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diff --git a/.gitattributes b/.gitattributes new file mode 100644 index 0000000..d7b82bc --- /dev/null +++ b/.gitattributes @@ -0,0 +1,4 @@ +*.txt text eol=lf +*.htm text eol=lf +*.html text eol=lf +*.md text eol=lf diff --git a/LICENSE.txt b/LICENSE.txt new file mode 100644 index 0000000..6312041 --- /dev/null +++ b/LICENSE.txt @@ -0,0 +1,11 @@ +This eBook, including all associated images, markup, improvements, +metadata, and any other content or labor, has been confirmed to be +in the PUBLIC DOMAIN IN THE UNITED STATES. + +Procedures for determining public domain status are described in +the "Copyright How-To" at https://www.gutenberg.org. + +No investigation has been made concerning possible copyrights in +jurisdictions other than the United States. Anyone seeking to utilize +this eBook outside of the United States should confirm copyright +status under the laws that apply to them. diff --git a/README.md b/README.md new file mode 100644 index 0000000..9f396ea --- /dev/null +++ b/README.md @@ -0,0 +1,2 @@ +Project Gutenberg (https://www.gutenberg.org) public repository for +eBook #52312 (https://www.gutenberg.org/ebooks/52312) diff --git a/old/52312-8.txt b/old/52312-8.txt deleted file mode 100644 index 597674e..0000000 --- a/old/52312-8.txt +++ /dev/null @@ -1,1173 +0,0 @@ -Project Gutenberg's The Methods and Scope of Genetics, by W. Bateson - -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/license - - -Title: The Methods and Scope of Genetics - An inaugural lecture delivered 23 October 1908 - -Author: W. Bateson - -Release Date: June 12, 2016 [EBook #52312] - -Language: English - -Character set encoding: ISO-8859-1 - -*** START OF THIS PROJECT GUTENBERG EBOOK THE METHODS AND SCOPE OF GENETICS *** - - - - -Produced by Chris Curnow, Martin Pettit and the Online -Distributed Proofreading Team at http://www.pgdp.net (This -file was produced from images generously made available -by The Internet Archive) - - - - - - -THE METHODS AND SCOPE OF GENETICS - - -CAMBRIDGE UNIVERSITY PRESS -London: FETTER LANE, E.C. -C. F. CLAY, MANAGER - -[Illustration: Logo] - -Edinburgh: 100, PRINCES STREET -Berlin: A. ASHER AND CO. -Leipzig: F. A. BROCKHAUS -New York: G. P. PUTNAM'S SONS -Bombay and Calcutta: MACMILLAN AND CO., LTD. - -_All rights reserved_ - - - - -THE METHODS AND SCOPE OF GENETICS - -_AN INAUGURAL LECTURE DELIVERED 23 OCTOBER 1908_ - -by -W. BATESON, M.A., F.R.S. - -PROFESSOR OF BIOLOGY IN THE UNIVERSITY OF CAMBRIDGE - -Cambridge: -at the University Press -1912 - - -_First Edition 1908_ -_Reprinted 1912_ - - - - -PREFATORY NOTE - - -The Professorship of Biology was founded in 1908 for a period of five -years partly by the generosity of an anonymous benefactor, and partly by -the University of Cambridge. The object of the endowment was the -promotion of inquiries into the physiology of Heredity and Variation, a -study now spoken of as Genetics. - -It is now recognized that the progress of such inquiries will chiefly be -accomplished by the application of experimental methods, especially -those which Mendel's discovery has suggested. The purpose of this -inaugural lecture is to describe the outlook over this field of research -in a manner intelligible to students of other parts of knowledge. - -W. B. - -_28 October, 1908_ - - - - -THE METHODS AND SCOPE OF GENETICS - - -The opportunity of addressing fellow-students pursuing lines of inquiry -other than his own falls seldom to a scientific man. One of these rare -opportunities is offered by the constitution of the Professorship to -which I have had the honour to be called. That Professorship, though -bearing the comprehensive title "of Biology," is founded with the -understanding that the holder shall apply himself to a particular class -of physiological problems, the study of which is denoted by the term -Genetics. The term is new; and though the problems are among the oldest -which have vexed the human mind, the modes by which they may be -successfully attacked are also of modern invention. There is therefore -a certain fitness in the employment of this occasion for the deliverance -of a discourse explaining something of the aims of Genetics and of the -methods by which we trust they may be reached. - -You will be aware that the claims put forward in the name of Genetics -are high, but I trust to be able to show you that they are not high -without reason. It is the ambition of every one who in youth devotes -himself to the search for natural truth, that his work may be found -somewhere in the main stream of progress. So long only as he keeps -something of the limitless hope with which his voyage of discovery -began, will his courage and his spirit last. The moment we most dread is -one in which it may appear that, after all, our effort has been spent in -exploring some petty tributary, or worse, a backwater of the great -current. It is because Genetic research is still pushing forward in the -central undifferentiated trunk of biological science that we confess no -guilt of presumption in declaring boldly that whatever difficulty may be -in store for those who cast in their lot with us, they need fear no -disillusionment or misgiving that their labour has been wasted on a -paltry quest. - -In research, as in all business of exploration, the stirring times come -when a fresh region is suddenly unlocked by the discovery of a new key. -Then conquest is easy and there are prizes for all. We are happy in that -during our own time not a few such territories have been revealed to the -vision of mankind. I do not dare to suggest that in magnitude or -splendour the field of Genetics may be compared with that now being -disclosed to the physicist or the astronomer; for the glory of the -celestial is one and the glory of the terrestrial is another. But I -will say that for once to the man of ordinary power who cannot venture -into those heights beyond, Mendel's clue has shown the way into a realm -of nature which for surprising novelty and adventure is hardly to be -excelled. - -It is no hyperbolical figure that I use when I speak of Mendelian -discovery leading us into a new world, the very existence of which was -unsuspected before. - -The road thither is simple and easy to follow. We start from a common -fact, familiar to everyone, that all the ordinary animals and plants -began their individual life by the union of two cells, the one male, the -other female. Those cells are known as germ-cells or _gametes_, that is -to say, "marrying" cells. - -Now obviously the diversity of form which is characteristic of the -animal and plant world must be somehow represented in the gametes, since -it is they which bring into each organism all that it contains. I am -aware that there is interplay between the organism and the circumstances -in which it grows up, and that opportunity given may bring out a -potentiality which without that opportunity must have lain dormant. But -while noting parenthetically that this question of opportunity has an -importance, which some day it may be convenient to estimate, the one -certain fact is that all the powers, physical and mental that a living -creature possesses were contributed by one or by both of the two -germ-cells which united in fertilisation to give it existence. The fact -that _two_ cells are concerned in the production of all the ordinary -forms of life was discovered a long while ago, and has been part of the -common stock of elementary knowledge of all educated persons for about -half a century. The full consequences of this double nature seem -nevertheless to have struck nobody before Mendel. Simple though the -fact is, I have noticed that to many it is difficult to assimilate as a -working idea. We are accustomed to think of a man, a butterfly, or an -apple tree as each _one_ thing. In order to understand the significance -of Mendelism we must get thoroughly familiar with the fact that they are -each _two_ things, double throughout every part of their composition. -There is perhaps no better exercise as a preparation for genetic -research than to examine the people one meets in daily life and to try -in a rough way to analyse them into the two assemblages of characters -which are united in them. That we are assemblages or medleys of our -parental characteristics is obvious. We all know that a man may have his -father's hair, his mother's colour, his father's voice, his mother's -insensibility to music, and so on, but that is not enough. - -Such an analysis is true, inasmuch as the various characters _are_ -transmitted independently, but it misses the essential point. For in -each of these respects the individual is double; and so to get a true -picture of the composition of the individual we have to think how _each_ -of the two original gametes was provided in the matter of height, hair, -colour, mathematical ability, nail-shape, and the other features that go -to make the man we know. The contribution of each gamete in each respect -has thus to be separately brought to account. If we could make a list of -all the ingredients that go to form a man and could set out how he is -constituted in respect of each of them, it would not suffice to give one -column of values for these ingredients, but we must rule two columns, -one for the ovum and one for the spermatozoon, which united in -fertilisation to form that man, and in each column we must represent how -that gamete was supplied in respect of each of the ingredients in our -list. When the problem of heredity is thus represented we can hardly -avoid discovering, by mere inspection, one of the chief conclusions to -which genetic research has led. For it is obvious that the contributions -of the male and female gametes may in respect of any of the ingredients -be either the same, or different. In any case in which the contribution -made by the two cells is the same, the resulting organism--in our -example the man--is, as we call it, _pure-bred_ for that ingredient, and -in all respects in which the contribution from the two sides of the -parentage is dissimilar the resulting organism is _cross-bred_. - -To give an intelligible account of the next step in the analysis without -having recourse to precise and technical language is not very easy. - -We have got to the point of view from which we see the individual made -up of a large number of distinct ingredients, contributed from two -sources, and in respect of any of them he may have received two similar -portions or two dissimilar portions. We shall not go far wrong if we -extend and elaborate our illustration thus. Let us imagine the contents -of a gamete as a fluid made by taking a drop from each of a definite -number of bottles in a chest, containing tinctures of the several -ingredients. There is one such chest from which the male gamete is to be -made up, and a similar chest containing a corresponding set of bottles -out of which the components of the female gamete are to be taken. But in -either chest one or more of the bottles may be empty; then nothing goes -in to represent that ingredient from that chest, and if corresponding -bottles are empty in both chests, then the individual made on -fertilisation by mixing the two collections of drops together does not -contain the missing ingredient at all. It follows therefore that an -individual may thus be "pure-bred," namely alike on both sides of his -composition as regards each ingredient in one of two ways, either by -having received the ingredient from the male chest and from the female, -or in having received it from neither. Conversely in respect of any -ingredient he may be "cross-bred," receiving the presence of it from one -gamete and the absence of it from the other. - -The second conception with which we have now to become thoroughly -familiar is that of the individual as composed of what we call presences -and absences of all the possible ingredients. It is the basis of all -progress in genetic analysis. Let me give you two illustrations. A blue -eye is due to the absence of a factor which forms pigment on the front -of the iris. Two blue-eyed parents therefore, as Hurst has proved, do -not have dark-eyed children. The dark eye is due to either a single or -double dose of the factor missing from the blue eye. So dark-eyed -persons may have families all dark-eyed, or families composed of a -mixture of dark and light-eyed children in certain proportions which on -the average are definite. - -Two plants of _Oenothera_ which I exhibit illustrate the same thing. One -of them is the ordinary _Lamarckiana_. I bend its stem. It will not -break, or only breaks with difficulty on account of the tough fibres it -contains. The stem of the other, one of de Vries' famous mutations, -snaps at once like short pastry, because it does not contain the factor -for the formation of the fibres. Such plants may be sister-plants -produced by the self-fertilisation of one parent, but they are distinct -in their composition and properties--and this distinction turns on the -presence or absence of elements which are treated as definite entities -when the germ-cells are formed. When we speak of such qualities as the -formation of pigment in an eye, or the development of fibres in a stem, -as due to transmitted elements or factors, you will perhaps ask if we -have formed any notion as to the actual nature of those factors. For my -own part as regards that ulterior question I confess to a disposition to -hold my fancy on a tight rein. It cannot be very long before we shall -_know_ what some of the factors are, and we may leave guessing till -then. Meanwhile however there is no harm in admitting that several of -them behave much as if they were ferments, and others as if they -constructed the substances on which the ferments act. But we must not -suppose for a moment that it is the ferment, or the objective substance, -which is transmitted. The thing transmitted can only be the power or -faculty to produce the ferment or the objective substance. - -So far we have been considering the synthesis of the individual from -ingredients brought into him by the two gametes. In the next step of our -consideration we reverse the process, and examine how the ingredients of -which he was originally compounded are distributed among the gametes -that are eventually budded off from him. - -Take first the case of the components in respect of which he is -pure-bred. Expectation would naturally suggest that all the germ-cells -formed from him would be alike in respect of those ingredients, and -observation shows, except in the rare cases of originating variations, -the causation of which is still obscure, that this expectation is -correct. - -Hitherto though without experimental evidence no one could have been -certain that the facts were as I have described them, yet there is -nothing altogether contrary to common expectation. But when we proceed -to ask how the germ-cells will be constituted in the case of an -individual who is cross-bred in some respect, containing that is to say, -an ingredient from the one side of his parentage and not from the other, -the answer is entirely contrary to all the preconceptions which either -science or common sense had formed about heredity. For we find definite -experimental proof in nearly all the cases which have been examined, -that the germ-cells formed by such individuals do either contain or not -contain a representation of the ingredient, just as the original gametes -did or did not contain it. - -If _both_ parent-gametes brought a certain quality in, then all the -daughter gametes have it; if neither brought it in, then none of the -daughter gametes have it. If it came in from one side and not from the -other, then on an average in half the resulting gametes it will be -present and from half it will be absent. This last phenomenon, which is -called segregation, constitutes the essence of Mendel's discovery. - -So recurring to the simile of the man as made by the mixing of -tinctures, the process of redistribution of his characters among the -germ-cells may be represented as a sorting back of the tinctures again -into a double row of bottles, a pair corresponding to each ingredient; -and each of the germ-cells as then made of a drop from one or other -bottle of each pair: and in our model we may represent the phenomenon of -segregation in a crude way by supposing that the bottles having no -tincture in them, instead of being empty contained an inoperative fluid, -say water, with which the tincture would not mix. When the new -germ-cells are formed, the two fluids instead of diluting each other -simply separate again. It is this fact which entitles us to speak of the -purity of germ-cells. They are pure in the possession of an ingredient, -or in not possessing it; and the ingredients, or factors, as we -generally call them, are units because they are so treated in the -process of formation of the new gametes and because they come out of the -process of segregation in the same condition as they went in at -fertilisation. - -As a consequence of these facts it follows that however complex may be -the origin of two given parents the composition of the offspring they -can produce is limited. There is only a limited number of types to be -made by the possible recombinations of the parental ingredients, and the -relative numbers in which each type will be represented are often -predicable by very simple arithmetical rules. - -For example, if neither parent possesses a certain factor at all, then -none of the offspring will have it. If either parent has two doses of -the factor then all the children will have it; and if either parent has -one dose of the factor and the other has none, then on an average half -the family will have it, and half be without it. - -To know whether the parent possesses the factor or not may be difficult -for reasons which will presently appear, but often it is quite easy and -can be told at once, for there are many factors which cannot be present -in the individual without manifesting their presence. I may illustrate -the descent of such a factor by the case of a family possessing a -peculiar form of night-blindness. The affected individuals marrying with -those unaffected have a mixture of affected and unaffected children, but -their unaffected children not having the responsible ingredient cannot -pass it on[1]. - -In such an observation two things are strikingly exemplified, (1) the -fact of the permanence of the unit, and (2) the fact that a _mixture_ of -types in the family means that one or other parent is cross-bred in some -respect, and is giving off gametes of more than one type. - -The problem of heredity is thus a problem primarily analytical. We have -to detect and enumerate the factors out of which the bodies of animals -and plants are built up, and the laws of their distribution among the -germ-cells. All the processes of which I have spoken are accomplished by -means of cell-divisions, and in the one cell-union which occurs in -fertilisation. If we could watch the factors segregating from each -other in cell-division, or even if by microscopic examination we could -recognize this multitudinous diversity of composition that must -certainly exist among the germ-cells of all ordinary individuals, the -work of genetics would be much simpler than it is. - -But so far no such direct method of observation has been discovered. In -default we are obliged to examine the constitution of the germ-cells by -experimental breeding, so contrived that each mating shall test the -composition of an individual in one or more chosen respects, and, so to -speak, sample its germ-cells by counting the number of each kind of -offspring which it can produce. But cumbersome as this method must -necessarily be, it enables us to put questions to Nature which never -have been put before. She, it has been said, is an unwilling witness. -Our questions must be shaped in such a way that the only possible -answer is a direct "Yes" or a direct "No." By putting such questions we -have received some astonishing answers which go far below the surface. -Amazing though they be, they are nevertheless true; for though our -witness may prevaricate, she cannot lie. Piecing these answers together, -getting one hint from this experiment, and another from that, we begin -little by little to reconstruct what is going on in that hidden world of -gametes. As we proceed, like our brethren in other sciences, we -sometimes receive answers which seem inconsistent or even contradictory. -But by degrees a sufficient body of evidence can be attained to show -what is the rule and what the exception. My purpose today must be to -speak rather of the regular than of the irregular. - -One clear exception I may mention. Castle finds that in a cross between -the long-eared lop-rabbit and a short-eared breed, ears of intermediate -length are produced: and that these intermediates breed approximately -true. - -Exceptions in general must be discussed elsewhere. Nevertheless if I may -throw out a word of counsel to beginners, it is: Treasure your -exceptions! When there are none, the work gets so dull that no one cares -to carry it further. Keep them always uncovered and in sight. Exceptions -are like the rough brickwork of a growing building which tells that -there is more to come and shows where the next construction is to be. - -You will readily understand that the presentation here given of the -phenomena is only the barest possible outline. Some of the details we -may now fill in. For example, I have spoken of the characters of the -organism, its colour, shape, and the like, as if they were due each to -one ingredient or factor. Some of them are no doubt correctly so -represented; but already we know numerous bodily features which need the -concurrence of several factors to produce them. Nevertheless though the -character only appears when all the complementary ingredients are -together present, each of these severally and independently follows, as -regards its transmission, the simple rules I have described. - -This complementary action may be illustrated by some curious results -that Mr Punnett and I have encountered when experimenting with the -height of Sweet Peas. There are two dwarf varieties, one the prostrate -"Cupid," the other the half-dwarf or "Bush" Sweet Peas. Crossed together -they give a cross-bred of full height. There is thus some element in the -Cupid which when it meets the complementary element from the Bush, -produces the characteristic length of the ordinary Sweet Pea. We may -note in passing that such a fact demonstrates at once the nature of -Variation and Reversion. The Reversion occurs because the two factors -that made the _height_ of the old Sweet Pea again come together after -being parted: and the Variations by which each of the dwarfs came into -existence must have taken place by the dropping out of one of these -elements or of the other. - -Conversely there are factors which by their presence can prevent or -inhibit the development and appearance of others present and -unperceived. - -For example, all the factors for pigmentation may be present in a plant -or an animal; but in addition there may be another factor present which -keeps the individual white, or nearly so. - -There are cases in which the action of the factors is superposed one on -top of the other, and not until each factor is removed in turn can the -effects of the underlying factors be perceived. So in the mouse if no -other colour-factor is present, the fur is chocolate. If the next factor -in the series be there, it is black. If still another factor be added, -it has the brownish grey of the common wild mouse. Conversely, by the -variation which dropped out the top factor, a black mouse came into -existence. By the loss of the black factor, the chocolate mouse was -created, and for aught we can tell there may be still more possibilities -hidden beneath. - -In the disentanglement of the properties and interactions of these -elementary factors, the science we must call to our aid is Physiological -Chemistry. The relations of Genetics with the other branches of biology -are close. Such work can only be conducted by those who have the good -fortune to be able to count upon continual help and advice from -specialists in the various branches of Zoology, Physiology, and Botany. -Often we have questions with which only a cytologist can deal, and -often it is the experience of a systematist we must invoke. The school -of Genetics in Cambridge starts under happy auspices in that we are -surrounded by colleagues qualified, and as we have often found, willing -to give us such aid unstinted. But with chemical physiology, we stand in -an even closer relation; and from the little I have dared to say -respecting the action and interaction of factors, it is evident that for -their disentanglement there must one day be an intimate and enduring -partnership arranged with the physiological chemists. - -Now, as the whole of the elaborate process by which the various elements -are apportioned among the gametes must be got through in a few -cell-divisions at most, and perhaps in one division only, it is not -surprising that there is sometimes an interaction between factors that -have quite distinct rôles to perform. These interactions are probably -of several kinds. One, which I shall illustrate presently, is probably -to be represented as a repulsion between two factors. As a consequence -of its operations when the various factors are sorted out into the -gametes, if the individual be cross-bred in respect of the _two_ -repelling factors, having received so to speak only a single dose of -each, then the gametes are made up in such a way that each takes one or -other of the two repelling factors, not both. - -Mutual repulsions of this kind probably play a significant part in the -phenomena of heredity. A single concrete case which Mr Punnett and I -have been investigating for some years will illustrate several of these -principles. We crossed together a pure white Sweet Pea having an erect -standard, with another pure white Sweet Pea having a hooded standard. -The result is, as you see, a purple flower with an erect standard. The -colour comes from the concurrence of complementary elements. A dose of a -certain ingredient from one parent meets a dose of another ingredient -from the other parent and the two make pigment in the flower. From other -experiments we know that the _purple_ colour of the pigment is due to a -dose of a third ingredient brought in from the hooded parent; and that -in the absence of that blue factor, as we may call it, the flower would -be red. The standard is erect because it contains a dose of the -erectness-factor from the erect parent, and the hooded parent can -readily be proved to owe its peculiar shape to the absence of that -element. - -Our purple plant is thus cross-bred for four factors, containing only -one dose of each. - -We let it fertilise itself, and its offspring show all the possible -combinations of the four different factors and their absences which the -genetic constitution of the plant can make. - -Note that one of the combinations we expect to find is missing. There -are white erect and white hooded--white because they are lacking one or -other of the complementary ingredients necessary to the production of -pigment. There are purple erect and purple hooded, of which the purple -erect must perforce contain all the four factors, and the purple hooded -must similarly contain all of them except that for erectness. But when -we turn to the red class we are surprised to find that they are all -erect, none hooded. One of the possible combinations is missing. If you -examine this series of facts you will find there is only one possible -interpretation: namely that the ingredient which turns the flower -purple--alkalinity, perhaps we may call it--never goes into the same -germ-cell as the ingredient which makes the standard erect. There are -plenty of ways of testing the truth of this interpretation. For example, -it follows that the purple erects from such a family will in perpetuity -have offspring 1 purple hooded: 2 purple erect: 1 red erect; also that -all the white hooded crossed with pure reds will give purples, and so -on. These experiments have been made and the result has in each case -been conformable to expectation. - -Between these two factors, the purpleness and the erectness of standard, -some antagonism or repulsion must exist. In some way therefore the -chemical and the geometrical phenomena of heredity must be -inter-related. - -Some one will say perhaps this is all very well as a scientific -curiosity, but it has nothing to do with real life. The right answer to -such criticism is of course the lofty one that science and its -applications are distinct: that the investigator fixes his gaze solely -on the search for truth and that his attention must not be distracted by -trivialities of application. But while we make this answer and at least -try to work in the spirit it proclaims, we know in our hearts that it is -a counsel of perfection. I suspect that even the astronomer who at his -spectroscope is analysing the composition of Vega or Capella has still -an eye sometimes free for the affairs of this planet, and at least the -fact that his discoveries may throw light on our destinies does not -diminish his zeal in their pursuit. And surely to the study of Heredity, -preeminently among all the sciences, we are looking for light on human -destiny. To pretend otherwise would be mere hypocrisy. So while -reserving the higher line of defence I will reply that again and again -in our experimental work we come very near indeed to human affairs. -Sometimes this is obvious enough. No practical dog-breeder or seeds-man -can see the results of Mendelian recombination without perceiving that -here is a bit of knowledge he can immediately apply. No sociologist can -examine the pedigrees illustrating the simple descent of a deformity or -a congenital disease, and not see that the new knowledge gives a solid -basis for practical action by which the composition of a race could be -modified if society so chose. More than this: we know for certain in one -case, from the work of Professor Biffen, that the power to resist a -disease caused by the invasion of a pathogenic organism, wheat-rust, is -due to the absence of one of the simple factors or ingredients of which -I have spoken, and what we know to be true in that one case we are -beginning to suspect to be true of resistance to certain other diseases. -No pathologist can see such an experiment as this of Professor Biffen's -without realizing that here is a contribution of the first importance -to the physiology of disease. - -There is no lack of utility and direct application in the study of -Genetics. I have alluded to some strictly practical results. If we want -to raise mangels that will not run to seed, or to breed a cow that will -give more milk in less time, or milk with more butter and less water, we -can turn to Genetics with every hope that something can be done in these -laudable directions. But here I would plead what I cannot but regard as -a higher usefulness in our work. Genetic inquiry aims at providing -knowledge that may bring, and I think will bring, certainty into a -region of human affairs and concepts which might have been supposed -reserved for ages to be the domain of the visionary. We have long known -that it was believed by some that our powers and conduct were dependent -on our physical composition, and that other schools have maintained -that nurture not nature, to use Galton's antithesis, has a -preponderating influence on our careers; but so soon as it becomes -common knowledge--not a philosophical speculation, but a certainty--that -liability to a disease, or the power of resisting its attack, addiction -to a particular vice, or to superstition, is due to the presence or -absence of a specific ingredient; and finally that these characteristics -are transmitted to the offspring according to definite, predicable -rules, then man's views of his own nature, his conceptions of justice, -in short his whole outlook on the world, must be profoundly changed. Yet -as regards the more tangible of these physical and mental -characteristics there can be little doubt that before many years have -passed the laws of their transmission will be expressible in simple -formulae. - -The blundering cruelty we call criminal justice will stand forth -divested of natural sanction, a relic of the ferocious inventions of -the savage. Well may such justice be portrayed as blind. Who shall say -whether it is crime or punishment which has wrought the greater -suffering in the world? We may live to know that to the keen satirical -vision of Sam Butler on the pleasant mountains of Erewhon there was -revealed a dispensation, not kinder only, but wiser than the terrific -code which Moses delivered from the flames of Sinai. - -If there are societies which refuse to apply the new knowledge, the -fault will not lie with Genetics. I think it needs but little -observation of the newer civilisations to foresee that _they_ will apply -every scrap of scientific knowledge which can help them, or seems to -help them in the struggle, and I am good enough Selectionist to know -that in that day the fate of the recalcitrant communities is sealed. - -The thrill of discovery is not dulled by a suspicion that the discovery -can be applied. No harm is done to the investigator if he can resist the -temptation to deviate from his aim. With rarest exceptions the -discoveries which have formed the basis of physical progress have been -made without any thought but for the gratification of curiosity. Of this -there can be few examples more conspicuous than that which Mendel's work -presents. Untroubled by any itch to make potatoes larger or bread -cheaper, he set himself in the quiet of a cloister garden to find out -the laws of hybridity, and so struck a mine of truth, inexhaustible in -brilliancy and profit. - -I will now suggest to you that it is by no means unlikely that even in -an inquiry so remote as that which I just described in the case of the -Sweet Pea, we may have the clue to a mystery which concerns us all in -the closest possible way. I mean the problem of the physiological nature -of Sex. In speaking of the interpretation of sexual difference -suggested by our experimental work as of some practical moment, I do not -imply that as in the other instances I have given, the knowledge is -likely to be of immediate use to our species; but only that if true it -makes a contribution to the stock of human ideas which no one can regard -as insignificant. - -In the light of Mendelian knowledge, when a family consists of more than -one type the fact means that the germ-cells of one or other parent must -certainly be of more than one kind. In the case of sex the members of -the family are thus of two kinds, and the presumption is overwhelming -that this distinction is due to a difference among the germ-cells. Next, -since for all practical purposes the numbers of the two sexes produced -are approximately equal, sex exhibits the special case in which a family -consists of two types represented in equal numbers, half being male, -half female. But I called your attention to the fact that equality of -types results when _one_ parent was cross-bred in the character -concerned, having received one dose only of the factor on which it -depends. So we may feel fairly sure that the distinction between the -sexes depends on the presence in one or other of them of an unpaired -factor. This conclusion appears to me to follow so immediately on all -that we have learnt of genetic physiology that with every confidence we -may accept it as representing the actual fact. - -The question which of the two sexes contains the unpaired factor is less -easy to answer, but there are several converging lines of evidence which -point to the deduction that in Vertebrates at least, and in some other -types, it is the female, and I feel little doubt that we shall succeed -in proving that in them femaleness is a definite Mendelian factor -absent from the male and following the ordinary Mendelian rules. - -Before showing you how the Sweet Pea phenomenon aids in this inquiry I -must tell you of some other experimental results. The first concerns the -common currant moth, _Abraxas grossulariata_. It has a definite pale -variety called _lacticolor_. With these two forms Doncaster has made a -remarkable series of experiments. When he began, _lacticolor_ was only -known as a female form. This was crossed with the _grossulariata_ male -and gave _grossulariata_ only, showing that the male was pure to type. -The hybrids bred together gave _grossulariata_ males and females and -_lacticolor_ females only. But the hybrid males bred to _lacticolor_ -females produced all four combinations, _grossulariata_ males and -females, and _lacticolor_ males and females. When the _lacticolor_ males -were bred to _grossulariata_ females, whether hybrid, or wild from a -district where _lacticolor_ does not exist, the result was that all the -males were _grossulariata_ and all the females _lacticolor_! It is -difficult to follow the course of such an experiment on once hearing and -all I ask you to remember is first that there is a series of matings -giving very curious distributions of the characters of type and variety -among the two sexes. And then, what is perhaps the most singular fact of -all, that the wild typical _grossulariata_ female can when crossed with -the _lacticolor_ male produce all females _lacticolor_. This last fact -can, we know, mean only one thing, namely that these wild females are in -reality hybrids of _lacticolor_; though since the males are pure -_grossulariata_, that fact would in the natural course of things never -be revealed. - -When we encounter such a series of phenomena as this, our business is to -find a means of symbolical expression which will represent all the -factors involved, and show how each behaves in descent. Such a system or -scheme we have at length discovered, and I incline to think that it must -be the true one. If you study this case you will find that there are -nine distinct kinds of matings that can be made between the variety, the -type and the hybrid, and the scheme fits the whole group of results. It -is based on two suppositions: - -1. That the female is cross-bred, or as we call it heterozygous for -femaleness-factor, the male being without that factor. The eggs are thus -each destined from the first to become either males or females, but as -regards sex the spermatozoa are alike in being non-female. - -2. That there is a repulsion between the femaleness-factor and the -_grossulariata_ factor. - -Such a repulsion between two factors we are justified in regarding as -possible because we have had proof of the occurrence of a similar -repulsion in the case of the two factors in the Sweet Pea. - -If the case of this moth stood alone it would be interesting, but its -importance is greatly increased by the fact that we know two cases in -birds which are closely comparable. The simpler case to which alone I -shall refer has been observed in the Canary. Like the Currant moth it -has a kind of albino, called Cinnamon, and males of this variety when -mated with ordinary dark green hen canaries produce dark males and -Cinnamons which are always hens; while the green male and the Cinnamon -hen produce nothing but greens of both sexes. This case, which has been -experimentally studied by Miss Durham, offers a certain complication, -but in its main outlines it is exactly like that of the moth, and the -same interpretation is applicable to both. - -The particular interpretation may be imperfect and even partially -wrong; but that we are at last able to form a working idea of the course -of such phenomena at all is a most encouraging fact. If we are right, as -I am strongly inclined to believe, we get a glimpse of the significance -of the popular idea that in certain respects daughters are apt to -resemble their fathers and sons their mothers; a phenomenon which is -certainly sometimes to be observed. - -There are several collateral indications that we are on the right track -in our theory of the nature of sex. One of these, derived from the -peculiar inheritance of colour-blindness, is especially interesting. -That affection is common in men, rare in women. Men who are colour-blind -can transmit the affection but men who have normal vision cannot. Women -however who are ostensibly normal may have colour-blind sons; and women -who are colour-blind have, so far as we know, no sons who are not -colour-blind[2]. - -Mendelian analysis of these facts shows that colour-blindness is due, -not, as might have been supposed, to the absence of something from the -composition of the body, but to the presence of something which affects -the sight. Just as nicotine-poisoning can paralyse the colour sense, so -may we conceive the development of a secretion in the body which has a -similar action. The comparative exemption of the woman must therefore -mean that there is in her a positive factor which counteracts the -colour-blindness factor, and it is not improbable that the counteracting -element is no other than the femaleness-factor itself[3]. - -I think I have said enough to prove that after all, those curiosities -collected from observation of Sweet Peas and Canaries have no remote -bearing on some very fascinating problems of human life. - -Lastly I suppose it is self-evident that they have a bearing on the -problem of Evolution. The facts of heredity and variation are the -materials out of which all theories of Evolution are constructed. At -last by genetic methods we are beginning to obtain such facts of -unimpeachable quality, and free from the flaws that were inevitable in -older collections. From a survey of these materials we see something of -the changes which will have to be made in the orthodox edifice to admit -of their incorporation, but he must be rash indeed who would now attempt -a comprehensive reconstruction. The results of genetic research are so -bewilderingly novel that we need time and an exhaustive study of their -inter-relations before we can hope to see them in proper value and -perspective. In all the discussions of the stability and fitness of -species who ever contemplated the possibility of a wild species having -one of its sexes permanently hybrid? When I spoke of adventures to be -encountered in genetic research I was thinking of such astonishing -discoveries as that. - -There are others no less disconcerting. Who would have supposed it -possible that the pollen-cells of a plant could be all of one type, and -its egg-cells of two types? Yet Miss Saunders' experiments have provided -definite proof that this is the condition of certain Stocks, of which -the pollen grains all bear doubleness, while the egg-cells are some -singles and some doubles. We cannot think yet of interpreting these -complex phenomena in terms of a common plan. All that we know is that -there is now open for our scrutiny a world of varied, orderly and -specific physiological wonders into which we have as yet only peeped. To -lay down positive propositions as to the origin and inter-relation of -species in general, now, would be a task as fruitless as that of a -chemist must have been who had tried to state the relationship of the -elements before their properties had been investigated. - -For the first time _Variation_ and _Reversion_ have a concrete, palpable -meaning. Hitherto they have stood by in all evolutionary debates, -convenient genii, ready to perform as little or as much as might be -desired by the conjuror. That vaporous stage of their existence is over; -and we see Variation shaping itself as a definite, physiological event, -the addition or omission of one or more definite elements; and Reversion -as that particular addition or subtraction which brings the total of the -elements back to something it had been before in the history of the -race. - -The time for discussion of Evolution as a problem at large is closed. We -face that problem now as one soluble by minute, critical analysis. Lord -Acton in his inaugural lecture said that in the study of history we are -at the beginning of the documentary age. No one will charge me with -disrespect to the great name we commemorate this year, if I apply those -words to the history of Evolution: Darwin, it was, who first showed us -that the species have a history that can be read at all. If in the new -reading of that history, there be found departures from the text laid -down in his first recension, it is not to his fearless spirit that they -will bring dismay. - - -FOOTNOTES: - -[1] The investigation of this remarkable family was made originally by -Cunier. The facts have been reexamined and the pedigree much extended by -Nettleship. The numerical results are somewhat irregular, but it is -especially interesting as being the largest pedigree of human disease or -defect yet made. It contains 2121 persons, extending over ten -generations. Of these persons, 135 are known to have been night-blind. -In no single case was the peculiarity transmitted through an unaffected -member. It should be mentioned that for night-blindness such a system of -descent is peculiar. More usually it follows the scheme described for -colour-blindness. It is not known wherein the peculiarity of this family -consists. - -[2] We have knowledge now of seven colour-blind women, having, in all, -17 sons who are all colour-blind. Most of these cases have been -collected by Mr Nettleship. - -[3] An alternative and perhaps more satisfactory interpretation of the -same facts has been proposed by Doncaster (_Jour. Genetics_ I, Pt 4, p. -377). Until more progress has been made with the analysis of sexual -differentiation it is not possible to decide which of the two -interpretations is correct. The numerical results predicted on both -systems are the same; but by introducing a more complicated though quite -reasonable formula for the representation of the sex-differences -Doncaster's method shows that colour-blindness may be a _recessive_ due -to the absence of a factor which produces normal colour-vision. - - -Cambridge: -PRINTED BY JOHN CLAY, M.A. -AT THE UNIVERSITY PRESS. - - - - - -End of Project Gutenberg's The Methods and Scope of Genetics, by W. 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Bateson, M.A., F.R.S. - </title> - <style type="text/css"> - - p { margin-top: .75em; - text-align: justify; - margin-bottom: .75em; - } - - p.bold {text-align: center; font-weight: bold;} - p.bold2 {text-align: center; font-weight: bold; font-size: 150%;} - - h1,h2,h3,h4,h5,h6 { - text-align: center; /* all headings centered */ - clear: both; - } - h1 span, h2 span { display: block; text-align: center; } - #id1 { font-size: smaller } - - - hr { - width: 33%; - margin-top: 2em; - margin-bottom: 2em; - margin-left: 33.5%; - margin-right: 33.5%; - clear: both; - } - - body{margin-left: 10%; - margin-right: 10%; - } - - .pagenum { /* uncomment the next line for invisible page numbers */ - /* visibility: hidden; */ - position: absolute; - left: 92%; - font-size: smaller; - text-align: right; - text-indent: 0px; - } /* page numbers */ - - .center {text-align: center;} - .smcap {font-variant: small-caps;} - .space-above {margin-top: 3em;} - .right {text-align: right;} - - </style> - </head> -<body> - - -<pre> - -Project Gutenberg's The Methods and Scope of Genetics, by W. Bateson - -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/license - - -Title: The Methods and Scope of Genetics - An inaugural lecture delivered 23 October 1908 - -Author: W. Bateson - -Release Date: June 12, 2016 [EBook #52312] - -Language: English - -Character set encoding: ISO-8859-1 - -*** START OF THIS PROJECT GUTENBERG EBOOK THE METHODS AND SCOPE OF GENETICS *** - - - - -Produced by Chris Curnow, Martin Pettit and the Online -Distributed Proofreading Team at http://www.pgdp.net (This -file was produced from images generously made available -by The Internet Archive) - - - - - - -</pre> - - -<div class="center"><a name="cover.jpg" id="cover.jpg"></a><img src="images/cover.jpg" alt="cover" /></div> - -<hr /> - -<p class="bold2">THE METHODS AND SCOPE<br />OF<br />GENETICS</p> - -<hr /> - -<p class="bold">CAMBRIDGE UNIVERSITY PRESS<br /> -London: FETTER LANE, E.C.<br /> -<span class="smcap">C. F. CLAY, Manager</span></p> - -<div class="center"><img src="images/logo.jpg" alt="logo" /></div> - -<p class="bold">Edinburgh: 100, PRINCES STREET<br /> -Berlin: A. ASHER AND CO.<br /> -Leipzig: F. A. BROCKHAUS<br /> -New York: G. P. PUTNAM'S SONS<br /> -Bombay and Calcutta: <span class="smcap">MACMILLAN AND CO., Ltd.</span></p> - -<p class="bold space-above"><i>All rights reserved</i></p> - -<hr /> - -<h1>THE METHODS AND SCOPE<br />OF<br />GENETICS</h1> - -<p class="bold space-above"><i>AN INAUGURAL LECTURE DELIVERED<br /> -23 OCTOBER 1908</i></p> - -<p class="bold space-above">by<br />W. BATESON, M.A., F.R.S.<br />PROFESSOR OF BIOLOGY IN THE UNIVERSITY OF CAMBRIDGE</p> - -<p class="bold space-above">Cambridge:<br />at the University Press<br />1912</p> - -<hr /> - -<p class="center"><i>First Edition 1908</i><br /><i>Reprinted 1912</i></p> - -<hr /> - -<h2>PREFATORY NOTE</h2> - -<p>The Professorship of Biology was founded in 1908 for a period of five -years partly by the generosity of an anonymous benefactor, and partly by -the University of Cambridge. The object of the endowment was the -promotion of inquiries into the physiology of Heredity and Variation, a -study now spoken of as Genetics.</p> - -<p>It is now recognized that the progress of such inquiries will chiefly be -accomplished by the application of experimental methods, especially -those which Mendel's discovery has suggested. The purpose of this -inaugural lecture is to describe the outlook over this field of research -in a manner intelligible to students of other parts of knowledge.</p> - -<p class="right">W. B.</p> - -<p><i>28 October, 1908</i></p> - - -<hr /> - -<p><span class='pagenum'><a name="Page_1" id="Page_1">[Pg 1]</a></span></p> - -<h2>THE METHODS AND SCOPE OF<br />GENETICS</h2> - -<p>The opportunity of addressing fellow-students pursuing lines of inquiry -other than his own falls seldom to a scientific man. One of these rare -opportunities is offered by the constitution of the Professorship to -which I have had the honour to be called. That Professorship, though -bearing the comprehensive title "of Biology," is founded with the -understanding that the holder shall apply himself to a particular class -of physiological problems, the study of which is denoted by the term -Genetics. The term is new; and though the problems are among the oldest -which have vexed the human mind, the modes by which they may be -successfully attacked<span class='pagenum'><a name="Page_2" id="Page_2">[Pg 2]</a></span> are also of modern invention. There is therefore -a certain fitness in the employment of this occasion for the deliverance -of a discourse explaining something of the aims of Genetics and of the -methods by which we trust they may be reached.</p> - -<p>You will be aware that the claims put forward in the name of Genetics -are high, but I trust to be able to show you that they are not high -without reason. It is the ambition of every one who in youth devotes -himself to the search for natural truth, that his work may be found -somewhere in the main stream of progress. So long only as he keeps -something of the limitless hope with which his voyage of discovery -began, will his courage and his spirit last. The moment we most dread is -one in which it may appear that, after all, our effort has been spent in -exploring some petty tributary, or worse, a backwater of the great -current. It is because<span class='pagenum'><a name="Page_3" id="Page_3">[Pg 3]</a></span> Genetic research is still pushing forward in the -central undifferentiated trunk of biological science that we confess no -guilt of presumption in declaring boldly that whatever difficulty may be -in store for those who cast in their lot with us, they need fear no -disillusionment or misgiving that their labour has been wasted on a -paltry quest.</p> - -<p>In research, as in all business of exploration, the stirring times come -when a fresh region is suddenly unlocked by the discovery of a new key. -Then conquest is easy and there are prizes for all. We are happy in that -during our own time not a few such territories have been revealed to the -vision of mankind. I do not dare to suggest that in magnitude or -splendour the field of Genetics may be compared with that now being -disclosed to the physicist or the astronomer; for the glory of the -celestial is one and the glory of the terrestrial is another. But I -will<span class='pagenum'><a name="Page_4" id="Page_4">[Pg 4]</a></span> say that for once to the man of ordinary power who cannot venture -into those heights beyond, Mendel's clue has shown the way into a realm -of nature which for surprising novelty and adventure is hardly to be -excelled.</p> - -<p>It is no hyperbolical figure that I use when I speak of Mendelian -discovery leading us into a new world, the very existence of which was -unsuspected before.</p> - -<p>The road thither is simple and easy to follow. We start from a common -fact, familiar to everyone, that all the ordinary animals and plants -began their individual life by the union of two cells, the one male, the -other female. Those cells are known as germ-cells or <i>gametes</i>, that is -to say, "marrying" cells.</p> - -<p>Now obviously the diversity of form which is characteristic of the -animal and plant world must be somehow represented in the gametes, since -it is they which bring into each organism<span class='pagenum'><a name="Page_5" id="Page_5">[Pg 5]</a></span> all that it contains. I am -aware that there is interplay between the organism and the circumstances -in which it grows up, and that opportunity given may bring out a -potentiality which without that opportunity must have lain dormant. But -while noting parenthetically that this question of opportunity has an -importance, which some day it may be convenient to estimate, the one -certain fact is that all the powers, physical and mental that a living -creature possesses were contributed by one or by both of the two -germ-cells which united in fertilisation to give it existence. The fact -that <i>two</i> cells are concerned in the production of all the ordinary -forms of life was discovered a long while ago, and has been part of the -common stock of elementary knowledge of all educated persons for about -half a century. The full consequences of this double nature seem -nevertheless to have struck nobody before Mendel. Simple though<span class='pagenum'><a name="Page_6" id="Page_6">[Pg 6]</a></span> the -fact is, I have noticed that to many it is difficult to assimilate as a -working idea. We are accustomed to think of a man, a butterfly, or an -apple tree as each <i>one</i> thing. In order to understand the significance -of Mendelism we must get thoroughly familiar with the fact that they are -each <i>two</i> things, double throughout every part of their composition. -There is perhaps no better exercise as a preparation for genetic -research than to examine the people one meets in daily life and to try -in a rough way to analyse them into the two assemblages of characters -which are united in them. That we are assemblages or medleys of our -parental characteristics is obvious. We all know that a man may have his -father's hair, his mother's colour, his father's voice, his mother's -insensibility to music, and so on, but that is not enough.</p> - -<p>Such an analysis is true, inasmuch as the various characters <i>are</i> -transmitted<span class='pagenum'><a name="Page_7" id="Page_7">[Pg 7]</a></span> independently, but it misses the essential point. For in -each of these respects the individual is double; and so to get a true -picture of the composition of the individual we have to think how <i>each</i> -of the two original gametes was provided in the matter of height, hair, -colour, mathematical ability, nail-shape, and the other features that go -to make the man we know. The contribution of each gamete in each respect -has thus to be separately brought to account. If we could make a list of -all the ingredients that go to form a man and could set out how he is -constituted in respect of each of them, it would not suffice to give one -column of values for these ingredients, but we must rule two columns, -one for the ovum and one for the spermatozoon, which united in -fertilisation to form that man, and in each column we must represent how -that gamete was supplied in respect of each of the ingredients in our -list. When the problem of<span class='pagenum'><a name="Page_8" id="Page_8">[Pg 8]</a></span> heredity is thus represented we can hardly -avoid discovering, by mere inspection, one of the chief conclusions to -which genetic research has led. For it is obvious that the contributions -of the male and female gametes may in respect of any of the ingredients -be either the same, or different. In any case in which the contribution -made by the two cells is the same, the resulting organism—in our -example the man—is, as we call it, <i>pure-bred</i> for that ingredient, and -in all respects in which the contribution from the two sides of the -parentage is dissimilar the resulting organism is <i>cross-bred</i>.</p> - -<p>To give an intelligible account of the next step in the analysis without -having recourse to precise and technical language is not very easy.</p> - -<p>We have got to the point of view from which we see the individual made -up of a large number of distinct ingredients,<span class='pagenum'><a name="Page_9" id="Page_9">[Pg 9]</a></span> contributed from two -sources, and in respect of any of them he may have received two similar -portions or two dissimilar portions. We shall not go far wrong if we -extend and elaborate our illustration thus. Let us imagine the contents -of a gamete as a fluid made by taking a drop from each of a definite -number of bottles in a chest, containing tinctures of the several -ingredients. There is one such chest from which the male gamete is to be -made up, and a similar chest containing a corresponding set of bottles -out of which the components of the female gamete are to be taken. But in -either chest one or more of the bottles may be empty; then nothing goes -in to represent that ingredient from that chest, and if corresponding -bottles are empty in both chests, then the individual made on -fertilisation by mixing the two collections of drops together does not -contain the missing ingredient at all. It follows<span class='pagenum'><a name="Page_10" id="Page_10">[Pg 10]</a></span> therefore that an -individual may thus be "pure-bred," namely alike on both sides of his -composition as regards each ingredient in one of two ways, either by -having received the ingredient from the male chest and from the female, -or in having received it from neither. Conversely in respect of any -ingredient he may be "cross-bred," receiving the presence of it from one -gamete and the absence of it from the other.</p> - -<p>The second conception with which we have now to become thoroughly -familiar is that of the individual as composed of what we call presences -and absences of all the possible ingredients. It is the basis of all -progress in genetic analysis. Let me give you two illustrations. A blue -eye is due to the absence of a factor which forms pigment on the front -of the iris. Two blue-eyed parents therefore, as Hurst has proved, do -not have dark-eyed children. The dark eye<span class='pagenum'><a name="Page_11" id="Page_11">[Pg 11]</a></span> is due to either a single or -double dose of the factor missing from the blue eye. So dark-eyed -persons may have families all dark-eyed, or families composed of a -mixture of dark and light-eyed children in certain proportions which on -the average are definite.</p> - -<p>Two plants of <i>Oenothera</i> which I exhibit illustrate the same thing. One -of them is the ordinary <i>Lamarckiana</i>. I bend its stem. It will not -break, or only breaks with difficulty on account of the tough fibres it -contains. The stem of the other, one of de Vries' famous mutations, -snaps at once like short pastry, because it does not contain the factor -for the formation of the fibres. Such plants may be sister-plants -produced by the self-fertilisation of one parent, but they are distinct -in their composition and properties—and this distinction turns on the -presence or absence of elements which are treated as definite entities -when the germ-cells are<span class='pagenum'><a name="Page_12" id="Page_12">[Pg 12]</a></span> formed. When we speak of such qualities as the -formation of pigment in an eye, or the development of fibres in a stem, -as due to transmitted elements or factors, you will perhaps ask if we -have formed any notion as to the actual nature of those factors. For my -own part as regards that ulterior question I confess to a disposition to -hold my fancy on a tight rein. It cannot be very long before we shall -<i>know</i> what some of the factors are, and we may leave guessing till -then. Meanwhile however there is no harm in admitting that several of -them behave much as if they were ferments, and others as if they -constructed the substances on which the ferments act. But we must not -suppose for a moment that it is the ferment, or the objective substance, -which is transmitted. The thing transmitted can only be the power or -faculty to produce the ferment or the objective substance.</p> - -<p><span class='pagenum'><a name="Page_13" id="Page_13">[Pg 13]</a></span></p><p>So far we have been considering the synthesis of the individual from -ingredients brought into him by the two gametes. In the next step of our -consideration we reverse the process, and examine how the ingredients of -which he was originally compounded are distributed among the gametes -that are eventually budded off from him.</p> - -<p>Take first the case of the components in respect of which he is -pure-bred. Expectation would naturally suggest that all the germ-cells -formed from him would be alike in respect of those ingredients, and -observation shows, except in the rare cases of originating variations, -the causation of which is still obscure, that this expectation is -correct.</p> - -<p>Hitherto though without experimental evidence no one could have been -certain that the facts were as I have described them, yet there is -nothing altogether contrary to<span class='pagenum'><a name="Page_14" id="Page_14">[Pg 14]</a></span> common expectation. But when we proceed -to ask how the germ-cells will be constituted in the case of an -individual who is cross-bred in some respect, containing that is to say, -an ingredient from the one side of his parentage and not from the other, -the answer is entirely contrary to all the preconceptions which either -science or common sense had formed about heredity. For we find definite -experimental proof in nearly all the cases which have been examined, -that the germ-cells formed by such individuals do either contain or not -contain a representation of the ingredient, just as the original gametes -did or did not contain it.</p> - -<p>If <i>both</i> parent-gametes brought a certain quality in, then all the -daughter gametes have it; if neither brought it in, then none of the -daughter gametes have it. If it came in from one side and not from the -other, then on an average in half the resulting gametes it will be -present and from half it will be<span class='pagenum'><a name="Page_15" id="Page_15">[Pg 15]</a></span> absent. This last phenomenon, which is -called segregation, constitutes the essence of Mendel's discovery.</p> - -<p>So recurring to the simile of the man as made by the mixing of -tinctures, the process of redistribution of his characters among the -germ-cells may be represented as a sorting back of the tinctures again -into a double row of bottles, a pair corresponding to each ingredient; -and each of the germ-cells as then made of a drop from one or other -bottle of each pair: and in our model we may represent the phenomenon of -segregation in a crude way by supposing that the bottles having no -tincture in them, instead of being empty contained an inoperative fluid, -say water, with which the tincture would not mix. When the new -germ-cells are formed, the two fluids instead of diluting each other -simply separate again. It is this fact which entitles us to speak of the -purity of germ-cells. They<span class='pagenum'><a name="Page_16" id="Page_16">[Pg 16]</a></span> are pure in the possession of an ingredient, -or in not possessing it; and the ingredients, or factors, as we -generally call them, are units because they are so treated in the -process of formation of the new gametes and because they come out of the -process of segregation in the same condition as they went in at -fertilisation.</p> - -<p>As a consequence of these facts it follows that however complex may be -the origin of two given parents the composition of the offspring they -can produce is limited. There is only a limited number of types to be -made by the possible recombinations of the parental ingredients, and the -relative numbers in which each type will be represented are often -predicable by very simple arithmetical rules.</p> - -<p>For example, if neither parent possesses a certain factor at all, then -none of the offspring will have it. If either parent has two doses of -the factor then all the children will<span class='pagenum'><a name="Page_17" id="Page_17">[Pg 17]</a></span> have it; and if either parent has -one dose of the factor and the other has none, then on an average half -the family will have it, and half be without it.</p> - -<p>To know whether the parent possesses the factor or not may be difficult -for reasons which will presently appear, but often it is quite easy and -can be told at once, for there are many factors which cannot be present -in the individual without manifesting their presence. I may illustrate -the descent of such a factor by the case of a family possessing a -peculiar form of night-blindness. The affected individuals marrying with -those unaffected have a mixture of affected and unaffected children, but -their unaffected children not having the responsible ingredient cannot -pass it on<a name="FNanchor_1_1" id="FNanchor_1_1"></a><a href="#Footnote_1_1" class="fnanchor">[1]</a>.</p> - -<p><span class='pagenum'><a name="Page_18" id="Page_18">[Pg 18]</a></span></p><p>In such an observation two things are strikingly exemplified, (1) the -fact of the permanence of the unit, and (2) the fact that a <i>mixture</i> of -types in the family means that one or other parent is cross-bred in some -respect, and is giving off gametes of more than one type.</p> - -<p>The problem of heredity is thus a problem primarily analytical. We have -to detect and enumerate the factors out of which the bodies of animals -and plants are built up, and the laws of their distribution among the -germ-cells. All the processes of which I have spoken are accomplished by -means of cell-divisions, and in the one cell-union which occurs in -fertilisation. If we could watch<span class='pagenum'><a name="Page_19" id="Page_19">[Pg 19]</a></span> the factors segregating from each -other in cell-division, or even if by microscopic examination we could -recognize this multitudinous diversity of composition that must -certainly exist among the germ-cells of all ordinary individuals, the -work of genetics would be much simpler than it is.</p> - -<p>But so far no such direct method of observation has been discovered. In -default we are obliged to examine the constitution of the germ-cells by -experimental breeding, so contrived that each mating shall test the -composition of an individual in one or more chosen respects, and, so to -speak, sample its germ-cells by counting the number of each kind of -offspring which it can produce. But cumbersome as this method must -necessarily be, it enables us to put questions to Nature which never -have been put before. She, it has been said, is an unwilling witness. -Our questions must be shaped in such a way that<span class='pagenum'><a name="Page_20" id="Page_20">[Pg 20]</a></span> the only possible -answer is a direct "Yes" or a direct "No." By putting such questions we -have received some astonishing answers which go far below the surface. -Amazing though they be, they are nevertheless true; for though our -witness may prevaricate, she cannot lie. Piecing these answers together, -getting one hint from this experiment, and another from that, we begin -little by little to reconstruct what is going on in that hidden world of -gametes. As we proceed, like our brethren in other sciences, we -sometimes receive answers which seem inconsistent or even contradictory. -But by degrees a sufficient body of evidence can be attained to show -what is the rule and what the exception. My purpose today must be to -speak rather of the regular than of the irregular.</p> - -<p>One clear exception I may mention. Castle finds that in a cross between -the long-eared lop-rabbit and a short-eared<span class='pagenum'><a name="Page_21" id="Page_21">[Pg 21]</a></span> breed, ears of intermediate -length are produced: and that these intermediates breed approximately -true.</p> - -<p>Exceptions in general must be discussed elsewhere. Nevertheless if I may -throw out a word of counsel to beginners, it is: Treasure your -exceptions! When there are none, the work gets so dull that no one cares -to carry it further. Keep them always uncovered and in sight. Exceptions -are like the rough brickwork of a growing building which tells that -there is more to come and shows where the next construction is to be.</p> - -<p>You will readily understand that the presentation here given of the -phenomena is only the barest possible outline. Some of the details we -may now fill in. For example, I have spoken of the characters of the -organism, its colour, shape, and the like, as if they were due each to -one ingredient or factor. Some of them are no doubt correctly so<span class='pagenum'><a name="Page_22" id="Page_22">[Pg 22]</a></span> -represented; but already we know numerous bodily features which need the -concurrence of several factors to produce them. Nevertheless though the -character only appears when all the complementary ingredients are -together present, each of these severally and independently follows, as -regards its transmission, the simple rules I have described.</p> - -<p>This complementary action may be illustrated by some curious results -that Mr Punnett and I have encountered when experimenting with the -height of Sweet Peas. There are two dwarf varieties, one the prostrate -"Cupid," the other the half-dwarf or "Bush" Sweet Peas. Crossed together -they give a cross-bred of full height. There is thus some element in the -Cupid which when it meets the complementary element from the Bush, -produces the characteristic length of the ordinary Sweet Pea. We may -note in passing that such a fact demonstrates<span class='pagenum'><a name="Page_23" id="Page_23">[Pg 23]</a></span> at once the nature of -Variation and Reversion. The Reversion occurs because the two factors -that made the <i>height</i> of the old Sweet Pea again come together after -being parted: and the Variations by which each of the dwarfs came into -existence must have taken place by the dropping out of one of these -elements or of the other.</p> - -<p>Conversely there are factors which by their presence can prevent or -inhibit the development and appearance of others present and -unperceived.</p> - -<p>For example, all the factors for pigmentation may be present in a plant -or an animal; but in addition there may be another factor present which -keeps the individual white, or nearly so.</p> - -<p>There are cases in which the action of the factors is superposed one on -top of the other, and not until each factor is removed in turn can the -effects of the underlying<span class='pagenum'><a name="Page_24" id="Page_24">[Pg 24]</a></span> factors be perceived. So in the mouse if no -other colour-factor is present, the fur is chocolate. If the next factor -in the series be there, it is black. If still another factor be added, -it has the brownish grey of the common wild mouse. Conversely, by the -variation which dropped out the top factor, a black mouse came into -existence. By the loss of the black factor, the chocolate mouse was -created, and for aught we can tell there may be still more possibilities -hidden beneath.</p> - -<p>In the disentanglement of the properties and interactions of these -elementary factors, the science we must call to our aid is Physiological -Chemistry. The relations of Genetics with the other branches of biology -are close. Such work can only be conducted by those who have the good -fortune to be able to count upon continual help and advice from -specialists in the various branches of Zoology, Physiology, and Botany. -Often we have<span class='pagenum'><a name="Page_25" id="Page_25">[Pg 25]</a></span> questions with which only a cytologist can deal, and -often it is the experience of a systematist we must invoke. The school -of Genetics in Cambridge starts under happy auspices in that we are -surrounded by colleagues qualified, and as we have often found, willing -to give us such aid unstinted. But with chemical physiology, we stand in -an even closer relation; and from the little I have dared to say -respecting the action and interaction of factors, it is evident that for -their disentanglement there must one day be an intimate and enduring -partnership arranged with the physiological chemists.</p> - -<p>Now, as the whole of the elaborate process by which the various elements -are apportioned among the gametes must be got through in a few -cell-divisions at most, and perhaps in one division only, it is not -surprising that there is sometimes an interaction between factors that -have quite distinct rôles<span class='pagenum'><a name="Page_26" id="Page_26">[Pg 26]</a></span> to perform. These interactions are probably -of several kinds. One, which I shall illustrate presently, is probably -to be represented as a repulsion between two factors. As a consequence -of its operations when the various factors are sorted out into the -gametes, if the individual be cross-bred in respect of the <i>two</i> -repelling factors, having received so to speak only a single dose of -each, then the gametes are made up in such a way that each takes one or -other of the two repelling factors, not both.</p> - -<p>Mutual repulsions of this kind probably play a significant part in the -phenomena of heredity. A single concrete case which Mr Punnett and I -have been investigating for some years will illustrate several of these -principles. We crossed together a pure white Sweet Pea having an erect -standard, with another pure white Sweet Pea having a hooded standard. -The result is, as you see,<span class='pagenum'><a name="Page_27" id="Page_27">[Pg 27]</a></span> a purple flower with an erect standard. The -colour comes from the concurrence of complementary elements. A dose of a -certain ingredient from one parent meets a dose of another ingredient -from the other parent and the two make pigment in the flower. From other -experiments we know that the <i>purple</i> colour of the pigment is due to a -dose of a third ingredient brought in from the hooded parent; and that -in the absence of that blue factor, as we may call it, the flower would -be red. The standard is erect because it contains a dose of the -erectness-factor from the erect parent, and the hooded parent can -readily be proved to owe its peculiar shape to the absence of that -element.</p> - -<p>Our purple plant is thus cross-bred for four factors, containing only -one dose of each.</p> - -<p>We let it fertilise itself, and its offspring<span class='pagenum'><a name="Page_28" id="Page_28">[Pg 28]</a></span> show all the possible -combinations of the four different factors and their absences which the -genetic constitution of the plant can make.</p> - -<p>Note that one of the combinations we expect to find is missing. There -are white erect and white hooded—white because they are lacking one or -other of the complementary ingredients necessary to the production of -pigment. There are purple erect and purple hooded, of which the purple -erect must perforce contain all the four factors, and the purple hooded -must similarly contain all of them except that for erectness. But when -we turn to the red class we are surprised to find that they are all -erect, none hooded. One of the possible combinations is missing. If you -examine this series of facts you will find there is only one possible -interpretation: namely that the ingredient which turns the flower -purple—alkalinity,<span class='pagenum'><a name="Page_29" id="Page_29">[Pg 29]</a></span> perhaps we may call it—never goes into the same -germ-cell as the ingredient which makes the standard erect. There are -plenty of ways of testing the truth of this interpretation. For example, -it follows that the purple erects from such a family will in perpetuity -have offspring 1 purple hooded: 2 purple erect: 1 red erect; also that -all the white hooded crossed with pure reds will give purples, and so -on. These experiments have been made and the result has in each case -been conformable to expectation.</p> - -<p>Between these two factors, the purpleness and the erectness of standard, -some antagonism or repulsion must exist. In some way therefore the -chemical and the geometrical phenomena of heredity must be -inter-related.</p> - -<p>Some one will say perhaps this is all very well as a scientific -curiosity, but it has nothing to do with real life. The right answer to -such criticism is of course the lofty one that science<span class='pagenum'><a name="Page_30" id="Page_30">[Pg 30]</a></span> and its -applications are distinct: that the investigator fixes his gaze solely -on the search for truth and that his attention must not be distracted by -trivialities of application. But while we make this answer and at least -try to work in the spirit it proclaims, we know in our hearts that it is -a counsel of perfection. I suspect that even the astronomer who at his -spectroscope is analysing the composition of Vega or Capella has still -an eye sometimes free for the affairs of this planet, and at least the -fact that his discoveries may throw light on our destinies does not -diminish his zeal in their pursuit. And surely to the study of Heredity, -preeminently among all the sciences, we are looking for light on human -destiny. To pretend otherwise would be mere hypocrisy. So while -reserving the higher line of defence I will reply that again and again -in our experimental work we come very near indeed to human affairs. -Sometimes this is obvious<span class='pagenum'><a name="Page_31" id="Page_31">[Pg 31]</a></span> enough. No practical dog-breeder or seeds-man -can see the results of Mendelian recombination without perceiving that -here is a bit of knowledge he can immediately apply. No sociologist can -examine the pedigrees illustrating the simple descent of a deformity or -a congenital disease, and not see that the new knowledge gives a solid -basis for practical action by which the composition of a race could be -modified if society so chose. More than this: we know for certain in one -case, from the work of Professor Biffen, that the power to resist a -disease caused by the invasion of a pathogenic organism, wheat-rust, is -due to the absence of one of the simple factors or ingredients of which -I have spoken, and what we know to be true in that one case we are -beginning to suspect to be true of resistance to certain other diseases. -No pathologist can see such an experiment as this of Professor Biffen's -without realizing<span class='pagenum'><a name="Page_32" id="Page_32">[Pg 32]</a></span> that here is a contribution of the first importance -to the physiology of disease.</p> - -<p>There is no lack of utility and direct application in the study of -Genetics. I have alluded to some strictly practical results. If we want -to raise mangels that will not run to seed, or to breed a cow that will -give more milk in less time, or milk with more butter and less water, we -can turn to Genetics with every hope that something can be done in these -laudable directions. But here I would plead what I cannot but regard as -a higher usefulness in our work. Genetic inquiry aims at providing -knowledge that may bring, and I think will bring, certainty into a -region of human affairs and concepts which might have been supposed -reserved for ages to be the domain of the visionary. We have long known -that it was believed by some that our powers and conduct were dependent -on our physical composition, and that other schools<span class='pagenum'><a name="Page_33" id="Page_33">[Pg 33]</a></span> have maintained -that nurture not nature, to use Galton's antithesis, has a -preponderating influence on our careers; but so soon as it becomes -common knowledge—not a philosophical speculation, but a certainty—that -liability to a disease, or the power of resisting its attack, addiction -to a particular vice, or to superstition, is due to the presence or -absence of a specific ingredient; and finally that these characteristics -are transmitted to the offspring according to definite, predicable -rules, then man's views of his own nature, his conceptions of justice, -in short his whole outlook on the world, must be profoundly changed. Yet -as regards the more tangible of these physical and mental -characteristics there can be little doubt that before many years have -passed the laws of their transmission will be expressible in simple -formulae.</p> - -<p>The blundering cruelty we call criminal justice will stand forth -divested of natural<span class='pagenum'><a name="Page_34" id="Page_34">[Pg 34]</a></span> sanction, a relic of the ferocious inventions of -the savage. Well may such justice be portrayed as blind. Who shall say -whether it is crime or punishment which has wrought the greater -suffering in the world? We may live to know that to the keen satirical -vision of Sam Butler on the pleasant mountains of Erewhon there was -revealed a dispensation, not kinder only, but wiser than the terrific -code which Moses delivered from the flames of Sinai.</p> - -<p>If there are societies which refuse to apply the new knowledge, the -fault will not lie with Genetics. I think it needs but little -observation of the newer civilisations to foresee that <i>they</i> will apply -every scrap of scientific knowledge which can help them, or seems to -help them in the struggle, and I am good enough Selectionist to know -that in that day the fate of the recalcitrant communities is sealed.</p> - -<p>The thrill of discovery is not dulled<span class='pagenum'><a name="Page_35" id="Page_35">[Pg 35]</a></span> by a suspicion that the discovery -can be applied. No harm is done to the investigator if he can resist the -temptation to deviate from his aim. With rarest exceptions the -discoveries which have formed the basis of physical progress have been -made without any thought but for the gratification of curiosity. Of this -there can be few examples more conspicuous than that which Mendel's work -presents. Untroubled by any itch to make potatoes larger or bread -cheaper, he set himself in the quiet of a cloister garden to find out -the laws of hybridity, and so struck a mine of truth, inexhaustible in -brilliancy and profit.</p> - -<p>I will now suggest to you that it is by no means unlikely that even in -an inquiry so remote as that which I just described in the case of the -Sweet Pea, we may have the clue to a mystery which concerns us all in -the closest possible way. I mean the problem of the physiological nature -of Sex. In speaking<span class='pagenum'><a name="Page_36" id="Page_36">[Pg 36]</a></span> of the interpretation of sexual difference -suggested by our experimental work as of some practical moment, I do not -imply that as in the other instances I have given, the knowledge is -likely to be of immediate use to our species; but only that if true it -makes a contribution to the stock of human ideas which no one can regard -as insignificant.</p> - -<p>In the light of Mendelian knowledge, when a family consists of more than -one type the fact means that the germ-cells of one or other parent must -certainly be of more than one kind. In the case of sex the members of -the family are thus of two kinds, and the presumption is overwhelming -that this distinction is due to a difference among the germ-cells. Next, -since for all practical purposes the numbers of the two sexes produced -are approximately equal, sex exhibits the special case in which a family -consists of two types represented in equal numbers, half being<span class='pagenum'><a name="Page_37" id="Page_37">[Pg 37]</a></span> male, -half female. But I called your attention to the fact that equality of -types results when <i>one</i> parent was cross-bred in the character -concerned, having received one dose only of the factor on which it -depends. So we may feel fairly sure that the distinction between the -sexes depends on the presence in one or other of them of an unpaired -factor. This conclusion appears to me to follow so immediately on all -that we have learnt of genetic physiology that with every confidence we -may accept it as representing the actual fact.</p> - -<p>The question which of the two sexes contains the unpaired factor is less -easy to answer, but there are several converging lines of evidence which -point to the deduction that in Vertebrates at least, and in some other -types, it is the female, and I feel little doubt that we shall succeed -in proving that in them femaleness is a definite Mendelian factor<span class='pagenum'><a name="Page_38" id="Page_38">[Pg 38]</a></span> -absent from the male and following the ordinary Mendelian rules.</p> - -<p>Before showing you how the Sweet Pea phenomenon aids in this inquiry I -must tell you of some other experimental results. The first concerns the -common currant moth, <i>Abraxas grossulariata</i>. It has a definite pale -variety called <i>lacticolor</i>. With these two forms Doncaster has made a -remarkable series of experiments. When he began, <i>lacticolor</i> was only -known as a female form. This was crossed with the <i>grossulariata</i> male -and gave <i>grossulariata</i> only, showing that the male was pure to type. -The hybrids bred together gave <i>grossulariata</i> males and females and -<i>lacticolor</i> females only. But the hybrid males bred to <i>lacticolor</i> -females produced all four combinations, <i>grossulariata</i> males and -females, and <i>lacticolor</i> males and females. When the <i>lacticolor</i> males -were bred to <i>grossulariata</i> females, whether hybrid, or wild<span class='pagenum'><a name="Page_39" id="Page_39">[Pg 39]</a></span> from a -district where <i>lacticolor</i> does not exist, the result was that all the -males were <i>grossulariata</i> and all the females <i>lacticolor</i>! It is -difficult to follow the course of such an experiment on once hearing and -all I ask you to remember is first that there is a series of matings -giving very curious distributions of the characters of type and variety -among the two sexes. And then, what is perhaps the most singular fact of -all, that the wild typical <i>grossulariata</i> female can when crossed with -the <i>lacticolor</i> male produce all females <i>lacticolor</i>. This last fact -can, we know, mean only one thing, namely that these wild females are in -reality hybrids of <i>lacticolor</i>; though since the males are pure -<i>grossulariata</i>, that fact would in the natural course of things never -be revealed.</p> - -<p>When we encounter such a series of phenomena as this, our business is to -find a means of symbolical expression which will<span class='pagenum'><a name="Page_40" id="Page_40">[Pg 40]</a></span> represent all the -factors involved, and show how each behaves in descent. Such a system or -scheme we have at length discovered, and I incline to think that it must -be the true one. If you study this case you will find that there are -nine distinct kinds of matings that can be made between the variety, the -type and the hybrid, and the scheme fits the whole group of results. It -is based on two suppositions:</p> - -<p>1. That the female is cross-bred, or as we call it heterozygous for -femaleness-factor, the male being without that factor. The eggs are thus -each destined from the first to become either males or females, but as -regards sex the spermatozoa are alike in being non-female.</p> - -<p>2. That there is a repulsion between the femaleness-factor and the -<i>grossulariata</i> factor.</p> - -<p>Such a repulsion between two factors we are justified in regarding as -possible because<span class='pagenum'><a name="Page_41" id="Page_41">[Pg 41]</a></span> we have had proof of the occurrence of a similar -repulsion in the case of the two factors in the Sweet Pea.</p> - -<p>If the case of this moth stood alone it would be interesting, but its -importance is greatly increased by the fact that we know two cases in -birds which are closely comparable. The simpler case to which alone I -shall refer has been observed in the Canary. Like the Currant moth it -has a kind of albino, called Cinnamon, and males of this variety when -mated with ordinary dark green hen canaries produce dark males and -Cinnamons which are always hens; while the green male and the Cinnamon -hen produce nothing but greens of both sexes. This case, which has been -experimentally studied by Miss Durham, offers a certain complication, -but in its main outlines it is exactly like that of the moth, and the -same interpretation is applicable to both.</p> - -<p><span class='pagenum'><a name="Page_42" id="Page_42">[Pg 42]</a></span></p><p>The particular interpretation may be imperfect and even partially -wrong; but that we are at last able to form a working idea of the course -of such phenomena at all is a most encouraging fact. If we are right, as -I am strongly inclined to believe, we get a glimpse of the significance -of the popular idea that in certain respects daughters are apt to -resemble their fathers and sons their mothers; a phenomenon which is -certainly sometimes to be observed.</p> - -<p>There are several collateral indications that we are on the right track -in our theory of the nature of sex. One of these, derived from the -peculiar inheritance of colour-blindness, is especially interesting. -That affection is common in men, rare in women. Men who are colour-blind -can transmit the affection but men who have normal vision cannot. Women -however who are ostensibly normal may have colour-blind sons; and<span class='pagenum'><a name="Page_43" id="Page_43">[Pg 43]</a></span> women -who are colour-blind have, so far as we know, no sons who are not -colour-blind<a name="FNanchor_2_2" id="FNanchor_2_2"></a><a href="#Footnote_2_2" class="fnanchor">[2]</a>.</p> - -<p>Mendelian analysis of these facts shows that colour-blindness is due, -not, as might have been supposed, to the absence of something from the -composition of the body, but to the presence of something which affects -the sight. Just as nicotine-poisoning can paralyse the colour sense, so -may we conceive the development of a secretion in the body which has a -similar action. The comparative exemption of the woman must therefore -mean that there is in her a positive factor which counteracts the -colour-blindness factor, and it is not improbable that the counteracting -element is no other than the femaleness-factor itself<a name="FNanchor_3_3" id="FNanchor_3_3"></a><a href="#Footnote_3_3" class="fnanchor">[3]</a>.</p> - -<p><span class='pagenum'><a name="Page_44" id="Page_44">[Pg 44]</a></span></p><p>I think I have said enough to prove that after all, those curiosities -collected from observation of Sweet Peas and Canaries have no remote -bearing on some very fascinating problems of human life.</p> - -<p>Lastly I suppose it is self-evident that they have a bearing on the -problem of Evolution. The facts of heredity and variation are the -materials out of which all theories of Evolution are constructed. At -last by genetic methods we are beginning to obtain such facts of -unimpeachable quality, and free from the flaws that were inevitable in -older collections. From a survey of these materials we see something of -the changes which will have to be made in the orthodox edifice to<span class='pagenum'><a name="Page_45" id="Page_45">[Pg 45]</a></span> admit -of their incorporation, but he must be rash indeed who would now attempt -a comprehensive reconstruction. The results of genetic research are so -bewilderingly novel that we need time and an exhaustive study of their -inter-relations before we can hope to see them in proper value and -perspective. In all the discussions of the stability and fitness of -species who ever contemplated the possibility of a wild species having -one of its sexes permanently hybrid? When I spoke of adventures to be -encountered in genetic research I was thinking of such astonishing -discoveries as that.</p> - -<p>There are others no less disconcerting. Who would have supposed it -possible that the pollen-cells of a plant could be all of one type, and -its egg-cells of two types? Yet Miss Saunders' experiments have provided -definite proof that this is the condition of certain Stocks, of which -the pollen grains<span class='pagenum'><a name="Page_46" id="Page_46">[Pg 46]</a></span> all bear doubleness, while the egg-cells are some -singles and some doubles. We cannot think yet of interpreting these -complex phenomena in terms of a common plan. All that we know is that -there is now open for our scrutiny a world of varied, orderly and -specific physiological wonders into which we have as yet only peeped. To -lay down positive propositions as to the origin and inter-relation of -species in general, now, would be a task as fruitless as that of a -chemist must have been who had tried to state the relationship of the -elements before their properties had been investigated.</p> - -<p>For the first time <i>Variation</i> and <i>Reversion</i> have a concrete, palpable -meaning. Hitherto they have stood by in all evolutionary debates, -convenient genii, ready to perform as little or as much as might be -desired by the conjuror. That vaporous stage of their existence is over; -and we see Variation shaping itself as<span class='pagenum'><a name="Page_47" id="Page_47">[Pg 47]</a></span> a definite, physiological event, -the addition or omission of one or more definite elements; and Reversion -as that particular addition or subtraction which brings the total of the -elements back to something it had been before in the history of the -race.</p> - -<p>The time for discussion of Evolution as a problem at large is closed. We -face that problem now as one soluble by minute, critical analysis. Lord -Acton in his inaugural lecture said that in the study of history we are -at the beginning of the documentary age. No one will charge me with -disrespect to the great name we commemorate this year, if I apply those -words to the history of Evolution: Darwin, it was, who first showed us -that the species have a history that can be read at all. If in the new -reading of that history, there be found departures from the text laid -down in his first recension, it is not to his fearless spirit that they -will bring dismay.</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> The investigation of this remarkable family was made -originally by Cunier. The facts have been reexamined and the pedigree -much extended by Nettleship. The numerical results are somewhat -irregular, but it is especially interesting as being the largest -pedigree of human disease or defect yet made. It contains 2121 persons, -extending over ten generations. Of these persons, 135 are known to have -been night-blind. In no single case was the peculiarity transmitted -through an unaffected member. It should be mentioned that for -night-blindness such a system of descent is peculiar. More usually it -follows the scheme described for colour-blindness. It is not known -wherein the peculiarity of this family consists.</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> We have knowledge now of seven colour-blind women, having, -in all, 17 sons who are all colour-blind. Most of these cases have been -collected by Mr Nettleship.</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> An alternative and perhaps more satisfactory interpretation -of the same facts has been proposed by Doncaster (<i>Jour. Genetics</i> I, Pt -4, p. 377). Until more progress has been made with the analysis of -sexual differentiation it is not possible to decide which of the two -interpretations is correct. The numerical results predicted on both -systems are the same; but by introducing a more complicated though quite -reasonable formula for the representation of the sex-differences -Doncaster's method shows that colour-blindness may be a <i>recessive</i> due -to the absence of a factor which produces normal colour-vision.</p></div></div> - -<p><span class='pagenum'><a name="Page_48" id="Page_48">[Pg 48]</a></span></p> - -<p class="center space-above">Cambridge:<br />PRINTED BY JOHN CLAY, M.A.<br />AT THE UNIVERSITY PRESS.</p> - - - - - - - - -<pre> - - - - - -End of Project Gutenberg's The Methods and Scope of Genetics, by W. 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