a I B HAR.Y OF THE UN IVLR.SITY OF ILLINOIS 610 . no i- 3 KLMOIb. SIORAC Digitized by the Internet Archive in 2011 with funding from University of Illinois Urbana-Champaign http://www.archive.org/details/isocortexofman612bail 41 I The Isocortex ofMan By PERCIVAL BAILEY and GERHARDT VON BONIN Urbana, 1951 UNIVERSITY OF ILLINOIS PRESS ILLINOIS MONOGRAPHS IN THE MEDICAL SCIENCES ' is a general title used to comprehend a series of contributions from the fundamental and clinical departments of the Chicago colleges of the University—College of Medicine, College of Dentistry, and College of Pharmacy. Each issue is an independent publication. For the convenience of bibliographers and librarians, each year's output is called a volume. Each volume consists of approximately 450 pages, priced at four dollars (by subscription) for paperbound copies. A volume may consist of one, two, three, or four individual monographs. Prices of individual numbers are indicated in the list of titles given on the back cover. Requests for exchanges should be addressed to the Exchange Department, University of Illinois Library, Urbana, Illinois. All communications concerning sale or subscription should be addressed to the Director, University of Illinois Press, Urbana, Illinois. Volumes I, II, and III of this series were issued under the title Illinois Medical and Dental Monographs. THE ISOCORTEX OF MAN ILLINOIS MONOGRAPHS IN THE MEDICAL SCIENCES: Vol. VI NoS. 1, 2 <' -*5 8 s o S 2 S u - 3 ° o S- § "3 CO 1 •45 —>> n3 CD £ 3 2= Pi & cu i >> O p, CO =2 _g t- CD 3 + bo >, + ^ + + _c M CD | "3 C 3 3, >1 '3,3 II II II II CD CD tar P a "3, H is § Li a O CD 3 t- - - >— fcOWO S~ Qi X CD X a. +3 CD e h += .< O t- O CD o ^ X ji CD <u o> X ll 00 -f^ - qj cd — m a -2s c o eg tn •--; m CD h ° s O be s '3 o3 O c3 CD o 2 j<! O CD CD 3 3 tf >h pq &h o H B CEC fc K I E- -S 3 3 » Mi. O o be c o £r pj >h CQ o The Isocortex ofMan By PERCIVAL BAILEY and GERHARDT VON BONIN Urbana, 1951 UNIVERSITY OF ILLINOIS PRESS Copyright 1951 by the university of Illinois press, printed in the united states of America. Board of Editors: a. c. ivy, o. f. KAMPMEIER, ISAAC SCHOUR, AND E. R. SERLES. >C/ REMOTE STORAGE To the memory of PAUL FLECHSIG * (4 Therefore the surgeon in seeking a thoroughly reliable knowl- edge must duly -prepare a dead body. .It should be the body of someone who had neither been excessively old, nor one who died of poison, nor of a protracted disease. Susruta (III, 5) PREFACE It is safe to say that interest in cortical architecture has not been widespread since the appearance of Brodmann's Vergleichende Lokalisationslehre der Grosshirnrinde in 1909. Brodmann's contention (p. 305), frequently reiterated by the Vogts, that each cortical area is an organ in itself, tied up cortical architectonics with a theory of cortical function that threatened, under the influence of holistic views in psy- chology and biology, to become more and more outmoded. The minute panellation of the cortex by the Yogt school, moreover, made this theory less and less tenable. The subject became not only difficult and abstruse, but doubts about the re- liability of cytoarchitectonics grew until, "Unless the criteria are clearly stated and objectively verifiable . architectonic charts of the cortex represent little more than the whim of the individual student," could be written, amidst widespread applause, by Lashley and Clark (1946). In 1927, G. Elliot Smith, who twenty years previously had published a brain map of his own, in reviewing the work of Economo and Koskinas, wrote: "It can be confidently said that such a survey will not have to be done again, and that this work will become a standard treatise of reference on the topography of the human cerebral cortex. No doubt in the future much detailed work will be done upon particular regions, perhaps introducing small points of modification, but the work as a whole is bound to stand as a permanent achievement." The work is a standard reference and a permanent achievement, yet there seem good reasons why the human cerebral cortex should be investigated once more, why it would be wr orth a considerable outlay of time and money to go over a ground that seemingly has been worked over so carefully. The number of brains that have actually been examined is not very large, nor was the state of their preservation always as good as one might have wished. Campbell (1905), by his own account, examined three hemispheres completely for nerve cells and nerve fibers, and two others incompletely. Brodmann never gives precise information about his material. The Brain Institute under Vogt's direction possessed, when Brockhaus (1940) pub- lished his paper, at least seven hemispheres. We find, in the last two volumes (49, 50) of the J . f. Psychol, u. Neurol, mention of eleven normal human brains serially sectioned for anatomical study. These brains were removed about ten hours postmortem (C. & O. Vogt, 1942) and put into a 10 per cent neutral formalin solution. When it is realized that one brain was not fixed until twenty-four hours after death, and when one of Vogt's pupils (Strasburger, 1937) mentions that the staining occasionally had to be corrected by means of a camel's-hair brush, one's desire for corroboration grows. Economo and Koskinas evidently had three brains, but took "most" of their photographs from one and the same hemisphere. Their material was fixed by immersion into formalin at some unstated time after death. Only for the occipital lobe (Economo, 1930a) the supratemporal plane (Economo and Horn, 1930; Kakeshita, 1925) and a few other areas do we have more plentiful information except for that which is contained in the studies of the Brain Institute of Moscow (only the first four volumes of which were available to us), difficult to — viii The Isocortex of Man obtain and in Russian. According to Blinkow and Poliakov (1938) the Moscow Institute possessed at that time "over 30 series of complete microscopic sections through the entire brain hemispheres of an adult." Whatever might be the meaning of this ambiguous statement, the study of yet another brain, properly fixed and stained, seemed desirable in any event. The importance of proper material for anatomical study was known already to the ancient Hindus. Through the kindness of Colonel J. E. Ash, then Director of the Armed Forces Institute of Pathology, we came into possession of a particularly perfect brain removed from a young healthy adult male, who died of an accident without injury to the head. The brain was removed within one hour of death after perfusion through the arteries with a 4 per cent aqueous solution of formaldehyde; it weighed, after fixation, 1,347 gm.; and will be referred to henceforth by the sym- bol H. Subsequent study proved the brain to be uniformly fixed throughout. The unsuitability of the brains removed many hours after death, and fixed in the usual way by immersion in formalin, should be evident to anyone, and the histological preparations made from them show clear evidences of pathological alteration. We might cite, as an example, the drawing of the cortex given by Ram6n y Cajal (1911, tome II, Fig. 334, p. 522). It could serve well, without alteration, in a textbook of neuropathology to illustrate severe acute degenerative changes of the ganglion cells. The superiority of brains immediately injected through the arterial system over those removed after the body has been kept in refrigeration for twenty- four hours, or even those removed immediately and immersed in the fixing fluid, has been recognized by M. Rose (1929), Bartelmez and Hoerr (1933), Barnard (1940), and many others. The immediate reason why we decided to embark on this renewed investigation of the human cerebral cortex is found in our previous work on macaque and chim- panzee. That work was undertaken to lend precision to the results of physiological neuronography performed on these primates and to be applied, so it was hoped, to man. But, in the process of our studies, it became clear that the precise homology of which we had dreamed could be achieved only when maps of all three primates man, chimpanzee, and macaque—had been drawn up by the same observers. As our work on cytoarchitectonics progressed, it became increasingly apparent to us that more than local patchings were required, as Lashley and Clark have also found. What was needed was to state as objectively as possible those architectural types that can be clearly distinguished, but to refrain from giving quite secondary, obviously unimportant, details the same critical value. Anybody can see, to give an example, the difference between Brodmann's areas 17 and 18. But the differences between his 18 and 19 are quite tenuous and very difficult to recognize. To draw a map on which these three areas are given three different markings—such as dots, cross-hatchings, and broken lines—is to create an entirely misleading impression. Useful as such maps are for the description of corticocortical connections, they do not translate accurately cytoarchitectonic data. The frontispiece of this monograph is drawn on a different principle (see opposite title page) from those used for the macaque (Bonin and Bailey, 1947) and chimpanzee (Baile}', Bonin, and McCulloch, 1950). In those maps colors were used mainly to facilitate the portrayal of regions of transitional structure and many areas were distinguished largely for the purpose Preface ix of describing the findings of the method of physiological neuronography without much regard for degree of differentiation.