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 (PKINS HOSPITAL REPOR 
 
 THE JOHNS HOPKINS HOSPITAL REPORTS 
 MONOGRAPHS. NEW SERIES No. VI 
 
 THE NUCLEI TUBERIS LATERALES 
 
 AND THE SO-CALLED GANGLION 
 
 OPTICUM BASALE 
 
 BY 
 EDWARD F.ImALONE 
 
 (From the Anatomical Laboratory of the University of Cincinnati) 
 Illustrated— 15 Plates 
 
 BALTIMORE 
 
 THE JOHNS HOPKINS PRESS 
 
 1914 
 
 fCopyright, 1914. by The Johns Hopkiiu Press]
 
 «!• 
 
 
 CONTENTS. 
 
 PAGE 
 
 I. Introduction 1 
 
 II. Material 3 
 
 III. Pars Mammillaris Hypothalami 4 
 
 IV. Ganglion Opticum Basale 7 
 
 A. Location and extent of tlie ganglion opticum basale 
 
 1. Man 8 
 
 2. Macacus rhesus 11 
 
 3. Lemur rufus 11 
 
 4. Cat 12 
 
 B. Cell type of the basal optic ganglion 13 
 
 C. Separation of the basal optic ganglion from surround- 
 
 ing cell groups through differences in cell char- 
 acter 14 
 
 V. Nuclei Tuberis Laterales 18 
 
 A. Location and extent of the nuclei tuberis laterales 
 
 1. Man 19 
 
 2. Macacus rhesus 21 
 
 B. Separation of the nuclei tuberis laterales from sur- 
 
 rounding cell groups through diiterences in cell 
 
 character 21 
 
 VI. Relationship of the Various Cell Groups to One Another 23 
 
 VII. Conclusion 27 
 
 VIII. Literature 31 
 
 IX. Bibliography 38 
 
 X. Explanation of Illustrations 39
 
 THE NUCLEI TUBEEIS LATERALES AND THE SO-CALLED 
 GANGLION OPTICUM BASALE. 
 
 By EDWARD F. MALONE, 
 
 (From the Anatomical Laboratory of the UniverHty of Cincinnati.) 
 
 There is perhaps no region of the mammalian brain concerning 
 whose cell groups less is known than that portion of the telencephalon 
 known as the pars optica hypothalami. The cell groups which consti- 
 tute the subject of this article are so vague and so hopelessly confused 
 in the literature that it is not even possible to determine whether the 
 names basal optic ganglion and nuclei tuberis should be applied to 
 different cell groups or to different portions of the same cell group. 
 In this article it will be shown that these names should be applied to 
 entirely different cell groups ; moreover the location and extent of the 
 two cell groups will be described in different mammals, together with a 
 consideration of their relation to surrounding groups of cells, and it 
 will be shown that these two cell groups under consideration are com- 
 posed of cells of radically different character through which each group 
 may be readily distinguished from the other and from the surrounding 
 cell groups. In addition the cell groups of the pars mammillaris of the 
 hypothalamus will be described. 
 
 When we consider the methods employed by most of those who study 
 the anatomy of the mammalian brain perhaps it will not be surprising 
 to realize that the best description of the basal optic ganglion and the 
 nuclei tuberis is that of Kolliker published in 1896, however faulty and 
 confusing this description may be. The reason for this lack -of infor- 
 mation concerning these cell groups is that most workers are interested 
 in the course of fiber tracts, and their interest in the structure of the 
 nervous system is for the most part limited to the largely mechanical 
 subdivisions revealed by fiber stains ; in such preparations the course of 
 fiber systems may often be followed, or at least they reveal certain 
 architectural differences of various regions which serve the purpose of 
 orientation. When preparations in which the cells are stained are 
 studied these preparations are for the most part regarded as showing 
 merely the negative picture of the fiber preparations, and the same
 
 2 Edward F. M alone. 
 
 topograpliical regions are observed, the cells being stained instead of 
 the fiber;;. When the cell character is noted it is done usually in a 
 superficial manner and principally for the purpose of orientation, and 
 with no purpose of bringing the cell character into relation with a 
 definite function. 
 
 The same lack of attention to the cell tyjie is apparent even in the 
 work of the relatively few investigators who employ the experimental 
 methods of Xissl or von Gudden. The following instance is a good 
 illustration of the value of carefully noting the cell character of various 
 cell groups. In his excellent study of the dorsal (sympathetic) nucleus 
 of the vagus nerve Molhant had shown that all the cells of this cell 
 column give origin to all of the vagus fibers which supply smooth 
 muscle and heart muscle, and that certain portions of this cell column 
 supply smooth muscle while a definite portion supplies heart muscle, 
 but he did not attempt to show that a definite type of cell was involved 
 in the innervation of each of these two types of muscle ; after studying 
 the vagus sympathetic nucleus I was able to show that the portions of 
 the nucleus which supply smooth muscle and heart muscle may be 
 readily distinguished by the fact that they are composed of cells of 
 difi'erent types, and that just as heart muscle is histologically inter- 
 mediate between smooth muscle and striated muscle, just so the cells 
 of the vagiis sympathetic nucleus which supply heart muscle are of a 
 histological character intermediate between that of the cells which 
 supply smooth muscle and striated muscle. Such an observation of the 
 relation of cell type to cell function enables us to locate accurately a 
 functional center even though its cells be mixed with those having a 
 different function, and homologous centers may be recognized in 
 different animal fonns; at the same time it emphasizes the necessity 
 of carefully noting the cell character of each cell group, so that even if 
 the function is imknown the presence of a definite cell type will offer a 
 problem for future experimental work, and in the meanwhile preclude 
 the possibility of confusing this cell group with surrounding cells. 
 These differences in cell type are of course most definite in the higher 
 mammals and especially in man, where the various cell groups are 
 highly specialized. 
 
 The foregoing consideration of the failure of workers on the anatomy 
 of the mammalian brain to attach sufficient importance to the different 
 types of cells explains why such characteristic cell groups as the basal 
 optic ganglion and the nuclei tuberis have not been clearly separated
 
 Nuclei Tuberis Laterales and the Ganglion Opticum Basale. 3 
 
 from one another and from the surrounding cell groups. The unfor- 
 tunate condition which here exists is present also in many other regions 
 of the nervous system, and I have considered the tendency of the anatom- 
 ical work on the mammalian brain somewhat at length not merely as 
 an interesting explanation for our present want of definite information, 
 but because this tendency contains a serious defect whose results are 
 most evident and far reaching, a defect which should be corrected. 
 
 The tendency of neurological workers to attach little importance to 
 differences in cell character I have criticised, not only at this point but 
 also later in this article. On the other hand I have no intention what- 
 ever of disparaging the ability of other authors, nor am I unapprecia- 
 tive of the excellence, in many cases, of their work and of the valuable 
 results which they have contributed. My criticism is aimed exclusively 
 at the almost universal belief that the various differences in cell type 
 in the nervous system are, after all, of no great importance, and that 
 We should reserve our efforts to discover the connections of various 
 portions of the nervous system. It would be unfair to blame authors for 
 being influenced by a belief so general; moreover it is only compara- 
 tively recently that we have begun to employ methods capable of reveal- 
 ing the striking differences in cell character, and only in recent years 
 has the development of cytology given an added stimulus to the careful 
 study of the nerve cell. Above all I wish to disclaim any belief that the 
 method herein employed — that of setting aside groups of cells of an 
 identical type and of comparing such cell groups and of attempting to 
 discover the correlation between cell tj'pe and cell function — is of any 
 more value than many other methods of investigation. I advocate it not 
 to displace other methods, but to complement them and add to their 
 usefulness. 
 
 Material. 
 
 The material studied consists of the following complete series : four of 
 man, one of macacus rhesus, two of lemur rufus, and three of the cat. 
 The tissue was in each case fixed in 95 per cent alcohol, and after dehy- 
 dration and treatment with chloroform embedded in paraffin. The 
 sections were stained in a one per cent aqueous solution of toluidin blue 
 (Griibler), differentiated in 95 per cent alcohol, dehydrated in absolute 
 alcohol, cleared in xylol, and mounted in Canada balsam.
 
 4 Edward F. Malone. 
 
 Pars Mamsiillaris Hypothal.\mi. 
 
 Before proceeding to describe the cell groups of the pars optica hypo- 
 thalami (telencephalon) it will be necessary to consider those of the 
 pars mammillaris hypothalami (diencephalon), since the cell groups 
 of these two subdivisions of the hypothalamus are most intimately 
 related and cannot be intelligently studied separately. In 1910 
 appeared my monograph " Uber die Kerne des mensehlichen Dienceph- 
 alon " in which the cell groups of the pars mammillaris hypothalami 
 of man were described, together with a brief consideration of certain 
 closely related cell groups of the pars optica. Since the publication of 
 this monograph I have studied both portions of the hypothalamus in 
 the monkey, lemur and cat, and have compared the cell groups observed 
 in these animals with homologous groups found in man. The following 
 description of the cell groups of the pars mammillaris of the monkey, 
 lemur and cat is accordingly original and I should like to call especial 
 attention to the illustrations which clearly show for the first time the 
 different cell groups in macacus rhesus (Figs. 19 to 25). In Series 
 D of man these cell groups are also well sIiowti (Figs. 11 to 18). The 
 other series do not show much of the pars mammillaris, since it did not 
 appear advisable to add many illustrations to show exclusively portions 
 of the pars mammillaris which are not intimately related to the pars 
 optica. In the series of macacus and the Series D of man the plane of 
 section is such as to include this caudal portion of the pars mammil- 
 laris. Moreover the dorsal portion of the pars mammillaris has not 
 been illustrated, not only because it has no close relation to the pars 
 optica but also because the thalaanus of the three lower forms must 
 first be studied. Certain types of cells also which on account of their 
 location could not be confused with the cells of the pars optica have not 
 been illustrated. Accordingly it appears best to defer such numerous 
 and ekborate illustrations until the whole diencephalon can be included, 
 and to limit the illustrations of the present article to those portions of 
 the pars mammillaris which are closely related to the pars optica. 
 
 In studying the anatomy of the cell groups of the diencephalon 
 nowhere are more satisfactory results obtained than in the hypothal- 
 aniu.s (pars mammillaris). The great difficulties encountered in 
 attempting to homologize the cell groups of the thalamus of different 
 mammals disappears when we reach the hypothalamus. This is due to 
 the fact that the thalamus is phylogenetically a much more recent
 
 Nuclei Tuheris Lateral es and the Ganglion Opticum Basale. 5 
 
 portion of the brain and its development is largely dependent upon that 
 of the rapidly developing pallium. So small is the difference in the cell 
 groups of the hypothalamus of the various mammals studied that for 
 the purposes of this article one description will suffice for all. The pars 
 mammillaris hypothalami is divided into the following primary nuclei 
 (groups of cells having an identical histological character) : 
 
 1. Ganglion mediale corporis mammillaris. 
 
 8. Nucleus intercalatus corporis mammillaris. 
 
 3. Nucleus tubero-mammillaris (nucleus mammillo-infundibularis). 
 
 4. Nucleus paraventricularis hypothalami. 
 
 5. Corpus hypothalamicum (Luysii). 
 
 6. Substantia reticularis liypotlialami. 
 
 7. Substantia grisea ventriculi tertii. 
 
 At this point it is advisable to read the explanation of the various 
 kinds of illustrations as given on pp. 39-40 ; otherwise the figures, some 
 of which are immediately to be referred to, might not be correctly inter- 
 preted. 
 
 1. Gfniglion mediale corporis mammillaris. 
 
 This cell group constitutes the greater portion of the mammillary 
 body, and extends further caudally than the other two groups. It is 
 shown in the series of macacus (Figs. 19 and 20) and in the Series D 
 of man (Fig. 11) ; no description of its relations will be given. Con- 
 cerning its cells it will suffice to state that they are of a type readily 
 distinguishable from the other two groups of the corpus mammillare. 
 
 2.' Nucleus intercalatus corporis mammillaris. 
 This is a small circumscribed cell group, described by me in 1910, 
 which lies between the medial and so-called lateral ganglion of the 
 mammillary body. Its location may be seen in the same figures that 
 show the medial ganglion (see above). Later Friedemann found this 
 group in cercopithecus and adopted the name here employed. The cells 
 of the nucleus intercalatus are readily distinguished from those of the 
 two other groups of the mammillar}' body through their characteristic 
 type; especially in man they have the relatively large and discrete 
 Nissl granules characteristic of efferent nerve cells. This cell group 
 should not be confused with groups which are occasionally mechanically 
 split off from the medial ganglion by fiber miasses ; the difference in 
 cell type renders this distinction easy.
 
 6 Edward F. Malone. 
 
 S. Nucleus tubero-mammilla/ris. 
 
 The cells of the so-called lateral ganglion of the corpus mammillare 
 may be readily followed orally and laterally into the angle between the 
 pes pedunculi and the tractus opticus, whereas orally and dorsally they 
 may be followed through many sections accompanying the tractus 
 thalamo-mammillaris and the columna fornicis. To this complex of 
 cells having an identical histological character I gave (1910) the name 
 " nucleus mammillo-infundibularis," a name which Friedemann 
 (1911) has adopted in his article on the diencephalon of cercopithecus. 
 It is evident that this name is unsuitable, since the cell group is not 
 situated in the infundibulum, and I have therefore changed it to 
 " nucleus tubero-mammillaris." The nucleus tubero-mammillaris is 
 shown in all the series illustrated and its relation to the basal optic 
 ganglion and the nuclei tuberis may be readily seen. (Since the figures 
 were drawn and labeled before I decided to change the name of this 
 nucleus, it appears on tlie outline drawings as "• nucleus mammillo-in- 
 fundibularis," but wherever space has permitted the new name has 
 been added in parentheses. The explanation of each plate is so ar- 
 ranged as to remove any possibility of confusion.) The character of its 
 cells in the different forms is illustrated in Figs. 41, 47, 52 and 57. 
 The relation of this nucleus to the basal optic ganglion will be discussed 
 later. It should be noted that a portion of this nucleus is situated in 
 the pars optica. 
 
 Jf. Nucleus paraveniricularis hypothalami. 
 
 This characteristic column of cells is situated partly in the pars 
 mammillaris but principally in the pars optica. Its location is shown 
 in all the series, and the character of its cells in Figs. 43, 49, 54 and 
 58. In man this nucleus was first described by me in 1910, and the 
 fact that such a striking cell group had not been previously described 
 in man is probably due to the almost exclusive use of fiber stains. In 
 1911 Friedemann found this cell group in cercopithecus. It is certainly 
 homologous with the group described by Cajal in rodents under the 
 name of " niicleo subventricular,"' and ^^^th that described by Ziehen 
 in marsupials as " Nucleus subcommissuralis." Homologous groups 
 are also possibly found in vertebrates even as low as the fishes. The 
 nucleus paraventricularis is intimately related to the basal optic gang- 
 lion and also to the nucleus tubero-mammillaris as tliese three nuclei
 
 Nuclei Tuheris Laterales and the Ganglion Opticum Basale. 1 
 
 appear to be components of one large cell complex, and while differing 
 from one another, differ much more from the surrounding cells. These 
 relations will be discussed later in detail. 
 
 5. Corpus hypothulamicum (Luysii) 
 
 This is such a compact, isolated cell mass, and is situated at such a 
 distance from the basal optic ganglion and the nuclei tuberis that no 
 description is necessary ; it has been included in the illustrations, how- 
 ever, for the purpose of orientation. 
 
 6. Substantia reticularis hypothalami. 
 
 This group of cells on account of its position need not be considered 
 in this article. A description will be found in my monograph on the 
 diencephalon. Many of its cells are of a motor type of structure, and 
 these together with the cells of the nucleus intercalatus of the corpus 
 mammillare are the only cells of the entire diencephalon which show 
 the histological character peculiar to motor cells. 
 
 7. Substantia grisea ventricuK tertii. 
 
 This cell mass is shown in all series and the cell type in Figs. 40, 46, 
 51 and 56. The greater portion lies in the pars optica, and it should 
 be noted that dorsally and laterally the cells are less densely packed 
 than ventrally and medially. The substantia grisea is closely related 
 to the nuclei tuberis, and this relation will be fully discussed in connec- 
 tion with the description of the nuclei tuberis. 
 
 If we now observe in the illustrations the location and extent of the 
 different cell groups previously described and note also the character of 
 their cells in the different mammals, we shall be prepared to understand 
 the following description of the basal optic ganglion and the nuclei 
 tuberis and the consideration of the relations of the latter two nuclei to 
 the fonner. 
 
 Ganglion Opticdm Basale. 
 
 The name ganglion opticum basale was given with the intention of 
 implying a function which this cell group almost certainly does not 
 possess ; on the other hand this name effectively distinguishes this cell 
 group from the nuclei tuberis with which it has been confused. Since 
 the so-called basal optic ganglion is such a characteristic cell group, and
 
 8 Edward F. }[alone. 
 
 since the homologous groups in various mammals are so constant both 
 as to location, extent and cell character, it appears highly probable that 
 we shall not have to wait long before this name may be abandoned for 
 one which will indicate its true function, and until then it seems better 
 not to introduce a temporary name implying merely some morpholog- 
 ical character. In arriving at the function of a cell group the first 
 essential (and probably the most important of all) is to clearly dis- 
 tinguish it from the surrounding cell groups and to study carefully 
 its relations to such cell groups ; in this article I shall endeavor by 
 means of illustrations and description to make the resulting picture 
 of the so-called basal optic ganglion so definite that its confusion with 
 all other cell groups will be impossible, and if this result be obtained 
 there need be little concern as to the fitness of the name employed. 
 
 Location and extent of the ganglion opticuni basale. 
 
 MAX. 
 
 The basal optic ganglion in man is a mass of gray matter, super- 
 ficially situated partly in the tuber cinereum and partly in the anterior 
 perforated substance, which extends along both borders of the optic 
 tract. The main mass of cells forms in the anterior j>erforated 
 substance a column which is closely applied to the dorsal portion of the 
 oro-lateral surface of the optic tract. This mass is connected by a 
 comparatively small number of scattered cells situated directly dorsal 
 to the tract with another cell mass which follows in the tuber cinereum 
 the caudo-medial border of the optic tract ; in other words the basal optic 
 ganglion consists, generally speaking, of two parallel columns of cells 
 lying along either side of the optic tract, which are connected by 
 scattered cells of the same type lying dorsal to the optic tract. For the 
 second time I strongly recommend a consideration of pp. 39-40, where 
 it is pointed out just what each type of illustration is intended to show. 
 
 Taking up the first series of man (Series AC) the basal optic gang- 
 lion appears caudally in Fig. 4 as a few scattered cells situated on the 
 peripher}- of the tuber cinereum ; Figs. 5 and 6 show the development 
 of the cell mass as we pass orally. The cell masses shown in Figs. 4 
 and 5 are to be regarded as a caudal projection of the cell column lying 
 along the caudo-medial border of the optic tract, and are continuous 
 with the caudal portion of this cell column which appears first in Fig. 6. 
 Between Figs. 6 and 7 is a considerable gap, in which the following
 
 Xuclci Tiiberis Latcrales and the Ganglion Opticum Basale. 9 
 
 changes have occurred : as the optic tract approaches the median plane 
 and applies itself to the base of the brain the cell mass continues to 
 increase in size, extending further laterally and also to a less extent 
 further medially, and assumes a position as a column on the dorsal 
 surface of the caudo-medial aspect of the optic tract, or in other words 
 it extends in the tuber cinereum along the line where the tuber joins 
 the optic tract. Midway between Figs. 6 and 7 the medial pole of the 
 column crosses the median line and caudal to the optic chiasm unites 
 with the corresponding column of the other side by means, however, of 
 only a few scattered cells situated in the iirfundibulum. The next 
 series (D) shows this fact. But still another change has occurred 
 between Figs. 6 and 7, namely, after reaching its greatest development, 
 which, as previously described, occurs at the juncture of the tuber 
 cinereum with the caudo-medial border of the optic tract, the cell mass 
 becomes rapidly diminished to- only a few scattered cells which are 
 situated on the dorsal (or deep) surface of the optic tract. Fig. 7 shows 
 the extent of the basal optic ganglion slightly oral to this region of its 
 poorest development; the section passes through the optic tract near 
 its oro-lateral border and since the section is near this border the cell 
 mass has increased in extent. Between Figs. 7 and 8 it increases 
 steadily in size to reach its maximum in Fig. 8 ; this figure represents 
 a section through the cell column which extends in the anterior perfor- 
 ated substance along the oro-lateral border of the optic tract. In Fig. 9 
 the basal optic ganglion has diminished in size, and this is due to a 
 shortening of the lateral portion of the column ; that the lateral portion 
 of the column should be shortened is evident when we recall that the 
 cell column lies parallel to the optic tract, and that the course of the 
 optic tract is such that in the present plane of section as one proceeds 
 orally the lateral portion of the tract appears and disappears first (see 
 Figs. 1 to 10). Fig. 10 shows the most oral portion of the basal optic 
 ganglion. The fact that the portion of the ganglion situated caudal 
 (medial) to the optic tract does not extend as far laterally -as that por- 
 tion situated oral (lateral) to the tract should be correlated with the 
 difference of the relation of the two borders of the tract to the base of 
 the brain ; for while the oral (lateral) border of the tract is in intimate 
 relation to the anterior perforated substance from the chiasm to a point 
 far lateral from the median plane, the caudal (medial) border of the 
 tract is in intimate relation to the tuber cinereum, which extends for
 
 10 Edward F. M alone. 
 
 a shorter distance laterally, and then the tract passes over the cms 
 cerebri, where of course it is no longer in relation to gray matter. 
 
 To sum up. Series AC of man (Figs. 4 to 10) shows, in sections 
 almost parallel to the course- of the optic tract, the appearance of the 
 three parallel columns of cells which constitute the basal optic gang- 
 lion ; one column lies in the tuber cinereum along the medio-caudal 
 border of the optic tract, the second extends along the dorsal (deep) 
 surface of the tract and consists of only a few cells, while the third and 
 largest column extends in the anterior perforated substance along the 
 latero-oral border of the tract. The caudal column is continuous in the 
 infundibulum with the corresponding column of the opposite side. The 
 figures show the inevitable relation of three parallel columns in slightly 
 oblique section. 
 
 Series D of man (Figs. 11 to 18) shows essentially the same relations, 
 but the plane of section is different. In the previous series the plane 
 was almost parallel to the course of the optic tract, although slightly 
 approaching the plane of a cross section ; in Series D the plane of sec- 
 tion is almost at right angles to the course of the optic tract, and both 
 the caudo-medial and the oro-lateral borders are shown in all sections. 
 It is evident that a plane of section at right angles to the median plane 
 (which would cut both halves of the brain symmetrically) could not 
 pass through the optic tract at right angles to its course, since the two 
 tracts converge towards the median plane as they pass orally ; therefore 
 to obtain a cross section of the right tract the plane of section must lie 
 more oral on this side than on the left. The asymmetry of the plane 
 of section of Series D is not quite great enough to produce a cross sec- 
 tion of the right optic tract, although this condition is very nearly 
 attained. 
 
 When Fig. 11 of Series D is compared with Fig. 4 of Series AC the 
 location of the basal optic ganglion is rather confusing, for in the 
 latter series it appeared oaudally in the tuber cinereum near the medio- 
 caudal border of the optic tract, while in Fig. 11 it appears first in the 
 anterior perforated substance along the oro-lateral border; if we keep 
 in mind the asymmetry of the section, and the fact that the cell column 
 here shown extends further laterally than the other two columns of the 
 cell group and that in such asymmetrical sections the lateral portion is 
 the most oral, little difficulty should exist in realizing just why this 
 one of the three cell columns should appear first. Fig. 12 shows practi- 
 cally the game relations, except that the optic tract is nearer the median
 
 Nuclei Tuheris Laterales and the Ganglion Opticum Basale. 11 
 
 line. In Fig. 13 the cell column along the medic-caudal border of the 
 tract, as well as the scattered cells dorsal to the tract, appears, and 
 from here throughout the entire series these three columns are present. 
 Of course all columns are in cross section. In Fig. 18 the cell mass is 
 seen to extend across the median line, a condition which occurs also in 
 the three other series of man. Series D (Figs. 11 to 18) shows clearly 
 that the basal optic ganglion consists of two well developed parallel 
 columns of cells connected by an intermediate parallel column consisting 
 of only a few cells. As will be shown later the exact extent of the 
 basal optic ganglion in all the mammals studied is readily determined 
 by the distinctive histological character of its cells, and accordingly the 
 three parallel cell columns have been considered as merely subdivisions 
 of one nucleus, not because this seems convenient, but because the 
 identical character of their cells makes such a conclusion unavoidable. 
 
 MAC.A.CUS RHESUS. 
 
 In maeaeus rhesus the basal optic ganglion is shown first in Fig. 22, 
 although it extends slightly further caudally. In tliis figure the gang- 
 lion is situated just oral to the line of apposition between the tuber 
 cinereum and the optic tract. In the sections between Figs. 23 and 23, 
 as we pass orally, the cells become reduced in number and lie dorsal to 
 the tract, and as the oro-lateral border of the tract is reached increase 
 in number. In Fig. 23 appears the cell column which lies in the ante- 
 rior perforated substance along the oro-lateral border of the tract. The 
 further development of the ganglion (Figs. 21: and 25) needs no 
 description. In Fig. 22 the medial pole of the ganglion is seen to 
 approach the median line, and between Figs. 22 and 23, just caudal to 
 the optic chiasm, a very few widely scattered cells unite the ganglia of 
 opposite sides. As in man the basal optic ganglion in macacus is 
 seen to consist of tlie same three parallel columns of cells, of which the 
 oro-lateral column (situated in the anterior perforated substance) con- 
 stitutes the greater part of the ganglion. 
 
 LEMDR RUFUS. 
 
 The basal optic ganglion of the lemur does not differ essentially 
 from that of man and maoacus, except that whereas in the two latter 
 animals it consisted of two parallel cell columns loosely connected by 
 a third, in the lemur this connecting mass of scattered cells (dorsal to
 
 12 Edward F. M alone. 
 
 the optic tract) is missing; consequently the ganglion consists of two 
 entirely separate parallel columns of cells. Fig. 27 represents a section 
 between the two cell columns. As in man and macacus the ganglia of 
 opposite sides are united by a very few cells; this union occurs just 
 caudal to the optic chiasm (slightly caudal to the level represented 
 in Fig. 27). 
 
 CAT. 
 
 The basal optic ganglion in the cat (Figs. 31 to 35) has practically 
 the same location and extent as in the lemur. As in the lemur the gang- 
 lion consists of the same two parallel cell columns, which are completely 
 separate. Between Figs. 32 and 33 lies the region in which the gang- 
 lion is absent. Just caudal to the optic chiasm (caudal to level of Fig. 
 33) the ganglia of opposite sides are united in this series by a few 
 cells ; in the other two series of the cat no union was present. 
 
 Comparing the location and extent of the basal optic ganglion in all 
 four animals the following facts should be noted: 
 
 1. The basal optic ganglion in all four animals consists almost 
 exclusively (in some cases exclusively) of two compact, parallel cell 
 columns. The larger of these two columns lies superficially in the 
 anterior perforated substance along the line where this region becomes 
 continuous with the oro-lateral border of the optic tract ; the smaller 
 column lies superficially in the tuber cinereum along the line where it 
 joins the optic tract. 
 
 2. These two constant, parallel cell columns are united in man, and 
 to a less extent also in macacus, by more or less diffusely scattered cells 
 of the same type which lie dorsal to the optic tract. 
 
 3. In all four forms there occurs a union of the ganglia of opposite 
 sides by means of diffusely scattered cells located just caudal to the 
 optic chiasm. In man this union is very definite, in macacus less 
 definite, while in the other two animals it is rudimentary and due to the 
 presence of a very few widely scattered cells between the two ganglia. 
 In the cat even this feebly developed fusion is not present in all 
 individuals. 
 
 4. Although the phylogenetic series from man to the cat is too short 
 to be of much service in revealing the phylogenetic development of so 
 constant a cell group as the basal optic ganglion, it is apparent that 
 as we descend the series the two parallel cell columns become separate 
 (more closely united in man than in macacus, and separate in the lemur
 
 Nuclei Tuberis Laterales and the Ganglion Opticum Basale. 13 
 
 and cat) ; the same is trae as to the fusion of the ganglia of opposite 
 sides (definite in man, less so in macacus, barely present in the lemur 
 ajid not always present in the cat.) 
 
 Cell type of the basal optic ganglion. 
 
 It is not my intention to attempt to give in words the characteristics 
 of the cells of the basal optic ganglion which have been already satis- 
 factorily shown in the illustrations. I shall confine myself to pointing 
 out the fundamental characters of these cells, and when we have thus 
 become familiar with the main features of the cell picture we shall be 
 in a position to discuss the relations of these cells to those of other 
 groups. The water color reproductions of the cells of the basal optic 
 ganglion, and of all other cells thus illustrated (Figs. 38 to 58), were 
 all drawn from cross-sections of the brain. In Series D of man, in 
 which the plane of section differs widely from that of a cross-section, 
 I have been unable to observe any important difference in the appear- 
 ance of the cells of the basal optic ganglion, and it is my opinion that 
 the plane of section does not affect to any appreciable extent the appear- 
 ance of the majority of these cells. The cells of the basal optic gang- 
 lion and of all other groups illustrated are as characteristic in Series D 
 a.s in Series AC. However, I have not made a careful enough study 
 of this point to state definitely that the cell character is absolutely 
 unaffected by differences in the plane of section. The cells of the basal 
 optic ganglion in all four animal forms (Figs. 38, 44, 50 and 55) are 
 large polygonal cells which possess very coarse processes. These pro- 
 cesses, as is shown in Fig. 37, form an intercellular feltwork; under 
 low power one would hardly suppose that this feltwork was composed 
 of cell processes, and the ganglion appears to be characterized not only 
 by its typical cells but also by the presence of a distinctive intercellular 
 substance. These processes are practically colorless, and are in Fig. 
 37 represented as blue because at this magnification they could not be 
 shown in any other manner. The cell processes in Figs. 38, 44, 50 and 
 55 are not as long as they appear in the actual preparations. Another 
 fundajnental character of these cells is the distribution of the Nissl 
 substance, nearly all of which is massed on the periphery of the cell ; 
 this peripheral distribution is not equal in all portions of the periphery, 
 since the depth is much greater at certain points, and at other points 
 of the periphery the iNrissl substance may be almost entirely absent.
 
 14 , Edicard F. Malone. 
 
 Of course the distribution of the Xissl substance will appear different 
 according to the location of the optical section, and the figures show this 
 to some extent, although they represent to a certain extent a combination 
 of different optical sections. Although minor differences in cell char- 
 acter exist in different animals, the cell group is phylogenetically 
 so old that in the relatively brief interval between man and the cat no 
 changes in cell character have occurred which are sufficiently funda- 
 mental to permit of correlation with the phylogenetic position of the 
 corresponding animals ; this point will be discussed later in connection 
 with the nuclei tuberis, where such a relation between cell character 
 and the phylogenetic position of tlie corresponding animal actually 
 exists. The structure of the cells of the ganglion opticum basale is 
 such as to exclude the possibility of these cells being motor, whereas 
 the large size of the cells probably indicates (as Dr. Donaldson has 
 suggested to me) that they either receive impulses converging from 
 many sources or distribute impulses over an extensive region. 
 
 Separation of the hasal optic ganglion from surrounding cell groups 
 through differences in cell ch^aracier. 
 
 A subdivision of any portion of the nen'ous system based merely 
 upon the splitting up of gray matter through the mechanical agency of 
 fiber masses is, except in certain cases, valuable merely with reference 
 to orientation; such a subdivision should therefore be considered as a 
 crude (although necessary) beginning to be followed by a more careful 
 study of the region involved in which the cell character of the various 
 groups receives careful attention. Moreover it is highly unsatisfactory 
 for an author to state dogmatically that he recognizes the presence of 
 certain cell groups upon the basis of certain differences of cell char- 
 acter, concerning which he is either silent or else treats in a superficial 
 manner; for we are left in doubt as to whether these differences in cell 
 character are really fundamental, and as to the degrees of relationship 
 between the cell characters of various cell groups. When we have de- 
 scribed the location and extent of various cell groups and have made it 
 possible to recognize this by means of a definite cell type, and when we 
 have clearly shown the differences and similarities in cell character of 
 various cell groups, and have pointed out the phylogenetic development 
 and relations of these groups, then and not until then will there be a 
 basis for experimental work which will help solve many important
 
 Nuclei Tuheris Laterales and the Ganglion Opticum Basal e. 15 
 
 questions as to the elementary mechanisms of the nervous system. In 
 the case of the basal optic ganglion, therefore, I shall not be content 
 with having pointed out its location and extent, but shall proceed to 
 compare the character of its cells with that of the cells of surrounding 
 groups ; such a comparison serves not only to set aside this cell group 
 as different from other cell groups by virtue of differences in cell 
 character, but it enables us also to recognize certain similarities of cell 
 character between the cells of the ganglion and the cells of other groups, 
 so that different degrees of relationship between the different cell 
 groups may be provisionally stated. 
 
 I shall first point out the differences in cell character between the 
 cells of the basal optic ganglion and those of the surrounding groups 
 so .as to complete the picture of this cell group ; afterwards the relations 
 of various cell groups will be discussed (but not until the nuclei 
 tuberis have been described). There is no diiSculty in distinguishing 
 the cells of the basal optic ganglion from those of the nuclei tuberis 
 and of the substantia grisea ventriculi tertii. This is shown by a 
 reference to the corresponding figures 38, 39 and -iO; 4-1, 45 and 46; 50 
 and 51; 55 and 56; moreover the difference between the cells of the 
 basal optic ganglion and those of the substantia grisea is well shown in 
 Fig. 37. 
 
 The cells of the basal optic ganglion may be readily distinguished 
 also from those of the nucleus tubero-mammillaris, although both types 
 of cells have a certain similarity; the failure to recognize the unity of 
 the cell complex known as the nucleus tubero-mammillaris (mammillo- 
 infundibularis) and to clearly separate it from the basal optic ganglion 
 is one of the most important causes for our meager knowledge of this 
 region. Considering first the relations of these tvi'o cell groups as to 
 location, it is evident from the illustrations that they are closely related 
 only for a short distance. Moreover the cells of the nucleus tubero- 
 jnammillaris are for the most part rather diffusely scattered, while 
 those of the basal optic ganglion are densely packed together and the 
 cell group is further characterized by the intercellular feltwork formed 
 by the coarse colorless cell processes ; these two points may be observed 
 by comparing Figs. 36 and 37. Comparing the cell type of the two cell 
 groups the fundamental difference found in all four animals involves 
 the appearance of the Nissl substance ; in both forms the Nissl substance 
 is located principally on the periphery of the cell, but in the cells of the 
 basal optic ganglion the massing of the Nissl substance on the periphery 
 
 2
 
 16 Edward F. Malone. 
 
 is extreme, whereas in the cells of the nucleus tubero-mammillaris the 
 N"issl substance is not so densely packed together on the periphery and 
 more of it is present in the central portion of the cell. In other words 
 the Nissl substance in the cells of the nucleus tubero-mammillaris is 
 more diffusely distributed throughout the entire cell. The difference 
 in cell type obtains throughout the entire extent of both nuclei. A 
 study of the illustrations shows that while other differences in cell type 
 occur in certain animals the fundamental difference consists in the 
 mode of distribution of the Nissl substance (Figs. 38 and 41 ; 44 and 
 47; 50 and 52; 55 and 57). 
 
 The separation of the basal optic ganglion from the nucleus paraven- 
 tricularis hypothalami on the basis of cell type is difficult. Fortunately 
 both are in all four animals sharply circumscribed, and although the 
 extremities of these two cell columns approach one another (Fig. 10), 
 they never actually fuse. A study of the location of these two cell 
 groups will show their axes are almost at right angles to each other. 
 In a former article I made the statement that the cells of the basal 
 optic ganglion in man could not be distinguished from those of the 
 nucleus paraventricularis, but a more careful study of the cell groups 
 in man together with their study in other animals proves that this 
 statement is not correct, although not far from the truth. As a matter 
 of fact I have been unable to observe any one fundamental difference 
 between these two types of cells which is clearly shown in all four 
 animal forms, although in each animal some differences are present 
 which always make a distinction possible. There is one difference which 
 holds fairly well for all forms, and this is the same difference that was 
 so evident between the cells of the basal optic ganglion and those of the 
 nucleus tubero-mammillaris, namely, the Xissl substance is more 
 densely massed on the periphery of the cell in the case of the basal optic 
 ganglion, while in the cells of the nucleus paraventricularis the Nissl 
 substance is more uniformly distributed throughout the cell. In man 
 ( Figs. 38 and 43 ) this difference is not marked, but in the basal optic 
 ganglion (Fig. 38) the Xissl substance on the ]3eriphery forms in part 
 large, irregularly placed masses of granules, while in the cells of the 
 nucleus paraventricularis (Fig. 43) the granules are smaller, more 
 uniform in size and do not stain so intensely; at the same time the 
 c-entral portion of the cell is somewhat more deeply stained (due to more 
 Nissl substance) than in the case of the cells of the basal optic ganglion. 
 Other differences in man are as follows: the cells of the basal optic
 
 Nuclei Tuberis Laterales and the Ganglion Opticum Basale. 17 
 
 ganglion are somewhat larger (the smallest cell illustrated in Pig. 38 
 shows only a small portion of a cell in optical section), the nuclear 
 membrane is not so definite, and the nucleus is larger. In macacus 
 (Figs. 44 and 49) the difference in distribution of the Nissl substance 
 is less than in any of the other animals, but even here the Nissl sub- 
 stance in the cells of the l>asal optic ganglion is more nearly confined 
 to the periphery, is denser here and not so dense in the interior portion 
 of the cell as in the case of the other cell group; this results in a 
 sharper differentiation between peripheral and central portions of the 
 cell in the case of the ba&al optic ganglion. Other differences in 
 macacus are: the cells and cell nuclei of the basal optic ganglion are 
 smaller than those of the nucleus paraventricularis (the opposite is 
 true in man), and the cells of this latter nucleus are rather piriform. 
 In the lemur (Figs. 50 and 54) the difference in distribution of the 
 Nissl substance is marked, the differentiation of the cell into peripheral 
 and central portions being much sharper in the cells of the basal optic 
 ganglion. In the cat (Figs. 55 and 58) the difference between per- 
 iphery and center of the cell is again more marked in the cells of the 
 basal optic ganglion, since in the cells of the other group the central 
 portion of the cell (including the cell nucleus) stains deeply (Fig. 58). 
 Note also in the cat that, just as in man, the cells of the basal optic 
 ganglion are larger, but unlike those of man have smaller cell nuclei 
 than the cells of the nucleus paraventricularis. 
 
 The resemblance of the cell type in the case of the basal optic gang- 
 lion and the nucleus paraventricularis hypothalami is therefore very 
 close, but in all four animals studied the cells of the basal optic gang- 
 lion were found to be more sharply differentiated into a peripheral zone 
 containing dense masses of Nissl substance and an inner portion rela- 
 tively free from this substance, whereas in the cells of the nucleus par- 
 aventricularis hypothalami the Nissl substance was not so sharply 
 differentiated as to its distribution. It will be recalled that within the 
 cells of the nucleus tubero-mammillaris this differentiation was even 
 less marked than within those of the nucleus paraventricularis. 
 
 Another cell group must be distinguished from the basal optic gang- 
 lion. This is the nucleus ansae peduncularis of Meynert (ganglion 
 basale of Kolliker) . It occurs in man, macacus and the lemur, but I 
 was unable to find it in any of my three series of the cat. Its location 
 and extent may be seen in the different illustrations. In all three 
 animals in which it occurs it is readily distinguished from the basal
 
 18 Edward F. Malone. 
 
 optic ganglion by tlie large size of its cells. In man, as was pointed 
 out by Kolliker, the cells of the ganglion basale may be readily separated 
 from those of the basal optic ganglion by the fact that they are heavily 
 pigmented (Fig. 42). In a previous article I have discussed the value 
 of pigmentation as a basis of distinguishing different cell types. 
 Yellow pigment occurs in certain nerve cells of the adult human and 
 increases with age, but this occurrence and increase in amount is not 
 errntic, but behaves differently with respect to different types of cells. 
 Some types of cells never contain pigment, others always contain it, 
 while still other types may contain none or under other conditions may 
 contain a small or even a moderate amount. But however the total 
 amount of yellow pigment in the brain of different adults may vary, 
 the amount in each specific type of cell retains the same relation to that 
 of every other type of cell ; in other words, the relative amount of pig- 
 ment in the different types of cells is constant in all individuals. Note 
 that this pigmentation of the cells of the ganglion basale in man is 
 accompanied by an additional characteristic (large size) and that the 
 homologous cells of macacus and the lemur are entirely different from 
 those of other cell groups, although pigmentation is of course lacking. 
 This question of the relative amount of yellow pigment as a character- 
 istic of certain types of cells which differ also in other respects will be 
 noted again in connection with the nuclei tuberis. This yellow pigment 
 should not be confounded with the brown pigment which occurs in the 
 cells of the substantia nigra and elsewhere. 
 
 Now that the differences in cell type between the cells of the basal 
 optic ganglion and those of surrounding groups have been studied it 
 woiild be desirable to consider the relationship of these various cell 
 groups from the standpoint of similarities in cell character. This 
 consideration, as well as that of the literature, must however be post- 
 poned until the nuclei tuberis have been described, since such a com- 
 parison of the various cell groups demands a familiarity with the 
 nuclei tuberis and the substantia grisea of the third ventricle. 
 
 Nuclei Tuberis Later-^les. 
 
 Under the name of nuclei tuberis laterales I shall describe certain 
 cell groups of the pars optica hypothalami (telenceplialon) which I 
 have formerly considered very briefly in my monograph on the human 
 dieneephalon. Although a number of authors have written of nuclei
 
 Nuclei Tuheris Laterales and the Ganglion Opticuin Basale. 19 
 
 tuberis, except in one case, there is absolutely no reason to believe that 
 any of them have had in mind the characteristic nests of cells which 
 will now be described. Kolliker undoubtedly saw these nuclei, 
 although, as will appear later, his description is faulty and his figures 
 incorrectly labeled ; the result is so confusing that a careful study of 
 this region is necessaiy to appreciate the value of Kolliker's observa- 
 tions. In the previous brief description I employed the name nuclei 
 tuberis, but in the present article adopt the name nuclei tuberis later- 
 ales. The name nuclei tuberis is nor distinctive and might be used to 
 indicate any cell group whatsoever located in the tuber cinereum; as 
 a matter of fact it has been so used, and the resulting confusion may be 
 imagined when one considers how many different cell groups lie in the 
 tuber cinereum. This confusion is not one merely of names, but 
 involves the failure to distinguish (regardless of names) various cell 
 groups of the tuber cinereum which in properly prepared material may 
 be readily distinguished not only through differences in location but 
 also through striking differences in cell type. 
 
 Location and extent of the nuclei tuberis laierales. 
 
 The nuclei tuberis laterales consist in man and macacus of several 
 nests of cells located on the periphery of the tuber cinereum. In the 
 lemur in this location one sees the beginning differentiation of these 
 cell groups from the cells which surround them, although these differ- 
 ences are so slight that I have not attempted to show them in the illus- 
 trations. In the cat these cell groups are absent. 
 
 MAN. 
 
 In Series AC of man the nuclei tuberis laterales are shown in Figs. 
 1 to 8; the size of the cells, however, has been so much reduced that 
 they should be examined with the aid of a hand lens. In Pig. 1 two 
 indentations are shown on the base of the brain which should be 
 noted. One indentation marks off on the base of the brain the medial 
 boundary of the pes pedunculi, while the more medial indentation 
 forms the superficial boundary between the tuber cinereum and the 
 nucleus tubero-mammillaris (nucleus mammillo-infundibularis), this 
 nucleus lying between these two fissures. The area bounded by these 
 two fissures ventrally, and by the columna fornicis dorsally, is of 
 especial importance in describing the location of the nuclei tuberis.
 
 20 Edward F. Malone. 
 
 On examining Figs. 1 to 6 these two fissures are seen to be practically 
 parallel, and the area bounded by them and by the fornix column under- 
 goes as we pass orally the following change : in the caudal portion (Fig. 
 1 ) this area is occupied by the cells of the nucleus tubero-mammillaris, 
 whose caudal pole constitutes the lateral ganglion of the maniraillary 
 body; passing further orally these large cells are replaced (especially 
 those situated ventrally and medially) by the small cells of the nuclei 
 tuberis laterales and of the substantia grisea. Accordingly the oaudal 
 portion of this region is continuous with the lateral ganglion of the 
 mammillary body, whereas orally it gradually becomes the lateral 
 portion of the tuber cinereum. In this region included between these 
 two fissures ventrally, the pes pedunculi laterally, and the fornix 
 column dorsally, lies by far the greater portion of the cells of the nuclei 
 tuberis laterales. Referring to Figs. 1 to 8 the location and extent of 
 these cell groups are evident, and only a few comments will be necessary. 
 The nuclei tuberis laterales consist of a variable number of well circum- 
 scribed nests of small cells, and can be distinguished even without the 
 aid of their characteristic cell type. These different cell nests are 
 more or less connected with one another, and their separation is to be 
 regarded probably as dependent upon the mechanical influence of fiber 
 masses, since their partial fusion, identical cell type, -and variable 
 number would seem to exclude the possibility of these separate groups 
 having different functions. The cell nests of the nuclei tuberis later- 
 ales are practically free from cells of surrounding groups. 
 
 In considering the second series of man (Series D, Figs. 11 to 15) it 
 is absolutely essential to understand the plane of sectionr; this has 
 already been described in considering the basal optic ganglion. It will 
 suffice to point out that Series D differs from Series AC in that the 
 plane of section of the former passes more caudally as it passes from 
 dorsal to ventral ; and that in it the opposite sides of the brain are cut 
 asymmetrically, in that the lateral portion of each section of Series D 
 is situated further oral than the medial portion of the corresponding 
 section. As was previously noted the plane of section of Scries D is the 
 only one which can pass through the right optic tract at right angles 
 to its long 'axis ; this relation to the optic tract will render a clear picture 
 of the plane of section relatively easy. Bearing this plane o^ section 
 in mind a study (with the aid of a hand lens) of Figs. 11 to 15 will 
 show that the location of the nuclei tuberis laterales in Series D is 
 practically the same as in the preceding Series AC, Figs. 1 to 8. Of
 
 Nuclei Tuberis Laterales and the Ganglion Opticum Basale. 21 
 
 the two fissures referred to previously, the lateral (just medial to the 
 pes pedunculi) does not occur, since the obliquity of the plane of section 
 is such that it appears only in sections situated further caudally. The 
 medial one of these two fissures, however, appears in every section from 
 Figs. 11 to 15, and as in the other series, forms superficially the medial 
 boundary of by far the greater portion of the nuclei tuberis laterales. 
 
 MACACUS RHESUS. 
 
 In macacus the location of the nuclei tuberis laterales is similar to 
 that in man, except that in most sections only one cell group is shown, 
 and the cell mass does not invade the medio-ventral portion of the 
 tuber. A comparison of Figs. 19 to 22 (macacus) with Figs. 1 to 8 
 (man) will show that in both animals the nuclei differ but little as to 
 location. The two parallel furrows on the base of the brain are shown 
 in Figs. 19 and 20. 
 
 In both man and macacus the nuclei tuberis laterales have, accord- 
 ingly, practically the same location, and by far the greater portion is 
 situated in the previously described region; this region, bounded ven- 
 trally by the two parallel furrows, laterally by the pes pedunculi, and 
 dorsally by the fornix column, is occupied caudally by the nucleus 
 tubero-mammillaris, and the nuclei tuberis laterales displace this 
 nucleus as one passes orally, and after the optic tract has fused with 
 the tuber, the nuclei tuberis are continued orally in the angle between 
 the pes pedunculi and the optic tract. Since the purpose of this paper 
 is in part to give a foundation for experimental study, it should be 
 noted that the main cell mas^ of fue nuclei tuberis laterales may be 
 accurately loaded in the intact brain just beneath the surface of the 
 most lateral portion of the tuber cinereum, between the two parallel 
 furrows and just caudal (medial) to the optic tract. In toluidin blue 
 sections the nuclei may be distinguished as circumscribed, lightly 
 staining areas by means of the unaided eye. As previously stated no 
 trace of the nuclei tuberis laterales is present in the cat, while in the 
 lemur they are indicated so faintly that it seems best not to attempt to 
 represent them in the illustrations. 
 
 Separation of the nuclei tuberis laterales from surrounding cell groups 
 through differences in cell character. 
 The nuclei tuberis laterales are readily distinguished from the sur- 
 rounding cell groups not only in that they are sharply circumscribed 
 cell masses having a constant location, but also through their charac-
 
 22 Edward F. Malone. 
 
 teristic cell type. A reference to Fig. 36 will serve as an introduction 
 to the further consideration of the differences in cell character. On 
 comparing the cell character of the nuclei tuheris laterales of man and 
 macacus (Figs. 39 and 45) with that of the other cell groups shown 
 in the illustrations it will at once be evident that this cell type differs 
 radically from all others except that of the substantia grisea of the 
 third ventricle (Figs. 40 and 46) ; this similarity is of importance, 
 and its significance w'ill be considered when the relationship of the 
 various cell types is discussed. Comparing the cell type of the nuclei 
 tuberis laterales in man (Fig. 39) and in maoacus (Fig. 45) it is 
 evident that aside from the 2>resence of pigmentation in man (of course 
 not to be expected in macacus) there is no essential diiference. The 
 fundamental characteristic of this cell type in both forms lies in the 
 appearance of the cytoplasm, which contains an extremely small amount 
 of Xissl substance in the form of fine granules ; the relation of cell 
 nucleus to cytoplasm in respect to volume is 'also practically the same, 
 although in macacus the cytoplasm is relatively less, a change that is 
 to be expected, as will appear later. In describing the cells of the 
 nucleus ansae peduncularis (p. 18) I pointed out the fact that if two 
 cell groups differ as to the amount of yellow pigment in their cells they 
 will also differ in other respects, and that homologous groups in lower 
 animals (where all pigmentation is absent) will also appear different. 
 Accordingly the presence of densely packed yellow granules in the cells 
 of the nuclei tuberis laterales of man is not the only distinctive charac- 
 teristic of these cells, but is connected with other characteristics, 
 common also to the homologous cells of macacus ; these characteristics 
 involve the appearance of the cytoplasm and make it possible to separate 
 these cells without difficulty from those of surrounding groups. If we 
 study the cell type of the substantia grisea in all four fonns (Figs. 40, 
 46, 51, and 56) we find the cell type practically unchanged, since in all 
 fonns the cells of this group are characterized by the extremely small 
 amount of cytoplasm and relatively large nucleus; in many cells the 
 cytoplasm is almost absent, and one sees only a large nucleus and 
 perhaps one or more cell processes. 
 
 Accordingly the cells of the substantia grisea may be readily separated 
 from those of all portions of the nuclei tuberis Laterales not only 
 through their smaller size, but also through the minute amount of 
 cytoplasm in relation to the size of the cell nucleus; however, on the 
 border line between these two types of cells occur occasionally transi-
 
 Nuclei Tuberis Laterales and the Ganglion Opticum Basale. 33 
 
 tioD types, whose significance will be discussed later. The nuclei 
 tuberis laterales must be sliarply distinguished from certain regions of 
 the substantia grisea in whicli the cells are more densely packed 
 together than in other portions of the substantia grisea. Differences 
 in the density of different portions of the substantia grisea may be seen 
 in many sections, and very definite areas of closely packed cells are 
 shown in Figs. 17, 23 and 34. While in such dense masses the cells 
 are possibly somewhat larger than the remaining cells of the substantia 
 grisea, I do not consider the difference in cell type clear enough to 
 justify one in setting aside such dense cell masses as distinct nuclei. 
 Whatever the significance of these denser cell masses of the substantia 
 grisea, one fact is certain : they diifer both in location and in cell 
 type from the nuclei tuberis laterales; the radical difference between 
 these two kinds of cell groups should be clearly recognized. Of course 
 the structure of the cells of the nuclei tuberis laterales as well as that 
 of the cells of the substantia grisea is radically different from that of 
 motor cells, while the small size of both types of cells would indicate 
 that the connections of each cell were very limited; this correlation 
 between cell size and the extent of the connections of each cell has 
 already been referred to. 
 
 Eelationship Of The Various Cell Groups To Oxe Another. 
 
 From the previous description of the various cell groups together 
 with a reference to the figures showing their cell types, it will be evident 
 that the cell groups herein considered fall naturally into two classes: 
 
 1. Those composed of small cells, including the substantia grisea, 
 and the nuclei tuberis laterales, and 
 
 3. Those composed of large cells, including the basal optic gang- 
 lion, the nucleus paraventricularis hypothalami, the nucleus tubero- 
 mammillaris, and perhaps also the nucleus ansae peduncularis (gang- 
 lion basale). 
 
 Taking up the first class of nuclei a most interesting relation will 
 be found to exist between the substantia grisea and the nuclei tuberis 
 laterales. The cells of the substantia grisea are the most primitive 
 of the hypothalamus. The constancy of cell type in all four animal 
 forms would suggest this, but the strongest evidence is derived from the 
 nature of the cell tyjie. The cell type of the substantia grisea, ^Wth its 
 scanty cytoplasm, resembles that of embryonic cells; it occurs often in
 
 24 Edward F. Malone. 
 
 invertebrates ; it is the only tj'pe in the hypothalamus which approaches 
 that of the neuroglia cells, and transition forms occur concerning which 
 one is in doubt as to whether they are neuroglia cells or cells of the sub- 
 stantia grisea ; the cells are most abundant near the ventricle from 
 whose border all cells of the hypothalamus have arisen, while laterally 
 they are replaced by more highly differentiated types of cells. It is by 
 no means certain that all or even any of the cells of the substantia 
 grisea are functional. That the cells of the nuclei tuberis laterales have 
 arisen from those of the substantia grisea is certain ; if we look in the 
 lemur at the regions where in higher forms the nuclei tuberis laterales 
 occur, we see that the type of cell is somewhat different from the sur- 
 rounding cells of the substantia grisea ; this difference increases in 
 maeaeus, and still more in man, while in the cat there is no indication of 
 any new group. It is thus evident that the most recent cell groups of 
 the hypothalamus (nuclei tuberis laterales) have arisen directly from 
 the oldest cell group (substantia grisea), and that new cell groups are 
 not necessarily formed by the further histological differentiation of 
 portions of cell groups which are already highly developed, thus result- 
 ing in a very highly differentiated cell type, but that on the contrary 
 new cell groups may arise from cell masses which have remained primi- 
 tive. In other words the fact tliat a cell group is of recent origin does 
 not give any information as to the extent of its histological differentia- 
 tion, since this differentiation depends also upon the development of the 
 cell group from which the new group arises. 
 
 In considering the second class of cell groups in the hypothalamus, 
 all of which are composed of large cells, the nucleus ansae peduncularis 
 requires only a brief consideration. The cell t}-pe of this cell group 
 differs considerably from that of other groups, and the nucleus is prob- 
 ably of much more recent phylogenetic origin, since I have not as yet 
 been able to find it in the cat ; in order to consider the relations of this 
 cell group a study of the neighboring regions of the telencephalon will be 
 necessary, and I shall content myself with having shown that it is clearly 
 distinguishable from the basal optic ganglion, and with having given 
 the relative location of both groups of cells. 
 
 Concerning the other three cell groups in this class (basal optic gang- 
 lion, nucleus paraventricularis hypothalami, and nucleus tubero-mam- 
 millaris), the phylogenetic series from the cat to man is far too short 
 to give such satisfactory results as in the ease of the nuclei tuberis 
 laterales, since the former three groups are of much older origin : certain
 
 Nuclei Tuberis Laterales and the Ganglion Opticum Basale. 25 
 
 relationships may, however, be established. Of these three cell groups 
 the basal optic ganglion is composed of cells whose type is by far the 
 most constant in the different animal forms, and the volume of this 
 cell group is reduced in the lower forms to a less extent than is true of 
 the two other nuclei; moreover, while the cell tj-pe of the basal optic 
 ganglion approaches closely that of the nucleus paraventricularis. there 
 are no transition forms between its cell type and that of the nucleus 
 tubero-mammillaris or that of the substantia grisea; finally the basal 
 optic ganglion is almost entirely free from the invasion of neighboring 
 cells. All of these considerations make it seem probable that of the 
 three nuclei the basal optic ganglion is the oldest and most highly 
 developed. Closely related to the basal optic ganglion is the nucleus 
 paraventricularis ; it is more reduced in size in the lower forms than 
 is the basal optic ganglion ; between its cell type and that of the nucleus 
 tubero-mammillaris transition forms occur, and between its cells lie 
 many cells of the substantia grisea. While these facts make it probable 
 that the nucleus paraventricularis is possibly not so old a cell group as 
 the basal optic ganglion, they enable one to state positively that the 
 nucleus paraventricularis is related on the one hand to the basal optic 
 ganglion and on the other to the nucleus tubero-mammillaris. The third 
 cell group, the nucleus tubero-mammillaris. is like the nucleus paraven- 
 tricularis much reduced in size in the lower animals ; it differs strik- 
 ingly from both the other cell groups in being not sharply circum- 
 scribed, but on the contran- is very diffuse,"^ extending through a large 
 portion of the h}-pothalamus ; it is invaded accordingly by the cells of 
 the substantia grisea to a much greater extent than is the nucleus para- 
 ventricularis. Finally the cell type of the nucleus tubero-mammillaris 
 shows a transition on the one hand to that of the nucleus paraventric- 
 ularis and on the other to that of the substantia grisea. Therefore the 
 nucleus tubero-mammillaris, while related to the nucleus paraventricu- 
 laris, is related to the substantia grisea in three ways : transition forms 
 of cells occur, the cells of both groups are intimately intermingled, and 
 both cell groups are widely and diffusely distributed. The nucleus 
 tubero-mammillaris is therefore the least highly developed of the three 
 groups (owing to its relation to the substantia grisea), but whether it is 
 also the most recent group cannot be decided here: it might just as 
 well be an old and relatively stationary group, and to decide this point 
 a longer phylogenetic series must be studied.
 
 26 Edivard F. M alone. 
 
 The relationship of these various cell groups of the hypothalamus 
 may be summarized as follows : 
 
 1. The substantia grisea ventriculi tertii is the oldest, least highly 
 developed, and the most diffusely distributed cell group of the hypothal- 
 amus. From it have been differentiated at least two (and probably all) 
 of the four cell groups that lie more or less embedded in it. Certain 
 portions of the substantia grisea appear slightly different from the 
 rest, in that the cells are here crowded closely together. Whether any or 
 all of the cells of the substantia grisea take part in conducting nervous 
 impulses, or whether this group serves merely as material from which 
 more highly differentiated cell groups are formed, is not known. 
 
 2. From the substantia grisea at least two different cell groups have 
 arisen ; these two groujjs of cells have developed along diverging lines. 
 One of these two types constitutes the nuclei tuberis laterales ; it is by 
 far the youngest cell group in the hypothalamus, the first indication of 
 its presence occurring in the lemur. The second cell group wiiich has 
 developed from the substantia grisea is the nucleus tubero^mammillaris ; 
 its cell type differs radically from that of the nuclei tuberis laterales, 
 and it is a much older cell group. 
 
 3. Along the same line as the nucleus tubero-mammillaris, and to a 
 higher degree of histological differentiation, have developed the nucleus 
 paraventricularis and the basal optic ganglion ; the nucleus paraventric- 
 ularis is intermediate between the other two nuclei. From such a 
 short phylogenetic seines as that from man to the cat it is impossible to 
 determine whether all three of these nuclei have developed iiidej)en- 
 dently from the substantia grisea or whether the other two nuclei liave 
 developed from the nucleus tubero-mammillaris and thus only indirectly 
 from the substantia grisea. 
 
 4. A more extensive series of animal forms should enable us to under- 
 stand such relations better; but a study of such material cannot be 
 expected to yield satisfactory results unless the different cell groups 
 be distinguished and unless the question of the relationship as ex- 
 pressed by the cell type be borne in mind. 
 
 5. Valuable results may be expected also from a study of the histo- 
 genesis of these various cell groups as revealed in the later stages of 
 development of the human brain; little, however, is to be expected 
 from such material unless it be fixed and stained witli reference to 
 the demonstration of cell structure in the nervous system, and the 
 methods usually employed are not well adapted to this purpose.
 
 Nuclei Tuberis Laterales and tJie Ganglion Opticum Basale. 27 
 
 Conclusion. 
 
 I have described with tlie aid of illustrations the location ^and extent 
 of the basal optic ganglion and nuclei tuberis laterales in four forms 
 of manxiuals, and have showii their relations to the surrounding cell 
 groups of the hypothalamus; moreover the cell types of these two cell 
 groups have been shown to differ radically, and the differences in cell 
 character have been pointed out through which one can readily and 
 accurately distinguish each of these two cell groups from those which 
 surround it. Accordingly the principal purpose of this study has been 
 accomplished, namely, to prepare a foundation of such a character as 
 to make further study possible. It is now possible to study in the same 
 manner neighboring portions of the telencephalon, or to e.vtend with 
 relative ease the present study of the hypothalamus through a much 
 longer phylogenetic series than the critical nature of this study has 
 permitted ; the relations of the various cell groups of the hypothalamus 
 can be studied also from the standpoint of histogenesis ; or the nature 
 of the cell processes and the character of the nerve endings around the 
 cells of each group may be determined. But the greatest use of this 
 study is that it makes possible intelligent experimental investigation 
 of the hypothalamus. Experimental workers have been inclined to 
 pass over differences in cell character and to make little or no attempt 
 to correlate cell function with cell character, and fail to clearly recognize 
 a fact to which I have, in a previous article, called attention in the 
 following words : " The histological character of a nerve cell is an indi- 
 cation of its function. Differences in connections with portions of the 
 organism which differ merely in spatial relations do not involve a 
 difference in the character of the nerve cells, but are associated merely 
 with the location of the nerve cell ; for instance, arm and leg muscles, 
 flexors and extensors are all innervated by the same tyix; of cell, 
 although such differences in peripheral connections correspond to the 
 differences in the position of the corresponding nerve cells." It is there- 
 fore evident that experimental work which detemiines the connections 
 of various portions of a given region with different portions of the 
 organism, without taking into consideration differences of cell character, 
 fails to distinguish between differences of connection dependent merely 
 upon spatial differences and those differences of connection which 
 involve differences in cell activity, such cell activity being indicated, 
 not only in the nervous system but in all portions of the entire organism.
 
 28 Edward F. Malone. 
 
 by a definite type of cell character. That these two aspects of the func- 
 tion of the nervous system might be successfully studied in the hypothal- 
 amus by the use of experimental methods has been the controlling 
 factor in determining the nature of this article. 
 
 But in addition to preparing a foundation for exj3erimental work an 
 attempt has been made to partially analyze the different elements in- 
 volved in the histological complex of each cell type and to note in 
 homologous cell groups of various animals what elements of cell charac- 
 ter are constant ; moreover in different cell groups of the same animal 
 those elements of the entire complex of cell character which are common 
 to the cells of two or more groups have been noted, and by this means an 
 attempt has been made to form a provisional, although crude, idea of the 
 degree of functional relationship among the different cell groups. At 
 first sight such an attempt might appear premature, or it might even 
 seem as though the task of the histologist were finished when he had 
 pointed out the existence and location of the various cell groups, and 
 that the determination of the functional relations of different cell groups 
 might be safely left to the experimental worker. Such a conclusion, 
 however, is not justified. Many regions of the brain are so complicated 
 as to make the results of experimental investigation extremely vague, 
 and in more suitable regions it is only under fortunate conditions that 
 we are informed of the functional relations of a cell group except that it 
 plays some unknown part in a complex mechanism underlying a com- 
 plex function. In solving the relations of the different components of 
 the various mechanisms of the nervous system histology must play an 
 important and perhaps the principal part, since in the nervous system, 
 as in the rest of the organism, it enables us to locate and to correctly 
 state the function of every cell group of one definite cell type, if the func- 
 tion of this cell type has in other cell groups been previously clearly 
 shown ; that, however, it is not self-sufficient is evident. 
 
 But the neuro-histologist has a much more difficult task than that of 
 interpreting in a general way the differences and similarities of cell 
 character in different cell types. The cell types of certain cell groups 
 resemble one another through possessing certain features of cell char- 
 acter common to all, and on the other hand differ in respect to other 
 features of cell character; that such similarities and differences of cell 
 type are an indication of corresponding similarities and differences of 
 function I have already clearly shown in two cases. I have shown (Am. 
 Jour. Anat., 1913) that although the three different types of muscle are
 
 Nuclei Tuheris Laterales and the Ganglion Opticum Basale. 29 
 
 supplied by three distinct types of nerve cells (p. 2), these three types 
 of nerve cells possess a fundamental similarity of cell character ; accord- 
 ingly the cell type varies as the cell function. In the second place I 
 have shown (Anat. Rec, 1913) that all cells supplying striated muscle, 
 whether directly (anterior horn cells, etc.) or more or less indirectly 
 (large pyramidal cells, cells of Deiters' nucleus, etc.) possess certain 
 fundamental features of cell character in common. This fundamental 
 character makes its appearance in the first cell of any efferent chain 
 which is set aside for the innervation of striated muscle and is present 
 in all cells of such a series, however many neurones may intervene 
 between the cell in question and the striated muscle fiber; on the other 
 hand the cells in such a voluntary motor chain differ in relation to their 
 position in the chain. It is thus evident that the particular group of 
 elements of cell type which corresponds to the motor function of the 
 cell may be distinguished from other elements of cell type which must 
 correspond to the other known or unknown influences. The feature 
 of cell character which corresponds to motor function is the arrangement 
 of Nissl substance in relatively coarse, discrete bodies; while the large 
 size (which is usually supposed to be common to motor cells) is not 
 directly related to the motor function, but is an element of cell character 
 which is probably (as stated previously) an indication of the extensive 
 connections of the cell, in that such a large cell receives impulses from 
 (sphere of reception) or sends impulses to (sphere of influence) a large 
 territory. 
 
 Accordingly each type of nerve cell should be regarded not as distinct 
 from all other types, but as the result of different influences (due to 
 the various relations of the nerve cell to other neurones and to the other 
 portions of the organism) ; when so viewed the manifold variations of 
 cell type, including cells of the most diverse types, as well as those of 
 extremely similar types, do not appear to offer insuperable difficulties, 
 but on the contrary these confusing relations of cell types will enable 
 us eventually to work out the details of various nervous mechanisms. 
 After the simultaneous analysis, by the aid of many methods, of the 
 histological characteristics and the functional relations of difi'erent 
 types of cells, and after the recognition of the result of any given 
 functional relation upon the histological character of the cell, the neuro- 
 histologist will be able to take a further step. He will not only deter- 
 mine one portion of the functional relation through the presence of 
 certain features of cell character (which is already possible in the ease
 
 30 Edward F. Malone. 
 
 of various types of motor cells), but he tvill actttaUy reconstruct the 
 greater portion of the entire complex of cell function upon the basis of 
 the presence in the cell of certain groups of histological characters the 
 meaning of each of which has been determined independently in differ- 
 ent cells. A beginning has already been made in that we are able to dis- 
 sociate in motor cells the characteristic structure (corresponding to the 
 motor activity) and the cell size (corresponding to the spheres of 
 influence and reception of the cell) ; these two dissociated elements of 
 cell character may be employed independently in examining any type of 
 cell in the entire nervous system. Such results can be accomplished 
 only through work which demands the exercise of the greatest critical 
 ability, but that they will be attained I am firmly convinced. 
 
 It is evident that no such results are yet possible in the case of the 
 hypothalamus, since no infonnation is available concerning the func- 
 tions of any of its cell groups; I have been comixdled therefore to 
 draw very general conclusions as to the nearness of functional relation- 
 ship between different cell groups, since these conclusions could at 
 present be based merely upon greater or lesser resemblance of cell type. 
 One conclusion as to the function of these cell groups is, however, 
 possible. The basal optic ganglion, the nuclei tuberis laterales, the 
 nucleus paraventricularis hypothalami and the nucleus tubero-mammil- 
 laris are composed of cells whose histological character indicates that 
 they are not efferent, but are concerned in receiving and correlating 
 incoming impidses; these cells do not possess the relatively large, dis- 
 crete Nissl bodies characteristic of efferent cells. Moreover (as sug- 
 gested by Dr. Donaldson) the small size of such cells as those of the 
 nuclei tuberis laterales would seem to indicate that the connections of 
 these cells were less extensive than those of the cells of the other three 
 nuclei. But now that these various cell groups have been described 
 experimental workers will be in a position to attack the problems 
 involving the connections and general functional significance of each 
 of these cell groups, realizing that differences of cell t}T)e in different 
 cell groups cannot be ignored, since such differences, as in every- portion 
 of the organism, correspond to differences in cell function ; we may then 
 expect results in this region upon which may be based in turn the con- 
 structive neuro-histology outlined above. 
 
 To those who prefer to employ several different histological methods 
 I shall point out the fact that the Nissl method as employed by me 
 brings out very clearly the differences and similarities of cell type, and
 
 Nuclei Tuberis Laterales and the Ganglion Opticum Basale. 31 
 
 accordingly should with the aid of experimental work and of other 
 histological methods suifice to give much valuable knowledge as to the 
 real relations involved in any nervous mechanism. Every method of 
 studying the nervous system has its advantages, and I have employed 
 the Nissl method merely because of all methods it is the most valuable 
 in giving a complete picture of the cell groups of the entire nervous 
 system. To this peculiar advantage of this method, through which we 
 can compare the histological character of all cell groups of the nervous 
 system, I should like to direct especial attention. Such a complete 
 picture of the cell groups of the nervous system would seem to offer 
 a most favorable ground-work for assimilating to itself isolated facts 
 observed by means of all other methods of studying the nervous system ; 
 without other methods (histological, experimental, etc.) it is of course 
 of little value. (See last paragraph of introduction, p. 3.) 
 
 In my opinion the demands of such work are so exacting that each in- 
 vestigator who attempts to gain a picture of the cell groups of a large 
 portion of the nervous system should confine himself, at least for the 
 most part, to one method, and of all methods that of Nissl shows the 
 cell structure in the clearest manner. It is not isolated cytological 
 details obtained by many methods, nor is it the proof or disproof of the 
 actual existence in the living cell of certain features of cell structure 
 that will be of most importance in the above outlined constructive 
 neuro-histology, but a thorough familiarity with the location, extent 
 and cell type of the different cell groups of practically the entire nervous 
 system, together with a most critical appreciation of the differences, 
 similarities and transitions of cell type of all these cell groups ; without 
 such broad knowledge of the various cell groups, which must of course 
 assimilate to itself isolated facts regarding the function and the connec- 
 tions of various cell groups directly and indirectly with one another, and 
 facts regarding tlie number, character, and mode of termination of the 
 cell processes of various cell types, without such a comprehensive knowl- 
 edge upon which to build we could not ho]w to penetrate far into the 
 exact relations of the individual neurones of the nervous system. 
 
 LlTER.\TUl!E. 
 
 The literature on the cell groups of the hypothalamus has been of 
 practically no assistance to me. In the first place I did not read it 
 until I had worked out these groups in the human brain, and upon 
 3
 
 32 Edward F. Malone. 
 
 reading it found no reason for modifying my conception of this region. 
 In tile second place some of tlie articles are concerned with animal forms 
 so far removed from those upon which I have worked that the recogni- 
 tion of homologous groups is by no means certain. Finally the 
 descriptions of cell groups are often so inadequate and so little attention 
 is given to the differences in cell type as to make the literature, except in 
 rare cases, almost unintelligible ; this is true especially in those articles 
 which are concerned with the brain of man and the higher mammals. 
 Accordingly the results of other workers on the cell groups of the hypo- 
 thalamus have not been incorporated to any great extent into the body of 
 this article, but are here appended for the sake of completeness ; to treat 
 the greater portion of the literature otherwise would render the subject 
 of this article needlessly obscure. The following consideration of the 
 literature is given for the benefit of those who wish to make a careful 
 study of the region in question ; accordingly it appears advisable not to 
 give an abstract of the various articles, since the complete accounts must 
 be read by those for whom the following consideration is intended. On 
 the contraiT I shall for the most part confine myself to a consideration 
 of certain points, showing wherein certain accounts are correct and 
 wherein erroneous, and wherein some descriptions are altogether unin- 
 telligible. 
 
 The description of tlie cell groups of the hypothalamus contained in my 
 monograph on the human diencephalon has been referred to in numerous 
 places in the text and a further reference is unnecessary. 
 
 In Strieker's Manual of Histology (American translation) Meynert 
 gives a brief but fairly clear description of the basal optic ganglion; on 
 p. 688 he says: "At the lateral border of the tuber cinereum lies the 
 inferior optic ganglion, which is 1.5 mm. broad, and contains spindle-shaped 
 cells 30 /x in length and 15 ii in breadth. It begins just above the optic 
 commissure and stretches along immediately over the tractus opticus as far 
 as the posterior border of the tuber cinereum, a distance of more than a 
 centimeter. I regard, with Luys, this optic ganglion as a part of the tuber 
 cinereum, because it projects downwards, in company with the latter, into 
 the lamina cinerea, beyond the surface of the lamina perforata anterior, of 
 which J. Wagner considers it to be a part, and because it extends farther 
 backward than the latter. Like the tractus itself, however, it certainly 
 follows the inner border of the anterior pert, space. On profile sections 
 (Fig. 270, IT) this ganglion has a sickle-like shape, the concavity looking 
 forward. According to Luys, the two ganglia touch at the median line, 
 a fact which I have not been able to verify, etc." From this description 
 and from a study of Fig. 268 it is clear that Meynert had in mind a cell 
 group which has much in common with that described under the same name
 
 Nuclei Tuheris Laterales and the Ganrjlwn Opticum Basale. 33 
 
 in the present article. The exact extent is, however, not clear, and we have 
 no assurance that he distinguished between this cell group and the nucleus 
 tubero-manimillaris, or that he included all portions of the basal optic 
 ganglion. Moreover he was unable to observe the union of the ganglia of 
 opposite sides, a fact which makes it almost certain that he failed to include 
 the medial portion of each ganglion. 
 
 When we attempt to understand the article of von Lenhossek, in which 
 he divides, in man, the basal optic ganglion of Meynert into a nucleus supra- 
 opticus and two nuclei tuheris, we are confronted with an impossible 
 task. We are informed that all three nuclei are composed of small, spindle- 
 shaped, multipolar nerve cells, as well as neuroglia; this is a most disturb- 
 ing statement and renders it absolutely impossible to form any idea as to 
 what cell groups the author had in mind. For by no means can the cells 
 of the basal optic ganglion be termed small, since they measure up to 35 m 
 in diameter, and compared with the cells of the nuclei tuheris and the 
 substantia grisea their large size is most evident (Fig. 37). Accordingly 
 the word " small " would seem to make it certain that Lenhossek's nucleus 
 supraopticus cannot be a portion of Meynert's basal optic ganglion: and 
 yet after Meynert's description and illustrations it would seem remarkable 
 for anyone to overlook such a prominent cell group as the oral portion of 
 the basal optic ganglion. Again we are informed that the nucleus anterior 
 of the tuber cinereum is the largest of all three nuclei (nucleus supra- 
 opticus and the anterior and postero-lateral nuclei of the tuber) " und 
 Mldet eigentUch den Hauptbestandteil des Tuber." and that apparently it 
 reaches the median line. What can this mean? Can the " small " cells of 
 this anterior nucleus be the large cells of the nucleus tubero-mammillaris, 
 or is this nucleus a portion of the substantia grisea? One is almost tempted 
 to infer from Lenhossek's description that his three portions of Meynert's 
 basal optic ganglion are a complex of what I have described as the basal 
 optic ganglion and portions of the nucleus tubero-mammillaris, all of which 
 are composed of large cells, and that the cells of the substantia grisea he 
 may possibly have considered neuroglia; since he employed the Weigert 
 stain such a mistake is by no means improbable. But just what the three 
 nuclei of Lenhossek represent is absolutely impossible to determine. 
 
 Kolliker's description of the nuclei tuberis and the basal optic ganglion I 
 have in a previous article termed vortrefflich ; this implies a uniform 
 excellence which this description by no means possesses. It were more 
 fitting to term Kolliker's account of these nuclei " remarkable," for it is 
 remarkable as to insight into this complicated region, and at the same time 
 remarkable for the contradictions, incorrectly labeled figures, and other 
 faults which detract from its value. Only to one who has made a thorough 
 study of this region is the difference apparent between the unintelligible 
 description of von Lenhossek and the essential excellence of that of 
 KoUiker, and from 1S96 until 1910 no one has made himself sufficiently 
 familiar with this region to realize the many excellent points in Kolliker's 
 work. The real contributions of Kolliker are as follows:
 
 34 Edward F. Malone. 
 
 Es miissen 'Nuclei tuberis von den Nuclei supraoptici unterschieden wer- 
 den (p. 602) . . .Die Nuclei tuberis kommen mehr in den medialen Gegenden 
 vor, besitzen kleine Nervenzellen, etc. (p. 603) . . .Die Nuclei supraoptici ha- 
 ben grossere Zellen, etc. (p. 603). Moreover Figs. 704 and 705 are correctly 
 labeled and entirely intelligible. Fig. 702 shows the nuclei tuberis laterales 
 excellently, but unfortunately they are termed "ganglia optica basalia"; 
 on p. 599 these groups are correctly termed nuclei tuberis, while on p. 519 
 they are incorrectly termed (in text) "ganglia optica basalia." This 
 uncorrected error in Fig. 702 is unfortunately characteristic of the incon- 
 sistent nature of Kolliker's description. But in Fig. 631 we meet a further 
 difficulty, since the cell mass of the tuber which in tig. 631 (rabbit) is termed 
 " Ganglion opticum basale " is stated on p. 603 to represent the " Ganglia 
 tuberis"; in reality this cell mass is simply a portion of the substantia 
 grisea and is not the nuclei tuberis laterales (which are shown in Figs. 
 702, G. o. b., and 705, G. t). On p. 599 he has incorrectly stated that the 
 nuclei tuberis are continued more or less definitely into two of the groups 
 of the corpus mammillare, nucleus intercalatus (or possibly a separated 
 portion of the medial nucleus as discussed in my monograph on the 
 diencephalon) and the nucleus tubero-mammillaris; this proves beyond 
 doubt that Kolliker's nuclei tuberis are to be regarded as topographical 
 groups rather than determined by a definite cell character. On the other 
 hand he has confused a portion of the nucleus tubero-mammillaris with 
 the basal optic ganglion; on p. 600 he says: Dieser Kern ist schon in der 
 Fig. 703 in der Grenzlage zwi-schen dem Tract us opticus, dem Reste des 
 Pes pedunculi und dem Nucleus Tuberis lateralis in den ersten Anfdngen 
 vorhanden und zwar me in der Linsenkernschlinge drin, wird aber nach 
 vorn immer grosser, etc. Accordingly Kolliker has included portions of 
 the nucleus tubero-mammillaris with the nuclei tuberis on the one 
 hand, and on the other with the basal optic ganglion. The vacant space 
 surrounding the third ventricle in his figures raises the question as to how 
 he regarded the substantia grisea of the third ventricle; its relations to the 
 other nuclei he has failed to point out, and in how far he confused this 
 cell mass with the nuclei tuberis we cannot determine, except in Fig. 631 
 in case of the rabbit, where this error is apparent; this cell group of the 
 rabbit is the substantia grisea and does not correspond to the nuclei tuberis 
 as shown in his figures of man. In conclusion, Kolliker's description and 
 figures are often very faulty, but on the other hand in his figures may be 
 recognized in many cases cell groups correctly represented. 
 
 Cajal (pp. 756-757) describes in rodents a cell group under the name 
 ganglia peri-kiasmatico o tangencial which without doubt is homologous 
 with the basal optic ganglion of higher mammals. He is in doubt as to 
 whether it corresponds to the supraoptic nucleus of Lenhossek and the 
 supraoptic complex of Kolliker. As previously explained it is impossible to 
 identify the supraoptic nucleus of Lenhossek, especially since he says that 
 it is composed of " small " cells; the description of Cajal as well as his 
 excellent illustration (Fig. 640) shows that the ganglio peri-kiasmiitico is
 
 Nuclei Tuheris Laterales and the Ganglion Opticum Basale. 35 
 
 composed of large cells. Cajal further states that there is a great difference 
 in form, position and extent between this ganglion of rodents and the above 
 human centers. While I have not yet studied the basal optic ganglion in 
 rodents I am strongly of the opinion that this ganglion in rodents does not 
 differ essentially either in form, position or extent from the basal optic 
 ganglion in man; I base this conclusion on the constancy of this ganglion 
 from man to the cat, and upon Cajal's description and illustration. There 
 is no difficulty in recognizing that the centers, which Cajal describes in the 
 tuber under the names of anterior, posterior and superior nuclei of the 
 tuber, are portions of the substantia grisea and have nothing to do with 
 the nuclei tuberis laterales of man and macacus; the difference in loca- 
 tion of these three nuclei and the fact that there is every reason to believe 
 that the nuclei laterales are not present in the rodents (appearing first 
 in the lemur) renders confusion impossible. Under the name niicleo 
 subventricular Cajal describes (p. 731) a cell column which is beyond 
 doubt homologous with the nucleus paraventricularis hypothalami, which 
 I have described in man and the higher mammals; Cajal's description and 
 illustration (Fig. 604, T) place this fact beyond the possibility of a doubt. 
 Accordingly Cajal's description of the hypothalamus enables us to recognize 
 positively two characteristic cell groups in rodents one of which is homolo- 
 gous with the basal optic ganglion and the other with the nucleus paraven- 
 tricularis hypothalami of higher animals. Concerning the complex of the 
 nucleus tubero-mammillaris we are left in doubt. 
 
 The presence of the basal optic ganglion and the nucleus paraventricularis 
 is certain not only in rodents but even in the marsupials, as is seen from 
 Ziehen's description of the brain of Pseudochirus peregrinus. In describing 
 Fig. 28 he says (p. 713) : Sehr schim ausgepriigt ist beiderseits das Gang- 
 lion opUciim basale (nucleus supraopticus). Rechts sendet es einen lang- 
 gestreckten AusUiufer in das Pedamentum laterale. Ingesammt erstreckt 
 es sich ilber 1.3 mm. in sagittaler Richtung. Seine Zellen messen 21 fi. 
 Frontalwiirts reicht es noch ein wenig iiber den vorderen Chias-marand 
 hinaus. Einen Zerfall in mehrere Zellgruppen, wie ihn Lenhossck und 
 Kolliker bei dem Menschen beschrieben haben. vermijchte ich nicht sicker 
 nachzuweisen. On p. 714 in describing this same section (Fig. 28) Ziehen 
 says: In der grauen Masse zu beiden Seiten des 3 Ventrikcls kann man — in 
 der Reihenfolge von untcn nach oben — folgende The.ile unterscheiden. 
 Vnmittelbar oberhalb des Chiasmas folgt der kleinzellige Nucleus tuberis 
 (evidently the substantia grisea), auf diesen ein gross-zelUger lateraler 
 Kern, in icelchem der Fornixsiiule eingebettet ist (a portion of my nucleus 
 tubero-mammillaris), und ein eben so grosszelUger medialer Kern, welcher 
 der Yentrikelwand unmittelbar anliegt. This latter nucleus is evidently 
 homologous with the nucleus paraventricularis hypothalami, and I 
 thoroughly agree with Ziehen when he continues as follows: Bei der sehr 
 starken Auspriigung des letzeren war ich sehr erstaunt, eine iihnliche 
 Bildung bei anderen Siiugern nirgends beschrieben zu linden. Ich mil den 
 Kern einstweilen als Nucleus subcommissuralis bezeichnen. Ziehen's
 
 36 Edward F. M alone. 
 
 illustrations unfortunately show almost nothing ot the various cell groups, 
 but his concise description is most valuable. 
 
 Worthy of special notice is the valuable contribution of Friedemann 
 entitled Die Cytoarchitektonik des Zivisvhenhirns der Cercopitheken mit 
 besonderer Beriicksichtigung des Thalamus opticus. As the title indicates 
 more attention is paid to the thalamus than to the hypothalamus, although 
 both the pars mammillaris and the pars optica are by no means neglected. 
 In this article Friedemann takes into consideration my monograph on the 
 human diencephalon. To consider intelligently Friedemann's account of the 
 cell groups of the hypothalamus it is necessary to realize that his chief 
 purpose is to subdivide this region into topographical cell groups, which 
 are not necessarily composed exclusively ot one type of cell, but may repre- 
 sent complex centers formed by the overlapping of several primary nuclei; 
 on the other hand he has occasionally divided a cell mass, composed of cells 
 of the same type, into subgroups whenever factors other than cell character 
 (such as number and grouping of cells) would permit. In this he has con- 
 sistently endeavored to give us a topographical subdivision into as many 
 areas as may possibly be distinguished in cell preparations. It is accord- 
 ingly evident that my cell groups (primary nuclei i have been set aside in 
 virtue of the possession of one type of cell which corresponds to some known 
 or unknown cell function or functions and that the topographical value of 
 my work is primarily concerned with the location and distribution of 
 such a single cell type; and on the other hand it is evident that Friede- 
 mann's subdivision, not being confined to this one criterion of cell character, 
 results in a much more minute subdivision which is of great value in exact 
 orientation, and that his cell groups may represent primary nuclei, portions 
 of such nuclei, or complex nuclei, according to the criteria employed in the 
 establishment of each cell group. Apparently Friedemann has not grasped 
 this fundamental difference in our aims, since (p. 312) he says: So xvichtig 
 auch u. a. die von Malone festgesteijte Tatsaehc ist, dass im Thalamus 
 opticus keine ZeUen vom viotorischen Typus vorkomm,en, so sehr wir auch 
 in vielen Einzelheiten mit diesem Autor iibereinstimmen, als topographis- 
 ches Einteilungsprinzip konnen wir seine Methode nicht anerkennen. 
 It is certainly my earnest desire that no one should consider the method 
 I have employed as a topographisches Einteilungsprinzip. since I myself 
 have never so considered it, and had supposed that I had made this fact 
 evident. But Friedemann's work is something more than a pure topo- 
 graphical subdivision, since he has informed us as to whether a cell group 
 has been recognized on account of its cell type, or whether it is differen- 
 tiated from the surrounding cells on the basis of other criteria; in other 
 words he has, in giving us this minute and most valuable topographical 
 subdivision, not lost sight of the distribution of the various cell types. The 
 fact that Friedemann and myself have subdivided the diencephalon from 
 different points of view is a fortunate circumstance, and the difference in 
 results is to be expected and welcomed. Other subdivisions from other 
 standpoints are desirable, but it is essential that we be accurately informed
 
 Nuclei Tuberis Laterales and the Ganglion Opticum Basale. 37 
 
 just what criteria have been employed in setting aside each cell group: and 
 moreover we should carefully distinguish between such cell groups of 
 different authors as are identical and such as do not entirely correspond. 
 
 Taking up the various cell groups of the hypothalamus, Friedemann des- 
 cribes the nucleus intercalatus corporis mammillaris and the nucleus 
 paraventricularis just as I had found them in man, and he adopts my names 
 for them. He also adopts my name "nucleus mammillo-infundibularis "; 
 I have previously in this article pointed out that this name must be 
 changed, and have adopted instead that of nucleus " tubero-mammillaris." 
 But it is not clear to me that Friedemann has included all the cells in this 
 group which I have assigned to it; unfortunately the illustrations (photo- 
 graphs) are of little value in showing the distribution of each cell type. 
 It is certain that his nucleus posterior pedamenti lateralis (Fig. 9, Pip) 
 is a portion of my nucleus tubero-mammillaris; moreover Friedemann 
 has called attention to the fact (p. 367) that his lamina grisea separans 
 (Fig. 9, Igs) is a portion of this same nucleus. Such subdivisions are of 
 course desirable for their topographical value, but the common cell charac- 
 ter should be kept in mind. In considering the basal optic ganglion Friede- 
 mann has fallen into the same error that I did in my monograph on the 
 diencephalon, and for the same reason, since both of us were not primarily 
 interested in the telencephalon; this error consists in considering merely 
 the cell mass which follows the oro-lateral border of the optic tract as 
 constituting this ganglion. It is certain that to the basal optic ganglion of 
 cercopithecus must be added Friedemann's " nucleus anterior pedamenti 
 lateralis " (p. 370 and Fig. 11, Pla) as well as his " stratum supraopticum " 
 (p. 369 and Fig. 9, Sso.). That these two cell groups belong to the basal 
 optic ganglion is clear both from the descriptions and illustrations. It is 
 unfortunate that neither the cross sections nor the horizontal sections 
 illustrated give the region just caudal to the optic chiasm. Friedemann's 
 cross sections may be best compared with my Series D of man (Figs. 11 to 
 18) since the plane of section is similar: in comparing these two series it is 
 evident that the most oral section (Fig. 11) of Friedemann stops short 
 of a most important region of the basal optic ganglion. Again his first 
 horizontal section (Fig. 12) likewise fails to give this region near the optic 
 chiasm. (The omission of this region is only natural, since Friedemann's 
 article deals primarily with the diencephalon.) Accordingly I see no reason 
 to believe that the basal optic ganglion in cercopithecus is essentially 
 different from that in other animals from man to the marsupials. 
 
 In Friedemann's description of cercopithecus I am unable to find any 
 cell groups which would correspond to the nuclei tuberis laterales, 
 and unfortunately the illustrations are of such a nature (photographs) as 
 not to reveal the presence of such groups. It is not probable that the nuclei 
 tuberis laterales are wanting in cercopithecus, since they are present in 
 macacus and are indicated even in the lemur; their presence, if indeed they 
 occur in cercopithecus, might easily be overlooked unless one had seen 
 them in man, where these nests of cells are very definite. The fact, which
 
 38 Edward F. Malone. 
 
 Friedemann himself has pointed out (p. 367-8), should be most carefully 
 noted: Friedemann's nuclei of the tuber cinereum are different portions 
 of the substantia grisea; he calls attention moreover to the fact (p. 367) 
 that the substantia grisea has been divided into these nuclei rather on 
 account of topographical relations and differences in the crowding together 
 of cells than on account of differences in cell form. Accordingly these 
 topographical cell groups of the substantia grisea must not be confused 
 with the nuclei tuberis laterales, since the latter are composed of cells of 
 another type. 
 
 Bibliography. 
 
 Cajal, S. R. y: Textura del sistema nervioso, etc., 1904, vol. 2. 
 FoREL, A. : Beitrage zur Kenntnis des Thalamus opticus, etc. 
 Gesammelte hirnaiiatomische Abhandlungen, 1872, pp. 17-43. 
 
 (Published originally in Bd. 66 der Sitzb. der k. Akad. d. Wis- 
 sensch. in Wien, dritte Abteilung, Juni.) 
 Friedemaxn, M. : Die Cjtoarchitcktonik des Zwischenhirns der Cer- 
 
 copitheken, etc. Jl. f. Psychologie u. Neurol., 1911, XVIII, 
 
 Erganzungsheft 2. 
 Ganser, L. : Yergleichende anatomische Studien iiber das Gehirn des 
 
 Maulwurfs; Morpholog. Jahrbuch, 1882, VII. 
 KoLLiKEE, A. : Handbuch der Gewebelehre, 1896, II. 
 VON Lenhossek, M. : Beobaclitungeu am richirn des Menschen, Anat. 
 
 Anz., 1887, II, Nr. 14. 
 Malone, E. : Uber die Kerne des menschlichen Diencephalon. .\us 
 
 dem Anhang zu den Abhandlungen der konigl. preuss. Akad. d. 
 
 Wissensch., 1910. 
 
 Observations concerning the comparative anatomy of the dien- 
 cephalon. Anat. Eec, 1912, VI, No. 7. 
 
 Eecognition of members of the somatic motor chain of nerve 
 
 cells, etc. Anat. Kec, 1913, VII, No. 3. 
 
 The nucleus cardiacus nervi vagi, etc. Am. Jour. Anat., 1913, 
 
 XV, No. 1. 
 JIeyxert, Th.: The brain of mammals. Strieker's manual of his- 
 tology (American translation), 1872. 
 Ziehen, Th. : Das Centralnervensystem der Monotremen und Mar- 
 
 supialier. Teil 2, erster Abschnitt. From Semon's Forschungs- 
 
 reisen. III.
 
 Nuclei Tuberis Laierales and the Ganglion Opticum Basale. 39 
 
 Explanation of Illustrations. 
 
 Figs. 1 to 35 and lA to 35A have been prepared in the following manner: 
 The entire section was in each case projected at a magnification of ten 
 diameters, and the outlines together with as many details as possible were 
 drawn in pencil; even bloodvessels, tears in the section, particles of detritus, 
 etc., were drawn in so as to serve the purpose of orientation. Under the 
 microscope at the same magnification (ten diameters) the drawing was 
 completed, with the occasional aid of higher magnification. (Of course the 
 drawings were not begun until I had a clear idea of the location, extent, 
 cell character, and relations of the various cell groups.) This completed 
 drawing is not used for publication but serves as the basis for two others. 
 In the first place an outline drawing is made (by tracing) and the various 
 structures labeled (Figs. lA to 35 A) ; in these drawings a rectangular area 
 is indicated which contains the region in which we are interested. A second 
 series of drawings (Figs. 1 to 35) are traced from that portion of the original 
 included within the rectangular area, and the details filled in, under control 
 of the microscope. Finally these two series of drawings have been repro- 
 duced in parallel columns; in order to do this the outline drawings have 
 been more greatly reduced in size than the detailed drawings, the resulting 
 magnification being indicated in the explanation of each plate. Figs. 1 to IS 
 have been so much reduced that they should be examined with the aid of a 
 hand lens; the cells of the nuclei tuberis laterales are especially difficult to 
 recognize without such aid. 
 
 It is important to understand just what these drawings are intended to 
 illustrate. The outline series (Figs. lA to 35A) needs no explanation, except 
 the statement that the size and relations of the structures shown have 
 been accurately reproduced. In the other series (Figs. 1 to 35) the distri- 
 bution of each cell group is accurately shown, each being indicated by one 
 type of cell which may be distinguished in all drawings from the cells of 
 all other cell groups; thus different cell groups may be distinguished when 
 their cells are intermingled, and it is thereby possible to reproduce the 
 actual extent of each cell type without resorting to an arbitrary boundary. 
 On the other hand in order to indicate differences in cell type it has been 
 necessary to make the cells much larger than they appear at such a low 
 magnification, and this in turn involves a reduction in the number of cells. 
 However, the relative density of each cell mass in its various portions is 
 shown, and the relative size of different cell types has been maintained, 
 although this difference in size has in some cases been somewhat exag- 
 gerated in order to make it more obvious. For a true picture of each cell 
 type one must study Figs. 38 to 58, in which each cell type has been 
 reproduced exactly as it appears. 
 
 The exact cell size and distance between the cells of the nuclei tuberis 
 laterales and the basal optic ganglion, as well as that of the surrounding 
 cell groups, has been reproduced exactly in Figs. 36 and 37, which also show 
 the different cell types as they appear at a relatively low magnification. 
 These two illustrations (Figs. 36 and 37) will be explained on the pages 
 facing them.
 
 40 Edward F. Malone. 
 
 Figures 3S to 58 were drawn as follows: 
 
 With the aid of the camera lucida the cell outlines and as many details 
 as possible were drawn in pencil. Then under control of the microscope 
 the exact appearance of the cells was reproduced in water color. Each 
 cell is to be regarded as an individual and the spatial relations of the cells 
 in each figure do not correspond to those between the cells of the corres- 
 ponding cell group in the preparations. In these figures the cells have 
 been reproduced by combining to a certain extent different optical sections 
 of the same cell. All cells have been drawn at a magnification of 580 diame- 
 ters, and have been reproduced without reduction. Figs. 38 to 58 were all 
 drawn from cross-sections of the brains of the various animals. In Series 
 D of man the plane of section differs widely from that of a cross-section, 
 and yet (without making a careful study of this point) I was unable to 
 observe that this difference in the plane of section was correlated with any 
 difference in the appearance of the cell types here illustrated. On the con- 
 trary I can state positively that each of the above types of cells was as 
 characteristic in Series D as in Series AC, and that differences in the plans 
 of section could not possibly lead to a confusion of the different cell types.
 
 42 Edward F. Malone. 
 
 PLATE I. 
 
 Homo. Seeies AC. 
 
 Magnification of Figs. 1 to 4 = 5 diameters. 
 Magnification of Figs. lA to 4A = 2 diameters. 
 
 • = Cells of the nucleus tubero-mammillaris (formerly known as nucleus 
 
 mammillo-infundibularis) . 
 
 • = Cells of the substantia grisea ventriculi tertii. 
 
 A = Cells of the nuclei tuberis laterales (n. t. 1.). (Examine with baud 
 
 lens ) . 
 O = Cells of the ganglion opticum basale (g. o. b.). 
 Q = Cells of the corpus hypothalamicum (Luysii).
 
 THE JOHNS HOPKINS HOSPITAL REPORTS. 
 
 PLATE I.
 
 4A Edward F. Malone. 
 
 PLATE II. 
 
 Homo. Series AC (Continued). 
 
 Magnification of Figs. 5 to 7 = 5 diameters. 
 
 Magnification of Figs. 5A to 7A = 2 diameters. 
 
 • = Cells of the nucleus tubero-mammillaris (nucleus mammillo-infundib- 
 
 ularis). 
 • = Cells of the substantia grisea ventriculi tertii. 
 A = Cells of the nuclei tuberis laterales (n. t. 1.). (Examine with hand 
 
 lens). 
 O = Cells of the ganglion opticum basale (g. o. b.). 
 O = Cells of the corpus hypothalamicum (Luysii).
 
 THE JOHNS HOPKINS HOSPITAL REPORTS. 
 
 PLATE II. 
 
 Fio. 7. 
 
 Fig. 7.\.
 
 46 Edward F. Malone. 
 
 PLATE III. 
 
 Homo. Series AC (Co.ncluded). 
 
 Magnification of Figs. 8 to 10 = 5 diameters. 
 Magnification of Figs. 8A to lOA = 2 diameters. 
 
 {Cells of tlie nucleus tubero-mammillaris (nucleus mammillo-infun- 
 dibularis). 
 Cells of the nucleus paraventricularis hypothalami. 
 • = Cells of the substantia grisea ventriculi tertii. 
 A = Cells of the nuclei tuberis laterales (n. t. 1.). (Examine with hand 
 
 lens ) . 
 O = Cells of the ganglion opticum basale (g. o. b. ). 
 
 O = Cells of the nucleus ansae peduncularis (ganglion basale of Kolliker). 
 Note that the cells of the nucleus tubero-mammillaris and those of the 
 nucleus paraventricularis are represented by the same type of cell; more- 
 over the same type of cell has been employed to represent the cells of the 
 corpus subthalamicum and those of the nucleus ansae peduncularis. It 
 seems advisable to use one type of cell to represent the cells of two different 
 groups, when these two groups are clearly distinguishable topographically, 
 rather than to multiply the number of symbols used, and thus render it 
 more diflBcult to distinguish between them. The outline drawings will 
 make a confusion of such cell groups impossible.
 
 THE JOHNS HOPKINS HOSPITAL REPORTS. 
 
 PLATE III. 
 
 Fig. 9. 
 
 Fig. 9a. 
 
 Fig. 10. 
 
 Fig. 10a.
 
 48 Edward F. Malone. 
 
 PLATE IV. 
 
 Homo. Series D. 
 
 Magnification of Figs. 11 to 14 = 5 diameters. 
 Magnification of Figs. IIA to 14A = 2 diameters. 
 
 = Cells of the nucleus tubero-mammillaris (nucleus mammillo-lnfundib- 
 ularis). 
 
 • = Cells of the substantia grisea ventriculi tertii. 
 
 [> = Cells of the nuclei tuberis laterales (n. t. 1.). (Examine with hand 
 
 lens). 
 O = Cells of the ganglion opticum basale (g. o. b.). 
 Q = Cells of the nucleus ansae peduncularis. 
 
 ° = Cells of the ganglion mediale corporis mammillaris. 
 
 ^ = Cells of the nucleus intercalatus corporis mammillaris. 
 
 . = Cells of the cortex cerebri. 
 
 >- = Cells of the globus pallidus nuclei lentiformis.
 
 THE JOHNS HOPKINS HOSPITAL REPORTS. 
 
 PLATE IV. 
 
 Fig. 11. 
 
 Fig. 11a. 
 
 Fig. 14. 
 
 Fig. 14a.
 
 50 Edward F. Malonc. 
 
 PLATE V. 
 Homo. Series D (Co.ntixued). 
 
 Magnification of Figs. 15 to 18 = 5 diameters. 
 Magnification of Figs. 15A to ISA = 2 diameters. 
 
 f Cells of the nucleus tubero-mammillaris (nucleus mammillo infun- 
 = .J dibularis). 
 
 I Cells of the nucleus paraventricularis hypothalami. 
 • = Cells of the substantia grisea ventriculi tertii. 
 A = Cells of the nuclei tuberis laterales (n. t. 1.). (Examine with hand 
 
 lens). 
 O = Cells of the ganglion opticum basale ( g. o. b. ). 
 Q = Cells of the nucleus ansae peduncularis. 
 . = Cells of the cortex cerebri. 
 >- = Cells of the globus pallidus nuclei lentiformis. 
 
 Note in Fig. 17 the closely packed group of cells of the substantia grisea 
 situated just lateral to the ventral tip of the third vtntricle (see text). 
 
 Note in Fig. IS the cells of the basal optic ganglion crossing mid line 
 (see text).
 
 THE JOHNS HOPKINS HOSPITAL REPORTS. 
 
 PLATE V. 
 
 Fig. 15. 
 
 Fig. ].5a. 
 
 Fig. 16. 
 
 Fig. 16a. 
 
 
 Fig. 17. 
 
 Fig. 17a. 
 
 
 Fig. 18. 
 
 Fig. 18a.
 
 52 Edward F. Malone. 
 
 PLATE VI. 
 
 Macacus Rhesus. Sebies A. 
 
 Magnification of Figs. 19 to 22 = 6% diameters. 
 Magnification of Figs. 19A to 22A = 2V4 diameters. 
 
 # = Cells of the nucleus tubero-mammillaris (nucleus mammillo-infundib- 
 ularis). 
 
 • = Cells of the substantia grisea ventriculi tertii. 
 A = Cells of the nuclei tuberis laterales (n. t. I.). 
 O = Cells of the ganglion opticum basale (g. o. b. ). 
 JCells of the corpus hypothalamicum (Luysil). 
 Q~ \ Cells of the nucleus ansae peduncularis. 
 
 ° = Cells of the ganglion mediale corporis mammillaris. 
 
 ^ = Cells of the nucleus intercalatus corporis mammillaris. 
 
 . = Cells of the cortex cerebri. 
 
 >- = Cells of the globus pallidus nuclei lentiformis. 
 
 Note in Fig. 22 the group of densely packed cells of the substantia grisea 
 located just lateral to the third ventricle (see text).
 
 THE JOHNS HOPKINS HOSPITAL REPORTS. 
 
 PLATE VI. 
 
 Fig. 19. 
 
 FiQ. 19a. 
 
 '■v;m 
 
 Fig. 20. 
 
 Fig. 20a. 
 
 Fig. 22. 
 
 Fig. 22a.
 
 54 Edward F. M alone. 
 
 PLATE VII. 
 
 Macacus Rhesus. Sebies A (Continxied). 
 
 Magnification of Figs. 23 to 25 = 6% diameters. 
 Magnification of Figs. 23A to 25A = 2% diameters. 
 
 {Cells of the nucleus tubero-mammillaris (nucleus mammillo-infun- 
 dibularis). 
 Cells of the nucleus paraventricularis hypothalami. 
 • = Cells of the substantia grisea ventriculi tertii. 
 O = Cells of the ganglion opticum basale (g. o. b. ). 
 = Cells of the nucleus ansae peduncularis (ganglion basale of Kolliker). 
 . = Cells of the cortex cerebri. 
 >- = Cells of the globus pallidus nuclei lentiformis. 
 
 Note in Fig. 24 lateral from the third ventricle the densely packed group 
 of cells of the substantia grisea; such masses of the substantia grisea should 
 not be confused with the nuclei tuberis laterales.
 
 THE JOHNS HOPKINS HOSPITAL REPORTS. 
 
 PLATE VII. 
 
 Fig. 2a 
 
 Fig. 23a. 
 
 .W 
 
 
 FiC. 24. 
 
 
 
 9, S • .-' 
 
 '^'*«??o- * *- .':•'.--•-":'■*•••• 
 
 'i^ct"";.-^ ;.:.:.•..■.■:■••.■.•. v. 
 
 Fig. 25. 
 
 FlG. 20A.
 
 56 Edward F. Malone. 
 
 PLATE VIII. 
 Lemur Rdfus. Series A. 
 
 Magnification of Figs. 26 to 30 = SVa diameters. 
 Magnification of Figs. 26A to 30A = 3% diameters. 
 
 rCells of the nucleus tubero-mammillaris (nucleus mammillo-infun- 
 = -i dibularis). 
 
 I Cells of the nucleus paraventricularis hypothalami. 
 • = Cells of the substantia grisea ventriculi tertii. 
 O = Cells of the ganglion opticum basale (g. o. b.). 
 >- = Cells of the globus pallidus nuclei lentiformis. 
 Note in Fig. 27 the absence of the basal optic ganglion (see text).
 
 THE JOHNS HOPKINS HOSPITAL REPORTS. 
 
 PLATE VIII. 
 
 Fig. 27. 
 
 . • ■ • • 
 . . ■ ••••••■ 
 
 if*-'--' .*•» " 
 
 •.•;.•••■»'.■••.•• 
 
 v.\ /■.•■.•.•:•■.'••;■ ■ ■ . • * 
 
 •••I I. .••.•:.■■.•••• •• ■ * > 
 
 "•.■:\ /.••■•.v-.v.^.--..--. • »£,•; .°^o • • 
 
 Fig. 28. 
 
 t!t .■••.••• ♦. 
 
 ■:::•.■.•:■:,■■»■ ■ 
 
 ■'■■ r' 
 
 Fig. 29. 
 
 
 Fig. 30. 
 
 Fig. 2Sa. 
 
 Nuc\e<(5 Piraverttr'ic 
 u\*r>5 liyPoth»Uw'\ 
 
 optiCo-m 
 
 ©■fi 
 
 T^- 
 
 C h iASITiA
 
 58 Edward F. Malone. 
 
 PLATE IX. 
 
 Cat. Series C. 
 
 Magnification of Figs. 31 to 35 = Sy^ diameters. 
 Magnification of Figs. 31A to 35A =: 3';; diameters. 
 
 (Cells of the nucleus tubero-mammillaris (nucleus mammillolufun- 
 dibularis). 
 Cells of the nucleus paraventricularis hypothalami. 
 • = Cells of the substantia grisea ventriculi tertii. 
 O = Cells of the ganglion opticum basale (g. o. b.). 
 . = Cells of the cortex cerebri. 
 )- = Cells of the globus pallidus nuclei lentiformis.
 
 THE 
 
 JOHNS HOPKINS HOSPITAL REPORTS 
 ■••.»• • • • . 
 
 :-;•■>• •••■ • .T • • • • Vv- • .■ 
 
 PLATE IX. 
 
 ' * • 
 
 
 ■••;•.•••>••"•*.♦• 
 ••■•'.>■•• i* • • • 
 
 < .• 
 
 P«M--: -..•' .4K-:S> 
 
 Fig. 32. 
 
 l»»:.v«V4:«.— 
 
 
 /•.••■•:••••••.• 
 
 »•••: •■•:••'■•• v". ■>. » . » ■* 
 l^•■•^^•: •".".•■.• • *• * *■'. ' 
 
 » Fig. 34a
 
 60 Edward F. M alone. 
 
 PLATE X. 
 Homo. SsatLES AC. Nuclei Tubesis Laterales. 
 
 Fig. 36 represents the actual appearance of a portion of the preparation 
 from which Fig. 4 (PI. I) was drawn. The preparation was projected at 
 a magnification of 83% diameters and every cell outlined in pencil. Then 
 under the microscope each cell (as far as this was possible) was identified 
 and reproduced in water color exactly as it appeared. This Illustration 
 is accordingly an accurate reproduction of the appearance of the preparation 
 except that bloodvessels and neuroglia have been omitted. 
 
 Magnification (after y^ reduction) = 55% diameters. 
 
 Comparing Fig. 36 with Figs. 4 and 4A it is evident that the region illus- 
 trated is the ventro-lateral portion of the tuber cinereum. At the right 
 upper corner is seen a portion of the pes pedunculi, while at the left upper 
 corner appears a portion of the fornix column. 
 
 The two groups of small pigmented cells are the nuclei tuberis laterales. 
 Note that they are partly circumscribed by a zone containing few cells. 
 
 The large cells are those of the nucleus tubero-mammillaris (nucleus 
 mammillo-intundibularis ) . 
 
 The small pale blue cells are those of the substantia grisea ventrlculi 
 tertii. Note transition types between these cells and the large cells of the 
 nucleus tubero-mammillaris; this transition involves both the size of the 
 cells and the intensity of the stain, so that some cells occur which could not 
 be positively assigned to either of these two cell groups. 
 
 Compare the various cell groups of Fig. 36 with the corresponding cell 
 types of PI. XII, where the cells (man) are highly magnified.
 
 JOHNS HOPKINS HOS 
 
 VOLUME XVil.
 
 JOHNS HOPKINS HOSPITAL REPORTS. 
 
 VOLUME XVII 
 
 /^ 
 
 ( 
 
 C
 
 62 Edward F. M alone. 
 
 PLATE XI. 
 Homo. Series AC. Ganglion Optkum Basale. 
 
 Fig. 37 shows the actual appearance of a portion of the basal optic gang- 
 lion in the preparation from which Fig. 9 (Plate III) was drawn. As in the 
 case of Fig. 36 the preparation was projected (but at a magnification of 
 145 diameters instead of S3%) and every cell outlined in pencil. Then 
 under the microscope every cell (except in rare instances where this was 
 impossible) was identified and reproduced accurately in water color. 
 
 Magnification (after % reduction) = 96% diameters. 
 
 Fig. 37 represents the whole breadth of the basal optic ganglion near its 
 medial pole (Fig. 9, PI. Ill), and shows the relation of this nucleus to the 
 substantia grisea of the third ventricle. To the right is seen the ventro- 
 lateral surface of the brain (anterior perforated substance). 
 
 The large deeply staining cells are those of the basal optic ganglion. 
 Note that it is sharply separated from the substantia grisea. The long 
 coarse processes of the cells of the basal optic ganglion are here represented 
 as blue, but in reality they are almost colorless; this figure, however, repre- 
 sents correctly the peculiar appearance of an intercellular feltwork, to 
 which these processes give rise. 
 
 The small pale cells are those of the substantia grisea ventriculi tertii. 
 Note the difference in appearance of the intercellular substance from that 
 within the basal optic ganglion, due to the absence of coarse cell processes. 
 
 As in Fig. 36 bloodvessels and neuroglia have not been drawn. 
 
 See PI. XII for the appearance of these two cell types (in man) under 
 higher magnification.
 
 JOHNS HOPKINS HOS 
 
 VOLUME XVII.
 
 JOHNS HOPKINS HOSPITAL REPORTS. 
 
 VOLUME XVII, 
 
 
 '• • * 
 
 
 ^ 
 
 ' 1 i 
 
 • e 
 
 \« 
 
 ^'^ 
 
 1* ' 
 
 
 . v•.,,»>,.v^\ v.- -: 
 
 v,» 
 
 
 
 i I 
 
 C)
 
 64 Edward F. Malone. 
 
 PLATE XII. 
 Homo. MAOiNiFicAxio.N = 580 Diameters. 
 
 38. Cells of the ganglion opticum basale. 
 
 39. Cells of the nuclei tuberis laterales. 
 
 40. Cells of the substantia grisea ventriculi tertii. 
 
 41. Cells of the nucleus tubero-niammillaris (formerly known as nucleus 
 
 mammillo-infundibularis). 
 
 42. Cells of the nucleus ansae peduncularis (also known as ganglion basale 
 
 of Kolliker). 
 
 43. Cells of the nucleus paraventricularis hypothalami.
 
 JOHNS HOPKINS HOSPITAL REPORTS. 
 
 VOLUME XVII. 
 
 38 
 
 41 
 
 
 
 ■^» 
 
 42 
 
 39 
 
 
 fes^ 
 
 40 
 
 •)
 
 66 Edward F. Malone. 
 
 PLATE XIII. 
 
 Macacus Rhesus. Magnification = 580 Diameters. 
 
 44. Cells of the ganglion opticum basale. 
 
 45. Cells of the nuclei tuberis laterales. 
 
 46. Cells of the substantia grisea ventriculi tertii. 
 
 47. Cells of the nucleus tubero-mammillaris (formerly known as nucleus 
 
 mammillo-infundibularis). 
 
 48. Cells of the nucleus ansae peduncularis (also known as the ganglion 
 
 basale of Kolliker). 
 
 49. Cells of the nucleus paraventricularis hypothalami.
 
 JOHNS HOPKINS HOSPITAL REPORTS. 
 
 VOLUME XVII. 
 
 44 
 
 47 
 
 : . . • it-- 
 
 J'-^--^ 
 
 V,'- \ 
 
 48 
 
 45 
 
 ^ 
 
 
 t^'- 
 
 46 
 
 49 
 
 wm 
 
 ,^ 
 
 r.
 
 i
 
 68 ' Edward F. Malone. 
 
 PLATE XIV. 
 
 Lemur Rufus. Maomficatiox = 5S0 Diametebs. 
 
 50. Cells of the ganglion opticum basale. 
 
 51. Cells of the substantia grisea ventriculi tertii. 
 
 52. Cells of the nucleus tubero-mammillaris (formerly known as the 
 
 nucleus mammillo-infundibularis ) . 
 
 53. Cells of the nucleus ansae peduncularis (also known as the ganglion 
 
 basale of Kolliker). 
 
 54. Cells of the nucleus paraventricularis hypothalami.
 
 JOHNS HOPKINS HOSPITAL REPORTS. 
 
 VOLUME XVI 
 
 50 
 
 "^^ 
 
 52 
 
 53 
 
 * J^J^ 
 
 r*\ 
 
 "■^ 
 
 f < * 
 
 .^1 
 
 54 
 
 51 
 
 
 -i) 
 
 '^
 
 70 Edward F. Malone. 
 
 PLATE XV. 
 Cat. Magnification — 5S0 Dia.metebs. 
 
 55. Cells of the ganglion opticum basale. 
 
 56. Cells of the substantia grisea ventriculi tertii. 
 
 57. Cells of the nucleus tubero-manimillaris (formerly known as the 
 
 nucleus mammillo-intundibularis). 
 5S. Cells of the nucleus paraventricularis hypothalami.
 
 JOHNS HOPKINS HOSPITAL REPORTS. 
 
 VOLUME XVII. 
 
 55 
 
 ilC 
 
 iW^" 
 
 56 
 
 57 
 
 
 58 
 
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