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Famous Russian brains: historical attempts to understand intelligence

Alla A. Vein, Marion L. C. Maat-Schieman
DOI: http://dx.doi.org/10.1093/brain/awm326 583-590 First published online: 8 January 2008


Russian scientists are certainly among those who contributed actively to the search for the neuroanatomical basis of exceptional mental capacity and talent. Research into brain anatomy was one of the topics of special interest in various Russian universities. A number of independent reports on the study of famous Russian brains appeared both in Russia and abroad. Collecting and mapping brains of elite Russians in a structured manner began in Moscow in 1924 with the brain of V. I. Lenin. In 1928, the Moscow Brain Research Institute was founded, the collection of which includes the brains of several prominent Russian neuroscientists, including V. M. Bekhterev, G. I. Rossolimo, L. S. Vygotsky and I. P. Pavlov. The fact that the brain of two of the most outstanding scholars of Russian neurology and psychiatry, A. Ya. Kozhevnikov (1836–1902) and S. S. Korsakov (1854–1900), have been studied is largely unknown. A report of the results of this study was published by A. A. Kaputsin in 1925 providing a detailed neuroanatomical assessment of the brains. A considerable weight, a predominance of the left hemisphere and a particularly complex convolution of the frontal and parietal lobes of both brains were reported, the assumption being that these brain parameters can serve as an indicator of mental capacity. The names Kozhevnikov and Korsakov are among those most cherished by Russian neuroscientists; they are also familiar to Western colleagues. The (re)discovery of the records of the brain autopsies is meaningful, maybe not so much from a neuroanatomical point of view as from a historical perspective.

  • neuroanatomy
  • elite brains
  • intelligence
  • Russia


‘Periculosum est credere et non credere’ – it is dangerous to believe as well as to disbelieve.’                  Blaise Pascal The search for the biological roots of extraordinary capacity has been going on for many centuries. From the 18th through the late 20th century, there was particular interest in the brain and its remarkable convolutions. Generations of prominent scientists, from Flechsig and Retzius to Vogt and Economo, were advocates of a structural–morphological approach to brain research (Hagner, 2004). Russian scientists are certainly among those who contributed actively to the search for the neuroanatomical basis of exceptional mental capacity and talent. Conventionally, it is thought that collecting and mapping the brains of famous Russians began in Moscow in 1924, starting with V. I. Lenin's brain and the foundation in 1928 of the Moscow Brain Research Institute (Spivak, 2001; Richter, 2007). In fact, it began much earlier in the universities. Collections in medical faculties included (among those of other celebrities) the brains of medical professors, many of whom had bequeathed this organ for scientific purposes. This was a tradition not unknown in other European countries and in the 19th century it was quite a respectable thing to do. Many eminent men in Europe willingly acceded to this service to science by giving permission for their brains to be removed, weighed and studied (Gere, 2003; Taylor, 1996). The systemic investigation of brains of geniuses started in Europe after 1830. The main parameters for brain examination were weight and patterns of gyral convolutions, an approach introduced in 1836 by the French psychiatrist, J. B. M. Parchappe (1800–67). He emphasized ‘exact measurement’ as opposed to the vague inspections of phrenologists (Hagner, 2003). In the 1850s, Rudolf Wagner (1805–64) carried out a methodological study of the brains of deceased Göttingen professors and this became the actual starting point for the dissection of the brains of numerous eminent scientists, writers, artists and politicians (Bentivoglio, 1998; Hagner, 2003). In 1876, French scientists founded the Mutual Autopsy Society of Paris (Societé d’autopsie mutuelle) in order that their donated bodies would be dissected by other members, and that psychological and biological characteristics would be correlated (Hecht, 2003; Dronkers et al., 2007). In 1901, during the International Association of Academies in Paris, the initiative was taken to establish a Brain Commission after which Brain Institutes appeared in Europe (Madrid, Leipzig, Vienna, Zurich, Frankfurt/Main, Budapest), in the USA (Philadelphia) and also in St Petersburg (Richter, 2000; Hagner, 2004).

The fact that the brains of two of the most outstanding scholars of Russian neurology and psychiatry, A. Ya. Kozhevnikov (1836–1902) and S. S. Korsakov (1854–1900), were studied is largely unknown. This paper will deal with the history and the results of this work in the context of a brief history of the study of elite brains in Russia.

Interest in brain anatomy in Russia

In Russia there was always a vivid interest in anatomy; in fact the very first state public museum was The Kunstkamera in St Petersburg founded by Tsar Peter the Great (1714), displaying some 2000 anatomical specimens purchased in Holland (Gokhman and Kozintsev, 1980). Research into brain anatomy was one of the topics of special interest being performed in various Russian universities including Moscow, St Petersburg, Kiev, Kazan and Tomsk, etc. In 1783, the Anatomical Museum of Moscow University was founded. D. N. Zernov (1843–1917), a Moscow University professor, who made a huge contribution to the Anatomical Museum by creating a vast collection of brain preparations. He produced the best classification of fissures and sulci of the brain for the time. He also demonstrated the absence of differences in the structure of the brains of representatives of different nationalities and races and was a fervent opponent of Cesare Lombroso (1836–1909) (Zernov, 1887; Sapin, 1986; Etingen, 1997). One of the most prominent scientists in this field was V. A. Betz (1834–94), professor at Kiev University. He described a particular type of pyramidal neurons located within the fifth layer of the primary motor cortex, which were later named after him—Betz cells. Betz introduced a new principle of cellular structure in relation to the laminar division of the cerebral cortex and marked the beginning of research into the cytoarchitectonics of the cerebral cortex (Sapin, 1986; Richter, 2007). Along with numerous centres, Moscow and St Petersburg (psycho)neurological schools maintained a leading position in anatomical research of the brain. The Moscow neurological school was founded in 1869 by A. Ya. Kozhevnikov (Fig. 1). He enjoyed a broad range of scientific interests, though his early work was primarily in the fields of neuroanatomy and neuromorphology, concentrating on amyotrophic lateral sclerosis, aphasia, myasthenia, familial spastic paralysis and cysticercosis of the brain (Vein, 2007). Kozhevnikov gathered a—for that time—unique collection of anatomical, histological and comparative anatomical specimens of the nervous system. These served as the basis for the Neurological Museum that opened in 1892 (Grashchenkov, 1960; Archangel'skii, 1965). The psychoneurological school in St Petersburg was dominated by the authority of V. M. Bekhterev (1857–1927). He authored fundamental works on anatomy and physiology of the brain. Bekhterev had formulated and popularized the idea that man should be regarded as ‘a single whole,’ ‘a biosocial entity’ whose understanding requires the study of human consciousness and psychology. From his attempts to combine research in neurology, psychiatry, neurophysiology, neuroanatomy, neuropsychology, neurosurgery, etc., one might consider him to be the founder of the multidisciplinary approach to brain exploration, to which end a Psychoneurological Institute was founded in St Petersburg in 1907 (Akimenko, 2007).

Fig. 1

A. Y a. Kozhevnikov (1836–1902) in the audience of Moscow Clinic for Nervous Diseases.Photo taken March 19, 1898—photo from Historical Museum of Moscow Medical Academy (with permission).

Russian elite brains

Bekhterev was interested in correlating the special features of a brain and the brilliant qualities of its owner. An example of this approach was the study of the brain of the eminent Russian chemist, the creator of the periodic table of elements, D. I. Mendeleev (1834–1907) (Fig. 2), first reported at the scientific session of the Psychoneurological Institute (May 1, 1908) and published in 1909 (Bekhterev and Weinberg, 1908; Bechterew & and Weinberg, 1909). The authors presented an exceedingly detailed description of Mendeleev's brain, weighing 1570 g (Fig. 3), the main conclusion being that there was a strong development of the left frontal and parietal lobes compared to the rest of the brain. Interestingly, the authors compared his brain with those of two famous Russian musicians which were available to them—the composer A. P. Borodin (1833–87), and the pianist, composer and conductor, A. G. Rubinstein (1829–94). The anterior part of the left gyrus temporalis superior of Mendeleev's brain was far less developed in comparison with this region in the brains of these musicians. According to the authors, this was a sign of his modest musical capacity (Bekhterev and Weinberg, 1908). The brains of Borodin and Rubinstein are still preserved in the Anatomical Museum of the Military Medical Academy in St Petersburg (Etingen, 1997).

Fig. 2

D. I. Mendeleev (1834–1907). Photo from Bechterew W. von, Weinberg R. Das Gehirn des Chemikers D. J. Mendelejew. In: Roux W, editor. Anatomische und entwicklungsgeschichtliche monographien. Heft 1. Leipzig: Verlag von Wilhelm Engelmann; 1909. p 22.

Fig. 3

Brain of D. I. Mendeleev. Photo from Bechterew W. von and Weinberg R. Das Gehirn des Chemikers D. J. Mendelejew. In: Roux W, editor. Anatomische und entwicklungsgeschichtliche monographien. Heft 1. Leipzig: Verlag von Wilhelm Engelmann; 1909. p 22.

There are several other independent reports on the study of famous Russians brains both in Russia and abroad. On dissecting the brain of the legendary general Skobelev [M. D. Skobelev (1843–82) was a Russian general famous for his conquest of Central Asia and his heroism during the Russian–Turkish War of 1877–78], Zernov reported finding no extraordinary features in the convolutions (Zernov, 1887). There were reports on the brain of the Russian novelist, Ivan Turgenev (1818–83), the weight of which reached an incredible 2021 g, and of the mathematician, Sofia Kovalevskaya [Sofia Kovalevskaya (1850–91) was the first major Russian female mathematician and a student of Karl Weierstrass in Berlin. In 1884, she was appointed professor at Stockholm University, the third woman in Europe to become a professor], the first famous woman whose brain was examined (Retzius, 1900; Bentivoglio, 1998). One-half of the brain of a leading Russian satirical writer of the 19th century, M. E. Saltykov-Schedrin (1826–89), the brains of two eminent Russian scientists N. N. Zinin (1812–80) and V. V. Pashutin (1845–1901); and the brains of the poets, H. Tumanyan (1869–1923) and V. Bryusov (1873–1924), were examined in 1915 and in 1924–25, respectively, and the results reported (Smirnov, 1915a, b; Etingen, 1997; Spivak, 2001). In the years immediately following the Bolshevik Revolution, some very daring, almost reckless, scientific projects were financed. Soviet scientists promised a victory over sleep, old age and even death. Various efforts were undertaken to ‘diagnose genius’. During his lifetime, V. I. Lenin (1870–1924) had been considered a genius. The undertaking to perform a detailed examination of his brain was launched immediately after his death. The work was started in 1925 by the famous German neuroanatomists, Oskar and Cécile Vogt, who analysed the brain's cytoarchitecture, based on the discovery made by the Russian neuroanatomist, Betz (Richter, 2007). All Russian participants in this project were representatives of the Kozhevnikov neurological school and associates of Moscow University Neurological Institute.

In 1925, the director of the Moscow University Neurological Institute, Professor G. I. Rossolimo (1860–1928), was honoured for his 40 years of work. Afterwards, a massive volume of collected works of the participants of this meeting was published with papers in Russian, French or German, containing sections on various topics, including social psychoneurology; morphology; physiology; psychology; pathology; clinical neurology, etc.

Among the published papers, there were two reports on the study of the brains of six prominent professors of Moscow University, which were kept in the collection of the Neurological Institute (Kapustin, 1925; Gindze, 1925a, b). With the permission of Rossolimo, A. A. Kapustin reported on the study of the brains of the founders of Russian neurology and psychiatry, Kozhevnikov and Korsakov. In 1926, the same report was published in the journal Clinical Archive of Genius and Talent (of Europathology) (Kapustin, 1926; Sirotkina, 2002).

Description of the brains of Kozhevnikov and Korsakov

Alexey Yakovlevich Kozhevnikov

Kozhevnikov is mainly known for his description in 1894 of Epilepsia partialis continua, or so-called ‘Kozhevnikov syndrome’ (Kozhevnikov, 1894). He was the head of the first Russian independent Department for Neurological and Mental diseases (1869) and later (1890) the very first specialized Clinic for Nervous Diseases in Russia and Europe. Kozhevnikov was the author of the first textbook of neurology in Russia in 1883. The creation of the Moscow Neurology School may be considered Kozhevnikov's greatest achievement. An elite core of Russian neurologists and psychiatrists gathered around him, including S. S. Korsakov, V. K. Rot, G. I. Pribytkov, V. A. Muratov, G. I. Rossolimo, L. S. Minor, L. O. Darkshevich and many others. During the last 3 years of his life, Kozhevnikov suffered from prostate cancer. He died on January 10, 1902 (Lisitsin, 1961).

Brain of Kozhevnikov

[This is a short literal translation from Russian by one of the authors (A.A.V.)]

On dissection, Kozhevnikov's brain (Fig. 4) weighed 1520 g. All measurements revealed a noticeable dominance of the left hemisphere over the right one. There were a huge number of ‘sulci’ of the third category in the regions of the frontal lobes and part of the parietal lobes. In addition to the basic sulci, including the sulcus frontalis medius (Eberstaller), sulcus frontalis obliquus (Marchand), sulcus frontomarginalis (Wernicke) and sulcus supraorbitalis (Broca), the left frontal lobe showed more than 20 small sulci creating a very complicated convoluted aspect. Many small sulci (up to 30) were located in the region of the right frontal lobe. Similarly, in the region of the parietal lobes a large number of small sulci could be distinguished. The angulus Rolandicus of the right hemisphere was equal to 70° and that of the left hemisphere 75°. When measured along the fissura pallii, the distance from the frontal pole of the right hemisphere to the occipital pole equalled 29 cm, and the distance from the frontal pole to the sulcus Rolandicus was 16.5 cm. Accordingly, the length of the frontal lobe represented 56.8% of the right hemisphere. For the left hemisphere these measurements amounted to 30 and 17.5 cm, respectively, thus the length of the left frontal lobe was 58.3%, superior to that of the right one, i.e. the left frontal lobe exceeded the right one in length by 1.5%.

Fig. 4

Brain of A. Ya. Kozhevnikov. Photo from Clin Arch Genius Talent (of Europathol) 1926; 2: 107–14.

Sergey Sergeievich Korsakov

Korsakov (Fig. 5) was one of Kozhevnikov's most outstanding pupils. He was 33 years old when Kozhevnikov appointed him director of the psychiatric division of his department, thus making him the first professor of Psychiatry in Russia. Korsakov was the author of numerous works in psychiatry, neuropathology, forensic medicine and a textbook on psychiatry. He studied the effects of alcoholism on the nervous system and described alcoholic polyneuritis with distinctive mental symptoms (‘cerebropathia psychica tokaemica’), later called ‘Korsakov's syndrome’. He was the first to produce a clear description of paranoia. Korsakov was among the leaders of more humane patient management by applying no-restraint principles. Until his premature death, he was the head of the Moscow University Clinic of Psychiatry, and is considered to be the founder of the Moscow psychiatric school (Ovsyannikov and Ovsyannikov, 2007).

Fig. 5

S. S. Korsakov (1854–1900). Photo from Historical Museum of Moscow Medical Academy (with permission).

After two heart attacks at the age of 44, Korsakov consulted a specialist in Vienna in 1898. Hypertrophy of heart associated with obesity and myocarditis was established. Korsakov died from heart failure at the age of 46 (Banshchikov, 1967).

Brain of Korsakov

On dissection, Korsakov's brain (Fig. 6) weighed 1603 g. At a second measurement on January 26, 1924, the weight was 1355 g. The angulus Rolandicus of the right hemisphere equalled 80° and that of the left 85°. On all measurements, a noticeable superiority of the left hemisphere was observed. In the region of the left frontal lobe, 25 small sulci were discernible in addition to the four main gyri. The fissura centralis anterior of the right frontal lobe was interrupted by more than 30 small fissures. The surface of the parietal lobes showed a similar complexity and the same peculiarities of the configuration of the sulci and gyri. The distance from the frontal pole to the occipital pole along the fissura pallii was 27 cm and the distance from the frontal pole to the sulcus Rolandicus along the same line equalled 15 cm. Accordingly, the length of the frontal lobe represented 55.5% of that of the right hemisphere. For the left hemisphere, the same distances were 28 and 16 cm, respectively. Thus, the length of the left frontal lobe amounted to 57.1% of that of the left hemisphere. The length of the left frontal lobe exceeded that of the right one by 1.6%.

Fig. 6

Brain of S. S. Korsakov. Photo from Clin Arch Genius Talent (of Europathol) 1926; 2: 107–14.

The author concluded that both brains had common features, including considerable weight, predominance of the left hemisphere, and complicated convolutions of the frontal and parietal lobes. There was speculation on the correlation between intellectual capacities and size and structure of the brain.

Pantheon of brains

In 1927, Bekhterev came up with a plan to organize ‘The Pantheon of Brains’ in Leningrad in order to collect elite brains (Vol’fson, 1928; Richter, 2007). It was a severe irony of fate that precisely when the question about creating the Pantheon had been positively solved, the very initiator of this creation, Bekhterev, suddenly passed away. The circumstances are still questionable. On December 17, 1927, the First All-Union Congress of Neuropathologists and Psychiatrists was held in Moscow. Bekhterev, along with L. S. Minor and G. I. Rossolimo, was elected as honourable chairmen of the congress. On December 23rd, the last day of the congress, Bekhterev gave a presentation during the afternoon session. In the evening, symptoms of a gastrointestinal disorder started and 24 hs later, Bekhterev died of (as officially stated) acute heart failure. Without any further post-mortem pathoanatomical investigation, his brain was removed, in accordance with his will, and his body was cremated the next day (Lerner et al., 2005). However, the idea did not fade away. In 1928, the neuroanatomical laboratory of Vogt and his Russian colleagues were reorganized into the Moscow Brain Research Institute (Fig. 7), where the structured collecting and mapping of the brains of famous Russians started. Bekhterev did not see his plan come to fruition, but his own brain enriched the collection of the Moscow Institute (the weight of his brain was 1720 g) (Spivak, 2001). The collection acquired the brains of Soviet politicians, famous writers, poets, musicians, etc. It is not surprising that these included the brains of prominent Russian neuroscientists, such as neurologist, G. I. Rossolimo (1860–1928)—1543 g; physiologist, I. P. Pavlov (1849–1936)—1517 g; neurologist, M. B. Kroll (1879–1939)—1520 g; psychiatrist, P. B. Gannushkin (1875–1933)—1495 g; psychologist, L. S. Vygotsky (1896–1934) (Bogolepova, 1993). During the Soviet period, the work of the Moscow Brain Research Institute continued behind closed doors. The collection was still expanding as recently as 1989, when it acquired the brain of A. D. Sakharov [A. D. Sakharov (1921–89) was an eminent Soviet nuclear physicist, dissident and human rights activist. He was an advocate of civil liberties and reforms in the Soviet Union. He was awarded the Nobel Peace Prize in 1975]—1440 g (Spivak, 2001).

Fig. 7

Museum of the Moscow Brain Research Institute (photo A. A. Vein, 2007).


Since the 19th century, scientists have attempted to establish why particular brains are especially productive. The results were unsatisfactory, none of the studies revealing a strong relationship between brain size or structure and function (Bentivoglio, 1998). Rather than being abandoned, however, this approach achieved high popularity in Russia, as well as in Europe and the USA.

The names of Kozhevnikov and Korsakov are among those most cherished by Russian neuroscientists. Kozhevnikov was the founder of the main journal of neurology and psychiatry in Russia—Zhurnal Nevropatologii i Psikhiatrii (1900). He named the new journal after his former student Korsakov, who died at a young age that same year. These two scholars are considered to be the ‘fathers of Russian neurology and psychiatry’. The (re)discovery of the records of the autopsies of their brains is, therefore, meaningful, perhaps not so much from a neuroanatomical point of view as from a historical perspective.

Although there are no records to this effect, it is quite plausible that it was the wish of Kozhevnikov and Korsakov to donate their brains to scientific study. When the detailed description of the gross anatomy of the brain of these outstanding scientists was published, similar works appeared in Russia, which had the same descriptive approach. This method was broadly used in Russia, Europe and the USA (Bekhterev and Weinberg, 1908, Smirnov 1915a, b; Gindze, 1925a, b; Hagner, 2004). Kapustin, along with other Russian neurologists, was presumably using two sources: Zernov's (1877) classification of brain anatomy and ‘The anatomy of the central nervous organs in health and in disease’ by H. Obersteiner, translated into Russian in 1888 and widely consulted (Obersteiner, 1888). In Russia, Basle Nomina Anatomica remained standard terminology until 1955 (Sapin, 1986). Most cerebral structures are easily recognizable, though there are few terms, e.g. ‘angulus Rolandicus’, which are difficult to trace nowadays. In the case of the brains of Kozhevnikov and Korsakov, their considerable weight, predominance of the left hemisphere and particularly complex convolutions of the frontal and parietal lobes were reported. The conclusion that these features were unusual was based on comparisons with other studies of weight and gross anatomy of the brain published in Russia and abroad. The author of the paper praised the achievements and the intellectual abilities of both scholars and subsequently made the assumption that the size and complexity of their brain anatomy could serve as an indicator of their mental capacity. These data were to a certain extent compatible with other reports on the brains of prominent scientists. The celebrated Swedish anatomist, G. M. Retzius (1842–1919), described unusual ‘secondary’ gyri in the frontal cortex and exceptional growth near the posterior part of the Sylvian fissure of the brains of outstanding scientists, including Kovalevskaya (Retzius, 1900; Finger, 1994). The American anatomist, E. A. Spitzka (1876–1922), who published the description of the brains of 137 famous individuals, showed that eminent people from ‘the exact sciences’ had the heaviest brains (Spitzka, 1905). The complex convolution of the frontal and parietal lobes of the left hemisphere was also described in scientists such as Kant (1694–1778), Gauss (1777–1855), Helmholtz (1821–94), Mendeleev (1834–1907), Haeckel (1834–1919), Lombroso (1835–1909), Gyldén (1841–96), Giacomini (1841–98) and von Monakow (1853–1930) (Bekhterev and Weinberg, 1909; Gindze, 1925a, b; Finger, 1994; Hagner, 2004). The data, however, lacked uniformity. As long ago as 1887, Zernov gave a visionary warning to avoid prejudice when examining the brains of the famous, an opinion close to the conclusions reached by Rudolf Wagner in his pioneering work (Zernov 1887, Finger, 1994). In 1925, another professor at Moscow University, Gindze, pointed out that both in Russia and abroad, descriptions and categorizations of the brains of gifted individuals were highly inconsistent (Gindze, 1925a). Nevertheless, the interest in this problem was not abandoned completely. Thus, there was a great deal of scientific and public interest in the anatomical structure of Albert Einstein's brain. The posterior parts of the parietal lobes of Einstein's brain were extensively developed, but none of the other neuroanatomical features of his brain proved to be exceptional (Diamond et al., 1985; Witelson et al., 1999; Hagner, 2004). At the end of the 20th century, there was still considerable interest in the macroanatomical investigation of the brain. Zilles et al. (1988) analysed the degree of cortical folding, using a gyrification index (GI), which happened to be greatest in the prefrontal and the parieto-temporo-occipital association cortex in human. The interaction between association cortices within parietal and frontal brain regions was suggested to be essential in understanding the biology of intelligence (Jung and Haier, 2007). The work of the Moscow Brain Research Institute continued, although hardly any results of the research on elite brains were revealed. It was not until the 1990s that a series of publications from the Institute appeared describing the brains of gifted individuals (In most of the Russian papers of the 90s the gifted individuals are coded, without identifying the names). Although no difference was demonstrated in brain weight, the cytoarchitectonics of some cortical fields was reported to be more complicated in the group of gifted individuals (Bogolepova, 1993, 1996; Bogolepova and Bogolepov, 1997; Bogolepova and Malofeeva, 2004). Moreover, recent publications on brain size and cognitive abilities have appeared revealing some positive correlations (van Valen, 1974; Gibson, 2002). A post-mortem study of the brains of 100 non-neurological cancer patients showed positive correlations between verbal intelligence and cerebral volume in right-handed men and women as well as in non-right-handed women (Witelson et al., 2006). Tisserand et al. (2001) report smaller head size to be associated with lower intelligence, lower general cognitive functioning and slower speed of information processing though no relation was found between head size and memory function. A recent meta-analysis of 37 neuroimaging studies demonstrates a consistent relationship between brain volume and intelligence (McDaniel, 2005).

Nevertheless, there is still no competent explanation for how the brain provides for exceptional mental capacities. This, therefore, remains a subject of interest, not only from a historical perspective.


The author of this paper (A.A.V.) has recently (2007) undertaken an attempt to trace the dissected brains of Kozhevnikov and Korsakov. Requests and personal visits were made to different departments of Moscow Medical Academy (previously Moscow University Medical Faculty): Historical Museum, Department of Anatomy and Anatomical Museum, Clinic for Nervous diseases, Clinic for Mental (Psychiatric) diseases as well as to the Institute of Brain Research and its museum. Sadly, no traces of these preparations could be found; in all probability, they are lost forever.


The authors are extremely grateful to Prof J. Voogd for his critical advice and help in the preparation of this manuscript.


  • Abbreviations:
    gyrification index


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