Anatomy. - "The relative weight of the braincortex in human races and in some animals and the asymmetry of the hemispheres." By Dr. C. U . ARIËNS KAPPERS.

(Communicated at the meeting of September 26. 1925).

Although many investigations have been made on the weight of the cortex. at least in man. th ere are great controversies in the results. As the methods. by which the investigators have tried to solve this problem. always were indirect ones. this is not surprising.

50. using the figures . given by W AGNER I) and CALORI 2) concerning the surface of the brain. DONALDSON 3) (l.c. p. 202-204) calculated this weight by multiplying the surface with the average depth of the human cortex and its specific weight.

On account of his own measurements (I. c. p. 203) he estimated the average dep th of the cortex. which. as we know. is locally varying in man (according to BRODMANN 4) from 1.8 tot 4.0 m.M.) to be 2.9 m.M. With OBERSTEINER 5) he took the S. W . of the human cortex to be 1.034.

DONALDSON. improving the researches of WAGNER by introducing a correction 6) for the alcohol ic shrinkage of the brainsurface so found the following figures :

Cases I Fresh T. B. W .7) I T. W . hem. 8) T . W . c. 9) Proc. C. hem.

FUCHS H99 gr. 1312gr. 743 gr. 56.6%

GAUSS H92 .. 1306 .. 731 .. 55.9%

Workman KREBS 1273 "

1114 .. 630 .. 56 .5%

Woman 29 years I 1185 .. 1037 .. 667 .. 64 .3%

Average . . . . ·1 58.3%

I) Masshestimmungen der OherBäche des groszen Gehirns. Inaugur. Diss. Göttingen 1864. 2) Del cervello nei due tipi brachicefalo e dolicocefalo italiani. Memorie dell'Accademia

delle Scienze dellïstituto di Bologna. Serie seconda, Tomo X. 1870 (p. H2-H5). 3) The growth of the brain. A study of the nervous system in relation to education .

London, Walte't Scott Itm, 1895. 4) Ueher Rindenmessungen. Centralblatt f. Nervenh. und Psych. 1908 (p. 790) und

Lokalisationslehre der Groszhirnrinde. Joh . Ambrosius Barth, Leipzig, 1909. 5) The average S.W. found by OBERSTEINER for the fresh cortex of man is properly

1.03305. Conf. OBERSTEINER. Anleitung heim Studium des Baues der nervösen Zentral­organe, 5te AuBage Wien, 1912, pag. 167 and Centr.bl. f. Nerv. h. k. 1894.

The average of the S .W . figures found by DANILEWSKY (already hefore OBERSTEINER) are also a little less. For the fresh human cortex this investigator found 1.0326. (Die Quantitativen Bestimmungen der grauen und weiszen Substanzen im Gehirn ; Centralbl. f. d. mediz. Wissensch. NO. H, 1880).

6) The figures found by WAGNER for the surface of both hemispheres together are : FUCHS 2210 cM2, GAUSS 2195 cM2, KREBS 1876 cM2, and woman 2041 cM2.

7) T . B. W. total brain weight to the calamus (without pia) . 8) T . W. Hem. = total weight of both hemispheres. 9) T . W . C. = total weight of the cortex of both hemispheres.

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The following table was composed by DONALDSON on account of the surface extension. found by CALORI in 22 brachycephalic and 19 doli­chocephalic Italians. measured chiefly in the same way as W AGNER did (see below).

I

-

Number of cases Aver. Aver. nver' l

Aver. perc . T. B. W . T.W. hem. T . W . C . C. per hem.

19 Brach. Ital. e! 1336 gr. 1169 gr. 731 gr.

I 62.5 %

16 Dolich. .. e! 1299 .. 1135 .. 690 .. 60.7 % 3 Brach. .. c;' 1148 .. 1005 " 635 " 63.2 %

3 Dolich. n 1110 971 595 61.2 % " + " " "

Average . . . . I 61 .85%

From this we see that. on account of the measurements of W AGNER. DONALDSON concludes at an average cortex weight percentage of 58.3. on account of those of CALORI of 61.85 per hemisphere.

It is not without interest to point out. what is meant by weight of hemispheres in such cases . Two methods are used to sever the hemispheres from the remainder of the brain. We understand by hemispheres either all that lies before the corp. quadrigemina ant. or cut them olf behind the corpora quadrigemina posteriora (immediately before the pons). The fir~t way is used by BOYO (see bel ow) and then the average weight of the hemispheres is about 87.5 % of that of the whole encephalon. The last method (mostly used) is followed by CLAPHAM, BROWNE and others (including myself) .

The determination of the surface has been made by W AGNER and CALORI al most in the same manner.

The outer surface of the brain was covered by triangles and parallelograms of goldleaf or tinfoil. These again were spread out. measured and counted. The cortex in the dep th of the sulci (which is almost 2/3 of the whole surface) was determined by measuring the length and the average depth of the sulci and multiplying those figures with two. It is evident that this method contains a source óf errors. Moreover the correction of the alcoholic shrinkage causes difficulties. N evertheless such a correction is necessary. because the volume and the surface shrink considerably in alcohol.

This method. determining the cortical surface in alcohol material. was also applied by JENSEN 1) on brains of a paralytic. a melancholica and three idiots. He compared his results with those of W AGNER and found that the total cortex surface of his patients varied from 94 010 to 75.3 Ofo from the figures found by W AGNER for GAUSS. the lowest percentage being that of the brain of an idiot and the other percentages. although remaining below those of GAUSS and FUCHS. falling within the group of the woman and the workman examined by W AGNER (I. c. p. 752).

1) Untersuchungen über die Beziehungen zwisschen Groszhirn und Geistesstöhrungen an sechs Gehirnen geisteskranker Individuen. Arch. f. Psychiatrie 1875 Bnd. V, Heft 3.

55 Proceedings Royal Acad. Amsterdam. Vol. XXVIII.

846

His absolute figures are of no avail for my purpose. since he examined pathological material exclusively and his figures have been derived (as the original ones of W AONER) from alcohol material.

His informations surely would have been of more value. if he himself had also measured the surface of some normal brains ; comparisons of two authors being more reliable. since different authors rarely make the same mistakes. It is selfevident that with the relative smaller surface of these brains. also a smaller volume and weight of cortex has been calculated in such cases. as is made probable also by the researches of JÄOER and HENNEBERO on pathological material (1. c. infra).

While the method of all these authors was based on measuring the surface. the average depth and on multiplying these with the S.W. of the cortex. we see that besides the objections mentioned above. this method contains still another source of errors. viz. the question how to flnd the average cortical depth. which was estimated by DONALDSON to be 2.9. by JENSEN 2.98. and by DANILEWSKY and HENNEBERO 2.5 m.M.

JÄOER 1) has avoided the use of an average depth flgure. determining the volume of the cortex directly with the compensation planimeter (already used by ANTON 2)) in the following way:

JÄOER cut the brain. af ter it had been put for a short timOe in 5010 formaline. in slices of 1 c.M.

He th en flrst measured the whole circumference of the brain (this being the external border of the cortex) with the planimeter and after~ wards the circumference of the white substance (this being the internal limit of the cortex). The latter he substracted from the flrst and by this means determined the transverse surface of the cortex of each slice. This flgure he multiplied with 1 c.M. thus calculating the volume of the cortex for each slice. For the poles he made corrections.

By multiplying the cortex volume thus found for the total brain with the S.W. 1.034. I determined the weight of the cortex. As. however. in

1) R. JÄGER. Planimetrische Messungen der Rinden- und Marksubstanz des Groszhirns. Inaugural Dissertation. Halle. 1910.

2) G. ANTON. Gehirnmessungen mittels des Kompensationsplanimeters. Wiener Klinische Rundschau : 1903. See also : Zur Kenntnisz der Störungen im Oberflächenwachstum des menschlichen Groszhirns. lste Mitteilung. Ibidem. Bnd. IX. 1888.

The planimetrie method had been applied already in 1881 by CONTI. who. however. did not determine the whole mass of the cortex. but the relation between the white and the grey substance on three levels. the first before the basal ganglia. the second through. the third behind the basal ganglia. in 1 men and 2 women (Rapport entre la substance grise et blanche du cerveau humain : Internationale Monatschrift für Anatomie und Physio­logie Bnd. 1. 1881).

Moreover Dr. ANITA TAPT has in 157 cases determined planimetrieally the relation between the grey and white substance in the transverse level of the optie chiasm. She attained the interesting result that In children. microcephalies and mongoloid idiots the relation was in favour of the grey substance. in normal cases. manie depressives and praecox patients in favour of the white substance. See Journalof nervous and mental diseases. Vol. 17. 1918.

847

his article the weight of the hemispheres is mentioned in one case only. the weightpercentage of the cortex of the hemispheres could be calculated in that case only. In a normal. four years old boy with a total brain~ weight of 1140 gr. and a weight of both hemispheres together = 997.5 gr. it amounted to 549 gr. or 55 Ofo of the weight of the hemispheres (in an adult normal woman he found 557 gr. cortex).

It appears that these figures of JÄOER are lower than those calculated by DONALDSON from the surface measurements of W AONER and CALORI.

DONALDSON I) himself also tried to determine the weight of the cortex. in still another way. viz. by making use of the figures found by DE REOIBUS for the waterpercentage and the S.W. of grey and white substance. If a hemisphere (in casu cortex and white substance without striatum) is weighed and its volume known. we may calculate its average S.W. Knowing that the S.W. of the cortex is 1.034 and that of the white substance 1.041 . we may calculate in a given case the relation between the amount of white and grey substance (in casu cortex).

From the figures of DE REOIBUS 2) . concerning the waterpercentage of grey and white substance (whïch closely coheres with its S.W.). DONALDSON calculated the quantity of cortex and came to a lower figure than calculated by him from the surface tables of W AONER and CALORJ.

He stated in his way an average of 54.8 °10 only. whïch is considerably lower than the 58.3°/0 derived from the surface measurements of WAONER and the 61.85 % calculated from those of CALORI. and closely resem bles JÄOER's (vide supra) result.

NO. of the bra in Weight 2 hem. Weight cortex Perc. cortex per hem.

NO. 1 1277 gr. 720 gr. 56.3%

.. 2 1194 .. 661 .. 55 .3%

.. 3 1152 .. 633 .. 54 .9%

.. 4 1067 .. 562 .. 52.6%

Average

This makes us suppose that the figures found by W AONER and CALORI for the surface are too high, a supposition that is con6rmed by HENNEBERO's 3) work, who determined the surface of the brain in a more exact way. I shall mention HENNEBERO's method below, but already

1) l.c. p . 204. 2) Published in Giacomini : Guida allo studio delle circonvoluzioni cerebrali deU'uomo

Torino, 1884. 3) HENNEBERG. Messung der Oberflächenausdehnung der Grosshirnrinde. Journal für

Psychologie und Neurologie, Bnd. 17, 1910-11.

848

will mention at once that, whereas DONALDSON (l.c. p. 205) on account of W AGNER' s figures finds a surface of 2352 CM2 in a brain of 1360 gr. T. B. W. and CALORI in a brain of 1325 gr. a surface of 2439 CM2 .• HENNEBERG only finds a . surface of 2052 CM2 in a brain of similar weight (1320 gr.). consequently 15 % less I).

Probably the surface measures of HENNEBERG are more exact than those of W AGNER and CALORI. Whereas these authors divided the brain only in hemispheres or lobes, determining the surface hidden in the sulci only indirectly, HENNEBERG measured this hidden surface also directly. by cutting the cortex (af ter fixation of the brain in 10 0J0 for~

maline) in pieces of 1 to 3 cM). In this way all the cortexsurfaces could be reached easily and measured directly.

For this purpose he used pieces of silkpaper. which does not swell in water. and then he measured all the pieces together by lifting them under water from the cortex and putting them on milligraphpaper. As objects he used (one case excepted) only the left hemispheres.

In this way he examined three Hannoverians, one Hottentot. one ' Herero and one Javanese. The cortexweight is calculated by takirig the S.W . = 1.034 and the average depth with HENNEBERG 2) and DANILEWSKY (l.c.) = 2.5.

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I T . B.w·1 Cases Average Surfage Cortex OfoCortex left hem. left hem. left hem. hem.

German I rJ 45 j. _3) 525 gr. 1082 cM2 279.5 53.4%

.. " rJ 22 j . 1510 gr. 655 .. 1240 .. 320 .5 48.9%

.. 111 '( 26 j . 1320 .. 565 .. 1016 .. 262 .6 46.5%

Average .9.6%

Hottentot rJ _3) 615 .. 1119 .. 289.0 47.0%

Herero rJ 1215 .. 530 .. 996 .. 257 .5 48.6%

Javanese c;:' 1230 .. 535 .. 1050 .. 271.5 50.7%

Hence results that HENNEBERG found a still lower average percentage for Europeans (49.6 %) than DONALDSON calculated from the S. W .. but also that in his Germans he found variations from 46.5 % to 53.4 0/0 '

I) TRAM ER who determined the surface of the right hemisphere of a normal woman of 35 years came to a figure still 20 % lower than HENNEBERG found for a woman of 26 years. As, however, TRAMER does not mention the T . B. W . of his woman, this figure is uncontrolable. It seems too small even for an average female T. B. W . of 1250 gr. (HENNEBERG's woman had a high brainweight though, th is being 1320 gr. which however seems to occur more in Hannover; KRAUSE) . See TRAM ER : Messung und Entwicklung der Rindenoberfläche des menschlichen Groszhirns. Arbeiten aus dem neurologischen Institute Zürich. Bnd X, 1916.

2) HENNEBERG himself made his calculations estimating the S . W . = 1.03; I made them using a S. W . = 1.034, this being more exact. The average difference is 0,3 % per hemisphere and 0.12 % per T . B. W .

3) In this case the total brainweight is not mentioned by HENNEBERG.

849

While all these methods of determining the cortexweight are indirect on es and so. probably are more open for errors than a direct method. I made direct weighings of the cortex in the following way:

Brains. hardened in formaline I). were weighed and afterwards halved in the sagittal diameter. Then the cerebellum and the pons were cut off along the corpora quadrigemina posteriora and the hemispheres were weighed. Afterwards these hemispheres were cut by the macrotome of REICHERT. in slices of 2 to 3 m.m. During 3-6 hours these slices were coloured in an aqueous nigrosine solution (1 : 1000). upon which the grey substance marked out almost black against the almost uncoloured white substance.

These slices I put on glass plates and severed the cortex by means of a thin knife. Then I weighed the cortex. af ter all the pieces (cortex and rest of hemisphere) had evaporated so much that the cortex plus the rest represented again the same weight the hemisphere had before being coloured in the nigrosine solution . This drying is necessary. as by their large surface the pieces absorb a large quantity of water.

In total 6 brains (12 hemispheres) were treated in this way : three Dutch and three Chinese brains from the Dutch East-Indies (formaline material).

My results are expressed not only in percentages of the hemisphere weight. but also in percentages of the T. B. W .• the latter giving a more reliable figure for comparison of the left cortex and the right one.

LEPT HEMISPHERES

Cases IT.B.~ Weight r Weight Weight oio cortex oio cortex I. hem. I. cortex I. rest p. hem. p. T.B.W .

Dutch. X. cf' 29 y. 1068 gr. -474.5 gr. 237 gr. 237 . 5gr. 49.99% 22.2 °lc

.. Gr. cf' 42 y. 1375 .. +618 .. +299 .. 319 .. 48 .38% +21.74%

.. Bo. cf' 18 y. 1360'/2 .. - 621 .. 331 .. 290 .. 53 . 3 % 24 .32%

Average I. 50.8 %

Chinese I. NO. 11 1014.5 .. -430 .. +218 .. 212 .. 50.0 % +21.49%

.. I. NO. 10 1344 .. -576 .. +300 .. 276 .. 52.1 % +22.32%

.. I. NO. 12 1425.5 .. -615.5 .. 313.5 .. 302 .. 50.9 % 22.0 010

Average 1.1 151.0 % 1

') That as weil the weight as the volume increase a littJe (± 1010 FLATAU) in formaline. does alter the celation between cortexweight and hemisphereweight only in a very small proportion . The same avails for the aqueous I Oio nigrosinesolution. The inHuence. exer­cised by the formaline -fixation. in my opinion is even smaller than the greater or smaller quantity of blood in the brain. as the S . W . of the blood is considerable and the vascu­larisation of grey and white substance differs much .

850

RIGHT HEMISPHERES

Cases T . B.W. Weight Weight Weight 0/ 0 cortex 0/ 0 cortex r. hem. r. cortex r. rest p.hem. p. T .B.W.

Dutch X.rJ 29y. 1068 gr. +477.5 gr. +238 .5 gr. 239 gr. 49 .94% +22 .33%

.. Gr. rJ 42y. 1375 .. -613 .. 297 .. 316 .. 48.4 % 21.16 %

.. Bo. rJ 18 y. 1360112 .. +622 .. +336 .. 286 .. 54.0 % +24.6 %

Average r. 50.6 %

Chinese I. NO. 11 1014.5 .. +437.5 .. 212.5 .. 225 .. 48.61 % 20.94%

.. I. NO. 10 1344 .. +584 .. 297 .. 287 .. 50 .86% 22.1 %

.. I. NO. 12 1425.5 .. +638 .. +320 .. 318 .. 50.11 % +22.45%

Average r. I I I I

49 .9 ~/o

Average for all hemispheres of the Dutch 50 .65%

.. .. .. .. .. .. Chinese 50.45%

The halving of the brain being never exact the cortexpercentage per hemisphere is influenced by the inexact halving of the brain. whereas we use the same constant. in expressing the cortexpercentage (right and left) in the total brain weight.

Doing so I found the average cortexpercentage per hem is ph ere to be 50.65 Ofo in the Dutch. this being al most 1 Ofo more than the figures. found by HENNEBERG in his Hannoverians. This amount (1 010) however falls within the individual variations in the West European races. showing an upperlimit of 54.0 Ofo.

The percentage in the Chinese. mentioned above differs only 0.2 010 from the Dutch. which is very little. and falls within the errors inevitable in such examinations. This result is in striking contrast to CLAPHAM's 1). who. estimating the depth of the braincortex in Chinese coolies of Hongkong (and of Pelewislanders) concluded : "although destitute of any means of accurately measuring the depth of the grey matter of the cerebral convolutions I am convinced that it was appreciably shallower than in the same structure in the average European" .

As to an eventual difference between the left and right cortex. my results show that once the right. another time the left cortex was. or seemed to be a little heavier. I value this difference the less. because it depends mostlyon absorbed water. If we all ow the pieces of cortex

1) CROCHLEY CLAPHAM. On the brain weights of some Chinese and Pelewislanders. Journalof the Anthropological Society of Great Britain and Ireland. Vol. VII 1878. pag. 92.

851

half a day to evaporate. the difference between right and left becomes a minimal.

Much has been written on an eventual asymmetry. BROCA I) found in the average the right hemisphere of men 1.93 gr.

heavier than the left. in women the same hemisphere 0.03 gr. heavier; BOYD 2). however. says: "it is a singular fact confirmed by the exami~ nation of nearly 200 cases at St. Marylebone in which the hemispheres were weighed separately that almost invariably the weight of the left hemisphere exceeded the weight of the right one by at least the eighth of an ounce". (a very small difference indeed. the eight of an ounce being 3.7 gr. so the difference in an average brain weight of 1360 gramme would be only 0.27 Ofo; DONALDSON).

Dr. THURNAM (quoted by BROWNE) in the contrary again found the right one heavier. According to CRICHTON BROWNE 3) it is a question of age. In 400 lunaties he found the right hemisphere predominating with the exception of women between 40 and 60 and men between 50 and 60. in as much as here the left hemisphere would be a little heavier. He found the greatest differences to exist under 20 years and then always in favour of the right hemisphere. this also being the case with pe-ople older than 70 years.

In contrary to the results above mentioned FRANCESCHI 4) found in

157 male brains in 49 cases the left hemisphere and in 51 the right one heavier. while in 57 cases they practically were of equal weight.

In 144 female brains he found in 43 cases the left hemisphere. in 46 eases the right one heavier and in 55 cases both hemispheres of equal weight (or varying in decimals of grammes). Also W AGNER (l.c.) and BRAUNE 5) sometimes found the left. sometimes the right hemisphere a little heavier.

DONALDSON supposes that. if the brain is always halved in the same position e. g. on the base. the occipital pole directed to the cutter. a righthanded anatomist will allways make the same error.

This personal error. in halving the brain. vanishes in weighing only the cortex. as the cortex of both hemispheres nowhere meets. the cortex of eaeh hemisphere being isolated. So the comparison of the quantity of cortex in the right hemisphere and the left one gives a more trustworthy criterium for an eventual asymmetry. especially if we express both in the same constant. viz. in °10 of T. B. W. The figures found by myself also prove

I) Cited by TOPINARO in his Eléments d'Anthropologie générale p. 582-583. Paris 1885. 2) BOYo. Tables of weight of the human body and its internal organs etc. Philosophical

Transactions of the Royal Society of London. Vol. 151. 1861. 3) CRICHTON BROWNE. The weight of the brain and its component parts in the

insane. Brain Vol. I 1879 p. 516. 4) Bollettino della Societa di Scienze mediche di Bologna 1888. Quoted af ter DONALOSON. 5) BRAUNE. Das Gewichtsverhältnisz der rechten und linken Hirnhälfte heim Menschen.

Arch . f. Anat. und Physiol. 1891.

852

that none of the hemispheres is constantly superior in cortexpercentage. I) . In three cases (marked in my table by t) the right cortex was a little

heavier. in three other cases (marked also by t) the left. HENNEBERG. though exammmg chiefly left hemispheres. has in one case (his European No. 111 with a T. B. W. = 1320 gr.) compared the surfaces of both hemispheres and found on the right 1035 C.M2 .. on the left 1026 C.M2. the result (multiplied with 2.5 as average depth and 1.034 as S. W .) being only a difference of fully five gram mes (0.3 Ofo per T. B. W.) in favour of the right cortex. This figure is within the radius of errors. as is also the average surplus of the right cortex in some of my brains (4.3 gr.) and of the left one in others (3.5 gr.).

My results show at best that the left cortex and the right one never are exactly equal in weight and that once the left and an other time the right one is or seems a little heavier. Nor in animals did I find a con~

stant preponderance (see next page). This accords with the results of FRANCESCHJ. W AGNER and BRAuNE. who too. in the hemispheres. found sometimes the left. sometimes the right one a Iittle heavier.

Moreover my results show that haiving the brain is a very poor way indeed. to determine any superiority of one hemisphere. Whereas of my six brains in one case only. the left hemisphere was heavier. in the other five cases the right one. the cortex determinations show that in three cases the left cortex percentage per T.B.W . was heavier. in the other three the right one. So in two cases (Chinese 11 and 10) in which the left hemisphere seemed to be lighter the cortex was nevertheless heavier.

Here already I should Iike to remark. that the calculation of cortexpercentage per T . B. W. is of value only for the mutual comparison of the right cortex and left one of one and the same cerebrum, not for the cortex percentages of different individuals. as in another paper (see the following issue of these proceedings) I shall prove. that the weight of the cerebellum (being a n important factor also in the T .B.W .) shows very large personal variations. as weil in the Dutch as in the Chinese (from 8 to 12°10 of the T . B. W .).

To conclude some remarks on the cortexweight in animaIs. Nor here direct weighings have ever been made. W AGNER measured the surface of the cortex in an Orang. and found

it to be 533.5 C.M2. and HENNEBERG found in a~ Ateles (with a T.B.W . = 142.5 gr. and a hemisphere weight = 64.5 gr.) a surface of 172 C.M.2 As. however. neither the average depth. nor the S.W. of the cortex is known in these animais. we cannot conclude from these figures to a weight percentage.

Supposing the S. W . in WAGNER 's case also to be 1.034. and the average depth = ~.47 m.M 2). the Drang of WAGNER would have a cortexweight al most equal to the cortexweight of my Drang .

1) This does not eo ipso exc1ude (areal) local weight asymmetries. although 1 doubt their c~nstancy. See however also MELLus. Anat. Récord . V . 1911.

2) According to MARBURG the average depth of the cortex. calculated from the frontal. central. temporal and occipital lobus. is even a Iittle bit larger (2.53 m.m) l.c. p. 585.

853

I weighed the cortex of a Marsupial (Macropus robustus), an Edentate (Choloepus didactylus), an Ungulate (horse), a Carnivore (dog), two kata­rrhine apes (Semnopithecus cephalopterus and Hylobates syndactylus) and an Anthropoid (Orang). The figures found for these animais, are shown in the following tabie.

Examining these figures more closely we observe that, in each of these animals the average cortexpercentage of both hemispheres is as large, or in most cases (Choloepus, Dog, Hylobates and Orang) even larger than in Man, th is being a consequence of the fact (see below) that smaller brains generally have more grey matter than white matter. So we also understand the observation of Miss TAFT, who found in children, microcephalics (and mongoloid idiots) a cortexpercentage, arger than in normals (l.c.) .

Next to this statement two more facts are str*ing in this tabie.

LEPT HEMISPHERES

-

I T . B. w · 1 Anima ls Weight Weight Weight 0/ 0 cortex 0/ 0 cortex l. hem. l. cortex l. rest per hem. p. T.B.W .

Macropus rob. 55 gr. 20 .5 gr .. 10 gr. 10.5 gr. i8.7 % 18 .2 %

Choloepus did. 3i .. 13 . 1 .. 7.1 .. 6 .. 53 .8 % +20.88%

Canls fam . 78 .5 .. 3i.1 .. 21.1 .. 13 .. 61.9 % +26.95%

Equus cab. 510 .. 187.2 96 .. 2 .. 91 "

51.3 % +18.86%

Semnopith . ceph. 65 .. 27 .6 13 .. 1 .. 11 .5 .. i7.3 % 20 . 15%

Hylobates syn. 105 .. 13 .0 .. 25 .. 18 .. 0 .. 58.11% 23 .8 %

Simia satyrus 293 .. 122.7 .. 67 .. 2 .. 55 .. 5 .. 5i .77% 22.87%

RIGHT HEMISPHERES

Animals lT. B. w · 1 Weight Weight Weight I % cortex ~o cortex r. hem. r . cortex r. rest p. hem. p. T. B.W.

Macropus rob. 55 gr. 20.5 gr. 10 gr. 10 .5 gr. i8 .7 % 18 .2 %

Choloepus did. 3i 13 7 6 53 .8 % 20.58%

Canis fam . 78.5 .. 33 20.5 .. 10 .5 .. 62.1 % 26.18%

Canis f. collie 77.7 .. 32 .35 .. 20.0 .. 12.35 .. 61.9 % 25.71%

Equus cab. 510 185 96 89 51.9 % 18.82%

Semnopith . ceph . 65 26 .5 .. 13 .5 .. 13 51.0 % +20 .77%

Hylobates syn . 105 11 25 .5 .. 18 .5 .. 58 % +2i.3 %

Sim ia satyrus 1

293 122.5 .. 69 .5 .. 53 .0 .. 56 .73% +23.61%

854

Firstly the high cortexweight per T . B. W. in Canis is astonishing I). especially in comparison with Semnopithecus. which has a smaller brainweight and a higher standing in the phylogenetic series. With both these facts we should rather expect a larger cortex percentage in Sem~ nopithecus. the larger development of the brain in apes causing the pallium to be developed more than the stem.

That a heavier cerebrum ought to have less cortex relatively than a lighter one (at least in the same group or in related animais) is explained by ERNST DE VRIES 2) and Hovy 3) who showed that. if the brainvolume increases (in related groups). the white matter must increase relatively more than the grey one 'l

N ow dogs and apes are not related but still both are gyrencephalic animals and by the lower standing of the dog as weil as by his larger brainvolume we would expect Canis to have a lower cortexpercentage than Semnopithecus.

This not being the case, but in the contrary Canis having a surprisingly high cortexpercentage. consequently that can be explained only by the large development of the mantle and the depth of the sulci in this anima!.

It would not surprise me. if the relation between the superficial cortex and the hidden cortex (being in man in the average 1 : i and in Drang 1 : 1.56) would be larger in Canis.

That the mande of the brain is exceedingly developed in the first dog. also results from the fact th at his hemispheres amount to 86.8 % of the T. B. W. (in the collie 83.3 %). in my apes the average being 82.9 % only.

A second remarkable fact is the high cortexpercentage in Hylobates. Although the cortex percentage in the Orang as weil as in Hylobates is high. in the latter it is still higher than in the former.

We cannot be surprised that in the Orang the cortexpercentage per T.B.W. is higher than in Semnopithecus. since in anthropoids the mande of the cerebrum is more developed than in Semnopithecus.

That however Hylobates has a higher cortexpercentage than the Orang cannot be explained by its smaller brainvolume alone. since from this standpoint the cortexpercentage in the still smaller brain of Semnopithecus ought to be still higher than in Hylobates. Which is not so.

That Hylobates has a greater percentage than both other apes can be only explained by a larger depth of the cortex in Hylobates than in the other apes mentioned above.

I) A second examination of the right hemisphere of a collie confirmed th is facto which besides is also in harmony with the results of DANILEWSKY (I. c.).

2) ERNST DE VRIES. Das Corpus striatum der Säugetiere. Anat. Anzeiger Bnd. 37.1910. 3) HOVY. On the relation between the quantity of white and grey substance in the

central nervous system. These Proceedings 16. p. 311. 1913 .) The lower cortexpercentage in Equus falls within th is group.

855

This explanation ag rees with the measurements of Marburg I). This author also emphasizes .. dasz die Rindenbr~ite des Hylobates absolut gröszer ist als die des Orangs. letzere aber gröszer als die der anderen untersuchten Affen." (I. c. p . 596).

This is in perfect harmony with my results. At last still one remark. Comparing my results with those of DONALDSON 2) for the Norwegian

rat. we find (I. c. p. 87) the cortexvolume (XX): being 393.1 5 m.m .. the spec. weight = J .05 (l.c. p. 76). the cortexweight to be 0.4128 gr. in the rat. The brainweight in this group being 2.054 gr. (l.c. p. 87). the relation of the T. W . C. to the total brainweight = 20.1 0J0 or in one hemisphere 10.05 0J0 per T. W. B.

This is a much smaller percentage than in my animals and in man. We know. however. that in most gyrencephalic animals more cortex is found in the sulci than on the surface. and ought to consider that the rat is lissencephalic. Were apes and man also lissencephalic. i.e. were there no cortex hidden in the sulci. the cortexpercentage of a hemisphere per T. W. B. would be even less than in the rat and amount to 7 0/0 scarcely. whereas it actually is in man and the higher apes more than 22% per T.B.W.

So also by my figures we may realize the great importance and necessity of fissuration for higher cortical development.

I) MARBURG. Beiträge zur Kenntnisz der Groszhirnrinde der Affen. Arbeiten aus dem Neurolog . Institut in Wien, 1907.

2) DONALDSON. The rat. 2d. ed . Memoirs Wistar Instit. Philadelphia 1924.