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+T.H. Huxley's On the Brain [from Descent of Man by Charles Darwin]
+
+The complete title is:
+Note on the Resemblances and Differences in the Structure and the
+Development of the Brain in Man and Apes
+
+#3 in our series by T.H. Huxley
+
+#8 in our series by Charles Darwin
+
+Our Darwin editor says this should be list under both names.
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+Note on the Resemblances and Differences in the Structure and the
+Development of the Brain in Man and Apes
+
+by Professor T.H. Huxley, F.R.S.
+
+From The Descent of Man by Charles Darwin
+
+October, 2000 [Etext #2354]
+
+
+T.H. Huxley's On the Brain [from Descent of Man by Charles Darwin]
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+
+NOTE ON THE RESEMBLANCES AND DIFFERENCES IN THE STRUCTURE AND THE
+DEVELOPMENT OF THE BRAIN IN MAN AND APES
+
+BY
+
+PROFESSOR T.H. HUXLEY, F.R.S.
+
+
+[This essay is taken from 'The Descent of Man and Selection in
+relation to Sex' by Charles Darwin where it appears at the end of
+Chapter VII which is also the end of Part I. Footnotes are
+numbered as they appear in 'The Descent of Man.']
+
+
+The controversy respecting the nature and the extent of the
+differences in the structure of the brain in man and the apes,
+which arose some fifteen years ago, has not yet come to an end,
+though the subject matter of the dispute is, at present, totally
+different from what it was formerly. It was originally asserted
+and re-asserted, with singular pertinacity, that the brain of all
+the apes, even the highest, differs from that of man, in the
+absence of such conspicuous structures as the posterior lobes of
+the cerebral hemispheres, with the posterior cornu of the lateral
+ventricle and the hippocampus minor, contained in those lobes,
+which are so obvious in man.
+
+But the truth that the three structures in question are as well
+developed in apes' as in human brains, or even better; and that
+it is characteristic of all the Primates (if we exclude the
+Lemurs) to have these parts well developed, stands at present on
+as secure a basis as any proposition in comparative anatomy.
+Moreover, it is admitted by every one of the long series of
+anatomists who, of late years, have paid special attention to the
+arrangement of the complicated sulci and gyri which appear upon
+the surface of the cerebral hemispheres in man and the higher
+apes, that they are disposed after the very same pattern in him,
+as in them. Every principal gyrus and sulcus of a chimpanzee's
+brain is clearly represented in that of a man, so that the
+terminology which applies to the one answers for the other. On
+this point there is no difference of opinion. Some years since,
+Professor Bischoff published a memoir (70. 'Die Grosshirn-
+Windungen des Menschen;' 'Abhandlungen der K. Bayerischen
+Akademie,' B. x. 1868.) on the cerebral convolutions of man and
+apes; and as the purpose of my learned colleague was certainly
+not to diminish the value of the differences between apes and men
+in this respect, I am glad to make a citation from him.
+
+"That the apes, and especially the orang, chimpanzee and gorilla,
+come very close to man in their organisation, much nearer than to
+any other animal, is a well known fact, disputed by nobody.
+Looking at the matter from the point of view of organisation
+alone, no one probably would ever have disputed the view of
+Linnaeus, that man should be placed, merely as a peculiar
+species, at the head of the mammalia and of those apes. Both
+shew, in all their organs, so close an affinity, that the most
+exact anatomical investigation is needed in order to demonstrate
+those differences which really exist. So it is with the brains.
+The brains of man, the orang, the chimpanzee, the gorilla, in
+spite of all the important differences which they present, come
+very close to one another" (loc. cit. p. 101).
+
+There remains, then, no dispute as to the resemblance in
+fundamental characters, between the ape's brain and man's: nor
+any as to the wonderfully close similarity between the
+chimpanzee, orang and man, in even the details of the arrangement
+of the gyri and sulci of the cerebral hemispheres. Nor, turning
+to the differences between the brains of the highest apes and
+that of man, is there any serious question as to the nature and
+extent of these differences. It is admitted that the man's
+cerebral hemispheres are absolutely and relatively larger than
+those of the orang and chimpanzee; that his frontal lobes are
+less excavated by the upward protrusion of the roof of the
+orbits; that his gyri and sulci are, as a rule, less
+symmetrically disposed, and present a greater number of secondary
+plications. And it is admitted that, as a rule, in man, the
+temporo-occipital or "external perpendicular" fissure, which is
+usually so strongly marked a feature of the ape's brain is but
+faintly marked. But it is also clear, that none of these
+differences constitutes a sharp demarcation between the man's and
+the ape's brain. In respect to the external perpendicular
+fissure of Gratiolet, in the human brain for instance, Professor
+Turner remarks: (71. 'Convolutions of the Human Cerebrum
+Topographically Considered,' 1866, p. 12.)
+
+"In some brains it appears simply as an indentation of the margin
+of the hemisphere, but, in others, it extends for some distance
+more or less transversely outwards. I saw it in the right
+hemisphere of a female brain pass more than two inches outwards;
+and on another specimen, also the right hemisphere, it proceeded
+for four-tenths of an inch outwards, and then extended downwards,
+as far as the lower margin of the outer surface of the
+hemisphere. The imperfect definition of this fissure in the
+majority of human brains, as compared with its remarkable
+distinctness in the brain of most Quadrumana, is owing to the
+presence, in the former, of certain superficial, well marked,
+secondary convolutions which bridge it over and connect the
+parietal with the occipital lobe. The closer the first of these
+bridging gyri lies to the longitudinal fissure, the shorter is
+the external parieto-occipital fissure" (loc. cit. p. 12).
+
+The obliteration of the external perpendicular fissure of
+Gratiolet, therefore, is not a constant character of the human
+brain. On the other hand, its full development is not a constant
+character of the higher ape's brain. For, in the chimpanzee, the
+more or less extensive obliteration of the external perpendicular
+sulcus by "bridging convolutions," on one side or the other, has
+been noted over and over again by Prof. Rolleston, Mr. Marshall,
+M. Broca and Professor Turner. At the conclusion of a special
+paper on this subject the latter writes: (72. Notes more
+especially on the bridging convolutions in the Brain of the
+Chimpanzee, 'Proceedings of the Royal Society of Edinburgh,'
+1865-6.)
+
+"The three specimens of the brain of a chimpanzee, just
+described, prove, that the generalisation which Gratiolet has
+attempted to draw of the complete absence of the first connecting
+convolution and the concealment of the second, as essentially
+characteristic features in the brain of this animal, is by no
+means universally applicable. In only one specimen did the
+brain, in these particulars, follow the law which Gratiolet has
+expressed. As regards the presence of the superior bridging
+convolution, I am inclined to think that it has existed in one
+hemisphere, at least, in a majority of the brains of this animal
+which have, up to this time, been figured or described. The
+superficial position of the second bridging convolution is
+evidently less frequent, and has as yet, I believe, only been
+seen in the brain (A) recorded in this communication. The
+asymmetrical arrangement in the convolutions of the two
+hemispheres, which previous observers have referred to in their
+descriptions, is also well illustrated in these specimens" (pp.
+8, 9).
+
+Even were the presence of the temporo-occipital, or external
+perpendicular, sulcus, a mark of distinction between the higher
+apes and man, the value of such a distinctive character would be
+rendered very doubtful by the structure of the brain in the
+Platyrrhine apes. In fact, while the temporo-occipital is one of
+the most constant of sulci in the Catarrhine, or Old World, apes,
+it is never very strongly developed in the New World apes; it is
+absent in the smaller Platyrrhini; rudimentary in Pithecia (73.
+Flower, 'On the Anatomy of Pithecia Monachus,' 'Proceedings of
+the Zoological Society,' 1862.); and more or less obliterated by
+bridging convolutions in Ateles.
+
+A character which is thus variable within the limits of a single
+group can have no great taxonomic value.
+
+It is further established, that the degree of asymmetry of the
+convolution of the two sides in the human brain is subject to
+much individual variation; and that, in those individuals of the
+Bushman race who have been examined, the gyri and sulci of the
+two hemispheres are considerably less complicated and more
+symmetrical than in the European brain, while, in some
+individuals of the chimpanzee, their complexity and asymmetry
+become notable. This is particularly the case in the brain of a
+young male chimpanzee figured by M. Broca. ('L'ordre des
+Primates,' p. 165, fig. 11.)
+
+Again, as respects the question of absolute size, it is
+established that the difference between the largest and the
+smallest healthy human brain is greater than the difference
+between the smallest healthy human brain and the largest
+chimpanzee's or orang's brain.
+
+Moreover, there is one circumstance in which the orang's and
+chimpanzee's brains resemble man's, but in which they differ from
+the lower apes, and that is the presence of two corpora
+candicantia--the Cynomorpha having but one.
+
+In view of these facts I do not hesitate in this year 1874, to
+repeat and insist upon the proposition which I enunciated in
+1863: (74. 'Man's Place in Nature,' p. 102.)
+
+"So far as cerebral structure goes, therefore, it is clear that
+man differs less from the chimpanzee or the orang, than these do
+even from the monkeys, and that the difference between the brain
+of the chimpanzee and of man is almost insignificant when
+compared with that between the chimpanzee brain and that of a
+Lemur."
+
+In the paper to which I have referred, Professor Bischoff does
+not deny the second part of this statement, but he first makes
+the irrelevant remark that it is not wonderful if the brains of
+an orang and a Lemur are very different; and secondly, goes on to
+assert that, "If we successively compare the brain of a man with
+that of an orang; the brain of this with that of a chimpanzee; of
+this with that of a gorilla, and so on of a Hylobates,
+Semnopithecus, Cynocephalus, Cercopithecus, Macacus, Cebus,
+Callithrix, Lemur, Stenops, Hapale, we shall not meet with a
+greater, or even as great a, break in the degree of development
+of the convolutions, as we find between the brain of a man and
+that of an orang or chimpanzee."
+
+To which I reply, firstly, that whether this assertion be true or
+false, it has nothing whatever to do with the proposition
+enunciated in 'Man's Place in Nature,' which refers not to the
+development of the convolutions alone, but to the structure of
+the whole brain. If Professor Bischoff had taken the trouble to
+refer to p. 96 of the work he criticises, in fact, he would have
+found the following passage: "And it is a remarkable
+circumstance that though, so far as our present knowledge
+extends, there IS one true structural break in the series of
+forms of Simian brains, this hiatus does not lie between man and
+the manlike apes, but between the lower and the lowest Simians,
+or in other words, between the Old and New World apes and monkeys
+and the Lemurs. Every Lemur which has yet been examined, in
+fact, has its cerebellum partially visible from above; and its
+posterior lobe, with the contained posterior cornu and
+hippocampus minor, more or less rudimentary. Every marmoset,
+American monkey, Old World monkey, baboon or manlike ape, on the
+contrary, has its cerebellum entirely hidden, posteriorly, by the
+cerebral lobes, and possesses a large posterior cornu with a
+well-developed hippocampus minor."
+
+This statement was a strictly accurate account of what was known
+when it was made; and it does not appear to me to be more than
+apparently weakened by the subsequent discovery of the relatively
+small development of the posterior lobes in the Siamang and in
+the Howling monkey. Notwithstanding the exceptional brevity of
+the posterior lobes in these two species, no one will pretend
+that their brains, in the slightest degree, approach those of the
+Lemurs. And if, instead of putting Hapale out of its natural
+place, as Professor Bischoff most unaccountably does, we write
+the series of animals he has chosen to mention as follows: Homo,
+Pithecus, Troglodytes, Hylobates, Semnopithecus, Cynocephalus,
+Cercopithecus, Macacus, Cebus, Callithrix, Hapale, Lemur,
+Stenops, I venture to reaffirm that the great break in this
+series lies between Hapale and Lemur, and that this break is
+considerably greater than that between any other two terms of
+that series. Professor Bischoff ignores the fact that long
+before he wrote, Gratiolet had suggested the separation of the
+Lemurs from the other Primates on the very ground of the
+difference in their cerebral characters; and that Professor
+Flower had made the following observations in the course of his
+description of the brain of the Javan Loris: (75. 'Transactions
+of the Zoological Society,' vol. v. 1862.)
+
+"And it is especially remarkable that, in the development of the
+posterior lobes, there is no approximation to the Lemurine, short
+hemisphered brain, in those monkeys which are commonly supposed
+to approach this family in other respects, viz. the lower members
+of the Platyrrhine group."
+
+So far as the structure of the adult brain is concerned, then,
+the very considerable additions to our knowledge, which have been
+made by the researches of so many investigators, during the past
+ten years, fully justify the statement which I made in 1863. But
+it has been said, that, admitting the similarity between the
+adult brains of man and apes, they are nevertheless, in reality,
+widely different, because they exhibit fundamental differences in
+the mode of their development. No one would be more ready than I
+to admit the force of this argument, if such fundamental
+differences of development really exist. But I deny that they do
+exist. On the contrary, there is a fundamental agreement in the
+development of the brain in men and apes.
+
+Gratiolet originated the statement that there is a fundamental
+difference in the development of the brains of apes and that of
+man--consisting in this; that, in the apes, the sulci which first
+make their appearance are situated on the posterior region of the
+cerebral hemispheres, while, in the human foetus, the sulci first
+become visible on the frontal lobes. (76. "Chez tous les singes,
+les plis posterieurs se developpent les premiers; les plis
+anterieurs se developpent plus tard, aussi la vertebre occipitale
+et la parietale sont-elles relativement tres-grandes chez le
+foetus. L'Homme presente une exception remarquable quant a
+l'epoque de l'apparition des plis frontaux, qui sont les premiers
+indiques; mais le developpement general du lobe frontal, envisage
+seulement par rapport a son volume, suit les memes lois que dans
+les singes:" Gratiolet, 'Memoire sur les plis cerebres de
+l'Homme et des Primateaux,' p. 39, Tab. iv, fig. 3.)
+
+This general statement is based upon two observations, the one of
+a Gibbon almost ready to be born, in which the posterior gyri
+were "well developed," while those of the frontal lobes were
+"hardly indicated" (77. Gratiolet's words are (loc. cit. p. 39):
+"Dans le foetus dont il s'agit les plis cerebraux posterieurs
+sont bien developpes, tandis que les plis du lobe frontal sont a
+peine indiques." The figure, however (Pl. iv, fig. 3), shews the
+fissure of Rolando, and one of the frontal sulci plainly enough.
+Nevertheless, M. Alix, in his 'Notice sur les travaux
+anthropologiques de Gratiolet' ('Mem. de la Societe
+d'Anthropologie de Paris,' 1868, page 32), writes thus:
+"Gratiolet a eu entre les mains le cerveau d'un foetus de Gibbon,
+singe eminemment superieur, et tellement rapproche de l'orang,
+que des naturalistes tres-competents l'ont range parmi les
+anthropoides. M. Huxley, par exemple, n'hesite pas sur ce point.
+Eh bien, c'est sur le cerveau d'un foetus de Gibbon que Gratiolet
+a vu LES CIRCONVOLUTIONS DU LOBE TEMPORO-SPHENOIDAL DEJA
+DEVELOPPEES LORSQU'IL N'EXISTENT PAS ENCORE DE PLIS SUR LE LOBE
+FRONTAL. Il etait donc bien autorise a dire que, chez l'homme
+les circonvolutions apparaissent d'a en w, tandis que chez les
+singes elles se developpent d'w en a."), and the other of a human
+foetus at the 22nd or 23rd week of uterogestation, in which
+Gratiolet notes that the insula was uncovered, but that
+nevertheless "des incisures sement de lobe anterieur, une
+scissure peu profonde indique la separation du lobe occipital,
+tres-reduit, d'ailleurs des cette epoque. Le reste de la surface
+cerebrale est encore absolument lisse."
+
+Three views of this brain are given in Plate II, figs. 1, 2, 3,
+of the work cited, shewing the upper, lateral and inferior views
+of the hemispheres, but not the inner view. It is worthy of note
+that the figure by no means bears out Gratiolet's description,
+inasmuch as the fissure (antero-temporal) on the posterior half
+of the face of the hemisphere is more marked than any of those
+vaguely indicated in the anterior half. If the figure is
+correct, it in no way justifies Gratiolet's conclusion: "Il y a
+donc entre ces cerveaux [those of a Callithrix and of a Gibbon]
+et celui du foetus humain une difference fondamental. Chez
+celui-ci, longtemps avant que les plis temporaux apparaissent,
+les plis frontaux, ESSAYENT d'exister."
+
+Since Gratiolet's time, however, the development of the gyri and
+sulci of the brain has been made the subject of renewed
+investigation by Schmidt, Bischoff, Pansch (78. 'Ueber die
+typische Anordnung der Furchen und Windungen auf den Grosshirn-
+Hemispharen des Menschen und der Affen,' 'Archiv fur
+Anthropologie,' iii. 1868.), and more particularly by Ecker (79.
+'Zur Entwicklungs Geschichte der Furchen und Windungen der
+Grosshirn-Hemispharen im Foetus des Menschen.' 'Archiv fur
+Anthropologie,' iii. 1868.), whose work is not only the latest,
+but by far the most complete, memoir on the subject.
+
+The final results of their inquiries may be summed up as
+follows:--
+
+1. In the human foetus, the sylvian fissure is formed in the
+course of the third month of uterogestation. In this, and in the
+fourth month, the cerebral hemispheres are smooth and rounded
+(with the exception of the sylvian depression), and they project
+backwards far beyond the cerebellum.
+
+2. The sulci, properly so called, begin to appear in the
+interval between the end of the fourth and the beginning of the
+sixth month of foetal life, but Ecker is careful to point out
+that, not only the time, but the order, of their appearance is
+subject to considerable individual variation. In no case,
+however, are either the frontal or the temporal sulci the
+earliest.
+
+The first which appears, in fact, lies on the inner face of the
+hemisphere (whence doubtless Gratiolet, who does not seem to have
+examined that face in his foetus, overlooked it), and is either
+the internal perpendicular (occipito-parietal), or the calcarine
+sulcus, these two being close together and eventually running
+into one another. As a rule the occipito-parietal is the earlier
+of the two.
+
+3. At the latter part of this period, another sulcus, the
+"posterio-parietal," or "Fissure of Rolando" is developed, and it
+is followed, in the course of the sixth month, by the other
+principal sulci of the frontal, parietal, temporal and occipital
+lobes. There is, however, no clear evidence that one of these
+constantly appears before the other; and it is remarkable that,
+in the brain at the period described and figured by Ecker (loc.
+cit. pp. 212-213, Taf. II, figs. 1, 2, 3, 4), the antero-temporal
+sulcus (scissure parallele) so characteristic of the ape's brain,
+is as well, if not better developed than the fissure of Rolando,
+and is much more marked than the proper frontal sulci.
+
+Taking the facts as they now stand, it appears to me that the
+order of the appearance of the sulci and gyri in the foetal human
+brain is in perfect harmony with the general doctrine of
+evolution, and with the view that man has been evolved from some
+ape-like form; though there can be no doubt that form was, in
+many respects, different from any member of the Primates now
+living.
+
+Von Baer taught us, half a century ago, that, in the course of
+their development, allied animals put on at first, the characters
+of the greater groups to which they belong, and, by degrees,
+assume those which restrict them within the limits of their
+family, genus, and species; and he proved, at the same time, that
+no developmental stage of a higher animal is precisely similar to
+the adult condition of any lower animal. It is quite correct to
+say that a frog passes through the condition of a fish, inasmuch
+as at one period of its life the tadpole has all the characters
+of a fish, and if it went no further, would have to be grouped
+among fishes. But it is equally true that a tadpole is very
+different from any known fish.
+
+In like manner, the brain of a human foetus, at the fifth month,
+may correctly be said to be, not only the brain of an ape, but
+that of an Arctopithecine or marmoset-like ape; for its
+hemispheres, with their great posterior lobster, and with no
+sulci but the sylvian and the calcarine, present the
+characteristics found only in the group of the Arctopithecine
+Primates. But it is equally true, as Gratiolet remarks, that, in
+its widely open sylvian fissure, it differs from the brain of any
+actual marmoset. No doubt it would be much more similar to the
+brain of an advanced foetus of a marmoset. But we know nothing
+whatever of the development of the brain in the marmosets. In
+the Platyrrhini proper, the only observation with which I am
+acquainted is due to Pansch, who found in the brain of a foetal
+Cebus Apella, in addition to the sylvian fissure and the deep
+calcarine fissure, only a very shallow antero-temporal fissure
+(scissure parallele of Gratiolet).
+
+Now this fact, taken together with the circumstance that the
+antero-temporal sulcus is present in such Platyrrhini as the
+Saimiri, which present mere traces of sulci on the anterior half
+of the exterior of the cerebral hemispheres, or none at all,
+undoubtedly, so far as it goes, affords fair evidence in favour
+of Gratiolet's hypothesis, that the posterior sulci appear before
+the anterior, in the brains of the Platyrrhini. But, it by no
+means follows, that the rule which may hold good for the
+Platyrrhini extends to the Catarrhini. We have no information
+whatever respecting the development of the brain in the
+Cynomorpha; and, as regards the Anthropomorpha, nothing but the
+account of the brain of the Gibbon, near birth, already referred
+to. At the present moment there is not a shadow of evidence to
+shew that the sulci of a chimpanzee's, or orang's, brain do not
+appear in the same order as a man's.
+
+Gratiolet opens his preface with the aphorism: "Il est dangereux
+dans les sciences de conclure trop vite." I fear he must have
+forgotten this sound maxim by the time he had reached the
+discussion of the differences between men and apes, in the body
+of his work. No doubt, the excellent author of one of the most
+remarkable contributions to the just understanding of the
+mammalian brain which has ever been made, would have been the
+first to admit the insufficiency of his data had he lived to
+profit by the advance of inquiry. The misfortune is that his
+conclusions have been employed by persons incompetent to
+appreciate their foundation, as arguments in favour of
+obscurantism. (80. For example, M. l'Abbe Lecomte in his
+terrible pamphlet, 'Le Darwinisme et l'origine de l'Homme,'
+1873.)
+
+But it is important to remark that, whether Gratiolet was right
+or wrong in his hypothesis respecting the relative order of
+appearance of the temporal and frontal sulci, the fact remains;
+that before either temporal or frontal sulci, appear, the foetal
+brain of man presents characters which are found only in the
+lowest group of the Primates (leaving out the Lemurs); and that
+this is exactly what we should expect to be the case, if man has
+resulted from the gradual modification of the same form as that
+from which the other Primates have sprung.
+
+
+
+
+
+End of T.H. Huxley's On the Brain [from Descent of Man by Charles Darwin]
+