and convenient
+expression, but have since changed the term to "transformative
+heredity" (as distinguished from conservative). This term is
+preferable, as inherited regressive modifications (degeneration,
+retrograde metamorphosis, etc.) come under the same head.</p>
+
+<p>Transformative heredity&mdash;or the transmission of acquired
+characters&mdash;is one of the most important principles in evolutionary
+science. Unless we admit it most of the facts of comparative anatomy
+and physiology are inexplicable. That was the conviction of Darwin no
+less than of Lamarck, of Spencer as well as Virchow, of Huxley as well
+as Gegenbaur, indeed of the great majority of speculative biologists.
+This fundamental principle was for the first time called in question
+and assailed in 1885 by August Weismann of Freiburg, the eminent
+zoologist to whom the theory of evolution owes a great deal of
+valuable support, and who has attained distinction by his extension of
+the theory of selection. In explanation of the phenomena of heredity
+he introduced a new theory, the "theory of the continuity of the
+germ-plasm." According to him the living substance in all organisms
+consists of two quite distinct kinds of plasm, somatic and germinal.
+The permanent germ-plasm, or the active substance of the two
+germ-cells (egg-cell and sperm-cell), passes unchanged through a
+series of generations, and is not affected by environmental
+influences. The environment modifies only the soma-plasm, the organs
+and tissues of the body. The modifications that these parts undergo
+through<span class='pagenum'><a name="Page_150" id="Page_150">[150]</a></span> the influence of the environment or their own activity (use
+and habit), do not affect the germ-plasm, and cannot therefore be
+transmitted.</p>
+
+<p>This theory of the continuity of the germ-plasm has been expounded by
+Weismann during the last twenty-four years in a number of able
+volumes, and is regarded by many biologists, such as Mr. Francis
+Galton, Sir E. Ray Lankester, and Professor J. Arthur Thomson (who has
+recently made a thorough-going defence of it in his important work
+<i>Heredity</i>),<a name="FNanchor_129_129" id="FNanchor_129_129"></a><a href="#Footnote_129_129" class="fnanchor">[129]</a> as the most striking advance in evolutionary
+science. On the other hand, the theory has been rejected by Herbert
+Spencer, Sir W. Turner, Gegenbaur, K&ouml;lliker, Hertwig, and many others.
+For my part I have, with all respect for the distinguished Darwinian,
+contested the theory from the first, because its whole foundation
+seems to me erroneous, and its deductions do not seem to be in accord
+with the main facts of comparative morphology and physiology.
+Weismann's theory in its entirety is a finely conceived molecular
+hypothesis, but it is devoid of empirical basis. The notion of the
+absolute and permanent independence of the germ-plasm, as
+distinguished from the soma-plasm, is purely speculative; as is also
+the theory of germinal selection. The determinants, ids, and idants,
+are purely hypothetical elements. The experiments that have been
+devised to demonstrate their existence really prove nothing.</p>
+
+<p>It seems to me quite improper to describe this hypothetical structure
+as "Neodarwinism." Darwin was just as convinced as Lamarck of the
+transmission of acquired characters and its great importance in the
+scheme of evolution. I had the good fortune to visit Darwin at Down
+three times and discuss with him the main principles of his system,
+and on each occasion we were fully agreed as to the incalculable
+importance of what I may call transformative inheritance. It is only
+proper to point out that Weismann's theory of the germ-plasm is in
+express contradiction to the fundamental principles of Darwin and
+Lamarck. Nor is it more acceptable <span class='pagenum'><a name="Page_151" id="Page_151">[151]</a></span>in what one may call its
+"ultradarwinism"&mdash;the idea that the theory of selection explains
+everything in the evolution of the organic world. This belief in the
+"omnipotence of natural selection" was not shared by Darwin himself.
+Assuredly, I regard it as of the utmost value, as the process of
+natural selection through the struggle for life affords an explanation
+of the mechanical origin of the adapted organisation. It solves the
+great problem: how could the finely adapted structure of the animal or
+plant body be formed unless it was built on a preconceived plan? It
+thus enables us to dispense with the teleology of the metaphysician
+and the dualist, and to set aside the old mythological and poetic
+legends of creation. The idea had occurred in vague form to the great
+Empedocles 2000 years before the time of Darwin, but it was reserved
+for modern research to give it ample expression. Nevertheless, natural
+selection does not of itself give the solution of all our evolutionary
+problems. It has to be taken in conjunction with the transformism of
+Lamarck, with which it is in complete harmony.</p>
+
+<p>The monumental greatness of Charles Darwin, who surpasses every other
+student of science in the nineteenth century by the loftiness of his
+monistic conception of nature and the progressive influence of his
+ideas, is perhaps best seen in the fact that not one of his many
+successors has succeeded in modifying his theory of descent in any
+essential point or in discovering an entirely new standpoint in the
+interpretation of the organic world. Neither N&auml;geli nor Weismann,
+neither De Vries nor Roux, has done this. N&auml;geli, in his
+<i>Mechanisch-Physiologische Theorie der Abstammungslehre</i><a name="FNanchor_130_130" id="FNanchor_130_130"></a><a href="#Footnote_130_130" class="fnanchor">[130]</a> which is
+to a great extent in agreement with Weismann, constructed a theory of
+the idioplasm, that represents it (like the germ-plasm) as developing
+continuously in a definite direction from internal causes. But his
+internal "principle of progress" is at the bottom just as teleological
+as the vital force of the Vitalists, and the micella <span class='pagenum'><a name="Page_152" id="Page_152">[152]</a></span>structure of the
+idioplasm is just as hypothetical as the "dominant" structure of the
+germ-plasm. In 1889 Moritz Wagner sought to explain the origin of
+species by migration and isolation, and on that basis constructed a
+special "migration-theory." This, however, is not out of harmony with
+the theory of selection. It merely elevates one single factor in the
+theory to a predominant position. Isolation is only a special case of
+selection, as I had pointed out in the fifteenth chapter of my
+<i>Natural history of creation</i>. The "mutation-theory" of De Vries,<a name="FNanchor_131_131" id="FNanchor_131_131"></a><a href="#Footnote_131_131" class="fnanchor">[131]</a>
+that would explain the origin of species by sudden and saltatory
+variations rather than by gradual modification, is regarded by many
+botanists as a great step in advance, but it is generally rejected by
+zoologists. It affords no explanation of the facts of adaptation, and
+has no causal value.</p>
+
+<p>Much more important than these theories is that of Wilhelm Roux<a name="FNanchor_132_132" id="FNanchor_132_132"></a><a href="#Footnote_132_132" class="fnanchor">[132]</a>
+of "the struggle of parts within the organism, a supplementation of
+the theory of mechanical adaptation." He explains the functional
+autoformation of the purposive structure by a combination of Darwin's
+principle of selection with Lamarck's idea of transformative heredity,
+and applies the two in conjunction to the facts of histology. He lays
+stress on the significance of functional adaptation, which I had
+described in 1866, under the head of cumulative adaptation, as the
+most important factor in evolution. Pointing out its influence in the
+cell-life of the tissues, he puts "cellular selection" above "personal
+selection," and shows how the finest conceivable adaptations in the
+structure of the tissue may be brought about quite mechanically,
+without preconceived plan. This "mechanical teleology" is a valuable
+extension of Darwin's monistic principle of selection to the whole
+field of cellular physiology and histology, and is wholly destructive
+of dualistic vitalism.</p>
+
+<p>The most important advance that evolution has made since Darwin and
+the most valuable amplification of his <span class='pagenum'><a name="Page_153" id="Page_153">[153]</a></span>theory of selection is, in my
+opinion, the work of Richard Semon: <i>Die Mneme als erhaltendes Prinzip
+im Wechsel des organischen Geschehens</i>.<a name="FNanchor_133_133" id="FNanchor_133_133"></a><a href="#Footnote_133_133" class="fnanchor">[133]</a> He offers a psychological
+explanation of the facts of heredity by reducing them to a process of
+(unconscious) memory. The physiologist Ewald Hering had shown in 1870
+that memory must be regarded as a general function of organic matter,
+and that we are quite unable to explain the chief vital phenomena,
+especially those of reproduction and inheritance, unless we admit this
+unconscious memory. In my essay <i>Die Perigenesis der Plastidule</i><a name="FNanchor_134_134" id="FNanchor_134_134"></a><a href="#Footnote_134_134" class="fnanchor">[134]</a>
+I elaborated this far-reaching idea, and applied the physical
+principle of transmitted motion to the plastidules, or active
+molecules of plasm. I concluded that "heredity is the memory of the
+plastidules, and variability their power of comprehension." This
+"provisional attempt to give a mechanical explanation of the
+elementary processes of evolution" I afterwards extended by showing
+that sensitiveness is (as Carl N&auml;geli, Ernst Mach, and Albrecht Rau
+express it) a general quality of matter. This form of panpsychism
+finds its simplest expression in the "trinity of substance."</p>
+
+<p>To the two fundamental attributes that Spinoza ascribed to
+substance&mdash;Extension (matter as occupying space) and Cogitation
+(energy, force)&mdash;we now add the third fundamental quality of Psychoma
+(sensitiveness, soul). I further elaborated this trinitarian
+conception of substance in the nineteenth chapter of my <i>Die
+Lebenswunder</i> (1904),<a name="FNanchor_135_135" id="FNanchor_135_135"></a><a href="#Footnote_135_135" class="fnanchor">[135]</a> and it seems to me well calculated to
+afford a monistic solution of many of the antitheses of philosophy.</p>
+
+<p>This important Mneme-theory of Semon and the luminous physiological
+experiments and observations associated with it not only throw
+considerable light on transformative inheritance, but provide a sound
+physiological foundation for <span class='pagenum'><a name="Page_154" id="Page_154">[154]</a></span>the biogenetic law. I had endeavoured to
+show in 1874, in the first chapter of my <i>Anthropogenie</i>,<a name="FNanchor_136_136" id="FNanchor_136_136"></a><a href="#Footnote_136_136" class="fnanchor">[136]</a> that
+this fundamental law of organic evolution holds good generally, and
+that there is everywhere a direct causal connection between ontogeny
+and phylogeny. "Phylogenesis is the mechanical cause of ontogenesis;"
+in other words, "The evolution of the stem or race is&mdash;in accordance
+with the laws of heredity and adaptation&mdash;the real cause of all the
+changes that appear, in a condensed form, in the development of the
+individual organism from the ovum, in either the embryo or the larva."</p>
+
+<p>It is now fifty years since Charles Darwin pointed out, in the
+thirteenth chapter of his epoch-making <i>Origin of Species</i>, the
+fundamental importance of embryology in connection with his theory of
+descent:</p>
+
+<p>"The leading facts in embryology, which are second to none in
+importance, are explained on the principle of variations in the many
+descendants from some one ancient progenitor, having appeared at a not
+very early period of life, and having been inherited at a
+corresponding period."<a name="FNanchor_137_137" id="FNanchor_137_137"></a><a href="#Footnote_137_137" class="fnanchor">[137]</a></p>
+
+<p>He then shows that the striking resemblance of the embryos and larvae
+of closely related animals, which in the mature stage belong to widely
+different species and genera, can only be explained by their descent
+from a common progenitor. Fritz M&uuml;ller made a closer study of these
+important phenomena in the instructive instance of the Crustacean
+larva, as given in his able work <i>F&uuml;r Darwin</i><a name="FNanchor_138_138" id="FNanchor_138_138"></a><a href="#Footnote_138_138" class="fnanchor">[138]</a> (1864). I then, in
+1872, extended the range so as to include all animals (with the
+exception of the unicellular Protozoa) and showed, by means of the
+theory of the Gastraea, that all multicellular, tissue-forming
+animals&mdash;all the Metazoa&mdash;develop in essentially the same way from the
+primary germ-layers.</p>
+
+<p><span class='pagenum'><a name="Page_155" id="Page_155">[155]</a></span></p><p>I conceived the embryonic form, in which the whole structure consists
+of only two layers of cells, and is known as the gastrula, to be the
+ontogenetic recapitulation, maintained by tenacious heredity, of a
+primitive common progenitor of all the Metazoa, the Gastraea. At a
+later date (1895) Monticelli discovered that this conjectural
+ancestral form is still preserved in certain primitive
+Coelenterata&mdash;Pemmatodiscus, Kunstleria, and the nearly-related
+Orthonectida.</p>
+
+<p>The general application of the biogenetic law to all classes of
+animals and plants has been proved in my <i>Systematische
+Phylogenie</i>.<a name="FNanchor_139_139" id="FNanchor_139_139"></a><a href="#Footnote_139_139" class="fnanchor">[139]</a> It has, however, been frequently challenged, both by
+botanists and zoologists, chiefly owing to the fact that many have
+failed to distinguish its two essential elements, palingenesis and
+cenogenesis. As early as 1874 I had emphasised, in the first chapter
+of my <i>Evolution of Man</i>, the importance of discriminating carefully
+between these two sets of phenomena:</p>
+
+<p>"In the evolutionary appreciation of the facts of embryology we must
+take particular care to distinguish sharply and clearly between the
+primary, palingenetic evolutionary processes and the secondary,
+cenogenetic processes. The palingenetic phenomena, or embryonic
+<i>recapitulations</i>, are due to heredity, to the transmission of
+characters from one generation to another. They enable us to draw
+direct inferences in regard to corresponding structures in the
+development of the species (e.g. the chorda or the branchial arches in
+all vertebrate embryos). The cenogenetic phenomena, on the other hand,
+or the embryonic <i>variations</i>, cannot be traced to inheritance from a
+mature ancestor, but are due to the adaption of the embryo or the
+larva to certain conditions of its individual development (e.g. the
+amnion, the allantois, and the vitelline arteries in the embryos of
+the higher vertebrates). These cenogenetic phenomena are later
+additions; we must not infer from them <span class='pagenum'><a name="Page_156" id="Page_156">[156]</a></span>that there were corresponding
+processes in the ancestral history, and hence they are apt to
+mislead."</p>
+
+<p>The fundamental importance of these facts of comparative anatomy,
+atavism, and the rudimentary organs, was pointed out by Darwin in the
+first part of his classic work, <i>The Descent of Man and Selection in
+Relation to Sex</i> (1871).<a name="FNanchor_140_140" id="FNanchor_140_140"></a><a href="#Footnote_140_140" class="fnanchor">[140]</a> In the "General summary and conclusion"
+(chap. xxi.) he was able to say, with perfect justice: "He who is not
+content to look, like a savage, at the phenomena of nature as
+disconnected, cannot any longer believe that man is the work of a
+separate act of creation. He will be forced to admit that the close
+resemblance of the embryo of man to that, for instance, of a dog&mdash;the
+construction of his skull, limbs, and whole frame on the same plan
+with that of other mammals, independently of the uses to which the
+parts may be put&mdash;the occasional reappearance of various structures,
+for instance of several muscles, which man does not normally possess,
+but which are common to the Quadrumana&mdash;and a crowd of analogous
+facts&mdash;all point in the plainest manner to the conclusion that man is
+the co-descendant with other mammals of a common progenitor."</p>
+
+<p>These few lines of Darwin's have a greater scientific value than
+hundreds of those so-called "anthropological treatises," which give
+detailed descriptions of single organs, or mathematical tables with
+series of numbers and what are claimed to be "exact analyses," but are
+devoid of synoptic conclusions and a philosophical spirit.</p>
+
+<p>Charles Darwin is not generally recognised as a great anthropologist,
+nor does the school of modern anthropologists regard him as a leading
+authority. In Germany, especially, the great majority of the members
+of the anthropological societies took up an attitude of hostility to
+him from the very beginning of the controversy in 1860. <i>The Descent
+of Man</i> was not merely rejected, but even the discussion of it was
+forbidden on the ground that it was "unscientific."</p>
+
+<p>The centre of this inveterate hostility for thirty years&mdash;especially
+<span class='pagenum'><a name="Page_157" id="Page_157">[157]</a></span>after 1877&mdash;was Rudolph Virchow of Berlin, the leading investigator
+in pathological anatomy, who did so much for the reform of medicine by
+his establishment of cellular pathology in 1858. As a prominent
+representative of "exact" or "descriptive" anthropology, and lacking a
+broad equipment in comparative anatomy and ontogeny, he was unable to
+accept the theory of descent. In earlier years, and especially during
+his splendid period of activity at W&uuml;rzburg (1848-1856), he had been a
+consistent free-thinker, and had in a number of able articles
+(collected in his <i>Gesammelte Abhandlungen</i>)<a name="FNanchor_141_141" id="FNanchor_141_141"></a><a href="#Footnote_141_141" class="fnanchor">[141]</a> upheld the unity of
+human nature, the inseparability of body and spirit. In later years at
+Berlin, where he was more occupied with political work and sociology
+(especially after 1866), he abandoned the positive monistic position
+for one of agnosticism and scepticism, and made concessions to the
+dualistic dogma of a spiritual world apart from the material frame.</p>
+
+<p>In the course of a Scientific Congress at Munich in 1877 the conflict
+of these antithetic views of nature came into sharp relief. At this
+memorable Congress I had undertaken to deliver the first address
+(September 18th) on the subject of "Modern evolution in relation to
+the whole of science." I maintained that Darwin's theory not only
+solved the great problem of the origin of species, but that its
+implications, especially in regard to the nature of man, threw
+considerable light on the whole of science, and on anthropology in
+particular. The discovery of the real origin of man by evolution from
+a long series of mammal ancestors threw light on his place in nature
+in every respect, as Huxley had already shown in his excellent
+lectures of 1863. Just as all the organs and tissues of the human body
+had originated from those of the nearest related mammals, certain
+ape-like forms, so we were bound to conclude that his mental qualities
+also had been derived from those of his extinct primate ancestor. <span class='pagenum'><a name="Page_158" id="Page_158">[158]</a></span></p>
+
+<p>This monistic view of the origin and nature of man, which is now
+admitted by nearly all who have the requisite acquaintance with
+biology, and approach the subject without prejudice, encountered a
+sharp opposition at that time. The opposition found its strongest
+expression in an address that Virchow delivered at Munich four days
+afterwards (September 22nd), on "The freedom of science in the modern
+State." He spoke of the theory of evolution as an unproved hypothesis,
+and declared that it ought not to be taught in the schools, because it
+was dangerous to the State. "We must not," he said, "teach that man
+has descended from the ape or any other animal." When Darwin, usually
+so lenient in his judgment, read the English translation of Virchow's
+speech, he expressed his disapproval in strong terms. But the great
+authority that Virchow had&mdash;an authority well founded in pathology and
+sociology&mdash;and his prestige as president of the German Anthropological
+Society, had the effect of preventing any member of the Society from
+raising serious opposition to him for thirty years. Numbers of
+journals and treatises repeated his dogmatic statement: "It is quite
+certain that man has descended neither from the ape nor from any other
+animal." In this he persisted till his death in 1902. Since that time
+the whole position of German anthropology has changed. The question is
+no longer whether man was created by a distinct supernatural act or
+evolved from other mammals, but to which line of the animal hierarchy
+we must look for the actual series of ancestors. The interested reader
+will find an account of this "battle of Munich" (1877) in my three
+Berlin lectures (April, 1905), <i>Der Kampf um die
+Entwickelungs-Gedanken</i>.<a name="FNanchor_142_142" id="FNanchor_142_142"></a><a href="#Footnote_142_142" class="fnanchor">[142]</a></p>
+
+<p>The main points in our genealogical tree were clearly recognised by
+Darwin in the sixth chapter of the <i>Descent of Man</i>. Lowly organised
+fishes, like the lancelot (Amphioxus), are descended from lower
+invertebrates resembling the larvae of an existing Tunicate
+(Appendicularia). From these <span class='pagenum'><a name="Page_159" id="Page_159">[159]</a></span>primitive fishes were evolved higher
+fishes of the ganoid type and others of the type of Lepidosiren
+(Dipneusta). It is a very small step from these to the Amphibia:</p>
+
+<p>"In the class of animals the steps are not difficult to conceive which
+led from the ancient Monotremata to the ancient Marsupials; and from
+these to the early progenitors of the placental mammals. We may thus
+ascend to the Lemuridae; and the interval is not very wide from these
+to the Simiadae. The Simiadae then branched off into two great stems,
+the New World and Old World monkeys; and from the latter, at a remote
+period, Man, the wonder and glory of the Universe, proceeded."<a name="FNanchor_143_143" id="FNanchor_143_143"></a><a href="#Footnote_143_143" class="fnanchor">[143]</a></p>
+
+<p>In these few lines Darwin clearly indicated the way in which we were
+to conceive our ancestral series within the vertebrates. It is fully
+confirmed by all the arguments of comparative anatomy and embryology,
+of palaeontology and physiology; and all the research of the
+subsequent forty years have gone to establish it. The deep interest in
+geology which Darwin maintained throughout his life and his complete
+knowledge of palaeontology enabled him to grasp the fundamental
+importance of the palaeontological record more clearly than
+anthropologists and zoologists usually do.</p>
+
+<p>There has been much debate in subsequent decades whether Darwin
+himself maintained that man was descended from the ape, and many
+writers have sought to deny it. But the lines I have quoted <i>verbatim</i>
+from the conclusion of the sixth chapter of the <i>Descent of Man</i>
+(1871) leave no doubt that he was as firmly convinced of it as was his
+great precursor Jean Lamarck in 1809. Moreover, Darwin adds, with
+particular explicitness, in the "general summary and conclusion"
+(chap. xxi.) of that standard work:<a name="FNanchor_144_144" id="FNanchor_144_144"></a><a href="#Footnote_144_144" class="fnanchor">[144]</a></p>
+
+<p>"By considering the embryological structure of man&mdash;the homologies
+which he presents with the lower animals,&mdash;the rudiments which he
+retains,&mdash;and the reversions to which he is liable, we can partly
+recall in imagination the <span class='pagenum'><a name="Page_160" id="Page_160">[160]</a></span>former condition of our early progenitors;
+and can approximately place them in their proper place in the
+zoological series. We thus learn that man is descended from a hairy,
+tailed quadruped, probably arboreal in its habits, and an inhabitant
+of the Old World. This creature, if its whole structure had been
+examined by a naturalist, would have been classed amongst the
+Quadrumana, as surely as the still more ancient progenitor of the Old
+and New World monkeys."</p>
+
+<p>These clear and definite lines leave no doubt that Darwin&mdash;so critical
+and cautious in regard to important conclusions&mdash;was quite as firmly
+convinced of the descent of man from the apes (the Catarrhinae, in
+particular) as Lamarck was in 1809 and Huxley in 1863.</p>
+
+<p>It is to be noted particularly that, in these and other observations
+on the subject, Darwin decidedly assumes the monophyletic origin of
+the mammals, including man. It is my own conviction that this is of
+the greatest importance. A number of difficult questions in regard to
+the development of man, in respect of anatomy, physiology, psychology,
+and embryology, are easily settled if we do not merely extend our
+<i>progonotaxis</i> to our nearest relatives, the anthropoid apes and the
+tailed monkeys from which these have descended, but go further back
+and find an ancestor in the group of the Lemuridae, and still further
+back to the Marsupials and Monotremata. The essential identity of all
+the Mammals in point of anatomical structure and embryonic
+development&mdash;in spite of their astonishing differences in external
+appearance and habits of life&mdash;is so palpably significant that modern
+zoologists are agreed in the hypothesis that they have all sprung from
+a common root, and that this root may be sought in the earlier
+Palaeozoic Amphibia.</p>
+
+<p>The fundamental importance of this comparative morphology of the
+Mammals, as a sound basis of scientific anthropology, was recognised
+just before the beginning of the nineteenth century, when Lamarck
+first emphasised (1794) the division of the animal kingdom into
+Vertebrates and In<span class='pagenum'><a name="Page_161" id="Page_161">[161]</a></span>vertebrates. Even thirteen years earlier (1781),
+when Goethe made a close study of the mammal skeleton in the
+Anatomical Institute at Jena, he was intensely interested to find that
+the composition of the skull was the same in man as in the other
+mammals. His discovery of the <i>os inter-maxillare</i> in man (1784),
+which was contradicted by most of the anatomists of the time, and his
+ingenious "vertebral theory of the skull," were the splendid fruit of
+his morphological studies. They remind us how Germany's greatest
+philosopher and poet was for many years ardently absorbed in the
+comparative anatomy of man and the mammals, and how he divined that
+their wonderful identity in structure was no mere superficial
+resemblance, but pointed to a deep internal connection. In my
+<i>Generelle Morphologie</i> (1866), in which I published the first
+attempts to construct phylogenetic trees, I have given a number of
+remarkable theses of Goethe, which may be called "phyletic
+prophecies." They justify us in regarding him as a precursor of
+Darwin.</p>
+
+<p>In the ensuing forty years I have made many conscientious efforts to
+penetrate further along that line of anthropological research that was
+opened up by Goethe, Lamarck, and Darwin. I have brought together the
+many valuable results that have constantly been reached in comparative
+anatomy, physiology, ontogeny, and palaeontology, and maintained the
+effort to reform the classification of animals and plants in an
+evolutionary sense. The first rough drafts of pedigrees that were
+published in the <i>Generelle Morphologie</i> have been improved time after
+time in the ten editions of my <i>Nat&uuml;rlich Sch&ouml;pfungsgeschichte</i>
+(1868-1902).<a name="FNanchor_145_145" id="FNanchor_145_145"></a><a href="#Footnote_145_145" class="fnanchor">[145]</a> A sounded basis for my phyletic hypotheses, derived
+from a discriminating combination of the three great
+records&mdash;morphology, ontogeny, and palaeontology&mdash;was provided in the
+three volumes of my <i>Systematische Phylogenie</i><a name="FNanchor_146_146" id="FNanchor_146_146"></a><a href="#Footnote_146_146" class="fnanchor">[146]</a> (1894 Protists and
+Plants, 1895 Vertebrates, 1896 Invertebrates).</p>
+
+<p><span class='pagenum'><a name="Page_162" id="Page_162">[162]</a></span></p><p>In my <i>Anthropogenie</i><a name="FNanchor_147_147" id="FNanchor_147_147"></a><a href="#Footnote_147_147" class="fnanchor">[147]</a> I endeavoured to employ all the known
+facts of comparative ontogeny (embryology) for the purpose of
+completing my scheme of human phylogeny (evolution). I attempted to
+sketch the historical development of each organ of the body, beginning
+with the most elementary structures in the germ-layers of the
+Gastraea. At the same time I drew up a corrected statement of the most
+important steps in the line of our ancestral series.</p>
+
+<p>At the fourth International Congress of Zoology at Cambridge (August
+26th, 1898) I delivered an address on "Our present knowledge of the
+Descent of Man." It was translated into English, enriched with many
+valuable notes and additions, by my friend and pupil in earlier days
+Dr. Hans Gadow (Cambridge), and published under the title: <i>The Last
+Link: our present knowledge of the Descent of Man</i><a name="FNanchor_148_148" id="FNanchor_148_148"></a><a href="#Footnote_148_148" class="fnanchor">[148]</a> The
+determination of the chief animal forms that occur in the line of our
+ancestry is there restricted to thirty types, and these are
+distributed in six main groups.</p>
+
+<p>The first half of this "Progonotaxis hominis," which has no support
+from fossil evidence, comprises three groups: (i) Protista
+(unicellular organisms, 1-5): (ii) Invertebrate Metazoa (Coelenteria
+6-8, Vermalia 9-11): (iii) Monorrhine Vertebrates (Acrania 12-13,
+Cyclostoma 14-15). The second half, which is based on fossil records,
+also comprises three groups: (iv) Palaeozoic cold-blooded Craniota
+(Fishes 16-18, Amphibia 19, Reptiles 20): (v) Mesozoic Mammals
+(Monotrema 21, Marsupialia 22, Mallotheria 23): (vi) Cenozoic Primates
+(Lemuridae 24-25, Tailed Apes 26-27, Anthropomorpha 28-30). An
+improved and enlarged edition of this hypothetic "Progonotaxis
+hominis" was published in 1908, in my essay <i>Unsere Ahnenreihe</i>.<a name="FNanchor_149_149" id="FNanchor_149_149"></a><a href="#Footnote_149_149" class="fnanchor">[149]</a></p>
+
+<p>If I have succeeded in furthering, in some degree, by these
+<span class='pagenum'><a name="Page_163" id="Page_163">[163]</a></span>anthropological works, the solution of the great problem of Man's
+place in nature, and particularly in helping to trace the definite
+stages in our ancestral series, I owe the success, not merely to the
+vast progress that biology has made in the last half century, but
+largely to the luminous example of the great investigators who have
+applied themselves to the problem, with so much assiduity and genius,
+for a century and a quarter&mdash;I mean Goethe and Lamarck, Gegenbaur and
+Huxley, but, above all, Charles Darwin. It was the great genius of
+Darwin that first brought together that symmetrical temple of
+scientific knowledge, the theory of descent. It was Darwin who put the
+crown on the edifice by his theory of natural selection. Not until
+this broad inductive law was firmly established was it possible to
+vindicate the special conclusion, the descent of man from a series of
+other Vertebrates. By his illuminating discovery Darwin did more for
+anthropology than thousands of those writers, who are more
+specifically titled anthropologists, have done by their technical
+treatises. We may, indeed, say that it is not merely as an exact
+observer and ingenious experimenter, but as a distinguished
+anthropologist and far-seeing thinker, that Darwin takes his place
+among the greatest men of science of the nineteenth century.</p>
+
+<p>To appreciate fully the immortal merit of Darwin in connection with
+anthropology, we must remember that not only did his chief work, <i>The
+Origin of Species</i>, which opened up a new era in natural history in
+1859, sustain the most virulent and widespread opposition for a
+lengthy period, but even thirty years later, when its principles were
+generally recognised and adopted, the application of them to man was
+energetically contested by many high scientific authorities. Even
+Alfred Russel Wallace, who discovered the principle of natural
+selection independently in 1858, did not concede that it was
+applicable to the higher mental and moral qualities of man. Dr.
+Wallace still holds a spiritualist and dualist view of the nature of
+man, contending that he is composed of a material frame (descended
+from the apes)<span class='pagenum'><a name="Page_164" id="Page_164">[164]</a></span> and an immortal immaterial soul (infused by a higher
+power). This dual conception, moreover, is still predominant in the
+wide circles of modern theology and metaphysics, and has the general
+and influential adherence of the more conservative classes of society.</p>
+
+<p>In strict contradiction to this mystical dualism, which is generally
+connected with teleology and vitalism, Darwin always maintained the
+complete unity of human nature, and showed convincingly that the
+psychological side of man was developed, in the same way as the body,
+from the less advanced soul of the anthropoid ape, and, at a still
+more remote period, from the cerebral functions of the older
+vertebrates. The eighth chapter of the <i>Origin of Species</i>, which is
+devoted to instinct, contains weighty evidence that the instincts of
+animals are subject, like all other vital processes, to the general
+laws of historic development. The special instincts of particular
+species were formed by adaptation, and the modifications thus acquired
+were handed on to posterity by heredity; in their formation and
+preservation natural selection plays the same part as in the
+transformation of every other physiological function. The higher moral
+qualities of civilised man have been derived from the lower mental
+functions of the uncultivated barbarians and savages, and these in
+turn from the social instincts of the mammals. This natural and
+monistic psychology of Darwin's was afterwards more fully developed by
+his friend George Romanes in his excellent works <i>Mental Evolution in
+Animals</i> and <i>Mental Evolution in Man</i>.<a name="FNanchor_150_150" id="FNanchor_150_150"></a><a href="#Footnote_150_150" class="fnanchor">[150]</a></p>
+
+<p>Many valuable and most interesting contributions to this monistic
+psychology of man were made by Darwin in his fine work on <i>The Descent
+of Man and Selection in Relation to Sex</i>, and again in his
+supplementary work, <i>The Expression of the Emotions in Man and
+Animals</i>. To understand the historical development of Darwin's
+anthropology one must read his life and the introduction to <i>The
+Descent of Man</i>. From the moment that he was convinced of the <span class='pagenum'><a name="Page_165" id="Page_165">[165]</a></span>truth
+of the principle of descent&mdash;that is to say, from his thirtieth year,
+in 1838&mdash;he recognised clearly that man could not be excluded from its
+range. He recognised as a logical necessity the important conclusion
+that "man is the co-descendant with other species of some ancient,
+lower, and extinct form." For many years he gathered notes and
+arguments in support of this thesis, and for the purpose of showing
+the probable line of man's ancestry. But in the first edition of <i>The
+Origin of Species</i> (1859) he restricted himself to the single line,
+that by this work "light would be thrown on the origin of man and his
+history." In the fifty years that have elapsed since that time the
+science of the origin and nature of man has made astonishing progress,
+and we are now fairly agreed in a monistic conception of nature that
+regards the whole universe, including man, as a wonderful unity,
+governed by unalterable and eternal laws. In my philosophical book
+<i>Die Weltr&auml;tsel</i> (1899)<a name="FNanchor_151_151" id="FNanchor_151_151"></a><a href="#Footnote_151_151" class="fnanchor">[151]</a> and in the supplementary volume <i>Die
+Lebenswunder</i> (1904)<a name="FNanchor_152_152" id="FNanchor_152_152"></a><a href="#Footnote_152_152" class="fnanchor">[152]</a> I have endeavoured to show that this pure
+monism is securely established, and that the admission of the
+all-powerful rule of the same principle of evolution throughout the
+universe compels us to formulate a single supreme law&mdash;the
+all-embracing "Law of Substance," or the united laws of the constancy
+of matter and the conservation of energy. We should never have reached
+this supreme general conception if Charles Darwin&mdash;a "monistic
+philosopher" in the true sense of the word&mdash;had not prepared the way
+by his theory of descent by natural selection, and crowned the great
+work of his life by the association of this theory with a naturalistic
+anthropology.</p>
+
+<div class="footnotes"><h3>FOOTNOTES:</h3>
+
+<div class="footnote"><p><a name="Footnote_127_127" id="Footnote_127_127"></a><a href="#FNanchor_127_127"><span class="label">[127]</span></a> <i>Generelle Morphologie der Organismen</i>, 2 vols.,
+Berlin, 1866.</p></div>
+
+<div class="footnote"><p><a name="Footnote_128_128" id="Footnote_128_128"></a><a href="#FNanchor_128_128"><span class="label">[128]</span></a> Eng. transl.; <i>The History of Creation</i>, London, 1876.</p></div>
+
+<div class="footnote"><p><a name="Footnote_129_129" id="Footnote_129_129"></a><a href="#FNanchor_129_129"><span class="label">[129]</span></a> London, 1908.</p></div>
+
+<div class="footnote"><p><a name="Footnote_130_130" id="Footnote_130_130"></a><a href="#FNanchor_130_130"><span class="label">[130]</span></a> Munich, 1884.</p></div>
+
+<div class="footnote"><p><a name="Footnote_131_131" id="Footnote_131_131"></a><a href="#FNanchor_131_131"><span class="label">[131]</span></a> <i>Die Mutationstheorie</i>, Leipzig, 1903.</p></div>
+
+<div class="footnote"><p><a name="Footnote_132_132" id="Footnote_132_132"></a><a href="#FNanchor_132_132"><span class="label">[132]</span></a> <i>Der Kampf der Theile im Organismus</i>, Leipzig, 1881.</p></div>
+
+<div class="footnote"><p><a name="Footnote_133_133" id="Footnote_133_133"></a><a href="#FNanchor_133_133"><span class="label">[133]</span></a> Leipzig, 1904.</p></div>
+
+<div class="footnote"><p><a name="Footnote_134_134" id="Footnote_134_134"></a><a href="#FNanchor_134_134"><span class="label">[134]</span></a> Berlin, 1876.</p></div>
+
+<div class="footnote"><p><a name="Footnote_135_135" id="Footnote_135_135"></a><a href="#FNanchor_135_135"><span class="label">[135]</span></a> <i>Wonders of Life</i>, London and New York, 1904.</p></div>
+
+<div class="footnote"><p><a name="Footnote_136_136" id="Footnote_136_136"></a><a href="#FNanchor_136_136"><span class="label">[136]</span></a> Eng. transl.; <i>The Evolution of Man</i>, 2 vols., London,
+1879 and 1905.</p></div>
+
+<div class="footnote"><p><a name="Footnote_137_137" id="Footnote_137_137"></a><a href="#FNanchor_137_137"><span class="label">[137]</span></a> <i>Origin of Species</i> (6th edit.), p. 396.</p></div>
+
+<div class="footnote"><p><a name="Footnote_138_138" id="Footnote_138_138"></a><a href="#FNanchor_138_138"><span class="label">[138]</span></a> Eng. transl.; <i>Facts and Arguments for Darwin</i>, London,
+1869.</p></div>
+
+<div class="footnote"><p><a name="Footnote_139_139" id="Footnote_139_139"></a><a href="#FNanchor_139_139"><span class="label">[139]</span></a> 3 vols., Berlin, 1894-96.</p></div>
+
+<div class="footnote"><p><a name="Footnote_140_140" id="Footnote_140_140"></a><a href="#FNanchor_140_140"><span class="label">[140]</span></a> <i>Descent of Man</i> (Popular Edit.), p. 927.</p></div>
+
+<div class="footnote"><p><a name="Footnote_141_141" id="Footnote_141_141"></a><a href="#FNanchor_141_141"><span class="label">[141]</span></a> <i>Gesammelte Abhandlungen zur wissenschaftlichen
+Medizin</i>, Berlin, 1856.</p></div>
+
+<div class="footnote"><p><a name="Footnote_142_142" id="Footnote_142_142"></a><a href="#FNanchor_142_142"><span class="label">[142]</span></a> Eng. transl.; <i>Last Words on Evolution</i>, London, 1906.</p></div>
+
+<div class="footnote"><p><a name="Footnote_143_143" id="Footnote_143_143"></a><a href="#FNanchor_143_143"><span class="label">[143]</span></a> <i>Descent of Man</i>, (Popular Edit.), p. 255.</p></div>
+
+<div class="footnote"><p><a name="Footnote_144_144" id="Footnote_144_144"></a><a href="#FNanchor_144_144"><span class="label">[144]</span></a> <i>Descent of Man</i>, p. 930.</p></div>
+
+<div class="footnote"><p><a name="Footnote_145_145" id="Footnote_145_145"></a><a href="#FNanchor_145_145"><span class="label">[145]</span></a> Eng. transl.; <i>The History of Creation</i>, London, 1876.</p></div>
+
+<div class="footnote"><p><a name="Footnote_146_146" id="Footnote_146_146"></a><a href="#FNanchor_146_146"><span class="label">[146]</span></a> Berlin, 1894-96.</p></div>
+
+<div class="footnote"><p><a name="Footnote_147_147" id="Footnote_147_147"></a><a href="#FNanchor_147_147"><span class="label">[147]</span></a> Leipzig, 1874, 5th edit. 1905. Eng. transl.; <i>The
+Evolution of Man</i>, London, 1905.</p></div>
+
+<div class="footnote"><p><a name="Footnote_148_148" id="Footnote_148_148"></a><a href="#FNanchor_148_148"><span class="label">[148]</span></a> London, 1898.</p></div>
+
+<div class="footnote"><p><a name="Footnote_149_149" id="Footnote_149_149"></a><a href="#FNanchor_149_149"><span class="label">[149]</span></a> <i>Festschrift zur 350-j&auml;hrigen Jubelfeier der Th&uuml;ringer
+Universit&auml;t Jena</i>. Jena. 1908.</p></div>
+
+<div class="footnote"><p><a name="Footnote_150_150" id="Footnote_150_150"></a><a href="#FNanchor_150_150"><span class="label">[150]</span></a> London, 1885; 1888.</p></div>
+
+<div class="footnote"><p><a name="Footnote_151_151" id="Footnote_151_151"></a><a href="#FNanchor_151_151"><span class="label">[151]</span></a> <i>The Riddle of the Universe</i>, London and New York,
+1900.</p></div>
+
+<div class="footnote"><p><a name="Footnote_152_152" id="Footnote_152_152"></a><a href="#FNanchor_152_152"><span class="label">[152]</span></a> <i>The Wonders of Life</i>, London and New York, 1904.</p></div>
+</div>
+
+
+<hr style="width: 65%;" />
+<p><span class='pagenum'><a name="Page_166" id="Page_166">[166]</a></span></p>
+<h2><a name="VI" id="VI"></a>VI</h2>
+
+<h2>MENTAL FACTORS IN EVOLUTION</h2>
+<p class="center"><span class="smcap1">By C. Lloyd Morgan, LLD., F.R.S</span></p>
+
+<p>In developing his conception of organic evolution Charles Darwin was
+of necessity brought into contact with some of the problems of mental
+evolution. In <i>The Origin of Species</i> he devoted a chapter to "the
+diversities of instinct and of the other mental faculties in animals
+of the same class."<a name="FNanchor_153_153" id="FNanchor_153_153"></a><a href="#Footnote_153_153" class="fnanchor">[153]</a> When he passed to the detailed consideration
+of <i>The Descent of Man</i>, it was part of his object to show "that there
+is no fundamental difference between man and the higher mammals in
+their mental faculties."<a name="FNanchor_154_154" id="FNanchor_154_154"></a><a href="#Footnote_154_154" class="fnanchor">[154]</a> "If no organic being excepting man," he
+said, "had possessed any mental power, or if his powers had been of a
+wholly different nature, from those of the lower animals, then we
+should never have been able to convince ourselves that our high
+faculties had been gradually developed."<a name="FNanchor_155_155" id="FNanchor_155_155"></a><a href="#Footnote_155_155" class="fnanchor">[155]</a> In his discussion of
+<i>The Expression of the Emotions</i> it was important for his purpose
+"fully to recognise that actions readily become associated with other
+actions and with various states of the mind."<a name="FNanchor_156_156" id="FNanchor_156_156"></a><a href="#Footnote_156_156" class="fnanchor">[156]</a> His hypothesis of
+sexual selection is largely dependent upon the <span class='pagenum'><a name="Page_167" id="Page_167">[167]</a></span>exercise of choice on
+the part of the female and her preference for "not only the more
+attractive but at the same time the more vigourous and vicious
+males."<a name="FNanchor_157_157" id="FNanchor_157_157"></a><a href="#Footnote_157_157" class="fnanchor">[157]</a> Mental processes and physiological processes were for
+Darwin closely correlated; and he accepted the conclusion "that the
+nervous system not only regulates most of the existing functions of
+the body, but has indirectly influenced the progressive development of
+various bodily structures and of certain mental qualities."<a name="FNanchor_158_158" id="FNanchor_158_158"></a><a href="#Footnote_158_158" class="fnanchor">[158]</a></p>
+
+<p>Throughout his treatment, mental evolution was for Darwin incidental
+to and contributory to organic evolution. For specialised research in
+comparative and genetic psychology, as an independent field of
+investigation, he had neither the time nor the requisite training.
+None the less his writings and the spirit of his work have exercised a
+profound influence on this department of evolutionary thought. And,
+for those who follow Darwin's lead, mental evolution is still in a
+measure subservient to organic evolution. Mental processes are the
+accompaniments or concomitants of the functional activity of specially
+differentiated parts of the organism. They are in some way dependent
+on physiological and physical conditions. But though they are not
+physical in their nature, and though it is difficult or impossible to
+conceive that they are physical in their origin, they are, for Darwin
+and his followers, factors in the evolutionary process in its physical
+or organic aspect. By the physiologist within his special and
+well-defined universe of discourse they may be properly regarded as
+epiphenomena; but by the naturalist in his more catholic survey of
+nature they cannot be so regarded, and were not so regarded by Darwin.
+Intelligence has contributed to evolution of which it is in a sense a
+product.</p>
+
+<p>The facts of observation or of inference which Darwin accepted are
+these: Conscious experience accompanies some <span class='pagenum'><a name="Page_168" id="Page_168">[168]</a></span>of the modes of animal
+behaviour; it is concomitant with certain physiological processes;
+these processes are the outcome of development in the individual and
+evolution in the race; the accompanying mental processes undergo a
+like development. Into the subtle philosophical questions which arise
+out of the na&iuml;ve acceptance of such a creed it was not Darwin's
+province to enter; "I have nothing to do," he said,<a name="FNanchor_159_159" id="FNanchor_159_159"></a><a href="#Footnote_159_159" class="fnanchor">[159]</a> "with the
+origin of the mental powers, any more than I have with that of life
+itself." He dealt with the natural history of organisms, including not
+only their structure but their modes of behaviour; with the natural
+history of the states of consciousness which accompany some of their
+actions; and with the relation of behaviour to experience. We will
+endeavour to follow Darwin in his modesty and candour in making no
+pretence to give ultimate explanations. But we must note one of the
+implications of this self-denying ordinance of science. Development
+and evolution imply continuity. For Darwin and his followers the
+continuity is organic through physical heredity. Apart from
+speculative hypothesis, legitimate enough in its proper place but here
+out of court, we know nothing of continuity of mental evolution as
+such: consciousness appears afresh in each succeeding generation.
+Hence it is that for those who follow Darwin's lead, mental evolution
+is and must ever be, within his universe of discourse, subservient to
+organic evolution. Only in so far as conscious experience, or its
+neural correlate, effects some changes in organic structure can it
+influence the course of heredity; and conversely only in so far as
+changes in organic structure are transmitted through heredity, is
+mental evolution rendered possible. Such is the logical outcome of
+Darwin's teaching.</p>
+
+<p>Those who abide by the cardinal results of this teaching are bound to
+regard all behaviour as the expression of the functional activities of
+the living tissues of the organism, and all conscious experience as
+correlated with such activities. For the purposes of scientific
+treatment, mental processes <span class='pagenum'><a name="Page_169" id="Page_169">[169]</a></span>are one mode of expression of the same
+changes of which the physiological processes accompanying behaviour
+are another mode of expression. This is simply accepted as a fact
+which others may seek to explain. The behaviour itself is the adaptive
+application of the energies of the organism; it is called forth by
+some form of presentation or stimulation brought to bear on the
+organism by the environment. This presentation is always an individual
+or personal matter. But in order that the organism may be fitted to
+respond to the presentation of the environment it must have undergone
+in some way a suitable preparation. According to the theory of
+evolution this preparation is primarily racial and is transmitted
+through heredity. Darwin's main thesis was that the method of
+preparation is predominantly by natural selection. Subordinate to
+racial preparation, and always dependent thereon, is individual or
+personal preparation through some kind of acquisition; of which the
+guidance of behaviour through individually won experience is a typical
+example. We here introduce the mental factor because the facts seem to
+justify the inference. Thus there are some modes of behaviour which
+are wholly and solely dependent upon inherited racial preparation;
+there are other modes of behaviour which are also dependent, in part
+at least, on individual preparation. In the former case the behaviour
+is adaptive on the first occurrence of the appropriate presentation;
+in the latter case accommodation to circumstances is only reached
+after a greater or less amount of acquired organic modification of
+structure, often accompanied (as we assume) in the higher animals by
+acquired experience. Logically and biologically the two classes of
+behaviour are clearly distinguishable: but the analysis of complex
+cases of behaviour where the two factors co&ouml;perate, is difficult and
+requires careful and critical study of life-history.</p>
+
+<p>The foundations of the mental life are laid in the conscious
+experience that accompanies those modes of behaviour, dependent
+entirely on racial preparation, which may broadly be described as
+instinctive. In the eighth chapter of <i>The<span class='pagenum'><a name="Page_170" id="Page_170">[170]</a></span> Origin of Species</i> Darwin
+says,<a name="FNanchor_160_160" id="FNanchor_160_160"></a><a href="#Footnote_160_160" class="fnanchor">[160]</a> "I will not attempt any definition of instinct.... Every
+one understands what is meant, when it is said that instinct impels
+the cuckoo to migrate and to lay her eggs in other birds' nests. An
+action, which we ourselves require experience to enable us to perform,
+when performed by an animal, more especially by a very young one,
+without experience, and when performed by many individuals in the same
+way, without their knowing for what purpose it is performed, is
+usually said to be instinctive." And in the summary at the close of
+the chapter he says,<a name="FNanchor_161_161" id="FNanchor_161_161"></a><a href="#Footnote_161_161" class="fnanchor">[161]</a> "I have endeavoured briefly to show that the
+mental qualities of our domestic animals vary, and that the variations
+are inherited. Still more briefly I have attempted to show that
+instincts vary slightly in a state of nature. No one will dispute that
+instincts are of the highest importance to each animal. Therefore
+there is no real difficulty, under changing conditions of life, in
+natural selection accumulating to any extent slight modifications of
+instinct which are in any way useful. In many cases habit or use and
+disuse have probably come into play."</p>
+
+<p>Into the details of Darwin's treatment there is neither space nor need
+to enter. There are some ambiguous passages; but it may be said that
+for him, as for his followers to-day, instinctive behaviour is wholly
+the result of racial preparation transmitted through organic heredity.
+For the performance of the instinctive act no individual preparation
+under the guidance of personal experience is necessary. It is true
+that Darwin quotes with approval Huber's saying that "a little dose of
+judgment or reason often comes into play, even with animals low in the
+scale of nature."<a name="FNanchor_162_162" id="FNanchor_162_162"></a><a href="#Footnote_162_162" class="fnanchor">[162]</a> But we may fairly interpret his meaning to be
+that in behaviour, which is commonly called instinctive, some element
+of intelligent guidance is often combined. If this be conceded the
+<span class='pagenum'><a name="Page_171" id="Page_171">[171]</a></span>strictly instinctive performance (or part of the performance) is the
+outcome of heredity and due to the direct transmission of parental or
+ancestral aptitudes. Hence the instinctive response as such depends
+entirely on how the nervous mechanism has been built up through
+heredity; while intelligent behaviour, or the intelligent factor in
+behaviour, depends also on how the nervous mechanism has been modified
+and moulded by use during its development and concurrently with the
+growth of individual experience in the customary situations of daily
+life. Of course it is essential to the Darwinian thesis that what Sir
+E. Ray Lankester has termed "educability," not less than instinct, is
+hereditary. But it is also essential to the understanding of this
+thesis that the differentiae of the hereditary factor should be
+clearly grasped.</p>
+
+<p>For Darwin there were two modes of racial preparation, (<i>1</i>) natural
+selection, and (<i>2</i>) the establishment of individually acquired habit.
+He showed that instincts are subject to hereditary variation; he saw
+that instincts are also subject to modification through acquisition in
+the course of individual life. He believed that not only the
+variations but also, to some extent, the modifications are inherited.
+He therefore held that some instincts (the greater number) are due to
+natural selection but that others (less numerous) are due, or partly
+due, to the inheritance of acquired habits. The latter involve
+Lamarckian inheritance, which of late years has been the centre of so
+much controversy. It is noteworthy however that Darwin laid especial
+emphasis on the fact that many of the most typical and also the most
+complex instincts&mdash;those of neuter insects&mdash;do not admit of such an
+interpretation. "I am surprised," he says,<a name="FNanchor_163_163" id="FNanchor_163_163"></a><a href="#Footnote_163_163" class="fnanchor">[163]</a> "that no one has
+hitherto advanced this demonstrative case of neuter insects, against
+the well-known doctrine of inherited habit, as advanced by Lamarck."
+None the less Darwin admitted this doctrine as supplementary to that
+which was more distinctively his own&mdash;for example in the case of the
+<span class='pagenum'><a name="Page_172" id="Page_172">[172]</a></span>instincts of domesticated animals. Still, even in such cases, "it may
+be doubted," he says,<a name="FNanchor_164_164" id="FNanchor_164_164"></a><a href="#Footnote_164_164" class="fnanchor">[164]</a> "whether any one would have thought of
+training a dog to point, had not some one dog naturally shown a
+tendency in this line ... so that habit and some degree of selection
+have probably concurred in civilising by inheritance our dogs." But in
+the interpretation of the instincts of domesticated animals, a more
+recently suggested hypothesis, that of organic selection,<a name="FNanchor_165_165" id="FNanchor_165_165"></a><a href="#Footnote_165_165" class="fnanchor">[165]</a> may be
+helpful. According to this hypothesis any intelligent modification of
+behaviour which is subject to selection is probably coincident in
+direction with an inherited tendency to behave in this fashion. Hence
+in such behaviour there are two factors: (1) an incipient variation in
+the line of such behaviour, and (2) an acquired modification by which
+the behaviour is carried further along the same line. Under natural
+selection those organisms in which the two factors co&ouml;perate are
+likely to survive. Under artificial selection they are deliberately
+chosen out from among the rest.</p>
+
+<p>Organic selection has been termed a compromise between the more
+strictly Darwinian and the Lamarckian principles of interpretation.
+But it is not in any sense a compromise. The principle of
+interpretation of that which is instinctive and hereditary is wholly
+Darwinian. It is true that some of the facts of observation relied
+upon by Lamarckians are introduced. For Lamarckians however the
+modifications which are admittedly factors in survival, are regarded
+as the parents of inherited variations; for believers in organic
+selection they are only the foster-parents or nurses. It is because
+organic selection is the direct outcome of and a natural extension of
+Darwin's cardinal thesis that some reference to it here is
+justifiable. The matter may be put with the utmost brevity as follows:
+(1) Variations (V) occur, some of which are in the direction of
+increased adaptation (+), others in the direction of decreased
+adaptation (-).</p>
+
+<p><span class='pagenum'><a name="Page_173" id="Page_173">[173]</a></span></p><p>(2) Acquired modifications (M) also occur. Some of these are in the
+direction of increased accommodation to circumstances (+), while
+others are in the direction of diminished accommodation (-). Four
+major combinations are</p>
+<table summary="" class="tb1">
+  <tr>
+    <td>(<i>b</i>) + V with - M,</td>
+    <td>
+    (<i>c</i>) - V with + M,</td>
+  </tr>
+  <tr>
+    <td>(<i>a</i>) + V with + M,</td>
+    <td>(<i>d</i>) - V with - M.</td>
+  </tr>
+</table>
+
+
+
+<p>Of these (<i>d</i>) must inevitably be eliminated while (<i>a</i>) are selected.
+The predominant survival of (<i>a</i>) entails the survival of the adaptive
+variations which are inherited. The contributory acquisitions (+ M)
+are not inherited; but there are none the less factors in determining
+the survival of the coincident variations. It is surely abundantly
+clear that this is Darwinism and has no tincture of Lamarck's
+essential principle, the inheritance of acquired characters.</p>
+
+<p>Whether Darwin himself would have accepted this interpretation of some
+at least of the evidence put forward by Lamarckians is unfortunately a
+matter of conjecture. The fact remains that in his interpretation of
+instinct and in allied questions he accepted the inheritance of
+individually acquired modifications of behaviour and structure.</p>
+
+<p>Darwin was chiefly concerned with instinct from the biological rather
+than from the psychological point of view. Indeed it must be confessed
+that, from the latter standpoint, his conception of instinct as a
+"mental faculty" which "impels" an animal to the performance of
+certain actions, scarcely affords a satisfactory basis for genetic
+treatment. To carry out the spirit of Darwin's teaching it is
+necessary to link more closely biological and psychological evolution.
+The first step towards this is to interpret the phenomena of
+instinctive behaviour in terms of stimulation and response. It may be
+well to take a particular case. Swimming on the part of a duckling is,
+from the biological point of view, a typical example of instinctive
+behaviour. Gently lower a recently hatched bird into water:
+coordinated movements of the limbs follow in rhythmical sequence. The
+behaviour is new to the individual though it is no doubt closely
+related<span class='pagenum'><a name="Page_174" id="Page_174">[174]</a></span> to that of walking, which is no less instinctive. There is a
+group of stimuli afforded by the "presentation" which results from
+partial immersion: upon this there follows as a complex response an
+application of the functional activities in swimming; the sequence of
+adaptive application on the appropriate presentation is determined by
+racial preparation. We know, it is true, but little of the
+physiological details of what takes place in the central nervous
+system; but in broad outline the nature of the organic mechanism and
+the manner of its functioning may at least be provisionally
+conjectured in the present state of physiological knowledge. Similarly
+in the case of the pecking of newly-hatched chicks; there is a visual
+presentation, there is probably a co&ouml;perating group of stimuli from
+the alimentary tract in need of food, there is an adaptive application
+of the activities in a definite mode of behaviour. Like data are
+afforded in a great number of cases of instinctive procedure,
+sometimes occurring very early in life, not infrequently deferred
+until the organism is more fully developed, but all of them dependent
+upon racial preparation. No doubt there is some range of variation in
+the behaviour, just such variation as the theory of natural selection
+demands. But there can be no question that the higher animals inherit
+a bodily organisation and a nervous system, the functional working of
+which gives rise to those inherited modes of behaviour which are
+termed instinctive.</p>
+
+<p>It is to be noted that the term "instinctive" is here employed in the
+adjectival form as a descriptive heading under which may be grouped
+many and various modes of behaviour due to racial preparation. We
+speak of these as inherited; but in strictness what is transmitted
+through heredity is the complex of anatomical and physiological
+conditions under which, in appropriate circumstances, the organism so
+behaves. So far the term "instinctive" has a restricted biological
+connotation in terms of behaviour. But the connecting link between
+biological evolution and psychological evolution is to be sought,&mdash;as
+Darwin fully realised,&mdash;in<span class='pagenum'><a name="Page_175" id="Page_175">[175]</a></span> the phenomena of instinct, broadly
+considered. The term "instinctive" has also a psychological
+connotation. What is that connotation?</p>
+
+<p>Let us take the case of the swimming duckling or the pecking chick,
+and fix our attention on the first instinctive performance. Grant that
+just as there is, strictly speaking, no inherited behaviour, but only
+the conditions which render such behaviour under appropriate
+circumstances possible; so too there is no inherited experience, but
+only the conditions which render such experience possible; then the
+cerebral conditions in both cases are the same. The biological
+behaviour-complex, including the total stimulation and the total
+response with the intervening or resultant processes in the sensorium,
+is accompanied by an experience-complex including the initial
+stimulation-consciousness and resulting response-consciousness. In the
+experience-complex are comprised data which in psychological analysis
+are grouped under the headings of cognition, affective tone and
+conation. But the complex is probably experienced as an unanalysed
+whole. If then we use the term "instinctive" so as to comprise all
+congenital modes of behaviour which contribute to experience, we are
+in a position to grasp the view that the net result in consciousness
+constitutes what we may term the primary tissue of experience. To the
+development of this experience each instinctive act contributes. The
+nature and manner of organisation of this primary tissue of experience
+are dependent on inherited biological aptitudes; but they are from the
+outset onwards subject to secondary development dependent on acquired
+aptitudes. Biological values are supplemented by psychological values
+in terms of satisfaction or the reverse.</p>
+
+<p>In our study of instinct we have to select some particular phase of
+animal behaviour and isolate it so far as is possible from the life of
+which it is a part. But the animal is a going concern, restlessly
+active in many ways. Many instinctive performances, as Darwin pointed
+out,<a name="FNanchor_166_166" id="FNanchor_166_166"></a><a href="#Footnote_166_166" class="fnanchor">[166]</a> are serial <span class='pagenum'><a name="Page_176" id="Page_176">[176]</a></span>in their nature. But the whole of active life is
+a serial and coordinated business. The particular instinctive
+performance is only an episode in a life-history, and every mode of
+behaviour is more or less closely correlated with other modes. This
+coordination of behaviour is accompanied by a correlation of the modes
+of primary experience. We may classify the instinctive modes of
+behaviour and their accompanying modes of instinctive experience under
+as many heads as may be convenient for our purposes of interpretation,
+and label them instincts of self-preservation, of pugnacity, of
+acquisition, the reproductive instincts, the parental instincts, and
+so forth. An instinct, in this sense of the term (for example the
+parental instinct), may be described as a specialised part of the
+primary tissue of experience differentiated in relation to some
+definite biological end. Under such an instinct will fall a large
+number of particular and often well-defined modes of behaviour, each
+with its own peculiar mode of experience.</p>
+
+<p>It is no doubt exceedingly difficult as a matter of observation and of
+inference securely based thereon to distinguish what is primary from
+what is in part due to secondary acquisition&mdash;a fact which Darwin
+fully appreciated. Animals are educable in different degrees; but
+where they are educable they begin to profit by experience from the
+first. Only, therefore, on the occasion of the first instinctive act
+of a given type can the experience gained be regarded as <i>wholly</i>
+primary; all subsequent performance is liable to be in some degree,
+sometimes more, sometimes less, modified by the acquired disposition
+which the initial behaviour engenders. But the early stages of
+acquisition are always along the lines predetermined by instinctive
+differentiation. It is the task of comparative psychology to
+distinguish the primary tissue of experience from its secondary and
+acquired modifications. We cannot follow up the matter in further
+detail. It must here suffice to suggest that this conception of
+instinct as a primary form of experience lends itself better to
+natural history treatment than Darwin's conception of<span class='pagenum'><a name="Page_177" id="Page_177">[177]</a></span> an impelling
+force, and that it is in line with the main trend of Darwin's thought.</p>
+
+<p>In a characteristic work,&mdash;characteristic in wealth of detail, in
+closeness and fidelity of observation, in breadth of outlook, in
+candour and modesty,&mdash;Darwin dealt with <i>The Expression of the
+Emotions in Man and Animals</i>. Sir Charles Bell in his <i>Anatomy of
+Expression</i> had contended that many of man's facial muscles had been
+specially created for the sole purpose of being instrumental in the
+expression of his emotions. Darwin claimed that a natural explanation,
+consistent with the doctrine of evolution, could in many cases be
+given and would in other cases be afforded by an extension of the
+principles he advocated. "No doubt," he said,<a name="FNanchor_167_167" id="FNanchor_167_167"></a><a href="#Footnote_167_167" class="fnanchor">[167]</a> "as long as man and
+all other animals are viewed as independent creations, an effectual
+stop is put to our natural desire to investigate as far as possible
+the causes of Expression. By this doctrine, anything and everything
+can be equally well explained.... With mankind, some expressions ...
+can hardly be understood, except on the belief that man once existed
+in a much lower and animal-like condition. The community of certain
+expressions in distinct though allied species ... is rendered somewhat
+more intelligible, if we believe in their descent from a common
+progenitor. He who admits on general grounds that the structure and
+habits of all animals have been gradually evolved, will look at the
+whole subject of Expression in a new and interesting light."</p>
+
+<p>Darwin relied on three principles of explanation. "The first of these
+principles is, that movements which are serviceable in gratifying some
+desire, or in relieving some sensation, if often repeated, become so
+habitual that they are performed, whether or not of any service,
+whenever the same desire or sensation is felt, even in a very weak
+degree."<a name="FNanchor_168_168" id="FNanchor_168_168"></a><a href="#Footnote_168_168" class="fnanchor">[168]</a> The modes of expression which fall under this head have
+<span class='pagenum'><a name="Page_178" id="Page_178">[178]</a></span>become instinctive through the hereditary transmission of acquired
+habit. "As far as we can judge, only a few expressive movements are
+learnt by each individual; that is, were consciously and voluntarily
+performed during the early years of life for some definite object, or
+in imitation of others, and then became habitual. The far greater
+number of the movements of expression, and all the more important
+ones, are innate or inherited; and such cannot be said to depend on
+the will of the individual. Nevertheless, all those included under our
+first principle were at first voluntarily performed for a definite
+object,&mdash;namely, to escape some danger, to relieve some distress, or
+to gratify some desire."<a name="FNanchor_169_169" id="FNanchor_169_169"></a><a href="#Footnote_169_169" class="fnanchor">[169]</a></p>
+
+<p>"Our second principle is that of antithesis. The habit of voluntarily
+performing opposite movements under opposite impulses has become
+firmly established in us by the practice of our whole lives. Hence, if
+certain actions have been regularly performed, in accordance with our
+first principle, under a certain frame of mind, there will be a strong
+and involuntary tendency to the performance of directly opposite
+actions, whether or not these are of any use, under the excitement of
+an opposite frame of mind."<a name="FNanchor_170_170" id="FNanchor_170_170"></a><a href="#Footnote_170_170" class="fnanchor">[170]</a> This principle of antithesis has not
+been widely accepted. Nor is Darwin's own position easy to grasp.</p>
+
+<p>"Our third principle," he says,<a name="FNanchor_171_171" id="FNanchor_171_171"></a><a href="#Footnote_171_171" class="fnanchor">[171]</a> "is the direct action of the
+excited nervous system on the body, independently of the will, and
+independently, in large part, of habit. Experience shows that
+nerve-force is generated and set free whenever the cerebro-spinal
+system is excited. The direction which this nerve-force follows is
+necessarily determined by the lines of connection between the
+nerve-cells, with each other and with various parts of the body."</p>
+
+<p>Lack of space prevents our following up the details of Darwin's
+treatment of expression. Whether we accept or do <span class='pagenum'><a name="Page_179" id="Page_179">[179]</a></span>not accept his three
+principles of explanation we must regard his work as a masterpiece of
+descriptive analysis, packed full of observations possessing lasting
+value. For a further development of the subject it is essential that
+the instinctive factors in expression should be more fully
+distinguished from those which are individually acquired&mdash;a difficult
+task&mdash;and that the instinctive factors should be rediscussed in the
+light of modern doctrines of heredity, with a view to determining
+whether Lamarckian inheritance, on which Darwin so largely relied, is
+necessary for an interpretation of the facts.</p>
+
+<p>The whole subject as Darwin realised is very complex. Even the term
+"expression" has a certain amount of ambiguity. When the emotion is in
+full flood, the animal fights, flees, or faints. Is this full-tide
+effect to be regarded as expression; or are we to restrict the term to
+the premonitory or residual effects&mdash;the bared canine when the
+fighting mood is being roused, the ruffled fur when reminiscent
+representations of the object inducing anger cross the mind? Broadly
+considered both should be included. The activity of premonitory
+expression as a means of communication was recognised by Darwin; he
+might, perhaps, have emphasised it more strongly in dealing with the
+lower animals. Man so largely relies on a special means of
+communication, that of language, that he sometimes fails to realise
+that for animals with their keen powers of perception, and dependent
+as they are on such means of communication, the more strictly
+biological means of expression are full of subtle suggestiveness. Many
+modes of expression, otherwise useless, are signs of behaviour that
+may be anticipated,&mdash;signs which stimulate the appropriate attitude of
+response. This would not, however, serve to account for the utility of
+the organic accompaniments&mdash;heart-affection, respiratory changes,
+vaso-motor effects and so forth, together with heightened muscular
+tone,&mdash;on all of which Darwin lays stress<a name="FNanchor_172_172" id="FNanchor_172_172"></a><a href="#Footnote_172_172" class="fnanchor">[172]</a> under his third
+principle. The biological value of all this is, however, of great
+<span class='pagenum'><a name="Page_180" id="Page_180">[180]</a></span>importance, though Darwin was hardly in a position to take it fully
+into account.</p>
+
+<p>Having regard to the instinctive and hereditary factors of emotional
+expression we may ask whether Darwin's third principle does not alone
+suffice as an explanation. Whether we admit or reject Lamarckian
+inheritance it would appear that all hereditary expression must be due
+to pre-established connections within the central nervous system and
+to a transmitted provision for coordinated response under the
+appropriate stimulation. If this be so, Darwin's first and second
+principles are subordinate and ancillary to the third, an expression,
+so far as it is instinctive or heredity, being "the direct result of
+the constitution of the nervous system."</p>
+
+<p>Darwin accepted the emotions themselves as hereditary or acquired
+states of mind and devoted his attention to their expression. But
+these emotions themselves are genetic products and as such dependent
+on organic conditions. It remained, therefore, for psychologists who
+accepted evolution and sought to build on biological foundations to
+trace the genesis of these modes of animal and human experience. The
+subject has been independently developed by Professors Lange and
+James;<a name="FNanchor_173_173" id="FNanchor_173_173"></a><a href="#Footnote_173_173" class="fnanchor">[173]</a> and some modification of their view is regarded by many
+evolutionists as affording the best explanation of the facts. We must
+fix our attention on the lower emotions, such as anger or fear, and on
+their first occurrence in the life of the individual organism. It is a
+matter of observation that if a group of young birds which have been
+hatched in an incubator are frightened by an appropriate presentation,
+auditory or visual, they instinctively respond in special ways. If we
+speak of this response as the expression, we find that there are many
+factors. There are certain visible modes of behaviour, crouching at
+once, scattering and then crouching, remaining motionless, the braced
+muscles sustaining an attitude of arrest, and so forth, There are also
+certain visceral or organic effects, such as affections <span class='pagenum'><a name="Page_181" id="Page_181">[181]</a></span>of the heart
+and respiration. These can be readily observed by taking the young
+bird in the hand. Other effects cannot be readily observed; vaso-motor
+changes, affections of the alimentary canal, the skin and so forth.
+Now the essence of the James-Lange view, as applied to these
+congenital effects, is that though we are justified in speaking of
+them as effects of the stimulation, we are not justified, without
+further evidence, in speaking of them as effects of the emotional
+state. May it not rather be that the emotion as a primary mode of
+experience is the concomitant of the net result of the organic
+situation&mdash;the initial presentation, the instinctive mode of
+behaviour, the visceral disturbances?</p>
+
+<p>According to this interpretation the primary tissue of experience of
+the emotional order, felt as an unanalysed complex, is generated by
+the stimulation of the sensorium by afferent or incoming physiological
+impulses from the special senses, from the organs concerned in the
+responsive behaviour, from the viscere and vaso-motor system.</p>
+
+<p>Some psychologists, however, contend that the emotional experience is
+generated in the sensorium prior to, and not subsequent to, the
+behaviour-response and the visceral disturbances. It is a direct and
+not an indirect outcome of the presentation to the special senses. Be
+this as it may, there is a growing tendency to bring into the closest
+possible relation, or even to identify, instinct and emotion in their
+primary genesis. The central core of all such interpretations is that
+instinctive behaviour and experience, its emotional accompaniments,
+and its expression, are but different aspects of the outcome of the
+same organic occurrences. Such emotions are, therefore, only a
+distinguishable aspect of the primary tissue of experience and exhibit
+a like differentiation. Here again a biological foundation is laid for
+a psychological doctrine of the mental development of the individual.</p>
+
+<p>The intimate relation between emotion as a psychological mode of
+experience and expression as a group of organic conditions has an
+important bearing on biological interpretation. The emotion, as the
+psychological accompaniment<span class='pagenum'><a name="Page_182" id="Page_182">[182]</a></span> of orderly disturbances in the central
+nervous system, profoundly influences behaviour and often renders it
+more vigourous and more effective. The utility of the emotions in the
+struggle for existence can, therefore, scarcely be over-estimated.
+Just as keenness of perception has survival-value; just as it is
+obviously subject to variation; just as it must be enhanced under
+natural selection, whether individually acquired increments are
+inherited or not; and just as its value lies not only in this or that
+special perceptive act but in its importance for life as a whole; so
+the vigourous effectiveness of activity has survival-value; it is
+subject to variation; it must be enhanced under natural selection; and
+its importance lies not only in particular modes of behaviour but in
+its value for life as a whole. If emotion and its expression as a
+congenital endowment are but different aspects of the same biological
+occurrence; and if this is a powerful supplement to vigour
+effectiveness and persistency of behaviour, it must on Darwin's
+principles be subject to natural selection.</p>
+
+<p>If we include under the expression of the emotions not only the
+premonitory symptoms of the initial phases of the organic and mental
+state, not only the signs or conditions of half-tide emotion, but the
+full-tide manifestation of an emotion which dominates the situation,
+we are naturally led on to the consideration of many of the phenomena
+which are discussed under the head of sexual selection. The subject is
+difficult and complex, and it was treated by Darwin with all the
+strength he could summon to the task. It can only be dealt with here
+from a special point of view&mdash;that which may serve to illustrate the
+influence of certain mental factors on the course of evolution. From
+this point of view too much stress can scarcely be laid on the
+dominance of emotion during the period of courtship and pairing in the
+more highly organised animals. It is a period of maximum vigour,
+maximum activity, and, correlated with special modes of behaviour and
+special organic and visceral accompaniments, a period also of maximum
+emotional excitement. The com<span class='pagenum'><a name="Page_183" id="Page_183">[183]</a></span>bats of males, their dances and aerial
+evolutions, their elaborate behaviour and display, or the flood of
+song in birds, are emotional expressions which are at any rate
+coincident in time with sexual periodicity. From the combat of the
+males there follows on Darwin's principles the elimination of those
+which are deficient in bodily vigour, deficient in special structures,
+offensive or protective, which contribute to success, deficient in the
+emotional supplement of which persistent and whole-hearted fighting is
+the expression, and deficient in alertness and skill which are the
+outcome of the psychological development of the powers of perception.
+Few biologists question that we have here a mode of selection of much
+importance, though its influence on psychological evolution often
+fails to receive its due emphasis. Mr. Wallace<a name="FNanchor_174_174" id="FNanchor_174_174"></a><a href="#Footnote_174_174" class="fnanchor">[174]</a> regards it as "a
+form of natural selection"; "to it," he says, "we must impute the
+development of the exceptional strength, size, and activity of the
+male, together with the possession of special offensive and defensive
+weapons, and of all other characters which arise from the development
+of these or are correlated with them." So far there is little
+disagreement among the followers of Darwin&mdash;for Mr. Wallace, with fine
+magnanimity, has always preferred to be ranked as such,
+notwithstanding his right, on which a smaller man would have
+constantly insisted, to the claim of independent originator of the
+doctrine of natural selection. So far with regard to sexual selection
+Darwin and Mr. Wallace are agreed; so far and no farther. For Darwin,
+says Mr. Wallace,<a name="FNanchor_175_175" id="FNanchor_175_175"></a><a href="#Footnote_175_175" class="fnanchor">[175]</a> "has extended the principle into a totally
+different field of action, which has none of that character of
+constancy and of inevitable result that attaches to natural selection,
+including male rivalry; for by far the larger portion of the
+phenomena, which he endeavours to explain by the direct action of
+sexual selection, can only be so explained on the hypothesis that the
+immediate agency is female choice or preference. It is to this that he
+imputes the origin of <span class='pagenum'><a name="Page_184" id="Page_184">[184]</a></span>all secondary sexual characters other than
+weapons of offence and defence.... In this extension of sexual
+selection to include the action of female choice or preference, and in
+the attempt to give to that choice such wide-reaching effects, I am
+unable to follow him more than a very little way."</p>
+
+<p>Into the details of Mr. Wallace's criticisms it is impossible to enter
+here. We cannot discuss either the mode of origin of the variations in
+structure which have rendered secondary sexual characters possible or
+the modes of selection other than sexual which have rendered them,
+within narrow limits, specifically constant. Mendelism and mutation
+theories may have something to say on the subject when these theories
+have been more fully correlated with the basal principles of
+selection. It is noteworthy that Mr. Wallace says:<a name="FNanchor_176_176" id="FNanchor_176_176"></a><a href="#Footnote_176_176" class="fnanchor">[176]</a> "Besides the
+acquisition of weapons by the male for the purpose of fighting with
+other males, there are some other sexual characters which may have
+been produced by natural selection. Such are the various sounds and
+odours which are peculiar to the male, and which serve as a call to
+the female or as an indication of his presence. These are evidently a
+valuable addition to the means of recognition of the two sexes, and
+are a further indication, that the pairing season has arrived; and the
+production, intensification, and differentiation of these sounds and
+odours are clearly within the power of natural selection. The same
+remark will apply to the peculiar calls of birds, and even to the
+singing of the males." Why the same remark should not apply to their
+colours and adornments is not obvious. What is obvious is that "means
+of recognition" and "indication that the pairing season has arrived"
+are dependent on the perceptive powers of the female who recognises
+and for whom the indication has meaning. The hypothesis of female
+preference, stripped of the aesthetic surplusage which is
+psychologically both unnecessary and unproven, is really only
+different in degree from that which Mr. Wallace admits <span class='pagenum'><a name="Page_185" id="Page_185">[185]</a></span>in principle
+when he says that it is probable that the female is pleased or excited
+by the display.</p>
+
+<p>Let us for our present purpose leave on one side and regard as <i>sub
+judice</i> the question whether the specific details of secondary sexual
+characters are the outcome of female choice. For us the question is
+whether certain psychological accompaniments of the pairing situation
+have influenced the course of evolution and whether these
+psychological accompaniments are themselves the outcome of evolution.
+As a matter of observation, specially differentiated modes of
+behaviour, often very elaborate, frequently requiring highly developed
+skill, and apparently highly charged with emotional tone, are the
+precursors of pairing. They are generally confined to the males, whose
+fierce combats during the period of sexual activity are part of the
+emotional manifestation. It is inconceivable that they have no
+biological meaning; and it is difficult to conceive that they have any
+other biological end than to evoke in the generally more passive
+female the pairing impulse. They, are based on instinctive foundations
+ingrained in the nervous constitution through natural (or may we not
+say sexual?) selection in virtue of their profound utility. They are
+called into play by a specialised presentation such as the sight or
+the scent of the female at, or a little in advance of, a critical
+period of the physiological rhythm. There is no necessity that the
+male should have any knowledge of the end to which his strenuous
+activity leads up. In presence of the female there is an elaborate
+application of all the energies of behaviour, just because ages of
+racial preparation have made him biologically and emotionally what he
+is&mdash;a functionally sexual male that must dance or sing or go through
+hereditary movements of display, when the appropriate stimulation
+comes. Of course after the first successful courtship his future
+behaviour will be in some degree modified by his previous experience.
+No doubt during his first courtship he is gaining the primary data of
+a peculiarly rich experience, instinctive and emotional. But the
+biological foundations of the behaviour of<span class='pagenum'><a name="Page_186" id="Page_186">[186]</a></span> courtship are laid in the
+hereditary coordinations. It would seem that in some cases, not indeed
+in all, perhaps especially in those cases in which secondary sexual
+behaviour is most highly evolved,&mdash;correlative with the ardour of the
+male is a certain amount of reluctance in the female. The pairing act
+on her part only takes place after prolonged stimulation, for
+affording which the behaviour of male courtship is the requisite
+presentation. The most vigourous, defiant and mettlesome male is
+preferred just because he alone affords a contributory stimulation
+adequate to evoke the pairing impulse with its attendant emotional
+tone.</p>
+
+<p>It is true that this places female preference or choice on a much
+lower psychological plane than Darwin in some passages seems to
+contemplate where, for example, he says that the female appreciates
+the display of the male and places to her credit a taste for the
+beautiful. But Darwin himself distinctly states<a name="FNanchor_177_177" id="FNanchor_177_177"></a><a href="#Footnote_177_177" class="fnanchor">[177]</a> that "it is not
+probable that she consciously deliberates; but she is most excited or
+attracted by the most beautiful, or melodious, or gallant males." The
+view here put forward, which has been developed by Prof. Groos,<a name="FNanchor_178_178" id="FNanchor_178_178"></a><a href="#Footnote_178_178" class="fnanchor">[178]</a>
+therefore seems to have Darwin's own sanction. The phenomena are not
+only biological; there are psychological elements as well. One can
+hardly suppose that the female is unconscious of the male's presence;
+the final yielding must surely be accompanied by heightened emotional
+tone. Whether we call it choice or not is merely a matter of
+definition of terms. The behaviour is in part determined by
+supplementary psychological values. Prof. Groos regards the coyness of
+females as "a most efficient means of preventing the too early and too
+frequent yielding to the sexual impulse."<a name="FNanchor_179_179" id="FNanchor_179_179"></a><a href="#Footnote_179_179" class="fnanchor">[179]</a> Be that as it may, it
+is, in any case, if we grant the facts, a means through which male
+sexual behaviour with <span class='pagenum'><a name="Page_187" id="Page_187">[187]</a></span>all its biological and psychological
+implications, is raised to a level otherwise perhaps unattainable by
+natural means, while in the female it affords opportunities for the
+development in the individual and evolution in the race of what we may
+follow Darwin in calling appreciation, if we empty this word of the
+aesthetic implications which have gathered round it in the mental life
+of man.</p>
+
+<p>Regarded from this standpoint of sexual selection, broadly considered,
+has probably been of great importance. The psychological
+accompaniments of the pairing situation have profoundly influenced the
+course of biological evolution and are themselves the outcome of that
+evolution.</p>
+
+<p>Darwin makes only passing reference to those modes of behaviour in
+animals which go by the name of play. "Nothing," he says,<a name="FNanchor_180_180" id="FNanchor_180_180"></a><a href="#Footnote_180_180" class="fnanchor">[180]</a> "is
+more common than for animals to take pleasure in practising whatever
+instinct they follow at other times for some real good." This is one
+of the very numerous cases in which a hint of the master has served to
+stimulate research in his disciples. It was left to Prof. Groos to
+develop this subject on evolutionary lines and to elaborate in a
+masterly manner Darwin's suggestion. "The utility of play," he
+says,<a name="FNanchor_181_181" id="FNanchor_181_181"></a><a href="#Footnote_181_181" class="fnanchor">[181]</a> "is incalculable. This utility consists in the practice and
+exercise it affords for some of the more important duties of
+life,"&mdash;that is to say, for the performance of activities which will
+in adult life be essential to survival. He urges<a name="FNanchor_182_182" id="FNanchor_182_182"></a><a href="#Footnote_182_182" class="fnanchor">[182]</a> that "the play
+of young animals has its origin in the fact that certain very
+important instincts appear at a time when the animal does not
+seriously need them." It is, however, questionable whether any
+instincts appear at a time when they are not needed. And it is
+questionable whether the instinctive and emotional attitude of the
+play-fight, to take one example, can be identified with those which
+accompany fighting in earnest, though no doubt they are <span class='pagenum'><a name="Page_188" id="Page_188">[188]</a></span>closely
+related and have some common factors. It is probable that play, as
+preparatory behaviour, differs in biological detail (as it almost
+certainly does in emotional attributes) from the earnest of after-life
+and that it has been evolved through differentiation and integration
+of the primary tissue of experience, as a preparation through which
+certain essential modes of skill may be acquired&mdash;those animals in
+which the preparatory play-propensity was not inherited in due force
+and requisite amount being subsequently eliminated in the struggle for
+existence. In any case there is little question that Prof. Groos is
+right in basing the play-propensity on instinctive foundations.<a name="FNanchor_183_183" id="FNanchor_183_183"></a><a href="#Footnote_183_183" class="fnanchor">[183]</a>
+None the less, as he contends, the essential biological value of play
+is that it is a means of training the educable nerve-tissue, of
+developing that part of the brain which is modified by experience and
+which thus acquires new characters, of elaborating the secondary
+tissue of experience on the predetermined lines of instinctive
+differentiation and thus furthering the psychological activities which
+are included under the comprehensive term "intelligent."</p>
+
+<p>In <i>The Descent of Man</i> Darwin dealt at some length with intelligence
+and the higher mental faculties.<a name="FNanchor_184_184" id="FNanchor_184_184"></a><a href="#Footnote_184_184" class="fnanchor">[184]</a> His object, he says, is to show
+that there is no fundamental difference between man and the higher
+mammals in their mental faculties; that these faculties are variable
+and the variations tend to be inherited; and that under natural
+selection beneficial variations of all kinds will have been preserved
+and injurious ones eliminated.</p>
+
+<p>Darwin was too good an observer and too honest a man to minimise the
+"enormous difference" between the level of mental attainment of
+civilised man and that reached by any animal. His contention was that
+the difference, great as it is, is one of degree and not of kind. He
+realised that, in the development of the mental faculties of man, new
+factors <span class='pagenum'><a name="Page_189" id="Page_189">[189]</a></span>in evolution have supervened&mdash;factors which play but a
+subordinate and subsidiary part in animal intelligence.
+Intercommunication by means of language, approbation and blame, and
+all that arises out of reflective thought, are but foreshadowed in the
+mental life of animals. Still he contends that these may be explained
+on the doctrine of evolution. He urges<a name="FNanchor_185_185" id="FNanchor_185_185"></a><a href="#Footnote_185_185" class="fnanchor">[185]</a> "that man is variable in
+body and mind; and that the variations are induced, either directly or
+indirectly, by the same general causes, and obey the same general
+laws, as with the lower animals." He correlates mental development
+with the evolution of the brain.<a name="FNanchor_186_186" id="FNanchor_186_186"></a><a href="#Footnote_186_186" class="fnanchor">[186]</a> "As the various mental faculties
+gradually developed themselves, the brain would almost certainly
+become larger. No one, I presume, doubts that the large proportion
+which the size of man's brain bears to his body, compared to the same
+proportion in the gorilla or orang, is closely connected with his
+higher mental powers." "With respect to the lower animals," he
+says,<a name="FNanchor_187_187" id="FNanchor_187_187"></a><a href="#Footnote_187_187" class="fnanchor">[187]</a> "M. E. Lartet,<a name="FNanchor_188_188" id="FNanchor_188_188"></a><a href="#Footnote_188_188" class="fnanchor">[188]</a> by comparing the crania of tertiary and
+recent mammals belonging to the same groups, has come to the
+remarkable conclusion that the brain is generally larger and the
+convolutions are more complex in the more recent form."</p>
+
+<p>Sir E. Ray Lankester has sought to express in the simplest terms the
+implications of the increase in size of the cerebrum. "In what," he
+asks, "does the advantage of a larger cerebral mass consist?" "Man,"
+he replies, "is born with fewer ready-made tricks of the
+nerve-centres&mdash;these performances of an inherited nervous mechanism so
+often called by the ill-defined term 'instincts'&mdash;than are the monkeys
+or any other animal. Correlated with the absence of inherited
+ready-made mechanism, man has a greater capacity of developing in the
+course of his individual growth similar <span class='pagenum'><a name="Page_190" id="Page_190">[190]</a></span>nervous mechanisms (similar
+to but not identical with those of 'instinct') than any other
+animal.... The power of being educated&mdash;'educability' as we may term
+it&mdash;is what man possesses in excess as compared with the apes. I think
+we are justified in forming the hypothesis that it is this
+'educability' which is the correlative of the increased size of the
+cerebrum." There has been natural selection of the more educable
+animals, for "the character which we describe as 'educability' can be
+transmitted, it is a congenital character. But the <i>results</i> of
+education can <i>not</i> be transmitted. In each generation they have to be
+acquired afresh, and with increased 'educability' they are more
+readily acquired and a larger variety of them.... The fact is that
+there is no community between the mechanisms of instinct and the
+mechanisms of intelligence, and that the latter are later in the
+history of the evolution of the brain than the former and can only
+develop in proportion as the former become feeble and defective."<a name="FNanchor_189_189" id="FNanchor_189_189"></a><a href="#Footnote_189_189" class="fnanchor">[189]</a></p>
+
+<p>In this statement we have a good example of the further development of
+views which Darwin foreshadowed but did not thoroughly work out. It
+states the biological case clearly and tersely. Plasticity of
+behaviour in special accommodation to special circumstances is of
+survival value; it depends upon acquired characters; it is correlated
+with increase in size and complexity of the cerebrum; under natural
+selection therefore the larger and more complex cerebrum as the organ
+of plastic behaviour has been the outcome of natural selection. We
+have thus the biological foundations for a further development of
+genetic psychology.</p>
+
+<p>There are diversities of opinion, as Darwin showed, with regard to the
+range of instinct in man and the higher animals as contrasted with
+lower types. Darwin himself said<a name="FNanchor_190_190" id="FNanchor_190_190"></a><a href="#Footnote_190_190" class="fnanchor">[190]</a> that "Man, perhaps, has somewhat
+fewer instincts than those possessed by the animals which come next to
+him in the <span class='pagenum'><a name="Page_191" id="Page_191">[191]</a></span>series." On the other hand, Prof. Wm. James says<a name="FNanchor_191_191" id="FNanchor_191_191"></a><a href="#Footnote_191_191" class="fnanchor">[191]</a> that
+man is probably the animal with most instincts. The true position is
+that man and the higher animals have fewer complete and self-sufficing
+instincts than those which stand lower in the scale of mental
+evolution, but that they have an equally large or perhaps larger mass
+of instinctive raw material which may furnish the stuff to be
+elaborated by intelligent processes. There is, perhaps, a greater
+abundance of the primary tissue of experience to be refashioned and
+integrated by secondary modification; there is probably the same
+differentiation in relation to the determining biological ends, but
+there is at the outset less differentiation of the particular and
+specific modes of behaviour. The specialised instinctive performances
+and their concomitant experience-complexes are at the outset more
+indefinite. Only through acquired connections, correlated with
+experience, do they become definitely organised.</p>
+
+<p>The full working-out of the delicate and subtle relationship of
+instinct and educability&mdash;that is, of the hereditary and the acquired
+factors in the mental life&mdash;is the task which lies before genetic and
+comparative psychology. They interact throughout the whole of life,
+and their interactions are very complex. No one can read the chapters
+of <i>The Descent of Man</i> which Darwin devotes to a consideration of the
+mental characters of man and animals without noticing, on the one
+hand, how sedulous he is in his search for hereditary foundations,
+and, on the other hand, how fully he realises the importance of
+acquired habits of mind. The fact that educability itself has innate
+tendencies&mdash;is in fact a partially differentiated educability&mdash;renders
+the unravelling of the factors of mental progress all the more
+difficult.</p>
+
+<p>In his comparison of the mental powers of men and animals it was
+essential that Darwin should lay stress on points of similarity rather
+than on points of difference. Seeking to establish a doctrine of
+evolution, with its basal concept of continuity of process and
+community of character, he was <span class='pagenum'><a name="Page_192" id="Page_192">[192]</a></span>bound to render clear and to emphasise
+the contention that the difference in mind between man and the higher
+animals, great as it is, is one of degree and not of kind. To this end
+Darwin not only recorded a large number of valuable observations of
+his own, and collected a considerable body of information from
+reliable sources, he presented the whole subject in a new light and
+showed that a natural history of mind might be written and that this
+method of study offered a wide and rich field for investigation. Of
+course those who regarded the study of mind only as a branch of
+metaphysics smiled at the philosophical ineptitude of the mere man of
+science. But the investigation, on natural history lines, has been
+prosecuted with a large measure of success. Much indeed still remains
+to be done; for special training is required, and the workers are
+still few. Promise for the future is however afforded by the fact that
+investigation is prosecuted on experimental lines and that something
+like organised methods of research are taking form. There is now but
+little reliance on casual observations recorded by those who have not
+undergone the necessary discipline in these methods. There is also
+some change of emphasis in formulating conclusions. Now that the
+general evolutionary thesis is fully and freely accepted by those who
+carry on such researches, more stress is laid on the differentiation
+of the stages of evolutionary advance than on the fact of their
+underlying community of nature. The conceptual intelligence which is
+especially characteristic of the higher mental procedure of man is
+more firmly distinguished from the perceptual intelligence which he
+shares with the lower animals&mdash;distinguished now as a higher product
+of evolution, no longer as differing in origin or different in kind.
+Some progress has been made, on the one hand in rendering an account
+of intelligent profiting by experience under the guidance of pleasure
+and pain in the perceptual field, on lines predetermined by
+instinctive differentiation for biological ends, and on the other hand
+in elucidating the method of conceptual thought employed,<span class='pagenum'><a name="Page_193" id="Page_193">[193]</a></span> for
+example, by the investigator himself in interpreting the perceptual
+experience of the lower animals.</p>
+
+<p>Thus there is a growing tendency to realise more fully that there are
+two orders of educability&mdash;first an educability of the perceptual
+intelligence based on the biological foundation of instinct, and
+secondly an educability of the conceptual intelligence which
+refashions and rearranges the data afforded by previous inheritance
+and acquisition. It is in relation to this second and higher order of
+educability that the cerebrum of man shows so large an increase of
+mass and a yet larger increase of effective surface through its rich
+convolutions. It is through educability of this order that the human
+child is brought intellectually and affectively into touch with the
+ideal constructions by means of which man has endeavoured, with more
+or less success, to reach an interpretation of nature, and to guide
+the course of the further evolution of his race&mdash;ideal constructions
+which form part of man's environment.</p>
+
+<p>It formed no part of Darwin's purpose to consider, save in broad
+outline, the methods, or to discuss in any fulness of detail the
+results of the process by which a differentiation of the mental
+faculties of man from those of the lower animals has been brought
+about&mdash;a differentiation the existence of which he again and again
+acknowledges. His purpose was rather to show that, notwithstanding
+this differentiation, there is basal community in kind. This must be
+remembered in considering his treatment of the biological foundations
+on which man's systems of ethics are built. He definitely stated that
+he approached the subject "exclusively from the side of natural
+history."<a name="FNanchor_192_192" id="FNanchor_192_192"></a><a href="#Footnote_192_192" class="fnanchor">[192]</a> His general conclusion is that the moral sense is
+fundamentally identical with the social instincts, which have been
+developed for the good of the community; and he suggests that the
+concept which thus enables us to interpret the biological ground-plan
+of morals also enables us to frame a rational ideal of the moral end.
+"As the <span class='pagenum'><a name="Page_194" id="Page_194">[194]</a></span>social instincts," he says,<a name="FNanchor_193_193" id="FNanchor_193_193"></a><a href="#Footnote_193_193" class="fnanchor">[193]</a> "both of man and the lower
+animals have no doubt been developed by nearly the same steps, it
+would be advisable, if found practicable, to use the same definition
+in both cases, and to take as the standard of morality, the general
+good or welfare of the community, rather than the general happiness."
+But the kind of community for the good of which the social instincts
+of animals and primitive men were biologically developed may be
+different from that which is the product of civilisation, as Darwin no
+doubt realised. Darwin's contention was that conscience is a social
+instinct and has been evolved because it is useful to the tribe in the
+struggle for existence against other tribes. One the other hand J. S.
+Mill urged that the moral feelings are not innate but acquired, and
+Bain held the same view, believing that the moral sense is acquired by
+each individual during his life-time. Darwin, who notes<a name="FNanchor_194_194" id="FNanchor_194_194"></a><a href="#Footnote_194_194" class="fnanchor">[194]</a> their
+opinion with his usual candour, adds that "on the general theory of
+evolution this is at least extremely improbable." It is impossible to
+enter into the question here: much turns on the exact connotation of
+the terms "conscience" and "moral sense," and on the meaning we attach
+to the statement that the moral sense is fundamentally identical with
+the social instincts.</p>
+
+<p>Presumably the majority of those who approach the subjects discussed
+in the third, fourth, and fifth chapters of <i>The Descent of Man</i> in
+the full conviction that mental phenomena, not less than organic
+phenomena, have a natural genesis, would, without hesitation, admit
+that the intellectual and moral systems of civilised man are ideal
+constructions, the products of conceptual thought, and that as such
+they are, in their developed form, acquired. The moral sentiments are
+the emotional analogues of highly developed concepts. This does not
+however imply that they are outside the range of natural history
+treatment. Even though it may be desirable to differentiate the moral
+conduct of men from <span class='pagenum'><a name="Page_195" id="Page_195">[195]</a></span>the social behaviour of animals (to which some
+such term as "pre-moral" or "quasi-moral" may be applied), still the
+fact remains that, as Darwin showed, there is abundant evidence of the
+occurrence of such social behaviour&mdash;social behaviour which, even
+granted that it is in large part intelligently acquired, and is itself
+so far a product of educability, is of survival value. It makes for
+that integration without which no social group could hold together and
+escape elimination. Furthermore, even if we grant that such behaviour
+is intelligently acquired, that is to say arises through the
+modification of hereditary instincts and emotions, the fact remains
+that only through these instinctive and emotional data is afforded the
+primary tissue of the experience which is susceptible of such
+modification.</p>
+
+<p>Darwin sought to show, and succeeded in showing, that for the
+intellectual and moral life there are instinctive foundations which a
+biological treatment alone can disclose. It is true that he did not in
+all cases analytically distinguish the foundations from the
+superstructure. Even to-day we are scarcely in a position to do so
+adequately. But his treatment was of great value in giving an impetus
+to further research. This value indeed can scarcely be over-estimated.
+And when the natural history of the mental operations shall have been
+written, the cardinal fact will stand forth, that the instinctive and
+emotional foundations are the outcome of biological evolution and have
+been ingrained in the race through natural selection. We shall more
+clearly realise that educability itself is a product of natural
+selection, though the specific results acquired through cerebral
+modifications are not transmitted through heredity. It will, perhaps,
+also be realised that the instinctive foundations of social behaviour
+are, for us, somewhat out of date and have undergone but little change
+throughout the progress of civilisation, because natural selection has
+long since ceased to be the dominant factor in human progress. The
+history of human progress has been mainly the history of man's higher
+educability, the products of which he has projected<span class='pagenum'><a name="Page_196" id="Page_196">[196]</a></span> on to his
+environment. This educability remains on the average what it was a
+dozen generations ago; but the thought-woven tapestry of his
+surroundings is refashioned and improved by each succeeding
+generation. Few men have in greater measure enriched the
+thought-environment with which it is the aim of education to bring
+educable human beings into vital contact, than has Charles Darwin. His
+special field of work was the wide province of biology; but he did
+much to help us to realise that mental factors have contributed to
+organic evolution and that in man, the highest product of Evolution,
+they have reached a position of unquestioned supremacy.</p>
+
+<div class="footnotes"><h3>FOOTNOTES:</h3>
+
+<div class="footnote"><p><a name="Footnote_153_153" id="Footnote_153_153"></a><a href="#FNanchor_153_153"><span class="label">[153]</span></a> <i>Origin of Species</i> (6th edit.), p. 205.</p></div>
+
+<div class="footnote"><p><a name="Footnote_154_154" id="Footnote_154_154"></a><a href="#FNanchor_154_154"><span class="label">[154]</span></a> <i>Descent of man</i> (2nd edit. 1888), Vol. I. p. 99;
+Popular edit. p. 99.</p></div>
+
+<div class="footnote"><p><a name="Footnote_155_155" id="Footnote_155_155"></a><a href="#FNanchor_155_155"><span class="label">[155]</span></a> <i>Ibid.</i> p. 99.</p></div>
+
+<div class="footnote"><p><a name="Footnote_156_156" id="Footnote_156_156"></a><a href="#FNanchor_156_156"><span class="label">[156]</span></a> <i>The Expression of the Emotions</i> (2nd edit.), p. 32.</p></div>
+
+<div class="footnote"><p><a name="Footnote_157_157" id="Footnote_157_157"></a><a href="#FNanchor_157_157"><span class="label">[157]</span></a> <i>Descent of Man</i>, Vol. II. p. 435.</p></div>
+
+<div class="footnote"><p><a name="Footnote_158_158" id="Footnote_158_158"></a><a href="#FNanchor_158_158"><span class="label">[158]</span></a> <i>Ibid.</i> 437, 438.</p></div>
+
+<div class="footnote"><p><a name="Footnote_159_159" id="Footnote_159_159"></a><a href="#FNanchor_159_159"><span class="label">[159]</span></a> <i>Origin of Species</i> (6th edit.), p. 205.</p></div>
+
+<div class="footnote"><p><a name="Footnote_160_160" id="Footnote_160_160"></a><a href="#FNanchor_160_160"><span class="label">[160]</span></a> <i>Origin of Species</i> (6th edit.), p. 205.</p></div>
+
+<div class="footnote"><p><a name="Footnote_161_161" id="Footnote_161_161"></a><a href="#FNanchor_161_161"><span class="label">[161]</span></a> <i>Ibid.</i> p. 233.</p></div>
+
+<div class="footnote"><p><a name="Footnote_162_162" id="Footnote_162_162"></a><a href="#FNanchor_162_162"><span class="label">[162]</span></a> <i>Ibid.</i> p. 205.</p></div>
+
+<div class="footnote"><p><a name="Footnote_163_163" id="Footnote_163_163"></a><a href="#FNanchor_163_163"><span class="label">[163]</span></a> <i>Origin of Species</i> (6th edit.), p. 233.</p></div>
+
+<div class="footnote"><p><a name="Footnote_164_164" id="Footnote_164_164"></a><a href="#FNanchor_164_164"><span class="label">[164]</span></a> <i>Origin of Species</i>, pp. 210, 211.</p></div>
+
+<div class="footnote"><p><a name="Footnote_165_165" id="Footnote_165_165"></a><a href="#FNanchor_165_165"><span class="label">[165]</span></a> Independently suggested, on somewhat different lines,
+by Profs. J. Mark Baldwin, Henry F. Osborn and the writer.</p></div>
+
+<div class="footnote"><p><a name="Footnote_166_166" id="Footnote_166_166"></a><a href="#FNanchor_166_166"><span class="label">[166]</span></a> <i>Origin of Species</i> (6th edit.), p. 206.</p></div>
+
+<div class="footnote"><p><a name="Footnote_167_167" id="Footnote_167_167"></a><a href="#FNanchor_167_167"><span class="label">[167]</span></a> <i>Expression of the Emotions</i>, p. 13. The passage is
+here somewhat condensed.</p></div>
+
+<div class="footnote"><p><a name="Footnote_168_168" id="Footnote_168_168"></a><a href="#FNanchor_168_168"><span class="label">[168]</span></a> <i>Ibid.</i> p. 368.</p></div>
+
+<div class="footnote"><p><a name="Footnote_169_169" id="Footnote_169_169"></a><a href="#FNanchor_169_169"><span class="label">[169]</span></a> <i>Expression of the Emotions</i>, pp. 373, 374.</p></div>
+
+<div class="footnote"><p><a name="Footnote_170_170" id="Footnote_170_170"></a><a href="#FNanchor_170_170"><span class="label">[170]</span></a> <i>Ibid.</i> p. 368.</p></div>
+
+<div class="footnote"><p><a name="Footnote_171_171" id="Footnote_171_171"></a><a href="#FNanchor_171_171"><span class="label">[171]</span></a> <i>Ibid.</i> p. 369.</p></div>
+
+<div class="footnote"><p><a name="Footnote_172_172" id="Footnote_172_172"></a><a href="#FNanchor_172_172"><span class="label">[172]</span></a> <i>Expression of the Emotions</i>, pp. 65 ff.</p></div>
+
+<div class="footnote"><p><a name="Footnote_173_173" id="Footnote_173_173"></a><a href="#FNanchor_173_173"><span class="label">[173]</span></a> Cf. William James, <i>Principles of Psychology</i>, Vol. <span class="smcap">ii</span>.
+Chap. <span class="smcap">xxv</span>, New York, 1890.</p></div>
+
+<div class="footnote"><p><a name="Footnote_174_174" id="Footnote_174_174"></a><a href="#FNanchor_174_174"><span class="label">[174]</span></a> <i>Darwinism</i>, pp. 282, 283, London, 1889.</p></div>
+
+<div class="footnote"><p><a name="Footnote_175_175" id="Footnote_175_175"></a><a href="#FNanchor_175_175"><span class="label">[175]</span></a> <i>Ibid.</i> p. 283.</p></div>
+
+<div class="footnote"><p><a name="Footnote_176_176" id="Footnote_176_176"></a><a href="#FNanchor_176_176"><span class="label">[176]</span></a> <i>Darwinism</i>, pp. 283, 284.</p></div>
+
+<div class="footnote"><p><a name="Footnote_177_177" id="Footnote_177_177"></a><a href="#FNanchor_177_177"><span class="label">[177]</span></a> <i>Descent of Man</i> (2nd edit.), Vol. <span class="smcap">ii</span>. pp. 136, 137;
+(Popular edit.), pp. 642, 643.</p></div>
+
+<div class="footnote"><p><a name="Footnote_178_178" id="Footnote_178_178"></a><a href="#FNanchor_178_178"><span class="label">[178]</span></a> <i>The Play of Animals</i>, p. 244, London, 1898.</p></div>
+
+<div class="footnote"><p><a name="Footnote_179_179" id="Footnote_179_179"></a><a href="#FNanchor_179_179"><span class="label">[179]</span></a> <i>Ibid.</i> p. 283.</p></div>
+
+<div class="footnote"><p><a name="Footnote_180_180" id="Footnote_180_180"></a><a href="#FNanchor_180_180"><span class="label">[180]</span></a> <i>Descent of Man</i>, Vol. <span class="smcap">ii</span>. p. 60; (Popular edit.), p.
+566.</p></div>
+
+<div class="footnote"><p><a name="Footnote_181_181" id="Footnote_181_181"></a><a href="#FNanchor_181_181"><span class="label">[181]</span></a> <i>The Play of Animals</i>, p. 76.</p></div>
+
+<div class="footnote"><p><a name="Footnote_182_182" id="Footnote_182_182"></a><a href="#FNanchor_182_182"><span class="label">[182]</span></a> <i>Ibid.</i> p. 75.</p></div>
+
+<div class="footnote"><p><a name="Footnote_183_183" id="Footnote_183_183"></a><a href="#FNanchor_183_183"><span class="label">[183]</span></a> <i>The Play of Animals</i> p. 24.</p></div>
+
+<div class="footnote"><p><a name="Footnote_184_184" id="Footnote_184_184"></a><a href="#FNanchor_184_184"><span class="label">[184]</span></a> <i>Descent of Man</i> (1st edit.), Chaps. <span class="smcap">ii</span>, <span class="smcap">iii</span>, <span class="smcap">v</span>; (2nd
+edit.), Chaps. <span class="smcap">iii</span>, <span class="smcap">iv</span>, <span class="smcap">v</span>.</p></div>
+
+<div class="footnote"><p><a name="Footnote_185_185" id="Footnote_185_185"></a><a href="#FNanchor_185_185"><span class="label">[185]</span></a> <i>Descent of Man</i>, Vol. <span class="smcap">i</span>. pp. 70, 71; (Popular edit.),
+pp. 70, 71.</p></div>
+
+<div class="footnote"><p><a name="Footnote_186_186" id="Footnote_186_186"></a><a href="#FNanchor_186_186"><span class="label">[186]</span></a> <i>Ibid.</i> p. 81.</p></div>
+
+<div class="footnote"><p><a name="Footnote_187_187" id="Footnote_187_187"></a><a href="#FNanchor_187_187"><span class="label">[187]</span></a> <i>Ibid.</i> (Popular edit.), p. 82.</p></div>
+
+<div class="footnote"><p><a name="Footnote_188_188" id="Footnote_188_188"></a><a href="#FNanchor_188_188"><span class="label">[188]</span></a> <i>Comptes Rendus des Sciences</i>, June 1, 1868.</p></div>
+
+<div class="footnote"><p><a name="Footnote_189_189" id="Footnote_189_189"></a><a href="#FNanchor_189_189"><span class="label">[189]</span></a> <i>Nature</i>, Vol. <span class="smcap">lxi</span>. pp. 624, 625 (1900).</p></div>
+
+<div class="footnote"><p><a name="Footnote_190_190" id="Footnote_190_190"></a><a href="#FNanchor_190_190"><span class="label">[190]</span></a> <i>Descent of Man</i>, Vol. <span class="smcap">i</span>. p. 100.</p></div>
+
+<div class="footnote"><p><a name="Footnote_191_191" id="Footnote_191_191"></a><a href="#FNanchor_191_191"><span class="label">[191]</span></a> <i>Principles of Psychology</i>, Vol. <span class="smcap">ii</span>. p. 289.</p></div>
+
+<div class="footnote"><p><a name="Footnote_192_192" id="Footnote_192_192"></a><a href="#FNanchor_192_192"><span class="label">[192]</span></a> <i>Descent of Man</i>, Vol. <span class="smcap">i</span>. p. 149.</p></div>
+
+<div class="footnote"><p><a name="Footnote_193_193" id="Footnote_193_193"></a><a href="#FNanchor_193_193"><span class="label">[193]</span></a> <i>Descent of Man</i>, p. 185.</p></div>
+
+<div class="footnote"><p><a name="Footnote_194_194" id="Footnote_194_194"></a><a href="#FNanchor_194_194"><span class="label">[194]</span></a> <i>Ibid.</i> p. 150 (footnote).</p></div>
+</div>
+
+
+<hr style="width: 65%;" />
+<p><span class='pagenum'><a name="Page_197" id="Page_197">[197]</a></span></p>
+<h2><a name="VII" id="VII"></a>VII</h2>
+
+<h2>THE INFLUENCE OF THE CONCEPTION OF EVOLUTION ON MODERN PHILOSOPHY</h2>
+<p class="center"><span class="smcap1">By H. H&ouml;ffding</span></p>
+<h4><i>Professor of Philosophy in the University of Copenhagen</i></h4>
+<h3>I</h3>
+<p>It is difficult to draw a sharp line between philosophy and natural
+science. The naturalist who introduces a new principle, or
+demonstrates a fact which throws a new light on existence, not only
+renders an important service to philosophy but is himself a
+philosopher in the broader sense of the word. The aim of philosophy in
+the stricter sense is to attain points of view from which the
+fundamental phenomena and the principles of the special sciences can
+be seen in their relative importance and connection. But philosophy in
+this stricter sense has always been influenced by philosophy in the
+broader sense. Greek philosophy came under the influence of logic and
+mathematics, modern philosophy under the influence of natural science.
+The name of Charles Darwin stands with those of Galileo, Newton, and
+Robert Mayer&mdash;names which denote new problems and great alterations in
+our conception of the universe.</p>
+
+<p>First of all we must lay stress on Darwin's own personality. His deep
+love of truth, his indefatigable inquiry, his wide horizon, and his
+steady self-criticism make him a scientific model, even if his results
+and theories should eventually come to possess mainly an historical
+interest. In the intel<span class='pagenum'><a name="Page_198" id="Page_198">[198]</a></span>lectual domain the primary object is to reach
+high summits from which wide surveys are possible, to reach them
+toiling honestly upwards by the way of experience, and then not to
+turn dizzy when a summit is gained. Darwinians have sometimes turned
+dizzy, but Darwin never. He saw from the first the great importance of
+his hypothesis, not only because of its solution of the old problem as
+to the value of the concept of species, not only because of the grand
+picture of natural evolution which it unrolls, but also because of the
+life and inspiration its method would impart to the study of
+comparative anatomy, of instinct and of heredity, and finally because
+of the influence it would exert on the whole conception of existence.
+He wrote in his note-book in the year 1837: "My theory would give zest
+to recent and fossil comparative anatomy; it would lead to the study
+of instinct, heredity, and mind-heredity, whole [of]
+metaphysics."<a name="FNanchor_195_195" id="FNanchor_195_195"></a><a href="#Footnote_195_195" class="fnanchor">[195]</a></p>
+
+<p>We can distinguish four main points in which Darwin's investigations
+possess philosophical importance.</p>
+
+<p>The evolution hypothesis is much older than Darwin; it is, indeed, one
+of the oldest guessings of human thought. In the eighteenth century is
+was put forward by Diderot and Lamettrie and suggested by Kant (1786).
+As we shall see later, it was held also by several philosophers in the
+first half of the nineteenth century. In his preface to <i>The Origin of
+Species</i>, Darwin mentions the naturalists who were his forerunners.
+But he has set forth the hypothesis of evolution in so energetic and
+thorough a manner that it perforce attracts the attention of all
+thoughtful men in a much higher degree than it did before the
+publication of the <i>Origin</i>.</p>
+
+<p>And further, the importance of his teaching rests on the fact that he,
+much more than his predecessors, even than Lamarck, sought a
+foundation for his hypothesis in definite facts. Modern science began
+by demanding&mdash;with Kepler and Newton&mdash;evidence of <i>varae causae</i>; this
+demand Darwin industriously set himself to satisfy&mdash;hence the wealth
+of <span class='pagenum'><a name="Page_199" id="Page_199">[199]</a></span>material which he collected by his observations and his
+experiments. He not only revived an old hypothesis, but he saw the
+necessity of verifying it by facts. Whether the special cause on which
+he founded the explanation of the origin of species&mdash;Natural
+Selection&mdash;is sufficient, is now a subject of discussion. He himself
+had some doubt in regard to this question, and the criticisms which
+are directed against his hypothesis hit Darwinism rather than Darwin.
+In his indefatigable search for empirical evidence he is a model even
+for his antagonists: he has compelled them to approach the problems of
+life along other lines than those which were formerly followed.</p>
+
+<p>Whether the special cause to which Darwin appealed is sufficient or
+not, at least to it is probably due the greater part of the influence
+which he has exerted on the general trend of thought. "Struggle for
+existence" and "natural selection" are principles which have been
+applied, more or less, in every department of thought. Recent
+research, it is true, has discovered greater empirical
+discontinuity&mdash;leaps, "mutations"&mdash;whereas Darwin believed in the
+importance of small variations slowly accumulated. It has also been
+shown by the experimental method, which in recent biological work has
+succeeded Darwin's more historical method, that types once constituted
+possess great permanence, the fluctuations being restricted within
+clearly defined boundaries. The problem has become more precise, both
+as to variation and as to heredity. The inner conditions of life have
+in both respects shown a greater independence than Darwin had supposed
+in his theory, though he always admitted that the cause of variation
+was to him a great enigma, "a most perplexing problem," and that the
+struggle for life could only occur where variation existed. But, at
+any rate, it was of the greatest importance that Darwin gave a living
+impression of the struggle for life which is everywhere going on, and
+to which even the highest forms of existence must be amenable. The
+philosophical importance of these ideas does not stand or fall with
+the answer<span class='pagenum'><a name="Page_200" id="Page_200">[200]</a></span> to the question, whether natural selection is a sufficient
+explanation of the origin of species or not it has an independent,
+positive value for everyone who will observe life and reality with an
+unbiased mind.</p>
+
+<p>In accentuating the struggle for life Darwin stands as a
+characteristically English thinker: he continues a train of ideas
+which Hobbes and Malthus had already begun. Moreover in his critical
+views as to the conception of species he had English forerunners; in
+the middle ages Occam and Duns Scotus, in the eighteenth century
+Berkeley and Hume. In his moral philosophy, as we shall see later, he
+is an adherent of the school which is represented by Hutcheson, Home
+and Adam Smith. Because he is no philosopher in the stricter sense of
+the term, it is of great interest to see that his attitude of mind is
+that of the great thinkers of his nation.</p>
+
+<p>In considering Darwin's influence on philosophy we will begin with an
+examination of the attitude of philosophy to the conception of
+evolution at the time when <i>The Origin of Species</i> appeared. We will
+then examine the effects which the theory of evolution, and especially
+the idea of the struggle for life, has had, and naturally must have,
+on the discussion of philosophical problems.</p>
+
+
+<p>II</p>
+
+<p>When <i>The Origin of Species</i> appeared fifty years ago Romantic
+speculation, Schelling's and Hegel's philosophy, still reigned on the
+continent, while in England Positivism, the philosophy of Comte and
+Stuart Mill, represented the most important trend of thought. German
+speculation had much to say on evolution, it even pretended to be a
+philosophy of evolution. But then the word "evolution" was to be taken
+in an ideal, not in a real, sense. To speculative thought the forms
+and types of nature formed a system of ideas, within which any form
+could lead us by continuous transitions to any other. It was a
+classificatory system which was re<span class='pagenum'><a name="Page_201" id="Page_201">[201]</a></span>garded as a divine world of thought
+or images, within which metamorphoses could go on&mdash;a condition
+comparable with that in the mind of the poet when one image follows
+another with imperceptible changes. Goethe's ideas of evolution, as
+expressed in his <i>Metamorphosen der Pflanzen und der Thiere</i>, belong
+to this category; it is, therefore, incorrect to call him a forerunner
+of Darwin. Schelling and Hegel held the same idea; Hegel expressly
+rejected the conception of a real evolution in time as coarse and
+materialistic. "Nature," he says, "is to be considered as a <i>system of
+stages</i>, the one necessarily arising from the other, and being the
+nearest truth of that from which it proceeds; but not in such a way
+that the one is <i>naturally</i> generated by the other; on the contrary
+[their connection lies] in the inner idea which is the ground of
+nature. The <i>metamorphosis</i> can be ascribed only to the notion as
+such, because it alone is evolution.... It has been a clumsy idea in
+the older as well as in the newer philosophy of nature, to regard the
+transformation and the transition from one natural form and sphere to
+a higher as an outward and actual production."<a name="FNanchor_196_196" id="FNanchor_196_196"></a><a href="#Footnote_196_196" class="fnanchor">[196]</a></p>
+
+<p>The only one of the philosophers of Romanticism who believed in a
+real, historical evolution, a real production of new species, was
+Oken.<a name="FNanchor_197_197" id="FNanchor_197_197"></a><a href="#Footnote_197_197" class="fnanchor">[197]</a> Danish philosophers, such as Treschow (1812) and Sibbern
+(1846), have also broached the idea of an historical evolution of all
+living beings from the lowest to the highest. Schopenhauer's
+philosophy has a more realistic character than that of Schelling's and
+Hegel's, his diametrical opposites, although he also belongs to the
+romantic school of thought. His philosophical and psychological views
+were greatly influenced by French naturalists and philosophers,
+especially by Cabanis and Lamarck. He praises the "ever memorable
+Lamarck," because he laid so much stress on the "will to live." But he
+repudiates as a "wonderful error" the idea that the organs of animals
+should <span class='pagenum'><a name="Page_202" id="Page_202">[202]</a></span>have reached their present perfection through a development in
+time, during the course of innumerable generations. It was, he said, a
+consequence of the low standard of contemporary French philosophy,
+that Lamarck came to the idea of the construction of living beings in
+time through succession!<a name="FNanchor_198_198" id="FNanchor_198_198"></a><a href="#Footnote_198_198" class="fnanchor">[198]</a></p>
+
+<p>The positivistic stream of thought was not more in favour of a real
+evolution than was the Romantic school. Its aim was to adhere to
+positive facts: it looked with suspicion on far-reaching speculation.
+Comte laid great stress on the discontinuity found between the
+different kingdoms of nature, as well as within each single kingdom.
+As he regarded as unscientific every attempt to reduce the number of
+physical forces, so he rejected entirely the hypothesis of Lamarck
+concerning the evolution of species; the idea of species would in his
+eyes absolutely lose its importance if a transition from species to
+species under the influence of conditions of life were admitted. His
+disciples (Littr&eacute;, Robin) continued to direct against Darwin the
+polemics which their master had employed against Lamarck. Stuart Mill,
+who, in the theory of knowledge, represented the empirical or
+positivistic movement in philosophy&mdash;like his English forerunners from
+Locke to Hume&mdash;founded his theory of knowledge and morals on the
+experience of the single individual. He sympathised with the theory of
+the original likeness of all individuals and derived their
+differences, on which he practically and theoretically laid much
+stress, from the influence both of experience and education, and,
+generally, of physical and social causes. He admitted an individual
+evolution, and, in the human species, an evolution based on social
+progress; but no physiological evolution of species. He was afraid
+that the hypothesis of heredity would carry us back to the old theory
+of "innate" ideas.</p>
+
+<p>Darwin was more empirical than Comte and Mill; experience disclosed to
+him a deeper continuity than they could <span class='pagenum'><a name="Page_203" id="Page_203">[203]</a></span>find; closer than before the
+nature and fate of the single individual were shown to be interwoven
+in the great web binding the life of the species with nature as a
+whole. And the continuity which so many idealistic philosophers could
+find only in the world of thought, he showed to be present in the
+world of reality.</p>
+
+
+<p>III</p>
+
+<p>Darwin's energetic renewal of the old idea of evolution has its chief
+importance in strengthening the conviction of this real continuity in
+the world, of continuity in the series of form and events. It was a
+great support for all those who were prepared to base their conception
+of life on scientific grounds. Together with the recently discovered
+law of the conservation of energy, it helped to produce the great
+realistic movement which characterises the last third of the
+nineteenth century. After the decline of the Romantic movement people
+wished to have firmer ground under their feet and reality now asserted
+itself in a more emphatic manner than in the period of Romanticism. It
+was easy for Hegel to proclaim that "the real" was "the rational," and
+that "the rational" was "the real": reality itself existed for him
+only in the interpretation of ideal reason, and if there was anything
+which could not be merged in the higher unity of thought, then it was
+only an example of the "impotence of nature to hold to the idea." But
+now concepts are to be founded on nature and not on any system of
+categories too confidently deduced <i>&agrave; priori</i>. The new devotion to
+nature had its recompense in itself, because the new points of view
+made us see that nature could indeed "hold to ideas," though perhaps
+not to those which we had cogitated beforehand.</p>
+
+<p>A most important question for philosophers to answer was whether the
+new views were compatible with an idealistic conception of life and
+existence. Some proclaimed that we have now no need of any philosophy
+beyond the principles of the conservation of matter and energy and the
+principle of natural evolution: existence should and could be
+definitely<span class='pagenum'><a name="Page_204" id="Page_204">[204]</a></span> and completely explained by the laws of material nature.
+But abler thinkers saw that the thing was not so simple. They were
+prepared to give the new views their just place and to examine what
+alterations the old views must undergo in order to be brought into
+harmony with the new data.</p>
+
+<p>The realistic character of Darwin's theory was shown not only in the
+idea of natural continuity, but also, and not least, in the idea of
+the cause whereby organic life advances step by step. This idea&mdash;the
+idea of the struggle for life&mdash;implied that nothing could persist, if
+it had no power to maintain itself under the given conditions. Inner
+value alone does not decide. Idealism was here put to its hardest
+trial. In continuous evolution it could perhaps still find an analogy
+to the inner evolution of ideas in the mind; but in the demand for
+power in order to struggle with outward conditions Realism seemed to
+announce itself in its most brutal form. Every form of Idealism had to
+ask itself seriously how it was going to "struggle for life" with this
+new Realism.</p>
+
+<p>We will now give a short account of the position which leading
+thinkers in different countries have taken up in regard to this
+question.</p>
+
+<p>I. Herbert Spencer was the philosopher whose mind was best prepared by
+his own previous thinking to admit the theory of Darwin to a place in
+his conception of the world. His criticism of the arguments which had
+been put forward against the hypothesis of Lamarck, showed that
+Spencer, as a young man, was an adherent to the evolution idea. In his
+<i>Social Statics</i> (1850) he applied this idea to human life and moral
+civilisation. In 1852 he wrote an essay on <i>The Development
+Hypothesis</i>, in which he definitely stated his belief that the
+differentiation of species, like the differentiation within a single
+organism, was the result of development. In the first edition of his
+<i>Psychology</i> (1855) he took a step which put him in opposition to the
+older English school (from Locke to Mill): he acknowledged "innate
+ideas" so far as to admit the tendency of acquired habits to<span class='pagenum'><a name="Page_205" id="Page_205">[205]</a></span> be
+inherited in the course of generations, so that the nature and
+functions of the individual are only to be understood through its
+connection with the life of the species. In 1857, in his essay on
+<i>Progress</i>, he propounded the law of differentiation as a general law
+of evolution, verified by examples from all regions of experience, the
+evolution of species being only one of these examples. On the effect
+which the appearance of <i>The Origin of Species</i> had on his mind he
+writes in his <i>Autobiography</i>: "Up to that time ... I held that the
+sole cause of organic evolution is the inheritance of
+functionally-produced modifications. The <i>Origin of Species</i> made it
+clear to me that I was wrong, and that the larger part of the facts
+cannot be due to any such cause.... To have the theory of organic
+evolution justified was of course to get further support for that
+theory of evolution at large with which ... all my conceptions were
+bound up."<a name="FNanchor_199_199" id="FNanchor_199_199"></a><a href="#Footnote_199_199" class="fnanchor">[199]</a> Instead of the metaphorical expression "natural
+selection," Spencer introduced the term "survival of the fittest,"
+which found favour with Darwin as well as with Wallace.</p>
+
+<p>In working out his ideas of evolution, Spencer found that
+differentiation was not the only form of evolution. In its simplest
+form evolution is mainly a concentration, previously scattered
+elements being integrated and losing independent movement.
+Differentiation is only forthcoming when minor wholes arise within a
+greater whole. And the highest form of evolution is reached when there
+is a harmony between concentration and differentiation, a harmony
+which Spencer calls equilibration and which he defines as a moving
+equilibrium. At the same time this definition enables him to
+illustrate the expression "survival of the fittest." "Every living
+organism exhibits such a moving equilibrium&mdash;a balanced set of
+functions constituting its life; and the overthrow of this balanced
+set of functions or moving equilibrium is what we call death. Some
+individuals in a species are so constituted that their moving
+equilibria are less easily <span class='pagenum'><a name="Page_206" id="Page_206">[206]</a></span>overthrown than those of other
+individuals; and these are the fittest which survive, or, in Mr.
+Darwin's language, they are the select which nature preserves."<a name="FNanchor_200_200" id="FNanchor_200_200"></a><a href="#Footnote_200_200" class="fnanchor">[200]</a>
+Not only in the domain of organic life, but in all domains, the summit
+of evolution is, according to Spencer, characterised by such a
+harmony&mdash;by a moving equilibrium.</p>
+
+<p>Spencer's analysis of the concept of evolution, based on a great
+variety of examples, has made this concept clearer and more definite
+than before. It contains the three elements; integration,
+differentiation and equilibration. It is true that a concept which is
+to be valid for all domains of experience must have an abstract
+character, and between the several domains there is, strictly
+speaking, only a relation of analogy. So there is only analogy between
+psychical and physical evolution. But this is no serious objection,
+because general concepts do not express more than analogies between
+the phenomena which they represent. Spencer takes his leading forms
+from the material world in defining evolution (in the simplest form)
+as integration of matter and dissipation of movement; but as he&mdash;not
+always quite consistently<a name="FNanchor_201_201" id="FNanchor_201_201"></a><a href="#Footnote_201_201" class="fnanchor">[201]</a>&mdash;assumed a correspondence of mind and
+matter, he could very well give these terms an indirect importance for
+psychical evolution. Spencer has always, in my opinion with full
+right, repudiated the ascription of materialism. He is no more a
+materialist than Spinoza. In his <i>Principles of Psychology</i> (&sect; 63) he
+expressed himself very clearly: "Though it seems easier to translate
+so-called matter into so-called spirit, than to translate so-called
+spirit into so-called matter&mdash;which latter is indeed wholly
+impossible&mdash;yet no translation can carry us beyond our symbols." These
+words lead us naturally to a group of thinkers whose starting-point
+was psychical evolution. But we have still one aspect of Spencer's
+philosophy to mention.</p>
+
+<p>Spencer founded his "laws of evolution" on an inductive <span class='pagenum'><a name="Page_207" id="Page_207">[207]</a></span>basis, but he
+was convinced that they could be deduced from the law of the
+conservation of energy. Such a deduction is, perhaps, possible for the
+more elementary forms of evolution, integration and differentiation;
+but it is not possible for the highest form, the equilibration, which
+is a harmony of integration and differentiation. Spencer can no more
+deduce the necessity for the eventual appearance of "moving
+equilibria" of harmonious totalities than Hegel could guarantee the
+"higher unities" in which all contradictions should be reconciled. In
+Spencer's hands the theory of evolution acquired a more decidedly
+optimistic character than in Darwin's; but I shall deal later with the
+relation of Darwin's hypothesis to the opposition of optimism and
+pessimism.</p>
+
+<p>II. While the starting-point of Spencer was biological or
+cosmological, psychical evolution being conceived as in analogy with
+physical, a group of eminent thinkers&mdash;in Germany Wundt, in France
+Fouill&eacute;e, in Italy Ardig&ograve;&mdash;took, each in his own manner, their
+starting-point in psychical evolution as an original fact and as a
+type of all evolution, the hypothesis of Darwin coming in as a
+corroboration and as a special example. They maintain the continuity
+of evolution; they find this character most prominent in psychical
+evolution, and this is for them a motive to demand a corresponding
+continuity in the material, especially in the organic domain.</p>
+
+<p>To Wundt and Fouill&eacute;e the concept of will is prominent. They see the
+type of all evolution in the transformation of the life of will from
+blind impulse to conscious choice; the theories of Lamarck and Darwin
+are used to support the view that there is in nature a tendency to
+evolution in steady reciprocity with external conditions. The struggle
+for life is here only a secondary fact. Its apparent prominence is
+explained by the circumstance that the influence of external
+conditions is easily made out, while inner conditions can be verified
+only through their effects. For Ardig&ograve; the evolution of thought was
+the starting-point and the type: in the<span class='pagenum'><a name="Page_208" id="Page_208">[208]</a></span> evolution of a scientific
+hypothesis we see a progress from the indefinite (<i>indistinto</i>) to the
+definite (<i>distinto</i>), and this is a characteristic of all evolution,
+as Ardig&ograve; has pointed out in a series of works. The opposition between
+<i>indistinto</i> and <i>distinto</i> corresponds to Spencer's opposition
+between homogeneity and heterogeneity. The hypothesis of the origin of
+differences of species from more simple forms is a special example of
+the general law of evolution.</p>
+
+<p>In the views of Wundt and Fouill&eacute;e we find the fundamental idea of
+idealism psychical phenomena as expressions of the innermost nature of
+existence. They differ from the older Idealism in the great stress
+which they lay on evolution as a real, historical process which is
+going on through steady conflict with external conditions. The
+Romantic dread of reality is broken. It is beyond doubt that Darwin's
+emphasis on the struggle for life as a necessary condition of
+evolution has been a very important factor in carrying philosophy back
+to reality from the heaven of pure ideas. The philosophy of Ardig&ograve;, on
+the other side, appears more as a continuation and deepening of
+positivism, though the Italian thinker arrived at his point of view
+independently of French-English positivism. The idea of continuous
+evolution is here maintained in opposition to Comte's and Mill's
+philosophy of discontinuity. From Wundt and Fouill&eacute;e Ardig&ograve; differs in
+conceiving psychical evolution not as an immediate revelation of the
+innermost nature of existence, but only as a single, though the most
+accessible example, of evolution.</p>
+
+<p>III. To the French philosophers Boutroux and Bergson, evolution proper
+is continuous and qualitative, while outer experience and physical
+science give us fragments only, sporadic processes and mechanical
+combinations. To Bergson, in his recent work <i>L'Evolution Cr&eacute;atrice</i>,
+evolution consists in an <i>&eacute;lan de vie</i> which to our fragmentary
+observation and analytic reflexion appears as broken into a manifold
+of elements and processes. The concept of matter in its scientific
+form is the result of this breaking asunder, essen<span class='pagenum'><a name="Page_209" id="Page_209">[209]</a></span>tial for all
+scientific reflexion. In these conceptions the strongest opposition
+between inner and outer conditions of evolution is expressed: in the
+domain of internal conditions spontaneous development of qualitative
+forms&mdash;in the domain of external conditions discontinuity and
+mechanical combination.</p>
+
+<p>We see, then, that the theory of evolution has influenced philosophy
+in a variety of forms. It has made idealistic thinkers revise their
+relation to the real world; it has led positivistic thinkers to find a
+closer connection between the facts on which they based their views;
+it has made us all open our eyes for new possibilities to arise
+through the <i>prima facie</i> inexplicable "spontaneous" variations which
+are the condition of all evolution. This last point is one of peculiar
+interest. Deeper than speculative philosophy and mechanical science
+saw in the days of their triumph, we catch sight of new streams, whose
+sources and laws we have still to discover. Most sharply does this
+appear in the theory of mutation, which is only a stronger
+accentuation of a main point in Darwinism. It is interesting to see
+that an analogous problem comes into the foreground in physics through
+the discovery of radioactive phenomena, and in psychology through the
+assumption of psychical new formations (as held by Boutroux, William
+James and Bergson). From this side, Darwin's ideas, as well as the
+analogous ideas in other domains, incite us to renewed examination of
+our first principles, their rationality and their value. On the other
+hand, his theory of the struggle for existence challenges us to
+examine the conditions and discuss the outlook as to the persistence
+of human life and society and of the values that belong to them. It is
+not enough to hope (or fear?) the rising of new forms; we have also to
+investigate the possibility of upholding the forms and ideals which
+have hitherto been the bases of human life. Darwin has here given his
+age the most earnest and most impressive lesson. This side of Darwin's
+theory is of peculiar interest to some special philosophical problems
+to which I now pass.<span class='pagenum'><a name="Page_210" id="Page_210">[210]</a></span></p>
+
+
+<p>IV</p>
+
+<p>Among philosophical problems the problem of knowledge has in the last
+century occupied a foremost place. It is natural, then, to ask how
+Darwin and the hypothesis whose most eminent representative he is,
+stand to this problem.</p>
+
+<p>Darwin started an hypothesis. But every hypothesis is won by inference
+from certain presuppositions, and every inference is based on the
+general principles of human thought. The evolution hypothesis
+presupposes, then, human thought and its principles. And not only the
+abstract logical principles are thus pre-supposed. The evolution
+hypothesis purports to be not only a formal arrangement of phenomena,
+but to express also the law of a real process. It supposes, then, that
+the real data&mdash;all that in our knowledge which we do not produce
+ourselves, but which we in the main simply receive&mdash;are subject to
+laws which are at least analogous to the logical relations of our
+thoughts; in other words, it assumes the validity of the principle of
+causality. If organic species could arise without cause there would be
+no use in framing hypotheses. Only if we assume the principle of
+causality, is there a problem to solve.</p>
+
+<p>Though Darwinism has had a great influence on philosophy considered as
+a striving after a scientific view of the world, yet here is a point
+of view&mdash;the epistemological&mdash;where philosophy is not only independent
+but reaches beyond any result of natural science. Perhaps it will be
+said: the powers and functions of organic beings only persist (perhaps
+also only arise) when they correspond sufficiently to the conditions
+under which the struggle of life is to go on. Human thought itself is,
+then, a variation (or a mutation) which has been able to persist and
+to survive. Is not, then, the problem of knowledge solved by the
+evolution hypothesis? Spencer had given an affirmative answer to this
+question before the appearance of <i>The Origin of Species</i>. For the
+individual, he said, there is an <i>&agrave; priori</i>, original, basis (or
+<i>Anlage</i>) for all mental life; but in the species all<span class='pagenum'><a name="Page_211" id="Page_211">[211]</a></span> powers have
+developed in reciprocity with extendal conditions. Knowledge is here
+considered from the practical point of view, as a weapon in the
+struggle for life, as an "organon" which has been continuously in use
+for generations. In recent years the economic or pragmatic
+epistemology, as developed by Avenarius and Mach in Germany, and by
+James in America, points in the same direction. Science, it is said,
+only maintains those principles and presuppositions which are
+necessary to the simplest and clearest orientation be applied to
+experience and to practical work, will successively be eliminated.</p>
+
+<p>In these views a striking and important application is made of the
+idea of struggle for life to the development of human thought. Thought
+must, as all other things in the world, struggle for life. But this
+whole consideration belongs to psychology, not to the theory of
+knowledge (epistemology), which is concerned only with the validity of
+knowledge, not with its historical origin. Every hypothesis to explain
+the origin of knowledge must submit to cross-examination by the theory
+of knowledge, because it works with the fundamental forms and
+principles of human thought. We cannot go further back than these
+forms and principles, which it is the aim of epistemology to ascertain
+and for which no further reason can be given.<a name="FNanchor_202_202" id="FNanchor_202_202"></a><a href="#Footnote_202_202" class="fnanchor">[202]</a></p>
+
+<p>But there is another side of the problem which is, perhaps, of more
+importance and which epistemology generally overlooks. If new
+variations can arise, not only in organic but perhaps also in
+inorganic nature, new tasks are placed before the human mind. The
+question is, then, if it has forms in which there is room for the new
+matter? We are here touching a possibility which the great master of
+epistemology did not bring to light. Kant supposed confidently that no
+other matter of knowledge could stream forth from the <span class='pagenum'><a name="Page_212" id="Page_212">[212]</a></span>dark source
+which he called "the thing-in-itself," than such as could be
+synthesised in our existing forms of knowledge. He mentions the
+possibility of other forms than the human, and warns us against the
+dogmatic assumption that the human conception of existence should be
+absolutely adequate. But he seems to be quite sure that the
+thing-in-itself works constantly, and consequently always gives us
+only what our powers can master. This assumption was a consequence of
+Kant's rationalistic tendency, but one for which no warrant can be
+given. Evolutionism and systematism are opposing tendencies which can
+never be absolutely harmonised one with the other. Evolution may at
+any time break some form which the system-monger regards as finally
+established. Darwin himself felt a great difference in looking at
+variation as an evolutionist and as a systematist. When he was working
+at his evolution theory, he was very glad to find variations; but they
+were a hindrance to him when he worked as a systematist, in preparing
+his work on Cirripedia. He says in a letter: "I had thought the same
+parts of the same species more resemble (than they do anyhow in
+Cirripedia) objects cast in the same mould. Systematic work would be
+easy were it not for this confounded variation, which, however, is
+pleasant to me as a speculatist, though odious to me as a
+systematist."<a name="FNanchor_203_203" id="FNanchor_203_203"></a><a href="#Footnote_203_203" class="fnanchor">[203]</a> He could indeed be angry with variations even as an
+evolutionist; but then only because he could not explain them, not
+because he could not classify them. "If, as I must think, external
+conditions produce little <i>direct</i> effect, what the devil determines
+each particular variation?"<a name="FNanchor_204_204" id="FNanchor_204_204"></a><a href="#Footnote_204_204" class="fnanchor">[204]</a> What Darwin experienced in this
+particular domain holds good of all knowledge. All knowledge is
+systematic, in so far as it strives to put phenomena in quite definite
+relations, one to another. But the systematisation can never be
+complete. And here Darwin has contributed much to widen the world, for
+us. He has shown us forces and tendencies in <span class='pagenum'><a name="Page_213" id="Page_213">[213]</a></span>nature which make
+absolute systems impossible, at the same time that they give us new
+objects and problems. There is still a place for what Lessing called
+"the unceasing striving after truth," while "absolute truth" (in the
+sense of a closed system) is unattainable so long as life and
+experience are going on.</p>
+
+<p>There is here a special remark to be made. As we have seen above,
+recent research has shown that natural selection or struggle for life
+is no explanation of variations. Hugo de Vries distinguishes between
+partial and embryonal variations, or between variations and mutations,
+only the last-named being heritable, and therefore of importance for
+the origin of new species. But the existence of variations is not only
+of interest for the problem of the origin of species; it has also a
+more general interest. An individual does not lose its importance for
+knowledge, because its qualities are not heritable. On the contrary,
+in higher beings at least, individual peculiarities will become more
+and more independent objects of interest. Knowledge takes account of
+the biographies not only of species, but also of individuals: it seeks
+to find the law of development of the single individual.<a name="FNanchor_205_205" id="FNanchor_205_205"></a><a href="#Footnote_205_205" class="fnanchor">[205]</a> As
+Leibnitz said long ago, individuality consists in the law of the
+changes of a being: "La loi du changement fait l'individualit&eacute; de
+chaque substance." Here is a world which is almost new for science,
+which till now has mainly occupied itself with general laws and forms.
+But these are ultimately only means to understand the individual
+phenomena, in whose nature and history a manifold of laws and forms
+always co&ouml;perate. The importance of this remark will appear in the
+sequel. <span class='pagenum'><a name="Page_214" id="Page_214">[214]</a></span></p>
+
+
+<p>V</p>
+
+<p>To many people the Darwinian theory of natural selection or struggle
+for existence seemed to change the whole conception of life, and
+particularly all the conditions on which the validity of ethical ideas
+depends. If only that has persistence which can be adapted to a given
+condition, what will then be the fate of our ideals, of our standards
+of good and evil? Blind force seems to reign, and the only thing that
+counts seems to be the most heedless use of power. Darwinism, it was
+said, has proclaimed brutality. No other difference seems permanent
+save that between the sound, powerful and happy on the one side, the
+sick, feeble and unhappy on the other; and every attempt to alleviate
+this difference seems to lead to general enervation. Some of those who
+interpreted Darwinism in this manner felt an aesthetic delight in
+contemplating the heedlessness and energy of the great struggle for
+existence and anticipated the realisation of a higher human type as
+the outcome of it: so Nietzsche and his followers. Others recognising
+the same consequences in Darwinism regarded these as one of the
+strongest objections against it; so D&uuml;hring and Kropotkin (in his
+earlier works).</p>
+
+<p>This interpretation of Darwinism was frequent in the interval between
+the two main works of Darwin&mdash;<i>The Origin of Species</i> and <i>The Descent
+of Man</i>. But even during this interval it was evident to an attentive
+reader that Darwin himself did not found his standard of good and evil
+on the features of the life of nature he had emphasised so strongly.
+He did not justify the ways along which nature reached its ends; he
+only pointed them out. The "real" was not to him, as to Hegel, one
+with the "rational." Darwin has, indeed, by his whole conception of
+nature, rendered a great service to ethics in making the difference
+between the life of nature and the ethical life appear in so strong a
+light. The ethical problem could now be stated in a sharper form than
+before. But this was not the first time that the idea of the struggle<span class='pagenum'><a name="Page_215" id="Page_215">[215]</a></span>
+for life was put in relation to the ethical problem. In the
+seventeenth century Thomas Hobbes gave the first impulse to the whole
+modern discussion of ethical principles in his theory of <i>bellum
+omnium contra omnes</i>. Men, he taught, are in the state of nature
+enemies one of another, and they live either in fright or in the glory
+of power. But it was not the opinion of Hobbes that this made ethics
+impossible. On the contrary, he found a standard for virtue and vice
+in the fact that some qualities and actions have a tendency to bring
+us out of the state of war and to secure peace, while other qualities
+have a contrary tendency. In the eighteenth century even Immanuel
+Kant's ideal ethics had&mdash;so far as can be seen&mdash;a similar origin.
+Shortly before the foundation of his definitive ethics, Kant wrote his
+<i>Idee zu einer allgemeinen Weltgeschichte</i> (1784), where&mdash;in a way
+which reminds us of Hobbes, and is prophetic of Darwin&mdash;he describes
+the forward-driving power of struggle in the human world. It is here
+as with the struggle of the trees for light and air, through which
+they compete with one another in height. Anxiety about war can only be
+allayed by an ordinance which gives everyone his full liberty under
+acknowledgment of the equal liberty of others. And such ordinance and
+acknowledgment are also attributes of the content of the moral law, as
+Kant proclaimed it in the year after the publication of his essay
+(1785).<a name="FNanchor_206_206" id="FNanchor_206_206"></a><a href="#Footnote_206_206" class="fnanchor">[206]</a> Kant really came to his ethics by the way of evolution,
+though he afterwards disavowed it. Similarly the same line of thought
+may be traced in Hegel though it has been disguised in the form of
+speculative dialectics.<a name="FNanchor_207_207" id="FNanchor_207_207"></a><a href="#Footnote_207_207" class="fnanchor">[207]</a> And in Schopenhauer's theory of the blind
+will to live and its abrogation by the ethical feeling, which is
+founded on universal sympathy, we have a more individualistic form of
+the same idea.</p>
+
+<p>It was, then, not entirely a foreign point of view which <span class='pagenum'><a name="Page_216" id="Page_216">[216]</a></span>Darwin
+introduced into ethical thought, even if we take no account of the
+poetical character of the word "struggle" and of the more direct
+adaptation, through the use and non-use of power, which Darwin also
+emphasised. In <i>The Descent of Man</i> he has devoted a special
+chapter<a name="FNanchor_208_208" id="FNanchor_208_208"></a><a href="#Footnote_208_208" class="fnanchor">[208]</a> to a discussion of the origin of the ethical
+consciousness. The characteristic expression of this consciousness he
+found, just as Kant did, in the idea of "ought"; it was the origin of
+this new idea which should be explained. His hypothesis was that the
+ethical "ought" has its origin in the social and parental instincts,
+which, as well as other instincts (e.g. the instinct of
+self-preservation), lie deeper than pleasure and pain. In many
+species, not least in the human species, these instincts are fostered
+by natural selection; and when the powers of memory and comparison are
+developed, so that single acts can be valued according to the claims
+of the deep social instinct, then consciousness of duty and remorse
+are possible. Blind instinct has developed to conscious ethical will.</p>
+
+<p>As already stated, Darwin, as a moral philosopher belongs to the
+school that was founded by Shaftesbury, and was afterwards represented
+by Hutcheson, Hume, Adam Smith, Comte and Spencer. His merit is,
+first, that he has given this tendency of thought a biological
+foundation, and that he has stamped on it a doughty character in
+showing that ethical ideas and sentiments, rightly conceived, are
+forces which are at work in the struggle for life.</p>
+
+<p>There are still many questions to solve. Not only does the ethical
+development within the human species contain features still
+unexplained;<a name="FNanchor_209_209" id="FNanchor_209_209"></a><a href="#Footnote_209_209" class="fnanchor">[209]</a> but we are confronted by the great problem whether
+after all a genetic historical theory can be of decisive importance
+here. To every consequent ethical consciousness there is a standard of
+value, a primordial value which determines the single ethical
+judgments as <span class='pagenum'><a name="Page_217" id="Page_217">[217]</a></span>their last presupposition, and the "rightness" of this
+basis, the "value" of this value can as little be discussed as the
+"rationality" of our logical principles. There is here revealed a
+possibility of ethical scepticism which evolutionistic ethics (as well
+as intuitive or rationalistic ethics) has overlooked. No demonstration
+can show that the results of the ethical development are definitive
+and universal. We meet here again with the important opposition of
+systematisation and evolution. There will, I think, always be an open
+question here, though comparative ethics, of which we have so far only
+the first attempts, can do much to throw light on it.</p>
+
+<p>It would carry us too far to discuss all the philosophical works on
+ethics, which have been influenced directly or indirectly by
+evolutionism. I may, however, here refer to the book of C. M.
+Williams, <i>A Review of the Systems of Ethics founded on the Theory of
+Evolution</i>,<a name="FNanchor_210_210" id="FNanchor_210_210"></a><a href="#Footnote_210_210" class="fnanchor">[210]</a> in which, besides Darwin, the following authors are
+reviewed: Wallace, Haeckel, Spencer, Fiske, Rolph, Barratt, Stephen,
+Carneri, H&ouml;ffding, Gizycki, Alexander, R&eacute;e. As works which criticise
+evolutionistic ethics from an intuitive point of view and in an
+instructive way, may be cited: Guyau, <i>La morale anglaise
+contemporaine</i>,<a name="FNanchor_211_211" id="FNanchor_211_211"></a><a href="#Footnote_211_211" class="fnanchor">[211]</a> and Sorley, <i>Ethics of Naturalism</i>. I will only
+mention some interesting contributions to ethical discussion which can
+be found in Darwinism besides the idea of struggle for life.</p>
+
+<p>The attention which Darwin has directed to variations has opened our
+eyes to the differences in human nature as well as in nature
+generally. There is here a fact of great importance for ethical
+thought, no matter from what ultimate premiss it starts. Only from a
+very abstract point of view can different individuals be treated in
+the same manner. The most eminent ethical thinkers, men such as Jeremy
+Bentham and Immanuel Kant, who discussed ethical questions from very
+opposite standpoints, agreed in regarding all <span class='pagenum'><a name="Page_218" id="Page_218">[218]</a></span>men as equal in respect
+of ethical endowment. In regard to Bentham, Leslie Stephen remarks:
+"He is determined to be thoroughly empirical, to take men as he found
+them. But his utilitarianism supposed that men's views of happiness
+and utility were uniform and clear, and that all that was wanted was
+to show them the means by which their ends could be reached."<a name="FNanchor_212_212" id="FNanchor_212_212"></a><a href="#Footnote_212_212" class="fnanchor">[212]</a> And
+Kant supposed that every man would find the "categorical imperative"
+in his consciousness, when he came to sober reflexion, and that all
+would have the same qualifications to follow it. But if continual
+variations, great or small, are going on in human nature, it is the
+duty of ethics to make allowance for them, both in making claims, and
+in valuing what is done. A new set of ethical problems have their
+origin here.<a name="FNanchor_213_213" id="FNanchor_213_213"></a><a href="#Footnote_213_213" class="fnanchor">[213]</a> It is an interesting fact that Stuart Mill's book
+<i>On Liberty</i> appeared in the same year as <i>The Origin of Species</i>.
+Though Mill agreed with Bentham about the original equality of all
+men's endowments, he regarded individual differences as a necessary
+result of physical and social influences, and he claimed that free
+play shall be allowed to differences of character so far as is
+possible without injury to other men. It is a condition of individual
+and social progress that a man's mode of action should be determined
+by his own character and not by tradition and custom, nor by abstract
+rules. This view was to be corroborated by the theory of Darwin.</p>
+
+<p>But here we have reached a point of view from which the criticism,
+which in recent years has often been directed against Darwin&mdash;that
+small variations are of no importance in the struggle for life&mdash;is of
+no weight. From an ethical standpoint, and particularly from the
+ethical standpoint of Darwin himself, it is a duty to foster
+individual differences that can be valuable, even though they can
+neither be of service for physical preservation nor be physically
+inherited. <span class='pagenum'><a name="Page_219" id="Page_219">[219]</a></span>The distinction between variation and mutation is here
+without importance. It is quite natural that biologists should be
+particularly interested in such variations as can be inherited and
+produce new species. But in the human world there is not only a
+physical, but also a mental and social heredity. When an ideal human
+character has taken form, then there is shaped a type, which through
+imitation and influence can become an important factor in subsequent
+development, even if it cannot form a species in the biological sense
+of the word. Spiritually strong men often succumb in the physical
+struggle for life; but they can nevertheless be victorious through the
+typical influence they exert, perhaps on very distant generations, if
+the remembrance of them is kept alive, be it in legendary or in
+historical form. Their very failure can show that a type has taken
+form which is maintained at all risks, a standard of life which is
+adhered to in spite of the strongest opposition. The question "to be
+or not to be" can be put from very different levels of being: it has
+too often been considered a consequence of Darwinism that this
+question is only to be put from the lowest level. When a stage is
+reached, where ideal (ethical, intellectual, aesthetic) interests are
+concerned, the struggle for life is a struggle for the preservation of
+this stage. The giving up of a higher standard of life is a sort of
+death; for there is not only a physical, there is also a spiritual,
+death.</p>
+
+
+<p>VI</p>
+
+<p>The Socratic character of Darwin's mind appears in his wariness in
+drawing the last consequences of his doctrine, in contrast both with
+the audacious theories of so many of his followers and with the
+consequences which his antagonists were busy in drawing. Though he, as
+we have seen, saw from the beginning that his hypothesis would
+occasion "a whole of metaphysics," he was himself very reserved as to
+the ultimate questions, and his answers to such questions were
+extorted from him.<span class='pagenum'><a name="Page_220" id="Page_220">[220]</a></span></p>
+
+<p>As to the question of optimism and pessimism, Darwin held that though
+pain and suffering were very often the ways by which animals were led
+to pursue that course of action which is most beneficial to the
+species, yet pleasurable feelings were the most habitual guides. "We
+see this in the pleasure from exertion, even occasionally from great
+exertion of the body or mind, in the pleasure of our daily meals, and
+especially in the pleasure derived from sociability, and from loving
+our families." But there was to him so much suffering in the world
+that it was a strong argument against the existence of an intelligent
+First Cause.<a name="FNanchor_214_214" id="FNanchor_214_214"></a><a href="#Footnote_214_214" class="fnanchor">[214]</a></p>
+
+<p>It seems to me that Darwin was not so clear on another question, that
+of the relation between improvement and adaptation. He wrote to Lyell:
+"When you contrast natural selection and 'improvement,' you seem
+always to overlook ... that every step in the natural selection of
+each species implies improvement in that species <i>in relation to its
+condition of life</i>.... Improvement implies, I suppose, <i>each form
+obtaining many parts or organs</i>, all excellently adapted for their
+functions." "All this," he adds, "seems to me quite compatible with
+certain forms fitted for simple conditions, remaining unaltered, or
+being, degraded."<a name="FNanchor_215_215" id="FNanchor_215_215"></a><a href="#Footnote_215_215" class="fnanchor">[215]</a> But the great question is, if the conditions of
+life will in the long run favour "improvement" in the sense of
+differentiation (or harmony of differentiation and integration). Many
+beings are best adapted to their conditions of life if they have few
+organs and few necessities. Pessimism would not only be the
+consequence, if suffering outweighed happiness, but also if the most
+elementary forms of happiness were predominant, or if there were a
+tendency to reduce the standard of life to the simplest possible, the
+contentment of inertia or stable equilibrium. There are animals which
+are very highly differentiated and active in their young state, but
+later lose their complex organisation and concentrate themselves on
+the one function of nutrition. In the human world analogies <span class='pagenum'><a name="Page_221" id="Page_221">[221]</a></span>to this
+sort of adaptation are not wanting. Young "idealists" very often end
+as old "Philistines." Adaptation and progress are not the same.</p>
+
+<p>Another question of great importance in respect to human evolution is,
+whether there will be always a possibility for the existence of an
+impulse to progress, an impulse to make great claims on life, to be
+active and to alter the conditions of life instead of adapting to them
+in a passive manner. Many people do not develop because they have too
+few necessities, and because they have no power to imagine other
+conditions of life than those under which they live. In his remarks on
+"the pleasure from exertion" Darwin has a point of contact with the
+practical idealism of former times&mdash;with the ideas of Lessing and
+Goethe, of Condorcet and Fichte. The continual striving which was the
+condition of salvation to Faust's soul, is also the condition of
+salvation to mankind. There is a holy fire which we ought to keep
+burning, if adaptation is really to be improvement. If, as I have
+tried to show in my <i>Philosophy of Religion</i>, the innermost core of
+all religion is faith in the persistence of value in the world, and if
+the highest values express themselves in the cry "Excelsior!" then the
+capital point is, that this cry should always be heard and followed.
+We have here a corollary of the theory of evolution in its application
+to human life.</p>
+
+<p>Darwin declared himself an agnostic, not only because he could not
+harmonise the large amount of suffering in the world with the idea of
+a God as its first cause, but also because he "was aware that if we
+admit a first cause, the mind still craves to know whence it came and
+how it arose."<a name="FNanchor_216_216" id="FNanchor_216_216"></a><a href="#Footnote_216_216" class="fnanchor">[216]</a> He saw, as Kant had seen before him and expressed
+in his <i>Kritik der Urtheilskraft</i>, that we cannot accept either of the
+only two possibilities which we are able to conceive: chance (or brute
+force) and design. Neither mechanism nor teleology can give an
+absolute answer to ultimate questions. The universe, and especially
+the organic life in it, can neither <span class='pagenum'><a name="Page_222" id="Page_222">[222]</a></span>be explained as a mere
+combination of absolute elements nor as the effect of a constructing
+thought. Darwin concluded, as Kant, and before him Spinoza, that the
+oppositions and distinctions which our experience presents, cannot
+safely be regarded as valid for existence in itself. And, with Kant
+and Fichte, he found his stronghold in the conviction that man has
+something to do, even if he cannot solve all enigmas. "The safest
+conclusion seems to me that the whole subject is beyond the scope of
+man's intellect; but man can do his duty."<a name="FNanchor_217_217" id="FNanchor_217_217"></a><a href="#Footnote_217_217" class="fnanchor">[217]</a></p>
+
+<p>Is this the last word of human thought? Does not the possibility, that
+man can do his duty, suppose that the conditions of life allow of
+continuous ethical striving, so that there is a certain harmony
+between cosmic order and human ideals? Darwin himself has shown how
+the consciousness of duty can arise as a natural result of evolution.
+Moreover there are lines of evolution which have their end in ethical
+idealism, in a kingdom of values, which must struggle for life as all
+things in the world must do, but a kingdom which has its firm
+foundation in reality.</p>
+
+<div class="footnotes"><h3>FOOTNOTES:</h3>
+
+<div class="footnote"><p><a name="Footnote_195_195" id="Footnote_195_195"></a><a href="#FNanchor_195_195"><span class="label">[195]</span></a> <i>Life and Letters of Charles Darwin</i>, Vol. <span class="smcap">i</span>. p. 8.</p></div>
+
+<div class="footnote"><p><a name="Footnote_196_196" id="Footnote_196_196"></a><a href="#FNanchor_196_196"><span class="label">[196]</span></a> <i>Encyclop&auml;die der philosophischen Wissenschaften</i> (4th
+edit.), Berlin, 1845, &sect; 249.</p></div>
+
+<div class="footnote"><p><a name="Footnote_197_197" id="Footnote_197_197"></a><a href="#FNanchor_197_197"><span class="label">[197]</span></a> <i>Lehrbuch der Naturphilosophie</i>, Jena, 1809.</p></div>
+
+<div class="footnote"><p><a name="Footnote_198_198" id="Footnote_198_198"></a><a href="#FNanchor_198_198"><span class="label">[198]</span></a> <i>Ueber den Willen in der Natur</i> (2nd edit.), Frankfurt
+a. M., 1854, pp. 41-43.</p></div>
+
+<div class="footnote"><p><a name="Footnote_199_199" id="Footnote_199_199"></a><a href="#FNanchor_199_199"><span class="label">[199]</span></a> Spencer, <i>Autobiography</i>, Vol. <span class="smcap">ii</span>. p. 50, London and
+New York, 1904.</p></div>
+
+<div class="footnote"><p><a name="Footnote_200_200" id="Footnote_200_200"></a><a href="#FNanchor_200_200"><span class="label">[200]</span></a> <i>Autobiography</i>, Vol. <span class="smcap">ii</span>. p. 100.</p></div>
+
+<div class="footnote"><p><a name="Footnote_201_201" id="Footnote_201_201"></a><a href="#FNanchor_201_201"><span class="label">[201]</span></a> Cf. my letter to him 1876, now printed in Duncan's
+<i>Life and Letters of Herbert Spencer</i>, p. 178. London, 1908.</p></div>
+
+<div class="footnote"><p><a name="Footnote_202_202" id="Footnote_202_202"></a><a href="#FNanchor_202_202"><span class="label">[202]</span></a> The present writer, many years ago, in his <i>Psychology</i>
+(Copenhagen, 1882; Eng. transl. London, 1891), criticised the
+evolutionistic treatment of the problem of knowledge from the Kantian
+point of view.</p></div>
+
+<div class="footnote"><p><a name="Footnote_203_203" id="Footnote_203_203"></a><a href="#FNanchor_203_203"><span class="label">[203]</span></a> <i>Life and Letters</i>, Vol. II. p. 37.</p></div>
+
+<div class="footnote"><p><a name="Footnote_204_204" id="Footnote_204_204"></a><a href="#FNanchor_204_204"><span class="label">[204]</span></a> <i>Ibid.</i> p. 232.</p></div>
+
+<div class="footnote"><p><a name="Footnote_205_205" id="Footnote_205_205"></a><a href="#FNanchor_205_205"><span class="label">[205]</span></a> The new science of Ecology occupies an intermediate
+position between the biography of species and the biography of
+individuals. Compare <i>Congress of Arts and Science</i>, St. Louis, Vol.
+<span class="smcap">v</span>. 1906 (The Reports of Drude and Robinson) and the work of my
+colleague, E. Warming.</p></div>
+
+<div class="footnote"><p><a name="Footnote_206_206" id="Footnote_206_206"></a><a href="#FNanchor_206_206"><span class="label">[206]</span></a> Cf. my <i>History of Modern Philosophy</i> (Eng. transl.
+London, 1900), <span class="smcap">i</span>. pp. 76-79.</p></div>
+
+<div class="footnote"><p><a name="Footnote_207_207" id="Footnote_207_207"></a><a href="#FNanchor_207_207"><span class="label">[207]</span></a> "Herrschaft und Knechtschaft," <i>Ph&ouml;nomenologie des
+Geistes</i>, <span class="smcap">iv</span>. A., Leiden, 1907.</p></div>
+
+<div class="footnote"><p><a name="Footnote_208_208" id="Footnote_208_208"></a><a href="#FNanchor_208_208"><span class="label">[208]</span></a> <i>The Descent of Man</i>, Vol. <span class="smcap">i</span>. Ch. iii.</p></div>
+
+<div class="footnote"><p><a name="Footnote_209_209" id="Footnote_209_209"></a><a href="#FNanchor_209_209"><span class="label">[209]</span></a> The works of Westermarck and Hobhouse throw new light
+on many of these features.</p></div>
+
+<div class="footnote"><p><a name="Footnote_210_210" id="Footnote_210_210"></a><a href="#FNanchor_210_210"><span class="label">[210]</span></a> New York and London, 1893.</p></div>
+
+<div class="footnote"><p><a name="Footnote_211_211" id="Footnote_211_211"></a><a href="#FNanchor_211_211"><span class="label">[211]</span></a> Paris, 1879.</p></div>
+
+<div class="footnote"><p><a name="Footnote_212_212" id="Footnote_212_212"></a><a href="#FNanchor_212_212"><span class="label">[212]</span></a> <i>English literature and society in the eighteenth
+century</i>, London, 1904, p. 187.</p></div>
+
+<div class="footnote"><p><a name="Footnote_213_213" id="Footnote_213_213"></a><a href="#FNanchor_213_213"><span class="label">[213]</span></a> Cf. my paper, "The law of relativity in Ethics,"
+<i>International Journal of Ethics</i>, Vol. <span class="smcap">i</span>. 1891, pp. 37-62.</p></div>
+
+<div class="footnote"><p><a name="Footnote_214_214" id="Footnote_214_214"></a><a href="#FNanchor_214_214"><span class="label">[214]</span></a> <i>Life and Letters</i>, Vol. I. p. 310.</p></div>
+
+<div class="footnote"><p><a name="Footnote_215_215" id="Footnote_215_215"></a><a href="#FNanchor_215_215"><span class="label">[215]</span></a> <i>Ibid.</i> Vol. II. p. 177.</p></div>
+
+<div class="footnote"><p><a name="Footnote_216_216" id="Footnote_216_216"></a><a href="#FNanchor_216_216"><span class="label">[216]</span></a> <i>Life and Letters</i>, Vol. 1. p. 306.</p></div>
+
+<div class="footnote"><p><a name="Footnote_217_217" id="Footnote_217_217"></a><a href="#FNanchor_217_217"><span class="label">[217]</span></a> <i>Life and Letters</i>, p. 307.</p></div>
+</div>
+
+
+<hr style="width: 65%;" />
+<p><span class='pagenum'><a name="Page_223" id="Page_223">[223]</a></span></p>
+<h2><a name="VIII" id="VIII"></a>VIII</h2>
+
+<h2>THE INFLUENCE OF DARWIN UPON RELIGIOUS THOUGHT</h2>
+<p class="center"><span class="smcap1">By P. N. Waggett, M.A., S.S.J.E.</span></p>
+<h3>I</h3>
+<p>The object of this paper is first to point out certain elements of the
+Darwinian influence upon Religious thought, and then to show reason
+for the conclusion that it has been, from a Christian point of view,
+satisfactory. I shall not proceed further to urge that the Christian
+apologetic in relation to biology has been successful. A variety of
+opinions may be held on this question, without disturbing the
+conclusion that the movements of readjustment have been beneficial to
+those who remain Christians, and this by making them more Christian
+and not only more liberal. The theologians may sometimes have
+retreated, but there has been an advance of theology. I know that this
+account incurs the charge of optimism. It is not the worst that could
+be made. The influence has been limited in personal range, unequal,
+even divergent, in operation, and accompanied by the appearance of
+waste and mischievous products. The estimate which follows requires
+for due balance a full development of many qualifying considerations.
+For this I lack space, but I must at least distinguish my view from
+the popular one that our difficulties about religion and natural
+science have come to an end.</p>
+
+<p>Concerning the older questions about origins&mdash;the origin<span class='pagenum'><a name="Page_224" id="Page_224">[224]</a></span> of the
+world, of species, of man, of reason, conscience, religion&mdash;a large
+measure of understanding has been reached by some thoughtful men. But
+meanwhile new questions have arisen, questions about conduct,
+regarding both the reality of morals and the rule of right action for
+individuals and societies. And these problems, still far from
+solution, may also be traced to the influence of Darwin. For they
+arise from the renewed attention to heredity, brought about by the
+search for the causes of variation, without which the study of the
+selection of variations has no sufficient basis.</p>
+
+<p>Even the existing understanding about origins is very far from
+universal. On these points there were always thoughtful men who denied
+the necessity of conflict, and there are still thoughtful men who deny
+the possibility of a truce.</p>
+
+<p>It must further be remembered that the earlier discussion now, as I
+hope to show, producing favourable results, created also for a time
+grave damage, not only in the disturbance of faith and the loss of
+men&mdash;a loss not repaired by a change in the currents of debate&mdash;but in
+what I believe to be a still more serious respect. I mean the
+introduction of a habit of facile and untested hypothesis in religious
+as in other departments of thought.</p>
+
+<p>Darwin is not responsible for this, but he is in part the cause of it.
+Great ideas are dangerous guests in narrow minds; and thus it has
+happened that Darwin&mdash;the most patient of scientific workers, in whom
+hypothesis waited upon research, or if it provisionally outstepped it
+did so only with the most scrupulously careful acknowledgment&mdash;has led
+smaller and less conscientious men in natural science, in history, and
+in theology to an over-eager confidence in probable conjecture and a
+loose grip upon the facts of experience. It is not too much to say
+that in many quarters the age of materialism was the least
+matter-of-fact age conceivable, and the age of science the age which
+showed least of the patient temper of inquiry.</p>
+
+<p>I have indicated, as shortly as I could, some losses and<span class='pagenum'><a name="Page_225" id="Page_225">[225]</a></span> dangers
+which in a balanced account of Darwin's influence would be discussed
+at length.</p>
+
+<p>One other loss must be mentioned. It is a defect in our thought which,
+in some quarters, has by itself almost cancelled all the advantages
+secured. I mean the exaggerated emphasis on uniformity or continuity;
+the unwillingness to rest any part of faith or of our practical
+expectation upon anything that from any point of view can be called
+exceptional. The high degree of success reached by naturalists in
+tracing, or reasonably conjecturing, the small beginnings of great
+differences, has led the inconsiderate to believe that anything may in
+time become anything else.</p>
+
+<p>It is true that this exaggeration of the belief in uniformity has
+produced in turn its own perilous reaction. From refusing to believe
+whatever can be called exceptional, some have come to believe whatever
+can be called wonderful.</p>
+
+<p>But, on the whole, the discontinuous or highly various character of
+experience received for many years too little deliberate attention.
+The conception of uniformity which is a necessity of scientific
+description has been taken for the substance of history. We have
+accepted a postulate of scientific method as if it were a conclusion
+of scientific demonstration. In the name of a generalisation which,
+however just on the lines of a particular method, is the prize of a
+difficult exploit of reflexion, we have discarded the direct
+impressions of experience; or, perhaps it is more true to say, we have
+used for the criticism of alleged experiences a doctrine of uniformity
+which is only valid in the region of abstract science. For every
+science depends for its advance upon limitation of attention, upon the
+selection out of the whole content of consciousness of that part or
+aspect which is measurable by the method of the science. Accordingly
+there is a science of life which rightly displays the unity underlying
+all its manifestations. But there is another view of life, equally
+valid, and practically sometimes more important, which recognises the
+immediate and lasting effect of crisis, difference, and revolution.
+Our ar<span class='pagenum'><a name="Page_226" id="Page_226">[226]</a></span>dour for the demonstration of uniformity of process and of
+minute continuous change needs to be balanced by a recognition of the
+catastrophic element in experience, and also by a recognition of the
+exceptional significance for us of events which may be perfectly
+regular from an impersonal point of view.</p>
+
+<p>An exorbitant jealousy of miracle, revelation, and ultimate moral
+distinctions has been imported from evolutionary science into
+religious thought. And it has been a damaging influence, because it
+has taken men's attention from facts, and fixed them upon theories.</p>
+
+
+<p>II</p>
+
+<p>With this acknowledgment of important drawbacks, requiring many words
+for their proper description, I proceed to indicate certain results of
+Darwin's doctrine which I believe to be in the long run wholly
+beneficial to Christian thought. These are:</p>
+
+<p>The encouragement in theology of that evolutionary method of
+observation and study, which has shaped all modern research:</p>
+
+<p>The recoil of Christian apologetics towards the ground of religious
+experience, a recoil produced by the pressure of scientific criticism
+upon other supports of faith:</p>
+
+<p>The restatement, or the recovery of ancient forms of statement, of the
+doctrines of Creation and of divine Design in Nature, consequent upon
+the discussion of evolution and of natural selection as its guiding
+factor.</p>
+
+<p>(1) The first of these is quite possibly the most important of all. It
+was well defined in a notable paper read by Dr. Gore, now Bishop of
+Birmingham, to the Church Congress at Shrewsbury in 1896. We have
+learnt a new caution both in ascribing and in denying significance to
+items of evidence, in utterance or in event. There has been, as in
+art, a study of values, which secures perspective and solidity in our
+representation of facts. On the one hand, a given utter<span class='pagenum'><a name="Page_227" id="Page_227">[227]</a></span>ance or event
+cannot be drawn into evidence as if all items were of equal
+consequence, like sovereigns in a bag. The question whence and whither
+must be asked, and the particular thing measured as part of a series.
+Thus measured it is not less truly important, but it may be important
+in a lower degree. On the other hand, and for exactly the same reason,
+nothing that is real is unimportant. The "failures" are not mere
+mistakes. We see them, in St. Augustine's words, as "scholar's faults
+which men praise in hope of fruit."</p>
+
+<p>We cannot safely trace the origin of the evolutionistic method to the
+influence of natural science. The view is tenable that theology led
+the way. Probably this is a case of alternate and reciprocal debt.
+Quite certainly the evolutionist method in theology, in Christian
+history and in the estimate of scripture, has received vast
+reinforcement from biology, in which evolution has been the ever
+present and ever victorious conception.</p>
+
+<p>(2) The second effect named is the new willingness of Christian
+thinkers to take definite account of religious experience. This is
+related to Darwin through the general pressure upon religious faith of
+scientific criticism. The great advance of our knowledge of organisms
+has been an important element in the general advance of science. It
+has acted, by the varied requirements of the theory of organisms, upon
+all other branches of natural inquiry, and it held for a long time
+that leading place in public attention which is now occupied by
+speculative physics. Consequently it contributed largely to our
+present estimation of science as the supreme judge in all matters of
+inquiry,<a name="FNanchor_218_218" id="FNanchor_218_218"></a><a href="#Footnote_218_218" class="fnanchor">[218]</a> to the supposed destruction of mystery and the
+disparagement of metaphysics which marked the last age, as well as to
+the just recommendation of scientific method in branches of learning
+where the direct acquisitions of natural science had no place.</p>
+
+<p>Besides this, the new application of the idea of law and <span class='pagenum'><a name="Page_228" id="Page_228">[228]</a></span>mechanical
+regularity to the organic world seemed to rob faith of a kind of
+refuge. The romantics had, as Berthelot<a name="FNanchor_219_219" id="FNanchor_219_219"></a><a href="#Footnote_219_219" class="fnanchor">[219]</a> shows, appealed to life
+to redress the judgments drawn from mechanism. Now, in Spencer,
+evolution gave us a vitalist mechanic or mechanical vitalism, and the
+appeal seemed cut off. We may return to this point later when we
+consider evolution; at present I only endeavour to indicate that
+general pressure of scientific criticism which drove men of faith to
+seek the grounds of reassurance in a science of their own; in a method
+of experiment, of observation, of hypothesis checked by known facts.
+It is impossible for me to do more than glance across the threshold of
+this subject. But it is necessary to say that the method is in an
+elementary stage of revival. The imposing success that belongs to
+natural science is absent: we fall short of the unchallengeable
+unanimity of the Biologists on fundamentals. The experimental method
+with its sure repetitions cannot be applied to our subject-matter. But
+we have something like the observational method of palaeontology and
+geographical distribution; and in biology there are still men who
+think that the large examination of varieties by way of geography and
+the search of strata is as truly scientific, uses as genuinely the
+logical method of difference, and is as fruitful in sure conclusions
+as the quasi-chemical analysis of Mendelian laboratory work, of which
+last I desire to express my humble admiration. Religion also has its
+observational work in the larger and possibly more arduous manner.</p>
+
+<p>But the scientific work in religion makes its way through difficulties
+and dangers. We are far from having found the formula of its
+combination with the historical elements of our apologetic. It is
+exposed, therefore, to a damaging fire not only from unspiritualist
+psychology and pathology but also from the side of scholastic dogma.
+It is hard to admit on equal terms a partner to the old undivided rule
+of books and learning. With Charles Lamb, we cry in some distress,
+<span class='pagenum'><a name="Page_229" id="Page_229">[229]</a></span>"must knowledge come to me, if it come at all, by some awkward
+experiment of intuition, and no longer by this familiar process of
+reading?"<a name="FNanchor_220_220" id="FNanchor_220_220"></a><a href="#Footnote_220_220" class="fnanchor">[220]</a> and we are answered that the old process has an
+imperishable value, only we have not yet made clear its connection
+with other contributions. And all the work is young, liable to be
+drawn into unprofitable excursions, side-tracked by self-deceit and
+pretence; and it fatally attracts, like the older mysticism, the
+curiosity and the expository powers of those least in sympathy with
+it, ready writers who, with all the air of extended research, have
+been content with narrow grounds for induction. There is a danger,
+besides, which accompanies even the most genuine work of this science
+and must be provided against by all its serious students. I mean the
+danger of unbalanced introspection both for individuals and for
+societies; of a preoccupation comparable to our modern social
+preoccupation with bodily health; of reflexion upon mental states not
+accompanied by exercise and growth of the mental powers; the danger of
+contemplating will and neglecting work, of analysing conviction and
+not criticising evidence.</p>
+
+<p>Still, in spite of dangers and mistakes, the work remains full of
+hopeful indications, and, in the best examples,<a name="FNanchor_221_221" id="FNanchor_221_221"></a><a href="#Footnote_221_221" class="fnanchor">[221]</a> it is truly
+scientific in its determination to know the very truth, to tell what
+we think, not what we think we ought to think,<a name="FNanchor_222_222" id="FNanchor_222_222"></a><a href="#Footnote_222_222" class="fnanchor">[222]</a> truly scientific
+in its employment of hypothesis and verification, and in growing
+conviction of the reality of its subject-matter through the repeated
+victories of a mastery which advances, like science, in the Baconian
+road of <span class='pagenum'><a name="Page_230" id="Page_230">[230]</a></span>obedience. It is reasonable to hope that progress in this
+respect will be more rapid and sure when religious study enlists more
+men affected by scientific desire and endowed with scientific
+capacity.</p>
+
+<p>The class of investigating minds is a small one, possibly even smaller
+than that of reflecting minds. Very few persons at any period are able
+to find out anything whatever. There are few observers, few
+discoverers, few who even wish to discover truth. In how many
+societies the problems of philology which face every person who speaks
+English are left unattempted! And if the inquiring or the successfully
+inquiring class of minds is small, much smaller, of course, is the
+class of those possessing the scientific aptitude in an eminent
+degree. During the last age this most distinguished class was to a
+very great extent absorbed in the study of phenomena, a study which
+had fallen into arrears. For we stood possessed, in rudiment, of means
+of observation, means for travelling and acquisition, qualifying men
+for a larger knowledge than had yet been attempted. These were now to
+be directed with new accuracy and ardour upon the fabric and behaviour
+of the world of sense. Our debt to the great masters in physical
+science who overtook and almost outstripped the task cannot be
+measured; and, under the honourable leadership of Ruskin, we may all
+well do penance if we have failed "in the respect due to their great
+powers of thought, or in the admiration due to the far scope of their
+discovery."<a name="FNanchor_223_223" id="FNanchor_223_223"></a><a href="#Footnote_223_223" class="fnanchor">[223]</a> With what miraculous mental energy and divine good
+fortune&mdash;as Romans said of their soldiers&mdash;did our men of curiosity
+face the apparently impenetrable mysteries of nature! And how natural
+it was that immense accessions of knowledge, unrelated to the
+spiritual facts of life, should discredit Christian faith, by the
+apparent superiority of the new work to the feeble and unprogressive
+knowledge of Christian believers! The day is coming when men of this
+mental character and rank, of this curiosity, this energy and this
+good fortune in investigation, <span class='pagenum'><a name="Page_231" id="Page_231">[231]</a></span>will be employed in opening mysteries
+of a spiritual nature. They will silence with masterful witness the
+over-confident denials of naturalism. They will be in danger of the
+widespread recognition which thirty years ago accompanied every
+utterance of Huxley, Tyndall, Spencer. They will contribute, in spite
+of adulation, to the advance of sober religious and moral science.</p>
+
+<p>And this result will be due to Darwin, first because by raising the
+dignity of natural science, he encouraged the development of the
+scientific mind; secondly because he gave to religious students the
+example of patient and ardent investigation; and thirdly because by
+the pressure of naturalistic criticism the religious have been driven
+to ascertain the causes of their own convictions, a work in which they
+were not without the sympathy of men of science.<a name="FNanchor_224_224" id="FNanchor_224_224"></a><a href="#Footnote_224_224" class="fnanchor">[224]</a></p>
+
+<p>In leaving the subject of scientific religious inquiry, I will only
+add that I do not believe it receives any important help&mdash;and
+certainly it suffers incidentally much damaging interruption&mdash;from the
+study of abnormal manifestations or abnormal conditions of
+personality.</p>
+
+<p>(3) Both of the above effects seem to me of high, perhaps the very
+highest, importance to faith and to thought. But, under the third
+head, I name two which are more directly traceable to the personal
+work of Darwin, and more definitely characteristic of the age in which
+his influence was paramount: viz. the influence of the two conceptions
+of evolution and natural selection upon the doctrine of creation and
+of design respectively. <span class='pagenum'><a name="Page_232" id="Page_232">[232]</a></span></p>
+
+<p>It is impossible here, though it is necessary for a complete sketch of
+the matter, to distinguish the different elements and channels of this
+Darwinian influence; in Darwin's own writings, in the vigourous
+polemic of Huxley, and strangely enough, but very actually for popular
+thought, in the teaching of the definitely anti-Darwinian evolutionist
+Spencer.</p>
+
+<p>Under the head of the directly and purely Darwinian elements I should
+class as preeminent the work of Wallace and of Bates; for no two sets
+of facts have done more to fix in ordinary intelligent minds a belief
+in organic evolution and in natural selection as its guiding factor
+than the facts of geographical distribution and of protective colour
+and mimicry. The facts of geology were difficult to grasp and the
+public and theologians heard more often of the imperfection than of
+the extent of the geological record. The witness of embryology,
+depending to a great extent upon microscopic work, was and is beyond
+the appreciation of persons occupied in fields of work other than
+biology.</p>
+
+
+<p>III</p>
+
+<p>From the influence in religion of scientific modes of thought we pass
+to the influence of particular biological conceptions. The former
+effect comes by way of analogy, <span class='pagenum'><a name="Page_233" id="Page_233">[233]</a></span>example, encouragement and challenge;
+inspiring or provoking kindred or similar modes of thought in the
+field of theology; the latter by a collision of opinions upon matters
+of fact or conjecture which seem to concern both science and religion.</p>
+
+<p>In the case of Darwinism the story of this collision is familiar, and
+falls under the heads of evolution and natural selection, the doctrine
+of descent with modification, and the doctrine of its guidance or
+determination by the struggle for existence between related varieties.
+These doctrines, though associated and interdependent, and in popular
+thought not only combined but confused, must be considered separately.
+It is true that the ancient doctrine of Evolution, in spite of the
+ingenuity and ardour of Lamarck, remained a dream tantalising the
+intellectual ambition of naturalists, until the day when Darwin made
+it conceivable by suggesting the machinery of its guidance. And,
+further, the idea of natural selection has so effectively opened the
+door of research and stimulated observation in a score of principal
+directions that, even if the Darwinian explanation became one day much
+less convincing than, in spite of recent criticism, it now is, yet its
+passing, supposing it to pass, would leave the doctrine of Evolution
+immeasurably and permanently strengthened. For in the interests of the
+theory of selection, "F&uuml;r Darwin," as M&uuml;ller wrote, facts have been
+collected which remain in any case evidence of the reality of descent
+with modification.</p>
+
+<p>But still, though thus united in the modern history of convictions,
+though united and confused in the collision of biological and
+traditional opinion, yet evolution and natural selection must be
+separated in theological no less than in biological estimation.
+Evolution seemed inconsistent with Creation; natural selection with
+Providence and Divine design.</p>
+
+<p>Discussion was maintained about these points for many years and with
+much dark heat. It ranged over many par<span class='pagenum'><a name="Page_234" id="Page_234">[234]</a></span>ticular topics and engaged
+minds different in tone, in quality, and in accomplishment. There was
+at most times a degree of misconception. Some naturalists attributed
+to theologians in general a poverty of thought which belonged really
+to men of a particular temper or training. The "timid theism"
+discerned in Darwin by so cautious a theologian as Liddon<a name="FNanchor_225_225" id="FNanchor_225_225"></a><a href="#Footnote_225_225" class="fnanchor">[225]</a> was
+supposed by many biologists to be the necessary foundation of an
+honest Christianity. It was really more characteristic of devout
+<i>naturalists</i> like Philip Henry Gosse, than of religious believers as
+such.<a name="FNanchor_226_226" id="FNanchor_226_226"></a><a href="#Footnote_226_226" class="fnanchor">[226]</a> The study of theologians more considerable and even more
+typically conservative than Liddon does not confirm the description of
+religious intolerance given in good faith, but in serious ignorance,
+by a disputant so acute, so observant and so candid as Huxley.
+Something hid from each other's knowledge the devoted pilgrims in two
+great ways of thought. The truth may be, that naturalists took their
+view of what creation was from Christian men of science who naturally
+looked in their own special studies for the supports and illustrations
+of their religious belief. Of almost every labourious student it may
+be said: "<i>Hic ab arte sua non recessit</i>." And both the believing and
+the denying naturalists, confining habitual attention to a part of
+experience, are apt to affirm and deny with trenchant vigour and
+something of a narrow clearness "<i>Qui respiciunt ad pauca, de facili
+pronunciant</i>."<a name="FNanchor_227_227" id="FNanchor_227_227"></a><a href="#Footnote_227_227" class="fnanchor">[227]</a></p>
+
+<p>Newman says of some secular teachers that "they <span class='pagenum'><a name="Page_235" id="Page_235">[235]</a></span>persuade the world of
+what is false by urging upon it what is true." Of some early opponents
+of Darwin it might be said by a candid friend that, in all sincerity
+of devotion to truth, they tried to persuade the world of what is true
+by urging upon it what is false. If naturalists took their version of
+orthodoxy from amateurs in theology, some conservative Christians,
+instead of learning what evolution meant to its regular exponents,
+took their view of it from celebrated persons, not of the front rank
+in theology or in thought, but eager to take account of public
+movements and able to arrest public attention.</p>
+
+<p>Cleverness and eloquence on both sides certainly had their share in
+producing the very great and general disturbance of men's minds in the
+early days of Darwinian teaching. But by far the greater part of that
+disturbance was due to the practical novelty and the profound
+importance of the teaching itself, and to the fact that the
+controversy about evolution quickly became much more public than any
+controversy of equal seriousness had been for many generations.</p>
+
+<p>We must not think lightly of that great disturbance because it has, in
+some real sense, done its work, and because it is impossible in days
+of more coolness and light, to recover a full sense of its very real
+difficulties.</p>
+
+<p>Those who would know them better should add to the calm records of
+Darwin<a name="FNanchor_228_228" id="FNanchor_228_228"></a><a href="#Footnote_228_228" class="fnanchor">[228]</a> and to the story of Huxley's impassioned championship, all
+that they can learn of George Romanes.<a name="FNanchor_229_229" id="FNanchor_229_229"></a><a href="#Footnote_229_229" class="fnanchor">[229]</a> For his life was absorbed
+in this very struggle and reproduced its stages. It began in a certain
+assured simplicity of biblical interpretation; it went on, through the
+glories and adventures of a paladin in Darwin's train, to the darkness
+and dismay of a man who saw all his most cherished beliefs rendered,
+as he thought, incredible.<a name="FNanchor_230_230" id="FNanchor_230_230"></a><a href="#Footnote_230_230" class="fnanchor">[230]</a> He lived <span class='pagenum'><a name="Page_236" id="Page_236">[236]</a></span>to find the freer faith for
+which process and purpose are not irreconcilable, but necessary to one
+another. His development, scientific, intellectual and moral, was
+itself of high significance; and its record is of unique value to our
+own generation, so near the age of that doubt and yet so far from it;
+certainly still much in need of the caution and courage by which past
+endurance prepares men for new emergencies. We have little enough
+reason to be sure that in the discussions awaiting us we shall do as
+well as our predecessors in theirs. Remembering their endurance of
+mental pain, their ardour in mental labour, the heroic temper and the
+high sincerity of controversialists on either side, we may well speak
+of our fathers in such words of modesty and self-judgment as Drayton
+used when he sang the victors of Agincourt. The progress of biblical
+study, in the departments of Introduction and Exegesis, resulting in
+the recovery of a point of view anciently tolerated if not prevalent,
+has altered some of the conditions of that discussion. In the years
+near 1858, the witness of Scripture was adduced both by Christian
+advocates and their critics as if unmistakably irreconcilable with
+Evolution.</p>
+
+<p>Huxley<a name="FNanchor_231_231" id="FNanchor_231_231"></a><a href="#Footnote_231_231" class="fnanchor">[231]</a> found the path of the blameless naturalist everywhere
+blocked by "Moses": the believer in revelation was generally held to
+be forced to a choice between revealed cosmogony and the scientific
+account of origins. It is not clear how far the change in Biblical
+interpretation is due to natural science, and how far to the vital
+movements of theological study which have been quite independent of
+the controversy about species. It belongs to a general renewal of
+Christian movement, the recovery of a heritage. "Special
+Creation"&mdash;really a biological rather than a <span class='pagenum'><a name="Page_237" id="Page_237">[237]</a></span>theological
+conception,&mdash;seems in its rigid form to have been a recent element
+even in English biblical orthodoxy.</p>
+
+<p>The Middle Ages had no suspicion that religious faith forbad inquiry
+into the natural origination of the different forms of life.
+Bartholomaeus Anglicus, an English Franciscan of the thirteenth
+century, was a mutationist in his way, as Aristotle, "the Philosopher"
+of the Christian Schoolmen, had been in his. So late as the
+seventeenth century, as we learn not only from early proceedings of
+the Royal Society, but from a writer so homely and so regularly pious
+as Walton, the variation of species and "spontaneous" generations had
+no theological bearing, except as instances of that various wonder of
+the world which in devout minds is food for devotion.</p>
+
+<p>It was in the eighteenth century that the harder statement took shape.
+Something in the preciseness of that age, its exaltation of law, its
+cold passion for a stable and measured universe, its cold denial, its
+cold affirmation of the power of God, a God of ice, is the occasion of
+that rigidity of religious thought about the living world which Darwin
+by accident challenged, or rather by one of those movements of genius
+which, Goethe<a name="FNanchor_232_232" id="FNanchor_232_232"></a><a href="#Footnote_232_232" class="fnanchor">[232]</a> declares, are "elevated above all earthly control."</p>
+
+<p>If religious thought in the eighteenth century was aimed at a fixed
+and nearly finite world of spirit, it followed in all these respects
+the secular and critical lead. "La philosophie r&eacute;formatrice du
+XVIII<sup>e</sup> si&egrave;cle<a name="FNanchor_233_233" id="FNanchor_233_233"></a><a href="#Footnote_233_233" class="fnanchor">[233]</a> ramenait la nature et la soci&eacute;t&eacute; &agrave; des
+m&eacute;canismes que la pens&eacute;e r&eacute;fl&eacute;chie peut concevoir et r&eacute;composer." In
+fact, religion in a mechanical age is condemned if it takes any but a
+mechanical tone. Butler's thought was too moving, too vital, too
+evolutionary, for the sceptics of his time. In a rationalist,
+encyclopaedic period, religion also must give hard outline to its
+facts, it must be <span class='pagenum'><a name="Page_238" id="Page_238">[238]</a></span>able to display its secret to any sensible man in
+the language used by all sensible men. Milton's prophetic genius
+furnished the eighteenth century, out of the depth of the passionate
+age before it, with the theological tone it was to need. In spite of
+the austere magnificence of his devotion, he gives to smaller souls a
+dangerous lead. The rigidity of Scripture exegesis belonged to this
+stately but imperfectly sensitive mode of thought. It passed away with
+the influence of the older rationalists whose precise denials matched
+the precise and limited affirmations of the static orthodoxy.</p>
+
+<p>I shall, then, leave the specially biblical aspect of the
+debate&mdash;interesting as it is and even useful, as in Huxley's
+correspondence with the Duke of Argyll and others in 1892<a name="FNanchor_234_234" id="FNanchor_234_234"></a><a href="#Footnote_234_234" class="fnanchor">[234]</a>&mdash;in
+order to consider without complication the permanent elements of
+Christian thought brought into question by the teaching of evolution.</p>
+
+<p>Such permanent elements are the doctrine of God as Creator of the
+universe, and the doctrine of man as spiritual and unique. Upon both
+the doctrine of evolution seemed to fall with crushing force.</p>
+
+<p>With regard to Man I leave out, acknowledging a grave omission, the
+doctrine of the Fall and of Sin. And I do so because these have not
+yet, as I believe, been adequately treated: here the fruitful reaction
+to the stimulus of evolution is yet to come. The doctrine of sin,
+indeed, falls principally within the scope of that discussion which
+has followed or displaced the Darwinian; and without it the Fall
+cannot be usefully considered. For the question about the Fall is a
+question not merely of origins, but of the interpretation of moral
+facts whose moral reality must first be established.</p>
+
+<p>I confine myself therefore to Creation and the dignity of man.</p>
+
+<p>The meaning of evolution, in the most general terms, is that the
+differentiation of forms is not essentially separate from their
+behaviour and use; that if these are within the <span class='pagenum'><a name="Page_239" id="Page_239">[239]</a></span>scope of study, that
+is also; that the world has taken the form we see by movements not
+unlike those we now see in progress; that what may be called proximate
+origins are continuous in the way of force and matter, continuous in
+the way of life, with actual occurrences and actual characteristics.
+All this has no revolutionary bearing upon the question of ultimate
+origins. The whole is a statement about process. It says nothing to
+metaphysicians about cause. It simply brings within the scope of
+observation or conjecture that series of changes which has given their
+special characters to the different parts of the world we see. In
+particular, evolutionary science aspires to the discovery of the
+process or order of the appearance of life itself: if it were to
+achieve its aim it could say nothing of the cause of this or indeed of
+the most familiar occurrences. We should have become spectators or
+convinced historians of an event which, in respect of its cause and
+ultimate meaning, would be still impenetrable.</p>
+
+<p>With regard to the origin of species, supposing life already
+established, biological science has the well founded hopes and the
+measure of success with which we are all familiar. All this has, it
+would seem, little chance of collision with a consistent theism, a
+doctrine which has its own difficulties unconnected with any
+particular view of order or process. But when it was stated that
+species had arisen by processes through which new species were still
+being made, evolutionism came into collision with a statement,
+traditionally religious, that species were formed and fixed once for
+all and long ago.</p>
+
+<p>What is the theological import of such a statement when it is regarded
+as essential to belief in God? Simply that God's activity, with
+respect to the formation of living creatures, ceased at some point in
+past time.</p>
+
+<p>"God rested" is made the touchstone of orthodoxy. And when, under the
+pressure of the evidences, we found ourselves obliged to acknowledge
+and assert the present and persistent power of God, in the maintenance
+and in the continued formation of "types," what happened was the
+abolition of a time-<span class='pagenum'><a name="Page_240" id="Page_240">[240]</a></span>limit. We were forced only to a bolder claim, to
+a theistic language less halting, more consistent, more thorough in
+its own line, as well as better qualified to assimilate and modify
+such schemes as Von Hartmann's philosophy of the Unconscious&mdash;a
+philosophy, by the way, quite intolerant of a merely mechanical
+evolution.<a name="FNanchor_235_235" id="FNanchor_235_235"></a><a href="#Footnote_235_235" class="fnanchor">[235]</a></p>
+
+<p>Here was not the retrenchment of an extravagant assertion, but the
+expansion of one which was faltering and inadequate. The traditional
+statement did not need paring down so as to pass the meshes of a new
+and exacting criticism. It was itself a net meant to surround and
+enclose experience; and we must increase its size and close its mesh
+to hold newly disclosed facts of life. The world, which had seemed a
+fixed picture or model, gained first perspective and then solidity and
+movement. We had a glimpse of organic <i>history</i>; and Christian thought
+became more living and more assured as it met the larger view of life.</p>
+
+<p>However unsatisfactory the new attitude might be to our critics, to
+Christians the reform was positive. What was discarded was a
+limitation, a negation. The movement was essentially conservative,
+even actually reconstructive. For the language disused was a language
+inconsistent with the definitions of orthodoxy; it set bounds to the
+infinite, and by implication withdrew from the creative rule all such
+processes as could be brought within the descriptions of research. It
+ascribed fixity and finality to that "creature" in which an apostle
+taught us to recognise the birth-struggles of an unexhausted progress.
+It tended to banish mystery from the world we see, and to confine it
+to a remote first age.</p>
+
+<p>In the reformed, the restored, language of religion, Creation became
+again not a link in a rational series to complete a circle of the
+sciences, but the mysterious and permanent relation between the
+infinite and the finite, between the moving changes we know in part,
+and the Power, after the fashion <span class='pagenum'><a name="Page_241" id="Page_241">[241]</a></span>of that observation, unknown, which
+is itself "unmoved all motion's source."<a name="FNanchor_236_236" id="FNanchor_236_236"></a><a href="#Footnote_236_236" class="fnanchor">[236]</a></p>
+
+<p>With regard to man it is hardly necessary, even were it possible, to
+illustrate the application of this bolder faith. When the record of
+his high extraction fell under dispute, we were driven to a
+contemplation of the whole of his life, rather than of a part and that
+part out of sight. We remembered again, out of Aristotle, that the
+result of a process interprets its beginnings. We were obliged to read
+the title of such dignity as we may claim, in results and still more
+in aspirations.</p>
+
+<p>Some men still measure the value of great present facts in
+life&mdash;reason and virtue and sacrifice&mdash;by what a self-disparaged
+reason can collect of the meaner rudiments of these noble gifts. Mr.
+Balfour has admirably displayed the discrepancy, in this view, between
+the alleged origin and the alleged authority of reason. Such an
+argument ought to be used not to discredit the confident reason, but
+to illuminate and dignify its dark beginnings, and to show that at
+every step in the long course of growth a Power was at work which is
+not included in any term or in all the terms of the series.</p>
+
+<p>I submit that the more men know of actual Christian teaching, its
+fidelity to the past, and its sincerity in face of discovery, the more
+certainly they will judge that the stimulus of the doctrine of
+evolution has produced in the long run vigour as well as flexibility
+in the doctrine of Creation and of man.</p>
+
+<p>I pass from Evolution in general to Natural Selection.</p>
+
+<p>The character in religious language which I have for short called
+mechanical was not absent in the argument from design as stated before
+Darwin. It seemed to have reference to a world conceived as fixed. It
+pointed, not to the plastic capacity and energy of living matter, but
+to the fixed <span class='pagenum'><a name="Page_242" id="Page_242">[242]</a></span>adaptation of this and that organ to an unchanging place
+or function.</p>
+
+<p>Mr. Hobhouse has given us the valuable phrase "a niche of organic
+opportunity." Such a phrase would have borne a different sense in
+non-evolutionary thought. In that thought, the opportunity was an
+opportunity for the Creative Power, and Design appeared in the
+preparation of the organism to fit the niche. The idea of the niche
+and its occupant growing together from simpler to more complex mutual
+adjustment was unwelcome to this teleology. If the adaptation was
+traced to the influence, through competition, of the environment, the
+old teleology lost an illustration and a proof. For the cogency of the
+proof in every instance depended upon the absence of explanation.
+Where the process of adaptation was discerned, the evidence of Purpose
+or Design was weak. It was strong only when the natural antecedents
+were not discovered, strongest when they could be declared
+undiscoverable.</p>
+
+<p>Paley's favourite word is "Contrivance"; and for him contrivance is
+most certain where production is most obscure. He points out the
+physiological advantage of the <i>valvulae conniventes</i> to man, and the
+advantage for teleology of the fact that they cannot have been formed
+by "action and pressure." What is not due to pressure may be
+attributed to design, and when a "mechanical" process more subtle than
+pressure was suggested, the case for design was so far weakened. The
+cumulative proof from the multitude of instances began to disappear
+when, in selection, a natural sequence was suggested in which all the
+adaptations might be reached by the motive power of life, and
+especially when, as in Darwin's teaching, there was full recognition
+of the reactions of life to the stimulus of circumstance. "The
+organism fits the niche," said the teleologist, "because the Creator
+formed it so as to fit." "The organism fits the niche," said the
+naturalist, "because unless it fitted it could not exist." "It was
+fitted to survive," said the theologian. "It survives because it
+fits," said the selectionist. The two forms of state<span class='pagenum'><a name="Page_243" id="Page_243">[243]</a></span>ment are not
+incompatible; but the new statement, by provision of an ideally
+universal explanation of process, was hostile to a doctrine of purpose
+which relied upon evidences always exceptional however numerous.
+Science persistently presses on to find the universal machinery of
+adaptation in this planet; and whether this be found in selection, or
+in direct-effect, or in vital reactions resulting in large changes, or
+in a combination of these and other factors, it must always be opposed
+to the conception of a Divine Power here and there but not everywhere
+active.</p>
+
+<p>For science, the Divine must be constant, operative everywhere and in
+every quality and power, in environment and in organism, in stimulus
+and in reaction, in variation and in struggle, in hereditary
+equilibrium, and in "the unstable state of species"; equally present
+on both sides of every strain, in all pressures and in all
+resistances, in short in the general wonder of life and the world. And
+this is exactly what the Divine Power must be for religious faith.</p>
+
+<p>The point I wish once more to make is that the necessary readjustment
+of teleology, so as to make it depend upon the contemplation of the
+whole instead of a part, is advantageous quite as much to theology as
+to science. For the older view failed in courage. Here again our
+theism was not sufficiently theistic.</p>
+
+<p>Where results seemed inevitable, it dared not claim them as God-given.
+In the argument from Design it spoke not of God in the sense of
+theology, but of a Contriver, immensely, not infinitely wise and good,
+working within a world, the scene, rather than the ever dependent
+outcome, of His Wisdom; working in such emergencies and opportunities
+as occurred, by forces not altogether within His control, towards an
+end beyond Himself. It gave us, instead of the awful reverence due to
+the Cause of all substance and form, all love and wisdom, a
+dangerously detached appreciation of an ingenuity and benevolence
+meritorious in aim and often surprisingly successful in contrivance.</p>
+
+<p>The old teleology was more useful to science than to reli<span class='pagenum'><a name="Page_244" id="Page_244">[244]</a></span>gion, and
+the design-naturalists ought to be gratefully remembered by
+Biologists. Their search for evidences led them to an eager study of
+adaptations and of minute forms, a study such as we have now an
+incentive to in the theory of Natural Selection. One hardly meets with
+the same ardour in microscopical research until we come to modern
+workers. But the argument from Design was never of great importance to
+faith. Still, to rid it of this character was worth all the stress and
+anxiety of the gallant old war. If Darwin had done nothing else for
+us, we are to-day deeply in his debt for this. The world is not less
+venerable to us now, not less eloquent of the causing mind, rather
+much more eloquent and sacred. But our wonder is not that "the
+underjaw of the swine works under the ground" or in any or all of
+those particular adaptations which Paley collected with so much skill,
+but that a purpose transcending, though resembling, our own purposes,
+is everywhere manifest; that what we live in is a whole, mutually
+sustaining, eventful and beautiful, where the "dead" forces feed the
+energies of life, and life sustains a stranger existence, able in some
+real measure to contemplate the whole, of which, mechanically
+considered, it is a minor product and a rare ingredient. Here, again,
+the change was altogether positive. It was not the escape of a vessel
+in a storm with loss of spars and rigging, not a shortening of sail to
+save the masts and make a port of refuge. It was rather the emergence
+from narrow channels to an open sea. We had propelled the great ship,
+finding purchase here and there for slow and uncertain movement. Now,
+in deep water, we spread large canvas to a favouring breeze.</p>
+
+<p>The scattered traces of design might be forgotten or obliterated. But
+the broad impression of Order became plainer when seen at due distance
+and in sufficient range of effect, and the evidence of love and wisdom
+in the universe could be trusted more securely for the loss of the
+particular calculation of their machinery.</p>
+
+<p>Many other topics of faith are affected by modern biology.<span class='pagenum'><a name="Page_245" id="Page_245">[245]</a></span> In some of
+these we have learnt at present only a wise caution, a wise
+uncertainty. We stand before the newly unfolded spectacle of
+suffering, silenced; with faith not scientifically reassured but still
+holding fast certain other clues of conviction. In many important
+topics we are at a loss. But in others, and among them those I have
+mentioned, we have passed beyond this negative state and find faith
+positively strengthened and more fully expressed.</p>
+
+<p>We have gained also a language and a habit of thought more fit for the
+great and dark problems that remain, less liable to damaging
+conflicts, equipped for more rapid assimilation of knowledge. And by
+this change biology itself is a gainer. For, relieved of fruitless
+encounters with popular religion, it may advance with surer aim along
+the path of really scientific life-study which was reopened for modern
+men by the publication of <i>The Origin of Species</i>.</p>
+
+<p>Charles Darwin regretted that, in following science, he had not done
+"more direct good"<a name="FNanchor_237_237" id="FNanchor_237_237"></a><a href="#Footnote_237_237" class="fnanchor">[237]</a> to his fellow-creatures. He has, in fact,
+rendered substantial service to interests bound up with the daily
+conduct and hopes of common men; for his work has led to improvements
+in the preaching of the Christian faith.</p>
+
+<div class="footnotes"><h3>FOOTNOTES:</h3>
+
+<div class="footnote"><p><a name="Footnote_218_218" id="Footnote_218_218"></a><a href="#FNanchor_218_218"><span class="label">[218]</span></a> F. R. Tennant: "The Being of God in the light of
+Physical Science," in <i>Essays on some theological questions of the
+day</i>. London, 1905.</p></div>
+
+<div class="footnote"><p><a name="Footnote_219_219" id="Footnote_219_219"></a><a href="#FNanchor_219_219"><span class="label">[219]</span></a> <i>Evolutionisme et Platonisme</i>, pp. 45, 46, 47. Paris,
+1908.</p></div>
+
+<div class="footnote"><p><a name="Footnote_220_220" id="Footnote_220_220"></a><a href="#FNanchor_220_220"><span class="label">[220]</span></a> <i>Essays of Elia</i>, "New Year's Eve," p. 41; Ainger's
+edition. London, 1899.</p></div>
+
+<div class="footnote"><p><a name="Footnote_221_221" id="Footnote_221_221"></a><a href="#FNanchor_221_221"><span class="label">[221]</span></a> Such an example is given in Baron F. von H&uuml;gel's
+recently finished book, the result of thirty years' research: <i>The
+Mystical Element of Religion, as studied in Saint Catherine of Genoa
+and her Friends</i>. London, 1908.</p></div>
+
+<div class="footnote"><p><a name="Footnote_222_222" id="Footnote_222_222"></a><a href="#FNanchor_222_222"><span class="label">[222]</span></a> G. Tyrrell, in <i>Mediaevalism</i>, has a chapter which is
+full of the important <i>moral</i> element in a scientific attitude. "The
+only infallible guardian of truth is the spirit of truthfulness."
+<i>Mediaevalism</i>, p. 182, London, 1908.</p></div>
+
+<div class="footnote"><p><a name="Footnote_223_223" id="Footnote_223_223"></a><a href="#FNanchor_223_223"><span class="label">[223]</span></a> <i>Queen of the Air</i>, Preface, p. vii. London, 1906.</p></div>
+
+<div class="footnote"><p><a name="Footnote_224_224" id="Footnote_224_224"></a><a href="#FNanchor_224_224"><span class="label">[224]</span></a> The scientific rank of its writer justifies the
+insertion of the following letter from the late Sir John
+Burdon-Sanderson to me. In the lecture referred to I had described the
+methods of Professor Moseley in teaching Biology as affording a
+suggestion of the scientific treatment of religion.
+</p><p>
+<span class="smcap">Oxford</span>,
+</p><p>
+<i>April 30, 1902</i>.
+</p><p>
+<span class="smcap">Dear Sir</span>:
+</p>
+<div class="blockquot"><p>I feel that I must express to you my thanks for the
+discourse which I had the pleasure of listening to yesterday
+afternoon.
+</p><p>
+I do not mean to say that I was able to follow all that you
+said as to the identity of Method in the two fields of
+Science and Religion, but I recognise that the "mysticism"
+of which you spoke gives us the only way by which the two
+fields can be brought into relation.
+</p><p>
+Among much that was memorable, nothing interested me more
+than what you said of Moseley.
+</p><p>
+No one, I am sure, knew better than you the value of his
+teaching and in what that value consisted. </p></div>
+<p>
+Yours faithfully,
+</p><p>
+J. BURDON-SANDERSON.
+</p></div>
+
+<div class="footnote"><p><a name="Footnote_225_225" id="Footnote_225_225"></a><a href="#FNanchor_225_225"><span class="label">[225]</span></a> H. P. Liddon, <i>The Recovery of S. Thomas</i>; a sermon
+preached in St. Paul's, London, on April 23rd, 1882 (the Sunday after
+Darwin's death).</p></div>
+
+<div class="footnote"><p><a name="Footnote_226_226" id="Footnote_226_226"></a><a href="#FNanchor_226_226"><span class="label">[226]</span></a> Dr. Pusey (<i>Unscience not Science adverse to Faith</i>,
+1878) writes: "The questions as to 'species,' of what variations the
+animal world is capable, whether the species be more or fewer, whether
+accidental variations may become hereditary ... and the like,
+naturally fall under the province of science. In all these questions
+Mr. Darwin's careful observations gained for him a deserved
+approbation and confidence."</p></div>
+
+<div class="footnote"><p><a name="Footnote_227_227" id="Footnote_227_227"></a><a href="#FNanchor_227_227"><span class="label">[227]</span></a> Aristotle, in Bacon, quoted by Newman in his <i>Idea of a
+University</i>, p. 78. London, 1873.</p></div>
+
+<div class="footnote"><p><a name="Footnote_228_228" id="Footnote_228_228"></a><a href="#FNanchor_228_228"><span class="label">[228]</span></a> <i>Life and Letters</i> and <i>More Letters of Charles
+Darwin.</i></p></div>
+
+<div class="footnote"><p><a name="Footnote_229_229" id="Footnote_229_229"></a><a href="#FNanchor_229_229"><span class="label">[229]</span></a> <i>Life and Letters</i>, London, 1896. <i>Thoughts on
+Religion</i>, London, 1895. <i>Candid Examination of Theism</i>, London,
+1878.</p></div>
+
+<div class="footnote"><p><a name="Footnote_230_230" id="Footnote_230_230"></a><a href="#FNanchor_230_230"><span class="label">[230]</span></a> "Never in the history of man has so terrific a calamity
+befallen the race as that which all who look may now (viz. in
+consequence of the scientific victory of Darwin) behold advancing as a
+deluge black with destruction, resistless in might, uprooting our most
+cherished hopes, engulphing our most precious creed, and burying our
+highest life in mindless destruction."&mdash;<i>A Candid Examination of
+Theism</i>, p. 51.</p></div>
+
+<div class="footnote"><p><a name="Footnote_231_231" id="Footnote_231_231"></a><a href="#FNanchor_231_231"><span class="label">[231]</span></a> <i>Science and Christian Tradition.</i> London, 1904.</p></div>
+
+<div class="footnote"><p><a name="Footnote_232_232" id="Footnote_232_232"></a><a href="#FNanchor_232_232"><span class="label">[232]</span></a> "No productiveness of the highest kind ... is in the
+power of anyone."&mdash;<i>Conversations of Goethe with Eckermann and Soret</i>.
+London, 1850.</p></div>
+
+<div class="footnote"><p><a name="Footnote_233_233" id="Footnote_233_233"></a><a href="#FNanchor_233_233"><span class="label">[233]</span></a> Berthelot, <i>Evolutionisme et Platonisme</i>, Paris, 1908,
+p. 45.</p></div>
+
+<div class="footnote"><p><a name="Footnote_234_234" id="Footnote_234_234"></a><a href="#FNanchor_234_234"><span class="label">[234]</span></a> <i>Times</i>, 1892, <i>passim.</i></p></div>
+
+<div class="footnote"><p><a name="Footnote_235_235" id="Footnote_235_235"></a><a href="#FNanchor_235_235"><span class="label">[235]</span></a> See Von Hartmann's <i>Wahrheit und Irrthum in
+Darwinismus</i>. Berlin, 1875.</p></div>
+
+<div class="footnote"><p><a name="Footnote_236_236" id="Footnote_236_236"></a><a href="#FNanchor_236_236"><span class="label">[236]</span></a> Hymn of the Church&mdash;
+</p>
+<div class="poem"><div class="stanza">
+<span class="i0">Rerum Deus tenax vigor,<br /></span>
+<span class="i0">Immotus in te permanens.<br /></span>
+</div></div>
+</div>
+
+<div class="footnote"><p><a name="Footnote_237_237" id="Footnote_237_237"></a><a href="#FNanchor_237_237"><span class="label">[237]</span></a> <i>Life and Letters</i>, Vol. <span class="smcap">iii</span>. p. 359.</p></div>
+</div>
+
+
+<hr style="width: 65%;" />
+<p><span class='pagenum'><a name="Page_246" id="Page_246">[246]</a></span></p>
+<h2><a name="IX" id="IX"></a>IX</h2>
+
+<h2>DARWINISM AND HISTORY</h2>
+<p class="center"><span class="smcap1">By J. B. Bury, LITT.D., LL.D.</span></p>
+<h4><i>Regius Professor of Modern History in the University of Cambridge</i></h4>
+
+<p>1. Evolution, and the principles associated with the Darwinian theory,
+could not fail to exert a considerable influence on the studies
+connected with the history of civilised man. The speculations which
+are known as "philosophy of history," as well as the sciences of
+anthropology, ethnography, and sociology (sciences which though they
+stand on their own feet are for the historian auxiliary), have been
+deeply affected by these principles. Historiographers, indeed, have
+with few exceptions made little attempt to apply them; but the growth
+of historical study in the nineteenth century has been determined and
+characterised by the same general principle which has underlain the
+simultaneous developments of the study of nature, namely the <i>genetic
+idea</i>. The "historical" conception of nature, which has produced the
+history of the solar system, the story of the earth, the genealogies
+of telluric organisms, and has revolutionised natural science, belongs
+to the same order of thought as the conception of human history as a
+continuous, genetic, causal process&mdash;a conception which has
+revolutionised historical research and made it scientific. Before
+proceeding to consider the application of evolutional principles, it
+will be pertinent to notice the rise of this new view.</p>
+
+<p>2. With the Greeks and Romans history had been either a descriptive
+record or had been written in practical inter<span class='pagenum'><a name="Page_247" id="Page_247">[247]</a></span>ests. The most eminent
+of the ancient historians were pragmatical; that is, they regarded
+history as an instructress in statesmanship, or in the art of war, or
+in morals. Their records reached back such a short way, their
+experience was so brief, that they never attained to the conception of
+continuous process, or realised the significance of time; and they
+never viewed the history of human societies as a phenomenon to be
+investigated for its own sake. In the middle ages there was still less
+chance of the emergence of the ideas of progress and development. Such
+notions were excluded by the fundamental doctrines of the dominant
+religion which bounded and bound men's minds. As the course of history
+was held to be determined from hour to hour by the arbitrary will of
+an extra cosmic person, there could be no self-contained causal
+development, only a dispensation imposed from without. And as it was
+believed that the world was within no great distance from the end of
+this dispensation, there was no motive to take much interest in
+understanding the temporal, which was to be only temporary.</p>
+
+<p>The intellectual movements of the fifteenth and sixteenth centuries
+prepared the way for a new conception, but it did not emerge
+immediately. The historians of the Renaissance period simply reverted
+to the ancient pragmatical view. For Machiavelli, exactly as for
+Thucydides and Polybius, the use of studying history was instruction
+in the art of politics. The Renaissance itself was the appearance of a
+new culture, different from anything that had gone before; but at the
+time men were not conscious of this; they saw clearly that the
+traditions of classical antiquity had been lost for a long period, and
+they were seeking to revive them, but otherwise they did not perceive
+that the world had moved, and that their own spirit, culture, and
+conditions were entirely unlike those of the thirteenth century. It
+was hardly till the seventeenth century that the presence of a new
+age, as different from the middle ages as from the ages of Greece and
+Rome, was fully realised. It was then that the triple division of
+ancient, medieval, and modern was first applied<span class='pagenum'><a name="Page_248" id="Page_248">[248]</a></span> to the history of
+western civilisation. Whatever objections may be urged against this
+division, which has now become almost a category of thought, it marks
+a most significant advance in man's view of his own past. He has
+become conscious of the immense changes in civilisation which have
+come about slowly in the course of time, and history confronts him
+with a new aspect. He has to explain how those changes have been
+produced, how the transformations were effected. The appearance of
+this problem was almost simultaneous with the rise of rationalism, and
+the great historians and thinkers of the eighteenth century, such as
+Montesquieu, Voltaire, Gibbon, attempted to explain the movement of
+civilisation by purely natural causes. These brilliant writers
+prepared the way for the genetic history of the following century. But
+in the spirit of the <i>Aufkl&auml;rung</i>, that eighteenth-century
+Enlightenment to which they belonged, they were concerned to judge all
+phenomena before the tribunal of reason; and the apotheosis of
+"reason" tended to foster a certain superior <i>a priori</i> attitude,
+which was not favourable to objective treatment and was incompatible
+with a "historical sense." Moreover the traditions of pragmatical
+historiography had by no means disappeared.</p>
+
+<p>3. In the first quarter of the nineteenth century the meaning of
+genetic history was fully realised. "Genetic" perhaps is as good a
+word as can be found for the conception which in this century was
+applied to so many branches of knowledge in the spheres both of nature
+and of mind. It does not commit us to the doctrine proper of
+evolution, nor yet to any teleological hypothesis such as is implied
+in "progress." For history it meant that the present condition of the
+human race is simply and strictly the result of a causal series (or
+set of causal series)&mdash;a continuous succession of changes, where each
+state arises causally out of the preceding; and that the business of
+historians is to trace this genetic process, to explain each change,
+and ultimately to grasp' the complete development of the life of
+humanity. Three influential writers, who appeared at this stage and<span class='pagenum'><a name="Page_249" id="Page_249">[249]</a></span>
+helped to initiate a new period of research, may specially be
+mentioned. Ranke in 1824 definitely repudiated the pragmatical view
+which ascribes to history the duties of an instructress, and with no
+less decision renounced the function, assumed by the historians of the
+<i>Aufkl&auml;rung</i>, to judge the past; it was his business, he said, merely
+to show how things really happened. Niebuhr was already working in the
+same spirit and did more than any other writer to establish the
+principle that historical transactions must be related to the ideas
+and conditions of their age. Savigny about the same time founded the
+"historical school" of law. He sought to show that law was not the
+creation of an enlightened will, but grew out of custom and was
+developed by a series of adaptations and rejections, thus applying the
+conception of evolution. He helped to diffuse the notion that all the
+institutions of a society or a nation are as closely interconnected as
+the parts of a living organism.</p>
+
+<p>4. The conception of the history of man as a causal development meant
+the elevation of historical inquiry to the dignity of a science. Just
+as the study of bees cannot become scientific so long as the student's
+interest in them is only to procure honey or to derive moral lessons
+from the labours of "the little busy bee," so the history of human
+societies cannot become the object of pure scientific investigation so
+long as man estimates its value in pragmatical scales. Nor can it
+become a science until it is conceived as lying entirely within a
+sphere in which the law of cause and effect has unreserved and
+unrestricted dominion. On the other hand, once history is envisaged as
+a causal process, which contains within itself the explanation of the
+development of man from his primitive state to the point which he has
+reached, such a process necessarily becomes the object of scientific
+investigation and the interest in it is scientific curiosity.</p>
+
+<p>At the same time, the instruments were sharpened and refined. Here
+Wolf, a philologist with historical instinct, was a pioneer. His
+<i>Prolegomena</i> to Homer (1795) an<span class='pagenum'><a name="Page_250" id="Page_250">[250]</a></span>nounced new modes of attack.
+Historical investigation was soon transformed by the elaboration of
+new methods.</p>
+
+<p>5. "Progress" involves a judgment of value, which is not involved in
+the conception of history as a genetic process. It is also an idea
+distinct from that of evolution. Nevertheless it is closely related to
+the ideas which revolutionised history at the beginning of the last
+century; it swam into men's ken simultaneously; and it helped
+effectively to establish the notion of history as a continuous process
+and to emphasise the significance of time. Passing over earlier
+anticipations, I may point to a <i>Discours</i> of Turgot (1750), where
+history is presented as a process in which "the total mass of the
+human race" "marches continually though sometimes slowly to an ever
+increasing perfection." That is a clear statement of the conception
+which Turgot's friend Condorcet elaborated in the famous work,
+published in 1795, <i>Esquisse d'un tableau historique des progr&egrave;s de
+l'esprit humain</i>. This work first treated with explicit fulness the
+idea to which a leading role was to fall in the ideology of the
+nineteenth century. Condorcet's book reflects the triumphs of the
+<i>Tiers &eacute;tat</i>, whose growing importance had also inspired Turgot; it
+was the political changes in the eighteenth century which led to the
+doctrine, emphatically formulated by Condorcet, that the masses are
+the most important element in the historical process. I dwell on this
+because, though Condorcet had no idea of evolution, the predominant
+importance of the masses was the assumption which made it possible to
+apply evolutional principles to history. And it enabled Condorcet
+himself to maintain that the history of civilisation, a progress still
+far from being complete, was a development conditioned by general
+laws.</p>
+
+<p>6. The assimilation of society to an organism, which was a governing
+notion in the school of Savigny, and the conception of progress,
+combined to produce the idea of an organic development, in which the
+historian has to determine the central principle or leading character.
+This is illustrated by the apotheosis of democracy in Tocqueville's<span class='pagenum'><a name="Page_251" id="Page_251">[251]</a></span>
+<i>D&eacute;mocratie en Am&eacute;rique</i>, where the theory is maintained that "the
+gradual and progressive development of equality is at once the past
+and the future of the history of men." The same two principles are
+combined in the doctrine of Spencer (who held that society is an
+organism, though he also contemplated its being what he calls a
+"super-organic aggregate"),<a name="FNanchor_238_238" id="FNanchor_238_238"></a><a href="#Footnote_238_238" class="fnanchor">[238]</a> that social evolution is a
+progressive change from militarism to industrialism.</p>
+
+<p>7. The idea of development assumed another form in the speculations of
+German idealism. Hegel conceived the successive periods of history as
+corresponding to the ascending phases or ideas in the self-evolution
+of his Absolute Being. His <i>Lectures on the Philosophy of History</i>
+were published in 1837 after his death. His philosophy had a
+considerable effect, direct and indirect, on the treatment of history
+by historians, and although he was superficial and unscientific
+himself in dealing with historical phenomena, he contributed much
+towards making the idea of historical development familiar. Ranke was
+influenced, if not by Hegel himself, at least by the Idealistic
+philosophies of which Hegel's was the greatest. He was inclined to
+conceive the stages in the process of history as marked by
+incarnations, as it were, of ideas, and sometimes speaks as if the
+ideas were independent forces, with hands and feet. But while Hegel
+determined his ideas by <i>a priori</i> logic, Ranke obtained his by
+induction&mdash;by a strict investigation of the phenomena; so that he was
+scientific in his method and work, <span class='pagenum'><a name="Page_252" id="Page_252">[252]</a></span>and was influenced by Hegelian
+prepossessions only in the kind of significance which he was disposed
+to ascribe to his results. It is to be noted that the theory of Hegel
+implied a judgment of value; the movement was a progress towards
+perfection.</p>
+
+<p>8. In France, Comte approached the subject from a different side, and
+exercised, outside Germany, a far wider influence than Hegel. The 4th
+volume of his <i>Cours de philosophie positive</i>, which appeared in 1839,
+created sociology and treated history as a part of this new science,
+namely as "social dynamics." Comte sought the key for unfolding
+historical development, in what he called the social-psychological
+point of view, and he worked out the two ideas which had been
+enunciated by Condorcet: that the historian's attention should be
+directed not, as hitherto, principally to eminent individuals, but to
+the collective behaviour of the masses, as being the most important
+element in the process; and that, as in nature, so in history, there
+are general laws, necessary and constant, which condition the
+development. The two points are intimately connected, for it is only
+when the masses are moved into the foreground that regularity,
+uniformity, and law can be conceived as applicable. To determine the
+social-psychological laws which have controlled the development is,
+according to Comte, the task of sociologists and historians.</p>
+
+<p>9. The hypothesis of general laws operative in history was carried
+further in a book which appeared in England twenty years later and
+exercised an influence in Europe far beyond its intrinsic merit,
+Buckle's <i>History of Civilisation in England</i> (1857-61). Buckle owed
+much to Comte, and followed him, or rather outdid him, in regarding
+intellect as the most important factor conditioning the upward
+development of man, so that progress, according to him, consisted in
+the victory of the intellectual over the moral laws.</p>
+
+<p>10. The tendency of Comte and Buckle to assimilate history to the
+sciences of nature by reducing it to general "laws," derived stimulus
+and plausibility from the vista offered by<span class='pagenum'><a name="Page_253" id="Page_253">[253]</a></span> the study of statistics,
+in which the Belgian Quetelet, whose book <i>Sur l'homme</i> appeared in
+1835, discerned endless possibilities. The astonishing uniformities
+which statistical inquiry disclosed led to the belief that it was only
+a question of collecting a sufficient amount of statistical material,
+to enable us to predict how a given social group will act in a
+particular case. Bourdeau, a disciple of this school, looks forward to
+the time when historical science will become entirely quantitative.
+The actions of prominent individuals, which are generally considered
+to have altered or determined the course of things, are obviously not
+amenable to statistical computation or explicable by general laws.
+Thinkers like Buckle sought to minimise their importance or explain
+them away.</p>
+
+<p>11. These indications may suffice to show that the new efforts to
+interpret history which marked the first half of the nineteenth
+century were governed by conceptions closely related to those which
+were current in the field of natural science and which resulted in the
+doctrine of evolution. The genetic principle, progressive development,
+general laws, the significance of time, the conception of society as
+an organic aggregate, the metaphysical theory of history as the
+self-evolution of spirit,&mdash;all these ideas show that historical
+inquiry had been advancing independently on somewhat parallel lines to
+the sciences of nature. It was necessary to bring this out in order to
+appreciate the influence of Darwinism.</p>
+
+<p>12. In the course of the dozen years which elapsed between the
+appearances of <i>The Origin of Species</i> (observe that the first volume
+of Buckle's work was published just two years before) and of <i>The
+Descent of Man</i> (1871), the hypothesis of Lamarck that man is the
+co-descendant with other species of some lower extinct form was
+admitted to have been raised to the rank of an established fact by
+most thinkers whose brains were not working under the constraint of
+theological authority.</p>
+
+<p>One important effect of the discovery of this fact (I am not speaking
+now of the Darwinian explanation) was to assign<span class='pagenum'><a name="Page_254" id="Page_254">[254]</a></span> to history a definite
+place in the coordinated whole of knowledge, and relate it more
+closely to other sciences. It had indeed a defined logical place in
+systems such as Hegel's and Comte's; but Darwinism certified its
+standing convincingly and without more ado. The prevailing doctrine
+that man was created <i>ex abrupto</i> had placed history in an isolated
+position, disconnected with the sciences of nature. Anthropology,
+which deals with the animal <i>anthropos</i>, now comes into line with
+zoology, and brings it into relation with history.<a name="FNanchor_239_239" id="FNanchor_239_239"></a><a href="#Footnote_239_239" class="fnanchor">[239]</a> Man's
+condition at the present day is the result of a series of
+transformations, going back to the most primitive phase of society,
+which is the ideal (unattainable) beginning of history. But that
+beginning had emerged without any breach of continuity from a
+development which carries us back to a quadrimane ancestor, still
+further back (according to Darwin's conjecture) to a marine animal of
+the ascidian type, and then through remoter periods to the lowest form
+of organism. It is essential in this theory that though links have
+been lost there was no break in the gradual development; and this
+conception of a continuous progress in the evolution of life,
+resulting in the appearance of uncivilised Anthropos, helped to
+reinforce, and increase a belief in, the conception of the history of
+civilised Anthropos as itself also a continuous progressive
+development.</p>
+
+<p>13. Thus the diffusion of the Darwinian theory of the origin of man,
+by emphasising the idea of continuity and breaking down the barriers
+between the human and animal <span class='pagenum'><a name="Page_255" id="Page_255">[255]</a></span>kingdoms, has had an important effect in
+establishing the position of history among the sciences which deal
+with telluric development. The perspective of history is merged in a
+larger perspective of development. As one of the objects of biology is
+to find the exact steps in the genealogy of man from the lowest
+organic form, so the scope of history is to determine the stages in
+the unique causal series from the most rudimentary to the present
+state of human civilisation.</p>
+
+<p>It is to be observed that the interest in historical research implied
+by this conception need not be that of Comte. In the Positive
+Philosophy history is part of sociology; the interest in it is to
+discover the sociological laws. In the view of which I have just
+spoken, history is permitted to be an end in itself; the
+reconstruction of the genetic process is an independent interest. For
+the purpose of the reconstruction, sociology, as well as physical
+geography, biology, psychology, is necessary; the sociologist and the
+historian play into each other's hands; but the object of the former
+is to establish generalisations; the aim of the latter is to trace in
+detail a singular causal sequence.</p>
+
+<p>14. The success of the evolutional theory helped to discredit the
+assumption or at least the invocation of transcendent causes.
+Philosophically of course it is compatible with theism, but historians
+have for the most part desisted from invoking the naive conception of
+a "god in history" to explain historical movements. A historian may be
+a theist; but, so far as his work is concerned, this particular belief
+is otiose. Otherwise indeed (as was remarked above) history could not
+be a science; for with a <i>deus ex machina</i> who can be brought on the
+stage to solve difficulties scientific treatment is a farce. The
+transcendent element had appeared in a more subtle form through the
+influence of German philosophy. I noticed how Ranke is prone to refer
+to ideas as if they were transcendent existences manifesting
+themselves in the successive movements of history. It is intelligible
+to speak of certain ideas as controlling, in a given period,&mdash;for
+instance, the idea of nationality; but from the scientific point<span class='pagenum'><a name="Page_256" id="Page_256">[256]</a></span> of
+view, such ideas have no existence outside the minds of individuals
+and are purely psychical forces; and a historical "idea," if it does
+not exist in this form, is merely a way of expressing a synthesis of
+the historian himself.</p>
+
+<p>15. From the more general influence of Darwinism on the place of
+history in the system of human knowledge, we may turn to the influence
+of the principles and methods by which Darwin explained development.
+It had been recognised even by ancient writers (such as Aristotle and
+Polybius) that physical circumstances (geography, climate) were
+factors conditioning the character and history of a race or society.
+In the sixteenth century Bodin emphasised these factors, and many
+subsequent writers took them into account. The investigations of
+Darwin, which brought them into the foreground, naturally promoted
+attempts to discover in them the chief key to the growth of
+civilisation. Comte had expressly denounced the notion that the
+biological methods of Lamarck could be applied to social man. Buckle
+had taken account of natural influences, but had relegated them to a
+secondary plane, compared with psychological factors. But the
+Darwinian theory made it tempting to explain the development of
+civilisation in terms of "adaptation to environment," "struggle for
+existence," "natural selection," "survival of the fittest," etc.<a name="FNanchor_240_240" id="FNanchor_240_240"></a><a href="#Footnote_240_240" class="fnanchor">[240]</a></p>
+
+<p>The operation of these principles cannot be denied. Man is still an
+animal, subject to zoological as well as mechanical laws. The dark
+influence of heredity continues to be effective; and psychical
+development had begun in lower organic forms,&mdash;perhaps with life
+itself. The organic and the social struggles for existence are
+manifestations of the same principle. Environment and climatic
+influence must be called in to explain not only the differentiation of
+the great racial sections of humanity, but also the varieties within
+these sub-species and, it may be, the assimilation of distinct
+varieties. <span class='pagenum'><a name="Page_257" id="Page_257">[257]</a></span>Ritter's <i>Anthropogeography</i> has opened a useful line of
+research. But on the other hand, it is urged that, in explaining the
+course of history, these principles do not take us very far, and that
+it is chiefly for the primitive ultra-prehistoric period that they can
+account for human development. It may be said that, so far as concerns
+the actions and movements of men which are the subject of recorded
+history, physical environment has ceased to act mechanically, and in
+order to affect their actions must affect their wills first; and that
+this psychical character of the causal relations substantially alters
+the problem. The development of human societies, it may be argued,
+derives a completely new character from the dominance of the conscious
+psychical element, creating as it does new conditions (inventions,
+social institutions, etc.) which limit and counteract the operation of
+natural selection, and control and modify the influence of physical
+environment. Most thinkers agree now that the chief clews to the
+growth of civilisation must be sought in the psychological sphere.
+Imitation, for instance, is a principle which is probably more
+significant for the explanation of human development than natural
+selection. Darwin himself was conscious that his principles had only a
+very restricted application in this sphere, as is evident from his
+cautious and tentative remarks in the 5th chapter of his <i>Descent of
+Man</i>. He applied natural selection to the growth of the intellectual
+faculties and of the fundamental social instincts, and also to the
+differentiation of the great races or "sub-species" (Caucasian,
+African, etc.) which differ in anthropological character.<a name="FNanchor_241_241" id="FNanchor_241_241"></a><a href="#Footnote_241_241" class="fnanchor">[241]</a> <span class='pagenum'><a name="Page_258" id="Page_258">[258]</a></span></p>
+
+<p>16. But if it is admitted that the governing factors which concern the
+student of social development are of the psychical order, the
+preliminary success of natural science in explaining organic evolution
+by general principles encouraged sociologists to hope that social
+evolution could be explained on general principles also. The idea of
+Condorcet, Buckle, and others, that history could be assimilated to
+the natural sciences was powerfully reinforced, and the notion that
+the actual historical process, and every social movement involved in
+it, can be accounted for by sociological generalisations, so-called
+"laws," is still entertained by many, in one form or another.
+Dissentients from this view do not deny that the generalisations at
+which the sociologist arrives by the comparative method, by the
+analysis of social factors, and by psychological deduction may be an
+aid to the historian; but they deny that such uniformities are laws or
+contain an explanation of the phenomena. They can point to the element
+of chance coincidence. This element must have played a part in the
+events of organic evolution, but it has probably in a larger measure
+helped to determine events in social evolution. The collision of two
+unconnected sequences may be fraught with great results. The sudden
+death of a leader or a marriage without issue, to take simple cases,
+has again and again led to permanent political consequences. More
+emphasis is laid on the decisive actions of individuals, which cannot
+be reduced under generalisations and which deflect the course of
+events. If the significance of the individual will had been
+exaggerated to the neglect of the collective activity of the social
+aggregate before Condorcet, his doctrine tended to eliminate as
+unimportant the roles of prominent men, and by means of this
+elimination it was possible to found sociology. But it may be urged
+that it is patent on the face of history that its course has
+constantly been shaped and modified by the wills of individuals,<a name="FNanchor_242_242" id="FNanchor_242_242"></a><a href="#Footnote_242_242" class="fnanchor">[242]</a>
+which are by no <span class='pagenum'><a name="Page_259" id="Page_259">[259]</a></span>means always the expression of the collective will;
+and that the appearance of such personalities at the given moments is
+not a necessary outcome of the conditions and cannot be deduced. Nor
+is there any proof that, if such and such an individual had not been
+born, some one else would have arisen to do what he did. In some cases
+there is no reason to think that what happened need ever have come to
+pass. In other cases, it seems evident that the actual change was
+inevitable, but in default of the man who initiated and guided it, it
+might have been postponed, and, postponed or not, might have borne a
+different cachet. I may illustrate by an instance which has just come
+under my notice. Modern painting was founded by Giotto, and the
+Italian expedition of Charles VIII, near the close of the sixteenth
+century, introduced into France the fashion of imitating Italian
+painters. But for Giotto and Charles VIII, French painting might have
+been very different. It may be said that "if Giotto had not appeared,
+some other great imitator would have played a role analogous to his,
+and that without Charles VIII there would have been the commerce with
+Italy, which in the long run would have sufficed to place France in
+relation with Italian artists. But the equivalent of Giotto might have
+been deferred for a century and probably would have been different;
+and commercial relations would have required ages to produce the
+<i>rayonnement imitatif</i> of Italian art in France, which the expedition
+of the royal adventurer provoked in a few years."<a name="FNanchor_243_243" id="FNanchor_243_243"></a><a href="#Footnote_243_243" class="fnanchor">[243]</a> Instances
+furnished by political history are simply endless. Can we conjecture
+how events would have moved if the son of Philip of Macedon had been
+an incompetent? The aggressive <span class='pagenum'><a name="Page_260" id="Page_260">[260]</a></span>action of Prussia which astonished
+Europe in 1740 determined the subsequent history of Germany; but that
+action was anything but inevitable; it depended entirely on the
+personality of Frederick the Great.</p>
+
+<p>Hence it may be argued that the action of individual wills is a
+determining and disturbing factor, too significant and effective to
+allow history to be grasped by sociological formulae. The types and
+general forms of development which the sociologist attempts to
+disengage can only assist the historian in understanding the actual
+course of events. It is in the special domains of economic history and
+<i>Culturgeschichte</i> which have come to the front in modern times that
+generalisation is most fruitful, but even in these it may be contended
+that it furnishes only partial explanations.</p>
+
+<p>17. The truth is that Darwinism itself offers the best illustration of
+the insufficiency of general laws to account for historical
+development. The part played by coincidence, and the part played by
+individuals&mdash;limited by, and related to, general social
+conditions&mdash;render it impossible to deduce the course of the past
+history of man or to predict the future. But it is just the same with
+organic development. Darwin (or any other zoologist) could not deduce
+the actual course of evolution from general principles. Given an
+organism and its environment, he could not show that it must evolve
+into a more complex organism of a definite predetermined type; knowing
+what it has evolved into, he could attempt to discover and assign the
+determining causes. General principles do not account for a particular
+sequence; they embody necessary conditions; but there is a chapter of
+accidents too. It is the same in the case of history.</p>
+
+<p>18. Among the evolutional attempts to subsume the course of history
+under general syntheses, perhaps the most important is that of
+Lamprecht, whose "kulturhistorische" attempt to discover and assign
+the determining causes. German history, exhibits the (indirect)
+influence of the Comtist school. It is based upon psychology, which,
+in his views, holds among the sciences of mind
+(<i>Geisteswissenschaften</i>)<span class='pagenum'><a name="Page_261" id="Page_261">[261]</a></span> the same place (that of a
+<i>Grundwissenschaft</i>) which mechanics holds among the sciences of
+nature. History, by the same comparison, corresponds to biology, and,
+according to him, it can only become scientific if it is reduced to
+general concepts (<i>Begriffe</i>). Historical movements and events are of
+a psychical character, and Lamprecht conceives a given phase of
+civilisation as "a collective psychical condition (<i>seelischer
+Gesamtzustand</i>)" controlling the period, "a diapason which penetrates
+all psychical phenomena and thereby all historical events of the
+time."<a name="FNanchor_244_244" id="FNanchor_244_244"></a><a href="#Footnote_244_244" class="fnanchor">[244]</a> He has worked out a series of such phases, "ages of
+changing psychical diapason," in his <i>Deutsche Geschichte</i>, with the
+aim of showing that all the feelings and actions of each age can be
+explained by the diapason; and has attempted to prove that these
+diapasons are exhibited in other social developments, and are
+consequently not singular but typical. He maintains further that these
+ages succeed each other in a definite order; the principle being that
+the collective psychical development begins with the homogeneity of
+all the individual members of a society and, through heightened
+psychical activity, advances in the form of a continually increasing
+differentiation of the individuals (this is akin to the Spencerian
+formula). This process, evolving psychical freedom from psychical
+constraint, exhibits a series of psychical phenomena which define
+successive periods of civilisation. The process depends on two simple
+principles, that no idea can disappear without leaving behind it an
+effect or influence, and that all psychical life, whether in a person
+or a society, means change, the acquisition of new mental contents. It
+follows that the new have to come to terms with the old, and this
+leads to a synthesis which determines the character of a new age.
+Hence the ages of civilisation are defined as the "highest concepts
+for subsuming without exception all psychical phenomena of the
+development of human societies, that is, of all historical
+events."<a name="FNanchor_245_245" id="FNanchor_245_245"></a><a href="#Footnote_245_245" class="fnanchor">[245]</a> Lamprecht deduces the idea of a special <span class='pagenum'><a name="Page_262" id="Page_262">[262]</a></span>historical
+science, which might be called "historical ethnology," dealing with
+the ages of civilisation, and bearing the same relation to
+(descriptive or narrative) history as ethnology to ethnography. Such a
+science obviously corresponds to Comte's social dynamics, and the
+comparative method, on which Comte laid so much emphasis, is the
+principal instrument of Lamprecht.</p>
+
+<p>19. I have dwelt on the fundamental ideas of Lamprecht, because they
+are not yet widely known in England, and because his system is the
+ablest product of the sociological school of historians. It carries
+the more weight as its author himself is a historical specialist, and
+his historical syntheses deserve the most careful consideration. But
+there is much in the process of development which on such assumptions
+is not explained, especially the initiative of individuals. Historical
+development does not proceed in a right line, without the choice of
+diverging. Again and again, several roads are open to it, of which it
+chooses one&mdash;why? On Lamprecht's method, we may be able to assign the
+conditions which limit the psychical activity of men at a particular
+stage of evolution, but within those limits the individual has so many
+options, such a wide room for moving, that the definition of those
+conditions, the "psychical diapasons," is only part of the explanation
+of the particular development. The heel of Achilles in all historical
+speculations of this class has been the role of the individual.</p>
+
+<p>The increasing prominence of economic history has tended to encourage
+the view that history can be explained in terms of general concepts or
+types. Marx and his school based their theory of human development on
+the conditions of production, by which, according to them, all social
+movements and historical changes are entirely controlled. The leading
+part which economic factors play in Lamprecht's system is significant,
+illustrating the fact that economic changes admit most readily this
+kind of treatment, because they have been less subject to direction or
+interference by individual pioneers.</p>
+
+<p>Perhaps it may be thought that the conception of <i>social<span class='pagenum'><a name="Page_263" id="Page_263">[263]</a></span> environment</i>
+(essentially psychical), on which Lamprecht's "psychical diapasons"
+depend, is the most valuable and fertile conception that the historian
+owes to the suggestion of the science of biology&mdash;the conception of
+all particular historical actions and movements as (1) related to and
+conditioned by the social environment, and (2) gradually bringing
+about a transformation of that environment. But no given
+transformation can be proved to be necessary (predetermined). And
+types of development do not represent laws; their meaning and value
+lie in the help they may give to the historian, in investigating a
+certain period of civilisation, to enable him to discover the
+inter-relations among the diverse features which it presents. They
+are, as some one has said, an instrument of heuretic method.</p>
+
+<p>20. The man engaged in special historical researches&mdash;which have been
+pursued unremittingly for a century past, according to scientific
+methods of investigating evidence (initiated by Wolf, Niebuhr,
+Ranke)&mdash;have for the most part worked on the assumptions of genetic
+history or at least followed in the footsteps of those who fully
+grasped the genetic point of view. But their aim has been to collect
+and sift evidence, and determine particular facts; comparatively few
+have given serious thought to the lines of research and the
+speculations which have been considered in this paper. They have been
+reasonably shy of compromising their work by applying theories which
+are still much debated and immature. But historiography cannot
+permanently evade the questions raised by these theories. One may
+venture to say that no historical change or transformation will be
+fully understood until it is explained how social environment acted on
+the individual components of the society (both immediately and by
+heredity), and how the individuals reacted upon their environment. The
+problem is psychical, but it is analogous to the main problem of the
+biologist.</p>
+
+<div class="footnotes"><h3>FOOTNOTES:</h3>
+
+<div class="footnote"><p><a name="Footnote_238_238" id="Footnote_238_238"></a><a href="#FNanchor_238_238"><span class="label">[238]</span></a> A society presents suggestive analogies with an
+organism, but it certainly is not an organism, and sociologists who
+draw inferences from the assumption of its organic nature must fall
+into error. A vital organism and a society are radically distinguished
+by the fact that the individual components of the former, namely the
+cells, are morphologically as well as functionally differentiated,
+whereas the individuals which compose a society are morphologically
+homogeneous and only functionally differentiated. The resemblances and
+the differences are worked out in E. de Majewski's striking book, <i>La
+Science de la Civilisation</i>. Paris. 1908.</p></div>
+
+<div class="footnote"><p><a name="Footnote_239_239" id="Footnote_239_239"></a><a href="#FNanchor_239_239"><span class="label">[239]</span></a> It is to be observed that history is (not only
+different in scope but) not co-extensive with anthropology <i>in time</i>.
+For it deals only with the development of man in societies, whereas
+anthropology includes in its definition the proto-anthropic period
+when <i>anthropos</i> was still non-social, whether he lived in herds like
+the chimpanzee, or alone like the male ourang-outang. (It has been
+well shown by Majewski that congregations&mdash;herds, flocks, packs,
+&amp;c.&mdash;of animals are not <i>societies</i>; the characteristic of a society
+is differentiation of function. Bee hives, ant hills, may be called
+quasi-societies; but in their case the classes which perform distinct
+functions are morphologically different.)</p></div>
+
+<div class="footnote"><p><a name="Footnote_240_240" id="Footnote_240_240"></a><a href="#FNanchor_240_240"><span class="label">[240]</span></a> Recently O. Seeck has applied these principles to the
+decline of Graeco-Roman civilisation in his <i>Untergang der antiken
+Welt</i>, 2 vols., Berlin, 1895, 1901.</p></div>
+
+<div class="footnote"><p><a name="Footnote_241_241" id="Footnote_241_241"></a><a href="#FNanchor_241_241"><span class="label">[241]</span></a> Darwinian formulae may be suggestive by way of analogy.
+For instance, it is characteristic of social advance that a multitude
+of inventions, schemes and plans are framed which are never carried
+out, similar to, or designed for the same end as, an invention or plan
+which is actually adopted because it has chanced to suit better the
+particular conditions of the hour (just as the works accomplished by
+an individual statesman, artist or savant are usually only a residue
+of the numerous projects conceived by his brain). This process in
+which so much abortive production occurs is analogous to elimination
+by natural selection.</p></div>
+
+<div class="footnote"><p><a name="Footnote_242_242" id="Footnote_242_242"></a><a href="#FNanchor_242_242"><span class="label">[242]</span></a> We can ignore here the metaphysical question of
+freewill and determinism. For the character of the individual's brain
+depends in any case on ante-natal accidents and coincidences, and so
+it may be said that the role of individuals ultimately depends on
+chance,&mdash;the accidental coincidence of independent sequences.</p></div>
+
+<div class="footnote"><p><a name="Footnote_243_243" id="Footnote_243_243"></a><a href="#FNanchor_243_243"><span class="label">[243]</span></a> I have taken this example from G. Tarde's <i>La logique
+sociale</i> (p. 403), Paris, 1904, where it is used for quite a different
+purpose.</p></div>
+
+<div class="footnote"><p><a name="Footnote_244_244" id="Footnote_244_244"></a><a href="#FNanchor_244_244"><span class="label">[244]</span></a> <i>Die kulturhistorische Methode</i>, Berlin, 1900, p. 26.</p></div>
+
+<div class="footnote"><p><a name="Footnote_245_245" id="Footnote_245_245"></a><a href="#FNanchor_245_245"><span class="label">[245]</span></a> <i>Ibid.</i> pp. 28, 29.</p></div>
+</div>
+
+
+<hr style="width: 65%;" />
+<p><span class='pagenum'><a name="Page_264" id="Page_264">[264]</a></span></p>
+<h2><a name="X" id="X"></a>X</h2>
+
+<h2>DARWINISM AND SOCIOLOGY</h2>
+<p class="center"><span class="smcap1">By C. Bougl&eacute;</span></p>
+<h4><i>Professor of Social Philosophy in the University of Toulouse and
+  Deputy-Professor at the Sorbonne, Paris</i></h4>
+<p>How has our conception of social phenomena, and of their history, been
+affected by Darwin's conception of Nature and the laws of its
+transformation? To what extent and in what particular respects have
+the discoveries and hypotheses of the author of <i>The Origin of
+Species</i> aided the efforts of those who have sought to construct a
+science of society?</p>
+
+<p>To such a question it is certainly not easy to give any brief or
+precise answer. We find traces of Darwinism almost everywhere.
+Sociological systems differing widely from each other have laid claim
+to its authority; while, on the other hand, its influence has often
+made itself felt only in combination with other influences. The
+Darwinian thread is worked into a hundred patterns along with other
+threads.</p>
+
+<p>To deal with the problem, we must, it seems, first of all distinguish
+the more general conclusions in regard to the evolution of living
+beings, which are the outcome of Darwinism, from the particular
+explanations it offers of the ways and means by which that evolution
+is effected. That is to say, we must, as far as possible, estimate
+separately the influence of Darwin as an evolutionist and Darwin as a
+selectionist.<span class='pagenum'><a name="Page_265" id="Page_265">[265]</a></span></p>
+
+<p>The nineteenth century, said Cournot, has witnessed a mighty effort to
+"r&eacute;int&eacute;grer l'homme dans la nature." From divers quarters there has
+been a methodical reaction against the persistent dualism of the
+Cartesian tradition, which was itself the unconscious heir of the
+Christian tradition. Even the philosophy of the eighteenth century,
+materialistic as were for the most part the tendencies of its leaders,
+seemed to revere man as a being apart, concerning whom laws might be
+formulated <i>&agrave; priori</i>. To bring him down from his pedestal there was
+needed the marked predominance of positive researches wherein no
+account was taken of the "pride of man." There can be no doubt that
+Darwin has done much to familiarise us with this attitude. Take for
+instance the first part of <i>The Descent of Man</i>: it is an accumulation
+of typical facts, all tending to diminish the distance between us and
+our brothers, the lower animals. One might say that the naturalist had
+here taken as his motto, "Whosoever shall exalt himself shall be
+abased; and he that shall humble himself shall be exalted." Homologous
+structures, the survival in man of certain organs of animals, the
+rudiments in the animal of certain human faculties, a multitude of
+facts of this sort, led Darwin to the conclusion that there is no
+ground for supposing that the "king of the universe" is exempt from
+universal laws. Thus belief in the <i>imperium in imperio</i> has been, as
+it were, whittled away by the progress of the naturalistic spirit,
+itself continually strengthened by the conquests of the natural
+sciences. The tendency may, indeed, drag the social sciences into
+overstrained analogies, such, for instance, as the assimilation of
+societies to organisms. But it will, at least, have had the merit of
+helping sociology to shake off the pre-conception that the groups
+formed by men are artificial, and that history is completely at the
+mercy of chance. Some years before the appearance of <i>The Origin of
+Species</i>, August Comte had pointed out the importance, as regards the
+unification of positive knowledge, of the conviction that the social
+world, the last refuge of spiritualism, is itself subject to
+determinism. It cannot be doubted<span class='pagenum'><a name="Page_266" id="Page_266">[266]</a></span> that the movement of thought which
+Darwin's discoveries promoted contributed to the spread of this
+conviction, by breaking down the traditional barrier which cut man off
+from Nature.</p>
+
+<p>But Nature, according to modern naturalists, is no immutable thing: it
+is rather perpetual movement, continual progression. Their discoveries
+batter a breach directly into the Aristotelian notion of species; they
+refuse to see in the animal world a collection of immutable types,
+distinct from all eternity, and corresponding, as Cuvier said, to so
+many particular thoughts of the Creator. Darwin especially
+congratulated himself upon having been able to deal this doctrine the
+<i>coup de gr&acirc;ce</i>: immutability is, he says, his chief enemy; and he is
+concerned to show&mdash;therein following up Lyell's work&mdash;that everything
+in the organic world, as in the inorganic, is explained by insensible
+but incessant transformations. "Nature makes no leaps"&mdash;"Nature knows
+no gaps": these two <i>dicta</i> form, as it were, the two landmarks
+between which Darwin's idea of transformation is worked out. That is
+to say, the development of Darwinism is calculated to further the
+application of the philosophy of Becoming to the study of human
+institutions.</p>
+
+<p>The progress of the natural sciences thus brings unexpected
+reinforcements to the revolution which the progress of historical
+discipline had begun. The first attempt to constitute an actual
+science of social phenomena&mdash;that, namely, of the economists&mdash;had
+resulted in laws which were called natural, and which were believed to
+be eternal and universal, valid for all times and all places. But this
+perpetuality, brother, as Knies said, of the immutability of the old
+zoology, did not long hold out against the ever-swelling tide of the
+historical movement. Knowledge of the transformations that had taken
+place in language, of the early phases of the family, of religion, of
+property, had all favoured the revival of the Heraclitean view:
+&#960;&#7937;&#957;&#964;&#945; &#961;&#949;&#953;&#771;. As to the categories of political economy, it was
+soon to be recognised, as by Lasalle, that they too are only
+historical.<span class='pagenum'><a name="Page_267" id="Page_267">[267]</a></span> The philosophy of history, moreover, gave expression
+under various forms to the same tendency. Hegel declares that "all
+that is real is rational," but at the same time he shows that all that
+is real is ephemeral, and that for history there is nothing fixed
+beneath the sun. It is this sense of universal evolution that Darwin
+came with fresh authority to enlarge. It was in the name of biological
+facts themselves that he taught us to see only slow metamorphoses in
+the history of institutions, and to be always on the outlook for
+survivals side by side with rudimentary forms. Anyone who reads
+<i>Primitive Culture</i>, by Tylor,&mdash;a writer closely connected with
+Darwin&mdash;will be able to estimate the services which these cardinal
+ideas were to render to the social sciences when the age of
+comparative research had succeeded to that of <i>&agrave; priori</i> construction.</p>
+
+<p>Let us note, moreover, that the philosophy of Becoming in passing
+through the Darwinian biology became, as it were, filtered; it got rid
+of those traces of finalism, which, under different forms, it had
+preserved through all the systems of German Romanticism. Even in
+Herbert Spencer, it has been plausibly argued, one can detect
+something of that sort of mystic confidence in forces spontaneously
+directing life, which forms the very essence of those systems. But
+Darwin's observations were precisely calculated to render such an
+hypothesis futile. At first people may have failed to see this; and we
+call to mind the ponderous sarcasms of Flourens when he objected to
+the theory of Natural Selection that it attributed to nature a power
+of free choice. "Nature endowed with will! That was the final error of
+last century; but the nineteenth no longer deals in
+personifications."<a name="FNanchor_246_246" id="FNanchor_246_246"></a><a href="#Footnote_246_246" class="fnanchor">[246]</a> In fact Darwin himself put his readers on
+their guard against the metaphors he was obliged to use. The processes
+by which he explains the survival of the fittest are far from
+<span class='pagenum'><a name="Page_268" id="Page_268">[268]</a></span>affording any indication of the design of some transcendent breeder.
+Nor, if we look closely, do they even imply immanent effort in the
+animal; the sorting out can be brought about mechanically, simply by
+the action of the environment. In this connection Huxley could with
+good reason maintain that Darwin's originality consisted in showing
+how harmonies which hitherto had been taken to imply the agency of
+intelligence and will could be explained without any such
+intervention. So, when later on, objective sociology declares that,
+even when social phenomena are in question, all finalist
+preconceptions must be distrusted if a science is to be constituted,
+it is to Darwin that its thanks are due; he had long been clearing
+paths for it which lay well away from the old familiar road trodden by
+so many theories of evolution.</p>
+
+<p>This anti-finalist doctrine, when fully worked out, was, moreover,
+calculated to aid in the needful dissociation of two notions: that of
+evolution and that of progress. In application to society these had
+long been confounded; and, as a consequence, the general idea seemed
+to be that only one type of evolution was here possible. Do we not
+detect such a view in Comte's sociology, and perhaps even in Herbert
+Spencer's? Whoever, indeed, assumes an end for evolution is naturally
+inclined to think that only one road leads to that end. But those
+whose minds the Darwinian theory has enlightened are aware that the
+transformations of living beings depend primarily upon their
+conditions, and that it is these conditions which are the agents of
+selection from among individual variations. Hence, it immediately
+follows that transformations are not necessarily improvements. Here,
+Darwin's thought hesitated. Logically his theory proves, as Ray
+Lankester pointed out, that the struggle for existence may have as its
+outcome degeneration as well as amelioration: evolution may be
+regressive as well as progressive. Then, too&mdash;and this is especially
+to be borne in mind&mdash;each species takes its good where it finds it,
+seeks its own path and survives as best it can. Apply this notion to<span class='pagenum'><a name="Page_269" id="Page_269">[269]</a></span>
+society and you arrive at the theory of multilinear evolution.
+Divergencies will no longer surprise you. You will be forewarned not
+to apply to all civilisations the same measure of progress, and you
+will recognise that types of evolution may differ just as social
+species themselves differ. Have we not here one of the conceptions
+which mark off sociology proper from the old philosophy of history?</p>
+
+<hr style='width: 45%;' />
+
+<p>But if we are to estimate the influence of Darwinism upon sociological
+conceptions, we must not dwell only upon the way in which Darwin
+impressed the general notion of evolution upon the minds of thinkers.
+We must go into details. We must consider the influence of the
+particular theories by which he explained the mechanism of this
+evolution. The name of the author of <i>The Origin of Species</i> has been
+especially attached, as everyone knows, to the doctrines of "natural
+selection" and of "struggle for existence," completed by the notion of
+"individual variation." These doctrines were turned to account by very
+different schools of social philosophy. Pessimistic and optimistic,
+aristocratic and democratic, individualistic and socialistic systems
+were to war with each other for years by casting scraps of Darwinism
+at each other's heads.</p>
+
+<p>It was the spectacle of human contrivance that suggested to Darwin his
+conception of natural selection. It was in studying the methods of
+pigeon breeders that he divined the processes by which nature, in the
+absence of design, obtains analogous results in the differentiation of
+types. As soon as the importance of artificial selection in the
+transformation of species of animals was understood, reflection
+naturally turned to the human species, and the question arose, How far
+do men observe, in connection with themselves, those laws of which
+they make practical application in the case of animals? Here we come
+upon one of the ideas which guided the researches of Gallon, Darwin's
+cousin. The author of <i>Inquiries into Human Faculty and its
+Development</i>,<a name="FNanchor_247_247" id="FNanchor_247_247"></a><a href="#Footnote_247_247" class="fnanchor">[247]</a> has <span class='pagenum'><a name="Page_270" id="Page_270">[270]</a></span>often expressed his surprise that, considering
+all the precautions taken, for example, in the breeding of horses,
+none whatever are taken in the breeding of the human species. It seems
+to be forgotten that the species suffers when the "fittest" are not
+able to perpetuate their type. Ritchie, in his <i>Darwinism and
+Politics</i><a name="FNanchor_248_248" id="FNanchor_248_248"></a><a href="#Footnote_248_248" class="fnanchor">[248]</a> reminds us of Darwin's remark that the institution of
+the peerage might be defended on the ground that peers, owing to the
+prestige they enjoy, are enabled to select as wives "the most
+beautiful and charming women out of the lower ranks."<a name="FNanchor_249_249" id="FNanchor_249_249"></a><a href="#Footnote_249_249" class="fnanchor">[249]</a> But, says
+Galton, it is as often as not "heiresses" that they pick out, and
+birth statistics seem to show that these are either less robust or
+less fecund than others. The truth is that considerations continue to
+preside over marriage which are entirely foreign to the improvement of
+type, much as this is a condition of general progress. Hence the
+importance of completing Odin's and De Candolle's statistics which are
+designed to show how characters are incorporated in organisms, how
+they are transmitted, how lost, and according to what law eugenic,
+elements depart from the mean or return to it.</p>
+
+<p>But thinkers do not always content themselves with undertaking merely
+the minute researches which the idea of Selection suggests. They are
+eager to defend this or that thesis. In the name of this idea certain
+social anthropologists have recast the conception of the process of
+civilisation, and have affirmed that Social Selection generally works
+against the trend of Natural Selection. Vacher de Lapouge&mdash;following
+up an observation by Broca on the point&mdash;enumerates the various
+institutions, or customs, such as the celibacy of priests and military
+conscription, which cause elimination or sterilisation of the bearers
+of certain superior qualities, intellectual or physical. In a more
+general way he attacks the democratic movement, a movement, as P.
+Bourget says, which is "anti-physical" and contrary to the <span class='pagenum'><a name="Page_271" id="Page_271">[271]</a></span>natural
+laws of progress; though it has been inspired "by the dreams of that
+most visionary of all centuries, the eighteenth."<a name="FNanchor_250_250" id="FNanchor_250_250"></a><a href="#Footnote_250_250" class="fnanchor">[250]</a> The "Equality"
+which levels down and mixes (justly condemned, he holds, by the Comte
+de Gobineau), prevents the aristocracy of the blond dolichocephales
+from holding the position and playing the part which, in the interests
+of all, should belong to them. Otto Ammon, in his <i>Natural Selection
+in Man</i>, and in <i>The Social Order and its Natural Bases</i>,<a name="FNanchor_251_251" id="FNanchor_251_251"></a><a href="#Footnote_251_251" class="fnanchor">[251]</a>
+defended analogous doctrines in Germany; setting the curve
+representing frequency of talent over against that of income, he
+attempted to show that all democratic measures which aim at promoting
+the rise in the social scale of the talented are useless, if not
+dangerous; that they only increase the panmixia, to the great
+detriment of the species and of society.</p>
+
+<p>Among the aristocratic theories which Darwinism has thus inspired we
+must reckon that of Nietzsche. It is well known that in order to
+complete his philosophy he added biological studies to his
+philological; and more than once in his remarks upon the <i>Wille zur
+Macht</i> he definitely alludes to Darwin; though it must be confessed
+that it is generally in order to proclaim the insufficiency of the
+processes by which Darwin seeks to explain the genesis of species.
+Nevertheless, Nietzsche's mind is completely possessed by an ideal of
+Selection. He, too, has a horror of panmixia. The naturalists'
+conception of "the fittest" is joined by him to that of the "hero" of
+romance to furnish a basis for his doctrine of the Superman. Let us
+hasten to add, moreover, that at the very moment when support was
+being sought in the theory of Selection for the various forms of the
+aristocratic doctrine, those same forms were being battered down on
+another side by means of that very theory. Attention was drawn to the
+fact that by virtue of the laws which Darwin himself had <span class='pagenum'><a name="Page_272" id="Page_272">[272]</a></span>discovered
+isolation leads to etiolation. There is a risk that the privilege
+which withdraws the privileged elements of Society from competition
+will cause them to degenerate. In fact, Jacoby in his <i>Studies in
+Selection, in connexion with Heredity in Man</i>,<a name="FNanchor_252_252" id="FNanchor_252_252"></a><a href="#Footnote_252_252" class="fnanchor">[252]</a> concludes that
+"sterility, mental debility, premature death and, finally, the
+extinction of the stock were not specially and exclusively the fate of
+sovereign dynasties; all privileged classes, all families in
+exclusively elevated positions share the fate of reigning families,
+although in a minor degree and in direct proportion to the loftiness
+of their social standing. From the mass of human beings spring
+individuals, families, races, which tend to raise themselves above the
+common level; painfully they climb the rugged heights, attain the
+summits of power, of wealth, of intelligence, of talent, and then, no
+sooner are they there than they topple down and disappear in gulfs of
+mental and physical degeneracy." The demographical researches of
+Hansen<a name="FNanchor_253_253" id="FNanchor_253_253"></a><a href="#Footnote_253_253" class="fnanchor">[253]</a> (following up and completing Dumont's) tended, indeed, to
+show that urban as well as feudal aristocracies, burgher classes as
+well as noble castes, were liable to become effete. Hence it might
+well be concluded that the democratic movement, operating as it does
+to break down class barriers, was promoting instead of impeding human
+selection.</p>
+
+<hr style='width: 45%;' />
+
+<p>So we see that, according to the point of view, very different
+conclusions have been drawn from the application of the Darwinian idea
+of Selection to human society. Darwin's other central idea, closely
+bound up with this, that, namely, of the "struggle for existence" also
+has been diversely utilised. But discussion has chiefly centered upon
+its signification. And while some endeavour to extend its application
+to everything, we find others trying to limit its range. The
+conception of a "struggle for existence" has in the <span class='pagenum'><a name="Page_273" id="Page_273">[273]</a></span>present day been
+taken up into the social sciences from natural science, and adopted.
+But originally it descended from social science to natural. Darwin's
+law is, as he himself said, only Malthus' law generalised and extended
+to the animal world: a growing disproportion between the supply of
+food and the number of the living is the fatal order whence arises the
+necessity of universal struggle, a struggle which, to the great
+advantage of the species, allows only the best equipped individuals to
+survive. Nature is regarded by Huxley as an immense arena where all
+living beings are gladiators.<a name="FNanchor_254_254" id="FNanchor_254_254"></a><a href="#Footnote_254_254" class="fnanchor">[254]</a></p>
+
+<p>Such a generalisation was well adapted to feed the stream of
+pessimistic thought; and it furnished to the apologists of war, in
+particular, new arguments, weighted with all the authority which in
+these days attaches to scientific deliverances. If people no longer
+say, as Bonald did, and Moltke after him, that war is a providential
+fact, they yet lay stress on the point that it is a natural fact. To
+the peace party Dragomirov's objection is urged that its attempts are
+contrary to the fundamental laws of nature, and that no sea wall can
+hold against breakers that come with such gathered force.</p>
+
+<p>But in yet another quarter Darwinism was represented as opposed to
+philanthropic intervention. The defenders of the orthodox political
+economy found in it support for their tenets. Since in the organic
+world universal struggle is the condition of progress, it seemed
+obvious that free competition must be allowed to reign unchecked in
+the economic world. Attempts to curb it were in the highest degree
+imprudent. The spirit of Liberalism here seemed in conformity with the
+trend of nature: in this respect, at least, contemporary naturalism,
+offspring of the discoveries of the nineteenth century, brought
+reinforcements to the individualist doctrine, begotten of the
+speculations of the eighteenth: but only, it appeared, to turn mankind
+away for ever from humanitarian dreams. Would those whom such
+conclusions repelled be <span class='pagenum'><a name="Page_274" id="Page_274">[274]</a></span>content to oppose to nature's imperatives
+only the protests of the heart? There were some who declared, like
+Bruneti&egrave;re, that the laws in question, valid though they might be for
+the animal kingdom, were not applicable to the human. And so a return
+was made to the classic dualism. This indeed seems to be the line that
+Huxley took, when, for instance, he opposed to the cosmic process an
+ethical process which was its reverse.</p>
+
+<p>But the number of thinkers whom this antithesis does not satisfy grows
+daily. Although the pessimism which claims authorisation from Darwin's
+doctrines is repugnant to them, they still are unable to accept the
+dualism which leaves a gulf between man and nature. And their
+endeavour is to link the two by showing that while Darwin's laws
+obtain in both kingdoms, the conditions of their application are not
+the same: their forms, and, consequently, their results, vary with the
+varying mediums in which the struggle of living beings takes place,
+with the means these beings have at disposal, with the ends even which
+they propose to themselves.</p>
+
+<p>Here we have the explanation of the fact that among determined
+opponents of war partisans of the "struggle for existence" can be
+found: there are disciples of Darwin in the peace party. Novicow, for
+example, admits the "<i>combat universel</i>" of which Le Dantec<a name="FNanchor_255_255" id="FNanchor_255_255"></a><a href="#Footnote_255_255" class="fnanchor">[255]</a>
+speaks; but he remarks that at different stages of evolution, at
+different stages of life the same weapons are not necessarily
+employed. Struggles of brute force, armed hand to hand conflicts, may
+have been a necessity in the early phases of human societies.
+Nowadays, although competition may remain inevitable and
+indispensable, it can assume milder forms. Economic rivalries,
+struggles between intellectual influences, suffice to stimulate
+progress: the processes which these admit are, in the actual state of
+civilisation, the only ones which attain their end without waste, the
+only ones logical. From one end to the other of the ladder of life,
+struggle is the order of the day; <span class='pagenum'><a name="Page_275" id="Page_275">[275]</a></span>but more and more as the higher
+rungs are reached, it takes on characters which are proportionately
+more "humane."</p>
+
+<p>Reflections of this kind permit the introduction into the economic
+order of limitations to the doctrine of "laisser faire, laisser
+passer." This appeals, it is said, to the example of nature where
+creatures, left to themselves, struggle without truce and without
+mercy; but the fact is forgotten that upon industrial battlefields the
+conditions are different. The competitors here are not left simply to
+their natural energies: they are variously handicapped. A rich store
+of artificial resources exists in which some participate and others do
+not. The sides then are unequal; and as a consequence the result of
+the struggle is falsified. "In the animal world," said De
+Laveleye,<a name="FNanchor_256_256" id="FNanchor_256_256"></a><a href="#Footnote_256_256" class="fnanchor">[256]</a> criticising Spencer, "the fate of each creature is
+determined by its individual qualities; whereas in civilised societies
+a man may obtain the highest position and the most beautiful wife
+because he is rich and well-born, although he may be ugly, idle or
+improvident; and then it is he who will perpetuate the species. The
+wealthy man, ill constituted, incapable, sickly, enjoys his riches and
+establishes his stock under the protection of the laws." Haycraft in
+England and Jentsch in Germany have strongly emphasised these
+"anomalies," which nevertheless are the rule. That is to say that even
+from a Darwinian point of view all social reforms can readily be
+justified which aim at diminishing, as Wallace said, inequalities at
+the start.</p>
+
+<p>But we can go further still. Whence comes the idea that all measures
+inspired by the sentiment of solidarity are contrary to Nature's
+trend? Observe her carefully, and she will not give lessons only in
+individualism. Side by side with the struggle for existence do we not
+find in operation what Lanessan calls "association for existence."
+Long ago, Espinas had drawn attention to "societies of animals,"
+temporary or permanent, and to the kind of morality that arose in
+them. Since then, naturalists have often insisted upon the importance
+of various forms of symbiosis. Kropotkin <span class='pagenum'><a name="Page_276" id="Page_276">[276]</a></span>in <i>Mutual Aid</i> has chosen
+to enumerate many examples of altruism furnished by animals to
+mankind. Geddes and Thomson went so far as to maintain that "Each of
+the greater steps of progress is in fact associated with an increased
+measure of subordination of individual competition to reproductive or
+social ends, and of interspecific competition to co-operative,
+association."<a name="FNanchor_257_257" id="FNanchor_257_257"></a><a href="#Footnote_257_257" class="fnanchor">[257]</a> Experience shows, according to Geddes, that the
+types which are fittest to surmount great obstacles are not so much
+those who engage in the fiercest competitive struggle for existence,
+as those who contrive to temper it. From all these observations there
+resulted, along with a limitation of Darwinian pessimism, some
+encouragement for the aspirations of the collectivists.</p>
+
+<p>And Darwin himself would, doubtless, have subscribed to these
+rectifications. He never insisted, like his rival, Wallace, upon the
+necessity of the solitary struggle of creatures in a state of nature,
+each for himself and against all. On the contrary, in <i>The Descent of
+Man</i>, he pointed out the serviceableness of the social instincts, and
+corroborated Bagehot's statements when the latter, applying laws of
+physics to politics, showed the great advantage societies derived from
+intercourse and communion. Again, the theory of sexual evolution which
+makes the evolution of types depend increasingly upon preferences,
+judgments, mental factors, surely offers something to qualify what
+seems hard and brutal in the theory of natural selection.</p>
+
+<p>But, as often happens with disciples, the Darwinians had out-Darwined
+Darwin. The extravagances of social Darwinism provoked a useful
+reaction; and thus people were led to seek, even in the animal
+kingdom, for facts of solidarity which would serve to justify humane
+effort.</p>
+
+<hr style='width: 45%;' />
+
+<p>On quite another line, however, an attempt has been made to connect
+socialist tendencies with Darwinian principles. Marx and Darwin have
+been confronted; and writers have <span class='pagenum'><a name="Page_277" id="Page_277">[277]</a></span>undertaken to show that the work of
+the German philosopher fell readily into line with that of the English
+naturalist and was a development of it. Such has been the endeavour of
+Ferri in Italy and of Woltmann in Germany, not to mention others. The
+founders of "scientific socialism" had, moreover, themselves thought
+of this reconciliation. They make more than one allusion to Darwin in
+works which appeared after 1859. And sometimes they use his theory to
+define by contrast their own ideal. They remark that the capitalist
+system, by giving free course to individual competition, ends indeed
+in a <i>bellum omnium contra omnes</i>; and they make it clear that
+Darwinism, thus understood, is as repugnant to them as to D&uuml;hring.</p>
+
+<p>But it is at the scientific and not at the moral point of view that
+they place themselves when they connect their economic history with
+Darwin's work. Thanks to this unifying hypothesis, they claim to have
+constructed&mdash;as Marx does in his preface to <i>Das Kapital</i>&mdash;a veritable
+natural history of social evolution. Engels speaks in praise of his
+friend Marx as having discovered the true mainspring of history hidden
+under the veil of idealism and sentimentalism, and as having
+proclaimed in the <i>primum vivere</i> the inevitableness of the struggle
+for existence. Marx himself, in <i>Das Kapital</i>, indicated another
+analogy when he dwelt upon the importance of a general technology for
+the explanation of this psychology:&mdash;a history of tools which would be
+to social organs what Darwinism is to the organs of animal species.
+And the very importance they attach to tools, to apparatus, to
+machines, abundantly proves that neither Marx nor Engels were likely
+to forget the special characters which mark off the human world from
+the animal. The former always remains to a great extent an artificial
+world. Inventions change the face of its institutions. New modes of
+production revolutionise not only modes of government, but modes even
+of collective thought. Therefore it is that the evolution of society
+is controlled by laws special to it, of which the spectacle of nature
+offers no suggestion.<span class='pagenum'><a name="Page_278" id="Page_278">[278]</a></span></p>
+
+<p>If, however, even in this special sphere, it can still be urged that
+the evolution of the material conditions of society is in accord with
+Darwin's theory, it is because the influence of the methods of
+production is itself to be explained by the incessant strife of the
+various classes with each other. So that in the end Marx, like Darwin,
+finds the source of all progress is in struggle. Both are grandsons of
+Heraclitus:&mdash;&#960;&#8001;&#955;&#949;&#956;&#959;&#962; &#960;&#945;&#964;&#7968;&#961; &#960;&#7937;&#957;&#964;&#969;&#957;. It sometimes happens, in
+these days, that the doctrine of revolutionary socialism is contrasted
+as rude and healthy with what may seem to be the enervating tendency
+of "solidarist" philanthropy: the apologists of the doctrine then
+pride themselves above all upon their faithfulness to Darwinian
+principles.</p>
+
+<hr style='width: 45%;' />
+
+<p>So far we have been mainly concerned to show the use that social
+philosophies have made of the Darwinian laws for practical purposes:
+in order to orientate society towards their ideals each school tries
+to show that the authority of natural science is on its side. But even
+in the most objective of theories, those which systematically make
+abstraction of all political tendencies in order to study the social
+reality in itself, traces of Darwinism are readily to be found.</p>
+
+<p>Let us take for example Durkheim's theory of Division of Labour.<a name="FNanchor_258_258" id="FNanchor_258_258"></a><a href="#Footnote_258_258" class="fnanchor">[258]</a>
+The conclusions he derives from it are that whenever professional
+specialisation causes multiplication of distinct branches of activity,
+we get organic solidarity&mdash;implying differences&mdash;substituted for
+mechanical solidarity, based upon likenesses. The umbilical cord, as
+Marx said, which connects the individual consciousness with the
+collective consciousness is cut. The personality becomes more and more
+emancipated. But on what does this phenomenon, so big with
+consequences, itself depend? The author goes to social morphology for
+the answer: it is, he says, the growing density of population which
+brings with it this increasing differentiation of activities. But,
+again, why? Because the greater density, in thrusting men up against
+each other, <span class='pagenum'><a name="Page_279" id="Page_279">[279]</a></span>augments the intensity of their competition for the means
+of existence; and for the problems which society thus has to face
+differentiation of functions presents itself as the gentlest solution.</p>
+
+<p>Here one sees that the writer borrows directly from Darwin.
+Competition is at its maximum between similars, Darwin had declared;
+different species, not laying claim to the same food, could more
+easily coexist. Here lay the explanation of the fact that upon the
+same oak hundreds of different insects might be found. Other things
+being equal, the same applies to society. He who finds some unadopted
+specialty possesses a means of his own for getting a living. It is by
+this division of their manifold tasks that men contrive not to crush
+each other. Here we obviously have a Darwinian law serving as
+intermediary in the explanation of that progress of division of labour
+which itself explains so much in the social evolution.</p>
+
+<p>And we might take another example, at the other end of the series of
+sociological systems. G. Tarde is a sociologist with the most
+pronounced anti-naturalistic views. He has attempted to show that all
+application of the laws of natural science to society is misleading.
+In his <i>Opposition Universelle</i> he has directly combatted all forms of
+sociological Darwinism. According to him the idea that the evolution
+of society can be traced on the same plan as the evolution of species
+is chimerical. Social evolution is at the mercy of all kinds of
+inventions, which by virtue of the laws of imitation modify, through
+individual to individual, through neighbourhood to neighbourhood, the
+general state of those beliefs and desires which are the only
+"quantities" whose variation matters to the sociologist. But, it may
+be rejoined, that however psychical the forces may be, they are none
+the less subject to Darwinian laws. They compete with each other; they
+struggle for the mastery of minds. Between types of ideas, as between
+organic forms, selection operates. And though it may be that these
+types are ushered into the arena by unexpected discoveries, we yet
+recognise in the<span class='pagenum'><a name="Page_280" id="Page_280">[280]</a></span> psychological accidents, which Tarde places at the
+base of everything, near relatives of those small accidental
+variations upon which Darwin builds. Thus, accepting Tarde's own
+representations, it is quite possible to express in Darwinian terms,
+with the necessary transpositions, one of the most idealistic
+sociologies that have ever been constructed.</p>
+
+<p>These few examples suffice. They enable us to estimate the extent of
+the field of influence of Darwinism. It affects sociology not only
+through the agency of its advocates but through that of its opponents.
+The questions to which it has given rise have proved no less fruitful
+than the solutions it has suggested. In short, few doctrines, in the
+history of social philosophy, will have produced on their passage a
+finer crop of ideas.<span class='pagenum'><a name="Page_281" id="Page_281">[281]</a></span></p>
+
+<div class="footnotes"><h3>FOOTNOTES:</h3>
+
+<div class="footnote"><p><a name="Footnote_246_246" id="Footnote_246_246"></a><a href="#FNanchor_246_246"><span class="label">[246]</span></a> P. Flourens, <i>Examen du Livre de M. Darwin sur
+l'Origine des Esp&egrave;ces</i>, p. 53, Paris, 1864. See also Huxley,
+"Criticisms on the <i>Origin of Species," Collected Essays</i>, Vol. II, p.
+102, London, 1902.</p></div>
+
+<div class="footnote"><p><a name="Footnote_247_247" id="Footnote_247_247"></a><a href="#FNanchor_247_247"><span class="label">[247]</span></a> <i>Inquiries into Human Faculty</i>, pp. 1, 2, 3 sq.,
+London, 1883.</p></div>
+
+<div class="footnote"><p><a name="Footnote_248_248" id="Footnote_248_248"></a><a href="#FNanchor_248_248"><span class="label">[248]</span></a> <i>Darwinism and Politics</i>, pp. 9, 22, London, 1889.</p></div>
+
+<div class="footnote"><p><a name="Footnote_249_249" id="Footnote_249_249"></a><a href="#FNanchor_249_249"><span class="label">[249]</span></a> <i>Life and Letters of Charles Darwin</i>, II. p. 385.</p></div>
+
+<div class="footnote"><p><a name="Footnote_250_250" id="Footnote_250_250"></a><a href="#FNanchor_250_250"><span class="label">[250]</span></a> V. de Lapouge, <i>Les S&eacute;lections sociales</i>, p. 259,
+Paris, 1896.</p></div>
+
+<div class="footnote"><p><a name="Footnote_251_251" id="Footnote_251_251"></a><a href="#FNanchor_251_251"><span class="label">[251]</span></a> <i>Die nat&auml;rliche Auslese beim Menschen</i>, Jena, 1893; <i>Du
+Gesellschaftsordnung und ihre nat&uuml;rlichen Grundlagen. Entwurf einer
+Sozialanthropologie</i>, Jena, 1896.</p></div>
+
+<div class="footnote"><p><a name="Footnote_252_252" id="Footnote_252_252"></a><a href="#FNanchor_252_252"><span class="label">[252]</span></a> <i>Etudes sur la S&eacute;lection dans ses rapports avec
+l'h&eacute;r&eacute;dit&eacute; chez l'homme</i>, Paris, p. 481, 1881.</p></div>
+
+<div class="footnote"><p><a name="Footnote_253_253" id="Footnote_253_253"></a><a href="#FNanchor_253_253"><span class="label">[253]</span></a> <i>Die drei Bev&ouml;lkerungsstufen</i>, Munich, 1889.</p></div>
+
+<div class="footnote"><p><a name="Footnote_254_254" id="Footnote_254_254"></a><a href="#FNanchor_254_254"><span class="label">[254]</span></a> <i>Evolution and Ethics</i>, p. 200; <i>Collected Essays</i>,
+Vol. IX, London, 1894.</p></div>
+
+<div class="footnote"><p><a name="Footnote_255_255" id="Footnote_255_255"></a><a href="#FNanchor_255_255"><span class="label">[255]</span></a> <i>Les Luttes entre Soci&eacute;t&eacute;s humaines et leurs phases
+successives</i>, Paris, 1893.</p></div>
+
+<div class="footnote"><p><a name="Footnote_256_256" id="Footnote_256_256"></a><a href="#FNanchor_256_256"><span class="label">[256]</span></a> <i>Le socialisme contemporain</i>, p. 384 (6th edit.),
+Paris, 1891.</p></div>
+
+<div class="footnote"><p><a name="Footnote_257_257" id="Footnote_257_257"></a><a href="#FNanchor_257_257"><span class="label">[257]</span></a> Geddes and Thomson, <i>The Evolution of Sex</i>, p. 311,
+London, 1889.</p></div>
+
+<div class="footnote"><p><a name="Footnote_258_258" id="Footnote_258_258"></a><a href="#FNanchor_258_258"><span class="label">[258]</span></a> <i>De la Division du Travail social</i>, Paris. 1893.</p></div>
+</div>
+
+
+<hr style="width: 65%;" />
+<h2>INDEX</h2>
+
+
+<div class="index">
+<ul class="IX">
+<li><i>Abraxas grossulariata</i>, <a href="#Page_100">100</a></li>
+
+<li>Acquired characters, transmission of, <a href="#Page_20">20</a>, <a href="#Page_28">28</a>, <a href="#Page_42">42</a>, <a href="#Page_94">94</a>, <a href="#Page_120">120</a>, <a href="#Page_149">149</a>, <a href="#Page_171">171</a>, <a href="#Page_173">173</a></li>
+
+<li><i>Acraea johnstoni</i>, 290</li>
+<li>[Transcriber's Note: No such page number or reference seen.]</li>
+
+<li>Adaptation, <a href="#Page_24">24</a>, <a href="#Page_27">27</a>, <a href="#Page_34">34</a>, <a href="#Page_39">39</a>, <a href="#Page_42">42-45</a>, <a href="#Page_50">50</a>, <a href="#Page_58">58</a>, <a href="#Page_79">79-86</a>, <a href="#Page_106">106</a>, <a href="#Page_107">107</a></li>
+
+<li>Adloff, <a href="#Page_140">140</a></li>
+
+<li>Alexander, <a href="#Page_217">217</a></li>
+
+<li>Ameghino, <a href="#Page_132">132</a>, <a href="#Page_138">138</a></li>
+
+<li>Ammon, O., Works of, <a href="#Page_271">271</a></li>
+
+<li><i>Anaea divina</i>, <a href="#Page_69">69</a></li>
+
+<li>Anglicus, Bartholomaeus, <a href="#Page_237">237</a></li>
+
+<li>Ankyroderma, <a href="#Page_40">40</a></li>
+
+<li>Anomma, <a href="#Page_44">44</a></li>
+
+<li>Anthropops, <a href="#Page_132">132</a></li>
+
+<li>Ants, modifications of, <a href="#Page_43">43-46</a>, <a href="#Page_51">51</a></li>
+
+<li>Ardig&ograve;, <a href="#Page_207">207</a>, <a href="#Page_208">208</a></li>
+
+<li>Argyll, Huxley and the Duke of, <a href="#Page_238">238</a></li>
+
+<li>Aristotle, <a href="#Page_3">3</a>, <a href="#Page_237">237</a>, <a href="#Page_240">240</a></li>
+
+<li>Avenarius, <a href="#Page_211">211</a></li>
+</ul><ul class="IX">
+
+<li>Bacon, on mutability of species, <a href="#Page_4">4</a>, <a href="#Page_5">5</a></li>
+
+<li>Baehr, von, on Cytology, <a href="#Page_99">99</a></li>
+
+<li>Bain, <a href="#Page_194">194</a></li>
+
+<li>Baldwin, J. M., <a href="#Page_53">53</a>, <a href="#Footnote_165_165">Foot Note 165</a></li>
+
+<li>Balfour, A. J., <a href="#Page_241">241</a></li>
+
+<li>Barratt, <a href="#Page_217">217</a></li>
+
+<li>Bates, H. W., on Mimicry, <a href="#Page_70">70</a>, <a href="#Page_76">76</a>
+<ul class="IX">
+<li>  &mdash;<a href="#Page_232">232</a></li>
+</ul></li>
+<li><span class="smcap">Bateson</span>, W., on <i>Heredity and
+Variation in Modern Lights</i>, <a href="#Page_87">87-110</a>
+<ul class="IX">
+<li>  &mdash;on discontinuous evolution, <a href="#Page_30">30</a></li>
+</ul></li>
+<li>Bathmism, <a href="#Page_14">14</a></li>
+
+<li>Bells (Sir Charles) <i>Anatomy of Expression</i>, <a href="#Page_177">177</a></li>
+
+<li>Bentham, Jeremy, <a href="#Page_217">217</a>, <a href="#Page_218">218</a></li>
+
+<li>Bergson, H., <a href="#Page_208">208</a></li>
+
+<li>Berkeley, <a href="#Page_200">200</a></li>
+
+<li>Berthelot, <a href="#Page_228">228</a></li>
+
+<li>Bickford, E., experiments on degeneration by, <a href="#Page_52">52</a></li>
+
+<li>Biophores, <a href="#Page_47">47</a></li>
+
+<li>Blumenbach, <a href="#Page_89">89</a></li>
+
+<li>Bodin, <a href="#Page_256">256</a></li>
+
+<li>Bonald, on war, <a href="#Page_273">273</a></li>
+
+<li>Bonnet, <a href="#Page_6">6</a></li>
+
+<li><span class="smcap">Bougl&eacute;, C.</span>, on <i>Darwinism and Sociology</i>, <a href="#Page_264">264-280</a></li>
+
+<li>Bourdeau, <a href="#Page_253">253</a></li>
+
+<li>Bourget, P., <a href="#Page_270">270</a></li>
+
+<li>Boutroux, <a href="#Page_208">208</a></li>
+
+<li>Brassica, hybrids of, <a href="#Page_106">106</a></li>
+
+<li><i>Brassica Napus</i>, <a href="#Page_106">106</a></li>
+
+<li>Broca, <a href="#Page_137">137</a>, <a href="#Page_270">270</a></li>
+
+<li>Brock, on Kant, <a href="#Footnote_6_6">Foot Note 6</a><span class='pagenum'><a name="Page_282" id="Page_282">[282]</a></span></li>
+
+<li>Bruneti&egrave;re, <a href="#Page_274">274</a></li>
+
+<li>Bruno, on Evolution, <a href="#Page_4">4</a></li>
+
+<li>Buch, von, <a href="#Page_15">15</a></li>
+
+<li>Buckle, <a href="#Page_252">252</a>, <a href="#Page_253">253</a>, <a href="#Page_256">256</a>, <a href="#Page_258">258</a></li>
+
+<li>Buffon, <a href="#Page_6">6-15</a>, <a href="#Page_21">21</a>, <a href="#Page_88">88</a></li>
+
+<li>Burdon-Sanderson, J., letter from, <a href="#Footnote_224_224">Foot Note 224</a></li>
+
+<li><span class="smcap">Bury</span>, J. B., on <i>Darwinism and History</i>, <a href="#Page_246">246-263</a></li>
+
+<li>Butler, Samuel, <a href="#Page_9">9</a>, <a href="#Footnote_17_17">Foot Note 17</a>, <a href="#Footnote_57_57">Foot Note 57</a>, <a href="#Footnote_61_61">Foot Note 61</a>, <a href="#Page_94">94</a>, <a href="#Footnote_66_66">Foot Note 66</a>, <a href="#Page_107">107</a></li>
+
+<li>Butterflies, mimicry in, <a href="#Page_65">65-83</a>
+<ul class="IX">
+<li>  &mdash;sexual characters in, <a href="#Page_59">59-63</a></li>
+</ul></li>
+</ul><ul class="IX">
+
+<li>Cabanis, <a href="#Page_201">201</a></li>
+
+<li>Candolle, de, <a href="#Page_270">270</a></li>
+
+<li>Carneri, <a href="#Page_217">217</a></li>
+
+<li><i>Castnia linus</i>, <a href="#Page_76">76</a></li>
+
+<li>Caterpillars, variation in, <a href="#Page_36">36</a>, <a href="#Page_37">37</a></li>
+
+<li>Cesnola, experiments on Mantis by, <a href="#Page_65">65</a></li>
+
+<li>Chaerocampa, colouring of, <a href="#Page_68">68</a></li>
+
+<li>Chambers, R., <i>The Vestiges of Creation</i> by, <a href="#Page_15">15</a></li>
+
+<li>Chromosomes and Chromomeres, <a href="#Page_47">47</a>, <a href="#Page_96">96-100</a></li>
+
+<li>Chun, <a href="#Footnote_36_36">Foot Note 36</a></li>
+
+<li>Claus, <a href="#Footnote_21_21">Foot Note 21</a></li>
+
+<li>Clodd, E., <a href="#Footnote_13_13">Foot Note 13</a></li>
+
+<li>Coadaptation, <a href="#Page_41">41-54</a></li>
+
+<li><i>Colobopsis truncata</i>, <a href="#Page_44">44</a></li>
+
+<li>Colour, E. B. Poulton, in relation to Sexual Selection, <a href="#Page_61">61-65</a></li>
+
+<li>Comte, A., <a href="#Page_200">200-203</a>, <a href="#Page_252">252-255</a>, <a href="#Page_262">262</a>, <a href="#Page_265">265</a></li>
+
+<li>Condorcet, <a href="#Page_221">221</a>, <a href="#Page_250">250</a>, <a href="#Page_252">252</a>, <a href="#Page_258">258</a></li>
+
+<li>Cope, <a href="#Page_138">138</a></li>
+
+<li>Correlation of organisms, Darwin's idea of the, <a href="#Page_2">2</a></li>
+
+<li>Cournot, <a href="#Page_265">265</a></li>
+
+<li>Cuvier, <a href="#Page_9">9</a>, <a href="#Page_10">10</a>, <a href="#Page_266">266</a>, <a href="#Page_268">268</a></li>
+
+<li>Cytology and heredity, <a href="#Page_95">95</a>, <a href="#Page_96">96</a>, <a href="#Page_99">99</a>, <a href="#Page_100">100</a></li>
+</ul><ul class="IX">
+
+<li><i>Danaida chrysippus</i>, <a href="#Page_75">75</a></li>
+
+<li><i>Danaida genutia</i>, <a href="#Page_75">75</a></li>
+
+<li><i>D. Plexippus</i>, <a href="#Page_75">75</a></li>
+
+<li>Dantec, Le, <a href="#Page_274">274</a></li>
+
+<li>Darwin, Charles, as an Anthropologist, <a href="#Page_146">146-165</a>
+<ul class="IX">
+<li>  &mdash;on ants, <a href="#Page_44">44</a></li>
+<li>  &mdash;and S. Butler, <a href="#Footnote_61_61">Foot Note 61</a>, <a href="#Page_94">94</a></li>
+<li>  &mdash;on Cirripedia, <a href="#Page_212">212</a></li>
+<li>  &mdash;on the Descent of Man, <a href="#Page_111">111-145</a></li>
+<li>  &mdash;evolutionist authors referred to in the <i>Origin</i> by, <a href="#Page_9">9</a></li>
+</ul></li>
+<li>Darwin, Charles, and Haeckel, <a href="#Page_137">137</a>
+<ul class="IX">
+<li>  &mdash;and History, <a href="#Page_246">246-263</a></li>
+<li>  &mdash;and Huxley, <a href="#Page_112">112</a></li>
+<li>  &mdash;on Lamarck, <a href="#Page_28">28</a>, <a href="#Page_129">129</a></li>
+<li>  &mdash;on Language, <a href="#Page_124">124</a></li>
+<li>  &mdash;and Malthus, <a href="#Page_16">16</a>, <a href="#Page_24">24</a>, <a href="#Page_91">91</a></li>
+<li>  &mdash;on Patrick Matthew, <a href="#Page_19">19</a></li>
+<li>  &mdash;on mental evolution, <a href="#Page_166">166-196</a></li>
+<li>  &mdash;on Natural Selection, <a href="#Page_21">21</a>, <a href="#Page_41">41</a>, <a href="#Page_54">54</a>, <a href="#Page_55">55</a>, <a href="#Page_122">122</a></li>
+<li>  &mdash;a "Naturalist for Naturalists," <a href="#Page_87">87</a></li>
+<li>  &mdash;his personality, <a href="#Page_187">187</a></li>
+<li>  &mdash;his influence on Philosophy, <a href="#Page_197">197-222</a></li>
+<li>  &mdash;predecessors of, <a href="#Page_1">1-22</a></li>
+<li>  &mdash;his views on religion, etc., <a href="#Page_115">115</a>, <a href="#Page_116">116</a>, <a href="#Page_219">219-222</a></li>
+<li>  &mdash;his influence on religious thought, <a href="#Page_223">223-245</a></li>
+<li>  &mdash;causes of his success, <a href="#Page_10">10</a>, <a href="#Page_90">90</a><span class='pagenum'><a name="Page_283" id="Page_283">[283]</a></span></li>
+</ul></li>
+
+
+<li>Darwin, Charles, on the <i>Vestiges of Creation</i>, <a href="#Page_15">15</a>
+<ul class="IX">
+<li>  &mdash;and Wallace, <a href="#Page_23">23</a>, <a href="#Page_183">183</a></li>
+<li>  &mdash;on evolution, <a href="#Page_7">7-15</a>, <a href="#Page_88">88</a></li>
+<li>  &mdash;on Lamarckism, <a href="#Page_11">11</a></li>
+</ul></li>
+<li>Darwin, F., on Prichard's "Anticipations," <a href="#Page_21">21</a></li>
+
+<li>Darwinism, Sociology, Evolution and, <a href="#Page_17">17-18</a></li>
+
+<li>Degeneration, <a href="#Page_49">49-51</a>, <a href="#Page_93">93</a></li>
+
+<li>Deniker, <a href="#Page_137">137</a></li>
+
+<li>Descartes, <a href="#Page_4">4</a></li>
+
+<li>Descent, history of doctrine of, <a href="#Page_1">1</a></li>
+
+<li><i>Descent of Man</i>, G. Schwalbe on <i>The</i>, <a href="#Page_111">111-145</a>
+<ul class="IX">
+<li>  &mdash;rejection in Germany of <i>The</i>, <a href="#Page_156">156</a></li>
+</ul></li>
+<li>Diderot, <a href="#Page_6">6</a>, <a href="#Page_198">198</a></li>
+
+<li>Dimorphism, seasonal, <a href="#Page_30">30</a></li>
+
+<li><i>Dismorphia orise</i>, <a href="#Page_75">75</a></li>
+
+<li>Dragomirov, <a href="#Page_273">273</a></li>
+
+<li>Driesch, <a href="#Footnote_67_67">Foot Note 67</a></li>
+
+<li>Dryopithecus, <a href="#Page_132">132</a></li>
+
+<li>Dubois, E., on Pithecanthropus, <a href="#Page_132">132</a>, <a href="#Page_137">137</a></li>
+
+<li>D&uuml;hring, <a href="#Page_214">214</a>, <a href="#Page_277">277</a></li>
+
+<li>Duns Scotus, <a href="#Page_200">200</a></li>
+
+<li>Duret, C., <a href="#Page_6">6</a></li>
+
+<li>Durkheim, on division of labour, <a href="#Page_278">278</a></li>
+</ul><ul class="IX">
+
+<li><a name="Ecology" id="Ecology"></a>Ecology, <a href="#Footnote_205_205">Foot Note 205</a></li>
+
+<li>Eimer, <a href="#Page_109">109</a></li>
+
+<li><i>Elymnias undularis</i>, <a href="#Page_73">73</a>, <a href="#Page_75">75</a></li>
+
+<li>Embryology, the Origin of Species and, <a href="#Page_154">154</a>, <a href="#Page_155">155</a></li>
+
+<li>Empedocles, <a href="#Page_3">3</a>, <a href="#Page_27">27</a>, <a href="#Page_151">151</a></li>
+
+<li>Engels, <a href="#Page_277">277</a></li>
+
+<li>Environment, action of, <a href="#Page_12">12</a>, <a href="#Page_13">13</a>, <a href="#Page_15">15</a></li>
+
+<li>Epicurus, a poet of Evolution, <a href="#Page_4">4</a></li>
+
+<li>Eristalis, <a href="#Page_75">75</a></li>
+
+<li>Espinas, <a href="#Page_275">275</a></li>
+
+<li>Evolution, and creation, <a href="#Page_233">233</a>
+<ul class="IX">
+<li>  &mdash;conception of, <a href="#Page_3">3-5</a>, <a href="#Page_9">9</a>, <a href="#Page_148">148</a>, <a href="#Page_151">151</a>, <a href="#Page_198">198</a></li>
+<li>  &mdash;discontinuous, <a href="#Page_30">30</a></li>
+<li>  &mdash;experimental, <a href="#Page_5">5</a>, <a href="#Page_7">7</a></li>
+<li>  &mdash;factors of, <a href="#Page_11">11-15</a></li>
+<li>  &mdash;mental, <a href="#Page_194">194</a></li>
+<li>  &mdash;Lloyd Morgan on mental factors in, <a href="#Page_166">166-196</a></li>
+<li>  &mdash;Darwinism and Social, <a href="#Page_18">18</a></li>
+<li>  &mdash;Saltatory, <a href="#Page_29">29-32</a></li>
+<li>  &mdash;Herbert Spencer on, <a href="#Page_204">204-207</a></li>
+<li>  &mdash;Philosophers and modern methods of studying, <a href="#Page_4">4</a></li>
+</ul></li>
+<li>Expression of the Emotions, <a href="#Page_177">177-184</a></li>
+</ul><ul class="IX">
+
+<li>Ferri, <a href="#Page_277">277</a></li>
+
+<li>Ferrier, his work on the brain, 523</li>
+<li>[Transcriber's note: No such page number or reference seen]</li>
+
+<li>Fichte, <a href="#Page_222">222</a></li>
+
+<li>Flourens, <a href="#Page_267">267</a></li>
+
+<li>Flowers and Insects, <a href="#Page_61">61</a>, <a href="#Page_78">78</a></li>
+
+<li>Fouill&eacute;e, <a href="#Page_207">207</a>, <a href="#Page_208">208</a></li>
+
+<li>Fraipont, on skulls from Spy, <a href="#Page_134">134</a></li>
+</ul><ul class="IX">
+
+<li><span class="smcap">Gadow</span>, <a href="#Page_162">162</a></li>
+
+<li><i>Gallus bankiva</i>, <a href="#Page_102">102</a></li>
+
+<li>Gallon, F., <a href="#Page_125">125</a>, <a href="#Page_150">150</a>, <a href="#Page_269">269</a></li>
+
+<li>Geddes, P., <a href="#Page_17">17</a>, <a href="#Footnote_32_32">Foot Note 32</a></li>
+
+<li>Geddes, P. and A. W. Thomson, <a href="#Page_276">276</a><span class='pagenum'><a name="Page_284" id="Page_284">[284]</a></span></li>
+
+
+
+<li>Gegenbaur, <a href="#Page_150">150</a>, <a href="#Page_163">163</a></li>
+
+<li>Genetics, <a href="#Page_93">93</a>, <a href="#Page_96">96</a></li>
+
+<li><i>Germ-plasm</i>, continuity of, <a href="#Page_95">95</a>
+<ul class="IX">
+<li>&mdash;Weismann on, <a href="#Page_46">46-51</a></li>
+</ul></li>
+<li>Germinal Selection, <a href="#Page_36">36</a>, <a href="#Page_37">37</a>, <a href="#Page_46">46-51</a>, <a href="#Page_64">64</a></li>
+
+<li>Gibbon, <a href="#Page_248">248</a></li>
+
+<li>Giuffrida-Ruggeri, <a href="#Page_138">138</a>, <a href="#Page_140">140</a></li>
+
+<li>Giotto, <a href="#Page_259">259</a></li>
+
+<li>Gizycki, <a href="#Page_217">217</a></li>
+
+<li>Goethe and Evolution, <a href="#Page_8">8</a>, <a href="#Page_14">14</a>, <a href="#Page_15">15</a>, <a href="#Page_201">201</a>
+<ul class="IX">
+<li>&mdash;on the relation between Man and Mammals, <a href="#Page_161">161</a>, <a href="#Page_163">163</a></li>
+<li>&mdash;<a href="#Page_221">221</a></li>
+</ul></li>
+
+<li>Gore, Dr., <a href="#Page_226">226</a></li>
+
+<li>Gorjanovi&#269;-Kramberger, <a href="#Page_134">134</a></li>
+
+<li>Gosse, P. H., <a href="#Page_234">234</a></li>
+
+<li><i>Grapta C. album</i>, <a href="#Page_69">69</a></li>
+
+<li>Groos, <a href="#Page_187">187</a>, <a href="#Page_188">188</a></li>
+
+<li>Gulick, <a href="#Page_15">15</a>, <a href="#Page_53">53</a></li>
+
+<li>Guyau, <a href="#Page_217">217</a></li>
+</ul><ul class="IX">
+
+<li>Haberlandt, G., <a href="#Page_34">34</a></li>
+
+<li><span class="smcap">Haeckel</span>, E., on <i>Charles Darwin as an Anthropologist</i>, <a href="#Page_146">146-165</a>
+<ul class="IX">
+<li>  &mdash;and Darwin, <a href="#Page_135">135-151</a>, <a href="#Page_137">137</a>, <a href="#Page_146">146-165</a></li>
+<li>  &mdash;on the Descent of Man, <a href="#Page_137">137</a>, <a href="#Page_143">143</a></li>
+<li>  &mdash;on Lamarck, <a href="#Page_8">8</a>, <a href="#Footnote_21_21">Foot Note 21</a></li>
+<li>  &mdash;a leader in the Darwinian controversy, <a href="#Page_137">137</a></li>
+<li>  &mdash;<a href="#Page_217">217</a></li>
+</ul></li>
+<li>H&auml;cker, <a href="#Page_33">33</a></li>
+
+<li>Hansen, <a href="#Page_272">272</a></li>
+
+<li>Hartmann, von, <a href="#Page_240">240</a></li>
+
+<li>Harvey, <a href="#Page_4">4</a></li>
+
+<li>Haycraft, <a href="#Page_275">275</a></li>
+
+<li>Hegel, <a href="#Page_201">201</a>, <a href="#Page_203">203</a>, <a href="#Page_215">215</a>, <a href="#Page_251">251</a>, <a href="#Page_252">252</a>, <a href="#Page_255">255</a></li>
+
+<li>Heraclitus, <a href="#Page_278">278</a></li>
+
+<li>Herder, <a href="#Page_4">4</a>, <a href="#Page_5">5</a>, <a href="#Page_20">20</a></li>
+
+<li>Heredity and Cytology, <a href="#Page_95">95</a>, <a href="#Page_96">96</a>
+<ul class="IX">
+<li>  &mdash;Haeckel on, <a href="#Page_147">147</a>, <a href="#Page_148">148</a>, <a href="#Page_149">149</a>, <a href="#Page_153">153</a></li>
+<li>  &mdash;and Variation, <a href="#Page_87">87-110</a></li>
+<li>  &mdash;<a href="#Page_219">219</a>, <a href="#Page_224">224</a></li>
+</ul></li>
+<li>Hering, E., on Memory, <a href="#Page_153">153</a></li>
+
+<li>Hertwig, O., <a href="#Page_150">150</a></li>
+
+<li>History, Darwin and, <a href="#Page_246">246-263</a></li>
+
+<li>Hobbes, T., <a href="#Page_200">200</a>, <a href="#Page_215">215</a></li>
+
+<li>Hobhouse, <a href="#Page_242">242</a></li>
+
+<li><span class="smcap">H&ouml;ffding</span>, H., on <i>The Influence of the Conception of Evolution on Modern Philosophy</i>, <a href="#Page_197">197-222</a></li>
+
+<li>Holothurians, calcareous bodies in skin of, <a href="#Page_37">37-41</a></li>
+
+<li><i>Homo heidelbergensis</i>, <a href="#Footnote_118_118">Foot Note 118</a></li>
+
+<li><i>H. neandertalensis</i>, <a href="#Page_138">138</a></li>
+
+<li><i>H. pampaeus</i>, <a href="#Page_144">144</a></li>
+
+<li><i>H. primigenius</i>, <a href="#Page_133">133</a>, <a href="#Page_134">134</a>, <a href="#Page_138">138</a>, <a href="#Page_144">144</a></li>
+
+<li><i>Homunculus</i>, <a href="#Page_132">132</a></li>
+
+<li>Hooker, Sir J. D., and Darwin, <a href="#Page_23">23</a>, <a href="#Page_116">116</a></li>
+
+<li>Huber, <a href="#Page_170">170</a></li>
+
+<li>H&uuml;gel, F. von, <a href="#Footnote_221_221">Foot Note 221</a></li>
+
+<li>Hume, <a href="#Page_200">200</a></li>
+
+<li>Hutcheson, <a href="#Page_216">216</a></li>
+
+<li>Huxley, T. H., and Darwin, <a href="#Page_112">112</a>, <a href="#Page_116">116</a>, <a href="#Page_268">268</a>
+<ul class="IX">
+<li>  &mdash;and the Duke of Argyll, <a href="#Page_238">238</a></li>
+<li>  &mdash;on Lamarck, <a href="#Page_89">89</a></li>
+<li>  &mdash;on Man, <a href="#Page_111">111</a>, <a href="#Page_112">112</a>, <a href="#Page_137">137</a>, <a href="#Page_146">146</a>, <a href="#Page_156">156</a>, <a href="#Page_160">160</a>, <a href="#Page_163">163</a></li>
+<li>  &mdash;on Selection, <a href="#Page_24">24</a>, <a href="#Page_91">91</a>
+<span class='pagenum'><a name="Page_285" id="Page_285">[285]</a></span></li>
+<li>  &mdash;on transmission of acquired characters, <a href="#Page_149">149</a></li>
+<li>  &mdash;<a href="#Page_14">14</a>, <a href="#Page_24">24</a>, <a href="#Page_104">104</a>, <a href="#Page_231">231-236</a>, <a href="#Page_273">273</a>, <a href="#Page_274">274</a></li>
+</ul></li>
+<li>Hybrids, Sterility of, <a href="#Page_104">104</a>, <a href="#Page_105">105</a>, <a href="#Page_106">106</a></li>
+
+</ul><ul class="IX">
+<li>Inheritance of acquired characters, <a href="#Page_93">93</a>, <a href="#Page_94">94</a></li>
+
+<li>Insects and Flowers, <a href="#Page_60">60</a>, <a href="#Page_61">61</a>, <a href="#Page_78">78</a>, <a href="#Page_79">79</a></li>
+
+<li>Instinct, <a href="#Page_122">122</a>, <a href="#Page_172">172-175</a></li>
+
+<li>Irish Elk, an example of coadaptation, <a href="#Page_41">41</a>, <a href="#Page_42">42</a>, <a href="#Page_45">45</a></li>
+
+</ul><ul class="IX">
+<li>Jacoby, <i>Studies in Selection</i> by, <a href="#Page_272">272</a></li>
+
+<li>James, W., <a href="#Page_180">180</a>, <a href="#Page_191">191</a>, <a href="#Page_211">211</a></li>
+
+<li>Jentsch, <a href="#Page_275">275</a></li>
+
+</ul><ul class="IX">
+<li>Kallima, protective colouring of, <a href="#Page_35">35</a>, <a href="#Page_68">68</a>, <a href="#Page_70">70</a></li>
+
+<li><i>K. inachis</i>, <a href="#Page_68">68</a></li>
+
+<li>Kammerer's experiments on Salamanders, <a href="#Page_28">28</a></li>
+
+<li>Kant, I., <a href="#Page_4">4</a>, <a href="#Page_5">5</a>, <a href="#Page_6">6</a>, <a href="#Page_27">27</a>, <a href="#Page_198">198</a>, <a href="#Page_211">211</a>, <a href="#Page_212">212</a>, <a href="#Page_217">217</a>, <a href="#Page_221">221</a>, <a href="#Page_222">222</a></li>
+
+<li>Keane, on the Primates, <a href="#Page_138">138</a></li>
+
+<li>Keith, on Anthropoid Apes, <a href="#Page_138">138</a></li>
+
+<li>Kepler, <a href="#Page_198">198</a></li>
+
+<li>Klaatsch, on Ancestry of Man, <a href="#Page_140">140</a></li>
+
+<li>Klaatsch and Hauser, <a href="#Page_134">134</a></li>
+
+<li>Knies, <a href="#Page_266">266</a></li>
+
+<li>K&ouml;lliker, his views on Evolution, <a href="#Page_29">29</a>, <a href="#Page_150">150</a></li>
+
+<li>Kollmann, on origin of human races, <a href="#Page_144">144</a></li>
+
+<li>Korschinsky, <a href="#Page_31">31</a></li>
+
+<li>Krause, E., <a href="#Footnote_10_10">Foot Note 10</a>, <a href="#Page_13">13</a></li>
+
+<li>Kropotkin, <a href="#Page_214">214</a>, <a href="#Page_275">275</a></li>
+
+</ul><ul class="IX">
+<li>Lamarck, his division of the Animal Kingdom, <a href="#Page_160">160</a>, <a href="#Page_161">161</a>
+<ul class="IX">
+<li>  &mdash;Darwin's opinion of, <a href="#Page_129">129</a></li>
+<li>  &mdash;on Evolution, <a href="#Page_9">9-14</a>, <a href="#Page_21">21</a>, <a href="#Page_25">25</a>, <a href="#Page_171">171</a>, <a href="#Page_172">172</a>, <a href="#Page_173">173</a>, <a href="#Page_179">179</a>, <a href="#Page_180">180</a>, <a href="#Page_201">201</a>, <a href="#Page_202">202</a>, <a href="#Page_253">253</a></li>
+<li>  &mdash;on Man, <a href="#Page_146">146</a>, <a href="#Page_148">148</a>, <a href="#Page_160">160</a>, <a href="#Page_163">163</a></li>
+<li>  &mdash;<a href="#Page_89">89</a>, <a href="#Page_109">109</a>, <a href="#Page_201">201</a>, <a href="#Page_202">202</a>, <a href="#Page_233">233</a></li>
+</ul></li>
+<li>Lamarckian principle, <a href="#Page_28">28</a>, <a href="#Page_41">41-44</a>, <a href="#Page_50">50-54</a>, <a href="#Page_67">67</a>, <a href="#Page_84">84</a>, <a href="#Page_86">86</a></li>
+
+<li>Lamb, C., <a href="#Page_229">229</a></li>
+
+<li>Lamettrie, <a href="#Page_198">198</a></li>
+
+<li>Lamprecht, <a href="#Page_260">260-263</a></li>
+
+<li>Lanessan, J. L. de, <a href="#Footnote_17_17">Foot Note 17</a>, <a href="#Page_275">275</a></li>
+
+<li>Lang, <a href="#Footnote_21_21">Foot Note 21</a></li>
+
+<li>Lange, <a href="#Page_180">180</a></li>
+
+<li>Language, Darwin on, <a href="#Page_123">123</a>, <a href="#Page_124">124</a>
+<ul class="IX">
+<li>  &mdash;Evolution and the Science of, <a href="#Page_178">178</a>, <a href="#Page_179">179</a>, <a href="#Page_188">188</a></li>
+</ul></li>
+<li>Lankester, Sir E. Ray, on degeneration, <a href="#Page_268">268</a>
+<ul class="IX">
+<li>  &mdash;on educability, <a href="#Page_170">170</a>, <a href="#Page_189">189</a></li>
+</ul></li>
+<li>Lankester, Sir E. Ray, on the germ-plasm theory, <a href="#Page_150">150</a></li>
+
+<li>Lapouge, Vacher de, <a href="#Page_270">270</a></li>
+
+<li>Lartet, M. E., <a href="#Page_189">189</a></li>
+
+<li>Lasalle, <a href="#Page_266">266</a></li>
+
+<li>Laveleye, de, <a href="#Page_275">275</a></li>
+
+<li>Lawrence, W., <a href="#Page_89">89</a>, <a href="#Footnote_65_65">Foot Note 65</a></li>
+
+<li>Lehmann-Nitsche, <a href="#Page_138">138</a>, <a href="#Page_144">144</a></li>
+
+<li>Leibnitz, <a href="#Page_4">4</a>, <a href="#Page_5">5</a>, <a href="#Page_213">213</a></li>
+
+<li>Lepidoptera, variation in, <a href="#Page_37">37</a>, <a href="#Page_60">60-63</a></li>
+
+<li>Lessing, <a href="#Page_4">4</a>, <a href="#Page_221">221</a></li>
+
+<li>Liddon, H. P., <a href="#Page_234">234</a></li>
+
+<li><i>Limenitis archippus</i>, <a href="#Page_74">74</a></li>
+
+<li>Linnaeus, <a href="#Page_6">6</a></li>
+
+<li>Locy, W. A., <a href="#Footnote_15_15">Foot Note 15</a></li>
+
+<li>Lovejoy, <a href="#Footnote_56_56">Foot Note 56</a></li>
+
+<li>Lubbock, <a href="#Page_125">125</a>
+<span class='pagenum'><a name="Page_286" id="Page_286">[286]</a></span></li>
+
+
+<li>Lucretius, a poet of Evolution, <a href="#Page_4">4</a></li>
+
+<li>Lyell, Sir Charles, and Darwin, <a href="#Page_23">23</a>, <a href="#Page_116">116</a>
+<ul class="IX">
+<li>  &mdash;the uniformitarian teaching of, <a href="#Page_89">89</a></li>
+</ul></li>
+</ul><ul class="IX">
+
+<li>Macacus, ear of, <a href="#Page_119">119</a></li>
+
+<li>Mach, E., <a href="#Page_153">153</a>, <a href="#Page_211">211</a></li>
+
+<li>Mahoudeau, <a href="#Page_137">137</a></li>
+
+<li>Maillet, de, <a href="#Page_6">6</a></li>
+
+<li>Majewski, <a href="#Footnote_238_238">Foot Note 238</a>, <a href="#Footnote_239_239">Foot Note 239</a></li>
+
+<li>Malthus, his influence on Darwin, <a href="#Page_16">16-18</a>, <a href="#Page_21">21</a>, <a href="#Page_24">24</a>, <a href="#Page_91">91</a>
+<ul class="IX">
+<li>  &mdash;<a href="#Page_200">200</a>, <a href="#Page_273">273</a></li>
+</ul></li>
+<li>Man, Descent of, <a href="#Page_126">126</a>, <a href="#Page_127">127</a>, <a href="#Page_128">128</a>, <a href="#Page_131">131-145</a>, <a href="#Page_156">156-165</a>, <a href="#Page_189">189</a>, <a href="#Page_254">254</a>, <a href="#Page_265">265</a>
+<ul class="IX">
+<li>  &mdash;mental and moral qualities of animals and, <a href="#Page_122">122-126</a>, <a href="#Page_164">164</a>, <a href="#Page_188">188-192</a></li>
+<li>  &mdash;pre-Darwinian views on the Descent of, <a href="#Page_1">1</a></li>
+</ul></li>
+<li>Man, Tertiary flints worked by, <a href="#Page_136">136</a></li>
+
+<li><i>Man</i>, G. Schwalbe on Darwin's <i>Descent of</i>, <a href="#Page_111">111-145</a></li>
+
+<li>Manouvrier, <a href="#Page_137">137</a></li>
+
+<li><i>Mantis religiosa</i>, colour experiments on, <a href="#Page_65">65</a>, <a href="#Page_68">68</a></li>
+
+<li>Marx, <a href="#Page_262">262</a>, <a href="#Page_276">276-278</a></li>
+
+<li>Matthew, P., and Natural Selection, <a href="#Page_18">18</a>, <a href="#Page_19">19</a></li>
+
+<li>Maupertuis, <a href="#Page_6">6</a>, <a href="#Page_88">88</a>, <a href="#Page_103">103</a></li>
+
+<li>Mayer, R., <a href="#Page_197">197</a></li>
+
+<li><i>Mechanitis lysimnia</i>, <a href="#Page_77">77</a></li>
+
+<li><i>Melinaea ethra</i>, <a href="#Page_77">77</a></li>
+
+<li>Mendel, <a href="#Page_97">97-100</a>, <a href="#Page_184">184</a>, <a href="#Page_228">228</a></li>
+
+<li>Merz, J. T.,<a href="#Footnote_14_14">Foot Note 14</a></li>
+
+<li>Mesopithecus, <a href="#Page_132">132</a></li>
+
+<li>Mill, J. S., <a href="#Page_193">193</a>, <a href="#Page_200">200</a>, <a href="#Page_202">202</a>, <a href="#Page_218">218</a></li>
+
+<li>Mimicry, <a href="#Page_70">70-82</a></li>
+
+<li>Moltke, on war, <a href="#Page_273">273</a></li>
+
+<li>Monkeys, fossil, <a href="#Page_132">132</a></li>
+
+<li>Montesquieu, <a href="#Page_248">248</a></li>
+
+<li>Monticelli, <a href="#Page_155">155</a></li>
+
+<li><span class="smcap">Morgan, C. Lloyd</span>, on <i>Mental Factors in Evolution</i>, <a href="#Page_166">166-196</a>
+<ul class="IX">
+<li>  &mdash;on Organic Selection, <a href="#Page_53">53</a></li>
+</ul></li>
+<li>Morgan, T. H., <a href="#Page_99">99</a></li>
+
+<li>Morselli, <a href="#Page_138">138</a></li>
+
+<li>Mortillet, <a href="#Page_136">136</a></li>
+
+<li>Moseley, <a href="#Footnote_224_224">Foot Note 224</a></li>
+
+<li>Muller, Fritz, <i>F&uuml;r Darwin</i> by, <a href="#Page_154">154</a>
+<ul class="IX">
+<li>  &mdash;on Mimicry, <a href="#Page_233">233</a></li>
+<li>  &mdash;<a href="#Page_59">59</a>, <a href="#Page_77">77</a></li>
+</ul></li>
+<li>Muller, J., <a href="#Page_147">147</a></li>
+
+<li>M&uuml;ller, Max, on language, <a href="#Page_124">124</a></li>
+
+<li><a name="Mutations" id="Mutations"></a>Mutation, <a href="#Page_15">15</a>, <a href="#Page_31">31</a>, <a href="#Page_184">184</a>, <a href="#Page_199">199</a>, <a href="#Page_209">209</a></li>
+</ul><ul class="IX">
+
+<li>N&auml;geli, <a href="#Page_109">109</a>, <a href="#Page_151">151</a>, <a href="#Page_153">153</a></li>
+
+<li>Nathusius, <a href="#Page_103">103</a></li>
+
+<li><a name="Natural_Selection" id="Natural_Selection"></a>Natural Selection, Darwin's views on, <a href="#Page_90">90</a>, <a href="#Page_91">91</a>, <a href="#Page_122">122</a>, <a href="#Page_149">149</a>
+<ul class="IX">
+<li>  &mdash;Darwin and Wallace on, <a href="#Page_2">2</a>, <a href="#Page_163">163</a>, <a href="#Page_183">183</a></li>
+<li>  &mdash;and design, <a href="#Page_241">241</a>, <a href="#Page_242">242</a></li>
+<li>  &mdash;and educability, <a href="#Page_195">195</a></li>
+<li>  &mdash;and human development, <a href="#Page_125">125</a>, <a href="#Page_256">256</a>, <a href="#Page_257">257</a></li>
+<li>  &mdash;<a href="#Page_16">16-20</a>, <a href="#Page_25">25</a>, <a href="#Page_26">26</a>, <a href="#Page_41">41</a>, <a href="#Page_55">55-58</a>, <a href="#Page_64">64-86</a>, <a href="#Page_87">87-96</a>, <a href="#Page_199">199</a>, <a href="#Page_233">233</a></li>
+</ul></li>
+<li>Neandertal skulls, <a href="#Page_133">133</a>, <a href="#Page_134">134</a></li>
+
+<li>Neodarwinism, <a href="#Page_150">150</a></li>
+
+<li>Newton, A., <a href="#Footnote_59_59">Foot Note 59</a></li>
+
+<li>Newton, I., <a href="#Page_197">197</a>, <a href="#Page_198">198</a></li>
+
+<li>Niebuhr, <a href="#Page_249">249</a>, <a href="#Page_263">263</a></li>
+
+<li>Nietzsche, <a href="#Page_214">214</a>, <a href="#Page_271">271</a>
+<span class='pagenum'><a name="Page_287" id="Page_287">[287]</a></span></li>
+
+<li>Nitsche, <a href="#Page_119">119</a></li>
+
+<li>Novicow, <a href="#Page_274">274</a></li>
+
+<li>Nuttall, G. H. F., <a href="#Page_135">135</a></li>
+</ul><ul class="IX">
+
+<li>Occam, <a href="#Page_200">200</a></li>
+
+<li>Odin, <a href="#Page_270">270</a></li>
+
+<li>Oecology, see <a href="#Ecology">Ecology</a></li>
+
+<li><i>Oenothera lamarckiana</i>, <a href="#Page_32">32</a></li>
+
+<li>Oestergren, on Holothurians, <a href="#Page_37">37-39</a></li>
+
+<li>Oken, L., <a href="#Page_7">7</a>, <a href="#Page_201">201</a></li>
+
+<li>Organic Selection, <a href="#Page_53">53</a>, <a href="#Page_54">54</a>, <a href="#Page_172">172</a>, <a href="#Page_173">173</a></li>
+
+<li>Orthogenesis, <a href="#Page_109">109</a></li>
+
+<li>Osborn, H. F., <a href="#Page_53">53</a>, <a href="#Footnote_165_165">Foot Note 165</a>
+<ul class="IX">
+<li>  &mdash;<i>From the Greeks to Darwin</i> by, <a href="#Page_3">3-5</a>, <a href="#Page_12">12</a>, <a href="#Page_14">14</a>, <a href="#Page_20">20</a></li>
+</ul></li>
+<li><i>Ovibos moschatus</i>, <a href="#Page_67">67</a></li>
+
+<li>Owen, Sir Richard, <a href="#Page_111">111</a></li>
+</ul><ul class="IX">
+
+<li>Packard, A. S., <a href="#Footnote_12_12">Foot Note 12</a>, <a href="#Footnote_18_18">Foot Note 18</a></li>
+
+<li>Palaeopithecus, <a href="#Page_132">132</a></li>
+
+<li>Paley, <a href="#Page_18">18</a>, <a href="#Page_242">242</a>, <a href="#Page_244">244</a></li>
+
+<li>Panmixia, Weismann's principle of, <a href="#Page_54">54</a></li>
+
+<li><i>Papilio dardanus</i>, <a href="#Page_72">72</a>, <a href="#Page_73">73</a>, <a href="#Page_74">74</a></li>
+
+<li><i>P. meriones</i>, <a href="#Page_73">73</a></li>
+
+<li><i>P. merope</i>, <a href="#Page_72">72</a></li>
+
+<li>Pearson, K., <a href="#Footnote_7_7">Foot Note 7</a></li>
+
+<li>Penck, <a href="#Page_136">136</a></li>
+
+<li>Peridineae, <a href="#Page_33">33</a></li>
+
+<li>Perrier, E., <a href="#Footnote_21_21">Foot Note 21</a>, <a href="#Page_20">20</a></li>
+
+<li>Perthes, B. de, <a href="#Page_123">123</a></li>
+
+<li>Pfeffer, W., <a href="#Page_28">28</a></li>
+
+<li>Philosophy, influence of the conception of evolution on modern, <a href="#Page_197">197-222</a></li>
+
+<li>Pithecanthropus, <a href="#Page_133">133</a>, <a href="#Page_134">134</a>, <a href="#Page_138">138</a>, <a href="#Page_143">143</a></li>
+
+<li>Pitheculites, <a href="#Page_144">144</a></li>
+
+<li>Plate, <a href="#Footnote_37_37">Foot Note 37</a></li>
+
+<li>Pliopithecus, <a href="#Page_132">132</a></li>
+
+<li>Pouchet, G., <a href="#Footnote_3_3">Foot Note 3</a></li>
+
+<li><span class="smcap">Poulton, E. B.</span>, experiments on Butterflies by, <a href="#Page_65">65</a>
+<ul class="IX">
+<li>  &mdash;on J. C. Prichard, <a href="#Page_20">20</a></li>
+<li>  &mdash;on Mimicry, <a href="#Page_69">69</a>, <a href="#Page_71">71</a>, <a href="#Page_75">75</a>, <a href="#Page_78">78</a></li>
+<li>  &mdash;<a href="#Footnote_34_34">Foot Note 34</a>, <a href="#Footnote_43_43">Foot Note 43</a>, <a href="#Footnote_49_49">Foot Note 49</a>, <a href="#Footnote_55_55">Foot Note 55</a></li>
+</ul></li>
+<li>Prichard, J. C., <a href="#Page_20">20</a>, <a href="#Page_21">21</a>, <a href="#Page_89">89</a>, <a href="#Footnote_65_65">Foot Note 65</a></li>
+
+<li><i>Pronuba yuccasella</i>, <a href="#Page_79">79</a></li>
+
+<li>Protective resemblance, <a href="#Page_65">65-70</a></li>
+
+<li>Pusey, <a href="#Page_115">115</a></li>
+
+</ul><ul class="IX">
+<li>Quatrefages, A. de, <a href="#Footnote_21_21">Foot Note 21</a>, <a href="#Page_19">19</a></li>
+
+</ul><ul class="IX">
+<li>Radiolarians, <a href="#Page_33">33</a></li>
+
+<li>Ranke, <a href="#Page_249">249</a>, <a href="#Page_251">251</a>, <a href="#Page_255">255</a>, <a href="#Page_263">263</a></li>
+
+<li>Rau, A., <a href="#Page_153">153</a></li>
+
+<li>Ray, J., <a href="#Page_4">4</a></li>
+
+<li>Regeneration, <a href="#Footnote_71_71">Foot Note 71</a></li>
+
+<li>Religious thought, Darwin's influence on, <a href="#Page_223">223-245</a></li>
+
+<li>Reversion, <a href="#Page_120">120</a>, <a href="#Page_121">121</a></li>
+
+<li>Ridley, H. N., <a href="#Footnote_88_88">Foot Note 88</a></li>
+
+<li>Ritchie, <a href="#Page_270">270</a></li>
+
+<li>Robinet, <a href="#Page_6">6</a></li>
+
+<li>Rolph, <a href="#Page_217">217</a></li>
+
+<li>Romanes, G. J., <a href="#Footnote_3_3">Foot Note 3</a>, <a href="#Page_15">15</a>, <a href="#Page_32">32</a>, <a href="#Page_54">54</a>, <a href="#Page_164">164</a>, <a href="#Page_234">234</a></li>
+
+<li>Roux, <a href="#Page_151">151</a>, <a href="#Page_152">152</a></li>
+
+<li>Ruskin, <a href="#Page_230">230</a></li>
+
+<li>Rutot, <a href="#Page_136">136</a></li>
+
+</ul><ul class="IX">
+
+<li>Saint-Hilaire, E. G. de, <a href="#Page_8">8</a>, <a href="#Page_15">15</a>, <a href="#Page_20">20</a></li>
+
+<li>Saltatory Evolution, <a href="#Page_29">29-32</a> (see also <a href="#Mutations">Mutations</a>)</li>
+
+<li>Sanders, experiments on Vanessa by, <a href="#Page_65">65</a></li>
+
+<li>Savigny, <a href="#Page_249">249</a></li>
+
+<li>Schelling, <a href="#Page_4">4</a>, <a href="#Page_5">5</a>, <a href="#Page_200">200</a>, <a href="#Page_201">201</a>
+<span class='pagenum'><a name="Page_288" id="Page_288">[288]</a></span></li>
+
+<li>Schleiden and Schwann, Cell-theory of, <a href="#Page_147">147</a></li>
+
+<li>Schoetensack, on <i>Homo heidelbergensis</i>, <a href="#Footnote_118_118">Foot Note 118</a></li>
+
+<li>Sch&uuml;tt, <a href="#Page_23">23</a></li>
+
+<li><span class="smcap">Schwalbe, G.</span>, on <i>The Descent of Man</i>, <a href="#Page_111">111-145</a></li>
+
+<li>Seeck, O., <a href="#Footnote_240_240">Foot Note 240</a></li>
+
+<li>Segregation, <a href="#Page_97">97</a>, <a href="#Page_98">98</a></li>
+
+<li>Selection, artificial, <a href="#Page_24">24</a>, <a href="#Page_25">25</a>, <a href="#Page_26">26</a>, <a href="#Page_41">41</a>, <a href="#Page_45">45</a>, <a href="#Page_120">120</a>, <a href="#Page_269">269-272</a>
+<ul class="IX">
+<li>  &mdash;germinal, <a href="#Page_35">35</a>, <a href="#Page_36">36</a>, <a href="#Page_46">46-52</a>, <a href="#Page_64">64</a></li>
+</ul></li>
+<li>Selection, natural (see <a href="#Natural_Selection">Natural Selection</a>)
+<ul class="IX">
+<li>  &mdash;organic, <a href="#Page_53">53</a>, <a href="#Page_171">171</a>, <a href="#Page_172">172</a></li>
+<li>  &mdash;sexual, <a href="#Page_55">55-64</a>, <a href="#Page_117">117</a>, <a href="#Page_118">118</a></li>
+<li>  &mdash;social and natural, <a href="#Page_271">271</a></li>
+<li>  &mdash;<a href="#Page_23">23-86</a>, <a href="#Page_103">103</a>, <a href="#Page_129">129</a>, <a href="#Page_130">130</a></li>
+</ul></li>
+<li>Selenka, <a href="#Page_131">131</a></li>
+
+<li>Semnopithecus, <a href="#Page_132">132</a></li>
+
+<li>Semon, R., <a href="#Page_28">28</a>, <a href="#Page_153">153</a></li>
+
+<li>Sergi, <a href="#Page_138">138</a>, <a href="#Page_143">143</a></li>
+
+<li>Sex, recent investigations on, <a href="#Page_99">99</a>, <a href="#Page_100">100</a></li>
+
+<li>Sibbern, <a href="#Page_201">201</a></li>
+
+<li><i>Smerinthus ocellata</i>, <a href="#Page_38">38</a></li>
+
+<li><i>Smerinthus populi</i>, <a href="#Page_38">38</a></li>
+
+<li><i>S. tiliae</i>, <a href="#Page_38">38</a></li>
+
+<li>Smith, A., <a href="#Page_200">200</a></li>
+
+<li>Sociology,  Darwinism and, <a href="#Page_264">264-280</a>
+<ul class="IX">
+<li>  &mdash;History and, <a href="#Page_255">255</a></li>
+</ul></li>
+<li>Sollas, W. J., <a href="#Page_134">134</a></li>
+
+<li>Sorley, W. R., <a href="#Page_217">217</a></li>
+
+<li>Species and varieties, <a href="#Page_100">100</a></li>
+
+<li>Spencer, H., on evolution, <a href="#Page_204">204-209</a>
+<ul class="IX">
+<li>  &mdash;on the theory of Selection, <a href="#Page_41">41</a></li>
+</ul></li>
+<li>Spencer, H., on Sociology, <a href="#Page_268">268</a>
+<ul class="IX">
+<li>  &mdash;on the transmission of acquired characters, <a href="#Page_149">149</a></li>
+<li>  &mdash;on Weismann, <a href="#Page_41">41</a>, <a href="#Page_150">150</a></li>
+<li>  &mdash;<a href="#Page_2">2</a>, <a href="#Page_17">17</a>, <a href="#Page_217">217</a>, <a href="#Page_231">231</a>, <a href="#Page_268">268</a></li>
+</ul></li>
+<li>Sphingidae, variation in, <a href="#Page_37">37</a></li>
+
+<li>Spinoza, <a href="#Page_153">153</a>, <a href="#Page_206">206</a></li>
+
+<li>Standfuss, <a href="#Page_82">82</a></li>
+
+<li>Stephen, L., <a href="#Page_217">217</a></li>
+
+<li>Sterility in hybrids, <a href="#Page_104">104-106</a></li>
+
+<li>Sterne, C, <a href="#Footnote_10_10">Foot Note 10</a></li>
+
+<li>Struggle for existence, <a href="#Page_25">25</a>, <a href="#Page_26">26</a>, <a href="#Page_272">272-274</a></li>
+
+<li>Sutton, A. W., <a href="#Footnote_73_73">Foot Note 73</a></li>
+
+<li>Synapta, calcareous bodies in skin of, <a href="#Page_38">38-41</a></li>
+
+<li>Syrphus, <a href="#Page_75">75</a></li>
+</ul><ul class="IX">
+
+<li>Tarde, G., <a href="#Page_279">279</a></li>
+
+<li>Tennant, F. R., <a href="#Footnote_218_218">Foot Note 218</a></li>
+
+<li>Tetraprothomo, <a href="#Page_138">138</a>, <a href="#Page_144">144</a></li>
+
+<li><span class="smcap">Thomson, J. A.</span>, on <i>Darwin's Predecessors</i>, <a href="#Page_1">1-22</a>
+<ul class="IX">
+<li>  &mdash;<a href="#Page_150">150</a></li>
+<li>  &mdash;and P. Geddes, <a href="#Page_276">276</a></li>
+</ul></li>
+<li>Treschow, <a href="#Page_201">201</a></li>
+
+<li>Treviranus, <a href="#Page_8">8</a>, <a href="#Page_14">14</a>, <a href="#Page_15">15</a></li>
+
+<li>Turgot, <a href="#Page_249">249</a></li>
+
+<li>Turner, Sir W., <a href="#Page_150">150</a></li>
+
+<li>Tylor, <a href="#Page_267">267</a></li>
+
+<li>Tyndall, W., <a href="#Page_267">267</a></li>
+
+<li>Tyrrell, G, <a href="#Footnote_222_222">Foot Note 222</a></li>
+
+</ul><ul class="IX">
+<li>Uhlenhuth, on blood reactions, <a href="#Page_135">135</a></li>
+
+<li>Use and disuse, <a href="#Page_28">28</a>, <a href="#Page_41">41-43</a>, <a href="#Page_48">48-54</a>, <a href="#Page_94">94</a>, <a href="#Page_95">95</a>, <a href="#Page_119">119</a>, <a href="#Page_149">149</a>
+<span class='pagenum'><a name="Page_289" id="Page_289">[289]</a></span></li>
+
+</ul><ul class="IX">
+
+<li>Vanessa, <a href="#Page_63">63</a></li>
+
+<li><i>V. levana</i>, <a href="#Page_31">31</a></li>
+
+<li><i>V. polychloros</i>, <a href="#Page_82">82</a></li>
+
+<li><i>V. urticae</i>, <a href="#Page_65">65</a>, <a href="#Page_82">82</a></li>
+
+<li>Variability, Darwin's attention directed to, <a href="#Page_24">24</a>
+<ul class="IX">
+<li>  &mdash;W. Bateson on, <a href="#Page_87">87-110</a></li>
+<li>  &mdash;causes of, <a href="#Page_200">200</a></li>
+</ul></li>
+<li>Variation, Darwin's views as an evolutionist, and as a systematist, on, <a href="#Page_212">212</a>
+<ul class="IX">
+<li>  &mdash;and heredity, <a href="#Page_87">87-110</a></li>
+<li>  &mdash;minute, <a href="#Page_28">28-32</a></li>
+<li>  &mdash;in relation to species, <a href="#Page_100">100</a>, <a href="#Page_101">101</a></li>
+</ul></li>
+<li>Varigny, H. de, <a href="#Page_6">6</a>, <a href="#Page_19">19</a></li>
+
+<li>Verworn, <a href="#Page_136">136</a></li>
+
+<li><i>Vestiges of Creation</i>, Darwin on <i>The</i>, <a href="#Page_15">15</a></li>
+
+<li>Virchow, his opposition to Darwin, <a href="#Page_157">157</a>, <a href="#Page_158">158</a>
+<ul class="IX">
+<li>  &mdash;on the transmission of acquired characters, <a href="#Page_149">149</a></li>
+</ul></li>
+<li>Vogt, <a href="#Page_137">137</a></li>
+
+<li>Voltaire, <a href="#Page_248">248</a></li>
+
+<li><span class="smcap">Vries, H.</span> de, the Mutation theory of, <a href="#Page_31">31</a>, <a href="#Page_101">101</a>, <a href="#Page_151">151</a>, <a href="#Page_213">213</a></li>
+
+</ul><ul class="IX">
+<li><span class="smcap">Waggett</span>, Rev. P. N., on <i>The Influence of Darwin upon Religious Thought</i>, <a href="#Page_223">223-245</a></li>
+
+<li>Wallace, A. R., on Colour, <a href="#Page_63">63</a>, <a href="#Page_71">71</a>
+<ul class="IX">
+<li>  &mdash;and Darwin, <a href="#Footnote_7_7">Foot Note 7</a>, <a href="#Page_23">23</a>, <a href="#Page_183">183</a></li>
+<li>  &mdash;on the Descent of Man, <a href="#Page_116">116</a></li>
+<li>  &mdash;on Malthus, <a href="#Page_17">17</a></li>
+<li>  &mdash;on Natural Selection, <a href="#Page_2">2</a>, <a href="#Page_16">16</a>, <a href="#Page_163">163</a>, <a href="#Page_232">232</a></li>
+</ul></li>
+<li>Wallace, A. R., on social reforms, <a href="#Page_275">275</a>, <a href="#Page_276">276</a>
+<ul class="IX">
+<li>  &mdash;on Sexual Selection, <a href="#Page_183">183</a>, <a href="#Page_184">184</a></li>
+</ul></li>
+<li>Walton, <a href="#Page_237">237</a></li>
+
+<li>Watt, J., and Natural Selection, <a href="#Page_21">21</a></li>
+
+<li><span class="smcap">Weismann, A.</span>, on <i>The Selection Theory</i>, <a href="#Page_23">23-86</a>
+<ul class="IX">
+<li>  &mdash;his germ-plasm theory, <a href="#Page_46">46-51</a>, <a href="#Page_149">149</a>, <a href="#Page_150">150</a></li>
+<li>  &mdash;and Prichard, <a href="#Page_20">20</a></li>
+<li>  &mdash;and Spencer, <a href="#Page_42">42</a></li>
+</ul></li>
+<li>Weismann, A., on the transmission of acquired characters, <a href="#Page_93">93-95</a>
+<ul class="IX">
+<li>  &mdash;<a href="#Page_156">156</a></li>
+</ul></li>
+<li>Wells, W. C, and Natural Selection, <a href="#Page_18">18</a></li>
+
+<li>White, G., <a href="#Page_3">3</a></li>
+
+<li>Williams, C. M., <a href="#Page_217">217</a></li>
+
+<li>Wilson, E. B., on cytology, <a href="#Page_99">99</a></li>
+
+<li>Wolf, <a href="#Page_249">249</a></li>
+
+<li>Wollaston's, T. V., <i>Variation of Species</i>, <a href="#Footnote_59_59">Foot Note 59</a></li>
+
+<li>Woltmann, <a href="#Page_277">277</a></li>
+
+<li>Woolner, <a href="#Page_118">118</a></li>
+
+<li>Wundt, on language, <a href="#Page_207">207</a>, <a href="#Page_208">208</a></li>
+</ul><ul class="IX">
+
+<li><i>Xylina vetusta</i>, <a href="#Page_82">82</a></li>
+</ul><ul class="IX">
+
+<li>Yucca, fertilisation of, <a href="#Page_78">78</a>, <a href="#Page_79">79</a></li>
+
+</ul><ul class="IX">
+<li>Zeller, E., <a href="#Footnote_3_3">Foot Note 3</a></li>
+
+<li><i>Zoonomia</i>, Erasmus Darwin's, <a href="#Page_7">7</a></li>
+</ul>
+</div>
+<hr style="width: 65%;" />
+<p><i>The publishers will be pleased to send, upon request, an illustrated
+catalogue setting forth the purposes and ideals of The Modern Library,
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+
+
+
+
+
+<pre>
+
+
+
+
+
+End of the Project Gutenberg EBook of Evolution in Modern Thought, by 
+Ernst Haeckel and J. Arthur Thomson and August Weismann
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+The Project Gutenberg EBook of Evolution in Modern Thought, by
+Ernst Haeckel and J. Arthur Thomson and August Weismann
+
+This eBook is for the use of anyone anywhere at no cost and with
+almost no restrictions whatsoever.  You may copy it, give it away or
+re-use it under the terms of the Project Gutenberg License included
+with this eBook or online at www.gutenberg.org
+
+
+Title: Evolution in Modern Thought
+
+Author: Ernst Haeckel
+        J. Arthur Thomson
+        August Weismann
+
+Release Date: August 29, 2007 [EBook #22430]
+
+Language: English
+
+Character set encoding: ASCII
+
+*** START OF THIS PROJECT GUTENBERG EBOOK EVOLUTION IN MODERN THOUGHT ***
+
+
+
+
+Produced by Marilynda Fraser-Cunliffe, Sankar Viswanathan,
+and the Online Distributed Proofreading Team at
+http://www.pgdp.net
+
+
+
+
+
+
+
+
+
+                         EVOLUTION IN MODERN
+
+                               THOUGHT
+
+
+
+                    BY HAECKEL, THOMSON, WEISMANN
+
+                              AND OTHERS
+
+
+
+
+
+                          THE MODERN LIBRARY
+
+                      PUBLISHERS :: :: NEW YORK
+
+       *       *       *       *       *
+
+
+
+
+CONTENTS
+
+
+I    DARWIN'S PREDECESSORS
+
+     J. Arthur Thomson, Professor of Natural History in the University of
+       Aberdeen
+
+
+II   _The Selection Theory_
+
+     August Weismann, Professor of Zoology in the University of
+       Freiburg (Baden)
+
+
+III  HEREDITY AND VARIATION IN MODERN LIGHTS
+
+     W. Bateson, Professor of Biology in the University of Cambridge
+
+
+IV   "THE DESCENT OF MAN"
+
+     G. Schwalbe, Professor of Anatomy in the University of Strassburg
+
+
+V    CHARLES DARWIN AS AN ANTHROPOLOGIST
+
+     Ernst Haeckel, Professor of Zoology in the University of Jena
+
+
+VI   MENTAL FACTORS IN EVOLUTION
+
+     C. Lloyd Morgan, Professor of Psychology at University College,
+       Bristol
+
+
+VII  THE INFLUENCE OF THE CONCEPTION OF EVOLUTION ON MODERN PHILOSOPHY
+
+       H. Hoeffding, Professor of Philosophy in the University of Copenhagen
+
+
+VIII THE INFLUENCE OF DARWIN UPON RELIGIOUS THOUGHT
+
+     Rev. P. H. Waggett
+
+
+IX   DARWINISM AND HISTORY
+
+     J. B. Bury, Regious Professor of Modern History in the University
+       of Cambridge
+
+
+X    DARWINISM AND SOCIOLOGY
+
+     C. Bougle, Professor of Social Philosophy in the University of
+       Toulouse, and Deputy-Professor at the Sorbonne, Paris
+
+       *       *       *       *       *
+
+
+
+EVOLUTION IN MODERN THOUGHT
+
+
+I
+
+DARWIN'S PREDECESSORS
+
+BY J. ARTHUR THOMSON
+
+_Professor of Natural History in the University of Aberdeen_
+
+
+In seeking to discover Darwin's relation to his predecessors it is
+useful to distinguish the various services which he rendered to the
+theory of organic evolution.
+
+(I) As everyone knows, the general idea of the Doctrine of Descent is
+that the plants and animals of the present day are the lineal
+descendants of ancestors on the whole somewhat simpler, that these
+again are descended from yet simpler forms, and so on backwards
+towards the literal "Protozoa" and "Protophyta" about which we
+unfortunately know nothing. Now no one supposes that Darwin originated
+this idea, which in rudiment at least is as old as Aristotle. What
+Darwin did was to make it current intellectual coin. He gave it a form
+that commended itself to the scientific and public intelligence of the
+day, and he won widespread conviction by showing with consummate skill
+that it was an effective formula to work with, a key which no lock
+refused. In a scholarly, critical, and pre-eminently fair-minded way,
+admitting difficulties and removing them, foreseeing objections and
+forestalling them, he showed that the doctrine of descent supplied a
+modal interpretation of how our present-day fauna and flora have come
+to be.
+
+(II) In the second place, Darwin applied the evolution-idea to
+particular problems, such as the descent of man, and showed what a
+powerful organon it is, introducing order into masses of uncorrelated
+facts, interpreting enigmas both of structure and function, both
+bodily and mental, and, best of all, stimulating and guiding further
+investigation. But here again it cannot be claimed that Darwin was
+original. The problem of the descent or ascent of man, and other
+particular cases of evolution, had attracted not a few naturalists
+before Darwin's day, though no one [except Herbert Spencer in the
+psychological domain (1855)] had come near him in precision and
+thoroughness of inquiry.
+
+(III) In the third place, Darwin contributed largely to a knowledge of
+the factors in the evolution-process, especially by his analysis of
+what occurs in the case of domestic animals and cultivated plants, and
+by his elaboration of the theory of Natural Selection which Alfred
+Russel Wallace independently stated at the same time, and of which
+there had been a few previous suggestions of a more or less vague
+description. It was here that Darwin's originality was greatest, for
+he revealed to naturalists the many different forms--often very
+subtle--which natural selection takes, and with the insight of a
+disciplined scientific imagination he realised what a mighty engine of
+progress it has been and is.
+
+(IV) As an epoch-marking contribution, not only to AEtiology but to
+Natural History in the widest sense, we rank the picture which Darwin
+gave to the world of the web of life, that is to say, of the
+inter-relations and linkages in Nature. For the Biology of the
+individual--if that be not a contradiction in terms--no idea is more
+fundamental than that of the correlation of organs, but Darwin's most
+characteristic contribution was not less fundamental,--it was the idea
+of the correlation of organisms. This, again, was not novel; we find
+it in the works of naturalists like Christian Conrad Sprengel,
+Gilbert White, and Alexander von Humboldt, but the realisation of its
+full import was distinctly Darwinian.
+
+
+_As Regards the General Idea of Organic Evolution_
+
+While it is true, as Prof. H. F. Osborn puts it, that "'Before and
+after Darwin' will always be the _ante et post urbem conditam_ of
+biological history," it is also true that the general idea of organic
+evolution is very ancient. In his admirable sketch _From the Greeks to
+Darwin_,[1] Prof. Osborn has shown that several of the ancient
+philosophers looked upon Nature as a gradual development and as still
+in process of change. In the suggestions of Empedocles, to take the
+best instance, there were "four sparks of truth,--first, that the
+development of life was a gradual process; second, that plants were
+evolved before animals; third, that imperfect forms were gradually
+replaced (not succeeded) by perfect forms; fourth, that the natural
+cause of the production of perfect forms was the extinction of the
+imperfect."[2] But the fundamental idea of one stage giving origin to
+another was absent. As the blue AEgean teemed with treasures of beauty
+and threw many upon its shores, so did Nature produce like a fertile
+artist what had to be rejected as well as what was able to survive,
+but the idea of one species emerging out of another was not yet
+conceived.
+
+Aristotle's views of Nature[3] seem to have been more definitely
+evolutionist than those of his predecessors, in this sense, at least,
+that he recognised not only an ascending scale, but a genetic series
+from polyp to man and an age-long movement towards perfection. "It is
+due to the resistance of matter to form that Nature can only rise by
+degrees from lower to higher types." "Nature produces those things
+which, being continually moved by a certain principle contained in
+themselves, arrive at a certain end."
+
+To discern the outcrop of evolution-doctrine in the long interval
+between Aristotle and Bacon seems to be very difficult, and some of
+the instances that have been cited strike one as forced. Epicurus and
+Lucretius, often called poets of evolution, both pictured animals as
+arising directly out of the earth, very much as Milton's lion long
+afterwards pawed its way out. Even when we come to Bruno who wrote
+that "to the sound of the harp of the Universal Apollo (the World
+Spirit), the lower organisms are called by stages to higher, and the
+lower stages are connected by intermediate forms with the higher,"
+there is great room, as Prof. Osborn points out,[4] for difference of
+opinion as to how far he was an evolutionist in our sense of the term.
+
+The awakening of natural science in the sixteenth century brought the
+possibility of a concrete evolution theory nearer, and in the early
+seventeenth century we find evidences of a new spirit--in the
+embryology of Harvey and the classifications of Ray. Besides sober
+naturalists there were speculative dreamers in the sixteenth and
+seventeenth centuries who had at least got beyond static formulae,
+but, as Professor Osborn points out,[5] "it is a very striking fact,
+that the basis of our modern methods of studying the Evolution problem
+was established not by the early naturalists nor by the speculative
+writers, but by the Philosophers." He refers to Bacon, Descartes,
+Leibnitz, Hume, Kant, Lessing, Herder, and Schelling. "They alone were
+upon the main track of modern thought. It is evident that they were
+groping in the dark for a working theory of the Evolution of life, and
+it is remarkable that they clearly perceived from the outset that the
+point to which observation should be directed was not the past but the
+present mutability of species, and further, that this mutability was
+simply the variation of individuals on an extended scale."
+
+Bacon seems to have been one of the first to think definitely about
+the mutability of species, and he was far ahead of his age in his
+suggestion of what we now call a Station of Experimental Evolution.
+Leibnitz discusses in so many words how the species of animals may be
+changed and how intermediate species may once have linked those that
+now seem discontinuous. "All natural orders of beings present but a
+single chain".... "All advances by degrees in Nature, and nothing by
+leaps." Similar evolutionist statements are to be found in the works
+of the other "philosophers," to whom Prof. Osborn refers, who were,
+indeed, more scientific than the naturalists of their day. It must be
+borne in mind that the general idea of organic evolution--that the
+present is the child of the past--is in great part just the idea of
+human history projected upon the natural world, differentiated by the
+qualification that the continuous "Becoming" has been wrought out by
+forces inherent in the organisms themselves and in their environment.
+
+A reference to Kant[6] should come in historical order after Buffon,
+with whose writings he was acquainted, but he seems, along with Herder
+and Schelling, to be best regarded as the culmination of the
+evolutionist philosophers--of those at least who interested themselves
+in scientific problems. In a famous passage he speaks of "the
+agreement of so many kinds of animals in a certain common plan of
+structure" ... an "analogy of forms" which "strengthens the
+supposition that they have an actual blood-relationship, due to
+derivation from a common parent." He speaks of "the great Family of
+creatures, for as a Family we must conceive it, if the above-mentioned
+continuous and connected relationship has a real foundation." Prof.
+Osborn alludes to the scientific caution which led Kant, biology being
+what it was, to refuse to entertain the hope "that a Newton may one
+day arise even to make the production of a blade of grass
+comprehensible, according to natural laws ordained by no intention."
+As Prof. Haeckel finely observes, Darwin rose up as Kant's Newton.[7]
+
+The scientific renaissance brought a wealth of fresh impressions and
+some freedom from the tyranny of tradition, and the twofold stimulus
+stirred the speculative activity of a great variety of men from old
+Claude Duret of Moulins, of whose weird transformism (1609) Dr. Henry
+de Varigny[8] gives us a glimpse, to Lorenz Oken (1779-1851) whose
+writings are such mixtures of sense and nonsense that some regard him
+as a far-seeing prophet and others as a fatuous follower of
+intellectual will-o'-the-wisps. Similarly, for De Maillet, Maupertuis,
+Diderot, Bonnet, and others, we must agree with Professor Osborn that
+they were not actually in the main Evolution movement. Some have been
+included in the roll of honour on very slender evidence, Robinet for
+instance, whose evolutionism seems to us extremely dubious.[9]
+
+The first naturalist to give a broad and concrete expression to the
+evolutionist doctrine of descent was Buffon (1707-1788), but it is
+interesting to recall the fact that his contemporary Linnaeus
+(1707-1778), protagonist of the counter-doctrine of the fixity of
+species,[10] went the length of admitting (in 1762) that new species
+might arise by inter-crossing. Buffon's position among the pioneers of
+the evolution-doctrine is weakened by his habit of vacillating between
+his own conclusions and the orthodoxy of the Sorbonne, but there is no
+doubt that he had firm grasp of the general idea of "l'enchainment des
+etres."
+
+Erasmus Darwin (1731-1802), probably influenced by Buffon, was another
+firm evolutionist, and the outline of his argument in the
+_Zoonomia_[11] might serve in part at least to-day. "When we revolve
+in our minds the metamorphoses of animals, as from the tadpole to the
+frog; secondly, the changes produced by artificial cultivation, as in
+the breeds of horses, dogs, and sheep; thirdly, the changes produced
+by conditions of climate and of season, as in the sheep of warm
+climates being covered with hair instead of wool, and the hares and
+partridges of northern climates becoming white in winter: when,
+further, we observe the changes of structure produced by habit, as
+seen especially in men of different occupations; or the changes
+produced by artificial mutilation and prenatal influences, as in the
+crossing of species and production of monsters; fourth, when we
+observe the essential unity of plan in all warm-blooded animals,--we
+are led to conclude that they have been alike produced from a similar
+living filament".... "From thus meditating upon the minute portion of
+time in which many of the above changes have been produced, would it
+be too bold to imagine, in the great length of time since the earth
+began to exist, perhaps millions of years before the commencement of
+the history of mankind, that all warm-blooded animals have arisen from
+one living filament?"... "This idea of the gradual generation of all
+things seems to have been as familiar to the ancient philosophers as
+to the modern ones, and to have given rise to the beautiful
+hieroglyphic figure of the [Greek: proton oon], or first great egg,
+produced by night, that is, whose origin is involved in obscurity, and
+animated by [Greek: Eros], that is, by Divine Love; from whence
+proceeded all things which exist."
+
+Lamarck (1744-1829) seems to have become an evolutionist
+independently of Erasmus Darwin's influence, though the parallelism
+between them is striking. He probably owed something to Buffon, but he
+developed his theory along a different line. Whatever view be held in
+regard to that theory there is no doubt that Lamarck was a
+thorough-going evolutionist. Professor Haeckel speaks of the
+_Philosophie Zoologique_ as "the first connected and thoroughly
+logical exposition of the theory of descent."[12]
+
+Besides the three old masters, as we may call them, Buffon, Erasmus
+Darwin, and Lamarck, there were other quite convinced pre-Darwinian
+evolutionists. The historian of the theory of descent must take
+account of Treviranus whose _Biology or Philosophy of Animate Nature_
+is full of evolutionary suggestions; of Etienne Geoffroy St. Hilaire,
+who in 1830, before the French Academy of Sciences, fought with
+Cuvier, the fellow-worker of his youth, an intellectual duel on the
+question of descent; of Goethe, one of the founders of morphology and
+the greatest poet of Evolution--who, in his eighty-first year, heard
+the tidings of Geoffrey St. Hilaire's defeat with an interest which
+transcended the political anxieties of the time; and of many others
+who had gained with more or less confidence and clearness a new
+outlook on Nature. It will be remembered that Darwin refers to
+thirty-four more or less evolutionist authors in his Historical
+Sketch, and the list might be added to. Especially when we come near
+to 1858 do the numbers increase, and one of the most remarkable, as
+also most independent champions of the evolution-idea before that date
+was Herbert Spencer, who not only marshalled the arguments in a very
+forcible way in 1852, but applied the formula in detail in his
+_Principles of Psychology_ in 1855.[13]
+
+It is right and proper that we should shake ourselves free from all
+creationist appreciations of Darwin, and that we should recognise the
+services of pre-Darwinian evolutionists who helped to make the time
+ripe, yet one cannot help feeling that the citation of them is apt to
+suggest two fallacies. It may suggest that Darwin simply entered into
+the labours of his predecessors, whereas, as a matter of fact, he knew
+very little about them till after he had been for years at work. To
+write, as Samuel Butler did, "Buffon planted, Erasmus Darwin and
+Lamarck watered, but it was Mr. Darwin who said 'That fruit is ripe,'
+and shook it into his lap" ... seems to us a quite misleading version
+of the facts of the case. The second fallacy which the historical
+citation is a little apt to suggest is that the filiation of ideas is
+a simple problem. On the contrary, the history of an idea, like the
+pedigree of an organism, is often very intricate, and the evolution of
+the evolution-idea is bound up with the whole progress of the world.
+Thus in order to interpret Darwin's clear formulation of the idea of
+organic evolution and his convincing presentation of it, we have to do
+more than go back to his immediate predecessors, such as Buffon,
+Erasmus Darwin, and Lamarck; we have to inquire into the acceptance of
+evolutionary conceptions in regard to other orders of facts, such as
+the earth and the solar system;[14] we have to realise how the growing
+success of scientific interpretation along other lines gave confidence
+to those who refused to admit that there was any domain from which
+science could be excluded as a trespasser; we have to take account of
+the development of philosophical thought, and even of theological and
+religious movements; we should also, if we are wise enough, consider
+social changes. In short, we must abandon the idea that we can
+understand the history of any science as such, without reference to
+contemporary evolution in other departments of activity.
+
+While there were many evolutionists before Darwin, few of them were
+expert naturalists and few were known outside a small circle; what was
+of much more importance was that the genetic view of Nature was
+insinuating itself in regard to other than biological orders of facts,
+here a little and there a little, and that the scientific spirit had
+ripened since the days when Cuvier laughed Lamarck out of court. How
+was it that Darwin succeeded where others had failed? Because, in the
+first place, he had clear visions--"pensees de la jeunesse, executees
+par l'age mur"--which a University curriculum had not made impossible,
+which the _Beagle voyage_ made vivid, which an unrivalled British
+doggedness made real--visions of the web of life, of the fountain of
+change within the organism, of the struggle for existence and its
+winnowing, and of the spreading genealogical tree. Because, in the
+second place, he put so much grit into the verification of his
+visions, putting them to the proof in an argument which is of its
+kind--direct demonstration being out of the question--quite
+unequalled. Because, in the third place, he broke down the opposition
+which the most scientific had felt to the seductive modal formula of
+evolution by bringing forward a more plausible theory of the process
+than had been previously suggested. Nor can one forget, since
+questions of this magnitude are human and not merely academic, that he
+wrote so that all men could understand.
+
+
+_As Regards the Factors of Evolution_
+
+It is admitted by all who are acquainted with the history of biology
+that the general idea of organic evolution as expressed in the
+Doctrine of Descent was quite familiar to Darwin's grandfather and to
+others before and after him, as we have briefly indicated. It must
+also be admitted that some of these pioneers of evolutionism did more
+than apply the evolution-idea as a modal formula of becoming, they
+began to inquire into the factors in the process. Thus there were
+pre-Darwinian theories of evolution, and to these we must now briefly
+refer.[15]
+
+In all biological thinking we have to work with the categories
+Organism--Function--Environment, and theories of evolution may be
+classified in relation to these. To some it has always seemed that the
+fundamental fact is the living organism,--a creative agent, a striving
+will, a changeful Proteus, selecting its environment, adjusting itself
+to it, self-differentiating and self-adaptive. The necessity of
+recognising the importance of the organism is admitted by all
+Darwinians who start with inborn variations, but it is open to
+question whether the whole truth of what we might call the Goethian
+position is exhausted in the postulate of inherent variability.
+
+To others it has always seemed that the emphasis should be laid on
+Function,--on use and disuse, on doing and not doing. Practice makes
+perfect; _c'est a force de forger qu'on devient forgeron_. This is one
+of the fundamental ideas of Lamarckism; to some extent it met with
+Darwin's approval; and it finds many supporters to-day. One of the
+ablest of these--Mr. Francis Darwin--has recently given strong reasons
+for combining a modernised Lamarckism with what we usually regard as
+sound Darwinism.[16]
+
+To others it has always seemed that the emphasis should be laid on the
+Environment, which wakes the organism to action, prompts it to change,
+makes dints upon it, moulds it, prunes it, and finally, perhaps, kills it.
+It is again impossible to doubt that there is truth in this view, for even
+if environmentally induced "modifications" be not transmissible,
+environmentally induced "variations" are; and even if the direct influence
+of the environment be less important than many enthusiastic supporters of
+this view--may we call them Buffonians--think, there remains the indirect
+influence which Darwinians in part rely on,--the eliminative process. Even
+if the extreme view be held that the only form of discriminate elimination
+that counts is inter-organismal competition, this might be included under
+the rubric of the animate environment.
+
+In many passages Buffon[17] definitely suggested that environmental
+influences--especially of climate and food--were directly productive
+of changes in organisms, but he did not discuss the question of the
+transmissibility of the modifications so induced, and it is difficult
+to gather from his inconsistent writings what extent of transformation
+he really believed in. Prof. Osborn says of Buffon: "The struggle for
+existence, the elimination of the least-perfected species, the contest
+between the fecundity of certain species and their constant
+destruction, are all clearly expressed in various passages." He quotes
+two of these:[18]
+
+"Le cours ordinaire de la nature vivante, est en general toujours
+constant, toujours le meme; son mouvement, toujours regulier, roule
+sur deux points inebranlables: l'un, la fecondite sans bornes donnee a
+toutes les especes; l'autre, les obstacles sans nombre qui reduisent
+cette fecondite a une mesure determinee et ne laissent en tout temps
+qu'a peu pres la meme quantite d'individus de chaque espece" ... "Les
+especes les moins parfaites, les plus delicates, les plus pesantes,
+les moins agissantes, les moins armees, etc., ont deja disparu ou
+disparaitront.".
+
+Erasmus Darwin[19] had a firm grip of the "idea of the gradual
+formation and improvement of the Animal world," and he had his theory
+of the process. No sentence is more characteristic than this: "All
+animals undergo transformations which are in part produced by their
+own exertions, in response to pleasures and pains, and many of these
+acquired forms or propensities are transmitted to their posterity."
+This is Lamarckism before Lamarck, as his grandson pointed out. His
+central idea is that wants stimulate efforts and that these result in
+improvements which subsequent generations make better still. He
+realised something of the struggle for existence and even pointed out
+that this advantageously checks the rapid multiplication. "As Dr.
+Krause points out, Darwin just misses the connection between this
+struggle and the Survival of the Fittest."[20]
+
+Lamarck[21] (1744-1829) seems to have thought out his theory of
+evolution without any knowledge of Erasmus Darwin's which it closely
+resembled. The central idea of his theory was the cumulative
+inheritance of functional modifications. "Changes in environment bring
+about changes in the habits of animals. Changes in their wants
+necessarily bring about parallel changes in their habits. If new wants
+become constant or very lasting, they form new habits, the new habits
+involve the use of new parts, or a different use of old parts, which
+results finally in the production of new organs and the modification
+of old ones." He differed from Buffon in not attaching importance, as
+far as animals are concerned, to the direct influence of the
+environment, "for environment can effect no direct change whatever
+upon the organisation of animals," but in regard to plants he agreed
+with Buffon that external conditions directly moulded them.
+
+Treviranus[22] (1776-1837), whom Huxley ranked beside Lamarck, was on
+the whole Buffonian, attaching chief importance to the influence of a
+changeful environment both in modifying and in eliminating, but he was
+also Goethian, for instance in his idea that species like individuals
+pass through periods of growth, full bloom, and decline. "Thus, it is
+not only the great catastrophes of Nature which have caused
+extinction, but the completion of cycles of existence, out of which
+new cycles have begun." A characteristic sentence is quoted by Prof.
+Osborn: "In every living being there exists a capability of an endless
+variety of form-assumption; each possesses the power to adapt its
+organisation to the changes of the outer world, and it is this power,
+put into action by the change of the universe, that has raised the
+simple zoophytes of the primitive world to continually higher stages
+of organisation, and has introduced a countless variety of species
+into animate Nature."
+
+Goethe[23] (1749-1832), who knew Buffon's work but not Lamarck's, is
+peculiarly interesting as one of the first to use the evolution-idea
+as a guiding hypothesis, e.g. in the interpretation of vestigial
+structures in man, and to realise that organisms express an attempt to
+make a compromise between specific inertia and individual change. He
+gave the finest expression that science has yet known--if it has known
+it--of the kernel-idea of what is called "bathmism," the idea of an
+"inherent growth-force"--and at the same time he held that "the way of
+life powerfully reacts upon all form" and that the orderly growth of
+form "yields to change from externally acting causes."
+
+Besides Buffon, Erasmus Darwin, Lamarck, Treviranus, and Goethe,
+there were other "pioneers of evolution," whose views have been often
+discussed and appraised. Etienne Geoffroy Saint-Hilaire (1772-1884),
+whose work Goethe so much admired, was on the whole Buffonian,
+emphasising the direct action of the changeful _milieu_. "Species vary
+with their environment, and existing species have descended by
+modification from earlier and somewhat simpler species." He had a
+glimpse of the selection idea, and believed in mutations or sudden
+leaps--induced in the embryonic condition by external influences. The
+complete history of evolution-theories will include many instances of
+guesses at truth which were afterwards substantiated, thus the
+geographer von Buch (1773-1853) detected the importance of the
+Isolation factor on which Wagner, Romanes, Gulick and others have laid
+great stress, but we must content ourselves with recalling one other
+pioneer, the author of the _Vestiges of Creation_ (1844), a work which
+passed through ten editions in nine years and certainly helped to
+harrow the soil for Darwin's sowing. As Darwin said, "it did excellent
+service in this country in calling attention to the subject, in
+removing prejudice, and in thus preparing the ground for the reception
+of analogous views."[24] Its author, Robert Chambers (1802-1871) was
+in part a Buffonian--maintaining that environment moulded organisms
+adaptively, and in part a Goethian--believing in an inherent
+progressive impulse which lifted organisms from one grade of
+organisation to another.
+
+
+_As Regards Natural Selection_
+
+The only thinker to whom Darwin was directly indebted, so far as the
+theory of Natural Selection is concerned, was Malthus, and we may once
+more quote the well-known passage in the Autobiography: "In October,
+1838, that is, fifteen months after I had begun my systematic enquiry,
+I happened to read for amusement 'Malthus on Population,' and being
+well prepared to appreciate the struggle for existence which
+everywhere goes on from long-continued observation of the habits of
+animals and plants, it at once struck me that under these
+circumstances favourable variations would tend to be preserved, and
+unfavourable ones to be destroyed. The result of this would be the
+formation of new species."[25]
+
+Although Malthus gives no adumbration of the idea of Natural Selection
+in his exposition of the eliminative processes which go on in mankind,
+the suggestive value of his essay is undeniable, as is strikingly
+borne out by the fact that it gave to Alfred Russel Wallace also "the
+long-sought clue to the effective agent in the evolution of organic
+species."[26] One day in Ternate when he was resting between fits of
+fever, something brought to his recollection the work of Malthus which
+he had read twelve years before. "I thought of his clear exposition of
+'the positive checks to increase'--disease, accidents, war, and
+famine--which keep down the population of savage races to so much
+lower an average than that of more civilized peoples. It then occurred
+to me that these causes or their equivalents are continually acting in
+the case of animals also; and as animals usually breed much more
+rapidly than does mankind, the destruction every year from these
+causes must be enormous in order to keep down the numbers of each
+species, since they evidently do not increase regularly from year to
+year, as otherwise the world would long ago have been densely crowded
+with those that breed most quickly. Vaguely thinking over the enormous
+and constant destruction which this implied, it occurred to me to ask
+the question, Why do some die and some live? And the answer was
+clearly, that on the whole the best fitted live. From the effects of
+disease the most healthy escaped; from enemies the strongest, the
+swiftest, or the most cunning; from famine the best hunters or those
+with the best digestion; and so on. Then it suddenly flashed upon me
+that this self-acting process would necessarily _improve the race_,
+because in every generation the inferior would inevitably be killed
+off and the superior would remain--that is, _the fittest would
+survive_."[27] We need not apologise for this long quotation, it is a
+tribute to Darwin's magnanimous colleague, the Nestor of the
+evolutionist camp,--and it probably indicates the line of thought
+which Darwin himself followed. It is interesting also to recall the
+fact that in 1852, when Herbert Spencer wrote his famous _Leader_
+article on "The Development Hypothesis" in which he argued powerfully
+for the thesis that the whole animate world is the result of an
+age-long process of natural transformation, he wrote for _The
+Westminster Review_ another important essay, "A Theory of Population
+deduced from the General Law of Animal Fertility," towards the close
+of which he came within an ace of recognising that the struggle for
+existence was a factor in organic evolution. At a time when pressure
+of population was practically interesting men's minds, Darwin,
+Wallace, and Spencer were being independently led from a social
+problem to a biological theory. There could be no better illustration,
+as Prof. Patrick Geddes has pointed out, of the Comtian thesis that
+science is a "social phenomenon."
+
+Therefore, as far more important than any further ferreting out of
+vague hints of Natural Selection in books which Darwin never read, we
+would indicate by a quotation the view that the central idea in
+Darwinism is correlated with contemporary social evolution. "The
+substitution of Darwin for Paley as the chief interpreter of the order
+of nature is currently regarded as the displacement of an
+anthropomorphic view by a purely scientific one: a little reflection,
+however, will show that what has actually happened has been merely the
+replacement of the anthropomorphism of the eighteenth century by that
+of the nineteenth. For the place vacated by Paley's theological and
+metaphysical explanation has simply been occupied by that suggested to
+Darwin and Wallace by Malthus in terms of the prevalent severity of
+industrial competition, and those phenomena of the struggle for
+existence which the light of contemporary economic theory has enabled
+us to discern, have thus come to be temporarily exalted into a
+complete explanation of organic progress."[28] It goes without saying
+that the idea suggested by Malthus was developed by Darwin into a
+biological theory which was then painstakingly verified by being used
+as an interpretative formula, and that the validity of a theory so
+established is not affected by what suggested it, but the practical
+question which this line of thought raises in the mind is this: if
+Biology did thus borrow with such splendid results from social theory,
+why should we not more deliberately repeat the experiment?
+
+Darwin was characteristically frank and generous in admitting that the
+principle of Natural Selection had been independently recognised by
+Dr. W. C. Wells in 1813 and by Mr. Patrick Matthew in 1831, but he had
+no knowledge of these anticipations when he published the first
+edition of _The Origin of Species_. Wells, whose "Essay on Dew" is
+still remembered, read in 1813 before the Royal Society a short paper
+entitled "An Account of a White Female, part of whose skin resembles
+that of a Negro" (published in 1818). In this communication, as Darwin
+said, "he observes, firstly, that all animals tend to vary in some
+degree, and, secondly, that agriculturists improve their domesticated
+animals by selection; and then, he adds, but what is done in this
+latter case 'by art, seems to be done with equal efficacy, though more
+slowly, by nature, in the formation of varieties of mankind, fitted
+for the country which they inhabit.'"[29] Thus Wells had the clear
+idea of survival dependent upon a favourable variation, but he makes
+no more use of the idea and applies it only to man. There is not in
+the paper the least hint that the author ever thought of generalising
+the remarkable sentence quoted above.
+
+Of Mr. Patrick Matthew, who buried his treasure in an appendix to a
+work on _Naval Timber and Arboriculture_, Darwin said that "he clearly
+saw the full force of the principle of natural selection." In 1860
+Darwin wrote--very characteristically--about this to Lyell: "Mr.
+Patrick Matthew publishes a long extract from his work on _Naval
+Timber and Arboriculture_, published in 1831, in which he briefly but
+completely anticipates the theory of Natural Selection. I have ordered
+the book, as some passages are rather obscure, but it is certainly, I
+think, a complete but not developed anticipation. Erasmus always said
+that surely this would be shown to be the case some day. Anyhow, one
+may be excused in not having discovered the fact in a work on Naval
+Timber."[30]
+
+De Quatrefages and De Varigny have maintained that the botanist Naudin
+stated the theory of evolution by natural selection in 1852. He
+explains very clearly the process of artificial selection, and says
+that in the garden we are following Nature's method. "We do not think
+that Nature has made her species in a different fashion from that in
+which we proceed ourselves in order to make our variations." But, as
+Darwin said, "he does not show how selection acts under nature."
+Similarly it must be noted in regard to several pre-Darwinian pictures
+of the struggle for existence (such as Herder's, who wrote in 1790
+"All is in struggle ... each one for himself" and so on), that a
+recognition of this is only the first step in Darwinism.
+
+Profs. E. Perrier and H. F. Osborn have called attention to a
+remarkable anticipation of the selection-idea which is to be found in
+the speculations of Etienne Geoffroy Saint-Hilaire (1825-1828) on the
+evolution of modern Crocodilians from the ancient Teleosaurs. Changing
+environment induced changes in the respiratory system and far-reaching
+consequences followed. The atmosphere, acting upon the pulmonary
+cells, brings about "modifications which are favourable or destructive
+('funestes'); these are inherited, and they influence all the rest of
+the organisation of the animal because if these modifications lead to
+injurious effects the animals which exhibit them perish and are
+replaced by others of a somewhat different form, a form changed so as
+to be adapted to (a la convenance) the new environment."
+
+Prof. E. B. Poulton[31] has shown that the anthropologist James Cowles
+Prichard (1786-1848) must be included even in spite of himself among
+the precursors of Darwin. In some passages of the second edition of
+his _Researches into the Physical History of Mankind_ (1826), he
+certainly talks evolution and anticipates Prof. Weismann in denying
+the transmission of acquired characters. He is, however, sadly
+self-contradictory and his evolutionism weakens in subsequent
+editions--the only ones that Darwin saw. Prof. Poulton finds in
+Prichard's work a recognition of the operation of Natural Selection.
+"After inquiring how it is that 'these varieties are developed and
+preserved in connexion with particular climates and differences of
+local situation,' he gives the following very significant answer: 'One
+cause which tends to maintain this relation is obvious. Individuals
+and families, and even whole colonies perish and disappear in climates
+for which they are, by peculiarity of constitution, not adapted. Of
+this fact proofs have been already mentioned.'" Mr. Francis Darwin and
+Prof. A. C. Seward discuss Prichard's "anticipations" in _More Letters
+of Charles Darwin_, Vol. _I._ p. 43, and come to the conclusion that
+the evolutionary passages are entirely neutralised by others of an
+opposite trend. There is the same difficulty with Buffon.
+
+Hints of the idea of Natural Selection have been detected elsewhere.
+James Watt,[32] for instance, has been reported as one of the
+anticipators (1851). But we need not prolong the inquiry further,
+since Darwin did not know of any anticipations until after he had
+published the immortal work of 1859, and since none of those who got
+hold of the idea made any use of it. What Darwin did was to follow the
+clue which Malthus gave him, to realise, first by genius and
+afterwards by patience, how the complex and subtle struggle for
+existence works out a natural selection of those organisms which vary
+in the direction of fitter adaptation to the conditions of their life.
+So much success attended his application of the Selection-formula that
+for a time he regarded Natural Selection as almost the sole factor in
+evolution, variations being pre-supposed; gradually, however, he came
+to recognise that there was some validity in the factors which had
+been emphasised by Lamarck and by Buffon, and in his well known
+summing up in the sixth edition of the _Origin_ he says of the
+transformation of species: "This has been effected chiefly through the
+natural selection of numerous successive, slight, favourable
+variations; aided in an important manner by the inherited effects of
+the use and disuse of parts; and in an unimportant manner, that is, in
+relation to adaptive structures, whether past or present, by the
+direct action of external conditions, and by variations which seem to
+us in our ignorance to arise spontaneously."
+
+To sum up: the idea of organic evolution, older than Aristotle, slowly
+developed from the stage of suggestion to the stage of verification,
+and the first convincing verification was Darwin's; from being an _a
+priori_ anticipation it has become an interpretation of nature, and
+Darwin is still the chief interpreter; from being a modal
+interpretation it has advanced to the rank of a causal theory, the
+most convincing part of which men will never cease to call Darwinism.
+
+FOOTNOTES:
+
+[Footnote 1: _Columbia University Biological Series_, Vol. I. New York
+and London, 1894. We must acknowledge our great indebtedness to this
+fine piece of work.]
+
+[Footnote 2: _op. cit._ p. 41.]
+
+[Footnote 3: See G. J. Romanes, "Aristotle as a Naturalist,"
+_Contemporary Review_, Vol. lix. p. 275, 1891; G. Pouchet, _La
+Biologie Aristotelique_, Paris, 1885; E. Zeller, _A History of Greek
+Philosophy_, London, 1881, and "Ueber die griechischen Vorgaenger
+Darwin's," _Abhandl. Berlin Akad._ 1878, pp. 111-124.]
+
+[Footnote 4: _op. cit._ p. 81.]
+
+[Footnote 5: _op. cit._ p. 87.]
+
+[Footnote 6: See Brock, "Die Stellung Kant's zur Deszendenztheorie,"
+_Biol. Centralbl._ viii. 1889, pp. 641-648. Fritz Schultze, _Kant und
+Darwin_, Jena, 1875.]
+
+[Footnote 7: Mr. Alfred Russel Wallace writes: "We claim for Darwin
+that he is the Newton of natural history, and that, just so surely as
+that the discovery and demonstration by Newton of the law of
+gravitation established order in place of chaos and laid a sure
+foundation for all future study of the starry heavens, so surely has
+Darwin, by his discovery of the law of natural selection and his
+demonstration of the great principle of the preservation of useful
+variations in the struggle for life, not only thrown a flood of light
+on the process of development of the whole organic world, but also
+established a firm foundation for all future study of nature"
+(_Darwinism_, London, 1889, p. 9). See also Prof. Karl Pearson's
+_Grammar of Science_ (2nd edit.), London, 1900, p. 32. See Osborn,
+_op. cit._ p. 100.]
+
+[Footnote 8: _Experimental Evolution_. London, 1892. Chap. I. p. 14.]
+
+[Footnote 9: See J. Arthur Thomson, _The Science of Life_. London,
+1899, Chap. XVI. "Evolution of Evolution Theory."]
+
+[Footnote 10: See Carus Sterne (Ernst Krause), _Die allgemeine
+Weltanschauung in ihrer historischen Entwickelung_. Stuttgart, 1889.
+Chapter entitled "Bestaendigkeit oder Veraenderlichkeit der
+Naturwesen."]
+
+[Footnote 11: _Zoonomia, or the Laws of Organic Life_, 2 vols. London,
+1794; Osborn, _op. cit._ p. 145.]
+
+[Footnote 12: See Alpheus S. Packard, _Lamarck, the Founder of
+Evolution, His Life and Work, with Translations of his writings on
+Organic Evolution_. London, 1901.]
+
+[Footnote 13: See Edward Clodd, _Pioneers of Evolution_, London, p.
+161, 1897.]
+
+[Footnote 14: See Chapter ix. "The Genetic View of Nature" in J. T.
+Merz's _History of European Thought in the Nineteenth Century_, Vol.
+2, Edinburgh and London, 1903.]
+
+[Footnote 15: See Prof. W. A. Locy's _Biology and its Makers_. New
+York, 1908. Part II. "The Doctrine of Organic Evolution."]
+
+[Footnote 16: Presidential Address to the British Association meeting
+at Dublin in 1908.]
+
+[Footnote 17: See in particular Samuel Butler, _Evolution Old and
+New_, London, 1879; J. L. de Lanessan, "Buffon et Darwin," _Revue
+Scientifique_, XLIII. pp. 385-391, 425-432, 1889.]
+
+[Footnote 18: _op. cit._ p. 136.]
+
+[Footnote 19: See Ernest Krause and Charles Darwin, _Erasmus Darwin_,
+London, 1879.]
+
+[Footnote 20: Osborn, _op. cit._ p. 142.]
+
+[Footnote 21: See E. Perrier, _La Philosophie Zoologique avant
+Darwin_, Paris, 1884; A. de Quatrefages, _Darwin et ses Precurseurs
+Francais_, Paris, 1870; Packard, _op. cit._; also Claus, _Lamarck als
+Begruender der Descendenzlehre_, Wien, 1888; Haeckel, _Natural History
+of Creation_, Eng. transl. London, 1879; Lang, _Zur Charakteristik der
+Forschungswege von Lamarck und Darwin_, Jena, 1889.]
+
+[Footnote 22: See Huxley's article "Evolution in Biology,"
+_Encyclopaedia Britannica_ (9th edit.), 1879, pp. 744-751, and Sully's
+article, "Evolution in Philosophy," _ibid._ pp. 751-772.]
+
+[Footnote 23: See Haeckel, _Die Naturanschauung von Darwin, Goethe und
+Lamarck_, Jena, 1882.]
+
+[Footnote 24: _Origin of Species_ (6th edit.), p. xvii.]
+
+[Footnote 25: _The Life and Letters of Charles Darwin_, Vol. 1. p. 83.
+London, 1887.]
+
+[Footnote 26: A. R. Wallace, _My Life, a Record of Events and
+Opinions_, London, 1905, Vol. 1, p. 232.]
+
+[Footnote 27: _My Life_, Vol. 1. p. 361.]
+
+[Footnote 28: P. Geddes. article "Biology." _Chambers's
+Encyclopaedia._]
+
+[Footnote 29: _Origin of Species_ (6th edit.), p. xv.]
+
+[Footnote 30: _Life and Letters_, II, p. 301.]
+
+[Footnote 31: _Science Progress_, New Series, Vol. 1. 1897. "A
+Remarkable Anticipation of Modern Views on Evolution." See also Chap.
+VI. in _Essays on Evolution_, Oxford, 1908.]
+
+[Footnote 32: See Prof. Patrick Geddes's article "Variation and
+Selection," _Encyclopaedia Britannica_ (9th edit.) 1888.]
+
+
+
+
+II
+
+THE SELECTION THEORY
+
+BY AUGUST WEISMANN
+
+_Professor of Zoology in the University of Freiburg_ (_Baden_)
+
+
+I. THE IDEA OF SELECTION
+
+Many and diverse were the discoveries made by Charles Darwin in the
+course of a long and strenuous life, but none of them has had so
+far-reaching an influence on the science and thought of his time as
+the theory of selection. I do not believe that the theory of evolution
+would have made its way so easily and so quickly after Darwin took up
+the cudgels in favour of it if he had not been able to support it by a
+principle which was capable of solving, in a simple manner, the
+greatest riddle that living nature presents to us,--I mean the
+purposiveness of every living form relative to the conditions of its
+life and its marvellously exact adaptation to these.
+
+Everyone knows that Darwin was not alone in discovering the principle
+of selection, and that the same idea occurred simultaneously and
+independently to Alfred Russel Wallace. At the memorable meeting of
+the Linnean Society on 1st July, 1858, two papers were read
+(communicated by Lyell and Hooker) both setting forth the same idea of
+selection. One was written by Charles Darwin in Kent, the other by
+Alfred Wallace in Ternate, in the Malay Archipelago. It was a splendid
+proof of the magnanimity of these two investigators, that they thus in
+all friendliness and without envy, united in laying their ideas
+before a scientific tribunal: their names will always shine side by
+side as two of the brightest stars in the scientific sky.
+
+The idea of selection set forth by the two naturalists was at the time
+absolutely new, but it was also so simple that Huxley could say of it
+later, "How extremely stupid not to have thought of that." As Darwin
+was led to the general doctrine of descent, not through the labours of
+his predecessors in the early years of the century, but by his own
+observations, so it was in regard to the principle of selection. He
+was struck by the innumerable cases of adaptation, as, for instance,
+that of the woodpeckers and tree-frogs to climbing, or the hooks and
+feather-like appendages of seeds, which aid in the distribution of
+plants, and he said to himself that an explanation of adaptations was
+the first thing to be sought for in attempting to formulate a theory
+of evolution.
+
+But since adaptations point to _changes_ which have been undergone by
+the ancestral forms of existing species, it is necessary, first of
+all, to inquire how far species in general are _variable_. Thus
+Darwin's attention was directed in the first place to the phenomenon
+of variability, and the use man has made of this, from very early
+times, in the breeding of his domesticated animals and cultivated
+plants. He inquired carefully how breeders set to work, when they
+wished to modify the structure and appearance of a species to their
+own ends, and it was soon clear to him that _selection for breeding
+purposes_ played the chief part.
+
+But how was it possible that such processes should occur in free
+nature? Who is here the breeder, making the selection, choosing out
+one individual to bring forth offspring and rejecting others? That was
+the problem that for a long time remained a riddle to him.
+
+Darwin himself relates how illumination suddenly came to him. He had
+been reading, for his own pleasure, Malthus' book on Population, and,
+as he had long known from numerous observations, that every species
+gives rise to many more descendants than ever attain to maturity, and
+that, therefore, the greater number of the descendants of a species
+perish without reproducing, the idea came to him that the decision as
+to which member of a species was to perish and which was to attain to
+maturity and reproduction might not be a matter of chance, but might
+be determined by the constitution of the individuals themselves,
+according as they were more or less fitted for survival. With this
+idea the foundation of the theory of selection was laid.
+
+In _artificial selection_ the breeder chooses out for pairing only
+such individuals as possess the character desired by him in a somewhat
+higher degree than the rest of the race. Some of the descendants
+inherit this character, often in a still higher degree, and if this
+method be pursued throughout several generations, the race is
+transformed in respect of that particular character.
+
+_Natural selection_ depends on the same three factors as _artificial
+selection_: on _variability_, _inheritance_, and _selection for
+breeding_, but this last is here carried out not by a breeder but by
+what Darwin called the "struggle for existence." This last factor is
+one of the special features of the Darwinian conception of nature.
+That there are carnivorous animals which take heavy toll in every
+generation of the progeny of the animals on which they prey, and that
+there are herbivores which decimate the plants in every generation had
+long been known, but it is only since Darwin's time that sufficient
+attention has been paid to the facts that, in addition to this regular
+destruction, there exists between the members of a species a keen
+competition for space and food, which limits multiplication, and that
+numerous individuals of each species perish because of unfavourable
+climatic conditions. The "struggle for existence," which Darwin
+regarded as taking the place of the human breeder in free nature, is
+not a direct struggle between carnivores and their prey, but is the
+assumed competition for survival between individuals _of the same_
+species, of which, on an average, only those survive to reproduce
+which have the greatest power of resistance, while the others, less
+favourably constituted, perish early. This struggle is so keen, that,
+within a limited area, where the conditions of life have long remained
+unchanged, of every species, whatever be the degree of fertility, only
+two, _on an average_, of the descendants of each pair survive; the
+others succumb either to enemies, or to disadvantages of climate, or
+to accident. A high degree of fertility is thus not an indication of
+the special success of a species, but of the numerous dangers that
+have attended its evolution. Of the six young brought forth by a pair
+of elephants in the course of their lives only two survive in a given
+area; similarly, of the millions of eggs which two thread-worms leave
+behind them only two survive. It is thus possible to estimate the
+dangers which threaten a species by its ratio of elimination, or,
+since this cannot be done directly, by its fertility.
+
+Although a great number of the descendants of each generation fall
+victims to accident, among those that remain it is still the greater
+or less fitness of the organism that determines the "selection for
+breeding purposes," and it would be incomprehensible if, in this
+competition, it were not ultimately, that is, on an average, the best
+equipped which survive, in the sense of living long enough to
+reproduce.
+
+Thus the principle of natural selection is _the selection of the best
+for reproduction_, whether the "best" refers to the whole
+constitution, to one or more parts of the organism, or to one or more
+stages of development. Every organ, every part, every character of an
+animal, fertility and intelligence included, must be improved in this
+manner, and be gradually brought up in the course of generations to
+its highest attainable state of perfection. And not only may
+improvement of parts be brought about in this way, but new parts and
+organs may arise, since, through the slow and minute steps of
+individual or "fluctuating" variations, a part may be added here or
+dropped out there, and thus something new is produced.
+
+The principle of selection solved the riddle as to how what was
+purposive could conceivably be brought about without the intervention
+of a directing power, the riddle which animate nature presents to our
+intelligence at every turn, and in face of which the mind of a Kant
+could find no way out, for he regarded a solution of it as not to be
+hoped for. For, even if we were to assume an evolutionary force that
+is continually transforming the most primitive and the simplest forms
+of life into ever higher forms, and the homogeneity of primitive times
+into the infinite variety of the present, we should still be unable to
+infer from this alone how each of the numberless forms adapted to
+particular conditions of life should have appeared _precisely at the
+right moment in the history of the earth_ to which their adaptations
+were appropriate, and precisely at the proper place in which all the
+conditions of life to which they were adapted occurred: the
+humming-birds at the same time as the flowers; the trichina at the
+same time as the pig; the bark-coloured moth at the same time as the
+oak, and the wasp-like moth at the same time as the wasp which
+protects it. Without processes of selection we should be obliged to
+assume a "pre-established harmony" after the famous Leibnitzian model,
+by means of which the clock of the evolution of organisms is so
+regulated as to strike in exact synchronism with that of the history
+of the earth! All forms of life are strictly adapted to the conditions
+of their life, and can persist under these conditions alone.
+
+There must therefore be an intrinsic connection between the conditions
+and the structural adaptations of the organism, and, _since the
+conditions of life cannot be determined by the animal itself, the
+adaptations must be called forth by the conditions_.
+
+The selection theory teaches us how this is conceivable, since it
+enables us to understand that there is a continual production of what
+is non-purposive as well as of what is purposive, but the purposive
+alone survives, while the non-purposive perishes in the very act of
+arising. This is the old wisdom taught long ago by Empedocles.
+
+
+II. THE LAMARCKIAN PRINCIPLE
+
+Lamarck, as is well known, formulated a definite theory of evolution
+at the beginning of the nineteenth century, exactly fifty years before
+the Darwin-Wallace principle of selection was given to the world. This
+brilliant investigator also endeavoured to support his theory by
+demonstrating forces which might have brought about the
+transformations of the organic world in the course of the ages. In
+addition to other factors, he laid special emphasis on the increased
+or diminished use of the parts of the body, assuming that the
+strengthening or weakening which takes place from this cause during
+the individual life, could be handed on to the offspring, and thus
+intensified and raised to the rank of a specific character. Darwin
+also regarded this _Lamarckian principle_, as it is now generally
+called, as a factor in evolution, but he was not fully convinced of
+the transmissibility of acquired characters.
+
+As I have here to deal only with the theory of selection, I need not
+discuss the Lamarckian hypothesis, but I must express my opinion that
+there is room for much doubt as to the cooeperation of this principle
+in evolution. Not only is it difficult to imagine how the transmission
+of functional modifications could take place, but, up to the present
+time, notwithstanding the endeavours of many excellent investigators,
+not a single actual proof of such inheritance has been brought
+forward. Semon's experiments on plants are, according to the botanist
+Pfeffer, not to be relied on, and even the recent, beautiful
+experiments made by Dr. Kammerer on salamanders, cannot, as I hope to
+show elsewhere, be regarded as proof, if only because they do not deal
+at all with functional modifications, that is, with modifications
+brought about by use, and it is to these _alone_ that the Lamarckian
+principle refers.
+
+
+III. OBJECTIONS TO THE THEORY OF SELECTION
+
+
+(_a_) _Saltatory evolution_
+
+The Darwinian doctrine of evolution depends essentially on _the
+cumulative augmentation_ of minute variations in the direction of
+utility. But can such minute variations, which are undoubtedly
+continually appearing among the individuals of the same species,
+possess any selection-value; can they determine which individuals are
+to survive, and which are to succumb; can they be increased by natural
+selection till they attain to the highest development of a purposive
+variation?
+
+To many this seems so improbable that they have urged a theory of
+evolution by leaps from species to species. Koelliker, in 1872,
+compared the evolution of species with the processes which we can
+observe in the individual life in cases of alternation of generations.
+But a polyp only gives rise to a medusa because it has itself arisen
+from one, and there can be no question of a medusa ever having arisen
+suddenly and _de novo_ from a polyp-bud, if only because both forms
+are adapted in their structure as a whole, and in every detail to the
+conditions of their life. A sudden origin, in a natural way, of
+numerous adaptations is inconceivable. Even the degeneration of a
+medusoid from a free-swimming animal to a mere brood-sac (gonophore)
+is not sudden and saltatory, but occurs by imperceptible modifications
+throughout hundreds of years, as we can learn from the numerous stages
+of the process of degeneration persisting at the same time in
+different species.
+
+If, then, the degeneration to a simple brood-sac takes place only by
+very slow transitions, each stage of which may last for centuries, how
+could the much more complex _ascending_ evolution possibly have taken
+place by sudden leaps? I regard this argument as capable of further
+extension, for wherever in nature we come upon degeneration, it is
+taking place by minute steps and with a slowness that makes it not
+directly perceptible, and I believe that this in itself justifies us
+in concluding that _the same must be true of ascending_ evolution. But
+in the latter case the goal can seldom be distinctly recognised while
+in cases of degeneration the starting-point of the process can often
+be inferred, because several nearly related species may represent
+different stages.
+
+In recent years Bateson in particular has championed the idea of
+saltatory, or so-called discontinuous evolution, and has collected a
+number of cases in which more or less marked variations have suddenly
+appeared. These are taken for the most part from among domesticated
+animals which have been bred and crossed for a long time, and it is
+hardly to be wondered at that their much mixed and much influenced
+germ-plasm should, under certain conditions, give rise to remarkable
+phenomena, often indeed producing forms which are strongly suggestive
+of monstrosities, and which would undoubtedly not survive in free
+nature, unprotected by man. I should regard such cases as due to an
+intensified germinal selection--though this is to anticipate a
+little--and from this point of view it cannot be denied that they have
+a special interest. But they seem to me to have no significance as far
+as the transformation of species is concerned, if only because of the
+extreme rarity of their occurrence.
+
+There are, however, many variations which have appeared in a sudden
+and saltatory manner, and some of these Darwin pointed out and
+discussed in detail: the copper beech, the weeping trees, the oak with
+"fern-like leaves," certain garden-flowers, etc. But none of them have
+persisted in free nature, or evolved into permanent types.
+
+On the other hand, wherever enduring types have arisen, we find traces
+of a gradual origin by successive stages, even if, at first sight,
+their origin may appear to have been sudden. This is the case with
+_seasonal Dimorphism_, the first known cases of which exhibited marked
+differences between the two generations, the winter and the summer
+brood. Take for instance the much discussed and studied form
+_Vanessa_ (_Araschnia_) _levana-prorsa_. Here the differences between
+the two forms are so great and so apparently disconnected, that one
+might almost believe it to be a sudden mutation, were it not that old
+transition-stages can be called forth by particular temperatures, and
+we know other butterflies, as for instance our Garden Whites, in which
+the differences between the two generations are not nearly so marked;
+indeed, they are so little apparent that they are scarcely likely to
+be noticed except by experts. Thus here again there are small initial
+steps, some of which, indeed, must be regarded as adaptations, such as
+the green-sprinkled or lightly tinted under-surface which gives them a
+deceptive resemblance to parsley or to Cardamine leaves.
+
+Even if saltatory variations do occur, we cannot assume that these
+_have ever led to forms which are capable of survival under the
+conditions of wild life_. Experience has shown that in plants which
+have suddenly varied the power of persistence is diminished.
+Korschinsky attributes to them weaknesses of organisation in general;
+"they bloom late, ripen few of their seeds, and show great
+sensitiveness to cold." These are not the characters which make for
+success in the struggle for existence.
+
+We must briefly refer here to the views--much discussed in the last
+decade--of H. de Vries, who believes that the roots of transformation
+must be sought for in _saltatory variations arising from internal
+causes_, and distinguishes such _mutations_, as he has called them,
+from ordinary individual variations, in that they breed true, that is,
+with strict in-breeding they are handed on pure to the next
+generation. I have elsewhere endeavoured to point out the weaknesses
+of this theory,[33] and I am the less inclined to return to it here
+that it now appears[34] that the far-reaching conclusions drawn by de
+Vries from his observations on the Evening Primrose, _Oenothera
+lamarckiana_, rest upon a very insecure foundation. The plant from
+which de Vries saw numerous "species"--his "mutations"--arise was not,
+as he assumed, a _wild species_ that had been introduced to Europe
+from America, but was probably a hybrid form which was first
+discovered in the Jardin des Plantes in Paris, and which does not
+appear to exist anywhere in America as a wild species.
+
+This gives a severe shock to the "Mutation theory," for the other
+_actually wild_ species with which de Vries experimented showed no
+"mutations" but yielded only negative results.
+
+Thus we come to the conclusion that Darwin[35] was right in regarding
+transformations as taking place by minute steps, which, if useful, are
+augmented in the course of innumerable generations, because their
+possessors more frequently survive in the struggle for existence.
+
+
+(_b_) _Selection-value of the initial steps_
+
+Is it possible that the insignificant deviations which we know as
+"individual variations" can form the beginning of a process of
+selection? Can they decide which is to perish and which to survive? To
+use a phrase of Romanes, can they have _selection-value_?
+
+Darwin himself answered this question, and brought together many
+excellent examples to show that differences, apparently insignificant
+because very small, might be of decisive importance for the life of
+the possessor. But it is by no means enough to bring forward cases of
+this kind, for the question is not merely whether finished adaptations
+have selection-value, but whether the first beginnings of these, and
+whether the small, I might almost say minimal increments, which have
+led up from these beginnings to the perfect adaptation, have also had
+selection-value. To this question even one who, like myself, has been
+for many years a convinced adherent of the theory of selection, can
+only reply: _We must assume so, but we cannot prove it in any case_.
+It is not upon demonstrative evidence that we rely when we champion
+the doctrine of selection as a scientific truth; we base our argument
+on quite other grounds. Undoubtedly there are many apparently
+insignificant features, which can nevertheless be shown to be
+adaptations--for instance, the thickness of the basin-shaped shell of
+the limpets that live among the breakers on the shore. There can be no
+doubt that the thickness of these shells, combined with their flat
+forms, protects the animals from the force of the waves breaking upon
+them,--but how have they become so thick? What proportion of thickness
+was sufficient to decide that of two variants of a limpet one should
+survive, the other be eliminated? We can say nothing more than that we
+infer from the present state of the shell, that it must have varied in
+regard to differences in shell-thickness, and that these differences
+must have had selection-value,--no proof therefore, but an assumption
+which we must show to be convincing.
+
+For a long time the marvellously complex _radiate_ and _lattice-work_
+skeletons of Radiolarians were regarded as a mere outflow of "Nature's
+infinite wealth of form," as an instance of a purely morphological
+character with no biological significance. But recent investigations
+have shown that these, too, have an adaptive significance (Haecker).
+The same thing has been shown by Schuett in regard to the lowly
+unicellular plants, the Peridineae, which abound alike on the surface
+of the ocean and in its depths. It has been shown that the long
+skeletal processes which grow out from these organisms have
+significance not merely as a supporting skeleton, but also as an
+extension of the superficial area, which increases the contact with
+the water-particles, and prevents the floating organisms from sinking.
+It has been established that the processes are considerably shorter in
+the colder layers of the ocean, and that they may be twelve times as
+long[36] in the warmer layers, thus corresponding to the greater or
+smaller amount of friction which takes place in the denser and less
+dense layers of the water.
+
+The Peridineae of the warmer ocean layers have thus become long-rayed,
+those of the colder layers short-rayed, not through the direct effect
+of friction on the protoplasm, but through processes of selection,
+which favoured the longer rays in warm water, since they kept the
+organism afloat, while those with short rays sank and were eliminated.
+If we put the question as to selection-value in this case, and ask how
+great the variations in the length of processes must be in order to
+possess selection-value; what can we answer except that these
+variations must have been minimal, and yet sufficient to prevent too
+rapid sinking and consequent elimination? Yet this very case would
+give the ideal opportunity for a mathematical calculation of the
+minimal selection-value, although of course it is not feasible from
+lack of data to carry out the actual calculation.
+
+But even in organisms of more than microscopic size there must
+frequently be minute, even microscopic differences which set going the
+process of selection, and regulate its progress to the highest
+possible perfection.
+
+Many tropical trees possess thick, leathery leaves, as a protection
+against the force of the tropical raindrops. The _direct_ influence of
+the rain cannot be the cause of this power of resistance, for the
+leaves, while they were still thin, would simply have been torn to
+pieces. Their toughness must therefore be referred to selection, which
+would favour the trees with slightly thicker leaves, though we cannot
+calculate with any exactness how great the first stages of increase in
+thickness must have been. Our hypothesis receives further support from
+the fact that, in many such trees, the leaves are drawn out into a
+beak-like prolongation (Stahl and Haberlandt) which facilitates the
+rapid falling off of the rain water, and also from the fact that the
+leaves, while they are still young, hang limply down in bunches which
+offer the least possible resistance to the rain. Thus there are here
+three adaptations which can only be interpreted as due to selection.
+The initial stages of these adaptations must undoubtedly have had
+selection-value.
+
+But even in regard to this case we are reasoning in a circle, not
+giving "proofs," and no one who does not wish to believe in the
+selection-value of the initial stages can be forced to do so. Among
+the many pieces of presumptive evidence a particularly weighty one
+seems to me to be _the smallness of the steps of progress_ which we
+can observe in certain cases, as for instance in leaf-imitation among
+butterflies, and in mimicry generally. The resemblance to a leaf, for
+instance of a particular Kallima, seems to us so close as to be
+deceptive, and yet we find in another individual, or it may be in many
+others, a spot added which increases the resemblance, and which could
+not have become fixed unless the increased deceptiveness so produced
+had frequently led to the overlooking of its much persecuted
+possessor. But if we take the selection-value of the initial stages
+for granted, we are confronted with the further question which I
+myself formulated many years ago: How does it happen _that the
+necessary beginnings of a useful variation are always present_? How
+could insects which live upon or among green leaves become all green,
+while those that live on bark become brown? How have the desert
+animals become yellow and the Arctic animals white? Why were the
+necessary variations always present? How could the green locust lay
+brown eggs, or the privet caterpillar develop white and lilac-coloured
+lines on its green skin?
+
+It is of no use answering to this that the question is wrongly
+formulated[37] and that it is the converse that is true; that the
+process of selection takes place in accordance with the variations
+that present themselves. This proposition is undeniably true, but so
+also is another, which apparently negatives it: the variation required
+has in the majority of cases actually presented itself. Selection
+cannot solve this contradiction; it does not call forth the useful
+variation, but simply works upon it. The ultimate reason why one and
+the same insect should occur in green and in brown, as often happens
+in caterpillars and locusts, lies in the fact that variations towards
+brown presented themselves, and so also did variations towards green:
+_the kernel of the riddle lies in the varying_, and for the present we
+can only say, that small variations in different directions present
+themselves in every species. Otherwise so many different kinds of
+variations could not have arisen. I have endeavoured to explain this
+remarkable fact by means of the intimate processes that must take
+place within the germ-plasm, and I shall return to the problem when
+dealing with "germinal selection."
+
+We have, however, to make still greater demands on variation, for it
+is not enough that the necessary variation should occur in isolated
+individuals, because in that case there would be small prospect of its
+being preserved, notwithstanding its utility. Darwin at first
+believed, that even single variations might lead to transformation of
+the species, but later he became convinced that this was impossible,
+at least without the cooeperation of other factors, such as isolation
+and sexual selection.
+
+In the case of the _green caterpillars with bright longitudinal
+stripes_, numerous individuals exhibiting this useful variation must
+have been produced to start with. In all higher, that is,
+multicellular organisms, the germ-substance is the source of all
+transmissible variations, and this germ-plasm is not a simple
+substance but is made up of many primary constituents. The question
+can therefore be more precisely stated thus: How does it come about
+that in so many cases the useful variations present themselves in
+numbers just where they are required, the white oblique lines in the
+leaf-caterpillar on the under surface of the body, the accompanying
+coloured stripes just above them? And, further, how has it come about
+that in grass caterpillars, not oblique but longitudinal stripes,
+which are more effective for concealment among grass and plants, have
+been evolved? And finally, how is it that the same Hawk-moth
+caterpillars, which to-day show oblique stripes, possessed
+longitudinal stripes in Tertiary times? We can read this fact from the
+history of their development, and I have before attempted to show the
+biological significance of this change of colour.[38]
+
+For the present I need only draw the conclusion that one and the same
+caterpillar may exhibit the initial stages of both, and that it
+depends on the manner in which these marking elements are
+_intensified_ and _combined_ by natural selection whether whitish
+longitudinal or oblique stripes should result. In this case then the
+"useful variations" were actually "always there," and we see that in
+the same group of Lepidoptera, e.g. species of Sphingidae, evolution
+has occurred in both directions according to whether the form lived
+among grass or on broad leaves with oblique lateral veins, and we can
+observe even now that the species with oblique stripes have
+longitudinal stripes when young, that is to say, while the stripes
+have no biological significance. The white places in the skin which
+gave rise, probably first as small spots, to this protective marking
+could be combined in one way or another according to the requirements
+of the species. They must therefore either have possessed
+selection-value from the first, or, if this was not the case at their
+earliest occurrence, there must have been _some other factors_ which
+raised them to the point of selection-value. I shall return to this in
+discussing germinal selection. But the case may be followed still
+farther, and leads us to the same alternative on a still more secure
+basis.
+
+Many years ago I observed in caterpillars of _Smerinthus populi_ (the
+poplar hawk-moth), which also possess white oblique stripes, that
+certain individuals showed _red spots_ above these stripes; these
+spots occurred only on certain segments, and never flowed together to
+form continuous stripes. In another species (_Smerinthus tiliae_)
+similar blood-red spots unite to form a line-like coloured seam in the
+last stage of larval life, while in _S. ocellata_ rust-red spots
+appear in individual caterpillars, but more rarely than in _S.
+populi_, and they show no tendency to flow together.
+
+Thus we have here the origin of a new character, arising from small
+beginnings, at least in _S. tiliae_, in which species the coloured
+stripes are a normal specific character. In the other species, _S.
+populi_ and _S. ocellata_, we find the beginnings of the same
+variation, in one more rarely than in the other, and we can imagine
+that, in the course of time, in these two species, coloured lines over
+the oblique stripes will arise. In any case these spots are the
+elements of variation, out of which coloured lines _may_ be evolved,
+if they are combined in this direction through the agency of natural
+selection. In _S. populi_ the spots are often small, but sometimes it
+seems as though several had united to form large spots. Whether a
+process of selection in this direction will arise in _S. populi_ and
+_S. ocellata_, or whether it is now going on cannot be determined,
+since we cannot tell in advance what biological value the marking
+might have for these two species. It is conceivable that the spots may
+have no selection-value as far as these species are concerned, and may
+therefore disappear again in the course of phylogeny, or, on the other
+hand, that they may be changed in another direction, for instance
+towards imitation of the rust-red fungoid patches on poplar and willow
+leaves. In any case we may regard the smallest spots as the initial
+stages of variation, the larger as a cumulative summation of these.
+Therefore either these initial stages must already possess
+selection-value, or, as I said before: _There must be some other
+reason for their cumulative summation_. I should like to give one more
+example, in which we can infer, though we cannot directly observe, the
+initial stages.
+
+All the Holothurians or sea-cucumbers have in the skin calcereous
+bodies of different forms, usually thick and irregular, which make the
+skin tough and resistant. In a small group of them--the species of
+Synapta--the calcareous bodies occur in the form of delicate anchors
+of microscopic size. Up till 1897 these anchors, like many other
+delicate microscopic structures, were regarded as curiosities, as
+natural marvels. But a Swedish observer, Oestergren, has recently
+shown that they have a biological significance: they serve the
+footless Synapta as auxiliary organs of locomotion, since, when the
+body swells up in the act of creeping, they press firmly with their
+tips, which are embedded in the skin, against the substratum on which
+the animal creeps, and thus prevent slipping backwards. In other
+Holothurians this slipping is made impossible by the fixing of the
+tube-feet. The anchors act automatically, sinking their tips towards
+the ground when the corresponding part of the body thickens, and
+returning to the original position at an angle of 45 degrees to the
+upper surface when the part becomes thin again. The arms of the anchor
+do not lie in the same plane as the shaft, and thus the curve of the
+arms forms the outermost part of the anchor, and offers no further
+resistance to the gliding of the animal. Every detail of the anchor,
+the curved portion, the little teeth at the head, the arms, etc., can
+be interpreted in the most beautiful way, above all the form of the
+anchor itself, for the two arms prevent it from swaying round to the
+side. The position of the anchors, too, is definite and significant;
+they lie obliquely to the longitudinal axis of the animal, and
+therefore they act alike whether the animal is creeping backwards or
+forwards. Moreover, the tips would pierce through the skin if the
+anchors lay in the longitudinal direction. Synapta burrows in the
+sand; it first pushes in the thin anterior end, and thickens this
+again, thus enlarging the hole, then the anterior tentacles displace
+more sand, the body is worked in a little farther, and the process
+begins anew. In the first act the anchors are passive, but they begin
+to take an active share in the forward movement when the body is
+contracted again. Frequently the animal retains only the posterior end
+buried in the sand, and then the anchors keep it in position, and make
+rapid withdrawal possible.
+
+Thus we have in these apparently random forms of the calcereous
+bodies, complex adaptations in which every little detail as to
+direction, curve, and pointing is exactly determined. That they have
+selection-value in their present perfected form is beyond all doubt,
+since the animals are enabled by means of them to bore rapidly into
+the ground and so to escape from enemies. We do not know what the
+initial stages were, but we cannot doubt that the little improvements,
+which occurred as variations of the originally simple slimy bodies of
+the Holothurians, were preserved because they already possessed
+selection-value for the Synaptidae. For such minute microscopic
+structures whose form is so delicately adapted to the role they have
+to play in the life of the animal, cannot have arisen suddenly and as
+a whole, and every new variation of the anchor, that is, in the
+direction of the development of the two arms, and every curving of the
+shaft which prevented the tips from projecting at the wrong time, in
+short, every little adaptation in the modelling of the anchor must
+have possessed selection-value. And that such minute changes of form
+fall within the sphere of fluctuating variations, that is to say,
+_that they occur_ is beyond all doubt.
+
+In many of the Synaptidae the anchors are replaced by calcareous rods
+bent in the form of an S, which are said to act in the same way.
+Others, such as those of the genus Ankyroderma, have anchors which
+project considerably beyond the skin, and, according to Oestergren,
+serve "to catch plant-particles and other substances" and so mask the
+animal. Thus we see that in the Synaptidae the thick and irregular
+calcareous bodies of the Holothurians have been modified and
+transformed in various ways in adaptation to the footlessness of these
+animals, and to the peculiar conditions of their life, and we must
+conclude that the earlier stages of these changes presented themselves
+to the processes of selection in the form of microscopic variations.
+For it is as impossible to think of any origin other than through
+selection in this case as in the case of the toughness, and the
+"drip-tips" of tropical leaves. And as these last could not have been
+produced directly by the beating of the heavy raindrops upon them, so
+the calcareous anchors of Synapta cannot have been produced directly
+by the friction of the sand and mud at the bottom of the sea, and,
+since they are parts whose function is _passive_ the Lamarckian factor
+of use and disuse does not come into question. The conclusion is
+unavoidable, that the microscopically small variations of the
+calcareous bodies in the ancestral forms have been intensified and
+accumulated in a particular direction, till they have led to the
+formation of the anchor. Whether this has taken place by the action of
+natural selection alone, or whether the laws of variation and the
+intimate processes within the germ-plasm have cooeperated will become
+clear in the discussion of germinal selection. This whole process of
+adaptation has obviously taken place within the time that has elapsed
+since this group of sea-cucumbers lost their tube-feet, those
+characteristic organs of locomotion which occur in no group except the
+Echinoderms, and yet have totally disappeared in the Synaptidae. And
+after all what would animals that live in sand and mud do with
+tube-feet?
+
+
+(_c_) _Coadaptation_
+
+Darwin pointed out that one of the essential differences between
+artificial and natural selection lies in the fact that the former can
+modify only a few characters, usually only one at a time, while Nature
+preserves in the struggle for existence all the variations of a
+species, at the same time and in a purely mechanical way, if they
+possess selection-value.
+
+Herbert Spencer, though himself an adherent of the theory of selection,
+declared in the beginning of the nineties that in his opinion the range of
+this principle was greatly over-estimated, if the great changes which have
+taken place in so many organisms in the course of ages are to be
+interpreted as due to this process of selection alone, since no
+transformation of any importance can be evolved by itself; it is always
+accompanied by a host of secondary changes. He gives the familiar example
+of the Giant Stag of the Irish peat, the enormous antlers of which required
+not only a much stronger skull cap, but also greater strength of the
+sinews, muscles, nerves and bones of the whole anterior half of the animal,
+if their mass was not to weigh down the animal altogether. It is
+inconceivable, he says, that so many processes of selection should take
+place _simultaneously_, and we are therefore forced to fall back on the
+Lamarckian factor of the use and disuse of functional parts. And how, he
+asks, could natural selection follow two opposite directions of evolution
+in different parts of the body at the same time, as for instance in the
+case of the kangaroo, in which the forelegs must have become shorter, while
+the hind legs and the tail were becoming longer and stronger?
+
+Spencer's main object was to substantiate the validity of the
+Lamarckian principle, the cooeperation of which with selection had been
+doubted by many. And it does seem as though this principle, if it
+operates in nature at all, offers a ready and simple explanation of
+all such secondary variations. Not only muscles, but nerves, bones,
+sinews, in short all tissues which function actively, increase in
+strength in proportion as they are used, and conversely they decrease
+when the claims on them diminish. All the parts, therefore, which
+depend on the part that varied first, as for instance the enlarged
+antlers of the Irish Elk, must have been increased or decreased in
+strength, in exact proportion to the claims made upon them,--just as
+is actually the case.
+
+But beautiful as this explanation would be, I regard it as untenable,
+because it assumes the _transmissibility of functional modifications_
+(so-called "acquired" characters), and this is not only
+undemonstrable, but is scarcely theoretically conceivable, for the
+secondary variations which accompany or follow the first as
+correlative variations, occur also in cases in which the animals
+concerned are sterile and _therefore cannot transmit anything to their
+descendants_. This is true of _worker bees_, and particularly of
+_ants_, and I shall here give a brief survey of the present state of
+the problem as it appears to me.
+
+Much has been written on both sides of this question since the
+published controversy on the subject in the nineties between Herbert
+Spencer and myself. I should like to return to the matter in detail,
+if the space at my disposal permitted, because it seems to me that the
+arguments I advanced at that time are equally cogent to-day,
+notwithstanding all the objections that have since been urged against
+them. Moreover, the matter is by no means one of subordinate interest;
+it is the very kernel of the whole question of the reality and value
+of the principle of selection. For if selection alone does not suffice
+to explain "_harmonious adaptation_" as I have called Spencer's
+_Coadaptation_, and if we require to call in the aid of the Lamarckian
+factor it would be questionable whether selection would explain any
+adaptations whatever. In this particular case--of worker bees--the
+Lamarckian factor may be excluded altogether, for it can be
+demonstrated that here at any rate the effects of use and disuse
+cannot be transmitted.
+
+But if it be asked why we are unwilling to admit the cooeperation of
+the Darwinian factor of selection and the Lamarckian factor, since
+this would afford us an easy and satisfactory explanation of the
+phenomena, I answer: _Because the Lamarckian principle is fallacious,
+and because by accepting it we close the way towards deeper insight_.
+It is not a spirit of combativeness or a desire for self-vindication
+that induces me to take the field once more against the Lamarckian
+principle, it is the conviction that the progress of our knowledge is
+being obstructed by the acceptance of this fallacious principle, since
+the facile explanation it apparently affords prevents our seeking
+after a truer explanation and a deeper analysis.
+
+The workers in the various species of ants are sterile, that is to
+say, they take no regular part in the reproduction of the species,
+although individuals among them may occasionally lay eggs. In addition
+to this they have lost the wings, and the _receptaculum seminis_, and
+their compound eyes have degenerated to a few facets. How could this
+last change have come about through disuse, since the eyes of workers
+are exposed to light in the same way as are those of the sexual
+insects and thus in this particular case are not liable to "disuse" at
+all? The same is true of the _receptaculum seminis_, which can only
+have been disused as far as its glandular portion and its stalk are
+concerned, and also of the wings, the nerves tracheae and epidermal
+cells of which could not cease to function until the whole wing had
+degenerated, for the chitinous skeleton of the wing does not function
+at all in the active sense.
+
+But, on the other hand, the workers in all species have undergone
+modifications in a positive direction, as, for instance, the greater
+development of brain. In many species large workers have evolved,--the
+so-called _soldiers_, with enormous jaws and teeth, which defend the
+colony,--and in others there are _small_ workers which have taken over
+other special functions, such as the rearing of the young Aphides.
+This kind of division of the workers into two castes occurs among
+several tropical species of ants, but it is also present in the
+Italian species, _Colobopsis truncata_. Beautifully as the size of the
+jaws could be explained as due to the increased use made of them by
+the "soldiers," or the enlarged brain as due to the mental activities
+of the workers, the fact of the infertility of these forms is an
+insurmountable obstacle to accepting such an explanation. Neither jaws
+nor brain can have been evolved on the Lamarckian principle.
+
+The problem of coadaptation is no easier in the case of the ant than
+in the case of the Giant Stag. Darwin himself gave a pretty
+illustration to show how imposing the difference between the two kinds
+of workers in one species would seem if we translated it into human
+terms. In regard to the Driver ants (Anomma) we must picture to
+ourselves a piece of work, "for instance the building of a house,
+being carried on by two kinds of workers, of which one group was five
+feet four inches high, the other sixteen feet high."[39]
+
+Although the ant is a small animal as compared with man or with the
+Irish Elk, the "soldier" with its relatively enormous jaws is hardly
+less heavily burdened than the Elk with its antlers, and in the ant's
+case, too, a strengthening of the skeleton, of the muscles, the nerves
+of the head, and of the legs must have taken place parallel with the
+enlargement of the jaws. _Harmonious adaptation_ (coadaptation) has
+here been active in a high degree, and yet these "soldiers" are
+sterile! There thus remains nothing for it but to refer all their
+adaptations, positive and negative alike, to processes of selection
+which have taken place in the rudiments of the workers within the egg
+and sperm-cells of their parents. There is no way out of the
+difficulty except the one Darwin pointed out. He himself did not find
+the solution of the riddle at once. At first he believed that the case
+of the workers among social insects presented "the most serious
+special difficulty" in the way of his theory of natural selection; and
+it was only after it had become clear to him that it was not the
+sterile insects themselves but their parents that were selected,
+according as they produced more or less well adapted workers, that he
+was able to refer to this very case of the conditions among ants "_in
+order to show the power of natural selection_."[40] He explains his
+view by a simple but interesting illustration. Gardeners have
+produced, by means of long continued artificial selection, a variety
+of Stock, which bears entirely double, and therefore infertile
+flowers.[41] Nevertheless the variety continues to be reproduced from
+seed, because, in addition to the double and infertile flowers, the
+seeds always produce a certain number of single, fertile blossoms, and
+these are used to reproduce the double variety. These single and
+fertile plants correspond "to the males and females of an ant-colony,
+the infertile plants, which are regularly produced in large numbers,
+to the neuter workers of the colony."
+
+This illustration is entirely apt, the only difference between the
+two cases consisting in the fact that the variation in the flower is
+not a useful, but a disadvantageous one, which can only be preserved
+by artificial selection on the part of the gardener, while the
+transformations that have taken place parallel with the sterility of
+the ants are useful, since they procure for the colony an advantage in
+the struggle for existence, and they are therefore preserved by
+natural selection. Even the sterility itself in this case is not
+disadvantageous, since the fertility of the true females has at the
+same time considerably increased. We may therefore regard the sterile
+forms of ants, which have gradually been adapted in several directions
+to varying functions, _as a certain proof_ that selection really takes
+place in the germ-cells of the fathers and mothers of the workers, and
+that _special complexes of primordia_ (_ids_) are present in the
+workers and in the males and females, and these complexes contain the
+primordia of the individual parts (_determinants_). But since all
+living entities vary, the determinants must also vary, now in a
+favourable, now in an unfavourable direction. If a female produces
+eggs, which contain favourably varying determinants in the worker-ids,
+then these eggs will give rise to workers modified in the favourable
+direction, and if this happens with many females, the colony concerned
+will contain a better kind of worker than other colonies.
+
+I digress here in order to give an account of the intimate processes,
+which, according to my view, take place within the germ-plasm, and
+which I have called "_germinal selection_." These processes are of
+importance since they form the roots of variation, which in its turn
+is the root of natural selection. I cannot here do more than give a
+brief outline of the theory in order to show how the Darwin-Wallace
+theory of selection has gained support from it.
+
+With others, I regard the minimal amount of substance which is
+contained within the nucleus of the germ-cells, in the form of rods,
+bands, or granules, as the _germ-substance_ or _germ-plasm_, and I
+call the individual granules _ids_. There is always a multiplicity of
+such ids present in the nucleus, either occurring individually, or
+united in the form of rods or bands (chromosomes). Each id contains
+the primary constituents of a _whole_ individual, so that several ids
+are concerned in the development of a new individual.
+
+In every being of complex structure thousands of primary constituents
+must go to make up a single id; these I call _determinants_, and I
+mean by this name very small individual particles, far below the
+limits of microscopic visibility, vital units which feed, grow, and
+multiply by division. These determinants control the parts of the
+developing embryo,--in what manner need not here concern us. The
+determinants differ among themselves, those of a muscle are
+differently constituted from those of a nerve-cell or a glandular
+cell, etc., and every determinant is in its turn made up of minute
+vital units, which I call _biophores_, or the bearers of life.
+According to my view, these determinants not only assimilate, like
+every other living unit, but they _vary_ in the course of their
+growth, as every living unit does; they may vary qualitatively if the
+elements of which they are composed vary, they may grow and divide
+more or less rapidly, and their variations give rise to
+_corresponding_ variations of the organ, cell, or cell-group which
+they determine. That they are undergoing ceaseless fluctuations in
+regard to size and quality seems to me the inevitable consequence of
+their unequal nutrition; for although the germ-cell as a whole usually
+receives sufficient nutriment, minute fluctuations in the amount
+carried to different parts within the germ-plasm cannot fail to occur.
+
+Now, if a determinant, for instance of a sensory cell, receives for a
+considerable time more abundant nutriment than before, it will grow
+more rapidly--become bigger, and divide more quickly, and, later, when
+the id concerned develops into an embryo, this sensory cell will
+become stronger than in the parents, possibly even twice as strong.
+This is an instance of a _hereditary individual variation_, arising
+from the germ.
+
+The nutritive stream which, according to our hypothesis, favours the
+determinant _N_ by chance, that is, for reasons unknown to us, may
+remain strong for a considerable time, or may decrease again; but even
+in the latter case it is conceivable that the ascending movement of
+the determinant may continue, because the strengthened determinant now
+_actively_ nourishes itself more abundantly,--that is to say, it
+attracts the nutriment to itself, and to a certain extent withdraws it
+from its fellow-determinants. In this way, it may--as it seems to
+me--get into _permanent upward movement, and attain a degree of
+strength from which there is no falling back_. Then positive or
+negative selection sets in, favouring the variations which are
+advantageous, setting aside those which are disadvantageous.
+
+In a similar manner a _downward_ variation of the determinants may
+take place, if its progress be started by a diminished flow of
+nutriment. The determinants which are weakened by this diminished flow
+will have less affinity for attracting nutriment because of their
+diminished strength, and they will assimilate more feebly and grow
+more slowly, unless chance streams of nutriment help them to recover
+themselves. But, as will presently be shown, a change of direction
+cannot take place at _every_ stage of the degenerative process. If a
+certain critical stage of downward progress be passed, even favourable
+conditions of food-supply will no longer suffice permanently to change
+the direction of the variation. Only two cases are conceivable; if the
+determinant corresponds to a _useful_ organ, only its removal can
+bring back the germ-plasm to its former level; therefore personal
+selection removes the id in question, with its determinants, from the
+germ-plasm, by causing the elimination of the individual in the
+struggle for existence. But there is another conceivable case; the
+determinants concerned may be those of an organ which has become
+_useless_, and they will then continue unobstructed, but with
+exceeding slowness, along the downward path, until the organ becomes
+vestigial, and finally disappears altogether.
+
+The fluctuations of the determinants hither and thither may thus be
+transformed into a lasting ascending or descending movement; and _this
+is the crucial point of these germinal processes_.
+
+This is not a fantastic assumption; we can read it in the fact of the
+degeneration of disused parts. _Useless organs are the only ones which
+are not helped to ascend again by personal selection, and therefore in
+their case alone can we form any idea of how the primary constituents
+behave, when they are subject solely to intra-germinal forces_.
+
+The whole determinant system of an id, as I conceive it, is in a state
+of continual fluctuation upwards and downwards. In most cases the
+fluctuations will counteract one another, because the passive streams
+of nutriment soon change, but in many cases the limit from which a
+return is possible will be passed, and then the determinants concerned
+will continue to vary in the same direction, till they attain positive
+or negative selection-value. At this stage personal selection
+intervenes and sets aside the variation if it is disadvantageous, or
+favours--that is to say, preserves--it if it is advantageous. Only
+_the determinant of a useless organ is uninfluenced by personal
+selection_, and, as experience shows, it sinks downwards; that is, the
+organ that corresponds to it degenerates very slowly but
+uninterruptedly till, after what must obviously be an immense stretch
+of time, it disappears from the germ-plasm altogether.
+
+Thus we find in the fact of the degeneration of disused parts the
+proof that not all the fluctuations of a determinant return to
+equilibrium again, but that, when the movement has attained to a
+certain strength, it continues _in the same direction_. We have entire
+certainty in regard to this as far as the downward progress is
+concerned, and we must assume it also in regard to ascending
+variations, as the phenomena of artificial selection certainly justify
+us in doing. If the Japanese breeders were able to lengthen the
+tail-feathers of the cock to six feet, it can only have been because
+the determinants of the tail-feathers in the germ-plasm had already
+struck out a path of ascending variation, and this movement was taken
+advantage of by the breeder, who continually selected for reproduction
+the individuals in which the ascending variation was most marked. For
+all breeding depends upon the unconscious selection of germinal
+variations.
+
+Of course these germinal processes cannot be proved mathematically,
+since we cannot actually see the play of forces of the passive
+fluctuations and their causes. We cannot say how great these
+fluctuations are, and how quickly or slowly, how regularly or
+irregularly they change. Nor do we know how far a determinant must be
+strengthened by the passive flow of the nutritive stream if it is to
+be beyond the danger of unfavourable variations, or how far it must be
+weakened passively before it loses the power of recovering itself by
+its own strength. It is no more possible to bring forward actual
+proofs in this case than it was in regard to the selection-value of
+the initial stages of an adaptation. But if we consider that all
+heritable variations must have their roots in the germ-plasm, and
+further, that when personal selection does not intervene, that is to
+say, in the case of parts which have become useless, a degeneration of
+the part, and therefore also of its determinant must inevitably take
+place; then we must conclude that processes such as I have assumed are
+running their course within the germ-plasm, and we can do this with as
+much certainty as we were able to infer, from the phenomena of
+adaptation, the selection-value of their initial stages. The fact of
+the degeneration of disused parts seems to me to afford irrefutable
+proof that the fluctuations within the germ-plasm _are the real root
+of all hereditary variation_, and the preliminary condition for the
+occurrence of the Darwin-Wallace factor of selection. Germinal
+selection supplies the stones out of which personal selection builds
+her temples and palaces: _adaptations_. The importance for the theory
+of the process of degeneration of disused parts cannot be
+over-estimated, especially when it occurs in sterile animal forms,
+where we are free from the doubt as to the alleged _Lamarckian factor_
+which is apt to confuse our ideas in regard to other cases.
+
+If we regard the variation of the many determinants concerned in the
+transformation of the female into the sterile worker as having come
+about through the gradual transformation of the ids into worker-ids,
+we shall see that the germ-plasm of the sexual ants must contain three
+kinds of ids, male, female, and worker ids, or if the workers have
+diverged into soldiers and nest-builders, then four kinds. We
+understand that the worker-ids arose because their determinants struck
+out a useful path of variation, whether upward or downward, and that
+they continued in this path until the highest attainable degree of
+utility of the parts determined was reached. But in addition to the
+organs of positive or negative selection-value, there were some which
+were indifferent as far as the success and especially the functional
+capacity of the workers was concerned: wings, ovarian tubes,
+_receptaculum seminis_, a number of the facets of the eye, perhaps
+even the whole eye. As to the ovarian tubes it is is possible that
+their degeneration was an advantage for the workers, in saving energy,
+and if so selection would favour the degeneration; but how could the
+presence of eyes diminish the usefulness of the workers to the colony?
+or the minute _receptaculum seminis_, or even the wings? These parts
+have therefore degenerated _because they were of no further value to
+the insect_. But if selection did not influence the setting aside of
+these parts because they were neither of advantage nor of disadvantage
+to the species, then the Darwinian factor of selection is here
+confronted with a puzzle which it cannot solve alone, but which at
+once becomes clear when germinal selection is added. For the
+determinants of organs that have no further value for the organism,
+must, as we have already explained, embark on a gradual course of
+retrograde development.
+
+In ants the degeneration has gone so far that there are no
+wing-rudiments present in _any_ species, as is the case with so many
+butterflies, flies, and locusts, but in the larvae the imaginable
+discs of the wings are still laid down. With regard to the ovaries,
+degeneration has reached different levels in different species of
+ants, as has been shown by the researches of my former pupil,
+Elizabeth Bickford. In many species there are twelve ovarian tubes,
+and they decrease from that number to one; indeed, in one species no
+ovarian tube at all is present. So much at least is certain from what
+has been said, that in this case _everything_ depends on the
+fluctuations of the elements of the germ-plasm. Germinal selection,
+here as elsewhere, presents the variations of the determinants, and
+personal selection favours or rejects these, or,--if it be a question
+of organs which have become useless,--it does not come into play at
+all, and allows the descending variation free course.
+
+It is obvious that even the problem of _coadaptation in sterile
+animals_ can thus be satisfactorily explained. If the determinants are
+oscillating upwards and downwards in continual fluctuation, and
+varying more pronouncedly now in one direction now in the other,
+useful variations of every determinant will continually present
+themselves anew, and may, in the course of generations, be combined
+with one another in various ways. But there is one character of the
+determinants that greatly facilitates this complex process of
+selection, that, after a certain limit has been reached, they go on
+varying in the same direction. From this it follows that development
+along a path once struck out may proceed without the continual
+intervention of personal selection. This factor only operates, so to
+speak, at the beginning, when it selects the determinants which are
+varying in the right direction, and again at the end, when it is
+necessary to put a check upon further variation. In addition to this,
+enormously long periods have been available for all these adaptations,
+as the very gradual transition stages between females and workers in
+many species plainly show, and thus this process of transformation
+loses the marvellous and mysterious character that seemed at the first
+glance to invest it, and takes rank, without any straining, among the
+other processes of selection. It seems to me that, from the facts that
+sterile animal forms can adapt themselves to new vital functions,
+their superfluous parts degenerate, and the parts more used adapt
+themselves in an ascending direction, those less used in a descending
+direction, we must draw the conclusion that harmonious adaptation here
+comes about _without the cooeperation of the Lamarckian principle_.
+This conclusion once established, however, we have no reason to refer
+the thousands of cases of harmonious adaptation, which occur in
+exactly the same way among other animals or plants, to a principle,
+the _active intervention of which in the transformation of species is
+nowhere proved. We do not require it to explain the facts, and
+therefore we must not assume it._
+
+The fact of coadaptation, which was supposed to furnish the strongest
+argument against the principle of selection, in reality yields the
+clearest evidence in favour of it. We _must_ assume it, _because no
+other possibility of explanation is open to us, and because these
+adaptations actually exist, that is to say, have really taken place_.
+With this conviction I attempted, as far back as 1894, when the idea
+of germinal selection had not yet occurred to me, to make "harmonious
+adaptation" (coadaptation) more easily intelligible in some way or
+other, and so I was led to the idea, which was subsequently expounded
+in detail by Baldwin, and Lloyd Morgan, and also by Osborn, and Gulick
+as _Organic Selection_. It seemed to me that it was not necessary that
+all the germinal variations required for secondary variations should
+have occurred _simultaneously_, since, for instance, in the case of
+the stag, the bones, muscles, sinews, and nerves would be incited by
+the increasing heaviness of the antlers to greater activity in _the
+individual life_, and so would be strengthened. The antlers can only
+have increased in size by very slow degrees, so that the muscles and
+bones may have been able to keep pace with their growth in the
+individual life, until the requisite germinal variations presented
+themselves. In this way a disharmony between the increasing weight of
+the antlers and the parts which support and move them would be
+avoided, since time would be given for the appropriate germinal
+variations to occur, and so to set agoing the _hereditary_ variation
+of the muscles, sinews and bones.[42]
+
+I still regard this idea as correct, but I attribute less importance
+to "organic selection" than I did at that time, in so far that I do
+not believe that it _alone_ could effect complex harmonious
+adaptations. Germinal selection now seems to me to play the chief part
+in bringing about such adaptations. Something the same is true of the
+principle I have called _Panmixia_. As I became more and more
+convinced, in the course of years, that the _Lamarckian principle_
+ought not to be called in to explain the dwindling of disused parts, I
+believed that this process might be simply explained as due to the
+cessation of the conservative effect of natural selection. I said to
+myself that, from the moment in which a part ceases to be of use,
+natural selection withdraws its hand from it, and then it must
+inevitably fall from the height of its adaptiveness, because inferior
+variants would have as good a chance of persisting as better ones,
+since all grades of fitness of the part in question would be mingled
+with one another indiscriminately. This is undoubtedly true, as
+Romanes pointed out ten years before I did, and this mingling of the
+bad with the good probably does bring about a deterioration of the
+part concerned. But it cannot account for the steady diminution, which
+always occurs when a part is in process of becoming rudimentary, and
+which goes on until it ultimately disappears altogether. The process
+of dwindling cannot therefore be explained as due to panmixia alone:
+we can only find a sufficient explanation in germinal selection.
+
+
+IV. DERIVATIVES OF THE THEORY OF SELECTION
+
+The impetus in all directions given by Darwin through his theory of
+selection has been an immeasurable one, and its influence is still
+felt. It falls within the province of the historian of science to
+enumerate all the ideas which, in the last quarter of the nineteenth
+century, grew out of Darwin's theories, in the endeavour to penetrate
+more deeply into the problem of the evolution of the organic world.
+Within the narrow limits to which this paper is restricted, I cannot
+attempt to discuss any of these.
+
+
+V. ARGUMENTS FOR THE REALITY OF THE PROCESSES OF SELECTION
+
+
+(_a_) _Sexual Selection_
+
+Sexual selection goes hand in hand with natural selection. From the
+very first I have regarded sexual selection as affording an extremely
+important and interesting corroboration of natural selection, but,
+singularly enough, it is precisely against this theory that an adverse
+judgment has been pronounced in so many quarters, and it is only quite
+recently, and probably in proportion as the wealth of facts in proof
+of it penetrates into a wider circle, that we seem to be approaching a
+more general recognition of this side of the problem of adaptations.
+Thus Darwin's words in his preface to the second edition (1874) of his
+book, _The Descent of Man and Sexual Selection_, are being justified:
+"My conviction as to the operation of natural selection remains
+unshaken," and further, "If naturalists were to become more familiar
+with the idea of sexual selection, it would, I think, be accepted to a
+much greater extent, and already it is fully and favourably accepted
+by many competent judges." Darwin was able to speak thus because he
+was already acquainted with an immense mass of facts, which, taken
+together, yield overwhelming evidence of the validity of the principle
+of sexual selection.
+
+_Natural selection_ chooses out for reproduction the individuals that
+are best equipped for the struggle for existence, and it does so at
+every stage of development; it thus improves the species in all its
+stages and forms. _Sexual selection_ operates only on individuals
+that are already capable of reproduction, and does so only in relation
+to the attainment of reproduction. It arises from the rivalry of one
+sex, usually the male, for the possession of the other, usually the
+female. Its influence can therefore only _directly_ affect one sex, in
+that it equips it better for attaining possession of the other. But
+the effect may extend indirectly to the female sex, and thus the whole
+species may be modified, without, however, becoming any more capable
+of resistance in the struggle for existence, for sexual selection only
+gives rise to adaptations which are likely to give their possessor the
+victory over rivals in the struggle for possession of the female, and
+which are therefore peculiar to the wooing sex: the manifold
+"secondary sexual characters." The diversity of these characters is so
+great that I cannot here attempt to give anything approaching a
+complete treatment of them, but I should like to give a sufficient
+number of examples to make the principle itself, in its various modes
+of expression, quite clear.
+
+One of the chief preliminary postulates of sexual selection is the
+unequal number of individuals in the two sexes, for if every male
+immediately finds his mate there can be no competition for the
+possession of the female. Darwin has shown that, for the most part,
+the inequality between the sexes is due simply to the fact that there
+are more males than females, and therefore the males must take some
+pains to secure a mate. But the inequality does not always depend on
+the numerical preponderance of the males, it is often due to polygamy;
+for, if one male claims several females, the number of females in
+proportion to the rest of the males will be reduced. Since it is
+almost always the males that are the wooers, we must expect to find
+the occurrence of secondary sexual characters chiefly among them, and
+to find it especially frequent in polygamous species. And this is
+actually the case.
+
+If we were to try to guess--without knowing the facts--what means the
+male animals make use of to overcome their rivals in the struggle for
+the possession of the female, we might name many kinds of means, but
+it would be difficult to suggest any which is not actually employed in
+some animal group of other. I begin with the mere difference in
+strength, through which the male of many animals is so sharply
+distinguished from the female, as, for instance, the lion, walrus,
+"sea-elephant," and others. Among these the males fight violently for
+the possession of the female, who falls to the victor in the combat.
+In this simple case no one can doubt the operation of selection, and
+there is just as little room for doubt as to the selection-value of
+the initial stages of the variation. Differences in bodily strength
+are apparent even among human beings, although in their case the
+struggle for the possession of the female is no longer decided by
+bodily strength alone.
+
+Combats between male animals are often violent and obstinate, and the
+employment of the natural weapons of the species in this way has led
+to perfecting of these, e.g. the tusks of the boar, the antlers of the
+stag, and the enormous, antler-like jaws of the stag-beetle. Here
+again it is impossible to doubt that variations in these organs
+presented themselves, and that these were considerable enough to be
+decisive in combat, and so to lead to the improvement of the weapon.
+
+Among many animals, however, the females at first withdraw from the
+males; they are coy, and have to be sought out, and sometimes held by
+force. This tracking and grasping of the females by the males has
+given rise to many different characters in the latter, as, for
+instance, the larger eyes of the male bee, and especially of the males
+of the Ephemerids (May-flies), some species of which show, in addition
+to the usual compound eyes, large, so-called turban-eyes, so that the
+whole head is covered with seeing surfaces. In these species the
+females are very greatly in the minority (1-100), and it is easy to
+understand that a keen competition for them must take place, and that,
+when the insects of both sexes are floating freely in the air, an
+unusually wide range of vision will carry with it a decided
+advantage. Here again the actual adaptations are in accordance with
+the preliminary postulates of the theory. We do not know the stages
+through which the eye has passed to its present perfected state, but,
+since the number of simple eyes (facets) has become very much greater
+in the male than in the female, we may assume that their increase is
+due to a gradual duplication of the determinants of the ommatidium in
+the germ-plasm, as I have already indicated in regard to sense-organs
+in general. In this case, again, the selection-value of the initial
+stages hardly admits of doubt; better vision _directly_ secures
+reproduction.
+
+In many cases _the organ of smell_ shows a similar improvement. Many
+lower Crustaceans (Daphnidae) have better developed organs of smell in
+the male sex. The difference is often slight and amounts only to one
+or two olfactory filaments, but certain species show a difference of
+nearly a hundred of these filaments (Leptodora). The same thing occurs
+among insects.
+
+We must briefly consider the clasping or grasping organs which have
+developed in the males among many lower Crustaceans, but here natural
+selection plays its part along with sexual selection, for the union of
+the sexes is an indispensable condition for the maintenance of the
+species, and as Darwin himself pointed out, in many cases the two
+forms of selection merge into each other. This fact has always seemed
+to me to be a proof of natural selection, for, in regard to sexual
+selection, it is quite obvious that the victory of the best-equipped
+could have brought about the improvement only of the organs concerned,
+the factors in the struggle, such as the eye and the olfactory organ.
+
+We come now to the _excitants_; that is, to the group of sexual
+characters whose origin through processes of selection has been most
+frequently called in question. We may cite the _love-calls_ produced
+by many male insects, such as crickets and cicadas. These could only
+have arisen in animal groups in which the female did not rapidly flee
+from the male, but was inclined to accept his wooing from the first.
+Thus, notes like the chirping of the male cricket serve to entice the
+females. At first they were merely the signal which showed the
+presence of a male in the neighbourhood, and the female was gradually
+enticed nearer and nearer by the continued chirping. The male that
+could make himself heard to the greatest distance would obtain the
+largest following, and would transmit the beginnings, and, later, the
+improvement of his voice to the greatest number of descendants. But
+sexual excitement in the female became associated with the hearing of
+the love-call, and then the sound-producing organ of the male began to
+improve, until it attained to the emission of the long-drawn-out soft
+notes of the mole-cricket or the maenad-like cry of the cicadas. I
+cannot here follow the process of development in detail, but will call
+attention to the fact that the original purpose of the voice, the
+announcing of the male's presence, became subsidiary, and the exciting
+of the female became the chief goal to be aimed at. The loudest
+singers awakened the strongest excitement, and the improvement
+resulted as a matter of course. I conceive of the origin of bird-song
+in a somewhat similar manner, first as a means of enticing, then of
+exciting the female.
+
+One more kind of secondary sexual character must here be mentioned:
+the odour which emanates from so many animals at the breeding season.
+It is possible that this odour also served at first merely to give
+notice of the presence of individuals of the other sex, but it soon
+became an excitant, and as the individuals which caused the greatest
+degree of excitement were preferred, it reached as high a pitch of
+perfection as was possible to it. I shall confine myself here to the
+comparatively recently discovered fragrance of butterflies. Since
+Fritz Mueller found out that certain Brazilian butterflies gave off
+fragrance "like a flower," we have become acquainted with many such
+cases, and we now know that in all lands, not only many diurnal
+Lepidoptera but nocturnal ones also give off a delicate odour, which
+is agreeable even to man. The ethereal oil to which this fragrance is
+due is secreted by the skin-cells, usually of the wing, as I showed
+soon after the discovery of the _scent-scales_. This is the case in
+the males; the females have no _special_ scent-scales recognisable as
+such by their form, but they must, nevertheless, give off an extremely
+delicate fragrance, although our imperfect organ of smell cannot
+perceive it, for the males become aware of the presence of a female,
+even at night, from a long distance off, and gather round her. We may
+therefore conclude, that both sexes have long given forth a very
+delicate perfume, which announced their presence to others of the same
+species, and that in many species (_not in all_) these small
+beginnings become, in the males, particularly strong scent-scales of
+characteristic form (lute, brush, or lyre-shaped). At first these
+scales were scattered over the surface of the wing, but gradually they
+concentrated themselves, and formed broad, velvety bands, or strong,
+prominent brushes, and they attained their highest pitch of evolution
+when they became enclosed within pits or folds of the skin, which
+could be opened to let the delicious fragrance stream forth suddenly
+towards the female. Thus in this case also we see that characters, the
+original use of which was to bring the sexes together, and so to
+maintain the species, have been evolved in the males into means for
+exciting the female. And we can hardly doubt, that the females are
+most readily enticed to yield to the butterfly that sends out the
+strongest fragrance,--that is to say, that excites them to the highest
+degree. It is a pity that our organs of smell are not fine enough to
+examine the fragrance of male Lepidoptera in general, and to compare
+it with other perfumes which attract these insects.[43] As far as we
+can perceive them they resemble the fragrance of flowers, but there
+are Lepidoptera whose scent suggests musk. A smell of musk is also
+given off by several plants: it is a sexual excitant in the
+musk-deer, the musk-sheep, and the crocodile.
+
+As far as we know, then, it is perfumes similar to those of flowers
+that the male Lepidoptera give off in order to entice their mates and
+this is a further indication that animals, like plants, can to a large
+extent meet the claims made upon them by life, and produce the
+adaptations which are most purposive,--a further proof, too, of my
+proposition that the useful variations, so to speak, are _always
+there_. The flowers developed the perfumes which entice their
+visitors, and the male Lepidoptera developed the perfumes which entice
+and excite their mates.
+
+There are many pretty little problems to be solved in this connection,
+for there are insects, such as some flies, that are attracted by
+smells which are unpleasant to us, like those from decaying flesh and
+carrion. But there are also certain flowers, some orchids for
+instance, which give forth no very agreeable odour, but one which is
+to us repulsive and disgusting; and we should therefore expect that
+the males of such insects would give off a smell unpleasant to us, but
+there is no case known to me in which this has been demonstrated.
+
+In cases such as we have discussed, it is obvious that there is no
+possible explanation except through selection. This brings us to the
+last kind of secondary sexual characters, and the one in regard to
+which doubt has been most frequently expressed,--decorative colours
+and decorative forms, the brilliant plumage of the male pheasant, the
+humming-birds, and the bird of Paradise, as well as the bright colours
+of many species of butterfly, from the beautiful blue of our little
+Lycaenidae to the magnificent azure of the large Morphinae of Brazil.
+In a great many cases, though not by any means in all, the male
+butterflies are "more beautiful" than the females, and in the Tropics
+in particular they shine and glow in the most superb colours. I really
+see no reason why we should doubt the power of sexual selection, and I
+myself stand wholly on Darwin's side. Even though we certainly cannot
+assume that the females exercise a conscious choice of the
+"handsomest" mate, and deliberate like the judges in a court of
+justice over the perfections of their wooers, we have no reason to
+doubt that distinctive forms (decorative feathers), and colours have a
+particularly exciting effect upon the female, just as certain odours
+have among animals of so many different groups, including the
+butterflies. The doubts which existed for a considerable time, as a
+result of fallacious experiments, as to whether the colours of flowers
+really had any influence in attracting butterflies have now been set
+at rest through a series of more careful investigations; we now know
+that the colours of flowers are there on account of the butterflies,
+as Sprengel first showed, and that the blossoms of Phanerogams are
+selected in relation to them, as Darwin pointed out.
+
+Certainly it is not possible to bring forward any convincing proof of
+the origin of decorative colours through sexual selection, but there
+are many weighty arguments in favour of it, and these form a body of
+presumptive evidence so strong that it almost amounts to certainty.
+
+In the first place, there is the analogy with other secondary sexual
+characters. If the song of birds and the chirping of the cricket have
+been evolved through sexual selection, if the penetrating odours of
+male animals,--the crocodile, the musk-deer, the beaver, the
+carnivores, and, finally, the flower-like fragrances of the
+butterflies have been evolved to their present pitch in this way, why
+should decorative colours have arisen in some other way? Why should
+the eye be less sensitive to _specifically male_ colours and other
+_visible_ signs _enticing to the female_, than the olfactory sense to
+specifically male odours, or the sense of hearing to specifically male
+sounds? Moreover, the decorative feathers of birds are almost always
+spread out and displayed before the female during courtship. I have
+elsewhere[44] pointed out that decorative colouring and
+sweet-scentedness may replace one another in Lepidoptera as well as in
+flowers, for just as some modestly coloured flowers (mignonette and
+violet) have often a strong perfume, while strikingly coloured ones
+are sometimes quite devoid of fragrance, so we find that the most
+beautiful and gaily-coloured of our native Lepidoptera, the species of
+Vanessa, have no scent-scales, while these are often markedly
+developed in grey nocturnal Lepidoptera. Both attractions may,
+however, be combined in butterflies, just as in flowers. Of course, we
+cannot explain why both means of attraction should exist in one genus,
+and only one of them in another, since we do not know the minutest
+details of the conditions of life of the genera concerned. But from
+the sporadic distribution of scent-scales in Lepidoptera, and from
+their occurrence or absence in nearly related species, we may conclude
+that fragrance is a relatively _modern_ acquirement, more recent than
+brilliant colouring.
+
+One thing in particular that stamps decorative colouring as a product
+of selection is _its gradual intensification_ by the addition of new
+spots, which we can quite well observe, because in many cases the
+colours have been first acquired by the males, and later transmitted
+to the females by inheritance. The scent-scales are never thus
+transmitted, probably for the same reason that the decorative colours
+of many birds are often not transmitted to the females: because with
+these they would be exposed to too great elimination by enemies.
+Wallace was the first to point out that in species with concealed
+nests the beautiful feathers of the male occurred in the female also,
+as in the parrots, for instance, but this is not the case in species
+which brood on an exposed nest. In the parrots one can often observe
+that the general brilliant colouring of the male is found in the
+female, but that certain spots of colour are absent, and these have
+probably been acquired comparatively recently by the male and have not
+yet been transmitted to the female.
+
+Isolation of the group of individuals which is in process of varying
+is undoubtedly of great value in sexual selection, for even a solitary
+conspicuous variation will become dominant much sooner in a small
+isolated colony, than among a large number of members of a species.
+
+Any one who agrees with me in deriving variations from germinal
+selection will regard that process as an essential aid towards
+explaining the selection of distinctive courtship-characters, such as
+coloured spots, decorative feathers, horny outgrowths in birds and
+reptiles, combs, feather-tufts, and the like, since the beginnings of
+these would be presented with relative frequency in the struggle
+between the determinants within the germ-plasm. The process of
+transmission of decorative feathers to the female results, as Darwin
+pointed out and illustrated by interesting examples, in the
+_colour-transformation of a whole species_, and this process, as the
+phyletically older colouring of young birds shows, must, in the course
+of thousands of years, have repeated itself several times in a line of
+descent.
+
+If we survey the wealth of phenomena presented to us by secondary
+sexual characters, we can hardly fail to be convinced of the truth of
+the principle of sexual selection. And certainly no one who has
+accepted natural selection should reject sexual selection, for, not
+only do the two processes rest upon the same basis, but they merge
+into one another, so that it is often impossible to say how much of a
+particular character depends on one and how much on the other form of
+selection.
+
+
+(_b_) _Natural Selection_
+
+An actual proof of the theory of sexual selection is out of the
+question, if only because we cannot tell when a variation attains to
+selection-value. It is certain that a delicate sense of smell is of
+value to the male moth in his search for the female, but whether the
+possession of one additional olfactory hair, or of ten, or of twenty
+additional hairs leads to the success of its possessor we are unable
+to tell. And we are groping even more in the dark when we discuss the
+excitement caused in the female by agreeable perfumes, or by striking
+and beautiful colours. That these do make an impression is beyond
+doubt; but we can only assume that slight intensifications of them
+give any advantage, and we _must_ assume this _since otherwise
+secondary sexual characters remain inexplicable_.
+
+The same thing is true in regard to natural selection. It is not
+possible to bring forward any actual proof of the selection-value of
+the initial stages, and the stages in the increase of variations, as
+has been already shown. But the selection-value of a finished
+adaptation can in many cases be statistically determined. Cesnola and
+Poulton have made valuable experiments in this direction. The former
+attached forty-five individuals of the green, and sixty-five of the
+brown variety of the praying mantis (_Mantis religiosa_), by a silk
+thread to plants, and watched them for seven days. The insects which
+were on a surface of a colour Similar to their own remained uneaten,
+while twenty-five green insects on brown parts of plants had all
+disappeared in eleven days.
+
+The experiments of Poulton and Sanders[45] were made with 600 pupae of
+_Vanessa urticae_, the "tortoise-shell butterfly." The pupae were
+artificially attached to nettles, tree-trunks, fences, walls, and to
+the ground, some at Oxford, some at St. Helens in the Isle of Wight.
+In the course of a month 93% of the pupae at Oxford were killed,
+chiefly by small birds, while at St. Helens 68% perished. The
+experiments showed very clearly that the colour and character of the
+surface on which the pupa rests--and thus its own conspicuousness--are
+of the greatest importance. At Oxford only the four pupae which were
+fastened to nettles emerged; all the rest--on bark, stones and the
+like--perished. At St. Helens the elimination was as follows: on
+fences where the pupae were conspicuous, 92%; on bark, 66%; on walls,
+54%; and among nettles, 57%. These interesting experiments confirm our
+views as to protective coloration, and show further, _that the ratio
+of elimination in the species is a very high one, and that therefore
+selection must be very keen_.
+
+We may say that the process of selection follows as a logical
+necessity from the fulfilment of the three preliminary postulates of
+the theory: variability, heredity, and the struggle for existence,
+with its enormous ratio of elimination in all species. To this we must
+add a fourth factor, the _intensification_ of variations which Darwin
+established as a fact, and which we are now able to account for
+theoretically on the basis of germinal selection. It may be objected
+that there is considerable uncertainty about this _logical_ proof,
+because of our inability to demonstrate the selection-value of the
+initial stages and the individual stages of increase. We have
+therefore to fall back on _presumptive evidence_. This is to be found
+in _the interpretative value of the theory_. Let us consider this
+point in greater detail.
+
+In the first place it is necessary to emphasize what is often
+overlooked, namely, that the theory not only explains the
+_transformations_ of species, it also explains _their remaining the
+same_; in addition to the principle of varying, it contains within
+itself that of _persisting_. It is part of the essence of selection,
+that it not only causes a part to _vary_ till it has reached its
+highest pitch of adaptation, but that it _maintains it at this pitch.
+This conserving influence of natural selection_ is of great
+importance, and was early recognised by Darwin; it follows naturally
+from the principle of the survival of the fittest.
+
+We understand from this how it is that a species which has become
+fully adapted to certain conditions of life ceases to vary, but
+remains "constant," as long as the conditions of life _for_ it remain
+unchanged, whether this be for thousands of years, or for whole
+geological epochs. But the most convincing proof of the power of the
+principle of selection lies in the innumerable multitude of phenomena
+which cannot be explained in any other way. To this category belong
+all structures which are only _passively_ of advantage to the
+organism, because none of these can have arisen by the alleged
+_Lamarckian principle_. These have been so often discussed that we
+need do no more than indicate them here. Until quite recently the
+sympathetic coloration of animals--for instance, the whiteness of
+Arctic animals--was referred, at least in part, to the _direct_
+influence of external factors, but the facts can best be explained by
+referring them to the processes of selection, for then it is
+unnecessary to make the gratuitous assumption that many species are
+sensitive to the stimulus of cold and that others are not. The great
+majority of Arctic land-animals, mammals and birds, are white, and
+this proves that they were all able to present the variation which was
+most useful for them. The sable is brown, but it lives in trees, where
+the brown colouring protects and conceals it more effectively. The
+musk-sheep (_Ovibos moschatus_) is also brown, and contrasts sharply
+with the ice and snow, but it is protected from beasts of prey by its
+gregarious habit, and therefore it is of advantage to be visible from
+as great a distance as possible. That so many species have been able
+to give rise to white varieties does not depend on a special
+sensitiveness of the skin to the influence of cold, but to the fact
+that Mammals and Birds have a general tendency to vary towards white.
+Even with us, many birds--starlings, blackbirds, swallows,
+etc.--occasionally produce white individuals, but the white variety
+does not persist, because it readily falls a victim to the carnivores.
+This is true of white fawns, foxes, deer, etc. The whiteness,
+therefore, arises from internal causes, and only persists when it is
+useful. A great many animals living in a _green environment_ have
+become clothed in green, especially insects, caterpillars, and
+Mantidae, both persecuted and persecutors.
+
+That it is not the direct effect of the environment which calls forth
+the green colour is shown by the many kinds of caterpillar which rest
+on leaves and feed on them, but are nevertheless brown. These feed by
+night and betake themselves through the day to the trunk of the tree,
+and hide in the furrows of the bark. We cannot, however, conclude
+from this that they were _unable_ to vary towards green, for there are
+Arctic animals which are white only in winter and brown in summer
+(Alpine hare, and the ptarmigan of the Alps), and there are also green
+leaf-insects which remain green only while they are young and
+difficult to see on the leaf, but which become brown again in the last
+stage of larval life, when they have outgrown the leaf. They then
+conceal themselves by day, sometimes only among withered leaves on the
+ground, sometimes in the earth itself. It is interesting that in one
+genus, Chaerocampa, one species is brown in the last stage of larval
+life, another becomes brown earlier, and in many species the last
+stage is not wholly brown, a part remaining green. Whether this is a
+case of a double adaptation, or whether the green is being gradually
+crowded out by the brown, the fact remains that the same species, even
+the same individual, can exhibit both variations. The case is the same
+with many of the leaf-like Orthoptera, as, for instance, the praying
+mantis (_Mantis religiosa_) which we have already mentioned.
+
+But the best proofs are furnished by those of ten-cited cases in which
+the insect bears a deceptive resemblance to another object. We now
+know many such cases, such as the numerous imitations of green or
+withered leaves, which are brought about in the most diverse ways,
+sometimes by mere variations in the form of the insect and in its
+colour, sometimes by an elaborate marking, like that which occurs in
+the Indian leaf-butterflies, _Kallima inachis_. In the single
+butterfly-genus Anaea, in the woods of South America, there are about
+a hundred species which are all gaily coloured on the upper surface,
+and on the reverse side exhibit the most delicate imitation of the
+colouring and pattern of a leaf, generally without any indication of
+the leaf-ribs, but extremely deceptive nevertheless. Anyone who has
+seen only one such butterfly may doubt whether many of the
+insignificant details of the marking can really be of advantage to the
+insect. Such details are for instance the apparent holes and splits
+in the apparently dry or half-rotten leaf, which are usually due to
+the fact that the scales are absent on a circular or oval patch so
+that the colourless wing-membrane lies bare, and one can look through
+the spot as through a window. Whether the bird which is seeking or
+pursuing the butterflies takes these holes for dewdrops, or for the
+work of a devouring insect, does not affect the question; the
+mirror-like spot undoubtedly increases the general deceptiveness, for
+the same thing occurs in many leaf-butterflies, though not in all, and
+in some cases it is replaced in quite a peculiar manner. In one
+species of Anaea (_A. divina_), the resting butterfly looks exactly
+like a leaf out of the outer edge of which a large semi-circular piece
+has been eaten, possibly by a caterpillar; but if we look more closely
+it is obvious that there is no part of the wing absent, and that the
+semi-circular piece is of a clear, pale yellow colour, while the rest
+of the wing is of a strongly contrasted dark brown.
+
+But the deceptive resemblance may be caused in quite a different
+manner. I have often speculated as to what advantage the brilliant
+white C could give to the otherwise dusky-coloured "Comma butterfly"
+(_Grapta C. album_). Poulton's recent observations[46] have shown that
+this represents the imitation of a crack such as is often seen in dry
+leaves, and is very conspicuous because the light shines through it.
+
+The utility obviously lies in presenting to the bird the very familiar
+picture of a broken leaf with a clear shining slit, and we may
+conclude, from the imitation of such small details, that the birds are
+very sharp observers and that the smallest deviation from the usual
+arrests their attention and incites them to closer investigation. It
+is obvious that such detailed--we might almost say such
+subtle--deceptive resemblances could only have come about in the
+course of long ages through the acquirement from time to time of
+something new which heightened the already existing resemblance.
+
+In face of facts like these there can be no question of chance and no
+one has succeeded so far in finding any other explanation to replace
+that by selection. For the rest, the apparent leaves are by no means
+perfect copies of a leaf; many of them only represent the torn or
+broken piece, or the half or two-thirds of a leaf, but then the leaves
+themselves frequently do not present themselves to the eye as a whole,
+but partially concealed among other leaves. Even those butterflies
+which, like the species of Kallima and Anaea, represent the whole of a
+leaf with stalk, ribs, apex, and the whole breadth, are not actual
+copies which would satisfy a botanist; there is often much wanting. In
+Kallima the lateral ribs of the leaf are never all included in the
+markings; there are only two or three on the left side and at more
+four or five on the right, and in many individuals these are rather
+obscure, while in others they are comparatively distinct. This
+furnishes us with fresh evidence in favour of their origin through
+processes of selection, for a botanically perfect picture could not
+arise in this way; there could only be a fixing of such details as
+heightened the deceptive resemblance.
+
+Our postulate of origin through selection also enables us to
+understand why the leaf-imitation is on the lower surface of the wing
+in the diurnal Lepidoptera, and on the upper surface in the nocturnal
+forms, corresponding to the attitude of the wings in the resting
+position of the two groups.
+
+The strongest of all proofs of the theory, however, is afforded by
+cases of true "mimicry," those adaptations discovered by Bates in
+1861, consisting in the imitation of one species by another, which
+becomes more and more like its model. The model is always a species
+that enjoys some special protection from enemies, whether because it
+is unpleasant to taste, or because it is in some way dangerous.
+
+It is chiefly among insects and especially among butterflies that we
+find the greatest number of such cases. Several of these have been
+minutely studied and every detail has been investigated so that it is
+difficult to understand how there can still be disbelief in regard to
+them. If the many and exact observations which have been carefully
+collected and critically discussed for instance by Poulton[47] were
+thoroughly studied the arguments which are still frequently urged
+against mimicry would be found untenable; we can hardly hope to find
+more convincing proof of the actuality of the processes of selection
+than these cases put into our hands. The preliminary postulates of the
+theory of mimicry have been disputed, for instance, that diurnal
+butterflies are persecuted and eaten by birds, but observations
+specially directed towards this point in India, Africa, America and
+Europe have placed it beyond all doubt. If it were necessary I could
+myself furnish an account of my own observations on this point.
+
+In the same way it has been established by experiment and observation
+in the field that in all the great regions of distribution there are
+butterflies which are rejected by birds and lizards, their chief
+enemies, on account of their unpleasant smell or taste. These
+butterflies are usually gaily and conspicuously coloured and thus--as
+Wallace first interpreted it--are furnished with an easily
+recognisable sign: a sign of unpalatableness or _warning colours_. If
+they were not thus recognisable easily and from a distance, they would
+frequently be pecked at by birds, and then rejected because of their
+unpleasant taste; but as it is, the insect-eaters recognise them at
+once as unpalatable booty and ignore them. Such _immune_[48] species,
+wherever they occur, are imitated by other palatable species, which
+thus acquire a certain degree of protection.
+
+It is true that this explanation of the bright, conspicuous colours
+is only a hypothesis, but its foundations--unpalatableness, and the
+liability of other butterflies to be eaten,--are certain, and its
+consequences--the existence of mimetic palatable forms--conform it in
+the most convincing manner. Of the many cases now known I select one,
+which is especially remarkable, and which has been thoroughly
+investigated, _Papilla dardanus_ (_merope_), a large, beautiful,
+diurnal butterfly which ranges from Abyssinia throughout the whole of
+Africa to the south coast of Cape Colony.
+
+The males of this form are everywhere _almost_ the same in colour and
+in form of wings, save for a few variations in the sparse black
+markings on the pale yellow ground. But the females occur in several
+quite different forms and colourings, and one of these only, the
+Abyssinian form, is like the male, while the other three or four are
+_mimetic_, that is to say, they copy a butterfly of quite a different
+family the Danaids, which are among the _immune_ forms. In each region
+the females have thus copied two or three different immune species.
+There is much that is interesting to be said in regard to these
+species, but it would be out of keeping with the general tenor of this
+paper to give details of this very complicated case of polymorphism in
+_P. Dardanus_. Anyone who is interested in the matter will find a full
+and exact statement of the case in as far as we know it, in Poulton's
+_Essays on Evolution_ (pp. 373-375[49]). I need only add that three
+different mimetic female forms have been reared from the eggs of a
+single female in South Africa. The resemblance of the forms to their
+immune models goes so far that even the details of the _local_ forms
+of the models are copied by the mimetic species.
+
+It remains to be said that in Madagascar a butterfly,
+
+_Papilio meriones_, occurs, of which both sexes are very similar in
+form and markings to the non-mimetic male of _P. dardanus_, so that it
+probably represents the ancestor of this latter species.
+
+In face of such facts as these every attempt at another explanation
+must fail. Similarly all the other details of the case fulfil the
+preliminary postulates of selection, and leave no room for any other
+interpretation. That the males do not take on the protective colouring
+is easily explained, because they are in general more numerous, and
+the females are more important for the preservation of the species,
+and must also live longer in order to deposit their eggs. We find the
+same state of things in many other species, and in one case (_Elymnias
+undularis_) in which the male is also mimetically coloured, it copies
+quite a differently coloured immune species from the model followed by
+the female. This is quite intelligible when we consider that if there
+were _too many_ false immune types, the birds would soon discover that
+there were palatable individuals among those with unpalatable warning
+colours. Hence the imitation of different immune species by _Papilio
+dardanus_!
+
+I regret that lack of space prevents my bringing forward more examples
+of mimicry and discussing them fully. But from the case of _Papilio
+dardanus_ alone there is much to be learnt which is of the highest
+importance for our understanding of transformations. It shows us
+chiefly what I once called, somewhat strongly perhaps, _the
+omnipotence of natural selection_ in answer to an opponent who had
+spoken of its "inadequacy." We here see that one and the same species
+is capable of producing four or five different patterns of colouring
+and marking; thus the colouring and marking are not, as has often been
+supposed, a necessary outcome of the specific nature of the species,
+but a true adaptation, which cannot arise as a direct effect of
+climatic conditions, but solely through what I may call the sorting
+out of the variations produced by the species, according to their
+utility. That caterpillars may be either green or brown is already
+something more than could have been expected according to the old
+conception of species, but that one and the same butterfly should be
+now pale yellow, with black; now red with black and pure white; now
+deep black with large, pure white spots; and again black with a large
+ocheous-yellow spot, and many small white and yellow spots; that in
+one sub-species it may be tailed like the ancestral form, and in
+another tailless like its Danaid model,--all this shows a far-reaching
+capacity for variation and adaptation that we could never have
+expected if we did not see the facts before us. How it is possible
+that the primary colour-variations should thus be intensified and
+combined remains a puzzle even now; we are reminded of the modern
+three-colour printing,--perhaps similar combinations of the primary
+colours take place in this case; in any case the direction of these
+primary variations is determined by the artist whom we know as natural
+selection, for there is no other conceivable way in which the model
+could affect the butterfly that is becoming more and more like it. The
+same climate surrounds all four forms of female; they are subject to
+the same conditions of nutrition. Moreover, _Papilio dardanus_ is by
+no means the only species of butterfly which exhibits different kinds
+of colour-pattern on its wings. Many species of the Asiatic genus
+Elymnias have on the upper surface a very good imitation of an immune
+Euploeine (Danainae), often with a steel-blue ground-colour, while the
+under surface is well concealed when the butterfly is at rest,--thus
+there are two kinds of protective coloration each with a different
+meaning! The same thing may be observed in many non-mimetic
+butterflies, for instance in all our species of Vanessa, in which the
+under side shows a grey-brown or brownish-black protective coloration,
+but we do not yet know with certainty what may be the biological
+significance of the gaily coloured upper surface.
+
+In general it may be said that mimetic butterflies are comparatively
+rare species, but there are exceptions, for instance _Limenitis
+archippus_ in North America, of which the immune model (_Danaida
+plexippus_) also occurs in enormous numbers.
+
+In another mimicry-category the imitators are often more numerous than
+the models, namely in the case of the imitation of _dangerous insects_
+by harmless species. Bees and wasps are dreaded for their sting, and
+they are copied by harmless flies of the genera Eristalis and Syrphus,
+and these mimics often occur in swarms about flowering plants without
+damage to themselves or to their models; they are feared and are
+therefore left unmolested.
+
+In regard also to the _faithfulness of the copy_ the facts are quite
+in harmony with the theory, according to which the resemblance must
+have arisen and increased _by degrees_. We can recognise this in many
+cases, for even now the mimetic species show very _varying degrees of
+resemblance_ to their immune model. If we compare, for instance, the
+many different imitators of _Danaida chrysippus_ we find that, with
+their brownish-yellow ground-colour, and the position and size, and
+more or less sharp limitation of their clear marginal spots, they have
+reached very different degrees of nearness to their model. Or compare
+the female of _Elymnias undularis_ with its model _Danaida genutia_;
+there is a general resemblance, but the marking of the Danaida is very
+roughly imitated in Elymnias.
+
+Another fact that bears out the theory of mimicry is, that even when
+the resemblance in colour-pattern is very great, the _wing-venation_,
+which is so constant, and so important in determining the systematic
+position of butterflies, is never affected by the variation. The
+pursuers of the butterfly have no time to trouble about entomological
+intricacies.
+
+I must not pass over a discovery of Poulton's which is of great
+theoretical importance--that mimetic butterflies may reach the same
+effect by very different means.[50] Thus the glass-like transparency
+of the wing of a certain Ithomiine (Methona) and its Pierine mimic
+(_Dismorphia orise_) depends on a diminution in the size of the
+scales; in the Danaine genus Itune it is due to the fewness of the
+scales and in a third imitator, a moth (_Castnia linus var.
+heliconoides_) the glass-like appearance of the wing is due neither to
+diminution nor to absence of scales, but to their absolute
+colourlessness and transparency, and to the fact that they stand
+upright. In another moth mimic (Anthomyza) the arrangement of the
+transparent scales is normal. Thus it is not some unknown external
+influence that has brought about the transparency of the wing in these
+five forms, as has sometimes been supposed. Nor is it a hypothetical
+_internal_ evolutionary tendency, for all three vary in a different
+manner. The cause of this agreement can only lie in selection, which
+preserves and intensifies in each species the favourable variations
+that present themselves. The great faithfulness of the copy is
+astonishing in these cases, for it is not _the whole_ wing which is
+transparent; certain markings are black in colour, and these contrast
+sharply with the glass-like ground. It is obvious that the pursuers of
+these butterflies must be very sharp-sighted, for otherwise the
+agreement between the species could never have been pushed so far. The
+less the enemies see and observe, the more defective must the
+imitation be, and if they had been blind, no visible resemblance
+between the species which required protection could ever have arisen.
+
+A seemingly irreconcilable contradiction to the mimicry theory is
+presented in the following cases, which were known to Bates, who,
+however, never succeeded in bringing them into line with the principle
+of mimicry.
+
+In South America there are, as we have already said, many mimics of
+the immune Ithomiinae (or as Bates called them Heliconidae). Among
+these there occur not merely species which are edible, and thus
+require the protection of a disguise, but others which are rejected on
+account of their unpalatableness. How could the Ithomiine dress have
+developed in their case, and of what use is it, since the species
+would in any case be immune? In Eastern Brazil, for instance, there
+are four butterflies, which bear a most confusing resemblance to one
+another in colour, marking, and form of wing, and all four are
+unpalatable to birds. They belong to four different genera and three
+sub-families, and we have to inquire: Whence came this resemblance and
+what end does it serve? For a long time no satisfactory answer could
+be found, but Fritz Mueller,[51] seventeen years after Bates, offered a
+solution to the riddle, when he pointed out that young birds could not
+have an instinctive knowledge of the unpalatableness of the
+Ithomiines, but must learn by experience which species were edible and
+which inedible. Thus each young bird must have tasted at least one
+individual of each inedible species and discovered its unpalatability,
+before it learnt to avoid, and thus to spare the species. But if the
+four species resemble each other very closely the bird will regard
+them all as of the same kind, and avoid them all. Thus there developed
+a process of selection which resulted in the survival of the
+Ithomiine-like individuals, and in so great an increase of resemblance
+between the four species, that they are difficult to distinguish one
+from another even in a collection. The advantage for the four species,
+living side by side as they do e.g. in Bahia, lies in the fact that
+only one individual from the _mimicry-ring_ ("inedible association")
+need be tasted by a young bird, instead of at least four individuals,
+as would otherwise be the case. As the number of young birds is great,
+this makes a considerable difference in the ratio of elimination. The
+four Brazilian species are _Lycorea halia_ (Danainae), _Heliconius
+narcaea_ (_eucrate_) (Heliconinae), _Melinaea ethra_, and _Mechanitis
+lysimnia_ (Ithomiinae).
+
+These interesting mimicry-rings (trusts), which have much significance
+for the theory, have been the subject of numerous and careful
+investigations, and at least their essential features are now fully
+established. Mueller took for granted, without making any
+investigations, that young birds only learn by experience to
+distinguish between different kinds of victims. But Lloyd Morgan's[52]
+experiments with young birds proved that this is really the case, and
+at the same time furnished an additional argument against the
+_Lamarckian principle_.
+
+In addition to the mimicry-rings first observed in South America,
+others have been described from Tropical India by Moore, and by
+Poulton and Dixey from Africa, and we may expect to learn many more
+interesting facts in this connection. Here again the preliminary
+postulates of the theory are satisfied. And how much more that would
+lead to the same conclusion might be added!
+
+As in the case of mimicry many species have come to resemble one
+another through processes of selection, so we know whole classes of
+phenomena in which plants and animals have become adapted to one
+another, and have thus been modified to a considerable degree. I refer
+particularly to the relation between flowers and insects. Darwin has
+shown that the originally inconspicuous blossoms of the phanerogams
+were transformed into flowers through the visits of insects, and that,
+conversely, several large orders of insects have been gradually
+modified by their association with flowers, especially as regards the
+parts of their body actively concerned. Bees and butterflies in
+particular have become what they are through their relation to
+flowers. In this case again all that is apparently contradictory to
+the theory can, on closer investigation, be beautifully interpreted in
+corroboration of it. Selection can give rise only to what is of use to
+the organism actually concerned, never to what is of use to some other
+organism, and we must therefore expect to find that in flowers only
+characters of use to _themselves_ have arisen, never characters which
+are of use to insects only, and conversely that in the insects
+characters useful to them and not merely to the plants would have
+originated. For a long time it seemed as if an exception to this rule
+existed in the case of the fertilisation of the yucca blossoms by a
+little moth, _Pronuba yuccasella_. This little moth has a
+sickle-shaped appendage to its mouth-parts which occurs hi no other
+Lepidopteron, and which is used for pushing the yellow pollen into the
+opening of the pistil, thus fertilising the flower. Thus it appears as
+if a new structure, which is useful only to the plant, has arisen in
+the insect. But the difficulty is solved as soon as we learn that the
+moth lays its eggs in the fruit-buds of the Yucca, and that the
+larvae, when they emerge, feed on the developing seeds. In effecting
+the fertilisation of the flower the moth is at the same time making
+provision for its own offspring, since it is only after fertilisation
+that the seeds begin to develop. There is thus nothing to prevent our
+referring this structural adaptation in _Pronuba yuccasella_ to
+processes of selection, which have gradually transformed the maxillary
+palps of the female into the sickle-shaped instrument for collecting
+the pollen, and which have at the same time developed in the insect
+the instinct to press the pollen into the pistil.
+
+In this domain, then, the theory of selection finds nothing but
+corroboration, and it would be impossible to substitute for it any
+other explanation, which now that the facts are so well known, could
+be regarded as a serious rival to it. That selection is a factor, and
+a very powerful factor in the evolution of organisms, can no longer be
+doubted. Even although we cannot bring forward formal proofs of it _in
+detail_, cannot calculate definitely the size of the variations which
+present themselves, and their selection-value, cannot, in short,
+reduce the whole process to a mathematical formula, yet we must assume
+selection, because it is the only possible explanation applicable to
+whole classes of phenomena, and because, on the other hand, it is made
+up of factors which we know can be proved actually to exist, and
+which, _if_ they exist, must of logical necessity cooeperate in the
+manner required by the theory. _We must accept it because the
+phenomena of evolution and adaptation must have a natural basis, and
+because it is the only possible explanation of them._[53]
+
+Many people are willing to admit that selection explains adaptations,
+but they maintain that only a part of the phenomena are thus
+explained, because everything does not depend upon adaptation. They
+regard adaptation as, so to speak, a special effort on the part of
+Nature, which she keeps in readiness to meet particularly difficult
+claims of the external world on organisms. But if we look at the
+matter more carefully we shall find that adaptations are by no means
+exceptional, but that they are present everywhere in such enormous
+numbers, that it would be difficult in regard to any structure
+whatever, to prove that adaptation had _not_ played a part in its
+evolution.
+
+How often has the senseless objection been urged against selection
+that it can create nothing, it can only reject. It is true that it
+cannot create either the living substance or the variations of it;
+both must be given. But in rejecting one thing it preserves another,
+intensifies it, combines it, and in this way _creates_ what is new.
+_Everything_ in organisms depends on adaptation; that is to say,
+everything must be admitted through the narrow door of selection,
+otherwise it can take no part in the building up of the whole. But, it
+is asked, what of the direct effect of external conditions,
+temperature, nutrition, climate and the like? Undoubtedly these can
+give rise to variations, but they too must pass through the door of
+selection, and if they cannot do this they are rejected, eliminated
+from the constitution of the species.
+
+It may, perhaps, be objected that such external influences are often
+of a compelling power, and that every animal must submit to them, and
+that thus selection has no choice and can neither select nor reject.
+There may be such cases; let us assume for instance that the effect
+of the cold of the Arctic regions was to make all the mammals become
+black; the result would be that they would all be eliminated by
+selection, and that no mammals would be able to live there at all. But
+in most cases a certain percentage of animals resists these strong
+influences, and thus selection secures a foothold on which to work,
+eliminating the unfavourable variation, and establishing a useful
+colouring, consistent with what is required for the maintenance of the
+species.
+
+Everything depends upon adaptation! We have spoken much of adaptation
+in colouring, in connection with the examples brought into prominence
+by Darwin, because these are conspicuous, easily verified, and at the
+same time convincing for the theory of selection. But is it only
+desert and polar animals whose colouring is determined through
+adaptation? Or the leaf-butterflies, and the mimetic species, or the
+terrifying markings, and "warning-colours" and a thousand other kinds
+of sympathetic colouring? It is, indeed, never the colouring alone
+which makes up the adaptation; the structure of the animal plays a
+part, often a very essential part, in the protective disguise, and
+thus _many_ variations may cooperate towards _one_ common end. And it
+is to be noted that it is by no means only external parts that are
+changed; internal parts are _always_ modified at the same time--for
+instance, the delicate elements of the nervous system on which depend
+the _instinct_ of the insect to hold its wings, when at rest, in a
+perfectly definite position, which, in the leaf-butterfly, has the
+effect of bringing the two pieces on which the marking occurs on the
+anterior and posterior wing into the same direction, and thus
+displaying as a whole the fine curve of the midrib on the seeming
+leaf. But the wing-holding instinct is not regulated in the same way
+in all leaf-butterflies; even our indigenous species of Vanessa, with
+their protective ground-colouring, have quite a distinctive way of
+holding their wings so that the greater part of the anterior wing is
+covered by the posterior when the butterfly is at rest. But the
+protective colouring appears on the posterior wing and on the tip of
+the anterior, _to precisely the distance to which it is left
+uncovered_. This occurs, as Standfuss has shown, in different degrees
+in our two most nearly allied species, the uncovered portion being
+smaller in _V. urticae_ than in _V. polychloros_. In this case, as in
+most leaf-butterflies, the holding of the wing was probably the
+primary character; only after that was thoroughly established did the
+protective marking develop. In any case, the instinctive manner of
+holding the wings is associated with the protective colouring, and
+must remain as it is if the latter is to be effective. How greatly
+instincts may change, that is to say, may be adapted, is shown by the
+case of the Noctuid "shark" moth, _Xylina vetusta_. This form bears a
+most deceptive resemblance to a piece of rotten wood, and the
+appearance is greatly increased by the modification of the innate
+impulse to flight common to so many animals, which has here been
+transformed into an almost contrary instinct. This moth does not fly
+away from danger, but "feigns death," that is, it draws antennae, legs
+and wings close to the body, and remains perfectly motionless. It may
+be touched, picked up, and thrown down again, and still it does not
+move. This remarkable instinct must surely have developed
+simultaneously with the wood-colouring; at all events, both
+cooeperating variations are now present, and prove that both the
+external and the most minute internal structure have undergone a
+process of adaptation.
+
+The case is the same with all structural variations of animal parts,
+which are not absolutely insignificant. When the insects acquired
+wings they must also have acquired the mechanism with which to move
+them--the musculature, and the nervous apparatus necessary for its
+automatic regulation. All instincts depend upon compound reflex
+mechanisms and are just as indispensable as the parts they have to set
+in motion, and all may have arisen through processes of selection if
+the reasons which I have elsewhere given for this view are
+correct.[54]
+
+Thus there is no lack of adaptations within the organism, and
+particularly in its most important and complicated parts, so that we
+may say that there is no actively functional organ that has not
+undergone a process of adaptation relative to its function and the
+requirements of the organism. Not only is every gland structurally
+adapted, down to the very minutest histological details, to its
+function, but the function is equally minutely adapted to the needs of
+the body. Every cell in the mucous lining of the intestine is exactly
+regulated in its relation to the different nutritive substances, and
+behaves in quite a different way towards the fats, and towards
+nitrogenous substances, or peptones.
+
+I have elsewhere called attention to the many adaptations of the whale
+to the surrounding medium, and have pointed out--what has long been
+known, but is not universally admitted, even now--that in it a great
+number of important organs have been transformed in adaptation to the
+peculiar conditions of aquatic life, although the ancestors of the
+whale must have lived, like other hair-covered mammals, on land. I
+cited a number of these transformations--the fish-like form of the
+body, the hairlessness of the skin, the transformation of the
+fore-limbs to fins, the disappearance of the hind-limbs and the
+development of a tail fin, the layer of blubber under the skin, which
+affords the protection from cold necessary to a warm-blooded animal,
+the disappearance of the ear-muscles and the auditory passages, the
+displacement of the external nares to the forehead for the greater
+security of the breathing-hole during the brief appearance at the
+surface, and certain remarkable changes in the respiratory and
+circulatory organs which enable the animal to remain for a long time
+under water. I might have added many more, for the list of adaptations
+in the whale to aquatic life is by no means exhausted; they are found
+in the histological structure and in the minutest combinations in the
+nervous system. For it is obvious that a tail-fin must be used in
+quite a different way from a tail, which serves as a fly-brush in
+hoofed animals, or as an aid to springing in the kangaroo or as a
+climbing organ; it will require quite different reflex-mechanisms and
+nerve combinations in the motor centres.
+
+I used this example in order to show how unnecessary it is to assume a
+special internal evolutionary power for the phylogenesis of species,
+for this whole order of whales is, so to speak, _made up of
+adaptations_; it deviates in many essential respects from the usual
+mammalian type, and all the deviations are adaptations to aquatic
+life. But if precisely the most essential features of the organisation
+thus depend upon adaptation, what is left for a phyletic force to do,
+since it is these essential features of the structure it would have to
+determine? There are few people now who believe in a phyletic
+evolutionary power, which is not made up of the forces known to
+us--adaptation and heredity--but the conviction that _every_ part of
+an organism depends upon adaptation has not yet gained a firm footing.
+Nevertheless, I must continue to regard this conception as the correct
+one, as I have long done.
+
+I may be permitted one more example. The feather of a bird is a
+marvellous structure, and no one will deny that as a whole it depends
+upon adaptation. But what part of it _does not_ depend upon
+adaptation? The hollow quill, the shaft with its hard, thin, light
+cortex, and the spongy substance within it, its square section
+compared with the round section of the quill, the flat barbs, their
+short, hooked barbules which, in the flight-feathers, hook into one
+another with just sufficient firmness to resist the pressure of the
+air at each wing-beat, the lightness and firmness of the whole
+apparatus, the elasticity of the vane, and so on. And yet all this
+belongs to an organ which is only passively functional, and therefore
+can have nothing to do with the _Lamarckian principle_. Nor can the
+feather have arisen through some magical effect of temperature,
+moisture, electricity, or specific nutrition, and thus selection is
+again our only anchor of safety.
+
+But--it will be objected--the substance of which the feather consists,
+this peculiar kind of horny substance, did not first arise through
+selection in the course of the evolution of the birds, for it formed
+the covering of the scales of their reptilian ancestors. It is quite
+true that a similar substance covered the scales of the Reptiles, but
+why should it not have arisen among them through selection? Or in what
+other way could it have arisen, since scales are also passively useful
+parts? It is true that if we are only to call adaptation what has been
+acquired by the species we happen to be considering, there would
+remain a great deal that could not be referred to selection; but we
+are postulating an evolution which has stretched back through aeons,
+and in the course of which innumerable adaptations took place, which
+had not merely ephemeral persistence in a genus, a family or a class,
+but which was continued into whole Phyla of animals, with continual
+fresh adaptations to the special conditions of each species, family,
+or class, yet with persistence of the fundamental elements. Thus the
+feather, once acquired, persisted in all birds, and the vertebral
+column, once gained by adaptation in the lowest forms, has persisted
+in all the Vertebrates from Amphioxus upwards, although with constant
+readaptation to the conditions of each particular group. Thus
+everything we can see in animals is adaptation, whether of to-day, or
+of yesterday, or of ages long gone by; every kind of cell, whether
+glandular, muscular, nervous, epidermic, or skeletal, is adapted to
+absolutely definite and specific functions, and every organ which is
+composed of these different kinds of cells contains them in the proper
+proportions, and in the particular arrangement which best serves the
+function of the organ; it is thus adapted to its function.
+
+All parts of the organism are tuned to one another, that is, _they are
+adapted to one another_, and in the same way _the organism as a whole
+is adapted to the conditions of its life, and it is so at every stage
+of its evolution._
+
+But all adaptations _can_ be referred to selection; the only point
+that remains doubtful is whether they all _must_ be referred to it.
+
+However that may be, whether the _Lamarckian principle_ is a factor
+that has cooeperated with selection in evolution, or whether it is
+altogether fallacious, the fact remains, that selection is the cause
+of a great part of the phyletic evolution of organisms on our earth.
+Those who agree with me in rejecting the _Lamarckian principle_ will
+regard selection as the only _guiding_ factor in evolution, which
+creates what is new out of the transmissible variations, by ordering
+and arranging these, selecting them in relation to their number and
+size, as the architect does his building-stones so that a particular
+style must result.[55] But the building-stones themselves, the
+variations, have their basis in the influences which cause variation
+in those vital units which are handed on from one generation to
+another, whether, taken together they form the _whole_ organism, as in
+Bacteria and other low forms of life, or only a germ-substance, as in
+unicellular and multicellular organisms.
+
+FOOTNOTES:
+
+[Footnote 33: _Vortraege ueber Descendenztheorie_, Jena, 1904, II. 269.
+Eng. Transl. London, 1904, II. p. 317.]
+
+[Footnote 34: See Poulton, _Essays on Evolution_, Oxford, 1908. pp.
+xix-xxii.]
+
+[Footnote 35: _Origin of Species_ (6th edit), pp. 176 _et seq._]
+
+[Footnote 36: Chun, _Reise der Valdivia_, Leipzig, 1904.]
+
+[Footnote 37: Plate, _Selektionsprinzip u. Probleme der Artbildung_
+(3rd edit.), Leipzig, 1908.]
+
+[Footnote 38: _Studien zur Descendenz-Theorie_ II., "Die Enstehung der
+Zeichnung bei den Schmetterlings-raupen," Leipzig, 1876.]
+
+[Footnote 39: _Origin of Species_ (6th edit.), p. 232.]
+
+[Footnote 40: _Origin of Species_, p. 233; see also edit. 1, p. 242.]
+
+[Footnote 41: _Ibid._ p. 230.]
+
+[Footnote 42: _The Effect of External Influences upon Development_,
+Romanes Lecture, Oxford, 1894.]
+
+[Footnote 43: See Poulton, _Essays on Evolution_, 1908, pp. 316, 317.]
+
+[Footnote 44: _The Evolution Theory_, London, 1904, I. p. 219.]
+
+[Footnote 45: _Report of the British Association_ (Bristol, 1898),
+London, 1899, pp. 906-909.]
+
+[Footnote 46: _Proc. Ent. Soc._, London, May 6, 1903.]
+
+[Footnote 47: _Essays on Evolution_, 1889-1907, Oxford, 1908,
+_passim_, e.g. p. 269.]
+
+[Footnote 48: The expression does not refer to all the enemies of this
+butterfly; against ichneumon-flies, for instance, their unpleasant
+smell usually gives no protection.]
+
+[Footnote 49: Professor Poulton has corrected some wrong descriptions
+which I had unfortunately overlooked in the Plates of my book
+_Vortraege ueber Descendenztheorie_, and which refer to _Papilio
+dardanus_ (_merope_). These mistakes are of no importance as far as an
+understanding of the mimicry-theory is concerned, but I hope shortly
+to be able to correct them in a later edition.]
+
+[Footnote 50: _Journ. Linn. Soc. London_ (_Zool._), Vol. xxvi. 1898,
+pp. 598-602.]
+
+[Footnote 51: In _Kosmos_, 1879, p. 100.]
+
+[Footnote 52: _Habit and Instinct_, London. 1896.]
+
+[Footnote 53: This has been discussed in many of my earlier works. See
+for instance _The All-Sufficiency of Natural Selection, a reply to
+Herbert Spencer_, London, 1893.]
+
+[Footnote 54: _The Evolution Theory_, London, 1904, p. 144.]
+
+[Footnote 55: _Variation under Domestication_, 1875, II. pp. 426,
+427.]
+
+
+
+
+III
+
+HEREDITY AND VARIATION IN MODERN LIGHTS
+
+BY W. BATESON, M.A., F.R.S.
+
+_Professor of Biology in the University of Cambridge_
+
+
+Darwin's work has the property of greatness in that it may be admired
+from more aspects than one. For some the perception of the principle
+of Natural Selection stands out as his most wonderful achievement to
+which all the rest is subordinate. Others, among whom I would range
+myself, look up to him rather as the first who plainly distinguished,
+collected, and comprehensively studied that new class of evidence from
+which hereafter a true understanding of the process of Evolution may
+be developed. We each prefer our own standpoint of admiration; but I
+think that it will be in their wider aspect that his labours will most
+command the veneration of posterity.
+
+A treatise written to advance knowledge may be read in two moods. The
+reader may keep his mind passive, willing merely to receive the
+impress of the writer's thought; or he may read with his attention
+strained and alert, asking at every instant how the new knowledge can
+be used in a further advance, watching continually for fresh footholds
+by which to climb higher still. Of Shelley it has been said that he
+was a poet for poets: so Darwin was a naturalist for naturalists. It
+is when his writings are used in the critical and more exacting spirit
+with which we test the outfit for our own enterprise that we learn
+their full value and strength. Whether we glance back and compare his
+performance with the efforts of his predecessors, or look forward
+along the course which modern research is disclosing, we shall honour
+most in him not the rounded merit of finite accomplishment, but the
+creative power by which he inaugurated a line of discovery endless in
+variety and extension. Let us attempt thus to see his work in true
+perspective between the past from which it grew, and the present which
+is its consequence. Darwin attacked the problem of Evolution by
+reference to facts of three classes: Variation; Heredity; Natural
+Selection. His work was not as the laity suppose, a sudden and
+unheralded revelation, but the first fruit of a long and hitherto
+barren controversy. The occurrence of variation from type, and the
+hereditary transmission of such variation had of course been long
+familiar to practical men, and inferences as to the possible bearing
+of those phenomena on the nature of specific difference had been from
+time to time drawn by naturalists. Maupertuis, for example, wrote: "Ce
+qui nous reste a examiner, c'est comment d'un seul individu, il a pu
+naitre tant d'especes si differentes." And again: "La Nature contient
+le fonds de toutes ces varietes: mais le hasard ou l'art les mettent
+en oeuvre. C'est ainsi que ceux dont l'industrie s'applique a
+satisfaire le gout des curieux, sont, pour ainsi dire, createurs
+d'especes nouvelles."[56]
+
+Such passages, of which many (though few so emphatic) can be found in
+eighteenth century writers, indicate a true perception of the mode of
+Evolution. The speculations hinted at by Buffon,[57] developed by
+Erasmus Darwin, and independently proclaimed above all by Lamarck,
+gave to the doctrine of descent a wide renown. The uniformitarian
+teaching which Lyell deduced from geological observation had gained
+acceptance. The facts of geographical distribution[58] had been shown
+to be obviously inconsistent with the Mosaic legend. Prichard, and
+Lawrence, following the example of Blumenbach, had successfully
+demonstrated that the races of Man could be regarded as different
+forms of one species, contrary to the opinion up till then received.
+These treatises all begin, it is true, with a profound obeisance to
+the sons of Noah, but that performed, they continue on strictly modern
+lines. The question of the mutability of species was thus prominently
+raised.
+
+Those who rate Lamarck no higher than did Huxley in his contemptuous
+phrase "_buccinator tantum_," will scarcely deny that the sound of the
+trumpet had carried far, or that its note was clear. If then there
+were few who had already turned to evolution with positive conviction,
+all scientific men must at least have known that such views had been
+promulgated; and many must, as Huxley says, have taken up his own
+position of "critical expectancy."[59]
+
+Why, then, was it, that Darwin succeeded where the rest had failed?
+The cause of that success was twofold. First, and obviously, in the
+principle of Natural Selection he had a suggestion which would work.
+It might not go the whole way, but it was true as far as it went.
+Evolution could thus in great measure be fairly represented as a
+consequence of demonstrable processes. Darwin seldom endangers the
+mechanism he devised by putting on it strains much greater than it can
+bear. He at least was under no illusion as to the omnipotence of
+Selection; and he introduces none of the forced pleading which in
+recent years has threatened to discredit that principle.
+
+
+
+For example, in the latest text of the _Origin_[60] we find him
+saying:
+
+     "But as my conclusions have lately been much misrepresented,
+     and it has been stated that I attribute the modification of
+     species exclusively to natural selection, I may be permitted
+     to remark that in the first edition of this work, and
+     subsequently, I placed in a most conspicuous
+     position--namely, at the close of the Introduction--the
+     following words: 'I am convinced that natural selection has
+     been the main but not the exclusive means of modification.'"
+
+But apart from the invention of this reasonable hypothesis, which may
+well, as Huxley estimated, "be the guide of biological and
+psychological speculation for the next three or four generations,"
+Darwin made a more significant and imperishable contribution. Not for
+a few generations, but through all ages he should be remembered as the
+first who showed clearly that the problems of Heredity and Variation
+are soluble by observation, and laid down the course by which we must
+proceed to their solution.[61] The moment of inspiration did not come
+with the reading of Malthus, but with the opening of the "first
+note-book on Transmutation of Species."[62] Evolution is a process of
+Variation and Heredity. The older writers, though they had some vague
+idea that it must be so, did not study Variation and Heredity. Darwin
+did, and so begat not a theory, but a science.
+
+The extent to which this is true, the scientific world is only
+beginning to realise. So little was the fact appreciated in Darwin's
+own time that the success of his writings was followed by an almost
+total cessation of work in that special field. Of the causes which led
+to these remarkable consequences I have spoken elsewhere. They
+proceeded from circumstances peculiar to the time; but whatever the
+causes there is no doubt that this statement of the result is
+historically exact, and those who make it their business to collect
+facts elucidating the physiology of Heredity and Variation are well
+aware that they will find little to reward their quest in the leading
+scientific Journals of the Darwinian epoch.
+
+In those thirty years the original stock of evidence current and in
+circulation even underwent a process of attrition. As in the story of
+the Eastern sage who first wrote the collected learning of the
+universe for his sons in a thousand volumes and by successive
+compression and burning reduced them to one and from this by further
+burning distilled the single ejaculation of the Faith "There is no god
+but God and Mohammed is the Prophet of God," which was all his maturer
+wisdom deemed essential:--so in the books of that period do we find
+the _corpus_ of genetic knowledge dwindle to a few prerogative
+instances and these at last to the brief formula of an unquestioned
+creed.
+
+And yet in all else that concerns biological science this period was,
+in very truth, our Golden Age, when the natural history of the earth
+was explored as never before; morphology and embryology were
+exhaustively ransacked; the physiology of plants and animals began to
+rival chemistry and physics in precision of method and in the rapidity
+of its advances; and the foundations of pathology were laid.
+
+In contrast with this immense activity elsewhere the neglect which
+befel the special physiology of Descent, or Genetics as we now call
+it, is astonishing. This may of course be interpreted as meaning that
+the favoured studies seemed to promise a quicker return for effort,
+but it would be more true to say that those who chose these other
+pursuits did so without making any such comparison; for the idea that
+the physiology of Heredity and Variation was a coherent science,
+offering possibilities of extraordinary discovery, was not present to
+their minds at all. In a word, the existence of such a science was
+well nigh forgotten. It is true that in ancillary periodicals, as for
+example those that treat of entomology or horticulture, or in the
+writings of the already isolated systematists,[63] observations with
+this special bearing were from time to time related, but the class of
+fact on which Darwin built his conceptions of Heredity and Variation
+was not seen in the highways of biology. It formed no part of the
+official curriculum of biological students, and found no place among
+the subjects which their teachers were investigating.
+
+During this period nevertheless one distinct advance was made, that
+with which Weismann's name is prominently connected. In Darwin's
+genetic scheme the hereditary transmission of parental experience and
+its consequences played a considerable role. Exactly how great that
+role was supposed to be, he with his habitual caution refrained from
+specifying, for the sufficient reason that he did not know.
+Nevertheless much of the process of Evolution, especially that by
+which organs have become degenerate and rudimentary, was certainly
+attributed by Darwin to such inheritance, though since belief in the
+inheritance of acquired characters fell into dispute, the fact has
+been a good deal overlooked. The _Origin_ without "use and disuse"
+would be a materially different book. A certain vacillation is
+discernible in Darwin's utterances on this question, and the fact gave
+to the astute Butler an opportunity for his most telling attack. The
+discussion which best illustrates the genetic views of the period
+arose in regard to the production of the rudimentary condition of the
+wings of many beetles in the Madeira group of islands, and by
+comparing passages from the _Origin_[64] Butler convicts Darwin of
+saying first that this condition was in the main the result of
+Selection, with disuse aiding, and in another place that the main
+cause of degeneration was disuse, but that Selection had aided. To
+Darwin however I think the point would have seemed one of dialetics
+merely. To him the one paramount purpose was to show that somehow an
+Evolution by means of Variation and Heredity might have brought about
+the facts observed, and whether they had come to pass in the one way
+or the other was a matter of subordinate concern.
+
+To us moderns the question at issue has a diminished significance. For
+over all such debates a change has been brought by Weismann's
+challenge for evidence that use and disuse have any transmitted
+effects at all. Hitherto the transmission of many acquired
+characteristics had seemed to most naturalists so obvious as not to
+call for demonstration.[65] Weismann's demand for facts in support of
+the main proposition revealed at once that none having real cogency
+could be produced. The time-honoured examples were easily shown to be
+capable of different explanations. A few certainly remain which cannot
+be so summarily dismissed, but--though it is manifestly impossible
+here to do justice to such a subject--I think no one will dispute that
+these residual and doubtful phenomena, whatever be their true nature,
+are not of a kind to help us much in the interpretation of any of
+those complex cases of adaptation which on the hypothesis of unguided
+Natural Selection are especially difficult to understand. Use and
+disuse were invoked expressly to help us over these hard places; but
+whatever changes can be induced in offspring by direct treatment of
+the parents, they are not of a kind to encourage hope of real
+assistance from that quarter. It is not to be denied that through the
+collapse of this second line of argument the Selection hypothesis has
+had to take an increased and perilous burden. Various ways of meeting
+the difficulty have been proposed, but these mostly resolve themselves
+into improbable attempts to expand or magnify the powers of Natural
+Selection.
+
+Weismann's interpellation, though negative in purpose, has had a
+lasting and beneficial effect, for through his thorough demolition of
+the old loose and distracting notions of inherited experience, the
+ground has been cleared for the construction of a true knowledge of
+heredity based on experimental fact.
+
+In another way he made a contribution of a more positive character,
+for his elaborate speculations as to the genetic meaning of
+cytological appearances have led to a minute investigation of the
+visible phenomena occurring in those cell-divisions by which
+germ-cells arise. Though the particular views he advocated have very
+largely proved incompatible with the observed facts of heredity, yet
+we must acknowledge that it was chiefly through the stimulus of
+Weismann's ideas that those advances in cytology were made; and though
+the doctrine of the continuity of germ-plasm cannot be maintained in
+the form originally propounded, it is in the main true and
+illuminating.[66] Nevertheless in the present state of knowledge we
+are still as a rule quite unable to connect cytological appearances
+with any genetic consequences and save in one respect (obviously of
+extreme importance--to be spoken of later) the two sets of phenomena
+might, for all we can see, be entirely distinct.
+
+I cannot avoid attaching importance to this want of connection between
+the nuclear phenomena and the features of bodily organisation. All
+attempts to investigate Heredity by cytological means lie under the
+disadvantage that it is the nuclear changes which can alone be
+effectively observed. Important as they must surely be, I have never
+been persuaded that the rest of the cell counts for nothing. What we
+know of the behaviour and variability of chromosomes seems in my
+opinion quite incompatible with the belief that they alone govern
+form, and are the sole agents responsible in heredity.[67]
+
+If, then, progress was to be made in Genetics, work of a different
+kind was required. To learn the laws of Heredity and Variation there
+is no other way than that which Darwin himself followed, the direct
+examination of the phenomena. A beginning could be made by collecting
+fortuitous observations of this class, which have often thrown a
+suggestive light, but such evidence can be at best but superficial and
+some more penetrating instrument of research is required. This can
+only be provided by actual experiments in breeding.
+
+The truth of these general considerations was becoming gradually clear
+to many of us when in 1900 Mendel's work was rediscovered.
+Segregation, a phenomenon of the utmost novelty, was thus revealed.
+From that moment not only in the problem of the origin of species, but
+in all the great problems of biology a new era began. So unexpected
+was the discovery that many naturalists were convinced it was untrue,
+and at once proclaimed Mendel's conclusions as either altogether
+mistaken, or if true, of very limited application. Many fantastic
+notions about the workings of Heredity had been asserted as general
+principles before: this was probably only another fancy of the same
+class.
+
+Nevertheless those who had a preliminary acquaintance with the facts
+of Variation were not wholly unprepared for some such revelation. The
+essential deduction from the discovery of segregation was that the
+characters of living things are dependent on the presence of definite
+elements or factors, which are treated as units in the processes of
+Heredity. These factors can thus be recombined in various ways. They
+act sometimes separately, and sometimes they interact in conduction
+with each other, producing their various effects. All this indicates a
+definiteness and specific order in heredity, and therefore in
+variation. This order cannot by the nature of the case be dependent on
+Natural Selection for its existence, but must be a consequence of the
+fundamental chemical and physical nature of living things. The study
+of Variation had from the first shown that an orderliness of this kind
+was present. The bodies and the properties of livings things are
+cosmic, not chaotic. No matter how low in the scale we go, never do we
+find the slightest hint of a diminution in that all-pervading
+orderliness, nor can we conceive an organism existing for a moment in
+any other state. Moreover not only does this order prevail in normal
+forms, but again and again it is to be seen in newly-sprung varieties,
+which by general consent cannot have been subjected to a prolonged
+Selection. The discovery of Mendelian elements admirably coincided
+with and at once gave a rationale of these facts. Genetic Variation is
+then primarily the consequence of additions to, or omissions from, the
+stock of elements which the species contains. The further
+investigation of the species-problem must thus proceed by the
+analytical method which breeding experiments provide.
+
+In the nine years which have elapsed since Mendel's clue became
+generally known, progress has been rapid. We now understand the
+process by which a polymorphic race maintains its polymorphism. When a
+family consists of dissimilar members, given the numerical proportions
+in which these members are occurring, we can represent their
+composition symbolically and state what types can be transmitted by
+the various members. The difficulty of the "swamping effects of
+inter-crossing" is practically at an end. Even the famous puzzle of
+sex-limited inheritance is solved, at all events in its more regular
+manifestations, and we know now how it is brought about that the
+normal sisters of a colour-blind man can transmit the colour-blindness
+while his normal brothers cannot transmit it.
+
+We are still only on the fringe of the inquiry. It can be seen
+extending and ramifying in many directions. To enumerate these here
+would be impossible. A whole new range of possibilities is being
+brought into view by study of the inter-relations between the simple
+factors. By following up the evidence as to segregation, indications
+have been obtained which can only be interpreted as meaning that when
+many factors are being simultaneously redistributed among the
+germ-cells, certain of them exert what must be described as a
+repulsion upon other factors. We cannot surmise whither this discovery
+may lead.
+
+In the new light all the old problems wear a fresh aspect. Upon the
+question of the nature of Sex, for example, the bearing of Mendelian
+evidence is close. Elsewhere I have shown that from several sets of
+parallel experiments the conclusion is almost forced upon us that, in
+the types investigated, of the two sexes the female is to be regarded
+as heterozygous in sex, containing one unpaired dominant element,
+while the male is similarly homozygous in the absence of that
+element.[68] It is not a little remarkable that on this point--which
+is the only one where observations of the nuclear processes of
+gameto-genesis have yet been brought into relation with the visible
+characteristics of the organisms themselves--there should be
+diametrical opposition between the results of breeding experiments and
+those derived from cytology.
+
+Those who have followed the researches of the American school will be
+aware that, after it had been found in certain insects that the
+spermatozoa were of two kinds according as they contained or did not
+contain the accessory chromosome, E. B. Wilson succeeded in proving
+that the sperms possessing this accessory body were destined to form
+_females_ on fertilisation, while sperms without it form males, the
+eggs being apparently indifferent. Perhaps the most striking of all
+this series of observations is that lately made by T. H. Morgan,[69]
+since confirmed by von Baehr, that in a Phylloxeran two kinds of
+spermatids are formed, respectively with and without an accessory (in
+this case, _double_) chromosome. Of these, only those possessing the
+accessory body become functional spermatozoa, the others degenerating.
+We have thus an elucidation of the puzzling fact that in these forms
+fertilisation results in the formation of _females_ only. How the
+males are formed--for of course males are eventually produced by the
+parthenogenetic females--we do not know.
+
+If the accessory body is really to be regarded as bearing the factor
+for femaleness, then in Mendelian terms female is DD and male is DR.
+The eggs are indifferent and the spermatozoa are each male, _or_
+female. But according to the evidence derived from a study of the
+sex-limited descent of certain features in other animals the
+conclusion seems equally clear that in them female must be regarded as
+DR and male as RR. The eggs are thus each either male or female and
+the spermatozoa are indifferent. How this contradictory evidence is to
+be reconciled we do not yet know. The breeding work concerns fowls,
+canaries, and the Currant moth (_Abraxas grossulariata_). The
+accessory chromosome has been now observed in most of the great
+divisions of insects,[70] except, as it happens, Lepidoptera. At first
+sight it seems difficult to suppose that a feature apparently so
+fundamental as sex should be differently constituted in different
+animals, but that seems at present the least improbable inference. I
+mention these two groups of facts as illustrating the nature and
+methods of modern genetic work. We must proceed by minute and specific
+analytical investigation. Wherever we look we find traces of the
+operation of precise and specific rules.
+
+In the light of present knowledge it is evident that before we can
+attack the Species-problem with any hope of success there are vast
+arrears to be made up. He would be a bold man who would now assert
+that there was no sense in which the term Species might not have a
+strict and concrete meaning in contradistinction to the term Variety.
+We have been taught to regard the difference between species and
+variety as one of degree. I think it unlikely that this conclusion
+will bear the test of further research. To Darwin the question, What
+is a variation? presented no difficulties. Any difference between
+parent and offspring was a variation. Now we have to be more precise.
+First we must, as de Vries has shown, distinguish real, genetic,
+variation from _fluctuational_ variations, due to environmental and
+other accidents, which cannot be transmitted. Having excluded these
+sources of error the variations observed must be expressed in terms of
+the factors to which they are due before their significance can be
+understood. For example, numbers of the variations seen under
+domestication, and not a few witnessed in nature, are simply the
+consequence of some ingredient being in an unknown way omitted from
+the composition of the varying individual. The variation may on the
+contrary be due to the addition of some new element, but to prove that
+it is so is by no means an easy matter. Casual observation is useless,
+for though these latter variations will always be dominants, yet many
+dominant characteristics may arise from another cause, namely the
+meeting of complementary factors, and special study of each case in
+two generations at least is needed before these two phenomena can be
+distinguished.
+
+When such considerations are fully appreciated it will be realised
+that medleys of most dissimilar occurrences are all confused together
+under the term Variation. One of the first objects of genetic analysis
+is to disentangle this mass of confusion.
+
+To those who have made no study of heredity it sometimes appears that
+the question of the effect of conditions in causing variation is one
+which we should immediately investigate, but a little thought will
+show that before any critical inquiry into such possibilities can be
+attempted, a knowledge of the working of heredity under conditions as
+far as possible uniform must be obtained. At the time when Darwin was
+writing, if a plant brought into cultivation gave off an albino
+variety, such an event was without hesitation ascribed to the change
+of life. Now we see that albino _gametes_, germs, that is to say,
+which are destitute of the pigment-forming factor, may have been
+originally produced by individuals standing an indefinite number of
+generations back in the ancestry of the actual albino, and it is
+indeed almost certain that the variation to which the appearance of
+the albino is due cannot have taken place in a generation later than
+that of the grandparents. It is true that when a new _dominant_
+appears we should feel greater confidence that we were witnessing the
+original variation, but such events are of extreme rarity, and no such
+case has come under the notice of an experimenter in modern times, as
+far as I am aware. That they must have appeared is clear enough.
+Nothing corresponding to the Brown-breasted Game fowl is known wild,
+yet that colour is a most definite dominant, and at some moment since
+_Gallus bankiva_ was domesticated, the element on which that special
+colour depends must have at least once been formed in the germ-cell of
+a fowl; but we need harder evidence than any which has yet been
+produced before we can declare that this novelty came through
+over-feeding, or change of climate, or any other disturbance
+consequent on domestication. When we reflect on the intricacies of
+genetic problems as we must now conceive them there come moments when
+we feel almost thankful that the Mendelian principles were unknown to
+Darwin. The time called for a bold pronouncement, and he made it, to
+our lasting profit and delight. With fuller knowledge we pass once
+more into a period of cautious expectation and reserve.
+
+In every arduous enterprise it is pleasanter to look back at
+difficulties overcome than forward to those which still seem
+insurmountable, but in the next stage there is nothing to be stained
+by disguising the fact that the attributes of living things are not
+what we used to suppose. If they are more complex in the sense that
+the properties they display are throughout so regular[71] that the
+Selection of minute random variations is an unacceptable account of
+the origin of their diversity, yet by virtue of that very regularity
+the problem is limited in scope and thus simplified.
+
+To begin with, we must relegate Selection to its proper place.
+Selection permits the viable to continue and decides that the
+non-viable shall perish; just as the temperature of our atmosphere
+decides that no liquid carbon shall be found on the face of the earth:
+but we do not suppose that the form of the diamond has been gradually
+achieved by a process of Selection. So again, as the course of descent
+branches in the successive generations, Selection determines along
+which branch Evolution shall proceed, but it does not decide what
+novelties that branch shall bring forth. "_La Nature contient le fonds
+de toutes ces varietes, mais le hazard ou l'art les mettent en
+oeuvre_," as Maupertuis most truly said.
+
+Not till knowledge of the genetic properties of organisms has attained
+to far greater completeness can evolutionary speculations have more
+than a suggestive value. By genetic experiment, cytology and
+physiological chemistry aiding, we may hope to acquire such knowledge.
+In 1872 Nathusius wrote:[72] "Das Gesetz der Vererbung ist noch nicht
+erkannt; der Apfel ist noch nicht vom Baum der Erkenntniss gefallen,
+welcher, der Sage nach, Newton auf den rechten Weg zur Ergruendung der
+Gravitationsgesetze fuehrte." We cannot pretend that the words are not
+still true, but in Mendelian analysis the seeds of that apple-tree at
+last are sown.
+
+If we were asked what discovery would do most to forward our inquiry,
+what one bit of knowledge would more than any other illuminate the
+problem, I think we may give the answer without hesitation. The
+greatest advance that we can foresee will be made when it is found
+possible to connect the geometrical phenomena of development with the
+chemical. The geometrical symmetry of living things is the key to a
+knowledge of their regularity, and the forces which cause it. In the
+symmetry of the dividing cell the basis of that resemblance we call
+Heredity is contained. To imitate the morphological phenomena of life
+we have to devise a system which can divide. It must be able to
+divide, and to segment as--grossly--a vibrating plate or rod does, or
+as an icicle can do as it becomes ribbed in a continuous stream of
+water; but with this distinction, that the distribution of chemical
+differences and properties must simultaneously be decided and disposed
+in orderly relation to the pattern of the segmentation. Even if a
+model which would do this could be constructed it might prove to be a
+useful beginning.
+
+This may be looking too far ahead. If we had to choose some one piece
+of more proximate knowledge which we would more especially like to
+acquire, I suppose we should ask for the secret of interracial
+sterility. Nothing has yet been discovered to remove the grave
+difficulty, by which Huxley in particular was so much oppressed, that
+among the many varieties produced under domestication--which we all
+regard as analogous to the species seen in nature--no clear case of
+interracial sterility has been demonstrated. The phenomenon is
+probably the only one to which the domesticated products seem to
+afford no parallel. No solution of the difficulty can be offered which
+has positive value, but it is perhaps worth considering the facts in
+the light of modern ideas. It should be observed that we are not
+discussing incompatibility of two species to produce offspring (a
+totally distinct phenomenon), but the sterility of the offspring which
+many of them do produce.
+
+When two species, both perfectly fertile severally, produce on crossing a
+sterile progeny, there is a presumption that the sterility is due to the
+development in the hybrid of some substance which can only be formed by the
+meeting of two complementary factors. That some such account is correct in
+essence may be inferred from the well-known observation that if the hybrid
+is not totally sterile but only partially so, and thus is able to form some
+good germ-cells which develop into new individuals, the sterility of these
+daughter-individuals is sensibly reduced or may be entirely absent. The
+fertility once re-established, the sterility does not return in the later
+progeny, a fact strongly suggestive of segregation. Now if the sterility of
+the cross-bred be really the consequence of the meeting of two
+complementary factors, we see that the phenomenon could only be produced
+among the divergent offspring of one species by the acquisition of at least
+_two_ new factors; for if the acquisition of a single factor caused
+sterility the line would then end. Moreover each factor must be separately
+acquired by distinct individuals, for if both were present together, the
+possessors would by hypothesis be sterile. And in order to imitate the case
+of species each of these factors must be acquired by distinct breeds. The
+factors need not, and probably would not, produce any other perceptible
+effects; they might, like the colour-factors present in white flowers, make
+no difference in the form or other characters. Not till the cross was
+actually made between the two complementary individuals would either factor
+come into play, and the effects even then might be unobserved until an
+attempt was made to breed from the cross-bred.
+
+Next, if the factors responsible for sterility were acquired, they
+would in all probability be peculiar to certain individuals and would
+not readily be distributed to the whole breed. Any member of the breed
+also into which _both_ the factors were introduced would drop out of
+the pedigree by virtue of its sterility. Hence the evidence that the
+various domesticated breeds say of dogs or fowls can when mated
+together produce fertile offspring, is beside the mark. The real
+question is, Do they ever produce sterile offspring? I think the
+evidence is clearly that sometimes they do, oftener perhaps than is
+commonly supposed. These suggestions are quite amenable to
+experimental tests. The most obvious way to begin is to get a pair of
+parents which are known to have had any sterile offspring, and to find
+the proportions in which these steriles were produced. If, as I
+anticipate, these proportions are found to be definite, the rest is
+simple.
+
+In passing, certain other considerations may be referred to. First,
+that there are observations favouring the view that the production of
+totally sterile cross-breds is seldom a universal property of two
+species, and that it may be a matter of individuals, which is just
+what on the view here proposed would be expected. Moreover, as we all
+know now, though incompatibility may be dependent to some extent on
+the degree to which the species are dissimilar, no such principle can
+be demonstrated to determine sterility or fertility in general. For
+example, though all our Finches can breed together, the hybrids are
+all sterile. Of Ducks some species can breed together without
+producing the slightest sterility; others have totally sterile
+offspring, and so on. The hybrids between several _genera_ of Orchids
+are perfectly fertile on the female side, and some on the male side
+also, but the hybrids produced between the Turnip (_Brassica napus_)
+and the Swede (_Brassica campestris_), which, according to our
+estimates of affinity, should be nearly allied forms, are totally
+sterile.[73] Lastly, it may be recalled that in sterility we are
+almost certainly considering a meristic phenomenon. _Failure to
+divide_ is, we may feel fairly sure, the immediate "cause" of the
+sterility. Now, though we know very little about the heredity of
+meristic differences, all that we do know points to the conclusion
+that the less-divided is dominant to the more-divided, and we are thus
+justified in supposing that there are factors which can arrest or
+prevent cell-division. My conjecture therefore is that in the case of
+sterility of cross-breds we see the effect produced by a complementary
+pair of such factors. This and many similar problems are now open to
+our analysis.
+
+The question is sometimes asked, Do the new lights on Variation and
+Heredity make the process of Evolution easier to understand? On the
+whole the answer may be given that they do. There is some appearance
+of loss of simplicity, but the gain is real. As was said above, the
+time is not ripe for the discussion of the origin of species. With
+faith in Evolution unshaken--if indeed the word faith can be used in
+application to that which is certain--we look on the manner and
+causation of adapted differentiation as still wholly mysterious. As
+Samuel Butler so truly said: "To me it seems that the 'Origin of
+Variation,' whatever it is, is the only true 'Origin of Species,'"[74]
+and of that Origin not one of us knows anything. But given
+Variation--and it is given: assuming further that the variations are
+not guided into paths of adaptation--and both to the Darwinian and to
+the modern school this hypothesis appears to be sound if unproven--an
+evolution of species proceeding by definite steps is more, rather than
+less, easy to imagine than an evolution proceeding by the accumulation
+of indefinite and insensible steps. Those who have lost themselves in
+contemplating the miracles of Adaptation (whether real or spurious)
+have not unnaturally fixed their hopes rather on the indefinite than
+on the definite changes. The reasons are obvious. By suggesting that
+the steps through which an adaptative mechanism arose were indefinite
+and insensible, all further trouble is spared. While it could be said
+that species arise by an insensible and imperceptible process of
+variation, there was clearly no use in tiring ourselves by trying to
+perceive that process. This labour-saving counsel found great favour.
+All that had to be done to develop evolution-theory was to discover
+the good in everything, a task which, in the complete absence of any
+control or test whereby to check the truth of the discovery, is not
+very onerous. The doctrine "_que tout est au mieux_" was therefore
+preached with fresh vigour, and examples of that illuminating
+principle were discovered with a facility that Pangloss himself might
+have envied, till at last even the spectators wearied of such dazzling
+performances.
+
+But in all seriousness, why should indefinite and unlimited variation
+have been regarded as a more probable account of the origin of
+Adaptation? Only, I think, because the obstacle was shifted one plane
+back, and so looked rather less prominent. The abundance of
+Adaptation, we all grant, is an immense, almost an unsurpassable
+difficulty in all non-Lamarckian views of Evolution; but if the steps
+by which that adaptation arose were fortuitious, to imagine them
+insensible is assuredly no help. In one most important respect indeed,
+as has often been observed, it is a multiplication of troubles. For
+the smaller the steps, the less could Natural Selection act upon them.
+Definite variations--and of the occurrence of definite variations in
+abundance we have now the most convincing proof--have at least the
+obvious merit that they can make and often do make a real difference
+in the chances of life.
+
+There is another aspect of the Adaptation problem to which I can
+allude very briefly. May not our present ideas of the universality and
+precision of Adaptation be greatly exaggerated? The fit of organism to
+its environment is not after all so very close--a proposition
+unwelcome perhaps, but one which could be illustrated by very copious
+evidence. Natural Selection is stern, but she has her tolerant moods.
+
+We have now most certain and irrefragable proof that much definiteness
+exists in living things apart from Selection, and also much that may
+very well have been preserved and so in a sense constituted by
+Selection. Here the matter is likely to rest. There is a passage in
+the sixth edition of the _Origin_ which has I think been overlooked.
+On page 70 Darwin says, "The tuft of hair on the breast of the wild
+turkey-cock cannot be of any use, and it is doubtful whether it can be
+ornamental in the eyes of the female bird." This tuft of hair is a
+most definite and unusual structure, and I am afraid that the remark
+that it "cannot be of any use" may have been made inadvertently; but
+it may have been intended, for in the first edition the usual
+qualification was given and must therefore have been deliberately
+excised. Anyhow I should like to think that Darwin did throw over that
+tuft of hair, and that he felt relief when he had done so. Whether
+however we have his great authority for such a course or not, I feel
+quite sure that we shall be rightly interpreting the facts of nature
+if we cease to expect to find purposefulness wherever we meet with
+definite structures or patterns. Such things are, as often as not, I
+suspect rather of the nature of tool-marks, mere incidents of
+manufacture, benefiting their possessor not more than the wire-marks
+in a sheet of paper, or the ribbing on the bottom of an oriental plate
+renders those objects more attractive in our eyes.
+
+If Variation may be in any way definite, the question once more
+arises, may it not be definite in direction? The belief that it is has
+had many supporters, from Lamarck onwards, who held that it was guided
+by need, and others who, like Naegeli, while laying no emphasis on
+need, yet were convinced that there was guidance of some kind. The
+latter view under the name of "Orthogenesis," devised I believe by
+Eimer, at the present day commends itself to some naturalists. The
+objection to such a suggestion is of course that no fragment of real
+evidence can be produced in its support. On the other hand, with the
+experimental proof that variation consists largely in the unpacking
+and repacking of an original complexity, it is not so certain as we
+might like to think that the order of these events is not
+predetermined.
+
+For instance the original "pack" may have been made in such a way that
+at the _n_th division of the germ-cells of a Sweet Pea a colour-factor
+might be dropped, and that at the _n_+_n_th division the hooded
+variety be given off, and so on. I see no ground whatever for holding
+such a view, but in fairness the possibility should not be forgotten,
+and in the light of modern research it scarcely looks so absurdly
+improbable as before.
+
+No one can survey the work of recent years without perceiving that
+evolutionary orthodoxy developed too fast, and that a great deal has
+got to come down; but this satisfaction at least remains, that in the
+experimental methods which Mendel inaugurated, we have means of
+reaching certainty in regard to the physiology of Heredity and
+Variation upon which a more lasting structure may be built.
+
+FOOTNOTES:
+
+[Footnote 56: _Venus Physique, contenant deux Dissertations, l'une sur
+l'origine des Hommes et des Animaux_; _Et l'autre sur l'origine des
+Noirs_, La Haye, 1746, pp. 124 and 129. For an introduction to the
+writings of Maupertuis I am indebted to an article by Professor
+Lovejoy in _Popular Sci. Monthly_, 1902.]
+
+[Footnote 57: For the fullest account of the views of these pioneers
+of Evolution, see the works of Samuel Butler, especially _Evolution,
+Old and New_ (2nd edit.) 1882. Butler's claims on behalf of Buffon
+have met with some acceptance; but after reading what Butler has said,
+and a considerable part of Buffon's own works, the word "hinted" seems
+to me a sufficiently correct description of the part he played. It is
+interesting to note that in the chapter on the Ass, which contains
+some of his evolutionary passages, there is a reference to "_plusieurs
+idees tres-elevees sur la generation_" contained in the Letters of
+Maupertuis.]
+
+[Footnote 58: See especially W. Lawrence, _Lectures on Physiology_,
+London, 1823, pp. 213 f.]
+
+[Footnote 59: See the chapter contributed to the _Life and Letters of
+Charles Darwin_, II. p. 195. I do not clearly understand the sense in
+which Darwin wrote (Autobiography, _ibid._ I. p. 87): "It has
+sometimes been said that the success of the _Origin_ proved 'that the
+subject was in the air,' or 'that men's minds were prepared for it.' I
+do not think that this is strictly true, for I occasionally sounded
+not a few naturalists, and never happened to come across a single one
+who seemed to doubt about the permanence of species." This experience
+may perhaps have been an accident due to Darwin's isolation. The
+literature of the period abounds with indications of "critical
+expectancy." A most interesting expression of that feeling is given in
+the charming account of the "Early Days of Darwinism" by Alfred
+Newton, _Macmillan's Magazine_, LVII. 1888, p. 241. He tells how in
+1858 when spending a dreary summer in Iceland, he and his friend, the
+ornithologist John Wolley, in default of active occupation, spent
+their days in discussion. "Both of us taking a keen interest in
+Natural History, it was but reasonable that a question, which in those
+days was always coming up wherever two or more naturalists were
+gathered together, should be continually recurring. That question was,
+'What is a species?' and connected therewith was the other question,
+'How did a species begin?'... Now we were of course fairly well
+acquainted with what had been published on these subjects." He then
+enumerates some of these publications, mentioning among others T.
+Vernon Wollaston's _Variation of Species_--a work which has in my
+opinion never been adequately appreciated. He proceeds: "Of course we
+never arrived at anything like a solution of these problems, general
+or special, but we felt very strongly that a solution ought to be
+found, and that quickly, if the study of Botany and Zoology was to
+make any great advance." He then describes how on his return home he
+received the famous number of the _Linnean Journal_ on a certain
+evening. "I sat up late that night to read it; and never shall I
+forget the impression it made upon me. Herein was contained a
+perfectly simple solution of all the difficulties which had been
+troubling me for months past.... I went to bed satisfied that a
+solution had been found."]
+
+[Footnote 60: _Origin_, 6th edit. (1882), p. 421.]
+
+[Footnote 61: Whatever be our estimate of the importance of Natural
+Selection, in this we all agree. Samuel Butler, the most brilliant,
+and by far the most interesting of Darwin's opponents--whose works are
+at length emerging from oblivion--in his Preface (1882) to the 2nd
+edition of _Evolution, Old and New_, repeats his earlier expression of
+homage to one whom he had come to regard as an enemy: "To the end of
+time, if the question be asked, 'Who taught people to believe in
+Evolution?' the answer must be that it was Mr. Darwin. This is true,
+and it is hard to see what palm of higher praise can be awarded to any
+philosopher."]
+
+[Footnote 62: _Life and Letters_, I. pp. 276 and 83.]
+
+[Footnote 63: This isolation of the systematists is the one most
+melancholy sequela of Darwinism. It seems an irony that we should read
+in the peroration to the _Origin_ that when the Darwinian view is
+accepted "Systematists will be able to pursue their labours as at
+present; but they will not be incessantly haunted by the shadowy doubt
+whether this or that form be a true species. This, I feel sure, and I
+speak after experience, will be no slight relief. The endless disputes
+whether or not some fifty species of British brambles are good species
+will cease." _Origin_, 6th edit. (1882), p. 425. True they have ceased
+to attract the attention of those who lead opinion, but anyone who
+will turn to the literature of systematics will find that they have
+not ceased in any other sense. Should there not be something
+disquieting in the fact that among the workers who come most into
+contact with specific differences, are to be found the only men who
+have failed to be persuaded of the unreality of those differences?]
+
+[Footnote 64: 6th edit. pp. 109 and 401. See Butler, _Essays on Life,
+Art, and Science_, p. 265, reprinted 1908, and _Evolution, Old and
+New_, chap. XXII. (2nd edit.), 1882.]
+
+[Footnote 65: W. Lawrence was one of the few who consistently
+maintained the contrary opinion. Prichard, who previously had
+expressed himself in the same sense, does not, I believe, repeat these
+views in his later writings, and there are signs that he came to
+believe in the transmission of acquired habits. See Lawrence, _Lect.
+Physiol._ 1823, pp. 436-437, 447. Prichard, Edin. Inaug. Disp. 1808
+[not seen by me], quoted _ibid._ and _Nat. Hist. Man_, 1843, pp. 34
+f.]
+
+[Footnote 66: It is interesting to see how nearly Butler was led by
+natural penetration, and from absolutely opposite conclusions, back to
+this underlying truth: "So that each ovum when impregnate should be
+considered not as descended from its ancestors, but as being a
+continuation of the personality of every ovum in the chain of its
+ancestry, which every ovum _it actually is_ quite as truly as the
+octogenarian _is_ the same identity with the ovum from which he has
+been developed. This process cannot stop short of the primordial cell,
+which again will probably turn out to be but a brief resting-place. We
+therefore prove each one of us to _be actually_ the primordial cell
+which never died nor dies, but has differentiated itself into the life
+of the world, all living beings whatever, being one with it and
+members one of another," _Life and Habit_, 1878, p. 86.]
+
+[Footnote 67: This view is no doubt contrary to the received opinion.
+I am however interested to see it lately maintained by Driesch
+(_Science and Philosophy of the Organism_, London, 1907, p. 233), and
+from the recent observations of Godlewski it has received distinct
+experimental support.]
+
+[Footnote 68: In other words, the ova are each _either_ female, _or
+male_ (i.e. non-female), but the sperms are all non-female.]
+
+[Footnote 69: Morgan, _Proc. Soc. Exp. Biol. Med._ V. 1908, and von
+Baehr, _Zool. Anz._ XXXII. p. 507, 1908.]
+
+[Footnote 70: As Wilson has proved, the unpaired body is not a
+universal feature even in those orders in which it has been observed.
+Nearly allied types may differ. In some it is altogether unpaired. In
+others it is paired with a body of much smaller size, and by selection
+of various types all gradations can be demonstrated ranging to the
+condition in which the members of the pair are indistinguishable from
+each other.]
+
+[Footnote 71: I have in view, for example, the marvellous and specific
+phenomena of regeneration, and those discovered by the students of
+"_Entwicklungsmechanik_." The circumstances of its occurrence here
+preclude any suggestion that this regularity has been brought about by
+the workings of Selection. The attempts thus to represent the
+phenomena have resulted in mere parodies of scientific reasoning.]
+
+[Footnote 72: _Vortraege ueber Viehzucht und Rassenerkenntniss_, p. 120,
+Berlin, 1872.]
+
+[Footnote 73: See Sutton, A. W., _Journ. Linn. Soc._ XXXVIII. p. 341,
+1908.]
+
+[Footnote 74: _Life and Habit_, London, p. 263, 1878]
+
+
+
+
+IV
+
+"THE DESCENT OF MAN"
+
+BY G. SCHWALBE
+
+_Professor of Anatomy in the University of Strassburg_
+
+
+The problem of the origin of the human race, of the descent of man, is
+ranked by Huxley in his epoch-making book _Man's Place in Nature_, as
+the deepest with which biology has to concern itself, "the question of
+questions,"--the problem which underlies all others. In the same
+brilliant and lucid exposition, which appeared in 1863, soon after the
+publication of Darwin's _Origin of Species_, Huxley stated his own
+views in regard to this great problem. He tells us how the idea of a
+natural descent of man gradually grew up in his mind. It was
+especially the assertions of Owen in regard to the total difference
+between the human and the simian brain that called forth strong
+dissent from the great anatomist Huxley, and he easily succeeded in
+showing that Owen's supposed differences had no real existence; he
+even established, on the basis of his own anatomical investigations,
+the proposition that the anatomical differences between the Marmoset
+and the Chimpanzee are much greater than those between the Chimpanzee
+and Man.
+
+But why do we thus introduce the study of Darwin's _Descent of Man_,
+which is to occupy us here, by insisting on the fact that Huxley had
+taken the field in defence of the descent of man in 1863, while
+Darwin's book on the subject did not appear till 1871? It is in order
+that we may clearly understand how it happened that from this time
+onwards Darwin and Huxley followed the same great aim in the most
+intimate association.
+
+Huxley and Darwin working at the same _Problema maximum_! Huxley
+fiery, impetuous, eager for battle, contemptuous of the resistance of
+a dull world, or energetically triumphing over it. Darwin calm,
+weighing every problem slowly, letting it mature thoroughly,--not a
+fighter, yet having the greater and more lasting influence by virtue
+of his immense mass of critically sifted proofs. Darwin's friend,
+Huxley, was the first to do him justice, to understand his nature, and
+to find in it the reason why the detailed and carefully considered
+book on the descent of man made its appearance so late. Huxley, always
+generous, never thought of claiming priority for himself. In
+enthusiastic language he tells how Darwin's immortal work, _The Origin
+of Species_, first shed light for him on the problem of the descent of
+man; the recognition of a _vera causa_ in the transformation of
+species illuminated his thoughts as with a flash. He was now content
+to leave what perplexed him, what he could not yet solve, as he says
+himself, "in the mighty hands of Darwin." Happy in the bustle of
+strife against old and deep-rooted prejudices, against intolerance and
+superstition, he wielded his sharp weapons on Darwin's behalf; wearing
+Darwin's armour he joyously overthrew adversary after adversary.
+Darwin spoke of Huxley as his "general agent."[75] Huxley says of
+himself "I am Darwin's bulldog."[76]
+
+Thus Huxley openly acknowledged that it was Darwin's _Origin of
+Species_ that first set the problem of the descent of man in its true
+light, that made the question of the origin of the human race a
+pressing one. That this was the logical consequence of his book Darwin
+himself had long felt. He had been reproached with intentionally
+shirking the application of his theory to Man. Let us hear what he
+says on this point in his autobiography: "As soon as I had become, in
+the year 1837 or 1838, convinced that species were mutable
+productions, I could not avoid the belief that man must come under the
+same law. Accordingly I collected notes on the subject for my own
+satisfaction, and not for a long time with any intention of
+publishing. Although in the 'Origin of Species' the derivation of any
+particular species is never discussed, yet I thought it best, in order
+_that no honourable man should accuse me of concealing my views_,[77]
+to add that by the work 'light would be thrown on the origin of man
+and his history.' It would have been useless and injurious to the
+success of the book to have paraded, without giving any evidence, my
+conviction with respect to his origin."[78]
+
+In a letter written in January, 1860, to the Rev. L. Blomefield,
+Darwin expresses himself in similar terms. "With respect to man, I am
+very far from wishing to obtrude my belief; but I thought it dishonest
+to quite conceal my opinion."[79]
+
+The brief allusion in the _Origin of Species_ is so far from prominent
+and so incidental that it was excusable to assume that Darwin had not
+touched upon the descent of man in this work. It was solely the desire
+to have his mass of evidence sufficiently complete, solely Darwin's
+great characteristic of never publishing till he had carefully weighed
+all aspects of his subject for years, solely, in short, his most
+fastidious scientific conscience that restrained him from challenging
+the world in 1859 with a book in which the theory of the descent of
+man was fully set forth. Three years, frequently interrupted by
+ill-health, were needed for the actual writing of the book:[80] the
+first edition, which appeared in 1871, was followed in 1874 by a much
+improved second edition, the preparation of which he very reluctantly
+undertook.[81]
+
+This, briefly, is the history of the work, which, with the _Origin of
+Species_, marks an epoch in the history of biological sciences--the
+work with which the cautious, peace-loving investigator ventured forth
+from his contemplative life into the arena of strife and unrest, and
+laid himself open to all the annoyances that deep-rooted belief and
+prejudice, and the prevailing tendency of scientific thought at the
+time could devise.
+
+Darwin did not take this step lightly. Of great interest in this
+connection is a letter written to Wallace on Dec. 22, 1857,[82] in
+which he says, "You ask me whether I shall discuss 'man.' I think I
+shall avoid the whole subject, as so surrounded with prejudices;
+though I fully admit that it is the highest and most interesting
+problem for the naturalist." But his conscientiousness compelled him
+to state briefly his opinion on the subject in the _Origin of Species_
+in 1859. Nevertheless he did not escape reproaches for having been so
+reticent. This is unmistakably apparent from a letter to Fritz Mueller
+dated Feb. 22 [1869?], in which he says: "I am thinking of writing a
+little essay on the Origin of Mankind, as I have been taunted with
+concealing my opinions."[83]
+
+It might be thought that Darwin behaved thus hesitatingly, and was so
+slow in deciding on the full publication of his collected material in
+regard to the descent of man, because he had religious difficulties to
+overcome.
+
+But this was not the case, as we can see from his admirable confession
+of faith, the publication of which we owe to his son Francis.[84]
+Whoever wishes really to understand the lofty character of this great
+man should read these immortal lines in which he unfolds to us in
+simple and straightforward words the development of his conception of
+the universe. He describes how, though he was still quite orthodox
+during his voyage round the world on board the _Beagle_, he came
+gradually to see, shortly afterwards (1836-1839) that the Old
+Testament was no more to be trusted than the Sacred Books of the
+Hindoos; the miracles by which Christianity is supported, the
+discrepancies between the accounts in the different Gospels, gradually
+led him to disbelieve in Christianity as a divine revelation. "Thus,"
+he writes,[85] "disbelief crept over me at a very slow rate, but was
+at last complete. The rate was so slow that I felt no distress." But
+Darwin was too modest to presume to go beyond the limits laid down by
+science. He wanted nothing more than to be able to go, freely and
+unhampered by belief in authority or in the Bible, as far as human
+knowledge could lead him. We learn this from the concluding words of
+his chapter on religion "The mystery of the beginning of all things is
+insoluble by us; and I for one must be content to remain an
+Agnostic."[86]
+
+Darwin was always very unwilling to give publicity to his views in
+regard to religion. In a letter to Asa Gray on May 22, 1860,[87] he
+declares that it is always painful to him to have to enter into
+discussion of religious problems. He had, he said, no intention of
+writing atheistically.
+
+Finally, let us cite one characteristic sentence from a letter from
+Darwin to C. Ridley[88] (Nov. 28, 1878). A clergyman, Dr. Pusey, had
+asserted that Darwin had written the _Origin of Species_ with some
+relation to theology. Darwin writes emphatically, "Many years ago when
+I was collecting facts for the 'Origin,' my belief in what is called a
+personal God was as firm as that of Dr. Pusey himself, and as to the
+eternity of matter I never troubled myself about such insoluble
+questions." The expression "many years ago" refers to the time of his
+voyage round the world, as has already been pointed out. Darwin means
+by this utterance that the views which had gradually developed in his
+mind in regard to the origin of species were quite compatible with the
+faith of the Church.
+
+If we consider all these utterances of Darwin in regard to religion
+and to his outlook on life (Weltanschauung), we shall see at least so
+much, that religious reflection could in no way have influenced him in
+regard to the writing and publishing of his book on _The Descent of
+Man_. Darwin had early won for himself freedom of thought, and to this
+freedom he remained true to the end of his life, uninfluenced by the
+customs and opinions of the world around him.
+
+Darwin was thus inwardly fortified and armed against the host of
+calumnies, accusations, and attacks called forth by the publication of
+the _Origin of Species_, and to an even greater extent by the
+appearance of the _Descent of Man_. But in his defence he could rely
+on the aid of a band of distinguished auxiliaries of the rarest
+ability. His faithful confederate, Huxley, was joined by the botanist
+Hooker, and, after longer resistance, by the famous geologist Lyell,
+whose "conversion" afforded Darwin peculiar satisfaction. All three
+took the field with enthusiasm in defence of the natural descent of
+man. From Wallace, on the other hand, though he shared with him the
+idea of natural selection, Darwin got no support in this matter.
+Wallace expressed himself in a strange manner. He admitted everything
+in regard to the morphological descent of man, but maintained, in a
+mystic way, that something else, something of a spiritual nature must
+have been added to what man inherited from his animal ancestors.
+Darwin, whose esteem for Wallace was extraordinarily high, could not
+understand how he could give utterance to such a mystical view in
+regard to man; the idea seemed to him so "incredibly strange" that he
+thought some one else must have added these sentences to Wallace's
+paper.
+
+Even now there are thinkers who, like Wallace, shrink from applying to
+man the ultimate consequences of the theory of descent. The idea that
+man is derived from ape-like forms is to them unpleasant and
+humiliating.
+
+So far I have been depicting the development of Darwin's work on the
+descent of man. In what follows I shall endeavour to give a condensed
+survey of the contents of the book.
+
+It must at once be said that the contents of Darwin's work fall into
+two parts, dealing with entirely different subjects. _The Descent of
+Man_ includes a very detailed investigation in regard to secondary
+sexual characters in the animal series, and on this investigation
+Darwin founded a new theory, that of sexual selection. With
+astonishing patience he gathered together an immense mass of material,
+and showed, in regard to Arthropods and Vertebrates, the wide
+distribution of secondary characters, which develop almost exclusively
+in the male, and which enable him, on the one hand, to get the better
+of his rivals in the struggle for the female by the greater perfection
+of his weapons, and, on the other hand, to offer greater allurements
+to the female through the higher development of decorative characters,
+of song, or of scent-producing glands. The best equipped males will
+thus crowd out the less well-equipped in the matter of reproduction,
+and thus the relevant characters will be increased and perfected
+through sexual selection. It is, of course, a necessary assumption
+that these secondary sexual characters may be transmitted to the
+female, although perhaps in rudimentary form.
+
+As we have said, this story of sexual selection takes up a great deal
+of space in Darwin's book, and it need only be considered here in so
+far as Darwin applied it to the descent of man. To this latter problem
+the whole of Part I is devoted, while Part III contains a discussion
+of sexual selection in relation to man, and a general summary. Part
+II treats of sexual selection in general, and may be disregarded in
+our present study. Moreover, many interesting details must necessarily
+be passed over in what follows, for want of space.
+
+The first part of the _Descent of Man_ begins with an enumeration of
+the proofs of the animal descent of man taken from the structure of
+the human body. Darwin chiefly emphasises the fact that the human body
+consists of the same organs and of the same tissues as those of the
+other mammals; he shows also that man is subject to the same diseases
+and tormented by the same parasites as the apes. He further dwells on
+the general agreement exhibited by young embryonic forms, and he
+illustrates this by two figures placed one above the other, one
+representing a human embryo, after Ecker, the other a dog embryo,
+after Bischoff.[89]
+
+Darwin finds further proofs of the animal origin of man in the reduced
+structures, in themselves extremely variable, which are either
+absolutely useless to their possessors, or of so little use that they
+could never have developed under existing conditions. Of such vestiges
+he enumerates: the defective development of the _panniculus carnosus_
+(muscle of the skin) so widely distributed among mammals, the
+ear-muscles, the occasional persistence of the animal ear-point in
+man, the rudimentary nictitating membrane (_plica semilunaris_) in the
+human eye, the slight development of the organ of smell, the general
+hairiness of the human body, the frequently defective development or
+entire absence of the third molar (the wisdom tooth), the vermiform
+appendix, the occasional reappearance of a bony canal (_foramen
+supracondyloideum_) at the lower end of the humerus, the rudimentary
+tail of man (the so-called taillessness), and so on. Of these
+rudimentary structures the occasional occurrence of the animal
+ear-point in man is most fully discussed. Darwin's attention was
+called to this interesting structure by the sculptor Woolner. He
+figures such a case observed in man, and also the head of an alleged
+orang-foetus, the photograph of which he received from Nitsche.
+
+Darwin's interpretation of Woolner's case as having arisen through a
+folding over of the free edge of a pointed ear has been fully borne
+out by my investigations on the external ear.[90] In particular, it
+was established by these investigations that the human foetus, about
+the middle of its embryonic life, possesses a pointed ear somewhat
+similar to that of the monkey genus Macacus. One of Darwin's
+statements in regard to the head of the orang-foetus must be
+corrected. A _large_ ear with a point is shown in the photograph,[91]
+but it can easily be demonstrated--and Deniker has already pointed
+this out--that the figure is not that of an orang foetus at all, for
+that form has much smaller ears with no point; nor can it be a
+gibbon-foetus, as Deniker supposes, for the gibbon ear is also without
+a point. I myself regard it as that of a Macacus-embryo. But this
+mistake, which is due to Nitsche, in no way affects the fact
+recognised by Darwin, that ear-forms showing the point characteristic
+of the animal ear occur in man with extraordinary frequency.
+
+Finally, there is a discussion of those rudimentary structures which
+occur only in _one_ sex, such as the rudimentary mammary glands in the
+male, the vesicula prostatica, which corresponds to the uterus of the
+female, and others. All these facts tell in favour of the common
+descent of man and all other vertebrates. The conclusion of this
+section is characteristic: "_It is only our natural prejudice, and
+that arrogance which made our forefathers declare that they were
+descended from demi-gods, which leads us to demur to this conclusion.
+But the time will before long come, when it will be thought wonderful
+that naturalists, who were well acquainted with the comparative
+structure and development of man, and other mammals, should have
+believed that each was the work of a separate act of creation._"[92]
+
+In the second chapter there is a more detailed discussion, again based
+upon an extraordinary wealth of facts, of the problem as to the manner
+in which, and the causes through which, man evolved from a lower form.
+Precisely the same causes are here suggested for the origin of man, as
+for the origin of species in general. Variability, which is a
+necessary assumption in regard to all transformations, occurs in man
+to a high degree. Moreover, the rapid multiplication of the human race
+creates conditions which necessitate an energetic struggle for
+existence, and thus afford scope for the intervention of natural
+selection. Of the exercise of _artificial_ selection in the human
+race, there is nothing to be said, unless we cite such cases as the
+grenadiers of Frederick William I, or the population of ancient
+Sparta. In the passages already referred to and in those which follow,
+the transmission of acquired characters, upon which Darwin does not
+dwell, is taken for granted. In man, direct effects of changed
+conditions can be demonstrated (for instance in regard to bodily
+size), and there are also proofs of the influence exerted on his
+physical constitution by increased use or disuse. Reference is here
+made to the fact, established by Forbes, that the Quechua Indians of
+the high plateaus of Peru show a striking development of lungs and
+thorax, as a result of living constantly at high altitudes.
+
+Such special forms of variation as arrests of development
+(microcephalism) and reversion to lower forms are next discussed.
+Darwin himself felt[93] that these subjects are so nearly related to
+the cases mentioned in the first chapter, that many of them might as
+well have been dealt with there. It seems to me that it would have
+been better so, for the citation of additional instances of reversion
+at this place rather disturbs the logical sequence of his ideas as to
+the conditions which have brought about the evolution of man from
+lower forms. The instances of reversion here discussed are
+microcephalism, which Darwin wrongly interpreted as atavistic,
+supernumerary mammae, supernumerary digits, bicornuate uterus, the
+development of abnormal muscles, and so on. Brief mention is also made
+of correlative variations observed in man.
+
+Darwin next discusses the question as to the manner in which man
+attained to the erect position from the state of a climbing quadruped.
+Here again he puts the influence of Natural Selection in the first
+rank. The immediate progenitors of man had to maintain a struggle for
+existence in which success was to the more intelligent, and to those
+with social instincts. The hand of these climbing ancestors, which had
+little skill and served mainly for locomotion, could only undergo
+further development when some early member of the Primate series came
+to live more on the ground and less among trees.
+
+A bipedal existence thus became possible, and with it the liberation
+of the hand from locomotion, and the one-sided development of the
+human foot. The upright position brought about correlated variations
+in the bodily structure; with the free use of the hand it became
+possible to manufacture weapons and to use them; and this again
+resulted in a degeneration of the powerful canine teeth and the jaws,
+which were then no longer necessary for defence. Above all, however,
+the intelligence immediately increased, and with it skull and brain.
+The nakedness of man, and the absence of a tail (rudimentariness of
+the tail vertebrae) are next discussed. Darwin is inclined to
+attribute the nakedness of man, not to the action of natural selection
+on ancestors who originally inhabited a tropical land, but to sexual
+selection, which, for aesthetic reasons, brought about the loss of the
+hairy covering in man, or primarily in woman. An interesting
+discussion of the loss of the tail, which, however, man shares with
+the anthropoid apes, some other monkeys and lemurs, forms the
+conclusion of the almost superabundant material which Darwin worked
+up in the second chapter. His object was to show that some of the most
+distinctive human characters are in all probability directly or
+indirectly due to natural selection. With characteristic modesty he
+adds:[94] "Hence, if I have erred in giving to natural selection great
+power, which I am very far from admitting, or in having exaggerated
+its power, which is in itself probable, I have at least, as I hope,
+done good service in aiding to overthrow the dogma of separate
+creations." At the end of the chapter he touches upon the objection as
+to man's helpless and defenceless condition. Against this he urges his
+intelligence and social instincts.
+
+The two following chapters contain a detailed discussion of the
+objections drawn from the supposed great differences between the
+mental powers of men and animals. Darwin at once admits that the
+differences are enormous, but not that any fundamental difference
+between the two can be found. Very characteristic of him is the
+following passage: "In what manner the mental powers were first
+developed in the lowest organisms, is as hopeless an enquiry as how
+life itself first originated. These are problems for the distant
+future, if they are ever to be solved by man."[95]
+
+After some brief observations on instinct and intelligence, Darwin
+brings forward evidence to show that the greater number of the
+emotional states, such as pleasure and pain, happiness and misery,
+love and hate are common to man and the higher animals. He goes on to
+give various examples showing that wonder and curiosity, imitation,
+attention, memory and imagination (dreams of animals), can also be
+observed in the higher mammals, especially in apes. In regard even to
+reason there are no sharply defined limits. A certain faculty of
+deliberation is characteristic of some animals, and the more
+thoroughly we know an animal the more intelligence we are inclined to
+credit it with. Examples are brought forward of the intelligent and
+deliberate actions of apes, dogs and elephants. But although no
+sharply defined differences exist between man and animals, there is,
+nevertheless, a series of other mental powers which are
+characteristics usually regarded as absolutely peculiar to man. Some
+of these characteristics are examined in detail, and it is shown that
+the arguments drawn from them are not conclusive. Man alone is said to
+be capable of progressive improvement; but against this must be placed
+as something analogous in animals, the fact that they learn cunning
+and caution through long continued persecution. Even the use of tools
+is not in itself peculiar to man (monkeys use sticks, stones and
+twigs), but man alone fashions and uses implements _designed for a
+special purpose_. In this connection the remarks taken from Lubbock in
+regard to the origin and gradual development of the earliest flint
+implements will be read with interest; these are similar to the
+observations on modern eoliths, and their bearing on the development
+of the stone industry. It is interesting to learn from a letter to
+Hooker,[96] that Darwin himself at first doubted whether the stone
+implements discovered by Boucher de Perthes were really of the nature
+of tools. With the relentless candour as to himself which
+characterised him, he writes four years later in a letter to Lyell in
+regard to this view of Boucher de Perthes' discoveries: "I know
+something about his errors, and looked at his book many years ago, and
+am ashamed to think that I concluded the whole was rubbish! Yet he has
+done for man something like what Agassiz did for glaciers."[97]
+
+To return to Darwin's further comparisons between the higher mental powers
+of man and animals; He takes much of the force from the argument that man
+alone is capable of abstraction and self-consciousness by his own
+observations on dogs. One of the main differences between man and animals,
+speech, receives detailed treatment. He points out that various animals
+(birds, monkeys, dogs) have a large number of different sounds for
+different emotions, that, further, man produces in common with animals a
+whole series of inarticulate cries combined with gestures, and that dogs
+learn to understand whole sentences of human speech. In regard to human
+language, Darwin expresses a view contrary to that held by Max Mueller:[98]
+"I cannot doubt that language owes its origin to the imitation and
+modification of various natural sounds, the voices of other animals, and
+man's own instinctive cries, aided by signs and gestures." The development
+of actual language presupposes a higher degree of intelligence than is
+found in any kind of ape. Darwin remarks on this point:[99] "The fact of
+the higher apes not using their vocal organs for speech no doubt depends on
+their intelligence not having been sufficiently advanced."
+
+The sense of beauty, too, has been alleged to be peculiar to man. In
+refutation of this assertion Darwin points to the decorative colours
+of birds, which are used for display. And to the last objection, that
+man alone has religion, that he alone has a belief in God, it is
+answered "that numerous races have existed, and still exist, who have
+no idea of one or more gods, and who have no words in their languages
+to express such an idea."[100]
+
+The result of the investigations recorded in this chapter is to show
+that, great as the difference in mental powers between man and the
+higher animals may be, it is undoubtedly only a difference "of degree
+and not of kind."[101]
+
+In the fourth chapter Darwin deals with the _moral sense_ or
+_conscience_, which is the most important of all differences between
+man and animals. It is a result of social instincts, which lead to
+sympathy for other members of the same society, to non-egoistic
+actions for the good of others. Darwin shows that social tendencies
+are found among many animals, and that among these love and
+kin-sympathy exist, and he gives examples of animals (especially dogs)
+which may exhibit characters that we should call moral in man (e.g.
+disinterested self-sacrifice for the sake of others). The early
+ape-like progenitors of the human race were undoubtedly social. With
+the increase of intelligence the moral sense develops farther; with
+the acquisition of speech public opinion arises, and finally, moral
+sense becomes habit. The rest of Darwin's detailed discussions on
+moral philosophy may be passed over.
+
+The fifth chapter may be very briefly summarised. In it Darwin shows
+that the intellectual and moral faculties are perfected through
+natural selection. He inquires how it can come about that a tribe at a
+low level of evolution attains to a higher, although the best and
+bravest among them often pay for their fidelity and courage with their
+lives without leaving any descendants. In this case it is the
+sentiment of glory, praise and blame, the admiration of others, which
+bring about the increase of the better members of the tribe. Property,
+fixed dwellings, and the association of families into a community are
+also indispensable requirements for civilisation. In the longer second
+section of the fifth chapter Darwin acts mainly as recorder. On the
+basis of numerous investigations, especially those of Greg, Wallace,
+and Galton, he inquires how far the influence of natural selection can
+be demonstrated in regard to civilised nations. In the final section,
+which deals with the proofs that all civilised nations were once
+barbarians, Darwin again uses the results gained by other
+investigators, such as Lubbock and Tylor. There are two sets of facts
+which prove the proposition in question. In the first place, we find
+traces of a former lower state in the customs and beliefs of all
+civilised nations, and in the second place, there are proofs to show
+that savage races are independently able to raise themselves a few
+steps in the scale of civilisation, and that they have thus raised
+themselves.
+
+In the sixth chapter of the work, Morphology comes into the foreground
+once more. Darwin first goes back, however, to the argument based on
+the great difference between the mental powers of the highest animals
+and those of man. That this is only quantitative, not qualitative, he
+has already shown. Very instructive in this connection is the
+reference to the enormous difference in mental powers in another
+class. No one would draw from the fact that the cochineal insect
+(Coccus) and the ant exhibit enormous differences in their mental
+powers, the conclusion that the ant should therefore be regarded as
+something quite distinct, and withdrawn from the class of insects
+altogether.
+
+Darwin next attempts to establish the _specific_ genealogical tree of
+man, and carefully weighs the differences and resemblances between the
+different families of the Primates. The erect position of man is an
+adaptive character, just as are the various characters referable to
+aquatic life in the seals, which, notwithstanding these, are ranked as
+a mere family of the carnivores. The following utterance is very
+characteristic of Darwin:[102] "If man had not been his own
+classifier, he would never have thought of founding a separate order
+for his own reception." In numerous characters not mentioned in
+systematic works, in the features of the face, in the form of the
+nose, in the structure of the external ear, man resembles the apes.
+The arrangement of the hair in man has also much in common with the
+apes; as also the occurrence of hair on the forehead of the human
+embryo, the beard, the convergence of the hair of the upper and under
+arm towards the elbow, which occurs not only in the anthropoid apes,
+but also in some American monkeys. Darwin here adopts Wallace's
+explanation of the origin of the ascending direction of the hair in
+the forearm of the orang,--that it has arisen through the habit of
+holding the hands over the head in rain. But this explanation cannot
+be maintained when we consider that this disposition of the hair is
+widely distributed among the most different mammals, being found in
+the dog, in the sloth, and in many of the lower monkeys.
+
+After further careful analysis of the anatomical characters Darwin
+reaches the conclusion that the New World monkeys (Platyrrhine) may be
+excluded from the genealogical tree altogether, but that man is an
+offshoot from the Old World monkeys (Catarrhine) whose progenitors
+existed as far back as the Miocene period. Among these Old World
+monkeys the forms to which man shows the greatest resemblance are the
+anthropoid apes, which, like him, possess neither tail nor ischial
+callosities. The platyrrhine and catarrhine monkeys have their
+primitive ancestor among extinct forms of the Lemuridae. Darwin also
+touches on the question of the original home of the human race and
+supposes that it may have been in Africa, because it is there that
+man's nearest relatives, the gorilla and the chimpanzee, are found.
+But he regards speculation on this point as useless. It is remarkable
+that, in this connection, Darwin regards the loss of the hair-covering
+in man as having some relation to a warm climate, while elsewhere he
+is inclined to make sexual selection responsible for it. Darwin
+recognises the great gap between man and his nearest relatives, but
+similar gaps exist at other parts of the mammalian genealogical tree:
+the allied forms have become extinct. After the extermination of the
+lower races of mankind, on the one hand, and of the anthropoid apes on
+the other, which will undoubtedly take place, the gulf will be greater
+than ever, since the baboons will then bound it on the one side, and
+the white races on the other. Little weight need be attached to the
+lack of fossil remains to fill up this gap, since the discovery of
+these depends upon chance. The last part of the chapter is devoted to
+a discussion of the earlier stages in the genealogy of man. Here
+Darwin accepts in the main the genealogical tree, which had meantime
+been published by Haeckel, who traces the pedigree back through
+Monotrems, Reptiles, Amphibians, and Fishes, to Amphioxus.
+
+Then follows an attempt to reconstruct, from the atavistic characters,
+a picture of our primitive ancestor who was undoubtedly an arboreal
+animal. The occurrence of rudiments of parts in one sex which only
+come to full development in the other is next discussed. This state of
+things Darwin regards as derived from an original hermaphroditism. In
+regard to the mammary glands of the male he does not accept the theory
+that they are vestigial, but considers them rather as not fully
+developed.
+
+The last chapter of Part I deals with the question whether the
+different races of man are to be regarded as different species, or as
+sub-species of a race of monophyletic origin. The striking differences
+between the races are first emphasised, and the question of the
+fertility or infertility of hybrids is discussed. That fertility is
+the more usual is shown by the excessive fertility of the hybrid
+population of Brazil. This, and the great variability of the
+distinguishing characters of the different races, as well as the fact
+that all grades of transition stages are found between these, while
+considerable general agreement exists, tell in favour of the unity of
+the races and lead to the conclusion that they all had a common
+primitive ancestor.
+
+Darwin therefore classifies all the different races as sub-species of
+_one and the same species_. Then follows an interesting inquiry into
+the reasons for the extinction of human races. He recognises as the
+ultimate reason the injurious effects of a change of the conditions of
+life, which may bring about an increase in infantile mortality, and a
+diminished fertility. It is precisely the reproductive system, among
+animals also, which is most susceptible to changes in the environment.
+
+The final section of this chapter deals with the formation of the
+races of mankind. Darwin discusses the question how far the direct
+effect of different conditions of life, or the inherited effects of
+increased use or disuse may have brought about the characteristic
+differences between the different races. Even in regard to the origin
+of the colour of the skin he rejects the transmitted effects of an
+original difference of climate as an explanation. In so doing he is
+following his tendency to exclude Lamarckian explanations as far as
+possible. But here he makes gratuitous difficulties from which, since
+natural selection fails, there is no escape except by bringing in the
+principle of sexual selection, to which, he regarded it as possible,
+skin-colouring, arrangement of hair, and form of features might be
+traced. But with his characteristic conscientiousness he guards
+himself thus: "I do not intend to assert that sexual selection will
+account for all the differences between the races."[103]
+
+I may be permitted a remark as to Darwin's attitude towards Lamarck.
+While, at an earlier stage, when he was engaged in the preliminary
+labours for his immortal work, _The Origin of Species_, Darwin
+expresses himself very forcibly against the views of Lamarck, speaking
+of Lamarckian "nonsense,"[104] and of Lamarck's "absurd, though clever
+work"[105] and expressly declaring, "I attribute very little to the
+direct action of climate, etc."[106] yet in later life he became more
+and more convinced of the influence of external conditions. In 1876,
+that is, two years after the appearance of the second edition of _The
+Descent of Man_, he writes with his usual candid honesty: "In my
+opinion the greatest error which I have committed, has been not
+allowing sufficient weight to the direct action of the environment,
+i.e. food, climate, etc. independently of a natural selection."[107]
+It is certain from this change of opinion that, if he had been able to
+make up his mind to issue a third edition of _The Descent of Man_, he
+would have ascribed a much greater influence to the effect of
+external conditions in explaining the different characters of the
+races of man than he did in the second edition. He would also
+undoubtedly have attributed less influence to sexual selection as a
+factor in the origin of the different bodily characteristics, if
+indeed he would not have excluded it altogether.
+
+In Part III of the _Descent_ two additional chapters are devoted to
+the discussion of sexual selection in relation to man. These may be
+very briefly referred to. Darwin here seeks to show that sexual
+selection has been operative on man and his primitive progenitor.
+Space fails me to follow out his interesting arguments. I can only
+mention that he is inclined to trace back hairlessness, the
+development of the beard in man, and the characteristic colour of the
+different human races to sexual selection. Since bareness of the skin
+could be no advantage, but rather a disadvantage, this character
+cannot have been brought about by natural selection. Darwin also
+rejected a direct influence of climate as a cause of the origin of the
+skin-colour. I have already expressed the opinion, based on the
+development of his views as shown in his letters, that in a third
+edition Darwin would probably have laid more stress on the influence
+of external environment. He himself feels that there are gaps in his
+proofs here, and says in self-criticism: "The views here advanced, on
+the part which sexual selection has played in the history of man, want
+scientific precision."[108] I need here only point out that it is
+impossible to explain the graduated stages of skin-colour by sexual
+selection, since it would have produced races sharply defined by their
+colour and not united to other races by transition stages, and this,
+it is well known, is not the case. Moreover, the fact established by
+me,[109] that in all races the ventral side of the trunk is paler than
+the dorsal side, and the inner surface of the extremities paler than
+the outer side, cannot be explained by sexual selection in the
+Darwinian sense.
+
+With this I conclude my brief survey of the rich contents of Darwin's
+book. I may be permitted to conclude by quoting the magnificent final
+words of _The Descent of Man_: "We must, however, acknowledge, as it
+seems to me, that man, with all his noble qualities, with sympathy
+which feels for the most debased, with benevolence which extends not
+only to other men but to the humblest living creature, with his
+god-like intellect which has penetrated into the movements and
+constitution of the solar system--with all these exalted powers--Man
+still bears in his bodily frame the indelible stamp of his lowly
+origin."[110]
+
+What has been the fate of Darwin's doctrines since his great
+achievement? How have they been received and followed up by the
+scientific and lay world? And what do the successors of the mighty
+hero and genius think now in regard to the origin of the human race?
+
+At the present time we are incomparably more favourably placed than
+Darwin was for answering this question of all questions. We have at
+our command an incomparably greater wealth of material than he had at
+his disposal. And we are more fortunate than he in this respect, that
+we now know transition-forms which help to fill up the gap, still
+great, between the lowest human races and the highest apes. Let us
+consider for a little the more essential additions to our knowledge
+since the publication of _The Descent of Man_.
+
+Since that time our knowledge of animal embryos has increased
+enormously. While Darwin was obliged to content himself with comparing
+a human embryo with that of a dog, there are now available the
+youngest embryos of monkeys of all possible groups (Orang, Gibbon,
+Semnopithecus, Macacus), thanks to Selenka's most successful tour in
+the East Indies in search of such material. We can now compare
+corresponding stages of the lower monkeys and of the Anthropoid apes
+with human embryos, and convince ourselves of their great resemblance
+to one another, thus strengthening enormously the armour prepared by
+Darwin in defence of his view on man's nearest relatives. It may be
+said that Selenka's material fills up the blanks in Darwin's array of
+proofs in the most satisfactory manner.
+
+The deepening of our knowledge of comparative anatomy also gives us much
+surer foundations than those on which Darwin was obliged to build. Just of
+late there have been many workers in the domain of the anatomy of apes and
+lemurs, and their investigations extend to the most different organs. Our
+knowledge of fossil apes and lemurs has also become much wider and more
+exact since Darwin's time: the fossil lemurs have been especially worked up
+by Cope, Forsyth Major, Ameghino, and others. Darwin knew very little about
+fossil monkeys. He mentions two or three anthropoid apes as occurring in
+the Miocene of Europe,[111] but only names _Dryopithecus_, the largest form
+from the Miocene of France. It was erroneously supposed that this form was
+related to _Hylobates_. We now know not only a form that actually stands
+near to the gibbon (_Pliopithecus_), and remains of other anthropoids
+(_Pliohylobates_ and the fossil chimpanzee, _Palaeopithecus_), but also
+several lower catarrhine monkeys, of which _Mesopithecus_, a form nearly
+related to the modern Sacred Monkeys (a species of _Semnopithecus_) and
+found in strata of the Miocene period in Greece, is the most important.
+Quite recently, too, Ameghino's investigations have made us acquainted with
+fossil monkeys from South America (_Anthropops_, _Homunculus_), which,
+according to their discoverer, are to be regarded as in the line of human
+descent.
+
+What Darwin missed most of all--intermediate forms between apes and
+man--has been recently furnished. E. Dubois, as is well known,
+discovered in 1893, near Trinil in Java, in the alluvial deposits of
+the river Bengawan, an important form represented by a skull-cap, some
+molars, and a femur. His opinion--much disputed as it has been--that
+in this form, which he named _Pithecanthropus_, he has found a
+long-desired transition-form is shared by the present writer. And
+although the geological age of these fossils, which, according to
+Dubois, belong to the uppermost Tertiary series, the Pliocene has
+recently been fixed at a later date (the older Diluvium), the
+_morphological value_ of these interesting remains, that is, the
+intermediate position of _Pithecanthropus_, still holds good. Volz
+says with justice,[112] that even if _Pithecanthropus_ is not _the_
+missing link, it is undoubtedly _a_ missing link.
+
+As on the one hand there has been found in _Pithecanthropus_ a form
+which, though intermediate between apes and man, is nevertheless more
+closely allied to the apes, so on the other hand, much progress has
+been made since Darwin's day in the discovery and description of the
+oldest human remains. Since the famous roof of a skull and the bones
+of the extremities belonging to it were found in 1856 in the
+Neandertal near Duesseldorf, the most varied judgments have been
+expressed in regard to the significance of the remains and of the
+skull in particular. In Darwin's _Descent of Man_ there is only a
+passing allusion to them[113] in connection with the discussion of the
+skull-capacity, although the investigations of Schaaffhausen, King,
+and Huxley were then known. I believe I have shown, in a series of
+papers, that the skull in question belongs to a form different from
+any of the races of man now living, and, with King and Cope, I regard
+it as at least a different species from living man, and have therefore
+designated it _Homo primigenius_. The form unquestionably belongs to
+the older Diluvium, and in the later Diluvium human forms already
+appear, which agree in all essential points with existing human races.
+
+As far back as 1886 the value of the Neandertal skull was greatly
+enhanced by Fraipont's discovery of two skulls and skeletons from Spy
+in Belgium. These are excellently described by their discoverer,[114]
+and are regarded as belonging to the same group of forms as the
+Neandertal remains. In 1899 and the following years came the discovery
+by Gorjanovic-Kramberger of different skeletal parts of at least
+ten individuals in a cave near Krapina in Croatia.[115] It is in
+particular the form of the lower jaw which is different from that of
+all recent races of man, and which clearly indicates the lowly
+position of _Homo primigenius_, while, on the other hand, the
+long-known skull from Gibraltar, which I[116] have referred to _Homo
+primigenius_, and which has lately been examined in detail by
+Sollas,[117] has made us acquainted with the surprising shape of the
+eye-orbit, of the nose, and of the whole upper part of the face.
+Isolated lower jaws found at La Naulette in Belgium, and at Malarnaud
+in France, increase our material which is now as abundant as could be
+desired. The most recent discovery of all is that of a skull dug up in
+August of this year [1908] by Klaatsch and Hauser in the lower grotto
+of the Le Moustier in Southern France, but this skull has not yet been
+fully described. Thus _Homo primigenius_ must also be regarded as
+occupying a position in the gap existing between the highest apes and
+the lowest human races, _Pithecanthropus_, standing in the lower part
+of it, and _Homo primigenius_ in the higher, near man. In order to
+prevent misunderstanding, I should like here to emphasise that in
+arranging this structural series--anthropoid apes, _Pithecanthropus_,
+_Homo primigenius_, _Homo sapiens_--I have no intention of
+establishing it as a direct genealogical series. I shall have
+something to say in regard to the genetic relations of these forms,
+one to another, when discussing the different theories of descent
+current at the present day.[118]
+
+In quite a different domain from that of morphological relationship,
+namely in the physiological study of the blood, results have recently
+been gained which are of the highest importance to the doctrine of
+descent. Uhlenhuth, Nuttall, and others have established the fact that
+the blood-serum of a rabbit which has previously had human blood
+injected into it, forms a precipitate with human blood. This
+biological reaction was tried with a great variety of mammalian
+species, and it was found that those far removed from man gave no
+precipitate under these conditions. But as in other cases among
+mammals all nearly related forms yield an almost equally marked
+precipitate, so the serum of a rabbit treated with human blood and
+then added to the blood of an anthropoid ape gives _almost_ as marked
+a precipitate as in human blood; the reaction to the blood of the
+lower Eastern monkeys is weaker, that to the Western monkeys weaker
+still; indeed in this last case there is only a slight clouding after
+a considerable time and no actual precipitate. The blood of the
+Lemuridae (Nuttall) gives no reaction or an extremely weak one, that
+of the other mammals none whatever. We have in this not only a proof
+of the literal blood relationship between man and apes, but the degree
+of relationship with the different main groups of apes can be
+determined beyond possibility of mistake.
+
+Finally, it must be briefly mentioned that in regard to remains of
+human handicraft also, the material at our disposal has greatly
+increased of late years, that, as a result of this, the opinions of
+archaeologists have undergone many changes, and that, in particular,
+their views in regard to the age of the human race have been greatly
+influenced. There is a tendency at the present time to refer the
+origin of man back to Tertiary times. It is true that no remains of
+Tertiary man have been found, but flints have been discovered which,
+according to the opinion of most investigators, bear traces either of
+use, or of very primitive workmanship. Since Rutot's time, following
+Mortillet's example, investigators have called these "eoliths," and
+they have been traced back by Verworn to the Miocene of the Auvergne,
+and by Rutot even to the upper Oligocene. Although these eoliths are
+even nowadays the subject of many different views, the preoccupation
+with them has kept the problem of the age of the human race
+continually before us.
+
+Geology, too, has made great progress since the days of Darwin and
+Lyell, and has endeavoured with satisfactory results to arrange the
+human remains of the Diluvial period in chronological order (Penck). I
+do not intend to enter upon the question of the primitive home of the
+human race; since the space at my disposal will not allow of my
+touching even very briefly upon all the departments of science which
+are concerned in the problem of the descent of man. How Darwin would
+have rejoiced over each of the discoveries here briefly outlined! What
+use he would have made of the new and precious material, which would
+have prevented the discouragement from which he suffered when
+preparing the second edition of _The Descent of Man_! But it was not
+granted to him to see this progress towards filling up the gaps in his
+edifice of which he was so painfully conscious.
+
+He did, however, have the satisfaction of seeing his ideas steadily
+gaining ground, notwithstanding much hostility and deep-rooted
+prejudice. Even in the years between the appearance of _The Origin of
+Species_ and of the first edition of the _Descent_, the idea of a
+natural descent of man, which was only briefly indicated in the work
+of 1859, had been eagerly welcomed in some quarters. It has been
+already pointed out how brilliantly Huxley contributed to the defence
+and diffusion of Darwin's doctrines, and how in _Man's Place in
+Nature_ he has given us a classic work as a foundation for the
+doctrine of the descent of man. As Huxley was Darwin's champion in
+England, so in Germany Carl Vogt, in particular, made himself master
+of the Darwinian ideas. But above all it was Haeckel who, in energy,
+eagerness for battle, and knowledge may be placed side by side with
+Huxley, who took over the leadership in the controversy over the new
+conception of the universe. As far back as 1866, in his _Generelle
+Morphologie_, he had inquired minutely into the question of the
+descent of man, and not content with urging merely the general theory
+of descent from lower animal forms, he drew up for the first time
+genealogical trees showing the close structural relationships of the
+different animal groups; the last of these illustrated the
+relationships of Mammals, and among them of all groups of the
+Primates, including man. It was Haeckel's genealogical trees that
+formed the basis of the special discussion of the relationships of
+man, in the sixth chapter of Darwin's _Descent of Man_.
+
+In the last section of this essay I shall return to Haeckel's
+conception of the special descent of man, the main features of which
+he still upholds, and rightly so. Haeckel has contributed more than
+any one else to the spread of the Darwinian doctrine.
+
+I can only allow myself a few words as to the spread of the theory of
+the natural descent of man in other countries. The Parisian
+anthropological school, founded and guided by the genius of Broca,
+took up the idea of the descent of man, and made many notable
+contributions to it (Broca, Manouvrier, Mahoudeau, Deniker and
+others). In England itself Darwin's work did not die. Huxley took care
+of that, for he, with his lofty and unprejudiced mind, dominated and
+inspired English biology until his death on June 29, 1895. He had the
+satisfaction shortly before his death of learning of Dubois'
+discovery, which he illustrated by a humourous sketch.[119] But there
+are still many followers in Darwin's footsteps in England. Keane has
+worked at the special genealogical tree of the Primates; Keith has
+inquired which of the anthropoid apes has the greatest number of
+characters in common with man; Morris concerns himself with the
+evolution of man in general, especially with his acquisition of the
+erect position. The recent discoveries of _Pithecanthropus_ and _Homo
+primigenius_ are being vigorously discussed; but the present writer is
+not in a position to form an opinion of the extent to which the idea
+of descent has penetrated throughout England generally.
+
+In Italy independent work in the domain of the descent of man is being
+produced, especially by Morselli; with him are associated, in the
+investigation of related problems, Sergi and Giuffrida-Ruggeri. From
+the ranks of American investigators we may single out in particular
+the eminent geologist Cope, who championed with much decision the idea
+of the specific difference of _Homo neandertalensis_ (_primigenius_)
+and maintained a more direct descent of man from the fossil Lemuridae.
+In South America too, in Argentina, new life is stirring in this
+department of science. Ameghino in Buenos Ayres has awakened the
+fossil primates of the Pampas formation to new life; he even believes
+that in his _Tetraprothomo_, represented by a femur, he has discovered
+a direct ancestor of man. Lehmann-Nitsche is working at the other side
+of the gulf between apes and man, and he describes a remarkable first
+cervical vertebra (atlas) from Monte Hermoso as belonging to a form
+which may bear the same relation to _Homo sapiens_ in South America as
+_Homo primigenius_ does in the Old World. After a minute investigation
+he establishes a human species _Homo neogaeus_, while Ameghino
+ascribes this atlas vertebra to his _Tetraprothomo_.
+
+Thus throughout the whole scientific world there is arising a new
+life, an eager endeavour to get nearer to Huxley's _problema maximum_,
+to penetrate more deeply into the origin of the human race. There are
+to-day very few experts in anatomy and zoology who deny the animal
+descent of man in general. Religious considerations, old prejudices,
+the reluctance to accept man, who so far surpasses mentally all other
+creatures, as descended from "soulless" animals, prevent a few
+investigators from giving full adherence to the doctrine. But there
+are very few of these who still postulate a special act of creation
+for man. Although the majority of experts in anatomy and zoology
+accept unconditionally the descent of man from lower forms, there is
+much diversity of opinion among them in regard to the special line of
+descent.
+
+In trying to establish any special hypothesis of descent, whether by
+the graphic method of drawing up genealogical trees or otherwise, let
+us always bear in mind Darwin's words[120] and use them as a critical
+guiding line: "As we have no record of the lines of descent, the
+pedigree can be discovered only by observing the degrees of
+resemblance between the beings which are to be classed." Darwin
+carries this further by stating "that resemblances in several
+unimportant structures, in useless and rudimentary organs, or not now
+functionally active, or in an embryological condition, are by far the
+most serviceable for classification."[121] It has also to be
+remembered that _numerous_ separate points of agreement are of much
+greater importance than the amount of similarity or dissimilarity in a
+few points.
+
+The hypotheses as to descent current at the present day may be divided
+into two main groups. The first group seeks for the roots of the human
+race not among any of the families of the apes--the anatomically
+nearest forms--nor among their very similar but less specialised
+ancestral forms, the fossil representatives of which we can know only
+in part, but, setting the monkeys on one side, it seeks for them lower
+down among the fossil Eocene Pseudo-lemuridae or Lemuridae (Cope), or
+even among the primitive pentadactylous Eocene forms, which may
+either have led directly to the evolution of man (Adloff), or have
+given rise to an ancestral form common to apes and men (Klaatsch,[122]
+Giuffrida-Ruggeri). The common ancestral form, from which man and apes
+are thus supposed to have arisen independently, may explain the
+numerous resemblances which actually exist between them. That is to
+say, all the characters upon which the great structural resemblance
+between apes and man depends must have been present in their common
+ancestor. Let us take an example of such a common character. The bony
+external ear-passage is in general as highly developed in the lower
+Eastern monkeys and the anthropoid apes as in man. This character
+must, therefore, have already been present in the common primitive
+form. In that case it is not easy to understand why the Western
+monkeys have not also inherited the character, instead of possessing
+only a tympanic ring. But it becomes more intelligible if we assume
+that forms with a primitive tympanic ring were the original type, and
+that from these were evolved, on the one hand, the existing New World
+monkeys with persistent tympanic ring, and on the other an ancestral
+form common to the lower Old World monkeys, the anthropoid apes and
+man. For man shares with these the character in question, and it is
+also one of the "unimportant" characters required by Darwin. Thus we
+have two divergent lines arising from the ancestral form, the Western
+monkeys (Platyrrhine) on the one hand, and an ancestral form common to
+the lower Eastern monkeys, the anthropoid apes, and man, on the other.
+But considerations similar to those which showed it to be impossible
+that man should have developed from an ancestor common to him and the
+monkeys, yet outside of and parallel with these, may be urged also
+against the likelihood of a parallel evolution of the lower Eastern
+monkeys, the anthropoid apes, and man. The anthropoid apes have in
+common with man many characters which are not present in the lower
+Old World monkeys. These characters must therefore have been present
+in the ancestral form common to the three groups. But here, again, it
+is difficult to understand why the lower Eastern monkeys should not
+also have inherited these characters. As this is not the case, there
+remains no alternative but to assume divergent evolution from an
+indifferent form. The lower Eastern monkeys are carrying on the
+evolution in one direction--I might almost say towards a blind
+alley--while anthropoids and men have struck out a progressive path,
+at first in common, which explains the many points of resemblance
+between them, without regarding man as derived directly from the
+anthropoids. Their many striking points of agreement indicate a common
+descent, and cannot be explained as phenomena of convergence.
+
+I believe I have shown in the above sketch that a theory which derives
+man directly from lower forms without regarding apes as
+transition-types leads _ad absurdum_. The close structural
+relationship between man and monkeys can only be understood if both
+are brought into the same line of evolution. To trace man's line of
+descent directly back to the old Eocene mammals, alongside of, but
+with no relation to these very similar forms, is to abandon the method
+of exact comparison, which, as Darwin rightly recognised, alone
+justifies us in drawing up genealogical trees on the basis of
+resemblances and differences. The farther down we go the more does the
+ground slip from beneath our feet. Even the Lemuridae show very
+numerous divergent conditions, much more so the Eocene mammals
+(Creodonta, Condylarthra), the chief resemblance of which to man
+consists in the possession of pentadactylous hands and feet! Thus the
+farther course of the line of descent disappears in the darkness of
+the ancestry of the mammals. With just as much reason we might pass by
+the Vertebrates altogether, and go back to the lower Invertebrates,
+but in that case it would be much easier to say that man has arisen
+independently, and has evolved, without relation to any animals, from
+the lowest primitive form to his present isolated and dominant
+position. But this would be to deny all value to classification, which
+must after all be the ultimate basis of a genealogical tree. We can,
+as Darwin rightly observed, only infer the line of descent from the
+degree of resemblance between single forms. If we regard man as
+directly derived from primitive forms very far back, we have no way of
+explaining the many points of agreement between him and the monkeys in
+general, and the anthropoid apes in particular. These must remain an
+inexplicable marvel.
+
+I have thus, I trust, shown that the first class of special theories
+of descent, which assumes that man has developed, parallel with the
+monkeys, but without relation to them, from very low primitive forms
+cannot be upheld, because it fails to take into account the close
+structural affinity of man and monkeys. I cannot but regard this
+hypothesis as lamentably retrograde, for it makes impossible any
+application of the facts that have been discovered in the course of
+the anatomical and embryological study of man and monkeys, and indeed
+prejudges investigations of that class as pointless. The whole method
+is perverted; an unjustifiable theory of descent is first formulated
+with the aid of the imagination, and then we are asked to declare that
+all structural relations between man and monkeys, and between the
+different groups of the latter, are valueless,--the fact being that
+they are the only true basis on which a genealogical tree can be
+constructed.
+
+So much for this most modern method of classification, which has
+probably found adherents because it would deliver us from the
+relationship to apes which many people so much dislike. In contrast to
+it we have the second class of special hypotheses of descent, which
+keeps strictly to the nearest structural relationship. This is the
+only basis that justifies the drawing up of a special hypothesis of
+descent. If this fundamental proposition be recognised, it will be
+admitted that the doctrine of special descent upheld by Haeckel, and
+set forth in Darwin's _Descent of Man_, is still valid to-day. In the
+genealogical tree, man's place is quite close to the anthropoid apes;
+these again have as their nearest relatives the lower Old World
+monkeys, and their progenitors must be sought among the less
+differentiated Platyrrhine monkeys, whose most important characters
+have been handed on to the present day New World monkeys. How the
+different genera are to be arranged within the general scheme
+indicated depends in the main on the classificatory value attributed
+to individual characters. This is particularly true in regard to
+_Pithecanthropus_, which I consider as the root of a branch which has
+sprung from the anthropoid ape root and has led up to man; the latter
+I have designated the family of the Hominidae.
+
+For the rest, there are, as we have said, various possible ways of
+constructing the narrower genealogy within the limits of this branch
+including men and apes, and these methods will probably continue to
+change with the accumulation of new facts. Haeckel himself has
+modified his genealogical tree of the Primates in certain details
+since the publication of his _Generelle Morphologie_ in 1866, but its
+general basis remains the same.[123] All the special genealogical
+trees drawn up on the lines laid down by Haeckel and Darwin--and that
+of Dubois may be specially mentioned--are based, in general, on the
+close relationship of monkeys and men, although they may vary in
+detail. Various hypotheses have been formulated on these lines, with
+special reference to the evolution of man. _Pithecanthropus_ is
+regarded by some authorities as the direct ancestor of man, by others
+as a side-track failure in the attempt at the evolution of man. The
+problem of the monophyletic or polyphyletic origin of the human race
+has also been much discussed. Sergi[124] inclines towards the
+assumption of a polyphyletic origin of the three main races of man,
+the African primitive form of which has given rise also to the
+gorilla and chimpanzee, the Asiatic to the Orang, the Gibbon, and
+_Pithecanthropus_. Kollmann regards existing human races as derived
+from small primitive races (pigmies), and considers that _Homo
+primigenius_ must have arisen in a secondary and degenerative manner.
+
+But this is not the place, nor have I the space to criticise the
+various special theories of descent. One, however, must receive
+particular notice. According to Ameghino, the South American monkeys
+(_Pitheculites_) from the oldest Tertiary of the Pampas are the forms
+from which have arisen the existing American monkeys on the one hand,
+and on the other, the extinct South American Homunculidae, which are
+also small forms. From these last, anthropoid apes and man have, he
+believes, been evolved. Among the progenitors of man, Ameghino reckons
+the form discovered by him (_Tetraprothomo_), from which a South
+American primitive man, _Homo pampaeus_, might be directly evolved,
+while on the other hand all the lower Old World monkeys may have
+arisen from older fossil South American forms (Clenialitidae), the
+distribution of which may be explained by the bridge formerly existing
+between South America and Africa, as may be the derivation of all
+existing human races from _Homo pampaeus_.[125] The fossil forms
+discovered by Ameghino deserve the most minute investigation, as does
+also the fossil man from South America of which Lehmann-Nitsche[126]
+has made a thorough study.
+
+It is obvious that, notwithstanding the necessity for fitting man's
+line of descent into the genealogical tree of the Primates, especially
+the apes, opinions in regard to it differ greatly in detail. This
+could not be otherwise, since the different Primate forms, especially
+the fossile forms, are still far from being exhaustively known. But
+one thing remains certain,--the idea of the close relationship between
+man and monkeys set forth in Darwin's _Descent of Man_. Only those who
+deny the many points of agreement, the sole basis of classification,
+and thus of a natural genealogical tree, can look upon the position of
+Darwin and Haeckel as antiquated, or as standing on an insufficient
+foundation. For such a genealogical tree is nothing more than a
+summarised representation of what is known in regard to the degree of
+resemblance between the different forms.
+
+Darwin's work in regard to the descent of man has not been surpassed;
+the more we immerse ourselves in the study of the structural
+relationships between apes and man, the more is our path illumined by
+the clear light radiating from him, and through his calm and
+deliberate investigation, based on a mass of material in the
+accumulation of which he has never had an equal. Darwin's fame will be
+bound up for all time with the unprejudiced investigation of the
+question of all questions, the descent of the human race.
+
+FOOTNOTES:
+
+[Footnote 75: _Life and Letters of Thomas Henry Huxley_, Vol. I. p.
+171, London, 1900.]
+
+[Footnote 76: _Ibid._, p. 363.]
+
+[Footnote 77: No italics in original.]
+
+[Footnote 78: _Life and Letters of Charles Darwin_, Vol. I. p. 93.]
+
+[Footnote 79: _Ibid._ Vol. II. p. 263.]
+
+[Footnote 80: _Ibid._ Vol. I. p. 94.]
+
+[Footnote 81: _Life and Letters_, Vol. III. p. 175.]
+
+[Footnote 82: _Ibid._ Vol. II. p. 109.]
+
+[Footnote 83: _Ibid._ Vol. III. p. 112.]
+
+[Footnote 84: _Ibid._ Vol. I. pp. 304-317.]
+
+[Footnote 85: _Life and Letters_, Vol. I. p. 309.]
+
+[Footnote 86: _Loc. cit._ p. 313.]
+
+[Footnote 87: _Ibid._ Vol. II. p. 310.]
+
+[Footnote 88: _Ibid._ Vol. III. p. 236. ["C. Ridley," Mr. Francis
+Darwin points out to me, should be H. N. Ridley. A.C.S.]]
+
+[Footnote 89: _Descent of Man_ (Popular Edit., 1901), fig. 1, p. 14.]
+
+[Footnote 90: G. Schwalbe, "Das Darwin'sche Spitzohr beim menschlichen
+Embryo," _Anatom. Anzeiger_, 1889, pp. 176-189, and other papers.]
+
+[Footnote 91: _Descent of Man_, fig. 3, p. 24.]
+
+[Footnote 92: _Descent of man_, p. 6.]
+
+[Footnote 93: _Ibid._ p. 54.]
+
+[Footnote 94: _Descent of Man_, p. 92.]
+
+[Footnote 95: _Ibid._ p. 100.]
+
+[Footnote 96: _Life and letters_, Vol. II. p. 161, June 22, 1859.]
+
+[Footnote 97: _Ibid._ Vol. III. p. 15, March 17, 1863.]
+
+[Footnote 98: _Descent of Man_, p. 132.]
+
+[Footnote 99: _Ibid._ pp. 136, 137.]
+
+[Footnote 100: _Ibid._ p. 143.]
+
+[Footnote 101: _Ibid._ p. 193.]
+
+[Footnote 102: _Descent of Man_, p. 231.]
+
+[Footnote 103: _Descent of Man_, p. 308.]
+
+[Footnote 104: _Life and Letters_, Vol. II. p. 23.]
+
+[Footnote 105: _Loc. cit._ p. 39.]
+
+[Footnote 106: _Loc. cit._ (1856), p. 82.]
+
+[Footnote 107: _Ibid._ Vol. III p. 159.]
+
+[Footnote 108: _Descent of Man_, p. 924.]
+
+[Footnote 109: "Die Hautfarbe des Menschen," _Mitteilungen der
+Anthropologischen Gessellschaft in Wien_, Vol. XXXIV. pp. 331-352.]
+
+[Footnote 110: _Ibid._ p. 947.]
+
+[Footnote 111: _Descent of Man_, p. 240.]
+
+[Footnote 112: "Das geologische Alter der Pithecanthropus-Schichten
+bei Trinil, Ost-Java." _Neues Jahrb. f. Mineralogie_. Festband, 1907.]
+
+[Footnote 113: _Descent of Man_, p. 82.]
+
+[Footnote 114: "La race humaine de Neanderthal ou de Canstatt en
+Belgique." _Arch. de Biologie_, VII. 1887.]
+
+[Footnote 115: Gorjanovic-Kramberger. _Der diluviale Mensch van
+Krapina in Kroatien_, 1906.]
+
+[Footnote 116: _Studien zur Vorgeschichte des Menschen_, 1906, pp. 154
+ff.]
+
+[Footnote 117: "On the cranial and facial characters of the Neandertal
+Race." _Trans. R. Soc._ London, vol. 199, 1908, p. 281.]
+
+[Footnote 118: Since this essay was written Schoetensack has
+discovered near Heidelberg and briefly described an exceedingly
+interesting lower jaw from rocks between the Pliocene and Diluvial
+beds. This exhibits interesting differences from the forms of lower
+jaw of _Homo primigenius_. (Schoetensack, _Der Unterkiefer des Homo
+heidelbergensis_, Leipzig, 1908.) G. S.]
+
+[Footnote 119: _Life and Letters of Thomas Henry Huxley_, Vol. II. p.
+394.]
+
+[Footnote 120: _Descent of Man_, p. 229.]
+
+[Footnote 121: _Loc. cit._]
+
+[Footnote 122: Klaatsch in his last publications speaks in the main
+only of an ancestral form common to men and anthropoid apes.]
+
+[Footnote 123: Haeckels latest genealogical tree is to be found in his
+most recent work, _Unsere Ahnenreihe_. Jena, 1908.]
+
+[Footnote 124: Sergi, G. _Europa_, 1908.]
+
+[Footnote 125: _See_ Ameghino's latest paper, "_Notas preliminaries
+sobre el Tetraprothomo argentinus_," etc. _Anales del Museo nacional
+de Buenos Aires_, XVI. pp. 107-242, 1907.]
+
+[Footnote 126: "Nouvelles recherches sur la formation pampeenne et
+l'homme fossile de la Republique Argentine." _Rivista del Museo de la
+Plata_, T. XIV. pp. 193-488.]
+
+
+
+
+V
+
+CHARLES DARWIN AS AN ANTHROPOLOGIST
+
+BY ERNST HAECKEL
+
+_Professor of Zoology in the University of Jena_
+
+
+The great advance that anthropology has made in the second half of the
+nineteenth century is due, in the first place, to Darwin's discovery
+of the origin of man. No other problem in the whole field of research
+is so momentous as that of "Man's place in nature," which was justly
+described by Huxley (1863) as the most fundamental of all questions.
+Yet the scientific solution of this problem was impossible until the
+theory of descent had been established.
+
+It is now a hundred years since the great French biologist Jean
+Lamarck published his _Philosophie Zoologique_. By a remarkable
+coincidence the year in which that work was issued, 1809, was the year
+of the birth of his most distinguished successor, Charles Darwin.
+Lamarck had already recognised that the descent of man from a series
+of other Vertebrates--that is, from a series of Ape-like Primates--was
+essentially involved in the general theory of transformation which he
+had erected on a broad inductive basis; and he had sufficient
+penetration to detect the agencies that had been at work in the
+evolution of the erect bimanous man from the arboreal and quadrumanous
+ape. He had, however, few empirical arguments to advance in support of
+his hypothesis, and it could not be established until the further
+development of the biological sciences--the founding of comparative
+embryology by Baer (1828) and of the cell-theory by Schleiden and
+Schwann (1838), the advance of physiology under Johannes Mueller
+(1833), and the enormous progress of palaeontology and comparative
+anatomy between 1820 and 1860--provided this necessary foundation.
+Darwin was the first to coordinate the ample results of these lines of
+research. With no less comprehensiveness than discrimination he
+consolidated them as a basis of a modified theory of descent, and
+associated with them his own theory of natural selection, which we
+take to be distinctive of "Darwinism" in the stricter sense. The
+illuminating truth of these cumulative arguments was so great in every
+branch of biology that, in spite of the most vehement opposition, the
+battle was won within a single decade, and Darwin secured the general
+admiration and recognition that had been denied to his forerunner,
+Lamarck, up to the hour of his death (1829).
+
+Before, however, we consider the momentous influence that Darwinism
+has had in anthropology, we shall find it useful to glance at its
+history in the course of the last half century, and notice the various
+theories that have contributed to its advance. The first attempt to
+give extensive expression to the reform of biology by Darwin's work
+will be found in my _Generelle Morphologie_ (1866)[127] which was
+followed by a more popular treatment of the subject in my _Natuerliche
+Schoepfungsgeschichte_ (1868),[128] a compilation from the earlier
+work. In the first volume of the _Generelle Morphologie_ I endeavoured
+to show the great importance of evolution in settling the fundamental
+questions of biological philosophy, especially in regard to
+comparative anatomy. In the second volume I dealt broadly with the
+principle of evolution, distinguishing ontogeny and phylogeny as its
+two coordinate main branches, and associating the two in the
+Biogenetic Law. The Law may be formulated thus: "Ontogeny (embryology
+or the development of the individual) is a concise and compressed
+recapitulation of phylogeny (the palaeontological or genealogical
+series) conditioned by laws of heredity and adaptation." The
+"Systematic introduction to general evolution," with which the second
+volume of the _Generelle Morphologie_ opens, was the first attempt to
+draw up a natural system of organisms (in harmony with the principles
+of Lamarck and Darwin) in the form of a hypothetical pedigree, and was
+provisionally set forth in eight genealogical tables.
+
+In the nineteenth chapter of the _Generelle Morphologie_--a part of
+which has been republished, without any alteration, after a lapse of
+forty years--I made a critical study of Lamarck's theory of descent
+and of Darwin's theory of selection, and endeavoured to bring the
+complex phenomena of heredity and adaptation under definite laws for
+the first time. Heredity I divided into conservative and progressive:
+adaptation into indirect (or potential) and direct (or actual). I then
+found it possible to give some explanation of the correlation of the
+two physiological functions in the struggle for life (selection), and
+to indicate the important laws of divergence (or differentiation) and
+complexity (or division of labor), which are the direct and inevitable
+outcome of selection. Finally, I marked off dysteleology as the
+science of the aimless (vestigial, abortive, atrophied, and useless)
+organs and parts of the body. In all this I worked from a strictly
+monistic standpoint, and sought to explain all biological phenomena on
+the mechanical and naturalistic lines that had long been recognised in
+the study of inorganic nature. Then (1866), as now, being convinced of
+the unity of nature, the fundamental identity of the agencies at work
+in the inorganic and the organic worlds, I discarded vitalism,
+teleology, and all hypotheses of a mystic character.
+
+It was clear from the first that it was essential, in the monistic
+conception of evolution, to distinguish between the laws of
+conservative and progressive heredity. Conservative heredity maintains
+from generation to generation the enduring characters of the species.
+Each organism transmits to its descendants a part of the morphological
+and physiological qualities that it has received from its parents and
+ancestors. On the other hand, progressive heredity brings new
+characters to the species--characters that were not found in preceding
+generations. Each organism may transmit to its offspring a part of the
+morphological and physiological features that it has itself acquired,
+by adaptation, in the course of its individual career, through the use
+or disuse of particular organs, the influence of environment, climate,
+nutrition, etc. At that time I gave the name of "progressive heredity"
+to this inheritance of acquired characters, as a short and convenient
+expression, but have since changed the term to "transformative
+heredity" (as distinguished from conservative). This term is
+preferable, as inherited regressive modifications (degeneration,
+retrograde metamorphosis, etc.) come under the same head.
+
+Transformative heredity--or the transmission of acquired
+characters--is one of the most important principles in evolutionary
+science. Unless we admit it most of the facts of comparative anatomy
+and physiology are inexplicable. That was the conviction of Darwin no
+less than of Lamarck, of Spencer as well as Virchow, of Huxley as well
+as Gegenbaur, indeed of the great majority of speculative biologists.
+This fundamental principle was for the first time called in question
+and assailed in 1885 by August Weismann of Freiburg, the eminent
+zoologist to whom the theory of evolution owes a great deal of
+valuable support, and who has attained distinction by his extension of
+the theory of selection. In explanation of the phenomena of heredity
+he introduced a new theory, the "theory of the continuity of the
+germ-plasm." According to him the living substance in all organisms
+consists of two quite distinct kinds of plasm, somatic and germinal.
+The permanent germ-plasm, or the active substance of the two
+germ-cells (egg-cell and sperm-cell), passes unchanged through a
+series of generations, and is not affected by environmental
+influences. The environment modifies only the soma-plasm, the organs
+and tissues of the body. The modifications that these parts undergo
+through the influence of the environment or their own activity (use
+and habit), do not affect the germ-plasm, and cannot therefore be
+transmitted.
+
+This theory of the continuity of the germ-plasm has been expounded by
+Weismann during the last twenty-four years in a number of able
+volumes, and is regarded by many biologists, such as Mr. Francis
+Galton, Sir E. Ray Lankester, and Professor J. Arthur Thomson (who has
+recently made a thorough-going defence of it in his important work
+_Heredity_),[129] as the most striking advance in evolutionary
+science. On the other hand, the theory has been rejected by Herbert
+Spencer, Sir W. Turner, Gegenbaur, Koelliker, Hertwig, and many others.
+For my part I have, with all respect for the distinguished Darwinian,
+contested the theory from the first, because its whole foundation
+seems to me erroneous, and its deductions do not seem to be in accord
+with the main facts of comparative morphology and physiology.
+Weismann's theory in its entirety is a finely conceived molecular
+hypothesis, but it is devoid of empirical basis. The notion of the
+absolute and permanent independence of the germ-plasm, as
+distinguished from the soma-plasm, is purely speculative; as is also
+the theory of germinal selection. The determinants, ids, and idants,
+are purely hypothetical elements. The experiments that have been
+devised to demonstrate their existence really prove nothing.
+
+It seems to me quite improper to describe this hypothetical structure
+as "Neodarwinism." Darwin was just as convinced as Lamarck of the
+transmission of acquired characters and its great importance in the
+scheme of evolution. I had the good fortune to visit Darwin at Down
+three times and discuss with him the main principles of his system,
+and on each occasion we were fully agreed as to the incalculable
+importance of what I may call transformative inheritance. It is only
+proper to point out that Weismann's theory of the germ-plasm is in
+express contradiction to the fundamental principles of Darwin and
+Lamarck. Nor is it more acceptable in what one may call its
+"ultradarwinism"--the idea that the theory of selection explains
+everything in the evolution of the organic world. This belief in the
+"omnipotence of natural selection" was not shared by Darwin himself.
+Assuredly, I regard it as of the utmost value, as the process of
+natural selection through the struggle for life affords an explanation
+of the mechanical origin of the adapted organisation. It solves the
+great problem: how could the finely adapted structure of the animal or
+plant body be formed unless it was built on a preconceived plan? It
+thus enables us to dispense with the teleology of the metaphysician
+and the dualist, and to set aside the old mythological and poetic
+legends of creation. The idea had occurred in vague form to the great
+Empedocles 2000 years before the time of Darwin, but it was reserved
+for modern research to give it ample expression. Nevertheless, natural
+selection does not of itself give the solution of all our evolutionary
+problems. It has to be taken in conjunction with the transformism of
+Lamarck, with which it is in complete harmony.
+
+The monumental greatness of Charles Darwin, who surpasses every other
+student of science in the nineteenth century by the loftiness of his
+monistic conception of nature and the progressive influence of his
+ideas, is perhaps best seen in the fact that not one of his many
+successors has succeeded in modifying his theory of descent in any
+essential point or in discovering an entirely new standpoint in the
+interpretation of the organic world. Neither Naegeli nor Weismann,
+neither De Vries nor Roux, has done this. Naegeli, in his
+_Mechanisch-Physiologische Theorie der Abstammungslehre_[130] which is
+to a great extent in agreement with Weismann, constructed a theory of
+the idioplasm, that represents it (like the germ-plasm) as developing
+continuously in a definite direction from internal causes. But his
+internal "principle of progress" is at the bottom just as teleological
+as the vital force of the Vitalists, and the micella structure of the
+idioplasm is just as hypothetical as the "dominant" structure of the
+germ-plasm. In 1889 Moritz Wagner sought to explain the origin of
+species by migration and isolation, and on that basis constructed a
+special "migration-theory." This, however, is not out of harmony with
+the theory of selection. It merely elevates one single factor in the
+theory to a predominant position. Isolation is only a special case of
+selection, as I had pointed out in the fifteenth chapter of my
+_Natural history of creation_. The "mutation-theory" of De Vries,[131]
+that would explain the origin of species by sudden and saltatory
+variations rather than by gradual modification, is regarded by many
+botanists as a great step in advance, but it is generally rejected by
+zoologists. It affords no explanation of the facts of adaptation, and
+has no causal value.
+
+Much more important than these theories is that of Wilhelm Roux[132]
+of "the struggle of parts within the organism, a supplementation of
+the theory of mechanical adaptation." He explains the functional
+autoformation of the purposive structure by a combination of Darwin's
+principle of selection with Lamarck's idea of transformative heredity,
+and applies the two in conjunction to the facts of histology. He lays
+stress on the significance of functional adaptation, which I had
+described in 1866, under the head of cumulative adaptation, as the
+most important factor in evolution. Pointing out its influence in the
+cell-life of the tissues, he puts "cellular selection" above "personal
+selection," and shows how the finest conceivable adaptations in the
+structure of the tissue may be brought about quite mechanically,
+without preconceived plan. This "mechanical teleology" is a valuable
+extension of Darwin's monistic principle of selection to the whole
+field of cellular physiology and histology, and is wholly destructive
+of dualistic vitalism.
+
+The most important advance that evolution has made since Darwin and
+the most valuable amplification of his theory of selection is, in my
+opinion, the work of Richard Semon: _Die Mneme als erhaltendes Prinzip
+im Wechsel des organischen Geschehens_.[133] He offers a psychological
+explanation of the facts of heredity by reducing them to a process of
+(unconscious) memory. The physiologist Ewald Hering had shown in 1870
+that memory must be regarded as a general function of organic matter,
+and that we are quite unable to explain the chief vital phenomena,
+especially those of reproduction and inheritance, unless we admit this
+unconscious memory. In my essay _Die Perigenesis der Plastidule_[134]
+I elaborated this far-reaching idea, and applied the physical
+principle of transmitted motion to the plastidules, or active
+molecules of plasm. I concluded that "heredity is the memory of the
+plastidules, and variability their power of comprehension." This
+"provisional attempt to give a mechanical explanation of the
+elementary processes of evolution" I afterwards extended by showing
+that sensitiveness is (as Carl Naegeli, Ernst Mach, and Albrecht Rau
+express it) a general quality of matter. This form of panpsychism
+finds its simplest expression in the "trinity of substance."
+
+To the two fundamental attributes that Spinoza ascribed to
+substance--Extension (matter as occupying space) and Cogitation
+(energy, force)--we now add the third fundamental quality of Psychoma
+(sensitiveness, soul). I further elaborated this trinitarian
+conception of substance in the nineteenth chapter of my _Die
+Lebenswunder_ (1904),[135] and it seems to me well calculated to
+afford a monistic solution of many of the antitheses of philosophy.
+
+This important Mneme-theory of Semon and the luminous physiological
+experiments and observations associated with it not only throw
+considerable light on transformative inheritance, but provide a sound
+physiological foundation for the biogenetic law. I had endeavoured to
+show in 1874, in the first chapter of my _Anthropogenie_,[136] that
+this fundamental law of organic evolution holds good generally, and
+that there is everywhere a direct causal connection between ontogeny
+and phylogeny. "Phylogenesis is the mechanical cause of ontogenesis;"
+in other words, "The evolution of the stem or race is--in accordance
+with the laws of heredity and adaptation--the real cause of all the
+changes that appear, in a condensed form, in the development of the
+individual organism from the ovum, in either the embryo or the larva."
+
+It is now fifty years since Charles Darwin pointed out, in the
+thirteenth chapter of his epoch-making _Origin of Species_, the
+fundamental importance of embryology in connection with his theory of
+descent:
+
+"The leading facts in embryology, which are second to none in
+importance, are explained on the principle of variations in the many
+descendants from some one ancient progenitor, having appeared at a not
+very early period of life, and having been inherited at a
+corresponding period."[137]
+
+He then shows that the striking resemblance of the embryos and larvae
+of closely related animals, which in the mature stage belong to widely
+different species and genera, can only be explained by their descent
+from a common progenitor. Fritz Mueller made a closer study of these
+important phenomena in the instructive instance of the Crustacean
+larva, as given in his able work _Fuer Darwin_[138] (1864). I then, in
+1872, extended the range so as to include all animals (with the
+exception of the unicellular Protozoa) and showed, by means of the
+theory of the Gastraea, that all multicellular, tissue-forming
+animals--all the Metazoa--develop in essentially the same way from the
+primary germ-layers.
+
+I conceived the embryonic form, in which the whole structure consists of
+only two layers of cells, and is known as the gastrula, to be the
+ontogenetic recapitulation, maintained by tenacious heredity, of a
+primitive common progenitor of all the Metazoa, the Gastraea. At a later
+date (1895) Monticelli discovered that this conjectural ancestral form is
+still preserved in certain primitive Coelenterata--Pemmatodiscus,
+Kunstleria, and the nearly-related Orthonectida.
+
+The general application of the biogenetic law to all classes of
+animals and plants has been proved in my _Systematische
+Phylogenie_.[139] It has, however, been frequently challenged, both by
+botanists and zoologists, chiefly owing to the fact that many have
+failed to distinguish its two essential elements, palingenesis and
+cenogenesis. As early as 1874 I had emphasised, in the first chapter
+of my _Evolution of Man_, the importance of discriminating carefully
+between these two sets of phenomena:
+
+"In the evolutionary appreciation of the facts of embryology we must
+take particular care to distinguish sharply and clearly between the
+primary, palingenetic evolutionary processes and the secondary,
+cenogenetic processes. The palingenetic phenomena, or embryonic
+_recapitulations_, are due to heredity, to the transmission of
+characters from one generation to another. They enable us to draw
+direct inferences in regard to corresponding structures in the
+development of the species (e.g. the chorda or the branchial arches in
+all vertebrate embryos). The cenogenetic phenomena, on the other hand,
+or the embryonic _variations_, cannot be traced to inheritance from a
+mature ancestor, but are due to the adaption of the embryo or the
+larva to certain conditions of its individual development (e.g. the
+amnion, the allantois, and the vitelline arteries in the embryos of
+the higher vertebrates). These cenogenetic phenomena are later
+additions; we must not infer from them that there were corresponding
+processes in the ancestral history, and hence they are apt to
+mislead."
+
+The fundamental importance of these facts of comparative anatomy,
+atavism, and the rudimentary organs, was pointed out by Darwin in the
+first part of his classic work, _The Descent of Man and Selection in
+Relation to Sex_ (1871).[140] In the "General summary and conclusion"
+(chap. xxi.) he was able to say, with perfect justice: "He who is not
+content to look, like a savage, at the phenomena of nature as
+disconnected, cannot any longer believe that man is the work of a
+separate act of creation. He will be forced to admit that the close
+resemblance of the embryo of man to that, for instance, of a dog--the
+construction of his skull, limbs, and whole frame on the same plan
+with that of other mammals, independently of the uses to which the
+parts may be put--the occasional reappearance of various structures,
+for instance of several muscles, which man does not normally possess,
+but which are common to the Quadrumana--and a crowd of analogous
+facts--all point in the plainest manner to the conclusion that man is
+the co-descendant with other mammals of a common progenitor."
+
+These few lines of Darwin's have a greater scientific value than
+hundreds of those so-called "anthropological treatises," which give
+detailed descriptions of single organs, or mathematical tables with
+series of numbers and what are claimed to be "exact analyses," but are
+devoid of synoptic conclusions and a philosophical spirit.
+
+Charles Darwin is not generally recognised as a great anthropologist,
+nor does the school of modern anthropologists regard him as a leading
+authority. In Germany, especially, the great majority of the members
+of the anthropological societies took up an attitude of hostility to
+him from the very beginning of the controversy in 1860. _The Descent
+of Man_ was not merely rejected, but even the discussion of it was
+forbidden on the ground that it was "unscientific."
+
+The centre of this inveterate hostility for thirty years--especially
+after 1877--was Rudolph Virchow of Berlin, the leading investigator
+in pathological anatomy, who did so much for the reform of medicine by
+his establishment of cellular pathology in 1858. As a prominent
+representative of "exact" or "descriptive" anthropology, and lacking a
+broad equipment in comparative anatomy and ontogeny, he was unable to
+accept the theory of descent. In earlier years, and especially during
+his splendid period of activity at Wuerzburg (1848-1856), he had been a
+consistent free-thinker, and had in a number of able articles
+(collected in his _Gesammelte Abhandlungen_)[141] upheld the unity of
+human nature, the inseparability of body and spirit. In later years at
+Berlin, where he was more occupied with political work and sociology
+(especially after 1866), he abandoned the positive monistic position
+for one of agnosticism and scepticism, and made concessions to the
+dualistic dogma of a spiritual world apart from the material frame.
+
+In the course of a Scientific Congress at Munich in 1877 the conflict
+of these antithetic views of nature came into sharp relief. At this
+memorable Congress I had undertaken to deliver the first address
+(September 18th) on the subject of "Modern evolution in relation to
+the whole of science." I maintained that Darwin's theory not only
+solved the great problem of the origin of species, but that its
+implications, especially in regard to the nature of man, threw
+considerable light on the whole of science, and on anthropology in
+particular. The discovery of the real origin of man by evolution from
+a long series of mammal ancestors threw light on his place in nature
+in every respect, as Huxley had already shown in his excellent
+lectures of 1863. Just as all the organs and tissues of the human body
+had originated from those of the nearest related mammals, certain
+ape-like forms, so we were bound to conclude that his mental qualities
+also had been derived from those of his extinct primate ancestor.
+
+This monistic view of the origin and nature of man, which is now admitted
+by nearly all who have the requisite acquaintance with biology, and
+approach the subject without prejudice, encountered a sharp opposition at
+that time. The opposition found its strongest expression in an address that
+Virchow delivered at Munich four days afterwards (September 22nd), on "The
+freedom of science in the modern State." He spoke of the theory of
+evolution as an unproved hypothesis, and declared that it ought not to be
+taught in the schools, because it was dangerous to the State. "We must
+not," he said, "teach that man has descended from the ape or any other
+animal." When Darwin, usually so lenient in his judgment, read the English
+translation of Virchow's speech, he expressed his disapproval in strong
+terms. But the great authority that Virchow had--an authority well founded
+in pathology and sociology--and his prestige as president of the German
+Anthropological Society, had the effect of preventing any member of the
+Society from raising serious opposition to him for thirty years. Numbers of
+journals and treatises repeated his dogmatic statement: "It is quite
+certain that man has descended neither from the ape nor from any other
+animal." In this he persisted till his death in 1902. Since that time the
+whole position of German anthropology has changed. The question is no
+longer whether man was created by a distinct supernatural act or evolved
+from other mammals, but to which line of the animal hierarchy we must look
+for the actual series of ancestors. The interested reader will find an
+account of this "battle of Munich" (1877) in my three Berlin lectures
+(April, 1905), _Der Kampf um die Entwickelungs-Gedanken_.[142]
+
+The main points in our genealogical tree were clearly recognised by
+Darwin in the sixth chapter of the _Descent of Man_. Lowly organised
+fishes, like the lancelot (Amphioxus), are descended from lower
+invertebrates resembling the larvae of an existing Tunicate
+(Appendicularia). From these primitive fishes were evolved higher
+fishes of the ganoid type and others of the type of Lepidosiren
+(Dipneusta). It is a very small step from these to the Amphibia:
+
+"In the class of animals the steps are not difficult to conceive which
+led from the ancient Monotremata to the ancient Marsupials; and from
+these to the early progenitors of the placental mammals. We may thus
+ascend to the Lemuridae; and the interval is not very wide from these
+to the Simiadae. The Simiadae then branched off into two great stems,
+the New World and Old World monkeys; and from the latter, at a remote
+period, Man, the wonder and glory of the Universe, proceeded."[143]
+
+In these few lines Darwin clearly indicated the way in which we were
+to conceive our ancestral series within the vertebrates. It is fully
+confirmed by all the arguments of comparative anatomy and embryology,
+of palaeontology and physiology; and all the research of the
+subsequent forty years have gone to establish it. The deep interest in
+geology which Darwin maintained throughout his life and his complete
+knowledge of palaeontology enabled him to grasp the fundamental
+importance of the palaeontological record more clearly than
+anthropologists and zoologists usually do.
+
+There has been much debate in subsequent decades whether Darwin
+himself maintained that man was descended from the ape, and many
+writers have sought to deny it. But the lines I have quoted _verbatim_
+from the conclusion of the sixth chapter of the _Descent of Man_
+(1871) leave no doubt that he was as firmly convinced of it as was his
+great precursor Jean Lamarck in 1809. Moreover, Darwin adds, with
+particular explicitness, in the "general summary and conclusion"
+(chap. xxi.) of that standard work:[144]
+
+"By considering the embryological structure of man--the homologies
+which he presents with the lower animals,--the rudiments which he
+retains,--and the reversions to which he is liable, we can partly
+recall in imagination the former condition of our early progenitors;
+and can approximately place them in their proper place in the
+zoological series. We thus learn that man is descended from a hairy,
+tailed quadruped, probably arboreal in its habits, and an inhabitant
+of the Old World. This creature, if its whole structure had been
+examined by a naturalist, would have been classed amongst the
+Quadrumana, as surely as the still more ancient progenitor of the Old
+and New World monkeys."
+
+These clear and definite lines leave no doubt that Darwin--so critical
+and cautious in regard to important conclusions--was quite as firmly
+convinced of the descent of man from the apes (the Catarrhinae, in
+particular) as Lamarck was in 1809 and Huxley in 1863.
+
+It is to be noted particularly that, in these and other observations
+on the subject, Darwin decidedly assumes the monophyletic origin of
+the mammals, including man. It is my own conviction that this is of
+the greatest importance. A number of difficult questions in regard to
+the development of man, in respect of anatomy, physiology, psychology,
+and embryology, are easily settled if we do not merely extend our
+_progonotaxis_ to our nearest relatives, the anthropoid apes and the
+tailed monkeys from which these have descended, but go further back
+and find an ancestor in the group of the Lemuridae, and still further
+back to the Marsupials and Monotremata. The essential identity of all
+the Mammals in point of anatomical structure and embryonic
+development--in spite of their astonishing differences in external
+appearance and habits of life--is so palpably significant that modern
+zoologists are agreed in the hypothesis that they have all sprung from
+a common root, and that this root may be sought in the earlier
+Palaeozoic Amphibia.
+
+The fundamental importance of this comparative morphology of the
+Mammals, as a sound basis of scientific anthropology, was recognised
+just before the beginning of the nineteenth century, when Lamarck
+first emphasised (1794) the division of the animal kingdom into
+Vertebrates and Invertebrates. Even thirteen years earlier (1781),
+when Goethe made a close study of the mammal skeleton in the
+Anatomical Institute at Jena, he was intensely interested to find that
+the composition of the skull was the same in man as in the other
+mammals. His discovery of the _os inter-maxillare_ in man (1784),
+which was contradicted by most of the anatomists of the time, and his
+ingenious "vertebral theory of the skull," were the splendid fruit of
+his morphological studies. They remind us how Germany's greatest
+philosopher and poet was for many years ardently absorbed in the
+comparative anatomy of man and the mammals, and how he divined that
+their wonderful identity in structure was no mere superficial
+resemblance, but pointed to a deep internal connection. In my
+_Generelle Morphologie_ (1866), in which I published the first
+attempts to construct phylogenetic trees, I have given a number of
+remarkable theses of Goethe, which may be called "phyletic
+prophecies." They justify us in regarding him as a precursor of
+Darwin.
+
+In the ensuing forty years I have made many conscientious efforts to
+penetrate further along that line of anthropological research that was
+opened up by Goethe, Lamarck, and Darwin. I have brought together the many
+valuable results that have constantly been reached in comparative anatomy,
+physiology, ontogeny, and palaeontology, and maintained the effort to
+reform the classification of animals and plants in an evolutionary sense.
+The first rough drafts of pedigrees that were published in the _Generelle
+Morphologie_ have been improved time after time in the ten editions of my
+_Natuerlich Schoepfungsgeschichte_ (1868-1902).[145] A sounded basis for my
+phyletic hypotheses, derived from a discriminating combination of the three
+great records--morphology, ontogeny, and palaeontology--was provided in the
+three volumes of my _Systematische Phylogenie_[146] (1894 Protists and
+Plants, 1895 Vertebrates, 1896 Invertebrates).
+
+In my _Anthropogenie_[147] I endeavoured to employ all the known
+facts of comparative ontogeny (embryology) for the purpose of
+completing my scheme of human phylogeny (evolution). I attempted to
+sketch the historical development of each organ of the body, beginning
+with the most elementary structures in the germ-layers of the
+Gastraea. At the same time I drew up a corrected statement of the most
+important steps in the line of our ancestral series.
+
+At the fourth International Congress of Zoology at Cambridge (August
+26th, 1898) I delivered an address on "Our present knowledge of the
+Descent of Man." It was translated into English, enriched with many
+valuable notes and additions, by my friend and pupil in earlier days
+Dr. Hans Gadow (Cambridge), and published under the title: _The Last
+Link: our present knowledge of the Descent of Man_[148] The
+determination of the chief animal forms that occur in the line of our
+ancestry is there restricted to thirty types, and these are
+distributed in six main groups.
+
+The first half of this "Progonotaxis hominis," which has no support
+from fossil evidence, comprises three groups: (i) Protista
+(unicellular organisms, 1-5): (ii) Invertebrate Metazoa (Coelenteria
+6-8, Vermalia 9-11): (iii) Monorrhine Vertebrates (Acrania 12-13,
+Cyclostoma 14-15). The second half, which is based on fossil records,
+also comprises three groups: (iv) Palaeozoic cold-blooded Craniota
+(Fishes 16-18, Amphibia 19, Reptiles 20): (v) Mesozoic Mammals
+(Monotrema 21, Marsupialia 22, Mallotheria 23): (vi) Cenozoic Primates
+(Lemuridae 24-25, Tailed Apes 26-27, Anthropomorpha 28-30). An
+improved and enlarged edition of this hypothetic "Progonotaxis
+hominis" was published in 1908, in my essay _Unsere Ahnenreihe_.[149]
+
+If I have succeeded in furthering, in some degree, by these
+anthropological works, the solution of the great problem of Man's
+place in nature, and particularly in helping to trace the definite
+stages in our ancestral series, I owe the success, not merely to the
+vast progress that biology has made in the last half century, but
+largely to the luminous example of the great investigators who have
+applied themselves to the problem, with so much assiduity and genius,
+for a century and a quarter--I mean Goethe and Lamarck, Gegenbaur and
+Huxley, but, above all, Charles Darwin. It was the great genius of
+Darwin that first brought together that symmetrical temple of
+scientific knowledge, the theory of descent. It was Darwin who put the
+crown on the edifice by his theory of natural selection. Not until
+this broad inductive law was firmly established was it possible to
+vindicate the special conclusion, the descent of man from a series of
+other Vertebrates. By his illuminating discovery Darwin did more for
+anthropology than thousands of those writers, who are more
+specifically titled anthropologists, have done by their technical
+treatises. We may, indeed, say that it is not merely as an exact
+observer and ingenious experimenter, but as a distinguished
+anthropologist and far-seeing thinker, that Darwin takes his place
+among the greatest men of science of the nineteenth century.
+
+To appreciate fully the immortal merit of Darwin in connection with
+anthropology, we must remember that not only did his chief work, _The
+Origin of Species_, which opened up a new era in natural history in
+1859, sustain the most virulent and widespread opposition for a
+lengthy period, but even thirty years later, when its principles were
+generally recognised and adopted, the application of them to man was
+energetically contested by many high scientific authorities. Even
+Alfred Russel Wallace, who discovered the principle of natural
+selection independently in 1858, did not concede that it was
+applicable to the higher mental and moral qualities of man. Dr.
+Wallace still holds a spiritualist and dualist view of the nature of
+man, contending that he is composed of a material frame (descended
+from the apes) and an immortal immaterial soul (infused by a higher
+power). This dual conception, moreover, is still predominant in the
+wide circles of modern theology and metaphysics, and has the general
+and influential adherence of the more conservative classes of society.
+
+In strict contradiction to this mystical dualism, which is generally
+connected with teleology and vitalism, Darwin always maintained the
+complete unity of human nature, and showed convincingly that the
+psychological side of man was developed, in the same way as the body,
+from the less advanced soul of the anthropoid ape, and, at a still
+more remote period, from the cerebral functions of the older
+vertebrates. The eighth chapter of the _Origin of Species_, which is
+devoted to instinct, contains weighty evidence that the instincts of
+animals are subject, like all other vital processes, to the general
+laws of historic development. The special instincts of particular
+species were formed by adaptation, and the modifications thus acquired
+were handed on to posterity by heredity; in their formation and
+preservation natural selection plays the same part as in the
+transformation of every other physiological function. The higher moral
+qualities of civilised man have been derived from the lower mental
+functions of the uncultivated barbarians and savages, and these in
+turn from the social instincts of the mammals. This natural and
+monistic psychology of Darwin's was afterwards more fully developed by
+his friend George Romanes in his excellent works _Mental Evolution in
+Animals_ and _Mental Evolution in Man_.[150]
+
+Many valuable and most interesting contributions to this monistic
+psychology of man were made by Darwin in his fine work on _The Descent
+of Man and Selection in Relation to Sex_, and again in his
+supplementary work, _The Expression of the Emotions in Man and
+Animals_. To understand the historical development of Darwin's
+anthropology one must read his life and the introduction to _The
+Descent of Man_. From the moment that he was convinced of the truth
+of the principle of descent--that is to say, from his thirtieth year,
+in 1838--he recognised clearly that man could not be excluded from its
+range. He recognised as a logical necessity the important conclusion
+that "man is the co-descendant with other species of some ancient,
+lower, and extinct form." For many years he gathered notes and
+arguments in support of this thesis, and for the purpose of showing
+the probable line of man's ancestry. But in the first edition of _The
+Origin of Species_ (1859) he restricted himself to the single line,
+that by this work "light would be thrown on the origin of man and his
+history." In the fifty years that have elapsed since that time the
+science of the origin and nature of man has made astonishing progress,
+and we are now fairly agreed in a monistic conception of nature that
+regards the whole universe, including man, as a wonderful unity,
+governed by unalterable and eternal laws. In my philosophical book
+_Die Weltraetsel_ (1899)[151] and in the supplementary volume _Die
+Lebenswunder_ (1904)[152] I have endeavoured to show that this pure
+monism is securely established, and that the admission of the
+all-powerful rule of the same principle of evolution throughout the
+universe compels us to formulate a single supreme law--the
+all-embracing "Law of Substance," or the united laws of the constancy
+of matter and the conservation of energy. We should never have reached
+this supreme general conception if Charles Darwin--a "monistic
+philosopher" in the true sense of the word--had not prepared the way
+by his theory of descent by natural selection, and crowned the great
+work of his life by the association of this theory with a naturalistic
+anthropology.
+
+FOOTNOTES:
+
+[Footnote 127: _Generelle Morphologie der Organismen_, 2 vols.,
+Berlin, 1866.]
+
+[Footnote 128: Eng. transl.; _The History of Creation_, London, 1876.]
+
+[Footnote 129: London, 1908.]
+
+[Footnote 130: Munich, 1884.]
+
+[Footnote 131: _Die Mutationstheorie_, Leipzig, 1903.]
+
+[Footnote 132: _Der Kampf der Theile im Organismus_, Leipzig, 1881.]
+
+[Footnote 133: Leipzig, 1904.]
+
+[Footnote 134: Berlin, 1876.]
+
+[Footnote 135: _Wonders of Life_, London and New York, 1904.]
+
+[Footnote 136: Eng. transl.; _The Evolution of Man_, 2 vols., London,
+1879 and 1905.]
+
+[Footnote 137: _Origin of Species_ (6th edit.), p. 396.]
+
+[Footnote 138: Eng. transl.; _Facts and Arguments for Darwin_, London,
+1869.]
+
+[Footnote 139: 3 vols., Berlin, 1894-96.]
+
+[Footnote 140: _Descent of Man_ (Popular Edit.), p. 927.]
+
+[Footnote 141: _Gesammelte Abhandlungen zur wissenschaftlichen
+Medizin_, Berlin, 1856.]
+
+[Footnote 142: Eng. transl.; _Last Words on Evolution_, London, 1906.]
+
+[Footnote 143: _Descent of Man_, (Popular Edit.), p. 255.]
+
+[Footnote 144: _Descent of Man_, p. 930.]
+
+[Footnote 145: Eng. transl.; _The History of Creation_, London, 1876.]
+
+[Footnote 146: Berlin, 1894-96.]
+
+[Footnote 147: Leipzig, 1874, 5th edit. 1905. Eng. transl.; _The
+Evolution of Man_, London, 1905.]
+
+[Footnote 148: London, 1898.]
+
+[Footnote 149: _Festschrift zur 350-jaehrigen Jubelfeier der Thueringer
+Universitaet Jena_. Jena. 1908.]
+
+[Footnote 150: London, 1885; 1888.]
+
+[Footnote 151: _The Riddle of the Universe_, London and New York,
+1900.]
+
+[Footnote 152: _The Wonders of Life_, London and New York, 1904.]
+
+
+
+
+VI
+
+MENTAL FACTORS IN EVOLUTION
+
+BY C. LLOYD MORGAN, LLD., F.R.S
+
+
+In developing his conception of organic evolution Charles Darwin was
+of necessity brought into contact with some of the problems of mental
+evolution. In _The Origin of Species_ he devoted a chapter to "the
+diversities of instinct and of the other mental faculties in animals
+of the same class."[153] When he passed to the detailed consideration
+of _The Descent of Man_, it was part of his object to show "that there
+is no fundamental difference between man and the higher mammals in
+their mental faculties."[154] "If no organic being excepting man," he
+said, "had possessed any mental power, or if his powers had been of a
+wholly different nature, from those of the lower animals, then we
+should never have been able to convince ourselves that our high
+faculties had been gradually developed."[155] In his discussion of
+_The Expression of the Emotions_ it was important for his purpose
+"fully to recognise that actions readily become associated with other
+actions and with various states of the mind."[156] His hypothesis of
+sexual selection is largely dependent upon the exercise of choice on
+the part of the female and her preference for "not only the more
+attractive but at the same time the more vigourous and vicious
+males."[157] Mental processes and physiological processes were for
+Darwin closely correlated; and he accepted the conclusion "that the
+nervous system not only regulates most of the existing functions of
+the body, but has indirectly influenced the progressive development of
+various bodily structures and of certain mental qualities."[158]
+
+Throughout his treatment, mental evolution was for Darwin incidental
+to and contributory to organic evolution. For specialised research in
+comparative and genetic psychology, as an independent field of
+investigation, he had neither the time nor the requisite training.
+None the less his writings and the spirit of his work have exercised a
+profound influence on this department of evolutionary thought. And,
+for those who follow Darwin's lead, mental evolution is still in a
+measure subservient to organic evolution. Mental processes are the
+accompaniments or concomitants of the functional activity of specially
+differentiated parts of the organism. They are in some way dependent
+on physiological and physical conditions. But though they are not
+physical in their nature, and though it is difficult or impossible to
+conceive that they are physical in their origin, they are, for Darwin
+and his followers, factors in the evolutionary process in its physical
+or organic aspect. By the physiologist within his special and
+well-defined universe of discourse they may be properly regarded as
+epiphenomena; but by the naturalist in his more catholic survey of
+nature they cannot be so regarded, and were not so regarded by Darwin.
+Intelligence has contributed to evolution of which it is in a sense a
+product.
+
+The facts of observation or of inference which Darwin accepted are
+these: Conscious experience accompanies some of the modes of animal
+behaviour; it is concomitant with certain physiological processes;
+these processes are the outcome of development in the individual and
+evolution in the race; the accompanying mental processes undergo a
+like development. Into the subtle philosophical questions which arise
+out of the naive acceptance of such a creed it was not Darwin's
+province to enter; "I have nothing to do," he said,[159] "with the
+origin of the mental powers, any more than I have with that of life
+itself." He dealt with the natural history of organisms, including not
+only their structure but their modes of behaviour; with the natural
+history of the states of consciousness which accompany some of their
+actions; and with the relation of behaviour to experience. We will
+endeavour to follow Darwin in his modesty and candour in making no
+pretence to give ultimate explanations. But we must note one of the
+implications of this self-denying ordinance of science. Development
+and evolution imply continuity. For Darwin and his followers the
+continuity is organic through physical heredity. Apart from
+speculative hypothesis, legitimate enough in its proper place but here
+out of court, we know nothing of continuity of mental evolution as
+such: consciousness appears afresh in each succeeding generation.
+Hence it is that for those who follow Darwin's lead, mental evolution
+is and must ever be, within his universe of discourse, subservient to
+organic evolution. Only in so far as conscious experience, or its
+neural correlate, effects some changes in organic structure can it
+influence the course of heredity; and conversely only in so far as
+changes in organic structure are transmitted through heredity, is
+mental evolution rendered possible. Such is the logical outcome of
+Darwin's teaching.
+
+Those who abide by the cardinal results of this teaching are bound to
+regard all behaviour as the expression of the functional activities of
+the living tissues of the organism, and all conscious experience as
+correlated with such activities. For the purposes of scientific
+treatment, mental processes are one mode of expression of the same
+changes of which the physiological processes accompanying behaviour
+are another mode of expression. This is simply accepted as a fact
+which others may seek to explain. The behaviour itself is the adaptive
+application of the energies of the organism; it is called forth by
+some form of presentation or stimulation brought to bear on the
+organism by the environment. This presentation is always an individual
+or personal matter. But in order that the organism may be fitted to
+respond to the presentation of the environment it must have undergone
+in some way a suitable preparation. According to the theory of
+evolution this preparation is primarily racial and is transmitted
+through heredity. Darwin's main thesis was that the method of
+preparation is predominantly by natural selection. Subordinate to
+racial preparation, and always dependent thereon, is individual or
+personal preparation through some kind of acquisition; of which the
+guidance of behaviour through individually won experience is a typical
+example. We here introduce the mental factor because the facts seem to
+justify the inference. Thus there are some modes of behaviour which
+are wholly and solely dependent upon inherited racial preparation;
+there are other modes of behaviour which are also dependent, in part
+at least, on individual preparation. In the former case the behaviour
+is adaptive on the first occurrence of the appropriate presentation;
+in the latter case accommodation to circumstances is only reached
+after a greater or less amount of acquired organic modification of
+structure, often accompanied (as we assume) in the higher animals by
+acquired experience. Logically and biologically the two classes of
+behaviour are clearly distinguishable: but the analysis of complex
+cases of behaviour where the two factors cooeperate, is difficult and
+requires careful and critical study of life-history.
+
+The foundations of the mental life are laid in the conscious
+experience that accompanies those modes of behaviour, dependent
+entirely on racial preparation, which may broadly be described as
+instinctive. In the eighth chapter of _The Origin of Species_ Darwin
+says,[160] "I will not attempt any definition of instinct.... Every
+one understands what is meant, when it is said that instinct impels
+the cuckoo to migrate and to lay her eggs in other birds' nests. An
+action, which we ourselves require experience to enable us to perform,
+when performed by an animal, more especially by a very young one,
+without experience, and when performed by many individuals in the same
+way, without their knowing for what purpose it is performed, is
+usually said to be instinctive." And in the summary at the close of
+the chapter he says,[161] "I have endeavoured briefly to show that the
+mental qualities of our domestic animals vary, and that the variations
+are inherited. Still more briefly I have attempted to show that
+instincts vary slightly in a state of nature. No one will dispute that
+instincts are of the highest importance to each animal. Therefore
+there is no real difficulty, under changing conditions of life, in
+natural selection accumulating to any extent slight modifications of
+instinct which are in any way useful. In many cases habit or use and
+disuse have probably come into play."
+
+Into the details of Darwin's treatment there is neither space nor need
+to enter. There are some ambiguous passages; but it may be said that
+for him, as for his followers to-day, instinctive behaviour is wholly
+the result of racial preparation transmitted through organic heredity.
+For the performance of the instinctive act no individual preparation
+under the guidance of personal experience is necessary. It is true
+that Darwin quotes with approval Huber's saying that "a little dose of
+judgment or reason often comes into play, even with animals low in the
+scale of nature."[162] But we may fairly interpret his meaning to be
+that in behaviour, which is commonly called instinctive, some element
+of intelligent guidance is often combined. If this be conceded the
+strictly instinctive performance (or part of the performance) is the
+outcome of heredity and due to the direct transmission of parental or
+ancestral aptitudes. Hence the instinctive response as such depends
+entirely on how the nervous mechanism has been built up through
+heredity; while intelligent behaviour, or the intelligent factor in
+behaviour, depends also on how the nervous mechanism has been modified
+and moulded by use during its development and concurrently with the
+growth of individual experience in the customary situations of daily
+life. Of course it is essential to the Darwinian thesis that what Sir
+E. Ray Lankester has termed "educability," not less than instinct, is
+hereditary. But it is also essential to the understanding of this
+thesis that the differentiae of the hereditary factor should be
+clearly grasped.
+
+For Darwin there were two modes of racial preparation, (_1_) natural
+selection, and (_2_) the establishment of individually acquired habit.
+He showed that instincts are subject to hereditary variation; he saw
+that instincts are also subject to modification through acquisition in
+the course of individual life. He believed that not only the
+variations but also, to some extent, the modifications are inherited.
+He therefore held that some instincts (the greater number) are due to
+natural selection but that others (less numerous) are due, or partly
+due, to the inheritance of acquired habits. The latter involve
+Lamarckian inheritance, which of late years has been the centre of so
+much controversy. It is noteworthy however that Darwin laid especial
+emphasis on the fact that many of the most typical and also the most
+complex instincts--those of neuter insects--do not admit of such an
+interpretation. "I am surprised," he says,[163] "that no one has
+hitherto advanced this demonstrative case of neuter insects, against
+the well-known doctrine of inherited habit, as advanced by Lamarck."
+None the less Darwin admitted this doctrine as supplementary to that
+which was more distinctively his own--for example in the case of the
+instincts of domesticated animals. Still, even in such cases, "it may
+be doubted," he says,[164] "whether any one would have thought of
+training a dog to point, had not some one dog naturally shown a
+tendency in this line ... so that habit and some degree of selection
+have probably concurred in civilising by inheritance our dogs." But in
+the interpretation of the instincts of domesticated animals, a more
+recently suggested hypothesis, that of organic selection,[165] may be
+helpful. According to this hypothesis any intelligent modification of
+behaviour which is subject to selection is probably coincident in
+direction with an inherited tendency to behave in this fashion. Hence
+in such behaviour there are two factors: (1) an incipient variation in
+the line of such behaviour, and (2) an acquired modification by which
+the behaviour is carried further along the same line. Under natural
+selection those organisms in which the two factors cooeperate are
+likely to survive. Under artificial selection they are deliberately
+chosen out from among the rest.
+
+Organic selection has been termed a compromise between the more
+strictly Darwinian and the Lamarckian principles of interpretation.
+But it is not in any sense a compromise. The principle of
+interpretation of that which is instinctive and hereditary is wholly
+Darwinian. It is true that some of the facts of observation relied
+upon by Lamarckians are introduced. For Lamarckians however the
+modifications which are admittedly factors in survival, are regarded
+as the parents of inherited variations; for believers in organic
+selection they are only the foster-parents or nurses. It is because
+organic selection is the direct outcome of and a natural extension of
+Darwin's cardinal thesis that some reference to it here is
+justifiable. The matter may be put with the utmost brevity as follows:
+(1) Variations (V) occur, some of which are in the direction of
+increased adaptation (+), others in the direction of decreased
+adaptation (-).
+
+(2) Acquired modifications (M) also occur. Some of these are in the
+direction of increased accommodation to circumstances (+), while
+others are in the direction of diminished accommodation (-). Four
+major combinations are
+
+   (_b_) + V with - M,      (_c_) - V with + M,
+
+   (_a_) + V with + M,      (_d_) - V with - M.
+
+Of these (_d_) must inevitably be eliminated while (_a_) are selected.
+The predominant survival of (_a_) entails the survival of the adaptive
+variations which are inherited. The contributory acquisitions (+ M)
+are not inherited; but there are none the less factors in determining
+the survival of the coincident variations. It is surely abundantly
+clear that this is Darwinism and has no tincture of Lamarck's
+essential principle, the inheritance of acquired characters.
+
+Whether Darwin himself would have accepted this interpretation of some
+at least of the evidence put forward by Lamarckians is unfortunately a
+matter of conjecture. The fact remains that in his interpretation of
+instinct and in allied questions he accepted the inheritance of
+individually acquired modifications of behaviour and structure.
+
+Darwin was chiefly concerned with instinct from the biological rather
+than from the psychological point of view. Indeed it must be confessed
+that, from the latter standpoint, his conception of instinct as a
+"mental faculty" which "impels" an animal to the performance of
+certain actions, scarcely affords a satisfactory basis for genetic
+treatment. To carry out the spirit of Darwin's teaching it is
+necessary to link more closely biological and psychological evolution.
+The first step towards this is to interpret the phenomena of
+instinctive behaviour in terms of stimulation and response. It may be
+well to take a particular case. Swimming on the part of a duckling is,
+from the biological point of view, a typical example of instinctive
+behaviour. Gently lower a recently hatched bird into water:
+coordinated movements of the limbs follow in rhythmical sequence. The
+behaviour is new to the individual though it is no doubt closely
+related to that of walking, which is no less instinctive. There is a
+group of stimuli afforded by the "presentation" which results from
+partial immersion: upon this there follows as a complex response an
+application of the functional activities in swimming; the sequence of
+adaptive application on the appropriate presentation is determined by
+racial preparation. We know, it is true, but little of the
+physiological details of what takes place in the central nervous
+system; but in broad outline the nature of the organic mechanism and
+the manner of its functioning may at least be provisionally
+conjectured in the present state of physiological knowledge. Similarly
+in the case of the pecking of newly-hatched chicks; there is a visual
+presentation, there is probably a cooeperating group of stimuli from
+the alimentary tract in need of food, there is an adaptive application
+of the activities in a definite mode of behaviour. Like data are
+afforded in a great number of cases of instinctive procedure,
+sometimes occurring very early in life, not infrequently deferred
+until the organism is more fully developed, but all of them dependent
+upon racial preparation. No doubt there is some range of variation in
+the behaviour, just such variation as the theory of natural selection
+demands. But there can be no question that the higher animals inherit
+a bodily organisation and a nervous system, the functional working of
+which gives rise to those inherited modes of behaviour which are
+termed instinctive.
+
+It is to be noted that the term "instinctive" is here employed in the
+adjectival form as a descriptive heading under which may be grouped
+many and various modes of behaviour due to racial preparation. We
+speak of these as inherited; but in strictness what is transmitted
+through heredity is the complex of anatomical and physiological
+conditions under which, in appropriate circumstances, the organism so
+behaves. So far the term "instinctive" has a restricted biological
+connotation in terms of behaviour. But the connecting link between
+biological evolution and psychological evolution is to be sought,--as
+Darwin fully realised,--in the phenomena of instinct, broadly
+considered. The term "instinctive" has also a psychological
+connotation. What is that connotation?
+
+Let us take the case of the swimming duckling or the pecking chick,
+and fix our attention on the first instinctive performance. Grant that
+just as there is, strictly speaking, no inherited behaviour, but only
+the conditions which render such behaviour under appropriate
+circumstances possible; so too there is no inherited experience, but
+only the conditions which render such experience possible; then the
+cerebral conditions in both cases are the same. The biological
+behaviour-complex, including the total stimulation and the total
+response with the intervening or resultant processes in the sensorium,
+is accompanied by an experience-complex including the initial
+stimulation-consciousness and resulting response-consciousness. In the
+experience-complex are comprised data which in psychological analysis
+are grouped under the headings of cognition, affective tone and
+conation. But the complex is probably experienced as an unanalysed
+whole. If then we use the term "instinctive" so as to comprise all
+congenital modes of behaviour which contribute to experience, we are
+in a position to grasp the view that the net result in consciousness
+constitutes what we may term the primary tissue of experience. To the
+development of this experience each instinctive act contributes. The
+nature and manner of organisation of this primary tissue of experience
+are dependent on inherited biological aptitudes; but they are from the
+outset onwards subject to secondary development dependent on acquired
+aptitudes. Biological values are supplemented by psychological values
+in terms of satisfaction or the reverse.
+
+In our study of instinct we have to select some particular phase of
+animal behaviour and isolate it so far as is possible from the life of
+which it is a part. But the animal is a going concern, restlessly
+active in many ways. Many instinctive performances, as Darwin pointed
+out,[166] are serial in their nature. But the whole of active life is
+a serial and coordinated business. The particular instinctive
+performance is only an episode in a life-history, and every mode of
+behaviour is more or less closely correlated with other modes. This
+coordination of behaviour is accompanied by a correlation of the modes
+of primary experience. We may classify the instinctive modes of
+behaviour and their accompanying modes of instinctive experience under
+as many heads as may be convenient for our purposes of interpretation,
+and label them instincts of self-preservation, of pugnacity, of
+acquisition, the reproductive instincts, the parental instincts, and
+so forth. An instinct, in this sense of the term (for example the
+parental instinct), may be described as a specialised part of the
+primary tissue of experience differentiated in relation to some
+definite biological end. Under such an instinct will fall a large
+number of particular and often well-defined modes of behaviour, each
+with its own peculiar mode of experience.
+
+It is no doubt exceedingly difficult as a matter of observation and of
+inference securely based thereon to distinguish what is primary from
+what is in part due to secondary acquisition--a fact which Darwin
+fully appreciated. Animals are educable in different degrees; but
+where they are educable they begin to profit by experience from the
+first. Only, therefore, on the occasion of the first instinctive act
+of a given type can the experience gained be regarded as _wholly_
+primary; all subsequent performance is liable to be in some degree,
+sometimes more, sometimes less, modified by the acquired disposition
+which the initial behaviour engenders. But the early stages of
+acquisition are always along the lines predetermined by instinctive
+differentiation. It is the task of comparative psychology to
+distinguish the primary tissue of experience from its secondary and
+acquired modifications. We cannot follow up the matter in further
+detail. It must here suffice to suggest that this conception of
+instinct as a primary form of experience lends itself better to
+natural history treatment than Darwin's conception of an impelling
+force, and that it is in line with the main trend of Darwin's thought.
+
+In a characteristic work,--characteristic in wealth of detail, in
+closeness and fidelity of observation, in breadth of outlook, in
+candour and modesty,--Darwin dealt with _The Expression of the
+Emotions in Man and Animals_. Sir Charles Bell in his _Anatomy of
+Expression_ had contended that many of man's facial muscles had been
+specially created for the sole purpose of being instrumental in the
+expression of his emotions. Darwin claimed that a natural explanation,
+consistent with the doctrine of evolution, could in many cases be
+given and would in other cases be afforded by an extension of the
+principles he advocated. "No doubt," he said,[167] "as long as man and
+all other animals are viewed as independent creations, an effectual
+stop is put to our natural desire to investigate as far as possible
+the causes of Expression. By this doctrine, anything and everything
+can be equally well explained.... With mankind, some expressions ...
+can hardly be understood, except on the belief that man once existed
+in a much lower and animal-like condition. The community of certain
+expressions in distinct though allied species ... is rendered somewhat
+more intelligible, if we believe in their descent from a common
+progenitor. He who admits on general grounds that the structure and
+habits of all animals have been gradually evolved, will look at the
+whole subject of Expression in a new and interesting light."
+
+Darwin relied on three principles of explanation. "The first of these
+principles is, that movements which are serviceable in gratifying some
+desire, or in relieving some sensation, if often repeated, become so
+habitual that they are performed, whether or not of any service,
+whenever the same desire or sensation is felt, even in a very weak
+degree."[168] The modes of expression which fall under this head have
+become instinctive through the hereditary transmission of acquired
+habit. "As far as we can judge, only a few expressive movements are
+learnt by each individual; that is, were consciously and voluntarily
+performed during the early years of life for some definite object, or
+in imitation of others, and then became habitual. The far greater
+number of the movements of expression, and all the more important
+ones, are innate or inherited; and such cannot be said to depend on
+the will of the individual. Nevertheless, all those included under our
+first principle were at first voluntarily performed for a definite
+object,--namely, to escape some danger, to relieve some distress, or
+to gratify some desire."[169]
+
+"Our second principle is that of antithesis. The habit of voluntarily
+performing opposite movements under opposite impulses has become
+firmly established in us by the practice of our whole lives. Hence, if
+certain actions have been regularly performed, in accordance with our
+first principle, under a certain frame of mind, there will be a strong
+and involuntary tendency to the performance of directly opposite
+actions, whether or not these are of any use, under the excitement of
+an opposite frame of mind."[170] This principle of antithesis has not
+been widely accepted. Nor is Darwin's own position easy to grasp.
+
+"Our third principle," he says,[171] "is the direct action of the
+excited nervous system on the body, independently of the will, and
+independently, in large part, of habit. Experience shows that
+nerve-force is generated and set free whenever the cerebro-spinal
+system is excited. The direction which this nerve-force follows is
+necessarily determined by the lines of connection between the
+nerve-cells, with each other and with various parts of the body."
+
+Lack of space prevents our following up the details of Darwin's
+treatment of expression. Whether we accept or do not accept his three
+principles of explanation we must regard his work as a masterpiece of
+descriptive analysis, packed full of observations possessing lasting
+value. For a further development of the subject it is essential that
+the instinctive factors in expression should be more fully
+distinguished from those which are individually acquired--a difficult
+task--and that the instinctive factors should be rediscussed in the
+light of modern doctrines of heredity, with a view to determining
+whether Lamarckian inheritance, on which Darwin so largely relied, is
+necessary for an interpretation of the facts.
+
+The whole subject as Darwin realised is very complex. Even the term
+"expression" has a certain amount of ambiguity. When the emotion is in
+full flood, the animal fights, flees, or faints. Is this full-tide
+effect to be regarded as expression; or are we to restrict the term to
+the premonitory or residual effects--the bared canine when the
+fighting mood is being roused, the ruffled fur when reminiscent
+representations of the object inducing anger cross the mind? Broadly
+considered both should be included. The activity of premonitory
+expression as a means of communication was recognised by Darwin; he
+might, perhaps, have emphasised it more strongly in dealing with the
+lower animals. Man so largely relies on a special means of
+communication, that of language, that he sometimes fails to realise
+that for animals with their keen powers of perception, and dependent
+as they are on such means of communication, the more strictly
+biological means of expression are full of subtle suggestiveness. Many
+modes of expression, otherwise useless, are signs of behaviour that
+may be anticipated,--signs which stimulate the appropriate attitude of
+response. This would not, however, serve to account for the utility of
+the organic accompaniments--heart-affection, respiratory changes,
+vaso-motor effects and so forth, together with heightened muscular
+tone,--on all of which Darwin lays stress[172] under his third
+principle. The biological value of all this is, however, of great
+importance, though Darwin was hardly in a position to take it fully
+into account.
+
+Having regard to the instinctive and hereditary factors of emotional
+expression we may ask whether Darwin's third principle does not alone
+suffice as an explanation. Whether we admit or reject Lamarckian
+inheritance it would appear that all hereditary expression must be due
+to pre-established connections within the central nervous system and
+to a transmitted provision for coordinated response under the
+appropriate stimulation. If this be so, Darwin's first and second
+principles are subordinate and ancillary to the third, an expression,
+so far as it is instinctive or heredity, being "the direct result of
+the constitution of the nervous system."
+
+Darwin accepted the emotions themselves as hereditary or acquired
+states of mind and devoted his attention to their expression. But
+these emotions themselves are genetic products and as such dependent
+on organic conditions. It remained, therefore, for psychologists who
+accepted evolution and sought to build on biological foundations to
+trace the genesis of these modes of animal and human experience. The
+subject has been independently developed by Professors Lange and
+James;[173] and some modification of their view is regarded by many
+evolutionists as affording the best explanation of the facts. We must
+fix our attention on the lower emotions, such as anger or fear, and on
+their first occurrence in the life of the individual organism. It is a
+matter of observation that if a group of young birds which have been
+hatched in an incubator are frightened by an appropriate presentation,
+auditory or visual, they instinctively respond in special ways. If we
+speak of this response as the expression, we find that there are many
+factors. There are certain visible modes of behaviour, crouching at
+once, scattering and then crouching, remaining motionless, the braced
+muscles sustaining an attitude of arrest, and so forth, There are also
+certain visceral or organic effects, such as affections of the heart
+and respiration. These can be readily observed by taking the young
+bird in the hand. Other effects cannot be readily observed; vaso-motor
+changes, affections of the alimentary canal, the skin and so forth.
+Now the essence of the James-Lange view, as applied to these
+congenital effects, is that though we are justified in speaking of
+them as effects of the stimulation, we are not justified, without
+further evidence, in speaking of them as effects of the emotional
+state. May it not rather be that the emotion as a primary mode of
+experience is the concomitant of the net result of the organic
+situation--the initial presentation, the instinctive mode of
+behaviour, the visceral disturbances?
+
+According to this interpretation the primary tissue of experience of
+the emotional order, felt as an unanalysed complex, is generated by
+the stimulation of the sensorium by afferent or incoming physiological
+impulses from the special senses, from the organs concerned in the
+responsive behaviour, from the viscere and vaso-motor system.
+
+Some psychologists, however, contend that the emotional experience is
+generated in the sensorium prior to, and not subsequent to, the
+behaviour-response and the visceral disturbances. It is a direct and
+not an indirect outcome of the presentation to the special senses. Be
+this as it may, there is a growing tendency to bring into the closest
+possible relation, or even to identify, instinct and emotion in their
+primary genesis. The central core of all such interpretations is that
+instinctive behaviour and experience, its emotional accompaniments,
+and its expression, are but different aspects of the outcome of the
+same organic occurrences. Such emotions are, therefore, only a
+distinguishable aspect of the primary tissue of experience and exhibit
+a like differentiation. Here again a biological foundation is laid for
+a psychological doctrine of the mental development of the individual.
+
+The intimate relation between emotion as a psychological mode of
+experience and expression as a group of organic conditions has an
+important bearing on biological interpretation. The emotion, as the
+psychological accompaniment of orderly disturbances in the central
+nervous system, profoundly influences behaviour and often renders it
+more vigourous and more effective. The utility of the emotions in the
+struggle for existence can, therefore, scarcely be over-estimated.
+Just as keenness of perception has survival-value; just as it is
+obviously subject to variation; just as it must be enhanced under
+natural selection, whether individually acquired increments are
+inherited or not; and just as its value lies not only in this or that
+special perceptive act but in its importance for life as a whole; so
+the vigourous effectiveness of activity has survival-value; it is
+subject to variation; it must be enhanced under natural selection; and
+its importance lies not only in particular modes of behaviour but in
+its value for life as a whole. If emotion and its expression as a
+congenital endowment are but different aspects of the same biological
+occurrence; and if this is a powerful supplement to vigour
+effectiveness and persistency of behaviour, it must on Darwin's
+principles be subject to natural selection.
+
+If we include under the expression of the emotions not only the
+premonitory symptoms of the initial phases of the organic and mental
+state, not only the signs or conditions of half-tide emotion, but the
+full-tide manifestation of an emotion which dominates the situation,
+we are naturally led on to the consideration of many of the phenomena
+which are discussed under the head of sexual selection. The subject is
+difficult and complex, and it was treated by Darwin with all the
+strength he could summon to the task. It can only be dealt with here
+from a special point of view--that which may serve to illustrate the
+influence of certain mental factors on the course of evolution. From
+this point of view too much stress can scarcely be laid on the
+dominance of emotion during the period of courtship and pairing in the
+more highly organised animals. It is a period of maximum vigour,
+maximum activity, and, correlated with special modes of behaviour and
+special organic and visceral accompaniments, a period also of maximum
+emotional excitement. The combats of males, their dances and aerial
+evolutions, their elaborate behaviour and display, or the flood of
+song in birds, are emotional expressions which are at any rate
+coincident in time with sexual periodicity. From the combat of the
+males there follows on Darwin's principles the elimination of those
+which are deficient in bodily vigour, deficient in special structures,
+offensive or protective, which contribute to success, deficient in the
+emotional supplement of which persistent and whole-hearted fighting is
+the expression, and deficient in alertness and skill which are the
+outcome of the psychological development of the powers of perception.
+Few biologists question that we have here a mode of selection of much
+importance, though its influence on psychological evolution often
+fails to receive its due emphasis. Mr. Wallace[174] regards it as "a
+form of natural selection"; "to it," he says, "we must impute the
+development of the exceptional strength, size, and activity of the
+male, together with the possession of special offensive and defensive
+weapons, and of all other characters which arise from the development
+of these or are correlated with them." So far there is little
+disagreement among the followers of Darwin--for Mr. Wallace, with fine
+magnanimity, has always preferred to be ranked as such,
+notwithstanding his right, on which a smaller man would have
+constantly insisted, to the claim of independent originator of the
+doctrine of natural selection. So far with regard to sexual selection
+Darwin and Mr. Wallace are agreed; so far and no farther. For Darwin,
+says Mr. Wallace,[175] "has extended the principle into a totally
+different field of action, which has none of that character of
+constancy and of inevitable result that attaches to natural selection,
+including male rivalry; for by far the larger portion of the
+phenomena, which he endeavours to explain by the direct action of
+sexual selection, can only be so explained on the hypothesis that the
+immediate agency is female choice or preference. It is to this that he
+imputes the origin of all secondary sexual characters other than
+weapons of offence and defence.... In this extension of sexual
+selection to include the action of female choice or preference, and in
+the attempt to give to that choice such wide-reaching effects, I am
+unable to follow him more than a very little way."
+
+Into the details of Mr. Wallace's criticisms it is impossible to enter
+here. We cannot discuss either the mode of origin of the variations in
+structure which have rendered secondary sexual characters possible or
+the modes of selection other than sexual which have rendered them,
+within narrow limits, specifically constant. Mendelism and mutation
+theories may have something to say on the subject when these theories
+have been more fully correlated with the basal principles of
+selection. It is noteworthy that Mr. Wallace says:[176] "Besides the
+acquisition of weapons by the male for the purpose of fighting with
+other males, there are some other sexual characters which may have
+been produced by natural selection. Such are the various sounds and
+odours which are peculiar to the male, and which serve as a call to
+the female or as an indication of his presence. These are evidently a
+valuable addition to the means of recognition of the two sexes, and
+are a further indication, that the pairing season has arrived; and the
+production, intensification, and differentiation of these sounds and
+odours are clearly within the power of natural selection. The same
+remark will apply to the peculiar calls of birds, and even to the
+singing of the males." Why the same remark should not apply to their
+colours and adornments is not obvious. What is obvious is that "means
+of recognition" and "indication that the pairing season has arrived"
+are dependent on the perceptive powers of the female who recognises
+and for whom the indication has meaning. The hypothesis of female
+preference, stripped of the aesthetic surplusage which is
+psychologically both unnecessary and unproven, is really only
+different in degree from that which Mr. Wallace admits in principle
+when he says that it is probable that the female is pleased or excited
+by the display.
+
+Let us for our present purpose leave on one side and regard as _sub
+judice_ the question whether the specific details of secondary sexual
+characters are the outcome of female choice. For us the question is
+whether certain psychological accompaniments of the pairing situation
+have influenced the course of evolution and whether these
+psychological accompaniments are themselves the outcome of evolution.
+As a matter of observation, specially differentiated modes of
+behaviour, often very elaborate, frequently requiring highly developed
+skill, and apparently highly charged with emotional tone, are the
+precursors of pairing. They are generally confined to the males, whose
+fierce combats during the period of sexual activity are part of the
+emotional manifestation. It is inconceivable that they have no
+biological meaning; and it is difficult to conceive that they have any
+other biological end than to evoke in the generally more passive
+female the pairing impulse. They, are based on instinctive foundations
+ingrained in the nervous constitution through natural (or may we not
+say sexual?) selection in virtue of their profound utility. They are
+called into play by a specialised presentation such as the sight or
+the scent of the female at, or a little in advance of, a critical
+period of the physiological rhythm. There is no necessity that the
+male should have any knowledge of the end to which his strenuous
+activity leads up. In presence of the female there is an elaborate
+application of all the energies of behaviour, just because ages of
+racial preparation have made him biologically and emotionally what he
+is--a functionally sexual male that must dance or sing or go through
+hereditary movements of display, when the appropriate stimulation
+comes. Of course after the first successful courtship his future
+behaviour will be in some degree modified by his previous experience.
+No doubt during his first courtship he is gaining the primary data of
+a peculiarly rich experience, instinctive and emotional. But the
+biological foundations of the behaviour of courtship are laid in the
+hereditary coordinations. It would seem that in some cases, not indeed
+in all, perhaps especially in those cases in which secondary sexual
+behaviour is most highly evolved,--correlative with the ardour of the
+male is a certain amount of reluctance in the female. The pairing act
+on her part only takes place after prolonged stimulation, for
+affording which the behaviour of male courtship is the requisite
+presentation. The most vigourous, defiant and mettlesome male is
+preferred just because he alone affords a contributory stimulation
+adequate to evoke the pairing impulse with its attendant emotional
+tone.
+
+It is true that this places female preference or choice on a much
+lower psychological plane than Darwin in some passages seems to
+contemplate where, for example, he says that the female appreciates
+the display of the male and places to her credit a taste for the
+beautiful. But Darwin himself distinctly states[177] that "it is not
+probable that she consciously deliberates; but she is most excited or
+attracted by the most beautiful, or melodious, or gallant males." The
+view here put forward, which has been developed by Prof. Groos,[178]
+therefore seems to have Darwin's own sanction. The phenomena are not
+only biological; there are psychological elements as well. One can
+hardly suppose that the female is unconscious of the male's presence;
+the final yielding must surely be accompanied by heightened emotional
+tone. Whether we call it choice or not is merely a matter of
+definition of terms. The behaviour is in part determined by
+supplementary psychological values. Prof. Groos regards the coyness of
+females as "a most efficient means of preventing the too early and too
+frequent yielding to the sexual impulse."[179] Be that as it may, it
+is, in any case, if we grant the facts, a means through which male
+sexual behaviour with all its biological and psychological
+implications, is raised to a level otherwise perhaps unattainable by
+natural means, while in the female it affords opportunities for the
+development in the individual and evolution in the race of what we may
+follow Darwin in calling appreciation, if we empty this word of the
+aesthetic implications which have gathered round it in the mental life
+of man.
+
+Regarded from this standpoint of sexual selection, broadly considered,
+has probably been of great importance. The psychological
+accompaniments of the pairing situation have profoundly influenced the
+course of biological evolution and are themselves the outcome of that
+evolution.
+
+Darwin makes only passing reference to those modes of behaviour in
+animals which go by the name of play. "Nothing," he says,[180] "is
+more common than for animals to take pleasure in practising whatever
+instinct they follow at other times for some real good." This is one
+of the very numerous cases in which a hint of the master has served to
+stimulate research in his disciples. It was left to Prof. Groos to
+develop this subject on evolutionary lines and to elaborate in a
+masterly manner Darwin's suggestion. "The utility of play," he
+says,[181] "is incalculable. This utility consists in the practice and
+exercise it affords for some of the more important duties of
+life,"--that is to say, for the performance of activities which will
+in adult life be essential to survival. He urges[182] that "the play
+of young animals has its origin in the fact that certain very
+important instincts appear at a time when the animal does not
+seriously need them." It is, however, questionable whether any
+instincts appear at a time when they are not needed. And it is
+questionable whether the instinctive and emotional attitude of the
+play-fight, to take one example, can be identified with those which
+accompany fighting in earnest, though no doubt they are closely
+related and have some common factors. It is probable that play, as
+preparatory behaviour, differs in biological detail (as it almost
+certainly does in emotional attributes) from the earnest of after-life
+and that it has been evolved through differentiation and integration
+of the primary tissue of experience, as a preparation through which
+certain essential modes of skill may be acquired--those animals in
+which the preparatory play-propensity was not inherited in due force
+and requisite amount being subsequently eliminated in the struggle for
+existence. In any case there is little question that Prof. Groos is
+right in basing the play-propensity on instinctive foundations.[183]
+None the less, as he contends, the essential biological value of play
+is that it is a means of training the educable nerve-tissue, of
+developing that part of the brain which is modified by experience and
+which thus acquires new characters, of elaborating the secondary
+tissue of experience on the predetermined lines of instinctive
+differentiation and thus furthering the psychological activities which
+are included under the comprehensive term "intelligent."
+
+In _The Descent of Man_ Darwin dealt at some length with intelligence
+and the higher mental faculties.[184] His object, he says, is to show
+that there is no fundamental difference between man and the higher
+mammals in their mental faculties; that these faculties are variable
+and the variations tend to be inherited; and that under natural
+selection beneficial variations of all kinds will have been preserved
+and injurious ones eliminated.
+
+Darwin was too good an observer and too honest a man to minimise the
+"enormous difference" between the level of mental attainment of
+civilised man and that reached by any animal. His contention was that
+the difference, great as it is, is one of degree and not of kind. He
+realised that, in the development of the mental faculties of man, new
+factors in evolution have supervened--factors which play but a
+subordinate and subsidiary part in animal intelligence.
+Intercommunication by means of language, approbation and blame, and
+all that arises out of reflective thought, are but foreshadowed in the
+mental life of animals. Still he contends that these may be explained
+on the doctrine of evolution. He urges[185] "that man is variable in
+body and mind; and that the variations are induced, either directly or
+indirectly, by the same general causes, and obey the same general
+laws, as with the lower animals." He correlates mental development
+with the evolution of the brain.[186] "As the various mental faculties
+gradually developed themselves, the brain would almost certainly
+become larger. No one, I presume, doubts that the large proportion
+which the size of man's brain bears to his body, compared to the same
+proportion in the gorilla or orang, is closely connected with his
+higher mental powers." "With respect to the lower animals," he
+says,[187] "M. E. Lartet,[188] by comparing the crania of tertiary and
+recent mammals belonging to the same groups, has come to the
+remarkable conclusion that the brain is generally larger and the
+convolutions are more complex in the more recent form."
+
+Sir E. Ray Lankester has sought to express in the simplest terms the
+implications of the increase in size of the cerebrum. "In what," he
+asks, "does the advantage of a larger cerebral mass consist?" "Man,"
+he replies, "is born with fewer ready-made tricks of the
+nerve-centres--these performances of an inherited nervous mechanism so
+often called by the ill-defined term 'instincts'--than are the monkeys
+or any other animal. Correlated with the absence of inherited
+ready-made mechanism, man has a greater capacity of developing in the
+course of his individual growth similar nervous mechanisms (similar
+to but not identical with those of 'instinct') than any other
+animal.... The power of being educated--'educability' as we may term
+it--is what man possesses in excess as compared with the apes. I think
+we are justified in forming the hypothesis that it is this
+'educability' which is the correlative of the increased size of the
+cerebrum." There has been natural selection of the more educable
+animals, for "the character which we describe as 'educability' can be
+transmitted, it is a congenital character. But the _results_ of
+education can _not_ be transmitted. In each generation they have to be
+acquired afresh, and with increased 'educability' they are more
+readily acquired and a larger variety of them.... The fact is that
+there is no community between the mechanisms of instinct and the
+mechanisms of intelligence, and that the latter are later in the
+history of the evolution of the brain than the former and can only
+develop in proportion as the former become feeble and defective."[189]
+
+In this statement we have a good example of the further development of
+views which Darwin foreshadowed but did not thoroughly work out. It
+states the biological case clearly and tersely. Plasticity of
+behaviour in special accommodation to special circumstances is of
+survival value; it depends upon acquired characters; it is correlated
+with increase in size and complexity of the cerebrum; under natural
+selection therefore the larger and more complex cerebrum as the organ
+of plastic behaviour has been the outcome of natural selection. We
+have thus the biological foundations for a further development of
+genetic psychology.
+
+There are diversities of opinion, as Darwin showed, with regard to the
+range of instinct in man and the higher animals as contrasted with
+lower types. Darwin himself said[190] that "Man, perhaps, has somewhat
+fewer instincts than those possessed by the animals which come next to
+him in the series." On the other hand, Prof. Wm. James says[191] that
+man is probably the animal with most instincts. The true position is
+that man and the higher animals have fewer complete and self-sufficing
+instincts than those which stand lower in the scale of mental
+evolution, but that they have an equally large or perhaps larger mass
+of instinctive raw material which may furnish the stuff to be
+elaborated by intelligent processes. There is, perhaps, a greater
+abundance of the primary tissue of experience to be refashioned and
+integrated by secondary modification; there is probably the same
+differentiation in relation to the determining biological ends, but
+there is at the outset less differentiation of the particular and
+specific modes of behaviour. The specialised instinctive performances
+and their concomitant experience-complexes are at the outset more
+indefinite. Only through acquired connections, correlated with
+experience, do they become definitely organised.
+
+The full working-out of the delicate and subtle relationship of
+instinct and educability--that is, of the hereditary and the acquired
+factors in the mental life--is the task which lies before genetic and
+comparative psychology. They interact throughout the whole of life,
+and their interactions are very complex. No one can read the chapters
+of _The Descent of Man_ which Darwin devotes to a consideration of the
+mental characters of man and animals without noticing, on the one
+hand, how sedulous he is in his search for hereditary foundations,
+and, on the other hand, how fully he realises the importance of
+acquired habits of mind. The fact that educability itself has innate
+tendencies--is in fact a partially differentiated educability--renders
+the unravelling of the factors of mental progress all the more
+difficult.
+
+In his comparison of the mental powers of men and animals it was
+essential that Darwin should lay stress on points of similarity rather
+than on points of difference. Seeking to establish a doctrine of
+evolution, with its basal concept of continuity of process and
+community of character, he was bound to render clear and to emphasise
+the contention that the difference in mind between man and the higher
+animals, great as it is, is one of degree and not of kind. To this end
+Darwin not only recorded a large number of valuable observations of
+his own, and collected a considerable body of information from
+reliable sources, he presented the whole subject in a new light and
+showed that a natural history of mind might be written and that this
+method of study offered a wide and rich field for investigation. Of
+course those who regarded the study of mind only as a branch of
+metaphysics smiled at the philosophical ineptitude of the mere man of
+science. But the investigation, on natural history lines, has been
+prosecuted with a large measure of success. Much indeed still remains
+to be done; for special training is required, and the workers are
+still few. Promise for the future is however afforded by the fact that
+investigation is prosecuted on experimental lines and that something
+like organised methods of research are taking form. There is now but
+little reliance on casual observations recorded by those who have not
+undergone the necessary discipline in these methods. There is also
+some change of emphasis in formulating conclusions. Now that the
+general evolutionary thesis is fully and freely accepted by those who
+carry on such researches, more stress is laid on the differentiation
+of the stages of evolutionary advance than on the fact of their
+underlying community of nature. The conceptual intelligence which is
+especially characteristic of the higher mental procedure of man is
+more firmly distinguished from the perceptual intelligence which he
+shares with the lower animals--distinguished now as a higher product
+of evolution, no longer as differing in origin or different in kind.
+Some progress has been made, on the one hand in rendering an account
+of intelligent profiting by experience under the guidance of pleasure
+and pain in the perceptual field, on lines predetermined by
+instinctive differentiation for biological ends, and on the other hand
+in elucidating the method of conceptual thought employed, for
+example, by the investigator himself in interpreting the perceptual
+experience of the lower animals.
+
+Thus there is a growing tendency to realise more fully that there are
+two orders of educability--first an educability of the perceptual
+intelligence based on the biological foundation of instinct, and
+secondly an educability of the conceptual intelligence which
+refashions and rearranges the data afforded by previous inheritance
+and acquisition. It is in relation to this second and higher order of
+educability that the cerebrum of man shows so large an increase of
+mass and a yet larger increase of effective surface through its rich
+convolutions. It is through educability of this order that the human
+child is brought intellectually and affectively into touch with the
+ideal constructions by means of which man has endeavoured, with more
+or less success, to reach an interpretation of nature, and to guide
+the course of the further evolution of his race--ideal constructions
+which form part of man's environment.
+
+It formed no part of Darwin's purpose to consider, save in broad
+outline, the methods, or to discuss in any fulness of detail the
+results of the process by which a differentiation of the mental
+faculties of man from those of the lower animals has been brought
+about--a differentiation the existence of which he again and again
+acknowledges. His purpose was rather to show that, notwithstanding
+this differentiation, there is basal community in kind. This must be
+remembered in considering his treatment of the biological foundations
+on which man's systems of ethics are built. He definitely stated that
+he approached the subject "exclusively from the side of natural
+history."[192] His general conclusion is that the moral sense is
+fundamentally identical with the social instincts, which have been
+developed for the good of the community; and he suggests that the
+concept which thus enables us to interpret the biological ground-plan
+of morals also enables us to frame a rational ideal of the moral end.
+"As the social instincts," he says,[193] "both of man and the lower
+animals have no doubt been developed by nearly the same steps, it
+would be advisable, if found practicable, to use the same definition
+in both cases, and to take as the standard of morality, the general
+good or welfare of the community, rather than the general happiness."
+But the kind of community for the good of which the social instincts
+of animals and primitive men were biologically developed may be
+different from that which is the product of civilisation, as Darwin no
+doubt realised. Darwin's contention was that conscience is a social
+instinct and has been evolved because it is useful to the tribe in the
+struggle for existence against other tribes. One the other hand J. S.
+Mill urged that the moral feelings are not innate but acquired, and
+Bain held the same view, believing that the moral sense is acquired by
+each individual during his life-time. Darwin, who notes[194] their
+opinion with his usual candour, adds that "on the general theory of
+evolution this is at least extremely improbable." It is impossible to
+enter into the question here: much turns on the exact connotation of
+the terms "conscience" and "moral sense," and on the meaning we attach
+to the statement that the moral sense is fundamentally identical with
+the social instincts.
+
+Presumably the majority of those who approach the subjects discussed
+in the third, fourth, and fifth chapters of _The Descent of Man_ in
+the full conviction that mental phenomena, not less than organic
+phenomena, have a natural genesis, would, without hesitation, admit
+that the intellectual and moral systems of civilised man are ideal
+constructions, the products of conceptual thought, and that as such
+they are, in their developed form, acquired. The moral sentiments are
+the emotional analogues of highly developed concepts. This does not
+however imply that they are outside the range of natural history
+treatment. Even though it may be desirable to differentiate the moral
+conduct of men from the social behaviour of animals (to which some
+such term as "pre-moral" or "quasi-moral" may be applied), still the
+fact remains that, as Darwin showed, there is abundant evidence of the
+occurrence of such social behaviour--social behaviour which, even
+granted that it is in large part intelligently acquired, and is itself
+so far a product of educability, is of survival value. It makes for
+that integration without which no social group could hold together and
+escape elimination. Furthermore, even if we grant that such behaviour
+is intelligently acquired, that is to say arises through the
+modification of hereditary instincts and emotions, the fact remains
+that only through these instinctive and emotional data is afforded the
+primary tissue of the experience which is susceptible of such
+modification.
+
+Darwin sought to show, and succeeded in showing, that for the
+intellectual and moral life there are instinctive foundations which a
+biological treatment alone can disclose. It is true that he did not in
+all cases analytically distinguish the foundations from the
+superstructure. Even to-day we are scarcely in a position to do so
+adequately. But his treatment was of great value in giving an impetus
+to further research. This value indeed can scarcely be over-estimated.
+And when the natural history of the mental operations shall have been
+written, the cardinal fact will stand forth, that the instinctive and
+emotional foundations are the outcome of biological evolution and have
+been ingrained in the race through natural selection. We shall more
+clearly realise that educability itself is a product of natural
+selection, though the specific results acquired through cerebral
+modifications are not transmitted through heredity. It will, perhaps,
+also be realised that the instinctive foundations of social behaviour
+are, for us, somewhat out of date and have undergone but little change
+throughout the progress of civilisation, because natural selection has
+long since ceased to be the dominant factor in human progress. The
+history of human progress has been mainly the history of man's higher
+educability, the products of which he has projected on to his
+environment. This educability remains on the average what it was a
+dozen generations ago; but the thought-woven tapestry of his
+surroundings is refashioned and improved by each succeeding
+generation. Few men have in greater measure enriched the
+thought-environment with which it is the aim of education to bring
+educable human beings into vital contact, than has Charles Darwin. His
+special field of work was the wide province of biology; but he did
+much to help us to realise that mental factors have contributed to
+organic evolution and that in man, the highest product of Evolution,
+they have reached a position of unquestioned supremacy.
+
+FOOTNOTES:
+
+[Footnote 153: _Origin of Species_ (6th edit.), p. 205.]
+
+[Footnote 154: _Descent of man_ (2nd edit. 1888), Vol. I. p. 99;
+Popular edit. p. 99.]
+
+[Footnote 155: _Ibid._ p. 99.]
+
+[Footnote 156: _The Expression of the Emotions_ (2nd edit.), p. 32.]
+
+[Footnote 157: _Descent of Man_, Vol. II. p. 435.]
+
+[Footnote 158: _Ibid._ 437, 438.]
+
+[Footnote 159: _Origin of Species_ (6th edit.), p. 205.]
+
+[Footnote 160: _Origin of Species_ (6th edit.), p. 205.]
+
+[Footnote 161: _Ibid._ p. 233.]
+
+[Footnote 162: _Ibid._ p. 205.]
+
+[Footnote 163: _Origin of Species_ (6th edit.), p. 233.]
+
+[Footnote 164: _Origin of Species_, pp. 210, 211.]
+
+[Footnote 165: Independently suggested, on somewhat different lines,
+by Profs. J. Mark Baldwin, Henry F. Osborn and the writer.]
+
+[Footnote 166: _Origin of Species_ (6th edit.), p. 206.]
+
+[Footnote 167: _Expression of the Emotions_, p. 13. The passage is
+here somewhat condensed.]
+
+[Footnote 168: _Ibid._ p. 368.]
+
+[Footnote 169: _Expression of the Emotions_, pp. 373, 374.]
+
+[Footnote 170: _Ibid._ p. 368.]
+
+[Footnote 171: _Ibid._ p. 369.]
+
+[Footnote 172: _Expression of the Emotions_, pp. 65 ff.]
+
+[Footnote 173: Cf. William James, _Principles of Psychology_, Vol. II.
+Chap. XXV, New York, 1890.]
+
+[Footnote 174: _Darwinism_, pp. 282, 283, London, 1889.]
+
+[Footnote 175: _Ibid._ p. 283.]
+
+[Footnote 176: _Darwinism_, pp. 283, 284.]
+
+[Footnote 177: _Descent of Man_ (2nd edit.), Vol. II. pp. 136, 137;
+(Popular edit.), pp. 642, 643.]
+
+[Footnote 178: _The Play of Animals_, p. 244, London, 1898.]
+
+[Footnote 179: _Ibid._ p. 283.]
+
+[Footnote 180: _Descent of Man_, Vol. II. p. 60; (Popular edit.), p.
+566.]
+
+[Footnote 181: _The Play of Animals_, p. 76.]
+
+[Footnote 182: _Ibid._ p. 75.]
+
+[Footnote 183: _The Play of Animals_ p. 24.]
+
+[Footnote 184: _Descent of Man_ (1st edit.), Chaps. II, III, V; (2nd
+edit.), Chaps. III, IV, V.]
+
+[Footnote 185: _Descent of Man_, Vol. I. pp. 70, 71; (Popular edit.),
+pp. 70, 71.]
+
+[Footnote 186: _Ibid._ p. 81.]
+
+[Footnote 187: _Ibid._ (Popular edit.), p. 82.]
+
+[Footnote 188: _Comptes Rendus des Sciences_, June 1, 1868.]
+
+[Footnote 189: _Nature_, Vol. LXI. pp. 624, 625 (1900).]
+
+[Footnote 190: _Descent of Man_, Vol. I. p. 100.]
+
+[Footnote 191: _Principles of Psychology_, Vol. II. p. 289.]
+
+[Footnote 192: _Descent of Man_, Vol. I. p. 149.]
+
+[Footnote 193: _Descent of Man_, p. 185.]
+
+[Footnote 194: _Ibid._ p. 150 (footnote).]
+
+
+
+
+VII
+
+THE INFLUENCE OF THE CONCEPTION OF EVOLUTION ON MODERN PHILOSOPHY
+
+BY H. HOeFFDING
+
+_Professor of Philosophy in the University of Copenhagen_
+
+
+I
+
+It is difficult to draw a sharp line between philosophy and natural
+science. The naturalist who introduces a new principle, or
+demonstrates a fact which throws a new light on existence, not only
+renders an important service to philosophy but is himself a
+philosopher in the broader sense of the word. The aim of philosophy in
+the stricter sense is to attain points of view from which the
+fundamental phenomena and the principles of the special sciences can
+be seen in their relative importance and connection. But philosophy in
+this stricter sense has always been influenced by philosophy in the
+broader sense. Greek philosophy came under the influence of logic and
+mathematics, modern philosophy under the influence of natural science.
+The name of Charles Darwin stands with those of Galileo, Newton, and
+Robert Mayer--names which denote new problems and great alterations in
+our conception of the universe.
+
+First of all we must lay stress on Darwin's own personality. His deep love
+of truth, his indefatigable inquiry, his wide horizon, and his steady
+self-criticism make him a scientific model, even if his results and
+theories should eventually come to possess mainly an historical interest.
+In the intellectual domain the primary object is to reach high summits
+from which wide surveys are possible, to reach them toiling honestly
+upwards by the way of experience, and then not to turn dizzy when a summit
+is gained. Darwinians have sometimes turned dizzy, but Darwin never. He saw
+from the first the great importance of his hypothesis, not only because of
+its solution of the old problem as to the value of the concept of species,
+not only because of the grand picture of natural evolution which it
+unrolls, but also because of the life and inspiration its method would
+impart to the study of comparative anatomy, of instinct and of heredity,
+and finally because of the influence it would exert on the whole conception
+of existence. He wrote in his note-book in the year 1837: "My theory would
+give zest to recent and fossil comparative anatomy; it would lead to the
+study of instinct, heredity, and mind-heredity, whole [of]
+metaphysics."[195]
+
+We can distinguish four main points in which Darwin's investigations
+possess philosophical importance.
+
+The evolution hypothesis is much older than Darwin; it is, indeed, one
+of the oldest guessings of human thought. In the eighteenth century is
+was put forward by Diderot and Lamettrie and suggested by Kant (1786).
+As we shall see later, it was held also by several philosophers in the
+first half of the nineteenth century. In his preface to _The Origin of
+Species_, Darwin mentions the naturalists who were his forerunners.
+But he has set forth the hypothesis of evolution in so energetic and
+thorough a manner that it perforce attracts the attention of all
+thoughtful men in a much higher degree than it did before the
+publication of the _Origin_.
+
+And further, the importance of his teaching rests on the fact that he,
+much more than his predecessors, even than Lamarck, sought a
+foundation for his hypothesis in definite facts. Modern science began
+by demanding--with Kepler and Newton--evidence of _varae causae_; this
+demand Darwin industriously set himself to satisfy--hence the wealth
+of material which he collected by his observations and his
+experiments. He not only revived an old hypothesis, but he saw the
+necessity of verifying it by facts. Whether the special cause on which
+he founded the explanation of the origin of species--Natural
+Selection--is sufficient, is now a subject of discussion. He himself
+had some doubt in regard to this question, and the criticisms which
+are directed against his hypothesis hit Darwinism rather than Darwin.
+In his indefatigable search for empirical evidence he is a model even
+for his antagonists: he has compelled them to approach the problems of
+life along other lines than those which were formerly followed.
+
+Whether the special cause to which Darwin appealed is sufficient or not, at
+least to it is probably due the greater part of the influence which he has
+exerted on the general trend of thought. "Struggle for existence" and
+"natural selection" are principles which have been applied, more or less,
+in every department of thought. Recent research, it is true, has discovered
+greater empirical discontinuity--leaps, "mutations"--whereas Darwin
+believed in the importance of small variations slowly accumulated. It has
+also been shown by the experimental method, which in recent biological work
+has succeeded Darwin's more historical method, that types once constituted
+possess great permanence, the fluctuations being restricted within clearly
+defined boundaries. The problem has become more precise, both as to
+variation and as to heredity. The inner conditions of life have in both
+respects shown a greater independence than Darwin had supposed in his
+theory, though he always admitted that the cause of variation was to him a
+great enigma, "a most perplexing problem," and that the struggle for life
+could only occur where variation existed. But, at any rate, it was of the
+greatest importance that Darwin gave a living impression of the struggle
+for life which is everywhere going on, and to which even the highest forms
+of existence must be amenable. The philosophical importance of these ideas
+does not stand or fall with the answer to the question, whether natural
+selection is a sufficient explanation of the origin of species or not it
+has an independent, positive value for everyone who will observe life and
+reality with an unbiased mind.
+
+In accentuating the struggle for life Darwin stands as a
+characteristically English thinker: he continues a train of ideas
+which Hobbes and Malthus had already begun. Moreover in his critical
+views as to the conception of species he had English forerunners; in
+the middle ages Occam and Duns Scotus, in the eighteenth century
+Berkeley and Hume. In his moral philosophy, as we shall see later, he
+is an adherent of the school which is represented by Hutcheson, Home
+and Adam Smith. Because he is no philosopher in the stricter sense of
+the term, it is of great interest to see that his attitude of mind is
+that of the great thinkers of his nation.
+
+In considering Darwin's influence on philosophy we will begin with an
+examination of the attitude of philosophy to the conception of
+evolution at the time when _The Origin of Species_ appeared. We will
+then examine the effects which the theory of evolution, and especially
+the idea of the struggle for life, has had, and naturally must have,
+on the discussion of philosophical problems.
+
+
+II
+
+When _The Origin of Species_ appeared fifty years ago Romantic
+speculation, Schelling's and Hegel's philosophy, still reigned on the
+continent, while in England Positivism, the philosophy of Comte and
+Stuart Mill, represented the most important trend of thought. German
+speculation had much to say on evolution, it even pretended to be a
+philosophy of evolution. But then the word "evolution" was to be taken
+in an ideal, not in a real, sense. To speculative thought the forms
+and types of nature formed a system of ideas, within which any form
+could lead us by continuous transitions to any other. It was a
+classificatory system which was regarded as a divine world of thought
+or images, within which metamorphoses could go on--a condition
+comparable with that in the mind of the poet when one image follows
+another with imperceptible changes. Goethe's ideas of evolution, as
+expressed in his _Metamorphosen der Pflanzen und der Thiere_, belong
+to this category; it is, therefore, incorrect to call him a forerunner
+of Darwin. Schelling and Hegel held the same idea; Hegel expressly
+rejected the conception of a real evolution in time as coarse and
+materialistic. "Nature," he says, "is to be considered as a _system of
+stages_, the one necessarily arising from the other, and being the
+nearest truth of that from which it proceeds; but not in such a way
+that the one is _naturally_ generated by the other; on the contrary
+[their connection lies] in the inner idea which is the ground of
+nature. The _metamorphosis_ can be ascribed only to the notion as
+such, because it alone is evolution.... It has been a clumsy idea in
+the older as well as in the newer philosophy of nature, to regard the
+transformation and the transition from one natural form and sphere to
+a higher as an outward and actual production."[196]
+
+The only one of the philosophers of Romanticism who believed in a
+real, historical evolution, a real production of new species, was
+Oken.[197] Danish philosophers, such as Treschow (1812) and Sibbern
+(1846), have also broached the idea of an historical evolution of all
+living beings from the lowest to the highest. Schopenhauer's
+philosophy has a more realistic character than that of Schelling's and
+Hegel's, his diametrical opposites, although he also belongs to the
+romantic school of thought. His philosophical and psychological views
+were greatly influenced by French naturalists and philosophers,
+especially by Cabanis and Lamarck. He praises the "ever memorable
+Lamarck," because he laid so much stress on the "will to live." But he
+repudiates as a "wonderful error" the idea that the organs of animals
+should have reached their present perfection through a development in
+time, during the course of innumerable generations. It was, he said, a
+consequence of the low standard of contemporary French philosophy,
+that Lamarck came to the idea of the construction of living beings in
+time through succession![198]
+
+The positivistic stream of thought was not more in favour of a real
+evolution than was the Romantic school. Its aim was to adhere to
+positive facts: it looked with suspicion on far-reaching speculation.
+Comte laid great stress on the discontinuity found between the
+different kingdoms of nature, as well as within each single kingdom.
+As he regarded as unscientific every attempt to reduce the number of
+physical forces, so he rejected entirely the hypothesis of Lamarck
+concerning the evolution of species; the idea of species would in his
+eyes absolutely lose its importance if a transition from species to
+species under the influence of conditions of life were admitted. His
+disciples (Littre, Robin) continued to direct against Darwin the
+polemics which their master had employed against Lamarck. Stuart Mill,
+who, in the theory of knowledge, represented the empirical or
+positivistic movement in philosophy--like his English forerunners from
+Locke to Hume--founded his theory of knowledge and morals on the
+experience of the single individual. He sympathised with the theory of
+the original likeness of all individuals and derived their
+differences, on which he practically and theoretically laid much
+stress, from the influence both of experience and education, and,
+generally, of physical and social causes. He admitted an individual
+evolution, and, in the human species, an evolution based on social
+progress; but no physiological evolution of species. He was afraid
+that the hypothesis of heredity would carry us back to the old theory
+of "innate" ideas.
+
+Darwin was more empirical than Comte and Mill; experience disclosed to
+him a deeper continuity than they could find; closer than before the
+nature and fate of the single individual were shown to be interwoven
+in the great web binding the life of the species with nature as a
+whole. And the continuity which so many idealistic philosophers could
+find only in the world of thought, he showed to be present in the
+world of reality.
+
+
+III
+
+Darwin's energetic renewal of the old idea of evolution has its chief
+importance in strengthening the conviction of this real continuity in
+the world, of continuity in the series of form and events. It was a
+great support for all those who were prepared to base their conception
+of life on scientific grounds. Together with the recently discovered
+law of the conservation of energy, it helped to produce the great
+realistic movement which characterises the last third of the
+nineteenth century. After the decline of the Romantic movement people
+wished to have firmer ground under their feet and reality now asserted
+itself in a more emphatic manner than in the period of Romanticism. It
+was easy for Hegel to proclaim that "the real" was "the rational," and
+that "the rational" was "the real": reality itself existed for him
+only in the interpretation of ideal reason, and if there was anything
+which could not be merged in the higher unity of thought, then it was
+only an example of the "impotence of nature to hold to the idea." But
+now concepts are to be founded on nature and not on any system of
+categories too confidently deduced _a priori_. The new devotion to
+nature had its recompense in itself, because the new points of view
+made us see that nature could indeed "hold to ideas," though perhaps
+not to those which we had cogitated beforehand.
+
+A most important question for philosophers to answer was whether the
+new views were compatible with an idealistic conception of life and
+existence. Some proclaimed that we have now no need of any philosophy
+beyond the principles of the conservation of matter and energy and the
+principle of natural evolution: existence should and could be
+definitely and completely explained by the laws of material nature.
+But abler thinkers saw that the thing was not so simple. They were
+prepared to give the new views their just place and to examine what
+alterations the old views must undergo in order to be brought into
+harmony with the new data.
+
+The realistic character of Darwin's theory was shown not only in the
+idea of natural continuity, but also, and not least, in the idea of
+the cause whereby organic life advances step by step. This idea--the
+idea of the struggle for life--implied that nothing could persist, if
+it had no power to maintain itself under the given conditions. Inner
+value alone does not decide. Idealism was here put to its hardest
+trial. In continuous evolution it could perhaps still find an analogy
+to the inner evolution of ideas in the mind; but in the demand for
+power in order to struggle with outward conditions Realism seemed to
+announce itself in its most brutal form. Every form of Idealism had to
+ask itself seriously how it was going to "struggle for life" with this
+new Realism.
+
+We will now give a short account of the position which leading
+thinkers in different countries have taken up in regard to this
+question.
+
+I. Herbert Spencer was the philosopher whose mind was best prepared by his
+own previous thinking to admit the theory of Darwin to a place in his
+conception of the world. His criticism of the arguments which had been put
+forward against the hypothesis of Lamarck, showed that Spencer, as a young
+man, was an adherent to the evolution idea. In his _Social Statics_ (1850)
+he applied this idea to human life and moral civilisation. In 1852 he wrote
+an essay on _The Development Hypothesis_, in which he definitely stated his
+belief that the differentiation of species, like the differentiation within
+a single organism, was the result of development. In the first edition of
+his _Psychology_ (1855) he took a step which put him in opposition to the
+older English school (from Locke to Mill): he acknowledged "innate ideas"
+so far as to admit the tendency of acquired habits to be inherited in the
+course of generations, so that the nature and functions of the individual
+are only to be understood through its connection with the life of the
+species. In 1857, in his essay on _Progress_, he propounded the law of
+differentiation as a general law of evolution, verified by examples from
+all regions of experience, the evolution of species being only one of these
+examples. On the effect which the appearance of _The Origin of Species_ had
+on his mind he writes in his _Autobiography_: "Up to that time ... I held
+that the sole cause of organic evolution is the inheritance of
+functionally-produced modifications. The _Origin of Species_ made it clear
+to me that I was wrong, and that the larger part of the facts cannot be due
+to any such cause.... To have the theory of organic evolution justified was
+of course to get further support for that theory of evolution at large with
+which ... all my conceptions were bound up."[199] Instead of the
+metaphorical expression "natural selection," Spencer introduced the term
+"survival of the fittest," which found favour with Darwin as well as with
+Wallace.
+
+In working out his ideas of evolution, Spencer found that
+differentiation was not the only form of evolution. In its simplest
+form evolution is mainly a concentration, previously scattered
+elements being integrated and losing independent movement.
+Differentiation is only forthcoming when minor wholes arise within a
+greater whole. And the highest form of evolution is reached when there
+is a harmony between concentration and differentiation, a harmony
+which Spencer calls equilibration and which he defines as a moving
+equilibrium. At the same time this definition enables him to
+illustrate the expression "survival of the fittest." "Every living
+organism exhibits such a moving equilibrium--a balanced set of
+functions constituting its life; and the overthrow of this balanced
+set of functions or moving equilibrium is what we call death. Some
+individuals in a species are so constituted that their moving
+equilibria are less easily overthrown than those of other
+individuals; and these are the fittest which survive, or, in Mr.
+Darwin's language, they are the select which nature preserves."[200]
+Not only in the domain of organic life, but in all domains, the summit
+of evolution is, according to Spencer, characterised by such a
+harmony--by a moving equilibrium.
+
+Spencer's analysis of the concept of evolution, based on a great
+variety of examples, has made this concept clearer and more definite
+than before. It contains the three elements; integration,
+differentiation and equilibration. It is true that a concept which is
+to be valid for all domains of experience must have an abstract
+character, and between the several domains there is, strictly
+speaking, only a relation of analogy. So there is only analogy between
+psychical and physical evolution. But this is no serious objection,
+because general concepts do not express more than analogies between
+the phenomena which they represent. Spencer takes his leading forms
+from the material world in defining evolution (in the simplest form)
+as integration of matter and dissipation of movement; but as he--not
+always quite consistently[201]--assumed a correspondence of mind and
+matter, he could very well give these terms an indirect importance for
+psychical evolution. Spencer has always, in my opinion with full
+right, repudiated the ascription of materialism. He is no more a
+materialist than Spinoza. In his _Principles of Psychology_ (Sec. 63) he
+expressed himself very clearly: "Though it seems easier to translate
+so-called matter into so-called spirit, than to translate so-called
+spirit into so-called matter--which latter is indeed wholly
+impossible--yet no translation can carry us beyond our symbols." These
+words lead us naturally to a group of thinkers whose starting-point
+was psychical evolution. But we have still one aspect of Spencer's
+philosophy to mention.
+
+Spencer founded his "laws of evolution" on an inductive basis, but he
+was convinced that they could be deduced from the law of the
+conservation of energy. Such a deduction is, perhaps, possible for the
+more elementary forms of evolution, integration and differentiation;
+but it is not possible for the highest form, the equilibration, which
+is a harmony of integration and differentiation. Spencer can no more
+deduce the necessity for the eventual appearance of "moving
+equilibria" of harmonious totalities than Hegel could guarantee the
+"higher unities" in which all contradictions should be reconciled. In
+Spencer's hands the theory of evolution acquired a more decidedly
+optimistic character than in Darwin's; but I shall deal later with the
+relation of Darwin's hypothesis to the opposition of optimism and
+pessimism.
+
+II. While the starting-point of Spencer was biological or
+cosmological, psychical evolution being conceived as in analogy with
+physical, a group of eminent thinkers--in Germany Wundt, in France
+Fouillee, in Italy Ardigo--took, each in his own manner, their
+starting-point in psychical evolution as an original fact and as a
+type of all evolution, the hypothesis of Darwin coming in as a
+corroboration and as a special example. They maintain the continuity
+of evolution; they find this character most prominent in psychical
+evolution, and this is for them a motive to demand a corresponding
+continuity in the material, especially in the organic domain.
+
+To Wundt and Fouillee the concept of will is prominent. They see the
+type of all evolution in the transformation of the life of will from
+blind impulse to conscious choice; the theories of Lamarck and Darwin
+are used to support the view that there is in nature a tendency to
+evolution in steady reciprocity with external conditions. The struggle
+for life is here only a secondary fact. Its apparent prominence is
+explained by the circumstance that the influence of external
+conditions is easily made out, while inner conditions can be verified
+only through their effects. For Ardigo the evolution of thought was
+the starting-point and the type: in the evolution of a scientific
+hypothesis we see a progress from the indefinite (_indistinto_) to the
+definite (_distinto_), and this is a characteristic of all evolution,
+as Ardigo has pointed out in a series of works. The opposition between
+_indistinto_ and _distinto_ corresponds to Spencer's opposition
+between homogeneity and heterogeneity. The hypothesis of the origin of
+differences of species from more simple forms is a special example of
+the general law of evolution.
+
+In the views of Wundt and Fouillee we find the fundamental idea of
+idealism psychical phenomena as expressions of the innermost nature of
+existence. They differ from the older Idealism in the great stress
+which they lay on evolution as a real, historical process which is
+going on through steady conflict with external conditions. The
+Romantic dread of reality is broken. It is beyond doubt that Darwin's
+emphasis on the struggle for life as a necessary condition of
+evolution has been a very important factor in carrying philosophy back
+to reality from the heaven of pure ideas. The philosophy of Ardigo, on
+the other side, appears more as a continuation and deepening of
+positivism, though the Italian thinker arrived at his point of view
+independently of French-English positivism. The idea of continuous
+evolution is here maintained in opposition to Comte's and Mill's
+philosophy of discontinuity. From Wundt and Fouillee Ardigo differs in
+conceiving psychical evolution not as an immediate revelation of the
+innermost nature of existence, but only as a single, though the most
+accessible example, of evolution.
+
+III. To the French philosophers Boutroux and Bergson, evolution proper
+is continuous and qualitative, while outer experience and physical
+science give us fragments only, sporadic processes and mechanical
+combinations. To Bergson, in his recent work _L'Evolution Creatrice_,
+evolution consists in an _elan de vie_ which to our fragmentary
+observation and analytic reflexion appears as broken into a manifold
+of elements and processes. The concept of matter in its scientific
+form is the result of this breaking asunder, essential for all
+scientific reflexion. In these conceptions the strongest opposition
+between inner and outer conditions of evolution is expressed: in the
+domain of internal conditions spontaneous development of qualitative
+forms--in the domain of external conditions discontinuity and
+mechanical combination.
+
+We see, then, that the theory of evolution has influenced philosophy
+in a variety of forms. It has made idealistic thinkers revise their
+relation to the real world; it has led positivistic thinkers to find a
+closer connection between the facts on which they based their views;
+it has made us all open our eyes for new possibilities to arise
+through the _prima facie_ inexplicable "spontaneous" variations which
+are the condition of all evolution. This last point is one of peculiar
+interest. Deeper than speculative philosophy and mechanical science
+saw in the days of their triumph, we catch sight of new streams, whose
+sources and laws we have still to discover. Most sharply does this
+appear in the theory of mutation, which is only a stronger
+accentuation of a main point in Darwinism. It is interesting to see
+that an analogous problem comes into the foreground in physics through
+the discovery of radioactive phenomena, and in psychology through the
+assumption of psychical new formations (as held by Boutroux, William
+James and Bergson). From this side, Darwin's ideas, as well as the
+analogous ideas in other domains, incite us to renewed examination of
+our first principles, their rationality and their value. On the other
+hand, his theory of the struggle for existence challenges us to
+examine the conditions and discuss the outlook as to the persistence
+of human life and society and of the values that belong to them. It is
+not enough to hope (or fear?) the rising of new forms; we have also to
+investigate the possibility of upholding the forms and ideals which
+have hitherto been the bases of human life. Darwin has here given his
+age the most earnest and most impressive lesson. This side of Darwin's
+theory is of peculiar interest to some special philosophical problems
+to which I now pass.
+
+
+IV
+
+Among philosophical problems the problem of knowledge has in the last
+century occupied a foremost place. It is natural, then, to ask how
+Darwin and the hypothesis whose most eminent representative he is,
+stand to this problem.
+
+Darwin started an hypothesis. But every hypothesis is won by inference
+from certain presuppositions, and every inference is based on the
+general principles of human thought. The evolution hypothesis
+presupposes, then, human thought and its principles. And not only the
+abstract logical principles are thus pre-supposed. The evolution
+hypothesis purports to be not only a formal arrangement of phenomena,
+but to express also the law of a real process. It supposes, then, that
+the real data--all that in our knowledge which we do not produce
+ourselves, but which we in the main simply receive--are subject to
+laws which are at least analogous to the logical relations of our
+thoughts; in other words, it assumes the validity of the principle of
+causality. If organic species could arise without cause there would be
+no use in framing hypotheses. Only if we assume the principle of
+causality, is there a problem to solve.
+
+Though Darwinism has had a great influence on philosophy considered as
+a striving after a scientific view of the world, yet here is a point
+of view--the epistemological--where philosophy is not only independent
+but reaches beyond any result of natural science. Perhaps it will be
+said: the powers and functions of organic beings only persist (perhaps
+also only arise) when they correspond sufficiently to the conditions
+under which the struggle of life is to go on. Human thought itself is,
+then, a variation (or a mutation) which has been able to persist and
+to survive. Is not, then, the problem of knowledge solved by the
+evolution hypothesis? Spencer had given an affirmative answer to this
+question before the appearance of _The Origin of Species_. For the
+individual, he said, there is an _a priori_, original, basis (or
+_Anlage_) for all mental life; but in the species all powers have
+developed in reciprocity with extendal conditions. Knowledge is here
+considered from the practical point of view, as a weapon in the
+struggle for life, as an "organon" which has been continuously in use
+for generations. In recent years the economic or pragmatic
+epistemology, as developed by Avenarius and Mach in Germany, and by
+James in America, points in the same direction. Science, it is said,
+only maintains those principles and presuppositions which are
+necessary to the simplest and clearest orientation be applied to
+experience and to practical work, will successively be eliminated.
+
+In these views a striking and important application is made of the
+idea of struggle for life to the development of human thought. Thought
+must, as all other things in the world, struggle for life. But this
+whole consideration belongs to psychology, not to the theory of
+knowledge (epistemology), which is concerned only with the validity of
+knowledge, not with its historical origin. Every hypothesis to explain
+the origin of knowledge must submit to cross-examination by the theory
+of knowledge, because it works with the fundamental forms and
+principles of human thought. We cannot go further back than these
+forms and principles, which it is the aim of epistemology to ascertain
+and for which no further reason can be given.[202]
+
+But there is another side of the problem which is, perhaps, of more
+importance and which epistemology generally overlooks. If new
+variations can arise, not only in organic but perhaps also in
+inorganic nature, new tasks are placed before the human mind. The
+question is, then, if it has forms in which there is room for the new
+matter? We are here touching a possibility which the great master of
+epistemology did not bring to light. Kant supposed confidently that no
+other matter of knowledge could stream forth from the dark source
+which he called "the thing-in-itself," than such as could be
+synthesised in our existing forms of knowledge. He mentions the
+possibility of other forms than the human, and warns us against the
+dogmatic assumption that the human conception of existence should be
+absolutely adequate. But he seems to be quite sure that the
+thing-in-itself works constantly, and consequently always gives us
+only what our powers can master. This assumption was a consequence of
+Kant's rationalistic tendency, but one for which no warrant can be
+given. Evolutionism and systematism are opposing tendencies which can
+never be absolutely harmonised one with the other. Evolution may at
+any time break some form which the system-monger regards as finally
+established. Darwin himself felt a great difference in looking at
+variation as an evolutionist and as a systematist. When he was working
+at his evolution theory, he was very glad to find variations; but they
+were a hindrance to him when he worked as a systematist, in preparing
+his work on Cirripedia. He says in a letter: "I had thought the same
+parts of the same species more resemble (than they do anyhow in
+Cirripedia) objects cast in the same mould. Systematic work would be
+easy were it not for this confounded variation, which, however, is
+pleasant to me as a speculatist, though odious to me as a
+systematist."[203] He could indeed be angry with variations even as an
+evolutionist; but then only because he could not explain them, not
+because he could not classify them. "If, as I must think, external
+conditions produce little _direct_ effect, what the devil determines
+each particular variation?"[204] What Darwin experienced in this
+particular domain holds good of all knowledge. All knowledge is
+systematic, in so far as it strives to put phenomena in quite definite
+relations, one to another. But the systematisation can never be
+complete. And here Darwin has contributed much to widen the world, for
+us. He has shown us forces and tendencies in nature which make
+absolute systems impossible, at the same time that they give us new
+objects and problems. There is still a place for what Lessing called
+"the unceasing striving after truth," while "absolute truth" (in the
+sense of a closed system) is unattainable so long as life and
+experience are going on.
+
+There is here a special remark to be made. As we have seen above,
+recent research has shown that natural selection or struggle for life
+is no explanation of variations. Hugo de Vries distinguishes between
+partial and embryonal variations, or between variations and mutations,
+only the last-named being heritable, and therefore of importance for
+the origin of new species. But the existence of variations is not only
+of interest for the problem of the origin of species; it has also a
+more general interest. An individual does not lose its importance for
+knowledge, because its qualities are not heritable. On the contrary,
+in higher beings at least, individual peculiarities will become more
+and more independent objects of interest. Knowledge takes account of
+the biographies not only of species, but also of individuals: it seeks
+to find the law of development of the single individual.[205] As
+Leibnitz said long ago, individuality consists in the law of the
+changes of a being: "La loi du changement fait l'individualite de
+chaque substance." Here is a world which is almost new for science,
+which till now has mainly occupied itself with general laws and forms.
+But these are ultimately only means to understand the individual
+phenomena, in whose nature and history a manifold of laws and forms
+always cooeperate. The importance of this remark will appear in the
+sequel.
+
+
+V
+
+To many people the Darwinian theory of natural selection or struggle
+for existence seemed to change the whole conception of life, and
+particularly all the conditions on which the validity of ethical ideas
+depends. If only that has persistence which can be adapted to a given
+condition, what will then be the fate of our ideals, of our standards
+of good and evil? Blind force seems to reign, and the only thing that
+counts seems to be the most heedless use of power. Darwinism, it was
+said, has proclaimed brutality. No other difference seems permanent
+save that between the sound, powerful and happy on the one side, the
+sick, feeble and unhappy on the other; and every attempt to alleviate
+this difference seems to lead to general enervation. Some of those who
+interpreted Darwinism in this manner felt an aesthetic delight in
+contemplating the heedlessness and energy of the great struggle for
+existence and anticipated the realisation of a higher human type as
+the outcome of it: so Nietzsche and his followers. Others recognising
+the same consequences in Darwinism regarded these as one of the
+strongest objections against it; so Duehring and Kropotkin (in his
+earlier works).
+
+This interpretation of Darwinism was frequent in the interval between
+the two main works of Darwin--_The Origin of Species_ and _The Descent
+of Man_. But even during this interval it was evident to an attentive
+reader that Darwin himself did not found his standard of good and evil
+on the features of the life of nature he had emphasised so strongly.
+He did not justify the ways along which nature reached its ends; he
+only pointed them out. The "real" was not to him, as to Hegel, one
+with the "rational." Darwin has, indeed, by his whole conception of
+nature, rendered a great service to ethics in making the difference
+between the life of nature and the ethical life appear in so strong a
+light. The ethical problem could now be stated in a sharper form than
+before. But this was not the first time that the idea of the struggle
+for life was put in relation to the ethical problem. In the
+seventeenth century Thomas Hobbes gave the first impulse to the whole
+modern discussion of ethical principles in his theory of _bellum
+omnium contra omnes_. Men, he taught, are in the state of nature
+enemies one of another, and they live either in fright or in the glory
+of power. But it was not the opinion of Hobbes that this made ethics
+impossible. On the contrary, he found a standard for virtue and vice
+in the fact that some qualities and actions have a tendency to bring
+us out of the state of war and to secure peace, while other qualities
+have a contrary tendency. In the eighteenth century even Immanuel
+Kant's ideal ethics had--so far as can be seen--a similar origin.
+Shortly before the foundation of his definitive ethics, Kant wrote his
+_Idee zu einer allgemeinen Weltgeschichte_ (1784), where--in a way
+which reminds us of Hobbes, and is prophetic of Darwin--he describes
+the forward-driving power of struggle in the human world. It is here
+as with the struggle of the trees for light and air, through which
+they compete with one another in height. Anxiety about war can only be
+allayed by an ordinance which gives everyone his full liberty under
+acknowledgment of the equal liberty of others. And such ordinance and
+acknowledgment are also attributes of the content of the moral law, as
+Kant proclaimed it in the year after the publication of his essay
+(1785).[206] Kant really came to his ethics by the way of evolution,
+though he afterwards disavowed it. Similarly the same line of thought
+may be traced in Hegel though it has been disguised in the form of
+speculative dialectics.[207] And in Schopenhauer's theory of the blind
+will to live and its abrogation by the ethical feeling, which is
+founded on universal sympathy, we have a more individualistic form of
+the same idea.
+
+It was, then, not entirely a foreign point of view which Darwin
+introduced into ethical thought, even if we take no account of the
+poetical character of the word "struggle" and of the more direct
+adaptation, through the use and non-use of power, which Darwin also
+emphasised. In _The Descent of Man_ he has devoted a special
+chapter[208] to a discussion of the origin of the ethical
+consciousness. The characteristic expression of this consciousness he
+found, just as Kant did, in the idea of "ought"; it was the origin of
+this new idea which should be explained. His hypothesis was that the
+ethical "ought" has its origin in the social and parental instincts,
+which, as well as other instincts (e.g. the instinct of
+self-preservation), lie deeper than pleasure and pain. In many
+species, not least in the human species, these instincts are fostered
+by natural selection; and when the powers of memory and comparison are
+developed, so that single acts can be valued according to the claims
+of the deep social instinct, then consciousness of duty and remorse
+are possible. Blind instinct has developed to conscious ethical will.
+
+As already stated, Darwin, as a moral philosopher belongs to the
+school that was founded by Shaftesbury, and was afterwards represented
+by Hutcheson, Hume, Adam Smith, Comte and Spencer. His merit is,
+first, that he has given this tendency of thought a biological
+foundation, and that he has stamped on it a doughty character in
+showing that ethical ideas and sentiments, rightly conceived, are
+forces which are at work in the struggle for life.
+
+There are still many questions to solve. Not only does the ethical
+development within the human species contain features still
+unexplained;[209] but we are confronted by the great problem whether
+after all a genetic historical theory can be of decisive importance
+here. To every consequent ethical consciousness there is a standard of
+value, a primordial value which determines the single ethical
+judgments as their last presupposition, and the "rightness" of this
+basis, the "value" of this value can as little be discussed as the
+"rationality" of our logical principles. There is here revealed a
+possibility of ethical scepticism which evolutionistic ethics (as well
+as intuitive or rationalistic ethics) has overlooked. No demonstration
+can show that the results of the ethical development are definitive
+and universal. We meet here again with the important opposition of
+systematisation and evolution. There will, I think, always be an open
+question here, though comparative ethics, of which we have so far only
+the first attempts, can do much to throw light on it.
+
+It would carry us too far to discuss all the philosophical works on
+ethics, which have been influenced directly or indirectly by
+evolutionism. I may, however, here refer to the book of C. M.
+Williams, _A Review of the Systems of Ethics founded on the Theory of
+Evolution_,[210] in which, besides Darwin, the following authors are
+reviewed: Wallace, Haeckel, Spencer, Fiske, Rolph, Barratt, Stephen,
+Carneri, Hoeffding, Gizycki, Alexander, Ree. As works which criticise
+evolutionistic ethics from an intuitive point of view and in an
+instructive way, may be cited: Guyau, _La morale anglaise
+contemporaine_,[211] and Sorley, _Ethics of Naturalism_. I will only
+mention some interesting contributions to ethical discussion which can
+be found in Darwinism besides the idea of struggle for life.
+
+The attention which Darwin has directed to variations has opened our
+eyes to the differences in human nature as well as in nature
+generally. There is here a fact of great importance for ethical
+thought, no matter from what ultimate premiss it starts. Only from a
+very abstract point of view can different individuals be treated in
+the same manner. The most eminent ethical thinkers, men such as Jeremy
+Bentham and Immanuel Kant, who discussed ethical questions from very
+opposite standpoints, agreed in regarding all men as equal in respect
+of ethical endowment. In regard to Bentham, Leslie Stephen remarks:
+"He is determined to be thoroughly empirical, to take men as he found
+them. But his utilitarianism supposed that men's views of happiness
+and utility were uniform and clear, and that all that was wanted was
+to show them the means by which their ends could be reached."[212] And
+Kant supposed that every man would find the "categorical imperative"
+in his consciousness, when he came to sober reflexion, and that all
+would have the same qualifications to follow it. But if continual
+variations, great or small, are going on in human nature, it is the
+duty of ethics to make allowance for them, both in making claims, and
+in valuing what is done. A new set of ethical problems have their
+origin here.[213] It is an interesting fact that Stuart Mill's book
+_On Liberty_ appeared in the same year as _The Origin of Species_.
+Though Mill agreed with Bentham about the original equality of all
+men's endowments, he regarded individual differences as a necessary
+result of physical and social influences, and he claimed that free
+play shall be allowed to differences of character so far as is
+possible without injury to other men. It is a condition of individual
+and social progress that a man's mode of action should be determined
+by his own character and not by tradition and custom, nor by abstract
+rules. This view was to be corroborated by the theory of Darwin.
+
+But here we have reached a point of view from which the criticism,
+which in recent years has often been directed against Darwin--that
+small variations are of no importance in the struggle for life--is of
+no weight. From an ethical standpoint, and particularly from the
+ethical standpoint of Darwin himself, it is a duty to foster
+individual differences that can be valuable, even though they can
+neither be of service for physical preservation nor be physically
+inherited. The distinction between variation and mutation is here
+without importance. It is quite natural that biologists should be
+particularly interested in such variations as can be inherited and
+produce new species. But in the human world there is not only a
+physical, but also a mental and social heredity. When an ideal human
+character has taken form, then there is shaped a type, which through
+imitation and influence can become an important factor in subsequent
+development, even if it cannot form a species in the biological sense
+of the word. Spiritually strong men often succumb in the physical
+struggle for life; but they can nevertheless be victorious through the
+typical influence they exert, perhaps on very distant generations, if
+the remembrance of them is kept alive, be it in legendary or in
+historical form. Their very failure can show that a type has taken
+form which is maintained at all risks, a standard of life which is
+adhered to in spite of the strongest opposition. The question "to be
+or not to be" can be put from very different levels of being: it has
+too often been considered a consequence of Darwinism that this
+question is only to be put from the lowest level. When a stage is
+reached, where ideal (ethical, intellectual, aesthetic) interests are
+concerned, the struggle for life is a struggle for the preservation of
+this stage. The giving up of a higher standard of life is a sort of
+death; for there is not only a physical, there is also a spiritual,
+death.
+
+
+VI
+
+The Socratic character of Darwin's mind appears in his wariness in
+drawing the last consequences of his doctrine, in contrast both with
+the audacious theories of so many of his followers and with the
+consequences which his antagonists were busy in drawing. Though he, as
+we have seen, saw from the beginning that his hypothesis would
+occasion "a whole of metaphysics," he was himself very reserved as to
+the ultimate questions, and his answers to such questions were
+extorted from him.
+
+As to the question of optimism and pessimism, Darwin held that though
+pain and suffering were very often the ways by which animals were led
+to pursue that course of action which is most beneficial to the
+species, yet pleasurable feelings were the most habitual guides. "We
+see this in the pleasure from exertion, even occasionally from great
+exertion of the body or mind, in the pleasure of our daily meals, and
+especially in the pleasure derived from sociability, and from loving
+our families." But there was to him so much suffering in the world
+that it was a strong argument against the existence of an intelligent
+First Cause.[214]
+
+It seems to me that Darwin was not so clear on another question, that
+of the relation between improvement and adaptation. He wrote to Lyell:
+"When you contrast natural selection and 'improvement,' you seem
+always to overlook ... that every step in the natural selection of
+each species implies improvement in that species _in relation to its
+condition of life_.... Improvement implies, I suppose, _each form
+obtaining many parts or organs_, all excellently adapted for their
+functions." "All this," he adds, "seems to me quite compatible with
+certain forms fitted for simple conditions, remaining unaltered, or
+being, degraded."[215] But the great question is, if the conditions of
+life will in the long run favour "improvement" in the sense of
+differentiation (or harmony of differentiation and integration). Many
+beings are best adapted to their conditions of life if they have few
+organs and few necessities. Pessimism would not only be the
+consequence, if suffering outweighed happiness, but also if the most
+elementary forms of happiness were predominant, or if there were a
+tendency to reduce the standard of life to the simplest possible, the
+contentment of inertia or stable equilibrium. There are animals which
+are very highly differentiated and active in their young state, but
+later lose their complex organisation and concentrate themselves on
+the one function of nutrition. In the human world analogies to this
+sort of adaptation are not wanting. Young "idealists" very often end
+as old "Philistines." Adaptation and progress are not the same.
+
+Another question of great importance in respect to human evolution is,
+whether there will be always a possibility for the existence of an
+impulse to progress, an impulse to make great claims on life, to be
+active and to alter the conditions of life instead of adapting to them
+in a passive manner. Many people do not develop because they have too
+few necessities, and because they have no power to imagine other
+conditions of life than those under which they live. In his remarks on
+"the pleasure from exertion" Darwin has a point of contact with the
+practical idealism of former times--with the ideas of Lessing and
+Goethe, of Condorcet and Fichte. The continual striving which was the
+condition of salvation to Faust's soul, is also the condition of
+salvation to mankind. There is a holy fire which we ought to keep
+burning, if adaptation is really to be improvement. If, as I have
+tried to show in my _Philosophy of Religion_, the innermost core of
+all religion is faith in the persistence of value in the world, and if
+the highest values express themselves in the cry "Excelsior!" then the
+capital point is, that this cry should always be heard and followed.
+We have here a corollary of the theory of evolution in its application
+to human life.
+
+Darwin declared himself an agnostic, not only because he could not
+harmonise the large amount of suffering in the world with the idea of
+a God as its first cause, but also because he "was aware that if we
+admit a first cause, the mind still craves to know whence it came and
+how it arose."[216] He saw, as Kant had seen before him and expressed
+in his _Kritik der Urtheilskraft_, that we cannot accept either of the
+only two possibilities which we are able to conceive: chance (or brute
+force) and design. Neither mechanism nor teleology can give an
+absolute answer to ultimate questions. The universe, and especially
+the organic life in it, can neither be explained as a mere
+combination of absolute elements nor as the effect of a constructing
+thought. Darwin concluded, as Kant, and before him Spinoza, that the
+oppositions and distinctions which our experience presents, cannot
+safely be regarded as valid for existence in itself. And, with Kant
+and Fichte, he found his stronghold in the conviction that man has
+something to do, even if he cannot solve all enigmas. "The safest
+conclusion seems to me that the whole subject is beyond the scope of
+man's intellect; but man can do his duty."[217]
+
+Is this the last word of human thought? Does not the possibility, that
+man can do his duty, suppose that the conditions of life allow of
+continuous ethical striving, so that there is a certain harmony
+between cosmic order and human ideals? Darwin himself has shown how
+the consciousness of duty can arise as a natural result of evolution.
+Moreover there are lines of evolution which have their end in ethical
+idealism, in a kingdom of values, which must struggle for life as all
+things in the world must do, but a kingdom which has its firm
+foundation in reality.
+
+FOOTNOTES:
+
+[Footnote 195: _Life and Letters of Charles Darwin_, Vol. I. p. 8.]
+
+[Footnote 196: _Encyclopaedie der philosophischen Wissenschaften_ (4th
+edit.), Berlin, 1845, Sec. 249.]
+
+[Footnote 197: _Lehrbuch der Naturphilosophie_, Jena, 1809.]
+
+[Footnote 198: _Ueber den Willen in der Natur_ (2nd edit.), Frankfurt
+a. M., 1854, pp. 41-43.]
+
+[Footnote 199: Spencer, _Autobiography_, Vol. II. p. 50, London and
+New York, 1904.]
+
+[Footnote 200: _Autobiography_, Vol. II. p. 100.]
+
+[Footnote 201: Cf. my letter to him 1876, now printed in Duncan's
+_Life and Letters of Herbert Spencer_, p. 178. London, 1908.]
+
+[Footnote 202: The present writer, many years ago, in his _Psychology_
+(Copenhagen, 1882; Eng. transl. London, 1891), criticised the
+evolutionistic treatment of the problem of knowledge from the Kantian
+point of view.]
+
+[Footnote 203: _Life and Letters_, Vol. II. p. 37.]
+
+[Footnote 204: _Ibid._ p. 232.]
+
+[Footnote 205: The new science of Ecology occupies an intermediate
+position between the biography of species and the biography of
+individuals. Compare _Congress of Arts and Science_, St. Louis, Vol.
+V. 1906 (The Reports of Drude and Robinson) and the work of my
+colleague, E. Warming.]
+
+[Footnote 206: Cf. my _History of Modern Philosophy_ (Eng. transl.
+London, 1900), I. pp. 76-79.]
+
+[Footnote 207: "Herrschaft und Knechtschaft," _Phoenomenologie des
+Geistes_, IV. A., Leiden, 1907.]
+
+[Footnote 208: _The Descent of Man_, Vol. I. Ch. iii.]
+
+[Footnote 209: The works of Westermarck and Hobhouse throw new light
+on many of these features.]
+
+[Footnote 210: New York and London, 1893.]
+
+[Footnote 211: Paris, 1879.]
+
+[Footnote 212: _English literature and society in the eighteenth
+century_, London, 1904, p. 187.]
+
+[Footnote 213: Cf. my paper, "The law of relativity in Ethics,"
+_International Journal of Ethics_, Vol. I. 1891, pp. 37-62.]
+
+[Footnote 214: _Life and Letters_, Vol. I. p. 310.]
+
+[Footnote 215: _Ibid._ Vol. II. p. 177.]
+
+[Footnote 216: _Life and Letters_, Vol. 1. p. 306.]
+
+[Footnote 217: _Life and Letters_, p. 307.]
+
+
+
+
+VIII
+
+THE INFLUENCE OF DARWIN UPON RELIGIOUS THOUGHT
+
+BY P. N. WAGGETT, M.A., S.S.J.E.
+
+
+I
+
+The object of this paper is first to point out certain elements of the
+Darwinian influence upon Religious thought, and then to show reason
+for the conclusion that it has been, from a Christian point of view,
+satisfactory. I shall not proceed further to urge that the Christian
+apologetic in relation to biology has been successful. A variety of
+opinions may be held on this question, without disturbing the
+conclusion that the movements of readjustment have been beneficial to
+those who remain Christians, and this by making them more Christian
+and not only more liberal. The theologians may sometimes have
+retreated, but there has been an advance of theology. I know that this
+account incurs the charge of optimism. It is not the worst that could
+be made. The influence has been limited in personal range, unequal,
+even divergent, in operation, and accompanied by the appearance of
+waste and mischievous products. The estimate which follows requires
+for due balance a full development of many qualifying considerations.
+For this I lack space, but I must at least distinguish my view from
+the popular one that our difficulties about religion and natural
+science have come to an end.
+
+Concerning the older questions about origins--the origin of the
+world, of species, of man, of reason, conscience, religion--a large
+measure of understanding has been reached by some thoughtful men. But
+meanwhile new questions have arisen, questions about conduct,
+regarding both the reality of morals and the rule of right action for
+individuals and societies. And these problems, still far from
+solution, may also be traced to the influence of Darwin. For they
+arise from the renewed attention to heredity, brought about by the
+search for the causes of variation, without which the study of the
+selection of variations has no sufficient basis.
+
+Even the existing understanding about origins is very far from
+universal. On these points there were always thoughtful men who denied
+the necessity of conflict, and there are still thoughtful men who deny
+the possibility of a truce.
+
+It must further be remembered that the earlier discussion now, as I
+hope to show, producing favourable results, created also for a time
+grave damage, not only in the disturbance of faith and the loss of
+men--a loss not repaired by a change in the currents of debate--but in
+what I believe to be a still more serious respect. I mean the
+introduction of a habit of facile and untested hypothesis in religious
+as in other departments of thought.
+
+Darwin is not responsible for this, but he is in part the cause of it.
+Great ideas are dangerous guests in narrow minds; and thus it has
+happened that Darwin--the most patient of scientific workers, in whom
+hypothesis waited upon research, or if it provisionally outstepped it
+did so only with the most scrupulously careful acknowledgment--has led
+smaller and less conscientious men in natural science, in history, and
+in theology to an over-eager confidence in probable conjecture and a
+loose grip upon the facts of experience. It is not too much to say
+that in many quarters the age of materialism was the least
+matter-of-fact age conceivable, and the age of science the age which
+showed least of the patient temper of inquiry.
+
+I have indicated, as shortly as I could, some losses and dangers
+which in a balanced account of Darwin's influence would be discussed
+at length.
+
+One other loss must be mentioned. It is a defect in our thought which,
+in some quarters, has by itself almost cancelled all the advantages
+secured. I mean the exaggerated emphasis on uniformity or continuity;
+the unwillingness to rest any part of faith or of our practical
+expectation upon anything that from any point of view can be called
+exceptional. The high degree of success reached by naturalists in
+tracing, or reasonably conjecturing, the small beginnings of great
+differences, has led the inconsiderate to believe that anything may in
+time become anything else.
+
+It is true that this exaggeration of the belief in uniformity has
+produced in turn its own perilous reaction. From refusing to believe
+whatever can be called exceptional, some have come to believe whatever
+can be called wonderful.
+
+But, on the whole, the discontinuous or highly various character of
+experience received for many years too little deliberate attention.
+The conception of uniformity which is a necessity of scientific
+description has been taken for the substance of history. We have
+accepted a postulate of scientific method as if it were a conclusion
+of scientific demonstration. In the name of a generalisation which,
+however just on the lines of a particular method, is the prize of a
+difficult exploit of reflexion, we have discarded the direct
+impressions of experience; or, perhaps it is more true to say, we have
+used for the criticism of alleged experiences a doctrine of uniformity
+which is only valid in the region of abstract science. For every
+science depends for its advance upon limitation of attention, upon the
+selection out of the whole content of consciousness of that part or
+aspect which is measurable by the method of the science. Accordingly
+there is a science of life which rightly displays the unity underlying
+all its manifestations. But there is another view of life, equally
+valid, and practically sometimes more important, which recognises the
+immediate and lasting effect of crisis, difference, and revolution.
+Our ardour for the demonstration of uniformity of process and of
+minute continuous change needs to be balanced by a recognition of the
+catastrophic element in experience, and also by a recognition of the
+exceptional significance for us of events which may be perfectly
+regular from an impersonal point of view.
+
+An exorbitant jealousy of miracle, revelation, and ultimate moral
+distinctions has been imported from evolutionary science into
+religious thought. And it has been a damaging influence, because it
+has taken men's attention from facts, and fixed them upon theories.
+
+
+II
+
+With this acknowledgment of important drawbacks, requiring many words
+for their proper description, I proceed to indicate certain results of
+Darwin's doctrine which I believe to be in the long run wholly
+beneficial to Christian thought. These are:
+
+The encouragement in theology of that evolutionary method of
+observation and study, which has shaped all modern research:
+
+The recoil of Christian apologetics towards the ground of religious
+experience, a recoil produced by the pressure of scientific criticism
+upon other supports of faith:
+
+The restatement, or the recovery of ancient forms of statement, of the
+doctrines of Creation and of divine Design in Nature, consequent upon
+the discussion of evolution and of natural selection as its guiding
+factor.
+
+(1) The first of these is quite possibly the most important of all. It
+was well defined in a notable paper read by Dr. Gore, now Bishop of
+Birmingham, to the Church Congress at Shrewsbury in 1896. We have
+learnt a new caution both in ascribing and in denying significance to
+items of evidence, in utterance or in event. There has been, as in
+art, a study of values, which secures perspective and solidity in our
+representation of facts. On the one hand, a given utterance or event
+cannot be drawn into evidence as if all items were of equal
+consequence, like sovereigns in a bag. The question whence and whither
+must be asked, and the particular thing measured as part of a series.
+Thus measured it is not less truly important, but it may be important
+in a lower degree. On the other hand, and for exactly the same reason,
+nothing that is real is unimportant. The "failures" are not mere
+mistakes. We see them, in St. Augustine's words, as "scholar's faults
+which men praise in hope of fruit."
+
+We cannot safely trace the origin of the evolutionistic method to the
+influence of natural science. The view is tenable that theology led
+the way. Probably this is a case of alternate and reciprocal debt.
+Quite certainly the evolutionist method in theology, in Christian
+history and in the estimate of scripture, has received vast
+reinforcement from biology, in which evolution has been the ever
+present and ever victorious conception.
+
+(2) The second effect named is the new willingness of Christian
+thinkers to take definite account of religious experience. This is
+related to Darwin through the general pressure upon religious faith of
+scientific criticism. The great advance of our knowledge of organisms
+has been an important element in the general advance of science. It
+has acted, by the varied requirements of the theory of organisms, upon
+all other branches of natural inquiry, and it held for a long time
+that leading place in public attention which is now occupied by
+speculative physics. Consequently it contributed largely to our
+present estimation of science as the supreme judge in all matters of
+inquiry,[218] to the supposed destruction of mystery and the
+disparagement of metaphysics which marked the last age, as well as to
+the just recommendation of scientific method in branches of learning
+where the direct acquisitions of natural science had no place.
+
+Besides this, the new application of the idea of law and mechanical
+regularity to the organic world seemed to rob faith of a kind of
+refuge. The romantics had, as Berthelot[219] shows, appealed to life
+to redress the judgments drawn from mechanism. Now, in Spencer,
+evolution gave us a vitalist mechanic or mechanical vitalism, and the
+appeal seemed cut off. We may return to this point later when we
+consider evolution; at present I only endeavour to indicate that
+general pressure of scientific criticism which drove men of faith to
+seek the grounds of reassurance in a science of their own; in a method
+of experiment, of observation, of hypothesis checked by known facts.
+It is impossible for me to do more than glance across the threshold of
+this subject. But it is necessary to say that the method is in an
+elementary stage of revival. The imposing success that belongs to
+natural science is absent: we fall short of the unchallengeable
+unanimity of the Biologists on fundamentals. The experimental method
+with its sure repetitions cannot be applied to our subject-matter. But
+we have something like the observational method of palaeontology and
+geographical distribution; and in biology there are still men who
+think that the large examination of varieties by way of geography and
+the search of strata is as truly scientific, uses as genuinely the
+logical method of difference, and is as fruitful in sure conclusions
+as the quasi-chemical analysis of Mendelian laboratory work, of which
+last I desire to express my humble admiration. Religion also has its
+observational work in the larger and possibly more arduous manner.
+
+But the scientific work in religion makes its way through difficulties
+and dangers. We are far from having found the formula of its
+combination with the historical elements of our apologetic. It is
+exposed, therefore, to a damaging fire not only from unspiritualist
+psychology and pathology but also from the side of scholastic dogma.
+It is hard to admit on equal terms a partner to the old undivided rule
+of books and learning. With Charles Lamb, we cry in some distress,
+"must knowledge come to me, if it come at all, by some awkward
+experiment of intuition, and no longer by this familiar process of
+reading?"[220] and we are answered that the old process has an
+imperishable value, only we have not yet made clear its connection
+with other contributions. And all the work is young, liable to be
+drawn into unprofitable excursions, side-tracked by self-deceit and
+pretence; and it fatally attracts, like the older mysticism, the
+curiosity and the expository powers of those least in sympathy with
+it, ready writers who, with all the air of extended research, have
+been content with narrow grounds for induction. There is a danger,
+besides, which accompanies even the most genuine work of this science
+and must be provided against by all its serious students. I mean the
+danger of unbalanced introspection both for individuals and for
+societies; of a preoccupation comparable to our modern social
+preoccupation with bodily health; of reflexion upon mental states not
+accompanied by exercise and growth of the mental powers; the danger of
+contemplating will and neglecting work, of analysing conviction and
+not criticising evidence.
+
+Still, in spite of dangers and mistakes, the work remains full of
+hopeful indications, and, in the best examples,[221] it is truly
+scientific in its determination to know the very truth, to tell what
+we think, not what we think we ought to think,[222] truly scientific
+in its employment of hypothesis and verification, and in growing
+conviction of the reality of its subject-matter through the repeated
+victories of a mastery which advances, like science, in the Baconian
+road of obedience. It is reasonable to hope that progress in this
+respect will be more rapid and sure when religious study enlists more
+men affected by scientific desire and endowed with scientific
+capacity.
+
+The class of investigating minds is a small one, possibly even smaller
+than that of reflecting minds. Very few persons at any period are able
+to find out anything whatever. There are few observers, few
+discoverers, few who even wish to discover truth. In how many
+societies the problems of philology which face every person who speaks
+English are left unattempted! And if the inquiring or the successfully
+inquiring class of minds is small, much smaller, of course, is the
+class of those possessing the scientific aptitude in an eminent
+degree. During the last age this most distinguished class was to a
+very great extent absorbed in the study of phenomena, a study which
+had fallen into arrears. For we stood possessed, in rudiment, of means
+of observation, means for travelling and acquisition, qualifying men
+for a larger knowledge than had yet been attempted. These were now to
+be directed with new accuracy and ardour upon the fabric and behaviour
+of the world of sense. Our debt to the great masters in physical
+science who overtook and almost outstripped the task cannot be
+measured; and, under the honourable leadership of Ruskin, we may all
+well do penance if we have failed "in the respect due to their great
+powers of thought, or in the admiration due to the far scope of their
+discovery."[223] With what miraculous mental energy and divine good
+fortune--as Romans said of their soldiers--did our men of curiosity
+face the apparently impenetrable mysteries of nature! And how natural
+it was that immense accessions of knowledge, unrelated to the
+spiritual facts of life, should discredit Christian faith, by the
+apparent superiority of the new work to the feeble and unprogressive
+knowledge of Christian believers! The day is coming when men of this
+mental character and rank, of this curiosity, this energy and this
+good fortune in investigation, will be employed in opening mysteries
+of a spiritual nature. They will silence with masterful witness the
+over-confident denials of naturalism. They will be in danger of the
+widespread recognition which thirty years ago accompanied every
+utterance of Huxley, Tyndall, Spencer. They will contribute, in spite
+of adulation, to the advance of sober religious and moral science.
+
+And this result will be due to Darwin, first because by raising the
+dignity of natural science, he encouraged the development of the
+scientific mind; secondly because he gave to religious students the
+example of patient and ardent investigation; and thirdly because by
+the pressure of naturalistic criticism the religious have been driven
+to ascertain the causes of their own convictions, a work in which they
+were not without the sympathy of men of science.[224]
+
+In leaving the subject of scientific religious inquiry, I will only
+add that I do not believe it receives any important help--and
+certainly it suffers incidentally much damaging interruption--from the
+study of abnormal manifestations or abnormal conditions of
+personality.
+
+(3) Both of the above effects seem to me of high, perhaps the very
+highest, importance to faith and to thought. But, under the third
+head, I name two which are more directly traceable to the personal
+work of Darwin, and more definitely characteristic of the age in which
+his influence was paramount: viz. the influence of the two conceptions
+of evolution and natural selection upon the doctrine of creation and
+of design respectively.
+
+It is impossible here, though it is necessary for a complete sketch of
+the matter, to distinguish the different elements and channels of this
+Darwinian influence; in Darwin's own writings, in the vigourous
+polemic of Huxley, and strangely enough, but very actually for popular
+thought, in the teaching of the definitely anti-Darwinian evolutionist
+Spencer.
+
+Under the head of the directly and purely Darwinian elements I should
+class as preeminent the work of Wallace and of Bates; for no two sets
+of facts have done more to fix in ordinary intelligent minds a belief
+in organic evolution and in natural selection as its guiding factor
+than the facts of geographical distribution and of protective colour
+and mimicry. The facts of geology were difficult to grasp and the
+public and theologians heard more often of the imperfection than of
+the extent of the geological record. The witness of embryology,
+depending to a great extent upon microscopic work, was and is beyond
+the appreciation of persons occupied in fields of work other than
+biology.
+
+
+III
+
+From the influence in religion of scientific modes of thought we pass
+to the influence of particular biological conceptions. The former
+effect comes by way of analogy, example, encouragement and challenge;
+inspiring or provoking kindred or similar modes of thought in the
+field of theology; the latter by a collision of opinions upon matters
+of fact or conjecture which seem to concern both science and religion.
+
+In the case of Darwinism the story of this collision is familiar, and
+falls under the heads of evolution and natural selection, the doctrine
+of descent with modification, and the doctrine of its guidance or
+determination by the struggle for existence between related varieties.
+These doctrines, though associated and interdependent, and in popular
+thought not only combined but confused, must be considered separately.
+It is true that the ancient doctrine of Evolution, in spite of the
+ingenuity and ardour of Lamarck, remained a dream tantalising the
+intellectual ambition of naturalists, until the day when Darwin made
+it conceivable by suggesting the machinery of its guidance. And,
+further, the idea of natural selection has so effectively opened the
+door of research and stimulated observation in a score of principal
+directions that, even if the Darwinian explanation became one day much
+less convincing than, in spite of recent criticism, it now is, yet its
+passing, supposing it to pass, would leave the doctrine of Evolution
+immeasurably and permanently strengthened. For in the interests of the
+theory of selection, "Fuer Darwin," as Mueller wrote, facts have been
+collected which remain in any case evidence of the reality of descent
+with modification.
+
+But still, though thus united in the modern history of convictions,
+though united and confused in the collision of biological and
+traditional opinion, yet evolution and natural selection must be
+separated in theological no less than in biological estimation.
+Evolution seemed inconsistent with Creation; natural selection with
+Providence and Divine design.
+
+Discussion was maintained about these points for many years and with
+much dark heat. It ranged over many particular topics and engaged
+minds different in tone, in quality, and in accomplishment. There was
+at most times a degree of misconception. Some naturalists attributed
+to theologians in general a poverty of thought which belonged really
+to men of a particular temper or training. The "timid theism"
+discerned in Darwin by so cautious a theologian as Liddon[225] was
+supposed by many biologists to be the necessary foundation of an
+honest Christianity. It was really more characteristic of devout
+_naturalists_ like Philip Henry Gosse, than of religious believers as
+such.[226] The study of theologians more considerable and even more
+typically conservative than Liddon does not confirm the description of
+religious intolerance given in good faith, but in serious ignorance,
+by a disputant so acute, so observant and so candid as Huxley.
+Something hid from each other's knowledge the devoted pilgrims in two
+great ways of thought. The truth may be, that naturalists took their
+view of what creation was from Christian men of science who naturally
+looked in their own special studies for the supports and illustrations
+of their religious belief. Of almost every labourious student it may
+be said: "_Hic ab arte sua non recessit_." And both the believing and
+the denying naturalists, confining habitual attention to a part of
+experience, are apt to affirm and deny with trenchant vigour and
+something of a narrow clearness "_Qui respiciunt ad pauca, de facili
+pronunciant_."[227]
+
+Newman says of some secular teachers that "they persuade the world of
+what is false by urging upon it what is true." Of some early opponents
+of Darwin it might be said by a candid friend that, in all sincerity
+of devotion to truth, they tried to persuade the world of what is true
+by urging upon it what is false. If naturalists took their version of
+orthodoxy from amateurs in theology, some conservative Christians,
+instead of learning what evolution meant to its regular exponents,
+took their view of it from celebrated persons, not of the front rank
+in theology or in thought, but eager to take account of public
+movements and able to arrest public attention.
+
+Cleverness and eloquence on both sides certainly had their share in
+producing the very great and general disturbance of men's minds in the
+early days of Darwinian teaching. But by far the greater part of that
+disturbance was due to the practical novelty and the profound
+importance of the teaching itself, and to the fact that the
+controversy about evolution quickly became much more public than any
+controversy of equal seriousness had been for many generations.
+
+We must not think lightly of that great disturbance because it has, in
+some real sense, done its work, and because it is impossible in days
+of more coolness and light, to recover a full sense of its very real
+difficulties.
+
+Those who would know them better should add to the calm records of
+Darwin[228] and to the story of Huxley's impassioned championship, all
+that they can learn of George Romanes.[229] For his life was absorbed
+in this very struggle and reproduced its stages. It began in a certain
+assured simplicity of biblical interpretation; it went on, through the
+glories and adventures of a paladin in Darwin's train, to the darkness
+and dismay of a man who saw all his most cherished beliefs rendered,
+as he thought, incredible.[230] He lived to find the freer faith for
+which process and purpose are not irreconcilable, but necessary to one
+another. His development, scientific, intellectual and moral, was
+itself of high significance; and its record is of unique value to our
+own generation, so near the age of that doubt and yet so far from it;
+certainly still much in need of the caution and courage by which past
+endurance prepares men for new emergencies. We have little enough
+reason to be sure that in the discussions awaiting us we shall do as
+well as our predecessors in theirs. Remembering their endurance of
+mental pain, their ardour in mental labour, the heroic temper and the
+high sincerity of controversialists on either side, we may well speak
+of our fathers in such words of modesty and self-judgment as Drayton
+used when he sang the victors of Agincourt. The progress of biblical
+study, in the departments of Introduction and Exegesis, resulting in
+the recovery of a point of view anciently tolerated if not prevalent,
+has altered some of the conditions of that discussion. In the years
+near 1858, the witness of Scripture was adduced both by Christian
+advocates and their critics as if unmistakably irreconcilable with
+Evolution.
+
+Huxley[231] found the path of the blameless naturalist everywhere
+blocked by "Moses": the believer in revelation was generally held to
+be forced to a choice between revealed cosmogony and the scientific
+account of origins. It is not clear how far the change in Biblical
+interpretation is due to natural science, and how far to the vital
+movements of theological study which have been quite independent of
+the controversy about species. It belongs to a general renewal of
+Christian movement, the recovery of a heritage. "Special
+Creation"--really a biological rather than a theological
+conception,--seems in its rigid form to have been a recent element
+even in English biblical orthodoxy.
+
+The Middle Ages had no suspicion that religious faith forbad inquiry
+into the natural origination of the different forms of life.
+Bartholomaeus Anglicus, an English Franciscan of the thirteenth
+century, was a mutationist in his way, as Aristotle, "the Philosopher"
+of the Christian Schoolmen, had been in his. So late as the
+seventeenth century, as we learn not only from early proceedings of
+the Royal Society, but from a writer so homely and so regularly pious
+as Walton, the variation of species and "spontaneous" generations had
+no theological bearing, except as instances of that various wonder of
+the world which in devout minds is food for devotion.
+
+It was in the eighteenth century that the harder statement took shape.
+Something in the preciseness of that age, its exaltation of law, its
+cold passion for a stable and measured universe, its cold denial, its
+cold affirmation of the power of God, a God of ice, is the occasion of
+that rigidity of religious thought about the living world which Darwin
+by accident challenged, or rather by one of those movements of genius
+which, Goethe[232] declares, are "elevated above all earthly control."
+
+If religious thought in the eighteenth century was aimed at a fixed
+and nearly finite world of spirit, it followed in all these respects
+the secular and critical lead. "La philosophie reformatrice du
+XVIII^{e} siecle[233] ramenait la nature et la societe a des
+mecanismes que la pensee reflechie peut concevoir et recomposer." In
+fact, religion in a mechanical age is condemned if it takes any but a
+mechanical tone. Butler's thought was too moving, too vital, too
+evolutionary, for the sceptics of his time. In a rationalist,
+encyclopaedic period, religion also must give hard outline to its
+facts, it must be able to display its secret to any sensible man in
+the language used by all sensible men. Milton's prophetic genius
+furnished the eighteenth century, out of the depth of the passionate
+age before it, with the theological tone it was to need. In spite of
+the austere magnificence of his devotion, he gives to smaller souls a
+dangerous lead. The rigidity of Scripture exegesis belonged to this
+stately but imperfectly sensitive mode of thought. It passed away with
+the influence of the older rationalists whose precise denials matched
+the precise and limited affirmations of the static orthodoxy.
+
+I shall, then, leave the specially biblical aspect of the
+debate--interesting as it is and even useful, as in Huxley's
+correspondence with the Duke of Argyll and others in 1892[234]--in
+order to consider without complication the permanent elements of
+Christian thought brought into question by the teaching of evolution.
+
+Such permanent elements are the doctrine of God as Creator of the
+universe, and the doctrine of man as spiritual and unique. Upon both
+the doctrine of evolution seemed to fall with crushing force.
+
+With regard to Man I leave out, acknowledging a grave omission, the
+doctrine of the Fall and of Sin. And I do so because these have not
+yet, as I believe, been adequately treated: here the fruitful reaction
+to the stimulus of evolution is yet to come. The doctrine of sin,
+indeed, falls principally within the scope of that discussion which
+has followed or displaced the Darwinian; and without it the Fall
+cannot be usefully considered. For the question about the Fall is a
+question not merely of origins, but of the interpretation of moral
+facts whose moral reality must first be established.
+
+I confine myself therefore to Creation and the dignity of man.
+
+The meaning of evolution, in the most general terms, is that the
+differentiation of forms is not essentially separate from their
+behaviour and use; that if these are within the scope of study, that
+is also; that the world has taken the form we see by movements not
+unlike those we now see in progress; that what may be called proximate
+origins are continuous in the way of force and matter, continuous in
+the way of life, with actual occurrences and actual characteristics.
+All this has no revolutionary bearing upon the question of ultimate
+origins. The whole is a statement about process. It says nothing to
+metaphysicians about cause. It simply brings within the scope of
+observation or conjecture that series of changes which has given their
+special characters to the different parts of the world we see. In
+particular, evolutionary science aspires to the discovery of the
+process or order of the appearance of life itself: if it were to
+achieve its aim it could say nothing of the cause of this or indeed of
+the most familiar occurrences. We should have become spectators or
+convinced historians of an event which, in respect of its cause and
+ultimate meaning, would be still impenetrable.
+
+With regard to the origin of species, supposing life already
+established, biological science has the well founded hopes and the
+measure of success with which we are all familiar. All this has, it
+would seem, little chance of collision with a consistent theism, a
+doctrine which has its own difficulties unconnected with any
+particular view of order or process. But when it was stated that
+species had arisen by processes through which new species were still
+being made, evolutionism came into collision with a statement,
+traditionally religious, that species were formed and fixed once for
+all and long ago.
+
+What is the theological import of such a statement when it is regarded
+as essential to belief in God? Simply that God's activity, with
+respect to the formation of living creatures, ceased at some point in
+past time.
+
+"God rested" is made the touchstone of orthodoxy. And when, under the
+pressure of the evidences, we found ourselves obliged to acknowledge
+and assert the present and persistent power of God, in the maintenance
+and in the continued formation of "types," what happened was the
+abolition of a time-limit. We were forced only to a bolder claim, to
+a theistic language less halting, more consistent, more thorough in
+its own line, as well as better qualified to assimilate and modify
+such schemes as Von Hartmann's philosophy of the Unconscious--a
+philosophy, by the way, quite intolerant of a merely mechanical
+evolution.[235]
+
+Here was not the retrenchment of an extravagant assertion, but the
+expansion of one which was faltering and inadequate. The traditional
+statement did not need paring down so as to pass the meshes of a new
+and exacting criticism. It was itself a net meant to surround and
+enclose experience; and we must increase its size and close its mesh
+to hold newly disclosed facts of life. The world, which had seemed a
+fixed picture or model, gained first perspective and then solidity and
+movement. We had a glimpse of organic _history_; and Christian thought
+became more living and more assured as it met the larger view of life.
+
+However unsatisfactory the new attitude might be to our critics, to
+Christians the reform was positive. What was discarded was a
+limitation, a negation. The movement was essentially conservative,
+even actually reconstructive. For the language disused was a language
+inconsistent with the definitions of orthodoxy; it set bounds to the
+infinite, and by implication withdrew from the creative rule all such
+processes as could be brought within the descriptions of research. It
+ascribed fixity and finality to that "creature" in which an apostle
+taught us to recognise the birth-struggles of an unexhausted progress.
+It tended to banish mystery from the world we see, and to confine it
+to a remote first age.
+
+In the reformed, the restored, language of religion, Creation became
+again not a link in a rational series to complete a circle of the
+sciences, but the mysterious and permanent relation between the
+infinite and the finite, between the moving changes we know in part,
+and the Power, after the fashion of that observation, unknown, which
+is itself "unmoved all motion's source."[236]
+
+With regard to man it is hardly necessary, even were it possible, to
+illustrate the application of this bolder faith. When the record of
+his high extraction fell under dispute, we were driven to a
+contemplation of the whole of his life, rather than of a part and that
+part out of sight. We remembered again, out of Aristotle, that the
+result of a process interprets its beginnings. We were obliged to read
+the title of such dignity as we may claim, in results and still more
+in aspirations.
+
+Some men still measure the value of great present facts in
+life--reason and virtue and sacrifice--by what a self-disparaged
+reason can collect of the meaner rudiments of these noble gifts. Mr.
+Balfour has admirably displayed the discrepancy, in this view, between
+the alleged origin and the alleged authority of reason. Such an
+argument ought to be used not to discredit the confident reason, but
+to illuminate and dignify its dark beginnings, and to show that at
+every step in the long course of growth a Power was at work which is
+not included in any term or in all the terms of the series.
+
+I submit that the more men know of actual Christian teaching, its
+fidelity to the past, and its sincerity in face of discovery, the more
+certainly they will judge that the stimulus of the doctrine of
+evolution has produced in the long run vigour as well as flexibility
+in the doctrine of Creation and of man.
+
+I pass from Evolution in general to Natural Selection.
+
+The character in religious language which I have for short called
+mechanical was not absent in the argument from design as stated before
+Darwin. It seemed to have reference to a world conceived as fixed. It
+pointed, not to the plastic capacity and energy of living matter, but
+to the fixed adaptation of this and that organ to an unchanging place
+or function.
+
+Mr. Hobhouse has given us the valuable phrase "a niche of organic
+opportunity." Such a phrase would have borne a different sense in
+non-evolutionary thought. In that thought, the opportunity was an
+opportunity for the Creative Power, and Design appeared in the
+preparation of the organism to fit the niche. The idea of the niche
+and its occupant growing together from simpler to more complex mutual
+adjustment was unwelcome to this teleology. If the adaptation was
+traced to the influence, through competition, of the environment, the
+old teleology lost an illustration and a proof. For the cogency of the
+proof in every instance depended upon the absence of explanation.
+Where the process of adaptation was discerned, the evidence of Purpose
+or Design was weak. It was strong only when the natural antecedents
+were not discovered, strongest when they could be declared
+undiscoverable.
+
+Paley's favourite word is "Contrivance"; and for him contrivance is
+most certain where production is most obscure. He points out the
+physiological advantage of the _valvulae conniventes_ to man, and the
+advantage for teleology of the fact that they cannot have been formed
+by "action and pressure." What is not due to pressure may be
+attributed to design, and when a "mechanical" process more subtle than
+pressure was suggested, the case for design was so far weakened. The
+cumulative proof from the multitude of instances began to disappear
+when, in selection, a natural sequence was suggested in which all the
+adaptations might be reached by the motive power of life, and
+especially when, as in Darwin's teaching, there was full recognition
+of the reactions of life to the stimulus of circumstance. "The
+organism fits the niche," said the teleologist, "because the Creator
+formed it so as to fit." "The organism fits the niche," said the
+naturalist, "because unless it fitted it could not exist." "It was
+fitted to survive," said the theologian. "It survives because it
+fits," said the selectionist. The two forms of statement are not
+incompatible; but the new statement, by provision of an ideally
+universal explanation of process, was hostile to a doctrine of purpose
+which relied upon evidences always exceptional however numerous.
+Science persistently presses on to find the universal machinery of
+adaptation in this planet; and whether this be found in selection, or
+in direct-effect, or in vital reactions resulting in large changes, or
+in a combination of these and other factors, it must always be opposed
+to the conception of a Divine Power here and there but not everywhere
+active.
+
+For science, the Divine must be constant, operative everywhere and in
+every quality and power, in environment and in organism, in stimulus
+and in reaction, in variation and in struggle, in hereditary
+equilibrium, and in "the unstable state of species"; equally present
+on both sides of every strain, in all pressures and in all
+resistances, in short in the general wonder of life and the world. And
+this is exactly what the Divine Power must be for religious faith.
+
+The point I wish once more to make is that the necessary readjustment
+of teleology, so as to make it depend upon the contemplation of the
+whole instead of a part, is advantageous quite as much to theology as
+to science. For the older view failed in courage. Here again our
+theism was not sufficiently theistic.
+
+Where results seemed inevitable, it dared not claim them as God-given.
+In the argument from Design it spoke not of God in the sense of
+theology, but of a Contriver, immensely, not infinitely wise and good,
+working within a world, the scene, rather than the ever dependent
+outcome, of His Wisdom; working in such emergencies and opportunities
+as occurred, by forces not altogether within His control, towards an
+end beyond Himself. It gave us, instead of the awful reverence due to
+the Cause of all substance and form, all love and wisdom, a
+dangerously detached appreciation of an ingenuity and benevolence
+meritorious in aim and often surprisingly successful in contrivance.
+
+The old teleology was more useful to science than to religion, and
+the design-naturalists ought to be gratefully remembered by
+Biologists. Their search for evidences led them to an eager study of
+adaptations and of minute forms, a study such as we have now an
+incentive to in the theory of Natural Selection. One hardly meets with
+the same ardour in microscopical research until we come to modern
+workers. But the argument from Design was never of great importance to
+faith. Still, to rid it of this character was worth all the stress and
+anxiety of the gallant old war. If Darwin had done nothing else for
+us, we are to-day deeply in his debt for this. The world is not less
+venerable to us now, not less eloquent of the causing mind, rather
+much more eloquent and sacred. But our wonder is not that "the
+underjaw of the swine works under the ground" or in any or all of
+those particular adaptations which Paley collected with so much skill,
+but that a purpose transcending, though resembling, our own purposes,
+is everywhere manifest; that what we live in is a whole, mutually
+sustaining, eventful and beautiful, where the "dead" forces feed the
+energies of life, and life sustains a stranger existence, able in some
+real measure to contemplate the whole, of which, mechanically
+considered, it is a minor product and a rare ingredient. Here, again,
+the change was altogether positive. It was not the escape of a vessel
+in a storm with loss of spars and rigging, not a shortening of sail to
+save the masts and make a port of refuge. It was rather the emergence
+from narrow channels to an open sea. We had propelled the great ship,
+finding purchase here and there for slow and uncertain movement. Now,
+in deep water, we spread large canvas to a favouring breeze.
+
+The scattered traces of design might be forgotten or obliterated. But
+the broad impression of Order became plainer when seen at due distance
+and in sufficient range of effect, and the evidence of love and wisdom
+in the universe could be trusted more securely for the loss of the
+particular calculation of their machinery.
+
+Many other topics of faith are affected by modern biology. In some of
+these we have learnt at present only a wise caution, a wise
+uncertainty. We stand before the newly unfolded spectacle of
+suffering, silenced; with faith not scientifically reassured but still
+holding fast certain other clues of conviction. In many important
+topics we are at a loss. But in others, and among them those I have
+mentioned, we have passed beyond this negative state and find faith
+positively strengthened and more fully expressed.
+
+We have gained also a language and a habit of thought more fit for the
+great and dark problems that remain, less liable to damaging
+conflicts, equipped for more rapid assimilation of knowledge. And by
+this change biology itself is a gainer. For, relieved of fruitless
+encounters with popular religion, it may advance with surer aim along
+the path of really scientific life-study which was reopened for modern
+men by the publication of _The Origin of Species_.
+
+Charles Darwin regretted that, in following science, he had not done
+"more direct good"[237] to his fellow-creatures. He has, in fact,
+rendered substantial service to interests bound up with the daily
+conduct and hopes of common men; for his work has led to improvements
+in the preaching of the Christian faith.
+
+FOOTNOTES:
+
+[Footnote 218: F. R. Tennant: "The Being of God in the light of
+Physical Science," in _Essays on some theological questions of the
+day_. London, 1905.]
+
+[Footnote 219: _Evolutionisme et Platonisme_, pp. 45, 46, 47. Paris,
+1908.]
+
+[Footnote 220: _Essays of Elia_, "New Year's Eve," p. 41; Ainger's
+edition. London, 1899.]
+
+[Footnote 221: Such an example is given in Baron F. von Huegel's
+recently finished book, the result of thirty years' research: _The
+Mystical Element of Religion, as studied in Saint Catherine of Genoa
+and her Friends_. London, 1908.]
+
+[Footnote 222: G. Tyrrell, in _Mediaevalism_, has a chapter which is
+full of the important _moral_ element in a scientific attitude. "The
+only infallible guardian of truth is the spirit of truthfulness."
+_Mediaevalism_, p. 182, London, 1908.]
+
+[Footnote 223: _Queen of the Air_, Preface, p. vii. London, 1906.]
+
+[Footnote 224: The scientific rank of its writer justifies the
+insertion of the following letter from the late Sir John
+Burdon-Sanderson to me. In the lecture referred to I had described the
+methods of Professor Moseley in teaching Biology as affording a
+suggestion of the scientific treatment of religion.
+
+OXFORD,
+
+_April 30, 1902_.
+
+DEAR SIR:
+
+     I feel that I must express to you my thanks for the
+     discourse which I had the pleasure of listening to yesterday
+     afternoon.
+
+     I do not mean to say that I was able to follow all that you
+     said as to the identity of Method in the two fields of
+     Science and Religion, but I recognise that the "mysticism"
+     of which you spoke gives us the only way by which the two
+     fields can be brought into relation.
+
+     Among much that was memorable, nothing interested me more
+     than what you said of Moseley.
+
+     No one, I am sure, knew better than you the value of his
+     teaching and in what that value consisted.
+
+Yours faithfully,
+
+J. BURDON-SANDERSON.
+
+]
+
+[Footnote 225: H. P. Liddon, _The Recovery of S. Thomas_; a sermon
+preached in St. Paul's, London, on April 23rd, 1882 (the Sunday after
+Darwin's death).]
+
+[Footnote 226: Dr. Pusey (_Unscience not Science adverse to Faith_,
+1878) writes: "The questions as to 'species,' of what variations the
+animal world is capable, whether the species be more or fewer, whether
+accidental variations may become hereditary ... and the like,
+naturally fall under the province of science. In all these questions
+Mr. Darwin's careful observations gained for him a deserved
+approbation and confidence."]
+
+[Footnote 227: Aristotle, in Bacon, quoted by Newman in his _Idea of a
+University_, p. 78. London, 1873.]
+
+[Footnote 228: _Life and Letters_ and _More Letters of Charles
+Darwin._]
+
+[Footnote 229: _Life and Letters_, London, 1896. _Thoughts on
+Religion_, London, 1895. _Candid Examination of Theism_, London,
+1878.]
+
+[Footnote 230: "Never in the history of man has so terrific a calamity
+befallen the race as that which all who look may now (viz. in
+consequence of the scientific victory of Darwin) behold advancing as a
+deluge black with destruction, resistless in might, uprooting our most
+cherished hopes, engulphing our most precious creed, and burying our
+highest life in mindless destruction."--_A Candid Examination of
+Theism_, p. 51.]
+
+[Footnote 231: _Science and Christian Tradition._ London, 1904.]
+
+[Footnote 232: "No productiveness of the highest kind ... is in the
+power of anyone."--_Conversations of Goethe with Eckermann and Soret_.
+London, 1850.]
+
+[Footnote 233: Berthelot, _Evolutionisme et Platonisme_, Paris, 1908,
+p. 45.]
+
+[Footnote 234: _Times_, 1892, _passim._]
+
+[Footnote 235: See Von Hartmann's _Wahrheit und Irrthum in
+Darwinismus_. Berlin, 1875.]
+
+[Footnote 236: Hymn of the Church--
+
+    Rerum Deus tenax vigor,
+    Immotus in te permanens.
+
+]
+
+[Footnote 237: _Life and Letters_, Vol. III. p. 359.]
+
+
+
+
+IX
+
+DARWINISM AND HISTORY
+
+BY J. B. BURY, LITT.D., LL.D.
+
+_Regius Professor of Modern History in the University of Cambridge_
+
+
+1. Evolution, and the principles associated with the Darwinian theory,
+could not fail to exert a considerable influence on the studies
+connected with the history of civilised man. The speculations which
+are known as "philosophy of history," as well as the sciences of
+anthropology, ethnography, and sociology (sciences which though they
+stand on their own feet are for the historian auxiliary), have been
+deeply affected by these principles. Historiographers, indeed, have
+with few exceptions made little attempt to apply them; but the growth
+of historical study in the nineteenth century has been determined and
+characterised by the same general principle which has underlain the
+simultaneous developments of the study of nature, namely the _genetic
+idea_. The "historical" conception of nature, which has produced the
+history of the solar system, the story of the earth, the genealogies
+of telluric organisms, and has revolutionised natural science, belongs
+to the same order of thought as the conception of human history as a
+continuous, genetic, causal process--a conception which has
+revolutionised historical research and made it scientific. Before
+proceeding to consider the application of evolutional principles, it
+will be pertinent to notice the rise of this new view.
+
+2. With the Greeks and Romans history had been either a descriptive
+record or had been written in practical interests. The most eminent
+of the ancient historians were pragmatical; that is, they regarded
+history as an instructress in statesmanship, or in the art of war, or
+in morals. Their records reached back such a short way, their
+experience was so brief, that they never attained to the conception of
+continuous process, or realised the significance of time; and they
+never viewed the history of human societies as a phenomenon to be
+investigated for its own sake. In the middle ages there was still less
+chance of the emergence of the ideas of progress and development. Such
+notions were excluded by the fundamental doctrines of the dominant
+religion which bounded and bound men's minds. As the course of history
+was held to be determined from hour to hour by the arbitrary will of
+an extra cosmic person, there could be no self-contained causal
+development, only a dispensation imposed from without. And as it was
+believed that the world was within no great distance from the end of
+this dispensation, there was no motive to take much interest in
+understanding the temporal, which was to be only temporary.
+
+The intellectual movements of the fifteenth and sixteenth centuries
+prepared the way for a new conception, but it did not emerge
+immediately. The historians of the Renaissance period simply reverted
+to the ancient pragmatical view. For Machiavelli, exactly as for
+Thucydides and Polybius, the use of studying history was instruction
+in the art of politics. The Renaissance itself was the appearance of a
+new culture, different from anything that had gone before; but at the
+time men were not conscious of this; they saw clearly that the
+traditions of classical antiquity had been lost for a long period, and
+they were seeking to revive them, but otherwise they did not perceive
+that the world had moved, and that their own spirit, culture, and
+conditions were entirely unlike those of the thirteenth century. It
+was hardly till the seventeenth century that the presence of a new
+age, as different from the middle ages as from the ages of Greece and
+Rome, was fully realised. It was then that the triple division of
+ancient, medieval, and modern was first applied to the history of
+western civilisation. Whatever objections may be urged against this
+division, which has now become almost a category of thought, it marks
+a most significant advance in man's view of his own past. He has
+become conscious of the immense changes in civilisation which have
+come about slowly in the course of time, and history confronts him
+with a new aspect. He has to explain how those changes have been
+produced, how the transformations were effected. The appearance of
+this problem was almost simultaneous with the rise of rationalism, and
+the great historians and thinkers of the eighteenth century, such as
+Montesquieu, Voltaire, Gibbon, attempted to explain the movement of
+civilisation by purely natural causes. These brilliant writers
+prepared the way for the genetic history of the following century. But
+in the spirit of the _Aufklaerung_, that eighteenth-century
+Enlightenment to which they belonged, they were concerned to judge all
+phenomena before the tribunal of reason; and the apotheosis of
+"reason" tended to foster a certain superior _a priori_ attitude,
+which was not favourable to objective treatment and was incompatible
+with a "historical sense." Moreover the traditions of pragmatical
+historiography had by no means disappeared.
+
+3. In the first quarter of the nineteenth century the meaning of
+genetic history was fully realised. "Genetic" perhaps is as good a
+word as can be found for the conception which in this century was
+applied to so many branches of knowledge in the spheres both of nature
+and of mind. It does not commit us to the doctrine proper of
+evolution, nor yet to any teleological hypothesis such as is implied
+in "progress." For history it meant that the present condition of the
+human race is simply and strictly the result of a causal series (or
+set of causal series)--a continuous succession of changes, where each
+state arises causally out of the preceding; and that the business of
+historians is to trace this genetic process, to explain each change,
+and ultimately to grasp' the complete development of the life of
+humanity. Three influential writers, who appeared at this stage and
+helped to initiate a new period of research, may specially be
+mentioned. Ranke in 1824 definitely repudiated the pragmatical view
+which ascribes to history the duties of an instructress, and with no
+less decision renounced the function, assumed by the historians of the
+_Aufklaerung_, to judge the past; it was his business, he said, merely
+to show how things really happened. Niebuhr was already working in the
+same spirit and did more than any other writer to establish the
+principle that historical transactions must be related to the ideas
+and conditions of their age. Savigny about the same time founded the
+"historical school" of law. He sought to show that law was not the
+creation of an enlightened will, but grew out of custom and was
+developed by a series of adaptations and rejections, thus applying the
+conception of evolution. He helped to diffuse the notion that all the
+institutions of a society or a nation are as closely interconnected as
+the parts of a living organism.
+
+4. The conception of the history of man as a causal development meant
+the elevation of historical inquiry to the dignity of a science. Just
+as the study of bees cannot become scientific so long as the student's
+interest in them is only to procure honey or to derive moral lessons
+from the labours of "the little busy bee," so the history of human
+societies cannot become the object of pure scientific investigation so
+long as man estimates its value in pragmatical scales. Nor can it
+become a science until it is conceived as lying entirely within a
+sphere in which the law of cause and effect has unreserved and
+unrestricted dominion. On the other hand, once history is envisaged as
+a causal process, which contains within itself the explanation of the
+development of man from his primitive state to the point which he has
+reached, such a process necessarily becomes the object of scientific
+investigation and the interest in it is scientific curiosity.
+
+At the same time, the instruments were sharpened and refined. Here
+Wolf, a philologist with historical instinct, was a pioneer. His
+_Prolegomena_ to Homer (1795) announced new modes of attack.
+Historical investigation was soon transformed by the elaboration of
+new methods.
+
+5. "Progress" involves a judgment of value, which is not involved in
+the conception of history as a genetic process. It is also an idea
+distinct from that of evolution. Nevertheless it is closely related to
+the ideas which revolutionised history at the beginning of the last
+century; it swam into men's ken simultaneously; and it helped
+effectively to establish the notion of history as a continuous process
+and to emphasise the significance of time. Passing over earlier
+anticipations, I may point to a _Discours_ of Turgot (1750), where
+history is presented as a process in which "the total mass of the
+human race" "marches continually though sometimes slowly to an ever
+increasing perfection." That is a clear statement of the conception
+which Turgot's friend Condorcet elaborated in the famous work,
+published in 1795, _Esquisse d'un tableau historique des progres de
+l'esprit humain_. This work first treated with explicit fulness the
+idea to which a leading role was to fall in the ideology of the
+nineteenth century. Condorcet's book reflects the triumphs of the
+_Tiers etat_, whose growing importance had also inspired Turgot; it
+was the political changes in the eighteenth century which led to the
+doctrine, emphatically formulated by Condorcet, that the masses are
+the most important element in the historical process. I dwell on this
+because, though Condorcet had no idea of evolution, the predominant
+importance of the masses was the assumption which made it possible to
+apply evolutional principles to history. And it enabled Condorcet
+himself to maintain that the history of civilisation, a progress still
+far from being complete, was a development conditioned by general
+laws.
+
+6. The assimilation of society to an organism, which was a governing
+notion in the school of Savigny, and the conception of progress,
+combined to produce the idea of an organic development, in which the
+historian has to determine the central principle or leading character.
+This is illustrated by the apotheosis of democracy in Tocqueville's
+_Democratie en Amerique_, where the theory is maintained that "the
+gradual and progressive development of equality is at once the past
+and the future of the history of men." The same two principles are
+combined in the doctrine of Spencer (who held that society is an
+organism, though he also contemplated its being what he calls a
+"super-organic aggregate"),[238] that social evolution is a
+progressive change from militarism to industrialism.
+
+7. The idea of development assumed another form in the speculations of
+German idealism. Hegel conceived the successive periods of history as
+corresponding to the ascending phases or ideas in the self-evolution
+of his Absolute Being. His _Lectures on the Philosophy of History_
+were published in 1837 after his death. His philosophy had a
+considerable effect, direct and indirect, on the treatment of history
+by historians, and although he was superficial and unscientific
+himself in dealing with historical phenomena, he contributed much
+towards making the idea of historical development familiar. Ranke was
+influenced, if not by Hegel himself, at least by the Idealistic
+philosophies of which Hegel's was the greatest. He was inclined to
+conceive the stages in the process of history as marked by
+incarnations, as it were, of ideas, and sometimes speaks as if the
+ideas were independent forces, with hands and feet. But while Hegel
+determined his ideas by _a priori_ logic, Ranke obtained his by
+induction--by a strict investigation of the phenomena; so that he was
+scientific in his method and work, and was influenced by Hegelian
+prepossessions only in the kind of significance which he was disposed
+to ascribe to his results. It is to be noted that the theory of Hegel
+implied a judgment of value; the movement was a progress towards
+perfection.
+
+8. In France, Comte approached the subject from a different side, and
+exercised, outside Germany, a far wider influence than Hegel. The 4th
+volume of his _Cours de philosophie positive_, which appeared in 1839,
+created sociology and treated history as a part of this new science,
+namely as "social dynamics." Comte sought the key for unfolding
+historical development, in what he called the social-psychological
+point of view, and he worked out the two ideas which had been
+enunciated by Condorcet: that the historian's attention should be
+directed not, as hitherto, principally to eminent individuals, but to
+the collective behaviour of the masses, as being the most important
+element in the process; and that, as in nature, so in history, there
+are general laws, necessary and constant, which condition the
+development. The two points are intimately connected, for it is only
+when the masses are moved into the foreground that regularity,
+uniformity, and law can be conceived as applicable. To determine the
+social-psychological laws which have controlled the development is,
+according to Comte, the task of sociologists and historians.
+
+9. The hypothesis of general laws operative in history was carried
+further in a book which appeared in England twenty years later and
+exercised an influence in Europe far beyond its intrinsic merit,
+Buckle's _History of Civilisation in England_ (1857-61). Buckle owed
+much to Comte, and followed him, or rather outdid him, in regarding
+intellect as the most important factor conditioning the upward
+development of man, so that progress, according to him, consisted in
+the victory of the intellectual over the moral laws.
+
+10. The tendency of Comte and Buckle to assimilate history to the
+sciences of nature by reducing it to general "laws," derived stimulus
+and plausibility from the vista offered by the study of statistics,
+in which the Belgian Quetelet, whose book _Sur l'homme_ appeared in
+1835, discerned endless possibilities. The astonishing uniformities
+which statistical inquiry disclosed led to the belief that it was only
+a question of collecting a sufficient amount of statistical material,
+to enable us to predict how a given social group will act in a
+particular case. Bourdeau, a disciple of this school, looks forward to
+the time when historical science will become entirely quantitative.
+The actions of prominent individuals, which are generally considered
+to have altered or determined the course of things, are obviously not
+amenable to statistical computation or explicable by general laws.
+Thinkers like Buckle sought to minimise their importance or explain
+them away.
+
+11. These indications may suffice to show that the new efforts to
+interpret history which marked the first half of the nineteenth
+century were governed by conceptions closely related to those which
+were current in the field of natural science and which resulted in the
+doctrine of evolution. The genetic principle, progressive development,
+general laws, the significance of time, the conception of society as
+an organic aggregate, the metaphysical theory of history as the
+self-evolution of spirit,--all these ideas show that historical
+inquiry had been advancing independently on somewhat parallel lines to
+the sciences of nature. It was necessary to bring this out in order to
+appreciate the influence of Darwinism.
+
+12. In the course of the dozen years which elapsed between the
+appearances of _The Origin of Species_ (observe that the first volume
+of Buckle's work was published just two years before) and of _The
+Descent of Man_ (1871), the hypothesis of Lamarck that man is the
+co-descendant with other species of some lower extinct form was
+admitted to have been raised to the rank of an established fact by
+most thinkers whose brains were not working under the constraint of
+theological authority.
+
+One important effect of the discovery of this fact (I am not speaking
+now of the Darwinian explanation) was to assign to history a definite
+place in the coordinated whole of knowledge, and relate it more
+closely to other sciences. It had indeed a defined logical place in
+systems such as Hegel's and Comte's; but Darwinism certified its
+standing convincingly and without more ado. The prevailing doctrine
+that man was created _ex abrupto_ had placed history in an isolated
+position, disconnected with the sciences of nature. Anthropology,
+which deals with the animal _anthropos_, now comes into line with
+zoology, and brings it into relation with history.[239] Man's
+condition at the present day is the result of a series of
+transformations, going back to the most primitive phase of society,
+which is the ideal (unattainable) beginning of history. But that
+beginning had emerged without any breach of continuity from a
+development which carries us back to a quadrimane ancestor, still
+further back (according to Darwin's conjecture) to a marine animal of
+the ascidian type, and then through remoter periods to the lowest form
+of organism. It is essential in this theory that though links have
+been lost there was no break in the gradual development; and this
+conception of a continuous progress in the evolution of life,
+resulting in the appearance of uncivilised Anthropos, helped to
+reinforce, and increase a belief in, the conception of the history of
+civilised Anthropos as itself also a continuous progressive
+development.
+
+13. Thus the diffusion of the Darwinian theory of the origin of man,
+by emphasising the idea of continuity and breaking down the barriers
+between the human and animal kingdoms, has had an important effect in
+establishing the position of history among the sciences which deal
+with telluric development. The perspective of history is merged in a
+larger perspective of development. As one of the objects of biology is
+to find the exact steps in the genealogy of man from the lowest
+organic form, so the scope of history is to determine the stages in
+the unique causal series from the most rudimentary to the present
+state of human civilisation.
+
+It is to be observed that the interest in historical research implied
+by this conception need not be that of Comte. In the Positive
+Philosophy history is part of sociology; the interest in it is to
+discover the sociological laws. In the view of which I have just
+spoken, history is permitted to be an end in itself; the
+reconstruction of the genetic process is an independent interest. For
+the purpose of the reconstruction, sociology, as well as physical
+geography, biology, psychology, is necessary; the sociologist and the
+historian play into each other's hands; but the object of the former
+is to establish generalisations; the aim of the latter is to trace in
+detail a singular causal sequence.
+
+14. The success of the evolutional theory helped to discredit the
+assumption or at least the invocation of transcendent causes.
+Philosophically of course it is compatible with theism, but historians
+have for the most part desisted from invoking the naive conception of
+a "god in history" to explain historical movements. A historian may be
+a theist; but, so far as his work is concerned, this particular belief
+is otiose. Otherwise indeed (as was remarked above) history could not
+be a science; for with a _deus ex machina_ who can be brought on the
+stage to solve difficulties scientific treatment is a farce. The
+transcendent element had appeared in a more subtle form through the
+influence of German philosophy. I noticed how Ranke is prone to refer
+to ideas as if they were transcendent existences manifesting
+themselves in the successive movements of history. It is intelligible
+to speak of certain ideas as controlling, in a given period,--for
+instance, the idea of nationality; but from the scientific point of
+view, such ideas have no existence outside the minds of individuals
+and are purely psychical forces; and a historical "idea," if it does
+not exist in this form, is merely a way of expressing a synthesis of
+the historian himself.
+
+15. From the more general influence of Darwinism on the place of
+history in the system of human knowledge, we may turn to the influence
+of the principles and methods by which Darwin explained development.
+It had been recognised even by ancient writers (such as Aristotle and
+Polybius) that physical circumstances (geography, climate) were
+factors conditioning the character and history of a race or society.
+In the sixteenth century Bodin emphasised these factors, and many
+subsequent writers took them into account. The investigations of
+Darwin, which brought them into the foreground, naturally promoted
+attempts to discover in them the chief key to the growth of
+civilisation. Comte had expressly denounced the notion that the
+biological methods of Lamarck could be applied to social man. Buckle
+had taken account of natural influences, but had relegated them to a
+secondary plane, compared with psychological factors. But the
+Darwinian theory made it tempting to explain the development of
+civilisation in terms of "adaptation to environment," "struggle for
+existence," "natural selection," "survival of the fittest," etc.[240]
+
+The operation of these principles cannot be denied. Man is still an
+animal, subject to zoological as well as mechanical laws. The dark
+influence of heredity continues to be effective; and psychical
+development had begun in lower organic forms,--perhaps with life
+itself. The organic and the social struggles for existence are
+manifestations of the same principle. Environment and climatic
+influence must be called in to explain not only the differentiation of
+the great racial sections of humanity, but also the varieties within
+these sub-species and, it may be, the assimilation of distinct
+varieties. Ritter's _Anthropogeography_ has opened a useful line of
+research. But on the other hand, it is urged that, in explaining the
+course of history, these principles do not take us very far, and that
+it is chiefly for the primitive ultra-prehistoric period that they can
+account for human development. It may be said that, so far as concerns
+the actions and movements of men which are the subject of recorded
+history, physical environment has ceased to act mechanically, and in
+order to affect their actions must affect their wills first; and that
+this psychical character of the causal relations substantially alters
+the problem. The development of human societies, it may be argued,
+derives a completely new character from the dominance of the conscious
+psychical element, creating as it does new conditions (inventions,
+social institutions, etc.) which limit and counteract the operation of
+natural selection, and control and modify the influence of physical
+environment. Most thinkers agree now that the chief clews to the
+growth of civilisation must be sought in the psychological sphere.
+Imitation, for instance, is a principle which is probably more
+significant for the explanation of human development than natural
+selection. Darwin himself was conscious that his principles had only a
+very restricted application in this sphere, as is evident from his
+cautious and tentative remarks in the 5th chapter of his _Descent of
+Man_. He applied natural selection to the growth of the intellectual
+faculties and of the fundamental social instincts, and also to the
+differentiation of the great races or "sub-species" (Caucasian,
+African, etc.) which differ in anthropological character.[241]
+
+16. But if it is admitted that the governing factors which concern the
+student of social development are of the psychical order, the
+preliminary success of natural science in explaining organic evolution
+by general principles encouraged sociologists to hope that social
+evolution could be explained on general principles also. The idea of
+Condorcet, Buckle, and others, that history could be assimilated to
+the natural sciences was powerfully reinforced, and the notion that
+the actual historical process, and every social movement involved in
+it, can be accounted for by sociological generalisations, so-called
+"laws," is still entertained by many, in one form or another.
+Dissentients from this view do not deny that the generalisations at
+which the sociologist arrives by the comparative method, by the
+analysis of social factors, and by psychological deduction may be an
+aid to the historian; but they deny that such uniformities are laws or
+contain an explanation of the phenomena. They can point to the element
+of chance coincidence. This element must have played a part in the
+events of organic evolution, but it has probably in a larger measure
+helped to determine events in social evolution. The collision of two
+unconnected sequences may be fraught with great results. The sudden
+death of a leader or a marriage without issue, to take simple cases,
+has again and again led to permanent political consequences. More
+emphasis is laid on the decisive actions of individuals, which cannot
+be reduced under generalisations and which deflect the course of
+events. If the significance of the individual will had been
+exaggerated to the neglect of the collective activity of the social
+aggregate before Condorcet, his doctrine tended to eliminate as
+unimportant the roles of prominent men, and by means of this
+elimination it was possible to found sociology. But it may be urged
+that it is patent on the face of history that its course has
+constantly been shaped and modified by the wills of individuals,[242]
+which are by no means always the expression of the collective will;
+and that the appearance of such personalities at the given moments is
+not a necessary outcome of the conditions and cannot be deduced. Nor
+is there any proof that, if such and such an individual had not been
+born, some one else would have arisen to do what he did. In some cases
+there is no reason to think that what happened need ever have come to
+pass. In other cases, it seems evident that the actual change was
+inevitable, but in default of the man who initiated and guided it, it
+might have been postponed, and, postponed or not, might have borne a
+different cachet. I may illustrate by an instance which has just come
+under my notice. Modern painting was founded by Giotto, and the
+Italian expedition of Charles VIII, near the close of the sixteenth
+century, introduced into France the fashion of imitating Italian
+painters. But for Giotto and Charles VIII, French painting might have
+been very different. It may be said that "if Giotto had not appeared,
+some other great imitator would have played a role analogous to his,
+and that without Charles VIII there would have been the commerce with
+Italy, which in the long run would have sufficed to place France in
+relation with Italian artists. But the equivalent of Giotto might have
+been deferred for a century and probably would have been different;
+and commercial relations would have required ages to produce the
+_rayonnement imitatif_ of Italian art in France, which the expedition
+of the royal adventurer provoked in a few years."[243] Instances
+furnished by political history are simply endless. Can we conjecture
+how events would have moved if the son of Philip of Macedon had been
+an incompetent? The aggressive action of Prussia which astonished
+Europe in 1740 determined the subsequent history of Germany; but that
+action was anything but inevitable; it depended entirely on the
+personality of Frederick the Great.
+
+Hence it may be argued that the action of individual wills is a
+determining and disturbing factor, too significant and effective to
+allow history to be grasped by sociological formulae. The types and
+general forms of development which the sociologist attempts to
+disengage can only assist the historian in understanding the actual
+course of events. It is in the special domains of economic history and
+_Culturgeschichte_ which have come to the front in modern times that
+generalisation is most fruitful, but even in these it may be contended
+that it furnishes only partial explanations.
+
+17. The truth is that Darwinism itself offers the best illustration of
+the insufficiency of general laws to account for historical
+development. The part played by coincidence, and the part played by
+individuals--limited by, and related to, general social
+conditions--render it impossible to deduce the course of the past
+history of man or to predict the future. But it is just the same with
+organic development. Darwin (or any other zoologist) could not deduce
+the actual course of evolution from general principles. Given an
+organism and its environment, he could not show that it must evolve
+into a more complex organism of a definite predetermined type; knowing
+what it has evolved into, he could attempt to discover and assign the
+determining causes. General principles do not account for a particular
+sequence; they embody necessary conditions; but there is a chapter of
+accidents too. It is the same in the case of history.
+
+18. Among the evolutional attempts to subsume the course of history under
+general syntheses, perhaps the most important is that of Lamprecht, whose
+"kulturhistorische" attempt to discover and assign the determining causes.
+German history, exhibits the (indirect) influence of the Comtist school. It
+is based upon psychology, which, in his views, holds among the sciences of
+mind (_Geisteswissenschaften_) the same place (that of a
+_Grundwissenschaft_) which mechanics holds among the sciences of nature.
+History, by the same comparison, corresponds to biology, and, according to
+him, it can only become scientific if it is reduced to general concepts
+(_Begriffe_). Historical movements and events are of a psychical character,
+and Lamprecht conceives a given phase of civilisation as "a collective
+psychical condition (_seelischer Gesamtzustand_)" controlling the period,
+"a diapason which penetrates all psychical phenomena and thereby all
+historical events of the time."[244] He has worked out a series of such
+phases, "ages of changing psychical diapason," in his _Deutsche
+Geschichte_, with the aim of showing that all the feelings and actions of
+each age can be explained by the diapason; and has attempted to prove that
+these diapasons are exhibited in other social developments, and are
+consequently not singular but typical. He maintains further that these ages
+succeed each other in a definite order; the principle being that the
+collective psychical development begins with the homogeneity of all the
+individual members of a society and, through heightened psychical activity,
+advances in the form of a continually increasing differentiation of the
+individuals (this is akin to the Spencerian formula). This process,
+evolving psychical freedom from psychical constraint, exhibits a series of
+psychical phenomena which define successive periods of civilisation. The
+process depends on two simple principles, that no idea can disappear
+without leaving behind it an effect or influence, and that all psychical
+life, whether in a person or a society, means change, the acquisition of
+new mental contents. It follows that the new have to come to terms with the
+old, and this leads to a synthesis which determines the character of a new
+age. Hence the ages of civilisation are defined as the "highest concepts
+for subsuming without exception all psychical phenomena of the development
+of human societies, that is, of all historical events."[245] Lamprecht
+deduces the idea of a special historical science, which might be called
+"historical ethnology," dealing with the ages of civilisation, and bearing
+the same relation to (descriptive or narrative) history as ethnology to
+ethnography. Such a science obviously corresponds to Comte's social
+dynamics, and the comparative method, on which Comte laid so much emphasis,
+is the principal instrument of Lamprecht.
+
+19. I have dwelt on the fundamental ideas of Lamprecht, because they
+are not yet widely known in England, and because his system is the
+ablest product of the sociological school of historians. It carries
+the more weight as its author himself is a historical specialist, and
+his historical syntheses deserve the most careful consideration. But
+there is much in the process of development which on such assumptions
+is not explained, especially the initiative of individuals. Historical
+development does not proceed in a right line, without the choice of
+diverging. Again and again, several roads are open to it, of which it
+chooses one--why? On Lamprecht's method, we may be able to assign the
+conditions which limit the psychical activity of men at a particular
+stage of evolution, but within those limits the individual has so many
+options, such a wide room for moving, that the definition of those
+conditions, the "psychical diapasons," is only part of the explanation
+of the particular development. The heel of Achilles in all historical
+speculations of this class has been the role of the individual.
+
+The increasing prominence of economic history has tended to encourage
+the view that history can be explained in terms of general concepts or
+types. Marx and his school based their theory of human development on
+the conditions of production, by which, according to them, all social
+movements and historical changes are entirely controlled. The leading
+part which economic factors play in Lamprecht's system is significant,
+illustrating the fact that economic changes admit most readily this
+kind of treatment, because they have been less subject to direction or
+interference by individual pioneers.
+
+Perhaps it may be thought that the conception of _social environment_
+(essentially psychical), on which Lamprecht's "psychical diapasons"
+depend, is the most valuable and fertile conception that the historian
+owes to the suggestion of the science of biology--the conception of
+all particular historical actions and movements as (1) related to and
+conditioned by the social environment, and (2) gradually bringing
+about a transformation of that environment. But no given
+transformation can be proved to be necessary (predetermined). And
+types of development do not represent laws; their meaning and value
+lie in the help they may give to the historian, in investigating a
+certain period of civilisation, to enable him to discover the
+inter-relations among the diverse features which it presents. They
+are, as some one has said, an instrument of heuretic method.
+
+20. The man engaged in special historical researches--which have been
+pursued unremittingly for a century past, according to scientific
+methods of investigating evidence (initiated by Wolf, Niebuhr,
+Ranke)--have for the most part worked on the assumptions of genetic
+history or at least followed in the footsteps of those who fully
+grasped the genetic point of view. But their aim has been to collect
+and sift evidence, and determine particular facts; comparatively few
+have given serious thought to the lines of research and the
+speculations which have been considered in this paper. They have been
+reasonably shy of compromising their work by applying theories which
+are still much debated and immature. But historiography cannot
+permanently evade the questions raised by these theories. One may
+venture to say that no historical change or transformation will be
+fully understood until it is explained how social environment acted on
+the individual components of the society (both immediately and by
+heredity), and how the individuals reacted upon their environment. The
+problem is psychical, but it is analogous to the main problem of the
+biologist.
+
+FOOTNOTES:
+
+[Footnote 238: A society presents suggestive analogies with an
+organism, but it certainly is not an organism, and sociologists who
+draw inferences from the assumption of its organic nature must fall
+into error. A vital organism and a society are radically distinguished
+by the fact that the individual components of the former, namely the
+cells, are morphologically as well as functionally differentiated,
+whereas the individuals which compose a society are morphologically
+homogeneous and only functionally differentiated. The resemblances and
+the differences are worked out in E. de Majewski's striking book, _La
+Science de la Civilisation_. Paris. 1908.]
+
+[Footnote 239: It is to be observed that history is (not only
+different in scope but) not co-extensive with anthropology _in time_.
+For it deals only with the development of man in societies, whereas
+anthropology includes in its definition the proto-anthropic period
+when _anthropos_ was still non-social, whether he lived in herds like
+the chimpanzee, or alone like the male ourang-outang. (It has been
+well shown by Majewski that congregations--herds, flocks, packs,
+&c.--of animals are not _societies_; the characteristic of a society
+is differentiation of function. Bee hives, ant hills, may be called
+quasi-societies; but in their case the classes which perform distinct
+functions are morphologically different.)]
+
+[Footnote 240: Recently O. Seeck has applied these principles to the
+decline of Graeco-Roman civilisation in his _Untergang der antiken
+Welt_, 2 vols., Berlin, 1895, 1901.]
+
+[Footnote 241: Darwinian formulae may be suggestive by way of analogy.
+For instance, it is characteristic of social advance that a multitude
+of inventions, schemes and plans are framed which are never carried
+out, similar to, or designed for the same end as, an invention or plan
+which is actually adopted because it has chanced to suit better the
+particular conditions of the hour (just as the works accomplished by
+an individual statesman, artist or savant are usually only a residue
+of the numerous projects conceived by his brain). This process in
+which so much abortive production occurs is analogous to elimination
+by natural selection.]
+
+[Footnote 242: We can ignore here the metaphysical question of
+freewill and determinism. For the character of the individual's brain
+depends in any case on ante-natal accidents and coincidences, and so
+it may be said that the role of individuals ultimately depends on
+chance,--the accidental coincidence of independent sequences.]
+
+[Footnote 243: I have taken this example from G. Tarde's _La logique
+sociale_ (p. 403), Paris, 1904, where it is used for quite a different
+purpose.]
+
+[Footnote 244: _Die kulturhistorische Methode_, Berlin, 1900, p. 26.]
+
+[Footnote 245: _Ibid._ pp. 28, 29.]
+
+
+
+
+X
+
+DARWINISM AND SOCIOLOGY
+
+BY C. BOUGLE
+
+_Professor of Social Philosophy in the University of Toulouse and
+Deputy-Professor at the Sorbonne, Paris_
+
+
+How has our conception of social phenomena, and of their history, been
+affected by Darwin's conception of Nature and the laws of its
+transformation? To what extent and in what particular respects have
+the discoveries and hypotheses of the author of _The Origin of
+Species_ aided the efforts of those who have sought to construct a
+science of society?
+
+To such a question it is certainly not easy to give any brief or
+precise answer. We find traces of Darwinism almost everywhere.
+Sociological systems differing widely from each other have laid claim
+to its authority; while, on the other hand, its influence has often
+made itself felt only in combination with other influences. The
+Darwinian thread is worked into a hundred patterns along with other
+threads.
+
+To deal with the problem, we must, it seems, first of all distinguish
+the more general conclusions in regard to the evolution of living
+beings, which are the outcome of Darwinism, from the particular
+explanations it offers of the ways and means by which that evolution
+is effected. That is to say, we must, as far as possible, estimate
+separately the influence of Darwin as an evolutionist and Darwin as a
+selectionist.
+
+The nineteenth century, said Cournot, has witnessed a mighty effort to
+"reintegrer l'homme dans la nature." From divers quarters there has
+been a methodical reaction against the persistent dualism of the
+Cartesian tradition, which was itself the unconscious heir of the
+Christian tradition. Even the philosophy of the eighteenth century,
+materialistic as were for the most part the tendencies of its leaders,
+seemed to revere man as a being apart, concerning whom laws might be
+formulated _a priori_. To bring him down from his pedestal there was
+needed the marked predominance of positive researches wherein no
+account was taken of the "pride of man." There can be no doubt that
+Darwin has done much to familiarise us with this attitude. Take for
+instance the first part of _The Descent of Man_: it is an accumulation
+of typical facts, all tending to diminish the distance between us and
+our brothers, the lower animals. One might say that the naturalist had
+here taken as his motto, "Whosoever shall exalt himself shall be
+abased; and he that shall humble himself shall be exalted." Homologous
+structures, the survival in man of certain organs of animals, the
+rudiments in the animal of certain human faculties, a multitude of
+facts of this sort, led Darwin to the conclusion that there is no
+ground for supposing that the "king of the universe" is exempt from
+universal laws. Thus belief in the _imperium in imperio_ has been, as
+it were, whittled away by the progress of the naturalistic spirit,
+itself continually strengthened by the conquests of the natural
+sciences. The tendency may, indeed, drag the social sciences into
+overstrained analogies, such, for instance, as the assimilation of
+societies to organisms. But it will, at least, have had the merit of
+helping sociology to shake off the pre-conception that the groups
+formed by men are artificial, and that history is completely at the
+mercy of chance. Some years before the appearance of _The Origin of
+Species_, August Comte had pointed out the importance, as regards the
+unification of positive knowledge, of the conviction that the social
+world, the last refuge of spiritualism, is itself subject to
+determinism. It cannot be doubted that the movement of thought which
+Darwin's discoveries promoted contributed to the spread of this
+conviction, by breaking down the traditional barrier which cut man off
+from Nature.
+
+But Nature, according to modern naturalists, is no immutable thing: it
+is rather perpetual movement, continual progression. Their discoveries
+batter a breach directly into the Aristotelian notion of species; they
+refuse to see in the animal world a collection of immutable types,
+distinct from all eternity, and corresponding, as Cuvier said, to so
+many particular thoughts of the Creator. Darwin especially
+congratulated himself upon having been able to deal this doctrine the
+_coup de grace_: immutability is, he says, his chief enemy; and he is
+concerned to show--therein following up Lyell's work--that everything
+in the organic world, as in the inorganic, is explained by insensible
+but incessant transformations. "Nature makes no leaps"--"Nature knows
+no gaps": these two _dicta_ form, as it were, the two landmarks
+between which Darwin's idea of transformation is worked out. That is
+to say, the development of Darwinism is calculated to further the
+application of the philosophy of Becoming to the study of human
+institutions.
+
+The progress of the natural sciences thus brings unexpected
+reinforcements to the revolution which the progress of historical
+discipline had begun. The first attempt to constitute an actual
+science of social phenomena--that, namely, of the economists--had
+resulted in laws which were called natural, and which were believed to
+be eternal and universal, valid for all times and all places. But this
+perpetuality, brother, as Knies said, of the immutability of the old
+zoology, did not long hold out against the ever-swelling tide of the
+historical movement. Knowledge of the transformations that had taken
+place in language, of the early phases of the family, of religion, of
+property, had all favoured the revival of the Heraclitean view:
+[Greek: panta rei]. As to the categories of political economy, it was
+soon to be recognised, as by Lasalle, that they too are only
+historical. The philosophy of history, moreover, gave expression
+under various forms to the same tendency. Hegel declares that "all
+that is real is rational," but at the same time he shows that all that
+is real is ephemeral, and that for history there is nothing fixed
+beneath the sun. It is this sense of universal evolution that Darwin
+came with fresh authority to enlarge. It was in the name of biological
+facts themselves that he taught us to see only slow metamorphoses in
+the history of institutions, and to be always on the outlook for
+survivals side by side with rudimentary forms. Anyone who reads
+_Primitive Culture_, by Tylor,--a writer closely connected with
+Darwin--will be able to estimate the services which these cardinal
+ideas were to render to the social sciences when the age of
+comparative research had succeeded to that of _a priori_ construction.
+
+Let us note, moreover, that the philosophy of Becoming in passing through
+the Darwinian biology became, as it were, filtered; it got rid of those
+traces of finalism, which, under different forms, it had preserved through
+all the systems of German Romanticism. Even in Herbert Spencer, it has been
+plausibly argued, one can detect something of that sort of mystic
+confidence in forces spontaneously directing life, which forms the very
+essence of those systems. But Darwin's observations were precisely
+calculated to render such an hypothesis futile. At first people may have
+failed to see this; and we call to mind the ponderous sarcasms of Flourens
+when he objected to the theory of Natural Selection that it attributed to
+nature a power of free choice. "Nature endowed with will! That was the
+final error of last century; but the nineteenth no longer deals in
+personifications."[246] In fact Darwin himself put his readers on their
+guard against the metaphors he was obliged to use. The processes by which
+he explains the survival of the fittest are far from affording any
+indication of the design of some transcendent breeder. Nor, if we look
+closely, do they even imply immanent effort in the animal; the sorting out
+can be brought about mechanically, simply by the action of the environment.
+In this connection Huxley could with good reason maintain that Darwin's
+originality consisted in showing how harmonies which hitherto had been
+taken to imply the agency of intelligence and will could be explained
+without any such intervention. So, when later on, objective sociology
+declares that, even when social phenomena are in question, all finalist
+preconceptions must be distrusted if a science is to be constituted, it is
+to Darwin that its thanks are due; he had long been clearing paths for it
+which lay well away from the old familiar road trodden by so many theories
+of evolution.
+
+This anti-finalist doctrine, when fully worked out, was, moreover,
+calculated to aid in the needful dissociation of two notions: that of
+evolution and that of progress. In application to society these had
+long been confounded; and, as a consequence, the general idea seemed
+to be that only one type of evolution was here possible. Do we not
+detect such a view in Comte's sociology, and perhaps even in Herbert
+Spencer's? Whoever, indeed, assumes an end for evolution is naturally
+inclined to think that only one road leads to that end. But those
+whose minds the Darwinian theory has enlightened are aware that the
+transformations of living beings depend primarily upon their
+conditions, and that it is these conditions which are the agents of
+selection from among individual variations. Hence, it immediately
+follows that transformations are not necessarily improvements. Here,
+Darwin's thought hesitated. Logically his theory proves, as Ray
+Lankester pointed out, that the struggle for existence may have as its
+outcome degeneration as well as amelioration: evolution may be
+regressive as well as progressive. Then, too--and this is especially
+to be borne in mind--each species takes its good where it finds it,
+seeks its own path and survives as best it can. Apply this notion to
+society and you arrive at the theory of multilinear evolution.
+Divergencies will no longer surprise you. You will be forewarned not
+to apply to all civilisations the same measure of progress, and you
+will recognise that types of evolution may differ just as social
+species themselves differ. Have we not here one of the conceptions
+which mark off sociology proper from the old philosophy of history?
+
+       *       *       *       *       *
+
+But if we are to estimate the influence of Darwinism upon sociological
+conceptions, we must not dwell only upon the way in which Darwin
+impressed the general notion of evolution upon the minds of thinkers.
+We must go into details. We must consider the influence of the
+particular theories by which he explained the mechanism of this
+evolution. The name of the author of _The Origin of Species_ has been
+especially attached, as everyone knows, to the doctrines of "natural
+selection" and of "struggle for existence," completed by the notion of
+"individual variation." These doctrines were turned to account by very
+different schools of social philosophy. Pessimistic and optimistic,
+aristocratic and democratic, individualistic and socialistic systems
+were to war with each other for years by casting scraps of Darwinism
+at each other's heads.
+
+It was the spectacle of human contrivance that suggested to Darwin his
+conception of natural selection. It was in studying the methods of
+pigeon breeders that he divined the processes by which nature, in the
+absence of design, obtains analogous results in the differentiation of
+types. As soon as the importance of artificial selection in the
+transformation of species of animals was understood, reflection
+naturally turned to the human species, and the question arose, How far
+do men observe, in connection with themselves, those laws of which
+they make practical application in the case of animals? Here we come
+upon one of the ideas which guided the researches of Gallon, Darwin's
+cousin. The author of _Inquiries into Human Faculty and its
+Development_,[247] has often expressed his surprise that, considering
+all the precautions taken, for example, in the breeding of horses,
+none whatever are taken in the breeding of the human species. It seems
+to be forgotten that the species suffers when the "fittest" are not
+able to perpetuate their type. Ritchie, in his _Darwinism and
+Politics_[248] reminds us of Darwin's remark that the institution of
+the peerage might be defended on the ground that peers, owing to the
+prestige they enjoy, are enabled to select as wives "the most
+beautiful and charming women out of the lower ranks."[249] But, says
+Galton, it is as often as not "heiresses" that they pick out, and
+birth statistics seem to show that these are either less robust or
+less fecund than others. The truth is that considerations continue to
+preside over marriage which are entirely foreign to the improvement of
+type, much as this is a condition of general progress. Hence the
+importance of completing Odin's and De Candolle's statistics which are
+designed to show how characters are incorporated in organisms, how
+they are transmitted, how lost, and according to what law eugenic,
+elements depart from the mean or return to it.
+
+But thinkers do not always content themselves with undertaking merely
+the minute researches which the idea of Selection suggests. They are
+eager to defend this or that thesis. In the name of this idea certain
+social anthropologists have recast the conception of the process of
+civilisation, and have affirmed that Social Selection generally works
+against the trend of Natural Selection. Vacher de Lapouge--following
+up an observation by Broca on the point--enumerates the various
+institutions, or customs, such as the celibacy of priests and military
+conscription, which cause elimination or sterilisation of the bearers
+of certain superior qualities, intellectual or physical. In a more
+general way he attacks the democratic movement, a movement, as P.
+Bourget says, which is "anti-physical" and contrary to the natural
+laws of progress; though it has been inspired "by the dreams of that
+most visionary of all centuries, the eighteenth."[250] The "Equality"
+which levels down and mixes (justly condemned, he holds, by the Comte
+de Gobineau), prevents the aristocracy of the blond dolichocephales
+from holding the position and playing the part which, in the interests
+of all, should belong to them. Otto Ammon, in his _Natural Selection
+in Man_, and in _The Social Order and its Natural Bases_,[251]
+defended analogous doctrines in Germany; setting the curve
+representing frequency of talent over against that of income, he
+attempted to show that all democratic measures which aim at promoting
+the rise in the social scale of the talented are useless, if not
+dangerous; that they only increase the panmixia, to the great
+detriment of the species and of society.
+
+Among the aristocratic theories which Darwinism has thus inspired we
+must reckon that of Nietzsche. It is well known that in order to
+complete his philosophy he added biological studies to his
+philological; and more than once in his remarks upon the _Wille zur
+Macht_ he definitely alludes to Darwin; though it must be confessed
+that it is generally in order to proclaim the insufficiency of the
+processes by which Darwin seeks to explain the genesis of species.
+Nevertheless, Nietzsche's mind is completely possessed by an ideal of
+Selection. He, too, has a horror of panmixia. The naturalists'
+conception of "the fittest" is joined by him to that of the "hero" of
+romance to furnish a basis for his doctrine of the Superman. Let us
+hasten to add, moreover, that at the very moment when support was
+being sought in the theory of Selection for the various forms of the
+aristocratic doctrine, those same forms were being battered down on
+another side by means of that very theory. Attention was drawn to the
+fact that by virtue of the laws which Darwin himself had discovered
+isolation leads to etiolation. There is a risk that the privilege
+which withdraws the privileged elements of Society from competition
+will cause them to degenerate. In fact, Jacoby in his _Studies in
+Selection, in connexion with Heredity in Man_,[252] concludes that
+"sterility, mental debility, premature death and, finally, the
+extinction of the stock were not specially and exclusively the fate of
+sovereign dynasties; all privileged classes, all families in
+exclusively elevated positions share the fate of reigning families,
+although in a minor degree and in direct proportion to the loftiness
+of their social standing. From the mass of human beings spring
+individuals, families, races, which tend to raise themselves above the
+common level; painfully they climb the rugged heights, attain the
+summits of power, of wealth, of intelligence, of talent, and then, no
+sooner are they there than they topple down and disappear in gulfs of
+mental and physical degeneracy." The demographical researches of
+Hansen[253] (following up and completing Dumont's) tended, indeed, to
+show that urban as well as feudal aristocracies, burgher classes as
+well as noble castes, were liable to become effete. Hence it might
+well be concluded that the democratic movement, operating as it does
+to break down class barriers, was promoting instead of impeding human
+selection.
+
+       *       *       *       *       *
+
+So we see that, according to the point of view, very different
+conclusions have been drawn from the application of the Darwinian idea
+of Selection to human society. Darwin's other central idea, closely
+bound up with this, that, namely, of the "struggle for existence" also
+has been diversely utilised. But discussion has chiefly centered upon
+its signification. And while some endeavour to extend its application
+to everything, we find others trying to limit its range. The
+conception of a "struggle for existence" has in the present day been
+taken up into the social sciences from natural science, and adopted.
+But originally it descended from social science to natural. Darwin's
+law is, as he himself said, only Malthus' law generalised and extended
+to the animal world: a growing disproportion between the supply of
+food and the number of the living is the fatal order whence arises the
+necessity of universal struggle, a struggle which, to the great
+advantage of the species, allows only the best equipped individuals to
+survive. Nature is regarded by Huxley as an immense arena where all
+living beings are gladiators.[254]
+
+Such a generalisation was well adapted to feed the stream of
+pessimistic thought; and it furnished to the apologists of war, in
+particular, new arguments, weighted with all the authority which in
+these days attaches to scientific deliverances. If people no longer
+say, as Bonald did, and Moltke after him, that war is a providential
+fact, they yet lay stress on the point that it is a natural fact. To
+the peace party Dragomirov's objection is urged that its attempts are
+contrary to the fundamental laws of nature, and that no sea wall can
+hold against breakers that come with such gathered force.
+
+But in yet another quarter Darwinism was represented as opposed to
+philanthropic intervention. The defenders of the orthodox political
+economy found in it support for their tenets. Since in the organic
+world universal struggle is the condition of progress, it seemed
+obvious that free competition must be allowed to reign unchecked in
+the economic world. Attempts to curb it were in the highest degree
+imprudent. The spirit of Liberalism here seemed in conformity with the
+trend of nature: in this respect, at least, contemporary naturalism,
+offspring of the discoveries of the nineteenth century, brought
+reinforcements to the individualist doctrine, begotten of the
+speculations of the eighteenth: but only, it appeared, to turn mankind
+away for ever from humanitarian dreams. Would those whom such
+conclusions repelled be content to oppose to nature's imperatives
+only the protests of the heart? There were some who declared, like
+Brunetiere, that the laws in question, valid though they might be for
+the animal kingdom, were not applicable to the human. And so a return
+was made to the classic dualism. This indeed seems to be the line that
+Huxley took, when, for instance, he opposed to the cosmic process an
+ethical process which was its reverse.
+
+But the number of thinkers whom this antithesis does not satisfy grows
+daily. Although the pessimism which claims authorisation from Darwin's
+doctrines is repugnant to them, they still are unable to accept the
+dualism which leaves a gulf between man and nature. And their
+endeavour is to link the two by showing that while Darwin's laws
+obtain in both kingdoms, the conditions of their application are not
+the same: their forms, and, consequently, their results, vary with the
+varying mediums in which the struggle of living beings takes place,
+with the means these beings have at disposal, with the ends even which
+they propose to themselves.
+
+Here we have the explanation of the fact that among determined
+opponents of war partisans of the "struggle for existence" can be
+found: there are disciples of Darwin in the peace party. Novicow, for
+example, admits the "_combat universel_" of which Le Dantec[255]
+speaks; but he remarks that at different stages of evolution, at
+different stages of life the same weapons are not necessarily
+employed. Struggles of brute force, armed hand to hand conflicts, may
+have been a necessity in the early phases of human societies.
+Nowadays, although competition may remain inevitable and
+indispensable, it can assume milder forms. Economic rivalries,
+struggles between intellectual influences, suffice to stimulate
+progress: the processes which these admit are, in the actual state of
+civilisation, the only ones which attain their end without waste, the
+only ones logical. From one end to the other of the ladder of life,
+struggle is the order of the day; but more and more as the higher
+rungs are reached, it takes on characters which are proportionately
+more "humane."
+
+Reflections of this kind permit the introduction into the economic
+order of limitations to the doctrine of "laisser faire, laisser
+passer." This appeals, it is said, to the example of nature where
+creatures, left to themselves, struggle without truce and without
+mercy; but the fact is forgotten that upon industrial battlefields the
+conditions are different. The competitors here are not left simply to
+their natural energies: they are variously handicapped. A rich store
+of artificial resources exists in which some participate and others do
+not. The sides then are unequal; and as a consequence the result of
+the struggle is falsified. "In the animal world," said De
+Laveleye,[256] criticising Spencer, "the fate of each creature is
+determined by its individual qualities; whereas in civilised societies
+a man may obtain the highest position and the most beautiful wife
+because he is rich and well-born, although he may be ugly, idle or
+improvident; and then it is he who will perpetuate the species. The
+wealthy man, ill constituted, incapable, sickly, enjoys his riches and
+establishes his stock under the protection of the laws." Haycraft in
+England and Jentsch in Germany have strongly emphasised these
+"anomalies," which nevertheless are the rule. That is to say that even
+from a Darwinian point of view all social reforms can readily be
+justified which aim at diminishing, as Wallace said, inequalities at
+the start.
+
+But we can go further still. Whence comes the idea that all measures
+inspired by the sentiment of solidarity are contrary to Nature's
+trend? Observe her carefully, and she will not give lessons only in
+individualism. Side by side with the struggle for existence do we not
+find in operation what Lanessan calls "association for existence."
+Long ago, Espinas had drawn attention to "societies of animals,"
+temporary or permanent, and to the kind of morality that arose in
+them. Since then, naturalists have often insisted upon the importance
+of various forms of symbiosis. Kropotkin in _Mutual Aid_ has chosen
+to enumerate many examples of altruism furnished by animals to
+mankind. Geddes and Thomson went so far as to maintain that "Each of
+the greater steps of progress is in fact associated with an increased
+measure of subordination of individual competition to reproductive or
+social ends, and of interspecific competition to co-operative,
+association."[257] Experience shows, according to Geddes, that the
+types which are fittest to surmount great obstacles are not so much
+those who engage in the fiercest competitive struggle for existence,
+as those who contrive to temper it. From all these observations there
+resulted, along with a limitation of Darwinian pessimism, some
+encouragement for the aspirations of the collectivists.
+
+And Darwin himself would, doubtless, have subscribed to these
+rectifications. He never insisted, like his rival, Wallace, upon the
+necessity of the solitary struggle of creatures in a state of nature,
+each for himself and against all. On the contrary, in _The Descent of
+Man_, he pointed out the serviceableness of the social instincts, and
+corroborated Bagehot's statements when the latter, applying laws of
+physics to politics, showed the great advantage societies derived from
+intercourse and communion. Again, the theory of sexual evolution which
+makes the evolution of types depend increasingly upon preferences,
+judgments, mental factors, surely offers something to qualify what
+seems hard and brutal in the theory of natural selection.
+
+But, as often happens with disciples, the Darwinians had out-Darwined
+Darwin. The extravagances of social Darwinism provoked a useful
+reaction; and thus people were led to seek, even in the animal
+kingdom, for facts of solidarity which would serve to justify humane
+effort.
+
+       *       *       *       *       *
+
+On quite another line, however, an attempt has been made to connect
+socialist tendencies with Darwinian principles. Marx and Darwin have
+been confronted; and writers have undertaken to show that the work of
+the German philosopher fell readily into line with that of the English
+naturalist and was a development of it. Such has been the endeavour of
+Ferri in Italy and of Woltmann in Germany, not to mention others. The
+founders of "scientific socialism" had, moreover, themselves thought
+of this reconciliation. They make more than one allusion to Darwin in
+works which appeared after 1859. And sometimes they use his theory to
+define by contrast their own ideal. They remark that the capitalist
+system, by giving free course to individual competition, ends indeed
+in a _bellum omnium contra omnes_; and they make it clear that
+Darwinism, thus understood, is as repugnant to them as to Duehring.
+
+But it is at the scientific and not at the moral point of view that
+they place themselves when they connect their economic history with
+Darwin's work. Thanks to this unifying hypothesis, they claim to have
+constructed--as Marx does in his preface to _Das Kapital_--a veritable
+natural history of social evolution. Engels speaks in praise of his
+friend Marx as having discovered the true mainspring of history hidden
+under the veil of idealism and sentimentalism, and as having
+proclaimed in the _primum vivere_ the inevitableness of the struggle
+for existence. Marx himself, in _Das Kapital_, indicated another
+analogy when he dwelt upon the importance of a general technology for
+the explanation of this psychology:--a history of tools which would be
+to social organs what Darwinism is to the organs of animal species.
+And the very importance they attach to tools, to apparatus, to
+machines, abundantly proves that neither Marx nor Engels were likely
+to forget the special characters which mark off the human world from
+the animal. The former always remains to a great extent an artificial
+world. Inventions change the face of its institutions. New modes of
+production revolutionise not only modes of government, but modes even
+of collective thought. Therefore it is that the evolution of society
+is controlled by laws special to it, of which the spectacle of nature
+offers no suggestion.
+
+If, however, even in this special sphere, it can still be urged that
+the evolution of the material conditions of society is in accord with
+Darwin's theory, it is because the influence of the methods of
+production is itself to be explained by the incessant strife of the
+various classes with each other. So that in the end Marx, like Darwin,
+finds the source of all progress is in struggle. Both are grandsons of
+Heraclitus:--[Greek: polemos pater panton]. It sometimes happens, in
+these days, that the doctrine of revolutionary socialism is contrasted
+as rude and healthy with what may seem to be the enervating tendency
+of "solidarist" philanthropy: the apologists of the doctrine then
+pride themselves above all upon their faithfulness to Darwinian
+principles.
+
+       *       *       *       *       *
+
+So far we have been mainly concerned to show the use that social
+philosophies have made of the Darwinian laws for practical purposes:
+in order to orientate society towards their ideals each school tries
+to show that the authority of natural science is on its side. But even
+in the most objective of theories, those which systematically make
+abstraction of all political tendencies in order to study the social
+reality in itself, traces of Darwinism are readily to be found.
+
+Let us take for example Durkheim's theory of Division of Labour.[258]
+The conclusions he derives from it are that whenever professional
+specialisation causes multiplication of distinct branches of activity,
+we get organic solidarity--implying differences--substituted for
+mechanical solidarity, based upon likenesses. The umbilical cord, as
+Marx said, which connects the individual consciousness with the
+collective consciousness is cut. The personality becomes more and more
+emancipated. But on what does this phenomenon, so big with
+consequences, itself depend? The author goes to social morphology for
+the answer: it is, he says, the growing density of population which
+brings with it this increasing differentiation of activities. But,
+again, why? Because the greater density, in thrusting men up against
+each other, augments the intensity of their competition for the means
+of existence; and for the problems which society thus has to face
+differentiation of functions presents itself as the gentlest solution.
+
+Here one sees that the writer borrows directly from Darwin.
+Competition is at its maximum between similars, Darwin had declared;
+different species, not laying claim to the same food, could more
+easily coexist. Here lay the explanation of the fact that upon the
+same oak hundreds of different insects might be found. Other things
+being equal, the same applies to society. He who finds some unadopted
+specialty possesses a means of his own for getting a living. It is by
+this division of their manifold tasks that men contrive not to crush
+each other. Here we obviously have a Darwinian law serving as
+intermediary in the explanation of that progress of division of labour
+which itself explains so much in the social evolution.
+
+And we might take another example, at the other end of the series of
+sociological systems. G. Tarde is a sociologist with the most
+pronounced anti-naturalistic views. He has attempted to show that all
+application of the laws of natural science to society is misleading.
+In his _Opposition Universelle_ he has directly combatted all forms of
+sociological Darwinism. According to him the idea that the evolution
+of society can be traced on the same plan as the evolution of species
+is chimerical. Social evolution is at the mercy of all kinds of
+inventions, which by virtue of the laws of imitation modify, through
+individual to individual, through neighbourhood to neighbourhood, the
+general state of those beliefs and desires which are the only
+"quantities" whose variation matters to the sociologist. But, it may
+be rejoined, that however psychical the forces may be, they are none
+the less subject to Darwinian laws. They compete with each other; they
+struggle for the mastery of minds. Between types of ideas, as between
+organic forms, selection operates. And though it may be that these
+types are ushered into the arena by unexpected discoveries, we yet
+recognise in the psychological accidents, which Tarde places at the
+base of everything, near relatives of those small accidental
+variations upon which Darwin builds. Thus, accepting Tarde's own
+representations, it is quite possible to express in Darwinian terms,
+with the necessary transpositions, one of the most idealistic
+sociologies that have ever been constructed.
+
+These few examples suffice. They enable us to estimate the extent of
+the field of influence of Darwinism. It affects sociology not only
+through the agency of its advocates but through that of its opponents.
+The questions to which it has given rise have proved no less fruitful
+than the solutions it has suggested. In short, few doctrines, in the
+history of social philosophy, will have produced on their passage a
+finer crop of ideas.
+
+FOOTNOTES:
+
+[Footnote 246: P. Flourens, _Examen du Livre de M. Darwin sur
+l'Origine des Especes_, p. 53, Paris, 1864. See also Huxley,
+"Criticisms on the _Origin of Species," Collected Essays_, Vol. II, p.
+102, London, 1902.]
+
+[Footnote 247: _Inquiries into Human Faculty_, pp. 1, 2, 3 sq.,
+London, 1883.]
+
+[Footnote 248: _Darwinism and Politics_, pp. 9, 22, London, 1889.]
+
+[Footnote 249: _Life and Letters of Charles Darwin_, II. p. 385.]
+
+[Footnote 250: V. de Lapouge, _Les Selections sociales_, p. 259,
+Paris, 1896.]
+
+[Footnote 251: _Die nataerliche Auslese beim Menschen_, Jena, 1893; _Du
+Gesellschaftsordnung und ihre natuerlichen Grundlagen. Entwurf einer
+Sozialanthropologie_, Jena, 1896.]
+
+[Footnote 252: _Etudes sur la Selection dans ses rapports avec
+l'heredite chez l'homme_, Paris, p. 481, 1881.]
+
+[Footnote 253: _Die drei Bevoelkerungsstufen_, Munich, 1889.]
+
+[Footnote 254: _Evolution and Ethics_, p. 200; _Collected Essays_,
+Vol. IX, London, 1894.]
+
+[Footnote 255: _Les Luttes entre Societes humaines et leurs phases
+successives_, Paris, 1893.]
+
+[Footnote 256: _Le socialisme contemporain_, p. 384 (6th edit.),
+Paris, 1891.]
+
+[Footnote 257: Geddes and Thomson, _The Evolution of Sex_, p. 311,
+London, 1889.]
+
+[Footnote 258: _De la Division du Travail social_, Paris. 1893.]
+
+
+
+
+INDEX
+
+
+_Abraxas grossulariata_, 100
+
+Acquired characters, transmission of, 20, 28, 42, 94, 120, 149, 171, 173
+
+_Acraea johnstoni_, 290
+[Transcriber's Note: No such page number or reference seen.]
+
+Adaptation, 24, 27, 34, 39, 42-45, 50, 58, 79-86, 106, 107
+
+Adloff, 140
+
+Alexander, 217
+
+Ameghino, 132, 138
+
+Ammon, O., Works of, 271
+
+_Anaea divina_, 69
+
+Anglicus, Bartholomaeus, 237
+
+Ankyroderma, 40
+
+Anomma, 44
+
+Anthropops, 132
+
+Ants, modifications of, 43-46, 51
+
+Ardigo, 207, 208
+
+Argyll, Huxley and the Duke of, 238
+
+Aristotle, 3, 237, 240
+
+Avenarius, 211
+
+
+Bacon, on mutability of species, 4, 5
+
+Baehr, von, on Cytology, 99
+
+Bain, 194
+
+Baldwin, J. M., 53, Foot Note 165
+
+Balfour, A. J., 241
+
+Barratt, 217
+
+Bates, H. W., on Mimicry, 70, 76
+  --232
+
+BATESON, W., on _Heredity and
+Variation in Modern Lights_, 87-110
+  --on discontinuous evolution, 30
+
+Bathmism, 14
+
+Bells (Sir Charles) _Anatomy of Expression_, 177
+
+Bentham, Jeremy, 217, 218
+
+Bergson, H., 208
+
+Berkeley, 200
+
+Berthelot, 228
+
+Bickford, E., experiments on degeneration by, 52
+
+Biophores, 47
+
+Blumenbach, 89
+
+Bodin, 256
+
+Bonald, on war, 273
+
+Bonnet, 6
+
+BOUGLE, C., on _Darwinism and Sociology_, 264-280
+
+Bourdeau, 253
+
+Bourget, P., 270
+
+Boutroux, 208
+
+Brassica, hybrids of, 106
+
+_Brassica Napus_, 106
+
+Broca, 137, 270
+
+Brock, on Kant, Foot Note 6
+
+Brunetiere, 274
+
+Bruno, on Evolution, 4
+
+Buch, von, 15
+
+Buckle, 252, 253, 256, 258
+
+Buffon, 6-15, 21, 88
+
+Burdon-Sanderson, J., letter from, Foot Note 224
+
+BURY, J. B., on _Darwinism and History_, 246-263
+
+Butler, Samuel, 9, Foot Note 17, Foot Note 57, Foot Note 61, 94, Foot Note 66, 107
+
+Butterflies, mimicry in, 65-83
+  --sexual characters in, 59-63
+
+
+Cabanis, 201
+
+Candolle, de, 270
+
+Carneri, 217
+
+_Castnia linus_, 76
+
+Caterpillars, variation in, 36, 37
+
+Cesnola, experiments on Mantis by, 65
+
+Chaerocampa, colouring of, 68
+
+Chambers, R., _The Vestiges of Creation_ by, 15
+
+Chromosomes and Chromomeres, 47, 96-100
+
+Chun, Foot Note 36
+
+Claus, Foot Note 21
+
+Clodd, E., Foot Note 13
+
+Coadaptation, 41-54
+
+_Colobopsis truncata_, 44
+
+Colour, E. B. Poulton, in relation to Sexual Selection, 61-65
+
+Comte, A., 200-203, 252-255, 262, 265
+
+Condorcet, 221, 250, 252, 258
+
+Cope, 138
+
+Correlation of organisms, Darwin's idea of the, 2
+
+Cournot, 265
+
+Cuvier, 9, 10, 266, 268
+
+Cytology and heredity, 95, 96, 99, 100
+
+
+_Danaida chrysippus_, 75
+
+_Danaida genutia_, 75
+
+_D. Plexippus_, 75
+
+Dantec, Le, 274
+
+Darwin, Charles, as an Anthropologist, 146-165
+  --on ants, 44
+  --and S. Butler, Foot Note 61, 94
+  --on Cirripedia, 212
+  --on the Descent of Man, 111-145
+  --evolutionist authors referred to in the _Origin_ by, 9
+
+Darwin, Charles, and Haeckel, 137
+  --and History, 246-263
+  --and Huxley, 112
+  --on Lamarck, 28, 129
+  --on Language, 124
+  --and Malthus, 16, 24, 91
+  --on Patrick Matthew, 19
+  --on mental evolution, 166-196
+  --on Natural Selection, 21, 41, 54, 55, 122
+  --a "Naturalist for Naturalists," 87
+  --his personality, 187
+  --his influence on Philosophy, 197-222
+  --predecessors of, 1-22
+  --his views on religion, etc., 115, 116, 219-222
+  --his influence on religious thought, 223-245
+  --causes of his success, 10, 90
+
+Darwin, Charles, on the _Vestiges of Creation_, 15
+  --and Wallace, 23, 183
+  --on evolution, 7-15, 88
+  --on Lamarckism, 11
+
+Darwin, F., on Prichard's "Anticipations," 21
+
+Darwinism, Sociology, Evolution and, 17-18
+
+Degeneration, 49-51, 93
+
+Deniker, 137
+
+Descartes, 4
+
+Descent, history of doctrine of, 1
+
+_Descent of Man_, G. Schwalbe on _The_, 111-145
+  --rejection in Germany of _The_, 156
+
+Diderot, 6, 198
+
+Dimorphism, seasonal, 30
+
+_Dismorphia orise_, 75
+
+Dragomirov, 273
+
+Driesch, Foot Note 67
+
+Dryopithecus, 132
+
+Dubois, E., on Pithecanthropus, 132, 137
+
+Duehring, 214, 277
+
+Duns Scotus, 200
+
+Duret, C., 6
+
+Durkheim, on division of labour, 278
+
+
+Ecology, Foot Note 205
+
+Eimer, 109
+
+_Elymnias undularis_, 73, 75
+
+Embryology, the Origin of Species and, 154, 155
+
+Empedocles, 3, 27, 151
+
+Engels, 277
+
+Environment, action of, 12, 13, 15
+
+Epicurus, a poet of Evolution, 4
+
+Eristalis, 75
+
+Espinas, 275
+
+Evolution, and creation, 233
+  --conception of, 3-5, 9, 148, 151, 198
+  --discontinuous, 30
+  --experimental, 5, 7
+  --factors of, 11-15
+  --mental, 194
+  --Lloyd Morgan on mental factors in, 166-196
+  --Darwinism and Social, 18
+  --Saltatory, 29-32
+  --Herbert Spencer on, 204-207
+  --Philosophers and modern methods of studying, 4
+
+Expression of the Emotions, 177-184
+
+
+Ferri, 277
+
+Ferrier, his work on the brain, 523
+[Transcriber's note: No such page number or reference seen]
+
+Fichte, 222
+
+Flourens, 267
+
+Flowers and Insects, 61, 78
+
+Fouillee, 207, 208
+
+Fraipont, on skulls from Spy, 134
+
+
+GADOW, 162
+
+_Gallus bankiva_, 102
+
+Gallon, F., 125, 150, 269
+
+Geddes, P., 17, Foot Note 32
+
+Geddes, P. and A. W. Thomson, 276
+
+Gegenbaur, 150, 163
+
+Genetics, 93, 96
+
+_Germ-plasm_, continuity of, 95
+--Weismann on, 46-51
+
+Germinal Selection, 36, 37, 46-51, 64
+
+Gibbon, 248
+
+Giuffrida-Ruggeri, 138, 140
+
+Giotto, 259
+
+Gizycki, 217
+
+Goethe and Evolution, 8, 14, 15, 201
+--on the relation between Man and Mammals, 161, 163
+--221
+
+Gore, Dr., 226
+
+Gorjanovic-Kramberger, 134
+
+Gosse, P. H., 234
+
+_Grapta C. album_, 69
+
+Groos, 187, 188
+
+Gulick, 15, 53
+
+Guyau, 217
+
+
+Haberlandt, G., 34
+
+HAECKEL, E., on _Charles Darwin as an Anthropologist_, 146-165
+  --and Darwin, 135-151, 137, 146-165
+  --on the Descent of Man, 137, 143
+  --on Lamarck, 8, Foot Note 21
+  --a leader in the Darwinian controversy, 137
+  --217
+
+Haecker, 33
+
+Hansen, 272
+
+Hartmann, von, 240
+
+Harvey, 4
+
+Haycraft, 275
+
+Hegel, 201, 203, 215, 251, 252, 255
+
+Heraclitus, 278
+
+Herder, 4, 5, 20
+
+Heredity and Cytology, 95, 96
+  --Haeckel on, 147, 148, 149, 153
+  --and Variation, 87-110
+  --219, 224
+
+Hering, E., on Memory, 153
+
+Hertwig, O., 150
+
+History, Darwin and, 246-263
+
+Hobbes, T., 200, 215
+
+Hobhouse, 242
+
+HOeFFDING, H., on _The Influence of the Conception of Evolution
+                             on Modern Philosophy_, 197-222
+
+Holothurians, calcareous bodies in skin of, 37-41
+
+_Homo heidelbergensis_, Foot Note 118
+
+_H. neandertalensis_, 138
+
+_H. pampaeus_, 144
+
+_H. primigenius_, 133, 134, 138, 144
+
+_Homunculus_, 132
+
+Hooker, Sir J. D., and Darwin, 23, 116
+
+Huber, 170
+
+Huegel, F. von, Foot Note 221
+
+Hume, 200
+
+Hutcheson, 216
+
+Huxley, T. H., and Darwin, 112, 116, 268
+  --and the Duke of Argyll, 238
+  --on Lamarck, 89
+  --on Man, 111, 112, 137, 146, 156, 160, 163
+  --on Selection, 24, 91
+  --on transmission of acquired characters, 149
+  --14, 24, 104, 231-236, 273, 274
+
+Hybrids, Sterility of, 104, 105, 106
+
+
+Inheritance of acquired characters, 93, 94
+
+Insects and Flowers, 60, 61, 78, 79
+
+Instinct, 122, 172-175
+
+Irish Elk, an example of coadaptation, 41, 42, 45
+
+
+Jacoby, _Studies in Selection_ by, 272
+
+James, W., 180, 191, 211
+
+Jentsch, 275
+
+
+Kallima, protective colouring of, 35, 68, 70
+
+_K. inachis_, 68
+
+Kammerer's experiments on Salamanders, 28
+
+Kant, I., 4, 5, 6, 27, 198, 211, 212, 217, 221, 222
+
+Keane, on the Primates, 138
+
+Keith, on Anthropoid Apes, 138
+
+Kepler, 198
+
+Klaatsch, on Ancestry of Man, 140
+
+Klaatsch and Hauser, 134
+
+Knies, 266
+
+Koelliker, his views on Evolution, 29, 150
+
+Kollmann, on origin of human races, 144
+
+Korschinsky, 31
+
+Krause, E., Foot Note 10, 13
+
+Kropotkin, 214, 275
+
+
+Lamarck, his division of the Animal Kingdom, 160, 161
+  --Darwin's opinion of, 129
+  --on Evolution, 9-14, 21, 25, 171, 172, 173, 179, 180, 201, 202, 253
+  --on Man, 146, 148, 160, 163
+  --89, 109, 201, 202, 233
+
+Lamarckian principle, 28, 41-44, 50-54, 67, 84, 86
+
+Lamb, C., 229
+
+Lamettrie, 198
+
+Lamprecht, 260-263
+
+Lanessan, J. L. de, Foot Note 17, 275
+
+Lang, Foot Note 21
+
+Lange, 180
+
+Language, Darwin on, 123, 124
+  --Evolution and the Science of, 178, 179, 188
+
+Lankester, Sir E. Ray, on degeneration, 268
+  --on educability, 170, 189
+
+Lankester, Sir E. Ray, on the germ-plasm theory, 150
+
+Lapouge, Vacher de, 270
+
+Lartet, M. E., 189
+
+Lasalle, 266
+
+Laveleye, de, 275
+
+Lawrence, W., 89, Foot Note 65
+
+Lehmann-Nitsche, 138, 144
+
+Leibnitz, 4, 5, 213
+
+Lepidoptera, variation in, 37, 60-63
+
+Lessing, 4, 221
+
+Liddon, H. P., 234
+
+_Limenitis archippus_, 74
+
+Linnaeus, 6
+
+Locy, W. A., Foot Note 15
+
+Lovejoy, Foot Note 56
+
+Lubbock, 125
+
+Lucretius, a poet of Evolution, 4
+
+Lyell, Sir Charles, and Darwin, 23, 116
+  --the uniformitarian teaching of, 89
+
+
+Macacus, ear of, 119
+
+Mach, E., 153, 211
+
+Mahoudeau, 137
+
+Maillet, de, 6
+
+Majewski, Foot Note 238, Foot Note 239
+
+Malthus, his influence on Darwin, 16-18, 21, 24, 91
+  --200, 273
+
+Man, Descent of, 126, 127, 128, 131-145, 156-165, 189, 254, 265
+  --mental and moral qualities of animals and, 122-126, 164, 188-192
+  --pre-Darwinian views on the Descent of, 1
+
+Man, Tertiary flints worked by, 136
+
+_Man_, G. Schwalbe on Darwin's _Descent of_, 111-145
+
+Manouvrier, 137
+
+_Mantis religiosa_, colour experiments on, 65, 68
+
+Marx, 262, 276-278
+
+Matthew, P., and Natural Selection, 18, 19
+
+Maupertuis, 6, 88, 103
+
+Mayer, R., 197
+
+_Mechanitis lysimnia_, 77
+
+_Melinaea ethra_, 77
+
+Mendel, 97-100, 184, 228
+
+Merz, J. T., Foot Note 14
+
+Mesopithecus, 132
+
+Mill, J. S., 193, 200, 202, 218
+
+Mimicry, 70-82
+
+Moltke, on war, 273
+
+Monkeys, fossil, 132
+
+Montesquieu, 248
+
+Monticelli, 155
+
+MORGAN, C. LLOYD, on _Mental Factors in Evolution_, 166-196
+  --on Organic Selection, 53
+
+Morgan, T. H., 99
+
+Morselli, 138
+
+Mortillet, 136
+
+Moseley, Foot Note 224
+
+Muller, Fritz, _Fuer Darwin_ by, 154
+  --on Mimicry, 233
+  --59, 77
+
+Muller, J., 147
+
+Mueller, Max, on language, 124
+
+Mutation, 15, 31, 184, 199, 209
+
+
+Naegeli, 109, 151, 153
+
+Nathusius, 103
+
+Natural Selection, Darwin's views on, 90, 91, 122, 149
+  --Darwin and Wallace on, 2, 163, 183
+  --and design, 241, 242
+  --and educability, 195
+  --and human development, 125, 256, 257
+  --16-20, 25, 26, 41, 55-58, 64-86, 87-96, 199, 233
+
+Neandertal skulls, 133, 134
+
+Neodarwinism, 150
+
+Newton, A., Foot Note 59
+
+Newton, I., 197, 198
+
+Niebuhr, 249, 263
+
+Nietzsche, 214, 271
+
+Nitsche, 119
+
+Novicow, 274
+
+Nuttall, G. H. F., 135
+
+
+Occam, 200
+
+Odin, 270
+
+Oecology, see Ecology
+
+_Oenothera lamarckiana_, 32
+
+Oestergren, on Holothurians, 37-39
+
+Oken, L., 7, 201
+
+Organic Selection, 53, 54, 172, 173
+
+Orthogenesis, 109
+
+Osborn, H. F., 53, Foot Note 165
+  --_From the Greeks to Darwin_ by, 3-5, 12, 14, 20
+
+_Ovibos moschatus_, 67
+
+Owen, Sir Richard, 111
+
+
+Packard, A. S., Foot Note 12, Foot Note 18
+
+Palaeopithecus, 132
+
+Paley, 18, 242, 244
+
+Panmixia, Weismann's principle of, 54
+
+_Papilio dardanus_, 72, 73, 74
+
+_P. meriones_, 73
+
+_P. merope_, 72
+
+Pearson, K., Foot Note 7
+
+Penck, 136
+
+Peridineae, 33
+
+Perrier, E., Foot Note 21, 20
+
+Perthes, B. de, 123
+
+Pfeffer, W., 28
+
+Philosophy, influence of the conception of evolution on modern, 197-222
+
+Pithecanthropus, 133, 134, 138, 143
+
+Pitheculites, 144
+
+Plate, Foot Note 37
+
+Pliopithecus, 132
+
+Pouchet, G., Foot Note 3
+
+POULTON, E. B., experiments on Butterflies by, 65
+  --on J. C. Prichard, 20
+  --on Mimicry, 69, 71, 75, 78
+  --Foot Note 34, Foot Note 43, Foot Note 49, Foot Note 55
+
+Prichard, J. C., 20, 21, 89, Foot Note 65
+
+_Pronuba yuccasella_, 79
+
+Protective resemblance, 65-70
+
+Pusey, 115
+
+
+Quatrefages, A. de, Foot Note 21, 19
+
+
+Radiolarians, 33
+
+Ranke, 249, 251, 255, 263
+
+Rau, A., 153
+
+Ray, J., 4
+
+Regeneration, Foot Note 71
+
+Religious thought, Darwin's influence on, 223-245
+
+Reversion, 120, 121
+
+Ridley, H. N., Foot Note 88
+
+Ritchie, 270
+
+Robinet, 6
+
+Rolph, 217
+
+Romanes, G. J., Foot Note 3, 15, 32, 54, 164, 234
+
+Roux, 151, 152
+
+Ruskin, 230
+
+Rutot, 136
+
+Saint-Hilaire, E. G. de, 8, 15, 20
+
+Saltatory Evolution, 29-32 (see also Mutations)
+
+Sanders, experiments on Vanessa by, 65
+
+Savigny, 249
+
+Schelling, 4, 5, 200, 201
+
+Schleiden and Schwann, Cell-theory of, 147
+
+Schoetensack, on _Homo heidelbergensis_, Foot Note 118
+
+Schuett, 23
+
+SCHWALBE, G., on _The Descent of Man_, 111-145
+
+Seeck, O., Foot Note 240
+
+Segregation, 97, 98
+
+Selection, artificial, 24, 25, 26, 41, 45, 120, 269-272
+  --germinal, 35, 36, 46-52, 64
+
+Selection, natural (see Natural Selection)
+  --organic, 53, 171, 172
+  --sexual, 55-64, 117, 118
+  --social and natural, 271
+  --23-86, 103, 129, 130
+
+Selenka, 131
+
+Semnopithecus, 132
+
+Semon, R., 28, 153
+
+Sergi, 138, 143
+
+Sex, recent investigations on, 99, 100
+
+Sibbern, 201
+
+_Smerinthus ocellata_, 38
+
+_Smerinthus populi_, 38
+
+_S. tiliae_, 38
+
+Smith, A., 200
+
+Sociology,  Darwinism and, 264-280
+  --History and, 255
+
+Sollas, W. J., 134
+
+Sorley, W. R., 217
+
+Species and varieties, 100
+
+Spencer, H., on evolution, 204-209
+  --on the theory of Selection, 41
+
+Spencer, H., on Sociology, 268
+  --on the transmission of acquired characters, 149
+  --on Weismann, 41, 150
+  --2, 17, 217, 231, 268
+
+Sphingidae, variation in, 37
+
+Spinoza, 153, 206
+
+Standfuss, 82
+
+Stephen, L., 217
+
+Sterility in hybrids, 104-106
+
+Sterne, C, Foot Note 10
+
+Struggle for existence, 25, 26, 272-274
+
+Sutton, A. W., Foot Note 73
+
+Synapta, calcareous bodies in skin of, 38-41
+
+Syrphus, 75
+
+
+Tarde, G., 279
+
+Tennant, F. R., Foot Note 218
+
+Tetraprothomo, 138, 144
+
+THOMSON, J. A., on _Darwin's Predecessors_, 1-22
+  --150
+  --and P. Geddes, 276
+
+Treschow, 201
+
+Treviranus, 8, 14, 15
+
+Turgot, 249
+
+Turner, Sir W., 150
+
+Tylor, 267
+
+Tyndall, W., 267
+
+Tyrrell, G, Foot Note 222
+
+
+Uhlenhuth, on blood reactions, 135
+
+Use and disuse, 28, 41-43, 48-54, 94, 95, 119, 149
+
+
+Vanessa, 63
+
+_V. levana_, 31
+
+_V. polychloros_, 82
+
+_V. urticae_, 65, 82
+
+Variability, Darwin's attention directed to, 24
+  --W. Bateson on, 87-110
+  --causes of, 200
+
+Variation, Darwin's views as an evolutionist, and as a systematist, on, 212
+  --and heredity, 87-110
+  --minute, 28-32
+  --in relation to species, 100, 101
+
+Varigny, H. de, 6, 19
+
+Verworn, 136
+
+_Vestiges of Creation_, Darwin on _The_, 15
+
+Virchow, his opposition to Darwin, 157, 158
+  --on the transmission of acquired characters, 149
+
+Vogt, 137
+
+Voltaire, 248
+
+VRIES, H. de, the Mutation theory of, 31, 101, 151, 213
+
+
+WAGGETT, Rev. P. N., on _The Influence of Darwin upon Religious
+                                    Thought_, 223-245
+
+Wallace, A. R., on Colour, 63, 71
+  --and Darwin, Foot Note 7, 23, 183
+  --on the Descent of Man, 116
+  --on Malthus, 17
+  --on Natural Selection, 2, 16, 163, 232
+
+Wallace, A. R., on social reforms, 275, 276
+  --on Sexual Selection, 183, 184
+
+Walton, 237
+
+Watt, J., and Natural Selection, 21
+
+WEISMANN, A., on _The Selection Theory_, 23-86
+  --his germ-plasm theory, 46-51, 149, 150
+  --and Prichard, 20
+  --and Spencer, 42
+
+Weismann, A., on the transmission of acquired characters, 93-95
+  --156
+
+Wells, W. C, and Natural Selection, 18
+
+White, G., 3
+
+Williams, C. M., 217
+
+Wilson, E. B., on cytology, 99
+
+Wolf, 249
+
+Wollaston's, T. V., _Variation of Species_, Foot Note 59
+
+Woltmann, 277
+
+Woolner, 118
+
+Wundt, on language, 207, 208
+
+
+_Xylina vetusta_, 82
+
+
+Yucca, fertilisation of, 78, 79
+
+
+Zeller, E., Foot Note 3
+
+_Zoonomia_, Erasmus Darwin's, 7
+
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