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®Rifiin<Tl Articles Anb Clinical Cases [JULY, 1912.] BEAIN PART I., VOL. 35. ®rifiin<tl Articles anb Clinical Cases. THE CYTO-AECHITECTUEE OF THE CEREBRAL CORTEX OF A HUMAN FCETUS OF EIGHTEEN WEEKS. BY JOSEPH SHAW BOLTON, M.D., D.Sc, F.R.O.P., AND • Downloaded from JOHN MURRAY MOYES, M.B., CH.B., D.P.M. PAGE I.—INTRODUCTION ... ... ... ... ... ... ... 1 II.—MATERIAL AND METHODS ... ... ... ... ... 3 by guest on January 7, 2015 III.—ANATOMICAL DESCRIPTION OP THE CEREBRUM ... ... ... 5 IV.—THE LOCALIZATION OF THE BETZ-CBLL AREA ... ... ... 8 V.—THE LOCALIZATION OF THE VISBO-SBNSOBY AREA... ... ... 10 VI.—THE CyTO-AKCHITECTURE OF— (1) The Precentral (Betz-oell) Cortex ... ... ... ... 12 (2) The Post-central Cortex ... ... ...• ... ... 15 (3) The Cortex of the Visuo-sensory Area ... ... ... 20 (4) The Cortex of the Gyrus Cinguli ... ... ... ... 21 (5) The Cortex of the Anterior Frontal Region... ... ... 22 VII.—SUMMARY ... ... ... ... ... ... ... 24 I.—INTRODUCTION. OP late years much progress has been made in the study of the cyto-architecture of the cortex cerebri and of cerebral function. In 1905 and 1907, Campbell [6] and Brodmann [4] respectively'published complete maps of the various areas into which the human adult cerebral cortex may be divided, and further details together with similar maps dealing with the cortex of certain orders of the mammalia have been published both by them and by other workers, notably by Mott [9], Mott and Kelley [10], Mott and Schuster [11 and 12], Watson [14]. As far as we are aware, however, no systematic study of the early evolution of the cyto-architecture of the human cortex cerebri has hitherto been attempted. DRAIN VOL. XXXV. / If L^ 1 2 . OBIGINAL AETICLES AND CLINICAL CASES On different occasions during the past twelve years, reference has been made by one of us (J. S. B.) to the characteristic features of the cortex of foetuses and infants, and to the manner in which this structure becomes laminated and develops. In 1900 [1] a description was given of the cortex of the visual area (both visuo-sensofy and visuo-psychic) of a child of 3 months, and of a case of anophthalmos of the age of 1 month. In both these cases the visuo-sensory area was mapped out. It was shown that inJthe case of the former the visuo-sensory area was relatively large in extent, though normal in distribution, whilst in the latter this area was deficient both in extent and in distribution, and its cortex exhibited a similar thinning of the outer layer of granules (Layer Ilia), and of the line of Gennari (Layer Illb), to that, which occurs in cases of long-standing blindness. It was further shown that an almost exact correspondence obtains between the thickness of the conjoined first and second layers of the cortex and the existing degree of amentia or dementia. • In 1903 [2] a description was given of the mode of development of Downloaded from the cortex of the prefrontal area, based on observations made on foetuses of different ages, stillborn children and infants. It was shown that definite lamination appears in the prefrontal region at about the by guest on January 7, 2015 sixth month of foetal life, by separation off of the innermost neuroblasts from the remainder. In other words the cortex develops from within outwards. When lamination is completed the cortex consists of the following five primary laminae, of which two are mainly fibre-laminse, and three mainly cell-laminae : (1) Outer fibre-lamina or superficial layer ; (2) outer cell-lamina or pyramidal layer of nerve-cells; (3) middle cell-lamina or layer of granule-cells ; (4) inner fibre-lamina or inner line of Baillarger, and (5) inner cell-lamina or layer of polymorphic cells. These observations were confirmed by Watson in 1907 [14], and Brodmann's monograph which was published in the year 1909 [5] contains certain illustrations of foetal cortex which agree in every histological respect with the features above referred to. In 1910 [3] this description was amplified and elaborated, and it was pointed out that, whilst the cortex generally evolves from within outwards, that of the visuo-sensory area (sensory projection cortex) develops before that of the visuo-psychic zone (cortex of lower associa- tion), and the latter evolves, before the cortex of the prefrontal region (cortex of higher association). Whilst it is not necessary for our present purpose to give further details with regard to these investigations, it may be remarked that CYTO-ARCHITECTURE OF THE CEREBRAL CORTEX OF A HUMAN FCETUS 3 in spite of their incompleteness they have served as a key for the elucidation of the structure of the cortex cerebri in the various types of mental disease, from the idiot on the one side of the scale to the dement on the other, and to enable a general pathology of mental disease to be indicated as a broad generalization of amentia and dementia on a basis of cortical sub-evolution and dissolution. Though not germane to the subject, reference must not be omitted to the classical work of Flechsig [8] on cortical localization, since this has so largely served as the basis of the more modern researches in this subject. Whilst «these myelinogenetic observations of Flechsig, and the more recent researches of the Vogts, have furnished valuable information for correlation with the results of the study of the cyto- architecture of the adult cortex, they are without bearing on the early evolution of the cerebral cortex, as lamination and histological differentia- tion occur at a much earlier date than does myelinization. The observations on the mode of evolution of the cortex of the foetus to which reference has just been made have enabled certain general Downloaded from laws of development to be stated. They are, however, not sufficiently complete to enable detailed statements to be made with regard to the early evolution of the cortex and to the histologically differentiable areas by guest on January 7, 2015 which first make their appearance. II.—MATERIAL AND METHODS. We have endeavoured in the present research to fill this gap from the aspects both of cyto-architecture and of cortical localization. The facts which determined our choice of a brain for these purposes are as follows: In a foetus of six months Betz-cells are present, and the pre- frontal region, which is of very late development, has begun to laminate. In a foetus of a little more than four months slight indications of differentiation of neuroblasts exist in the prefrontal region, but no definite evolution of lamination has occurred here. For the study of the early evolution of the cyto-architecture of the human cerebrum, a brain, therefore, from a foetus of about eighteen weeks—i.e., a trifle older than the last just referred to—seemed most likely to be productive of useful results. The evidence on which is based the age of the foetus we have employed is as follows:— The foetus when extended measured 25 cm. in length. When flexed as in utero the length from the vertex to the tip of the sacrum 4 ORIGINAL ARTICLES AND CLINICAL CASES was ]4"25 cm. The fingers were distinct and separate, and the nails had appeared. The scalp was covered with downy hair, but no hair whatever was apparent on the body. The mouth was open. The membrana pupillaris was present. Ossification had not begun in the os calcis or in the pubic bone. Small centres of ossification were found in the upper two sacral segments. The gall-bladder was formed. It was relatively thick, and the cavity was small, but apparent. A slight Downloaded from by guest on January 7, 2015 FIG. 1.—Two diameters. Uostripped left hemisphere of human foetus of the age of eighteen weeks. fold was noticeable "at the ileocsecal junction, but the valve was not present. The kidneys were of about the size of a small hazel-nut, and measured 23 mm. in length by 8 mm. in width. The sex was definitely determinable, the foetus being a female. As at this age the brain, even when hardened, is of the consistence of soft jelly or blancmange, considerable difficulties were met with which, without proving unsurmountable, nevertheless rendered the study of" the hemispheres less complete than was at first hoped. Unfortunately it is impossible in a research of this particular kind to CYTO-ARCHITECTURE OP THE CEREBRAL CORTEX OF A HUMAN PCETUS 5 fill in the gaps by employing another exactly similar brain. This is the case on the one hand because it would, to say the least, be very difficult to duplicate a brain of a particular age; and on the other because, as one of us has experienced, great differences exist in the degree of development of two brains which are apparently of exactly the same age. These are in fact so marked as to render futile any attempt to fix in an exact manner the various stages of early cortical development. The brain was hardened in formalin in situ, the skull bones of the vertex being first removed. After a preliminary hardening the brain was carefully floated into a separate dish of formalin. The examination was conducted by the only really satisfactory method in our experience, that of piecemeal blocking—the pia-arachnoid being retained as long as possible in order to keep the fragile hemispheres from breaking up, and the blocks being cut under fluid. The latter were embedded and cut in paraffin, and the sections were selected by counting and stained by the polychrome and iron-alum-hsematoxylin methods. The former Downloaded from method was found to demonstrate the Betz-cells in a beautiful manner, whereas the latter was of value owing to the permanent nature of the results obtained by means of it, since it is quite easy with experience by guest on January 7, 2015 to interpret such sections in terms of the more elegant but less permanent polychrome specimens.
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