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Note on the Resemblances and Differences in the Structure and the Development of the Brain in Man and Apes T.H. Huxley 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. Copyright laws are changing all over the world, be sure to check the copyright laws for your country before posting these files!! Please take a look at the important information in this header. We encourage you to keep this file on your own disk, keeping an electronic path open for the next readers. Do not remove this. **Welcome To The World of Free Plain Vanilla Electronic Texts** **Etexts Readable By Both Humans and By Computers, Since 1971** *These Etexts Prepared By Hundreds of Volunteers and Donations* Information on contacting Project Gutenberg to get Etexts, and further information is included below. We need your donations. 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] *******This file should be named huxbr10.txt or huxbr10.zip****** Corrected EDITIONS of our etexts get a new NUMBER, huxbr11.txt VERSIONS based on separate sources get new LETTER, huxbr10a.txt This etext was prepared by Sue Asscher <[email protected]>

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Note on the Resemblances and Differences in the Structureand the Development of the Brain in Man and Apes

T.H. Huxley

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 theDevelopment 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.

Copyright laws are changing all over the world, be sure to checkthe copyright laws for your country before posting these files!!

Please take a look at the important information in this header.We encourage you to keep this file on your own disk, keeping anelectronic path open for the next readers. Do not remove this.

**Welcome To The World of Free Plain Vanilla Electronic Texts**

**Etexts Readable By Both Humans and By Computers, Since 1971**

*These Etexts Prepared By Hundreds of Volunteers and Donations*

Information on contacting Project Gutenberg to get Etexts, andfurther information is included below. We need your donations.

Note on the Resemblances and Differences in the Structure and theDevelopment 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]*******This file should be named huxbr10.txt or huxbr10.zip******

Corrected EDITIONS of our etexts get a new NUMBER, huxbr11.txtVERSIONS based on separate sources get new LETTER, huxbr10a.txt

This etext was prepared by Sue Asscher <[email protected]>

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NOTE ON THE RESEMBLANCES AND DIFFERENCES IN THE STRUCTURE AND THEDEVELOPMENT 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 inrelation to Sex' by Charles Darwin where it appears at the end ofChapter VII which is also the end of Part I. Footnotes arenumbered as they appear in 'The Descent of Man.']

The controversy respecting the nature and the extent of thedifferences 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, totallydifferent from what it was formerly. It was originally assertedand re-asserted, with singular pertinacity, that the brain of allthe apes, even the highest, differs from that of man, in theabsence of such conspicuous structures as the posterior lobes ofthe cerebral hemispheres, with the posterior cornu of the lateralventricle 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 welldeveloped in apes' as in human brains, or even better; and thatit is characteristic of all the Primates (if we exclude theLemurs) to have these parts well developed, stands at present onas secure a basis as any proposition in comparative anatomy.Moreover, it is admitted by every one of the long series ofanatomists who, of late years, have paid special attention to thearrangement of the complicated sulci and gyri which appear uponthe surface of the cerebral hemispheres in man and the higherapes, that they are disposed after the very same pattern in him,as in them. Every principal gyrus and sulcus of a chimpanzee'sbrain is clearly represented in that of a man, so that theterminology which applies to the one answers for the other. Onthis point there is no difference of opinion. Some years since,

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Professor Bischoff published a memoir (70. 'Die Grosshirn-Windungen des Menschen;' 'Abhandlungen der K. BayerischenAkademie,' B. x. 1868.) on the cerebral convolutions of man andapes; and as the purpose of my learned colleague was certainlynot to diminish the value of the differences between apes and menin 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 toany other animal, is a well known fact, disputed by nobody.Looking at the matter from the point of view of organisationalone, no one probably would ever have disputed the view ofLinnaeus, that man should be placed, merely as a peculiarspecies, at the head of the mammalia and of those apes. Bothshew, in all their organs, so close an affinity, that the mostexact anatomical investigation is needed in order to demonstratethose differences which really exist. So it is with the brains.The brains of man, the orang, the chimpanzee, the gorilla, inspite of all the important differences which they present, comevery close to one another" (loc. cit. p. 101).

There remains, then, no dispute as to the resemblance infundamental characters, between the ape's brain and man's: norany as to the wonderfully close similarity between thechimpanzee, orang and man, in even the details of the arrangementof the gyri and sulci of the cerebral hemispheres. Nor, turningto the differences between the brains of the highest apes andthat of man, is there any serious question as to the nature andextent of these differences. It is admitted that the man'scerebral hemispheres are absolutely and relatively larger thanthose of the orang and chimpanzee; that his frontal lobes areless excavated by the upward protrusion of the roof of theorbits; that his gyri and sulci are, as a rule, lesssymmetrically disposed, and present a greater number of secondaryplications. And it is admitted that, as a rule, in man, thetemporo-occipital or "external perpendicular" fissure, which isusually so strongly marked a feature of the ape's brain is butfaintly marked. But it is also clear, that none of thesedifferences constitutes a sharp demarcation between the man's andthe ape's brain. In respect to the external perpendicularfissure of Gratiolet, in the human brain for instance, ProfessorTurner remarks: (71. 'Convolutions of the Human CerebrumTopographically Considered,' 1866, p. 12.)

"In some brains it appears simply as an indentation of the marginof the hemisphere, but, in others, it extends for some distancemore or less transversely outwards. I saw it in the righthemisphere of a female brain pass more than two inches outwards;and on another specimen, also the right hemisphere, it proceededfor four-tenths of an inch outwards, and then extended downwards,as far as the lower margin of the outer surface of thehemisphere. The imperfect definition of this fissure in themajority of human brains, as compared with its remarkabledistinctness in the brain of most Quadrumana, is owing to thepresence, in the former, of certain superficial, well marked,secondary convolutions which bridge it over and connect theparietal with the occipital lobe. The closer the first of thesebridging gyri lies to the longitudinal fissure, the shorter is

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the external parieto-occipital fissure" (loc. cit. p. 12).

The obliteration of the external perpendicular fissure ofGratiolet, therefore, is not a constant character of the humanbrain. On the other hand, its full development is not a constantcharacter of the higher ape's brain. For, in the chimpanzee, themore or less extensive obliteration of the external perpendicularsulcus by "bridging convolutions," on one side or the other, hasbeen noted over and over again by Prof. Rolleston, Mr. Marshall,M. Broca and Professor Turner. At the conclusion of a specialpaper on this subject the latter writes: (72. Notes moreespecially on the bridging convolutions in the Brain of theChimpanzee, 'Proceedings of the Royal Society of Edinburgh,'1865-6.)

"The three specimens of the brain of a chimpanzee, justdescribed, prove, that the generalisation which Gratiolet hasattempted to draw of the complete absence of the first connectingconvolution and the concealment of the second, as essentiallycharacteristic features in the brain of this animal, is by nomeans universally applicable. In only one specimen did thebrain, in these particulars, follow the law which Gratiolet hasexpressed. As regards the presence of the superior bridgingconvolution, I am inclined to think that it has existed in onehemisphere, at least, in a majority of the brains of this animalwhich have, up to this time, been figured or described. Thesuperficial position of the second bridging convolution isevidently less frequent, and has as yet, I believe, only beenseen in the brain (A) recorded in this communication. Theasymmetrical arrangement in the convolutions of the twohemispheres, which previous observers have referred to in theirdescriptions, is also well illustrated in these specimens" (pp.8, 9).

Even were the presence of the temporo-occipital, or externalperpendicular, sulcus, a mark of distinction between the higherapes and man, the value of such a distinctive character would berendered very doubtful by the structure of the brain in thePlatyrrhine apes. In fact, while the temporo-occipital is one ofthe most constant of sulci in the Catarrhine, or Old World, apes,it is never very strongly developed in the New World apes; it isabsent in the smaller Platyrrhini; rudimentary in Pithecia (73.Flower, 'On the Anatomy of Pithecia Monachus,' 'Proceedings ofthe Zoological Society,' 1862.); and more or less obliterated bybridging convolutions in Ateles.

A character which is thus variable within the limits of a singlegroup can have no great taxonomic value.

It is further established, that the degree of asymmetry of theconvolution of the two sides in the human brain is subject tomuch individual variation; and that, in those individuals of theBushman race who have been examined, the gyri and sulci of thetwo hemispheres are considerably less complicated and moresymmetrical than in the European brain, while, in someindividuals of the chimpanzee, their complexity and asymmetrybecome notable. This is particularly the case in the brain of ayoung male chimpanzee figured by M. Broca. ('L'ordre des

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Primates,' p. 165, fig. 11.)

Again, as respects the question of absolute size, it isestablished that the difference between the largest and thesmallest healthy human brain is greater than the differencebetween the smallest healthy human brain and the largestchimpanzee's or orang's brain.

Moreover, there is one circumstance in which the orang's andchimpanzee's brains resemble man's, but in which they differ fromthe lower apes, and that is the presence of two corporacandicantia--the Cynomorpha having but one.

In view of these facts I do not hesitate in this year 1874, torepeat and insist upon the proposition which I enunciated in1863: (74. 'Man's Place in Nature,' p. 102.)

"So far as cerebral structure goes, therefore, it is clear thatman differs less from the chimpanzee or the orang, than these doeven from the monkeys, and that the difference between the brainof the chimpanzee and of man is almost insignificant whencompared with that between the chimpanzee brain and that of aLemur."

In the paper to which I have referred, Professor Bischoff doesnot deny the second part of this statement, but he first makesthe irrelevant remark that it is not wonderful if the brains ofan orang and a Lemur are very different; and secondly, goes on toassert that, "If we successively compare the brain of a man withthat of an orang; the brain of this with that of a chimpanzee; ofthis 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 agreater, or even as great a, break in the degree of developmentof the convolutions, as we find between the brain of a man andthat of an orang or chimpanzee."

To which I reply, firstly, that whether this assertion be true orfalse, it has nothing whatever to do with the propositionenunciated in 'Man's Place in Nature,' which refers not to thedevelopment of the convolutions alone, but to the structure ofthe whole brain. If Professor Bischoff had taken the trouble torefer to p. 96 of the work he criticises, in fact, he would havefound the following passage: "And it is a remarkablecircumstance that though, so far as our present knowledgeextends, there IS one true structural break in the series offorms of Simian brains, this hiatus does not lie between man andthe manlike apes, but between the lower and the lowest Simians,or in other words, between the Old and New World apes and monkeysand the Lemurs. Every Lemur which has yet been examined, infact, has its cerebellum partially visible from above; and itsposterior lobe, with the contained posterior cornu andhippocampus minor, more or less rudimentary. Every marmoset,American monkey, Old World monkey, baboon or manlike ape, on thecontrary, has its cerebellum entirely hidden, posteriorly, by thecerebral lobes, and possesses a large posterior cornu with awell-developed hippocampus minor."

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This statement was a strictly accurate account of what was knownwhen it was made; and it does not appear to me to be more thanapparently weakened by the subsequent discovery of the relativelysmall development of the posterior lobes in the Siamang and inthe Howling monkey. Notwithstanding the exceptional brevity ofthe posterior lobes in these two species, no one will pretendthat their brains, in the slightest degree, approach those of theLemurs. And if, instead of putting Hapale out of its naturalplace, as Professor Bischoff most unaccountably does, we writethe 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 thisseries lies between Hapale and Lemur, and that this break isconsiderably greater than that between any other two terms ofthat series. Professor Bischoff ignores the fact that longbefore he wrote, Gratiolet had suggested the separation of theLemurs from the other Primates on the very ground of thedifference in their cerebral characters; and that ProfessorFlower had made the following observations in the course of hisdescription of the brain of the Javan Loris: (75. 'Transactionsof the Zoological Society,' vol. v. 1862.)

"And it is especially remarkable that, in the development of theposterior lobes, there is no approximation to the Lemurine, shorthemisphered brain, in those monkeys which are commonly supposedto approach this family in other respects, viz. the lower membersof the Platyrrhine group."

So far as the structure of the adult brain is concerned, then,the very considerable additions to our knowledge, which have beenmade by the researches of so many investigators, during the pastten years, fully justify the statement which I made in 1863. Butit has been said, that, admitting the similarity between theadult brains of man and apes, they are nevertheless, in reality,widely different, because they exhibit fundamental differences inthe mode of their development. No one would be more ready than Ito admit the force of this argument, if such fundamentaldifferences of development really exist. But I deny that they doexist. On the contrary, there is a fundamental agreement in thedevelopment of the brain in men and apes.

Gratiolet originated the statement that there is a fundamentaldifference in the development of the brains of apes and that ofman--consisting in this; that, in the apes, the sulci which firstmake their appearance are situated on the posterior region of thecerebral hemispheres, while, in the human foetus, the sulci firstbecome visible on the frontal lobes. (76. "Chez tous les singes,les plis posterieurs se developpent les premiers; les plisanterieurs se developpent plus tard, aussi la vertebre occipitaleet la parietale sont-elles relativement tres-grandes chez lefoetus. L'Homme presente une exception remarquable quant al'epoque de l'apparition des plis frontaux, qui sont les premiersindiques; mais le developpement general du lobe frontal, envisageseulement par rapport a son volume, suit les memes lois que dansles singes:" Gratiolet, 'Memoire sur les plis cerebres del'Homme et des Primateaux,' p. 39, Tab. iv, fig. 3.)

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This general statement is based upon two observations, the one ofa Gibbon almost ready to be born, in which the posterior gyriwere "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 posterieurssont bien developpes, tandis que les plis du lobe frontal sont apeine indiques." The figure, however (Pl. iv, fig. 3), shews thefissure of Rolando, and one of the frontal sulci plainly enough.Nevertheless, M. Alix, in his 'Notice sur les travauxanthropologiques de Gratiolet' ('Mem. de la Societed'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 lesanthropoides. M. Huxley, par exemple, n'hesite pas sur ce point.Eh bien, c'est sur le cerveau d'un foetus de Gibbon que Gratioleta vu LES CIRCONVOLUTIONS DU LOBE TEMPORO-SPHENOIDAL DEJADEVELOPPEES LORSQU'IL N'EXISTENT PAS ENCORE DE PLIS SUR LE LOBEFRONTAL. Il etait donc bien autorise a dire que, chez l'hommeles circonvolutions apparaissent d'a en w, tandis que chez lessinges elles se developpent d'w en a."), and the other of a humanfoetus at the 22nd or 23rd week of uterogestation, in whichGratiolet notes that the insula was uncovered, but thatnevertheless "des incisures sement de lobe anterieur, unescissure peu profonde indique la separation du lobe occipital,tres-reduit, d'ailleurs des cette epoque. Le reste de la surfacecerebrale 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 viewsof the hemispheres, but not the inner view. It is worthy of notethat the figure by no means bears out Gratiolet's description,inasmuch as the fissure (antero-temporal) on the posterior halfof the face of the hemisphere is more marked than any of thosevaguely indicated in the anterior half. If the figure iscorrect, it in no way justifies Gratiolet's conclusion: "Il y adonc entre ces cerveaux [those of a Callithrix and of a Gibbon]et celui du foetus humain une difference fondamental. Chezcelui-ci, longtemps avant que les plis temporaux apparaissent,les plis frontaux, ESSAYENT d'exister."

Since Gratiolet's time, however, the development of the gyri andsulci of the brain has been made the subject of renewedinvestigation by Schmidt, Bischoff, Pansch (78. 'Ueber dietypische Anordnung der Furchen und Windungen auf den Grosshirn-Hemispharen des Menschen und der Affen,' 'Archiv furAnthropologie,' iii. 1868.), and more particularly by Ecker (79.'Zur Entwicklungs Geschichte der Furchen und Windungen derGrosshirn-Hemispharen im Foetus des Menschen.' 'Archiv furAnthropologie,' 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 asfollows:--

1. In the human foetus, the sylvian fissure is formed in thecourse of the third month of uterogestation. In this, and in thefourth month, the cerebral hemispheres are smooth and rounded

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(with the exception of the sylvian depression), and they projectbackwards far beyond the cerebellum.

2. The sulci, properly so called, begin to appear in theinterval between the end of the fourth and the beginning of thesixth month of foetal life, but Ecker is careful to point outthat, not only the time, but the order, of their appearance issubject to considerable individual variation. In no case,however, are either the frontal or the temporal sulci theearliest.

The first which appears, in fact, lies on the inner face of thehemisphere (whence doubtless Gratiolet, who does not seem to haveexamined that face in his foetus, overlooked it), and is eitherthe internal perpendicular (occipito-parietal), or the calcarinesulcus, these two being close together and eventually runninginto one another. As a rule the occipito-parietal is the earlierof the two.

3. At the latter part of this period, another sulcus, the"posterio-parietal," or "Fissure of Rolando" is developed, and itis followed, in the course of the sixth month, by the otherprincipal sulci of the frontal, parietal, temporal and occipitallobes. There is, however, no clear evidence that one of theseconstantly 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-temporalsulcus (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 theorder of the appearance of the sulci and gyri in the foetal humanbrain is in perfect harmony with the general doctrine ofevolution, and with the view that man has been evolved from someape-like form; though there can be no doubt that form was, inmany respects, different from any member of the Primates nowliving.

Von Baer taught us, half a century ago, that, in the course oftheir development, allied animals put on at first, the charactersof the greater groups to which they belong, and, by degrees,assume those which restrict them within the limits of theirfamily, genus, and species; and he proved, at the same time, thatno developmental stage of a higher animal is precisely similar tothe adult condition of any lower animal. It is quite correct tosay that a frog passes through the condition of a fish, inasmuchas at one period of its life the tadpole has all the charactersof a fish, and if it went no further, would have to be groupedamong fishes. But it is equally true that a tadpole is verydifferent 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, butthat of an Arctopithecine or marmoset-like ape; for itshemispheres, with their great posterior lobster, and with nosulci but the sylvian and the calcarine, present thecharacteristics found only in the group of the Arctopithecine

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Primates. But it is equally true, as Gratiolet remarks, that, inits widely open sylvian fissure, it differs from the brain of anyactual marmoset. No doubt it would be much more similar to thebrain of an advanced foetus of a marmoset. But we know nothingwhatever of the development of the brain in the marmosets. Inthe Platyrrhini proper, the only observation with which I amacquainted is due to Pansch, who found in the brain of a foetalCebus Apella, in addition to the sylvian fissure and the deepcalcarine fissure, only a very shallow antero-temporal fissure(scissure parallele of Gratiolet).

Now this fact, taken together with the circumstance that theantero-temporal sulcus is present in such Platyrrhini as theSaimiri, which present mere traces of sulci on the anterior halfof the exterior of the cerebral hemispheres, or none at all,undoubtedly, so far as it goes, affords fair evidence in favourof Gratiolet's hypothesis, that the posterior sulci appear beforethe anterior, in the brains of the Platyrrhini. But, it by nomeans follows, that the rule which may hold good for thePlatyrrhini extends to the Catarrhini. We have no informationwhatever respecting the development of the brain in theCynomorpha; and, as regards the Anthropomorpha, nothing but theaccount of the brain of the Gibbon, near birth, already referredto. At the present moment there is not a shadow of evidence toshew that the sulci of a chimpanzee's, or orang's, brain do notappear in the same order as a man's.

Gratiolet opens his preface with the aphorism: "Il est dangereuxdans les sciences de conclure trop vite." I fear he must haveforgotten this sound maxim by the time he had reached thediscussion of the differences between men and apes, in the bodyof his work. No doubt, the excellent author of one of the mostremarkable contributions to the just understanding of themammalian brain which has ever been made, would have been thefirst to admit the insufficiency of his data had he lived toprofit by the advance of inquiry. The misfortune is that hisconclusions have been employed by persons incompetent toappreciate their foundation, as arguments in favour ofobscurantism. (80. For example, M. l'Abbe Lecomte in histerrible pamphlet, 'Le Darwinisme et l'origine de l'Homme,'1873.)

But it is important to remark that, whether Gratiolet was rightor wrong in his hypothesis respecting the relative order ofappearance of the temporal and frontal sulci, the fact remains;that before either temporal or frontal sulci, appear, the foetalbrain of man presents characters which are found only in thelowest group of the Primates (leaving out the Lemurs); and thatthis is exactly what we should expect to be the case, if man hasresulted from the gradual modification of the same form as thatfrom which the other Primates have sprung.

End of T.H. Huxley's On the Brain [from Descent of Man by Charles Darwin]