Paper - An iconometrographic representation of the growth of the central nervous system in man
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Grenell RG. and Scanimon RE. An iconometrographic representation of the growth of the central nervous system in man. (1943) J. Comp. Neural. 79(3): 329-
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- 1 An Iconometrographic Representation of the Growth of the Central Nervous System in Man
- 1.1 Historical Note
- 1.2 Part I. The Spinal Cord Introduction
- 1.3 Part II. The Brain
- 1.4 Literature Cited Introductory Note
- 1.5 A. The Brain
- 1.6 B. The Spinal Card
- 1.7 Literature Cited
An Iconometrographic Representation of the Growth of the Central Nervous System in Man
Robert G. Grenell and Richard E. Scanimon
Department of Anatomy and the Graduate School, Uniners-ity of Minnesota, Minneapolis
Robert G. Grenell - Now at the Laboratory of Physiology, Yale University School of Medicine, New Haven, Conn.
In connection with the general discussion of the purposes of iconometrography given in this pa.per, a few notes may be presented on the development of this powerful and ﬂexible method.
Iconometrography is not new. It may be said to have been used in a simple form by A. Diirer (1528, 1557). It was certainly employed in one fugitive plate by Criséstomo Martinez, probably before 1690. This plate was reproduced in 1692 and in 1780. A modern copy has been published by Frank (’20).
Hueter (1865) made use of the method in a study on the growth of the thoracic skeleton. W. His (1880-1885) seems to have been the ﬁrst investigator to make use of both line-ar and areal iconometrographs. The former illustrates the development of the digestive tract. The latter consists of determinations of the area of the upper and lower extremities. The method was employed by Langer (1884) to the study of the growth of the skull, by H01] (1898) to the growth of the facial skeleton, and with great success by Merkel (1882, ’03) to the study of the growth of the skeleton.
The method has been greatly extended in recent years by the development of volumetric or tridimensional iconometrographs, such as are used in this paper. The application of iconometrography to serial or ranked observations is a separate branch of the method that need not be discussed here.
The term iconometrography (from atmbv, a likeness, picture or, more particularly, a conventionalized image; uéroov, the measure; and vodgjsiv, to draw) is used to replace the earlier term iconography proposed by Scammon (’29). The shorter is properly applied to certain forms of ecclesiastic art.
CAMPER, P. 1791 Dissertation physique . . . ur les diiférences réelles que présentent les traits «in visage ehez les hommes de dilférents pays et de ditferents ages . . . Traduite du Hollandois . . . Utrecht.
DiiR.ER., A. 1557 Les quatre livres d’Albert Diirer . . . Paris.
FRANK, M. 1920 History and bibliography of anatomic illustration . . . by Ludwig Choulant. Translated and edited by Mortimer Frank. Chicago: The University of Chicago Press.
HIS, W. 1880-1885 Anatomic menschlicher Embryonen. 3 vol. + Atlas. Leipzig.
HOLL, M. 1898 Ueber Gesichtsbildung. Mitteil. d. anthrop. Gesellsch. z. Wien, Bd. 28, s. 57-100.
HUETER, C. 1865 Die Formentwickelung am Skelett des mensehlichen Thorax. Leipzig.
LANGE-R, C. 1884 Anatomic der ausseren Formen des menschlichen Kiirpers. Wien.
MERKEL, F. 1882 Beitrag zur Kenntniss der postembryonalen Entwicklung des menschlichen Schadels. In: Beitréige zur Anatornie und Embryologie als Festgabe Jacob Henle . . . pp. 164-183 + pl. 14-20. Bonn.
1903 Bemerkungen zum Beckenwachstum. Anat. Hefte, Bd. 20, S. 123-150.
SCAMMON, R. R. 1929 The measurement and analysis of human growth. (Third conference on Research in Child Development, University of Toronto, Toronto, Canada, May 2-4, 1929, p. 10 et seq.) Washington, D. C.: Division of Anthropology and Psychology, National Research Council.
Part I. The Spinal Cord Introduction
Growth is one type of change. It is the result of the activities of fundamental processes, past and present—processes which form a continuum that may be regarded as the growth phenomenon, or which may be said to give rise to growth. It is not a static, but a dynamic condition, which Varies (in its relationships to time, space, etc.) from individual to individual, from structure to structure within one individual, and from tissue to tissue and cell to cell within a single structure itself. The growth of a structure may be spoken of, therefore, as relative (when compared with that of other structures), absolute or both. It is different from the changes of a strictly inert body with respect to the dimension of time. A cell, existing in time, will be exposed to forces that are an integral part of its external environment and its metabolism, and will of necessity respond. As these inﬂuences become greater in number and more complex, or during those limited periods of time when they are more active or powerful, the cell is subjected to change after change. The resultant series of changes may be said to be growth.
From its very essence, growth is something which has deﬁnite properties. It cannot be discussed without considering its relation to rate (a change with time, either relative or absolute, which may be expressed mathematically or otherwise), or without one’s being aware of it’s being the product of a continuous series of reactions, and also as a synthesis of known and unknown mechanisms of physical or chemical nature.
Growth can be studied from one of two points of view; either we investigate the metabolic and other causative factors affecting it, and postulate what may presumably occur, or we make observations, recordings and comparisons of a structure at various intervals up to and including its full-grown state.
From the latter point of view, the growth entity can be further analyzed in three different ways: qualitatively (differentiation), quantitatively (changes in dimensions, texture and composition) and functionally (physiological), each of which may or may not have a bearing on the others. The problem at hand is one of a quantitative nature. How, then, are we to present our measurements and observations, by a simple technique, which will nonetheless take into account the properties mentioned above? In order to meet these requirements and demonstrate the growth of the spinal cord, iconometrographs have been employed, as containing all the characteristics necessary to illustrate the essential points mentioned in the foregoing discussion. An iconometrograph, as used here, then, may be deﬁned as a series of pictorial representations of a structure or structures, based on quantitative data, which portrays Various properties or dimensions of such structures in proportion, in relation to each other or to other dimensions of the body, and which takes into account their variations with time.
We wish to express our gratitude to Dr. Edith Boyd, for her valuable suggestions through the investiga.tion and the preparation of the manuscript; and to Miss Jean Hirsch, Miss Evelyn Erickson a.nd Mr. Carl Bauer for their interest in and capable renditions of the plates.
Materials and Methods
The data used in the compilation of the accompanying ﬁgures were obtained from measurements in the published works of Lassek and Rasmussen (’38, ’39) and Miller (’13). Lassek and Rasmussen incorporated some values modiﬁed from Miller, and as they state, their ﬁgures compare favorably with those of Donaldson and Davis (’03), and Baistrocchi (1884). From the data volumes of square prisms and spheres were calculated by Euclidian formulae to ﬁt the volume of a square prism in ﬁgure 2, and to ﬁt the volume of a sphere in ﬁgures 3 and 4 (the formula for ﬁgure 2 was, V = A'L; for ﬁgures 3 and 4, V = 4.189r3). In ﬁgure 5 the ﬁgures in the form of percentages were represented as radians, and the diagrams drawn accordingly. The ﬁgures in ﬁgure 1 were adapted from Eternod (1899), von Koelliker (1861), Retzius (1896) and Sobotta-McMurrich (’33).
The plates were prepared to show a simple means by which the quantitative aspects of growth may be analyzed, namely, the iconometrographic method.
Observations and Discussion
An investigation of the growth of the nervous system is of interest because of the intrinsic nature of the subject, for the functions of which the embryo or fetus is capable at various times are determined by the growth of the nervous structures. Moreover, any investigation of the central nervous system will help us to understand man’s mental abilities of inﬂuencing or reacting to his environment. Such an investigation is also of interest because_it enables us to compare the mode of growth of the nervous structures with that of other organs. As expressed by Scammon (’33):
- “The growth and development of the central nervous system of man exhibit certain striking characteristics that distinguish its history from that of most parts of the body. The increase in mass of the central nervous system is precocious and is completed long before the body as a whole reaches its deﬁnitive size. The major features of its form are all established at a relatively early period and undergo little change thereafter; and the neurones that constitute its essential elements are present in almost their full quotaat a remarkably early stage in development.”
Before breaking down the spinal cord into its several component parts for analytical study, it is well to glance brieﬂy at it and at the associated structures of the central nervous system, and to note the more general changes (in the spinal cord) that may be observed throughout the growth period. The central nervous system forms from a medullary plate. The plate folds into a tube (after differentiation into parts that may be termed brain and spinal cord). The caudal portion of the plate (or later, the tube) represents future spinal cord. It extends to the inferior tip of the body until approximately the end of the second month of embryonic life. The caudal part of the cord, at this stage, will later form the ﬁlum terminale (this division may be recognized by the middle of the third fetal month), which will increase in length with time due to increase in rate of growth of the vertebral column. At birth, the cord terminates at about the inferior border of the second lumbar vertebra; in the adult, about one vertebra higher. This relationship has been discussed by Lassek and Rasmussen (’38) who found that the spinal cord at birth is proportionately longer to body length than in the adult, being 1 to 3.2 in the former, and 1 to 4.1 in the latter.
The volume changes of the central nervous system as a whole have been depicted as a curve. Dunn (’21) states that,
- “The curve of the absolute volume of the central nervous system, when based upon crown—heel or total body length, is concave like all other curves of volume growth of the fetal organs. . . . When calculated according to age in fetal months, the absolute volume of the central nervous system is found to be 2 cc. at the beginning of the third month, rising to 36 cc. at the beginning of the sixth month, and to about 340 cc. at birth. . . . The growth curve of the central nervous system is analogous in both its character and its slope to the growth curves of nearly all the fetal viscera. . . . It is a complex of four distinct subtypes of growth, all of which are dominated by the growth of the bulky cerebral hemispheres.” The four subtypes referred to are cerebral growth, brain-stem and cord growth, cerebellum growth, and so-called compound growth, which “represents merely the combined effect of two or more of the above types predominated by the mass of the cerebral hemispheres.”
Figure 1 illustrates these general observations in actual stages of development of a 2.1-mm. embryo, by the use of the iconometrograph; the embryo is depicted at 3 months, 4 months, 6 months, 7 months, newborn and adult. The central nervous system has been held to the same length throughout, so that we are enabled to see the changes which have taken place, but which would have been concealed if each had been represented in its actual length. We can readily observe the lengthening of the ﬁlum terminals, and also, in each case, the mass and other relationships of spinal cord to brain, and of the several parts of the brain to each other. This method of representation, then, allows us to observe the general growth picture from embryo to adult, and gives us a basis upon which to establish a more detailed analysis.
What are the changes in dimensions of the spinal cord with age, and when and where do they occur? Figures 2 to 5 present answers to these questions. It was deemed simplest to begin by portraying the cord as consisting of square prisms; that is, a square prism of white matter within which is shown a square prism of gray matter. In ﬁgure 2, the ﬁrst row of prisms represents the actual dimensions (drawn to scale) of spinal cords in an 11-mm. embryo (ca. 33 days), 15-cm. fetus (3.9 lunar months), 32-cm. fetus (6.5 lunar months), newborn and adult. There are decided increases in length, in volume and in area (any two of which determine uniquely the third). The length augmentation from 11-mm. embryo to newborn is approximately 2.5 times; to adult 58.5 times; thus, from birth to maturity, the cord almost triples in length. The multiple increment in volume from 11-mm. embryo to 15 cm. fetus is about 67; from 15-cm. fetus to newborn almost 10; and from newborn to adult slightly more than 10 (this is also shown in ﬁgure 3 in a different form).
Figure 1 Squat: Pums urumrrma Vouuur. or Smuu. Cow Ft-an embryo to «ink i
There is also a change in the volumetric relationships of gray matter to white matter. In the earlier stages, this ratio is much smaller than at later periods (this may also be seen in ﬁgures 3, 4 and 5), due to a greater increase in white matter than in gray.
The remaining ﬁgures in ﬁgure '2 are attempts to demonstrate the essence of the foregoing changes, by assuming that they do not occur. That is, if we hold any one dimension constant, (which in the normal course of events increases with age) through all the stages of growth, the constructions obtained reveal the changes and Variations that were obscured by its own growth during the period when it was not constant (i.e., when it was increasing). The second row of ﬁgures results when the length has been held constant to the length of the newborn throughout the series. Thus, in the prenatal stages, where the length has been increased over the normal value, the area decreases, whereas after birth the process is reversed. In the third row, where the volume has been held constant, the exact opposite to what resulted with constant length, occurs. This would be expected. Finally, where area has been held constant, the expected increase in length is evidenced.
If, then, we synthesize all the observations made when we hold a dimension constant, we can mentally reproduce the ﬁrst row of structures on ﬁgure 2——the actual changes.
The questions as to when and where the above changes take place, still remain to be answered. Figure 3 consists of spheres representing the volumes of gray matter. white matter and total cord from fetus of about 4 lunar months (15 cm. CH) to ma.turity. The greatest part of the increase in volume of the cord appears to lie in the growth of the white matter, most of which occurs after birth. In this instance, the multiple increment from 15-cm. fetus (ca. 4 lunar months) to newborn is slightly more than 14 and that from newborn to adult about 15. In distinction the respective increments for gray matter are approximately 9 and 6 (this being true, even though the white volume is less than the gray during the first half of gestation). The important increase in volume, then, occurs for the most part after birth in the white matter.
Figure 4; represents the regional volumes of gray and white matter of the cord. Here, two major points can be clearly distinguished. Firstly, the increase in volume of the cord with age is for the most part a postnatal one in the white substance. Secondly, within the white matter itself the largest percentage increase is in the thoracic region, with the cervical region next in importance (the cervical augmenting approximately 24 times from 15—cm. fetus to adult — slightly less than the total cord— and the thoracic about 38 times). Furthermore, both gray and White matter grow more rapidly during the last half of the prenatal period, with the white rapidly increasing over the gray; and during all prenatal stages, the cervical region contains the greatest amount of both gray and white matter. Both the absolute and relative changes in the lumbar and sacral segments of the cord are small.
In order to portray these relationships more clearly, a series of sixteen discs has been drawn up to show the regional percentage of volumes of gray matter, white matter and of the length of the total cord. These are shown in ﬁgure 5. They present a recapitulation of many of the points mentioned above as well as some additional ones. The Volume of gray matter shows little prenatal change in regional relationships, but all four regions increase in the postnatal period; the thoracic the most. followed by cervical, lumbar and sacral in that sequence. The cervical region has the largest volume of gray matter in both the prenatal and postnatal periods. The growth in the thoracic region is again apparent in the white matter, where the postnatal increase is 1970%. On the other hand, the white matter of the cervical segments increases 1071%, that of the lumbar 1061%, that of the sacral 829% and the total white matter 1410% (Donaldson and Davis, ’03). Similar results can be seen regarding the percentage volumes and length of the total cord. The thoracic cord shows the greatest increase in all dimensions, of all regions, with cervical and lumbar following. The least change occurs in the sacral region.
Concerning brain changes in the prenatal period, Dunn (’21) states that, “The growth of the spinal cord resembles that of the brain and particularly that. of the pons and medulla.” He also notes that, “At the middle of the second month the cord volume is equal to 4.4% of the total brain volume. It then drops rapidly at ﬁrst and afterward more slowly to about 0.85% of the total brain volume at birth.”
Certain major conclusions may be drawn from the foregoing material:
1. The spinal cord increases in length, area and Volume from the early embryonic period to birth, through infancy and probably through the ﬁrst decade.
2. The greatest increase takes place not only in the white matter, but in a speciﬁc region of the white matter~the thoracic.
3. The largest absolute changes occur after birth.
4. The iconometrographic method presents the development of the spinal cord in a concise, simple and systematic manner. Its major advantage over other methods of exposition lies in its use of the third dimension. This use of the third dimension enables one to see what is taking place with greater ease and clarity, without resort to verbal, mathema.tical or other complex methods. The iconometrograph, then, is a way of assembling, analyzing a.nd synthesizing quantitative changes, in order to study the essence of the more fundamental qualitative factors underlying the phenomenon of growth. It is a. means of representing data and gleaning information as to what basic physiological processes are, by what they do.
Part II. The Brain
Many investigators have reported their ﬁndings concerning various characteristics of growth of the brain and its parts. The changes in this organ. with growth, are of interest because it is desirable to know both the “normal” structure and its “normal” age changes which give a key to pathologic occurrences a.nd may be correlated with the development of general nervous coordination of the body, reﬂex activity, etc., and perhaps with the deVelopment of intelligence. GROVVTH or THE NERVOUS SYSTEM 343
Numerous studies can be found concerning brain weight, brain volume and shape, myelinization, changes in size and proportion of divisions or regions of the brain with age, and formulae for, or curves of, brain growth. Apparently, however, no attempt has been made for decades to summarize such information. The present investigation is an attempt to accomplish this by means of the iconometrographic method.
Material and Methods
The data employed in designing the accompanying plates were calculated from ﬁgures of Jenkins (’21), Dunn (’21), Scammon (unpubl). and Scammon and Hesdorffer (’36). All the spheres in ﬁgures 6, 7 and 8 were determined by recal culating (according to the formula V = 4.189r"; 4.189 =—:—“)
the recorded volumes of various parts of the brain, to ﬁt the volume of a sphere. The spheres are placed one inside the other for the following reasons:
1. It permits a representation resembling the situation in the actual brain as much as possible in such a schema.
2. It is possible to grasp quickly, easily and clearly, the changes that occur in volume because of the simplicity of the geometric ﬁgure, and the use of the third dimension.
The remaining ﬁgures, i.e.. those of the schema in ﬁgure 6, are diagrammatic representations of percentage volume, with respect to the brain as a whole, obtained by merely converting
percentage ﬁgures to radians, and drawing the segments to the
proper angles. OBSERVATIONS
The ﬁrst set of ﬁgures in ﬁgure 6 represents the volumes of the brain-stem, cerebellum, cerebrum and of the total brain in ﬁve prenatal stages: 12.3 cm. CH, 22.3 cm., 32.2 cm., 42.8 cm. and 52.2 cm. CR (all are mean body length). In ﬁgure 7, the volumes of the same divisions are represented during the ﬁrst postnatal year, again showing ﬁve stages: birth, 2 months, 3 months, 6 months and 12 months. The spheres, arranged one within the other, demonstrate not only the growth of these re344 ROBERT G. GRENELL AND RICHARD E. SCAMMON
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Figure 6 GROWTH or THE NERVOUS SYSTEM 3-15
gions at ten different stages, but what is equally if not more important, allow a comparison of their rates of growth, for they show the relation of one volume to another at each stage. This method, then, illustrates the actual absolute a.nd relative growth of the brain and its parts in the fetal period and infancy.
There is a steady, rapid increase in volume in all four spheres. The brain-stem volume in the embryonic and fetal periods (from 12.3 cm. to 52.2 cm. CH), increases from 0.52 cc., to 7.62 cc.; the cerebellum volume, from 0.26 cc., to 2134 cc.; the cerebrum volume, from 5.2ecc., to 334.4 (20,; and the total brain volume from 5.4 cc., to 367.5 cc. It is also obvious (especially if the ﬁgures in ﬁgure 6 are compared with those in ﬁgure 7) that there are differences in the relative increase in volume of the parts of the brain. In the embryo, the cerebrum and cerebellum are relatively small, the brain-stem large. With the passage of time, the cerebrum shows a slight relative increase (with respect to volume of the total brain) to the terminal stages of the prenatal period. This is followed by a deﬁnite decrease. The cerebellum shows a continuous and marked relative increase in volume. The cerebellar increase, therefore. is one of relative rate of growth. This is of great interest and will be mentioned later. The brainstem, which was comparatively large in the embryo. remains so in early fetal stages, but shows a steady relative decrease in volume through the later fetal period and the ﬁrst postnatal year. (It has been shown to have a slight increase at the end of infancy.) No marked relative changes occur after the tenth year}
The second set of ﬁgures in ﬁgure 6 represents one of the distinct and interesting changes occurring in the cerebrum in the prenatal period. It demonstrates one of the few cases of human ontogeny recapitulating phylogeny, and explains the small size of the olfactory system in man and the presence of so—called vestigial ﬁber paths. The archipallium (olfactory) is quite large as ‘compared with the neopallium in early stages, but shows a steady decrease in volume throughout the fetal period, as opposed to a marked increase of the (phylogenetically speaking) more recently developed part of the cerebrum. By the end of the prenatal period, the archipallium is small and has decreased very markedly (both absolutely and relatively), while the neopallium has been rapidly developing. According to Jenkins (’21),
1 It is possible that the small relative increase between 1 and 10 years is due to the small sample of material available for this period.
in a 16-mm. embryo, the archipallium comprises about 18% of the total brain weight; the neopallium slightly more than 3%.
In a 367-mm. fetus, the percentages of archipallium and neopallium are 1.03 and 83.92 respectively.
Figure 8 demonstrates the volumes of the several parts of the cerebrum from birth to maturity. The upper four ﬁgures represent gray matter (cerebral cortex, caudate nucleus, lenticular nucleus, globus pallidus, putamen, claustrum, amygdaloid nucleus, hippocampus, thalamus, mamillary bodies and geniculate bodies), and the lower four, the white matter (subcortical white substance, corona radiata, corpus callosum, anterior commissure, internal capsule, ansa peduncularis, fornix and optic tract), with respect to total gray substance and total white substance respectively. (No values were obtainable for mamillary body volume at birth and 0.44 years, or for the claustrum at birth.) The various increases in volume may be seen in table 1. It will be noted that the greatest relative increase of gray substance is in
Volumes in t'€'Ilti’rVl€t6"7'8 of parts of the cerebrum as shown in ﬁgure 8 at birth and 32 years of age. Data from R. E. Scammon.
1 2 3 4 5 6 Birth . . . . . . . . . . . . . . 195.2 5.3 0.8 5.2+ 0.2 206.7 32 Years . . . . . . . . . . . 495.2 28.9 3.3 12.7 0.3 540.4
1 2 3 4 5 6 Birth . . . . . . . . . . . . . . 75.8 1.9 3.2 0.3 (0.0—0.1) 81.2 32 Years . . . . . . . . , . 468.3 21.8 17.9 3.1 0.6 511.7
the basal nuclei; of white substance, in the fornix. Moreover, although the gray substance more than doubles from newborn to adult (Hesdorﬁer and Scammon, ’36, state that the cerebrum as a whole increases about two and one-half times in volume in postnatal life; the basal nuclei nearly threefold), the whit.e substance increases to a much greater extent, so that most of the postnatal growth of the cerebrum may be said to occur in the white matter. This increase in white matter has been thought to be a result of myelinization.
Discussion and Conclusions
Certain advantages of iconometrographic representation are evident when it is compared with other methods. Various curves, graphs, tables and other expressions (mathematical and graphic) concerned with some phase of growth of the brain, may be found throughout the literature (e.g., the studies of Dunn, F ranceschi, Jenkins, etc.). These attempt to show the changes in structure or relationships which characterize the growth of an organ or organs. However, although these techniques convey a great deal of information, this information is limited in amount and kind. and, moreover, necessitates speciﬁc knowledge on the part of the reader, concomitant with a mass of long and frequently involved explanations. All this material, for the brain and its major parts. can be summarized in one or two iconometrographs, along with the further advantage (not possible with the other methods) of representing the third dimension at any one time and also through an extended time period—a factor of the utmost importance for the visualization of changes occurring in a body in Euclidian space.
Several points of interest concerning the growth of the cerebellum are evident in the accompanying plates. The rapid rate of absolute growth. with its marked relative increase——even during a postnatal period of absolute decrease—immediately arouse interest, especially in the light of evidence such as that described by Tyler and van Harreveld (’41) and Kabat and Pusin (’41). The former workers have found, in the rat, that in the ﬁrst three weeks of postnatal life the cerebellum (and medulla) takes up most of the oxygen used by the nervous system (i.e.. it has the highest rate of oxygen consumption per milligram wet weight). Tyler and van Harreveld also found that even in the adult mouse the oxygen uptake of the cerebellum is the highest per unit volume. Kabat and Pusin studied cerebellar development in dogs. They noted the growth of the external granular layer, and observed the occurrence of postnatal mitosis. These findings (plus those evident in the accompanying plates), suggest that a study of human cerebellar growth in the prenatal and early part of the postnatal periods might help to furnish valuable information concerning problems of normal structure and function and of cerebellar pathology.
Literature Cited Introductory Note
Even a fairly complete list of the published records on the quantitative growth of the central nervous system would require more space than here available. The history of the accumulation of these records begins with occasional notations in the eighteenth century, of the size of the brain and spinal cord in the developmental period. There are numerous, although short, tabulations of quantitative records in the first half of the nineteenth century. The following’ outline is a very brief and condensed summary of the subject:
A. The Brain
1. Quantitative studies of postnatal growth
Brain as a whole. lﬁeginning with the early tabular records of Tiedemann (1816), Sims (1835), Peacock (1851), Huschke (1854) and others. The data were greatly improved by the publication of extensive tables by R. Boyd (1861), von Bischoff (1880), Oppenheimer (1889) and H. Vierordt (1890). These were, and still are being increased by numerous collections of records, including those of Retzius (1896), Marchand (’02), Hultgren (’12), Roessle and Roulet ('32) and more recent contributors.
There are at least twenty graphic representations of these data in a variety of forms, extending from the early work of Oppenheimer (1889), Vierordt (1890) and Keith (1895), to the curves of Roessle and Roulet ('32).
Formulae for the expression of the relation of brain weight to age in postnatal life have been developed by Pearl (’06), Scammon and Dunn (’24), Dunn (’25) and \Vcinbach (’38).
Brain pavrts. Extensive records of the growth of the major divisions of the brain (cerebellum, cerebrum and brain stem) were ﬁrst published by R. Boyd (1861), although there were several minor col— lections of data published prior to this date. They have been extended by Pﬁster (’ 3a) and other observers. Empirical formulae with their graphic presentation were published by Scammon and Dunn (’24).
Cerebral parts. The study of the postnatal growth of the parts of the cerebrum may be said to have begun with the determination of cerebral surface by R. \\'ag'ner (1860) and H. \Vag11er (1864). Systematic planimetric measurements of serial thick sections of the cerebrum, by Anton (’03), Jaeger (’10), Dahlberg (’23) and a num— ber of later workers furnished data for estimating the respective volumes of the cerebral cortex a11d medulla. Improvements in the thick section and planimetric techniques by Dunn (’25), enabled him to compute the volume of the basal nuclei as a whole and certain of their divisions as well as that of certain divisions of the white matter. His studies cover the period from birth to maturity. Hesdorffer and Scammon (’36) using the material of Dunn, made thick»plate wax reconstructions illustrating the postnatal growth of the basal nuclei and the cortex. The basal nuclei have also been studied quantitatively from dissections by Francesehi (1888).
2. Quantitative studies of prenatal growth
Brain as a whole. Data on the growth of the brain in prenatal life were published in the latter part of the nineteenth century by Arnovljevic (1884), Brandt (1886) and a number of other observers. Jackson (’09) published extensive tables of the volume, absolute and relative, of the brain in prenatal life. Jenkins (’21) and Dunn (’21, ’26) published extensive data on the subject, and these have been augmented by numerous other studies.
Graphics of the growth of the brain in the prenatal period have been published by Jenkins (’19) Zangemeister (’11) and Dunn (’21, ’26).
Empirical formulae have been proposed by Dunn (’21, ’26).
Bra/in parts. The data on this subject are less extensive, those of Jenkins (’19) and Dunn (’21, ’26), being apparently the most extensive.
Graphics are included in the work of Jenkins and of Dunn, and Dunn has proposed two series of empirical formulae.
There seem to be no quantitative data on the prenatal growth of the parts of the cerebrum.
B. The Spinal Card
1. Quantitative studies of postnatal growth
The older quantitative studies of the cord were conﬁned mainly to measurements of its length. The chief sources of data on its postnatal growth in mass are the studies of Mies (1893), Pﬁster (’03b), Lassek (’35) and Lassek and Rasmussen ( ’38). The work of Lassek and Rasmussen furnished data on the gray and white matter. There are no extensive graphics or formulae for the postnatal growth of the spinal cord.
2. Quantitative studies of prenatal growth
The more important sources of data are the papers of Jackson (’09), Miller (’13), Dunn (’21, ’26) and Lassek and Rasmussen (’39). Miller and Lassek and Rasmussen give detailed data on the growth of the gray and white matter. . Graphics of the growth of the cord in prenatal life are given in the papers of J aekson, Miller, Dunn, and Lassek and Rasmussen. Dunn (’2.1, ’26) has proposed empirical formulae for expressing the prenatal» growth of the cord in length and volume.
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