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MARINE BIOLOGICAL LABORATORY. 


Received 
Accession No. 
Given by 


Place, 


*,* No book or pamphlet is to be removed from the Lab- 
oratory without the permission of the Trustees. 


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THE JOURNAL 


OF 


Comparative Neurology 


AOUWARTERLY PERTODICAL 


DEVOTED TO THE 


Comparative Study of the Nervous System. 


EDITED BY 
C. L. HERRICK, Professor of Biology, Denison University. 
ASSOCIATED WITH 


OLIVER S. STRONG, Columbia University, 
C. JUDSON HERRICK, Denison University. 


VOLUME VI, 1896. 


PUBLISHED BY THE EDITORS. 
GRANVILLE, OHIO, U. S. A, 
R. Friedlander & Son, Berlin, European Agents. 


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The Journal of Comparative Neurology. 


Contents of Vol. VI, 1896. 


MARCH, 1896. 
Pages 1-58, i-xliv. Plates I-VIII. 


ILLUSTRATIONS OF CENTRAL ATROPHY AFTER EYE INJURIES. By 
CoE Herricks With Platten 2 2 SS eae ae oe Soe ees 
LECTURE NOTES ON ATTENTION. AN ILLUSTRATION OF THE EmM- 
PLOYMENT OF NEUROLOGICAL ANALOGIES FOR PSYCHICAL PRO- 
BLEMS» |) BY GZ. Plerreck 2 ost Bea ee eee 
A NOTE ON THE CEREBRAL FISSURATION OF THE SEAL [PHOCA 
WELULINA- By erred. Lash) DuSe..) With Plate ili SS ae ee 
MoRPHOLOGY OF THE NERVOUS SYSTEM OF CypRIS. By C. . Tur- 
ers \ NV Plates DT Vil See ee eee a) Se eT NOUS 
PRELIMINARY NOTES ON THE CRANIAL NERVES OF CRYPTOBRANCHUS 
ALLEGHANIENSIS..” By V/e A McGrec0r (BiSGnes aan e een eee 
ON THE THREE POINTS IN THE NERVOUS ANATOMY OF AMPHIBIANS. 


By, 0/5) Ss) areestey., | ithe three) figures 2 se ee ae eee eee 


IYRERARY NOTICHSS ti boc 05 SS he heh a ek Oe es 
Effects of Alcohol on the Cortical Nerve Cell --_.--_.---- 
Structure of Nerve Cells after Electrocution --.-..------__ 
Cortical Pathology of Permanent Dementia -------_._----- 
The Fenetians of the’ Frontal) Ropessic 2 ie es 
Cerebral Reesivestion= = 22 22 ee eee Se ee 
Motor Functions of Dorsal Spinal Nerves ---.-...-------- 
Spiral Fibers in the Invertébratesuc ! 2222 222552250222 202 
Structure of ‘the Dhalamus 322 Oy eee eee ose ee 
he) Lateral) Line System oft Amphibia se) oso 
Brains; of Sallropsida isa ase aoe ee ee ee 
Frophte - Nerveses 225.555 sis ob an le oe a el 
Ampullo-oculo-motor Connections 222 sake se ase eke 
The Initial Stages of the Degeneration of Nerve Fibers___ 
Cortigal: Olfactory Appartus: 2.5202 be eee eee 
Fiber Connections of the Olfactory Lobe of Man-__-----. 
The Relation of Sensory and Motor Areas of the Cortex_. 
Is\.the Decorticated: Dog/Consetous/? x24 see ee 
The Paraphysis 
Be ING wy JOU a eee 
Structure of the Cytoplasm of the Nerve Cell....---.----- 
the Pons|:Varolii) of) Mansi. 2s. Sees. ee ee 


The Influence of the Cerebrum upon the Excretion of Ni- 


trogen ——___- ee 
Nerve) Permini‘of, the (Prostate "2252 sae eee ee 
Demonolatry of the Nineteenth’ Centuryi2222222e2se. ee noe 
he ‘Growth of ‘the (Brains 222 522-2 eee ee 
Recent Studies in the Forebrain of Reptiles__-__.-..___--. 


Modifications of Nerve Cells in Different Functional States_ 
‘The* American? Lobster 2222-22-02 0 eee eee ee 
Pain in the Pectoral Region Sympathetically Accompanying 

Irritation of; ithe) Horearmce a2 eee ee eee eee 
Corrigendum ea acne eee eee 


JUNE, 1896. 
Pages 59-132, xlv-liv. Plates IX-XIII. 


COMPARATIVE ANATOMY OF THE INSULA, By Tracy Earl Clark, B.S. 
With Plates) EX- Xo 82 SS ooo ee rte eee ee 
REVIEW OF THE GOLGI METHOD. By Olver S. Strong, Part I_----- 
THE DorsAL SACK, THE AULIX AND THE DIENCEHPHALIC FLEXURE. 
By, Bs iGs Wilder 2252 Ife Sea ea eee eee ek. wane 
THE EcTAL RELATIONS OF THE RIGHT AND LEFT PARIETAL AND 
PAROCCIPITAL, BISSURES., By (65) Goui2ideree ene ne a ae 
EDITORIAL. NFEURONYMIC PROGRESS IN AMERICA-.....- Seon k so seae 
EITERARY, (NOTICES 2o22--0--5— = a Bie Ae Serres CS ee ee 
The Development of the Brain and Sense-Organs in Elas- 
mobranchs 222002 222225, Boe ee eee eee 

the) Sensory) Physiology (of vActinianse= sor -ss-se- soe en ea 

Phe; Histology: of the) Myclone20e2 2 aes eee weno eee 
Anatomy, of the ‘Gerebrum of; Notonyctes/222=_-2--2=- eee 
Romanes on Weilsmannisim] = ee eee ae 
Germinal(/Selection: 22 [223-2 se ee ree aie os. 2 

the Psychology vofivAttentionsse === eee eee 

Studies from the Yale Psychological Laboratory_...-----_- 

The Embryology of the Medulla of the Rabbit-_.._______ 
Wranismis 222212 528 Sees Ne ie ae eek 


DECEMBER, 1896. Double Number. 
Pages 133-352. Plates XIV-XXII. 


THE BRAIN OF THE BEE. A PRELIMINARY CONTRIBUTION TO THE 
MORPHOLOGY OF THE NERVOUS SYSTEM OF THE ARTHROPODA. 
By #..C. Kenyon, Ph.D. With Plates XIV-XXIT_2....-=..-. 
THE ORIGIN AND GROWTH OF BRAIN CELLS IN THE ADULT Bopy. 
By \oward Ayers cts iin ots) COs ee ee oe 
THE INNERVATION OF THE AUDITORY EPITHELIUM IN MUSTELUS 
CANIS, DEIGAY.t4! Byres, (ore al hee a een ee eet ee 
NEURAL TERMS, INTERNATIONAL AND NATIONAL. By Burt G. 
Walder MOD sues Sees ee soe Ce ee Ap ee eae ee 


xxvi 
XXVil 
XXVii 
XXXVI 
XXXVili 
xli 

xliii 


xliv 
xliv 


58 
101 


128 


129 
131 


xlv 
xlv 

“xl vii 
xl viii 
xlix 


133 
211 
214 


216 


ILLUSTRATIONS OF CENTRAL ATROPHY AFTER 
EYE INJURIES. 


With Plate 1. 
Gb SHerrick: 


In the present paper it is not hoped to add anything new 
to the vast and.daily augmenting literature of the visual connec- 
tions but simply to place in accessible form some data accumu- 
lated, with the hope of ultimately making a careful study of the 
peripheral connections and particularly the central mechanism 
of accommodation. The difficulty of finding figures illustrating 
the actual results of the atrophies due to the destruction of the 
eye has disposed us to print our own diagrams with a running 
commentary. The material consists of serial sections of the 
brain of two rabbits whose eyes were operated on shortly after 
birth and before the eyes had opened. Series 292 is from a 
rabbit operated on August 25th, the right eye being removed. 
Killed November Ist, when nearly full grown. Series 297 is 
from a rabbit from which the left eye was removed September 
Igth, being then one day old. It was killed December 1gth. 
As these brains were intended to compare with control series of 
normal brains and were to be used primarily in the study of the 
cells, etc., they were stained with sublimate hematoxylin and 
fuchsin. In both cases the loss of vision was complete and the 
wound closed by first intention without suppuration, though in 
one the globe of the eye and the muscular attachments were 
less completely destroyed than in the other. Unfortunately 
there was no opportunity for careful post-mortem examination of 
these conditions which can be inferred from the state of atrophy 
found in the sections. It will be seen that the atrophy in the 
optic tracts was much more complete in No. 297, that is in the 
one operated upon within 24 hours after birth, than in the other 
which was operated upon a few days later. We begin with No. 
292, commencing behind the chiasm. The optic tract on the right 


2 JOURNAL OF ComPARATIVE NEUROLOGY. 


side is intact except that the innermost band of fibres is filled 
with granules to such an extent as to be decidedly darker than 
the ectal portions. The degenerate portion does not embrace 
more than a sixth of the whole. On the left side (that corres- 
ponding to the injured eye) the conditions are exactly reversed. 
The outer five-sixths is degenerate and filled with irregularly 
disposed granules, while the ental layer appears normal or near- 
ly so. Under a moderate power (1-5 in. obj.) the difference be- 
tween the degenerate and normal tracts is conspicuous. The 
unaltered portions have the nutritive granules arranged in regu- 
lar files of 6-8 between the fibres and these rows are disposed 
at approximately equal distances. In the degenerate portions 
the granules are not only relatively more numerous but they 
are dispersed with absolute irregularity. The fibres are no 
longer approximately parallel but are irregularly meshed. The 
sheaths are partly dissolved and the myelin completely ab- 
sorbed. (Cf. Fig. 11.) The tectum in this series is somewhat 
larger on the right side but there are no demonstrable histolog- 
ical differences. 

Passing caudad to the level of the exit of the oculo-motor, 
we find the right root entirely absent while its fellow is normal. 
The right nidulus of origin is also absent while the interpedun- 
cular niduli, which some writers persist, in spite of abundant 
evidence, in regarding as accessory third niduli, are intact on 
both sides. It is not possible to identify positively any of the 
elements of the missing nidulus though shrunken adendritic 
elements occur in the site. (Fig. 3.) 

At the level where the interpeduncular nidulus ends cau- 
dad, the cells of the fourth nidulus appear in the sections and 
are more numerous on the right than on the left side. Still 
there is no conspicuous difference in this subject between the 
two sides. The dorsal longitudinal fasciculus becomes more 
concentrated at this point and the nidulus lies directly dorsad of 
it in a stroma of deeply staining material. There are about the 


1In discussing the sections it may be premised that the cephalic aspect of 
the section is presented so that the right side of the section is the left side of 
the figures. 


HEeRRIcKk, Central Atrophy After Eye Injuries. 3 


same number of fibres in the root of the fourth nerve on either 
side. (See Fig. 4.) The external portion of the fourth nerve 
of the right side is apparently wholly degenerate while its fel- 
low of the opposite side is intact. For some reason, then, the 
degenerative process has destroyed the central tract and nidulus 
of the third nerve but not those of the fourth, though the ex- 
ternal part of the latter has disappeared. We find no central 
tracts degenerate between the fourth and sixth nerve niduli, but 
the latter has as completely disappeared as in the case of the 
oculo-motor. Nothing is left of root, tract or nidulus. The nid- 
ulus of the left side lies, as usual, just laterad of the genu of 
VII and its root tract turns sharply mesad beneath the VII 
fibres and then passes directly ventrad to the root. Some traces 
of the sheath may be found by careful search in the course of 
the root of the other side. 

The result of the operation, then, has been as follows: (1) 
degeneration of the lateral three fourths to five sixths of the 
left optic tract and of the mesal one fourth to one sixth of the 
right tract, (2) proportional atrophy of the optic tecta, most 
marked on the left side, (3) obliteration of the nidulus, root 
tracts and roots of III and of the external part and ‘roots of IV 
on the right side, (4) the obliteration of VI on the left side as 
completely as III. Negatively, the central portions of IV are 
unaltered and the interpeduncular niduli are intact. 

The rabbit of series 297 was operated on earlier and the 
results are rather of the nature of atrophies than of degenera- 
tions. The diameter of the left optic nerve is somewhat more 
than one fourth that of the right. At the chiasm, as seen in 
Fig. 7, the proportions are about the same. The degenerated 
portions are almost completely absorbed leaving only atrophic 
phenomena. Sections further caudad show very distinctly the 
atrophy in the tract. (Fig. 8.) 

At the proper level the corpus geniculatum of the left side 
appears with its tracts, but on the other side it seems quite ab- 
sent. There are no degenerative changes to enable us to trace 
the tracts cephalad to the cortex but there seems no reason to 
doubt that they are atrophic throughout. The reduction in the 


4 JOURNAL OF COMPARATIVE NEUROLOGY. 


tectum has gone on and perhaps nearly reached its limit. The 
left tectum is fully four times the size of the right but the inti- 
mate structure is not obviously different. 

Passing to the oculo-motor apparatus, the third nerve roots 
are both present and it requires close examination to discover 
anything unusual in them but the fibres are atrophied to some 
extent. The niduli, on the other hand, are nearly completely 
atrophied. The cells are nearly all shrunken and shrivelled and 
lie in large perilymph spaces. The contrast between these cells 
and the unaltered cells of the interpeduncular niduli is instruc- 
tive. (See Fig. 10.) In this case the fourth nerve has shared 
the fate of the third. While the atrophy has been more marked 
on the right side than on the left both are affected. The series 
unfortunately does not extend to include the sixth, but it is 


probable that the same condition existed in it. Our material is 
inadequate to illustrate the changes in the tracts and niduli of 


the second order. 
In attempting to analyse the various alterations with which 


we here have to do it is necessary to distinguish degeneration from 
atrophy and aplasy, the latter being a failure to develop asa 
result of operative interference. See Singer and Munzer, 
Denksch. Kaiserl. Akad. 2. Wien. 1890. 


DESCRIPTION OF FIGURES. 


Fig. z. Transection of rabbit brain (No. 292) immediately cephalad of 
the optic tecta, showing the atrophy of the tracts of the left side. The section 
is not exactly transverse. 

Fig. 2. Section through the tecta showing the atrophy of the left side and 
in the nidulus numbered ‘‘3.” 

Fig. 3. Section at the roots of the third nerves showing the absence of the 
right nidulus and root. 

Fig. 4. Portion of a section through the fourth nidulus showing that it is 
not affected by the atrophy. 

Fig. 5. Section at the exit of the sixth showing the absence of the fibres 
of the right side. 

Fig. 6. Part of a section further caudad showing the complete atrophy of 
the right nidulus. 

Fig. 7. Part of a transection of a rabbit brain (No. 297) from which the 
left eye was removed at birth, showing the degeneration of the right tract. 

Fig. 8. A-section further caudad. 

Fig. 9g. Section at the habenz showing atrophy in tract of second order 
on the right as well as in the optic tract. 

Fig. ro. Cells from the nidulus of the third (A) and from the interpedun- 
cular nidulus (B) to demonstrate the atrophic changes in the former. 

Fig. 11. Normal and degenerate portions of optic tract from 292, 8. 


LECTURE, NOTES “ON ATTENTION. 


An ILLUSTRATION OF THE EMPLOYMENT OF NEUROLOGICAL 
ANALOGIES FOR PsycHICAL PROBLEMS. 


C. L. HERRICK. 


At once a most difficult and a most important group of 
problems is offered in the study of attention. The importance 
grows, on one hand, out of the close connection of attention 
and its cultivation with the practical obligations of pedagogy, 
and, on the other, out of the theoretical relations of attention 
with fundamental mental endowments like volition and apper- 
ception. 

The source of the difficulty is also two-fold; first, the at- 
tempts at experimental study of attention suffer from the ambig- 
uity that, when all is done, the results are all apparently physi- 
ological rather than psychological. But this is only a conspicu- 
ous instance of what is true of experimental psychology as a 
whole. Care is necessary to discriminate the content in the 
mind from the physical accompaniments unless one is prepared 
to ignore that distinction entirely. The second difficulty is that 
different observers have different theoretical points of view and 
an unbiased stand-point is as impossible as it would be unpro- 
ductive. The purposeless and indiscriminate collection of facts 
rarely furthers science. Working hypotheses and a_ willing- 
ness to reject them when evidence requires are the theoretically 
desirable conditions, though in practice the willingness to reject 
is on the part of some other observer or.critic (none the less 
salutary in the result). 

It seems proper to divide attention into (A) external or 
sensory and (B) internal or cognitional. This is merely a work- 
ing division corresponding to the common experience that some- 
thing may ‘‘catch the eye,’ on one hand, and that we ‘‘con- 
centrate our thought”’ ona problem, on the other. It would 


b 


6 JOURNAL OF COMPARATIVE NEUROLOGY. . 


seem that popular language has crystallized an opinion that in 
one case the process is passive and in the other active. We are 
slow to reject the calm and impartial dicta of common sense 
—in fact, in some sense, this dictum is likely to be found cor- 
rect, though the meaning of active and passive in such connec- 
tions may suffer a good deal as the result of analysis. 

External Attention.—One of the first evidences of awaken- 
ing psychical life in the child is seen in the motions of the eye- 
balls as he attempts to follow moving objects, and one of the 
last evidences that is given that the departing spirit still hovers 
over the threshold is the same seeking motion of the eyes. As 
the most direct and unbiased avenue to consciousness, vision is 
the best sense from which to study the phenomena of external 
attention. Now if the eye be fixed upon a given field of view 
coinciding with the visual field of the retina it is evident that all 
parts of the field are capable of producing an impression on the 
retina and so on the nerve fibres and brain; in fact, each such 
spot does stimulate a point in the brain corresponding to the 
spot in the retina on which it falls. This neural irritation is not 
yet in consciousness but is a part of the ‘‘content of sense,” 
or sensory material for consciousness. Returning to the retina, 
if the field of view be uniform and uniformly illuminated and 
the gaze be fixed upon it under circumstances excluding other 
visual distractions a small disc will appear in the centre of 
the field of view which is somewhat more bright than the sur- 
rounding portions. : 

This little experiment is the physiological aspect of the 
fact that there is in the axial pole of the retina a spot in which 
only the cones are present and consequently the intensities of 
light and shade are more vivid than elsewhere in the retina. 
This relation would lose most of its significance were it not for 
another relation connected with it. 


1The descriptions of most of the experiments accompanying the lecture are 
omitted as they are of such a nature as will readily suggest themselves to the 
experienced reader. 


Herrick, Lecture Notes on Attention. 7 


Experiment.— Fix the single eye upon a faintly illuminated screen with per- 
forations near the margin of the field and a shutter behind in such a way that 
a ray of strong light may be revealed at will. Note that the axis of the eye in- 
voluntarily turns toward the ijluminated spot. Vary the experiment by using 
small colored images painted with transparent color and projected upon a trans- 
parent part of the screen, etc. Now determine to resist this tendency and keep 
the eyes immovably fixed on the original spot. Observe whether there is a 
feeling of tension in the eye, a strain on antagonistic muscles of the neck or in 
the scalp or even asense of tension in the brain. Interpret the results. Ob- 
serve what happens when the subject is encouraged to expect the new impres- 
sion on the right and then unexpectedly it appears on the left. A wider angle 
than the actual retinal one may profitably be used in these experiments while in 
other cases a diaphragm will be needed. In some it is an advantage to have 
the head fixed while it might be instructive in the deviation experiments in 
some cases to have the weight of the head counterpoised by braces under chin 
and occiput suspended to a swivel in the ceiling 

What is the result if the eye is centred on a bright, a varying, or a barely 
legible object? Is there a conflict of interest ? 

We have then in the eye a device for differentiating a part 
of the field of vision by reason of which that portion is more 
vividly reproduced in the brain and we have an anatomical adjust- 
ment of the muscles of the eyeball to enable this sensitive 
point to be turned upon any object which, by reason of the in- 
herent intensity, usurps the position. More than this, there is 
a very complicated system of nervous correlation reflexes which 
must have developed pavit passu with the anatomical structure 
by which the visual coordinations are affected anatomically.' 

It is of fundamental importance to decide whether the 
turning of the eye upon the vivid point in the margin of the 
field is due to volition or is spontaneous and a pure reflex. It 
seems certainly to be the latter. It may be noticed that in the 
fixing of both eyes on any field there is a complicated coordina- 
tion which has to be learned but passes out of voluntary con- 
trol, so far as the details of the process are concerned, very 
early. When any particular object is to be ‘‘ fixed ”’ in vision 
the two macule luteze must be properly adjusted as ‘‘identical’”’ 
spots of the retina and when this is done all other parts of the 
two images will fall on ‘‘identical” spots. Now when a bright 


or vivid spot appears near the margin of the field we might 


1 The instructor will review the visual-motor coédrdinating systems. 


8 JOURNAL OF COMPARATIVE NEUROLOGY. 


assume that the effect is the same as that of a false accommoda- 
tion of the maculz and the instinct to correct it has potence to 
make the new accommodation before an act of judgment can 
adjudicate the question. ' Whether this is an entirely satisfactory 
solution or not there is little doubt that external attention is of 
the nature of a reflex which may or may not retain a relation 
of subordinated connection with conscious processes. If care- 
ful attention is paid to what goes on during these involuntary 
acts of accommodation one may notice a certain sensation in 
the eyeball and muscles usually not felt at all. Now even 
where these sensational elements do not enter consciousness as 
such (or at all) they are stored up along with the retinal sensa- 
tion and may be employed in the formation of a sense of posi- 
tion—a concept of a spatial nature. However obscure the 
mechanism of this process, there is no doubt that it takes place. 
When we exercise ourselves in the effort to mentally estimate 
the position of various objects in the field of vision seen by aid 
of momentary illumination and discover that this too is possible, 
it simply seems to mean that associations have been fixed be- 
tween lateral cells in the retina and the muscle sensations pro- 
duced in bringing an image on the macula lutea through the 
necessary arc to impose it on that spot. 


Experiment.—Using a screen at a fixed distance from the eye the subject 
practices the estimation of distances until a degree of familiarity with the 
measures is obtained. Then in the dark try the same by instantaneous electric 
spark. Note accuracy. Tabulate results. Now try the same experiment vary- 
ing the duration of the illumination and observe whether there is an optimim 
period and, if possible, whether the eye is moved. Compare the two methods 
of estimation, 7. ¢., that where the eye motions are used directly and those 


where they are used indirectly. 

It is, then, obvious that the same class of sensations, viz. 
those derived from the act of accommodation, may be used in 
one case to produce asense of effort or may become, in another, 
data for localization. 


* Compare the labored attempt of ZIEHEN to explain this point. Lettfaden, 
p- 134 et seq. 


Herrick, Lecture Notes on Attention. fe) 


E-xperiment.—Glance casually over a page of print and note whether some 
phrase or word catches the eye. (The experiment is much more conclusive when 
done unintentionally). What word emerged? On repeating this experiment 
and glancing casually down the columns of the ‘*Times’’ the writer caught the 
word ‘‘ college”? and ‘‘university’’ many times also the word ‘‘ association ’’ 
several times and hardly another word. <A house keeper looking at the page of 
the ‘*Youth’s Companion ”’ 


? 


saw ‘* cooking” and ‘‘sweeping” first. 


The experiment may be conducted under scientific control in various ways. 
Arrange groups of figures similar in size and body of shade, most of them un- 
familiar, one or more familiar, and reveal them for ashort period (not too short), 
or produce familiarity by exposing squares covered by some one or more of the 
symbols as preparatory to the experiment. Note the effect of ‘‘ association.”’ 

The result of these experiments convinces that the selec- 
tion of one out of many impressions to be placed in the favored 
attitude in the sense organ is largely influenced by znterest 
and faézt. The word ‘‘interest’’ is here used in a popular and 
perhaps misleading sense. Probably in the sphere of the ex- 
ternal sense these effects are all the result of habit. For exam- 
ple, the reason that my eye caught the words ‘‘ college,” ‘‘ uni- 
versity,”’ and ‘‘association’’ out of their places among others 
was certainly not the superior interest which these words have 
for me, for they emerged before there was any consciousness of 
the interest. True, I had often associated in mind these very 
words in interesting connections and had consciously gone about 
to seek these words or passages containing them with the mind 
‘‘loaded,”’ so to speak, for this particular game. The result is 
a habit of selection which retains its unconscious power. The 
number of cotemporaneous sensations received by any sense 
is purely a matter of the structural adaptability of the organ or 
its natural field. All of the impressions of a given field of 
sense may become the content of that sense and so may exert 
their appropriate effects in infra-conscious spheres of association, 
etc., even though only part of them ever reach consciousness. 
The discussion as to the possible number of cotemporaneous 
sensations is based on a misconception. Though the content of 
sense may be diversified only one thing is ever in the focus of 
consciousness at a given time. 

So far we have been dealing with comparatively pure cases 
of external attention, but it is not to be denied that the con- 


10 JOURNAL OF COMPARATIVE NEUROLOGY. 


scious processes react on the sensory organs and modify their 
functions. It is a familiar commonplace that the degree of at- 
tention is very dependent on the feeling tone of a sensation. 
Two points are to be noted here: first, that the greater number 
of feelings associated with sense impressions have a practical 
value for the life of the individual and have so acquired an im- 
pulsive connection with external attention without calling con- 
sciousness into play; and, second, that the very nature of feel- 
. ings involves intensity of the stimulus (summation, on one hand, 
and irradiation, on the other,) so that this comes under the head 
of intensity as above. But let us turn to another form of this 
activity. I resolve to attend to the excitations of some sense 
or, aS we say, to keep eyes and ears open. This resolves itself 
into a more or less fruitless attempt to accommodate the organ 
voluntarily and results in for the most part ill-directed if not dis- 
turbing oscillations in the tension of the organ. Thus when in 
the night I strain the ear to catch a sound in no special locality I 
become conscious of the fact that the tensor tympani and stape- 
dius muscles are making castanets of my ears and audition is im- 
peded rather than facilitated ; so when I strain if perhaps I can 
make out another star in a vacant space I lose more than one 
‘‘dimension”’ before visible. One becomes conscious of a 
sense of effort in various organs, even in such as have no direct 
connection with vision and also of alterations in the somatic sen- 
sations. These sensations of effort, then, seem to us to prove 
the presence of attention, even voluntary attention, when there 
is absolutely no attention, for it is idle to speak of attention 
when nothing is attended to. It is only a cavil to say that in 
the case indicated there is attention to the sensations of effort. 
Such reasoning is an endless chain. When we really attend to 
something in the sensory sphere it is some specially selected an- 
ticipated thing. Even in the general case of listening for a 
noise we reproduce in thought one noise after another in ad- 
vance. ‘‘It will prove to be a creak, or a scratch, or the like.”’ 
Imagination meets the visual impression more than half way. 
‘Ts it a dagger that I see before me?”’ We are looking for 
something—our attention consists in the scrutiny of the content 


Herrick, Lecture Notes on Attention. II 


of sense with a reproduced vestige as a guide. The repro- 
duction may be of a very simple sort, as that of number—we 
may be repeating mentally, simply ‘‘ how many?” as we scru- 
tinize the sense content, but it is easy to see that the state 
of things here is very different from that in case of external at- 
tention—we are now dealing with a complicated central corre- 
lation across if not wholly within the field of consciousness. 
Internal Attention.—We have seen that internal attention 
popularly lays claim to active and conscious effort, but we have 
learned that such sense of mental effort may be, and sometimes 
is, simply the association of muscular sensations and that inter- 
nal attention implies a variety of coordinations in consciousness 
and we need not be blamed if we fall to doubting the validity 
of the sense of self-direction in internal attention. Internal at- 
tention is connected with the process of comparison either of 
the content of sense with some vestige or concept, or of two 
or more psychical elements along themselves. Thus we receive 
the advices from sense with a predetermined interrogatory, it 
may be, ‘‘Is the new impression the same as some other? 
Does it belong to the same class? Was it previously associated 
with A,B, .or'€.?) Is it as .big,.as loud) as \ persistent, jeter? 
In other words, we compare the new sensation with some stand- 
ard in reproduction and form more or less implicate judgements 
as to identity, similarity, difference etc., so that we find our- 
selves projected into regions in the very penetralia of the psy- 
chical life far from the supposed boundaries of mere attention. 
This experience is enough to show that attention is not a fac- 
ulty or independent activity of mind. Inner attention then 
may be but the measure of the vividness of consciousness 
under different conditions and attendant circumstances, whether 
of presentations of sense or central processes. But, it is proper 
to ask, what then becomes of the appearance of voluntary di- 
rection of attention? If it were true that we can force our 
minds to attend to any thing we choose, then this would become 
a matter of volition and need not require a separate treatment. 
Very brief reflection shows that we cannot attend to anything 
we please, for, in the first place, it is necessary that the thing 


12 JOURNAL OF COMPARATIVE NEUROLOGY. 


should be presented to consciousness. We cannot attend to a 
sensation not already a content of sense nor to an unreproduced 
reproduction. We can not manufacture a sensation to ‘‘attend 
to,’’ but can we perhaps reproduce a vestige not in conscious- 
ness? It is evidently absurd. In the case of the various sense 
contents we have seen that some have special attributes of in- 
tensity or special reinforcing collaterals to adapt them to affect the 
higher centres. May it not be true that the vestiges of con- 
scious processes (memory material) may have similar differential 
powers and special vehicles? The familiar fact that repetition 
facilitates reproduction shows that these vestiges are capable of 
being differentiated in their power of appeal to consciousness. 
Now experimentally let us undertake to recall the name of the 
man we met yesterday. ‘‘ Think’ as I may, no trace appears. 
I wrinkle the brow and strain all sorts of antagonistic muscles 
and set up a great sense of effort in the attempt to induce cere- 
bral innervation, in fact, I do succeed in forcing an unusual sup- 
ply of blood into the head and quickening the play of images 
but none of them is the right one. Try anew tack! The pre- 
sentations originally responsible for all the disturbance were of 
—a person—a man—met yesterday—and we have exhausted 
the associational images called up by ‘‘man” 
‘‘Jones, Jenkins, Jaynes, etc.’’ let us dismiss them and see 


—‘‘name ’’— 


what ‘‘ yesterday”’ has to offer. The vestiges are fresher—they 
rush upon me—more trouble to dismiss the undesired ones. 


Certainly here they are! ‘‘ Broadway—hospital ambulance— 
Dr. B’s clinic ’’—and, at last Dr. B. himself.- What could be 
more simple, I have ‘‘ recalled the name.’’ No, but was not 


what I really did the inhibiting of the undesired reproduction 
in order to let the chain of association follow its own laws of 
appearance? Thus we seem to have reached Wundt’s posi- 
tion that attention is essentially the results of inhibition. We 
cannot remain content with this conclusion for, first, practically, 
we are unable by introspection to find evidence of any such 
voluntary inhibition, second, theoretically, we seem to have 
gained nothing by such an assumption for the source and nature 
of the volition involved is as obscure as ever. What is certain is 


Herrick, Lecture Notes on Attention. 13 


that consciousness hovers from point to point in accordance with 
laws some of which we are able to formulate, if not to explain. 
Before leaving this point we meet the apparently legitimate crit- 
icism that what has been said applies rather to reproduction than 
to attention. Attention, it may be said, is not responsible for 
calling data to consciousness but for retaining them when there. 
However, observe that, if the conclusion that only one datum 
can be in consciousness at once be correct, experience shows 
that it is impossible to keep attention fixed for the fraction of a 
second on absolutely one presentation. There is a constant flux 
and alteration. Attention becomes then a set of rapidly re- 
peated reproductions. In thinking intently of one thing we 
limit the field of oscillation and cut off distractions as much as 
possible, but the oscillations with the various resulting associa- 
tions continue and give pregnancy to the meditation. Here 
again inhibition emerges as the agent of attention. Here too 
we are forced to reject the interpretation as above. The sense 
of voluntary effort in attention we may agree with Ziehen in 
ascribing to sensations of innervation, etc., that in no sense are 
a cause of the attention. But we would not stop there, but be- 
lieve that the sense of personal participation in our activities is 
of a more complex character than that suggested by the Miins- 
terberg-Ziehen school. 

To understand attention then one must understand the na- 
ture of consciousness. From its psychological side this is con- 
fessedly insoluble and we must be content either to pursue our 
search no further or to accept such suggestions as may come 
from neurolcgy and philosophy. In taking the latter course we 
are frankly entering a theoretical domain where only a balance 
of probabilities and reasoning from analogy can serve us. 

The organ of consciousness is the cerebrum and the great 
and distinguishing character of the hemispheres is the unlim- 
ited associational mechanism. All sorts of lower neural pro- 
cesses which are to influence consciousness have their offices 
here but these stations are all connected with each other. From 
all that we know of the analogies of nerve force it does not 
seem inherently impossible that a current from an eye centre 


14 JOURNAL OF COMPARATIVE NEUROLOGY. 


might if permitted to reach an auditory cortical station, produce 
some sensation—perhaps of a new kind of sound. ‘The anat- 
omical relations prevent an interference of this kind but the re- 
sults of the current’s action on the eye station may meet and 
modify a similar product of an ear station in a centre of a higher 
order. In the frontal lobes, perhaps, is a projection system of 
the highest order where the last fine adjustments occur. There 
must be, it would seem, a constant balance or resolution of 
forces in such a system. The centre of tension, if it could be 
seen, would be found flitting with greater rapidity that the re- 
flected disc of the electrometer of a testing outfit from one sta- 
tion to another in obedience to the various disturbances of equl- 
librium—due here to a new sense presentation, there to a nu- 
tritive adjustment removing a temporary inhibition. There is, 
to all appearance, only this sort of physiological unity possible. 
The only psychological unity is the unity of consciousness. Is 
there a connection between the two? Weare inclined to an- 
swer in the affirmative. While it seems necessary from consid- 
erations not necessary to enter upon here to consider conscious- 
ness itself a spontaneity of a higher than nervous nature, it is 
apparent that it finds its physical condition in this vortex or ner- 
vous ‘‘centre of gravity.”’ If this or something like this is the 
truth about consciousness, then attention is a name for the play 
of consciousness and a study of its laws reduces, on one hand, 
to the investigation of neural equilibrium and, on the other, to 
a natural history of consciousness. The conditions of inner at- 
tention are those of association and inhibition. 


XP NOEE ON, ViHip “CEREBRAL S PISSURATION °O 
THE SEAL) (Phocalvitulina)." 


PIERRE) A.) Fisa, 1D) Se; 


Bureau of Artal Industry, Washington, D.C, 


The specimen was not more than one year of age, and came 
into my possession in November, 1895. During the manipu- 
lation incident to the process of hardening some general resem- 
blance of the fissural pattern to that of the feline or canine type 
was noticed. The pia was removed and the fissures studied at 
an intermediate stage of the hardening before their walls had 
lost their pliancy. 

Dr. E. C. Spitzka (1890) in a preliminary paper (Ameri- 
ean Naturalist, XX VI. 115-122) refutes some rather grave mis- 
representions by Theodor, 1887 (Das Gehirn des Seehundes, 
Inaugural Dissertation) one of them being the existence of 
a so-called Commissura Suprema an elongated transverse mass 
of fibers lying dorsal to the Callosum, an artifact pure and sim- 
ple. Theodor further states that ‘‘ the Seals and (ordinary) 
Carnivora are in their cerebral organization today widely sep- 
arated and their common origin must be sought in a remote 
geological period.’”’ In regard to this Spitzka states that the 
examination of a series of brains beginning with the mink, the 
fresh and salt water otters and passing through the eared to the 
earless seals would show about as beautiful a transition asa 
morphologist could well desire. 

In the specimen at hand the frontal region of the brain is 
very much fore-shortened and gives the appearance of consider- 
able width. On the lateral aspect the Sylvian fissure is well 
marked and pursues its usual dorso-caudal course. Just in front 
of it and extending vertically (dorso-ventrally) is another un- 


' Read at the Meeting of the American .Anatomists Dec., 1895, Philadel- 
phia, Pa, 


16 JOURNAL OF COMPARATIVE NEUROLOGY. 


named fissure which superficially seems to run into the Sylvian 
at its base. This vertical fissure Spitzka has taken for the Syl- 
vian. He describes it as follows: ‘‘ It is the enormous hyper- 
trophy of this field (auditory cortical) which crowds the Syl- 
vian into its unusual vertical, nay anticlinal position.’”’ The 
blunt frontal end of the cerebrum is an additional reason for 
making the above condition possible but this view leaves a well 
marked fissure in the usual situation of the Sylvian unaccounted 
for. This opinion is still more difficult to accept when we come 
to ‘‘sound ”’ the fissures (examine their depths). The vertical 
and the true Sylvian fissures meet superficially at the latero-ven- 
tral angle of the cerebrum and if the sides of the Sylvian be sep- 
arated, it will be seen that the vertical fissure instead of directly 
joining the Sylvian becomesa submerged fissure, at this point 
corresponding to the preopercular area of the human brain, 
and crops out again on the ventral surface on the front or ceph- 
alic wall of the mouth of the Sylvian. In reality the vertical is 
an open surface fissure in its dorsal half and overlapped by the 
cortex (supergyre) in its ventral half. This condition is found 
on both sides. 

Tiedemann (Icones cerebri simiarum et quorundam mam- 
malium rariorum, Hidelbergia, 1821), ina figure of the base 
of the seal’s brain shows the ventral outcropping of a fissure in 
the cephalic wall at the base of the Sylvian which is none other 
than the terminus of the vertical. The present specimen con- 
firms this exactly. 

In the feline brain the fissure approximating most nearly 
to this vertical fissure of the seal would, in my opinion, be the 
anterior fissure; for if the frontal region of the cat’s brain 
could be moulded by any process of growth to the relatively 
fore-shortened condition of the seal’s brain the parts would as- 
sume very much the same relations. The fissuva postica of the 
cat’s brain does not seem to be represented in the seal. 

The super-Sylvian and the post-Sylvian are well marked. 
The two are continuous with each other at the surface on both 
sides; but at their union two branches are given off on the left 
hemicerebrum and one on the right hemicerebrum. The pres- 


Fisu, Cerebral Fissuration of the Seal. 17 


ence of subgyres at this point makes it very difficult to get ac- 
curate soundings ; and whether the union of these two fissures 
is anything more than superficial is therefore somewhat un- 
certain. 

The lateral fissure is long and tortuous and well on the dor- 
sal surface; its average distance from the intercerebral cleft 
being about one centimeter. On the left side it unites super- 
ficially with the super-Sylvian at the level of the vertical anter- 
ior fissure. On the right side the union is deeper and in ad- 
vance of the anterior. The caudal termination of the lateral 
fissure is at the extreme end of the crebrum and is in the shape 
of a T due to the union of a small transverse fissure, possibly 
the lunate of the cat. 

The ansate is a well developed fissure and is almost directly 
vertical in its course. It arises within less than a centimeter 
of the mesal surface caudal to the cruciate and extends ven- 
trally to about the level of the superficial confluence of the an- 
terior and Sylvian fissures. On each hemicerebrum there is a 
superficial union of the super-Sylvian with the ansate but in 
each there is evidence of a shallow (vadum) between them. - 

The diagonal of the cat is not distinctly represented in the 
seal, unless on account of the very much foreshortened con- 
dition it has become confused with the ansate or cephalic por- 
tion of the super-Sylvian. 

The super-orbital fissure arises from the base of the ol- 
factory bulb, which covers its origin, and extends in an oblique 
ventro-caudal direction to within one half a centimeter of the 
basi-Sylvian. 

The olfactory is a very short but deep fissure in which the 
delicate olfactory crus is almost completely imbedded. 

The cruciate fissure of the right side extends for a distance 
of a centimeter and a half on the dorso-lateral surface and has 
considerable depth. It also extends a centimeter and a half on 
the mesal surface. Very near the dorso-mesal margin a dorsal 
branch is given off and nearly opposite this is a ventral branch. 

The left cruciate fissure extends less than one centimeter 
on the lateral surface, but a minor fissure connecting with it 


18 JOURNAL OF COMPARATIVE NEUROLOGY. 


superficially makes it apparently extend over two centimeters. 
As with the right hemicerebrum a dorsal and a ventral branch 
are given off on the mesal surface, near the dorso-mesal margin. 
The cruciate continues superficially as far as the splenium with 
a fissure corresponding to the super-callosal. A super-callosal 
seems to be represented on the right side, but it does not join the 
cruciate. 

The splenial fissure arises at the dorso-mesal margin of the 
hemicerebrum at about the level of the splenium. It is oblique 
in its direction (ventro-caudal) and is more vertical than hori- 
zontal. It extends well on to the ventral aspect (about one 
centimeter). 

In the dog and cat cerebrum the fissures are for the most 
part isolated and distinct ; but in the seal there are numerous 
branches given off and rather an unusual amount of confluence 
of the larger fissures. The complexity is increased by the sinu- 
ous course and the presence, below the surface, of numerous sub- 
gyres or outgrowths along one wall, while the other wall overlaps 
and becomes more or less concave in adapting itself to this 
growth. This sinuosity at the depth of the fissure (much more 
marked than at the surface) makes an accurate sounding well- 
nigh impossible. The obliquity of the fissures and the over- 
lapping of one wall by the other, especially in the case of the 
ventral portion of the splenial amounts to almost an operculum 
or poma. These conditions in many regions render the accu- 
rate determination of a fissural integer very difficult. 

Underlying all there seems to be a fissural pattern not un- 
like that of the feline to which are added the various complex- 
ities above enumerated, but these are hardly sufficient to ob- 
scure the carnivore type. 


Fisu, Cerebral Fissuration of the Seal. 19 


PATE Uk: 

REFERENCE LETTERS FOR THE FISSURES. 
ans,—ansate. mi.—medilateral. 
ant,—anterior. Post.—postica. 
Cal.—callosum P. Syl.—post-Sylvian. 
Con.—confinis. rh.—rhinal. 
Cor.—coronal. sc.—supercallosal (?) 
Cr.—cruciate. so.—superorbital. 
Diag.— diagonal. spl.—splenial. 
7.—lunate. Sp. Syl.—super-Sylvian, 
Zat.—lateral. Syl.—Sylvian. 


m.—marginal. 


EXPLANATION OF FIGURES. 


In all of the figures the important fissures are drawn with heavy lines and 
the minor fissures with light ones. The first four figures were photographed 
and enlarged drawings made. A small cross indicates a shallow depth of the 
fissure at that point. The fissural names are based upon those used in the 
Anatomical Technology (Wilder and Gage). 


fig. z. Lateral view of the left hemicerebrum of the seal. 
Fig. 2. Mesal view of the right hemicrebrum. 

Fig. 3. Lateral view of the right cerebrum. 

Fig. 4. Mesal view of the left hemicerebrum. 


fig. 5. The base of the seal’s brain, after Tiedemann. This figure shows 
the ventral outcropping of the vertical anterior (?) fissure in the cephalic wall 
of the Sylvian. 


fig. 6. The dorsal aspect of the seal’s brain, after Tiedemann. 

Fig. 7. Lateral aspect of the left hemicerebrum of the cat (diagrammatic). 
Fig. 8. Mesal view of the cat's right hemicerebrum (diagrammatic). 

fig. g. Schematic type for brain of Carnivora, after Pausch. 

fig. ro. A cross section of a fissure, to show the obliquity. 


Fig. zr. A diagram to show the difference in the course of a fissure at its 
surface and depth. The heavy lines represent the fissural walls at the surface. 
The dotted lines and arrows represent the deep course of the fissure. 


MORPHOLOGY OF THE NERVOUS SYSTEM OF 
CYPRIs: 


GC, AHS SP URNER: 


With Plates ITI—V/1/I. 


Introduction. 


About three years ago, on publishing a, ‘‘ Preliminary 
Note on the Nervous System of the Genus Cyfrzs,”’ I promised 
to publish, in the near future, a histological monograph on the 
Ostracoda. This paper is intended as the first installment of 
such a monograph. In this communication it is intended to 
give a detailed discussion of the central and peripheral nervous 
systems of the Cypride. 

Owing to its large size, Cypris herrickt Turner has been 
selected as the type to study. To eliminate all mere size-vari- 
ations, Cyprinotus incongruens Ramdohr has been used as a 
check. For the study of the central nervous system, these 
two species have been considered representative types of the 
Cypride ; but, in studying the peripheral sense organs, use has 
been made of all accessible genera. These genera are: Can- 
dona, Cyclocypris, Cypria, Cypris, Cyprinotus, and Cypridopsas. 

This paper being the embodiment of neurological investi- 
gations, any prolonged discussion of extra-neural systems 
would be out of place; yet, in order to facilitate a comprehen- 
sive discussion of the nervous system, a few terse histological 
statements upon the associated systems have been made. 


BIBLIOGRAPHY. 
Craus: (C: 
1. Ueber die blassen Kolben und Cylinder an den An- 
tennen der Copepoden und Ostracoden. Wairzburg 


naturw: Let.’ 3, Bd;, 1860, Tat. Vil, Pisa. 


TURNER, Nervous System of Cypris. 21 


2. Ueber die Organization von die Cypridinen. Zeit. f. 
WISH ZOOL. daw LOO, LOD. AA 15, elaine Ne, 

3. Abs. do. in Anzerg. kats. Akad. wiss. Wien, Vol. VII, 
1890, pp. 55-60. 

ao trans. do: ine A Van ase Nol om uly, aps MOS: 

5. Neue Beobachtungen tber Cypridinen. Zezt. f. wrss. 
Z00ts, BC. XO, ppe 2 WI-277., spleens. 7,. 3) 

6. Das Medien-auge der Crustaceen. Ard. Inst. Wren, 
Bd EXS pp. 225-206; 4 Vat) 18091: 

7. Ueber den feineren Bau des Medien-auges der Crusta- 
ceen. Anz. Akad. Wien, p. 124-127, 1891. | 

LANG. 

8. Text-Book of Comparative Anatomy. Trans. by 

Bernard, p. 344. 


REHBERG, HERMANN. 
g. Beitrage zur Naturgeschichte niederer Crustaceen. Brve- 
men, 1884, p.. 11-13, Taf. II. 
TurRNER, C. H. 
10. Preliminary Note on the Nervous System of the Genus 
Cypris. Jour. of Comp. Neur., Vol. Ill, 1893, pp. 35- 
AO; pl Lil and liv: 
VAVRA, WENZEL. 
11. Monographie der Ostracoden Bohmens. rag, 1891, 


Pp: 16-10; Mis, 14: 


HISTORICAL RESUME. 


Among the legions of histological monographs that recent 
instruments and improved technique have called forth, those on 
the neurology of the Ostracoda form an all but insignificant part. 
To the best of my knowledge the first to write on this subject 
was Leydig. He described the peculiar seta found on the sec- 
ond joint of the antenna and called it a sensory seta. 

Then followed Dr. Carl Claus, who, in 1860 (1) discussed 
these same sete and called them ‘‘d/asse Kolben und Cylinder”’ 
or ‘‘ Leydische organe.”’ In 1865 (2) the same author discussed the 
eyes of Cypridina, laying special stress upon a simple eye 


22 JOURNAL OF COMPARATIVE NEUROLOGY. 


which occurs in front of the paired eyes, because he thought 
that the presence of that eye renders quite probable the 
hypothesis that the compound eyes of the Cypride 
are analogous to those of the Cladocera. According to 
Dr. Claus this median simple eye consists of three layers; Ist, 
an outer transparent lens; 2nd, an inner nervous layer ; 3rd, an 
intermediate pigment layer. Both in form and position the 
outer portion of this eye resembles the lens of the Heteropod eye. 
The non-pigmented striated portion, into the substance of which 
nerve fibres penetrate, contains elongated cells. Later Dr. 
Claus (5) showed that in Cyfrzdina the size and position of the 
paired eyes serve to differentiate the sexes. In the males, the 
paired eyes are almost in the middle of the shell, while the much 
larger, unpaired, median eye is imbedded ina swelling of the 
forehead. In 1891, Dr. C. Claus (6) stated a theory concerning 
the compound eyes of the Ostracoda. 

Dr. Rehberg (g) also has contributed something to further 
this cause. Among other things he has minutely and accurate- 
ly described the tip of the second foot of Cyprts. In his eyes, 
this member is something more than a leg functioning as an 
ovipositor, for in its tip it bears a sensory organ which is proba- 
bly auditory in function. In addition to this he has described, 
in the following manner, the sensory seta found on the third . 
joint of the antenna. This sensory seta consists of: Ist, a 
chitin-surrounded proximal portion; 2nd, a short transparent 
middle-piece ; 3rd, a granular knob-like distal portion. Its nerve 
enters through a hole in the chitinous exoskeleton. ; 

Later Dr. Lang (8) summarized the accessible literature 
on the central nervous system of the Os¢racoda in the following 
words: ‘‘ The ventral chain of Cythere, which follows the brain 
and circum-oesophageal commissures, is said to consist of an in- 
fra-oesophageal ganglion and four subsequent ventral ganglia. 
The infra-oesophageal ganglion is said to show its composition out 
of two ganglia and to innervate the jaws, while the three subse- 
quent ganglia give off nerves to the limbs, and the last ganglion, 
nerves to the most posterior divisions of the body and the genital 
apparatus. In contrast with the above, the ventral chain of Ha/o- 


Turner, Nervous System of Cyprts. 23 
J S) 


cypris appears much concentrated. It consists of an infra-oeso- 
phageal ganglion, with nerves to the jaws and maxillipeds, and 
a small ventral ganglion. Out of the latter arise two pairs of 
nerves, which probably innervate the musculature of the limbs 
and the abdomen. 

Finally comes Wenzel Vavra who, in his late work (11), has 
devoted three pages to the discussion of the nervous system of 
the Ostracoda. In that work he makes the following statements: 
Ist, the central nervous system consists of a brain and a five- 
ganglionic ventral chain; 2nd, the optic and antennulary nerves 
arise in the brain, but the antennary nerve arises in the pharyn- 
geal collar ; 3rd, in the five-ganglionic ventral chain which extends 
to the genital apparatus, the anterior three ganglia are closely 
approximated; 4th, from each ganglion a nerve passes to a pair 
of limbs and from the last ganglion a nerve passes to the genit- 
al apparatus ; 5th, what Dr. Claus says about the structure of the 
eye is correct; 6th, in Cyprza and Cyclocypris the eye is large 
and the pigment black, in Cazdona the eye is inconspicuous and 
the pigment reddish; 7th, in 7yphlocypris, although the eye of 
the embryo is pigmented yet the eye of the adult is a degener- 
ated, non-pigmented, sense organ; 8th, in Notodromas the three 
eyes are separated, but each of the paired eyes is connected, by 
means of a stalk, with the median eye, and each of these three 
large eyes receives a distinct nerve from the brain; oth, the 
sensory seta of the second antenna, which is longest in 7yphlo- 
cypris, Candonopsts and Cyclocypris, is an olfactory organ; 10th, 
the ‘‘blasse kolben”’ on the last segment of both the anten- 
nules and antennae are composed of two segments; 11th, on 
the fifth segment of the antenna of male and female specimens of 
Notodromas there occurs a special sensory seta; 12th, in Can- 
dona, Candonopsis, and Cypria the distal extremity of the 
fourth segment of the male antennae bears a characteristic seta. 


TECHNIQUE. 


For hardening and fixing Ostracodes alcoholic picro-sul- 


24 JOURNAL OF COMPARATIVE NEUROLOGY. 


phuric has proven the best fluid. This fluid is compounded in 
the following way: 

70 per cent. alcohol. ; : ; 100 vols, 

Sulphuric acid, conc. . : : : 2 vols. 

Picric acid, as much as will dissolve. 

For the above formula I am indebted to Dr. J. Playfair 
McMurrich, now of Ann Arbor. Since this fluid sometimes 
causes thin shells to curl, it is not so good for museum speci- 
mens as 70 per cent. alcohol ; but for histological work it is ex- 
cellent. It does not injure the tissues, it penetrates chitin easily, 
and it fixes the cell structure. The living specimens were 
placed in this fluid and allowed to remain for at least twenty- 
four hours. Then after being washed in 70 per cent. alcohol 
until all the picric acid had been removed, they were either 
transferred to the stain or else, after being hardened and sec- 
tioned in the usual way, they were stained on the slide. 

Among the numerous stains tried were: Kleinenberg’s he- 
matoxylin, Delafield’s hamatoxylin, Czokor’s alum cochineal, 
borax catmine, eosin, etc., but none of these gave satisfactory 
results. Finally Ehrlich’s hamatoxylin and tincture of alum 
cochineal were tried. Both of these stains gave good results. 
In using both of these stains the specimens were stained in toto. 
When Ehrlich’s haematoxylin was used, the specimens were over- 
stained and then washed out with acidulated 70 per cent. alco- 
hol; but when the tincture of cochinea] was used, after remain- 
ing in the stain for about twenty-four hours, the specimens were 
washed in 70 per cent. alcohol and then hardened and sectioned 
in the usual manner. 


PRELIMINARY. 


Although a lengthy discussion of the histology of extra- 
neural systems would be foreign to the purpose of this paper, 
yet, in order to facilitate a comprehensive description of the 
nervous system, it is thought best to give a brief description of 
the internal parts of the Ostracoda. 

All previous writers to the contrary notwithstanding, the 
shell of the Ostracoda consists of three layers: an ectal layer, 


TurRNER, Wervous System of Cypris. 25 


which has been hardened by the deposition in it of calcium car- 
bonate; an ental, thin, flexible layer; and an intermediate layer 
of connective tissue [fig. 1]. Where the shell is united to the 
body the ental layer is absent [fig. 1, 6]. The connective tis- 
sue in the shell is a continuation of the connective tissue of the 
body. Most of the space between the ental and ectal layers 
not occupied by the connective tissue, is filled by the gonads 
and lateral evaginations of the mid-gut. 

The body proper consists of two regions, a broad cepha- 
lothorax anda narrower abdomen. The abdomen is movable 
on the cephalothorax [fig. g]. Seven of the eight pairs of ap- 
pendages arise from the cephalothorax, while the remaining pair 
(the abdominal rami) arises from the caudal extremity of the 
abdomen. 

From near the dorsal extremity of the cephalic margin of 
this creature arise the antennules and from a little below the 
middle of the same margin arise the antennae. Passing caudad, 
from the ventral margin arise, in quick succession, the mandibles, 
Ist maxillae, 2nd maxillae, rst leg, 2nd leg. 

The mouth, which lies between the mandibles, is bordered 
by an unpaired upper and by paired lower lips. These, to- 
gether with the mandibles, not only triturate the food, but serve 
also as the locus of three pairs of sense organs [fig. 16]. The 
short oesophagus, which is bordered with large cells [fig. gc], 
extends almost vertically dorsad into the fore-stomach [fig. g]. 
In the walls of this grinding apparatus cells can be seen. Now 
follows the stomach, from the walls of which lateral evaginations 
project into the shell. The wall of this stomach is composed 
of a single layer of columnar cells [fig. 37,41]. The dorsal por- 
tion of the stomach is quite large, extending from the cephalic 
extremity of the cephalothorax to the beginning of the abdo- 
men [fig. 9,16]. The intestine extends through the abdomen 
to the dorsally located anus, which is situated near the abdomi- 
nal rami [fig. g]. 

The ovaries lie between the shell Jayers. After entering 
the body on the dorsal side just caudad of the abdomen and 
thence passing ventrad into the abdomen, the sexual ducts pass 


26 JoURNAL OF COMPARATIVE NEUROLOGY. 


ventro-meso-caudad to the median unpaired vagina, which lies 
on the ventral side near the abdominal rami. The walls of the 
abdominal portions of the sexual ducts are quite glandular. 
They secrete the chorion of the egg. 

Near the ventral side of the body there is a broad chitin- 
ous exoskeleton [fig. 1,5], to which the muscles of the extrem- 
ities are attached. 

Surrounding all the organs of the body (nervous system 
included) we find a mesenchyme-like connective tissue. This 
connective tissue consists of membranous and fibrous cells, 
among which are scattered larger and denser irregularly shaped 
cells [fig. 1,11]. 

It is important that no one should suppose that the above 
description is intended as an exhaustive histological study of 
the Ostracoda. Although it contains some points that have not 
yet been brought out by other investigators, nevertheless, it is 
given merely to facilitate an intelligent description of the ner- 
vous system and its relation to the extra-neural systems. 


The Central Nervous System. 


The central nervous system of Cyfris, like that of the 
higher crustacea (J/alacostraca) is composed of a supra-oesoph- 
ageal ganglion which is united by a pharyngeal collar to a multi- 
ganglionic ventral chain. This is true, not only of Cyprzs, but 
also of Cypridopsis, Cyclocypris and Candona. When we recall 
that Dr. Lang states that in Cythere and Hfalocypris this is also 
the case, we have grounds for believing that in all Ostracoda the 
central nervous system consists of a supra-oesophageal ganglion 
which is united to a ventral chain by a pharyngeal collar. 

Supra-oesophageal ganglion.—This ganglion lies about half 
way between the dorsal and ventral surfaces of the body and be- 
tween the oesophagus and the cephalic border of the body. It 
lies just below the place where the oesophagus enters the mid- 
gut. In my preliminary paper the statement was made that 
this ganglion was much nearer to the dorsal than to the ventral 
surface. This was a misprint; the ganglion usually lies nearer 
the ventral than the dorsal surface. This is a compound gan- 


TuRNER, Wervous System of Cypris. 27, 


glion. Although small, yet this supra-oesophageal ganglion is 
probably compounded out of seven distinct ganglia. These 
problematic ganglia are: three ganglia that have fused to form 
the unpaired optic ganglion, two antennulary ganglia, and two 
antennary ganglia. 

The optic ganglion is located in the roof (dorso-cephalic 
portion) of the supra-oesophageal ganglion. Although in the 
adult it is a median unpaired structure, yet its histology seems 
to indicate that it is a triune structure. The optic nerve arises 
from the apex of this ganglion. In Cypvzs and the allied forms 
examined by me, the optic nerve is a single unpaired nerve; but 
in Votodromas, according to Wenzel Vavra, three optic nerves 
arise from the brains The fact that in Motodromas three nerves 
arise from the optic ganglion lends support to the view that this 
is a triune ganglion. However, we must look to embryology 
for a final settlement of this question, and the necessary data 
are not at hand. Wenzel Vavra has called this portion of the 
supraoesophageal ganglion from which the optic nerve arises the 
fore-brain. 

The paired antennulary ganglia occupy the lateral portion 
of the supra-oesophageal ganglion [fig. 10]. Wenzel Vavra 
has called this portion of this ganglion the mid-brain. From - 
the dorsal portion of each side of this ganglion, arises a nerve. 

Further ventrad, but nearer the meson, in the lateral por- 
tion of the supra-oesophageal ganglion lie the paired antennary 
ganglia [fig. 10]. A portion of this ganglion lies in the phar- 
yngeal collar. Wenzel Vavra has called this portion of the gan- 
glion the hind brain. 

The supra-oesophageal ganglion is connected, by means of 
a pair of circum-oesophageal commissures, with a sub-oesopha- 
geal ganglion [fig. 16, 42]. These commissures pass obliquely 
backwards and downwards (ventro-caudad) from the lateral por- 
tion of the supra-oesophageal ganglion to the corresponding 
portions of the sub-oesophageal ganglion. These two commis- 
sures constitute the pharyngeal collar. This collar is not merely 
a commissure but it is also the locus of a nerve centre. All along 


28 JOURNAL OF COMPARATIVE NEUROLOGY. 


its front (cephalic) border extends a chain of ganglionic cells. 
The chain of cells is a portion of the antennary ganglion [fig. 3]. 

As in all invertebrate nervous systems, so here, the central 
nervous system is composed of two things: nerve cells and 


neuroglia (‘‘Punktsubstanz”’ of Leydig, ‘‘ Marksubstanz”’ of 
Dietels and Rawitz, ‘‘ central Nervennetz’’ of Bellonci and 


Haller). In the supra-oesophageal ganglia of Cyprts the nerve 
cells are aggregated in the dorsal, cephalic, ventral and lateral 
portions of the periphery, while the neuroglia is central. 

Ventral Nerve Cord.—In his comparative anatomy, Pro- 
fessor Lang has said: ‘‘ The ventral cord of Cythere which fol- 
lows the brain and oesophageal commissures is said to consist of 
an infra-oesophageal ganglion and four subsequent ventral gan- 
glia. The infra-oesophageal ganglion is said to show its com- 
position out of two ganglia and to innervate the jaws, while the 
three subsequent ganglia give off nerves to the limbs, and the 
last ganglion nerves to the most posterior division of the body 
and genital apparatus. In contrast with the above, the ventral 
chain of Hlalocypris appears much concentrated. It consists of 
an infra-oesophageal ganglion, with nerves to the jaws and max- 
illipeds, andasmall ventral ganglion. Out of the latter arise 
two pairs of nerves, which probably innervate the musculature of 
the limbs and abdomen.”’ 

Morphologically the ventral chain of Cypris is intermedi- 
ate between these two extremes. It is more concentrated than 
that of Cythere but not quite so compact as that of Halocypris. 
In this case the ventral chain consists of an infra-oesophageal 
ganglion and two subsequent ventral ganglia. All of these 
nerve ganglia are connected, not only by two longitudinal com- 
missures, but also by straggling chains of nerve cells [fig. 
32,42]. To repeat, the ventral chain of Cypris is composed of 
three pairs of ganglia, which are united by commissural fibres 
and nerve cells. 

Now Wenzel Vavra has recently stated that the ventral 
caiuia of Cypris is conjosed of five ganglia. At first blush 
these two views seem to be irreconcilable. However, when crit- 
ically compared, the two views are not so antagonistic as they 


TuRNER, Wervous System of Cypris. 29 


seem. Dr. Wenzel Vavra confesses that ‘‘ Die ersten dret Gan- 
glien stehen sehr gedrangt.’’ Although, in the adult the sub- 
oesophageal ganglion of Cypris is an indivisible unit, yet my 
investigations lead me to believe that it has been compounded 
out of at least three pairs of distinct ganglia. Now if by say- 
ing that the first three ganglia are closely compacted Wenzel 
Vavra means that they are compacted sufficiently to form a tri- 
une ganglion then our views harmonize. 

From this compound ganglion arise the following five pairs 
of nerves: nerve of the upper lip, mandibular nerve, nerve of 
the lower lip, thoracic nerve. 

The two halves of this ganglion are connected by three 
transverse Commissures. 

The first pair of ganglia behind (caudad of ) the sub-oesoph- 
ageal ganglion gives off a pair of nerves to the second maxillae. 

From the last ganglion arise two pairs of leg nerves and 
one unpaired abdominal nerve. 

In the region of the sub-oesophageal ganglion the ventral 
chain lies above (dorsad of ) the endoskeleton, while back of that 
region it lies below (ventrad of) the endoskeleton. 

In the region of the sub-oesophageal ganglion the nerve 
cells are confined to the ventral and lateral surfaces of the gan- 
glion, while in the rest of the chain, the nerve cells occupy all 
of the periphery and also the mesal plane. 

Root gangla.—At the origin of certain nerves there is a 
mass of ganglion cells. The ganglion is often quite intimately 
united with the central nerve chain. 


NERVES. 


The principal nerves of Cyfris are: the optic, the anten- 
nulary, the antennary, the labial, the mandibular, the labral, 
the two maxillary nerves, the thoracic, the two leg nerves and 
the abdominal nerve. 

Optic Nerve.—The optic nerve arises from the apex of the 
optic ganglion, which is also the apex of the brain. In Cypvis, 
Cyprinotus and allied forms examined by me, this is a median 
unpaired nerve. At a distance from the brain which varies 


30 JOURNAL OF COMPARATIVE NEUROLOGY. 


in different species, this nerve splits into three branches—one 
for each of the divisions of the median tripartite eye. Passing 
upwards (dorsad ) between the mid-gut and the cephalic border 
of the body these branches pass to the compound eye, which 
lies near the dorsal surface of the body. Indeed, this median 
eye lies against the shell, vertically above the supra-oesopahgeal 
ganglion | fig. 2, 9, 100, 420 |. 

In Notodromas, according to Wenzel Vavra, there are three 
optic nerves arising from the apex of the brain [ fig. 30]. Here 
also we find that the three parts of the eye are widely separated 
[ fig. 30]. It does not require a very active imagination to 
conclude that simultaneously with the divarication of the com- 
ponent parts of the triune eye there went a longitudinal tripar- 
tite splitting of the entire optic nerve. 

Antenaulary Nerves.—Vhe paired antennulary nerves arise 
one from near each lateral surface of the apex of the supra- 
oesophageal ganglion [fig. 10, 42]. After leaving this gan- 
glion the nerve passes in a caudally convex curve upwards 
(dorsad ), between the mid-gut and the cephalic border of the 
body, to the corresponding ganglion. There is no root gan- 
glion at the base of this nerve. 

Antennary Nerves.—Where each circum-oesophageal com- 
missure leaves the supra-oesophageal ganglion arises an anten- 
nary nerve [ fig. 3]. This nerve receives its fibres, not only 
from the brain, but also from a ganglion that extends in a strag- 
gling manner along the pharyngeal collar | fig. 2, 8 ]. 

Labial Nerves. —The labial nerve or nerve of the upper 
lip arises in the cephalic portion of the sub-oesophageal ganglion. 
Instead of leaving the central nerve chain immediately, this 
nerve passes upwards (cephalo-dorsad ) into the pharyngeal col- 
lar and thence forward (cephalad ) to the upper lip [ fig. 3, 16, 
42]. There is a large root ganglion at the origin of this nerve 
[fig. 3]. The labial nerve has two main branches. One 
branch innervates a pear-shaped sense organ situated in the 
front part of the body below [ ventrad of | the base of the an- 
tenna, while the other innervates a harp-shaped sense organ 
located in the lower [ventral] portion of the upper lip [fig. 16]. 


TuRNER, WVervous System of Cypris. 31 


Mandibular Nerves.—From each lateral portion of the sub- 
oesophageal ganglion, just below (ventro-caudad of) the place 
of fusion of the pharyngeal collar with this ganglion, a man- 
dibular nerve leaves the nervous chain. Just after entering the 
mandible this nerve branches, one branch passing upwards 
(cephalo-dorsad ) to the mandibular muscles, while the other 
passes downward (cephalo-ventrad ) to a sense organ at the 
base of the mandibular teeth [ fig. 7 mx, 40 md]. There is 
no root ganglion to this nerve. 

Labral Nerves.—From each lateral portion of the ventral 
aspect of this same ganglion, a short distance behind (caudad 
of ) the origin of the mandibular nerve, arises a labral nerve. 
Thence the nerve passes, in a caudally convex curve, downward 
( ventrad ) to the lower lips [ fig. 32, 42 ]. The chief function 
of this nerve is to innervate a harp-shaped sense organ located 
in the lower lip [ fig. 16 D]. This nerve has a root ganglion. 

first Maxillary Nerves.—A_ short distance caudad of the 
origin of the mandibular nerve, a maxillary nerve leaves each 
side of this same ganglion. This is the first maxillary nerve. 
It passes latero-caudad to the first maxilla [fig. 7 mx”, 42]. 

Thoracic Nerves.—Al\\ of the four pairs of nerves just de- 
scribed arise from the ventral side of the sub-oesophageal gan- 
glion. Wenow come toa pair of nerves that arise from the 
dorsal aspect of the same ganglion. These are what I beg per- 
mission to call the thoracic nerves. These nerves arise from 
the lateral portion of the dorsal aspect of this ganglion just 
back of (caudad of) the origin of the first maxillary nerve. 
Arising as it does from the dorsal aspect of the ventral chain, 
the root of this nerve lies immediately beneath the endoskeleton. 
This nerve passes obliquely upwards and backwards (dorso-latero- 
caudad), and, after branching, innervates the abductor muscles 
of the shell [fig. 327, 427]. 

To the best of the writer’s knowlege, neither this nerve nor 
the labial nor the labral nerves have been described by any pre- 
vious writer on the Ostracoda. 

From the cephalic extremity of the ventral nerve chain to 
the roots of the thoracic nerve the nerve cells are confined to 


32 JouRNAL OF CoMPARATIVE NEUROLOGY. 


the ventral and ventro-lateral surfaces of the cord. Behind 
(caudad of ) the roots the nerve cells entirely surround the gan- 
glia and occupy the mesal plane as well. 

Within the ventral chain, beneath the origin of the thor- 
acic nerves, there is a pair of ellipsoidal cavities. These might 
properly be called thoracic ventricles. 

Second Maxillary Nerves. —TVhis nerve arises from the lat- 
eral aspect of the first ganglion behind (caudad of) the sub- 
oesophageal ganglion. 

Leg Nerves.—From the last ventral ganglion of Cypras 
arise two paired nerves and one unpaired nerve. The paired 
nerves innervate the legs, while the unpaired nerve passes into 
the abdomen. The nerve of the first leg passes immediately 
latero-ventrad to the first leg; but the nerve of the second leg 
passes backwards (caudad) along with the abdominal nerve, a 
short distance before passing to the second leg [fig. 42]. 

Abdominal Nerve.—As has been stated, from the caudal 
portion of the last ventral ganglion a large unpaired median 
nerve passes upwards and backward (dorso-caudad) into the ab- 
domen. This is the abdominal nerve. After entering the ab- 
domen, this nerve becomes more and more attenuated, owing 
to the fact that it gives off fibres to the reproductive system 
[fig. 32. an, 42 Ad]. 

The above descriptions of the principal nerves of Cyprzs do 
not agree either with the descriptions of the few men who have 
written on this subjects or with the statements made in my pre- 
liminary paper. I cannot answer for the other men, but my 
error was in mistaking the labial nerve for the mandibular nerve. 
If you but recall the compactness of the anterior (cephalic) re- 
gion of the body of Cyfrzs, you can see how easy it would be 
to make such a mistake. However, I now have at hand sev- 
eral series which demonstrate, beyond the shadow of a doubt, 
that the nerves arise as stated in this paper. 

No doubt it is this same compactness of the cephalic por- 
tion of the body of Cypris which is responsible for the non-dis- 
covery of the labial, labral, and thoracic nerves by other 
observers. 


TurNER, Nervous System of Cyprts. 33 


Careful search has not yet revealed a sympathetic nervous 


system. 
Sense Organs. 


Compound Eye.—The most conspicuous sense organ of the 
genus Cypris is the compound eye. When viewed from above, 
this eye usually resembles a quadrilateral pigment spot [fig. 36]. 
In Cyclocypris and Cypria, this eye is quite largeand the pigment 
is intensely black; in Cypris and Cyprznotus, the eye is of medi- 
um size ; in Candona, the eye is inconspicuous and the pigment 
is reddish ; while in Zyphlocyprits, according to Wenzel Vavra, 
there is no eye in the adult. The eye of the embryonic 7yphlo- 
cypris, according to the same author, becomes metamorphosed 
into an inconspicuous non-optical sense organ. 

. This eye is a triune structure. It consists of a median and 
two lateral portions, each of which is supplied with a lens 
[fig. 2]. The lens of the median portion is on the front aspect 
[fig. 36], while the lenses of the lateral portions are on the 
sides [fig. 2, 36]. This median compound eye, which lies in 
about the same vertical transverse plane as the supra-oesopha- 
geal ganglion, is located where the dorsal border of the animal 
is united to the dorsal portion of the shell. Hence there are two 
planes, at right angles to each other, that pass through both the 
compound eye and the supra-oesophageal ganglion. These 
two planes are the mesal and a transverse plane. 

Histologically there are five parts to this compound eye: 
lens, retina, pigment, superficial epithelium, nerve [fig. 2]. 
Corresponding to each of the three divisions of the eye we 
have a retina and a lens [fig. 2] anda nerve. The pigment is 
deposited between and around the retina. The retinas are cell- 
ular structures, the cells being arranged with their longest axis 
perpendicular to the outer surface of the retina [fig. 2]. Each 
retina is supplied with a lens, and these are always on the outer 
surface. In the lateral portions these lenses are on the side, 
while in the median the lens is on the front aspect [fig. 36]. 
The median portion lies lower than the lateral. Excepting the 
central portions of the lenses, all of the surface of the eye, as 
well as the space between the retinas, is pigmented [fig. 2, 36]. 


34 JOURNAL OF COMPARATIVE NEUROLOGY. 


In Cypris herricki Turner the nerve fibres are united to the 
outer (peripheral) ends of the retinal cells. This agrees with 
what Dr. Claus found to be the state of things in the eyes of 
the Copepoda, Clodocera and Ostracoda examined by him [7]. 

There is a striking resemblance between the triune eyes of 
the Ostracoda and those of the Copepoda. In describing the 
compound eye of Calanella mediterranea, Dr. Grenacher remarks 
‘‘Each eye is composed of a pigment cup and a strongly re- 
fractive transparent ‘lens’ laid in and on it. The term lens is, 
however, not applicable. It is composed of several cells, each 
of which is connected, whether at its outer or inner side is not 
yet certain, with a fibre of the optic nerve, and must therefore 
be considered asa retinal cell.’’ Since the superficial resem- 
blance between the Ostvacod eye and the Copepod eye is so great, 
one is apt to conclude that the above words of Dr. Grenacher 
might be applied to the Ostvacod eye. Against such an assump- 
tion I desire to enter a most emphatic protest. As stated above, 
in the eye of Cypris we find both retinal cells and lenses, and 
the two structures are histologically quite distinct. On the out- 
side of the eye, covering lens and all, there is a nucleated epi- 
thelial layer [fig. 2]. 

Since the publication of the paper’ in which O. Butschli 
attempts to show how the lateral eyes of the vertebrates may 
have been derived from the median tripartite eye of Sa/pa, this 
tripartite eye, which is of almost universal occurrence among 
the entomostraca, becomes enhanced in interest. One at once 
begins to speculate on the possibility of this median triune eye 
of the entomostraca becoming transformed into the lateral eyes 
of the higher crustacea. Indeed, in his late work, Dr. Claus” 
has already stated that the lateral eyes of the Corycxide are 
laterally rotated portions of the median eyes. In this connec- 
tion it may be of interest to quote a portion of Dr. Giesbeck’s 


1 Rinige Bemerkungen ueber die Augen der Salpen, Zool. Anz., XV., 
Jahrg. 349. 

2 Ueber die feineren Bau des Medianauges der Crustaceen. Anzeiger Akad. 
Wien. 1891, pp. 124-127. Ref. in Zool. Jahrsbericht fiir 1891, Arthropoda, 29. 


TURNER, WVervous System of Cypris. 35 


resume of Dr. Claus’ paper: ‘‘ Die Sietenaugen der Coryceiden 
sind abgedruckte Theile des Medienauges, diejenigen der Pon- 
telliden entsprechen dagegen dem zusammengesetzten Arthro- 
podenauge, wahrend ihr ventrales Auge ein dreitheliges Me- 
dienauge dirfte ebenso wie die beiden frontalen Sinnesorgane 
der ersten anlage nach auf Zellengruppen der Scheitelplatte, 
vor deraus wir, . . die obern Schlundganglien der Glieder- 
thiere abzuleiten haben, zu beziehen sein; die drei Augentheile 
sind vielleicht mit den drei Punktaugen an der Scheitenplatte 
von Annelidenlarven phylogenetisch in Beziehung zu bringen ; 
die Lage ihrer Pigmentzellen und ihre inverse Form werden auf 
eine convergent nach einem Punkte gerichtete Drehung zuriick- 
gefuhrt, welche mit dem Herabriicken des Organs in die Tiefe 
verbunden war; die Secret und Cornealinsen werden von Hypo- 
dermiszellen abgeschieden, ahnlich wie Krystallkegel und Cor- 
neafacette des zusammengesetzen auges; das auge besitzt eine 
mesodermal Hille, die sich in das neurilemm des Opticus fort- 
setzt ; Die function des Medienauges war ursprunlich das Thier 
beztglich der Richtung der Lichtquelle zu orientiren; bei 
compliciterem Bau, schon bei den Calaniden, hat es wahrschein- 
lich auch die Fahigkeit einer beschrankten Bildperception. Am 
Medianauge der Malakostraken-larven ist der ventral Becher 
bisher incht beobdachtet, aber wohl auch vorhanden.” 

A comparative study of the eyes of the Ostracoda lends 
support to the view held by Dr. Claus. In Zyphlocypris there 
is practically no eye, in Candona the triune eye is small and in- 
conspicuous, in Cyp7zs it is somewhat larger, in Cyprza and Cyclo- 
cyprts it is large and conspicuous, while in Votodromas the paired 
portions of the compound eyes are so far separated that they look 
like a pair of simple eyes united toa median eye by means of a 
stalk [fig. 30]. 

Pear-shaped Sense Organs.—I\n the cephalic portion of the 
body, a short distance above (dorsad of ) the upper lip, there is 
a pair of peculiar pear-shaped sense organs [fig. 16, P]. To 
the best of my knowledge this organ has been overlooked by 
all other writers on the histology of Cyprzs. When viewed 


36 JOURNAL OF COMPARATIVE NEUROLOGY. 


from the side, this is a pear-shaped organ, with its base directed 
outwards (ectad) and its apex inwards (entad). 

In hematoxylin preparations, in the base of this pear- 
shaped organ, there is a densely stained circular disc. In Cy- 
prinotus tncongruens (Ramdohr) this disc is 12.92 micromillime- 
ters in diameter [fig. 34]. 

The apex of this organ is surrounded by a ganglion of 
nerve cells [fig. 34]. 

The length of this organ varies in different species. Ina 
general way, the larger the species the larger this sense organ. 
In Cyprinotus tncongruens Ramdohr, which is about 1.35 milli- 
meters in length, this organ is 35.53 micromillimeters long and 
32.3 micromillimeters wide ; while in Cyprzs herricki Turner, 
which is about three millimeters in length, this organ is 71.06 
micromillimeters long and 48.45 micromillimeters wide. In 
other words, in Cypris herricki, which is about twice as long as 
Cyprinotus incongruens, the pear-shaped sense organ is about 
twice as long as the corresponding organ of Cyprdnotus tn- 
conguens. 

This organ is situated a short distance below the base of 
the antenna and close to the outer wall of the body. The broad 
base lies in contact with the body wall [fig. 16, P]. 

Since this organ is much larger in Cypris herricki Turner 
than in any other form known to me, the following histological 
study of the organ is based upon sections of that species. 

From a histological standpoint, this pear-shaped organ is 
composed of a neuroglia-like matrix, in which are two transverse 
rows of nuclei. One of these rows of large nuclei is located 
about half way between the base and the apex, while the other 
row is situated near the apex. Near the base of this organ the 
matrix is denser than it is elsewhere [fig. 31]. The organ is 
surrounded by a nucleated epithelium. 

This organ is innervated by a branch of the labial nerve 
[fig. 16]. 

There is a pair of these organs, but they are so closely ap- 
proximated that a hasty glance might not reveal both members 
of the pair. 


TuRNER, (Vervous System of Cypris. 37 


As to the function of this organ, I have no definite proof; 
but I am inclined to think that it functions as an eye. In 
Cypridina, besides the usual compound eye, Dr. Claus has 
described a median, unpaired simple eye. Now I am inclined to 
think that this pear-shaped organ of Cyfrzs is homologous with 
the unpaired simple eye of Cypridina. At first blush, there 
are two facts that seem to militate against such an assumption 
Ist, the median eye described by Claus is unpaired, while this 
pear-shaped organ is distinctly paired; 2nd, the median eye 
described by Dr. Claus is near to the compound eye, while this 
pear-shaped organ is far removed from the compound eye. In 
spite of these opposing facts, there are several weighty consid- 
erations which lead me to hold to the statements made above. 
Ist, Cypridina is much less compact than Cypris; 2nd, in 
Cypridina the compound eye and the simple eye are situated 
near the centre of the shell; 3rd, in Cyprzdina the parts of the 
compound eye are widely separated, while in Cyprzs they are 
closely approximated. May it not be that in the ancestors of 
Cypris the compound and simple eyes lie near each other and 
near the middle of the shell, as it now is in Cyprzdina ? May it 
not be that when the compound eye migrated towards the 
dorsal surface, the simple eye migrated towards the ventral ? 
And when: we remember that in histology this pear-shaped 
organ resembles the invertebrate simple eye, we have, I think, 
sufficient grounds for calling this a simple eye. To be sure this 
pear-shaped organ is one of a pair while the simple eye of 
Cypridina is unpaired; but in this day, the phenomena of 
transformations due to fission and to fusion are too well known for 
this difference to merit even a passing consideration. The set- 
tlement of this question, however, must be left to embryology. 

Sense Organs of the Mouth.—In this same region of the 
body, there is another set of hitherto undescribed sense organs. 
From the caudal border of the upper lip, from the cephalic 
border of the lower lip, and from between the teeth of the 
mandible, arise numerous hairs [fig. 16]. These hairs are some- 
times plumose [fig. 35]. At the base of each of these sets 
of hairs there is a similar sense organ [fig. 16, B, C, D]. Each 


38 JOURNAL OF COMPARATIVE NEUROLOGY. 


of these organs is composed of oblong nucleated cells, which 
are arranged with their longest axes perpendicular to the 
hirsute surface [fig. 4]. The organs in the upper lip are 
innervated by a branch of the labial nerve, those in the man- 
dible by a branch of the mandibular nerve and those in the 
lower lip, by the labral nerve [fig. 16]. Thus we have, sur- 
rounding the mouth, three pairs of similar sense organs. 

I have no definite proof as to the function of this set of 
organs; but, since they surround the mouth, they probably 
function as food discriminators. Whether this discriminating 
sense is one of touch or taste I am not prepared to state. 

Auditory Organ.—After describing rather minutely what 
_ he considers to be a sense organ located at the tip of the second 
foot, Dr. Rehberg expresses his belief that that organ is an ear. 
Now I agree with Dr. Rehberg in believing that there is a sense 
organ located in the tip of the second foot; but I think that its 
function is not auditory but tactile. Ido not think that the 
structure of the tip of the second foot warrants Dr. Rehberg’s 
assumption. And beside, at the base each of the antennules I 
have found what I consider the auditory organ of Cyprzs [fig. 16, 
£]. This is an ellipsoidal body in the centre of which there is 
a sac. The space between the outer wall of the ellipsoid and 
the sac 1s occupied by a single layer of columnar cells. The 
nuclei of these cells are located near the periphery. These cells 
are best seen in transverse sections of the organ [fig. 33]. 
Often the cells contain large vacuoles [fig. 29]. Within the sac 
there is a small spherical body which I take to be an otolith 
[fig. 29]. Since in most of my preparations this sac has been 
filled with a deposit of the stain used [fig. 33] I am not sure 
that this spherical body that I find in other preparations is an 
otolith or merely a bit of stain. 

As to the innervation of this organ, I have no definite 
knowledge. However, the antennary nerve is so intimately 
associated with this organ [fig. 29a], that I am inclined to be- 
lieve that that nerve innervates the auditory organ. 

There are two patent facts that lead me to hold that this is 
an auditory organ: Ist, in histology it resembles an otocyst ; 


TuRNER, Wervous System of Cypris. 39 


2nd, in location it resembles the auditory organ of the higher 
crustacea (/alacostraca). 

Olfactory Organs.—On the third joint of the antenna of all 
the fresh-water Ostracoda, there exists a peculiar seta which all 
writers have called an olfactory organ. As has been remarked 
by other writers on this subject, this organ consists of the fol- 
lowing three segments: basal piece, middle piece, end piece 
[fig. 15]. In some cases the basal piece has been subdivided 
into two pieces [fig. 17], while in others no middle piece can be 
distinguished [fig. 14]. The nerve enters this organ through a 
perforation in its base. Thence it extends along the axis of this 
portion until it reaches the base of the middle-piece, where it 
terminates in a small knob [fig. 15, 17]. In Cypres herricki 
Turner this terminal knob is about three micromillimeters in 
diameter. 

The usual width of this organ is about three micro-milli- 
meters. However, in Cyprza exculpta Fischer it is only about 
two micromillimeters wide, while in Cyprzs herrickt Turner it is 
about eight eight micromillimeters wide. In length the organ 
varies from about 85 micromillimeters in Cypris herricki 
Turner to about 29 micromillimeters in Cypria ineguivalva 
Turner. In the same genera the length of the organ varies 
directly with the length of the body of the specimen—the 
longer the body the longer the organ. In those genera in 
which the swiming setae of the antenna are greatly developed 
(Cyclocypris, Cypria) this olfactory organ is relatively larger 
than in those genera in which the natatory sete of the antenna 
are more feebly developed (Cyprzs, Candona). 

In the same species the tip of this organ may be either 
blunt or pointed. Indeed, on one antenna of a specimen the 
apex of this organ may be blunt, while on the other the tip 
may be pointed. 

The shape of this organ varies in different species. A 
consultation of figures 12-15, 17-Ig and 21-22 will give a bet- 


ter idea of these various shapes than could any number of 
words. 


40 JouRNAL oF CoMPARATIVE NEUROLOGY. 


The following table has been compiled in order to facilitate 
a comparative study of this olfactory organ: 


P) 8) SES 5 

4 : . 2lea|else| Ss 

Table showing the relative size of Leydig’s = = é fas BS 
organ ina few Cypride. = as ae) Po 

6) 3)? 23]: » 

TPL: Belts 

NAME OF SPECIMEN. MICROMILLIMETERS 

Candona acuminata (Fischer) Q ------------ 3 | 10] 20 | 43 |1250 
Candona crogmani Turner 9, type ---------/ 19 6'| 23 | 48 |1520 

BC ae Fe exception_.-.-. 16| —]| 19} 35 
ee es i 6 ------------ 13| 6}| 26] 45 |1520 
Candona delawarensis Turner, Q ---------- 10} 3] 23] 36] 950 
GC ss + $ ----------- 14| 5] 20] 39 

Candona fabeformis (Fischer) Q ------------ 12] 6)| 19 | 37 {1030 
Cyclocypris levis (O. F. Muller) Sasa ee aa 29] 6] 32|67]| 570 
Cypria exculpta (Fischer) ------------------- 23! 3116] 42] 640 
Cypria ineguivalva Turner, 9 ---------- ---- 13/10] 6] 29] 520 
Cypria ophthalmica (Jurine) ----------------- Io | 10| 16] 36] 580 
Cypridopsis vidua (O. F. Miiller)------------ 19 | —]| 16] 35] 700 
Cyprinotus burlingtonensis, Turner, 9 --.---- 26 | 10 | 29 | 65 |1600 
Cyprinotus crena, Turner, 9 ---~----------- 19| 8| 23] 40/1230 
Cyprinotus incongruens (Ramdohr) ---------- 39| 6116] 61/1350 
Cypris fuscata (Fischer) ------ ues ples 2 eee 36| 7 | 20] 63 |1500 
Cypress therrickt, Wunders? eae a ee 2| 4149 | 85 |3000 


Sensory Sete.—In addition to tne sense organs already de- 
scribed there are several types of sensory sete that deserve at 
least a passing notice. Not having been able to trace a nerve 
into any of these sete, I have given but a brief notice of them. 

At the tip of the antennules and antennae of all the 
Cypride there occurs a two-segmented sensory seta [fig. 20]. 
This seta is longest in Zaphlocypris, where it is as long as the 
terminal claw. In Condona fabeformis Fischer this seta is 48 
micromillimeters long. In this case, the terminal segment is 
ten micromillimeters long. | Owing to its small size and to the 
number and size of the associated claws and filaments, this 
organ is a very difficult object to study. 

At the extremity of the fourth segment of the antenna of 
Notodromas there is a peculiar seta with a funnel-shaped tip 
[fig. 28]. 

At the tip of the fourth segment of the antenna of the male 
members of the genera Candona, Candonopsts and Cypris there 
is a peculiar two-jointed seta [fig. 24-27]. Dr. Wenzel Vavra 


TurRNER, Wervous System of Cypris. 4! 


considers this a rudimentary organ. Its morphology varies in 
different species. By Dr. Wenzel Vavra the variations that we 
meet with in this organ are considered to be of taxonomic value. 

On the tip of the third and fourth segments of the mandi- 
bular palp of certain Cyprzd@, there occurs a short dagger- 
shaped setose seta [fig. 23 D.S.]. This seta was first described 


) 2 ohis 'setanOCcurs ime: 


in the ‘‘ Entomostraca of Minnesota. 
Candona crogmant Turner, Candona delawarensis Turner, Cyprt- 
dopsis vidua O. F. Miller, Cyprzs herrickt Turner, Cyprinotus 


burling tonensts Turner. 
Recapitulation. 


1. The nervous system of Cyprzs consists of a supra- 
oesophageal ganglion, which is connected to a ventral chain by 
a pharyngeal collar. 

2. The ventral chain consists of an infra-oesophageal and 
two subsequent ganglia. The infra-oesophageal ganglion has 
probably been compounded out of at least three pairs of ganglia. 

3. From the supra-oesophageal ganglion arise one un- 
paired and two paired nerves. These nerves are: the optic, 
the antennulary, the antennary. The antennary nerve receives 
a portion of its fibres from a ganglion that lies within the phar- 
yngeal collar. The nerve itself arises from the point where 
the collar joins the supra-oesophageal ganglion. 

4. From the infra-oesophageal ganglion arise five pairs of 
nerves. Four of these pairs arise from the ventral portion of 
the ganglion, while the other arises from the dorsal. The four 
that arise from the ventral half are: the labial, the mandibular, 
the labral, and the first maxillary. The one that arises from the 
dorsal portion is the thoracic nerve. 

5. The labial nerve passes forward into the pharyngeal 
collar before leaving the central nerve chain. 

6. The thoracic nerve innervates the shell muscles. 


1C, L. Herrick and C. H. Turner. Synopsis of the Entomostraca of 
Minnesota with descriptions of related species comprising all known forms from 
the United States included in the orders Copepoda, Cladocera, Ostracoda. St. 
Paul, Minn., 1895. 


42 JOURNAL OF COMPARATIVE NEUROLOGY. 


7. The 2nd maxillary nerve arises from the first ganglion 
back of the infra-oesophageal ganglion. 

8. From the last thoracic ganglion arise one unpaired and 
two paired nerves. The paired nerves are the leg nerves; the 
unpaired nerve is the abdominal nerve. The abdominal nerve 
innervates the genital apparatus. 

g. There is a median compound triune eye. The eye is 
situated near the dorsal surface. Each of the three divisions 
of this eye is supplied with retinal cells and alens. The nerve 
enters the outer ends of the retinal cells. In most of the fresh- 
water Cyprid@ the component parts of this eye are closely ap- 
proximated. In Wotodromas, on the contrary, the three parts 
are widely separated. 

10. In most of the fresh-water Cyprzd@ the optic nerve 
is a median unpaired structure, which splits into three branches. 
In Notodromas, however, there are three distinct optic nerves. 

11. Inthe front part of the body, between the base of 
the antennae and the upper lip, there is a pair of pear-shaped 
sensory organs. These are probably simple eyes. This organ 
is innervated by a branch of the labial nerve. 

12. At the base of the antennule there is an auditory 
organ. 

13. Bordering the mouth there are three pairs of similar 
sense organs. These organs lie in the upper lip, at the base of 
the mandibular teeth, and in the lower lip. They are innervated 
by branches of the following nerves: labial, mandibular, and 
labral. 

14. On the third segment of the second antenna there is 
the so-called olfactory organ. This organ consists of three seg- 
ments; the basal, the middle, and the end piece. The nerve 
enters the basal portion and extends to the base of the middle 
piece, where it terminates in a knob-like swelling. 

15. In addition to the above sense organs there are sev- 
eral types of sensory setae. 


TURNER, Vervous System of Cypris. 43 


EXPLANATION OF PLATES. 


Excepting where otherwise stated, the plates were all made from camera 
drawings made by the writer. 


PEATE TIT: 


Fig. 1. Cypris herricki Turner, transverse section of body in the region 
of the posterior portion of the mid-gut. A, endoskeleton; n, nerve chain. 

fig. 2. Do., do., transverse section of median eye. 

Fig. 3. Do., do., longitudinal section of circum-oesophageal commissure. 
a’, antennary nerve; md, mandibular nerve. 

fig. 4. Do., do., transverse section of the sense organ of lower lip. 

fig. 5. Transverse section of the endoskeleton. 

fig. 6. Do., do., transverse section of the body in the region of the lat- 
era] diverticles of the mid-gut. 


PLATE IV. 


Fig. 7. Cypris herricki Turner, longitudinal section through the sub- 
oesophageal ganglion. Mx’, mandibular nerve; mx’, Ist maxillary nerve. 

fig. 8. Do., do., longitudinal section through the brain at the level of the 
origin of the circum-oesophageal commissures, A’, antennary nerve. 

fig. 9. Do., do., median longitudinal section of the entire body. S.g., 
brain; A, harp-shaped sense organ of the lower lip; B, ventral nerve chain; 
C, cells in wall of oesophagus. 

fig.1o. Do., do., transverse section through the brain, in the region of 
the optic and antennulary nerves. 

Fig. 11. Do., do., longitudinal section of a portion of the ventral chain. 


PLATE V. 


Fig. 12. Candona delawarensts Turner, olfactory seta from the third joint 
of the antenna. 

Fig. 13.  Candona facbformis (Fischer), do., do. 

Fig. 14. Cypridopsis vidua (O. F. Miiller), do., do. 

Fig. 15. Cypris herrickt, Turner, do., do. 

fig. 16. Cypris herricki Turner, longitudinal section through the body. 
A.N., antennulary nerve; B, harp-shaped sense organ of the upper lip; C, harp- 
shaped sense organ at the base of the mandibular teeth; D, harp-shaped sense 
organ in the lower lip; E, auditory organ; L.N., labial nerve; P, pear-shaped 
sense organ of the upper lip; S.G., supra-oesophageal ganglion. 

Fig. 17.  Cypris fuscata Jurine, olfactory seta from the third joint of the 
antenna. 

fig. 18. Cyclocypris laevis (0. F. Miiller), do. 

fig. 19. Cypria exculpta (Fischer), do. , 

Fig. 20. Candona fabeformis (Fischer), sensory seta from tip of first 
antenna. 

Fig. 21. Cypria ophthalmica (Jurine), sensory seta from third joint of 
antenna. 

fig. 22. Cyclocypris laevis (O. F. Miiller), do., do. 


44 JoURNAL OF COMPARATIVE NEUROLOGY. 


PLATE VI. 

Fig. 23. Cypris herricki Turner, mandibular palp. D.S., dagger-shaped 
seta. 

Fig. 24. Candonopsis pubescens, sensory seta from the four segments of the 
antenna of the male. 

Fig. 25. Condonopsis kingslet, do. 

Fig. 26. Candona fabeformis, do. 

Fig. 27. Candona candida, do. 

Fig. 28. Notodromas monacha, Sensory seta from the fifth segment of the 
antenna. 

Figures 24-28 have been copied from Wenzel Vavra. 


PLATE VII. 


Fig. 29. Cypris herricki Turner, auditory organ from a longitudinal per- 
pendicular section. A. antennulary nerve; m, muscle. 

fig. 30. Natodromas monacha, eyes (after Vavra). 

Fig. 31. Cypris herricki Turner, pear-shaped organ of the upper lip, sec- 
tional view. ; 

Fig. 32. Cyprinotus incongruens (Ramdohr), ventral chain. A, antennary 
nerve; A N, abdominal nerve; L, labial nerve; L’, branch of same to pear— 
shaped organ; L’’, branch of same to harp-shaped organ; La, labrum. 

Fig. 33. Cypris herricki, Turner, auditory organ, from a horizontal-longi- 
tudinal section. 

Fig. 34. Cyprinotus incongruens (Ramdohr), pear-shaped sense organ of 
upper lip, surface view. 

Fig. 35. Cypris herricki Turner, tip of mandible showing hairs that be- 
long to the sensory organ found in the mandible. 

Fig. 36. Cypris pubera, eye, surface view (after Wenzel Vavra). 


PLATE VIII. 


Fig. 37. Cypris herricki, Turner, wall of mid-gut. 

Fig. 38. Cypris herrickt, Turner, tip of second foot. 

Fig. 39. LDo., do.,wall of oesophagus. 

Fig. 40. Cyprinotus incongruens (Ramdohr), transverse section through 
the mandibular nerve. S, sub-oesophageal ganglion ; md, mandibular nerve. 

Fig. 41. Cypris herricki Turner, surface view of wall of mid-gut. 

fig. 42. Diagram of the central nervous system of Cyfrzs. A, atennu- 
lary nerve; Ab, abdominal nerve; At, antennary nerve; L, labial nerve; L’, 
branch of labial nerve passing to the pear-shaped sense organ; L’’, branch of 
labial nerve passing to the harp-shaped sense organ; Lb, labral nerve; Lg’, Ist 
leg nerve; Lg’’, second leg nerve ; Md, mandibular nerve; Mx’, Ist maxillary 
nerve; Mx’, second maxillary nerve; T, thoracic nerve; T. V., thoracic 
ventricle. 

Fig. 43. Nerve cell of Cypris. 


PRELIMINARY NOTES ON THE CRANIAL NERVES 
OF CRYPTOBRANCHUS ALLEGHANIENSIS. 


J He MeGrEcor;, )B-Se;, 


University Fellow in Biology, Columbia University. 


The present communication is by no means a complete 
description of the cranial nerves of Cryptobranchus, but is merely 
a somewhat cursory sketch of the distribution of most of the 
more important rami. It was the hope of the writer to have 
completed before this time a study of the brain and cranial 
nerves of this amphibian, working out the nerve components 
from serial sections, but the difficulty in obtaining embryonic 
material for sectioning has so delayed the work that it has been 
thought best to present a brief preliminary account based almost 
entirely upon dissections of adult animals. 

Recent researches on cranial nerve components render it 
possible to determine, even by dissection, some points of im- 
portance in nerve homologies, though, of course, reconstruction 
from sections will be necessary to ascertain the more intimate re- 
lations of the components. 

In general, no attempt has been made in the present paper 
to trace the relations between nerve-roots and the ultimate rami. 
So far as possible the writer has endeavored to indicate homol- 
ogies with other amphibians, basing the comparisons mainly 
upon recent investigations on Rana, Amblystoma and 
Salamandra. 

Nervus olfactorius.—This nerve is somewhat peculiar in 
Cryptobranchus from the fact that it divides into numerous 
branches before leaving the cranial cavity. Within the cranium 
these branches extend cephalad in two large bundles on each» 
side, giving each olfactory nerve the appearance of being 
double. 


46 JOURNAL OF COMPARATIVE NEUROLOGY. 


Nervus opticus.—The optic is very similar to that of uro- 
deles in general. There is a slight vestige of a lumen at the 
base of the nerve. 

The nerves of the eye-muscles are all present but very 
small. The oculomotor is the largest of the three and maintains 
an independent course throughout its whole extent, leaving the 
cranium by a special foramen slightly caudad of the optic fora- 
men. In part of its course this nerve lies very close to ramus 
ophthalmicus profundus of the fifth. The courses of the sroch- 
learis and abducens have not been determined. They are both 
extremely minute, as would be supposed from the very degen- 
erate condition of the eye-muscles. The point of origin of the 
abducens appears to be somewhat variable. It has been figured 
by Osborn as leaving the medulla on a level with the fifth 
nerve, but transverse sections prepared by the writer show the 
abducens taking its exit on a level with the cephalic roots of the 
glossopharyngeo-vagus complex as it does in Rana, Diemctylus, 
Salamandra, Amblystoma and other forms. 

Nervus trigeminus.—The distribution of the fifth nerve in 
Cryptobranchus has been described by Fischer and more re- 
cently by H. H. Wilder, though neither of these writers has 
given a complete description. The first ramus, the ~ ophthal- 
micus profundus, after leaving the ganglion, traverses a canal in 
the pterygoid bone, coming to the surface some distance cepha- 
lad of the other two rami. It trends cephalad, passing mesad 
of the eye-ball and dorsad of the optic nerve and all the eye 
muscles except the superior oblique and superior rectus. “The 
greater part of the ramus passes cephalad to a point cephalad 
of the eye, where it breaks up into numerous branches anasto- 
mosing with branches of the superior maxillary ramus. The 
nerve thus formed enters a groove in the superior maxillary 
bone and sends branches to the skin of the upper lip through 
three or four foramina anda small notch in the lateral border of 
the anterior nares. This branch has been called by Wilder 
ramus nasalts externus. 

The two branches to the nasal capsule have been well de- 
scribed by Wilder who designates the ventral and dorsal 


McGreoor, Cranial Nerves of Cryptobranchus. 47 


branches respectively as vamus nasalis externus I and ramus 
nasalts externus II, The former, as shown by Wilder, receives 
a branch from the ramus maxillaris V. 

In addition to the branches already described there is a 
small branch entering the eye-ball which is probably to be re- 
garded as the ciliary nerve, and another small branch which is 
traceable to the region of the antorbital fissure in the nasal 
capsule. The latter branch is probably distributed to the glands 
of the nasal region, but a study of serial sections will be neces- 
sary to settle this point. 

The ramus maxillaris supertoris of the Trigeminus breaks 
up into several branches immediately upon leaving the cranium. 
It runs dorsad of the masseter muscle then dips ventrad of the 
eye supplying numerous branches to the skin of the upper lip 
and anatomosing with the ramus nasalis externus of the deep 
ophthalmic. A branch also runs along the floor of the nasal 
capsule as described in connection with the ramus ophthalmicus 
profundus. The ramus maxillaris superioris is accompanied for 
some distance by the ramus ophthalmicus superficialis of the 
facial nerve, and the two are described together by Herrick in 
Amblystoma as ramus fronto-maxillaris. Though the two nerves 
run close together for a part of their course, they are separate 
as far back as the ganglion Gasseri. It is almost certain that 
one of the branches which appears to belong to the superior 
maxillary ramus represents the ramus buccalis VII (in the 
sense in which this name is used by Strong and the writers on 
fishes). This view is supported by the presence of well-devel- 
oped lateral line organs between the eye and the angle of the 
mouth, the so-called infra-orbital sense organs. 

The ramus mandibularis V is larger than either of the 
other two rami. At its point of exit from the skull it gives a 
small branch to m. temporalis, then, trending laterad, it pierces 
m. masseter to which it contributes several twigs. At about 
the middle of its course through m. masseter a fairly stout 
branch separates from the main ramus and runs latero-caudad 
passing through a slight notch under the lateral border of the 
squamosal bone. This branch sends a number of twigs ceph- 


48 THE JOURNAL OF COMPARATIVE NEUROLOGY. 


alad, supplying the skin covering m. masseter and the region 
of the angle of the jaw; it is evidently a general cutaneous 
branch and may represent Strong’s vamus accessorius V. The 
greater part of samus mandibularis V enters the inferior maxil- 
lary by a large foramen about one-third the length of the bone 
from the proximal end. Just before its entrance into the man- 
dible it divides into two branches, the caudal of which passes 
directly through the mandible, appearing on the ventral border 
almost opposite the point of entrance. The foramen of exit 
is included between the splenial and angular bones. This por- 
tion of the nerve innervates m. mylohyoideus anterior, and is 
traceable almost to the symphysis menti, breaking up into sev- 
eral branches on the ventral surface of the muscle. C. J. Her- 
rick finds this branch to contain sensory fibres in Amblystoma, 
and the same is probably true in the case of Cryptobranchus. 
The branch in question is called vamus mentalis by von Plessen 
_and Rabinovicz, but, as Herrick states, the name is more prop- 
erly applied to the ramus we shall next describe. 

The ramus mentalis V is the cephalic one of the two 
branches which enter the mandible. It divides almost imme- 
diately into two branches which are separated by the Meckelian 
cartilage ; the ental branch is the dental nerve and supples the 
alveolar region; the other, a larger branch, lies ectad of the 
Meckelian cartilege and traverses a canal in the dentary bone. 
Shortly beyond its point of separation from the inner branch it 
is joined by the ramus alveolaris VII, which enters a foramen 
in the ectal surface of the dentary bone. The fibres of the two 
nerves mingle and pass cephalad through the canal above men- 
tioned, extending almost to the symphysis menti and giving off 
some half-dozen clusters of small branches which pierce the 
ectal surface of the dentary bone at intervals and supply the 
skin of that region. It is not certain as yet whether these fibres 
to the skin belong to the ramus mentalis V or to the ramus 
alveolaris VII. 

Nervus Facialis.—The seventh nerve of Cryptobranchus 
does not exhibit any very important variations from the con- 
dition generally found in urodeles. In correlation to the well- 


McGrecor, Cranial Nerves of Cryptobranchus. 49 


marked lateral line sense-organs, we find the rami of the dorsal 
VII root, or VIIb. of Strong, well developed. The fibres of 
the Rr. ophthalmicus super fictalis and buccalis are very probably, 
judging from analogy with the frog larva, contained in a large 
root (VII wand / of Osborn) which enters the caudal border of 
the Gasserian ganglion. These two rami leave the ganglion in 
company with ramus maxillaris superioris passing with it dorsad 
of m. temporalis. Ramus ophthalmicus superficiais VIT runs 
cephalad to the nostril, innervating the supra-orbital sense-organs 
of the lateral line system. This ramus seems to have been en- 
tirely overlooked by Fischer, and several recent writers have 
called it ramus frontalis V. Its community of origin with other 
lateral line rami, as shown by Strong, and its distribution to 
the region of the supra-orbital line of sense-organs leaves no 
doubt as to its true nature. Ramus buccalis VII accompanies 
ramus maxillaris superiorts V as far as the eye and ends in the 
infra-orbital sense-organs. This is not the homologue of the 
ramus buccalts as described by von Plessen and Rabinovicz and 
by C. J. Herrick, but the ramus maxillaris V of these writers 
no doubt includes the ramus buccalis as the name is here used. 
The root marked d in the figures of Salamandra given by von 
Plessen and Rabinovicz and of Amblystoma by Herrick is prob- 
ably homologous with the VII w and / of Crytobranchus as fig- 
ured by Osborn and probably contains the fibres of the two 
lateral line rami here described. 

Besides the two rami of the VII already described there 
are four others, all of which were described by Fischer, who 
called them Rr. palatinus, mentalis, alveolaris and jugularts. 
Ramus palatinus is rather small; it passes ventrad through a 
foramen in the prootic ossification and is covered ventrally for 
some distance by the very broad parasphenoid bone. Near the 
level of the posterior nares it forms anastomoses with Rr. oph- 
thalmicus profundus V and maxillaris superioris V. Onaccount 
of its anastomoses the ultimate distribution of the facialis fibres 
can be determined only by serial sections through the head. 
The vamus alveolaris of Fischer runs laterad along the dorsal 
surface of the squamosal bone, then cephalad between m. masse- 


50 THE JOURNAL OF COMPARATIVE NEUROLOGY. 


ter and the skin, and enters a foramen in the lateral aspect of 
the dentary bone cephalad of the area of insertion of m, masse- 
ter. Within the bone it unites with a branch of ramus men- 
talis V, and pursues its course almost to the symphysis menti, 
supplying branches to the lower lip, through some six or eight 
foramina in the dentary bone. As stated in connection with 
ramus mentalis V, it is not certain which component supplies 
the fibres to the skin of the lower lip. It appears to me not 
improbable that thisramus alveolaris VII may correspond to the 
ramus mandibularis tnternus VII described by Strong in the tad- 
pole, and may therefore represent the chorda tympant. The 
dentary being a dermal bone, the nerve may have secondarily 
become covered by it. I have not been able to find all the 
branches described by von Plessen and Rabinovicz and by Her- 
rick in Salamandra and Amblystoma respectively. The ramus 
in question must be represented by the alveolaris, accessory 
hyo-mandibularis or buccalis of Salamandra and Amblystoma, 
in the sense in which Herrick and v. Plessen and Rabinovicz use 
these names. Herrick thinks, perhaps with good reason, that 
the hyo-mandibular ramus of Amblystoma is the homologue of 
Strong’s chorda tympani or ramus mandibularis internus. If 
my supposition regarding the homologies is correct, the latter 
name is preferable to vamus alveolarts. Fischer's ramus mentalis 
is really the lateral line ramus to the lower jaw and therefore 
the branch which Strong designates as vamus mandibularis ex- 
ternus in the tadpole. In Cryptobranchus this branch does not 
accompany the hyo-mandibular as in the frog, but is a separate 
ramus as far back as the ganglion. It ccurses laterad along the 
dorsal face of the squamosal bone, passes ventrad of the man- 
dible and cephalad along the ventral surface of the mylohyoid 
muscles close to the mandible, dividing into two branches which 
supply the gular and oral lines of lateral line sense-organs. 

The fourth branch mentioned by Fischer as ramus jugularts, 
is the rsamus hyo-mandibularis of later writers. As in other 
amphibians it receives the vamus communicans IX ad VII. 
The ramus hyo-mandibularis in Cyptobranchus does not contain 
all the elements demonstrated by Strong in the nerve of that 


McGreoor, Cranial Nerves of Cryptobranchus. 51 


name in the frog, since, as indicated above, the ramus mandibu- 
larts externus and ramus mandibularis internus are separate rami 
as far back as the facialis ganglion. Ramus hyo-mandibularis 
pierces the cephalic division of m. depressor maxillae inferioris 
to which it gives several small twigs, then trends ventrad break- 
ing up into numerous small branches in m. mylohyoideus pos- 
terior. The ramus is thus in the main motor but a few sensory 
branches pass to the skin over m. depressor maxillae inferioris, 
and probably there are sensory branches to the skin under the 
jaw. Strong has shown that in the frog these sensory fibres are 
derived from the ramus communicans IX ad VII. 

Nervus acusticus. The auditory nerve of Cryptobranchus 
has the usual close relation to the facial. Its distribution is 
beautifully described and figured by Retzius in Vol. I of Das 
Gehororgan der Wirbelthiere. 

Nervus glossopharyngeus. The most cephalic branch of the 
glossopharyngeus is ramus communicans [IX ad VII. This ramus 
trends latero-cephalad and unites with the ramus hyo-mandibu- 
laris at about the point of entrance of the latter into m. de- 
pressor maxillae inferioris. The next division consists of sev- 
eral slender branches passing dorsad to the skin on the dorsal 
surface. Since this division innervates the region of the occip- 
ital group of lateral-line sense-organs there is a possibility that 
it represents the ramus supratemporals of Strong. 

The ramus immediately caudad of the preceding is a slen- 
der branch which passes ventrad to the mucous membrane of the 
roof of the pharynx. It can be traced cephalad as far as the 
post-nasal process, lying close to a blood-vessel between the 
pterygoid bone and the mucous membrane of the roof of the 
mouth. Herrick finds a somewhat similar ramus in Am- 
blystoma which anastomoses with the ramus palatinus VII. 
This ramus seems to be the ramus pharyngeus of Fischer. 

The vamus linguals lies furthest caudad and is much the 
largest of the glossopharyngeus branches. After passing latero- 
caudad over the hyo-suspensorial ligament it gives off a slender 
cutaneous branch, which anastomoses with a somewhat similar 
branch of the vagus, supplying the skin cephalad of the branch- 


52 JOURNAL OF COMPARATIVE NEUROLOGY. 


ial cleft. After giving off the cutaneous branch the ramus en- 
ters m. hyobranchialis to which it gives a motor branch, then, 
emerging from the muscle, it passes dorsad of the hypo-hyal 
cartilage and enters the tongue, near its cephalad border. Oc- 
casionally the r. lingualis merely lies on the ventral surface of 
m. hyo-branchialis, only the motor branch penetrating the mus- 
cle. Thus the ramus generally called lingualis is a mixed 
nerve. 

Nervus vagus. As my study of the vagus is not yet as 
complete as it is possible to make it even by gross dissection, I 
shall not attempt a description of all its rami. 

The most cephalic branch gives fibres to the skin cephalad 
of the branchial cleft and anastomoses with the branch of the 
glossopharyngeus which innervates the same region. There is 
also a small motor branch running cephalad to m. constrictor 
arcuum branchialium, and to the caudal part of m. genio-cerat- 
oideus. Another slender branch passes to m. levator arcuum 
branchialium and to the skin caudad of the branchial cleft. 

The largest ramus includes the ramus visceralis and a large 
recurrent branch. This latter branch is the one figured by 
Fischer as the hypoglossal nerve, and he was probably at least 
partially correct, as this nerve receives an anastomosing branch 
not only from the first but also from the second and third spi-. 
nal nerves. The greater part of the fibres of this nerve are, 
however, plainly derived from the vagus. and probably repre- 
sent a ramus recurrens vagt. It gives branches to m. constrictor 
pharyngeus, m. depressor arcus branchialis posterioris and m. dor- 
so-laryngeus, then runs cephalad and appears to end in m. genio- 
hyoideus. A branch to m. sterno-hyoideus is also derived 
from some part of this vago-hypoglossal complex. The most 
caudal ramus of the vagus is a lateral line branch. 

The first spinal nerve has no dorsal root, its origin is almost 
at the level of the foramen magnum, and it takes its exit by a 
foramen in the first vertebra. It gives several small branches 
to the m. longissimus dorsi of the neck region, but the largest 
ramusis that which anastomoses with the ramus recurrens of the 
vagus. There are also anastomosing branches which unite this 


McGreoor, Cranial Nerves of Cryptobranchus. 53 


branch of the first spinal with the second and third spinal 
nerves. The last two have dorsal roots, and the third takes 
part in forming the brachial plexus. Communication between 
vagus and hypoglossus has been discovered in larval Anura but 
it has not been described, so far as 1 am aware, in Urodela. 
Most of the material at my disposal was injured in the neck re- 
gion so that the relations of these parts could not be clearly 
ascertained. Many points regarding the intimate relations of 
vagus and hypoglossal nerves will require a study of serial 
sections for their elucidation. 


ON THREE POINTS: IN THE NERVOUS ANATOR 
OF AMPHIBIANS. 


J. S. Kunecstey. 
With three figures. 


In my studies of the head of Amphiuma, which I hope to 
have completed at an early date, I have found it necessary to 
make comparisons with the results of other observers, and in 
some cases to repeat their observations, especially in those cases 
where Amphiuma presented features not easily reconciled with 
the conditions existing in other foms. 

For this purpose I have had to go over the results ob- 
tained by von Plessen and Rabinowicz in their classic ‘‘ Die 


d 


Kopfnerven von Salamandra maculata.” Studying latvae of 
this species, varying between two and a half and three centi- 
metres in length, these authors figure and describe a commis- 
sure as existing between the ramus palatinus of the facial and 
their supra-maxillaris superior of the trigeminal, a condition 
which is not easily reconcilable with what I find in Amphiuma 
nor with what other authors find to exist in the Urodeles. To 
test the question as to whether Salamandra maculata was unique 
in this respect I sectioned a larva of twenty six millimeters 
and plotted the sections in the same manner as these authors 
did. As awhole I can confirm their results, but there are a 
few features which need correction. In their figures they give 
a small nerve, designated by the letter z, forming a ramus com- 
municans between the mandibularis and the maxillary (their 
supra-maxillaris superior) nerves. In my studies this ramus 
communicans also occurs, but, instead of its being very small it 
is fully as large as the branch from the ‘‘nebenganglien”’ with 
which it connects. Detailed study of many Urodeles shows 
that the maxillaris must be regarded as a compound nerve, 
with both facial and trigeminal components. See fig. 1. 


KincsLey, Nervous Anatomy of Amplibrans. 55 


Still further in front von Plessen and Rabinowicz describe 
a nerve uniting the maxillaris with the palatine. As sucha 
ramus communicans occurs in no other Urodele which I have 


Figure 1. Wateral reconstruction of the proximal portions of the fifth and 
seventh nerves of a Salamandra maculata embryo, body length 26mm. ag, 
accessory ganglion of VII; g/, ganglion of facialis; gg, Gasserian ganglion ; 
md, ramus mandibularis ; mx, ramus maxillaris; of, ophthalmicus profundus ; 
os, ophthalmicus superficialis; vos, root of the ophthalmicus profundus; 7Vv, 
root of the trigeminal; vi, root of the facialis; *, ramus communicans be- 
tween the mandibularis and maxillaris nerves. x 30. 


studied, this condition was also investigated. In all other Uro- 
deles the connection occurs between the opthalmicus profundus 
(the nasalis of von Plessen and Rabinowicz) and the palatine, a 
condition which is not found in their figures or description. In 
this I think they have made a mistake and that a part of their 
error has arisen in this manner. In Salamandra, near the an- 
terior limit of the eye the maxillaris nerve comes to lie close 
against the inferior oblique muscle at its insertion on the ball 
while the middle branch (their 0) of the ophthalmicus pro- 
fundus lies against the same muscle near its origin. It there- 
fore appears possible that they have mistaken the muscle fora 
part of the ramus communicans, a mistake the more readily 
made since in many forms there is such connection between the 
maxillaris and the palatine. The true relationship may be seen 
from fig. 2, which shows the middle division of the ophthal- 
micus profundus connected with the palatine by a short 
transverse commissure, just as is the case in Amblystoma, 
(my studies confirming the results of Herrick in this respect) 
and in Amphiuma and other forms. In Salamandra maculata, 
in front of this ramus communicans, the middle branch of the 
opthalmicus profundus suddenly bends downwards and goes to 


7 


56 JOURNAL OF COMPARATIVE NEUROLOGY. 


the alveolar region of the upper jaw, while the palatine is con- 
tinued forward as described by these authors, passing just in- 
side the internal nares, and forward and beneath the premax- 
illary bone. 


Figure 2, Projection of fifth and seventh nerves of Salamandra maculata, 
viewed from above; letters as before excepting of”, of°, middle and outer divis- 
ions of the ophthalmicus profundus; {, ramus communicans between palatine 
and profundus nerves. x 30. 


In the Journal of Morphology Vol. XI, No. 2, Mr. Alvin 
Davison gives an account of some features in the anatomy of 
the young and of the adult Amphiuma. It is unnecessary to 
notice here the whole article but there is a point of nervous anat- 
omy mentioned which may fittingly be commented upon ina 
Journal devoted to neurological matters. Mr. Davison says 
(p. 401): ‘‘The most important and interesting structure is 
found below and external to the eye in my smallest specimen, 
seventy eight millimeters in length. There appears in this re- 


KinGsLey, Nervous Anatomy of Amphibians. 57 


gion a canal, one tenth of a millimeter long, which is walled by 
columnar epithelial cells extremely regular in outline. Exter- 
nal to the epithelial wall there is seen a thick layer inferiorly of 
degenerated tissue, which is bounded by a thin layer of fibrous 
connective tissue. In three other specimens, eighty eight, 
ninety and ninety two millimeters respectively, no trace of this 
degenerate canal could be discovered, and in the smallest spec- 
imen I was able to detect it on the right side only.”’ 

In all probability it is this structure to which he refers later 
in his paper (p. 405). After briefly summarising Wiedersheim’s 
observations on the tentacular apparatus of the Gymnophiona 
he says: ‘‘As I have already shown, there exists in my young- 
est specimen of Amphiuma the atrophied remnants of the ten- 
tacular apparatus. The columnar epithelial lining of the canal 
is very distinct in about one dozen transverse sections through 
the orbits. In some of the sections I have discovered what I 
believe to be the degenerated retractor muscle. This apparatus 
in Amphiuma has precisely the same relative location as in the 
Coecilians. For some unexplainable reason neither Hay nor 
Kingsley found this organ in the young embryo. . . The 
occurrence of this degenerated structure in the young Amphi- 
uma and its complete disappearance in the adult gives unmis- 
takable evidence of the relationship of the Coeciliidae and 
Amphiumide.’’ A single figure illustrates the point made. 

Were it true that Amphiuma possesses, either in the young 
or the adult, rudiments of a tentacular apparatus, the fact would 
prove of great value to those who would recognize in the Gym- 
nophiona only degenerate Amphiume. It is not, however, 
‘‘unexplainable”’ why neither Hay nor myself found such a 
structure for no tentacular apparatus exists in Amphiuma. 1 
have considerable material in the way of young Amphiumae— 
possibly forty specimens—embracing both younger and older 
material than that in the possession of Mr. Davison and includ- 
ing some from the same lot as the four specimens at his com- 
mand. In the first place the structures described and figured 
by him are not in the proper position for the tentacular appara- 
tus, as a little careful reading of the Sarasin’s monograph would 


58 JOURNAL OF COMPARATIVE NEUROLOGY. 


have shown him. Secondly had he carefully examined the eye 
muscles of his Amphiuma he would have found that all are 
present, while the Sarasins say that the retractor muscle of the 
tentacle is probably produced from the retractor bulbi. 


| f 0 
SOO oP 


“yn 


mx* 


Figure 3. Portion of section through the head of a larval Amphiuma ; let- 
ters as before excepting f, frontal bone; /, parietal bone; 2, ramus nasalis in- 
ternus of ophth. profundus ; osf, orbitosphenoid; ¢7, trabecula; fs, parasphenoid; 
9, optic nerve; m, maxillary bone; 4, blood vessel ; 6* and mx*, blood vessel and 
branch of maxillary nerve constituting the tentacular apparatus of Davison. 

On the other hand I find a canaland a solid structure lying 
immediately beneath it, in the very place where Mr. Davison’s 
tentacular apparatus appears, but I interpret the features differ- 
ently. The supposed degenerate muscle is a peripheral branch 
of the maxillary nerve, the tentacular canal is one of the branches 
of the sub-orbital blood vessels. The only objection to this is 
that our author describes the canal as lined by columnar 
epithelium, a condition possibly the result of contraction of the 
vessel and a consequent throwing of its intima into folds, or 
possibly he has mistaken the contained blood corpuscles for 
epithelial cells. The short course which he allows the canal is 
explained by the fact that distally it breaks up while more 
proximally it is compressed as it passes between two nerve 


twigs. 


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Pee COMPARATIVE ANATOMY VOR THE GINSULA 


Tracy Eart Crark, B.S. 
With Plates IX-XI1l, 


TABLE OF CONTENTS. 


Introductory Note, . F : : 4 ; ’ 5 59 
List of Specimens Examined, : : : : O 60 
Historical Sketch, : : : ; : : : a 62 
The Development of the Operculums, . : ° 63 
Early Development of the Insula, ‘ . 5 é 67 
The Adult Human Insula, ; : 7 . : 69 
Relation of the Insula to the Claustrum, ; ; 6 73 
The Insula in the Primates other than Man, ‘ 75 
The Insula in the Carnivora, : 5 ; ; : igh 
The Insula in the Proboscidia, : : : : 82 
The Insula in the Rodentia, : ‘ : c : 83 
The Insula in the Ungulata, : 4 : ; ‘ 83 
The Insula in the Cetacea, : : : : c go 
Summary, ; : 0 : 5 : D 5 : QI 
Synonymy, 3 5 : ; : : 5 . . 93 
Bibliography, : ° : A : . ; : 94 
Explanation and Description of Figures, é : : 97 


The importance attached to this region of the human brain 
and the wide divergence of opinion as to the existence of an 
homologous area in other mammals led the writer, at the sug- 
gestion of Professor Burt G. Wilder, to select it in 1890 as the 
subject for his baccalaureate thesis. The extensive neurologic 
literature and abundant material possessed by the Anatomical 
Department of the Cornell University soon convinced the writer 
of his inability to cover more than the macroscopic side of the 
subject and that upon a very few specimens. Later the kind 
offer of Professor Wilder to place at his disposal all of the ma- 


terial at hand led to the prosecution of the work somewhat 
further. 


60 JOURNAL OF COMPARATIVE NEUROLOGY. 


It is with a keen sense of gratitude that I now acknowl- 
edge my indebtedness to Professor Wilder whose advice has 
been frequently sought and who has always shown great interest 
in the work. I desire also to thank Professor Simon H. Gage 
and Instructor Pierre A. Fish for their interest and assistance, 
and Mrs. S. P. Gage for the loan of specimens. 

The following list, while incomplete, will give some idea 
of the extent and nature of the material upon which the writer 
has based his conclusions. The accession numbers are the serial 
numbers in the Accession Book of the Department of Verte- 
brate Zoology of the Cornell University. 


Aon 


No.) D3°* | Age Groups. 


SUB-CLASS I. PROTOTHERIA. 
ORDER I. MONOTREMATA. 
76|Male Ornithorhynchus anatinus, duck-bill. 


SUB-CLASS II. METATHERIA. 
ORDER II. MARSUPIALIA. 


Didelphys virginiana, common opossum. 
Several specimens. 


378 Macropus giganteus, giant kangaroo. 
Two specimens. 
oe me \ Hypsiprymnus moschatus, kangaroo-rat. 


SUB-CLASS IM. EUTHERTA: 
ORDER III. EDbDENTATA. 
No specimens. 
ORDER IV. SIRENIA. 
844| Male Adult Manatus americanus, American manatee. 
ORDER V. CETACEA. 
670| Male Adult Globtocephalus melas, pilot-whale, black-fish. Fig. 35. 


ORDER VI. UNGULATA. 
Camelus bactrianus, Bactrian camel. 


2177|Male 


2122|Male Camelus dromedarius dromedary. Fig. 22. 
776 Cartacus clama, fallow deer. 
g61|Male Cartacus virginiana, Virginia deer, red deer. Fig. 23. 


Ovis aries, domestic sheep. Figs. 32-34. 
Many specimens. 


3408 Female |Adult ] 
3409|Female Adult r Bos taurus, domestic cow. Figs. 26-30. 
3362| Female Juv. | 


Juv. 


2777 Lquus caballus, domestic horse. 

2840) 6 mo. “ec “6 66 ce 

2125 Maie Term ‘“ “ “ “ 

2095| Male Adult es Ge be es Fig. 25. 


2259|Male jJuv. Equus asinus, burro, Fig. 24. 


CLARK, 


Comparative Anatomy of the Insula. 61 


? 
ae Sex Age. Groups. 
ORDER VI. UnGuLatTa—Continued. 
Sus scrofa, domestic pig. 
Many specimens. 
2123 Tapirus malayanus, Malayan tapir. Fig. 31. 
ORDER VII. ToxoponmTia. 
Fossil. 
OrDER VIII. HyracorpeEa. 
No specimen. 
ORDER IX. RODENTIA. 
Arctomys monax, woodchuck, marmot. 
3075 Castor fiber, beaver. 
Lepus cuniculus, rabbit. 
Several specimens. 
Mus decumanus, brown rat. 
Mus musculus, common mouse. 
Scturus hudsonius, red squirrel. 
ORDER X. PROBOSCIDIA. 
2181|Female Elephas indicus, Indian elephant. Fig. 21. 
166 ORDER XI. CARNIVORA. 
3124 5-8 wk \ Canis familiar?s, domestic dog. Figs. 14-18. 
Many specimens. 
277|Male Lynx rufus, American wild-cat, lynx. 
157|Female |22 da. Felts leo, African lion. 
309|Female |18 yrs. felis concolor, puma, panther. Fig, Io. 
Felis pardalis, ocelot. 
felis domestica, domestic cat. Fig. 19. 
Many specimens. 
Flyaena striata, hyaena. 
2031|Male Procyon lotor, raccoon. Figs. 1, and 12. 
Several other specimens. 
Mephitis mephitica, skunk. Fig. 20. 
Several specimens. 
Putortus vison, mink. 
197 Putorius domestica, ferret. 
Phoca vitulina, seal. Fig. 9. 
a645,Male Juv. Ursus thibetianus, Thibet-bear. Fig. 13. 
ORDER XII. INSECTIVORA. 
3356 Condylura cristata, star-nose mole. 


265|Female |Juv. 


ages. 


Sorex platyrhinus, common shrew, shrew mouse. 


ORDER XIII. CHEIROPTERA. 


Vespertilio subulatus, common brown bat. 
Several specimens. 


ORDER XIV. PRIMATES. 
Anthropopithecus troglodytes, Chimpanzee. 
Cercocebus fuliginosus, 
Macacus. Fig. 8. 
Several specimens. 


Fig. 7. 


Numerous preparations of the human brain at various 
See Figs. 1-6. 


62 JOURNAL OF COMPARATIVE NEUROLOGY. 


HISTORICAL SKETCH. 


Vicq d’ Azyr (16, 1786) seems to have been the first to 
call attention to this area in the human brain. He refers 
(Pt. II, p. 26) to it as ‘‘ the convolutions situated between the 
Sylvian fissure and the corpus striatum” and again (Pt. II, 
p. 74) as ‘‘the convolutions which accompany the Sylvian fis- 
sure.”’ These references would seem to indicate that at that 
time all that was known was the fact of the existence of certain 
gyres in that region which, on account of their concealed posi- 
tion, had attracted but little attention; furthermore these are 
the only references made to this region in his volume of sev- 
eral hundred pages. 

Monro (42, 1788) pictured three gyres of the human in- 
sula seen after the removal of the ventral portion of the cere- 
brum, but he neither named nor described them. 

Hence it remained for Joh. Christ. Reil (1804-1806) to 
give the first description as follows: ‘‘ Die Insel hat eine 
langlichrunde Gestallt.”” Since that time the area has been 
known as the ‘‘insula”’ or ‘‘island of Reil,” although many 
other names have from time to time been applied to it (see Syn- 
onymy, p. 93.) , 

The insula seems to have attracted little attention during 
the next half century and it was not until about 1860 that the 
study of cerebral localization caused increased importance to be 
attached to it. Until several years later it was supposed that 
the insula governed, at least in part, the power of articulate 
speech. Authors had noted that in aphasics there was gen- 
erally a lesion of the@nsula and the left subfrontal gyre. They 
seem not, however, to have taken account of lesions confined 
strictly to either area and they assumed that the insula had been 
first affected and that later the lesion had extended into the ad- 
joining area. Broca ( 4 ) disproved this. By comparing the 
lesions found in postmortems upon a large number of aphasic 
patients he reached the conclusion as early as 1861 that the 
faculty of speech was located in the left subfrontal gyre, still 
often called ‘‘Broca’s convolution.”’ During the next seven 


CLARK, Comparative Anatomy of the Insula. 63 


years a large number of aphasics were examined for the pur- 
pose of verification, and in 1868 he published a résumé of the 
facts upon which he based his conclusion. This discovery nat- 
urally aroused great interest in the insula inasmuch as ‘‘ lesions 
of the left subfrontal gyre generally extend into the insula,” 
while those (2) ‘‘ confined strictly to the insula are very rare.” 
In 1868 Broca (4, 113) pointed out that in aphasics there is almost 
always a lesion in the caudal half of the subfrontal gyre of 
either the right or the left side, and that this lesion is sometimes 
confined exclusively to this area. However, in about one case in 
twenty this area is apparently perfectly sound, but a lesion ex- 
ists in the insula or an adjacent gyre with which the subfrontal 
is continuous. The lesion is located in about Ig out of 20 
times on the left side and the right side remains perfectly sound. 
He had never found a case in which the patient was possessed 
of this faculty and at the autopsy showed a lesion of both sub- 
frontal gyres. Speech has, however, persisted in some patients 
where an autopsy showed the total destruction of the right sub- 
frontal. He also said he had noticed several cases of idiots who 
had never been able to learn to read, where the subfrontal gyre 
was absent on both sides. 

So far as the writer has been able to learn, no satisfactory 
reason has ever been given for this strange association of the 
subfrontal region and the insula; yet if, as Meynert (gs) says, 
‘‘fibers fromthe subfrontal gyre pass just entad of the insular 
cortex,’ then it would seem very likely that a diseased condi- 
tion of those fibers sufficient to cause loss of speech would 
affect the overlying cortex of the insula producing lesions and 
perhaps a breaking down of the cellular cortex, as frequently 
happens. <A clot of blood, through pressure upon the insula, 
has also been said to cause loss of speech. 


THE DEVELOPMENT OF THE OPERCULUMS. 


In 1892, Cunningham (45, 78) said of the human brain: 
‘‘As Mihalkovics has pointed out, it is toward the end of the 
second month of development that the first signs of the Sylvian 
depression may be detected. At the same time, it is right to 


64 JOURNAL OF COMPARATIVE NEUROLOGY. 


state that its appearance is frequently delayed beyond this 
stage and I have observed hemispheres well on in the third 
‘month with hardly a trace of it.’”” What is later to become the 
Sylvian fissure is now only a broad shallow furrow deepest near 
the base of the brain and radiating in a fan-shape dorsad where 
it disappears upon the lateral surface. Gradually by the 
growth of what will later form the frontal, temporal and pari- 
etal lobes, an overlapping or folding occurs and the fossa grad- 
ually narrows until completely closed. What really happens 
is that by the more rapid enlargement of these lobes in an en- 
closed space, the intervening broad area- in the bottom of the 
fossa, which early becomes fixed in position, is finally covered 
and the lips of the Sylvian fissure are brought together. These 
overlapping portions are the so-called operculums and in the 
human brain are four in number, namely: the operculum arising 
from the frontal and parietal, the preoperculum and the suod- 
operculum from the frontal, and the fostoperculum from the 
temporal lobe. These overlapping portions do not differ struc- 
turally from the surrounding parts, being convoluted and cov- 
ered with cinerea on both the ectal and the ental surfaces, and 
the gyres of the ental surface are interdigitated with those of the 
insula. The dorso-caudal end of the Sylvian fissure is the part 
first formed and the first as well to become completely closed. 
This is due to the rapid growth of the operculum and postoper- 
culum. 

In 1890 Beer (4, 309 (ix) ) said: ‘‘ According to Ecker, 
Mihalkovics, etc., the Sylvian fissure of the adult is formed by 
the gradual narrowing of a flat broad depression on the lateral 
surface of the hemisphere, which is due to arrest of develop- 
ment of the zzsula Rel in the 3rd month of fetal life.” The 
conclusion that the insular area becomes depressed by a cessa- 
tion of growth, while favored by many neurologists, appears to 
me unwarranted. Presumably what happens as stated by 
Wilder is that there is probably no complete arrest of develop- 
ment, but that the surrounding parts develop much more rap- 
idly so that in a short time they project farther laterad, leaving 
the insula relatively depressed. While this seems to account 


CiaRK, Comparative Anatomy of the Insula. 65 


for all observed phenomena, the writer is inclined to the belief 
that one very important factor has been underestimated. Dur- 
ing the period when the insula first appears, the brain as a 
whole is enlarging very rapidly. This enlargement in the re- 
gion which will later become the Sylvian fissure it is assumed 
may be due more largely to the expansion of the paraccele 
than to the thickening of the paraccelian wall by the addition 
of true nervous substance, which of course is always interstitial. 
The anatomy of this region would seem to favor this latter 
conclusion. The intimate connection of the insula with the 
brain stem is such as to preclude any marked lateral extension 
and, as the wall of the paraccele is then but little more than 
membrane, any sudden increase of development, either by add- 
ition to the true nervous parietes or by the expansion of the 
paraccele, would cause the area caudad and dorsad to project 
laterad giving outline to the insula by the formation of the cir- 
cuminsular fissure and later the dorsal end of the Sylvian. It 
also suggests the possibility of finding in a fetus of the 
fourth to the seventh month an expansion of the paraccele 
corresponding in a general way to the outline of the rudimen- 
tary operculum. The conclusion given above agrees with ob- 
servations upon the development of this region as described by 
Cunningham, Ecker and others. It is also further believed that 
this effect of expansion does not stop here, but that the over- 
lapping of the insula and consequent formation of operculums 
may be due in part to the expansion of the paraccele, and that 
the subsequent thickening of the walls by the addition of true 
nervous substance combined with the effects of growth in an 
enclosed space may account in part for the continued conceal- 
ment of the insular area in the adults of Primates and some 
of the Carnivora. 

The exact time of the closing of the Sylvian fissure by the 
operculums is not known. In 1873, Ecker said it was from 
the ninth to the tenth month. 

In 1888 Mingazzini (41) said: ‘‘In one brain of the 
eighth month, I have found the fissure almost completely 
closed.”” The condition here presented I think is one of indi- 


66 JoURNAL OF COMPARATIVE NEUROLOGY. 


vidual difference rather than the rule, as I have examined sev- 
eral brains of this age and have not seen one closed or 
~ nearly so. 

Broca in 1888 (4, 545) said: ‘‘Thissuperficial position of the 
cephalic extremity of the insula has been observed, more or 
less, in man, upon a great number of new-born children or 
those of a few months old; after the second year it is rarely 
seen except upon the brains of idiots and imbeciles (and also in 
some deaf-mutes).”’ 

The determination of the exact time of closing of the Syl- 
vian fissure in the normal human brain is of considerable mor- 
phologic importance as it marks the completion of a period 
of remarkable growth of these lobes; and in making a collec- 
tion of brains between the eighth and twelfth months for this 
purpose as many as possible should be obtained so that individ- 
ual differences might not induce a wrong conclusion. 

The operculums do not always completely close the Sylvian 
fissure and hence the insula is exposed sometimes even in the 
adult human brain. In 1887 Rolleston (49, 35), in his descrip- 
tion of an adult male Australian brain, said: ‘‘ The island of 
Reil is exposed on the left side. . . The exposure of the 
island of Reil implies that the surrounding gyri are ill-devel- 
oped; Broca’s convolution is thus shown to be defective, a point 
of interest in an Australian savage whose language is primitive 
It is questionable if 
facts would bear out this conclusion, and while some of the gyres 


) 


as shown by its unclassified character.’ 


might in this specimen have been ‘‘ill-developed”’ it would 
have been interesting to learn if this had been observed on 
any other specimen. 

The brain of the philosopher Chauncey Wright has a part 
of the insula exposed. In 1889 Wilder (63,158) said: ‘‘ Whether 
any of this condition is due to pressure during hardening 
which may have occasioned also the peculiar roundness of the 
temporal lobe, it is impossible to determine and Dr. Dwight 
does not recall the condition of the parts when the brain 
was removed.” In others, as that of the Swedish carpenter 
(No. 318), the operculums not only conceal the insula but abso- 


CiaRK, Comparative Anatomy of the Insula. 67 


lutely overlap each other. These, withthe possible exception 
of the Australian, are all types of individual difference and their 
significance may be summed up thus: (1) if the insula is not 
wholly concealed, it shows that if the insula is normal, the sur- 
rounding parts have not reached their full development, while 
if the insula is abnormally large, the surrounding parts may or 
may not be fully developed; (2) if the insula is completely 
covered so that the operculums overlap each other, it would 
appear that, if the insula is normal, the surrounding gyri must 
be abnormally developed, while if the insula were unusually 
small, the normal operculums might even overlap each other. 


EARLY DEVELOPMENT OF THE HUMAN INSULA. 


We have seen that the rudiments of the Sylvian fissure 
become apparent toward the end of the second month of 
fetal life. There is then in the bottom of the shallow fossa 
an indefinitely-defined oval or somewhat triangular area which 
apparently develops more slowly than the surrounding lobes 
and which finally becomes a subarea. As development pro- 
ceeds, this area beeomes more elevated and hence more sharply 
defined by the formation of the circuminsular fissure. The re- 
gion thus mapped out is sometimes called the zzsular area be- 
cause it is the part which will eventually become the insula in 
man and presents a condition found in the adults of some 
animals. 

Mingazzini (49, 217, 1888) says: ‘‘ Until the end of the 
sixth month the insula is quite smooth; constantly there ap- 
pear in the seventh month those fissures [ circuminsular, 
Wilder; vzgoles, Broca] which circumscribe the future gyres 
of the insula. I can therefore not agree with Mikalkovics who 
asserts that the insula is smooth up to the beginning of the ninth 
month. I found this quite so once ina brain of the eighth 
month upon which only one small cortical fissure appeared. 
In the eighth month, they increase, oftentimes, to the number 
of four, among which is produced the caudal one which is larger 
and deeper than the others ; after the ninth month, there are, 


d 


as a rule, four fissures.’’ Mingazzini probably erred in the 


68 JOURNAL OF COMPARATIVE NEUROLOGY. 


above statement. The caudal one of the four fissures on the 
normal insula. is never, so far as the writer has been able to 
learn, ‘‘larger and deeper ’”’ than the transinsular fissure which 
is cephalad of it and the observations of Cunningham (44, 347- 
348) in 1891 seem to show pretty conclusively that even when 
three fissures are present on this area, the caudal one is usu- 
ally in the postinsula. 

In the spring of 1890 the writer became convinced, as the 
result of the study of the comparative anatomy of this region, 
that future investigation would show that, barring the circum- 
insular fissure, the transinsular, which is doubtless the most 
constant and best, defined of the fissures on this area, would be 
found to be the first to appear after the circuminsular and that 
its homologue exists in some other animals at least. The 
following table compiled from Cunningham, 1891 (14, 347-348) 
seems to prove that this really is the order of development in 
the human fetus. The brains enumerated below range from 
five to six and a half months. 


Transin- 

CEREBRUMS T 

State of Development of Insula. Total | sular f. 

Right | Left present. 
Iinsulasperfectly/smooth\ yee ee aaa 2 4 6 fo) 
oe wit hutransins ula tiss Ure see re 3 4 7 7 
a OC ‘Cfandsthe soneicep uplades=s==aas 6 2 8 8 
ae CG ‘* f. one cephalad and one f. caudad; 2 I 2) 3 
Totals): 2.2265 28eceo eee tha Sees 13 II 24 18 


This same author (14, 347) says: ‘‘ Furthermore, there is 
good reason to believe that in the development of the sulci and 
gyri the right insula is usually in advance of the left, and also 
that the process is greatly retarded in the female brain.”” The 
first part of the above statement of Cunningham, while appar- 
ently borne out by the table, seems remarkable inasmuch as 
the left insula in the normal adult as usually larger than the 
right. 

In 1892 Cunningham (15, 15 summary) said: ‘‘In the adult 
brain the insula is proportionately longer in the male than the 
female. At all periods of growth it would seem that the insula 
is relatively longer on the left side than on the right side. In 


Crark, Comparative Anatomy of the Insula. 69 


the negro brain, it would appear that the insula is relatively 
shorter than in the European brain.” 

The completion of the circuminsular fissure on the ventral 
side is usually not accomplished until about the time of birth. 
Prior to this completion of the circuminsular, which is prob- 
ably influenced to a considerable extent by the cephalic growth 
of the tip of the temporal lobe, the insula is continuous with 
the ventral surface of the cerebrum through the Sylvian fossa. 
It was probably this condition to which in 1885 Meynert (39, 4) 
called attention as follows: ‘‘the island of Reil is connected 
with a protuberance (the olfactory lobe) situated on the lower 
aspect of the frontal portion of the fore brain.” It will be 
seen (Figs. 1 and 2) that the lateral root of the olfactory tract 
extends into the Sylvian fossa (that. portion of the Sylvian fis- 
sure which does not close up). It is not believed, however, 
that the fibres of this tract extend over any portion of the in- 
sula, but on the contrary that they preserve approximately the 
same relation to the insula as in the sheep (Fig. 34), the skunk 
(Fig. 20), the cat (Fig. 19) and the mink, in each of which it is 
to be noted that the fibres do not pass dorsad of the bottom of 
the rhinal fissure, thus sharply demarcating the insula from the 
olfactory lobe. 


THE ADULT HUMAN INSULA. 


Usually the adult human insula is entirely concealed within 
the Sylvian fissure and can be seen only by divaricating the 
lips of the fissure or after cutting them entirely away. Broca 
(4, 651) says that it is not normally exposed after the second 
year. This occlusion is a condition found so far as known in 
all primates and in many of the imprimates. In all brains pos- 
‘sessing an insula, the cortex bends in from the lateral surface 
and passes without interruption under the fissures and over the 
surface of the insula completely covering it whether convoluted 
or not. In all brains observed by the writer, the gyres of the 
operculums are interdigitated with those of the insula. 

In 1889 Wilder (63, Prop. CXLI) wrote of this area in 
man, ‘‘the insula is a part of the cortex, which, at one period 


70 JOURNAL OF COMPARATIVE NEUROLOGY. 


wholly superficial, is gradually covered more or less completely 
by converging folds of the adjacent regions. The insula thus 
becomes a subgyre, while the operculum, preoperculum, sub- 
operculum and postoperculum are supergyres.”’ 

The adult human insula is always sharply demarcated by 
the surrounding circuminsular fissure (7zgoles, Broca). Its posi- 
tion is generally in a plane nearly parallel with the lateral sur- 
face as shown in Figs. 3 and 4. An exception is shown in Fig 5. 
In this specimen the gyres form a plane nearly perpendicular 
to the lateral surface. This is a condition found in the sheep, 
the pig (12,9), the calf, and some horses, and is believed to be 
due to some peculiarity in the development of the brain stem. 

In the adult, the insula is a large broad eminence, the sur- 
face of which is richly convoluted. It generally possesses from 
four to six gyres diverging cephalad, dorsad and caudad and 
producing an appearance not unlike that of a fan. The num- 
ber of gyres is subject to considerable variation even in the two 
insulas of the same brain. The brain of Guiteau, the assassin 
of Garfield, showed marked peculiarities in this respect. Ac- 
cording to Spitzka (55, 386) the left insula contained seven fis- 
sures and eight gyres, while the right had only five fissures and 
six gyres. It will be noticed that the right insula had its full 
complement of fissures and gyres while the left exceeded it by 
two. Another case markedly different from this one has also 
been reported by the same author. It, like the preceding, is 
that of a murderer. He says, the left insula had ‘‘ fewer and 
flatter gyri than that [six gyri] of the right side, and was 
not far different from that of an orang in my possession.”” In 
all normal brains, the left insula is said to be larger. 

The insula is usually quite deeply fissured and is always 
divided into two plainly unequal parts. This division is pro- 
duced by a deep fissure extending dorso-caudad from the ven- 
tral side—where it is deepest—to the dorsal margin where it 
connects again with the circuminsular. This forms a cephalic 
part, pretnsula, and a smaller caudal one, postensula. The pre- 
insula is not only larger but contains more gyres. All of the 
fissures of the insula communicate with the circuminsular dor- 


CLARK, Comparative Anatomy of the Insula. | 


sad but with the exception of the transinsular none of them 
reach the ventral margin. The area common to all the gyres 
of the preinsula, and from which they all appear to spring, is 
usually the part most elevated and is called the pole of the insula. 
Eberstaller (17, 741) says that the ‘‘limen insula ”’ (pole) is 
found only or in a certain form in the human brain. 

But for the unusually elevated pole in this specimen, Fig. 
5 might be said to represent a nearly typical insula. The pre- 
insula usually possesses three gyres of which the first (the one 
most cephalic) and the third are usually the best developed and 
these are frequently subdivided dorsad by short fissures. The 
postinsula seldom possesses more than two gyres the first of 
which is usually the larger. These gyres are united ventrad 
and are always longer than those of the preinsula. Eberstaller 
(17, 741) says the first gyre of the preinsula is the largest and 
strongest and that the second gyre is usually weakly developed. 
The postinsulas shown in Figs. 3 and 4 exhibit a peculiarity 
which is very striking when compared with the usually simple 
character of the area. The three small elevations of Fig. 3 
and the one in Fig. 4 dorsad belong to the postinsula. It is pos- 
sible that the shallow furrows separating these eminences may 
have been caused by arterial pressure. Fg. 3 also shows 
the relation of the insula to the medicornu. The specimen shown 
in Fig. 5 is remarkable for the unusual elevation of the pole. 
As previously stated this condition is probably the result of some 
peculiarity in the development of the brain stem. 

In 1887 Waldschmidt (61, 375) said the insula was larger in 
educated persons. There are many facts favorable to this view 
but it lacks confirmation. 

Eberstaller and Cunningham have each called attention to 
the apparently intimate relation of the transinsular and central 
fissures. These fissures are very constant in the human brain, 
they lie in the same plane, they take the same general direction 
and they appear in the fetus at about the same time (5-6 mo.). 
Eberstaller (17, 749, 1887) says: ‘‘ The cephalic insula is con- 
nected wholly with the frontal lobe, the caudal insula with that 
portion just caudad of the central fissure.” 


72 JOURNAL OF COMPARATIVE NEUROLOGY. 


In 1891 Cunningham (14, 338-9) said, that Guldberg had 
suggested for the transinsular fissure ‘‘the very appropriate 
-name of sulcus centralis insule which indicates net only its 
central position in the island of Reil, but also its relation to the 
central fissure on the outer surface of the hemisphere mantle.” 
This author then endeavors to establish a connection between 
gyres and fissures on either side of the transinsular with those 
on the same side of the central fissure. He further states that 
‘it is true that we cannot regard these corresponding convolu- ~ 
tions and sulci as being directly continuous with each other but 
still in many cases something which approaches very nearly to 
continuity occurs. Thus it is well known that the inferior pre- 
central sulcus, the fissures of Rolando and the intraparietal sul- 
cus are not infrequently carried downwards, so as to cut into 
the fronto-parietal operculum and open into the Sylvian fissure.”’ 
There are many facts which seem to show that Guldberg and 
Cunningham erred in their conception of the relation of these 
two areas : : 

1. The variations in the ventral extension of the central 
fissure are believed to be due to individual difference. 

2. The connection of the central fissure with the Sylvian, 
when it occurs, is almost always by a small, variable opercular 
fissure (Cunningham 14, 341-2). Wilder, in 1894, in the report on © 
the brain of an educated suicide, having apparently duplicate 
central fissures, says (65, 2): ‘‘On the left both centrals enter 
the Sylvian, with a depth of at least 5 mm. On the right, the 
first central approaches the precentral, the second joins the Syl- 
vian at a depth of about 3 mm.” So far as the writer has been 
able to learn, this connecting fissure never completely divides 
the operculum nor interrupts the circuminsular fissure in an 
otherwise normal brain. 

3. The absence of the central fissure has not been shown 
to produce an appreciable effect upon the transinsular and if 
either is absent it is more often the central fissure. 

4. The insula is always sharply defined and is believed to 
be a distinct lobe as sharply demarcated from the surrounding 
areas as any lobe of the cerebrum, and the fissures and 


Ciark, Comparative Anatomy of the Insula. 73 


gyres upon its surface are also distinct and not dependent upon 
those of the rest of the cerebrum. 

5. The central fissure is not known to exist outside of the 
primate brain, while, as will be shown, the transinsular fissure 
is present in many imprimates and the relation of the transinsu- 
lar to the Sylvian in them is sometimes as in man (see raccoon); 
but more often it coincides in position and direction with the 
Sylvian fissure. 

6. The transinsular fissure gradually becomes less deep as 
it approaches the circuminsular fissure dorsad, and it has never 
been shown to extend beyond this fissure, while in some im- 
primates it never reaches the dorsal margin of the circuminsular 
fissure. 

7. The central fissure is sometimes double; the writer 
has been unable to find a single recorded instance of the dupli- 
cation of the transinsular fissure. 

8. The reasons given by Cunningham (13, 287) to show 
that because the preoperculum is (as he thinks) absent in the an- 
thropoid it necessarily follows that a portion of the insula is 
also absent, are not well founded. The human insula has near- 
ly or quite the same form before as after the appearance of the 
preoperculum and the writer fails to see why it should be oth- 
erwise in the ape. 

g. The mononym ‘‘transinsular” fissure (Wilder) indi- 
cates better the relation of this fissure to the insula and is far 
preferable to the polyonym ‘‘sulcus centralis Reilii’? of Guld- 
berg and Cunningham. 

The blood-supply of the insula is derived from the branch- 
es of the medicerebral artery which ramify over its surface. 
The main branch lies in the transinsular fissure as shown in 
Fig. 4. From these branches small arteries penetrate the cor- 
tex directly. 


THE RELATION OF THE INSULA TO THE CLAUSTRUM. 


On account of its intimate connection with the brain-stem, 
the insula has been called the stem-lobe or Stammlappen. En- 
tad of the insular cortex (see Fig. 6) but separated from it by 


74 JOURNAL OF COMPARATIVE NEUROLOGY. 


a band of myelinic fibers, lies another cellular area called the 
claustrum (Burdach). These fibers, says Meynert, have their 
- origin partly from the subfrontal gyre and partly from the un- 
cus from which they radiate in a fan-shape separating the areas 
referred to above. The same author also says (37, 674): ‘‘The 
fibers of the fasciculus uncinatus traverse freely the substance of 
the claustrum as well as that of the zucleus amygdale. The an- 
terior cord-like and hook-shaped part of the fasceculus uncinatus is 
joined by bundles of fibers distributed, as it were, in layers 
which pass through and along the surface of the claustrum, con- 
stituting an important part of the medullary substance of the 
island (of Reil) and of the external capsule.”’ 

In 1889 Spitzka (56,172) said the claustrum ‘‘corres- 
ponds in extent nearly with the extent of the insula.” 

Obersteiner in 1890 (42, 68) said of the claustrum, ‘‘its 
lateral surface adapts itself to a certain extent to the cortex of 
the island of Reil, exhibiting similar small elevations and de- 
pressions.”’ 

Until recently, the claustrum has been assumed to be pecu- 
liar to the primate brain, and was an object of much specula- 
tion; but the researches cf Betz, Meynert, Duret, Browning and 
others seem now to prove beyond a doubt that it is really a part 
of the insula which became separated from the cortex proper 
by the growth of the fibers referred to above. In1879 Richet 
(48, 23) wrote: ‘‘It appears that in the brains of idiots, the 
white lamina, which separates the insular convolutions from the 
avant-mur is absent, and the avant-mur really becomes the in- 
ternal layer of the cortex cerebri (Betz).” The same author 
(48, 24) quoting from Duret, says the circulation is the same as 
that of the insula. Browning in 1889 (5, 234) agreed with Duret 
and stated that the blood supply of the claustrum passed 
through the cortex of the insula and not through the precrib- 
rum (anterior perforated space). This being established it would 
seem that the presence of a claustrum was indicative of the 
presence of an insula. It also suggests the possibility of these 
fibers passing so far entad as not to demarcate a claustrum as is 
probably the case in the sheep. The former inference regard- 


CLaRK, Comparative Anatomy of the Insula. 75 


ing the presence of a claustrum indicating the presence of an 
insula at first seemed plausible but later reflection has caused 
the idea to be abandoned (see discussion under Carnivora). 


THE INSULA IN THE PRIMATES OTHERK THAN MAN. 


The anthropoid apes, which in structure are so closely 
allied to man, possess a well developed insula. 

Leuret and Gratiolet seem to have been the first to recog- 
mize this area. They say (91, Il, 112):)* Le lobe central_parait 
particulier a homme et aux singes, peut-etre voit-on quelque 
chose d’analogue dans les makis, mais on ne voit rien de sem- 
blable chez les autres mammiféres.”’ 

In 1891 Cunningham (13, 287) said: ‘‘One of the most re- 
markable characters in the cerebrum of the orang and the chim- 
panzee is the fotal absence of the frontal and orbital opercula. 
The temporal and fronto-parietal opercula are alone present.” 
There is some diversity of opinion as to the entire ‘‘ absence ”’ 
of the suboperculum in these brains and Cunningham also admits 
the presence of a fissure which forms the cephalic boundary of 
the operculum but he does not believe it homologous with the 
presylvian fissure and hence does not recognize a suboperculum. 
Cunningham (13, 287) also says: ‘‘It is curious that while the 
anthropoid should be so absolutely destitute of a frontal and 
an orbital operculum, there are many of the lower apes (e. g. 
baboons, macaques, etc.) which show a faint trace of the orbital 
operculum.” 

In 1880), Parker, (46) ustated that the insula of) the 
chimpanzee ‘‘ is well developed and is marked by several con- 
spicuous radiating convolutions (gyri breves, Gall) and lies en- 
tirely concealed beneath a well-developed operculum.” 

In 1877 Major (34, 46) said the insula was relatively as 
well as absolutely smaller in the chimpanzee, etc., than in 
man. 

Hartmann (26, 195) in 1886 wrote that ‘In the gorilla, 
chimpanzee, and orang, the island of Reil is generally—at least, 
according to my experience—overlapped by the operculum, 
although there are instances in which this is not the case.” 


76 JOURNAL OF COMPARATIVE NEUROLOGY. 


In 1878 Broca (4, 645) said the insula of the gorilla bears 
‘‘three gyres as in the orang, while there are four in the chim- 
panzee.’’ Chapman in 1879 (g, 61) said: ‘‘The central lobe, or 
island of Reil, which is very slightly convoluted, is entire con- 
cealed in the chimpanzee.”’ 

Spitzka (56, 255) says that the gyres of the orang’s insula 
correspond as to their direction and relations to those of man, 
though less marked. He further states that ‘‘in every anthro- 
poid dissected by myself, I find these gyri and sulci; and one 
sulcus is a constant feature of even the Cynocephali.”’ 

The specimen here shown (Fig. 7) does not exhibit an 
area so well marked as those described by the above authors 
and this is probably due to the undeveloped state of the chimpan- 
zee brain at this age. Excepting the circuminsular, the fissures 
of the insula are very rudimentary and few in number. This 
insula was entirely concealed within the Sylvian fissure. There 
seems to have been a feeble attempt at producing a transinsular 
fissure but no apparent approximation to a pole. Broca (4, 369) 
says: ‘‘ L’insula est toujours simple chez les cébiens et les pith- 
éciens, ainsi que chez gibbons. Chez les grands anthropoides 
et chez l’‘homme, elle se subdivise en un certain nombre de plis 
qui convergent vers le pole et vont gagner successivement les 
divers points de la rigole supérieure, qu’ils traversent pour se 
jeter profondément dans le lobe frontal.”’ 

Marchand (36, 1893) says: ‘‘In the lower apes the isle 
of Reil isexposed. This is the case in microcephalic idiots.” 

Spitzka (56, 173) has shown that the claustrum in present 
in the baboon and that it is similarly located to that of man. 

The insula of the macaque monkey (Fig. 8) shows a well 
rounded eminence deeply imbedded in the Sylvian fissure. It 
is sharply demarcated by the circuminsular fissure but is other- 
wise unfissured and is’ about twice as long as broad. 

Cercocebus fuliginosus presents an insula very similar to that 
of the macaque. It, like the last-named, shows a well-rounded 
eminence and a well-defined circuminsular fissure. The insula 
was deeply imbedded in the Sylvian fissure. 

There are several lemurine brains in the Cornell Museum, 


CiarK, Comparative Anatomy of the Insula. 70 


but they were not available for examination at the time this 
thesis was prepared. 

In 1866, Flower (g1) said of the Javan loris (Stexops 
javanicus) on separating the lips of the Sylvian fissure ‘‘ no 
distinct median lobe, or insula, could be traced.” The same 
author also says ‘‘In the sulci of the outer face in the lemur, 
the Sylvian fissure is deeper especially at its anterior or lower 
part and conceals a small but distinctly marked insula or me- 
dian lobe.”’ 


ORDERS CHEIROPTERA AND INSECTIVORA, 


The specimens belonging to the above named orders, so far 
examined by the writer, showed no evidence of a macroscopic 
insula. 


THE INSULA IN THE CARNIVORA. 


The area which represents the insula in this order is, in 
general, not well developed, but in a more or less rudimentary 
form may be said to be fairly constant. As a macroscopic area, 
however, I have been unable to recognize it in the domestic cat, 
the Angora cat, Lyx rufus, the ocelot (Felis pardalis), the 
skunk (Mephitis mephitica), the mink (Putorius vison) and the 
ferret (Putorius domestica). My observations made in the 
latter part of 1893 upon the seal (Phoca vitulina) agree with 
the conclusion of .P. A. Fish (29 %, 16, 1896) who has 
shown that the Sylvian fissure has not suffered any 
marked displacement and that Spitzka (1890, Azmertcan Nat- 
uralist, XXVI, 115-122) erred in assuming a vertical com- 
municating fissure (v. 7. Fig. 9) as the true Sylvian. The posi- 
tion of the insula shows conclusively that the caudal one of 
these two fissures is the true Sylvian. Dr. Fish also says: ‘‘The 
vertical and the true Sylvian fissures meet superficially at the 
latero-ventral angle of the cerebrum and if the sides of the Syl- 
vian be separated, it will be seen that the vertical fissure in- 
stead of directly joining the Sylvian becomes a submerged fis- 
sure . . . andcrops out again on the ventral surface on 
the front or cephalic wall of the mouth of the Sylvian. 

This condition is found on both sides.’’ The insula is repre- 


78 JOURNAL OF COMPARATIVE NEUROLOGY. 


sented in the seal (Phoca vitulina ) and the panther (Fels con- 
color) in Figs. g and Io respectively. In each the area is devoid 
of fissures except the circuminsular, thus indicating its very rud- 
imentary character. The area is only slightly elevated and was 
entire concealed within the Sylvian fissure. It is present and 
of about the same rudimentary character in the hyena where it 
is flatter than in the preceding and is entirely concealed. It is 
concealed and rudimentary in the lion. In 1879 Pansch (45, 166) 
said that it is present but concealed in the fox. 

| Fig. 11 represents this area in the raccoon. The circum- 
insular fissure is interrupted at two points leaving the insula 
joined to the temporal lobe caudad and the cephalic boundary 
ill defined. This elongated and flattened area is crossed by a 
fissure which is believed to represent the transinsular, although 
it does not correspond exactly with the direction of the Sylvian 
(see Fig. 12), as is usually the case in the imprimates at least. 
Nevertheless it almost completely divides the area into a pre- 
insula and a postinsula, is deepest at its junction with the rhinal 
and at that point it corresponds in position with the Sylvian. 
This lack of conformity is strongly suggestive of a displacement 
of the Sylvian fissure as the result of growth in an inclosed 
space which may have carried the dorsal end of the Sylvian 
caudad while the bounding parts of the ventral end, which is 
more closely connected with the brain stem, have maintained 
their original position. Future investigation may also show that 
some peculiar development of the frontal or parietal lobes or of 
both has aided in bringing about this condition. The relation 
of the transinsular to the Sylvian fissure in this brain is strik- 
ingly similar to that in the human, but in the raccoon there is 
no evidence of the central fissure. The area is otherwise un- 
fissured, but slightly elevated and, with the exception of the 
cephalic third which lies concealed in the Sylvian fossa, it is 
completely hidden within the lips of the Sylvian fissure. As 
previously stated, the cephalic boundary is ill defined and the 
insular area graduates out into the frontal lobe as in the case of 
young pigs. The plane (plane of fissured surface) of the insula 
in this specimen forms an angle of more than 45° to the lateral 


CLaRK, Comparative Anatomy of the Insula. 79 


surface while that of the seal, panther, hyena and most primates 
is nearly parallel to the the lateral surface. 

The insula of the Thibet bear (Ursus thibctianus), Fig. 
13, is very different in its appearance from the preceding—as 
if, instead of the prolonged growth caudad, as in the raccoon, 
the surrounding areas which presumably have reached a higher 
state of, development, had produced a cephalo-caudal compres- 
sion of this area, resulting in the formation of two parallel gyres 
separated by the transinsular fissure. Relative to the length of 
the cerebrum, the insula in this specimen is much shorter than 
that in the raccoon. The transinsular fissure here is deep and 
coincides very closely both in direction and position with the 
Sylvian. The area is sharply demarcated by the circuminsular 
fissure except at the cephalic end where a slight depression 
probably marks the cephalic boundary. The area is well 
rounded and the plane of the insula is parallel to the lateral sur- 
face. The insula was not wholly concealed, a small portion of 
the caudo-ventral end of the cephalic subgyre being visible be- 
fore dissection. The area was almost entirely within the Syl- 
vian fissure. In 1888 Turner (59, 566) said: ‘‘ In the brain of 
the Polar Bear, I have shown that an entire arched convolution 
is concealed within that [Sylvian] fissure.”’ 

This area in the dog (Cans familiaris) is situated normally 
at the bottom of the Sylvian fissure and is entirely concealed. 
The plane is parallel to the lateral surface or nearly so. It isa 
well rounded, oval area somewhat triangular in form, is sharply 
defined by the circuminsular fissure and, as stated by Huxley, 
is usually unfissured. Figs. 14 and 15 represent an exceptional 
condition. The insula was exposed on both sides and was sit- 
uated almost wholly in the Sylvian fossa. The area was well 
rounded and its outline approximately rectangular. Another 
peculiar specimen is shown in Fig. 16. Here the insula, while 
entirely concealed within the Sylvian fissure, was crossed by a 
fissure which may represent the transinsular fissure and if so 
the preinsula is the smaller. In order to determine with greater 
certainty the nature of this fissure the area was removed and 
sectioned. The shallowness of the fissures (Fig. 17) plainly 


80 JoURNAL OF COMPARATIVE NEUROLOGY. 


shows that the fissure crossing this areais not a part of the 
circuminsular but an independent fissure, probably the transin- 
~ sular as previously stated. The small area cephalad undoubt- 
edly represents a preinsula. Owing to the normally simple 
character of this area in the dog, these individual differences 
are very striking. 

A transection of the cerebrum of a dog is shown in Fig. 
18. The insula appears as it normally exists concealed by the 
overlapping portions of the temporal and parietal lobes. The 
myelinic fibers of the olfactory tract readily distinguish the 
rhinal fissure and it will be noted that they do not extend be- 
yond the bottom of this fissure. A band of myelinic fibers 
extending apparently from the dorsal overlapping area of the pari- 
etal lobe, passes through the insular cortex cutting off the ental 
layers which constitute an area probably homologous with the 
claustrum. The study of sections caudad of this one shows 
that this band of fibers ventrad of the insula bends around into 
the temporal lobe. It would thus appear that a part of them at 
least might be association fibers. Fibers also given off from 
this band are distributed through the claustrum. Myelinic 
fibers are also found in the ectal dense layer of cortex of the 
insula and they extend for the most part parallel with the sur- 
face. 

The insula receives its blood-supply directly from minute 
branches of the medicerebral artery, which pass in from the sur- 
face and which are seen in longisection in Fig. 18. The writer 
has been unable in most cases to determine the origin of ves- 
sels found in the claustrum as they were seen only in transec- 
tion. A few however have been found which passed directly 
across the area as in the case of the one represented in the dor- 
sal end of the claustrum. These would seem to indicate that 
a part of the supply might be received directly from the surface 
arteries. In the vicinity of the fiber tracts, arteries, in general, 
seem to follow the direction of the length of the fibers and not 
to cross them. 

As previously stated, no macroscopic area appears at the bot- 
tom of the Sylvian fissure of either the cat (see Fig. 19) or the 


CiarK, Comparative Anatomy of the Insula. 81 


skunk (Fig. 20). In each the width of the Sylvian fissure is 
practically the same throughout its length and at the bottom as 
near the surface. The study of many transections of the Sylvian 
region confirms the statement that there is no marked widening 
of the Sylvian fissure at any point ; there is absolutely no sign 
of a circuminsular fissure and consequently no evidence of an 
ectal area homologous with the insula. Recourse was then had 
to the Weigert method with the result that it has been demon- 
strated in each that a cellular area had been separated from the 
cortex entad of the Sylvian fissure by a thin lamina of myelinic 
fibres. This area is similarly situated to the claustrum of the 
dog and in the skunk at least the fibers appear to arise from 
the area dorsad of the Sylvian and pass to some area ventrad or 
vice versa. In the cat, however, this lamina appears to connect 
the area ventrad of the Sylvian fissure with the central mass of 
myelinic fibers rather than with the area dorsad of the Sylvian. 
The origin of these fibers does not effect the question as to 
whether or not this is really a claustrum for, while it is known 
that fibers from the uncus and subfrontal gyri pass entad of the 
insula and assist in the demarcation of a claustrum in the human 
brain, it is not known but what fibers having a different origin 
also take part. Certain it is that if these myelinic fibers (which 
appear to be of the association type in the dog and skunk) were 
not there this area would remain a part of the cortex as has 
been shown is sometimes the case in the human brain. The 
fibers which make up this lamina are much fewer than in the 
dog and the number is still less in the skunk than in the cat. 
Some twelve days after I had thus independently reached the 
conclusion that this area was homologous with the claustrum, my 
attention was called by Professor Wilder to an article by 
Graeme M. Hammond, in 1881 (96). This article contains fig- 
ures showing that the author had noticed the area in the cat, 
sketched it though imperfectly and named it the claustrum. In 
1889, Spitzka (56, 173) also figured this area in the cat but did 
not name it. The writer was unable to recognize a claustrum 
in (Weigert) transections of the brain ofthe mink. This animal 
has no Sylvian fissure. 


82 JOURNAL OF COMPARATIVE NEUROLOGY. 


The significance of the presence of the claustrum is diffi- 
cult to determine. As previously stated, the existence of a 
~ claustrum might, in general, be taken to indicate the presence 
of an insula but such is not believed to be true in either of 
these instances. Presumably if a rudimentary insula did exist in 
the cat, then only a small portion of the claustrum would corres- 
pond therewith; and if only a part of the claustrum belongs to 
this area, then, as in this case where the claustrum is of consid- 
erable size, the larger part of the claustrum has necessarily no 
conection with the insula. Indeed it might exist quite as well 
without an insula. Hence it would appear that the only signifi- 
cance attached to the presence of a claustrum in such cases is 
that the cortex at the bottom of the Sylvian fissure is relatively 
farther entad than in those brains where the area is not so de- 
marcated, as in the sheep and mink. 

Turner in 1888 (59, 566) said: ‘‘In the true carnivora, the 
Sylvian convolution was, asa rule, superficial and on the cra- 
nial aspect, though in the otter and badger indications.of the 
depression of its anterior limb within the fissure were seen. In 
the seals and walrus, the concealment of this convolution was 
still more marked, so that the brains of these animals form ap- 
parently, in this particular, a transition to those of man and apes, 
in which the concealment of the island is complete.’ 


9 Nie 


THE INSULA IN THE PROBOSCIDIA. 


In the single specimen of the Indian elephant examined 
by the writer, the insula (Fig. 21) is only slightly developed. 
It consists of an almost flat area in the bottom of the Sylvian 
fissure and is made up of three gyres, two of which belong 
to the postinsula. This disposition of the insular area suggests 
that in the unusual dog (Fig. 16) except that in the latter the 
postinsula is at the same time more elevated. In the elephant 
the postinsula contains the greater number of gyres but the 
preinsula is the more elevated. The entire area is deeply 
imbedded in the Sylvian fissure and extends ventrad as far as 
the rhinal fissure. This observation is contrary to that of Owen, 
1868, who declares (44, 129) that ‘‘ the minor inter-Sylvian con- 


Ciark, Comparative Anatomy of the Insula. 83 


volutions are exposed in the sheep and elephant.” He also says 
(44, 123) ‘‘It is well marked and rises high up the Sylvian fis- 
sure in the Proboscidians.”’ In 1879 Spitzka (5g ) said that the in- 
sula in the elephant is completely covered. The largest branch of 
the medicerebral artery lies in what I have called the transinsular 
fissure but this artery does not reach the dorsal margin of thisarea. 
At about one-half or two-thirds of the distance from the ven- 
tral boundary of the insula, the artery leaves the area to pass 
dorsad into the Sylvian fissure. This peculiar condition seems 
to have been brought about by the very unusual growth of the 
area dorsad of the insula, which fold has been projected ventrad 
as a rudimentary subopercular fold. 


THE INSULA IN THE RODENTIA. 


None of the members of this order so far as examined by 
the writer showed any evidence of a macroscopic insula. Tran- 
sections of the Sylvian region prepared by the Weigert method 
failed to show the presence either of an insula or a claustrum in 
the woodchuck. | 

THE INSULA IN THE UNGULATA. 


With the exception of the primates, members of this order 
seem in general to have reached a higher state of insular devel- 
opment than those of any other order, a significant fact when it 
is considered that this order is usually placed so far down the list 
on account of other characters. The specimen of the camel here 
shown (Fig. 22) can hardly be taken to indicate much more 
than the existence of an insula in this animal. The area is 
nearly flat and presents no fissures other than the circuminsular. 
The insula is proportionally very small and shows on the whole 
a rudimentary state of development compared with the sur- 
rounding lobes. It was completely concealed within the Sylvian 
fissure, the ventral end scarcely reaching the rhinal fissure ; the 
wider end of the insula is dorsal. It is concealed and rudimen- 
tary in the bactrian camel (Camellus bactrianus). 

The deer (No. 961) presents a well defined area and at the 
same time one not indicating a high degree of development. 
The insula (Fig. 23) was completely concealed and is situated 


84 JOURNAL OF COMPARATIVE NEUROLOGY. 


almost wholly in the Sylvian fossa, only the caudal end extend- 
ing dorsad into the Sylvian fissure. With one exception the 
- fissures are merely broad shallow depressions which indicate a 
tendency to divide the area into five gyres. The caudal fissure 
probably represents the transinsular and is the only one indenting 
the lateral surface of this area. The plane of the insula is 
nearly perpendicular to the lateral surface; the surface is well 
rounded and throughout its entire length lies just dorsad of the 
olfactory tract. The insula is present also in the fallow deer 
(Cariacus clama) and exposed on both sides. At least three 
fourths of the insula is cephalad of the ventral end of the Syl- 
vian fissure. The transinsular fissure is rudimentary on the 
left side but better developed on the right. The difference in 
the position, relative development, form, and relations of the 
insula in the camel and deer are very great and more marked 
than one might expect in animals so closely related. 

The area which represents the insula in the sheep (Figs. 32- 
33) is, contrary to the rule, exposed. It is probably due to this 
fact that this area has not previously been treated by writers 
on the sheep brain. Only Leuret and Owen seem to have 
recognized it. The former states that, ‘except in) the wsheep: 
they [supplementary convolutions] are found ‘‘only in the brains 
of man, elephant and monkeys.’”’ Owen says: ‘‘the minor in- 
ter-Sylvian convolutions are exposed in the elephant and 
sheep.” 

This area, as has been rightly said, is exposed in the sheep 
and throughout its length lies just dorsad of the olfactory tract 
in the Sylvian fossa. The rhinal fissure which separates these 
areas is always well defined. In adults the lateral exposed sur- 
face is usually indented by two (apparently three) fissures—the 
most cephalic and caudal being found upon dissection to be 
parts of the circuminsular, while the median one is the lateral 
outcrop of what the writer considers the homologue of the 
transinsular fissure. Other indentations are frequently noticed 
but they are usually slight and are probably caused by pressure 
of branches of the medicerebral artery which lie in them. Up- 
on dissection, it becomes evident from the greater fissuration 


CrLarK, Comparative Anatomy of the Insula. 85 


that the dorsal and not the lateral surface is homologous to the 
lateral areas of most mammals and that the plane of the insula 
is perpendicular to the lateral aspect. Presumably in this brain 
as in the human the circuminsular fissure is the first developed 
and in adults is usually complete. In younger specimens 
however it is quite noticeable that this fissure is interrupted in 
one or more places as on the lateral surface at the cephalic end 
of Fig. 32 and at the dorso-caudal boundary of Fig. 33. With 
the exception of the circuminsular but one fissure of this area 
appears to be constant and is the only one extending upon the 
lateral as well as the dorsal surface. This fissure is continuous 
with the rhinal ; in it lies the largest branch of the medicere- 
bral artery which passes dorso-mesad into the Sylvian fissure ; 
it coincides throughout most of its course with the direction of 
the Sylvian, divides more or less completely the area into a 
preinsula anda postinsula and is the fissure which, in the dis- 
cussion of the lateral surface, was called the transinsular fissure. 
This branch of the medicerebral artery after traversing nearly 
two thirds of the length of the transinsular from the lateral sur- 
face mesad suddenly leaves it and passes up into the Sylvian 
fissure, thus giving the appearance as if the nearly equilateral 
triangular caudo-mesal portion of the caudal gyre of the pre- 
insula had grown caudo-ventrad of the artery. Presumably 
what has happened is that a displacement of the Sylvian fissure 
has occurred due toa greater development of the temporal or 
occipital lobes than the frontal and parietal; that this growth 
in an inclosed space has resulted in carrying cephalad the dor- 
sal end of the Sylvian and with it the artery, thus removing 
the artery from the mesal end of the transinular, where presum- 
ably it originally was lodged. The vertical position of the Syl- 
vian fissure would seem to indicate that such a displacement 
had occurred. The relation of the Sylvian to the transinsular 
fissure is very striking when compared with that in the raccoon, 
where the Sylvian was mostly caudad of the transinsular. In 
two instances only have fissures other than those above men- 
tioned been seen on the lateral surface of the sheep brain and in 
each case, as might be expected, they were in the preinsula. 


86 JOURNAL OF COMPARATIVE NEUROLOGY. 


The dorso-lateral aspect (Fig. 33) shows the preinsula to con- 
sist of three gyres while the postinsula has but one. The gyres 
- of the insula are in general simple. One specimen, however, 
showed the caudal gyre (postinsula) subdivided into three parts. 
The fissures grow deeper as they approach the meson except 
the transinsular but it is not known that any are constant. The 
sheep brain exhibits two unusual conditions shown in the tran- 
section of this brain (Fig. 34): first, that a bundle of myelinic 
fibers enters the insula from which it might be inferred that this 
area is functional, or at least assists in the performance of a 
function. These fibers seem to pass principally to the dorso- 
lateral part of this area, although it is evident that many are 
given off along its course to other parts of the area mostly 
dorsad. Secondly, there is no indication of a claustrum. It 
is not believed that any great significance can be attached 
to the apparent absence of this part in the sheep other than 
that the insula in this animal is relatively farther laterad than 
in those where the claustrum becomes so demarcated and hence 
the fibers from surrounding areas which in some other animals 
pass through the cortex here pass wholly mesad. The blood 
supply is derived wholly from branches of the medicerebral 
artery which extend over the surface. The position of the 
rhinal fissure and the relations of the insula to the olfactory - 
tract are clearly shown. The myelinic fibers of the latter ex- 
tend only to the bottom of the rhinal and in no case have fibers 
been found extending upon the insular area. This fiber area of 
the olfactory tract extends considerably farther ventrad than 
shown (Fig. 34) and at the ventral end the fibers are most 
numerous and collected into a bundle of considerable size. 

The insula of the cow (Figs. 26-29) is well developed, ex- 
posed on the lateral aspect and lies almost wholly within the 
Sylvian fossa. It presents in this animal peculiarities not pre- 
viously met with. The area in but one of five specimens was 
completely divided into a preinsula anda postinsula. In this 
one (3362) the transinsular fissure was well marked, was deepest 
at the latero-ventral end, and was continuous mesad with the 
circuminsular. The fissure cephalad of the transinsular also 


CiarK, Comparative Anatomy of the Insula. 87 


completely crossed the insula but was shallow at the ventral 
margin. The area is more often incompletely divided by the 
transinsular fissure which, while it is of considerable length and 
depth, is almost always confined to the lateral surface. The 
part most elevated in the five specimens examined lies in the 
postinsula always directly caudad or caudo-mesad of the dorsal 
end of the transinsular fissure. What has been assumed as the 
plane of the insula is at an angle somewhat greater than ninety 
degrees with the lateral surface. The caudal and caudo-lateral 
surfaces of this elevated portion slope very abruptly from the 
apex of this elevated portion to the temporal lobe. The slope 
toward the mesal boundary is always more gradual and becomes 
still more so as the cephalic boundary is approached. A cu- 
rious condition exists at the caudal end of the preinsula where, 
on the lateral surface, the appearance is as if a quite successful 
effort had been made to tuck that part caudo-mesad under 
the postinsula. This is more marked on the left side (see Fig. 
26) than on the right (Fig. 27), and the crowded appearance of 
this area seemed more marked in adult than in younger speci- 
mens. The surrounding lobes appear to be well developed, 
while the olfactory tract is enormous. The study of this brain 
seems to confirm the impression that these remarkable features 
are due to growth in an enclosed space. The insula shows the 
effects of compression in a marked degree. 

The circuminsular fissure of the right side (Figs. 27-28) sharp- 
ly defines the insular area except at the temporal lobe where the 
fissure is interrupted and a junction of the insula with the tem- 
poral lobe is formed at the lateral surface. This insula consists 
of seven gyres only one of which belongs to the postinsula. 
On the left side of the same specimen (Figs. 26 and 29) the 
circuminsular fissure is also interrupted with the formation of a 
similar junction as on the right side. Two interruptions of the 
circuminsular are found at the cephalic margin on the left side. 
The dorsal surface is divided into four gyres, three of which 
belong to the preinsula. A zygal fissure’ (Fig. 29, 2/) exists near 


1 See Wilder (64, 155). 


88 JOURNAL OF COMPARATIVE NEUROLOGY. 


the middle of this area, the base extending in the direction of 
the length of the area. While this surface is much less fissured 
than the corresponding surface of the right side, the lateral sur- 
face of the left side appears much more complicated. In the 
left insula of the cow (3137) the transinsular fissure was wholly 
exposed on the lateral surface; there was no interruption of the 
circuminsular at the cephalic margin, and the dorsal surface pre- 
sented five gyres, four of which belong to the preinsula. 

In 1890, Turner (61, ‘‘ Fig. 26’) gave a wrong idea of the 
Sylvian fissure (see Fig. 30) in the cow. He mistook the 
transinsular fissure for the Sylvian. 

In the insula of the cow, individual and lateral variations 
are perhaps more marked than in that of any other animal ex- 
amined. We have here exemplified also the peculiar folded 
appearance first noticed in the bear but more marked. 

The area representing the insula in the burro (Aguas asznus) 
is better developed than that in the deer. It is well rounded 
and, except at two points along the cephalic boundary, the area 
is sharply defined by the circuminsular fissure (Fig. 24). As in 
the deer, the area is divided into a preinsula and a postinsula by 
the transinsular fissure, the latter area containing but one, while 
the former consists of four gyres all well developed and sharply 
defined. There is no approximation to a pole asin man. The 
location of the area is similar to that of the deer, only the caudal 
one third lying in the Sylvian fissure. The two most cephalic 
gyres of the preinsula were exposed before dissection and the 
plane of the area makes an angle of not more than 60° with 
that of the lateral surface. 

In 1871, Lussana and Lemoigne (33) stated that the insula 
of the horse was of considerable size. The writer has exam- 
ined five brains, two of which, at least, are worthy of descrip- 
tion. The first was that of a work-horse (2777, R. half) which 
showed five gyres exposed previous to dissection. Upon dis- 
section the plane appeared almost perpendicular to the lateral 
surface, the gyres were six in number and the fifth (from the 
cephalic end) was subdivided by a small fissure at the mesal 
end, The fissure separating gyres 2 and 3 is deep, extends 


CiarK, Comparative Anatomy of the Insula. 89 


over the dorsal surface and almost completely divides the pre- 
insula upon the lateral surface. 

The insulas of the ‘‘Sage horse” (Fig. 25) consist of 
five gyres each, of which three belong to the preinsula and two 
to the postinsula. Before dissection the left insula showed four 
gyres exposed while the right showed three. One of the main 
branches of the medicerebral artery lies in what I have called 
the transinsular fissure (Fig. 25). This fissure is well marked 
on both sides. In the left insula the two most cephalic gyres 
are nearly perpendicular, while the other three are at an angle 
of about 60° to the lateral surface. On the right all the gyres 
are in about the same plane which lies nearly perpendicular to 
the lateral aspect. Those of the postinsular seem better devel- 
oped and the first gyre of the postinsula is the one most ele- 
vated. There seems to be an approximation to a pole in this 
specimen not found in any other animals than the primates. 
The second gyre is also the smallest as in man. 

The insular region of the tapir (Zapzrus malayanus, Fig. 
31) presents a very peculiar appearance, from the fact that be- 
tween the area which I have called the insula and the rhinal fis- 
sure there seems to have arisen a corrugated area which is not 
believed to belong to the insula at all. This area extends from 
the frontal to the extreme end of the occipital lobe and through- 
out its course lies just dorsad of the rhinal fissure which is in- 
terrupted at several points on the left side but is continuous on 
the right. The fissure dorsad of this area is nearly parallel to 
the rhinal except near the caudal end. The area included be- 
tween these two fissures is fissured, but only slightly elevated 
above the olfactory tract, and is continuous with the insular area. 
The insula in this specimen is only slightly elevated and con- 
tains but two gyres of which the more cephalic is the larger. 
The fissures are simple and very shallow exeept the circuminsu- 
lar at the cephalic end. The transinsular is not well marked 
near the mesal margin. The insula is entirely concealed. 

In 1878 Krueg (30, 325) said the insula was present in the 
Hippopotamide. Chapman in 1881 (9, 144) said: ‘‘The 


90 JouRNAL oF COMPARATIVE NEUROLOGY, 


Sylvian fissure in my Hippopotamus is quite evident and within 
it I noticed a rudimentary island of Reil.”’ 


THE INSULA IN THE CETACEA. 


Fig. 35 represents the insula of the porpoise (Globzoceph- 
alus melas). In this specimen the insula is completely con- 
cealed within the Sylvian fissure and the plane is parallel to 
the lateral surface. The area is proportionally very large but 
at the same time very flat and there is no evidence of a fis- 
sure homologous with the transinsular and no approximation to 
a pole. This specimen shows at least nine gyres, three of 
which are subdivided by fissures which are deepest ‘near the 
circuminsular. It is not known that any of the fissures of this 
area are constant. In 1879 Spitzka (53) says the ‘‘insula in the 
porpoise has four times as many convolutions and is twice as 
large as that in man and is completely covered by the opercu- 
The same author also says that one 


” 


lum and temporal lobe. 
specimen in his posssession has either thirteen or fifteen gyres. 
Wilder [lecture] said that the gyres of the porpoise insula ex- 
ceed in number those of any other mammal observed by him, 
but they are smaller, and he concluded that ‘‘in combined 
elevation and fissuration, the insula reaches its highest devel- 
opment in man.”’ 

Ziehen (31, 110, 1889) says of the white whale (Beluga 
lucas), ‘‘ The area between the fissura circularis interna and the 
fissura circularis externa, probably also a large part of the cov- 
ered portion of the floor of the Sylvian fossa ought to be desig- 
nated as insula.”” Of the Sylvian fissure in W/egaptera boops, he 
(31, 125) says ‘‘Ursprung vom aus der Fissura Rhinalis An- 
terior. Die MHaupttheil bildet die obere Begrenzung des 
Inseldreiecks.”’ 

None of the representatives examined of the remaining 
orders show any evidence of an insula. 


CiarKk, Comparative Anatomy of the Insula. fo 


SUMMARY. 

1. Neither the insula nor the claustrum is constant among 
mammals. 

2. The insula is present in the Primates, Carnivora, Pro- 
boscidia, Ungulata and Cetacea. It is apparently absent in the 
Cheiroptera, Insectivora, Rodentia, Sirenia, Marsupialia and 
Monotremata. 

3. The claustrum may include parts of areas other than 
the insula. 

4. Theinsula and the claustrum may generally be consid- 
ered as parts of the same cortical area; the claustrum may be 
present without the insula; both may be present or both may be 
absent. 

5. The apparent absence of the claustrum when the in- 
sula is present indicates that, relative to the surrounding areas, 
the insular cortex is farther laterad than in those specimens 
where it is so demarcated. The myelinic fibers of the sur- 
rounding areas then pass wholly entad of the insular cortex. 

6. The insula is, in general, a convoluted area. 

7. The insula is sometimes a subarea but quite often it is 
exposed upon the lateral surface of the cerebrum. 

8. The exposure of the insula may indicate either its own 
excess or the deficiency of the overlapping adjacent parts. 

O:)) bhe® eytes of the overlapping areas are always _ inter- 
calated with those of the insula. 

10. Individual and lateral variations upon this area are 
numerous and marked and are probably due in part to growth 
in an enclosed space. 

11. The insula in the Primates, the Carnivora, some of 
the Ungulata and Cetacea and perhaps others is a subarea or sub- 
gyres at the bottom of the Sylvian fissure. 

12. The insula frequently is almost wholly within the Syl- 
vian fossa. This condition is not believed to indicate in any 
way a different relation to the parts entad but rather a more uni- 
form development of the insula and the surrounding areas so 
that no lateral overlapping has taken place and hence the ven- 
tral end of the Sylvian fissure remains unclosed as a fossa, 


92 JOURNAL OF COMPARATIVE NEUROLOGY. 


13. The insular area, in the more generalized types of 
mammals, at least, does not belong primarily to the rhinenceph- 
-alon as held by Turner in regard to the human brain. 

14. The fissured surface which is usually the dorsal in the 
raccoon, the deer, the burro, the horse, the pig, and the sheep, 
is homologous with what appears on the lateral surface of the 
insula of the Primates, most of the Carnivora and some others. 

15. The variation in the plane of the fissured surface of 
the insula is probably due to some peculiarity in the develop- 
ment of the brain stem. 

16. Fissures on the insula are less deep and more open 
than on the surrounding parts; this is probably due to growth 
under pressure of overlapping parts. 

17. The lack of conformity between the Sylvian and 
transinsular fissures both in position and direction is probably 
due to the mechanical effect of growth in an enclosed space by 
which a displacement of the Sylvian has occurred. 

18. The difference in the relative mass of the insula, its 
position, plane and relative development in the camel and deer 
is very much greater than might be expected in two animals so 
closely related. 

19. The difference between the insula of the bear and the 
raccoon is very striking. 

20. It is noteworthy that the cat (7. domestica) and oce- 
lot (/. pardalis) and the lynx have no insula, while the panther 
(#. concolor) and the lion (/. /eo) have a rudimentary one. ° 

21. The overlapping of the insula in the human brain, 
and presumably in others, is not due to a cessation of growth 
of the insula but to the more rapid growth of the surrounding 
lobes and the rapid enlargement of the paracceles. 

22. The apparent approximation to a pole in the horse is 
noteworthy. 

23. The primitive insula, if such exists, is a somewhat 
elevated area of greater or less size surrounded by a circuminsu- 
lar fissure and located in the Sylvian fossa or in the fissure if 
the fissure is continuous with the rhinal. 


CLARK, Comparative Anatomy of the Insula. 93 


SYNONYMY—INSULA. 


1. Central lobe.—The application of this term to the insula is objection- 
able on account of its more common use in connection with a lobe of the cerebel- 
lum and also because the adjective, central, is often applied to fissures and lobes 
topographically related to the central fissure (f. of Rolando). 

2. Convolutions placed between the fissure of Sylvius and the corpus 
striatum (Vicq d’Azyr). 

3. Circonvolutions supplementaires (Leuret and Gratiolet). 

Entosylvian folds or tract (Owen). 
s. Fifth lobe of the brain. 
6. Gyri breves (Gall, Arnold). 
7.  Gyri operti. 
8. Gyri unciformes (Eberstaller). 
Opn elle: 

10. Insel (Reil). 

11. Insellappen. 

12. Insula. This name takes precedence over all others, being the Latin 
form of ‘‘Insel,’’ the name given by its first describer, Reil. The regular par- 
onyms of insula as given by Wilder (22, 530-31) are English, insula; German, 
Insel; French, insulé; Italian. isola. 

13. Insula de Reil. 


14. Insula del Reil. 
15. Insular lobe. 


16. Insula Reilii. 

17. Insula Sylvii (Mingazzini). 

18. Intersylvian convolutions (Owen). 

19. Intralobular gyri (Quain). 

20. Island of Reil (English Authors). 

Zien wsolar 

22. Limen insule. 

23. Lobe central (see note under 1). 

24. Lobe central de l’insula (see note under 1). 
25. Lobe de l’insula de Reil (Broca). 

26. Lobe intermediare. 

27. Lobe moyen. 

28. Lobettino centrale nel tipo pecorino (Tenchini and Negrini). 
29. Lobo centrale (Gratiolet). 

30. Lobo fondamentale (Lussana). 

31. Lobo insulare (Lussana). 

32. Lobule de l’insula. 

33. Lobule du corps strié. 

34. Lobule sous-sylvien (Broca). 

35. Lobulo del corps striato. 

36. Lobulo sotto-silvico (Tenchini and Negrini). 
37- Lobulus centralis (see note under 1). 

38. Lobulus corporis striati. 

39. Lobulus fissuree Sylvii. 


904 JOURNAL OF COMPARATIVE NEUROLOGY. 


40. Lobus caudicis (Burdach). 

41. Lobus insule. 

42. Lobus intermedius s. opertus. 

43. Lobus opertus (Arnold). 

44. Lobus retractus. 

45. Median lobe. 

46. Minor intersylvian convolutions (Owen). 

47. Quinto lobo delle cervello. 

48. Reil’s island. 

49. Stammlappen (Huschke), 

50. Subsylvian fold (Owen). 

51. Subsylvian lobe. 

52. Subsylvian tract (Owen). 

53. Versteckenlappen (Arnold). 

54. Zwischenlappen (Arnold). 

SYNONYMY—CLAUSTRUM. 

I, Avant-mur. 
2. Claustrum (Burdach). 


3. External wall (Huguenin, Charcot). 
4. Noyau rubané. 
5- Nucleus teenizformis (Arnold). 
6. Rampart. 
7. KRempart. 
8. Vormauer. 
SYNONYMY—TRANSINSULAR FISSURE. 
1. Fissura interinsularis (Eberstaller). 
2. Hauptfurche der Insel (Eberstaller). 
3. Inselfurche (Eberstaller). 
4. Le grand sillon de l’insula (Brissaud). 
5. Sulcus centralis insulze (Guldberg) 13. 
6. Sulcus centralis Reilii (Turner). 
= 7. Sulcus Insulee (Eberstaller). ¥ 
8. Transinsular fissure (Wilder). 
SYNONYMY—CIRCUMINSULAR FISSURE. 
1. Circuminsular fissure, (Wilder). 
2. Fissura circularis. 
3. Rigoles de ]’insula (Broca). 
4. Sulcus circularis RKeilii (Schwalbe) 43. 


BIBLIOGRAPHY. 


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Phys., XXIV ; new series, IV, 309 (IX-XI). 

2. 1882. BOURNEVILLE et BONNAIRE. Lesion Ancienne du Lobule de 
*Insula. Archives de Neurologie, III, 176-186. 


CLARK, Comparative Anatomy of the Insula. 95 


3. 1893. BrissAuD, E. Anatomie du cerveau de ’homme. Paris, Atlas 
and Text. 

4. 1888. Broca, PAuL. Memoires sur le cerveau de homme et des pri- 
mates. aris. 

5. 1889. BROWNING, WM. Vessels of the Brain. Ref. H’db’k, Med. Sct- 
ences, VIII, 231-244. 

6. 1890. BROWNING, WM. Vessels of the Brain. Ref. H’di’k. Med. 
Scez., IX, 138-141. 

7. 1819-1826. BurRpACH, KARL FRIEDRICH. Vom Baue und Leben 
des Gehirns. Lezpzig. 

§. 1879. CHAPMAN, HENRY C. On the Structure of the Chimpanzee, 
Proc. Acad. Nat. Sct., Philadelphia, 52-63. 

9. 1881. CHAPMAN, HENRY C. Observations on the Hippopotamus. 
Proc. Acad. Nat. Sct., Philadelphia, 126-148. 

10. 1892. CHAPMAN, HENRY C. Observations upon the Brain of the 
Gorilla. Proc. Acad. Nat. Sct., Philadelphia, 203-212. 

11. 1883. CHARcoT, J. M. Lectures on the Localization of Cerebral 
and Spinal Diseases. arts. Translated and edited by W. B. Hadden, 
London. 

12. 1893. CLARK, TRAcy E, The Insula of the Pig. Jour. Comp. Neur. 
March, 7-10, 1 plate. 

13. 1891. CUNNINGHAM, D. J. The Sylvian Fissure and the Island of 
Reil in the Primate Brain. Journ, Anat. and Phys., XXV, pt. 1, 286-291. 

14. 1891. CUNNINGHAM, D. J. The Development of the Gyri and Sulci 
on the Surface of the Island of Reil of the Human Brain. Journ. Anat. and 
Phys., XXV, pt. III, 338-348, New Series, Vol. V. 

15. 1892. CUNNINGHAM, D. J, Contribution to the Surface Anatomy of 
the Cerebral Hemispheres. Cunningham's, Memoirs, No. 7, R. 1. Acad., Dublin, 

16. 1786. D’AzyR, Vicg. Traité d’anatomie et de physiologie. ards. 

17. 1887. EBERSTALLER, Oscar. Zur Anatomie und Morphologie der 
Insula Reilii. Azat. Anz., II, 739-750. 

18. 1888. EBERSTALLER, OscAR. Noch einmal die Insula Reilii. Azar, 
Anz ils 52. 

19. 1885. FAmILIAntT, VicrortaA. Beitrage zur Vergleichung der Hirn- 
furchen bei den Carnivoren und den Primaten. Sern. 

20. 1886. FERE, CH. Traité elementaire d’anatomie médicale du systéme 
nerveux, 79-87. aris. 

20%. 1896. FIsH, PIERRE A. A Note on the Cerebral Fissuration of the 
Seal (Phoca vitulina). Journ. Comp. Neur., V1, 15-18, 1 plate. 

21. 1866. FLower, W. A. On the Brain of the Javan Loris (Stenops 
javanicus). Zvansactions Zoological Soc., V, 103-110. 

22. 1889. GAGE and WILDER. Anatomical Terminology. Ref. A’dd’h, 
Med. Scz.. VIII, 5§15-537- 

93. 1881. GIACOMINI, CARLO. Varieta delle circonvoluzioni cerebrali 
del’uomo. Zorino. 

24. 1884. GIACOMINI, CARLO. Guida allo Studio delle circonvoluzioni 
cerebrali dell’uomo. Torino, 92-99. 


96 JouRNAL OF COMPARATIVE NEUROLOGY. 


25. 1887. GuLpBerGc, Gustav A. Zur Morphologie der Insula Reilii. 
Anat. Anz., 11, No. 21, 659-665. 

26. 1881. Hammonp, GRAEME M. Anew Cortical Centre. Medical 
’ Record, March 19, Abstract in Scéence, April 16. Mew Vork. 

27. 1886. HARTMANN, RoBERT. Anthropoid Apes. Mew York. 

28. 1893. HerrRIcK, C. L. Recent Progress in the Comparative Anato- 
my of the Nervous System. Ref. H’dd’k. Med. Scz., 1X, 669-690. 

29. 1872. Huxtry, THos. H. A Manual of the Vertebrated Animals, 
New York. 

30. 1878. KRrueEG, JuLius. Ueber die Furchung der Grosshirnrinde der 
Ungulaten. Zettschrift f. Wiss. Zool., XXXI, 297-345. 

31. 1889. KUKENTHAL und ZIEHEN. Ueber das Centralnervensystem 
der Cetaceen. Vergleichend-anatomisch und Entwickelungsgeschichtliche Untersuch- 
ungen an Walthieren, Erster Theil. ena. 

32. 1839-56. LrURET et GRATIOLET. Anatomie comparée du systéme 
nerveux de l’homme et des primates. Tome II, 112. 

38. 1871. LussANA e LEMOIGNE. Fisiologia dei centri nervosi ence- 
falici. Padova. 

34. 1877. Major, HERBERT C. The Structure of the Island of Reil in 
Apes. Lancet, II. 

35. 1877. Major, HERBERT C. The Histology of the Island of Reil. 
Monthly Micoscopical Journal, July, XVIII, 14-21. London. 

36. 1893. MARCHAND, FELIX. Die Morphologie des Stirnlappens und 
der Insel der Anthropomorpben. /Jena, Monograph, 180 pp., 3 plates. Abstract 
in Jour. Comp. Neurol., 1894, IV, p. cx. 

37. 1864. MARSHALL, JOHN. Onthe Brain of a Bushwoman and on 
the Brains of two Idiots of European Descent. Phzlos. Transactions, 501-508. 
London, 

38. 1872. MEYNERT, THEODOR. The Brains of Mammals. Stricker’s 
Manual of Histology, 650-766. Translated by Buck, Mew York. 

39. 1885. MEYNERT, THEODOR. Psychiatry. Translated by Sachs, 
New York. 

40. 1886. Mitts, C. K, Arrested and Aberrant Development of the 
Fissures and Gyres in the Brains of Paranoiacs, Criminals, Idiots and Negroes. 
Journ, Nervous and Mental Disease, XIII Nos. 9 and to. 

41, 1888. MINGAZzINI, J. Ueber die Entwickelung der Furchen und 
Windungen des Menschlichen Gehirns. Gzessen. 

42, 1788. Monro, ALEXANDER. Observations on the Structure and 
Functions of the Nervous System. 

43, 1890. OBERSTEINER, HEINRICH. The Anatomy of the Central Ner- 
vous Organs. Translated by Hill, Ph2/adelphia. 

44, 1568. Owen, RICHARD. On the Anatomy of the Vertebrates, III. 
London. 

45. 1879. Panscu, ADOLF. Beitrige zur Morphologie des Grosshirns 
der Saiigethiere. Morphologisches Jahrbuch, Fiinfter Band. Leipzig. 

46. 1880. PARKER, ANDREW J. JMéedical Record, Jan. 10. 


CLARK, Comparative Anatomy of the Insula. 97 


47. 1804-6. ReIL, JoH. Curist. On the Sylvian Fissure and Insula, * 
Archiv fiir Physiologie, 1X, 196. 

48. 1879. RicHET, CHAs. Physiology and Histology of the Cerebral 
Convolutions. Translated by Fowler. Mew Vor. 

49. 1887. RoLiEston, H. D. Description of the Cerebral Hemispheres 
of an Adult Australian Male. Journ, Anthrop. Inst. G. B. and I., XVII, 32-43. 

50. 1893. SAcHs, HEINRICH. Vortrige iiber Bau und Thitigkeit des 
Grosshirns und die Lehre. Breslau. 

51. 1881. SCHWALBE, G. Lehrbuch der Neurologie. Zylangen. 

52. 1879. SpitzKa, E.C. Report of N. Y. Neurol. Soc. Meeting. 
Medical Record, May 25. New Vork. 

53. 1880. SpirzKA, E.C. The Brain of a Porpoise, Medical Record, 
XVII, 72, Jan. Mew York, 

54, 1880-81. SpirzKA, E.C. A Remarkable Peculiarity of an Anthro- 
poid Brain. Sctence, I, 25. Mew York. 

55. 1882. Spitzka, E. C. On the Autopsy of Guiteau. Amer. Journ. 
Neurol. and Psychiatry, 1, 381-392. 

56. 1883. SpitzKa, E. C. Contributions to Encephalic Anatomy. Amer, 
Journ. Neurol. and Psych., Il, 249-274. 

57. 1889. Spitzka, E.C. Histology of the Brain. Ref. A’di’k Med. 
Sc#., VIII, 164-189. 

58. 1885. STARR, M. ALLEN. Diagnosis of Local Lesions of the Brain. 
Ref. Hdb’k. Med. Sci., 1, 642-648. 

59. 1889. TENCHINI e NEGRINI. Sulla Corteccia Cerebrale degli Equini 
e Bovini. Parma. ; 

60. 1858. TURNER, SIR WM. Convolutions of the Seal and Walrus, 
Journ. Anat. and Phys., XXII, 566-568. 

61. 1890. TURNER, SIR WM. The Convolutions of the Brain. Journ, 
Anat. and Phys., XXV, 105-153. 

62. 1887. WALDSCHMIDT, JULIUS. Beitrige zur Anatomie des Taub- 
stummengehirns. Adlgemeine Zettsch. f. Psych. u. Psychische Gerichtliche Mea- 
zcin, XLIII, 373-379. 

63. 1886. WILDER and GaGE, Anatomical Technology, 2nd Ed. Mew 
York. 

64. 1889. WiLpER, BurTG. Brain, Gross or Macroscopic Anatomy. 
Ref, Hdb’k. Med. Scz., VIII, 107-164. 

65. 1893. WILDER, Burt G. Brain, Gross or Macroscopic Anatomy. 
Ref. Hdb’k. Med. Sci., UX, 99-111. 

66. 1894. WILDER, Burt G,. Exhibition of a Suicide’s Brain, with two 
Pistol-ball Wounds. Remarks on its Fissural Anomalies. Journ. of Nervous 
and Mental Disease, December, 1-4. 


EXPLANATION OF FIGURES. 


With the exception of figures 1, 2, 3, 4, 5, 6, 7, 8, 12, 17, 18, 19, 20, 21, 
25, 31 and 34, all the figures were produced as follows: The specimens were 
photographed natural size, with a vertical camera; from the photographs out- 


* The exact title of Reil’s article is not known by the writer. 


98 JOURNAL oF COMPARATIVE NEUROLOGY. 


lines were traced also with the camera. Liability to distortion due to the eu- 
largement of the parts of an uneven surface was materially reduced. Figures 
I, 2, 3, 4, 5, 6. 7, 21, 25, 31, were photographed natural size and enlarged from 
photographs or figures as indicated, with an enlarging apparatus. Figures 8, 
17, 18, 19, 20 and 34 were photographed as stated and traced from photographs, 
Fig. 12 was made from outlines traced from photographs of lateral views 
before and after dissection. 


ABBREVIATIONS. 

art.—artery. p. of.—postoperculum, 
circ. f.—circuminsular fissure. pretn.—preinsula. 
c/.—claustrum. Preop.—preoperculum. 
fi. olf. tr.—myelinic fibres of olfactory 7h. f.—rhinal fissure. 

tract. subop.—suboperculum. 
im.—insula. Sy. #.—Sylvian fissure. 
of.—operculum. tr. f.—transinsular fissure. 
P.—-Pole of the Insula. z. f.—zygal fissure. 
postin.—postinsula. v. f.—vertical fissure. 


DESCRIPTION OF FIGURES. 


The numbers are the serial numbers of the Museum of Vertebrate Zoology 
of Cornell University. 


PLATE IX. 


Fig. 1. UWateral aspect of the left half of the brain of a human fetus, size 
and age unknown. 2278; x 1 (after Wilder, 68, Fig, 4760). 

Fig. 2. Ventral aspect of the left cerebrum of a human fcetus, size and 
age unknown. 1820; x 1 (after Wilder, 64, Fig. 4782). 

Fig. 3. Wateral aspect of right human insula, age and sex unknown, adult, 
1805; x I. The insula was exposed by cutting away the surrounding lobes 
down to the level of the circuminsular fissure, thus exposing the medicornu and 
hippocamp. The three eminenges 1, 2 and 3 belong to the postinsula. The 
fissure marked 4 is too sharply outlined near the ventral end. At the point in- 
dicated it is a mere vasal furrow and does not communicate with the Sylyian 
fissure ventrad. 

Fig. 4. Lateral aspect of right human insula, age and sex unknown, 
young, 1823; x 1. This preparation shows the branches of the medicerebral ar- 
tery as it divides over the surface of the insula. The largest branch lies in the 
transinsular fissure for about two thirds of the distance across the insula, then 
leaves this fissure and bends caudo-dorsad as if towards the Sylvian. There is 
also shown a tendency toward the demarcation of an eminence similar to that 
shown in Fig. 3. 

Fig. 5. Dorso-lateral aspect of the right human insula, age 37 years, 
male, 376; x 1. A nearly normal insula except for the unusually élevated pole, 
P. The short furrow cephalad is represented too deep and too narrow. 

Fig. 6. Caudal aspect of a transection of the left cerebrum of man, after 
Wilder (64, 4732)- Shows the relation of the insula to the claustrum, the brain 
stem and operculums. 


CrLarK, Comparative Anatomy of the Insula. 99 


PLAT EHX. 


fig. 7. Lateral aspect of the right insula of chimpanzee, supposed age 4 
years, female, 265; x 1. The operculums have been cut away. 

Fig. 8. Lateral aspect of the right insula of a macaque monkey, age and 
sex unknown; x I. The lateral portion of the cerebrum was removed to the 
level of the ventral half of the circuminsular fissure. The remainder of the 
insula was exposed by cutting a groove parallel with the Sylvian fissure. 

fig. g. Lateral aspect of the left insula of a seal (Phoca vitulina); x 1. 
Operculums cut away. The black areas at the dorsal and ventral ends indicate 
broken surfaces where the parietes were removed, and are too black. 

fig. ro. Lateral aspect of the left insula of a panther (#elzs concolor), fe- 
male, 18 years old, 309; x 1. QOperculums cut away. 

fig. zr. Lateral aspect of the right insula of an adult raccoon (Procyon 
Jotor), male, age unknown, 2031; x I. The overlapping portions have been re- 
moved. 

fig,712, Diagram of the same, showing the relation of the Sylvian and 
transinsular fissures. 

fig. 13. Lateral aspect of the right insula of a Thibet bear (Ursus thibet- 
zanus), male, age at least 5 years; 645; x I. Shows well the relation of the 
transinsular and Sylvian fissures. 

fig. 14. Lateral aspect of insular region of a dog (cross between French 
poodle and Skye terrier), age more than 18 years, sex unknown, 166; x I. 
Shows the exposure of the insula before dissection. 

Fig. 15. Lateral aspect of the insula of the same with the operculums cut 
away; x 1%. 

Fig. 16. Lateral aspect of the right insula of a dog (Canis famzliarts), age 
unknown, probably adult, male; x I. 

fig. 17. Outline of a section of the above looking dorsad; x1. The 
dotted brace indicates the extent of the insula. 


PLATE XI. 


fig. 18. Transection of the cerebrum of a dog (C. familiaris), male, age 
a few weeks, 3124; x 21%. Section made according to Weigert’s method, after 
hardening the brain in 3% and 5% bichromate of potassium, imbedded in collo- 
dion and cut in oil; differentiates alba and cinerea. 

fig. 7g. ‘Transection (Weigert) of a portion of the left cerebrum across the 
Sylvian fissure of a cat (Hels domestica), age and sex unknown; x 2%. 

Fig. 20. Transection (Weigert) of a portion of the right cerebrum across 
the Sylvian fissure of a skunk (Mephitis mephitica), age atid sex unknown; x 2%. 

fig. 34. Transection of the right insular region of a sheep (Ov7s aries), 
age and sex unknown, looking cephalad ; x 4%. 


PLATE XII. 


Fig. 21. Lateral aspect of the left insula of an elephant (Z/ephas zndicus), 
age unknown, female, 2181; x 1. Overlapping portions have been dissected 
away, 


100 JOURNAL OF COMPARATIVE NEUROLOGY, 


Fig, 22. Lateral aspect of insulaof a camel (Camelus dromedarius), male, 
age 15 minutes, 2122; x I. The shaded portion at the right of the insular area 
is a broken surface and does not belong with the insula. 

Fig. 23. Lateral aspect of the left insula of a deer (Cariacus virginiana), 
male, age unknown, 961; x I. 

Fig. 24. Lateral aspect of the left insula of a Mexican burro (Zguus 
asinus), male, age unknown, 2259; x I. 

Fig\25. Lateral aspect of the left insula of a horse (#guzs caballus), male, 
adult, 2095; x I. 

Fig. 26, Wateral aspect of the left insular region of a cow (Bos taurus), 
adult, sex unknown, 3408; x I. Shows exposure of the insula. 

Fig. 27. Lateral aspect of the right insular region of a cow (Bos taurus) 
adult, sex unknown, 3408; x I, Shows exposure of insula before dissection. 

Fig, 28. WDorso-lateral aspect of the right insula of the same, exposed by 
cutting away the operculums. 


PLATE XIII: 


Fig, 29. Dorso-lateral aspect of the left insula of the same after dissection. 

Fig. 30. Lateral aspect of the cephalic half of the left cerebrum of 
a cow (Bos taurus), after Turner (61, Fig. 26). The fissure marked 5S, which 
Turner considers as the Sylvian, is the transinsular; the Sylvian is directly 
dorsad. 

Fig. 31. Lateral aspect of the left cerebrum of a Malayan tapir (Zapzrus 
malayanus), age and sex unknown, 2123; x 1. Dorsal portion of the cerebrum 
not shown ; insula exposed by removal of overlapping parts. 

Fig. 32. \ateral aspect of the left cerebrum of a sheep (Ov¢s artes), sex 
and age unknown, probably young; x 1. Shows the insula exposed before 
dissection. 

Fig. 33. Dorso-lateral aspect of the right insula of a sheep (Ov¢s arzes), 
sex unknown, probably young; x 1. Insula exposed by cutting away dorsal 
overlapping portions. 

Fig. 34. See description under Plate XI. 

Fig. 35. Lateral aspect of the insula of a Porpoise (Glodicocephalus melas), 
female, adult, 670; x 1. Insula exposed by cutting away of the operculums. 
Sy. f. designates the basisylvian fissure between the temporal and frontal 
lobes. 


REVIEW OF THE GOLGI METHOD. 


By OLIVER S. STRONG. 


The advent of the Golgi method in nerve histology has so 
ereatly enlarged our knowledge and altered our conceptions of 
the structure of the nervous system in many respects and the 
method, or methods, itself has such well defined peculiarities that 
it has been thought that a general review of it, from the techni- 
cal side would be of interest and perhaps of use, especially in 
view of the very considerable number of investigators now 
employing it. 

The review does not aim at any originality of treatment 
but is simply a compilation from available literature of its vari- 
ous modifications and applications. It may be stated that it 
does not include Golgi’s arsenic-gold chloride method nor even 
the application of the bichromate-silver methods to the structure 
of medullated nerve fibres. 

It has seemed most appropriate to begin the review with a 
translation of the technique of Golgi’s methods as given by Golgi 
himself, principally in his work ‘‘Studi sulla fina anatomia degli 
organi centrali del sistema nervoso,” pp. 181-208. The trans- 
lation is made, however, from the German edition of Golgi’s 
works ( ‘‘ Untersuchungen iiber den feineren Bau des centralen 
und peripherischen Nervensystems,” pp. 169-182, translated by 
R. Teuscher). Golgi’s own account of the technique is still 
the most complete nor does it seem to be by any means univer- 
sally understood how completely Golgi worked it out and how 
largely we owe not only the discovery but also the development 
of the method to him. It is for these reasons as well as for the 
many valuable hints contained therein that the translation of this 
rather extensive account of Golgi’s is here given. 

‘The particular methods to which I owe my most note- 
worthy success are the following: (1) The method of black- 
staining by successively treating the pieces (of brain tissue) 


102 JOURNAL OF COMPARATIVE NEUROLOGY. 


with bichromate of potassium or ammonium and silver nitrate. 
(2) The method of the successive action of a mixture of osmic 
acid and bichromate and of silver nitrate. (3) The method of 
the combined action of bichromate of potassium or ammonium 
and bichloride of mercury. (The stain appears black by trans- 
mitted, metallic white by reflected light). 

‘“(1) Zhe method of the combined action of bichromate of 
potassium and of silver nitrate. In the series of methods which 
I have especially employed this is, in a manner, the fundamen- 
talone. The others are only variations of this, devised to 
shorten the time of the preliminary treatment, to make the 
preparations more stable, to vary the results in various ways, 
especially to obtain a greater extension of the reaction and to 
cause the reaction to affect one or another species of the ele- 
ments or a part of them. 

‘‘T consider it to the point to call attention to the fact that 
the procedure of the microscopical technique which I will de- 
scribe, although it rests essentially upon the action of silver ni- 
trate, has nothing in common with the usual method of staining 
the intercellular substance of endothelium, epithelium and con- 
nective tissue brown or black. In the latter method dilute so- 
lutions of silver nitrate are applied immediately to the fresh 
tissue, exclusively to the surface of membranes or membranous 
tissues of slight thickness (aponeurotic plates, substance of the 
cornea, intima of vessels) and light exerts an important influ- 
ence upon the reaction whereby the blackening of the combina- 
tion which the silver salt forms with the ground substance is 
brought about. With my method the light has nothing to do 
and the reaction takes place through the gradual penetration of 
the silver salt into more or less voluminous pieces which have 
been previously treated with bichromate. The black-staining 
of the various elements composing the nervous tissue results 
froma reducing action which the elements themselves exert, 
under the influence of the bichromate, upon the silver salt. 

‘‘The procedure necessary to bring about the black-stain- 
ing of the elements of the central nervous system consists essen- 
tially of two parts. 


StronG, Review of the Golgi Method. 103 


‘‘(a) Hardening of pieces in a solution of potassium bt- 
chromate. 

‘<(b) Immersion of the hardened pieces in a solution of silver 
nitrate. 

‘‘ (a) Hardening in bichromate. Although there are no es- 
pecial rules for the hardening other than those which must usu- 
ally be followed to obtain a good uniform hardening, yet it is 
this part of the process which requires the most care. This is 
the more so because the time necessary to harden the pieces to 
the degree required for the action of the second reagent varies 
very considerably according to different circumstances and espe- 
cially according to the temperature. 

‘« For the first immersion of the pieces, I use either a sim- 
ple two per cent. solution of potassium bichromate or the usual 
formula of Miller. (The reagents should be pure.) There 
must be an abundant quantity of fluid in proportion to the quan- 
tity of pieces to be hardened. 

‘The part of the brain or spinal cord to be treated is cut 
into tolerably small pieces (about 1 to1% ccm.). It is important 
that the pieces be fresh; the fresher the pieces, the better the 
results. It is well to use, preferably, the brains of animals just 
killed, yet satisfactory results can also be obtained 24 to 48 hours 
after death. It is hardly necessary to say that the pieces must 
be cut regularly and in definite directions (according to the part 
to be studied) so as to permit orientation as to the part and the 
location of, the elements in the future study. 

‘« That the hardening may proceed with some rapidity and 
be uniform, it is well to successively increase the concentration 
of the fluid, raising the quantity of bichromate from 2g to 2%, 
3, 4and 5%. 

‘Whether the fluid is increased in strength in hardening 
the pieces, or remains the same strength, it is always necessary 
to change it from time to time to avoid the formation of moulds 
which, as is well known, develop abundantly in bichromate so- 
lution when the pieces are to some extent neglected. For the 
same reason it is advantageous to place in the vessels with the 
pieces a small quantity of some substance which will prevent 


104 JOURNAL OF COMPARATIVE NEUROLOGY. 


the growth of hyphomycetes, as camphor, salicylic acid, ete. 
The most important point and at the same time the most diff- 
cult to determine in order to obtain good results with this meth- 
od is the length of time during which the pieces must be kept 
in the bichromate solution before one passes on to the second 
part of the process, the reaction with the silver nitrate. 

‘‘The proper duration of immersion for the pieces to ob- 
tain that degree or particular kind of hardening which is best 
fitted to secure, when they are laid in the silver solution, a fine 
and diffused action upon the various elements of the nervous 
system varies according to various conditions. These are the 
strength of the fluid, the condition of the pieces, the quantity 
of fluid, temperature and, consequently, the time of the year. 

‘‘ The differences arising from the strength and quantity of 
the fluid may be eliminated by paying strict attention to the 
strength of the fluid, by using covered vessels and preserving 
the same ratio between the number of pieces and quantity of 
fluid. 

‘The influence of temperature upon the results of the re- 
action is more important, indeed practically all the uncertainties 
of the method depend upon this. For example, to mention 
extremes, good results (which, with the progressive changes, of 
which I shall speak later, continue to appear and extend) can be 
obtained in the warm season after an immersion of 15 to 20 days 
and seldom after 30 to 40 or 50 days; on the other hand in the 
cold season good results are scarcely obtainable after,an immer- 
sion in bichromate of less than 1 to 1% months. The reaction 
(with the progressive accompanying changes) may then con- 
tinue to manifest itself for 2, 3 or 4 months, provided, ‘of course, 
the pieces are preserved according to the rules given above. It 
is almost superfluous to say that during the gradual change from 
the warm to the cold season and vice versa corresponding 
changes in the appearance of the reaction take place. It is not 
easy to remedy these temperature changes, especially because 
these changes of environment are united with the other causes 
of uncertainty mentioned and so act that observations made 
upon one series of pieces, never agree closely with those made 


STRONG, Rewew of the Golgt Method. 105 


upon another series. A warm chamber, of which I shall speak 
later, cannot bring about the accuracy sought for. 

‘‘The surest means of remedying these inconveniences is 
the persevering repetition of the process, i. e. one must have a 
good number of pieces available, bring several from time to 
time into the silver solution and then ascertain whether they are 
in the desired condition. If a good reaction has taken place, 
one continues the trials at regular intervals in order to obtain 
all the stages of the reaction, which constitute an advantage of 
this method. It is self-evident that the different trials must fol- . 
low each other at intervals differing according to the time of the 
year. Inthe warm season, when the requisite hardening is 
reached much earlier, the trials must follow each other more 
quickly ; in the cold season, on the other hand, when the de- 
sired hardening is first reached after a month, the trials can be 
made at intervals of 8 to 1o days beginning with the time when 
one, according to my direction, has ground to assume that the 
tissue has begun to enter the desired condition. 

‘© (6) Transference of the hardened pieces into the solution of 
silver nitrate. Although the various conditions of which I have 
spoken make it impossible to state with complete accuracy for 
how many weeks or days the pieces must be brought from the 
bichromate into the solution of silver, this is no ground for con- 
cluding that the method is subject to excessive uncertainty. All 
difficulties are overcome and one can be absolutely sure of al- 
ways obtaining excellent results by the simple procedure of 
steadily extending the trials with every series of pieces. The 
difficulties are thus very like those which one encounters in the 
employment of all other impregnation and imbibition processes, 
not excepting the simple carmine staining, in which, as is well 
known, one only reaches quick and certain results after repeated 
trials when he has learned to know the nature of the staining 
fluid and of the pieces to be stained. 

‘‘T usually employ a 34 % silver solution; yet I will remark 
that it is not necessary to adhere closely to this formula to ob- 
tain the reaction. A slightly stronger or weaker solution does 
not affect the result. I will also-add that a slightly weaker so 


106 JOURNAL OF COMPARATIVE NEUROLOGY. 


lution (%%) appears to be somewhat more suitable (giving 
finer results though confined to fewer elements) so long as the 
pieces have not yet reached the complete hardening, while a 
slightly stronger solution (to 1%) appears better adapted for 
pieces whose hardening has progressed a little too far. 

‘« The quantity of the silver solution to be used must vary 
with the number and size of the pieces to be laid therein, but 
must be relatively abundant. For two or three pieces of about 
1 ccm. I use about half a beaker (dzcchzere) of the fluid. 

‘‘The moment the pieces are brought from the bichromate 
into the silver solution a copious yellowish precipitate of silver 
chromate results. The formation of this precipitate takes place, 
of course, at the expense of the strength of the fluid inasmuch 
as through the formation zz /oco of the insoluble precipitate a 
more or less considerable portion of the silver salt is deposited. 
This changes, naturally, the relation (osmotic as well) between 
the fluid which should penetrate into the piece and the inner 
portions of the piece. It might happen that the whole or the 
greatest part of the silver would be precipitated from the solu- 
tion, which would result in the more or less complete absence of 
the reaction. To avoid this mishap it is expedient to first wash 
the pieces in which the reaction is sought in a weaker solution 
of silver. I use for this purpose, from motives of economy, 
silver solutions which have already been used on other pieces 
without the silver having been fully neutralized. When this 
washing has been continued until the pieces cause no more pre- 
cipitate when brought into a clear solution’ they are finally 
placed in the fluid of the proper strength. From there on the 
preparation usually requires no especial attention, for if the 
solution is present in copious quantity it is sufficient to let the 
fluid penetrate into the interior of the piece. Yet it is well to 
consider that it is sometimes expedient, with pieces thoroughly 
saturated with bichromate through a long sojourn therein to 


1Several minutes should elapse to test this, inasmuch as the discoloration of 
the silver solution by the reddish precipitate sometimes takes place rather slow- 
ly, both in this and in the rapid method.— Writer. 


STRONG, Review of the Golgi Method. 107 


change the solution for a fresh one after the pieces have been 
in the first solution 6 to 8 hours. This must be done whenever 
the fluid assumes a yellow color, which shows that the silver 
nitrate is neutralized. In this case the reagent can no longer 
possess the necessary strength to penetrate to the interior of 
the pieces. 

‘‘T have already said that this reaction, through which 
the black staining of the elements is brought about, has nothing 
in common with that which stains the intercellular substance 
under the influence of light. I now need to add that it is en- 
tirely the same whether the pieces in our method are kept in 
the light or in the dark; the reaction which is brought about 
through the gradual penetration of the silver into the interior of 
the tissue takes place equally well in both cases, The only rule 
relating to keeping the pieces in the silver which experience has 
shown to be in some manner useful is that they should be kept 
in winter in a well-heated room. I place the vessel on a table 
which is not far from the stove of the laboratory. 

‘«The pieces must remain, asa rule, in the silver solution 
for 24 to 30 hours, in exceptionable cases 48 hours. The 
period of 24 to 30 hours must form the rule although, 
when the time of hardening has been correctly hit upon, the 
reaction may be well advinced in 2 to 3 hours. In such 
cases one may say that the reaction begins immediately, at 
least in the superficial layers, to extend gradually deeper 
with the deeper penetration of the fluid. In the exception- 
al cases when it is best to leave the pieces 48 hours and 
longer in the nitrate solution and where it is well to change the 
solution a second time, one must regulate his procedure by the 
results of a microscopal examination of some superficial sec- 
tions from which the condition of the reaction may be inferred. 
Moreover one can perceive from the yellowing of the fluid, 
whether the reagent is nearly neutralized. 

‘« As for the rest, it is to be remarked that an indefinite 
sojourn of the pieces in the silver solution lasting days, wecks 
or even months is in no way injurious to them; on the contrary 


108 JOURNAL OF COMPARATIVE NEUROLOGY. 


this is a suitable means of preservation for pieces destined for a 
particular investigation of long duration. 

‘‘One of the most interesting peculiarities of the process 
which I here describe consists in the fact that, while the brown- 
ish black stain acts quite similarly upon all elements of the 
nervous tissue (various kinds of ganglion cells, nerve-fibers, 
elements of the neuroglia and walls of vessels), yet in reality 
the staining of all these at one time forms an exception, i.e. 
when the elements are in a certain state of hardening which one 
only happens upon accidentally in a great number of trials. As 
a rule the reaction appears only partially, i.e. it affects only one 
or another layer with gradations and combinations which one 
may term endless. 

“This peculiarity does not detract from the method, but 
is rather among its advantages, for if the reaction affected all 
kinds of elements at the same time there would evidently arise 
such an inextricable confusion that it would be impossible to 
orient oneself in respect to the locations and relations of the in- 
dividual parts. When, for example, in one preparation the 
the cells especially are stained black, in another principally 
the neuroglia together with the vessels and some groups of 
nerve cells, it is evident that one can by the comparison of 
many preparations obtain a general view of the various pecu- 
liarities of the arrangement and relations to each other of the 
individual species of elements and of the connection of the 
structures of various regions. y 

‘‘ This is so much the more the case since these combina- 
tions and gradations also appear in certain layers and different 
zones into which one is accustomed to divide different regions 
of the nervous system. In the cortex, for example, the reaction 
appears, with the various combinations above mentioned, some- 
times in the superficial or middle, sometimes in the deep layers. 

«© A law undoubtedly exists governing the manner of de- 
velopment of the black stain and the succession of the reaction 
among the various kinds of elements and it would be interest- 
ing to learn to know this so.as to be able to bring about one or 
another result at will; but it is extremely difficult, if not im- 


STRONG, Review of the Golgi Method. 109 


possible, to attain this. This difficulty will be readily compre- 
hended when one reflects that the diversity of results is brought 
about not only by the conditions given already but also by the 
unequal hardening action of the bichromate so that the individ- 
ual layers of the pieces are in different conditions. In the 
individual pieces the degree of hardening may increase from 
center to periphery so that a number of the above combina- 
tions and gradations may appear in one piece. 

‘‘The following approximate rule, however, may be accept- 
ed for the way the reaction enters the various elements of the 
nervous tissue when a number of similar pieces are successively 
subjected to the action of the silver nitrate. There stain in the 
following order: 

“1, The bundles of nerve fibers. At the same time with 
the staining of these fibers some scattered ganglion cells which 
lie dispersed in the gray matter appear. 

‘‘The staining of the nerve fibers at the beginning shows 
little delicacy, the reaction being, so to speak, tumultuous, but 
gradually gains in fineness with progressive hardening (always, how- 
ever, after a more or less brief period of time). Then the individual 
fibers (axis-cylinders) composing the bundles can be well seen 
and also individual fibrillae streaming from the bundles, the finest 
details of whose course and branching can be seen at a glance. 

‘°2. The ganghon cells. VYhe ganglion cells of the su- 
perficial layers always stain first (e. g. in the cortex the small 
cells of the peripheral zone), but at the same time with them 
also some cells irregularly scattered in the inner layers. As the 
reaction progresses it affects the cells rather than the fibers 
and the tendency is for the stain of the cells to become more 
general and to extend from the periphery inwards. Then, 
too, while the reaction is becoming more complete among the 
cells of the deeper layers it becomes always more limited among 
those of the superficial layers. 

‘*With the cells as with the fibers the reaction is at first 
coarse and little fitted to bring to view certain interesting de- 
tails. For example, the nervous process is not stained at first 
to any great extent and usually only a short piece of it is to be 


110 JOURNAL OF COMPARATIVE NEUROLOGY. 


seen so that neither its course, direction nor its few or numerous 
branches can be perceived. With the gradual progress of the 
reaction the nerve cells are displayed more clearly and the finest 
subdivisions of their protoplasmic as well as their nervous pro- 
cesses appear. 

“©3, Cells of the icurogha. An interesting reaction oc- 
curs in the cells of the neuroglia; it may be said that it takes 
place in pieces suitably hardened in bichromate from the begin- 

ning of the phase to the end. In fact at both the time when 
~ the fibers predominate and when the cells predominate individ- 
ual neuroglia cells or groups of them are to be seen showing the 
characteristic reaction of the silver nitrate (coffee-brown or yel- 
_lowish). Besides, with this species of element the reaction only 
becomes fine and diffused in a somewhat advanced period of 
hardening so that their typical form and relations are plain. 
The reaction in neuroglia cells takes place for a long time be- 
yond the time favorable for staining nerve cells. 

The finest reaction for the nerve cells, especially for the 
nervous processes, occurs at a somewhat advanced stage 
of hardening, namely when, with the advance of the re- 
action among neuroglia cells, it is limited among the gan- 
glion cells. It is precisely among isolated blackened cells 
that the stain of the individual functional (axis-cylinder) 
processes is finest; one can observe the smallest details of 
their course and branching. I must again recall that the reac- 
tion must be produced in a series of pieces which have consecu- 
tively received suitable treatment in order to learn to know all 
its phases. 

‘« After we have so circumstantially laid down the funda- 
mental rules of procedure, it would be superfluous to go into 
particulars about the differences obtaining between the different 
provinces of the central nervous system (the cortex cerebri, the 
so-called ganglia of the base, the cerebellum, the spinal cord). 
I only remark here that, under similar conditions, pieces from 
the cortex reach in bichromate the suitable state of hardening 
somewhat sooner that than those from the cerebellar laminae, 
that the latter reach it a little later than pieces of the spinal cord 


StronG, Review of the Golgi Method. III 


and that finally the so-called ganglia of the stem reach the prop- 
er hardening still somewhat later than the parts named. 

‘‘A last remark. When the above-described peculiarities 
of the process are considered, it is intelligible how it often hap- 
pens that the reaction appears only in one part of the piece. 
For example, it is absent in the superficial layers, where there 
is, aS a matter of fact, more often than otherwise only an irregu- 
lar precipitate, and is present in the interior or vece versa. One 
must remember this and when, very likely, the first sections 
made near the surface show nothing of interest one must not 
thereupon conclude that the reaction has failed, for it often hap- 
pens that such preparations, in which only single, isolated cells 
are stained, are among the most instructive for details of the in- 
dividual elements. 

“« Tyeatment and Preservation of Preparations. Whether 
the black stain has turned out so that the piece is worth keep- 
ing for further investigation can be ascertained by means of trial 
sections examined in glycerine or in the reaction fluid itself. 
Then one must provide for the preservation of the piece and the 
microscopical sections. Although it is certain that a longer so- 
journ in the silver solution does no harm whatever and that such 
a sojourn may serve as a means of preservation, yet is expedi- 
dient, in order to have the pieces ready for further treatment, to 
transfer them to pure commercial alcoho]. This not only serves 
to harden the tissue farther but also to free it from the silver 
nitrate which, as I shall mention below, is very injurious to the 
preservation of the microscopical sections. To accomplish the 
latter the alchol should be changed two or three times till it re- 
mains transparent for a number of days after the piece is brought 
into it. In this way the pieces can be kepta longtime. I 
have kept them for about nine years in this way and can obtain 
from them, when I wish, preparations as clear as those obtained 
from them shortly after their preparation. 

‘«The further treatment of the microscopical sections cor- 
responds essentially with the usual procedure in obtaining 
anhydrous preparations except for some peculiarities necessary to 


112 JOURNAL OF COMPARATIVE NEUROLOGY. 


overcome some difficulties in the way of securing stable 
preparations. 

‘«The sections, before they are permanently brought into 
gum damar or Canada balsam, must first be treated successively, 
according to the classical method, with absolute alcohol and 
some clearing fluid. Each of these steps requires an especial 
care not necessary with ordinary preparations: 

‘*(a) Treatment with absolute alcohol. The only rule to be 


' especially noticed here is that the sections must be very care- 


fully dehydrated by bringing them into three or four changes 
of pure absolute alcohol. This is the only principal rule in or- 
der to obtain a long preservation, for the more accurately and 
carefully the dehydration is carried out, thereby freeing the 
tissue from the last trace of silver nitrate, the more one can rely 
upon the preparations remaining clear a long time. 

‘« (6) Clearing. The sections to be mounted must first be 
brought, for clearing, from absolute alcohol into creosote, 
where they remain some minutes, and then into turpen- 
tine. In the latter they can remain a long while. The selec- 
tion of these two substances and their consecutive use is another 
aid to securing a long preservation. Among many other sub- 
stances tried for clearing I have also found oleum origani' for 
my method very useful, but I have found no sufficient ground 
for abandoning the first mentioned fluids. The sections usually 
remain in turpentine only 10 to 15 minutes but may remain 
there longer. 

‘«(c) Completion of the microscopical preparations. For per- 
manent preservation the sections are brought from turpentine 
into damar which, after many comparative tests, I have found 
better adapted for this purpose than Canada balsam. I must 
here call attention especially to a peculiar treatment of the sec- 
tions ; contrary to the usual custom, I do not cover the prepar- 
ations with a cover glass. When the sections are covered in 
the usual way with a cover glass, they begin after a time to turn 


‘This oil, followed by washing in xylol instead of turpentine, is pre- 
ferred by the writer. 


StronG, Review of the Golgt Method. 113 


yellow (owing toa second impregnation which takes place), 
then the outline of the stained cell elements become obliterated, 
the whole tissue becomes opaque and, after a period of from 
two or three months to two years, the preparations, with few 
exceptions, become useless. On the contrary they may keep 
a long time thanks.to the repeated washing, of which I have 
spoken, and especially to the mode of mounting without a cover 
slip ina layer of damar. I can now state that the earlier la- 
mentable disadvantage that preparations made by my method 
soon spoiled is now almost completely remedied. I have many 
preparations made by me nine years ago which have not yet lost 
their original clearness. 

‘‘Tf the good preservation appears menaced by an incipi- 
ent yellowing, another longer bath, on the slide, in turpentine 
will restore transparency and freshness to the preparation. 

‘‘T have found it convenient to employ for this kind of 
mounting a special wooden slide with a square opening in 
which, by means of a groove, a glass plate (a cover slip of 
somewhat greater diameter than usual) is fitted and stuck fast 
with a solution of shellac in alcohol. This serves as a slide and 
the section adheres to it by means of the damar. 

‘‘This kind of slide not only enables the section to be ex- 
amined from both sides but also has the advantage of prevent- 
ing dust from fouling the object, to which this kind of mount 
would be especially exposed. To accomplish this it is only 
necessary to turn the side of the slide with the section down- 
wards as soon as the damar is hard enough, or to pile the prep- 
arations on top of each other. 

“‘T further remark that it is wise to shield the objects 
from the influence of light ; still this precaution is not entirely 
necessary if the repeated washing has been carefully performed. 
After fulfilling these conditions, I might expose preparations 
for days to the sun’s rays without injury to them. 

‘‘ This is not the place to lay stress upon the value of the 
results which can be attained by means of this method. The 
figures accompanying this work demonstrate it sufficiently. 
They display the forms to be observed in the preparations with 


114 JOURNAL OF COMPARATIVE NEUROLOGY. 


a fineness not only not exaggerated but inferior to the natural ob- 
ject. I will here only bring forward the disadvantages of the 
method, in order to give the means by which they are to be 
avoided, The long time between the placing of the pieces in 
bichromate and the appearance of the reaction (it not infre- 
quently happens that in consequence of this the pieces are for- 
gotten), the uncertainty about the extremely variable time re- 
quired to reach the proper hardening, the different conditions 
in which individual layers of the same piece are found, all these 
are disadvantages whose removal would be desirable. 

‘“‘T have sought by expedients to change my method in 
one way or another in order to secure greater certainty and ac- 
curacy in the results. Among the means tried by me I pre- 
sent the following which have yielded me a certain advantage. 

‘«(a) Injections of bichromate (solution to 2% %.)' It must 
be abundantly and constantly applied so that the whole paren- 
chyma of the part to be investigated is fully and uniformly 
penetrated by the hardening fluid. The fixation of the elements 
by the reagent, where possible, before the slightest post mortem 
change can take place is of the highest importance in securing 
a very delicate reaction. The action of the injection consists 
principally in giving a uniform hardening, furthermore in pre- 
venting, very likely, a slight post mortem change in the in- 
terior of the piece and finally in abbreviating the sojourn in 
bichromate. 

‘Tf ITmay draw a conclusion from some especially suc- 
cessful reactions accomplished in this way, I must declare that 
the injection is in these various respects actually of consider- 
able advantage. Some other experiments, not yet very ex- 
tended, have convinced me that a favorable influence is exerted 
in the same way by injecting, not a simple solution of bichro- 
mate, but one with gelatine added (244% bichromate, 100 cc. ; 
dry gelatine, which is dissolved in the usual way, 5 to 6 grams). 
This procedure appears to me especially fitted to give the 


! A stronger solution would probably be better, inasmuch as it undergoes 
dilution in the tissue.— Writer. 


STRONG, Review of the Golegt Method. 115 


pieces in less time that particular hardening most favorable to 
the best reaction with silver nitrate. I mention, for example, 
a case where I have obtained graduated reactions of surprising 
fineness on pieces 15 to 30 days after they were placed in bi- 
chromate at a temperature of 15° to 20° C. (in autumn), the 
pieces having been subjected to the above treatment. 

‘The injection is performed in the usual way (with a sim- 
ple syringe or with a siphon in which the pressure is regulated 
by the height of the vessel containing the injection fluid) either 
through the carotid, when one wishes to limit the hardening to 
the cerebrum and cerebellum, or through the aorta when the 
fluid should also extend to the spinal cord. 

‘‘It is superfluous to state that when the bichromate and 
gelatine is injected it must be warmed so that it will remain 
fluid. In this case it is especially important to perform the 
operation immediately after the death of the animal, before the 
tissues are cold. Only in this way does one secure the finest 
and most widespread injection. 

‘‘ After the injection the nervous parts are removed from 
their cavities, cut into pieces and brought as usual into bichro- 
mate where they are carefully treated as dealt with above. 

‘“(6) Hardening in bichromate at a constant temperature. The 
circumstance, pointed out several times, that the uncertainty 
about the time at which the pieces must be brought from the 
bichromate into the silver solution depends for the greater part 
upon the temperature of the medium leads to the idea that the 
best means of avoiding this inconvenience would be the em- 
ployment of a constant temperature for the bichromate in 
which the pieces lie. For this purpose the warm chambers used 
in investigations upon micro-organisms seem best adapted. 

‘‘T have used the chamber of Wiesnegg in which I main- 
tained a temperature of 20° to 25°. This had good success but 
only in the direction of considerably abbreviating the period of 
hardening in bichromate so that the reaction could be obtained 
much sooner than formerly and in a tolerably constant period 
of time. Thus the reaction in the warm chamber appeared 
after 8 to 10 days and proceeded to completion up to 15 to 20 


116 JOURNAL OF COMPARATIVE NEUROLOGY. 


days. This is, perhaps, an advantage in so far as one can with 
sureness obtain certain preparations for demonstrations in a tol- 
erably brief time. But the advantage is not extended to the 
fineness of the result since in all such preparations the reaction 
turns out rather coarse. I was not thereby encouraged to ex- 
tend experiments in this direction, especially as the abbrevia- 
tion of the time can be attained in other simpler ways and as 
the pieces in the chamber quickly pass by the period favorable 
to the success of the reaction without attaining the kind of 
hardening sought—which is a not insignificant disadvantage. 

‘‘(c) Hardening in Erlicki’s fuud (bichromate of potassium, 
2% g.; copper sulphate, % g.; distilled water, 100 g.). Re- 
garding this I confine myself to stating that the copper salt 
added to the bichromate did not prevent the reaction and that 
the Erlicki’s fluid possessed the same advantages and disadvan- 
tages as the preceding method (warm chamber). It accelerates 
the hardening so that ina few days (6 to 8 to 10) the black 
stain of various elements of the nervous system can be ob- 
tained by transferring to silver, but the result cannot be com- 
mended for fineness. Moreover the period advantageous for the 
reaction is very quickly passed over. 

‘« As it appeared to me that the limited and not very fine 
form of the reaction might be due in part to the rapid action of 
the hardening fluid, I weakened the same by mixing it in grad- 
ually increasing quatities with Miiller’s fluid (Erlicki 20% to 
50%, Miller 80% to 50%). The results obtained by means of 
this variation were decidedly good. After only 5 to 6 to 8 days 
immersion in such a fluid I obtained preparations which in re- 
gard to fineness of result had a certain worth. It thus appears 
to me that this variation can be recommended for the purpose 
of quick demonstrations of cell-forms. For the finest details, 
especially the relation of the functional processes of the gang- 
lion cells and the nerve fibers, I find that the first procedure is 
always to be preferred, or also the following : 

‘62. Method of the successive actions of a mixture of osmic 
acid with bichromate and of the silver nitrate. This procedure 
also is only a modification of the original but deserves a place in 


SrronG, Review of the Golgt Method. 117 


the exposition as a method by itself, partly because the not unim- 
portant changes of the results which it yields and the treatment 
which it requires are to be ascribed to the newly added re- 
agent, partly because the process so modified can remedy some 
inconveniences of the original method. 

‘«Tt can be applied in two ways, namely : 

‘‘(a)_ By laying small pieces of nervous tissue directly in 
a mixture of bichromate and osmic acid (2% to 2%% sol. of bi- 
chromate, 8 parts; 1% sol. of osmic acic, 2 parts). 

‘“‘The black stain is obtained the most quickly with this 
procedure. The black staining of a great number of nervous 
elements can be obtained by transferring into silver at the second 
or third day (see the directions for procedure in the description 
of the original method). The reaction extends itself on the im- 
mediately following days, then, as usual, diminishes and at the 
tenth or twelfth day entirely ceases. 

‘‘The treatment of the macro- and microscopical prepara- 
tions which are obtained in this way must be considerably modi- 
fied. Pieces prepared by this method differ trom those prepared 
by the first method inasmuch as when they are kept a long time 
for future use, they become diffusely blackened and thereby use- 
less. They must be kept in the same silver solution which has serv- 
ed for the reaction. Then they are brought into pure alcohol, 
which must be changed, where they remain not longer than two 
days, sectioned and subjected to the above described treatment 
(absolute alcohol with repeated washing, creosote, turpentine, 
damar) necessary for their permanent preservation as micro- 
scopical preparations. 

‘Although this application of the osmium-bichromate 
solution is certain and, as far as fineness is concerned, yields 
satisfactory results; yet I find that for a systematic study of any 
definite portion of the nervous system the following method is 
far preferable: 

‘““(b) Bringing of fresh pieces into the bichromate solution ; 
first transference into an osmium-bichromate solution ; second trans- 
ference into the silver solution. It is different with this second 
procedure than with the preceding, in which the series of pieces 


118 JOURNAL OF COMPARATIVE NEUROLOGY. 


of tissue to be examined are useless after a few days. Here the 
fresh pieces (with or without injection) are laid in the bichro- 
mate solution and remain, so to speak, in the hand of the inves- 
tigator. They can either immediately or later be tried i. e. 
during a period of from 3 or 4 to 25 or 30 days after the immer- 
sion. If one during this whole period transfers at intervals of 
2 to 3 or 4 days some pieces into the osmium-bichromate solu- 
tion, he thus possesses many secondary series of pieces which 
are brought singly (1 or 2 at a time) into the nitrate solution. 
These, from the third or fourth to the eighth or tenth day of 
their sojourn in the mixture, yield with certainty, when brought 
into the silver, preparations with all the consecutive gradations 
and combinations described in the original method and also 
possessing surprising fineness. 

‘“‘After-treatment. Preservation of the pieces in the silver 
solution; pure alcohol for 2 or 3 days, till one has time to un- 
dertake the examination; repeated washing out of the sec- 
tions with absolute alcohol; cresote, turpentine, damar, mount- 
ing without cover glass. 

‘This is the method which I at present prefer for the 
demonstration of the finest details in the structure of the central 
nervous system. The particular grounds for this preference are 
the following: (1) Certainty of obtaining the reaction in many 
gradations, if one makes use of a certain number of pieces. 
(2) The considerable length of time during which one can 
obtain the reaction—while one can also attain it in a few days. 
This renders an accurate investigation much easier. (3) The 
pieces are much more conveniently treated. (4) Finally, one 
obtains at the same time with the gentle gradation of the results 
also a greater fineness of the same, especially regarding the be- 
havior of the functional processes of the ganglion cells. 

“©3. Method of the consecutive actions of the bichromate 
of potassium and of bichloride of mercury. This can likewise 
yield valuable results whose value is not diminished because 
they in many respects conform to those obtained by the silver 
nitrate. Indeed, the particular purposes it can fulfil and_ its 
peculiar advantages are in and for themselves so important that 


SrroneG, Review of the Golgt Method. 1@ fe) 


it must be given a place of its own alongside the silver nitrate 
method. The clearness with which the various elements of the 
nervous system emerge in this reaction is not less than that 
brought about by the silver nitrate. The elements appear, 
when viewed under the microscope by transmitted light, com- 
pletely black after the action of the sublimate and for micro- 
scopal investigation the action is the same as when there is an 
actual black stain. But this stain is only an appearance due to 
the opacity of the elements upon which, probably owing toa 
reducing action, the mercury has precipitated. In reflected 
light one notices that the elements appear entirely white, indeed 
under stronger magnification they show plainly a metallic luster. 

“‘T will remark that the particular advantages of this 
method consist first in the fact that the reaction can take place 
in large pieces, further that its success is absolutely certain with- 
out being necessarily bound by strict rules as to the time of 
sojourn in the hardening fluid, and finally in the fact that the 
preparations which it yields require no especial precautions for 
their preparation, but can be treated in the usual way, like sec- 
tions stained with carmine. 

‘The mode of application of the sublimate method is only 
distinguished from the silver method by some unessential things. 
It likewise consists of two essential processes: 

‘‘(a) Hardening of the pieces in bichromate. 

‘‘(b) Transference of the same into a solution of bichlo- 
ride of mercury and sojourn in the latter. 

‘¢(a) The hardening in bichromate is done entirely in the 
usual way. (See the original method).: I only add that the 
reaction does not proceed in an essentially different manner 
if consecutively stronger solutions of I, 2, 3% are employed or 
if the pieces are immediately laid in Miuller’s fluid. In general 
it is expedient for the pieces to be small but this is not abso- 
lutely necessary. Good results are also obtained with large 
pieces, indeed with whole brains. In the latter case the pre- 
serving fluids require a long time to pentetrate by osmosis from 
the periphery into the interior and the central portions could 
spoil before they experienced the action of the fluid. It is nec- 


120 JOURNAL OF COMPARATIVE NEUROLOGY. 


essary therefore to make a careful preliminary injection of 
bichromate solution so that the reagent is well distributed 
throughout the organ. 

‘‘A few days (6 to 8 or less) sojourn in the bichromate 
solution is sufficient to obtain, by putting the pieces into subli- 
mate solution, an extended fine black stain of a greater or less 
number of cells (indeed one can obtain an indication of the reac- 
tion on the fresh brain which is placed immediately in the subli- 
_ mate solution). A more suitable period to obtain fine and 
extended results is from 20 to 30 days. A much longer hard- 
ening (from 2, 3, 4 months or more) is by no means unfavorable 
for the reaction. I remember, among other cases, to have 
obtained reactions of wonderful fineness in some whole brains 
which were in bichromate solution nearly a whole year. 

‘Tt will be perceived that this indefiniteness of the time 
constitutes a very advantageous circumstance since thereby 
pieces can be employed which would otherwise be useless. 

‘«(b) Transference of the pieces into the sublimate solution. 
The solution used by me contains %% of bichloride of mer- 
cury. I have satisfied myself that the method is equally suc- 
cessful when the solution is weaker (144%) or stronger (1%). 
The pieces are brought immediately from the bichromate into 
this solution. 

‘« The reaction throughout the thickness of the piece results 
much more slowly than with the silver nitrate. If the pieces 
- are suitably hardened, 24 to 48 hours suffices with the latter. 
With the sublimate, on the other hand, not less than 8 to Io 
days are necessary, in order that the reagent may penetrate 
throughout the piece, when the pieces are small and much more 
(2 months and upwards) when the pieces are large (whole 
brains). The period of action of the bichromate must also be 
considered ; the longer this has been, the longer must be the 
sojourn in the sublimate, but the more complete and delicate is 
the reaction. 

‘‘During the sojourn of the pieces in the sublimate solu- 
tion, the bichromate with which the tissue is saturated diffuses 
out and impairs the purity of the fluid, which assumes a yellow 


StrRonG, Review of the Golgi Method. 121 


color while the pieces become paler. For this reason the sub- 
limate solution must be changed daily, especially at the begin- 
ning of the immersion. Later the changes are made only when 
the solution becomes yellow. 

“‘It may be assumed that the reaction begins when the 
pieces are entirely decolorized, i. e. when the tissue is complete- 
ly freed from bichromate. If, beginning about this time, sec- 
tions are made and examined under the microscope daily, it 
will be noticed that the first traces of the reaction begin 3 or 4 
days after the immersion and that they can be known bya 
number of small black spots scattered here and there. After 
4 or § days more one sees the cell-forms gradually become more 
complete and numerous and the reaction thenceforward con- 
tinues to extend and complete itself. It even appears that 
further advantages are gained when the sojourn in the sublimate 
solution is extended indefinitely, the sublimate being changed as 
often as it becomes yellow through the presence of bichromate. 
With brains which have been long exposed to the action of the 
bichromate,——and such often yield the most beautiful results,— 
the sublimate solution must be changed during several months 
before this yellowing ceases. 

‘«The above constitutes a further difference from the manner 
of action of the silver nitrate, inasmuch as in the latter the 
whole action is completed in 24 to 48 hours, after which no 
further action is exerted, although the pieces can be kept in 
it longer. 

‘When the reaction has reached its maximum, the pieces 
remain colorless and have the appearance of fresh brain tissue 
which has been slightly washed in water. 

‘«The pieces may remain in the sublimate solution as long 
as one pleases, not only on account of the possibility of a fur- 
ther extension of the reaction but also because they thereby 
receive a hardening better adapted for making fine sections. 

‘« As to the manner in which the reaction extends to the 
different elements, I will merely remark that the reaction affects 
the ganglion cells in pieces which have reached that degree of 
hardening attained in the first month’s immersion in bichromate 


122 JOURNAL OF COMPARATIVE NEUROLOGY. 


and the reduction only extends itself gradually to the nerve 
fibers also. The reaction displays itself to the fullest extent in 
the nerve fibers almost exclusively in pieces which have lain 
a long time in bichromate and are very strongly hardened. 
I recall in this connection the brains which had been kept very 
nearly a year in bichromate: they showed an almost universal 
very fine stain of the bundles of nerve fibers and of their finest 
subdivision. 

‘* Treatment and preservation of microscopical preparations. 
The only special precaution required by preparations made by 
means of the sublimate reaction before they are mounted in 
glycerine or balsam is a careful washing in water. Without 
this precaution a precipitate in the form of a black powder or 
needle-shaped crystals is formed in the sections some days after 
mounting and if it does not entirely spoil, yet seriously mars 
them. As to the rest, the usual mode of preparation is employ- 
ed: mounting in glycerine, damar or Canada balsam after the 
necessary dehydration in absolute alcohol and clearing in -creo- 
sote or clove oil. No further precaution is necessary. 

‘«When I described this method the first time’ I expressed 
the conviction that it could be still further perfected so as to 
yield finer results than those hitherto attained by me. Practice 
has later led me to some modifications which have improved it. 
But it has experienced another important development owing to 
the perservering experiments of Dr. Mondino who succeeded in 
applying the process with remarkable success to nothing Jess 
than a whole human brain. I will here add the words them- 
selves in which this observer summarizes the advantages which 
one can gain from the use of the bichloride of mercury for the 
study of the central nervous system. 


ICAMILLO GOLGI, Di una nuova reazione apparentemente nera delle cel- 
lule nervose cerebrali ottenuta col bichluro di mercurio, Archivio per le Sc. 
med,. Vol. III. 


STRONG, Review of the Golgt Method. 123 


‘The following is Dr. Mondino’s Summary :' 

‘“«« A, The sublimate method is the first by means of 
which we can obtain the black stain of the nerve cells and their 
functional processes in the entire brain and enables us to follow 
these latter directly in their course through the brain. 

‘«« There is no doubt but that this technique fulfils the re- 
quirements of scientific accuracy better and puts us in a better 
position to obtain precise knowledge of the so much debated 
course of the fibers in the brain than all the methods hitherto 
tried. At the most one could only, with the aid of the latter, 
see whether numerous functional processes, collected into bun- 
dles, proceed in certain directions but with our technique one 
can examine them fiber by fiber and follow their anastomoses. 

‘««*B. In all other methods we must, in order to obtain 
consecutive series of brain sections, bring the individual sections 
into vessels with the staining fluid. “As one cannot provide so 
many vessels with fluid unless he possesses unusual means, sev- 
ral sections must be brought into one vessel and can therefore 
only be enumerated by groups and not singly. By the method 
here described this result can be attained with great ease. - 

“« °C. In the: other methods the sections: must’ be very 
thin and are liable to be torn in the various manipulations (from 
the microtome into the staining fluid, then to the slide, etc). 
As the sections are very thin they must also be much more nu- 
merous when a whole brain is sectioned ; hence greater expense, 
loss of time and more labor in making the preparations. In our 
method the sections need not be thin, they are therefore less 
numerous and exposed to fewer risks; whence little danger of 
losing sections, slight expense in the preparation and greater 
rapidity in the preparation of a whole brain. 

‘««D. Finally one must use in all other methods dyes, 
commercial and absolute alcohol and clove oil or turpentine, 
while we employ a little sublimate and creosote, which are 


IMONDINO. Sull’uso del bichloruro di mercurio nello studio degli organi 
centrali del sistema nervoso. Communic. fatta alla R. Accad. di Med. di 
Torino nella Seduta del 2 Genn., 1885, 


124 JoURNAL OF COMPARATIVE NEUROLOGY. 


very cheap and inexpensive. In the other methods we must 
use cover slips, because the high magnification which they re- 
quire—and then one does not see much—would not be applicable 
with the thick layers of damar. We do not require this and 


thereby escape not only expense but also the difficulty of 
avoiding bubbles of air under large coverslips whereby the pre- 
paration is often endangered.’ 

‘‘Tt appears to me, apart from all economy of material, 
time and labor, as well as the convenience of cutting pieces in 
the microtome so to speak at odd moments without injury to 
them from the long contact with water, that this method which 
enables us for the first time to follow in sections the course of 
nerve fibers through the whole brain shows an advance in the 
technique of the study of the central nervous system and takes 
precedence over all others. 

‘‘As I pass over the application for the macroscopical study 
of the brain which Dr. Mondino has also made of this method, 
I will here in conclusion again assert that the sublimate method 
takes a high place among the microscopical methods for the 
study of the nerve centers, alongside of the methods in which 
silver nitrate plays the chief role.” 

Additional technical notes in Golgi’s article, ‘‘Das diffuse 
nervose Netz der Centralorgane des Nervensystems. Seine phy- 
siologische Bedeutung ”’ (from the Rendiconti des R. Instituto 
Lombardo, Ser. II, Vol. XXIV, Fasc. 8 and 9), pp. 259 and 
260 of the German edition of Golgi’s works: 

‘‘The method which was most useful to me in the investi- 
gations described in the first part of this work, was the staining 
of the nervous elements with mercury sublimate, but with a 
modification which enhanced its demonstrative value without 
changing the fundamental procedure. The latter consists (1) 
in the hardening of the pieces in bichromate of potassium, 
(2) in the transference from this into a %% to 1% solution of 
bichloride of mercury. 

‘«Since I have given in another work (Studi sulla fina ana- 
tomia degli organi centrali del sistema nervoso, p. 202) a detailed 
description of what I call the fundamental part of the method, I 


Srrone, Review of the Golgt Method. 125 


consider it fitting to add that the best and finest reactions in the 
nerve fibers and the interstitial diffuse network was observed 
by me in pieces (from the spinal cord of the new-born kitten) 
which had lain a long time (in part over two years) in a one per 
cent. solution of sublimate after a long preceding sojourn ina 
bichromate solution (first, Muller's fluid, then, pure bichromate 
to 3%). Since they were pieces which had lain in the labora- 
tory in this way ready for examination but had not been used, I 
can, naturally, not tell what influence the long sojourn in the 
sublimate may have exerted. 

‘©The modification introduced by me, to which I must at- 
tribute a certain value for the clear demonstration of fine details 
and to which I call the attention of the observer, consists simply 
in a slight addition, viz., the blackening of the glistening white 
stain which the nerve elements receive by means of the mercury 
impregnation. 

‘‘ As is known, the elements treated with sublimate appear 
black in transmitted light on account of the opacity caused by 
the reaction but in reflected light they appear white. This dif- 
ference may be easily observed by turning off the mirror of the 
microscope. 

‘«This kind of appearance is satisfactory for observation 
with low or medium magnification, where less fine details are 
concerned, but it is otherwise with the finer details where 
stronger magnification is required. In this case the metallic 
luster of the fine parts, e. g. the finest divisions of the nerve 
fibers, evidently affects the observation unfavorably by giving 
the pictures a certain indistinctness. The black stain which re- 
places the white-metallic brings out better the outlines of the 
fibers and so increases the demonstrative value of the pre- 
paration. 

‘‘TInasmuch as the impregnation consists of metallic mer- 
cury, the transformation of the metallic white into deep black 
can be accomplished, according to the teaching of elementary 
chemistry, by means of a number of reagents. There can serve 
for this purpose: the sulphite and hyposulphite (particularly 
sodium sulphite and hyposulphite in 5% solution), the sulphide 


126 JOURNAL OF COMPARATIVE NEUROLOGY. 


(of potassium, sodium and ammonium, the first two in 1% to 
2%, the third in %% solutions), sulphuretted hydrogen (one 
part of the saturated solution and three parts of distilled water). 
One can also use with advantage the sulphocyanide (of potas- 
sium, sodium and ammonium in 2% solutions). 

‘« The solutions of sulphite and hyposulphite, especially the 
second, render necessary a careful watching of the preparations 
that they are not entirely destroyed through a disappearance of 
the metallic impregnation. 

‘The sulphide (of potassium and sodium) are easier to 
manage but the complete preservation of the preparations is not 
entirely certain with them. 

‘‘The sulphocyanide acts very well in bringing into view 
the smallest parts upon which the metallic impregnation has act- 
ed but it does not give a uniform black, only a brownish stain. 
Besides this the cells and fibers under the action of this reagent 
assume a punctate, almost pulverulent appearance. 

‘‘Sulphuretted hydrogen is very disagreeable on account 
of its offensive smell (a peculiarity which it has in common with 
ammonium sulphide) and it also has a tendency (as has the am- 
monium sulphide) to stain those parts containing no subli- 
mate brownish, which impairs very greatly the clearness of the 


preparation. 
‘‘ From all these grounds, and, particularly on account of 


its rapidity and certainty of action, on account of the intensity, 
uniformity and sharpness of the black stain obtained and on ac- 
count of the certain permanency of the preparations, the mix- 
ture used by photographers to stain and fix their pictures upon 
aristotype paper is to be preferred to all the other substances 
given here (naturally for the special purpose of the treatment of 
the sublimate preparations). 

‘‘From the many formulae of this kind which are found in 
books on photographic technique, I have adopted one which I 
repeat in a footnote.! 


For toning the two following solutions are separately prepared : 
(a) Water, 1 liter, 
Sodium hyposulphite, 175 g. 


StronG, Review of the Golgi Method. 127 


‘“«The modification which I have adopted with my subli- 
mate method is as follows: 

‘«The pieces which have been proven to be successfully 
impregnated are imbedded in celloidin in the usual way and 
cut with the microtome. The sections are then subjected to 
the following treatment : 

‘‘(1) Washing in distilled water. 

‘©(2) Immersion one or two minutes (they can also re- 
main several minutes without injury) in the above fixing and 
staining fluid. Several cubic centimeters of the fluid suffice 
for many sections. The blackening can be observed with the 
naked eye. 

‘©(3) Careful washing in distilled water. 


‘©(4) If desired, light carmine stain to bring out the cell 
bodies and nuclei in the fine interstitial nervous network. Acid 
carmine is best adapted for this according to my experience and 
I find especially suitable a dilution of this staining fluid with 
acetic acid and alcohol (equal parts). The fluid into which the 
sections are brought must have a deep red color. 


‘“(5) Repeated washing in water and then successive 
transference into alcohol and clove oil and finally mounting in 
Canada balsam or damar in the usual way. 

‘« Preparations treated in this way possess, besides the 
above mentioned advantages, the additional one that the fine 
powdery precipitate does not after a while appear. This pre- 
cipitate almost always, if there have not been previous repeated 
and long continued washings, at last spoils the preparations 
prepared according to the original method.”’ 


[ro BE CONTINUED. | 


Alum, 20 g. 
Ammonium sulphocyanide, Io g. 
Sodium chloride, 40 g. 
This mixture stands quiet for 8 days and is then filtered. 
(b) Water, roo g. 
Gold chloride, 1 g. 
To prepare the bath one mixes 
of solution (a), 60 ccm. 
6G ce (b), 7 ccm. 
old, combined bath, 40 ccm. 
For economy and convenience I use the fluid which has also served for ton- 
ing, thus for this purpose almost useless. 


THE DORSAL SACK, THE AULIX AND THE DEER 
CEPHALIC (RVUEXURE: 


B. G. WILDER. 
Abstract. 


In many of the lower vertebrates there is a pouch-like 
evagination of the diatela (membranous roof of the diacele, or 
‘third ventricle’’) between the epiphysis and the paraphysis. 
In Ceratodus and in Polyodon it is anteverted and very large. 
In the green turtle it projects dorsad. Its mammalian repre- 
sentative is crowded caudad by the cerebrum and rests upon the 
retroverted epiphysis. It is commonly ignored in figures and 
descriptions, but was shown by Reichert. The speaker, with- 
out recognizing its homology, figured it in the Mew York Medt- 
cal Journal, 1885, p. 320. It is large in the sheep and long in 
the cow and horse. The diaplexuses are continued into it. 

Aulx is the mononym proposed by the speaker in 1882 
for the ‘Sulcus Monroi” of Reichert—a sigmoid groove on 
the mesal surface of the thalamus just ventrad of the medicom- 
missure and connecting the orifice of the mesocele with the 
porta (foramen of Monro) at either side. His and C. S. Minot 
regard it as the diencephalic representative of the ‘‘ salcus inter- 
zonalis,” the primary furrow between the dorsal and ventral 
zones of the nervous axis. But recent observations in the 
neurologic laboratory at Cornell upon embryo kittens render this 
interpretation somewhat doubtful, and the matter is under inves- 
tigation. 

In 1889, in the ‘‘ Reference Hand-Book of the Medical 
Sciences”’ the speaker called attention to the sharp angle be- 
tween the prosencephalic and the diencephalic portions of the 
brain cavities. A recent review of the brains of all vertebrate 


'Read before the American Neurological Association, June 4, 1896. 


Wiper, Parietal and Paroccipital Fissures. 129 


classes warrants the generalization that a marked diencephalic 
flexure occurs in all Reptiles, Birds and Mammals, while in 
Amphibians and ‘‘fishes’”’ it is absent or insignificant. 


tit ECTAL RELATIONS OF Tie RIGHT AND- EERE 
PARIETAL AND) PAROCEIPITAL HISSURES? 


B. G. WILDER. 
Abstract. 


The parietal and paroccipital fissures may be either com- 
pletely separated by an isthmus, or apparently continuous. 
When so continuous ectally there may still be an ental and con- 
cealed vadum or shallow. Disregarding the vadum on the pres- 
ent occasion, the ectal relations of the two fissures may be des- 
ignated as either contenwty or separation. That continuity oc- 
curs more frequently on the left side has been noted by Ecker, 
Cunningham and the writer. Hitherto, however, statistics have 
included unmated hemicerebrums as well as mates from the 
same individuals. The following statement is based upon the 
cerebrums of 58 adults of both sexes and various nationalities 
and characters. The speaker has examined 48; the other 10 
have been accurately recorded by Bischoff, Dana, Jensen and 
Mills. 

The four possible combinations of right and left continuity 
and separation occurred as follows: 

I. Left continuity and right separation in 27; 46.5%. 

II. Right and left continuity in 22; 38 %. 

III. Right and left separation in 8; 13.8 %. 
IV. Left separation and right continuity in 1; 1.7%. 


1Read before the American Neurological Association, June 3, 1896. 


130 JOURNAL OF COMPARATIVE NEUROLOGY. 


When 5 groups of persons are recognized the combina- 

tions are as follows: 
~ A. In 8 moral and educated persons, combination I, 
62753 Th 25 Tit ras. 

B. In 23 ignorant or unknown, I, 56.5; II, 34.8; II, 8.7. 

Cin 2oinsane) 1,40. ieo5 il 20) al Vege 

De Mniaomurderers, Teo; Me 7s el be 

Bo ines mearoes, les emis oys 

So far as these 58 individuals are concerned, the most com- 
mon combination, vzz., left continuity and right separation, is 
decidedly the rule with the moral and educated, less frequent 
with the ignorant and unknown, the insane and negroes, and 
does not occur at all in the murderers. The only instance of 
the reverse combination (left separation and right continuity) is 
an insane Swiss woman. The only two known to be left-handed 
presented the more frequent combination I. 

These statistics suggest many special queries and problems, 
some of which were briefly indicated. But the speaker wished 
this to be regarded as a preliminary communication and asked 
the cooperation of other members in the effort to obtain satis- 
factory records of larger numbers, particularly of brains of well- 
born, moral and educated persons. For this purpose a blank 
form was outlined and will be sent upon application. 


EDITORIAL. 


NEURONYMIC PROGRESS IN AMERICA. 


The American Neurological Association, at its recent ses- 
sion in Philadelphia, took a step at once sure and decided toward 
the simplification and unification of neurologic nomenclature. 
The Committee on Neuronymy (Drs. H. H. Donaldson, L. 
GrGray,-Cy Ke Mills iC. Seguin, i. Ca-spitzka sandy Bs iG: 
Wilder, chairman) presented a report which was adopted 
unanimously by the Association, June 6. To give this impor- 
tant measure as wide a circulation as possible we publish the 
report of the committee as furnished us by its chairman. 

The first five sections are substantially identical with re- 
ports that were adopted unanimously by the Association of 
American Anatomists in 1889 and by the American Association 
for the Advancement of Science in 1890 and 1892. The re- 
commendations are as follows: 

1. That the adjectives borsAL and VENTRAL be employed 
in place of posterior and axterior as commonly used in human 
anatomy, and in place of upper and dower as sometimes used in 
comparative anatomy. 

2. That the cornua of the spinal cord, and the spinal 
nerve-roots, be designated as DORSAL and VENTRAL rather than 
as postertor and anterior. 

3. That the costiferous vertebre be called THORACIC rather 
than dorsal. 

4. That, other things being equal, Mononyms (single-word 
terms) be preferred to polyonyms (terms consisting of two or 
more words). 

5. That the Azppocampus minor be called cALCAR; the 
hippocampus major, HIPPOCAMPUS; the fous Varoliz, pons; the 
msula Retlit, INSULA; pra mater and dura mater, respectively PIA 
and DURA. 


132 JoURNAL OF COMPARATIVE NEUROLOGY. 


6. That the following be employed rather than their vari- 
ous synonyms: HYPOPHYSIS, EPIPHYSIS (for coxarimm and cor- 
pus pineale), CHIASMA, OBLONGATA, LEMNISCUS, MONTICULUS, TEG- 
MENTUM, PULVINAR, FALX, TENTORIUM, THALAMUS, CALLOSUM, 
STRIATUM, DENTATUM, MESENCEPHALON, PALLIUM, OLIVA, CLAVA, 
OPERCULUM, FISSURA CENTRALIS (for f. Rolando, etc.), F. CAL- 
CARINA, F. COLLATERALIS, F. HIPPOCAMPI, CUNEUS, PRAECUNEUS, 
CLAUSTRUM, FORNIX, INFUNDIBULUM, VERMIS. 

It will be noted that at least twenty (about half) of the 
names above recommended coincide with those adopted by the 
European committee in 1895, and that with some others, e. g. 
CALLOSUM, DURA and CALCAR, the difference is due merely to 
the elimination of superfluous words. 

The next number of the JouRNAL OF COMPARATIVE NEv- 
ROLOGY will contain an extended commentary upon the Report 
by the chairman of the committee. 


Journal of Comparative Neurology. Vol. VI. PLATE IX, 


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THE BRAIN OF THE BEE 


A PRELIMINARY CONTRIBUTION TO THE MORPHOLOGY OF THE 
NERVOUS SYSTEM OF THE ARTHROPODA. 


By F. C. Kenyon, Ph.D., 
Fellow in Biology, Clark University, Worcester, Mass. 


CONTENTS. 


Introduction. 


MATERIALS. 
METHODs, 


General description. 


External appearance. 
CEREBRAL NERVES, 
GENERAL INTERNAL STRUCTURE. 
TYPES OF NERVE CELLS. 
CONNECTION BETWEEN NERVE FIBERS. 
TRACHEA, 


Special description, the Proto-cerebron. 


THE MUSHROOM BODIES, 
The calyx-cup. 
The stalks of the mushroom bodies, 
The roots of the mushroom bodies. 
MINUTE STRUCTURE OF THE MUSHROOM BODIES. 
fibers ending in the calyx. 
Fibers ending in the stalks and roots. 
THE FUNCTION OF THE NERVE CELLS OF THE MUSHROOM BODIES, 
THE CENTRAL BODY. 
THE TUBERCLES OF THE CENTRAL BODY. 
THE FIBRILLAR ARCH. 
THE ASSOCIATION FIBERS OF THE PROTO-CEREBRON, 


The Deuto-cerebron. 


THE ANTENNAL LOBE, 
THE ROOT OF THE ANTENNO-SENSORY NERVE. 
THE ROOT OF THE ANTENNO-MOTOR NERVES, 


134 JOURNAL OF COMPARATIVE NEUROLOGY. 


The dorso-cerebral fiber tracts. 
THE CONNECTIONS WITH THE OPTIC LOBES. 
The anterior optic tract. 
The postero-superior optic tract. 
The antero-supertor optic tract. 
The antero-posterior optic tract. 
The posterior optic tracts. 
THE DORSO-CEREBRAL COMMISSURES. 
The superior dorso-cerebral commissure. 
The anterior commissure. 
The optic commussures. 
The inferior dorso-cerebral commissures. 
THE ANTENNO-CEREBRAL TRACTS. 
The inner antenno-cerebral tract. 
The middlé antenno-cerebral tract. 
The outer antenno-cerebral tract. 
THE DORSO-VENTRAL TRACT. 
The ocellar nerves. 


FIBERS FROM THE BRAIN TO THE OCELLAR GANGLIA. 
The commissural and ventro-cerebral region. 


The cell groups. 


THE CELLS OF THE DORSO-CEREBRON. 
THE CELLS OF THE VENTRO-CEREBRON, 


Conclusion. 
Bibliography. 
introduction. 


Since Viallanes’ (gs) monograph on the brain of the grass- 
hopper (Oedopoda and Caloptenus) and the smaller but as excel- 
lent work of Cuccati on the organization of the brain of So- 
momya appeared nothing has been added to our knowledge of 
the structure of the hexapod brain. In fact no one seems to 
have worked at the subject. Such being the case, my endeavor 
by an application of one of the more recent histo-neurological 
methods to bring this subject into line with what is now known 
relative to the stucture of the central nervous system of several 
other zxvertebrates and more especially of the vertebrates will 
doubtless be appreciated. Here, as elsewhere, an application 
of the chief of the two recent methods, namely that of Golgi, 
brings to light facts before known only from inference, and also 
often materially alters conceptions based wholly upon the older 


Kenyon, Zhe Brain of the Bee. 135 


methods of staining. A detailed history of the matter in hand 
is therefore needed, but since it would add too much to the 
length of the present paper, the pretentions of which as indi- 
cated in the title are very limited, little more than a reference 
here and there will be given. This course is doubtless excusa- 
ble since Retzius (99) has already given a summary to which the 
reader may be referred. Three papers that have appeared since 
the publication of this author’s work may, however, be noted. 

The first of these is the paper by Saint-Remy (99) on the 
brain of tracheate arthropoda. In this the author describes the 
brains studied histologically of myriopods, chilopods, arach- 
nids and Perzpatus; but although the work is comprehensive 
and fills a void in our knowledge of the arthropod nervous sys- 
tem, it is far from treating the subject with sufficient depth, a 
deficiency mostly due to the method of staining employed. 
The nerve cells, or cell bodies, following Dietl, he divides into 
two groups, one composed of large cells well supplied with ex- 
tra-nuclear protoplasm and with relatively small nuclei having 
few chromatin elements, the other of small cells having little or 
no extra-nuclear protoplasm and relatively large nuclei richly 
supplied with chromatin elements and restricted to certain cere- 
bral areas. To the latter he applies the term chromatin cells 
in preference to the ganglionary nuclei of Dietl and others. 

Two small nerves that he describes for the myriopods may 
be passed by with the remark that one, namely the nerve of 
Tomosvary appears to have no homologue in the hexapods un- 
less it be the small nerve mentioned by Newton ( 79) as arising 
from the front of the brain of the cockroach and terminating in 
a small organ near the base of the antenna. The other, or 
tegumentary, nerve may be the homologue of the tegumentary 
nerve described by Viallanes (gg) for the grasshoppers and con- 
sequently, as will be shown in subsequent pages, the homo- 
logue of the salivary nerve that I have traced out in the bee. 

The most interesting discovery recorded in the paper is 
that of structures found in the brain of Scutzgrva that he de- 
scribes as mushroom bodies. The term he bases upon their 
form and not upon an idea of their being the homologues of the 


136 JOURNAL OF COMPARATIVE NEUROLOGY. 


organs of the same name found in the hexapods. But from 
the fact that their cells are of the chromatic cell type and simi- 
larly situated andalso from the fact that he seems able to recognize 
nearly all the parts generally described in the organs in the hex- 
apods it may here be suggested that they may possibly be the 
homologues of these latter organs. There is needed, however, 
a more minute and painstaking study than St. Remy has given 
the structures before one should come to a definite conclusion as 
to their homologies. 

The second of the three papers is that by Viallanes (93) 
on ‘‘The neural centers and sensory organs of articulates ’’ and 
is mainly of interest here in showing this author’s conception of 
the minute structure of the nervous system. In his earlier pa- 
pers he did not touch upon the matter, but in his last paper 
one finds him following St. Remy’s grouping of the cells and 
adding a few ideas upon the finer structure of the fibrillar sub- 
stance. These ideas were doubtless founded more upon the 
general results obtained with the methylen blue and the Golgi 
methods, which by this time had become very generally known, 
than upon any observations of his own made by the aid of his 
methods of staining. 

He describes and figures diagrammatically three kinds of 
fibers. One of these, of large size, originates from large 
cells and passes outside of the central nervous system after first 
giving off in the latter a few short and fine branches. These 
he calls motor fibers. Another kind, of much smaller size, enter 
from cells outside the central system and branching more or less 
profusely connect with the small branchlets of the first, forming 
thus the terminations and connections of sensory fibers. 

The third kind are short, small fibers originating from 
the chromatin cells and, branching profusely, connect with 
the fine branchlets of the other two kinds of fibers. As will 
be seen in subsequent pages, this general idea is supported by 
facts, though, as indicated above, it was with Viallanes almost 
purely hypothetical. 

The last of the three papers is one by Binet (94). This 
author so far as the structure of the dorso-cerebron is concerned 


Kenyon, Zhe Brain of the Bee. 137 


seems to follow Viallanes without a question, and deals almost 
wholly with the ventral system, which he treats anatomically 
and physiologically. He seems to have added a few new facts 
to our knowledge of the subject treated, although he leaves 
very much still to be done. The points at which he seems to 
have erred or where his treatment is deficient will be noted, 
when they are closely related to the matter in hand, in subse- 
quent pages. Here it may be noted that he seems to have ob- 
tained the beginning of a correct idea of the constitution of the 
arthropod nervous system based upon the work of Retzius and 
others employing similar methods; but, failing completely in 
his use of the Cajal-Golgi method and obtaining but litttle better 
results with methylen blue, he concludes that, since in any one 
preparation neither of these methods brings out the whole structure 
they are not to be depended upon, and then passes on to a de- 
pendence upona modification of the old method of Weigert 
and becomes lost in the mass of detail that he so much desired 
to obtain. 

The results of other writers will be noted in connection 
with the descriptions of the structures with which their works 
deal. 

MATERIAL. 


The material used has been almost exclusively the common 
honey bee (Afzs mellifica), which was at hand in abundance, 
for which I owe many thanks to Dr. C. F. Hodge. Possibly a 
thousand or more brains were treated by several of the bichro- 
mate of silver methods, and of this number scarcely more than 
fifteen or twenty per cent. were found successfully impregnated. 


METHODS. 


Aside from impregnation with bichromate of silver, the 
methods that I have employed are various, and among these the 
one that has given the best results, brought details to light most 
beautifully, has been one using sulphate of copper and haema- 
toxylin with brains hardened in from 10 to 20% formol for 
twenty four hours or longer. Such preparations show the axis- 
cylinders as purplish-brown fibers within their surrounding 


138 JOURNAL OF COMPARATIVE NEUROLOGY. 


slightly bluish sheaths, the nuclei of the latter and of trachee, 
the tracts of nerve fibers and some of the association fibers, 
clearly marked off from the surrounding ground work of fine 
branchlets forming the Punktsubstanz of the older German 
writers and the medullary substance of Viallanes. The struc- 
ture of the cell bodies and their nuclei is also well brought out. 

In some cases the freshly excised brains were dropped into 
10 to 20% formol, in others they were placed in the following 
mixture, which gave just about as good results : 


10% Potassium bichromate, 40 parts. 
5% Sulphate of copper, —_40 parts. 
Formol, ° : 20 parts. 


This might be improved by leaving out the bichromate, 
which is rather unnecessary since the formol likewise has the 
property of rendering the components of animal tissues insolu- 
ble in water and alcohol. 

Brains hardened in this mixture were washed for from a 
little while to some hours in tap water, then for from a few to 
twenty four hours in 70% alcohol, after which they were de- 
hydrated, imbedded in paraffin, sectioned and stained on the 
slide in the hamatoxylin. 

In the other cases the sections of the formol-hardened 
brains were treated with the sulphate of copper (5%). Ina 
solution of this they were left for from a few to twenty four 
hours or in a warmed solution for from 20 to 30 minutes. They 
were then washed off in tap water and stained. 

The hematoxylin mixture that gives the best results is 
one containing phosphomolybdic acid, though the plain alcohol- 
water hematoxylin used in the method of Weigert and its 
modifications gave good results for certain details in the roots 
of the mushroom bodies. The former mixture is that suggested 
by Mallory and is composed as follows : 


10% Phosphomolybdic acid WC.c. 
Hematoxylin crystals I grm. 
Chloral hydrate : 6 to 10 grm. 


Water 4 : : 100 C.¢, 


Kenyon, Zhe Brain of the Bee. 139 


Enough of this was added to a dish of water to give the 
latter a black appearance, or it was diluted about I to 5. From 
a quarter of an hour to an hour is required for staining. The 
sections hardened in the potassium bichromate mixture require 
a longer time than the others. 

After staining, the sections were washed off with 70% 
alcohol and dehydrated or, if too deeply stained, left in the 70% 
alcohol for some time. With the dilute solution, however, there 
is not much danger of overstaining, if the process is watched. 

For impregnation with bichromate of silver the rapid 
method of Cajal was at first employed, but only later to be set 
aside when it was discovered that one in which the osmic acid is 
replaced by formol gives a much more transparent background 
for the darkened fibers and cells, thus allowing much thicker 
sections to be cut, while at the same time retaining the 
rapidity that has made the osmic acid mixture so excel- 
lent. At first the osmic acid was replaced by the same 
amount (one-fifth) of pure formol, later Strong’s suggestion 
of equal parts of 10% potassium bichromate and _ formol 
was followed. This was then modified to 5% bichromate 
and a smaller amount of formol as follows : 


5% bichromate, 8o cc. 
Formol, 20 cc. 


This gave impregnations as good as the stronger and a lit- 
tle better than the weaker combination while at the same time 
lessening the formation of a black precipitate that. is always 
found in formol-bichromate mixtures after they have stood some 
hours. This precipitate was avoided as much as possible by 
changing to fresh fluid every twenty-four hours. Impregnations 
may be obtained in brains left unchanged in the mixture until 
ready for the silver solution. The precipitate must however weak- 
en both the formol and the bichromate and hence operate disad- 
vantageously. I think my failures were more numerous with 
brains left in the unchanged fluid, (though no experiments were 
undertaken to settle the matter definitely,) and hence I am not 
inclined to agree wholly with Kopsch (96) when he asserts that 
the precipitate makes no material difference. 


140 JOURNAL OF COMPARATIVE NEUROLOGY. 


Like this writer, I also found that twenty-four hours in the 
formol-bichromate is sufficient for hardening and that tissues 
may then be transferred to pure bichromate. He employed a 
344% solution, however, while I, since it seemed to act more 
quickly, used a 5% solution. 

From three to four or five days immersion in formol-bi- 
chromate are necessary for obtaining impregnations of nerve 
fibers, and almost three for the cells, while the tracheee may 
be incrusted almost to the exclusion of nerve fibers after one 
or two days. 

The more or less inevitable precipitate and crystals were of 
course met with, but on the whole my preparations were com- 
paratively free from them. In fact, judging from the printed 
experiences of others, I seem to have been more favored than 
is usual. Only a few cases occurred out of the hundreds of 
brains sectioned where the precipitate at all resembled the fine 
branching of the nerve fibers, and in these there was no ground for 
a deception of the practiced eye, for the angular contours of the 
precipitate readily distinguished it from the rounded ones of the 
fibers. 

The precipitate gave the greatest annoyance in the layer of 
cell-bodies. This is so thin on the more exposed parts of the 
brain that it was usually completely obscured when the brains 
were brought into direct contact with the fluids. Attempts were 
made to obviate this difficulty. Formic acid was tried and dis- 
carded, as was also Berkeley’s (95) suggestion of adding a few 
drops of phosphomolybdic acid to the silver solution just before 
using. The latter can, in fact, do nothing more than weaken 
the solution of silver nitrate, and one may therefore as well take 
a weaker solution to begin with. When this is done and the 
tissue, after first quickly washing off the bichromate with water, 
is carefully rinsed in the weak solution, or one that has been 
previously used, before it is allowed to stand, one will be much 
less troubled with the incrustation. 

This method did not prove as good, however, as one in 
which the brain was left zz sztw in the head, enough only of the 
front, top, or side of which was cut away to allow the fluids to 


Kenyon, Zhe Brain of the Bee. I4!I 


enter. In such cases the part most exposed would be incrust- 
ed, while that opposite would be entirely free. Other methods 
were thought of and tried. That of wrapping a brain in filter 
paper was considered too rough a treatment ; covering a brain 
with celloidin failed. Had the one suggested by Retzius (9 4), 
namely, of dipping a brain in warm gelatine, been thought of or 
known in time, it would doubtless have given excellent results. 

The nitrate of silver was employed in strengths ranging 
from %% to 2%. The stronger solutions, though giving 
stronger impregnations, less evidence of beading, or fuller fibers, 
have the disadvantage of being more inclined than the weaker 
solutions to form artifacts. A solution of 1% strength was 
finally adopted for leaving the specimens in over night or until 
I was ready to section them. 

Generally only one or two immersions in the fluids was 
given the specimens, since it was desirable to have only a few 
fibers impregnated so that very thick sections could be cut and 
thus give the entire trajectory or as much as possible of a fiber 
unobscured by other details. 

The impregnated brains were transferred to absolute alco- 
hol and then to celloidin. Where they were entirely free from 
the head or from chitinous particles that would tend to prevent 
them from becoming fairly saturated or surrounded with the 
celloidin, sections wese cut within two hours from the removal 
from the silver solution. But where they were surrounded by 
the chitinous cephalic capsule from six to twenty-four hours 
were found necessary to form a celloidin block of a consistency 
sufficient to prevent tearing and breaking of the sections. 

Sections were cut all the way from twenty to two hundred 
and ten microns thick, thus, in the thickest, making only three 
or four frontal sections to a brain. 


General Description. 


External appearance. 


Under the general term ‘‘brain”’ I here understand the 
whole of the neural mass included within the head, excepting 
only the two small ganglia generally known as the stomatogastric 


142 JOURNAL OF COMPARATIVE NEUROLOGY. 


ganglia. The optic lobes and the subcesophageal ganglion are 
part of it. There is no real ground for leaving the latter por- 
tion out as is generally done under the term subcesophageal. 
Even when it is widely separated from the larger mass and is to 
be found crowded back into the pro-thorax, there is abundant 
reason, as shown by the physiological experiments of Binet 
(94) and others, for retaining this portion under the general 
term brain. Hence in subsequent pages it will be referred to 
as the ventro-cerebron in contrast to the super-intestinal portion 
or dorso-cerebron. 

The optic lobes will be omiited from this paper for consid- 
eration in a later one. 

The bee’s brain thus limited is, roughly speaking, when 
viewed from the front, a quadrilateral structure, with rounded 
angles. Setting aside the ventro-cerebron, which projects back- 
ward, it is considerably compressed antero-posteriorly (Pl. XX), 
and abuts closely against the middle upper portion of the pos- 
terior walls of the head, leaving a very considerable space in 
front filled with tracheal air-sacs and the whitish, often yellow- 
ish or orange racemose bunches of the salivary glands. These 
latter organs also fill in the smaller spaces behind the brain and 
are closely applied to it. On the dorsal side are three small 
protuberances (Pl. XVII, XX) with black tops, or the so-called 
ganglia of the ocelli. Below in the lower third the brain is 
pierced by a large oval opening, the cesophageal foramen, with 
its largest axis vertical. Near the roof of this is the longitu- 
dinal median nerve of the stomatogastric system. On either side 
and below the foramen are the rather conspicuous antennal 
lobes forming the greater portion of what has been described as 
the deuto-cerebron. The ventro-cerebron, as already mentioned, 
projects considerably backwards, but it is so closely united with 
the dorso-cerebron by extremely thick and short cesophageal 
commissures that it is very readily recognized as part of the 
brain. So compact is this deuto-trito-cerebral portion of the 
brain that were it not for the fact that the labral nerves are rec- 
ognized as arising from it, indications of the trito-cerebral lobe 


Kenyon, Zhe Brain of the Lee. 143 


would be recognizable only in sections. Even in these its boun- 
daries are indefinable. 


THE CEREBRAL NERVES, 


From the lower side of the antennal lobe there is given off 
the large antennal nerve, which passes forward and _ slightly 
downward to the-entrance of the antenne. From the lower 
surface behind the origin of the large nerve there originates 
a small one that may be readily traced in sections to the anten- 
nal muscles within the head. This, which I will denominate 
the antenno-motor internus, in contradistinction to the other, or az- 
tenno-motor externus, was first discovered and traced to its termi- 
nation in the muscles by Newton (79) in his study of the brain 
of the cockroach. It is described by Viallanes (7) in his 
paper on the brain of the wasp as the ‘‘nerf antennaire acces- 
soire’’ and was correctly identified by him with the nerve de- 
scribed by Newton, although he was unable to trace it to its 
termination in the muscles. Doubtless it may be found as a 
separate nerve in all tracheate arthropods. One occupying the 
same position I have recognized in Scalopendrella tmmaculata 
and traced into muscles that I supposed to belong to the an- 
tenne. 

Behind this internal antenno-motor nerve and immediately 
behind the base of the antennal lobe there is a somewhat larger 
nerve that I have traced towards the mouth-parts of the bee 
and from its position take to be the labral nerve. This arises 
from the trito-cerebron and may be seen in fragment in the pho- 
tograph (fig. 2, Pl. XIV), which is of a section slightly too far 
forward to show the lobe and the entering nerve root. 

From the lower side of the ventro-cerebron (PI. XX) there 
arise not far apart from each other three pairs of large nerves 
that like the one just described may be readily traced to the 
mouth parts. Since in most of my preparations these parts 
were cut away to allow a free access of the fixing fluids to the 
brain or to free the object as much as possible from chitinous 
parts to avoid tearing during the process of sectioning, I have 
not yet traced the nerves to their endings. From their posi- 


144 JOURNAL OF COMPARATIVE NEUROLOGY. 


tion, however, I infer them to-be the mandibular, maxillary, 
and labial nerves. From the posterior lower surface of the 
ventro-cerebron there arises a small seventh pair of nerves di- 
rected backwards and downward toward the region just below 
the foramen magnum. Just how they terminate or what their 
function may be I have not thus far been able to determine. 
They were seen by Viallanes (7) in the wasp, but were left as 
now still to be traced. 

In his monograph on the brain of the Orthoptera, Viallanes 
(8) described under the name of ‘‘nerfes tegumentaires”’ a 
pair of nerves emerging from the surface of the brain and pass- 
ing on to the roof of the head. These appear to be nothing 
more nor less than a pair of nerves that he found in the wasp 
the year before and described as nerves of unknown termina- 
tions. 

This eighth pair of nerves is readily seen and traced in the 
bee and one may be recognized in section in the lower photo- 
graph in plate XVI. Both in bichromate of silver and in hema- 
toxylin preparations I have been able to trace out its entire 
course and to show that Viallanes’ denomination of tegument- 
ary nerve is entirely a misnomer. It arises from the side of the 
ventro-cerebron where its root enters the latter, passes in, some- 
what downwards and backwards, towards the median line of the 
latter, makes a turn backwards and becomes lost in the general 
mass of fibrillar substance. The nerve (Pl. XVII) closely adher- 
ing to the posterior surface of the brain and beneath the gen- 
eral envelope of the latter passes outwards and upwards until it 
reaches the upper portion of the posterior surface of the inner 
or third optic fibrillar mass or ganglion, where it branches. One 
branch continues almost perpendicularly upwards, passes out of 
the brain envelope and branches among the lobes of the saliv- 
ary glands. In bichromate of silver preparations the branches 
may be followed into the lobes, but I have not yet seen the 
finer terminations. The other large branch continues outward 
along the supero-posterior surface of the optic lobe, branching 
and leaving the ‘envelope from time to time, and may be fol- 
lowed to the basement membrane of the retina where it finally 


Kenyon, The brain of the Bee. 145 


emerges to pass into the glandular lobes situated in proximity 
to that structure. The nerve, then, instead of having anything 
to do with the integument, is the nerve to the salivary glands, 
and is doubtless made up entirely of efferent fibres. 

The small pair of nerves described by Newton (79) as 
arising from the anterior surface of the brain in /atta and go- 
ing to the small white spot near the base of each antenna, 
are not, so far as I have been able to discover, represented in 
the bee. 

THE GENERAL INTERNAL STRUCTURE. 


Internally the brain is composed of a mass of very fine 
fibrils scarcely recognizable as such in ordinary preparations, 
which is very well called fibrillar substance. The upper portions 
of the anterior surface of this below the mushroom bodies to 
be described later and the corresponding part of the posterior 
portion of each of the two lateral lobes are directly covered by 
the membrane forming the perineurium. But in the region of 
the mushroom bodies, laterally, in the posterior median furrow, 
and below, about the greater part of the antennal lobes, fill- 
ing the space between these and the proto-cerebron, and the 
lower and lateral surface of the ventro-cerebron (the upper sur- 
face of the latter is bare) it is covered by heaps of cells which 
spread out into layers of only one or two cells deep. Processes 
from many of these cells are gathered into large bundles which, 
penetrating the fibrillar substance, give it~—both the brain and 
the ventral system as seen in sections—a lobular appearance that 
has been the subject of no inconsiderable amount of painstak- 
ing and lengthy and as tedious description, notably by Vial- 
lanes, and later by Binet (94) in his paper on the sub-intestinal 
nervous system of insects. The fibers thus entering become 
lost in the fibrillar substance by repeated branching or continue 
first to more distant parts, thus forming connecting tracts. This 
relation of the fibrillar substance and the fibers may be readily 
seen in the accompanying photographs, and in most prepara- 
tions by ordinary staining methods. The assistance of the 
method of impregnation with bichromate of silver or that of 


146 JOURNAL OF COMPARATIVE NEUROLOGY. 


intra-vitam staining with methylen blue is necessary to determine 
the relations of cells, fibers and fibrillar substance completely. 


THE TYPES (OF NERVE (CELES: 


By the assistance of the bichromate of silver method we 
may imagine a cell with a nerve fiber dividing at some distance 
from the cell body into two branches that form with ita Tora Y 
as probably the most typical of the nerve cells of the bee, or prob- 
ably of arthropods in general. The assertion does not hold 
‘good completely, for there are cells in the optic lobes and even 
in the central part of the brain in which all the branches given 
off appear as very small branches of large fibers. 

To classify for convenience of description the various mod- 
ifications met with in the brain one may divide them into : 

1. Afferent or sensory fibers in which the cell body is situ- 
ated somewhere in the neighborhood of the external (to the 
brain) organ of sense and sends its neurite into the brain and 
its dendrite into the sense organ. Such cells may be called bipolar, 
and in them, asin all other nerve cells, the only criterion of 
the difference between dendrite and neurite is in the direction 
of the passage of the neural impulse with reference to the cell 
body, or to the trunk of the fiber leading to the cell body. 
Such fibers do not penetrate far into the brain, where they may 
terminate in a mass of fine branchlets as in the case of the fibers 
from the antenne. In other fibers the terminal branching is 
more or less loose or not massed together, as in the case of 
whatever sensory fibers there are entering through the labral, 
mandibular, maxillary and labial nerves or from the ventral cord. 

In the accompanying diagrams this class of fibers and all 
others bearing sensory stimuli into the portion of the brain 
here considered, as from the optic lobes and the ventral cord, 
are colored blue. 

2. Connecting fibers in which the cell body is situated to 
one side of the track of the passing stimulus—assuming that it 
does not necessarily go to the cell body—and in which the 
dichotomous T- or Y-shaped branching is most easily recogniz- 
able. The peripheral of the two branches connects with the 


Kenyon, Zhe Brain of the Bee. 147 


terminations of the sensory fibers of the first class and, since it 
bears the transferred impulse inward, may be called the den- 
drite. Usually it is shorter than the other branch that, with 
others of its kind, passes as a tract of fibers through the fibrillar 
substance to the mushroom bodies, in which it breaks up 
into a bushy or arborescent mass of branches, or to other parts 
of the brain, where the branching is of a more open kind. 

The fibers connecting the sensory fibers of the antennal 
lobes with the mushroom bodies are most typical of this class. 
Others that must be classed along with them enter from the 
optic lobes and along with them are represented in violet in the 
diagrams. Others found connecting the central body, later to 
be described, are doubtless to be classed here, since this body 
seems to bein connection with the fibers of the ocelli; but 
owing to the unsatisfactory condition of my present understand- 
ing of this body have been left in black. The mode of branch- 
ing at the ends! of the dendrite is represented in fig. 15, PI. 
XXI, while the branching terminals in the mushroom bodies 
are shown in figs. 17, 18, 21, of the same plate. Most of these 
fibers appear to be provided with a nucleated sheath. 

The representation of fibers found in the central body in 
figs. 32, 37, Pl. XXII, by comparison with figs. 17 and 21 of the 
preceding plate, which beyond a doubt represent the receptive 
terminations, would seem to indicate that the former are den- 
dritic. Fig. 18, Pl. XXI, shows the terminations of the neurite- 
branches in the mushroom bodies as seen under a magnifica- 
tion of 586 diameters. 

3. The cells of the mushroom bodies. These fall under the 
head of Y-shaped cells and consequently follow the class just 
described, but will be left for consideration along with the 
bodies which they largely compose. They are represented 
in red. 

4. Efferent fibers which originate from cell bodies situated 
within the brain and bear efferent stimuli to muscles, glands or 
to cells outside of the brain are represented in the diagrams 
in brown. The cells of this order seem to depart widely from 
the simple T-shaped general type as is shown in fig. 36, Pl. 


148 JoURNAL OF COMPARATIVE NEUROLOGY. 


XXII, which represents the best specimen of motor cell that 
my preparations offered. That it is motor in nature is assumed 
from its general appearance, resembling to a certain extent the 
motor fibers described and figured by Bethe (95), and the fact 
that it is cut off near the cut end of a large fiber not figured 
that passes into the antenno-motor nerve and with which it ap- 
pears to have been continuous, but on account of a slight 
bend outside the plane of the razor was cut in two. This ap- 
peared in a nearly sagittal section. In a thick section cut in the 
frontal plane, or nearly transversely to the main axis of the 
animal’s body, fibers were seen, such as one represented in 
Plates XVII and XX, sending a process into a tract of fibers 
that appeared to pass down behind the antennal lobe, apparently 
into the root of the antennal nerve, which were therefore thought 
to be motor fibers. Combining these two with what was seen 
of fibers in other sections in about the same locality as the 
motor cell mentioned it would appear that the large motor cells 
send a process into the fibrillar substance, which almost imme- 
diately gives off small branches that may end at a considerable 
distance from their point of origin. The process becomes very 
much enlarged for a considerable extent of its course and may 
even send off branches of considerable size before passing on 
into the root of a nerve as a small fiber. 

5.  TDhe commissural fibers, which are doubtless of the gen- 
eral T-shaped form, though I have not been able to find their 
cells of origin. They pass from one side of the brain to the 
other and may be represented by the fibers composing the 
superior commissure. The fibers composing the so-called com- 
missure of the optic lobes may possibly also be classed here. 
Other fibers there are in plenty than those gathered into tracts 
passing between the two halves of the brain, but they are better 
classed with the next group. 

6. Association fibers seemingly serving the purpose of con- 
necting the elements already described with one another. These 
are mostly large profusely and irregularly branching forms 
some of which are shown in plate XXII. In some“cases I have 
been able to make out the cells of these, in others I have not. 


Kenyon, The Brain of the Bee. 149 


They all form a promiscuous group that might be, but for lack 
of a better knowledge of them and their cells of origin, may as 
well not be, subdivided. Almost all of them are known only 
from fragments of which those shown in plates XXI and XXII 
are the best examples. Some of them may possibly even be 
fragments of motor-fibers. 

To these groups may be appended two others, one repre- 
sented by the enormous fibers from the ocelli and the other by 
the equally enormous fibers situated near the dorsal surface of 
the ventral system and of the ventro-cerebron. They will be 
considered in connection with the ocellar nerves. 


THE CONNECTIONS BETWEEN NERVE FIBERS. 


What facts I am able to bring forth bearing upon the 
‘‘contact’’ idea and the old ‘‘ mesh-work ”’ idea of the fibrillar 
terminations, though more properly considered here, I will de- 
fer until I have described the mushroom bodies, in connection 
with which the facts mentioned occur. 


TRACHEA. 


Besides these neural elements and the sheaths surrounding 
some of them, there are to be found large numbers of trachea! 
fibers. In bichromate of silver preparations they are the first 
elements to become impregnated and then are brought plainly 
into view. The larger branches may be very readily recognized 
in hematoxylin preparations. Their general tapering form and 
the time of their taking up the bichromate of silver prevents 
their becoming confounded with nerve fibers. 

To explain their origin it must first be noted that the brain 
of the bee is surrounded by a number of tracheal sacs, the in- 
ner walls of which closely cover it. Here and there tracheal 
outgrowths from the sacs especially in the neighborhood of the 
cell clusters pass into the brain. They branch more or less pro- 
fusely among the cells and then send smaller branches into the 
fibrillar substance as shown in fig. 29, Pl. XXII, which repre- 
sents a camera drawing of the trachez of one of the mushroom 
bodies. Trachez almost identical in appearance with these, are 
found on all sides of the brain. Others, however, of a larger 


150 JOURNAL OF COMPARATIVE NEUROLOGY. 


size are also found that enter into the fibrillar substance more 
deeply, and gain their entrance along with some nerve fiber or 
bunch of fibers. 

In no case have I been able to make out a tracheal net- 
work such as described by Emil Holmgren (95) for the spin- 
ning glands of lepidopterous larvae. So far as bichromate of 
silver preparations thus far made show, the branchlets all termin- 
ate independently of one another. In some of my copper-hema- 
toxylin preparations there are, however, certain details that re- 
call Holmgren’s figures which lead me to suppose that a 
tracheal network may exist. 


Special Description—the Proto-cerebron. 


THE MUSHROOM BODIES. 


From this hasty general account of the brain and its con- 
stituent elements I pass to a more detailed consideration of its 
parts and choose for the first the mushroom bodies as being the 
most conspicuous and as the better serving for the orientation 
of other parts later to be described. 

These organs were first noted by Dujardin ( 50) in a study 
of transparent preparations zz ¢ofo. From their slightly lobular 
or folded appearance he compared them with the convolutions 
of the human brain and described them as ‘‘lobes 4 convolu- 
tions ”’ probably associated with the intelligence of the insect. 
As will be seen in the sequel, this supposition has very much 
more in its support than the crude or undeveloped morphologi: 
cal technique of that time was capable of bringing to light. 
Later they were again found by Leydig (64) who mentions 
them under the term ‘‘Gestieltes Korper.”” They were again 
described by Rabl-Rickhard (75) who, according to Viallanes 
(7), first correctly described the calyx. But it was not until 
Dietl ( 76) had applied microtomy and stained his sections with 
osmic acid that a complete description of their whole structure was 
obtained. Since that time they have been so often re-described 
for insects representing the different orders of hexapods that 
they may be considered as fairly well known structures very 


Kenyon, Zhe Brain of the Bee. I51 


characteristic of—especially the higher—hexapods. Special 
swellings found on the brains of certain of the crustacea 
have been compared with them, but it is to be seriously doubted, 
I think, whether such swellings or cellular heaps are properly 
to be homologized directly with them. In neither Retzius’ 
(90) figure of the brain of Astacus fluviatits nor in Bethe’s fig- 
ures of the brain of Carcinus menus can I find cells having the 
relations and the appearance of those that I find in the 
bee. I have noticed nothing resembling the structures in 
isopods, or amphipods, nor I have found indications of them in 
the brains of Pauropus, Polyxenus, juloid diplopods,  Scolo- 
pendrella, Lithobius, nor even in several forms of Thysanura 
that I have examined. If cells homologous with those filling 
the cup-like calyx of the mushroom bodies of the bee are at all 
present in these forms, they are so undifferentiated as to be in- 
distinguishable from the general mass of cells about them. 


The Calyx-Cup. 


The four bodies that in the bee and the Hymenoptera in 
general constitute what may be called the calyx-cup form the 
upper portion of the proto-cerebron, and are arranged side by 
side, a pair to each lateral lobe. In section, as shown in the 
photographs of Plates XIV and XV and in optical section in 
transparent unsectioned preparations, they are very noticeable 
from the curved masses of fibrillar substance that form the base 
and walls or calyx of each making them resemble optical sections 
of so many cups. Each cup holds a mass of cells that fill it to 
the brim. The walls of each lateral pair are nearly or quite 
contiguous, except distally where they diverge from one another 
leaving a space filled with a mass of small cells, as is also the 
space between each outer cup and the inner fibrillar mass of 
each optic lobe. Viewed from above they would appear elon- 
gated antero-posteriorly or with their included cells as four folds 
along the top of the brain, each extending backwards and in- 
wards toward the median line. The inner one of each pair is 
broadest anteriorly and somewhat overlaps its outer mate or 
appears to crowd it back. On the other hand the outer one is 


152 JouRNAL OF CoMPARATIVE NEUROLOGY. 


broadest and extends the farthest posteriorly. Other general 
details may be made out from the photographs. Below them 
antero-laterally, below their whole posterior extent, or occupy- 
ing the spaces between them and the rest of the proto-cerebron 
are to be found ganglionic cells of various sizes, but mostly of 
the small kind to be described elsewhere. There is also a small 
mass of cells continuous with the mass between the upper por- 
tion of each pair of cups, but which in the sections photo- 
graphed appear to be cut off from the larger mass. Each fibril- 
lar mass is outwardly clothed by a thin nucleated membrane. 
These are the bodies described by Dujardin as ‘‘lobes 


‘ ”? 


a convolutions. By later writers they are _ variously 
called. Leydig (64) described them as ‘‘Lappen mit Wind- 
ungen ;” Rabl-Riickhard (75) as ‘‘ Rind Korper ;” Dietl ( 76), 
Berger (7g) and Cuccati (gg) as ‘‘ Pilzhutformiger Kor- 
per ;” while Newton ( 79) introduced the name calyx, which was 
adopted by Packard (gg) and Viallanes (7 and gg). 

The bodies reach their highest development in the Hymen- 
optera and are much larger in the social wasps than in the 
honey bee. In Jlatta the lateral walls of the cups are much 
reduced, and in the Coleoptera the cup-like form is scarcely 
recognizable, while in Forficula and Acridum the fibrillar sub- 
stance only forms a broad plate. Even this is scarcely, if at 
all, recognizable in Dytzscus. In Zabanus and Somomya the four 
folds are reduced to two and in the former of these genera are 
scarcely to be distinguished by a comparison of their cells with 
those surrounding them. In the Hemiptera they are not dis- 
tinguishable at all. 


The Stalks of the Mushroom Bodtes. 


From near the middle of the lower surface of each cup 
a column of fibrillar substance passes into the main mass of the 
proto-cerebron. In the bee these are much shorter than in 
other forms. The inner one descends almost perpendicularly 
downward fora short distance where it is joined by the one 
from the outer cup, which is somewhat longer and has an in- 
ward and somewhat forward course. In transverse sections the 


Kenyon, Zhe Brain of the Bee. 153 


columns or stalks present a nearly circular outline and are 
distinctly delimited from the surrounding mass of fibrillar 
substance. 


The Roots of the Mushroom Bodies. 


At the place of union of the two stalks there are given off 
two masses of fibrillar substance of cylindrical shape, but with 
a diameter considerably greater than that of the stalks. One 
extends obliquely downwards towards the median plane of the 
brain where it nearly abuts against the one from the opposite 
side (fig. 2, Pl. XIV). Sometimes, as shown in the figure, 
the point of one may slightly overlap the other. Like the 
stalks, this structure is easily distinguishable from the surround- 
ing mass of fibrillar substance even in ordinary preparations. 

The other, which may be distinguished from the one just 
described, or inner root, by terming it the anterior root, ex- 
tends straight forwards and abuts against the peri-cerebral 
membrane. It is longer than the inner root, but has nearly the 
same diameter. In section it is nearly circular (fig. 1, Pl. XIV), 
and in the transparent preparations zz Zofo appears as a circular 
body in each proto-cerebral lobe, considerably above the an- 
tennal lobe. For most of its course it is well delimited from 
the surrounding mass, but anteriorly it becomes fused with the 
surrounding fibrillar substance on its lower inner side. In this 
region it may be seen to be entered by many irregularly branch- 
ing fibers, even in preparations stained with haematoxylin (fig. 
Tee ely SE): 


THE MINUTE STRUCTURE OF THE MUSHROOM BODIES. 


In sections cut in the frontal plane and treated with sul- 
phate of copper and hematoxylin the cells filling the cups of 
the mushroom bodies appear to be of two kinds, both arranged 
one above the other so as to present the appearance of tiers ar- 
ranged more or less radially with respect to the calyx as a cir- 
cumference. Laterally they are much larger than those in the 
middle, which in such sections forma dark colored triangle 
(fig. 1). Those at the side are much lighter. In sections 
of brains fixed and colored in von Rath’s platino-osmo-picric acid 


154 JOURNAL OF COMPARATIVE NEUROLOGY. 


mixture the contrast between the two kinds is noticeable only 
upon close inspection and then only as a difference in size. 
After staining with fuchsin brings out no further difference. 
The chromatin elements then appear more distinct and more 
numerous, but both seem equally well supplied. Nor is there 
a difference in the presence or absence of a small dark colored 
nucleolus in preparations treated after the copper-hematoxylin 
method. The slight chemical difference indicated by the differ- 
ence in depth of coloring shown by the latter method is, how- 
“ever, supported by a difference in the fibers given off from the 
cells, as will be seen a few lines further on. 

In similar sections of the brain of a wasp I find but one 
size in the cells of this region. But two sizes occur in the 
- cockroach according to Flogel (7g) and are doubtless to be 
found in other forms. But one or two kinds, the cells of the 
mushroom bodies in the hexapods generally, and of the regions 
compared with them in crustacea, are strongly distinguished 
from nerve cells elsewhere in the brain, and in the whole nerv- 
ous system for that matter, in being nearly devoid of extra-nu- 
clear protoplasm. This fact led Dietl to term them ganglionary 
nuclei, a term that was later altered by St. Remy (99) to the 
-equally characteristic one of chromatic cells. 

Between the tiers of cells fine fibrils and trachea make 
their way to the inner surface of the cup, the trachez also 
branching among the cells as already described or figured in 
Plate XXII (fig. 29). The nerve fibers may be traced, with 
considerable difficulty, however, tc the cells. In bichromate-of 
silver preparations both cells and fibers easily impregnate and 
where the section is in the right place or where too many elements 
have not become darkened, the fibers are readily traced into the 
fibrillar substance forming the calyx-cup (fig. 18, Pl. XXI), 

Here the fibers from the larger or marginal cells break up 
into a series of fine branches that again subdivide and produce 
a bushy or arborescent formation that reaches to the outer wall 
of the calyx and of which the drawing (fig. 18, Pl. XXI), al- 
though made with a camera, can give but a poor conception. 
The preparation from which this drawing was made is one of 


Kenyon, Zhe Brain of the Bee. 155 


the first that were fairly well impregnated, but not so well as 
later ones, and is far from showing the richness of branching 
seen in the latter. Further, the branches are delicate and the 
smaller ones covered with fine short processes that give the 
whole a more or less feathery appearance. The preparation is 
not black, but of a delicate reddish brown tint. The difficulty 
of representing them and the fact that the connections with the 
cell bodies are poorly represented in the other preparations 
caused me to be satisfied with the figure given. 

The processes of the smaller cells forming the pyramidal 
central heap, as shown in the same figure (fig. 18, Pl. XXI), 
also pass into and branch in the fibrillar calyx-cup. These 
branches resemble those of the other fibers, but, as shown, are 
far less numerous. Moreover the processes from the cells may 
be slightly smaller. 

As shown in the figure, the processes do not always follow 
the most: direct line to the fibrillar substance, but seem often to 
bend about the intervening cell-bodies. Consequently little de- 
pendence can be placed upon even the best of preparations by 
ordinary methods for tracing fibers for any considerable distance 
from the cell body towards the fibrillar substance. Fibers may 
readily be seen, however, passing from the cellular mass into 
the calyx, for between the two there is a narrow space but 
loosely filled with fibers, which may be seen more or less indis- 
tinctly in the photographs. 

In this space are to be seen small round deeply stained 
bodies, which upon close examination and by comparison with 
sections cut in a plane parallel to the inner walls or to the base of 
the calyx are seen to be composed of a large number of very fine 
fibers arranged parallel to one another and forming thus a close- 
ly crowded fiber tract. These radiate in all directions from the 
place of union of the calyx and the stalk from which they seem 
to come. Outwardly the fibers separate and gradually spread 
out over the whole inner surface of the cup. Such phenomena 
are best shown in bichromate of silver preparations, but may be 
seen in preparations by ordinary methods and have been fig- 
ured by earlier writers (Viallanes, Cuccati). 


156 JOURNAL OF COMPARATIVE NEUROLOGY. 


A further study of bichromate of silver preparations will 
show that each fiber from one of these bundles after its separa- 
tion from the rest becomes continuous with a fibrillar process 
entering the calyx-cup from one of the cells above. Thus there 
is formed a nerve cell of the general Y-type (fig. 18, Pl. XXI). 
No other nerve cells are found in the cups. 

Before continuing the consideration of the fiber just noted 
it may be well to mention the form of the cell bodies here and 
of those elsewhere. The larger number of successful impreg- 
nations of the cell bodies of the cups present a perfectly smooth 
outline. Very many, however, have from one to five or six 
projecting processes, which may branch. In this respect the 
smaller cells of the pyramidal group are exactly like the larger 
border cells, though such a condition did not happen to be 
shown in the section from which figure 18 was drawn. To de- 
termine whether they differed in this respect the sections of sev- 
eral brains were examined, resulting in the conclusion just given. 
Many of the cells resembled those shown in figure 19, Pl. XXI. 
Whether these processes are protoplasmic continuations of the 
cells to which they are attached or whether they represent 
merely filled up adjacent inter-cellular spaces, is a rather difficult 
question to answer. Here is one of the most serious difficulties 
presented by the bichromate of silver method. A few such 
cells I have seen in the optic lobes, but there as elsewhere cells 
with smooth contours seem to be the rule, and the rule might be 
taken to indicate an artifact nature for the processes in 
question. Further than this I find no reason for believing 
them to be artifacts. On the other hand, the fact that there 
was no evidence of undoubted artifacts in the immediate neigh- 
borhood of the cells described, together with the peculiar cir- 
cumstance of the extremely small amount of extra-nuclear pro- 
toplasm in the cells filling the cups and their generally crowded 
appearance, would indicate that probably the processes are real- 
ly parts of the cells with which they are connected. 

Returning to the fibers of these cells, it may be noted that 
it is the bichromate of silver method alone that is able to show 
their course and termination. Fortunately there is no difficulty, 


Kenyon, The Brain of the Bee. 157 


such as there is with the fibers of other regions, in determining 
their immediate relationships completely. The bundles of fibers 
already mentioned as radiating from the place of union of the 
calyx and stalk as though coming from the latter do so, or 
rather, to start from the parent cell, pass down into the stalk. 
At the junction of the stalk and the two roots each fiber divides 
into two branches one of which passes obliquely downward 
towards the median line, the other straight forward to the an- 
terior surface of the brain. Both terminate without further 
branching, and the inner group form the inner root, and that 
directed forward the anterior root of the mushroom bodies. 
Throughout their whole course from the entrance into the stalk 
to their termination in the inner and the anterior roots the fibers 
remain parallel, the only deviation from a straight line being 
a slight waving. They neither decrease nor increase in size 
but in the stalk they sometimes, though not always, ap- 
pear to be covered by fine short processes that can neither be 
described as thread-like nor tooth-like. These may be arti- 
facts, but I am inclined to think that their non-appearance 
might as well be accounted for by defective impregnation. 

There is found, then, in the mushroom bodies a nerve cell 
with a smooth or irregular body sending a process into the 
fibrillar substance forming the calyx, branching there profusely 
aud sending a second process from the first down through the 
stalk and forming an inner and an outer branch. As will be 
seen beyond all doubt a little farther on, the branching process 
in the calyx is the dendrite and the second process is the neu- 
rite. In the relation of its neurite to other fibers it recalls to 
some extent the relation between the neurites of the cells of 
the granular layer and the dendrites of the cells of Purkinje in 
the mammalian cerebellum, but as a whole the cell seems to be 
unique among all known nerve cells. 


Fibers Ending in the Calyx. 


In bichromate of silver preparations, besides the branch- 
ing dendrites of the cells just described, there may also be seen 
a large number of other fibers all somewhat coarser than the 


158 JouRNAL OF CoMPARATIVE NEUROLOGY. 


fibers of these cells and permeating and branching in the calyx 
in all directions (figs. 23, 26, 27, Pl. XXI). Often they are 
impregnated almost to the exclusion of the fibers from the cells 
above. At other times both appear, but then they most often 
obscure one another, making it necessary to section a large 
number of brains in order to find cases sufficiently free from a 
repetition of details to enable one to determine the form and 
relationships of the various fibers correctly. When such cases 
occur it is found that the fibers terminating in the calyx all 
.-make their entrance in the neighborhood of the junction of the 
calyx and stalk, or from the under surface of the cups. All 
appear to send a main fiber in a circle about the head of the 
stalk and from this to radiate numerous branches to the more 
distant parts. 

The difference in size and mode of branching of these en- 
tering fibers allows two kinds to be distinguished. In the 
smaller kind of branches radiated from the region of the stalk, 
one slender kind branch more or less irregularly and repeatedly, 
terminating finally in an irregular swelling such as shown in fig. 
27, which represents a magnification of about 583 diameters. 

The larger kind (fig. 23) radiate off from the portion en- 
circling the head of the stalk a number of stocky branches that 
after penetrating the surrounding mass of fibers for a relatively 
short distance branch out into a cymose head. Other portions 
aparently of this same fiber pass out into more distant parts of 
the calyx. The figure given was drawn with as much care as 
possible and well represents what was seen, but appearances 
are often deceptive, and I am inclined to think there are two 
kinds of fibers here instead of one as represented. It may 
happen that fibers overlie one another in such a way as to ap- 
pear as one continuous fiber, a deception that, especially in 
thick sections, the most careful focusing will not always avoid. 
The fibers shown in the figure as immediately surrounding the 
junction of the stalk and calyx appear to be restricted to the 
region covered by the median group of small cells. The others 
appear to have gained an entrance to one side of the stalk and 


Kenyon, The Brain of the Bee. 159 


take in a larger region, besides appearing to have a different 
mode of branching. 

The difference in size between figs. 23 and 26 is mostly a 
difference of magnification, which in the former is 121 and in 
the latter a 102 diameters, yet the fibers represented in fig. 23 
are actually the coarser. 

Just where either of these two kinds, the fine and the 
coarse fibers originate, or what other parts of the brain are con- 
nected with the mushroom bodies by them is yet to be deter- 
mined. Almost all that can be said is that six different tracts 
of fibers terminate here, two from the optic lobe, one from the 
ventral region, and three from the antennal lobes. To these 
may probably be added a seventh of a commissural nature. 
These tracts will be described further on. 


The Fibers Ending in the Stalk and the Roots. 


In certain places (fig. 1, Pl. XIV), as already pointed out, 
fibers entering the anterior roots of the mushroom bodies may 
be very readily seen in sections treated after the copper-hama- 
toxylin method. In preparations by the bichromate of silver 
method others may be distinguished. One large tract traced 
to its cells of origin behind and between the stalks of the mush- 
room bodies by Viallanes (87) in his study of the wasp spreads 
out on the side of the anterior half of the anterior root and 
branching sends one branch into it. The fibers of this 
pass nearly horizontally across or obliquely downwards to the 
opposite side giving off from their upper side several sub- 
branches (fig. 21, Pl. XXI) that after passing a greater or less 
distance upward break up into a thickly bushy head. These 
all occupy almost the same level so that often in specimens 
treated with osmic acid or with hematoxylin, especially the 
latter, the appearance is presented of several bands crossing 
transverse sections of the roots. Figures 3 and 4 show these 
bands excellently, and represent the two roots as seen in a 
slightly oblique section treated with copper sulphate and a so- 
lution of hamatoxylin made according to one of Weigert’s for- 
mulz. Bands of this sort occur throughout the whole extent 


160 JourNAL oF COMPARATIVE NEUROLOGY. 


of the root. In the inner root also similar bands, though less 
conspicuous, are found, but there instead of being transverse, 
they are longitudinal. 

The fibers just described are more or less coarse, consider- 
ably more so than fibers entering the lower outer side of the 
root behind them (fig. 28). The sub-branches of the second 
kind of entering fiber also take an upward course and break up 
into a series of branchlets, which do not, however, form a com- 
pact bushy head. Where their cells of origin are situated Iam 
unable to say further than that subsequent studies will probably 
show them to be situated somewhere beneath the calyx-cups, 
either to one side of or behind the stalks. Their outward-go- 
ing branches seem to penetrate into the lower lateral depths of 
the brain. A third fiber enters this same region at a lower 
level and appears to come from the lower lateral side (fig. 28). 

Behind these fibers near the junction of the roots, or in the 
plane shown in fig. 1, Pl. XIV, onthe left, fibers are found 
entering on the lower inner side which present a more delicate 
appearance and are otherwise different from those first described 
(fig. 28). They branch more or less dichotomously and end in 
a tuft of branchlets which in preparations with osmic acid or 
hematoxylin give rise to the broken appearance shown in the 
section of the left root in fig. 1, a comparison with which will 
show that fig. 28 is not properly oriented and that it should be 
viewed from the corner of the plate so as to make the branches 
crossing the dotted ring take a nearly horizontal course. 

In the region of the junction of the two roots and the two 
stalks rather coarse fibers are found (fig. 17) considerably resem- 
bling those first described, whose more or less bushy terminals 
along with those last described produced the knotted appear- 
ance described by Rabl-Rickhard (75) for the ant as ‘‘ 2-3 
small clefts rounded off above,” and the lenticular spots only 
faintly shown in fig. 2. The same appearance as that described 
by Rabl-Riickhard is found in the bases of the stalks in the bee, 
but the section represented in fig. 2 is too far back to show it. 
To imagine the figure, however, all that is necessary is to sup- 
pose the lenticular spots to be larger and united at their bases. 


Kenyon, The Brain of the Bee. 161 


The lighter places between the spots are filled by the fibers of 
the mushroom body cells alone (fig. 17). 

In the stalk my preparations show the existence of incom- 
ing fibers resembling the upper ones represented in fig. 28 
more than any of the others. But details are not sufficient to 
enable one to determine their origin or the destination of their 
outer branches any better than, in fact not so well as, those en- 
tering the anterior root. 

Very nearly the same thing may be said with respect to 
the fibers entering the inner root. Some of these appear to be 
connected with the central body later to be described, others 
seem to come from the opposite lower side of the brain. Their 
branches in the root are of the same general type as those rep- 
resented in fig. 21, but the sub-branches are much shorter, 
more uniform, and end in avery much more compact head of 
branchlets. These of course cause the longitudinal banded ap- 
pearance already mentioned. 

Besides these six or seven groups of entering branches 
there are to be found numerous fibers irregularly branching over 
the surface or near the surface of the roots, which may also send 
small branches into them (fig. 22, Pl. XXI), and here may be 
noted a fiber seen in one section only that is of no little in- 
terest. Whether it belongs to any of the regular groups I am 
unable to say definitely, but think from its appearance that it 
must be a branch of one of the irregular fibers just mentioned. 
It is figured on plate XXI (fig. 25). It enters the right ante- 
rior root near its terminus from some little distance in the lateral 
part of the brain and is there of considerable size. But almost im- 
mediately upon entering it decreases in volume and after branch- 
ing becomes connected with the neurites of the mushroom body 
cells. The junction appears as complete as though there existed 
here a veritable fusion. So fine are the fibers that the prepar- 
ation also seems to show that the parallel fibers from the mush- 
room body cells have branches in this place. The latter condi- 
tion cannot, however, be the true one, for in all other prepara- 
tions, which are numerous, no such branching is shown. With 
Lenhosseék ( 95) I agree that in general the bichromate of silver 


162 JOURNAL OF COMPARATIVE NEUROLOGY. 


method, usually showing only silhouettes, is defective and very 
apt to mislead in the matter of fibrillar connections. But in the 
specimen in question viewed with a Leitz ocular No. 2, and ob- 
jective No. 5, there is no more reason for doubting the contin- 
uity of the lines shown than there is in fibrils shown in prepar- 
ations stained with hematoxylin or any other histological color- 
ing matter. They are not superimposed, nor is there any reason 
for thinking them to be of tracheal nature. 

When we consider the nature of the method used, it should 
not appear strange that fibrils in very close contact should be so 
impregnated with bichromate of silver as to cover up all evi- 
dences of separation. In fact it seems to me that such apparent 
evidences of continuity are to be expected, without, however, 
even raising a doubt as to the complete validity of the evidence 
obtained from the generality of terminations shown and from 
histo-ontogeny. 

Before leaving the matter it must be noted that, although 
the figure given is a camera drawing made with the aid of care- 
ful focussing, it does not correspond with what appears when the 
camera is removed. Then the fibres a, d and ¢ are seen to belong 
to the parallel fibres which are continued upwards beyond their 
junction with the fibrillar branches of the incoming fiber. 


THE FUNCTION OF THE NERVE-CELLS OF THE MUSHROOM BODIES. 


Ever since Dujardin (59) discovered the mushroom bodies 
and pointed out the relation between their size and the develop- 
ment of insect intelligence, nearly every writer on the subject 
of the hexapod brain who has referred to the matter of intelli- 
gence has recognized the fact. Leydig (64) thought them con- 
nected with the intelligence of the animal, but added that they 
also seemed to bear some relation to sight, or to the ocelli. Rabl- 
Rickhard (75) thought the same, except that he pointed out 
that Leydig’s second supposition is incorrect, since in blind ants 
the bodies are found to be well developed, while the optic 
lobes and the nerves of the ocelli are absent. Forel (74) 
pointed out the same facts and added that the mushroom bodies 
are much the largest in the worker ant, and that those in the 


Kenyon, The Brain of the Bee. 163 


female are larger than those in the male. Two years later 
Brandt (76) affrmed the same condition to be true for the Hy- 
menoptera generally. Berger (78) agreed with Dujardin and 
the rest. Dietl (76) was inclined to dissent, and Flogel (7s) 
seemed to consider the roots and stalks as a framework (Geviis?) 
for the rest of the cerebral substance. Nevertheless the tabu- 
lar summary of his study on the different orders that he gives at 
the end of his paper is of no inconsiderable value in showing the 
truth of Dujardin’s inference, so far as comparative anatomy is 
able to do so. The physiological experiments of Faivre (57) 
and others, which have lately been repeated and perfected in the 
excellent work of Binet (94), are of great value in showing that 
the seat of intelligent control of an insect’s movements is in 
the dorso-cerebron. Thus far, however, no attempts seem to 
have been made to determine experimentally the function of the 
mushroom bodies as parts distinct from the rest of the dorso- 
cerebron. Experiments are badly needed to complete our 
knowledge of the structures. 

All that Iam able at present to offer is the evidence from 
the minute structure and the relationships of the fibers of these 
bodies. This seems to be of no inconsiderable weight in sup- 
port of the general idea started by Dujardin. For in connec- 
tion with what was made known by Flogel and those before him 
and has since been confirmed and extended by other writers, 
one is able to see that the cells of the bodies in question are 
much more specialized in structure and isolated from the gen- 
eral mass of nerve fibers in those insects where it is generally 
admitted complexity of action or intelligence is greatest. Con- 
sidering the calyx and the cells above it alone, one finds that 
according to Flogel neither are recognizable in the Hemiptera, 
that in Diptera (Zabanus) and Odonata (deschna) the bodies 
are recognizable from those in the neighborhood only by a 
slight difference in the size of the cells representing them ; that in 
Lepidoptera the calyx-cup is very small or entirely absent, that 
in the Coleoptera (Zenthrido, Cynips) the calyx is not so ex- 
tended as to be very well recognized asa cup, and that in the 
Orthoptera the lateral walls of the cup are either very small 


164 JOURNAL OF COMPARATIVE NEUROLOGY. 


(Blatta) or entirely absent (Forficula, Acridium, etc.) ; while in 
the Hymenoptera the calyx is folded upwards so as to give 
room for many more neural elements and connections. 

If one considers the little that has thus far been made known 
by means of the bichromate of silver and the methylen-blue 
methods concerning the cerebral nerve fibers in other inverte- 
brates, one finds that sensory stimuli are transferred to a num- 
ber of irregular cells or fibers distinguished roughly by the term 
association fibers and from these transferred to motor or other 

_fibers bearing efferent neural impulses. And from such a com- 
parison it appears that in the groups of insects certain of these 
association fibers have gradually been set apart from the rest so 
as to render it possible for an entering stimulus to become an 
efferent impulse by taking a direct or an indirect course. In 
the former case the course is to be compared with the course ofa 
stimulus producing reflex action in man. Later on it will be 
seen that the present assumption of the existence of fibers per- 
mitting such a course is fairly well founded. Thus a sensory 
impulse from an optic lobe or from an antenna may reach .the 
fibers going to the mouth parts through possibly but one asso- 
ciation fiber, 

In the other case it may take a special or somewhat indi- 
rect course to and through the fibers of the cells of the mush- 
room bodies and from them reach the efferent fibers through 
the processes of one or more association cells. From those facts 
it seems far within the bounds of reason to suppose that the 
nerve cells of the mushroom bodies rendered so prominent by 
their specialization of form and position are the elements that 
control or produce actions that one distinguishes by the term 
intelligent. 

An analysis of these actions is needed, but that is beyond 
the limits of this paper. Suffice it to point out that the exper- 
iments of Binet (94) and others show that when the connections 
between the dorso- and ventro-cerebron are destroyed the phe- 
nomena afterwards observed are similar to those seen ina 
pigeon or mammal when its cerebral hemispheres are removed. 
One difference is notable, namely, that the operation of eating 


Kenyon, The Brain of the Bee. 165 


in the case of the insect appears to be reflex ; for when one, vivi- 
sected as just indicated, is placed upon its food it begins and con- 
tinues to eat. In the other case the animal must be fed. The vari- 
ous bodily movements of the insect, however, after a time take it 
away from the food, and once away it is wholly unable to return. 
Such insects when passing food even almost within touch of their 
palpi are wholly unable to change their course so as to secure 
it, although the movements of their palpi plainly show that its 
presence has affected them. Whether this would still remain 
the case when the vivisection is of sucha nature as to allow 
sensory stimuli from the antenne or visual stimuli to act through 
the reflex channels already mentioned is as yet unknown. From 
a consideration of the facts that I have pointed out and the 
facts demonstrated by comparative anatomy and embryology 
relative to the homologies of the antenne one is warranted in 
expecting that the animal would still lack the power of drecting 
its movements. 


THE CENTRAL BODY. 


This peculiar body was first made known by Dietl ( 76) as 
‘the fan-shaped structure,’” and was two years later described by 
Flogel (78) as the central body, a term that has since been gen- 
erally employed. It lies above and behind the ends of the inner 
roots of the mushroom bodies and is wholly composed of large 
nerve fibers and fibrillar substance, plus, doubtless some tra- 
chealclements (tgs. 2,.5,°0,.7, 9, O,.G0.). ) Antew of its, con: 
nections I have been able to ascertain, but very much yet re- 
mains to be learned, so much in fact that any assertion as to its 
functions can be but little better than a mere guess. Viallanes 
(87) describes it as connected with all the surrounding parts of 
the brain, but that is an assertion too sweeping for the facts 
known to me after an application of the bichromate of silver, 
the copper-hematoxylin and platino-osmo-picric acid methods, 
and was much too sweeping for the actual facts known to Vial- 
lanes. Berger (78) recognized the body in Dytiscus as a lenti- 
cular body receiving a bundle of fibers from each side which 
entering broke up into its individual fibers, some going 


166 JOURNAL OF COMPARATIVE NEUROLOGY. 


out anteriorly, some posteriorly, or, in short, as a center of 
disassociation. 

The body throughout my diagrams and figures is distin- 
guished by the letters c. 0. 

In the honey bee this body as a whole has a very much 
distorted spherical shape. In frontal sections the fan-shape 
described by Dietl and Berger is apparent (fig. 5), though the 
outline presented might be better described as reniform. Viewed 
from above or in horizontal sections it has a somewhat trape- 
zoidal outline. 

_ Internally, as shown by sections in the frontal plane, it is 
divided by a space filled with nerve fibers and trachez, into an 
upper and larger portion covering a lower and smaller portion. 
Both are seen in preparations by ordinary methods to be cor- 
posed of fibrillar substance, but in frontal sections the lower 
usually stains more deeply (fig. 5). Antero-posteriorly the 
upper portion is also much the larger and considerably over- 
hangs the other, partly covering two masses of fibrillar substance 
a little farther on to be described as tubercles. : 

Taken as a whole, fibers seem to reach it from or leave it 
in nearly all directions ; but the two parts seem to be supplied 
somewhat differently. Those entering the lower are seen to 
originate from cells above the antennal lobes and upon reaching 
the lower lateral edges to take a transverse course below the body 
and send several branches upwards that subdivide arborescently 
producing a compact mass of branchlets that recall the arborescent 
and bushy terminations of the association fibers in the roots. of 
the mushroom bodies (figs. 32 and 37). As in the case of these 
latter fibers, it is to the compact branching mass of fibrills 
that is due the depth of color so noticeable in preparations 
stained with osmic acid or with hematoxylin. Other fibers 
either pass out or enter from the fibrillar substance of the brain 
immediately in front, while branches from association fibers in 
the anterior region seem to enter the anterior end and the 
posterior lower end of this portion of the body (fig. 40, PI. 
XXII). 

In the upper portion the same arborescent method of 


Kenyon, Zhe Brain of the Bee. 167 


branching also occurs, and gives rise to the wavy upper contour 
shown in figs. 5 and 6. But here the fibers originate from cells 
above the body in the median plane or from cells behind it and 
after passing behind the body enter the cleft separating the two 
portions, where many of them are gathered into bundles with an 
antero-posterior course. These recall the radiating bundles seen 
in the cups of the mushroom bodies. Other fibers also send their 
branches into it from its outer surface (Pl. XVIII). 

Just behind the inner roots of the mushroom bodies large 
numbers of fibers seem to be given off, some of which pass 
downwards to the neighborhood of the cesophageal foramen, 
while others take a more lateral course (fig. 5). It is this ap- 
pearance, which is very noticeable in preparations by ordinary 
methods, that doubtless led Viallanes to make the assertion re- 
ferred to. When impregnated with bichromate of silver these 
fibers seem to originate from cells situated somewhere behind 
the central body and, after passing over its surface and sending 
branches in to it, to take the course described and shown so well 
in fig. 5. Some of them are probably the neurites of a group 
of cells whose dendritic branches form a commissure immedi- 
diately behind the central body (fig. 9, Pl. XVI) and send their 
sub-branches into its posterior surface (Pls. XVII and XVIII, 
cells 5 and 6). 

Another group of cells lying above the body in the me- 
dian plane send their processes backwards over its surface and 
after sending a branch to what will soon be described as the 
fibrillar arch pass down behind and enter the space separating 
the upper and lower portions of the body and branch in the form- 
er portion (cell 2, Pl. XVIII). Fibers from cells situated behind 
the body and above the fibrillar arch also make the same con- 
nections. Another group of fibers originating from cells above 
the antennal lobes pass obliquely inwards to the space just be- 
hind the lower half of the body and passing upward in front of 
the glomerules end in the upper half. What connections are 
made by them I have not thus far been able to ascertain. No 
fibers passing between any portion of the central body and the 
calices of the mushroom bodies have been seen, and whatever 


168 JOURNAL OF COMPARATIVE NEUROLOGY. 


indications have been noticed of connections with the inner 
roots of these bodies are of such a fragmentary and otherwise 
imperfect nature as to render their existence very doubtful. All 
that may be safely said is that it is connected with the fibrillar 
arch, possibly the ocellar glomerule, with the mass of associa- 
tion fibers immediately in front, with the lower lateral regions 
of the proto-cerebron, and with the trito-cerebral region form- 
ing the cesophageal commissure, where its fibers may possibly 
connect with others entering from the ventral cord, or with 
' those belonging to the ventro-cerebron. 


THE TUBERCLES OF THE CENTRAL BODY. 


Behind the lower portion of the central body and partly 
covered by the overhanging upper portion (fig. 6, Pl. XV) are 
two small round, deeply staining masses of fibrillar substance to 
which. Viallanes (§7) applied the term of ‘‘tubercules du corps 
central.’ Just what their relations are to the central body, or 
what may be the origin of the fibrils forming them I have not 
been able to determine definitely. But from the fact that in 
thick unstained sections one can distinguish in the immediate 
neighborhood behind them masses of fibrillar substance of a sim- 
ilar globular appearance I am inclined to think them globules of 
the same kind and having the same or similar relationships as the 
latter. My preparations do not satisfactorily show fibers leading 
into or going from them. But there are indications that the mass 
of smaller glomerules is formed by terminations of the nerves 
from the ocelli very much as the so-called olfactory glomerules 
later to be described are produced by the terminations of the 
fibers from the antennae, and for that reason I choose to call 
them ocellar glomerulz. This particular region of the brain 
along with the central body needs considerably more light than 
I am at present able to throw upon it. 


THE FIBRILLAR ARCH. 


This peculiar structure may as well be considered here, 
since it is connected with the central body, behind which and 
somewhat below the level of whose upper surface it lies. It is 
a rod-like mass of fibrillar substance that upon either side rises 


Kenyon, Zhe Brain of the Bee. 169 


from but is scarcely connected with, the mass of fibrillar sub- 
stance behind the union of the roots and stalks of the mush- 
room bodies and passes directly across the median line through 
the large mass of cells filling the posterior median furrow, thus 
forming a sort of commissure described by Viallanes (7) as 


y 


‘‘le pont du lobes cerebraux.”” It does not form a commis- 
sure, such as is ordinarily understood by the term, even though 
it connects the two lateral halves of the brain, and seems much 
better denominated by the term ‘‘gabelformiger Korper’’ em- 
ployed by Cuccati (ss) or better still by the one that I have used. 
The central portion of the arch may be seen in figure 8, while 
in figure 9, several sections below, the pillars upon which it rises 
appear as two dark spots, one on either side of the enormous 
nerves from the ocelli. 

In preparations treated with copper-sulphate and hama- 
toxylin or by the platino-osmo-picric acid method of von Rath, 
large nerve fibers may be distinguished passing irregularly along 
its outer surface, and seemed to originate from cells situated on 
both sides of the brain near its ends. Viallanes (87) mentions 
fibers passing from it to the central body, a connection that as 
already intimated my preparations with bichromate of silver 
show to actually exist. A group of fibres from cells in the 
median plane above the central body send each a short branch 
that ends here in a tuft of branchlets (fig. 2, Pl. XVIII) and 
then continues on down into the central body as described when 
speaking of that structure. This short branch and its tuft-like 
ending without doubt represents the dendrite and the larger 
branch to the central body the neurite of the cells of this group. 
Other groups situated above and both before and behind reach 
the central body after branching in exactly the same manner. 
Vialianes (g7) supposed that possibly the structure received 
branches from the ocellar nerves, but was far from being sure. 
Whether his supposition is true my preparations thus far made 
do not enable me to decide definitely, but from the fact that 
the nerves in question seem to begin to break up or become 
less numerous in this region and from a few indications in bichro- 
mate of silver preparations I am inclined to think that possibly 


170 JOURNAL OF COMPARATIVE NEUROLOGY. 


his supposition will sometime be found to be true. What con- 
nections are made by the fibers mentioned as covering the outer 
surface of the structure, I am not prepared to say. 


THE ASSOCIATION FIBERS OF THE PROTO-CEREBRON. 


The fibrillar substance of the regions surrounding the 
bodies already described, or the rest of the central proto-cere- 
bron, is composed of the association fibers coming from, and the 
tracts of fibers going to, these bodies, together with bundles of 
, fibers entering it from various groups of cells and giving rise to 
a great number of irregular fibers to be classed under the gen- 
eral term of association fibers. In only a few cases have the 
cells from which these fibers originate been impregnated along 
with their fibers so as to make their relationships perfectly clear. 
What little has been learned I have endeavored to bring out in 
the diagrams that follow as well as in the camera sketches in 
the last two plates. 

The fibers, as may be readily seen in fig. 1, take various 
courses, binding the upper, the lower and the lateral parts to- 
gether. Fibers originating from cells above the antennal lobes 
pass up on the inner side of the anterior roots of the mush- 
room bodies and branch arborescently in the region just above 
them (fig. 31 and fiber 10 in the diagrams). Just below the 
root a small branch is given off that passes outward horizon- 
tally. Other fibers seen in preparations by the copper-hema- 
toxylin method in front of the plane of fig. 1 seem to originate 
from cells above the antennal lobes, and, after passing upwards 
near the median line as rather large fibers, bend over the ante- 
rior roots of the same side and become lost in the fibrillar sub- 
stance (Pl. XVII). The fibers of the cells in the median line 
under the edges of the calyx-cup and in front of the central body 
also seem to branch in this region, for I find no indications of 
their forming a chiasma and passing into the lower regions of 
the brain as described by Viallanes (84) for the Orthoptera, 
and by Cuccati (ss) for Somomya. Other fibers of unknown 
origin (fig. 40, Pl. XXII), but of a very conspicuous appear- 
ance branch rather profusely in this region of the fibrillar sub- 


Kenyon, The Brain of the Bee. Tear 


stance and send branches back into the central body. Others 
bend over the inner roots and send branches upwards and across 
to the region immediately above the base of the anterior root of 
the opposite side (fig. 32, and fiber 50, Pl. XVIII). Another 
seems to arrive, (whether from over the top of the central body 
or from somewhere beneath the calyx-cup, is difficult to decide, ) 
and branches near the junction of the two roots, one branch 
going toward the central body, the other straight downward 
(fiber 21, Pl. XVII). A similarly perpendicular fiber is also 
found on the outer side of the root (fiber 20, Pl. XVII). Still 
another (fig. 32) seems to arise as a small fiber from over the 
central body, passes downward to the opposite side, where it 
becomes much enlarged, and gives offnumerous branches, these 
even passing out under the anterior root. It may possibly be 
one of a group of fibers (fiber 49, Pl. XVIII) originating from 
a group of cells near the fibrillar arch and passing directly for- 
wards over the central body. 

A horizontal fiber (fig. 22, Pl. XXI) originates somewhere 
at the side of the brain and after giving off several branches 
that spread out over the top of the anterior root ends in an ar- 
borescent system of fine branchlets in the upper part of the 
middle portion of the cerebron just across the root. 

The fibers connecting the anterior and posterior region are 
most striking in appearance and are of considerable extent. 
One (fig. 30) apparently originates from cells in the region 
above and behind the antennal lobe and passes backward to one 
side of the median plane giving off a branch that subdivides 
and passes to the median anterior root, one passing up close to 
it and the other to a lower level. After giving off this branch 
it becomes very much smaller and passes on backwards, giving 
off several small branches meanwhile. Near the posterior 
wall it divides into two branches of small size, but of consider- 
able length, the one going directly upward and_ branching 
among the ocellar glomerule, the other directly downward into 
the cesophageal commissure and the ventro-cerebron. 

Another fibre (48) equally, if not more peculiar, is shown 
in figure 34. This appears to originate from a cell situated 


172 JOURNAL OF COMPARATIVE NEUROLOGY. 


behind the inner root of the mushroom bodies (providing one 
may judge from the appearance of the fiber and the direction 
of its branches). Just behind the inner root and to one side of 
the base of the central body it becomes very much enlarged, 
gives off two relatively short branches and then one long one 
that passes up over the inner root and over the anterior root 
just in front of the junction of the two with the stalks, as shown 
by the dotted lines in the figure. Another branch a little fur- 
ther forward passes up to and branches in the fibrillar region to 
‘one side of the central body, while in front of this a number of 
short branches are given off that branch very profusely in the 
region to one side of the base of the central body. The main 
fiber continues forward to the neighborhood of the antennal 
lobes giving evidence of further branching, but on account of 
being either unimpregnated or cut off can not be traced farther. 

The fiber shown in figure 35 is nearer the median plane 
and appears to come from the cesophageal commissure, passing 
upward and then directly forward under the central body, giv- 
ing off along its course several branches that run back into or 
towards the commissure and then four very long though small 
branches that pass upwards in front of the central body ending, 
so far as can be traced, above the level of the top of the latter. 
One of the four passes farther forward and gives off a branch 
that turns back into the neighboring region to one side. 

In figure 33 a more peculiarly twisted fiber (35) is shown, 
which was found in the lateral region of the proto-cerebron be- 
low and to one side of the anterior root and behind the optic 
body, near which it may possibly originate. Its base is 
at the right of the figure. It passes outwards for a short dis- 
tance then backwards, gives off a branch that turns upon itself, 
divides again several times, some of the branches going up- 
wards, the rest downwards and forward. There it passes down- 
ward for a short distance and divides into two large branches of 
almost equal size, one going backward, so far as traceable, ap- 
parently to the posterior region of the brain, the other forward, 
twisting above the parent stem and finally breaking up into a 
number of branches that terminate behind the central body. 


Kenyon, The Brain of the Bee. 173 


The Deuto.cerebron. 
4 THE ANTENNAL LOBE. 


The antennal lobes comprise by far the larger portion of 
this division of the insect brain, especially so in the honey bee, 
and the rest of the Hymenoptera for that matter, where the 
remaining small portion is so fused with the proto- and the 
trito-cerebron as to leave little or no trace of separating 
boundaries. 

Considering first the antennal lobes, one may say that sec- 
tions cut in various planes show that each lobe is composed 
internally of a nearly spherical mass of fibrillar substance con- 
nected with the rest of the brain by a small neck of the same 
substance and surrounded on all sides by a large number of cells, 
some of which belong to it and some to the proto- and the 
trito-cerebron, and fill in the deep spaces between it and these 
two parts. Inside of this sphere and near or composing its 
periphery one finds small globular masses that by any of the 
ordinary methods of staining take a much greater depth of 
color than the more internal part, and in them one recognizes 
the great non-nucleated cells of Leydig (64) and Rabl-Rick- 
hard (75), which were first more correctly described by Dietl 
(76) as olfactory bodies. Flogel (7s) claims the honor of point- 
ing out their true significance, for, he says, he called attention 
in an unpublished communication to the Kieler Physiologische 
Vereins, July 30, 1874, to the fact that they are not cells. 
Fig. 1 shows the bodies or glomerule very plainly. 

Between them even in ordinary preparations processes from 
the bordering cells may be traced to the inner fibrillar sub- 
stance, but it is doubtful whether branches entering them may 
be seen in such preparations. Cuccati (ss) regarded them 
along with the rest of the mass as made up for the most part 
of branches of these processes that by subdividing formed with 
whatever other fibers that might be there a sort of net work. 
The only difference that he supposed to exist between them and 
the central portion of the mass was in the greater fineness of 
the meshes of the net. But while his supposition with respect 


174 JOURNAL OF COMPARATIVE NEUROLOGY. 


to the relation of the glomerule to the surrounding cells is cor- 
rect and even though some of my preparation are so strongly 
impregnated with bichromate of silver as to present consider- 
able resemblance to a meshwork, I am inclined to believe other 
preparations showing a tuft-like system of branching illustrate 
the true condition (fig. 10 and 16, Pl. XXI), and that the idea 
of connection by contact holds as well here as elsewhere. As 
already pointed out cases might occur in such _ prepara- 
tions in which the darkened fibers might appear perfectly con- 
' tinuous, so as not even to be explained by Lenhossék’s sugges- 
tion of a silhouette of superimposed fibers, and still not be 
fused or continuous. Happily such deceptive appearaces 
seem to be rare except in too heavily impregnated preparations. 

When impregnated with bichromate of silver the glom- 
erules strongly recall the appearance of the olfactory glomerule 
figured by Retzius (92) for the vertebrates, and may perhaps be 
very well called olfactory glomerule. But exactly what their 
relations may be with the surrounding cells and the incoming 
fibers I am not able to say definitely. In none of my prepara- 
tions is an unbroken connection shown between them and the 
cells clothing the lobes, the fragmentary appearances, however, 
amply warrant the supposition that the processes of the cells 
make their way between the bodies and after passing some dis- 
tance in the middle mass of fibers send a branch into one of the 
glomerulz and another one into the proto-cerebron (fig. 10, 16 
and 24, Pl. XXI). Whether the process from a single cell sends 
branches to more than one body does not appear, but if one 
may judge from the form of the cells of the mushroom body, 
probably it does not. 


THE ROOT OF THE ANTENNO-SENSORY NERVE. 


Besides the fibers just described there are to be distin- 
guished two other kinds. One of these composes the root of 
the antenno-sensory nerve, which in preparations with osmic 
acid or by the copper-hematoxylin method may be traced from 
its entrance into the lobe on the lower anterior surface nearly to 
the posterior upper surface. Throughout its course it gives off 


Kenyon, Zhe Brain of the Bee. 175 


numerous fibers and branches. My preparations by the better 
neural method that are favorable for tracing it are not numer- 
ous. Fig. 16 represents one in which the fibers shown are all 
that are impregnated in this region. Here a fiber is seen com- 
ing in from the nerve and at some distance from its entrance 
bending aside to a glomerula from which another fiber passes 
out and then on towards the proto-cerebron. The other fiber 
soon after entering breaks up into three branches one of which 
terminates ina glomerula. The tuft-like termination was not 
seen when the drawing was made, but after the latter was fixed 
to the plate a re-examination of the specimen brought it to 
light, but with scarcely sufficient distinctness to be drawn with 
a camera. 

If one is to credit, as one must, the long list of writers 
since the time of Newport dealing with the antenne anatomi- 
cally and physiologically, one must conclude that these organs 
have at least two, and in many cases, if not always, three differ- 
ent functions. That they are both tactile and olfactory has 
long been known, and recently Child (94) has brought forth 
very good anatomical reasons for their being also auditory. 
Such being the case one might expect to find as many different 
kinds of terminations in the antennal morula. 

There is a striking resemblance between the tufts forming 
the glomerule and those forming the olfactory glomerulz in 
mammals as described and figured by Retzius ( 92 @); and, since 
in the ventro-cerebron where sensory fibers from the oral nerves 
undoubtedly terminate no such glomerule are found, one may 
be very easily led to the conclusion that the glomerule are for 
olfactory terminations. But Iam unable to produce evidence 
of other terminations and must for the present conclude that 
the glomerulze are formed by the terminations of all kinds of 
fibers from the antennze. This conclusion is supported by the 
fact that the tubercles of the central body, in which fibers from 
the ocellar nerves apparently terminate, and the ocellar glome- 
rule have a very similar appearance. 

Another kind of fiber is shown along with its cell in fig. 
15. This was found in the posterior ventral regions and bears 


176 JouRNAL OF COMPARATIVE NEUROLOGY. 


a slight resemblance to cells and fibers that are undoubtedly of 
efferent nature. It may bea portion of such a one that by 
branching among the olfactory glomerule as indicated in the 
figure makes a reflex course from the afferent antennal fibers to 
the antennal muscles. Still it may be only an association fiber 
of which there may be many in the antennal lobe, though their 
appearance does not distinguish them as such as readily as in 
the case of the association fibers of the rest of the brain. 
_ Whatever other fibers there are that might be classed as such 
are of small size. Some of them appear to pass into the neck 
already spoken of as connecting the lobe to the rest of the 
brain. 


THE ROOT OF THE ANTENNO-MOTOR NERVES. 


The small internal antenno-motor nerve supplying the an- 
tennal muscles within the head appears to be but a branch of 
the larger one passing on to the antenna and with it has a com- 
mon root. In sagittal sections treated by ordinary methods 
this may be readily followed on the lower side of the antennal 
lobe and somewhat to one side of its median line as a bundle of 
medullated fibers that passes through the group of cells filling 
the lower space between the cesophageal commissure and the 
globular mass of fibrillar substance of the lobe. It penetrates 
the fibrillar substance here and may still be followed through an 
inward, backward, and upward course for a considerable dis- 
tance. Gradually it decreases somewhat in size or loses its 
fibers, and is finally untraceable. In such sections of brains 
impregnated with bichromate of silver the fibers are frequently 
found impregnated and then form a large bunch of branching 
fibers spreading out in the commissure taking in the trito-cere- 
bron and to a slight extent the ventro-cerebron. Branches also 
pass upwards into the proto-cerebral region. In one case of a 
thick frontal section embracing the posterior part of the anten- 
nal lobe and nearly all of the proto-cerebron behind its plane a 
single fiber considerably smaller for a portion of its length can 
be seen above the level of the lobe, as shown in Plate XVII on 
the right where an attempt was made to reproduce it. Below 


Kenyon, Zhe Brain of the Bee. 177 


it decreases in size and joins several other fibers, the whole of 
which were not impregnated, and forms with them a band of 
fibers traceable down behind the lobe until hidden by the inter- 
vening strongly impregnated glomerule. As this band occu- 
pies the same position as the antenno-motor root seen in ordi- 
nary preparations I take it for granted that the fiber is motor 
and belongs to this nerve. The large part of the fiber inclines 
posteriorly in passing upward, terminating, so far as traceable, 
very close to the inner root of the mushroom body at some little 
distance from the median line. Three of its branches are im- 
pregnated. The lower one passes forwards towards the inner 
side of the antennal lobe and does not subdivide. From this 
fact and its apparent destination I have supposed that its cell of 
origin is situated among the cells filling this space between the 
lobe and the commissure. Higher up two branches are given 
off nearly together, one passing internally towards the median 
line, and the other, which is much longer, passing outward and 
downward in the trito-cerebral region and branching. But here 
it is obscured by other details. A short distance above this an- 
other long branch is given off. This goes diagonally backwards to 
the posterior-lateral angle of the proto-cerebron ending in several 
small branches. 

In other sections made in the horizontal plane fibers of a 
similar appearance were found taking a horizontal course and 
passing apparently beneath the antennal morula (Pl. XIX). 
These branch, as shown in the diagram, much more profusely 
than the other one just described, a fact that is doubtless due to 
their better impregnation. One sends a long slender process, 
or rather continues as such, backwards to the layer of cells 
clothing this part of the brain. But whether it actually arises 
from any of them I am unable to say. It is not impossible, and, 
if one considers the fiber shown on the opposite side of the 
plate, which after branching profusely and irregularly sends off 
a slender process in the direction of the antenno-motor nerve, 
it may not seem improbable. The details are not sufficient to 
settle the matter, however, and until they are shown to exist, I 
shall be inclined to believe that probably all the cells giving rise 


178 JOURNAL OF COMPARATIVE NEUROLOGY. 


to the antenno-motor nerve are situated close to or in the lobe 
on the anterior side of the brain. Fig. 39 represents a camera 
drawing of the two fibers just described and also another from 
the next section above them. 


The Dorso-Cerebral Fiber Tracts. 
THE CONNECTIONS WITH THE OPTIC LOBES. 


Tracts of fibers passing from the optic lobes into the cen- 

tral part of the proto-cerebron were early recognized by Ber- 
“ger (78) and Bellonci (s2), but the interpretations of these au- 
thors are very largely incorrect. Viallanes (7,88) seems to 
have described most of them much more correctly, and the 
same may be said of Cuccati (sg), so far as I am able, without 
an actual study of sections of Somomya, to homologize his results 
with my own. Certain peculiarities are figured and described 
by him that render a comparison difficult. ' 

Before describing the connections it should be noted that 
the masses of fibrillar substance forming what are usually known 
as the optic ganglia, but which are much more properly denom- 
inated fibrillar masses, are seen in frontal or horizontal sections 
treated by ordinary methods to be composed of two outer lay- 
ers of densely staining masses, and very much resemble in form 
two meniscus lenses placed one within the other. The space 
between them is filled with a loose mass of fibers, and the whole 
is so placed as to have its convex surface directed outward and 
its concave surface inward. 

The inner of these masses lies in hexapods close against 
the central proto-cerebron, being separated from it by a layer 
of cells and a few bands of fibers. In other words there is no 
optic nerve such as is found in the Crustacea. The attempts 
by the earlier writers to distinguish an optic nerve were long 
ago shown by Viallanes (87) to be more or less unsuccessful, 
since there is not one but several connecting tracts. These 
with the optic lobe sufficiently removed from the rest of the 
brain, might, however, produce the homologue of the crustacean 
optic nerve. 


Kenyon, Zhe Brain of the Bee. 179 
The Anterior Optic Tract. 


Even in ordinary preparations there are indications of fibers 
passing radially or nearly perpendicularly to its two surfaces, 
through the inner fibrillar mass. Emerging from the inner sur- 
face these are gathered into two bundles, the larger of which 
passes forward, and slightly downward along the outer surface 
of the central proto-cerebron and finally terminates in a small 
oval body called by Viallanes (7) the optic tubercle, situated 
immediately above the antennal lobe and to one side and below 
the terminus of the anterior root of the mushroom bodies of 
that side. In preparations with bichromate of silver the bundle 
is often very prominently shown, and in addition also the arbor- 
escent terminations of its fibers in the optic body, as shown in 
the diagrams that follow. This body in the bee is divided into 
a large inner mass and a very small outer mass, and into the 
latter some of the fibers of the tract send each a small branch 
before passing on to the larger body. 

Other fibers apparently arising from cells in the immediate 
neighborhood also branch in the body and connect it with other 
parts of the brain. One group passes to the opposite side and 
appears to terminate in the opposite optic body. Others pass 
backward, but no tracts appear to connect it with the mush- 
room bodies, although individual fibers are often found going 
in that direction. 


The Postero-superior Optic Tract. 


As just indicated, the fibers emerging from the convex sur- 
face of the inner fibrillar mass divide into a large anda small 
bundle. The latter may be considered the postero-superior 
optic tract, which seems to have escaped the notice of Viallanes 
(87,88). It may be easily traced in preparations either by the 
copper-hematoxylin method or by the _ platino-osmo-aceto- 
picric acid method of von Rath, and after leaving its larger 
companion, which is almost immediately, it passes upward along 
the outer surface of the central proto-cerebral mass, and joining 
the antero-superior tract, to be described a few lines further on, 
takes an inward course until, arrived close to the stalk of the 


180 JoURNAL OF COMPARATIVE NEUROLOGY. 


outer mushroom body, it leaves this other tract and passes be- 
hind the stalk close to the junction of the latter with the calyx. 
Whether any of its branches pass into the calyx here I am not 
able to say definitely, but apparently they do. In preparations 
with bichromate of silver some at least of its fibers may be fol- 
lowed across the intervening space to the opposite side of the 
stalk of the inner mushroom body. Whether any of them con- 
tinue on into the superior commissural tract my preparations 
thus far made do not show. 


The Antero-supertor Optic Tract. 


This as described by Viallanes (§7) and Cuccati (gg) starts 
from the outer fibrillar mass. Instead of originating like the 
two tracts just described from the radiating fibers of the mass, 
it arises from the fibers filling the space between its two lentic- 
ular portions. These fibers are gathered into a bundle and leave 
the mass at its antero-superior edge. Passing backward and 
inward through the intervening layer of cells, the bundle reaches 
the outer surface of the central proto-cerebral mass, then turning 
upward joins the postero-superior bundle. Just before reaching 
the stalk of the outer mushroom body it leaves its companion 
and passes in front of the stalk, close to its junction with the 
calyx, to the space between the two stalks where its fibers branch, 
one group passing into the outer, the other into the inner 
calyx. In one preparation I was able to determine the cells of 
origin of this tract and to follow it through several sections to 
its connection with the optic mass. According to the facts 
shown in this brain impregnated with bichromate of silver the 
description should be reversed and the union of the tract with 
the optic mass be spoken of as its terminus, for the cells from 
which it originates comprise a small group on the supero-ante- 
rior-lateral surface of the central proto-cerebral mass and send 
their processes as a bundle of fibers almost directly backward 
to the space between the two stalks, whence a branch passes 
into the calices and another to the fibrillar optic mass, as 
described. 


Kenyon, Zhe Brain of the Bee. 181 


The Antero-postertor Optic Tract. 


This has been well described by Viallanes (§7) in his me- 
moirs on the hexapod brain and also by Cuccati (gg) in his 
paper on Somomya erythrocephala. Like the tract last described 
it is formed from the fibers coming from the central portion of 
the outer optic fibrillar mass, which it leaves at the same point. 
Leaving the antero-superior tract at the optic mass it takes a 
backward and somewhat inward and downward course through 
the intervening mass of cells, passing over the anterior optic 
tract and between the inner optic mass and the central portion 
of the proto-cerebron and turns into the latter at its postero- 
lateral lower angle. After entering here the fibers separate 
from one another and branch, but may be followed for some 
distance towards the median line. Where its cells of origin are 
to be found I cannot say definitely. But in certain of my _ bi- 
chromate of silver preparations I have seen processes from cells 
in a group near the starting point of the tract entering the 
fibrillar mass, and it is not improbable that they are the cells of 
origin of the fibers of this tract. 


The Posterior Optic Tracts.* 


Of the remaining tracts connecting the optic lobe with the 
central proto-cerebral mass, there are several; but before pro- 


*The following shows to what extent my results correspond with those of 
Viallanes (87) who found four and I five tracts connecting the optic lobe with 
the central cerebral mass: 


A. Connections with the outer optic mass. 
I. Faisceau supéro-antereur = 
Antero-superior tract mzh#. 


2. Faisceau supéro-postereur = 


& 


Antero-posterior optic tract mzhz , 

B. Connections with the inner optic mass. 

3. Faisceau inféro-antereur = 
Anterior optic tract mhz. 
4. Faisceau inféro-postéreur, 2 tracts = 
2 of my posterior tracts. 

In his work on the grasshopper (88) he was unable to find the two tracts 
from the outer mass, while from the inner mass he found arising two tracts» 
one corresponding to my anterior optic tract, the other—his cordon commis- 
sural—to my lower optic commissure. 


182 JOURNAL OF COMPARATIVE NEUROLOGY. 


ceeding with their description it should be noted that the inner 
optic mass is so turned as to bring its posterior edge much closer 
to the central brain mass than its anterior margin. It is at this 
posterior margin that the entrance to the space between its two 
lenticular portions is situated. 

From this opening emerge a large number of fibers gath- 
ered into several bundles, the most of which terminate in the 
adjacent posterior lower portion of the central proto-cerebron. 

' Of these bundles Viallanes (7) distinguished four, and 
Cuccati (gg) seems to have found some of them in Somomya. 
' The inner optic region of this genus, as shown by the latter 
author, differs so much from what I find in the bee or have seen 
figured or described elsewhere that comparisons are very uncer- 
tain. What he calls the ovoid body (eiformige Korper) doubt- 
less corresponds to the inner optic mass as generally met with, 
but what may be the homologue of his S-form body I am ata 
loss to know unless it may possibly be the outer lenticular por- 
tion considerably separated from the inner one. But in case 
that were true, there would be an anomalous condition of fibers 
from the concave face of the outer optic mass passing through 
the space, a condition of affairs that is not true in the case of 
the bee nor probably in any other hexapod. He also figures 
and describes several tracts of fibers that apparently correspond 
to some of those to be described here a few lines ahead, but as 
originating from the outer surface of the S-form or of the ovoid 
body, which, I take it, is equivalent to describing them in the 
bee as arising from the outer surface of the inner optic mass, 
and this is not true. ; 

But to proceed with the optic tracts, there may first be 
described one, noted by both Viallanes (g7) and Cuccati (gs ), 
leaving the inner margin of the inner lenticular body and pass- 
ing into the central proto-cerebral mass a little above the point 
of entrance of the antero-posterior tract and continuing thence 
as a loose band of fibers across the posterior region of the 
brain and below the fibrillar arch to the inner optic mass of the 
opposite side, thus forming what may be termed the upper 
optic commissure. It does not pass directly across but is 


Kenyon, Zhe Brain of the Bee. 183 


arched slightly upward. It may be followed with little difficul- 
ty in ordinary preparations, and I have several times found its 
fibers impregnated in brains treated by the bichromate of silver 
method, Whether its fibers branch in the central brain I can- 
not say definitely. There are indications, however, that they 
do, and further, that their cells of origin may be situated among 
those forming the mass of cells behind the central body. 

Below this, in a plane taking in the anterior optic tracts 
and the optic body, and a little below the lower level of the in- 
ner roots of the mushroom bodies, there are three tracts, which 
may be seen in figure 10, Pl. XVI. These differ considerably 
according to the direction of the plane in which the section is 
‘cut, and also seem to differ somewhat in different individuals. In 
another brain from that from which the figure was taken there is 
a bundle that is not shown in the figure. This is described by 
Viallanes (7) as divided into a thicker and a thinner bundle, 
which is the case here. Both emerge from the inner edge of 
the inner lenticular body and terminate in the neighboring cen- 
tral region after penetrating no further than do the posterior two 
of those shown in the figure. The anterior of the two is con- 
siderably the larger and from its manner of staining may readi- 
ly be considered as composed of many very fine fibrils. The 
other is composed of fewer and coarser fibers and seems to 
arise more directly from the space between the two lenticular 
bodies. 

Below these are two more bundles, or the posterior two 
shown in the figure. These resemble the one just described 
and arise and terminate similarly. 

At a slightly lower level is the last of the series, which 
arises from near the margin of the inner lenticular body and 
passing directly into the central mass, bends slightly down- 
ward and then upward again to its former level and passes 
on as the lower optic commissure to the lobe of the opposite 
side. This is a much more compact and well marked bundle 
than the upper optic commissure and seemingly gives off no 
branches, although further study will probably show that it 
does. In one preparation by the copper-hematoxylin 


184 JOURNAL OF COMPARATIVE NEUROLOGY. 


method in which the sections-are in the horizontal plane I find 
fibers above it on each side of the brain that seem to run 
down to and join itin its middle portion. But in bichromate of 
silver preparations, although I have several times found some of 
its fibers impregnated I have not found sufficient evidence to 
warrant my saying that it branches. Just where its cells of or- 
igin are I have not been able to determine. 


THE DORSO-CEREBRAL COMMISSURES. 


Of the bands of fibers connecting the two lateral halves of 
the dorso-cerebron there are at least six, and of these two have 
already been noted in connection with the optic tracts. Some 
of them have long been known, but on account mostly of a 
lack of proper neurological methods have seldom been correctly 
understood. Viallanes (g7,88) seems to have made fewer, mis- 
takes than any other writer who has considered them. 


The Superior Dorso-cerebral Commissure. 


This band of fibers crossing the median line of the brain 
above the central body and between the two inner stalks of the 
mushroom bodies (fig. 2) was noted by Dietl (76) and Ber- 
ger (78) and considered as connecting the two optic lobes, a 
mistake perpetuated by Bellonci in his study of Grylotalpa. 
The fibers are brought out very prominently by most of the or- 
dinary methods and I have repeatedly found them impregnated 
in brains treated by the bichromate of silver method. They 
are all of small size and non-medullated. By a comparison of 
frontal and horizontal sections (figs. 2 and 7) they are seen. to 
form a band whose broadest extent is in the antero-posterior di- 
rection, and to be divided at either end into two parts—a divis- 
ion that can sometimes be distinguished throughout the length 
of the commissure—one of which passes behind and the other 
in front of the stalk. There does not seem to be a crossing of 
the fibers, but those passing in front of one stalk also pass in 
front of the other. In preparations with haematoxylin or 
with osmic acid the halves may be traced in horizontal sections 
around the inner stalks to the space between these and the outer 
ones and apparently into the calices. Since the tracts from the 


Kenyon, Zhe Brain of the Bee. 185 


optic and antennal lobes also enter this region, very much care 
is necessary in distinguishing the different tracts, but since they 
fuse to some extent upon entering the calices, it is impossible to 
follow any one of them singly. Viallanes probably correctly 
interpreted this bundle as a commissure between the two pairs 
of mushroom bodies. There is apparently very little probabil- 
ity of any of its fibers reaching the optic lobes, though none of 
the methods employed by me have thus far demonstrated con- 
clusively. that they do not, nor have I been able to determine 
their exact terminations in the calices nor the location of their 
cells of origin. In preparations with hematoxylin and with 
osmic acid processes from the cells behind the inner side of the 
inner stalks have been traced upwards and forwards, apparently 
passing into the commissure as it passes around the stalks, but 
no such fibers have been found impregnated in brains prepared 
according to the bichromate of silver method. This negative 
evidence is not, however, at al! conclusive since a sufficiently 
large number of brains have not been treated. 


The Anterior Commissure: 


The smal] band of fibers composing this commissure was 
first discovered by Viallanes (87) as connecting the two optic 
bodies. It may be followed in ordinary preparations without 
much difficulty from the posterior margin of one optic body 
across the anterior region of the brain and below the roots of the 
mushroom bodies to the optic body of the opposite side. Ap- 
parently its fibers originate from the group of cells immediately 
inside of and below the body. The processes from the cells 
seem to pass upward, and after sending a short process into the 
body where as seen in bichromate of silver preparations, it 
branches very profusely, and then passes across the median line. 
I have not been able to determine exactly its termination on the 
opposite side or whether it or any of its branches penetrate the 
optic body here. One bichromate of silver preparation where 
the fibers of the tract are most completely impregnated is incon- 
clusive from the very fact of the extent of the impregnation. 
From it one might conclude that branches pass into the optic 


186 JourNnaL OF CoMPARATIVE NEUROLOGY. 


body, but the greater number into the region immediately 
behind it. 
The Optic Commussures. 


The two tracts of fibers forming the one the upper and the 
other the lower optic commissure have already been described 
in connection with the optic tracts. As noted there, I am un- 
able to say definitely whether either give off branches to the 
brain in passing, and it may be added here that I have no evi- 
dence to show that a single nerve cell makes the entire con- 
nection between the lobes, if such be actually the function of 
the tracts. The question needs much further study. 


The Inferior Dorso-cerebral Commissures. 


The two groups of fibers connecting the lateral halves of 
the dorso-cerebron immediately above the cesophageal fora- 
men have been variously misunderstood. They are shown in 
fig. 10 where they appear as two short tracts in front of the 
larger lower optic commissure. The anterior of the two was 
supposed by the earlier writers to connect-the antennal lobes, 
and even the olfactory glomerulz of one side with those of the 
other. Of such a direct connection I find no evidence what- 
ever, though without doubt the two commissures may be con- 
sidered as connecting the two deuto-cerebral lobes. In ordinary 
preparations sectioned in the horizontal plane both commissures 
are found immediately above the foramen, in fact almost bord- 
ering upon it, and may be traced for a short distance into the 
fibrillar substance of either side where the fibers curve slightly 
downward and separating from the tract branch and disappear. 
In frontal sections the fibers may be followed a little farther 
than in those cut in a horizontal plane. In fig. 5 one of the 
tracts may be seen though somewhat indistinctly. 

In bichromate of silver preparations, although I have fre- 
quently found commissural fibers impregnated, I have not found 
sufficient details to warrant a definite assertion as to the exact 
relation of the tracts to the rest of the brain, nor as to the situ- 
ation of the cells giving rise to their fibers. What details there 
are, however, indicate that the anterior tract connects the two 


Kenyon, Zhe Brain of the Bee. 187 


regions, which may be called deuto-cerebral, immediately be- 
hind the antennal lobes. If it in any way connects the two 
lobes or their glomerulz, it is only by means of intervening as- 
sociation fibers. But such a connection is doubtful from phy- 
siological reasons. It seems much more probable that the com- 
missural fibers by the aid of association fibers of one side, 
should connect the glomerulz of that side with motor fibers of 
the other. Still the other supposition is not impossible, and 
one should rather follow the maxim of Hunter, ‘‘do not think, 
but go and see’, and wait until facts, physiological or other- 
wise, are obtained to decide the matter. 

The fibers of the posterior tract, which is separated from 
the other by the ascending tracts from the antennal lobes to 
the mushroom bodies, connect more posterior parts of the brain. 
The fibers shown in fig. 38, apparently belong to this tract 
since at the median line they occupy the same position. Here 
they are obscured by a mass of precipitate filling the roof of 
the foramen, but since there are no other fibers impregnated here, 
there can be little doubt that the fibers of the opposite side are 
a continuation of them. The branching and larger portion of 
them is found in the posterior lateral lower angle of the brain, 
and it is possible that they may originate from cells in this 
neighborhood. One of them, it may be noted, sends a slender 
branch forwards toward the antennal lobe. 


THE ANTENNO-CEREBRAL TRACTS. 


The existence of tracts of fibers passing upwards from the 
antennal lobes into the proto-cerebron has been known ever 
since Bellonci (gg) published his account of them in Gryllotalpa. 
But of the three pairs of such tracts he seems to -have distin- 
guished but two, and certainly misunderstood the relationships 
of these. For he describes them as antenno-optic connections 
passing from the antennal lobe upwards into the proto-cerebron 
and to the optic lobes by way of the superior commissure, which 
he recognized with Dietl and Berger as connecting the optic 
lobes. The same error was committed by Cuccati (8s) who 
found in Somomya what is unquestionably the homologue of 


188 JOURNAL OF COMPARATIVE NEUROLOGY. 


the inner tract to be described a little farther on, and which is 
also the main tract discovered by Bellonci. Viallanes (87) 
found the tract in the wasp and for the first time described it 
correctly. 

In certain of my preparations of the brain of the bee by 
the copper-hematoxylin method these antenno-cerebral tracts 
are very distinct and very easily followed. This is the case in 
sections cut in any o¢ the three planes. 


The Inner Antenno-cercbral Tract. 


_ The one first to be described is the largest and also the one 
first discovered. It arises in the neck of fibrillar substance con- 
necting the antennal morula with the rest of the brain and as- 
cending upwards, inwards and backwards (fig. 2) it passes be- 
tween the two ventral commissures of the dorso-cerebron (fig. 
10). Thence curving backwards somewhat it passes behind the 
root of the mushroom bodies and the outer hinder margin of 
the central body and in front of the fibrillar arch (figs. 8 and g) 
to the level of the superior commissure, behind which it may 
be seen in section in fig. 7, From here it bends forward and 
passing around in front of the inner stalk of the mushroom 
body enters the inner calyx at its junction with the stalk and on 
the antero-lateral side of the latter. 

In bichromate of silver preparations I have several times 
found the fibers of the tract impregnated for nearly their entire 
length. And in many cases different portions of the tract in 
successive sections were so well impregnated that their superim- 
position was a very easy matter. Fig. 24 represents the super- 
imposition of three such sections and shows the tract from 
the calyx above to the inside of the antennal morula below. In 
no case have I found fibers from a glomerula passing into the 
tract, but the arrangement of the many fiber-fragments that 
have been seen almost conclusively demonstrated that they enter 
it (fig. 24). The fiber from the glomerula shown in the figure 
offers avery good example. 

Nor have I found the fibers in cennection with cells, but, 
considering their connection with the glomerulz as_ practically 


Kenyon, Zhe Brain of the Bee. 189 


demonstrated, it may be remembered that very good reasons 
appear for considering the glomerulz as connected with the cells 
clothing the morula, and from this it is fairly evident that the 
same cells give rise to the fibers of the tract. Ai little further 
evidence will appear in connection with the outer tract. 


The Middle Antenno-cerebral Tract. 


The next tract to be considered is the smallest of the three 
and arises as a branch of the one just described at a point just 
above the level of the ventral commissures. It takes an out- 
ward course behind the mushroom body (fig. 6) and continues 
upward to the region behind and between the stalks and then 
bending forward somewhat passes into the outer calyx. 

This seems to be the branch figured by Bellonci as passing 
outward into the supposed tract between the optic lobes. In 
several of my bichromate of silver preparations it is very read- 
ily followed. 

The Outer Antenno-cerebral Tract. 


This tract is of considerable size and originates, as shown 
in bichromate of silver preparations (fig. 24), from cells above 
the antennal morula. From the morula it passes backward and 
outward towards the side of the central brain mass and gradu- 
ally turning upwards passes behind, but in contact with, the low- 
er optic commissure (fig. 6) and finally reaches the outer dorsal 
surface of the fibrillar substance and there, joining the antero-su- 
perior optic tract, passes in front of the outer stalk into the 
calyx above. Whether any of its fibers continue to the other 
calyx or not, I cannot say. 

In some frontal sections there appears to be given off from 
this tract below the lower optic commissure a small bundle of 
fibers that curves outward and apparently joins the commissure. 
But in horizontal sections it seems to follow the commissure to 
a point near the outer surface of the fibrillar substance, where it 
turns abruptly forward and enters the mass forming the antero- 
lateral lower angle of the central cerebral mass. 


190 JOURNAL OF COMPARATIVE NEUROLOGY. 


THE DORSO-VENTRAL TRACT. 


Five tracts of fibers have thus far been described as enter- 
ing the calices of the mushroom bodies and there still remains 
another. This was first brought to my notice in a frontal sec- 
tion of a brain impregnated with bichromate of silver, and was 
subsequently recognized in a similar section of a pupal brain 
treated by the copper-hematoxylin method. It begins as a 
small band of fibers in the region between the two stalks, where 
branches of it enter both calices, and passes downward be- 
neath the cells clothing the posterior surface of the brain to the 
cesophageal foramen, then continues anteriorly along the roof of 
this so as to reach the opposite side of the brain near the anterior 
surface. In its passage along the roof of the foramen its fibers 
send branches into the side from which the main fibers came, and 
these form an arborescent termination among branches of fibers 
coming from the ventro-cerebron and the ventral cord. Since 
these latter fibers cross the median line above the foramen an- 
teriorly, the passage of the tract to the opposite side brings it 
into close relation with them along their whole extent along the 
roof of the foramen. This relation is best shown in plate XX, 
which shows two fibers of the tract as seen in a single section. 
The cells of origin of the tract have not been seen. But suff- 
cient has been determined to show in what close relationship the 
cells of the mushroom bodies stand with the ventral nervous 
system. 


The Ocellar Nerves. 


In the bee the ocelli are situated close together, andas a 
consequence their so-called ganglia are contiguous and their 
nerves toa large extent fused. The nerve from the median ocel- 
lus passes downward and backward between the two inner calices 
and fusing with the nerves from the lateral ocelli forms with them 
a single mass that divides to pass around a large trachea piercing 
the brain in the median line between the calyx cups. Immedi- 
ately below this the fibers again come into close contact, but at 
a little lower level some of those coming originally from the 
median ocellus separate from the rest and follow a course that 


Kenyon, Zhe Brain of the Bee. IgI 


takes them down in front of the fibrillar arch to terminate final- 
ly, as apparently shown in copper-hzematoxylin preparations 
cut sagittally, in the tubercles of the central body. 

The others form a broad band of fibers passing down be- 
hind the fibrillar arch to which, as suggested by Cuccati (gs), 
some of them may give off branches. Certainly some of the 
smaller fibers cannot be traced below this level (Pl. XVI). 

Both the anterior and the posterior groups are composed of 
two kinds of fibers, one enormously large and the other of or- 
dinary size. The large fibers are very noticeable in prepara- 
tions by the copper-hematoxylin method from the fact that 
they stain but slightly and thus appear as light colored spots in 
a deeply colored surrounding mass of cells (fig. 7-9). Similar 
results are obtained with von Rath’s platino-osmo-aceto-picric acid 
mixture. The unstained fibers look very much like sections of 
tubes. But in preparations treated with Weigert’s hematoxylin, 
or in those after-stained with fuchsin the fibers stain more or less 
deeply and thus demonstrate that they are at least not empty 
tubes. Even in preparations by my copper-hematoxylin method 
staining sometimes takes place. In such cases the inner mass 
appears in section to be shrunken away from the surrounding 
fibrillar wall, but remaining connected with it by slender fila- 
ments. This would seem to show that they are not formed 
upon exactly the same plan as other nerve fibers. A careful 
histological study of their structure is necessary. 

The nerves thus constituted may be followed for some dis- 
tance below the fibrillar arch, but seem to branch and gradually 
decrease in size. This is noticeable in the figures, where in 
fig. 7, the anterior nerve appears with three large fibers and at 
a considerably lower level (fig. g) there is but one. The fibers 
of this nerve soon become untraceable and appear to pass into 
the fibrillar arch and the ocellar glomerule beneath. 

Some of the fibers of the other two nerves seem to termi- 
nate in the same way, but a large number of the larger fibers 
continue downward, as seen in both horizontal and frontal sec- 
tions, until they reach the neighborhood above the cesophageal 
foramen, where they separate. Some seem to terminate in this 


192 JoURNAL OF COMPARATIVE NEUROLOGY. 


region. The others come into close contact with the hinder 
surface of the brain and pass on downward in the upper sur- 
face of the cesophageal commissures to the ventro-cerebron, 
whence they pass on backward along the dorsal surface of the 
ventral cord, thus effectually disproving Binet’s (94) assertion 
that there are no large fibers in the dorsal region of the cord 
comparable to those found in crustacea. In many cases I have 
found these large fibers of the cord in bichromate of silver 
| preparations passing along the dorsal surface of the ventro- 
cerebron on into the region just below the level of the central 
body as shown in Pl. XX. What their significance may be I 
do not at present undertake to say. Had Binet persevered 
longer in his trial of either of the two methods preeminent in 
the study of the nervous system, he probably would have found 
them.. But while the existence of the large fibers is beyond 
question, there is still a chance that my description of them as 
coming from the ocelli is erroneous, for the reason that I have 
not been able, in specimens prepared by the bichromate of silver 
method or by any other, to follow a fiber through an unbroken or 
unsectioned course from the ocellar ganglion or the upper por- 
tion of an ocellar nerve to the ventral cord. Yet impregnated 
fibers have been followed without interruption from the copper- 
the level indicated above, and A the large fibers have been care- 
fully traced through sections treated according to my’ copper- 
hematoxylin method. It should be noted that Cuccati (8s) 
also traced them in Samomya, as I have done here, into the 
ventral cord. Viallanes (87,88) did not succeed in following 
them below the fibrillar mass behind the inner root of the 
mushroom bodies. 


THE FIBERS FROM THE BRAIN TO THE OCELLAR GANGLIA. 


Besides the fibers just described as constituting ‘the ocellar 
nerves there may, in preparations by the bichromate of silver 
method, be distinguished others of an efferent nature (fig. 42, 
Pl. XXII). These may be traced from below the level of the 
calices into the ocellar ganglia where they branch arborescently. 
None are large. Where they originate or what may be their 


Kenyon, Zhe Lrain of the Bee. 193 


fibrillar connections within the brain, I have not as yet been 
able to learn. Not many preparations were obtained show- 
ing them passing out of the ganglia, and in fact only one in 
which they could be traced from below the calices, and from 
this the two fibers of the figures were taken. 

As there are no muscular structures in the ganglia, their 
function is doubless to control the action of the pigment of the 
pigment cells. 


The Commissural and Ventro-cerebral Region. 


Since in the case of the bee they are so closely bound to 
it, the parts composing the cesophageal commissures may be 
considered along with the ventro-cerebron. 

As already pointed out, the trito-cerebral lobe is very much 
reduced and is mainly distinguishable externally by the labral 
nerve. This is not far behind the internal antenno-motor nerve 
and its root may be traced for a short distance into a small lobe 
just inside the neck of the antennal morula. Fig. 2 is just a 
little too far forward to show it, but asection of the nerve may 
be seen on the left at its point of entrance. In bichromate of 
silver preparations the fibers entering from it very quickly form 
a bushy tuft of branches just beneath the root of the antenno- 
motor nerve. Some of these reach back into the ventro- 
cerebron. 

The roots of the other oral nerves may be followed in or- 
dinary preparatious each as a light band of fibers that passes 
upward above the point of entrance of the nerve and then bends 
inward towards the median line and ends in a roll of fibers that 
is apparently much more easily distinguishable in other forms 
than in the bee, and which Binet (94) calls the ventral column. 
This passes backward into the ventral cord. The two roots that 
Binet describes for each crural and each ventro-cerebral nerve 
in Cerambyx and other beetles I do not find in my sections of 
the bee’s ventro-cerebron. This may be due to their being 
fused, but the attention that I have given this portion of the 
brain is as yet small in comparison to that given the dorso- 
cerebron. 


194 JouRNAL oF CoMPARATIVE NEUROLOGY. 


In bichromate of silver preparations some of the fibers en- 
tering from the nerves very quickly break up into branches, but 
reach a little above the lower half of the ganglion. In a few 
cases the roots were found impregnated for nearly their whole 
extent and did not appear to give off many branches to the 
ventral region. In one case two cells in front of the mandibu- 
lar nerve were each found connected with a fiber that passed 
out through the nerve, but whatever inner branches they may 
have had were not impregnated (Pl. XX). In another case 
‘a large cell was found on the lower lateral side of the brain 
sending a process into the region above, where it gave off a very 
extensive small branch (fig. 36). Near the broken end of the 
fiber another that is not shown in the figure began. This was 
apparéntly a continuation of it. The appearances of the sec- 
tion indicated that the fiber had been cut on account of its form- 
ing a slight bend outside the plane of the razor. The second 
fiber turning forwards gradually became swollen distally and 
then decreasing passed asa smaller fiber into the root of the 
antenno-motor nerve. The latter cell then is certainly motor, 
and taking for granted what has already been fairly well demon- 
strated by other observers; namely, that cells in the brain send- 
ing their neurites out of it are motor or bearers of efferent neu- 
ral impulses, and the fibers entering from external cells are sen- 
sory, it is just as surely true that the other cell is of a motor 
nature. 

This ventral position of motor cells here tends to indicate 
that the dorsal motor and ventral sensory area of ventral gang- 
lia, as distinguished by physiologal experiments, is considerably 
independent of the position of the cell. It may be pointed out 
that in dorsal lesions it is the connection of the dendrites of 
motor cells with fibers bearing stimuli to them from other 
parts of the nervous system that is broken; while in ventral le- 
sions it may be the terminations of sensory fibers or of associ- 
ation cells and their fibers or both that are destroyed. From 
this it may be seen how indefinite are apt to be results obtained 
by the physiological vivisection methods. Should experi- 
ments be repeated with a thorough knowledge of the location 


Kenyon, Zhe Brain of the Bee. 195 


of the cells about a ganglion so as to render it possible to rup- 
ture individual cells and not fibrillar connections, and sufficient 
time is allowed to elapse for the degeneration of the dendrites 
and neurite of the ruptured cell, it will doubtless be found that 
ventral and lateral portions of the ganglion are motor. 

Such careful experiments are needed and, performed upon 
the ventro-cerebron, they may, by the aid of either the methy- 
lin blue intra-vitam or the bichromate of silver method, be able 
to demonstrate which of the large number of cells located there 
perform the function of co-ordinating the movements of the 
body. 

The association fibers of this ventral region of the brain 
are numerous, and of the many fragments of them seen one of 
the best is shown in fig. 24. This, as is evident from the fig- 
ure, occupies the upper portion of the commissural region and 
sends the greater part of its branches and branchlets dorsally 
into the lateral region of the proto-cerebron in the dorso-ventral 
plane of the anterior root of the mushroom bodies. One branch 
passes forward inside the inner antenno-cerebral tract. 

Numerous fibers may be found connecting the antennal 
morula through its neck of fibrillar substance with the com- 
missure. A single fragment shown in plate XX (fiber 37) sends 
a process upward in the posterior region that branches below 
the level of the ocellar glomerulz, while another is sent forward 
apparently to the antennal lobe. Just before reaching this it 
sends a secondary branch downward into the ventro-cerebron. 
The fibers seen entering the ventro-cerebron from the ventral 
cord are in some cases remarkable for their size. These large 
fibers occupy the dorsal region and have already been spoken 
of in connection with the nerves of the ocelli. Fragments of 
them have been seen as far back as the first thoracic ganglion, 
and anteriorly, as before noted, they have been traced in bichro- 
mate of silver preparations into the proto-cerebron behind the 
ocellar glomerule. Branches are given off in this region that 
pass among the glomerule to the commissural region as shown 
in the plate. Much shorter ones are given off in the ventro- 
cerebron. Another large fiber (fiber 38) apparently ends in the 


196 JOURNAL OF COMPARATIVE NEUROLOGY. 


ventro-cerebral and commissural region. Lower down in the 
ventro-cerebron another fiber is shown (fiber 41) that branches 
among the fibers from the oral nerves and thence apparently 
passes backward into the ventral cord. A smaller fiber (40) 
makes the same connections. 

In some of my preparations there may be distinguished 
what may be described as a roll of fine fibers, the individual 
fibers of which cannot be traced, that passes through the ven- 
tro-cerebron and thence upward through each commissure; 
whence it crosses to the opposite lower side of the dorso-cere- 
bron or over the anterior part of the roof of the cesophageal 
foramen.’ I do not know what this can be unless it is a contin- 
uation of the ventral column described by Binet (94). Its pas- 
sage into the dorso-cerebron is directly contradictory of this au- 
thor’s assertion that ‘‘ il n’existe dans le ganglion sous cesopha- 
gien-aucun croisement des connectifs qui prennent leur origine 
dans le cerveau.’”’ But Viallanes (87,88) speaks of a fibrillar 
tract that passes from the cord to the opposite side of the dorso- 
cerebron. In close connection with this roll individual fibers 
may be followed from the cord to the same destination. . 

What is apparently a branch of this roll continues upward 
towards the central body in the commissure of the same side, 
or without crossing the median line. 

It is this roll of fibers that has already been mentioned in 
connection with the dorso-ventral tract, with the fibers of which 
it seems to come into contact. Assuming that, which is doubt- 
less true, it is the continuation of the ventral columns described | 
by Binet (94) and that the root of the crural and the alary 
nerves terminating in it are sensory, which is also probably 
true, there is then seen to be a direct sensory tract for external 
stimuli from all parts of the body to the calices of the mush- 
room bodies, a fact of no little importance in completing the 
chain of evidence demonstrating that the cells of these bodies 
are the ones that enable the animal to adapt itself to the vary- 
ing conditions of life. 

A similar group of fibers may be followed in copper-hama- 
toxylin preparations from the ventral cord through the ventro- 


Kenyon, Zhe Brain of the Bee. 197 


cerebron to the antennal lobe within the region of the inner ter- 
minations of the motor-fibers to the oral and ventral nerves. 
In such preparations fibers leaving the tract may be. readily seen 
and it is not a connection between the antenno-motor nerves 
and the ventral cord such as Cuccati (s8) seems to imply in his 
description of the same tract in Somomya. Connecting the 
pair of tracts in the ventro-cerebral region are four transverse 
bands of fibers. These have been noted several times in bi- 
chromate of silver preparations, but the longitudinal tracts to 
the antennal lobe were found impregnated but once. 

The transverse cesophageal commissure seems to have be- 
come completely fused with the ventro-cerebron in the bee, and 
is apparently represented by a few transverse fibers found in the 
lower floor of the cesophageal foramen. 


The Cell Groups. 


Having finished the description, incomplete though it be, 
of the central fibrillar substance of the bee brain, there still re- 
main the groups of cells clothing the mass to be discussed. 
The location of these, after recognizing the general form of a 
hexapod nerve cell, is of little physiological importance com- 
pared with the extent and connections of their fibers, but mor- 
phologically they may have considerable significance. 

As before mentioned, the cells of the brain are gathered 
into masses that completely cover the fibrillar substance in cer- 
tain regions and fill in the spaces between its lobes. Thus the 
spaces between the optic lobes, the antennal morula and the 
central cerebral mass are completely filled by them, as is also 
the deep furrow between the two lateral proto-cerebral lobes 
on the posterior side of the brain. 

These masses are, with certain exceptions, subdivided into 
small groups the fibers of which form a bundle in penetrating 
the central mass. These groups so far as I have been able to 
distinguish them may be designated by numbers as follows be- 
low. The most prominent of the exceptions to the rule of the 
formation of small bundles are the cells about the calices of the 
mushroom bodies. These are very similar in appearance to the 


198 JOURNAL OF COMPARATIVE NEUROLOGY. 


small cells filling the central portion of the calyx-cups and their 
general mass is in some places continuous with that filling the 
latter, their relation to which may be compared to the over- 
flowing contents of a cup. They are found in the space be- 
tween each pair of calices and in that between the latter and the 
fibrillar mass below, anteriorly and posteriorly, and grade into the 
mass of cells between the central mass and the fibrillar masses 
of the optic lobes. No processes from any of them to the por- 
tion of the brain beneath them have been seen in any of my 
preparations, but in all they seem to be in some way closely re- 
lated to the calices. In one instance a process was traced into 
a calyx from a cell in the space between the pair of cups, and 
this, branching there, exactly resembled the dendrites of the cells 
inside. Since this was one of the earlier of my discoveries, and 
since the boundaries of the cup are not always distinct in non- 
stained, or bichromate of silver preparations, and since at this 
moment I am unable to verify the matter, there is here a possi- 
bility of a mistake. If not, and the cells are of the same order 
as those inside the cup, their neurites must be looked for as 
passing inside to the radiating bundles of fibers. 

Between the posterior margins of the stalks and immedi- 
ately below the calices I have several times found larger cells 
impregnated, whose processes passed. in between the stalks and 
up between the calices. Perhaps these may be the cells giving 
rise to the fibers of the superior commissure, but the details to 
be seen are too poor to warrant a conclusion. 


THE CELLS OF THE DORSO-CEREBRON, 


I. To proceed with the enumeration of the groups of 
cells, there may first be noted a small group of medium sized 
cells, situated dorso-antero-laterally beneath the outer calyx, the 
processes of which pass almost directly backwards to the neigh- 
borhood of the inner surface of the outer stalk and form the 
processes giving rise to the fibers of the antero-superior optic 
tract. 

II. Behind and below these, but in close proximity, is an- 
other group of similar appearance, the processes of which form 


Kenyon, The Brain of the Bee. 199 


a tract passing into the fibrillar substance towards the median 
line of the brain. 

III. Below the anterior optic tract nearly half way between 
the inner optic mass and the optic body is a group of mod- 
erately large cells, the tract of fibers from which passes under 
the optic tract and upwards towards the middle superior surface 
of the anterior root of the mushroom body, above which they 
seem to branch. In all preparations where seen there is consid- 
erable difficulty in distinguishing the fibers of this tract from the 
outer branches of the tract from the posterior region of the 
calices to the root. 

IV. Immediately outside of and below the optic body is 
a similar group whose processes pass beneath the optic tract 
upward and inward, apparently, along the lower surface of the 
anterior root. 

V. Inside of and below the optic body is a group of cells 
of median size whose fibers pass upward along the outer side of 
the anterior root. 

VI. Below these and above the antennal morula is a 
group that, sends a tract of fibers upwards across the inner side 
of the root (figs. 24 and 31, fiber 10 of the diagrams). 

VII. Outside of these and above the morula are three 
small groups that send as many tracts of fibers inward to the 
central body (fiber 30). They pass below the latter structure 
and, turning upwards in front of the tubercles of the central 
body, enter the space separating its dorsal and ventral portions. 
Apparently they form the tract that is figured by Bellonci (sg) 
as connecting the inner antenno-cerebral tract with the superior 
commissure on the opposite side of the brain, or as forming a 
chiasma with its companion from the opposite side. 

VIII. Near the latter group is another whose processes 
form the outer antenno-cerebral tract (fig. 24). 

IX. Beneath the calices and in the median plane is a 
group of cells recognizable in fig. 1, the processes of which pass 
directly downward, and according to Viallanes (gs) enter the 
cesophageal commissure of the brain, thus forming a chiasma. 
In the wasp (Viallanes 87) and in Somomya (Cuccati gg) their 


200 JOURNAL OF COMPARATIVE NEUROLOGY. 


relationships seem to be the same as in the bee, in which I find 
no evidence of crossing. 

X. Immediately behind the latter group is another com- 
posed of cells of the same large size, the fibers of which pass 
backward over the central body and connect it with the fibrillar 
arch (fiber 2). 

XI. Considering the posterior region of the brain there 
may first be noted a small group situated below the outer calyx 
at a point where the outer antenno-cerebral tract reaches the 
. dorsal surface of the fibrillar substance. Its processes pass 
downwards into conjunction with the tract and might be taken 
for its processes of origin were it not for the fact that bichro- - 
mate of silver preparations show that its cells are situated in the 
antennal lobe. Where the fibers of this group go to I have not 
been able to determine. 

XII. Behind the stalks of the mushroom bodies is a small 
group of large cells whose processes pass between the stalks to 
the anterior root (fiber 7) giving rise to some of the association 
fibers in that structure. ; 

XIII. Behind the inner stalk there is a group of medium 
sized cells whose processes pass inward behind the inner anten- 
no-cerebral tract and behind the central body into which each 
fiber sends several branches (fiber 5). 

XIV. Behind the lateral margin of the central body and 
beneath the fibrillar arch is a companion group whose fibers 
pass directly forward and turning about in front of the inner 
anteno-cerebral tract pass behind and send branches into the 
central body (fiber 6). The two groups thus form a commis- 
sure behind this structure. As before mentioned it is possible 
that their neurites may be the fibers seen leaving the central 
body and passing into the lower lateral portions of the proto- 
cerebron and towards the commissural region (fig. 5). 

XV. Above these groups and between the central body 
and the fibrillar arch and below the latter is a small group 
whose processes form a bundle (figs. 2, 5, 6, 8) passing just in- 
side of the inner antenno-cerebral tract over the top of the 


Kenyon, Zhe Brain of the Bee. 201 


central body to the median anterior mass of fibrillar substance 
(fiber 49). | 

XVI. Above these and in front of the arch is a small 
group of cells, often staining more deeply than those surround- 
ing them, whose processes pass directly downwards outside of 
the tract just described and appear to enter the inner antenno- 
cerebral tract (fiber 3), 

XVII. Above the ends of the fibrillar arch is a large 
group of cells of large size whose fibers form a dorso-ventrally 
flattened band that passes forwards outside of the inner antenno- 
cerebral tract and above the superior commissure (figs. 2, 7, 8) 
and bending about the inner stalk distribute themselves in the 
fibrillar substance above and outside of the anterior root of the 
mushroom bodies (fiber 23). 

XVIII. Below the fibrillar arch and a little below the 
commissure of the central body is a loose group of cells whose 
processes in some cases appear to pass into the superior optic 
commissure. In others they pass across to the fibrillar mass 
behind the base of the junction of the roots and stalks of the 
mushroom bodies (fiber 4). 

XIX. Behind group XVI and separated from it by the 
fibrillar arch is one that appears to send its processes down- 
ward into the inner antenno-cerebral tract, but probably they 
are sent into the central body. 

XX. Another group below these and the fibrillar arch 
sends its processes downward and inward apparently under the 
superior optic commissure, in front of which they bend upwards 
and apparently pass to the calices along with the fibers of the 
superior commissure and those of the inner antenno-cerebral 
tract. 

XXI. The cells about the antennal morula are most nu- 
merous in the space between it and the restof the brain, although 
they cover almost its entire surface, and are perhaps gathered 
into as many groups as there are inter-glomerular spaces. One 
group that should have been noted before is situated in the in- 
ner superior side of the neck of the morula and sends its fibers 
upward through the anterior portion of the central mass of 


202 JOURNAL OF COMPARATIVE NEUROLOGY. 


fibrillar substance and near the median line. Near the lower 
surface of the inner calyx they turn outward over the anterior 
root. 

XXII. Three groups that may be considered as one on 
account of their situation below the neck of the morula have 
their processes directed upwards toward the neck. 

XXIII. In the lateral lower side of the central cerebral 
mass are several groups. One large one at the posterior lower 
edge of the inner optic mass sends its processes straight forward 
into the fibrillar mass forming the anterior lateral angle of the 
brain below the anterior optic tract. 

XXIV. Another group inside the last sends its fibers into 
the adjacent posterior angle. 

XXV._ A little above the group just noted is another that 
sends its fibers in among the terminations of the posterior optic 
tracts. 


THE CELLS OF THE VENTRO-CEREBRON, 


Excepting the dorsal surface the whole of the ventro-cere- 
bron is covered with a mass of cells all of which are of consid- 
erable size, but largest ventrally. Among them may be distin- 
guished seven principal groups. 

XXVI. A ventro-median group between the origin of 
the maxillary nerves sends a band of fibers directly through the 
mass above, passing behind a group of commissural fibers 
among which may be those of the transverse cesophageal com- 
missure to the dorsal surface. E 

XXVII. A second ventro-median group also occurs be- 
tween the origins of the labial nerves. Its fibers after reaching 
the neighborhood of the dorsal surface separate and apparently 
reach the latter. 

XXVIII. Between this group and the origin of the nerve 
is another of large cells whose processes form a distinct tract 
passing upward and outward just outside of the ventral column 
at the level of which it branches, one half continuing upward 
and forward towards the commissural region, the other turn- 


Kenyon, The Brain of the Bee. 203 


ing outward and becoming lost in the surrounding fibrillar sub- 
stance. 

XXIX. <A group outside of the last and behind the ori- 
gin of the labial nerve sends a band of fibers nearly half way 
through the mass behind the root of the nerve. 

XXX. Between the origins of the labial and the maxil- 
lary nerves and outside of them on the lower lateral surface is 
a large group whose fibers forming a well marked band pass 
inward and then upward near the root of the maxillary nerve 
and then upward and inward and apparently forward towards 
the commissural region. 

XXXI. Considerably above the last, or on the upper lat- 
eral surface and below the origin of the salivary nerve, is a group 
whose fibers pass directly inwards and join those of the last 
group near where these turn forwards. 

XXXII. Just above the one just described and in front of 
the plane of the origin of the maxillary nerves is a group that 
sends a band of fibers upwards into the commissural region. 


Conclusion, 


From the facts detailed in the preceeding pages it is evi- 
dent that even though there are more difficulties in the way of 
obtaining good results than with the vertebrates, a patient ap- 
plication of the bichromate of silver method will throw as much 
light upon the organization of the hexapod nervous system as 
it has upon that of the higher animals. By its aid during the 
past winter the minute structure of the so-called mushroom bod- 
ies has been brought to light and several links added that almost 
complete the chain of evidence demonstrating the function of 
these peculiar bodies to be that of enabling the insect to intelli- 
gently adapt itself to its surroundings. They are shown to be 
connected at their calices with two pairs of sensory tracts of 
fibers from the optic lobes, with three from the antennal lobes 
and with one, that is probably also sensory, from the ventral 
nervous system. Their roots are shown by fragmentary evi- 
dence, sufficient to warrant the conclusion, to be very 
probably connected with the inner terminals of motor, or posst- 


204 JOURNAL OF COMPARATIVE NEUROLOGY, 


bly of other efferent fibers, but the exact course of the connec- 
tion and the number of cellular elements composing it remains 
to be demonstrated. 

The central body is plainly shown to be connected with the 
fibrillar arch, and possibly the ocellar glomerule, and probably 
through these also with the nerves of the ocelli. Further it is 
connected with the fibrillar mass in front, and with that below 
it, but it does not appear to be connected with the mushroom 
bodies unless it be with their roots. It receives its fibers 
from cells situated above and behind it, and from some above 
the antennal lobes. 

The olfactory glomerule so-called, are shown to be formed 
by the tuft-like terminals of the fibers composing the antennal 
morulz, and in these the terminals of the fibers of the antenno- 
cerebral tracts seem to play the most prominent role. 

Several tracts of fibers from the optic lobes connect the 
latter with other parts of the brain than the mushroom bodies, 
and some of them doubtless form a part of an optic reflex 
tract with fibers from the ventro-cerebron and the ventral ‘cord. 

Such reflex tracts are not evident in connection with the 
antennal lobes, but the positions of fiber fragments seen indi- 
cate the possibility of reflex connections here also. 

Some of the fibers from the ocellar nerves terminate in the 
tubercles of the central body, others in the ocellar glomerule. 
Some also may connect with the fibrillar arch. Some of the 
large fibers pass downward and become the enormous fibers of 
the dorsal surface of the ventral cord. 

A tract of fibers, probably a continuation of what Binet 
calls the ventral column of the ventral cord, passes upward 
through the commissures and comes into connection above the 
cesophageal foramen with a dorso-ventral tract to the calices of 
the mushroom bodies. 

A tract from the cord that is probably a branch of ‘the 
ventral column passes forward to the posterior part of the an- 
tennal lobes giving off fibers on their branches along the way 
among the terminations of the fibers passing into the oral and 
the antenno-motor nerves. 


Kenyon, Zhe Brain of the Bee. 205 


In the ventro-cerebron of the bee there is recognizable but 
one root to each of the oral nerves. Of these that of the max- 
illary nerve is most prominently, and that of the mandibular 
nerve the most meagerly, developed. The roots connect with 
the ventral column as described by Binet (94) for one of the 
branches of the double root recognizable in the Coleoptera. 

Motor cells have been found in a ventral position in the 
ventro-cerebron, which does not accord with the distinction, 
based upon physiological experiments by Faivre (57) and Binet 
(94) of a dorsal motor anda ventral sensory area for each gan- 
glion of the ventral cord. This is reconciled by pointing out 
that it is the fibrillar connections that are destroyed in the le- 
sions produced dorsally, and the association cells and fibres and 
the terminations of sensory fibers in ventral lesions. If exper- 
iments be performed destroying individual cells and thus pro- 
ducing a consequent degeneration of their individual fibers, it 
will probably be found that the ventral surface of a ganglion 
may be described as a motor area. 


Bibliography. 


296. BrECHTEREW. Die Lehre von den Neuronen und die Entladungs- 
theorie. Neurol. Centralbl., 50-57 and 103-113. 

°82. BeLLoncr. Intorno alla struttura e alle connessione dei lobi olfattori 
negli artropodi superiori e nei vertebrati, Reale Acad, d. Lincet, 1881-2, Sr. 3, 
XIIL, 555-564. 

°78. BERGER. Untersuchungen iiber den Bau des Gehirns und der Ret- 
ina der Arthropoden, 47d. d. zool. Inst. Wien u. Triest, 1, 173-220. 


295. BERKLEY. Studies on the Lesions Produced by the Action of Cer- 
tain Poisons on the Cortical Nerve Cell. Bratz, LXXII, 473-496. 


°95. BrETHE. Studien iiber das Centralnervensystem von Carcinus mzenus 
nebst Angaben iiber ein neues Verfahren der Methylenblaufixation. Arch, ff. 
mtikr, Anat., XLIV, 579-622; Zetts. wiss. Mikr., XII, 130-132. 

°94. BINET. Contribution a l’étude du systéme nerveux sous-intestinale 
des insectes. Journ, 1 Anat. Phys., XXX, 449-580. 

°76. BRANDT. Anatomical and Morphological Researches on the Nervous 
System of Hymenopterous Insects. Ann. Mag. Nat. Hist., 4 Ser., XVIII, 504- 
506. 


°94. CHILD. The Antennal Sense Organs of Insects. Am, Wat., XXVII, 
608-609 ; Abstr. from Anat. Anz., XVII, 35. 


206 JOURNAL OF COMPARATIVE NEUROLOGY. 


°88. CuccaTiI. Ueber die Organization des Gehirns der Somomya ery- 
throcephala. Zezt. f. wtss. Zool., XLVI, 240-269. 

°76. DieTLt. Die Organization des Arthropodengehirns. Zezts. f. wiss. 
Zool., XXVIII, 488-517. 

°50. DuyARDIN. Mémoire,sur le system nerveux des insectes. Ann, Sci. 
IMT5 Bele DAO HOS Se SEG, 1 1%, 

°57. FAIvre. Du cerveau des Dytisques considéré dans ses reports avec 
la locomotion. Amn. Sct. Nat., VIII, 245-274. 

978. FLOEGEL. Ueber den einheitlichen Bau des Gehirns in den verschied- 
enen Insekten Ordnung. Ze?ts. f. wiss. Zool., XXX, Supplement, 556-592. 

9974. Foret. Les Fourmis de la Suisse. Lyon. 

995. HOoLMGREN. Die Tracheen Endverzweigungen bei den Spinndrtisen 
der Lepidopterenlarven. <Axat. Anz., XI. 340-346. 

°96. KopscH. Erfahrungen iiber die Verwendung des Formaldehyds bei 
der Chromsilber-Impregnation. Anat, Anz., XI, 727. 

°95. LENHOSSEK. Der feineren Bau des Nervensystems im Lichte neuester 
Forschungen. 2d Ed. 

°64. Leypic. Vom Bau des Thierischen KGrpers. 232 et seq. 

°79. Newton. On the Brain of the Cockroach. Quart. Journ. Micr, Sci., 
N.S., XIX, 340-356. 

980. PACKARD. The Brain of the Locust. Second Rep. U. S. Ent. Com., 
223-242. 

975. RasBL-RUECKHARD. Studien tiber Insektengehirne. Azchert and Du 
Bois-Raymond’s Arch. f. Anat., 488-499. 

990. RetTzius. Zur Kenntnis des Nervensystems der Crustaceen. fol. 
Ontersuch., N. F., I. No. 1. 

*92a. RetTzius. Die Endigungsweise des Reichnerven. Biol. Untersuch., 
Iie 18G5 WME INOS 25 ANNIE OS 

°92b. ReErzius. Ueber die sensiblen Nervenendigungen in den Epithe- 
lien bei Wirbelthieren. ol. Untersuch., N. F., 1V, No. 5, p. 40. 

995. RetTzius. Zur Kenntnis des Gehirnganglions und die sensiblen Ner- 
vensystems der Polychaeten. zol, Untersuch., N. F., VIII, No. 2. : 

990. SartntT-REMy. Contribution a étude des cerveaux chez les arthro- 
podes tracheates. Arch. Zool. Exper. et Gen., 2d Ser., V, 4th Memoir. 

987. VIALLANES. Le cerveau de la Guepe (Vespa crabro et vulgaris). 
Ann, Sci, Nat., 7 Ser., II, 5-100. 

988. VIALLANES. Le cerveau du Criquet (Oedipoda coerulescens et Cal- 
optenus italicus). Amn. Sct. Nat., 7 Ser., IV, 1-120. 

°93. VIALLANES, Etudes histologigues et organologiques sur les centres 
nerveux et les organs des sens des animaux articulés. Ann. Sct. Nat., 7 Ser., 
XIV, 405 et seq, 


Kenyon, Zhe Brain of the Bee. 


207 


EXPLANATIONSOF PLATES. 


EXPLANATION OF COLORS. 


Red. Cells and fibers of the mushroom bodies. 


Blue. 
Violet. 
parts of the nervous system. 
Green. 
Brown. 


Sensory fibers from cells outside of the central cerebral mass, 
Tracts connecting the calices of the mushroom bodies with other 


The superior dorso-cerebral commissure. 
Motor and other efferent fibers. 


Black.: Association and miscellaneous fibers and cells. 


EXPLANATION OF ABBREVIATIONS. 


a. c.—Anterior commissure. 

a 4.—Anteror roots of the mushroom 
bodies. 

a. t. c.—Antero-inferior dorso-cerebral 
commissure. 

A, Z.—Antennal lobe. 

a m.—External antenno-motor nerve. 

a. m',—Internal antenno-motor nerve. 

a, s,—Antenno-sensory fibers. 

c. 6 —Central body. 

c. ¢.—Fibrillar arch. 

c.cb.—Commissure of the central body. 

clx.—Calices of the mushroom bodies. 

d. v.—Dorso-ventral tract. 

e. clx.—Calices of the outer mushroom 
bodies. 

F,.—Oesophageal foramen. 

g. ch.—Tubercles of the central body. 

#. cle.—Calices of the inner mushroom 
bodies. 

z.4.—Inner roots of the mushroom 
bodies. 

#. o. c.—Inferior optic commissure. 

7,—Labial nerve. 

/.cbh,—Lower portion of the central 
body. 

7, t,2g.—The antero-posterior 
tract. 

ma,—Mandibular nerve. 

mx.—Maxillary nerve, 


optic 


o' o? and o’—Antenno-cerebral tracts. 
o' inner, o? middle, o? out- 
er. 

o, 6,—Olfactory bodies or glomerulz. 

o. c.—Ocelli, oc.! median; oc.? right, 
oc. left. 

oc. w.—Posterior group of fibers from 
the ocelli. 

op. 6.—Optic body. 

p-—Posterior nerve. 

p.t.c.—Postero-inferior dorso-cerebral 
commissure. 

p. o. t.—A posterior optic tract. 

r, 0. g.—Region of the ocellar glomer- 
ulze. 

s.—Stomatogastric nerve. 

s. n.—Nerve of the salivary glands. 

5. 0. ¢.—Superior optic commissure. 
5.@. g.—Ventro-cerebron or subceso- 
phageal ganglion. 

t. op. 6.—Anterior optic tract (to the 
optic body). 

st,—Stalks of the mushroom bodies. 

m, c.——The superior dorso-cerebral com- 
missure. 

wu. c. 6.—Upper portion of the central 
body. 

uw. ¢. 2¢.—Antero-superior optic tract. 

u. t. 3¢.—Postero-superior optic tract. 

v, ¢.—Ventral column. 

z.—Small cells of mushroom body. 


208 JOURNAL OF COMPARATIVE NEUROLOGY. 


PLATE XIV. 


Fig. 1. Frontal section several sections in front of the junction of the 
roots and stalks of the mushroom bodies, showing association fibers in the me- 
‘dian region, and the antennal morula. 7. Tract of fibers from cells behind the 
stalks to the anterior root of the mushroom bodies ; ¢., large trachea piercing 
the ocellar nerves and the dorsal region of the brain. s, Median nerve of the 
stomatogastric system; z, pyramid of small cells of the mushroom body. Pre- 
pared by the formol-copper-hzematoxylin method. 


Fig. 2, Frontal section through the stalks and inner roots of the mush- 
room bodies, embracing the superior and the antero-inferior dorso-cerebral com- 
missures. Same method as fig. I. 


Figs. 3 and 4. Frontal section. The two anterior roots of the mushroom 
bodies, showing the bands produced by the bushy terminations of the associa- 
tion fibers. By a modified Weigert method. 


PLATE XV. 


Fig. 5. Frontal section just behind the inner roots of the mushroom bod- 
ies, showing the fibers radiating from the central body. By the formol-copper- 
hematoxylin method. 


Fig. 6. <A frontal section further back, embracing the inferior optic com- 
missure and the middle and the outer antenno-cerebral tracts, Same method. 


PLATE XVI. 


Horizontal sections, all by the formol-copper-hzmatoxylin method. 


Fig. 7. Section cutting off the inner calices of the mushroom bodies and 
embracing the superior dorso-cerebral commissure and the top of the central 
body. c. cb, commissure of the central body. 


Fig. 8. A section at a lower level embracing the top of the fibrillar arch. 


Fig. 9. A section embracing the posterior commissure of the central body 
and passing just through the top of the lower portion of this. The ‘‘ pillars” 
of the fibrillar arch appear on either side of the light tract formed by the large 
fibers of the ocellar nerves. ~ 


Fig. ro. A section embracing the two inferior dorso-cerebral and the lower 
optic commissures, and two of the posterior optic tracts. The salivary nerve 
appears in the upper right hand corner as a light spot. 


PLATE XVII. 


Fig. 11. A frontal view of the brain exclusive of the outer portion of the 
optic lobes as a transparent object, showing the different bodies in dotted black 
outline and the fiber tracts in colors. For an explanation of the latter see expla- 
nation of colors. General outline and the outlines of the calices taken from a 
camera drawing of a brain mounted zz fofo in balsam. The outlines of the 
other bodies and the fiber tracts taken from sections prepared by the formol- 
copper-hzmatoxylin method and reconstructed by the aid of millimeter paper 
and by the superimposition of camera drawings. 


Kenyon, Zhe Brain of the Bee. 209 


PLATE XVIII. 


Fig. 12. A thick horizontal section embracing the region from the base of 
the inner mushroom bodies to the middle of the optic bodies. Outlines from 
formol-copper-hzmatoxylin sections as in fig. II. 


PLATE XIX. 


Fig. 13. The remainder of the brain below that represented in fig. 12. 
Outlines from sections as in fig. 12. 


PLATE XX, 


fig. 14. The lateral half of the brain viewed in optical sagittal section 
from the median plane outward. Outlines from thick sections prepared by the 
Golgi method. 


PEATE XXI. 


Cells and fibers from camera drawings of bichromate of silver preparations. 
Fig. 15. Cells and fibers from the antennal lobe. x 121. 


fig. 76. Fibers from the antennal nerve and an olfactory body. {In the 
section the right hand branch of the upper fiber was found, after the drawing 
here was made, to end in a small tuft of branches forming a portion of a glom- 
erulayx 20). 


fig. 17. Terminations of an association fiber in the junction of the stalks 
and roots of the mushroom bodies. x 121. 


fig. 78. Cells and fibers of the mushroom bodies. x 102. 
Fig. 19. Cell bodies of the same. x 121. 


Fig. 20. An olfactory glomerula formed by the termination of a fiber of 
one of the antenno-cerebral tracts. 


Fig. 21. The terminations of association fibers in the anterior root of the 
mushroom bodies. 


Fig. 22. A fiber passing across the top of one of the anterior roots and 
branching profusely in the region just inside of it. Some of the branches seem 
to penetrate the root. 


fig, 23. Fibers terminating in the calices entering about the origin of the 
stalk. 


Fig. 24. A drawing from three superimposed sagittal sections embracing 
the region from near the median line to the outer side of the anterior root. g, 
cells on the outside of the antennal lobe; above them and obscuring their fibers 
is a mass that may be composed of several over impregnated cells just above the 
antennal morula. For the rest see the diagrams in plates XVII to XX. 


fig. 25. Fiber entering an anterior root and apparently fusing with the 
parallel fibers. See text, p. 161. 


Fig. 26. The smaller kind of fibers terminating in the calices, x 102, 
See text, p. 157. 


210 JOURNAL OF COMPARATIVE NEUKOLOGY. 


Fig. 27. The terminations of the same magnified about 586 diameters. 


Fig. 28. Association fibers terminating in the anterior roots. This figure 
should be viewed from the adjacent corner of the plate. See text, p. 160. 


PLATE XXII. 


Fig. 29. Tracheal terminations in the calyx of the mushroom bodies, x 
Io2. Frontal section. 


Fig. 30. Fiber 36 of diagrams. x 121. Sagittal section. 

Fig. 37. Fibre 10 of the diagrams. x 102. It originates from a cell sit- 
uated above the antennal morula. Sagittal section. 

Fig. 32. Fibers from an oblique horizontal section, One with arborescent 
terminations in the central body. Two passing from one inner root to the re- 
gion above the anterior root of the opposite side. One passing forwards over 
the central body and branching in the region in front of it. 

Fig. 33. A twisted fiber from the region below and behind the optic body. 
Sagittal section. 

Fig. 34. Fiber from a sagittal section. x 121. It passes below an inner 
root and sends a branch up over the union of the same with the stalks. 

Fig. 35. Fiber passing antero-posteriorly below the central body and send- 
ing several branches up in front of the latter. x 121. 


Fig. 36. A portion of a motor fiber with its cell-body. See text, p. 174. 


Fig. 37. Fiber from the antennal region terminating arborescently in the 
central body. From a frontal section. x 121. 

Fig. 38. Fibers of the posterior inferior dorso-cerebral commissure. 
x 102. 

fig 39. Fibers apparently belonging to the antenno-motor nerve, and one 
commissural fiber of the ventro cerebron. 

Fig. go. Three fibers from two consecutive median sagittal sections. All 
three are seen to send a process each into the central body. A fragment in the 
lower part of the figure may be a portion of the outer or right hand fiber of the 
upper part of the figure. x 121. 

Fig. 41. A fiber from the base of the central body sending long branches 
down into the ventro-cerebron and to the anterior portion of the ventral cord. 
x 121. 


Fig. gz. Two fibers terminating in a so-called ocellar ganglion, x 121. 


Notg.—On page 174, line 8, for ‘“(p. )” read ‘‘(p. 161).”’ 
ERRATA, 


Page 133, line 5, add 1895-6. 
“184, ** 21, for Grylotalpa read Gryllotalpa. 
«192, ‘* 22, delete the copper-. 
Seo eee Ons. eco sani dl eae 
“192, ** 26, for Samomya read Somomya. 
210, delete the note at the foot of the page. 


THE ORIGIN AND GROWTH OF BRAIN CELLS IN 
PAP VADULT BODY: 


‘By Howarp AYERS. 


No part of our anatomical knowledge has had such slow 
growth as that pertaining to the origin and growth of brain cells 
and their ultimate relations to one another and to the rest of the 
body. 

The current theory holds that after the nerve cell once 
sends out its axis cylinder fiber and becomes a functional cell it 
is incapable of division or further reproduction. For example 
in the case of the human embryo it is stated that beginning 
with the fourth week the neuroblasts continue rapid growth up 
to the third month of fcetal life after which time no increase in 
numbers takes place. All increase in the brain or its parts 
after this period is due to the increase in size of the nerve cells 
(from one to 500 times their original size), the increase in the 
length and size of the sheaths of Schwann and the medullary 
matter, the growth of the blood vessels and the neuroglia ele- 
ments and the connective tissue additions to the brain. 


Double Auditory and Electric Ganghon Cells. 


In a former publication’ I figured and described some of 
the cells of the cochlear ganglion as multipolar and in some cases 
even as double cells. These latter cells were found to be com- 
posed of two nucleated cell bodies closely connected by a short 
connecting bar of protoplasm, in appearance like the peripheral 
and central fibers given off from the other ends of these two cells 
(Gi loc. ct, PIT, fig: 6, x). The origin of these double cells 
I was ‘unable to trace at that time but their occurrence in the 
ganglion was of interest to me in connection with the explana- 
tion of the origin and relations of the hair cells. In my recent 


1The Auditory or Hair Cells of the Ear and their Relations to the Auditory 
Nerve, /Journ. Morph., Vol. VIII, No. 3, 1893. 


212 JOURNAL OF COMPARATIVE NEUROLOGY. 


studies in the anatomy of the Torpedo brain similar double cells 
have again come under observation and it is the significance 
of these cells in the solution of the problems of brain 
-growth and especially the histogenetic processes which are in- 
volved to which I wish to invite special attention. I have been 
able to determine that these double cells in the Torpedo brain 
may by their complete separation from one another become the 
ordinary ganglion cells of the brain. In the figures which I 
shall publish soon, some of the cells are reproduced with great 
care. All the figures are camera drawings, and are taken from 
sections of the brain of an adult Zorpedo occidentalis, the sec- 
tions being stained by Weigert’s method. No single case of 
the karyokinetic process (in the nucleus) has been found in my 
preparations but the other relations and conditions of the cells 
leave no doubt that here in the Torpedo brain functional cells 
actually divide in the manner so well known for ordinary tissue 
cells. The stages of cell division which occur in the electric 
lobes are as follows: 

(1). Large motor cells, not to be distinguished from the 
ordinary functional cells except by the size of the nucleus and 
cell body. 

(2). Cells of the same size as (1) but with two nuclear 
bodies. Both may be close together in the centre of the cell 
or widely separated and lying near the periphery of the cell. 

(3). Cells showing an evident constriction of the proto- 
plasmic body between the nuclei as though about to divide. 

(4). Double cells with short connecting bars which. are 
usually large and band-shaped. 

(5). Double cells in which the connecting bar is drawn 
out into a thin filament, tapering conically from either cell body 
towards the other. 

(6). Since each nerve cell of the brain and ganglia has a 
peri-lymphatic capsule surrounding it, when the cell body is cut 
into two the peri-lymphatic space is not at once doubled but 
the two cells still lie in a common cavity. Because of this it is 
possible to trace the genetic relation of these electro-motor cells 
even after they have completely separated by the breaking of 


AYERS, Origin and Growth of Brain Cells. 213 


the connecting bands, as in those cases where the nerve cells 
become completely separated. Ultimately of course the lym- 
phatic spaces divide also by completing the capsular wall close 
about each cell. 


The Diviston of the Cell. 


Incisions of two very different kinds are found in the 
bodies of the electro-motor cells. Those produced by cell 
fission and those produced by unequal growth in response to 
local pressure upon some part of the cell by blood- Wessels or 
nerve fibers from some neighboring cell. 


Relation to Blood-vessels. 


The large ganglion cells of the Torpedo brain have appar- 
ently approached the limit of the size of nerve cells, which 
require, as we know, specially good facilities for nutrition 
(respiration and excretion included). The usual vertebrate 
brain is made up of cells varying in size from .002 m.m. to 
jum, m. and they are disposed about ' the) courses of (the 
blood-vessels in such a way as to admit of easy transfer of 
food to the cells and of the nitrogenous waste products away 
from the cells. 

In the Torpedo’s electric lobes, however, there is added to 
the usual arrangements a special feature in the relation of the 
cells to the blood-vessels. It consists in the migration of the 
over-grown ganglion cells to the walls of the arterial capillaries 
on the surfaces of which they spread themselves out and more 
or less completely surround the vascular tubes so as to present 
the appearance of being more or less deeply grooved or even 
perforated. The cell wall in immediate contact with the capil- 
lary cells is very thin and thus the very best facilities are pre- 
sented for the performance of the physiological activities of the 
brain cells. 

One of the anatomical causes of the increase in size of the 
electric lobe of the Torpedo is the production of new cells 
by the division of already functional electro-motor cells. 
This division occurs most abundantly at the ventricular sur- 
face of the lobe. 


214 JOURNAL OF COMPARATIVE NEUROLOGY. 


There is avery evident grouping of the cells in the elec- 
tric lobes in groups of 2, 3, 4, or larger numbers up to 12 or 
more. The axis cylinder processes of the members of a 
‘group converge towards a common centre, indicating, I think, a 
common origin from a single cell. 

While most cells of the electric lobe possess a_ single 
nucleus, the number having two nuclei is relatively large 
(500 in each lobe). Cells with three nuclei are occasionally 
found, 50 in each lobe. 

The centrosome is plainly visible in some of the cells. 


Morphological Laboratory, University of Missouri, Columbia, Mo., March 
10, 1896. 


THE INNERVATION OF THE AUDITORY EPG 
LIUM: IN MUSTEEUSHGANIS, DEAGAW; 


By Ao PDS MORRILT, 


I wish, at this time, to give a brief preliminary statement 
of the results obtained by the use of methylen blue in the study 
of the auditory epithelium of the ‘‘smooth dog-fish.”’ 

The fibers of the auditory nerve lose their medullary sheath 
on entering the sensory epithelium of the ampullae and branch 
at different levels, although the branches are much more num- 
erous at the base of the hair cells. From the latter branches, 
many of which extend horizontally, numerous extremely fine 
nerve fibers arise which end either in characteristic enlarge- 
ments in contact with the proximal portions of the hair-cells or 
pass between them to end free near the surface in similar but 
smaller enlargements. 

The nerve fiber often branches at the base of the hair-cell, 
the two portions closely adhering to the cell and ending in en- 
largements at nearly the same level on opposite sides of the 


MorriLi, /unervation of Auditory Epithelium. 215 


cell. In other cases the fiber can be traced to the cell and may 
end in a single enlargement at its very base. Occasional free 
nerve endings were found in the central portions of the 
auditory epithelium, at a considerable distance from the hair- 
cells, and end free in enlargements. 


Conclusions. 


(1) No continuation of the nerve into the cell was ob- 
served although the cells were semi-transparent. 

(2) No trace of Kaiser’s cup-like mass was found at the 
base of the hair-cells. 

(3) The triangular enlargements at the points where the 
nerves branch are due to the nerve sheath, through which the 
nerve fibers can be seen. 

(4) Satisfactory evidence of anastomosis of nerve fibers 
was not obtained. 

(5) There are two kinds of nerve endings in the audi- 
tory epithelium, the one being free near the surface and the 
other ending in knob-like structures in contact with the base of 
the hair-cells. 

(6) No varicosities were found on the nerve fibers in the 
best stained preparations, while the terminal enlargement was 
always present. 


Hamilton College, Clinton, N. Y. 


NEURAL TERMS, INTERNATIONAL AND 
NATIONAL.’ 


By Burt G. Wiper, M. D., 
Professor of Neurology, etc., Cornell University. 


Introduction. 


Five conditions have led to the preparation of this article 
at this time. 

$1. The American Neurological Association, at its session 
in Philadelphia, June 5, 1896, unanimously adopted the Report of 
the Committee on Neuronymy embodying the previous reports 
of three other American committees and extending the list of 
Latin terms recommended from eleven to forty ; see §8o. 

$2. The Anatomische Gesellschaft, at its session in Basel, 
April 19, 1895, adopted the Report of its Committee on Ana- 
tomische Nomenclatur, comprising a list of Latin names for all 
the visible parts of the human body (see Table VII), and pro- 
vided for its revision at intervals of three years. Presumably 
the Gesellschaft sanctioned the declarations of principles which 
had been published by the secretary of the committee (Krause, 
91, 94; see Part V.)? The list was published early in the 
summer of 1895 asa part of an article, ‘‘ Die Anatomisthe 
Nomenclatur,”’ by Professor Wilhelm His, constituting a ‘‘Sup- 
plement-Band” to the ‘‘Anatomische Abtheilung” of the 
Archiv fir Anatomie und Physiologie. Certain principles and 
certain portions of the list merit high commendation; others, 


1The more important parts of this article were embodied in a lecture, ‘‘The 
Present Aspects of the Nomenclature of the Brain,” delivered at the Marine 
Biological Laboratory, Aug. 3, 1896. 


2See the Bibliography, Part IX. The date after the name of a writer de- 
signates the year of publication. The joint works of S. H. Gage and myself 
are indicated by W. & G., ’82, ’86, ’89. 


Wiper, eural Terms. 217. 


in my opinion, are to be as deeply regretted. Among the least 
acceptable features are the designations and coordination of the 
encephalic segments and the assignment of parts thereto; 
able: We 


§3. In the official action of the Gesellschaft (Part V), and 
in arecent manual by the president of its committee, Professor 
Albert von Kolliker are declarations against the efforts of the 
American committees which may be due in part to ignorance 
or misapprehension of the facts. As chairman of two of the 
American committees and as secretary of a third, I may not 
inappropriately endeavor to remove the impediments to a clear- 
er comprehension of our position. I particularly desire to free 
the committees, their individual members, and the associations 
which they represent, from responsibilities not yet assumed by 
them. 


$4. In the article above mentioned Professor His not 
only evinces a failure to comprehend the aims of the Amer- 
ican committees, but also misrepresents what has been done 
by me as an individual. Such misrepresentations, unless 
corrected, might well, especially in Germany, impair the 
efficiency of my past and present utterances upon An- 
atomic Nomenclature. A correspondence begun in Decem- 
ber, 1895, has failed to adjust our disagreement, and it is most 
reluctantly submitted to other anatomists. In an experience of 
thirty-five years this is my first scientific controversy, and I 
trust it may be the last. 


$5. During the quarter of a century since my attention 
was first drawn to the defects of current anatomic terms my 
convictions may be assigned to five different stages dating re- 
spectively from 1871, 1880, 1884, 1889 and 1895. Beyond 
the last I now discern no opportunity for progress except- 
ing in the elaboration of details. It is my desire to devote the 
rest of my life to the study of the brain, and this seems to be 
a fitting time for submitting such statements of principle and 
suggestions of practice as may facilitate the labors of others 
upon Anatomic Nomenclature. 


218 JOURNAL OF CoMPARATIVE NEUROLOGY. 


§6. This article comprises nine parts as follows: 
I. Definitions of certain terms employed in the dis- 
cussion of Anatomic Nomenclature. 
| II. Stages of the writer's terminologic progress. 
III. Report of the Committee on Neuronymy of the 
American Neurological Association, with commentaries. 
IV. Discussion of the differences between certain terms in 
that report and those adopted by the Anatomische Gesellschaft. 
V. Reply to criticisms offered by the Anatomische 
Gesellschaft and by its members. 
VI. Correspondence with Professor Wilhelm His. 
VII. List of the Neural terms adopted by the Anatom- 
ische Gesellschaft and of those now preferred by the writer. 
VIII. Concluding remarks, 
IX. Bibliography. 


Part I. Definitions. 


$7. It is assumed that all who read this article seriously 
are familiar with the parts of the brain referred to, and that 
they have likewise a ‘‘ working knowledge” of ancient and 
modern languages. But space will be saved by the employ- 
ment of certain terms which are rather linguistic than anatomic, 
and some of which either are comparatively new, or are here 
used in new or restricted senses; these are therefore first briefly 
defined. The definitions of course apply to the Latin forms of 
the English words; the adjectives and other derivatives are 
self-explanatory. ’ 

$8. Onym.—From évepa, same as veya, a name.  Pro- 
posed by Coues (’82) in the sense of biologic name. It is 
seldom needed alone but is the essential element or base ($30) 
of many derivatives. 

§9. Toponym.—From onym and céxoz, place. A term 
indicating location or direction; e. g. /ateral, at the side; /aterad, 
toward the side ; ¢vansection, cutting across. 


1 Definitions may be found also in the more recent English and medical 
dictionaries. Some of the terms are discussed in my articles, ’85 and 8g. 


WILpER, Neural Terms. 219 


S10. Organonym.—The name of a part or organ; e. g. 
humerus. 


S11. NMeuronym.—The name of a part of the nervous 
system. 

S12. Polyonym.—A name consisting of more than one 
word; e. g. fissura centralts ; rostrum corporis callost; plexus 
choriotdea ventricult quarti ; iter a tertio ad quartum ventriculun. 
This use of the word jpolyonym is analogous to that of poly- 
anary, polygamy, etc.; see note to p. 518 0f W. &G, 89. 

§13. Dionym.—A term consisting of two words; e. g. 
vertebra thoracalts; arteria brachialis ; gyrus callosalis. Dionyms 
are perhaps the most common kind of polyonyms. They have 
a certain analogy with the technical names of animals and plants 
since the noun often indicates a group of similar or related 
parts and the adjective designates a specific member of the 
group. 

$14.  Trionym.—A term consisting of three words; e. g. 
vertebra thoracalis prima. Here, as with the so-called trinom- 
ials of zoology, the second adjective may be said to designate 
a subspecies. 

$15. Mononym.—A name consisting of a single word ; 
e. g. zsula. Strictly speaking a mononym is either a noun or 
some other word used asa noun. But the application may be 
conveniently extended as in the next definition. 

$16. Mononymic Qualifier.—A qualifying word, (adjective, 
participle or genitive), consisting of a single word; e. g. the 
second word in each of the following dionyms: Gyrus postcen- 
tralis (for G. centralts posterior); G. subfrontalis (for G. frontals 
mmfertor). 

$17. Ordinal Names.—These indicate the order or num- 
eric location of a member of a series; e. g. costa prima; verte- 
bra thoracalts prima." 


1 With any series extending lengthwise of the vertebrate body the member 
nearest the head is regarded as first. The only instance known to me of disre- 
gard of this conventional assignment is the enumeration of the segments of 
the brain in the schema of Prof. His as adopted by the German Committee. 


220 JOURNAL OF COMPARATIVE NEUROLOGY. 


$18. Attvibutive Names.—These refer, at least in part, to 
some real or fancied attribute; e. g. callosum; oblongata; 
vagus. 

$19. Szmle Names.—These express real or fancied re- 
semblances to other objects by means of the suffixes formzs or 
oides; e. g. restiformis, trapezoides. Most simile names might 
as well be converted into the corresponding metaphoric names ; 
e. g. restis, trapestum. 

$20. Melaphoric Names.—The names of non-anatomic 
objects are transferred to parts having some real or fancied 
resemblance thereto; e. g. pons, znsula, thalamus, falx. 

S21. Metaphoric Diminutives.—Since many parts are 
smaller than the more familiar objects whose names have been 
transferred to them the diminutive form is sometimes used; e. 
g. vallicula (from vallis); fasciculus (from fasczs) ; colliculus 
(from collts); clavicula (from clavis). Since, however, size is so 
variable and unessential an attribute, and since verbal diminu- 
tives are commonly longer than their originals, the latter might 
as well be employed. But this suggestion would not apply to 
a case where there are two of a general sort differing mainly in 
size; e. g. cerebrum and cerebellum ; falx (falx cerebrt) ; falcu- 
la (falx cerebellt). 

§22. Polychrestic Word.—One that does duty in many 
connections; e. g., occtpttalis, which in various combinations 
aids in designating at least twenty-five different parts. 

$23. Homonym.—A name applied to two or more differ- 
ent parts; an ambiguous term. An extreme case is that of os 
as signifying either a bone or an orifice; the oblique cases and 
derivatives of course distinguish them. J/edulla has been ap- 
plied to several parts. Afzphysis may designate the end of a 
bone or a part of the brain. Theoretically objectionable, the 
context commonly frees homonyms from serious ambiguity. 

$24. Jdionym.—A word which, at least in anatomy, re- 
fers to but one part; e. g., cerebellum; thalamus ; chiasma ; 
pons ; imsula. 

$25. Ldionyms by Recombination. Cornu posterius, as em- 
ployed by most anatomists, is a homonym, designating either a 


WILDER, Neural Terns. 22% 


cavity of the cerebrum ora feature of the myel (spinal cord). 
But fostcornu, as introduced by me in 1881, applies only to the 
cerebral cavity and is thus an idionym. 

§26. Contextual Explicituess.—For want ofa better phrase 
this may refer to the possibility of employing terms that might 
be ambiguous but for their association with others. A com- 
mon example is cord, which may be used in at least five senses, 
by the neurologist, the laryngologist, the surgeon, the obste- 
trician and the embryologist. When an entire publication or 
section of it refers to a group of organs of the same general 
character, then the generic element of their polyonymic desig- 
nations may be often omitted and the specific alone employed ; 
é. g., with arteries, fissures, gyres, cic. Indeed, to be abso- 
lutely explicit or idionymic in all cases would require many 
new names or the addition of genitives or other qualifiers to 
many already existing. 

$27. Locative Names.—TVhe location of a part is a general 
and comprehensive attribute and, as remarked by Owen, ‘‘sig- 
nifies its totality without calling prominently to mind any one 
particular quality, which is thereby apt to be deemed, undeser- 
vedly, more essential than the rest.” 

$28. Prepositional Locatives.—With these the qualifying 
prefix, a preposition or adverb, indicates the location of a part 
relatively to some other part, more important, more easily rec- 
ognized, or earlier designated. Praecuneus designates a cortical 
area just ‘‘in front of’ the cuneus. 

S29. Adjectival Locatives.—These indicate either the lo- 
cation of a part within some general region, or its membership 
of aseries. Vertebra thoracals designates a spinal segment in 
the thorax. Commassura anterior, cm. media, and cm. posterior 
distinguish members of a conventional series. J/esencephalon, 
prosencephalon and metencephalon designate members of a natural 
series, and the prepositions have the force of adjectives ; see 
S$$173-192. 

§30. Base (verbum basale).—The original or more essen- 
tial element of a derivative, as distinguished from prefixes, suf- 
fixes, inflective terminations, etc. 


222 JOURNAL OF COMPARATIVE NEUROLOGY. 


$31. Derivative.—A -word derived or formed either im- 
mediately or remotely from another; e. g., zworganic, organize, 
and organs are derivatives of organ. 

$32.  Correlative Names.—These are derivatives containing 
no obvious locative element but intended to indicate some re- 
lation between the part so designated and the part designated 
by the base; e. g., fissura calcarina associates an ectal fissure 
with the calcar, an ental ridge. 

$33. Lponyms.—Personal names, i. e., derived from the 
names of individuals; e. g., fissura Sylva; pons Varoli. Some 
objections to these are stated in W. & G., ’89, §59,' and as 
they are condemned by the German committee most of them 
will probable disappear. An exception perhaps should be 
jissura Sylvit (see $120). 

$34. Pectlonymy.?—Proposed by me (W. & G., ’89, §16) 
as a mononym for ferminologic variety or inconsistency within a 
single article or work; e. g., the use of fisswra and sulcus for 
the same cerebral furrow, of centralzs and Rolando for the same 
fissure. Between pp. 464 and 507 of Schwalbe’s ‘‘ Neurol- 
ogie”’ occur Crus fornicis (498), Fornix-schenkel (464), Forntx- 
saulchen (507), Gewolbe-schenkel (464). His (95) adopts Fora- 
men interventriculare but uses Foramen Monroi on p. 166 and 
‘* Monro'schen Loche’’ on p. 167. 

$35.  Daurect Pecilonymy.—In the cases mentioned above 
and others that might be adduced from nearly every work 
known to me, one and the same part is designated by two or 
more substantives or words used substantively. This is dzrect 
pecilonymy. A special variety of it occurs when different gen- 
eric names are applied to two homologous parts; e. g., in Hux- 
ley and Hawkins’ ‘‘ Comparative Osteology”’ the arm is called 
the ‘‘anterior exrtremity”’ the leg, the ‘‘ hind dd.” 


” 


1 There may be added the liability of misspelling ; oro has been spelled 
Monroe, Munro and Munroe; see my paper, ’8o, id. 


? From zozxilos, various, changeful, inconstant; compare zozxiAofovidos, 
of changeful counsel ; Zectlopoda, various footed. The unfamiliar term is per- 
haps the less objectionable in that it stands for a habit which may ere long be 
eradicated. 


WILDER, Neural Terms. 223 


836. Indirect Pectlonymy.—But when a certain substantive 
is used in one passage, and in another an adjective or other de- 
rivative from a different substantive, the pecilonymy is indirect 
or implied; e. g., ‘‘ certain fibers are called peduncular because 
they pass into the cvura cerebri.’’ Very commonly a certain 
fissure is named Rolando, but adjoining gyres, paracentral, 
anterior central, etc. 

$37. Pecilonymy by Permutation.—When a name, or the 
adjective part of a name, contains two or more elements of ap- 
proximately equal value, they are subject to accidental or in- 
tentional transpositions that may cause misapprehension. For 
example in his paper on the brain of Afzeles (Zool. Soc. Proc., 
1861) Huxley refers to the same fissure as occipito-temporal 
on p. 258 and as temporo-occipital on p. 260. One might 
infer that two different things were indicated just as, in chem- 
istry, Aydro-carbon and carbo-hydrate have different significations. 
Similar diversity of usage exists with regard to the occipital 
fissure, which is called by some occifito-partetal and by others 
parteto-occipital. Orbito-frontal and fronto-orbital constitute an- 
other instance. 

$38. <Abbreviational Pecilonymy.—The following is a good 
example of a bad system: in the translations of two of 
Meynert’s works occur corpus qguadrigeminum ; corp. quadrigem- 
mum; corp. quadrigem.; corp. quadrig.; corp. quad. 

$39. The Perpetration or Toleration of Fecilonymy may be 
ascribed to five mental conditions : 

A. Pure heedlessness. 

B. Indifference to the just claims of readers and especi- 
ally of students. 

C. Pride in the hardly gained familiarity with the syn- 
onymy of parts. 

D. Desire to avoid repetition, as in certain forms of lit- 
efahy,expresssion): see W. & G:, 80, $73, B, note: 

E. Unwillingness to commit oneself to a particular’ name. 

1 In some cases all the current titles of a part are so unacceptable that one 
recalls Shakespeare’s epigram as to the ‘‘ Small choice among rotten apples,” 


and the demand of the dissatisfied guest, ‘‘If this is tea, bring me coffee ; if it 
is coffee, bring me tea,” 


224 JouRNAL OF CoMPARATIVE NEUROLOGY. 


Such hesitation constitutes the only valid justification of pecil- 
onymy. But the same end might be gained by a simple de- 
claration, without the risk of confusing or misleading the reader. 

$40. Magnilogy.—The employment of lengthy or pon- 
derous terms when briefer would suffice. This is simply one 
form of what may be called anatomic esotery. Now that the 
choice is offered, the anatomist who deliberately says aponeurosis 
for fascia, anfractuosity for fissure, and convolution for gyre, 
thereby arrays himself with the village orator in whose turgid 
discourse a fire is always a conflagration. 

$41. Pertssology.—The following example of needless 
amplification occurs in a special article by a distinguished neu- 
rologist in a leading metropolitan medical journal: ‘‘ The an- 
terior column of gray matter extends throughout the spinal 
cord, and the upper enlarged intra-cranial end of the spinal cord, 
which is known as the oblong cord or medulla (medulla ob- 
longata).’”’ As shown in W. & G., 89, 529, §76, the informa- 
tion contained in these thirty-two words might have been given 
in fifteen. 

$42. LEguivalents, Synoyms, and Tsonyms.—Equivalents 
are terms meaning the same thing, e. g., pons, pons Varoliz, 
pont, and Lriicke. Strictly speaking, pous Varoliz isa synonym, 
or equivalent in the same language, while pout and Lriicke are 
isonyms or equivalents in other languages. But for simplicity 
all may be here regarded as synonyms, just as, in biology, sya- 
onymy embraces all the appellations of organisms, whatever 
their nationality. Hence one may recognize two groups of 
synonyms, viz., pavonyms and heteronyms. 

$43.  Paronyms and Heteronyms.—Excluding pons Varolit 
(the dionymic, eponymic synonym of fons), the other equivalents 
are the French font, the Italian ponte, the Spanish puente, the 
German JSviicke, and the English drzdge. Of these, the first 
three are obviously related to the Latin fous, while the last two 
have no such relationship. The former have been called by 
me paronyms,’ the latter, heteronyms; ’85, c,9; W. &G., ’86, 


1 Paronymy or paronymization includes what has been called word-adoption, 
word-appropriation, word-assumption, word-borrowing, etc. 


Wiper, Neural Terms. 225 


preface; W. & G., ’89, 519. A familiar illustration is the 
Latin canals, ot which canal is the English paronym, while 
heteronyms are Zube, passage, trough and water-course. The 
Greek épyavov might be rendered by part, tnsteument or agent, 
and these are its English heteronyms; but the Latin paronym 
is organum; the French, organe; the Italian, organo; the 
English, oxgan; and the German, Organ. Each of these is, 
so to speak, a geographic variety of the original or antecedent 
word; indeed it may be regarded as the same word modified in 
accordance with the genius of each language. The case may be 
compared with that of a traveller who maintains his essential 
identity notwithstanding ‘‘in Rome he does as the Romans 
do,’’ and in other countries conforms to the customs of the 
inhabitants in respect to garb and demeanor. 

$44. Methods of Paronymization.—For linguistic reasons 
paronymy is general and easy with the Romance languages, less 
so with the Germanic and with English. Still there are ex- 
amples enough (Tables II, 1V) to warrant the belief that into 
either may be adopted any Latin substantive or adjective.’ 
Paronymic methods vary with the language and with the word. 
Some of the modes of conversion of Latin words into their 
Anglo-paronyms are formulated in W. & G., ’89, §66. The 
instances there given involve more or less orthographic modifi- 
cation, ranging in extent from the case of fiver (from fibra) to 
that of alms (from eleemosyna.) These are changed paronyms. 

$45. Unchanged Paronyms.—But there are other evi- 
dences of paronymization, vzz., (a) Pronunciation, e. g., Parts, 
Detroit. (b) Hyphenation with a word unmistakably of another 
language; e. g. in Lalken-splentum, the hyphen indicates the adop- 
tion of the Latin sp/enzum asa German word. (c) Combination, 
e. g., Ponusfasern and other cases in Table IV. (d) Declaration 
that the writer regards the unmodified word as adopted.” (e) 
‘ ‘Also other and perhaps all parts of speech, but they do not concern us 
ere. 


*Were all foreign words italicized, then in a given case the non-italicization 
of a word would indicate its adoption. Since the Germans commonly capital- 
ize all nouns, that feature does not not necessarily signify that a word is regar- 
ded as an unchanged paronym; see Table. 


226 JouRNAL OF COMPARATIVE NEUROLOGY. 


Employment of the vernacular form of the plural or of an 
oblique case; e. g., the Latin plural of Jens is dentes, but the 
English is /enses; so atlas (atlantes) atlases; enema (enemata) 
enemas ; animal (animatia) animals: in the phrase ‘fibers of 
‘the callosum,” the last word might still be regarded as Latin; 
but if one said ‘‘callosum’s fibers’? the English possessive 
would indicate paronymization. 

$46. International and National Terms.—By general 
consent Latin constitutes a common or international language 
for scientists. National terms may be either unrelated to 
the Latin antecedents,’ hence heteronyms ; or obviously related 
thereto, hence paronyms. Seahorse, Cheval marin and See- 
pferd are synonyms (in the broader sense, $42) but to either an 
Englishman, a Frenchman or a German, two of them are for- 
eign words and unacceptable. Hzppocampus is distinctly a 
Latin word, and the frequent occurrence of such imparts a ped- 
antic character to either discourse or printed page. //zppocamp, 
hippocampe, hippocampo and HHippokamp are as distinctly nation- 
al forms of the common international antecedent (not to in- 
voke the original Greek izzézapzos), and are readily recognized 
by all, while yet conforming to the ‘‘genius”’ of each language. 

§47. The Paronymic Advantages of Mononyms.—The ob- 
ject of paronymy is to endow anatomic language with national- 
ity without obscuring its internationality. With mononyms 
the paronymic changes (if any) are slight, involving mostly the 
termination, or, with German, the capitalization of nouns and the 
occasional replacement of c by &. The word is readily recag- 
nized, and its abbreviation would be the same in any language. 
But with polyonyms the relative position of the substantive and 
the qualifier is commonly reversed in the two groups of lan- 
guages, Romaniform and Germaniform. In the former the 
noun more often precedes, in the latter it almost always follows.’ 
Hence there is a different aspect of the entire term, and the 


1Or related so remotely that the connection is obscure. 


2Notwithstanding the familiar exceptions, a/ma mater, pia mater, and 
notary public. 


Wiper, Neural Terms. 2277 


abbreviations are transposed. The Anglo-paronym of commis- 
sura posterior is posterior commissure, and the respective abbrevi- 
ations might be c, fg. and f, c.; but if the Latin dionym be mon- 
onymized into postcommissura, the English paronym Is postcom- 
missure, and the abbreviation fc. answers for both. See gg61-64. 

$48. Lemitatiors to Paronymy.—As already admitted 
Haniohe 
bids the rigid and universal application of the principle of par- 


with regard to mononymy (§88), the ‘‘nature of things 


onymy. Certain parts, so exposed or so vital as to have gained 
early and popular attention, have received vernacular names or 
heteronyms which are brief and generally understood. Such 
are head, hand, foot, heart and brain. Indeed the use of the 
Latin equivalent for either of these would impress most persons 
as pedantic. But this concession of, for example, the suffici- 
ency of dvazn instead of encephalon does not warrant the reten- 
tion or formation of an indefinite number of inflectives, deriva- 
tives and compounds from the heteronym. The same remark 
applies to other languages.’ 


Part Il. Stages of the Writer's Terminologic Progress. 


$49. The following summary of the changes of my views 
during a quarter of a century shows, I trust, a general advance 
in the comprehension of the subject and justifies me in com- 
menting upon the labors of others. 

§5o. I. 1871-79. Inan effort to confirm, extend and 
modify certain morphologic ideas of my teacher, Jeffries Wy- 
man, I enumerated (71, 172) the following requirements of 
technical terms: 1. Classic Derivation. 2. Capacity for In- 
flection. 3. Brevity. 4. Independence of Context for Signi- 
fication. 5. Non-ambiguity to the Ear as well as to the Eye. 
6. Previous Use in a Kindred Sense. 

Of the thirty-three names then adopted or proposed for 


1 Of the two German vernacles for encephalon, Gehérn is more commonly 
used alone and Arn in composition. On my list there; are 35 compounds of 
Gehirn and 106 of Hirn; moreover, of the former, one-half are duplicated among 
the latter. 


228 JOURNAL OF COMPARATIVE NEUROLOGY. 


the limbs, their segments and joints’ all were mononyms (§ 15). 
With most were given the ‘singular and plural nominative and 
genitive. Two, omozone and tschizone, were Anglicized (paro- 
_nymized). The designation of the fingers by the adjectives, z- 
dex, medius, annularts and minimus, without preposing the sub- 
stantive digitus, recognized Contextual Explicitness (§ 26) and 
the validity of adjectival nouns (§ 117). 

$51. Then, as now, the most desirable (yet not absolutely 
essential [$§ 67-70] ) attributes of technical terms seemed to 
me (1) Classic Derivation, (2) Capacity of Inflection. But 
both these had been adumbrated long before by Barclay (03) 
and Whewell (’40), and distinctly enunciated by Owen (’46, 171) 
in the immortal paragraph wherein sye/on was proposed : 

‘‘The fore part of the neural axis * * is called the brain or 
encephalon; the rest I term mye/on, (Greek pvedés marrow). As 
an apology for proposing a name, capable of being inflected 
adjectively, for a most important part [see W. & G., ’89, § 48] 
of the body which has hitherto received none, I may observe 
that, so long as the brief definitions, ‘marrow of the spine,’ 
‘chord of the spine,’ are substituted for a proper name, all pro- 
positions respecting it must continue to be periphrastic, e. g., 
‘diseases of the spinal marrow,’ ‘functions of the spinal chord,’ 
instead of myelonal [myelic]? diseases, myelonal functions; or 
if the pathologist speaks of ‘spinal disease,’ meaning disease 
of the spinal marrow, he is liable to be misunderstood as refer- 
ring to the disease of the spinal or vertebral column. But were 
the anatomist to speak of the canalin the spinal marrow of fishes 
as the ‘myelonal canal’ he would at once distinguish it from 
the canal of the spinal column. The generally accepted term 


‘ Among the few new terms were omos, for shoulder-joint, and omozona and 
éschizona, for shoulder-girdle and pelvic-girdle respectively. They still seem to 
me preferable to the ‘* Articulatio humeri,” ‘* Cingulum extremitatis supertoris” 
and ‘‘ Cingulum extremitatis inferiorts’’ of the German list. 


? On several previous occasions (’85, 354; ‘85, 12; ’89, 531) I have shown 
that analogy with words like angel and angelic (from 4yyeoc) calls for myel and 
myelic as the English nominative and adjective of myelon ; myelonal is clumsy, 
and analogy would involve the replacement of encephalic by encephalonal. 


WILpvER, Neural Terms. 229 


‘chorda’ or ‘chorda dorsalis,’ for the embryonic gelatinous 
basis of the spine, adds another source of confusion likely to 
arise from the use of the term ‘spinal chord,’ applied to the 
myelon, or albuminous contents of the spinal canal.’” 

§52. In 1873 (’73, 306) Owen’s examples of ectogluteus, 
mesogluteus and entogluteus \ed me to propose the locative mono- 
nyms ectopectoralis and entopectoralis for the two frequently 
named muscles whose relative proportions in most mammals 
are so misrepresented by the adjectives major and munor. 

$53.) Wh haveialneady(/85,.\¢, 51190; 2, 1) expressed my 
sense of obligation to Owen’s terminologic precepts and exam- 
ples, and my regret that the limits of the article, ‘‘ Anatomical 
Terminology”’ (W. & G., ’89) did not permit even more ex- 
tended selections than are embraced among the ‘‘ Aphorisms 
respecting Nomenclature”’ therein. 

$54. II. 1880-1883. While preparing a paper on the 
brain of the cat (’81, @) and (with S. H. Gage) a volume of 
directions for laboratory work (W. & G.,’82) I adopted from 
Barclay (’03) the unambiguous toponyms (§9) dorsal, dorsad, 
etc.; replaced his meszon by meson, the direct paronym of péoov; 
added ectal, ental, etc.; and simplified some organonyms, espe- 
cially muscular (W. & G., ’82, 207) and neural (’80, f; ’81, 4, a) 
in the following ways: (a) Dropping unessential adjectives 
( optecus from thalamus and chiasma); eponymic (§33 ) qualifiers 
(Varolit, Reilit, Rolando); and generic nouns (corpus, mater, and 
membrana ) from adjectives which were sufficiently distinctive 
and could be used as substantives (callosum, dura, mucosa) ; 
(b) substituting prepositions for adjectives (e. g., postcommius- 
sura for commissura posterior); (c) Replacing certain polyonyms 
by mononyms more or less nearly akin thereto (e. g., damina 
terminalis by terma); and (d) Abandoning the anthropotomic 
misnomers of the encephalic cavities in favor of mononyms co- 


1 The foregoing first appeared half a century ago; the mononym mye/on was 
employed consistently by Owen, and on at least one occasion by his rival, Hux- 
ley (’72, 65). These facts should secure for it the consideration due to high 
authority and moderate antiquity, and forestall any hasty proposition to employ . 
it in a different sense. 


230 JOURNAL OF COMPARATIVE NEUROLOGY. 


ordinated with the commonly accepted titles of the encephalic 
segments (e. g., Aguaeductus Sylvit and Iter a tertio ad quartum 
ventriculum for mesocoelia )' 

§55. In respect to terminologic practice, before the publi- 
cation of any paper or book there was made a list of the Latin 
terms to be employed, with abbreviations thereof, and these 
were adhered to throughout. 

§56. Notwithstanding their defects, these efforts to im- 
prove anatomic language elicited favorable comment, helpful 
criticism, and more or less actual adoption from Oliver Wendell 
Holmes (’81), Joseph Leidy (’85, ’89),? Henry F. Osborn 
(’83, ’84), E. C. Spitzka (’81) and R. Ramsay Wright (’85). 

$57. Ill. 1884-1888. Although now satisfied as to the 
correctness of the general system and as to the excellence of 
most of the individual terms, I began to realize more fully the 
magnitude and difficulty of the task and the necessity for coun- 
sel and cooperation. In the summer of 1884, at my sugges- 
tion, committees were appointed by the American Neurological 
Association and the American Association for the Advance- 
ment of Science. The constitution of these committees (§ 80) 
insured that no hasty action would be taken, and warranted the 
hope that any conclusions reached by them would be consid- 
ered seriously here and abroad. Personal conferences were held 
when practicable, but most of the work of comparing views and 
preparing preliminary reports was done by correspondence. 

$58. As collaborator on a medical dictionary (Foster, 
"88-'94), I undertook to obtain a list of names already applied 
to parts of the central nervous system. In 1888 the total was 
10500, distributed as follows, in round numbers: Latin, 3100; 


1 Nothing in my terminologic experience has been more gratifying and en- 
couraging than the approximate coincidence of a similar proposition by T. Jef- 
fery Parker (’82, ’84.) 


? While engaged upon the new edition of his ‘‘ Anatomy,” Professor Leidy 
wrote to me under date of Jan. 20, 1885: ‘‘I wish to aid in reforming the 
nomenclature of Anatomy, and in doing so propose to Anglicize the names to 
some extent (246). Will you please look over this list of muscles and tell me 
whether I can do better with any of the names.’”’ Ten days later he submitted 
a list of the many terms. Many of my suggestions were adopted. 


WILDER, WVeural Terms. 231 


English, 1800; French, 1800; Italian and Spanish, 900; Ger- 
man, 2900. Assuming the number of parts or features to be 
500-600, there were evidently many superfluous neuronyms, 
especially in Latin and German. ‘The excess in these two lan- 
guages might be accounted for in part by the international 
character of the former, and by the large number of publica- 
tions in the latter. 

§59. Buta careful scrutiny disclosed two other causes : 
(1) Many of the Latin names, especially the older, comprised 
so many words as to constitute descriptive phrases, and to fur- 
nish opportunity for conscious or unconscious abridgement and 
permutation (§ 37); each resultant combination had to be re- 
garded asaname. In W. &G., ’89 $56, are enumerated no 
less than twenty-three distinct Latin names for the fibrous 
bundle connecting the cerebellum with the oblongata; they 
average nearly 2.7 words each." 

§60. Of the German names a small proportion (58, or 
two per cent. of the total) had any obvious resemblance to 
equivalent Latin terms (/7ssur to fissura, Commussur to Com- 
missura, Centralcanal to Canalis centralis); the vast majority 
were vernacular translations (e. g., Briicke, Schenkel, Seepferd- 
efuss, Sehhiigelpolster).” Different writers made different trans- 
lations, and considerable variation occurred in different parts of 
the same publication (§ 34). Hence there arose a multitude of 
terms, acceptable and intelligible only to readers of the same 
nationality, and bearing no relation to the original or interna- 
tional Latin terms. Ina greater or less degree the same might 
be said of the other modern languages. 

S61. It will be seen that two opposing influences were 
operating. Each anatomist preferred to employ terms belong- 
ing to his own language ; at the same time he preferred that 
other anatomists should employ Latin terms with which he was 


1 All these might be replaced by the single word, postpedunculus. 


2 Without imputing even so worthy a motive as national self-satisfaction, 
the effect was asif certain neurologists had yielded to a desire to confer upon 
the printed page an obtrusively German aspect. 


232 JOURNAL OF COMPARATIVE NEUROLOGY. 


already familiar, or which were intelligible without an intimate 
acquaintance with other modern languages than his own. 

§62. With a view to reconcile these two opposing ten- 
~ dencies I formulated (’85, c) the distinction between heteronyms 
(§ 43) and paronyms (§ 44), and proposed that, with few ex- 
ceptions (§ 48) heteronyms should be discarded in favor of 
paronyms. ‘‘Since each paronym suggests the original Latin 
name, the latter forms a bond of intelligence between writers 
and readers of different nationalities.” 

§63. The international advantages of paronyms over het- 
eronyms have been distinctly recognized, and the principle in- 
dorsed, by the American branch of the International Committee 
of Biological Nomenclature, and by the American Association 
for the Advancement of Science (Proceedings, 1892, 233). 

$64. That mononyms are more readily and uniformly 
paronymized than polyonyms, and dionyms than other polyo- 
nyms, has been already mentioned (§ 47) and is indeed self- 
evident. 

$65. IV. 1889-1894. But the recognition of the nature 
and causes of neuronymic hypertrophy and deformity, and 
even the formulation of general principles of relief, still left un- 
accomplished the necessary operations of excision and correc- 
tion. My inability to decide in season which should be re- 
garded as the names, and which as merely synonyms, was one 
of the reasons for not accepting the invitation of Dr. Foster to 
frame the definitions in the dictionary above mentioned. Par- 
tial lists had been prepared in connection with the ‘‘ Anatomi- 
cal Technology” (W. & G., 82, 436-438) and the ‘‘Cartwright 
Lectures’ (’84, @). The latter list (84, 2) contained 115 names 
exclusive of the fissures and gyres and blood vessels. In con- 
nection with a paper, ‘‘Owen’s Nomenclature of the Brain ” 
(90, g), there was presented to the Association of American 
Anatomists a ‘‘ Macroscopic Vocabulary’ of about 200 names, 


1 The history of the process, and the names of my present and former col- 
leagues who so materially assisted me, are given in several publications (’85, ¢; 


W. & G., ’86; ’89). 


WILDER, Weural Terms. 233 


with synonyms and references (’90, %). The vessels, fissures 
and gyres were estimated at 140, and lists of them were pub- 
lished at various periods (85, ¢, A, g; ’86, g). 

$66. This made a total of about 340 parts or features of 
the central nervous system the designations of which I had se- 
lected or framed from among the vast accumulation of available 
terms. These names had already been found serviceable in the 
research and instruction carried on under my direction; and 
they were embodied in the articles on the gross anatomy of the 
brain (89, @, 0, c; 93, @, &, c); and questions involved in their 
adoption were discussed at length in ‘‘ Anatomical Terminol- 
ony  (W.)&'G., ’So). 

$67. V. 1895-1896. Among the requirements of tech- 
nical terms enumerated in 1871 was ‘‘Independence of Context 
for Signification.”” The rigid application of this would exclude 
all homonyms (§23,) and would require every term to be 
absolutely explicit. It was perhaps not unnatural for a com- 
parative beginner in the subject to make such a rule, and, hav- 
ing made it, to adhere to it somewhat persistently as in the 
following cases (S$68-70). 

$68. Of the three current appellations, conarium, epiphy- 
sis and corpus pineale, the last was rejected unhesitatingly asa 
polyonym, and the second as applying equally (without the 
qualifier cerebrz) to the separable end of a growing bone; as 
recently acknowledged (’96, 6) I long resisted the precept and 
example of H. F. Osborn and E. 'C. Spitzka in favor of epzph- 
ysts as correlative with hypophyszs, and failed to recognize the 
full force of Dall’s remark, ‘‘The human mind wearies of too 
many names and much more readily assimilates a new meaning 
for an old one,”’ although it was printed as Aphorism XV in 
Wer eerGi 89). 15 20; 

$69. Likewise, although favoring the general plan of ren- 
dering the Latin ae and oe by e in Anglicised (paronymized) 
words,’ I retained the diphthong in coea and its compounds 


In this country no medical writer has more persistently and vigorously 
urged this simplification than the former editor of the (Phil.) Medical News 
Gould, G. M., ’94, ’96. 


234 JOURNAL OF COMPARATIVE NEUROLOGY. 


(from xocdéa, a cavity) for the sake of distinguishing them from 
the derivatives of x/j7j, a tumor. I now frankly acknowledge 
the non-necessity of the diphthong even for the discrimination 
of encephalocele, the normal cavity of the brain, from the same 
word signifying an abnormal protrusion of the organ.' 

§70. In August, 1884 ('84, a, 114) I proposed to replace 
the common polyonym azzs cerebro-spinalis, and even Owen’s 
myelencephalon, by the brief mononym xeuron, warranted by nex- 
ralis, neurenterica, etc. and correlated with exteron (canahs al- 
mentaria) and axon (axis somatica). The term was used by 
Minot (’92, 606); Stowell (85); Waters (’91,362) and others. 
Its abandonment by me in favor of newraxis (’89, @) was due 
to two later observations: (a) the prior use of xeuraxzs* in the 
same sense: (0) the prior application of xeuvon to a part of an 
invertebrate eye. I have since been led to believe that I was 
unduly influenced by these considerations. Unfortunately the 
matter is now complicated by (a) the application of xeuron to 
the entire nerve-cell including its processes, and (0) the designa- 
tion of the ‘‘axis-cylinder process” by xeuraxon, easily con- 
founded with neuraxis.* I have already declared (’93,100, 
95, ¢’45) my lack of personal feeling in the matter, but the 
more I think of it the greater appear to me the advantages of 
neuron. In view of the practical efficiency of ‘‘contextual ex- 
plicitness” (§26), its ‘‘invertebrate’’ use may be ignored, and 
where there could be any doubt as to whether xeuvon referred 
to the entire nervous axis or only to one of its histologic con- 
stituents smacroneuron and wmucroneuron might be employed. 
Cases not strictly analogous, and yet worthy of note in this con- 
nection are the general use of Jody and belly for parts of a mus- 


1Those who are interested in vicissitudes of opinion may think I ought to 
admit that for a certain intermediate period the e alone was used in the paron- 
yms of coelia; 84, a. 


7In the ‘‘Dictionnaire de Medicin” of Robin and Littré occurs névraxe, the 
Galloparonym of a potential antecedent, mewraxis; but neither the propounder 
nor the first adopter is named. 


’For some history and discussion of these and kindred terms see the papers 
of Fish ('94) and Baker (’95). 


WILDER, WVeural Terms. 235 


cle; and of ¢arsws and cz/um in both macroscopic and micro- 
scopic senses. Whatever may be the outcome I shall always 
regret the confusion arising from what I now regard as a mani- 
festation of excessive conscientiousness. 

S71. Terms Withdrawn.—TVhrough ignorance, misappre- 
hension, timidity, or over-confidence, I have at various times pro- 
posed or employed needless or objectionable terms. Their for- 
mal withdrawal is here made in accordance with a conviction 
which was expressed (’91,a@) five years ago: ‘‘Since every 
one makes mistakes, the interests of all concerned would be 
subserved by the adoption of the custom of each correcting his 
own, either as soon as discovered or periodically ; a sort of sci- 
entific confession of sins. The natural corollary to this would 
be that each well-disposed discoverer of another’s fault would 
inform him privately so that he might make prompt correction. 
This plan I have followed in several cases, and have reason to 
believe it has served to avoid personal irritation and the need- 
less repetition of criticism.” 

§72. The following terms are hereby withdrawn: //yfo- 
campa (for hippocampus [mazjor]); W. & G., ’86, 4000; Spitzka, 
84; Vicq d’Azyr, 1786, 61, e¢ seg. Torus (for tuber [ctnereum)). 
Lenum (for torcular | Herophil’]|;’84).  Cerebrocortex (for cortex 
cerebri or cerebral cortex). Cerebellocortex (for cortex cerebelli or 
cerebellar cortex. Comimissure habenarum (for supracommissura); 
W. & G., 86,4008. Mediventricle (for ‘‘third ventricle’’); ’80, d 
Lativentricle (for ‘‘lateral ventricle”); 80 ad. Procele (for para- 
cele); ’84, c; W. &G., '86, 400. Coele and its compounds (for 
cele and its compounds); (§69). ‘ 

§73. If the foregoing list of my verbifactive sins appears 
damagingly large, let the critics scan their own records with 
equal closeness; I have at least been consistent within the lim- 
its of a single publication. 

$74. Fifteen years ago I was so oppressed by the length 
and obtrusive Latinity of the current names for the segments 
of the brain, prosencephalon, etc., as to suggest (’81, 4) that they 
might sometimes be abbreviated to prosen., etc. Three years 
later, in accordance with the analogies of quad (for guadrangle), 


236 JOURNAL OF COMPARATIVE NEUROLOGY. 


grog (for grogram), photo (for photograph), stereo (for stereotype), 
and consols \(for consolidated annuities), I ventured (84, 4) to 
drop the sign of abbreviation and to use these verba decaudata 
as real words. Their withdrawal a year afterward (854, 
354) may have been due partly to the horror of some anatom- 
ic friends in whose eyes they were needless and indefensible 
‘nicknames ;’’ but I was influenced mainly by the revulsion 
attendant upon the formulation of the principle of paronymy 
whereby the shorter prosencephal, etc., were legitimized. Yet 
I anticipate the eventual rehabilitation of the abridged forms. 

§75.  <Acknowledgements.—I have had more or less fre- 
quent conference or correspondence with nearly all the members 
of the four committees named elsewhere (§§80-84) and with 
other scientific or literary authorities. Only by investigators, 
teachers, and practitioners equally eminent, preoccupied, and fa- 
miliar with current terminology, can it be wholly realized what it 
meant for these men to give prompt and full attention to que- 
ries and propositions that threatened to disturb the verbal basis 
of their intercommunications. Reviewing the experience, I am 
amazed at the uniform readiness and kindliness of the responses? 
and can truly say that even when not wholly or directly encour- 
aging they were always fruitful. To four men are due par- 
ticular acknowledgements. | 

$76. As student (1873-1877), as assistant (1875-1880), as 
colleague (since 1880) and as collaborator (‘‘ Anatomical Tech- 
nology,’ 1880-1892; ‘‘ Anatomical Terminology,” 1888-1889) 
Simon H. Gage has been constantly and preeminently helpful. 

§77. Edward C. Spitzka (§80, note) one of the most 
learned, progressive and productive American neuro-anatomists, 
generously entertained the new terms (’81), adopted some, and 
for others proposed improvements; nay, this undaunted up- 
holder of an unpopular opinion in a period of intense political 


1The last case to meet my eye is Azppos for hippopotami (The Nation), 1896; 
verily ‘‘the slang of one age may become the purism of the next.” 


?Their nature made it the easier to meet with equanimity the few attempts 
to check terminologic progress by ridicule. For the response to one of these 
see my paper ’86, f. 


Witper, Neural Terms. 237 


excitement’ went so far as to say that some of my suggestions 
had been long in his own mind but that he had ‘‘lacked the 
courage to bring them before his colleagues.’’ Dr. Spitzka’s 
cordial interest has never abated, and I only lament that more 
practical duties leave him less time now than formerly for re- 
search in the anatomy of the brain. 

§78. Ihave already expressed (W. & G.,’89, §2 note) my 
appreciation of the erudition and kindness of my colleague in 
Comparative Philology, Benjamin I. Wheeler. Aside from in- 
formation imparted at personal interviews, the etymologic and 
linguistic points upon which he has enlightened me cover nearly 
one hundred of the ‘‘ correspondence slips.’” 

$79. To quote his own words, ‘‘The last thing an old 
teacher wants is a new set of terms for a familiar set of objects.” 
Yet this did not prevent Oliver Wendell Holmes, then for the 
third of a century professor of Anatomy in the Harvard Medi- 
cal School, from writing, May 3, 1881, a letter containing the 
following passages: 


‘«T have read carefully your paper [’81 4] on Nomenclature. I en- 
tirely approve it as an attempt. I am struck with the reasonableness 
of the system and the fitness of many of the special terms. ‘The plan 
is an excellent one; it is a new garment which will fit Science well, if 
that capricious and fantastic and old-fashioned dressing lady can only 
be induced to try it it on.” The entire letter is printed on pp. 11-12 
of (W. & G.,’82). It was asource of comfort tome and doubt- 
less led many to consider seriously suggestions that might otherwise 
have been ignored or repelled. 


Part ITI. Action of the American Neurological Association. 


$80. On the 5th of June, 1896, ata regular meeting in 
Philadelphia, the American Neurological Association adopted 


1As an expert at the trial of Guiteau he held the mental constitution of the 
assassin to be abnormal; see Alendst and Neurologést, 1883, April et seq. 


*For the use of slips in correspondence see W. and G., ’86, 52 and Sczence, 
Jan. 16, 1885, p. 44. Z 


238 JOURNAL OF COMPARATIVE NEUROLOGY. 


unanimously the ‘‘ Report,of the Committee on Neuronymy.””? 
The recommendations were as follows: 


1. That the adjectives DORSAL and VENTRAL be employed 
in place of posterior and anterior as commonly used in human 
anatomy, and in place of wpper and /ower as sometimes used in 
comparative anatomy. 


2. That the cornua of the spinal cord, and the spinal 
nerve-roots, be designated as DORSAL and VENTRAL rather than 
as posterior and anterior. 

3. That the costiferous vertebrae be called THORACIC rather 
than dorsal. 


4. That, other things being equal, Mononyms (single-word 
terms) be preferred to polyonyms (terms consisting of two or 
more words). 


5. That the Azppocampus minor be called CALCAR ; the /zp- 
pocampus major, HIPPOCAMPUS; the fous Varoliz, pons; the 
znsula Retlit, INSULA; pra mater and dura mater, respectively PIA 
and DURA. 


6. That the following be employed rather than their va- 
rious Synonyms: HYPOPHYSIS, EPIPHYSIS (for covarium and cor- 
pus pineale), CHIASMA, OBLONGATA, LEMNISCUS, MONTICULUS, TEG- 
MENTUM, PULVINAR, FALX, TENTORIUM, THALAMUS, CALLOSUM, 
STRIATUM, DENTATUM, MESENCEPHALON, PALLIUM, OLIVA, CLAVA, 
OPERCULUM, FISSURA CENTRALIS (for £. Rolando, etc.), F. CALCAR- 
INA, F. COLLATERALIS, F. HIPPOCAMPI, CUNEUS, PRAECUNEUS, 
CLAUSTRUM, FORNIX, INFUNDIBULUM, VERMIS. : 


1 The committee was appointed by the President of the Association, upon 
the suggestion of the writer, at the regular meeting in New York city, June 20, 
1884. One of the most interested of the original members, Dr. W. R. Birdsall, 
has since died. It now comprises Henry H. Donaldson, Ph.D., professor of 
Neurology, Chicago University; Landon Carter Gray. M.D., professor of Ner- 
vous and Mental Diseases, New York Polyclinic; Charles K. Mills, M.D., pro- 
fessor of Diseases of the Mind and Nervous System in the Philadelphia Poly- 
clinic; Edward C. Seguin, M.D., professor of Diseases of the Mind and Ner- 
vous System in the Medical Department of Columbia University ; Edward C, 
Spitzka, M.D., formerly professor of the Anatomy and Physiology of the Ner- 
vous System in the Post-graduate Medical School of New York city; and B. G, 
Wilder, Chairman. 


WitpveER, Neural Terms. 239 


$81. Sections 1, 2, 3, and 5 constituted the ‘Preliminary 
Report of the Committee on Anatomical Nomenclature”’ of the 
Association of American Anatomists which was adopted unani- 
mously by that body Dec. 27, 1889." 

$82. Section 4 is substantially identical with the second 
paragraph of the ‘‘Second Preliminary Report” of the same 
Committee,” viz., ‘‘ Your Committee recommend to anatom- 
ists that, other things being equal, terms consisting of a single 
word each be employed rather than terms consisting of two or 
more words.” Proceedings for 1895, p. 4. 

$83. Section 4 is also substantially represented in the 
‘‘Third Preliminary Report of the Committee on Anatomical 
Nomenclature with special reference to the Brain ” * which was 
adopted unanimously by the American Association for the Ad- 
vancement of Science, Sept. 2, ,1889:— ‘‘ They agree upon 
one point, v7z., the advantages, other things being equal, of 
mononyms (single-word terms) over polyonyms (terms consist- 
ing of two or more words).”” The report was published in 
the Proceedings for 1889, p. 26. 

Sections I, 2, 3, 5, occur verbatim in the Fourth Report of 
the same committee which was adopted unanimously by the 
Association Aug. 25, 1890 and printed in the Proceedings, p. 20. 


1The members of the committee at that time were Joseph Leidy, M.D., 
L.L.D., professor of Anatomy in the University of Pennsylvania, president ; 
Harrison Allen, M.D., formerly professor of Physiology in the U. of P.; Frank 
Baker, M.D., professor of Anatomy in the Medical Department of George. 
town University ; Thomas B. Stowell, Ph.D., principal of the Potsdam (N. Y.) 
Normal School; and B. G. Wilder, Secretary. Tothe committee, at the meet. 
ing, was added Thomas Dwight, M. D., professor of Anatomy in the Harvard 
Medical School. The report was published in the History and Records of the 
Association for 1838, 1889, 1890, p. 5. 


2 Upon the death of Dr. Leidy, Dr. Allen succeeded to the chairmanship 
of the Committee. The place of Dr. Stowell, resigned on account of pressing 
administrative duties, was filled by the appointment of F. H. Gerrish, M.D., 
professor of Anatomy in the Medical School of Maine. 


3 The Committee comprised, besides H. Allen, F. Baker, T. B. Stowell, 
and B. G. Wilder, chairman (see 381, note), Henry F. Osborn, Sc.D., professor 
of Biology in Columbia University. 


240 JOURNAL OF CoMPARATIVE NEUROLOGY. 


§84. The first five sections of the report of the Neuro- 
logical Committee are embodied verbatim in the ‘‘ Preliminary 
Contribution of the American Branch of the International 
Committee on Biological Nomenclature of the American As- 
sociation for the Advancement of Science’’* which was adopted 
unanimously by that body Aug. 23, 1892, and published in its 
Proceedings, p. 231.” 

$85. The report just mentioned is so clear, comprehen- 
sive, and concise that its main features are here summarized : 

a. ‘Terms relating to position and direction [toponyms, 

9] should be intrinsic rather than extrinsic ; that is, should re- 
fer to the organism itself rather than to the external world.” 


6. ‘So far as possible terms should be single, designa- 
tory words [mononyms, $15] rather than descriptive phrases.” 


c. Terms derived from the names of persons [eponyms, 
§33] should be avoided. 


dé. ‘*Each term should have a Latin [international, $46] 
form.”’ 

e. ‘Each term should have also a [national, §43] form in 
accordance with the genius of each modern language, e. g., 
a paronym [$44] of the original Latin form.” 

jf. The report gives due recognition of the labors of other 
committees and of individuals. 

$86. Returning to the report adopted by the American 
Neurological Association [$8o0| its recommendations may be in- 
dicated conveniently in the following Table I: ; 


1 The members are George L. Goodale, Ph.D., professor of Natural His- 
tory in Harvard University, chairman ; John M. Coulter, L.L.D., president of 
the State University of Indiana; Theodore Gill, Ph.D., Smithsonian Institution; 
Charles Sedgwick Minot, Ph.D., professor of Embryology in Harvard Univer- 
sity; Simon H. Gage, B.S., professor of Histology and Embryology in Cornell 
University, secretary. 


2 Reprints were distributed to biologists of all nationalities and may be ob- 
ained from the secretary. 


Wiper, Weural Terms. 241 


TABLE I. 


Forty Latin names recommended unanimously by the American Neurolog- 
ical Association, June 5, 1896. 


Mononyms  (stngle-word terms) Dionyms 
CALCAR HIPPOCAMPUS PIA CORNU DORSALE 
CALLOSUM HypopHysis PoNs CORNU VENTRALE 
CHIASMA INFUNDIBULUM PRAECUNEUS |FISSURA CALCARINA 
CLAUSTRUM |INSULA ; PULVINAR FISSURA CENTRALIS 
CLAVA LEMNISCUS STRIATUM FISSURA COLLATERALIS 
CUNEUS MESENCEPHALON TEGMENTUM |FISSURA HIPPOCAMPI 
DENTATUM |MONTICULUS TENTORIUM |RADIX DORSALIS 
DURA OBILONGATA THALAMUS’ |RADIX VENTRALIS 
EPIPHyYsIS OLIVA VERMIS VERTEBRA THORACALIS 
FALX OPERCULUM 
FORNIX PALLIUM 


$87. Cerebellum might well have been added to the list 
for there is no disagreement as to the application of the word. 
Cerebrum, unfortunately, is used in at least two senses, (a) as 
equivalent to the prosencephal, and (4) as including also two or 
three adjoining segments. 

$88. Of the forty terms in the above list thirty-one are 
mononyms (S15). The other nine, nearly one-fourth, are dio- 
nyms ($13). This fact is to be noted in connection with cer- 
tain animadversions of the German committee (Part V). 

$89. In Table II are given in parallel columns (1) the 
Latin (international) names adopted by the American Neuro- 
logical Association ; (2) their English paronyms; (3) the regu- 
lar Latin adjectives; (4) the English forms or paronyms of 
the Latin adjectives.’ 


1 Had space permitted I should have been glad to amplify this table after 
the pattern of that upon pp 530 and 531 of W.&G., ’89. Forty-five terms 
were there given substantially as in the following example: 


LATIN 
(Common polyonym in upper line.) 
Nom. singular | Genitive | Nom. plural | Adjective 
Thalamus nervt optict 


Thalamus | Zhalamt | Thalami | Thalamicus 
ENGLISH 
(Heteronym in upper line.) 
FRENCH 
Singular Plural Adjective GERMAN 
ITALIAN 
Optic bed 
Thalame 
THALAMUS | THALAMI | THALAMIC Thalamus 
Talamo 


242 


TABLE If. 


JOURNAL OF COMPARATIVE NEUROLOGY. 


Derivatives of the Terms adopted by the American Neurological Association. 


Sudbstantives Adjectives 
Latin English Latin English 
1|Calcar Calcar Calcarinus Calcarine 
2)|Callosum Callosum Callosalis | Callosal 
3)Chiasma Chiasma or chiasm Chiasmaticus Chiasmatic 
4|Claustrum Claustrum Claustralis Claustral 
5|Clava Clava Clavalis Claval 
6!Cornu dorsale —_|Dorsal cornu 
7|Cornu ventrale |Ventral cornu 
8}Cuneus Cuneus Cunealis Cuneal 
g|Dentatum Dentatum Dentatalis Dentatal 
10|Dura Dura Duralis Dural 
11|Epiphysis Epiphysis Epiphysialis Epiphysial 
12|Falx alx Falcialis Falcial 
13|F. calcarina Calcarine fissure 
14|F. centralis Central fissure 
15|F. collateralis |Collateral fissure 
16/F. hippocampi |Hippocampal fissure 
17|Fornix Fornix Fornicalis Fornical 
18|Hippocampus !Hippocamp or hippo-|Hippocampi (gen.)|/Hippocampal 
campus or hippocampalis 
19|Hypophysis Hypophysis Hypophysialis Hypophysial 
20|Infundibulum = _|Infundibulum Infundibularis Infundibular 
21\Insula Insula Insularis Insular 
22|Lemniscus Lemniscus Lemniscalis Lemniscal 
23|Mesencephalon |Mesencephal or mesen-|Mesencephalicvs |Mesencephalic 
cephalon 
24|Monticulus Monticulus Monticularis Monticular 
25|Oblongata Oblongata Oblongatalis Oblongatal 
26|Oliva Oliva or olive Olivaris Olivary 
27\|Operculum Operculum or opercle |Opercularis Opercular 
28} Pallium Pallium Pallialis Pallial 
29|Pia Pia Pialis Pial 
30| Pons Pons Pontilis Pontile 
31\Praecuneus Precuneus Praecunealis Precuneal 
32|Pulvinar Pulvinar Pulvinaris Pulvinar 
33) Striatum Striatum Striatalis Striatal 
34|Tegmentum Tegmentum or tegment|Tegmentalis Tegmental ~ 
35\Tentorium Tentorium Tentorialis Tentorial 
36|Thalamus Thalamus Thalamicus Thalamic 
37|Radix dorsalis |Dorsal root 
38|Radix ventralis |Ventral root 
39| Vermis Vermis Vermianus Vermian 
40|Vertebra_ thora-/Thoracic vertebra 


calis 


WILpvER, Neural Terms. 243 


§go. It should be borne in mind that only the Latin 
names in the first column have the sanction of the various As- 
sociations that have adopted them. The derivatives and the 
comments thereon do not constitute parts of the reports. In- 
deed, as will be seen, there is room for considerable latitude of 
opinion and usage; my own views may be imperfect and even 
inconsistent, but I think the analogies adduced are sound. 

S91. English Plurals.—The parts of the brain are so sel- 
dom named in the plural that a separate column is not given there- 
for. Analogy with crises, strata, fungi, algae and phonomena would 
justify the employment of the regular Latin plural in certain 
cases, e. g., thalami, epiphyses, hippocampi, cornua, striata and 
vertebrae. On the other hand, aveas, vistas, hernias, emporiums, 
lenses, geniuses, pianos, indexes, pericarps, angles, atlases, dt- 
plomas, and similes are precedents for calcars, chiasmas (or 
chiasms), falxes, lippocamps, insulas, mesencephals, ponses, ver- 
mises. Bonuses would even justify thalamuses but the length of 
the latter is objectionable. 

$92. Close resemblance of the Angloparonyms to the Latin 
originals.—This is so obvious as to hardly require mention. 
With more than half, the two forms are identical in spelling, fe) 
that the Latinity of the originals can only be indicated to the 
eye by italics, and to the ear by the pronunciation now com- 
monly adopted for Latin words.' 

§93.  Hippocamp.—For this, as the Angloparonym of 
hippocampus, there are many precedents, notably the follow- 
ing:—antepenult, digit, tmpediment, diagram, telegram (which was 
strenuously objected to when first introduced), epzcarp and perr- 
carp ; see my note as to peszcard, ’95, a. 

S94. Infundibulum.—lf the part so designated were fre- 
quently mentioned it is probable that either a shorter word 
would be found,or the present name be paronymized as z7fundt- 


1 The Angloparonyms of Latin words, even when orthographically unmod- 
ified, are Eaglish by adoption and are to be so pronounced; to pronounce ¢c/aus- 
trum, clowstroom in an English sentence would beas affectedas to say mamorarn- 
doom. As an English word od/ongata has the first a as in mate. 


244 JOURNAL OF COMPARATIVE NEUROLOGY. 


bule, after the analogy of vetucule, dwerticle, etc. The same may 
be said of smonticulus and monticule; see $167. 

$95.  Mesencephalon.—By itself and used occasionally the 
. Latin form is certainly euphonious and unobjectionable ; but in 
any discussion of the segmental constitution of the brain, 
whether written or spoken, the frequent recurrence of the ob- 
trusively Latin termination is pedantic and burdensome. Its 
omission is warranted by words like angel.' 

S96. Operculum and Opercle.—The Latin tetrasyllable is 
not commonly oppressive, but the compounds preoperculum etc., 
might well become so. The case is comparable with that of 
ultima ; with it, and even with penxultima, the last two syllables 
are endured; but when two more syllables are added at one end, 
then two are dropped from the other, leaving antepenult of only 
moderate length.. Preopercle, subopercle and postopercle are 
already applied to analogous parts of the fish’s head; see 
$$67-70. 

$97. Praecuneus.—Here the difference between the Latin 
antecedent and the Angloparonym consists in the replacement 
of the ae by e¢ as in preposition, pretext, preface, etc. 

S98.  Zentorium.—By analogy with ovary, aviary, granary, 
laboratory etc., the Angloparonym would be éentory, and this 
word has been used to designate the awning of atent. But 
tentorium is unobjectionable and likely to be retained as an un- 
changed paronym. 

S99. ontilis.—Unwarrantable forms of the English ad- 
jective from pons occur so frequently that there is here repro- 
duced a paragraph from my recent note on the subject, ’96, a. 
‘‘In the subtitle of the letter above mentioned the case is re- 
ferred to as one of ‘pontine hemorrhage.’ This form of the 
adjective is not uncommon in medical literature, and pontic and 
pontal have found their way into the dictionaries.. Now, as may 
be seen from any Latin lexicon, fovtal has no justification what- 
ever. /onticus, the Latin antecedent of foviic, is derived from 
pontus, the sea. Pontinus, the antecedent of pontine, was orig- 


' Respecting the possibility of further abridgement of these terms see 274. 


WILDER, Neural Terms. 245 


inally Pomptinus, and refers to a district of Italy. As already 
pointed out by me (article Anatomical Terminology, Buck's 
Reference Handbook of the Medical Sciences, VIII, 524, $50), 
the only legitimate Latin adjective from fozs is pontilts, and its 
Angloparonym is pontile. The use of any other form tends 
to cause confusion and to bring discredit upon medical 
scholarship.” 


Part IV. Comparison of the Terms Adopted by the American 
Neurological Association with those Adopted by the Anat- 
omische Gesellschaft. 


§roo. Inthe accompanying Table III are given in par- 
allel columns (1) the forty terms adopted by the A. N. A.; (2) 
the corresponding terms adopted by the Anatomische Gesell- 
schaft ; (3) some of the Latin synonyms. 

§1o1. Probably few will question the inferiority of the 
discarded synonyms in the third column; hence are here con- 
sidered mainly the relative merits of the two other sets. 

$102. The extent of agreement is impressive and en- 
couraging. With the following twenty-four terms there is abso- 
lute consensus between the American and the German commit- 
tees: Claustrum; Clava; Cuneus; Fissura calcarina; F. collat- 
eralis; F. hippocampi; Fornix; Hippocampus; Hypophysis ; 
Infundibulum ; Insula; Lemniscus ; Mesencephalon ; Monticu- 
lus; Oliva; Operculum ;' Pallium; Pons; Praecuneus; Pulvi- 
nar; Tegmentum; Thalamus; Vermis; Vertebra thoracalis. 

$103. With the following ten terms the differences lie 
merely in the retention by the Germans of certain words which 
the Americans regard as superfluous. In the following list these 
words are italicized: Calcar avs ($105); Corpus callosum ($111); 


1 The case of this term is peculiar. The German committee particularize 
three parts, frontal, partetal and temporal of a general operculum. The Neuro- 
logical Association regards the parietal portion as ¢#e operculum, the frontal and 
temporal being so specified. (By the present writer these are designated as 
pracoperculum and postoperculum, and the orbztal portion as suboperculum.) It will 
be seen therefore that while the word ofercu/um is identical with both commit- 
tees, its significance is general with the German and special with the American. 


246 JOURNAL OF COMPARATIVE NEUROLOGY. 


TABLE III. 


Comparison of the Terms adopted by the American Neurological Associa- 
tion (first column) with those of the German Committee (second column) and 
with certain Synonyms (third column). 


I|}\CALCAR 


2|/CALLOSUM 
CHIASMA 


3 
4|CLAUSTRUM 
5|CLAVA 

6|CORNU DORSALE 
7|\CORNU VENTRALE 
8|CUNEUS 


9| DENTATUM 
Io/DURA 
11) EPIPHYSIS 


12|FALXx 
13|FISSURA CALCARINA 
14| FISSURA CENTRALIS 


15 FISSURA COLLATERALIS 


16|FISSURA HIPPOCAMPI 


17| FORNIX 
18} HIPPOCAMPUS 


19] HYPOPHYSIS 


20| INFUNDIBULUM 
21|/INSULA 


22| LEMNISCUS 

23] MESENCEPHALON 
24) MONTICULUS 
25|OBLONGATA 
26/OLIVA 
27|OPERCULUM 


28) PALLIUM 
29|PIA 
30| Pons 


31] PRAECUNEUS 
32| PULVINAR 
33|STRIATUM 
34|TEGMENTUM 
35|TENTORIUM 
36/THALAMUS 


37) RADIX DORSALIS 
38] RADIX VENTRALIS 
39| VERMIS 


Calcar avis 


Corpus callosum 
Chiasma opticum 


Claustrum 

Clava 

Columna posterior 
Columna anterior 
Cuneus 


Nucleus dentatus 
Dura mater 
Corpus pineale 


Falx cerebri 
Fissura calcarina 
Sulcus centralis 


Fissura collateralis 


Fissura hippocampi 
Fornix 
Hippocampus 


Hypophysis 


Infundibulum 
Insula 


Lemniscus 

Mesencephalon 

Monticulus 

Medulla oblongata 

Oliva 

Operculum (pars par- 
ietalis) 

Pallium 

Pia mater 

Pons [Varolii] 


Praecuneus 
Pulvinar 

Corpus striatum 
Tegmentum 
Tentorium cerebelli 
Thalamus 


Radix posterior 
Radix anterior 
Vermis 


40' VERTEBRA THORACALIS! Vertebra thoracalis 


Hippocampus minor; Eminentia 
digitalis; Unguis 

Corpus callosum; Trabs cerebri 

Chiasma nervorum opticorum; 
Commissura optica 

Claustrum 

Processus clavatus 

Cornu posterius 

Cornu anterius 

Lobulus occipitalis internus; Lob- 
ulus cuneatus 

Corpus dentatum cerebelli 

Dura mater 

Conarium; Glandula pinealis ; 
Epiphysis cerebri 

Falx major; Processus falciformis 

Fissura occipitalis horizontalis 


Fissura Rolandica; S. postero- 
parietalis 
Sulcus occipito-temporalis; 5S. 


temporo-occipitalis 

Fissura hippocampi 

Fornix tricuspidalis 

Hippocampus major; Cornu Am- 
monis 

Glandula pituitaria ; Hypophysis 
cerebri 

Pedunculus hypophyseos 

Insula Reilli; Lobus centralis ; 
Median lobe 

Laqueus 

Mesencephalon 

Monticulus 

Bulbus rhachidicus 

Corpus olivare ; Nucleus olivaris 

Operculum parietale 


Mantellum 
Pia mater 
Pons Varolii ; 
nularis 
Lobus quadrilaterus 
Tuberculum [thalami] posterius 
Ganglion cerebri anterius 
Tegmentum caudicis 
Tentorium 
Thalamus opticus; T. nervi optici; 
Ganglion cerebri posterius 
Radix posterior 
Radix anterior 
Vermis bombycinus; 
vermiformis 
Vertebra dorsalis 


Protuberantia an- 


Processus 


Witper, Neural Terms. 247 


Chiasma opticum (§109) ; Wucleus dentatus (§123); Dura mater 
($116); Falx cerebri ($124); Medulla oblongata (§116); Pia 
mater ($117); Corpus striatum (§115); Tentorium cerebelli 
($125). 

$104. With the remaining six terms the differences are 
more or less radical (S$119-122, 127-133). 

§105. CALCAR versus calcar avis. ‘Thirty years ago, 
in connection with the controversy as to the cerebral , peculiari- 
ties of man, the term /zppocampus minor became familiar even 
to general readers. Nevertheless, probably influenced in some 
degree by Huxley’s proposition to replace Owen’s posthippo- 
campal and Henle’s occipitals horizontalis by calcarina,' anatom- 
ists have been more and more generally employing calcar avts, 
and this is adopted by the German committee in preference also 
to uuguis and eminentia digitalis. The advantages of correlated 
names ($32) for collocated parts are many and great, as illus- 
trated by Aippocampus [major] and fissura hippocampi; by emt- 
nentia collaterals and fissura collaterals. In the present case 
these advantages would have been gained equally had Huxley 
adopted Owen’s fosthippocampal for the fissure and proposed 
posthippocampus for the ental ridge corresponding thereto. In- 
deed this would have been in accordance with the general prin- 
ciple of locative names (§29) and learners would have been 
spared thereby some effort of memory. In this, however, as 
in so many other instances it is now idle to speculate upon the 
consequences of harmonious cooperation between the two lead- 
ers of English anatomy at that period. Assuming that calcar 
avis has general and decided preference over the other names 
enumerated, there need be stated here only the grounds upon 
which calcar has been unanimously adopted by four American 
Committees and by the three associations which they represent. 

§106. Briefly, the adoption of calcar is a logical corollary 
of the recommendation which is common to the reports of all 


1Pye-Smith wrote as follows nearly twenty years ago (’77): ‘‘Of all the syn- 
onyms of hippocampus minor, calcar avis is the most distinctive and brings it at 
once into relation with the calcarine fissure.” 


248 JOURNAL OF COMPARATIVE NEUKOLOGY. 


four American committees, viz., ‘‘Other things being equal, 
it is recommended that mononyms be preferred to polyonyms.”’ 
Calcar avis is a polyonym ; ca/car is a mononym. 

If it be said that wxguzs is also a mononym, the answer is 
that in this case ‘‘other things”’ would not be equal, because 
(1) no general preference has ever been shown for it or for any 
term of which it is a constituent; (2) there would be lost the 
advantage of the correlation now existing between the ental 
ridge and the fissure collocated therewith. 

§107. Two objections might be offered to the omission of 
the qualifying genitive, avzs. 

1. The original sense of the Latin calcar was spur, and 
its application to the sharp projection on the leg of the cock 
was metaphoric. This can hardly be entertained as a serious 
objection; indeed, although the modern spur has a toothed 
wheel or rowel, the primitive instrument was little more than a 
spike; hence the qualifying genitive is needless. 

2. Calcar has also been applied occasionally to two other 
parts, viz., the calcaneum (os calcis) and the styloid process of 
the temporal bone. But, (a) neither of these uses is sanctioned 
by the German committee, and (0) even if they were, the con- 
text would infallibly avert misapprehension ($§23, 67); indeed, 
the German committee apply c/vus without qualification to fea- 
tures of two adjacent cranial bones, the occipital and sphe- 
noidal. 

$108. Finally, the sufficiency of the mononymic substan- 
tive, calcar, is practically conceded by all who employ the mon- 
onymic adjective, calcariuus, in any of its Latin inflections, or 
in any of its national paronymic forms. The simplest require- 
ments of logic present the following dilemma: If calcarinus is 
sufficiently distinctive, so is calcar from which it is derived. 
But if calcar avis is essential, then the adjective should be cal 
cart-avianus or some such compound. See also under dura 
(115). There seems to have been little if any hesitation on the 
part of the German committee in adopting /fissuva calcarina 
(His, ’95, 170) and no reason for the maintenance of calcar avis 
has yet come under my notice. 


WiLverR, Neural Terms. 249 


S1o9g. CHIASMA vs. Chiasma opticum.—Meynert’s chi- 
asma nervi acustict is not retained by the German committee, 
and even if it were there is no likelihood of confusion with it 
or with Camper’s chiasma tendinum. The chiasma is and always 
will be that of the optic nerves. The use of any qual- 
ifier suggests undesirable variations like chiasma nervorum optic- 
orum and commussura optica. Furthermore, the sufficiency of 
the unincumbered mononym is practically conceded by the Ger- 
man committee in designating one of the subarachnoid spaces 
as cisterna cliasmatis; see also His, ’95, 171, line 8’. 

S110. ZTHALAMUS.—This term may naturally be men- 
tioned here. In the German list the adjective ofticus is omitted, 
and His makes the following remark ('95, 7, lines 1-3): ‘‘Wir 
stimmen unsererseits vollig bei, wenn das Wort Thalamus kurz- 
weg an die Stelle von Thalamus opticus gesetzt wird.’”’ But it is 
worthy of note that ¢a/amus is strictly'an idionym (§24), and 
that the only valid excuse for the addition of the adjective is a 
desire to aid the student’s memory by the association with the 
optic nerve. Asa matter of fact, no case of real advantage is 
known to me, and the frequent repetition of the adjective may 
easily become a burden, as pointed out by me in 1888 ('88, 8). 

S111. CALLOSUM vs. corpus callosum.—Corpus callosum 
is the most familiar type of a large group of anatomic names. In 
1889, including unusual synonyms, I recorded one hundred 
neural polyonyms of which corvpus constituted the initial word. 
Ten such remain upon the German list, (viz., corpus restiforme ; 
cp. trapezoideum ; cp. medullare ; cp. quadrigeminum ; cp. mamil- 
lare; cp. gentculatum mediale ; cp. gnc. laterale; cp. pineale ; cp. 
callosum ; cp. striatum), and their genitives are correspondingly 
in evidence. 

§112. It must be admitted that corpus callosum is rather 
attractively sonorous. It is easily pronounced and even, like 
quadrupedante, ‘‘runs trippingly from the tongue.’ But that is 


} The word chzasma is discussed at some length by Hyrtl, ’80, 105-106. 


? A similar concession has been made (Sczence, June, 22, 1888, editorial) to 
the claims of proper names like Johnny McWhorter which are euphonious and 
easily remembered. 


250 JOURNAL OF COMPARATIVE NEUROLOGY. 


no reason for the retention of a word which is not merely need- 
less, but really burdensome by reason of the frequency with 
which certain parts are mentioned. In one short paper (Srazn, 
’ October, 1885, 377-379) corpus callosum occurs twenty times, 
an average of once in five lines; corpus occupies 2.5 lines, one- 
fortieth of the entire paper. 

§113. The elimination of corpus from all neural names 
constituted one of the fundamental propositions of my first 
communication upon the general subject (80, 7), and since that 
time it has been consistently practised and persistently preached. 

§114. By the use of the genitive case, corporis callost, 
the German committee have designated the various divisions of 
the callosum, (splenium, genu, truncus and rostrum) ; also the 
sulcus along its dorsal margin. They have thus avoided the 
use of the secondary adjective cal/losals. But in expressly re- 
jecting pedunculus corporis callost in favor of gyrus subcallosus 
(His, ’95, 170-172) they practically concede the superfluity of 
the corpus. 

§115. Unless we are prepared to abandon all adjective- 
substantives (§118) there seems to be no reason for the further 
retention of corpus in any of the terms enumerated in §111. 
Corpus fornicis of the German list is not open to the objection 
that naturally arises against corpus corporis callost but truncus 
corpus callosi is a good precedent for truncus fornicis if the 
distinction be necessary. 

§116. DURA vs. dura mater.—This constitutes a type 
and test case for a considerable group of anatomic terms from 
which, for fifteen years, I have dropped the (here italicized) 
nouns, viz., pia mater; substantia alba; substantia cinerea; 
membrana (or tunica) serosa ; mb. (or tz.) mucosa; mb. (or tn.) 
submucosa; md. (or tz.) arachnoidea; medulla oblongata. From 
the group of ‘‘corpus”’ polyonyms they differ in that the elim- 
ination of the substantive leaves a feminine instead of a neuter 
adjective to be used substantively, and as a base for the forma- 
tion of secondary adjectives, dural, mucosal, cinereal, arach- 
noidal, ete. 

§117. Curiously enough the first precedent for this known 


WILpER, Veural Terms. 251 


to me dates back a hundred and fifty years. Inthe ‘‘ Medical 
Dictionary” of James (1743), in the article ‘‘cerebrum’”’, occurs 
the following sentence: ‘‘The superficial vessels of the cerebrum 
are lodged’ between the two laminae of the pia.” 

§118. The employment of the mononymic feminine ad- 
jectives as substantives, and of the secondary adjectives de- 
rived therefrom, has now become so general! that the matter 
would hardly need discussion but for the reactionary attitude of 
the German committee. Yet this attitude is really not main- 
tained consistently. Cornea is a feminine adjective. So is sclera. 
In arachnoidea encephah the feminine adjective is used as a noun. 
Muscularis mucosae and tela submucosa are warrants for mucosa, 
etc. Finally, although the useless noun is retained in dura 
mater spinalis and filum durae matris spinalis, the very next 
terms in their list, cavum epidurale and cavum subdurale, are indi- 
rect and probably unintended, yet none the less complete, pre- 
cedents for duva pure and simple, and for the substantive em- 
ployment of any and all feminine adjectives whatsoever. 

§1i9. EPIPHYSIS vs. corpus pineale.—His regards epiph- 
ysisas a ‘‘generelles Wort” (’95, 163) and theancient dionym 
is adopted by the Germancommittee.?- My own earlier preference 
was for conarzum, as stated in §68. I now realize the desirabil- 
ity of the verbal as well as the topographic correlation with 
hypophysis and paraphysis, and the inutility of maintaining in all 
cases the rigid doctrine of 1871 (§67). 

§120. FISSURA CENTRALIS vs. sulcus centrals (or 
fissura or sulcus Rolando).—By comparison of the three columns 
it will be seen that two distinct points are concerned, involving 
respectively the generic and the specific names of this feature of 
the lateral aspect of the cerebrum. If eponyms or personal 
names are to be abandoned, as decided by the German commit- 


1 In Foster’s Medical Dictionary, dura and fia, dural and pial, are major 
headings, dura mater and pia mater being merely synonyms. 


2 In the earlier publication (Scéence, July 17, 1896, p. 71) of the report of 
the Neurological Committee, the date, 7895, after epiphysis would indicate its 
adoption by the Germans. That was an error for which I must be held respon-. 
sible, and which was corrected as soon as possible after it was noted; ’96, 4. 


252 JOURNAL OF COMPARATIVE NEUROLOGY. 


tee and as advocated by me, since 1880,’ then all the derivatives 
of Rolando must be discarded in favor of cextralis and its deriva- 
tives. Those who prefer the eponym should show that Rolando’s 
- figure and description really merit such commemoration, and 
should be also at least‘consistent in the employment of deriva- 
tives. Paracentralis, praccentralis and postcentralis have no other 
justification than topographic reference to cextralis; yet itis by 
no means uncommon to find in oneand the same paper ‘‘fissure 
of Rolando” and ‘‘paracentral lobule.”’ 

§121. As tothe generic terms, fissuva and sulcus, the 
former has been consistently employed by me since 1880 for all 
linear depressions of the} cerebral surface, while the German 
committee restrict it to the sylvian (called by them cevebri lat- 
evalis), the collateral, the occipital (their parieto-occipitalis) the 
calcarine and the hippocampal, and name all the others sadcz. 
They regard the striatum as constituting an ental correlative of 
the sylvian (p. 170); hence it may be inferred that fisswra indi- 
cates a corrugation of the entire parietes, while sz/cws indicates a 
linear furrow not represented in the cavity by a corresponding 
elevation.” Fully conceding the desirability of recognizing the 
distinction between the two groups of cerebral furrows, the fol- 
lowing considerations lead me to question the advisability of 
employing the two generic words in the senses proposed by the 
German committee. 

(1). /2ssura and its various paronyms and heteronyms are 
already well established and commonly associated with cerebral 
topography. This subject, on account of its various relations, 
physiologic, pathologic, surgical and psychologic, has already 
gained much general interest. Sw/cus is a comparatively unfa- 
miliar word. It is distinctively Latin and technical. . Its Latin 
plural, sz/cz, is even more so. It does not readily lend itself to 


‘ With the exception of fissura Sy/vii and certain derivatives of sy/véana, 


*The two groups are sometimes distinguished as ¢ofal and partial, or as complete 
and zxcomplete. The former seem to be preferable, since with the total the to- 
tality of the parietes is involved, whereas complete and incomplete seem to im- 
ply differing degrees of relative perfection. 


WILvER, WVeural Terms. 253 


paronymization, sw/c and szulcuses being both somewhat un- 
acceptable. 

(2). Sulcus has recently been employed by Mrs. Gage 
(93), O. D. Humphrey (’94), P. A. Fish ('94), and B. F. Kings- 
bury (95), for ental (entocelian or intraventricular) depressions 
which are less likely than the cerebral furrows to become sub- 
jects of general interest. 

$122, (3). There isa practical difficulty that cannot be 
ignored. Nothing in the words fssura and sulcus, or in their 
ordinary associations, serves to admonish us as to the proposed 
distinction. Hence there is liability to misuse and confusion. 
Many actual instances of this might be cited but the following 
may suffice. Edinger (’95) apparently intends to apply fissura 
to the total fissures, and the occipital is so designated in the in- 
dex; but on Fig. 33 it is called sadcus. Kolliker (‘‘Entwickel- 
ungsgeschichte,”” p. 555) attributes salcus calcarinus to Huxley, 
who uses fissure as does Kolliker in the explanation of a figure. 
Flower (‘‘Proteles,”” Zool. Soc. Proc., 1869) applies to the supra- 
orbital, fissura and sulcus indifferently. Huxley (‘‘ Vertebrated 
Animals’) says that the cerebral surface becomes complicated 
by ridges and furrows ‘‘the gyri and sulci;” but the first of 
the ‘‘sulci’’ to be mentioned is the ‘‘ sylvian fissure”’ and the 
second ‘‘the fissure of Rolando,” the latter also being desig- 
nated on Fig. 21 asthe ‘‘sulcus of Rolando.” Flower and 
Lydekker (Mammals, p. 71) say ‘‘the sylvian fissure is one of 
the most constant of the sulci. In the last two cases the gen- 
eric designation of the shallower furrows is made to include 
both kinds, and curiously enough this usage is apparently sanc- 
tioned by the German committee in introducing gyvz cerebri and 
sulct cerebrt as comprehensive names and then specifying certain 
Sulcisand fissurae -seeulable VIL. IL -10;)07;/(26: 

§123. DENTATUM vs. Nucleus dentatus.—Two separate 
questions are involved in the choice between these terms: (a) 
The use of xucleus (with a masculine adjective) in place of 
corpus (with a neuter) ; (4) The employment of an adjective of 
either gender as a substantive. The latter is considered in con- 
nection with collosum and dura (§S11t-118). The substitution of 


254 JOURNAL OF COMPARATIVE NEUROLOGY. 


nucleus for corpus seems to the American committee to consti- 
tute a step backward, as tending to obscure the commonly ac- 
cepted distinction between the part in question, with the analo- 
gous part in the oliva on the one hand, and the ‘‘nuclei’”? of ori- 
gin of the various nerves on the other. 

$124. FALX vs. Falx cerelxi.—The German committee 
designate the slighter fold of dura between the two lateral masses 
of the cerebellum as falr cerebelli. The present writer prefers 
the diminutive, falcula. The American committee has not yet 
passed upon this case. Even should they retain falr cerebelli it 
would not prove a serious burden, because the part is hardly 
mentioned once while the cerebral septum is named ten times. 

$125. ZENTORIUM vs. tentorium cerebell.—This case is 
even stronger than that of falx, for tentorium is an idionym (§24). 

§126. SZTRIATUM vs. Corpus striatum.—See callosum, 
(S111). 

$127. CORNU DORSALEvs. columna ( grisea) postertor. 
—Two distinct issues are involved here: (a) toponymic, be- 
tween foster zor and dorsalis; (6) organonymic, between colwmna 
and cornu. The former will be considered in connection with 
cornu ventrale ($131) and radix dorsalis (§132). 

§128. CORNU vs. columna.—lIt is almost embarassing to 
find myself advocating the maintenance of ancient and general 
usage against one comparatively novel. Probably most ana- 
tomic teachers will sympathize with the German committee in 
their objection to the application of cornu to what is really one 
of several ridges of a deeply fluted column of gray nervous tis- 
sue constituting the core of the ‘‘ spinal cord;’ ridges that re- 
semble ‘‘horns”’ only when artificially exposed upon transec- 
tion. At least ten years ago I was so deeply impressed by this 
inappropriateness of cornu as to hunt up an architectural term, 
namely avrzs, signifying the ridge between two adjoining chan- 
nels of a Doric column. Whether or not it was derived from 
arista, it is excellent Latin in form, and acceptable in every re- 
spect save its novelty. 


1 The question of preference between mucleus, and nidus (Spitzka) and nidu- 
/us (C. L. Herrick), need not be considered upon the present occasion. 


Wiper, Neural Terms. 255 


§129. Yet I believe that I did well to refrain from its in- 
troduction ; for, after all , in nine cases out of ten, the artificial 
appearance presented upon section is what is first offered the 
student, and I have never known a case of misapprehension 
occasioned thereby. Upon the whole, this has seemed to the 
American committee a good case for the observance of Huxley’s 
aphorism (80, 16) as to the unadvisability of interfering with 
terms that are well established and have a definite connotation, 
even when they may be etymologically inadequate, e. g., cadlo- 
sum. Individually, I should feel that the case against cornu 
would be much stronger were it a word of half a dozen sylla- 
bles, or lacking in euphony. 

§130. The assignment of co/umna to the ridges of the myelic 
cinerea naturally involved the replacement of that word, as com- 
monly applied to the intervening masses of alba, by some other 
word ; the German committee selected fumzculus. If cornu be 
retained, columna will be available as hitherto. Even if a change 
be made, however, why not /uzzs instead of the longer diminu- 
tive, upon the grounds stated in $21? There could hardly be 
confusion with the same word as applied to the ‘‘ umbilical 
cord.” 

§131. CORNU VENTRALE.—As an objection to this 
term it might be urged that consistency would involve the ap- 
plication of the same words to the ‘‘ middle” or ‘“ descending” 
extension of the ‘‘lateral ventricle,’ which the German com- 
mittee call cornu infertus. What the American committee may 
do in this connection remains to be seen. There would be no 
real cause for ambiguity, however, since cornu temporale, c. 
Jrontale, and c. occipitale are perfect examples of a class of 
terms that suggest parts or regions already familiar. Personally 
I have never had any difficulty, the locative, mononymic, 
idionyms (§$25, 27) medicornu, praecornu and postcornu, having 
been consistently employed by me for fifteen years (’81, J. d). 

§132. RADIX DORSALIS vs. radix posterior.—Since, 
with this and with vadix ventralis (or anterior) the Americans 
and the Germans are at one as to the substantive element, there 
only recurs the toponymic difference already alluded to in con- 


256 JoURNAL OF COMPARATIVE NEUROLOGY. 


nection with the ridges of the myelic cinerea (§127). The differ- 
ence is far reaching and literally radical. As with the myelic 
sulci, columns, cornua and commissures, the folds of the axilla, 
the aspects of the thigh, the tubercles of the cervical vertebrae, 
the sides of the stomach and other viscera, the valves of the 
heart, there is exemplified one of the most undesirable features 
of the pernicious influence of anthropotomy upon anatomy at 
large.* 

§133. Upon this subject the position of the German com- 
mittee in 1895 is indicated by the following translation of pass- 
ages from His, (’95, 109-110): ‘‘ As mentioned above, Herr 
von Kolliker has proposed replacing generally the words anterior 
and posterior by ventralis and dorsalis where the relations to 
comparative anatomy, and especially to the anatomy of domes- 
ticated animals, render it desirable; that is, where the terms 
anterior and postertor apply only to the upright attitude of man. 
* * * We do not deny the merit of such strict usage, but 
the commission has not been able to decide upon its adoption. 
It involves all kinds of difficulties and inconveniences. * * * 
We leave time to determine whether or not we shall depart from 
the traditional usage associated with the erect attitude of man.” 

Had most of the members of the commission been investi- 
gators and teachers of zootomy rather than of anthropotomy, 
there would probably have been no hesitation in adopting terms 
that apply equally well to all vertebrates in any attitude. Let 
us hope that the distinguished President of the Commission 
may live to see his recommendations unanimously adopted.- 

$134. I close this discussion of the differences between 
the recommendations of the American and German committees 
with the remark that, strictly speaking, not one of the words 
in the first column of Table III can be imputed to us. All 
were in use for longer or shorter periods prior to 1880. Com- 


1 «The influence of the nomenclature of human anatomy, reflected down- 
ward upon the dawning structures of the lower animals which culminate in 
man, is nowhere more obstructive to a plain and true indication of the nature 
of parts than in regard to those of the brain.” Owen, ’61, I, 294, note. 


Wiper, WVeural Terms. 257 


parison with the second and third columns will show that in 
most cases our office was merely to disencumber the essential 
elements of prexisting terms from superfluous accessories. 


Part V. Reply to the Criticisms offered by the Anatomische 
Gesellschaft or its Members upon the Terminologic Propost- 
tions or Usage of American Committees or Individuals. 


§135. Inthe introduction to this article (§3) it is inti- 
mated that certain ‘‘ German declarations against the efforts of 
the American committees may be due in part to ignorance or 
How far these mental condi- 


” 


misapprehension of the facts. 
tions might have been avoided may be judged from the follow- 
ing statements. 

§136. As mentioned in Part II, even if my earlier formu- 
lation of principles and specific suggestions (1871-1879) be dis- 
regarded, in and after 1880 papers referring directly to anatomic 
or neurologic nomenclature were published in periodicals pre- 
sumably accessible to anatomists everywhere.’ The appoint- 
ments of the committees of the American Neurological Associ- 
ation and of the American Association for the Advancement of 
Science (§$§80-84) were duly announced in medical and scientific 
journals, and in the transactions of the two associations. A 
brief preliminary report of the A. A. A. S. committee was 
printed in the Proceedings of the Association for 1886 (p. 56). 

§137. The ‘‘Nomenclatur Commission” of the Anatom- 
ische Gesellschaft was appointed in 1889. Primarily German 
(His, ’95, 3), additions were made in the following year from 
England, France and Italy. According to His (’95, 6, 7) co- 
Operation was also asked from Thane, Duval and Testut. A\l- 
though the Anatomische Gesellschaft contains some American 


To what extent reprints of those papers were sent to German anatomists 
at that time, I'am unable to determine. 


2 Its final composition was as follows: A. von Kdlliker, president; O. Hert- 
wig, W. His, Kollman, Merkel, Schwalbe, Toldt, Waldeyer, Bardeleben, Henke, 
von Mihalkovics, Riidinger, von Kupffer, Turner, Cunningham, Leboucgq, 
Romiti, and W. Krause, secretary. This list was furnished me by the 
secretary in April, 1892. 


258 JoURNAL OF COMPARATIVE NEUROLOGY. 


associates, none, so far as I know, was invited to serve on the 
committee. Ifthe omission was remarked by others, they prob- 
ably, like myself, attributed it to the geographic interval. 

§138. In the fall or winter of 1890 I received from the 
secretary, Prof. W. Krause, a letter (since mislaid) reading 
substantially as follows: ‘‘I have an idea that something as to 
Anatomical Nomenclature has been done in America. Please 
to send me copies of Reports and other documents.”’ 

$139. In response there were promptly sent copies of all 
the Preliminary Reports together with my papers and lists up 
to date. Their receipt was acknowledged by Dr. Krause Dec. 
2g9th., 1890, and later in the following : 


Gottingen, Apr., 15, 1891. 
Dear Mr. Wilder : 

I have received your papers with many thanks; if possible 
please to send me some ten further copies of your two pages on polyo- 
nyms, heteronyms etc., of the brain; [‘* Handbook,” VIII, 530, 531]. 
You will receive upon my part after some weeks a complete copy of 
all the proceedings of the European committee on Anat. Nomenclature. 

With kind regards, 
W. KRAUSE. 
P.S. Ihave reprinted your Preliminary Report in the Monthly 
International Journal of Anat. 1891, Vol. VII, No. 5 a. 6, p. 239. 


§140. My answer was as follows: 


Ithaca, N. Y., Apr., 30, 1891. 
Prof. W. Krause, Dear Sir: 
; In response to yours of April r5th, I take pleasure in send- 
ing by bookpost copies of the sheet named, and also copies of printed 
lists, and of figures prepared for my students exemplifying the use of 
the names. We have commonly employed the English paronyms of 
the Latin words, but the latter are given in the ‘‘ Macroscopic Vocab- 
ulary” [’90, 2]. You do not speak of having seen my articles in the 
“‘ Reference Handbook.” Perhaps the work has not been introduced 
abroad: since, however, they present my last views as to the principles 
of Nomenclature, and also apply those principles to the brain, I desire 
that your committee should consult them. In a separate roll will go by 
bookpost a copy in sheets to be kept while of service to the committee. 
Permit me to ask attention especially to the paragraphs on pp. 522 
[$45] and 532 [$82.] Very cordially yours, 
B. G. WILDER. 

P.S. The photographs are of a column inthe same volume, 
VIII, of the Handbook, in W. Browning’s article on the vessels of the 
brain. He accepts and applies the principles of mononymy and 
paronymy. 


Wiper, Neural Terms. 259 


§141. Later, March 27, 1892, I wrote: 


‘¢TIn my letter of April 30, 1891, I announced that I should mail 
you a copy of the sheets of the article ‘‘ Anatomical Terminology ”’ 
from the eighth volume of the Reference Handbook of the Medical 
Sciences, published in 1889. I do not find record of its reception. If 
it was lost in the mail! I will try to lend you another. Iam still of the 
opinion that the article contains the essence of what has been and is to 
be done, and that its perusal will not only facilitate the labors of your 
committee but enable present workers in the matter to avoid neglect 
and injustice toward their predecessors from Chaussier down.” 


§142. To my earlier inquiry (March 20, 1892), as to 
whether ,jthe Handbook article had reached him, Dr. Krause 
replied under date of April 12: ‘‘The [second ?] copy of your 
mentioned article has been set in circulation among the mem- 
bers of the committee.” 

$143. The foregoing was my last communication from 
the secretary. ;A feature of the article of Prof. His (’95; see 
Part VI) led me to ask Dr. Krause (Dec., 1,\ 1895) whether 
the article had ever been transmitted to the former. To this 
query, which was repeated July 10, 1896, no reply has been 
meecived. See, however, the letterof Prof. His, Aug) 27, 1896; 
Part, VI, . 

§144. On the fourth of April, 1892, Dr. Krause had 
sent me a list of the then members of the committee (eighteen 
in number), and suggested that copies of my papers be sent to 
them direct. So far as practicable that has been done ever 
since. 

$145. The kind offer made in Dr. Krause’s letter of 
April 15, 1891, was duly fulfilled. From time to time there 
reached me copies of most of the several ‘‘Abstimmungs” and 
‘‘Schlussredactions”, the last arriving early in 1894. They 
comprise nine hundred and forty-two large pages, and constitute 
a monument to the learning and industry of the secretary of the 
committee. Although not published in the ordinary sense, 
the large number of the committee, their wide European distri- 
bution, and the commentaries of Prof. His (’95), lead me to 


1 Apparently this copy was lost in transmission ; a second was obtained 
from a member of an American Committee and mailed with the above letter. 


260 JOURNAL OF CoMPARATIVE NEUROLOGY. 


regard occasional reference to their contents as no breach of 
confidence. 

$146. Besides the reprint in the ‘‘ Monatschrift” as an- 
nounced in his letter of April 15, 1891, Dr. Krause reproduced 
the ‘‘Preliminary Report” of the committee of the Associa- 
tion of American Anatomists (1889, $81) upon p. 104 of the 
‘‘Osteologie, Abstimmung I,”’ and upon p. 105 the Table of 
arterial names from my paper, ’85,g. Upon p. 103 are com- 
ments, translated as follows: 


“In America also has been formed a committee upon Anatomical 
Nomenclature. ‘Their proceedings have so far led to the herewith 
published preliminary recommendations. At the beginning of Osteolo- 
gy the proposal to apply thoracic instead of dorsal to a region of the 
vertebral column would be taken into consideration. Wilder further 
proposes to choose if possible [wo méglich] mononyms instead of poly- 
onyms, i. e., to say, A. praecerebellaris instead of A. cerebelli superior. 
[This last sentence is most important]. In Wood’s Reference Hand- 
book of the Medical Sciences (VIII, p. 523) are discussed the advan- 
tages of mononyms (single word terms) over polyonyms. That such 
proposals cannot be practicable is evident from a glance at the ap- 
pended list of arteries, or by recalling the binominal system of designa- 
tion introduced by Linnaeus, which only made possible the develop- 
ment of descriptive Natural Science.” 


$147. Notwithstanding the fact that the implied accusa- 
tion as to my accepting no terms consisting of more than one 
word is refuted by the dionym quoted, A. (Arteria) praccerebel- 
larts, this self-raised specter of an exclusive dogma of monony- 
my so haunted the German committee that when their final Re- 
port was adopted, April 19, 1895, the Record of the Proceed- 
ings of the Anatomische Gesellschaft (Azatomischer Anzeizer, 
‘‘Erganzungsheft,”’ 1895, p. 162) contains the following extra- 
ordinary manifesto (here translated) : 


““On motion of the Nomenclature committee (signed by the 
members present at Basel, Herren von KGlliker, W. His, Leboucq, 
Toldt, Fr. Merkel, Schwalbe, Waldeyer, Romiti, von Bardeleben) the 
anatomical society makes the following declaration: The Anatomische 
Gesellschaft believes it should take a stand with regard to the endeav- 
ors of the American Nomenclature Committee. It acknowledges the 
usefulness of as short names as possible, and the suitableness of some 
propositions that have some from America. But it protests against the 
reckless |riicksichtlose] introduction of mononyms, and against the 
consequent radical remodelling of anatomical language as it has hither- 


WILpDER, Weural Terms. 261 


to existed. Consideration for the already established laws of common 
language formation, as also regard for the historic development of our 
own science, forbid the Anatomische Gesellschaft to follow the Ameri- 
can committee in this way. Should the formation of a peculiar ana- 
tomic vocabulary in America advance in the direction indicated, there 
would then be opened an impassable gulf between the representatives 
of anatomic and medical science, and consequently the codperation in 
scientific work hitherto existing would be deeply disturbed.” 

§148. In order to appreciate the full significance of the 
foregoing, there must be borne in mind certain facts: (qa) It is 
directed by the Anatomische Gesellschaft against the endeavors 
[Bestrebungen] of tie American Nomenclature Committee. 
(0) At that time (spring of 1895) there were four American 
committees ($$80-84) representing three distinct associations. 
(c) The committee of the Neurological Association had not 
then reported. (d) The reports of the other three committees 
were substantially embodied in that of the most comprehensive 
committee of the most comprehensive association, viz., the A. 
A. A. 5S., (§84), of which I was nota member. (¢) What had 
been done by any American committee up to the spring of 
1895, therefore, is represented by sections 1-5 of the Neuro- 
logical report (S80). (/) The recommendations therein con- 
tained had been adopted unanimously by the three committees 
and by the three associations, hence the reference of the Ana- 
tomische Gesellschaft to the American Committee, while strict- 
ly inaccurate, really did no injustice. 

$149. Having disposed of these historic details let us 
now see whether the real divergence between the American and 
German conclusions at that time was such as to warrant warn- 
ings and injunctions so solemn and sweeping as are contained 
in the declaration above quoted, (§147.) 

$150. The American recommendations may be grouped 
as (a) specific and (6):general. The specific refer to the follow- 
ing terms: (1) vertebrae thoracales ; (2) hippocampus ; (3) pons ; 
(4) zzsula; (5) calcar; (6) pra; (7) dura. Of these seven, 
four, a majority, coincide absolutely with the German adop- 
tions. 


$151. The other three are discussed in Part III, §§105, 


’ 


262 JOURNAL OF COMPARATIVE NEUROLOGY. 


116, 117). As there remarked, the German adoption of fis- 
sura calcarina ($108) and cavum subdurale (§118) practically 
concedes the adequacy of calcar, dura and pia. 

§152. The special application of dorsal and ventral to the 
spinal cornua and nerve roots is involved in the general recom- 
mendation of the American committees to employ those ex- 
plicit adjectives in place of the ambiguous toponyms of anthro- 
potomy, ($132). As already stated (133) the maintenance of 
the latter by the Anatomische Gesellschaft contravened the 
opinion of the oldest German anatomist, who was also chair- 
man of the ‘‘ Nomenclatur Commission.’’ Americans are not 
likely to repent of a step that has been sanctioned by such 
authority. 

$153. There remains the general recommendation includ- 
ed in section 4, viz., ‘‘That, other things being equal, mono- 


’ 


nyms be preferred to polyonyms.’’ So far as may be inferred 
from the official declarations of the Gesellschaft, and from the 
papers of its secretary and of Professor His, this constitutes the 
most substantial element of the terminologic phantasm. which 
the Germans have erected between themselves and the Ameri- 
can committees. 

$154. How unsubstantial even this really is may be seen 
from the following facts: (a) Of the eleven specific terms 
adopted by three American committees up to the spring of | 
1895, five, nearly one half, were polyonyms, viz., vertebra thor- 
acals, radix dorsalis, radix ventralis, cornu dorsale, and cornu 
ventrale. (b) Even among the forty specific terms adopted by 
the Neurological Association in 1896 (S80) nine are polyonyms, 
nearly one-fourth of the whole. (c) Among the (about) five 
hundred and forty neural terms adopted by the Gesellschaft, at 
least forty, about one-fourteenth, are mononyms, and there are 
others among the names of the other parts of the body. 

$155. Inshort, before condemning the American com- 
mittees for prefering CALCAR to calcar avis, and PIA and DURA 
to pia mater and dura mater, the Germans must justify their 
recommendation of the following mononymic substantives in 
place of polyonyms all of which,are perfectly legitimate, and 


Wiper, Weural Terms. 263 


some of which have been hithero more commonly employed: 
ATLAS (vertebra cervicalis prima); EPISTROPHEUS (Uv. ¢. secundas); 
MANDIBULA (0s maxillare inferius); THALAMUS (thalamus opticus); 
cuNEUS (lobulus cuneatus); PRAECUNEUS (lobulus quadrilaterus); 
UNCUS (gyrus uncinatus); HIPPOCAMPUS (ippocampus mazor); 
PONS (fous Varolit); INSULA (ensula Rel). 

§156. Likewise, even had any one of the committees 
sanctioned certain mononymic adjectives that have hitherto 
been employed only by individuals, before entering upon an 
Index expurgatorius such ‘‘harmless vocables”’ as ARTERIA 
POSTCEREBRALIS (for A. cerebralis posterior ), GYRUS SUBFRONT- 
ALIS (for G. frontalis inferior), and FisSURA PRAESYLVIANA (for 
Ramus anterior ascendens fissurae cerebri laterals | Sylviz }), they 
must account for the following instances upon their own list of 
the replacement of qualifying phrases in perfectly good and reg- 
ular standing by adjectives consisting of a single word each: 
MUSCULUS EPICRANIUS (occipito-frontalis); MuscULUS PLATYSMA 
(M. platysma myoides); SULCUS PRAECENTRALIS (S. centrals an- 
terior); and FIssURA CALCARINA (F, occipitalis horizontalts). 


§157. Notwithstanding, however, the above enumerated 
German examples of mononymy, it must be admitted that there 
still exist somewhat radical distinctions in principle between the 
committees of the two nations. Good words may be predicated 


) 


of both; but so may ‘‘ good works” be done sometimes by 


the ‘‘unjust”’ as well as by the ‘‘just.”” The real distinction 
lies not so much in specific deeds as in general preference. 


$158. Asearly as 1891, the principles upon which the 
German committee proposed to base their revision of Anatomic 
Nomenclature were formulated and announced as_ follows 
(Krause, “ol, 2): 


‘Die erwaihnten Grundsiatze sind nicht so zu verstehen, als ob 
sie etwa ausnahmefreien Naturgesetzen entsprechen sollten. Ihre For- 
mulirung ist um so mehr als eine provisorische zu betrachten, als es 
sich hier nur um die Myologie handelt. 

1. Jeder Kérpertheil darf nur einen einzigen lateinischen Namen 
haben. Es geht nicht an, wie es manche Handbiicher thun, sich mit 
einem sive zu helfen und die beiden Ausdriicke dann abwechselnd zu 
gebrauchen. Deutsche Benennungen, insofern sie nicht ohnehin 


264 JOURNAL OF COMPARATIVE NEUROLOGY. 


feststehen oder soweit sie iiberhaupt néthig, bleiben der freien Aus- 


wahl eines Jeden iiberlassen. 
2. Der Name soll ein kurzes sicheres Merkzeichen sein und 


weder eine Beschreibung noch eine speculative Betrachtung in sich 
einschliessen. 
3. Kein KOrpertheil soll einen unnéthig langen Namen fiihren. 
4. Kein K6rpertheil soll denselben Namen haben, den schon 
ein anderer fiihrt, mit Ausnahme etwa des Falles, wenn es sich um 


Homologien handelt. 

5. Die Namen sollen sprachlich und orthographisch richtig sein. 

6. Die Benennungen nach Personen werden so viel als thunlich 
vermieden, namentlich wenn sie betrachtliche ‘ historische Unrichtig- 
keiten enthalten. 

7. Im Ganzen will die Commission conservativ im weitesten 
Sinne verfahren.”’ 

§158. In the following translation the bracketed interpo- 
lations refer to definitions or commentaries in the present 
article. 

‘¢ The principles here stated are not to be understood as if meant 
for laws of nature free from exception. Their formulation is all the 
more to be regarded as provisional, since muscles alone are here in 
question : 

1. Each part of the body should have a single Latin name. One 
should not, asin many manuals, employ a sive, and then use both 
expressions alternately [pecilonymy, §34]. German designations 
[heteronyms, $43], in so far as they are not already established, or so 
far as they are really necessary, are left to the free choice of the 
individual. 


2. The name should be a short definite expression (sicheres 
Merkzeichen) and indicate neither description nor speculative obser- 
vation, [$51, from Owen]. 

3. No part shall have an unnecessarily long name [magniloquy, 

40]. : 
: iS No part shall have the same name that already belongs to an- 
other, except in the case of homologues, [homonyms, §§23, 67]. 
5- The name shall be grammatically and orthographically correct. 


6. Personal designations are to be avoided as much as possible, 
especially when they contain considerable historic errors; [eponyms, 


§33]- 


7. In general the commission intends to act conservatively in 
the widest sense of the word.” 


$159. In August of the same year Dr. Krause presented 
to the British Association for the Advancement of Science a 
brief statement of principles (Krause, 0) and commentary there 
on. The essential parts are here reproduced. 


Wiper, Neural Terms. 265 


‘‘ The state of things has every year become worse and worse ; 
in Germany especially it has become almost insupportable [$$58-60] 
* * %* In the same university sometimes different anatomical no- 
menclatures exist. * * * In two orthiee years we shall have 
finished and then we shall ask the anatomists of other countries to give 
their candid opinion on the whole. ' 

1. The name should be as short as possible. 

2. Personal nomenclature should not if possible be used. 

3. No part should have more than one name. 

4. This name shall always be a Latin one; every nation can af- 
terward easily translate it [$42] after its own fashion. Latin is the 
only real international language [$46], and by adopting it we hope to 
have a sound foundation.” 


§160. For the most part these principles are to be com- 
mended. Before suggesting qualifications of the fourth items 
I cannot refrain from calling attention to the absolute lack of 
intimation either that any of these principles had ever been 
enunciated before, or that any individual or committee had ever 
undertaken to effect an improvement in anatomic language. 
Granting the inutility of American precepts and examples that 
a German should present to an English scientific body proposi- 
tions as to terminologic reformation as if they were wholly 
original and without (so far as recorded) naming those apostles, 
Barclay, Whewell, Owen, and Pye-Smith (the last still living), 
was surely most incongruous. 

§161. It will be seen from the various passages above 
quoted that the Germans are at last ' in accord with the Amer- 
icans in recognizing the value of brevity as a feature of ana- 
tomic terms. But I have as yet failed to find in their publica- 
tions or private letters even the faintest glimmer of comprehen- 


1 T say at Jast, in view of the enormous number of lengthy terms, both Latin 
and vernacular, for whose continuance and even origia German anatomists are 
responsible ; 260. Some of the heteronyms are indeed ‘‘ fearfully and wonder- 
fully made,” and can be most fitly characterized as verbal ‘‘ tandems,”’ unman- 
ageable by persons not specially trained. As remarked by Owen, ‘‘ The happy 
facility for combination which the German language enjoys has long enabled 
the very eminent anatomists of that intellectual part of Europe to condense the 
definitions of anthropotomy into single words; but these combinations cannot 
become cosmopolitan; such terms as ‘Zwischenkiemendeckelstiick,’ are likely to 
be restricted to the anatomists of the country where the vocal powers are trained 
from infancy to their utterance.” 


266 JOURNAL OF CoMPARATIVE NEUROLOGY. 


sion of the more fundamental grounds upon which the Ameri- 
can committees prefer not merely that terms should be reason- 
ably short, but also, when practicable, consist of single words 
each, i. e., mononyms, or, at most, of two words each, a noun 
and a mononymic adjective. 

$162. In addition to the incidental previous re ferences to 
the subject in the present paper (§$25, 47, 50) the differences 
between the more and the less essential advantages of mono- 
nyms over polyonyms may be fairly indicated by extracts from 
two earlier articles. The first (’84, e) was presented to the 
American Association for the Advancement of Science in con- 
nection with my suggestion that a committee on Anatomic No- 
menclature be appointed by that body. Since it contains no 
reference to prior terminologists, it may be proper to add that, 
as published, it was a mere abstract, and that in both earlier 
and later papers of greater’ length (31, 0, c; W21GeNGpees 
’89), an effort was made to give due credit to those who had 
either indicated or smoothed the way. 


$163. ‘*7. Should not organonymic terms (terms of designation) 
be, as far as practicable, brief; capable of inflection; classic in deriva- 
tion and form; already used in a kindred sense? ! 

8. Since the length of aterm may depend upon not only the 
number of syllables and letters, but also upon the number of separate 
words, and since, properly, only single words are capable of inflection, 
is there not a twofold reason why the names of parts, weth certain self- 
evident exceptions, (nerves, etc.2) should be mononymic ? 

g. Is there much real analogy between the nomenclature of 
anatomy and that of zoology and botany ? [cf. $146, last line.] 

10. How far should priority be regarded in the selection of ex- 
isting names ? ; 

11. Can priority be claimed for terms which are vernacular or 
descriptive ? 

12. In considering all questions of terminologic reform, should 
we not regard less our present and personal convenience, than the in- 
terests of the vastly more numerous anatomical workers of the future?” 


\ [This note was a part of the paper.] ‘In later papers ('85, 4 and ¢) I have 
suggested that, in English works, so far as possible, the names be given an Eng- 
lish aspect by paronymization. For example, commissura becomes commissure ; 
pedunculus, peduncle, With many names no change is needed, as with Zorta, 
aulix, fornix, callosum, etc. Heteronyms or vernacular translations are regarded 
as objectionable. The same principle is applicable to other languages,” 


2. This phrase is italicized for a reason that will appear later, 2172. 


Wiper, Weural Terms. 267 


§164. The distinctions were more fully set forth seven 
years ago in the later article (W. & G., ’89, 523) from which 
is taken the following summary. 


A. From mononyms adjectives may be regularly derived. 

B. Mononyms are more readily compounded. 

C. Mononyms are constant in form, excepting for the regular 
derivatives, numbers, cases, adjectives, and paronyms. 

D. Polyonyms, on the contrary, are subject to variations of sev- 
eral kinds: (a) By omission of words. (4) By the conversion of gen- 
itives into adjectives. (c) By the substitution of totally different words. 
(2) By permutation ($37). 

E. Mononyms may be more uniformly abbreviated ($38). 

F. Mononyms are commonly shorter than the corresponding 
polyonyms. The exceptions are due to the fact that the absolute 
length of a term depends upon three factors, viz , (a) the number of 
words ; (6) the number of syllables in the words; (¢) the number of 
letters in the syllables. The written length of a term is affected by 
all three of the above named conditions ; but its spoken length is in- 
dependent of the number of letters. 

$165. But the essential characteristic and principal advantage of 
the Latin mononym is (G) that 772s capable of adoption into any other 
language, either unchanged, or with so slight amodtfication as not to hinder 
tts ready recognition by the anatomist of any nationality. 

$166. Between longer and shorter words is merely a difference 
in degree, e. g., crus and pedunculus; between a Latin word and its 
vernacular equivalent or heteronym (e. g., pedunculus and footlet) and 
its paronym (peduncle) the choice might depend upon individual pref- 
erence; but mononyms and polyonyms, terms of one word each and 
terms of two or more (e. g., pedunculus and pedunculus cerebelli) differ 
in kind; it is not a question of size or euphony, but of essential en- 
dowment and capacity. The mononym is to the polyonym as is the 
water to the earth or as is the bird to the tortoise. 

$167. The desirability of replacing polyonyms by mononynms is 
in direct ratio with the frequency with which the part is mentioned, 
and with the need of employing corresponding adjectives. Mypophy- 
sis (formerly pituitary body) and conarium (formerly pineal gland) are 
much more frequently named since the discovery that the former has 
peculiar developmental relations with the pharynx, and the latter even 
more remarkable connection with a vestigial organ of sight. So long 
as the thin sheet of nervous tissue just dorsad of the chiasma was re- 
garded as insignificant, and even as occasionally absent, either damina 
cinerea or lamina terminals might not be so objectionable ; but its now- 
admitted morphologic importance as the cephalic boundary of the en- 
cephalic cavities justifies the use of the mononym, éerma. The ort 
fice left by the removal of the hypophysis and infundibulum had ap- 
parently received no name up to 1880; foramen infundibult was a suf- 
ficiently appropriate descriptive term ; but the frequency of its men- 
tlon in notes respecting the preservation of the brain led me to discard 


268 JOURNAL OF COMPARATIVE NEUROLOGY. 


it in favor of Jura. The first two cervical vertebrae alone have mon- 
onymic titles ; but were there similar anatomical, physiological, or 
surgical occasions for specifying any other member of the vertebral 
series, a mononym would probably soon be proposed and generally 
accepted. 


§168. The German criticisms of the American methods 
and results quoted above (§$147-153) are comparatively brief 
and general. A more extended and particular arraignment is 
contained in the article of Prof. Wilhelm His (95, 6-7).  Al- 
though published under his name, yet its association with the 
official list of Momzna anatomica' as adopted by the Gesellschaft, 
the prominent part taken by him in the formation and work of 
the committee,and his high reputation based upon splendid con- 
tributions to science extending through many years, all endow 
his utterances upon the subject with not merely personal but 
semi-official significance. Indeed, in the absence (so far as I 
know) of any dissent therefrom, those utterances must be re- 
garded as representing the views of the older German anatom- 
ists?» On this account it is more to be regretted that some of 
his criticisms should exhibit in a marked degree the lack of in- 
formation and of comprehension mentioned in the introduction 
to the present article (§§3-4). 

$169. Criticism by Prof. Wilhelm His (’95, 6-7).° 


‘‘Die anatomische Gesellschaft Grossbritanniens hat 1893 eine 
eigene Commission zur Anpassung unserer Vorschlage an die englisch- 
en Bediirfnisse niedergesezt,4 und etwas friiher noch (1890) hat eine 
Gesellschaft amerikanischer Anatomen die Nomenclaturfrage in die 
Hand zu nehmen versucht. Die Aeusserungen der englischen Com- 
mission liegen noch nicht vor, dagegen haben die amerikanische 
Commission und deren sehr eifriges Mitglied, Hr. Wilder, bereits 


1It would be desirable to know whether the list may be obtained otherwise 
than as part of the article, as will be the case with Part VII of the present 
paper. 

2By some of the younger neurologists the American ideas have been more 
hospitably entertained and certain newer terms actually adopted. 

3The original is here reproduced verbatim et literatim et punctuatim, In the 
translation (7170) I have ventured to modify certain subordinate features that 
may have been due to haste upon the part of the author or proof-reader. 


4 Journal of Anatomy and Physiology, 1894. Vol. XXVII [XXVIII]. 


WiLvER, WVeural Terms. 269 


eine Reihe von kleineren Aufsitzen und Broschiiren ver6ffentlicht! 


‘‘Die Ziele, welche die amerikanische Commission verfolgt, 
sind dieselben wie die unsrigen, ihre Wege zum Ziel sind aber 
wesentlich andere, als die von uns gewahlten. Gemeinsam ist uns 
der Grundsatz, dass jeder Theil nur einen einzigen Namen haben 
soll, und dass dieser Name méglichst einfach und bezeichnend zu 
wahlen sei. Wilder und seine Collegen gehen aber weiter und verlang- 
en lauter ‘‘ Mononyme,” d. h. Substantive ohne ferneren Zusatz. 
Sie sagen z. B.: praecornu und postcornu satt cornu anterius und 
cornu posterius, postcava statt vena cava posterior u. a.m. _ Bis jetzt 
liegt von Wilder die Bearbeitung der Gehirnnomenclatur nach den vor- 
geschlagenen Grundsiatzen vor, und dieser erste Versuch erlaubt uns 
ein Urtheil iiber die Consequenzen, zu welchen das vorwiegende Bes- 
treben nach Mononymen hinfiihrt. Wir stimmen unsererseits vollig 
bei, wenn das wort Thalamus kurzweg an die Stelle von Thalamus op- 
ticus gesetzt wird, wir haben selbst nichts dagegen, wenn im taglichen 
Verkehr das Wort Dura fiir dura Mater encephali in allen den Fallen 
gebraucht wird, wo eine Verwechselung ausgeschlossen bleibt. Da- 
gegen kénnen wir nicht anerkennen, dass Worte wie ‘‘ Medipeduncu- 
lus” fiir Pedunculus cerebelli ad pontem einen sprachlichen oder 
praktischen Fortschritt bedeuten. Die Zusammenziehung verschied- 
ener Worte in ein einziges kann ja unter Umstanden eine Vereinfach- 
ung sein, Aehnlich dem allzu gedrungenen Telegraphenstil kann sie 
indessen auch zur Unklarheit fiihren, und dann ist ihr Nutzen ein neg- 
ativer, denn also gebildete Worte verlangen zu ihrem Verstandniss 
besondere Erlauterungen. Medipedunculus ohne Zusatz ist unver- 
standlich, es miisste schon Medipedunculus cerebelli heissen, und dem 
ware wiederum Pedunculus medius cerebelli vorzuziehen, weil das 
barbarisch gebildete Wort Medipedunculus ebensowohl fiir Pedunculus 
medius, als fiir Pedunculus medialis oder fiir Pars media oder medi- 
alis pedunculi gebraucht werden kénnte. Sprachwidrige Wortzusam- 
mensetzung enthalt aber Wilder’s Liste sehr viele, und man hat nicht 
néthig, ein philologischer Pedant zu sein, um in Worten wie Terma 
(anstatt Lamina terminalis), Postramus (fiir Ramus posterior arboris 
cerebelli) u. a.m. Aergerniss zu nehmen. Viele Worte, wie Cimbia 
(Tractus peduncularis transversus), coelia (fiir Cavitas encephali), 
Aulix (fiir Sulcus Monroi) u. a. m. sind iibrigens neu oder wie Isth- 
mus (fiir Gyrus annectens) in einem anderen, als dem bisherigen Sinn 
benutzt. Ich weiss nicht, wie weit der Kreis amerikanischer Fachge- 
nossen reicht, welchen Wilder hinter sich hat. Jedenfalls fiihrt das 


1Von den durch Wilder versandten Schriften citire ich als die hauptsachlich- 
sten. The fundamental principles of anatomical Nomenclature, by Burt C. 
Wilder MD. from the Medical News. 19. December 1891. 

Ferner: Fissural diagrams of the human brain. Macroscopical Vocabu- 
lary of the brain presented to the Assoc. of American anatomésts at Boston, Mass. 
29. Dec. 1890. American Reports upon Anatomical Nomenclature. 1889- 
1890, with Notes by Wilder, Cornell University, 5. February 1892. 


270 JouRNAL OF CoMPARATIVE NEUROLOGY. 


Vorgehen des letzteren zur Schaffung einer véllig neuen, grossentheils 
recht fremdartig klingendeti Sprache, und auf diesen Boden kann 
ihm unsere Commission, ohne Verleugnung ihrer historischen Grund- 
satze, nicht folgen.” 


§170. In the following translation of the foregoing extract 
from the article of Prof. His (’95, 6, 7), besides corrections of 
supposed typographic errors, and changes in the order of words 
in accordance with more usual English custom, there are intro- 
duced in brackets (2) words from the original that might have 
more than one meaning ; (2) references to sections of the pres- 
ent article; (c) letters to facilitate the assignment of subsequent 
commentaries to the special points involved. 


‘« The anatomical society of Great Britain appointed in 1893 its 
own commission to adapt our propositions to English needs,! and 
somewhat earlier still (1890), [a] a society of American anatomists 
[4] tried to take up the nomenclature question. The statement of the 
English committee is not at hand, but the American committee [c] 
and its very ardent member, Mr. Wilder [d ], have already published 
a series of small papers and dbrochures.2 The aims of the American 
committee are the same as our own, but their methods are essentially 
[7] different from those chosen by us. The fundamental idea is the 
same, that each part should have only a single [Latin] name (§46), 
and this the most simple possible [g]._ But Wilder and his colleagues 
[Z] go farther and desire [z] absoiute [verdangen wists mononyms, 2. é., 
substantives without further additions. They say [7], for example, 
praccornu and postcornu instead of cornu anterius and cornu posterius ; 
postcava [| instead of vena cava posterior, with many similar terms. 
We have already the treatment of encephalic nomenclature by Wilder 
in accordance with the proposed method, and this first attempt allows 
us [/] to form an opinion as to the results [7] to which the preponder- 
ating |vorwzegende| effort toward mononymy leads. On our part we 
assent fully when the word ¢halamus alone [7] takes the place of thal 
amus opticus ; we even do not object if, in ordinary communications, 


1 Journal of Anatomy and Physiology, XXVIII [not XXVII, as in original], 
pp. VII-IX, 1894. » 


2T cite here the chief [Aauptsdchlichsten] of Wilder’s [e] transmitted [ver- 
sandten| publications: ‘‘The fundamental principles of Anatomical Nomencla- 
ture,’”? The Medical News, Dec. 19, 1891, 708-710. ‘‘ Fissural Diagrams,” accom- 
panying ‘‘ Remarks on the Brain of Chauncey Wright, with Commentaries upon 
Fissural Diagrams,’’ Jour. of Nervous and Mental Disease, XVII, 1890, 753-754; 
Amer. Neurolog. Assoc. Transactions, 1890. ‘‘ Macroscopic Vocabulary of the 
Brain, with Synonyms and References ;”’ presented to the Association of Amer- 
ican Anatomists, Dec. 29, 1890; O., pp. 13. ‘‘ American Reports upon Ana- 
tomical Nomenclature, 1889-1890, with Notes.” O., pp. 3, Feb. 5, 1892. 


Wiper, Neural Terms. 27% 


[0] the word dura for dura mater encephalt is used when misapprehen- 
sion is impossible. But we cannot acknowledge that words like med¢- 
pedunculus |p| for pedunculus cerebelli ad pontem constitute a linguistic 
or practical improvement. The contraction of several words into a 
single one may be under certain circumstances [g] a simplification ; 
but, as with the too concise telegraph style [7], it may, on the other 
hand, involve lack of clearness, and then its advantage 1s negatived ; 
for words formed in this manner demand a special explanation for their 
comprehension. Medipedunculus alone is unintelligible [s]. It must 
surely be called medipedunculus cerebelli, and over this again should 
preference be given to pedunculus medius cerebellit, because the barbar- 
ously formed [/] word medtpedunculus could be used indifferently for 
pedunculus medtwus, for pedunculus medialis, or for pars media or medtalis 
peduncult [uw]. But Wilder’s list contains very many [v] ungrammatic 
verbal combinations, and one need not be a philologic pedant [zw] to 
take offense at words like ¢erma (instead of lamina terminalis) [x], post- 
ramus (for ramus postertor arboris cerebellt) |y| and others. Man 
words like cimbia (for tractus peduncularis transversus) [2], coelta (for 
cavitas encephalt) |a|, aulix (for sulcus Monrot ) ||, and others, are 
moreover new [7], or, like ¢sthmus (for gyrus annectens), used in another 
than the accepted sense. I know not how far may extend the circle 
of American collaborators supporting Wilder [0]. At all events, his 
proposals tend to create a language entirely new [<] and for the most 
part quite strange, and on this ground our commission cannot follow 
him, without renouncing its historic principles [y].” 


§171. Certain points in the foregoing extract have already 
become the subjects of the correspondence between Prof. His 
and myself referred to in the Introduction ($4) and constituting 
Part VI. Their inclusion in that later division of the present 
article is the more desirable in view of the hope based upon 
my last letter from Prof. His that, before that part is put in 
type, he may have discussed those points publicly. 

$172. It will be seen that, like the German committee 
and the entire Gesellschaft, Prof. His is disturbed by the Amer- 
ican preference for mononyms.. In one passage he character- 
izes it as the ‘‘vorwiegende Bestreben” [m]; in another [i] 
by ‘‘verlangen lauter.”” Now verlangen has two distinct mean- 
ings, viz., deserve or prefer, and demand or insist upon. As will 
appear in Part VI, it is now (Nov. 10, 1896) nearly a year since 
Prof. His had an opportunity to state in which of these senses 
the word is to be understood, and more than two months have 
elapsed since the ambiguity was brought expressly to his notice. 
The sweeping charge (§147) framed by the Gesellschaft in his 


272 JOURNAL OF COMPARATIVE NEUROLOGY. 


presence and apparently under his direction would justify the 
interpretation of verlangen as demand. But until assured to that 
effect by Prof. His himself, I hesitate to entertain so monstrous 
a perversion of the facts upon the following grounds: (a) In all 
the reports of the American committees the expression of pref- 
erence for mononyms is accompanied by the proviso ‘‘ other 
things being equal.’”’ (0) I have repeatedly conceded the im- 
possibility of the application of the principle to groups of parts, 
fissures, gyres, vertebrae, bloodvessels, muscles and nerves. 
The remark that there are ‘‘certain self-evident exceptions 
(nerves, etc.),” occurs in a paper ('84,¢; $163, 8) presented 
by me twelve years ago to the American Association for the 
Advancement of Science and printed in the Proceedings of that 
body (pp. 528-529). If Prof. His ever consults any American 
scientific publication it would seem not unnatural that the title of 
that paper, ‘‘Onsome Questions in Anatomical Nomenclature,” 
should have attracted the notice of one who, three years later, 
himself urged (’95, 1) upon the German anatomical society the 
need of action upon the subject, and who was occupied. there- 
with more or less for eight years afterward. 

$173. In order to eliminate so far as possible the personal 
element from the consideration of the special criticisms of Prof. 
His, I select as the first subject of rejoinder a term, Postcava, 
in which my interest is only indirect, as of one toward a child 
by adoption rather than by paternity. Omitting intervening 
phrases not affecting the interpretation, the complaint of Prof. 
His reads (translated) as follows :—‘‘ Wilder and his colleagues 
[4] * * * say [7] praccornu and postcornu for cornu anter tus 
and cornu posterius, postcava [k| for vena cava posterior, with 
many similar terms.’’ The implied disclaimer as to ‘‘ philologic 
pedantry ” [zw] can hardly embrace a toleration of misstatement ; 
hence, before discussing the intrinsic merits of the word se- 
lected, it may be well to dispose of minor points that might 
complicate the main issue. 

$174. In the text Prof. His refers only to ‘‘ Wilder,” and 
in note 2 (see the original, $169) an initial is wrong. Hence it 
is only just to state that my terminologic transgressions must 


Witper, Neural Terms. 273 


not be imputed to Harris H. Wilder, professor in Smith Col- 
lege, Northampton, Mass., whose researches, especially upon 
lungless salamanders,’ make me proud to claim him as a distant 
relative. 

$175. The objectionable words are attributed to ‘‘ Wilder 
and his colleagues.”” Not one of the three specified words, or 
of the ‘‘many similar terms,” has been sanctioned by either 
of the four committees, and few of the members thereof have 
adopted them. For the confusion and possible injustice here 
occasioned no adequate explanation can be offered. 

§176. The phrase ‘‘postcava statt vena cava posterior” 
would naturally imply that the latter is the name preferred by 
the German committee. Yet the official list contains (p. 77) 
only vena cava inferior.” 

$177. So far as appears in the article of Prof. His, fost- 
cava was coined by me. On the contrary, so far as I am aware, 
it (in the derivative, postcava/) was first introduced by Richar d 
Owen about the middle of the century, and employed by him 
consistently thereafter. 

§178. Whether or not the two historic facts just men- 
tioned® were known to Prof. His he alone can tell, and the fate 
of other queries does not encourage an effort to ascertain. 
Hence I am compelled to offer propositions which each reader 
must accept, or reject, or explain in accordance with his own 


” 


information and judgment. 

(1). Postcava, inthe form fostcaval (§177), occurs fre- 
quently in the writings of a leading English anatomist. 

(2). Those writings must be known and accessible to Prof. 
His. Hence there is no excuse for the erroneous intimation in 
the article. 


1 Anatomischer Anzeiger, 1X, Jan. 20, 1894, and XII, 182-192, 1896. 


+ In passing it may be remarked that the retention of superzor and inferior 
as the essential elements of the designations of these great vessels constitutes 
one of the many evidences of the non-emancipation of the German committee 
from anthropotomic enslavement; see 2133. 


3 My non-responsibility is certain; the responsibility of Owen is assumed 
in the absence of evidence to the contrary. 


274 JOURNAL OF CoMPARATIVE NEUROLOGY. 


(3). Whatever its source, postcava differs from the more 
usual terms in its comparative brevity, while at the same time 
not open to the charge of ambiguity. Why then was it not in- 
. Cluded in the column of synonyms from ‘‘ sonstigen Autoren” 
in the protocols of the German committee, as was a less com- 
mon and acceptable synonym, viz., ‘‘ vena cava _ inferior 
thoracica ?”’ 

(4). If the entire committee supposed me to be the au- 
thor of postcava, their action was consistent, since no term is 
credited to me in the column indicated. 

(5). But if any members of the committee knew that 
posicava originated with Richard Owen, their objections to the 
word might well have been waived out of respect for him. 

§178. The actual form employed by Owen is specified 
above, not merely for the sake of accuracy, but also in order to 
forestall criticism upon a point where disagreement is possible. 
It is, I think, a sound proposition that the zntroduction of any 
derivative, oblique case, or national paronym, practically renders 
the introducer responsible for the actual or potential Latin antece- 
dent of such words, in accordance with the usual rules of derivation 
and paronyny. 1 donot remember seeing the foregoing propo- 
sition distinctly formulated,’ but reflection will show its sound- 
ness. One of the wisest recommendations of the A. A. A. S. 
Committee on Biological Nomenclature ($85) was that the Latin 
(international ) form of a term should always be given, whether 
or not the national paronyms. Now cava is the feminine form 
of cavus, and vena cava was used (perhaps not in the specific 
modern sense ) by Cicero, ‘‘De Natura Deorum,” 2, 55, 38.” 
There seems to have been no classic adjective, although cavatus, 
the participle of cavo, was available as such. Analogy fully 
warrants (S116, et seq.) the acceptance of cava as a subsian- 
tive, and the derivation therefrom of a secondary adjective in the 
form of either cavatus or cavalis. The latter evidently was 


1It probabiy has been in purely linguistic connections. 


* For some discussion of cava see Hyrtl, ’80, 98, 99. 


Wiper, Neural Terms. 275 


chosen (constructively) by Owen when (in 1862, ‘*Onthe Aye- 
Aye,”’ Zool. Trans., V, 86, and perhaps earlier) he employed 
post-caval vein and pre-caval vei. Later, the hyphen was 
omitted, and in the ‘‘ Comparative Anatomy of Vertebrates ”’ 
occur ‘‘postcaval vein, postcaval trunk, postcaval orifice, and 
postcaval,) 0503-505; 11, 203); TIT, 552 et seq. ) Bending 
the discovery in Owen’s writings of some history of the stages 
by which the final reduction was effected, the following series 
is certainly thinkable:—(1) Vena cava posterior; (2) Posterior 
vena cava; (3) Posterior caval vein; (4) Post. caval vein; (5) 
Post-caval vein ; (6) Postcaval vein ; (7) Postcaval ; (8) Postcava. 
Whatever may have been the actual steps, never did Owen 
reach a more final terminologic result, and no case better 
exemplifies the unwisdom of the reactionary attitude of the 
German committee. 

$179. Since Professor His offers no specific objections to 
postcava their nature can only be inferred from his general re- 
marks and from his criticisms of medzpedunculus. Perhaps there- 
fore the simplest and most comprehensive rejoinder is to recap- 
itulate briefly the several attributes of the term, leaving each 
reader to estimate their value for himself. Besides references 
to this article, the initial G will indicate terms adopted by the 
Anatomische Gesellschaft, or attributes commended by that 
body. 

(a) Brevity; §§40, 50; G., §158, 3. (6) Latin form; 
§50; G., §158, 1. (¢) It isa mononym; §§15, 47. (d) It is 
a locative name; §27. (¢) It is an adjectival locative; §29. 
(f) It is capable of inflection, i.e., postcavalis, postcaval, post- 
cavals. (g) Its various national representatives (paronyms 
$$43-45) differ little or none from the international antecedent. 
(2) It has in the derivative, pos/caval, high authority (Richard 
Owen) and moderate antiquity (1862 or earlier). (¢) It is an 
idionym ($24) and not likely to be applied to any other part in 
any vertebrate. (#) It is sufficiently euphonious, and easily re- 
membered. (/) Like other euphonious and easily remembered 
mononyms it constitutes no bar to the progress of one who may 
never have heard the more common polyonyms. Those who 


276 JoURNAL OF COMPARATIVE NEUROLOGY. 


are familiar with those polyonyms, whether vexa cava infertor, 
vena cava ascendens, or vena cava posterior, could hardly fail to 
recognize its signification. Since 1881 no other term than Jost- 
cava has been used by me for the great vein in question. I 
have yet to learn of a single instance of misapprehension or 
other difficulty caused thereby among either general or special 
students. 

§180. There remains the question of the etymologic or- 
thodoxy of postcava, and this involves the much more comprehen- 
sive and difficult question as to the definition of etymologic 
orthodoxy. Without presuming to invade the jurisdiction of 
philologic experts, for the practical discussion of the case in 
point, precedents need be sought in only two periods, the classic 
and the recent. 

$181. I freely admit that there is known to me no in- 
stance in classic Latin literature of the employment of jos?, 
whether alone or in composition, with the force of an adjective 
and as equivalent to posterus or posterior. That this negative 
evidence is hardly conclusive may be seen froma single case 
among the scores that might be adduced. With the Romans 
ztem was anadverb. With us it is not only an adverb, but also 
a noun and a verb, and the basis of two derivatives, zdemzze and 
atemiser. 

§182. In recent times the precedents are partly direct and 
partly indirect. Among the former are fostabdomen, postact, 
postarytenoid, postfactor, postfurca, postpubis, postscapula. In all 
of these post has the force of an adjective, not of a preposition. 

$183. Indirect precedents are cases in which other prepo- 
sitions have the force of adjectives in composition. Such are 
preadaptation, precentor, preexistence, preformation, presternum ; 
also subgenus, subflavor, subfactor, submaster, subtitle." 

$184. Since, however, the German committee sanction 
none of the anatomic terms in the foregoing lists and avoid the 


1 Among analogous Greek words the following has been furnished me by 
my friend, L. L. Forman, Instructor in Greek at Cornell University : npogviag, 
an advance guard, 


WiILpER, Neural Terms. 277 


use of praesternum by retaining manubrium sternt, they would 
probably decline to regard them as adequate justifications for 
postcava. But can they consistently. condemn it or any sim- 
ilar terms? Let us see. 

§185. Prof. His, the German committee, and the Ana- 
tomische Gesellschaft, after several years deliberation and appar- 
ently without any disagreement, have adopted and recommended 
the names metencephalon and prosencephalon for certain segments 
of the brain. Now meta and pros are the English forms of the 
Greek perd and zpég. These are both prepositions. Like post 
and prae they are also adverbs. The terms into which they 
enter have no reference to a third part ‘‘behind’”’ which or 
‘before’ which the metencephal and prosencephal are situated. 
The German translation of prosencephalon is Vorderhirn, and the 
English, forebrain, both signifying the first or most cephalic 
member of the series of coordinate encephalic segments. With 
slight modifications the foregoing remarks apply equally toa 
third name adopted by the German committee, dencephalon, 
the preposition 6a having the force of an adjective. 

§186. Iam unable to recognize any distinction, logical or 
etymological, between the metencephalon and prosencephalon 
which the Germans commend, and the fostcava and praccava 
which Prof. His condemns. The irregular terms for which he 
is in part responsible may be few; but his virtuous denunciation 
of me for producing a larger number of the same sort is no 
more reasonable than the demand of the woman to be punished 
lightly for bringing forth an illegitimate child upon the ground 
that it was ‘‘such a little one.” 

§187. Strictly, however, even if the degree of oppro- 
brium to be cast upon the individual concerned were to be 
measured by the number of terms of a certain kind, this 
would have no bearing upon the question of the acceptability 
of agiven term. /Postcava and praecava are to be considered 
upon their merits as brief, convenient, and absolutely unambig- 
uous designations intended to replace inconvenient descriptive 
phrases. In favor of vena cava superior and vena cava inferior 


278 JOURNAL OF COMPARATIVE NEUROLOGY. 


antiquity alone can be urged; against praecava and postcava 
can be alleged only the sinfulness of comparative youth. 

_ $188. Inthe foregoing discussion I have refrained from 
following one line of argument that readily suggests itself and 
is, indeed, almost formulated in the hypothetic series between 
posterior vena cava and fostcava as stated in $178, wvzz., The 
prefix post might not unnaturally be regarded as the abbrevia- 
tion of posterior or postero. Were compounds of fost alone con- 
cerned, this simple line of argument might perhaps be adequate; 
but it will not serve for compounds of the correlative pvae, nor 
for those of the Greek prepositions, éx/, yerd, ixd, etc. 

§189. The straightforward way of dealing with the mat- 
ter is to assume that fostand prac, in composition, may have 
the force of the adjectives posterior and anterior respectively.' 
‘<Tf this be treason, make the most of it.” 

S190. It seems to me that the nature of the issue between 
postcava and vena cava inferior (or posterior) is such as to in- 
volve the acceptance or rejection of the following propositions. 

A. Language was made by and for man, and not the 
reverse. 

B. Grammatic rules are framed from time to time in order 
to maintain the uniformity that is acceptable and convenient. 

C. Like the roads we traverse, such rules are but means 
to ends, and have no intrinsic sanctity. 

D. Like a circuitous but familiar road, a commonly ac- 
cepted rule is not to be abandoned without reflection. On the 
other hand, no more is it to be laboriously travelled when new 
conditions render a ‘‘short cut”’ desirable. 

E. Extrinsic toponyms (2. ¢., terms of location or direc- 
tion that do not refer expressly to the recognized body-regions, 
dorsum, venter, etc.) should conform to the more usual verte- 
brate attitude rather than to the erect attitude of man; e. g., 


1 Tt is well understood in this country that the Vew York Medical Journal 
and the ‘‘ Encyclopaedic Medical Dictionary ”’ stand for the highest scholarship. 
Yet so long ago as 1885, when some of my simplified terms were submitted to 
him, their editor, Dr. F. P. Foster replied, * I think some of the words excel- 
lent, Araccommissura, for example.”’ 


Wiper, Neural Terms. 279 


posterior and antertor, supertor and inferzor, and their deriva- 
tives, compounds, and abbreviations, should have significations 
zootomic rather than anthropotomic. 

F. There now prevail and are likely to persist two con- 
ditions not merely unknown to the Paires anatomict, but prob- 
ably not imagined by them: (a) the enormous increase of ana- 
tomic and physiologic knowledge; (0) its general diffusion 
among the people.'' These two conditions” militate against the 
rigid maintenance of grammatic rules that might prevent the 
establishment of new and shorter channels, or the fabrication of 
new and briefer technical terms, the ‘‘tools of thought.”’ 
Terms like vena cava posterior are obtrusively Latin, and hence 
not acceptable to the laity; too much time and space are lost in 
speaking and writing them, and time and space are daily becom- 
ing more precious. 

§1g1. Consciously or unconsciously, for many years Eng- 
lish and American anatomists have been gradually simplifying 
their terminology in substantial accordance with the foregoing 
propositions. In Germany the signs of such improvements are 
as yet comparatively few. 

§192. Even if, however, the German committee were 
reconciled to the employment of certain prepositions in compo- 
sition with the force of adjectives, there would still remain’ 
special objection to fost as indicating toward the tail rather than 
toward the back. This objection is radical, and the conflict in- 
volved is irrepressible ; $$131-133. 

§193. Postramus.—To this, as a mononymic substitute 
for Ramus posterior arboris vitae cerebel, Prof. His offers no 


1 Tn fulfilment of the declaration of the elder Agassiz, ‘‘ Science must cease 
to be the property of the few; it must be woven into the common life of the 
world.” 


2 There is really a third condition, equally novel, but bearing less directly 
upon the present question, viz., the pursuit of anatomy by women. Whatever 
view may be taken of this in other respects, all decent men must rejoice that it 
has hastened the elimination of the needless Momina impudica which formerly 
defiled even the description of the brain. For further commentary upon this 
matter see W. & G., ’82, 27. 


3’ Excepting with the chairman, 2133. 


280 JOURNAL OF COMPARATIVE NEUROLOGY. 


specific objections (§170, v) but they may be inferred to be (a) 
that it isa post compound (§§174-191); (0) that the German 
list does not include any terms for the branch-like divisions of 
the cerebellar ‘‘tree.’’ If these branches no longer merit spec- 
ification, postramus and pracramus will vanish quietly with the 
polyonyms from which they were condensed. 

§194. Terma.—From the context (§170, #) it may be in- 
ferred that this is objected to upon etymologic grounds. Ter 
ma does not, it is true, occur in classic Latin lexicons. But 
neither does chzasma which is embraced within the German list. 
The latter is the ‘‘new Latin” paronym of yiacua, and ferma is 
merely somewhat newer Latin for répya. Why should we use 
the longer Latin terminus for terma any more than the longer 
Latin decussatio for chiasma? 

§195. Aulx.—This (§170, f) is the regular Latin paronym 
of avija& and signifies a furrow. It was proposed by me asa 
mononym for Reichert’s Szdcus Monrot, not merely as a mono- 
nym but because its resemblance to az/a’ readily recalls the 
fact that it connects the lateral orifice of the aula (the porta) 
across the face of the thalamus with the funnel-shaped orifice of 
the ‘‘aqueduct.”’ If, as apparently held by His and Minot, this 
furrow should prove to be only part of the general boundary 
line (Sulcus imitans ventriculorum or Sulcus interzonals) between 
the dorsal and ventral zones, then it might not need specifica- 
tion. But the later observations of Mrs. Gage (96, a), as stat- 
ed in my paper (’96, @), cast considerable doubt upon this inter- 
pretation of the ‘‘sulcus Monroi.”” In any caseit must be borne 
in mind that it extends from the aqueduct to the aula and not to 
the Recessus opticus as represented in one published figure. 

§196. Czmbia.—This architectural term, signifying a band, 
fillet or cincture, was"used by me in 1881 in discussing the very 
distinct ridge across the crus (cerebri) of the cat before I knew 
that it had been called Tvactus peduncularis transversus by Gud- 
den (1880) and still earlier (1861) fasczo transverso by an Italian 


1My mononym for pars ventriculi communis media,pars foraminis Monrot me- 
dia, etc.; 211. 


WILpDER, WVeural Terms. 281 


anatomist. Unless the first name is to be retained there can be 
no logical objection to replacing the second by a third. The 
most complete exemplification of the practical superiority of 
cimbia is supplied by the section concerning it in the latest 
edition of the ‘‘Gewebelehre”’ by the distinguished chairman of 
the German committee. On pp. 606-609 (really only three 
pages if the cuts be excluded) and in addition to the page-head 
and the section-title, Zvactus peduncularis transversus occurs ten 
times, occupying more than five fulllines. The occurrence of 
Tractus peduncularts in one place, and of TZvactus alone or in 
composition with German words in several, shows how burden- 
some the polyonym had become, and howirresistible the tempta- 
tion to vary. In the explanation of Fig. 707, the full title oc- 
curs once; also Zvact. ped and Tr. ped. ; compare §38. Tractus 
transversus pedunculi of Brissaud is declared to stand for a differ- 
ent bundle, but in the absence of such declaration the two 
names would almost inevitably be supposed to mean the same 
thing anatomically as they do etymologically. Again, since 
Kolliker concedes pedunculi also to the cerebellum (pp. 337, 371, 
etc.), and since many anatomists prefer to designate the fibrous 
masses between the pons and the optic tracts as the crura, there 
is ample opportunity for misapprehension upon the part of the 
student, unless, in accordance with the absolute explicitness in- 
sisted upon by His (§ 170, s), there be introduced the qualifier 
cerebri or cerebralis, All these objectionable conditions vanish 
with the adoption of czméza.' Even if this were rejected, ambi- 
guity could be avoided and brevity attained by designating the 
great fibrous masses above mentioned as crura rather than pe- 
dunculi, thus providing for Zvactus cruris transversus, aS sug- 
gested in W. and G., ’89, §57, note. 

§197.—Jsthmus.—Prof. His complains that this word is 
used by mein the sense of Gyrus annectens. This latter term 
does not occur in the German list, so I assume that Gyrus trans- 
wtivus is meant. No one of my terminologic propositions gives 


1That it is not classic Latin, and that it may even have been a corruption of 
cimbra, constitute no bar to its adoption into anatomy. 


2382 JouRNAL OF COMPARATIVE NEUROLOGY. 


me more satisfaction than that of replacing Gyrus annectens, 


bridging convolution, and pli de passage, by isthmus when the cor- 
tical area is visible at the surface, and by vadum when it is con- 
cealed; the occasional interruption of the central fissure is thus 
the /sthmus centrals ; that between the adjoining ends of the 
parietal and paroccipital fissures, the /s¢hmus parocctpitalts, etc. 
So far I cheerfully plead guilty tothe charge. But with what 
justice does Prof. His complain further that this employment 
of zsthmus is in an ‘‘unusual sense’ when his own list contains 
Isthmus gyri forncati? Indeed, even were this complaint well- 
founded, it comes with a poor grace from (a2) a German whose 
-fellow-countryman (Waldeyer) applied (1891) to the nerve-cell 
the term xeuvon, which had been introduced by me (’84) for the 
entire cerebro-spinal axis; from (6) a member of the Nomencla- 
tur Commission whose chairman (KGlliker) applied (1893) to 
the axis-cylinder process of a nerve-cell a term (#ewraxon) practi- 
cally identical with one (euvaxis) which occurs in a standard 
French Medical Dictionary for the cerebro-spinal axis; and.from 
(c) one who himself, upon altogether inadequate grounds," has 
made the term in question, zs¢/wmus, of segmental value, and who 
has needlessly and unjustifiably modified the scope of prosenceph- 
alon and reversed the hitherto commonly accepted sense of 
metencephalon; see Table VII. 
$198. Medipedunculus.—To this term Prof. His devotes 
one-fourth of his entire criticism (§170, ~-2). . Hence some re- 
joinder should be made although the objections impress me as 
either ill-founded in themselves or inconsistent upon the part of 
the objector. Asa word, medipedunculus is no more ‘‘ barbar- 
ous” than medttullium,* Mediterranean, or medieval. As a des- 
ignation rather than a description, it requires definition. The 
beginner would remember medipedunculus quite as easily as 
‘‘pedunculus cerebelli ad pontem”’;' and since experienced 
anatomists know that there are three cerebellar ‘‘stalks” on 
each side but only two ‘‘pedunculi cerebri,’’ one on each side, 


1 This term, by the way, does not occur in the German list, where appar- 
ently it is replaced by brachium pontis. 


Wiper, Neural Terms. 283 


he is not likely to infer that either of the latter is meant by 
medipedunculus. In fact, this term, as coined and defined by 
me,’ is now an idionym, applicable to but a single part of the 
brain. 

$199. In order to be absolutely explicit and independent 
of the context the following terms from the German list should 
be accompanied by the words here bracketed after them :— 
Chvus [occipitahs|; Chvus | sphenoidahs|; Pars cervicalis 
[medullae spinalis |; Sulcus lateralis anterior | medullae oblon- 
gatae |; Sulcus hinitans ventriculorum | encephal |; Pars centralts 
[ ventricule lateralis |; Ventriculus terminalis | medullae spinalrs |; 
Lamina terminalis { encephalr |. The identity of the adjective in 
the last two terms would lead the beginner to associate them 
topographically, and he certainly would never infer that they 
designate parts at opposite poles of the cerebro-spinal axis.’ 

§200. From the standpoint of Prof. His the foregoing 
must be regarded as serious blemishes upon the German list. 
From my point of view, although I might object to certain of 
the names as such, it would not be on account of their lack of 
explicitness. As has been said above (§26) in many instances 
explicitness is to be gained from the context. But with really 
the larger number, I am confident that well selected, brief, and 
fairly suggestive designatory names can and will be learned and 
remembered without any difficulty, especially if the study of the 
brain be begun at an early age; see Part VIII. 

$201. Coelia.—This word, in place of cavitas encephal s. 
ventriculus encephal, is one of the three cited by Prof. His (§170, 
a) as examples of my many terms that are objectionable be- 


1In this connection two remarks are naturally suggested: (1) Medipedun- 
culus is an adjectival locative, it and its correlatives, praepedunculus and fost: 
pedunculus, constituting one of the most perfect groups of that kind ; #29, 189. 
(2) The obtrusively Latin termination of these words, as well as the length of 
the words themselves, forced upon me in 1884 (262) the consideration of the 
whole subject of paronymy. 


2 In the absence of adequate context or prior definition, would any reader 
imagine that spongiocyte and spongioplasm refer to elements of the nervous 
tissue? 


284 JOURNAL OF CoMPARATIVE NEUROLOGY. 


cause they are ‘‘new.”’ In the lexicon ot Liddell and Scott 
xotdta éyxegddov is quoted as in good and regular standing among 
Greek medical writers. According to Burdach (‘‘vom Baue 
und Leben des Gehirns,”’ 1819-1822, II, 301, 378, 380), Galen 
designated the ‘‘fourth ventricle” as xo:déa éxcoStov eyxegahov, 
tetapty xothta and draovla xowhla ( ‘«De usu partium, fap Bric VIEL 
C.XII. p. 170); the ‘third ventricle” as péon tpiry xordéa (idem. 
IX, III, 172); and the ‘‘lateral ventricles” as zpoodéar xordéat 
( ‘‘ De odoratus instrumento,”’ II, 110). Coela is then certainly 
not ‘‘new.’’ Had Prof. His said wxusual, his objection would 
have been more nearly justified by the facts, although in 
recent encephalic literature on both sides of the water com- 
pounds of coe/a are more and more frequently encountered. 

$202. In favor of coeha (English ce/za or cele)’ in place of 
ventriculus may be urged the following: 

I. Its Greek origin renders it compoundable regularly 
and euphoniously with the characteristic prefixes already em- 
ployed in the segmental names, e. g., mesencephalon, etc. 

2. These compounds are mononyms and therefore capa- 
ble (§47) of inflection, (e. g., mesocoeliae), derivation (e. g., 
mesocoeliana), and adoption into other languages without ma- 
terial change; e. g., English, mesocele; French, mesocoele ; 
German, Mesokéle ; Italian, mesocelia. 

3. The various national paronyms thus formed are like- 
wise capable of derivation; e. g., mesocelian. 

4. There is classic authority for the use of coe/a in the 
sense of encephalic cavity ; §2o1. ; 

5. These ancient usages are assumed to be familiar to 
educated anatomists, who therefore should recognize the com- 
pounds with little or no hesitation. 

6. The compounds are so euphonious and so obviously 
correlated with the segmental names as to be learned and re- 


1 As already remarked (W. & G., ’89, 280, note), in an’ English sentence 
the word by itself might be either ambiguous or pedantic and the general term 
cavity commonly answers the purpose ; see 248, and (for the replacement of o¢ 
by ¢) 269. 


Wiper, Neural Terms. 285 


membered easily even by general students, and by those who 
may not have had a classical training.’ 

7. In recent times it has been independently proposed by 
two anatomists, teachers as well as investigators.” 

8. It has been adopted more or less completely by three 
of the older American neurologists, Henry F. Osborn, (82, 
’84, 88), E. C. Spitzka (’81, ’84), and R. Ramsay Wright 
(84, °85), and unreservedly by eight of the younger, W. 
Browning, I. E. Clark, P. A. Fish, | Mrs: S: PB. Gage,» O;.D: 
Humphrey, B. F. Kingsbury, T. B. Stowell, and B. B. Stroud. 

$203. It will be noted that among the advantages of coeka 
over ventriculus is not enumerated its freedom from ambiguity. 
Theoretically, of course, ventriculus (encephal) might be mis- 
taken for ventreculus (cardiae s. corde). Practically, however, 
the context would almost infallibly obviate misapprehension. * 
Hence from my point of view, the absolute unambiguity of 
coca and its compounds would not in itself justify its replace- 
ment of ventriculus. It would be a causa vera, but hardly a 
causa sufficcens. 

$204. The concluding remark of Professor His may be 
said to ‘‘cap the climax” of his ill-founded criticism. The 
characterizations, ‘‘vollig neuen” and ‘‘grossentheils recht 
fremdartig Klingenden,”’ could hardly have been more sweeping 


1 Among the hundreds of such students at Cornell University and at the 
Medical School of Maine who have gained their practical and theoretic knowl- 
edge of encephalic morphology by means of these compounds no special diffi- 
culty has ever been experienced. 


2 My propositions first appeared in the paper, ’81, 4, March Ig and 26, 
1881. On the fifteenth of August, 1882, Prof. T. Jeffery Parker read be- 
fore the the Otago Institute of New Zealand a paper (’82) in which mesocoele 
and similar compounds were introduced, although he was evidently quite una- 
ware of my prior publication. The terms were also employed in his ‘* Zootomy”’ 
(’84) and in a later paper (86). 


8My previous reference (2199) to the polyonymic derivative, sudcus limitans 
ventriculorum, was not for the sake of demonstrating the ambiguity of that term 
but to illustrate the inconsistency of the implied demand of Professor His (3170) 
that all terms must be self-explanatory and require no definition. 


286 JOURNAL OF CoMPARATIVE NEUROLOGY. 


had I proposed to replace Latin by Choctaw.’ Any anatomist, 
unprejudiced, and not above conceding the possibility that some 
good thing may come out of the American Nazareth, who will 
candidly compare the terms in Table VI (Part VII) will admit 
that in the second column a comparatively small number are 
new in the strict sense of the word, and that the large majority 
are either identical with those in the first, or differ therefrom 
merely in the omission of useless words, or in the replacement 
of adjectives by prefixes of like signification.? 

— $205. Comments and criticisms by Prof. A. von Kolliker. 
Early in 1892, about a year after the date of the first letters 
from the secretary of the German committee (§138) its chair- 
man spoke as follows (‘‘ Nervenzellen und Nervenfasern,” Bzo- 
log. Centralblatt, XII, 36, Jan. 30, 1892): 

**So haben amerikanische Gelehrte im letzten Jahre einen Ver- 
such zur Verbesserung einiger Teile der anatomischen Nomenklatur 
unternommen, und hoffen wir, dass dieselben unserer Einladung zu 
gemeinsamer Arbeit entgegenkommen werden. Sollte es gelingen, 
wenigstens die lateinischen Namen in den morphologischen Wissen- 


schaften zu einem Gemeingute aller zu machen, so ware hiermit offen- 
bar ein grosses Ziel erreicht.” 


$206. Following is a translation of the above: 


‘‘ During the past year American scientists have made an effort 
toward the improvement of some parts of Anatomic Nomenclature. 
We hope they will look favorably upon our invitation to join us in our 
work. Should we succeed at least in making the Latin names in the 
morphologic sciences common property, a great end would thereby be 
attained.” F 


$206. The foregoing implies that the codperation of 
American anatomists not only was desired but had been asked. 
I am not aware that other members of the American commit- 


1At that time, although my principal article on terminology had not been 
read by Professor His (see Part VI), the lists of terms preferred by me were in 
his hands (?140, note), so that no claim can be entertained that he referred 
merely to what he assumed my proposals ‘‘ tended ”’ to bring about. 


2Just as this goes to the printer (Nov. 13, 1896) there reach me several re- 
prints of the note by Professor His in the Amat, Anszeiger (XII, 446-448, Novem- 
ber, 1896) entitled “‘Herr Burt Wilder und die Anatomische Nomenclatur.” 
His remarks will be considered in connection with our correspondence, Part VI. 


WILDER, WVeural Terms. 287 


tees corresponded with members of the German committee upon 
the subject of anatomic nomenclature, and the request of Dr. 
Krause for our reports and for my papers could hardly be inter- 
preted as an official invitation to cooperate; see $137. 

$207. So faras I am aware, the only other utterance of 
Prof. Kolliker upon the subject constitutes a paragraph on p. 
814 of the second volume of his ‘‘Gewebelehre”’ in the part 
which was published, I believe, in the spring of 1896. It is as 
follows: 


‘‘Die Nomenklatur anlangend, habe ich mich selbstverstandlich in 
diesem §, wie in dem ganzen Buche, an die bekannten Beschliisse der 
Kommission der anatomischen Gesellschaft und den Vomenclator ana- 
tomicus derselben gehalten. Die in den letzten Jahren von Amerika 
ausgehende anatomische Nomenklatur halte ich fiir vollkommen ver. 
fehlt und fiir so unverwendbar, dass es mir unmdglich ist, Abhandlun- 
gen zu lesen, denen dieselbe zu Grunde gelegt ist. Man kann doch 
von einem Gelehrten, der eine ordentliche Schulbildung genossen hat, 
nicht verlangen, dass er die vielen Barbarismen dieser Nomenklatur, 
wie metatela, metaplexus, auliplexes, diaplexus, ectocinerea, cephalad, caud- 
ad, dorsad, cephalo-dorsad, ventro-caudad, dorso-caudad, hemtcerebrum u. 
s. w. ruhig annehme und bei einer grossen Anzahl anderer Worte, wie 
terma, proton, pero, prosoterma, dtaterma, supraplexus, aula, alba, crista, 
diacoele, mesocoele u. s. w. erst hingehe und lerne, was dieselben 
bedeuten sollen. Als altester deutscher Anatom wird es mir wohl er- 
laubt sein, den amerikanischen Kollegen den Rath zu geben, auf die- 
ser Bahn nicht weiter fortzuschreiten, sonst wiirde es im Laufe von 
einigen Jahren dazu kommen, dass man sich hiiben und driiben nicht 
mehr verstiinde und jeder wissenschaftliche Verkehr unmédglich 
wiirde.” 

§208. Following is the translation: 


‘*As regards nomenclature I have of course maintained in this sec- 
tion, as in the whole book, the known conclusions of the committee 
of the anatomische Gesellschaft and the omenclator anatomtcus there- 
of. The anatomic nomenclature coming from America in recent 
years, I regard as a complete failure, and as unavailable, because it is 
impossible for me to read articles based thereon. One cannot ask of 
a scholar who has received an accurate education that he accept 
quietly the many barbarisms of this nomenclature, such as metatela, 
metaplexus, auliplexus, diaplexus, ectocinerea, cephalad, caudad, dorsad, 
cephalo-dorsad, ventro-caudad, dorso-caudad, hemicerebrum, etc. and with 
a great number of other words like ¢ervma, proton, pero, prosoterma, 
diaterma, supraplexus, aula, alba, crista, diacoele, mesocoele, etc. first go 
to work to learn what they mean. As the oldest German anatomist, 
it may be permitted me to advise our American colleagues not to con- 
tinue farther in this direction lest, in the course of a few years, it come 


288 JouRNAL oF CoMPARATIVE NEUROLOGY. 


about that we cannot understand each other in the two countries, and 
hence all scientific intercourse will become impossible.” 


§209. The age of Prof. Kolliker, his magnificent services 
in the advancement and diffusion of science, and the evident 
sincerity of his regret at American departures from what he re- 
gards as terminologic rectitude, all demand serious rejoinder. 

$210. Among the terms that one must ‘‘ go to work to 
learn what they mean,” c/zsta ( fornicis ) designates a feature of 
the brain which, so far as I know, was first described by me in 
1880 (’80, g; W. & G., 82, §1214; Mrs. Gage, ’93, 283-284); 
as its discoverer, I may be permitted to assign ita name. evo 
was proposed by me (’81, 0) for the soft ectal layer of the 
Bulbus olfactorius which, in hardened animal brains, often peels 
off the firmer ental ‘‘ core”’ like a boot; but the word is seldom 
needed. If the ¢erma ($194) pertains to two encephalic seg- | 
ments, dzaterma and prosoterma seem to be both appropriate and in- 
telligible. JZesocoele and diacoele (Latin, mesocoelia and diacoela; 
English, mesocele and dracele) have been discussed directly or 
indirectly above (§201-203). Swpraplexus, (introduced, I think 
by Mrs. Gage ) would certainly be supposed to indicate a plexus 
in the roof of some part of the brain cavity. Alba could 
hardly suggest anything else than the substantia alba of the Ger- 
man list; see $116. 

S211. Aula.—After years of confusion, doubt, and even 
distress of mind, induced by the failure to reconcile the facts of 
development and comparative anatomy with the prevalent 
nomenclature of the brain, in 1880 (’80, d, ¢, f; ’81, 6, d) I 
proposed av/a upon grounds formulated two years later as fol- 
lows (W. and G., ’82, $1065) :— 


(1) ‘*To substitute brief single words for the phrases, ‘‘ventricu- 
lus communis,” ‘‘ventriculus lobi communis,” mesal part of the 
‘*common ventricular cavity,” ‘‘ foramen Monroi,” ete. 


(2) Because the phrase most commonly employed, /oramen 
Monrot, is used to designate at least three different cavities or orifices: 
(a2) The cavity by which either paracoelia [‘‘ lateral ventricle ”’] com- 
municates with the mesal series of cavities; (4) The two lateral ori- 
fices together with the intervening space; (c) The mesal [ cephalic ] 
orifice of the diacoelia. We have been unable to ascertain by whom 


¢ 


WILpDER, Neural Terms. 289 


the phrase was first employed, and the description by Monro secund- 
us (1783), in whose honor it was applied, is somewhat vague. 

(3) In order to indicate our opinion of the desirability of 
recognizing the aula as morphologically an important element of the 
series of encephalic cavities.”! 


$212. Proton.—This neuter noun was used by me (’93, a, 
$46, note) to designate the comparatively undifferentiated mass 
in which two or more parts might afterward be distinguishable. 
It is free from certain obvious and by no means inconsiderable 
objections that may be brought against Azlage and fundament 
as English words. It is subject to inflection, and may be 
adopted into any language. In many derivatives or compounds 
it is associated in the minds of all educated persons with the 
general idea of primitiveness. Its employment is in harmony 
with the following phrases from Aristotle cited for me by Prof. 
B. I. Wheeler:—t0 zp@tov; 4 xewty BAn; 4 cpwTy aitia, 

$213. In short, all my regrets for the errors already con- 
fessed (§ 71) and for others of which I may be convicted,  to- 
gether with all my doubts regarding certain of the terms not as 
yet acted upon by the American committees, shrink into the 
background of my mind as I reflect upon the nature and signifi- 
cance of aw/a and proton, and upon the advantages that have 
been and may be gained from their employment. 

$214. In the foregoing extract (§207) I suppose ‘‘Barbar- 
ismen’”’ means verba hybrida, i. e., words formed by the con- 
junction of Latin and Greek elements. If so, one of the six 
specified organonyms, viz., auliplexus, must be exempted from 
the stigma since aula, although derived from the Greek aij 
is a Latin word in good standing. To the remaining five I add 
five other neural mononymic hybrids for which I am respons- 
ible. Ina parallel column of Table IV are given the equivalents 
either as in the German list or as they would be regularly form- 
ed. In another (the first) column are placed an equal num- 
ber of hybrid words selected from the German list, and hence, 
I infer, sanctioned by the President of the Commission. With 


1 With some of the lower vertebrates (e. g., Chémaera, ’77, a,) the aula is 
much more extensive than either of the ‘‘lateral ventricles” with which it is 
connected through the two portae. 


290 JOURNAL OF COMPARATIVE NEUROLOGY. 


both groups I have italicized the Greek element which (so far as 
appears in a standard lexicon) failed to be adopted (by paronymy) 
into classic Latin. Of course the abstract etymologic immor- 
ality involved in the two sets of irregular connections is the 
same. The offspring of the second set surely compare favor- 
ably with the first in point of comeliness,' and their utility as 
savers of labor and time may be estimated by comparison with 
their polyonymic equivalents in the third column. 


TABLE ILy. 

Hybrid words con-|| Hybrid words for| Polyonymic equivalents of the words in 
tained in the Ger-|| which thewriter| the second column, mostly from the Ger- 
man List. zs responsrble, man List. 

Lpidurale Metatela “Lamina chorioidea epithelialis”’ [ven- 

triculi quarti] 

Mesovaricus Diatela ‘Lamina chorioidea epithelialis ”’ [ven- 

triculi tertii] 

Parumbilicales Paratela ‘* Lamina chorioidea epithelialis ” [ven- 

triculi lateralis] 

Parolfactorius Metaplexus ‘¢ Plexus chorioideus ventriculi quarti” 

Perichoriotdiale Diaplexus ‘a $s os tertii ”’ 

Suprachorozdea Paraplexus ss aS ee lateralis” 

Choriocapillaris Ectocinerea *¢ Substantia corticalis” 

Prerygopalatinus Entocinerea “¢Stratum griseum centrale ”’ 

Prerygomandibularis|| Zemzcerebrum |Dimidium cerebri laterale 

Phrentcocostalis Hemiseptum Dimidium septi pellucidi laterale 


$215. The reasonable view of hybrid terms seems to me to 
be embodied in the following remark of Barclay ('03): 


‘‘Notwithstanding the opprobrium attached by some to certain con- 
nections and intermarriages among harmless vocables, I should be in- 
clined not to reject the codperation of the two languages (Greek and 
Latin) where experience shows it to be convenient, useful or neces- 
sary.” 

Abstractly, we may all prefer horses to mules, but this need 
not hinder us from recognizing that, under certain circumstances, 
the latter are more efficient than the former, and that, ina given 
case, a horse may not be even so handsome as a mule. 

$216. The verdict of Prof. Kolliker that the nomenclature 


coming from America in recent years is a ‘‘complete failure”’ 


'The first four from the German list might have been replaced from the same 
source by the less acceptable ‘‘sphenopalatinum,”’ ‘‘sphenodccipitalis,” ‘‘occi- 
pitomastoidea” and ‘‘squamosomastoidea.”’ 


Wiper, Neural Terms. 291 


because he cannot read the articles based thereon, approximates 
what has been called ‘‘the erection of the limitations of one’s 
individual experience into objective laws of the universe.” I 
sincerely trust that he may some day concede the validity of 
these two propositions:—(1) A considerable number of investi- 
gators and advanced instructors on both sides of the ocean have 
employed the ‘‘American” system more or less systematically. 
(2) Judging from my own experience as learner and teacher, 
the hundreds of students, general and special, upon whom that 
system has been practised since 1880, have either saved so much 
time, or gained so much more information within a given time, 
as to make its employment ‘‘worth while’ even when the later 
environment proved unfavorable to its permanent use. 

$217. In concluding this response to the criticism of ‘‘the 
oldest German anatomist’’, I venture to call his attention to the 
different reception accorded my plans for terminologic simplifi- 
cation by two other anatomic teachers well advanced in years, 
viz., Joseph Leidy (§ 56, note) and Oliver Wendell Holmes 
($79). In order also that I may not appear unmindful of the 
fact that the assimilation of verbal novelties becomes less easy 
with increasing age,’ I reproduce the concluding paragraph of 
my second paper upon the subject (’81, 0): 

‘‘The beginner can learn the new terms even more easily than the 
eld, and at any rate he has nothing to forget. But the trained ana- 
tomist shrinks from an unfamiliar word as from an unworn boot; the 
trials of his own pupilage are but vaguely remembered; each day there 
seems more to be done, and less time in which to do it; nor is it to be 
expected that he will be attracted spontaneously toward the consider- 
ation that his own personal convenience and preferences. and even 
those of all his distinguished contemporaries, should be held of little 


moment as compared with the advantages which reform may insure to 
the vastly more numerous anatomical workers of the future.” 


1The argumentum ad hominem is ungracious at the best, and the occasions for 
its employment in this paper have been too numerous already. But when I re- 
call the delay and mystification inflicted upon me and my students by the va- 
riety and heterogeneity of terms, Latin and vernacular. with which most German 
treatises upon encephalic anatomy literally bristle (22 58-60, 169, note), I can- 
not but feel that, however sincere may be the repentance therefor among the 
anatomists of that nation, the needed reform should have been practised for a 
somewhat longer period before others were rebuked. 


292 JOURNAL OF COMPARATIVE NEUROLOGY. 


§218. Under the limitations of time, space and compe- 
tence, I have now responded to the criticisms of the Anatom- 
ische Gesellschaft, the German committee, and three individual 
members thereof. However unfounded, unwise, or even unjust 
some of those criticisms may appear to me, I am disposed to 
believe that they were prompted by a wish to further what the 
critics regard as the best interests of anatomic science. They 
will, therefore, I trust, credit me with a similar motive in urging 
upon them as anatomists, and especially as German anatomists, 
the earnest consideration of one of the topics discussed in the 
earlier portion of this paper, viz., Paronymy as opposed to Het- 
eronymy (S$ 43,85, e). 

§ 219. Unfortunately, upon this point (which seemed to 
me in 1885 so important that it was the subject of my address 
as president of the American Neurological Association ’85, c) I 
find myself unable as yet to determine what view is held by the 
German committee. Its secretary has declared ($159, 4) ‘‘that 
each nation can translate the Latin (international) name after its 
own fashion ;” also (§ 158, 1) that German designations [hetero- 
nyms] in so far as they are not already established, or so far as 
they are really necessary, are left to the free choice of the indi- 
vidual. 

$220. Whatever be the real or intended purport of the 
foregoing, the latest publications of the chairman of the commis- 
sion (‘‘Gewebelehre’’), and of the member who has most fully 
discussed the subject, contain many and even diverse German 
vernacular equivalents of the Latin terms recommended by 
them. The article of Prof. His ('95, @) yields at least a score of 
such which have no etymologic relation with the Latin terms. 
It seems to me that, with the exception of Gezrn and possibly 
a few others (§ 48), the systematic employment of paronyms 
(§ 46) would greatly facilitate the comprehension of German 
writings by anatomists of other nationalities, and especially by 
students, without materially embarassing the Germans them- 
selves. 


Some of the Latin 
terms adopted by 
the Amer. Neurol. 


Wiper, Weural Terms. 293 


$221. The feasibility of this method was exemplified in 
some degree in 1889 upon the ‘‘Table of Paronyms” (W. & 


Asssoctation. 


I 
2 


5 
4 
5 
8 
9 
) Ke) 
II 
12 
13 
14 


25 


16|F, hippocampi 


17 


18 
Ig 


20 
21 
22 
=) 


24 
25 
26 
27 


28 
29 
30 
31 
32 
33 
34 
35 
36 


Calcar 
Callosum 
Chiasma 
Claustrum 
Clava 
Cuneus 
Dentatum 
Dura 
Epiphysis 
Falx 

F. calcarina 
F. centralis 
F. collateralis 


Fornix 


Hippocampus 
Hy pophysis 


Infundibulum 
Insula 
Lemniscus 


Mesencephalon 


Monticulus 
Oblongata 
Oliva 
Operculum 


Pallium 

Pia 

Pons 
Praecuneus 
Pulvinar 
Striatum 
Tegmentum 
Tentorium 
Thalamus 


TABLE Y. 
Examples of the use of these paronyms, 
German paronyms. or of analogous ones, by German 
writers. 
Kalkar Kalkig 
Kallosum s Callosum 
Chiasma ‘‘Chiasmas” (Onufrowics, ’87, 12) 
Claustrum “‘Cerebellumgebiet” (His, ’80, 26) 
Clave Olive 
Cuneus 
Dentatum ‘‘Ablegat.” 
Dura s. Dur 
Epiphyse Krause, 779, 780 
Falx 
Kalkarinfissur 
Centralfissur “‘Centralcanal”’? (Henle, ’79 231.) 
Collateral fissur “‘Collateralfurche’”’ (Wernicke, ’82, I, 
I 
Hippokampalfissur 
Fornix ‘‘Fornixtaenien’’(His, ’95, 167) ‘‘For- 
nixschenkel”’ (Schwalbe, ’81, 464) 
Hippokamp 
fAlypophyse ‘‘Hypophysenblase” (Schwalbe, ’81, 
477) 
Infundibulum eC crabellmpebeie? (His, ’80, 26) 
Insel (Henle, ’79, 170) 
Lemniscus 


Mesenkephal s. Mesen-/‘Rhinencephalons”’ (His, ’95, 175) 
cephalon 


Montikel “‘Follikel’”’ (Schwalbe, ’81, 473) 

Oblongat Advocat 

Olive ‘‘Nebenolive” (Schwalbe, ’81, 615) 

Operculum “des Operculums,” Jen. Zettschr., 
XXIX, 25 

Pallium “des Spleniums” 

Pia ‘‘Piaplatte’’ (Edinger ? ?) 

Pons ‘“‘Ponsfasern” (Edinger ’89, 153) 

Pracuneus 

Pulvinar 

Striatum s. Striat ‘‘Ablegat”’ 

Tegmentum s. Tegment |«*«Moment”’ 

Tentorium 

Thalamus “Thalamuskern” (Schwalbe, ’81, 711) 


G., ’89, 530-531). Inow venture to offer in Table V what 
seem to me appropriate German paronyms of most of the Latin 
terms adopted by the American Neurological Association ($80). 
So far as possible either the actual paronyms, or analogous pre- 


294 JoURNAL OF COMPARATIVE NEUROLOGY. 


cedents therefor, have been’ selected from German writings. In 
the other cases the suggestions may not be always well-founded. 


Part VI. Correspondence with Professor Wilhelm Fits. 


§222. The general occasion for the existence of this Part 
was stated in the Introduction, §4. The hope, entertained at 
the time that was written and expressed at a later period ($171), 
that Prof. His might publicly correct all his errors of omission 
and commission, has not been realized in his article (just re- 
ceived in the Avatomischer Anzeiger, XII, 446-448, Nov., 1896, 
hence the correspondence is most regretfully submitted. The 
only changes consist in the omission of unessential paragraphs, 
and in the addition of notes, or of words in brackets. 

§223. From the writer to Prof. His, Dec. 3, 1895. 


Prof. W. His, Dear Sir:—Not until a week ago was able I to read 
your article in the Archiv f. Anatomie etc., 1895, ‘* Die anatomische 
Nomenclatur,” wherein, on pages 6 and 7, you do me the honor to 
discuss my views especially with reference to encephalic terms. 

It is my intention to present the subject at the coming meeting of 
the Association of American Anatomists, on the 26th of this month, 
and before doing so I desire to obtain from you answers to the sub- 
joined queries. It will be a favor to me if the answers, or as many | 
as possible of them, can reach me before the meeting, and it will be 
an advantage to our discussions if you permit them to be made 
public. 

Regretting to differ with you, and to ask you to perhaps incon- 
venience yourself in order to reach me in season, I have the honor to 
remain, Very respectfully yours, 

Burt G. WILDER. 


1. I see no reference to the writings of Richard Owen or Pye. 
Smith. The latter (Jour. of Anat. and Physiology, Oct. 1877) pub- 
lished an article of 22 pages and insisted upon the sufficiency of thala- 
mus which you concede. The former introduced fostcava and precava 
and (‘‘ Anat of Vertebrates,’’? III p. 136) gives a list of fissures, all 
the names being mononyms and some (cal/osal, supercallosal and sub- 
Jrontal) apparently unobjectionable. Why were these not included in 
the column of synonyms by ‘‘ various authors” [in the protocols of 
the committee ? §178, 3]. 

2. Excepting some compounds of excephalon, nearly all the mono- 
nyms upon yourlist of encephalic terms, about twenty-five in num- 
ber, had already been selected and adopted by me. Was not this 


Witpver, WVeural Terms. 295 


coincidence worth mention as indicating the possibility of some degree 
of harmonious codperation between us? 

3. You say ‘‘ Wilder und seine Collegen verlangen lauter Mon- 
onyme.” This is correct [but only in the sense of. prefer; see $172]. 
You then add, ‘‘Sie sagen, z.B. praecornu, pdstcornu und fpostcava.” 
These words are used by me, but not, so far as I know, by other 
members of the American Committees,! although they recommend the 
employment of CALCAR for Atppocampus minor, HIPPOCAMPUS for A. 
major, PONS for pons Varolit, INSULA for insula Retlii, and PIA and DURA 
for pia mater and dura mater respectively. 

4. You refer to the principle of mononymy. But you do not 
seem to have gathered, even from my ‘‘ Paronymy versus Heterony- 
my as Neuronymic Principles” [85, ¢], that mononyms are preferred 
by us to polyonyms not so much because they are usually shorter, but 
because, whatever their length, they are capable of two desirable 
modifications, vz., (a) inflection as adjectives (e g., thalamicus, callos- 
alts, duralis, etc.) and (4) adoption by paronymy into other languages, 
(e g, hippocampus, hippocampe, hippocamp, Hippokamp, hippocampo). 
Am I to infer that this feature of the matter was unknown to you, 
or regarded as slight in importance ? 

5. You say ‘‘Sprachwidrige Wortzusammensetzung enthalt aber 
Wilder’s Liste sehr viele”? It would not be without probability or 
precedent that errors should occur among so large a number of terms, 
but I must insist upon the specification of my ungrammatic verbal 
combinations In particular I ask fuller grounds of objection to med- 
ipedunculus |S198] 

6. You mention certain papers by me [as ‘‘eine Reihe von 
kleineren Aufsatzen und Broschiiren” ; $170, d, note 2]. Their few- 
ness [four], their brevity, and their recent dates [1890-1892] would in- 
dicate that T had done little on the subject and that my views are cor- 
respondingly unimportant. Yet my first paper on [encephalic] Nom- 
enclature was in 1880 and I have published something almost annual- 
ly since upon it. The article |W. and G, ’89] in the ‘‘ Reference 
Handbook ” [of the Medical Sciences, VIII, 515-533, 1889] was, so 
far as I know, the fullest discussion of late years; a copy was sent to 
Dr. Krause [secretary of your committee] about May, 1891, and his 
letter of April t2yarg92, informed me that it had been ‘‘set in circu- 
lation among the Committee.” It is also mentioned in most of my 
papers or documents printed since 1889. In 1892, the American As- 
sociation for the Advancement of Science adopted unanimously the 
Report of the Committee (of which I am not a member) on Biological 


1As stated in the Introduction, #3, I desire to free others from responsi- 
bilities which they have not assumed. So far as may be inferred from the entire 
absence of later reference to this point upon the part of Professor His, an alle- 
gation, absolutely unfounded and casting upon several American anatomists what 
they may regard as serious discredit, may be publicly made and neither sub- 
stantiated nor withdrawn. 


296 JouRNAL OF COMPARATIVE NEUROLOGY. 


Nomenclature [§85, 7, and note? ], including a characterization of the 
Handbook article ‘‘ as representing an epitome of the whole subject, 
with suggestions for future progress.” A copy of that Report was 
sent to you. Is your absolute silence as to the article to be interpreted 
as indicating that you not only never saw it, but never heard of it, or 
had no idea of its scope? 


§224. From Professor His to the writer, Dec. 15, 1895. 


SEHR GEEHRTER HERR CoLiecE: Ihr Brief vom 3d M. mit den 6 
fragen trifft erst heute bei mir ein, und es ist mir zweifelhaft, ob meine 
Antwort bis zum 26 in Ihren Handen sein kann. Ohne in Einzeln- 
heiten einzutreten, was wohl ziemlich nutzlos sein wiirde, hebe ich 
hervor, dass es sich in dieser Sache nicht um eine persdnliche Angel- 
egenheit handelt, da die anatomische Gesellschaft als solche zu Ihren 
Bestrebungen Stellung genommen hat. Sie finden die Erklarung der 
anatomischen Gesellschaft im Bericht [etc.; see this article $147]. 


Hochachtungsvoll Ihr ergebener, W, His: 
§225. Translation of the above letter from Professor His, 
Deew15, 1895. 


ESTEEMED COLLEAGUE: Yours of the 3d containing six queries 
was received to-day and I doubt whether my answer can reach you for 
the 26th. Without entering into details, which would very likely be 
useless, I point out that this is not a personal matter, since the Ana- 
tomische Gesellschaft as such is opposed to your efforts. You will 
find its declaration in the Report, etc. [see $147 of this article]. 

Respectfully yours, W. Hus: 


§226. From the writer to Professor His, Jan. 3, 1896. 


ProFessor His, DEAR SiR: Your favor of Dec. 15th reached 
Philadelphia after the close of the last session of the Association of 
American Anatomists. J infer that you had no objection to tts publica- 
tion. Pray accept my thanks for your promptness, and for the ref- 
erences to the published opinion of your committee. This I was ac- 
quainted with. It is practically a declaration of conservatism which 
I hope to find time to comment upon. I conclude that you regard 
the queries in my communication of Dec. 3, 1895, as covered by the 
committee’s opinion. Whatever view might be taken as to the first 
four, I feel justified in calling upon you personally for explicit re- 
sponses to the last two, vz.:—What ‘‘ ungrammatic verbal combina- 
tions” occur in my recent lists? What explanation have you for in- 
timating that my first publication on Anatomic Nomenclature was 
dated in 188g instead of 1880, and for ignoring the article ‘“‘ Anat. 
Terminology” in the ‘‘ Reference Handbook of the Medical Sciences,” 
1889 ?P 

The immediate question is, What is due to those continental 
anatomists who, in the absence of my lists of terms or of the publica- 
tions by myself and other Americans who have more or less complete- 


WILpER, Weural Terms. 297 


ly adopted them, may have been unduly influenced by the absence of 
those terms from the Committee’s lists of synonyms, and by the seri- 
ous omissions and errors embodied, however unintentionally, in your 
article ? 


It seems to me not too much to ask of you to supply the oppor- 
tunity for independent comparison and judgment by reproducing in 
your Archiv the accompanying List of Neurologic and Vasal Terms 
adopted by your Committee, the terms preferred by me in parallel 
columns, the accompanying commentaries by me and such as you may 
think best to add, and the correspondence of which this forms a part. 

I have the honor to remain, Very respectfully, 

Your obedient servant, Burt G. WILDER. 

§227. The further correspondence between Professor His 
and myself was as follows :—Second letter from him, Feb. 29, 
1896. Third from me, Aug. 11. Third from him, Aug. 27. 
Fourth from me, Sept. 28. Although my third contained the 
statement that I was editing our correspondence for publication, 
and the third from Professor His offered no objection thereto, 
after consultation with the editors of this journal it has been 
deemed best to summarize the later letters, and to include the 
article (His, ’96) already referred to (§222). As indicated at 
the outset of my second letter (Jan. 3, 1896; $226) I felt that 
he had tacitly granted my request for permission to publish his 
first letter (§224). 

§228. At this stage I should like it understood that, in 
originally addressing Professor His directly rather than in the 
columns of a journal, I had two motives, vzz., Furst, the belief 
that the impending discussion of Anatomic Nomenclature by 
the Association of American Anatomists would be facilitated 
by the information sought. Secondly, the sincere desire to avoid 
a public controversy by affording to one whom I regarded as 
mistaken the opportunity to modify his statements voluntarily. 
This desire was in accordance with the sentiment, long enter- 
tained, but first expressed six years ago in this journal (’91, 4, 
201-202) that, ‘‘since everyone makes mistakes, the interests of 
all concerned would be best subserved by the adoption of the 
custom of each correcting his own, either as soon as discovered 
or periodically ; a sort of scientific confession of sins. The 
natural corollary to this would be that each well-disposed dis- 
coverer of another’s faults would inform him privately so that 
he might make prompt correction. This plan I have followed in 
several cases, and have reason to believe it has served to avoid 
personal irritation and the needless repetition of criticism.’” 


1 The lamentable failure of the plan in the present case does not lessen my 
confidence in its essential soundness. 


298 JOURNAL OF COMPARATIVE NEUROLOGY. 


§229. The second letter of Professor His contained no re- 
plies to my questions, and evinced no disposition to discuss the 
subject publicly ; on the contrary it declared that he had given 
up all connection with nomenclature, and preferred to devote 
his time to other studies. 

$230. In order to appreciate the situation from my stand- 
point there should be borne in mind certain facts, and certain 
propositions that are unlikely to be contested:—(a@) Pro- 
fessor His not only originated the German movement for 
a revised nomenclature, but was a member of the special 
committee of three on the ‘‘ Redactionsausschuss ;” (0) he 
had set forth the aims, methods and results of the entire 
committee in an extended article ; (c) having assumed such re- 
sponsibilities, it was his duty to ascertain what had been done 
already ; (2) had my own labors been wholly ignored I should 
have contented myself with my associates in this respect, Bar- 
clay, Whewell, Owen, Pye-Smith, and others; (e) but, in con- 
nection with a somewhat extended animadversion upon my 
views, what purported to be a list of my ‘‘chief publications” 
upon nomenclature omitted that which was most comprehen- 
sive; (/) under date of April 12, 1892 ($142), the secretary 
of the German committee had assured me that a copy of my 
principal article had been set in circulation among the members 
of the committee ; (g) inquiries dated Dec. 1, 1895 and July 
10, 1896, as to whether it had actually been transmitted to Pro- 
fessor His, remained unanswered by the secretary; (/) the last 
of the six inquiries accompanying my first letter (§223, 6) might ~ 
have been answered at once and ina single word; (2) there 
would then have remained merely the explanation of his non- 
acquaintance with the article in question. 

§231. On the eleventh of August, after waiting. five 
months in the hope that reflection might convince Professor 
His that his previous letters did not meet the fundamental require- 
ments of the situation, I reminded him that the six points at 
issue between us fell into two distinct categories ; that points I, 
2 and 4 (indicated by the sections so numbered in my first 
letter) called for the exercise of a certain amount of profess- 
ional courtesy, and that he was clearly within his rights in de- 
clining to take the time required for their elucidation; but that 
the other three had no necessary connection with nomenclature, 
and that courtesy need not be invoked in dealing with them. 
He had brought a general indictment, wzz., as to the existence 
of ‘‘ many ungrammatic verbal combinations in my lists” (§223, 


WILDER, WVeural Terms. 299 


5), and he was bound to furnish specifications. He had uttered 
a charge (‘‘ Wilder und seine Collegen verlangen lauter Mono- 
nyme,’’ §§172, 223, 3) which was either ambiguous or unfounded, 
and he was bound to explain or withdraw it. While undertak- 
ing to enumerate my ‘‘ hauptsachlichste Schrifte,” he had failed 
to mention the principal one, and he was bound to account for 
either his ignorance of the facts or for his failure to state them. 

§232. To this Professor Hisreplied under date of Aug. 
27. His sentiments are expressed in his article (His, ’96) 
of the same date published two months later. The original is 
readily accessible, but a translation is here appended. 


‘“« Mr. Burt Wilder and Anatomic Nomenclature. 
By Wilhelm His. 


Some time after the publication of the Nomenclature adopt- 
ed by the Anatomical society at Basel, Professor Burt Wilder 
of Ithaca, N. Y., U. S. A., complained to me by letter concern- 
ing the manner in which his own efforts at simplification of ana- 
tomic nomenclature had been mentioned in the introduction to 
the BNA” Mr. Wilder clothed his complaints in the form 
of personal questions and furthermore expressed the desire to 
publish my answers at the same time with his questions. This 
is a somewhat inquisitorial proceeding, unusual hitherto in in- 
tercourse among scholars. In the present case, also, the 
question is not a personal difference between Mr. Wilder and 
myself but concerns matters that have had their formal settle- 
ment through the transactions at Basel. Moreover, the ana- 
tomical society has stated its views respecting the efforts of the 
American Nomenclature committee in a special declaration, 
signed by all the members of the commission’ and by the Pres- 


1 With the exception of this article by Professor His, all of the translatiuns 
of German extracts and letters in the present paper have been kindly revised by” 
my friend Fraulein Berthe J. Bartelmann, Instructor in German at Smith Col- 
lege, Northampton, Mass. 


? Explained in a note upon the reverse of the title leaf of His, ’95:—‘‘ Als 
abgekiirtze Bezeichnung der in Basel angenommenen Nomina anatomica wird 
vorgeschlagen [ BNA]’’. 


3 Proceedings of the Anatomische Gesellschaft, at the ninth meeting at Basel, 
edited by K. v. Bardeleben, Jena, 1895, p. 162. [It is doubtless through an 
oversight that this statement is not here qualified by the phrase, ‘‘ present at 
Basel,” as in the report quoted in 3147. Of the eighteen members of the com- 
mittee, nine were then present ]. 


300 JOURNAL OF COMPARATIVE NEUROLOGY. 


ident of the Society. In concert with my colleagues on the 
editorial committee, the Messrs. Waldeyer and W. Krause, I 
have refused consequently to answer the separate questions of 
Mr. Wilder, and have referred him to the above mentioned 
declaration. Likewise, I had to refuse a later and unreasonable 
demand to print in the Archiv fur Anatomie u. Physiol. his brain- 
terms together with those of the BNA. My colleagues and 
I were of the opinion that such a proceeding is not incumbent 
upon me, and that Mr. Wilder should be asked to present his 
amendments to the special revision committee of the Anatom- 
ical Society. 

‘¢A letter just received from Mr. Wilder, very objection- 
able in its form, shows me however that he believes himself to 
have been injured by me through the intentional ignoring of 
his earlier publications reaching back to 1880, as well as his 
more comprehensive article, ‘‘ Anatomical Terminology,” in 
the ‘‘Reference Handbook of the Medical Sciences,” VIII, 
T8890, p. 515-533.” 

‘¢To that I feel it my duty to offer an explanation. I cited 
such of Mr. Wilder’s papers, with the qualification as the most 
important, as at that time were at hand. The ‘‘ Reference 
Handbook,”’ little known in Germany, has never been in my 
hands, and by mischance the separate proof-sheets of the article 
referred to, which Mr. Wilder had sent to Mr. Krause in the 
year 1891 with a view to the information of our committee at 
that time, has not reached me. It was, Mr. Krause informs me, 
lost by one of the members. Indeed, I was not aware of the 
existence of this treatise when I wrote the introduction to the 
BNA., and an intentional neglect of the same is out of 
the question. 

‘‘In order however to avoid all reproach of injustice to- 
ward Mr. Wilder I present herewith his questions in the form 
revised by him [ $223, 1-6]. Each reader has then the basis 
for independent judgment. 

Saasgrund, Ct. Wallis, Schweiz, 27 August, 1896.” 

§233. To the letter written at the same time as the fore- 
going article I replied (Sept. 28) that the plainess and peremp- 
toriness of my previous communication might have been avoided 
had my earlier letters received due consideration. There has 
emanated from Professor His no sign of regret for his omis- 
sions. his errors, or his delay in their rectification. 


Witver, Neural Terms. 301 


Part VII. List of the Neural Terms, about Five Hundred and 
Forty in Number, Adopted by the Anatomische Gesell. 
schaft in 1895, together with Those now 
Preferred by the Writer. 


$234. The special names are arranged in seven groups, corre- 
sponding with the six definitive encephalic segments (see Table VII) 
and the myelon (spinal cord). In accordance with the general rule the 
series begins with the first (most cephalic or ‘‘ anterior”?) segment, and 
ends with the myelon. 


$235. First Column.—-These names are as in the German list 
published by Prof. His (’95, a, 80-87), excepting as follows :—(a@) Some 
typographic errors may have escaped detection. (4) It was found im- 
practicable to reproduce the original ‘‘ middle heads” in their various 
typographic forms; so far as possible, however, the subordination of 
less to more comprehensive terms is indicated by ‘‘ indents.” (c) In 
addition to words, mostly genitives of proper names, bracketed in the 
original, there are here introduced in brackets some entire terms to 
facilitate the recognition of certain parts apparently not specified in 
the German list. 


$236. Second Column.—These are the Latin terms now preferred 
by me. In some cases I have been in doubt as to the identity of parts 
indicated in the German list; in others as to the desirability of any 
designation at all. The capitalized names are those respecting the ex- 
cellence of which, both as designations and as terms, I feel most fully 
assured. 


$237. Zhird Column.—This contains the English forms, or An- 
glo-paronyms, of the Latin terms preferred by me. Often they are 
identical therewith, and in all cases the differences are so slight as not 
to hinder their recognition; §$44-48. 


$238. Last Column.—The signs refer to the adoptions of the Latin 
terms in the middle column. (a) The word gemeral signifies that the 
term is in common use. (4) The capitals N., G., A.,5., signify the 
formal adoption of the terms by the American Neurological Associa- 
tion (1896; §80); the Anatomische Gesellschaft (1895; $137); the 
Association of American Anatomists (1889; $81); and the American 
Association for the Advancement of Science (1892; $84). (c) The 
dates, ’80-’96, are those of my own adoption of the terms in the mid- 
dle column; see Part II and the Bibliography. 


302 JOURNAL OF COMPARATIVE NEUROLOGY. 


TABLE VI. 
Termini Neurologici Generales. 

German Committee. Present writer. Angloparonyms. | Adoptions, 
1|Nervus NERVUS Nerve General 
2|/Ganglion GANGLION Ganglion General 
3|Substantia alba ALBA Alba ’80; 2116 
4|Substantia grisea CINEREA Cinerea itefoy) OC 
5|Substantia gelatinosa |GELATINOSA Gelatinosa 96 
6|Taenia telarum RIPA Ripa SI 
7|Ependyma _ ventriculo-[ENDYMA Endyma "81 

rum 
8|[Liquor cerebrospinalis]};COELIOLYMPHA _ |Celiolymph 89 
9|Sulcus limitans ventric-|Sulcus interzonalis Interzonal  sul-|’96; 3195 

ulorum cus. 

1o|/Nuclei nervorum cere-|Nidi Nidi "89; 4123 
bralium 
11|Nuclei originis Nidi originis Nidi of origin |’89 
12|Nuclei terminales Nidi terminales Terminal nidi_ |’89 
13/Ramus communicans |RAMUS COMMUNI-|Communicating |General 
CANS ramus 
I4;/Ramus anastomoticus ;|RAMUS ANASTO-/Anastomotic ra-|General 
} MOTICUS mus. 
15|Nervus cutaneus NERVUS CUTAN -/Cutaneous nerve|General 
EUS 
16|Nervus muscularis NERVUS MUSCU-|Muscular nerve|General 
LARIS 
17|Nervus articularis N. ARTHRALIS Arthral nerve |’96 | 
18] Plexus nervorum spinal- PLEXUS NERVO-|Plexuses of the|General 
ium RUM SPINALIUM spinal nerves 
Termini Neurologici Proprii. 
I Rhinencephalon Rhinencephalon  |Rhinencephal °81 
1)Area parolfactoria [Bro-|Area parolfactoria Parolfactory area 

cae] 
2|Bulbus olfactorius Bulbus olfactorius Olfactory bulb /’81 
3\Gyrus subcallosus G. subcallosus Subcallosal gyre 
4|Limen insulae Limen Limen : 
5|Lobus olfactorius Lobus olfactorius Olfactory lobe |’81 
6|Pars anterior (rhinen- ? ? 

cephali) ? ? 
7\Pars posterior (rhinen- ? ? 

cephali) 
8|Stria intermedia Radix intermedia Middle root 81 
g|Stria medialis Radix mesalis Mesal root SI 
1o|Stria olfactoria lateralis|Radix lateralis Lateral root SI 
I1|Substantia perforata lat-- PRAECRIBRUM Precribrum 89 

eralis 
12/\Sulcus parolfactorius ? ? 

anterior 
13/Sulcus parolfactorius ? fe 

posterior 
14|Tractus olfactorius Tractus olfactorius Olfactory tract 


15|/Trigonum olfactorium |Trigonum olfactorium |Olfactory trigon 


WILDER, WVeural Terms. 


I Rhinencephalon 


“16|[Ventriculus olfacto-/RHINOCOELIA 


rius| 


Rhinencephalon 


17|{[Pars olfactoria foram-/Pars olfactoria aulae 


inis  interventricu- 


laris] 


18|[Pars anterior commis-|Pars olfactoria 


surae anterioris] 


Telencephalon 


1|Hemisphaerium 


Pallium 

Fissura longitudinalis 
cerebri 

F. transversa cerebri 


Gyri cerebri 

G, profundi 

G, transitivi. 

[G. operti] 

[G. operientes] 
Sulci cerebri 
[Fissurae cerebri] 
S. et F. operti 

S. et F. operientes 
Impressio petrosa 


Cnt Qt _ 


Xe) 


10 
II 
12 
13 
14 


commissurae 


Prosencephalon 


prae- 


Rhinencephal 


Rhinocele 


Olfactory part of 
the aula 


Olfactory part of)’ 


the precom- 
missure 


Prosencephal 


Hemicerebrum (cere- 
brum /e7e) 


Pallium 
F. intercerebralis 


Hiatus tentorli 


GYRI 

VADA 

ISTHMI 

Subgyri 

Supergyri 
FISSURAE 
FISSURAE 
Subfissurae 
Superfissurae 
Impressio petrosa 


15|[Impressio confluentis] |Impressio torcularis 


16|Fossa cerebri lateralis) FOSSA SYLVII 


17|Fissura cerebri lateralis) FISSURA SYLVII 


[Sylvii] 
[Syl vii] 
18} Ramus posterior 
19 
cendens 
20 
zontalis 
21 [Pars basilaris] 


22| Lobi cerebri 


23) Insula 
24| [Pars anterior] 
25 [Pars posterior] 
26 

sulae] 
27 Gyri insulae 
28] G. longus insulae 
29) Gyri breves insulae 


F. SYLVII (fre) 


Ramus anterior as-|F, PRAESYLVIANA 
Ramus anterior hori-|F. SUBSYLVIANA 


F. basisylviana 
Lobi cerebri 
INSULA 


Praeinsula 
Postinsula 


[Sulcus centralis in-/F, TRANSINSULAR- 


IS 
G. INSULAE 


G. longus insulae 


G. breves insulae 


Hemicere brum 
(cerebrum com- 
monly) 
Pallium 
Intercerebral fis- 
sure 
Tentorial 
val 
Gyres or gyri 
Vadums 
Isthmuses 
Subgyres 
Supergyres 
Fissures 
Fissures 
Subfissures 
Superfissures 
Petrosal impres- 
sion 
Torcular impres- 
sion 
Sylvian fossa 


inter- 


Sylvian fissure 


Sylvian f. (com-|’ 


monly) 
Presylvian f. 


Subsylvian f. 
Basisylvian f. 
Cerebral lobes 
Insula 
Preinsula 
Postinsula 


Transinsular f. 


Insular gyres 


Long insular 


gyre 


Short insular 


gyres 


89 
85 


General 
80, A. G. 
INES: 
89 
89 
89 


89 


304 


Telencephalon 


Prosencephalon 


30|S. circularis [Reili] 
31} [Polus insulae] 
32)Operculum 


33 Pars frontalis 
34 Pars parietalis 
35 Pars temporalis 
36} [Pars orbitalis] 


F. circuminsularis 
Polus insulae 
Opercula 
PRAEOPERCULUM 
OPERCULUM 
POSTOPERCULUM 
SUBOPERCULUM 


37| S. centralis [Rolandi] |F. CENTRALIS 


38] G. centralis anterior 


G. PRAECENTRAL- 
IS 


JOURNAL OF CoMPARATIVE NEUROLOGY. 


Prosencephal 


382 


Circuminsular f.|’89 


Insular pole 
Operculums 
Preoperculum 
Operculum 
Postoperculum 
Suboperculum 
Central fissure 
Precentral gyre 


39| G. centralis posterior |G. POSTCENTRALIS/Postcentral g. 


40/[S. centralis inferior] 


F. subcentralis 


41/[G. transitivus central--ISTHMUS CEN- 


s] 


TRALIS 


Subcentral f, 


Central isthmus 


i 
42|[G. transitivus profun--VADUM CENTRALE)|Central vadum 


dus centralis] 
43|Lobus frontalis 
44| [Pars postfrontalis] 
45| [Pars praefrontalis] 
46}Polus frontalis 
47\Sulcus 
[pars superior] 
48 
inferior] 
49|G. frontalis superior 


50| S. frontalis superior 
51|/G. frontalis medius 


Pars superior 
Pars inferior 


52 
53 


Lobus frontalis 
Pars postfrontalis 


Pars praefrontalis 


Polus frontalis 


praecentralis}F. PRAECENTRAL- 


IS 


S. praecentralis [pars|F. postfrontalis 


G. SUPERFRONT- 
ALIS 

F. SUPERFRONT- 
ALIS 

G. MEDIFRONTAL- 
IS 


Pars dorsalis 


Pars ventralis 


54/[Sulcus frontalis inter-|F. MEDIFRONTAL- 


medius | 


55\|Sulcus frontalis inferior 


56/G. frontalis inferior 


57 Pars opercularis 
58 Pars triangularis 
59 Pars orbitalis 


60)/G. rectus 

61|S. olfactorius 
62/Gyri orbitales 
63|Sulci orbitales 
64|Lobus temporalis 
65|Polus temporalis 


66|Sulci temporales trans- 


versi 


67|Gyri temporales trans- 


versi 


68|/G. temporalis superior 


69/S. temporalis superior |F. SUPERTEMPOR-|Supertemporal f. 


70|G, temporalis medius 


IS 
F. SUBFRONTALIS 
G. SUBFRONTALIS 
Pars opercularis 
Pars praeopercularis 
Pars subopercularis 
G. mesorbitalis 
F. olfactorius 
G. orbitales 
F. orbitales 
Lobus temporalis 
Polus temporalis 
F. transtemporales 


G, transtemporales 


gyres 
G. SUPERTEMPOR-|Supertemporal g. 


ALIS 
ALIS 


G.MEDITEMPORAL-|Meditemp’ral g. 
IS 


Frontal lobe 


Postfrontal por- 


tion 
Prefrontal 

tion 
Frontal pole 
Precentral f. 


Postfrontal f. 
Superfrontal f. 
Superfrontal f. 
Medifrontal g. 
Dorsal part 
Ventral part 
Medifrontal f. 
Subfrontal f. 


Subfrontal g. 
Opercular part 


Preop’rcular part 
Subop’rcul’r part 


Mesorbital g. 
Olfactory f. 
Orbital gyres 
Orbital fissures 


Temporal lobe 
Temporal pole 
Transtemporal 
fissures 
Transtemporal 


por- 


89; Z102 
85 
*Si5sWNie 
85 

85 

82, N.2120 
85 

89 

89 

89; 2197 
89; 2197 


General 


General 
8s 
8s 
8s 
8s 
Ss 


II 
71 


IS 
72|G. temporalis inferior |G. SUBTEMPORAL- 
IS 


73 
74 


75 


76 


77 
78 


79 
80 


81 
82 
83 


84 
85 
86 
87 


88 
89 
go 


91 
92 
93 
94 


95 
96 
97 
98 
99 
100 
101 
102 


103 
104 
105 
106 
107 


WILDER, WVeuval Terms. 


Telencephalon 


S. temporalis medius 


F. collateralis 
G. fusiformis 


G. lingualis 


Lobus occipitalis 

Polus occipitalis 

S. occipitalis transver- 
sus 

G.occipitales sup’riores 

S. oc. superiores 

G. oc. laterales 

Lobus parietalis 

Lobulus parietalis su- 
perior 

Lobulus parietalis in- 
ferior 

Sulcus interparietalis 


[Pars postcentralis su- 
perior] 
[Pars postcentralis in- 
ferior] 
[Pars horizontalis] 
[Pars occipitalis] 
[S. praeparoccipitalis] 


[S. postparoccipitalis] 


[Gyrus transitivus par- 
occipitalis] 

[G. transitivus opertus 
paroccipitalis] 


Prosencephalon 
F,MEDITEMPORAL- 


Prosencephal 


Subtemporal g. 


F. COLLATERALIS |Collateral f. 


G. SUBCOLLATER-|Subcollateral g. 
ALIS 
G. SUBCALCAR-|Subcalcarine g. 


INUS 

L. occipitalis 

P. occipitalis 

F. paroccipitalis 
parte) 


Occipital lobe 

Occipital pole 

Part of the par- 
occipital f. 


(Za 


: ? 
? ? 
> ? 


Parietal lobe 
Parietal gyre 


Lobus parietalis 
G. PARIETALIS 


G. SUBPARIETALIS|Subparietal g. 


Complexus fissuralis in-|Intraparietal fis- 
traparietalis (?) sural complex 

F, POSTCENTRALIS|Postcentral f. 

F, praeparietalis Preparietal f. 

F. PARIETALIS Parietal f. 

F. PAROCCIPITALIS|Paroccipital f. 

F. PRAEPAROCCIP-|Preparoccipital f. 
ITALIS 

F, POSTPAROCCIPI-|Postparoccipital 


Lobulus parietalis in- 
ferior 
G. supramarginalis 
G, angularis 
[S. intermedius] 
[S. exoccipitalis] 
[F. calcarina externa] 
5. corporis callosi 
S. cinguli 
Pars subfrontalis 


Pars marginalis 
S. subparietalis 
F. hippocampi 
G. fornicatus 


TALIS fissure 
ISTHMUS  PAROC-/Paroccipital isth- 
CIRTTATIS mus 
VADUM PAROCCIP-|Paroccipital va-|’ 
ITALE dum 
G. SUBPARIETALIS|Subparietal g. 
G. marginalis Marginal gyre 
G, angularis Angular g. 
F. intermedia (?) Intermedial f. 
F. exoccipitalis Exoccipital f. 
F. lambdoidalis Lambdoidal f. 
F. CALLOSALIS Callosal f. 
\[needless] 
F. SUPERCALLO- _ _|Supercallosal f. 
SALIS 


Paracentral f. 
Precuneal f, 
Hippocampal f. 


F. paracentralis 
F. praecunealis 

F, HIPPOCAMPI 
[needless] 


395 


°82 


Meditemporal f.|’85 


85 


85 
85 


85 


General 


General 
89 


89 


85 
96 
85 
86 
86 
86 


"86; 3197 


G. cinguli G. CALLOSALIS Callosal g. 
1o8|Isthmus gyri fornicati |Isthmus cinguli Isthmus of the|’96 
cingulum 
Un Uncus 


109 


I 


cus [gyri hippocam-/UNCUS 
i] 


82 


306 JOURNAL OF COMPARATIVE NEUROLOGY. 


’ 


II Telencephalon Prosencephalon Prosencephal 82 
I1o|Substantia reticularis |Reticularis alba Reticular alba 

‘ alba [Arnoldi] 

111/Lobulus paracentralis |G.PARACENTRALIS|Paracentral gyre|‘85 
112|Praecuneus PRAECUNEUS Precuneus 785 Gane 
113|F. parietooccipitalis |F. OCCIPITALIS Occipital fissure |’85 
114\F. calcarina F, CALCARINA Calcarine f. 85 G. N. 
I15|F. occipito-calcarina |F, occalcarina Occalearine f. |'89 
116)[S. postcalcarina] F. postcalcarina Postcalcarine f. |’85 
117|Cuneus CUNEUS Cuneus 89 G. N. 
118|[Isthmus cunei] ISTHMUS CUNEI  |Cuneal isthmus |’96 
119|[Isthmus opertus cunei]| Vadum cunei Cuneal vadum_ |'96 
120/[S. adoccipitalis] F. adoccipitalis Adoccipital f. — |’89 
121|[Cuneus anterior] CUNEOLUS Cuneolus 95 
122|[S. inflecta] F. inflecta Inflected f. 85 
123|[S. fronto-marginalis] |F. fronto-marginalis Fronto-marginal |’89 


124|S. oc. laterales ? ? 


il. B. Cava et Corum Parietes. 


II Telencephalon Prosencephalon Prosencephal | 82 
1\[Cavitas telencephali] |PROSOCOELIA Prosocele 86; 2201 
2\[Pars medialis foramin-|AULA (ere) Aula(commonly)|’80; 4211 

is interventricularis] 
3)[Pars lateralis foraminis| PORTA Porta "81; 2211 
interventricularis] 
4/Ventriculus lateralis PARACOELIA Paracele ’S9 
5 Pars centralis CELLA Cella 82 
6} Cornu anterius PRAECORNU Precornu 81; 7185 
7 Cornu posterius POSTCORNU Postcornu SI 
8] Cornu inferius MEDICORNU Medicornu "81 
9|Corpus striatum STRIATUM Striatum 80; N. 
10/Nucleus caudatus CAUDATUM Caudatum 80; Q11I 
It Caput nuclei caudati/CAPUT Caput ‘89; 726 
12 Cauda nuclei caudati/CAUDA Cauda iste) 
13/Stria terminalis Taenia Tenia 89 
14|Lamina affixa ? ? 
15|Taenia chorioidea RIPA Ripa 82a 
16|Lamina chorioidea epi-- PARATELA Paratela *89 
thelialis 
17|Calcar avis CALCAR Calcar (7106) ’81;A.N.S. 
18/(Bulbus cornu poste--EMINENTIA OCCIP- Occipital emi -|’89 
rioris) ITALIS nence 
19|Eminentia collateralis |EMINENTIA COL- [Collateral emi-|/General 
LATERALIS nence 
20/Trigonum collaterale |Trigonum collaterale |Collateral trigon|’96 
21|/Hippocampus HIPPOCAMPUS Hippocamp 780), AC Gre 
N.S 
22|Fimbria hippocampi |FIMBRIA Fimbria 81 
23/Taenia fimbriae RIPA Ripa SI 
24/Digitationes hippo-|Digitationes hip po-|Hippocampalj’96 
campi campi digitations 
25|Fascia dentata hippo-|Fasciola Fasciola SI 


campi 


Wiper, Neural Terms. 


Il 


6|Fasciculus longitudina- 
lis inferior 
7\Fasciculus longitudina- 
lis superior 
8}Fasciculus uncinatus 
g|Radiatio corporis cal- 
losi 


Fasciculus dorsalis 
Fasciculus ventralis 


Fasciculus uncinatus 


Radiatio callosi 


Telencephaloa Prosencephalon Prosencephal | ’82 
26/Commissura hippo- |Fornicommissura Fornicommis- |’89 
campi sure 
27|Corpus callosum CALLOSUM Callosum (Z111) 80 N. 
28} Splenium corporis |SPLENIUM Splenium 80 
callosi 
29} Truncus corporis |CALLOSUM (fere) Callosum (com-|’80 
callosi j monly) 
30| Genu corporis cal-\GENU '/Genu 81; 226 
losi 
31 Rostrum corporis |ROSTRUM Rostrum SI 
callosi 
32 Lamina rostralis Copula Copula ‘ 89 
8 Striae transversae |Striae transversae Transverse striae|’98 
34 Stria longitudinalis|Stria mesalis Mesal stria 96 
medialis i 
35)  Stria longitudinalis|Stria lateralis Lateral stria 96 
lateralis ; 
36 Fasciola cinerea Fasciola Fasciola 81 
37| [Indusium] Indusium Indusium 93 
38) Fornix FORNIX Fornix General 
39 Crus fornicis needless ; ; 
40 Corpus fornicis FORNIX (fre) Fornix (com-|’80 
monly) 
41 Taenia fornicis Ripa Ripa. 
2 Columna fornicis Fornicolumna Fornicolumn 84 
43| Pars libera colum-|Pars libera fornicolum-|Free part of the)’96 
nae fornicis nae fornicolumn 
44! Pars tecta columnae|Pars tecta fornicolum- Concealed part ’96 
fornicis nae of the forni- 
column 
45| [Crista] CRISTA Crista ’80; Z210 
46) [Carina] CARINA Carina 81 
47\Septum pellucidum |SEPTUM Septum 89; 726 
48 Lamina septi pellu-- HEMISEPTUM Hemiseptum 82 
cidi 
49| [Cavum septi pellu-/PSEUDOCOELIA Pseudocele SI 
cidi| 
Ii. €. Sectiones Prosencephali. 

I Telencephalon Prosencephalon Prosencephal 782 
1\Substantia corticalis |CORTEX Cortex SI 
2|Centrum semiovale MEDULLA Medulla "89; 726 
3|Decursus fibrarum cer- 

ebralium ? 
4|Fibrae arcuatae cerebri ? 
5/Cingulum Cingulum Cingulum 


Dorsal fasciculus 


Ventral fascicu- 
lus 

Uncinate 
culus 

Callosal rad ia- 
tion 


fasci- 


307 


308 JoURNAL OF COMPARATIVE NEUROLOGY. 
II Telencephalon Prosencephal Prosencephal 81 
Io} Pars frontalis Radiatio frontalis Frontal radia- 
tion 
11| Pars parietalis Radiatio parietalis Parietal radia- 
tion 
12| Pars temporalis Radiatio temporalis Temporal radia- 
tion 
13] Pars occipitalis Radiatio occipitalis Occipital radia- 
tion 
14) Tapetum Tapetum Tapetum General 
15|Nucleus lentiformis LENTICULA Lenticula 89; 19 
16| Putamen PUTAMEN Putamen General 
17| Globus pallidus Pallidum Pallidum 96 
18|Claustrum CLAUSTRUM Claustrum General 
19/Capsula externa ig ? 
20|Capsula interna Capsula Capsula 89 
21; Genu capsulae inter-/Genu Genu 
nae 
22| Pars frontalis capsu-|Pars frontalis Frontal part 
lae internae 
23| Pars occipitalis c. i. -|Pars occipitalis Occipital part 
24|Nucleus amygdalae Amygdala Amygdala 85 
25|Corona radiata CORONA Corona 89 
26| Pars frontalis Pars frontalis Frontal part 
27| Pars parietalis ijPars parietalis Parietal part 
28} Pars temporalis Pars temporalis Temporal part 
29| Pars occipitalis Pars occipitalis Occipital part 
30|Radiatio corporis striata|Radiatio striatalis Striatal radia- 
tion 
31|Radiatio occipitothal- |Radiatio thalamica Thalamic radia- 
amica [Gratioleti] tion 
32|Commissura anterior |PRAECOMMISSURA| Precommissure |’81 
[cerebri] 
33| Pars anterior (see I, 18) 
34| Pars posterior Pars temporalis Temporal portion 
ne ° - 
iil Whale odceohalon Diencephalon Diencephal Sl 
1|Thalamus THALAMUS Thalamus 80, G. N. 
2} Pulvinar PULVINAR Pulvinar General 
3} Tuberculum anterius|/Tuberculum thalami |Thalamic_tuber- 
thalami cle 

4| Taenia thalami RIPA Ripa 81; 3240 

5| Stria medullaris Stria medullaris Meduilary stria 

6| Lamina chorioidea {Pars marginalis dia-|Marginal part of 
epithelialis telae? diatela (needless) 

7|Metathalamus (Neediess) 

8| Corpus geniculatum]POSTGENICULUM |Postgeniculum |’89 
mediale 

9} Corpus geniculatum]/PRAEGENICULUM |Pregeniculum  |’89 
laterale 

10| Epithalamus (Needless) 96 

11| Corpus pineale EPIPHYSIS Epiphysis 95; 768 

12| Recessus pinealis Recessus epiphysialis |Epiphysialrecess 

13} Recessus suprapine-|Saccus dorsalis Dorsal sack 96, d 


alis 


Wiper, Neural Terms. 309 


Ill PREG EAEeEH AION Diencephalon Dience phal 81 

14 Habenula HABENA Habena SI 

15} Commissura habenu-/SUPRACOMMISSU- Supracommis-_ |’86 

larum RA (H. F. Osborn) sure 

16| Trigonum habenulae|Trigonum habenae Habenal trigon |’96 

17| Hypothalamus : (Needless) 

18|Pars mamillaris hypo- (Needless) 

thalami 

19} Corpus mamillare ALBICANS Albicans 81 
20|Pars optica hypothalami (Needless) 

21| Tuber cinereum Tuber Tuber 9 

22| Infundibulum INFUNDIBULUM Infundibulum /’S1, G.N. 
23| Hypophysis HYPOPHYSIS Hypophysis 80, G.N. 
24 Lobus anterior Praehypophysis Prehypophysis |’89 

25 Lobus posterior Posthypophysis Posthypophysis |’89 

26| Tractus opticus TRACTUS OPTICUS |Optic tract General 
27 Radix medialis Radix mesalis Mesal root SI 

28 Radix lateralis Radix lateralis Lateral root ‘$1 
29|Chiasma opticum CHIASMA Chiasma 80, N. 
30|Lamina terminalis TERMA Terma SI 


Ili. B. Cavitas Diencephali 


ll Diencephaion et 


Thalamencephalon Diencephalon Diencephal S81 
1}Ventriculus tertius DIACOELIA Diacele "SI 
2|Aditus ad aquaeduc- (Needless) 

tum cerebri 
3/Commissura posterior |POSTCOMMISSURA |Postcommissure |’81 
[cerebri] 
4|Foramen interventricu-|(Vide Prosencephalon |(See Prosenceph- 
lare [Monroi]| B, 2) al B, 2) 
5\Sulcus hypothalamicus) Aulix Aulix (2195) 84 
[Monroi] 
6| Massa intermedia ~ |MEDICOMMISSURA | Medicommissure|’80 
7\Recessus opticus RECESSUS OPTICUS]|Optic recess 82 
8}Recessus infundibuli |RECESSUS INFUN-|Infundibular re- 
DIBULI cess 
9|Commissura anterior |(Vide Prosencephalon |(See Prosenceph- 
[cerebri] C, 32) aly@ 32) 
10|Recessus triangularis |Recessus aulae Aulic recess (See|’81 


Prosen, B, 2; 
’ 2240 


310 JoURNAL OF ComPARATIVE NEUROLOGY. 
Hil. C. Sectiones Diencephali 
Jiencephaton et . : 
Til oN ee aan DPiencephalon Diencephal 81 
I1/Stratum zonale 
12)Nucleus anterior thal- 
ami 
13|N. medialis thalami 
14|N. lateralis thalami 
15!Laminae medullares 
thalami 
16} Nucleus corporis genic- 
culati medialis 
17|N. c. g. lateralis 
18|N habenulae Nidus habenae Habenal nidus 
I9|Fasciculus retroflexus |Fasciculus retroflexus |Fasciculus retro- 
[Meynerti] flexus i 
20|Nucleus hypothalami- 
cus [Corpus Luysi] 
21/Pars grisea hypothala- 
mi 
22|Commissura superior 
[Meynerti] 
23|Commissura inferior| Infracommissura Infracommissure 
[Guddeni] 
24|Nuclei corporis mamil-|Nidi albicantiae Albicantial nidi 
laris 
25|Fasciculus thalamoma- 
millaris [Vicq qd’ 
Azyri] 
26|Fasciculi pedunculo- 
mamillaris 
27| Pars tegmentalis 
28} Pars basilaris 
29|Ansa peduncularis 
30| Pedunculus thalami 
inferior 
31} Ansa lenticularis 
IV| Mesencephalon Mesencephalon Mesencephal |’8S1,Gen’l 
1;Aquaeductus cerebri |MESOCOELIA Mesocele SI 
[Sylvii] 
2|Basis pedunculi CRUSTA Crusta 89 
3\Brachium quadrigemi-|/POSTBRACHIUM Postbrachium — |’89 
num inferius 
4\Brachium quadrigem-;/PRAEBRACHIUM _ |Prebrachium 89 
inum superius 
5/Colliculus interior POSTGEMINUM Postgeminum  |’89 
6!Colliculus superior PRAEGEMINUM Pregeminum 89 
7\Corpora quadrigemina |QUADRIGEMINUM (|Quadrigeminum |'89 
8|Decussatio brachii con-|Decussatio brachii Brachial decus- 
junctivi sation 
9|Decussationes tegmen-|Decussationes tegmen-|Tegmental  de- 
torum torum cussations 
1o|Fasciculus longitudina-|Fasciculus longitudina-|Longitudinal 
lis medialis lis | fasciculus 


Wiper, Neural Terns. 


31g 


lV Mesencephaton Meseneephalon Meseneephal |°81,Gen‘l 
11|Formatio reticularis |Reticula Reticula 
12|Fossa interpeduncularis FOSSA INTERCRU-|Intercrural fossa)’89 
[Tarini] RALIS 


13| Lamina quadrigemina 

14;Lemniscus lateralis 

15|Lemniscus medialis 

16|Nucleus colliculi in- 
ferioris 

17|Nucleus nervi oculomo- 
torll 

18| Nucleus radicis descen- 


dentis n. trigemini 
19}Nucleus ruber 
20| Nuclei tegmenti 
21|Pedunculus cerebri 


22|Radix descendens n. 
trigemini 
23|Recessus anterior 
[of 12] 
24|Recessus posterior 
25/Stratum album profun- 
dum 
26|Stratum griseum cen- 
trale 


Stratum griseum collic- 
ull superioris 

Stratum zonale 

Substantia nigra 

Substantia periorata 
posterior 


27 


Lemniscus lateralis 


Lemniscus mesalis 


| 
( 


Nidus oculomotorius 


| RUBRUM 

Nidi tegmenti 

CRUS 

Recessus postcribralis 


Recessus praepontilis 


Entocinerea 


CAPPA 


INTERCALATUM 
POSTCRIBRUM 


31/Sulcus lateralis Sulcus lateralis 
32|Sulcus n. oculomotorii |Sulcus oculomotorius 
33) Tegmentum, TEGMENTUM 
34|[Tractus pedunculi “IMBIA 
transversus | 
35|Velum medullare ant. |)VALVULA 
36|Frenuium veli med. ant |FRENULUM 
Vv Metenvephalon Epencephalon 
1/[Pars metencephalica |\picoetia 
ventriculi quarti] 
2|[Pars metencephalica |Praeoblongata 
medullae oblongatae] 
3)Cerebellum CEREBELLUM 
4\Sulci cerebelli Sulci et rimulae 
5|Gyri cerebelli FOLIA 
6} Vallecula cerebelli VALLIS 
7\Incisura cerebelli pos- 
terior 


| 


Lateral lemnis-|’89, G. N. 


cus 
Mesal lemniscus 


Oculomotor ni- 


dus 


Rubrum 
Tegmental nidi 
Crus (2196) 


Postcribral re- | 
cess 


Prepontile recess 


Entocinerea 89 
Cappa "89 
Intercalatum 80; 2241. 
Posteribrum 89 
Lateral sulcus  |’89 
Oculomotor sul-|’89 

cus 
Tegmentum 89, G.N. 
Cimbia 81 
Valvula 81 
Frenulum 89; 2241 

Epencephal St 
Epicele Sr 
Preoblongata 85 
Cerebellum General 
(Sues andrimulas|'89; 3241 
Foliums *S9 
Vallis 906 


312 


¥ Metencephaton 
8/Sulcus horizontalis 
cerebelli 

go] Fissura transversa cere- 

belli 

Io] Vermis 

11|Lingula cerebelli 

12|Vincula lingulae cere- 

belli 

13|Lobus centralis 

14|Monticulus 

15/Culmen 

16|Declive 


17|Folium vermis 
18] Tuber vermis 
19|Pyramis [vermis] 
20|Uvula [vermis] 
21|Nodulus 
22| Hemisphaerium 
belli 
23|[Sulcus praeclivalis 
24|Ala lobuli centralis 
25|Lobulus quadrangularis 


cere- 


26) Pars anterior 

27| Pars posterior 

28|Lobulus semilunaris su- 
perior 

29! Lobulus semilunaris 
inferior 

3c| Lobulus biventer 

31{[Lobulus gracilis] 

32|Flocculus 


33|Flocculi secundarii 
34 ee oe 


ce ce 


36|Pedunculus flocculi 

37|Nidus avis 

38|[Folium cacuminis] 
> 


4o0|Brachium conjuncti- 
vum cerebelli 

41|[Fossa praepeduncu- 
laris| 

42| Fastigium 

43| Pons [Varoli] 


44| Sulcus basilaris 

45| Fasciculus obliquus 
pontis 

46| Fila lateralia pontis 

47| Brachium pontis 


. 


Epencephalon 


JOURNAL OF COMPARATIVE NEUROLOGY. 


Epencephal 


Sulcus peduncularis 


is 


VERMIS 
LINGULA 
Folia 


MONTICULUS 
CULMEN 
DECLIVE 


Tuber 

Pyramis 

UVULA 

NODULUS 

Pileum (B. B. Stroud) 


Sulcus furcalis 
Lobus quadrangularis 


Pars cephalica 
Pars caudalis 
Lobus praesemilunaris 


Lobus postsemilunaris 


Lobus cuneiformis 
Lobus gracilis 
FLOCCULUS 
Paraflocculus 
Supraflocculus 
Mediflocculus 
> 


? 
Cacumen 


POSTPEDUNCULUS 
PRAEPEDUNCULUS 


Fossa praepeduncularis 


FASTIGIUM 
PONS 


SULCUS BASILARIS 
Fasciculus obliquus 


? 
MEDIPEDUNCULUS 


81 


Peduncular sul-|’89 


cus 
? 


Vermis 
Lingula 
Foliums 7241 


Monticulus 
Cuimen 
Declive 


Tuber 
Pyramid 
Uvula 
Nodulus 
Pileum; 3241 


Furcal sulcus 


Quadrangular 
lobe 
Cephalic part 
Caudal part 
Presemilunar 
lobe 
Postsemilunar 
lobe 
Cuneiform lobe 
Slender lobe 
Flocculus 
Paraflocculus 
Supraflocculus 
Mediflocculus 


2241 
Cacumen 
Postpeduncle 
Prepeduncle 


Prepeduncular 

fossa 
Fastigium 
Pons 


Basilar sulcus 
Oblique fascicle 


Medipeduncle 


89 


89 
89 
89 
94 
"904 
"94 


89 
82 
7825-2241 


"89 


89 
80, A. 
G. N.S. 


81; 7198 


WILpvER, Neural Terms. 


Y. B. Sectiones Pontis et Cerebelli. 


Metencephalon Epencephalon 


Epencephal 


——— | _ eee Oe 


I 
2 


3 


Pars dorsalis pontis 
Raphe Rhaphe 
Nucleus n. abducentis |Nidus abducentis 


4|Nuclei motorii n. trig- 


5 


6 


7 


8 
9 


10 
II 
12 
13 
14 
15 
16 


17 


21|Lemniscus LEMNISCUS 
22| L. medialis sensitivus 
23| Lz. lateralis acusticus 
24|Pars basilaris pontis 
25|Fibrae pontis profundae|Fibrae pontis entales 
26|Fasciculi longitudinales 

[pyramidales] 
27|Nuclei pontis Nidi pontis 
28|Fibrae pontis superfic- |Fibrae pontis ectales 

jalis 

29/Corpus medullare Medulla 
30; Laminae medullares Laminae medullares 
31|/Arbor vitae ARBOR 
32| [Ramus posterior] Postramus 
33) [Ramus anterior] Praeramus 
34|Substantia corticalis CORTEX 
35| [Lamina basalis] [35, 30 and 37 are 
36) [Stratum cinereum] bracketed in the Ger- 
37| [Stratum gangliosum]} man list] 
38| Stratum granulosum 
39|Nucleus dentatus DENTATUM (6123) 
40| Nucleus fastigii Fastigatum 
41|Nucleus emboliformis |Embolus 


emini 
Radix descendens [mes- 
encephalica] n. trig- 
emini 
Tractus spinalis n. 
trigemini 
Nucleus tractus spinalis 
n. trigemini 
Nucleus n. facialis Nidus facialis 
Radix n. facialis Radix n. facialis 


Pars prima 
Genu [internum] 


Pars secunda 
Nuclei n. acustici Nidi acustici 
Nuclei n. cochlearis |Nidi n. cochlearis 
Nuclei n. vestibularis| Nidi n. vestibularis 
Nucleus olivaris supe- 
rior 
Nucleus lemnisci later- 
alis 
Fasciculus longitudi- |Fasciculus longitudi- 
nalis medialis nalis 
Formatio reticularis 
Corpus trapezoideum |TRAPEZIUM 


42|Nucleus globuliformis |Globulus 


Rhaphe 
Abducent nidus 


Facial nidus 
Root of the fac- 
ial nerve 


Acoustic nidi 
Cochlear nidi 
Vestibular nidi 


Longitudinal fas- 
ciculus 


Trapezium 
Lemniscus 


Ental fibers of 
the pons 


Pontile nidi 
Ectal fibers of 
the pons 
Medulla 
Medullary lami- 
nae 
Arbor 
Postramus 
Preramus 
Cortex 


Dentatum 
Fastigatum 
Embolus 
Globulus 


"81; 219 
89, G.N 


314 JouRNAL OF CoMPARATIVE NEUROLOGY. 


VI| Myelencephaion Metencephalon Metencephal °80 
1}Ventriculus quartus  |Metepicoelia Metepicele ’86; 7242 


[cavitas communis 
myelencephali et 
metencephali] 
2|Fossa rhomboidea 
3|Pars inferior fossae Calamus Calamus "82 
rhomboideae [cala- 
mus scriptorius] 
4|Pars intermedia f. r. 
5|Pars superior f. r. 
6\Sulcus limitans f. r. —_ |Sulcus interzonalis (?) |Interzonal sulcus 
7| Fovea inferior 2195 
8| Fovea superior 
9|/Trigonum n. hypoglossi | 


10|Striae medullares Striae acusticae Acoustic striae 
11|Eminentia medialis 
12|Colliculus facialis Lophius facialis Facial lophius 
13|Ala cinerea Ala cinerea 
14|Area acustica Area acustica 
15|Locus caeruleus Locus caeruleus 
16|[Hordea] Hordea [Spitzka] Hordeums 
17|Tegmen ventriculi 
quarti 
18/Velum medullare pos- |Kilos Kilos 
terius 9242 
19) Taenia ventriculi quarti|Ligula Ligula 89; 7239 
20|Obex OBEX Obex 89; 7242 
21)Lamina chorioidea epi-|Pars marginalis meta- |(Needless) 
thelialis telae 
22|(Apertura medialis ven-| METAPORUS Metapore 89; '93, « 


triculi quarti [Fora- 
men Magendii] ) 


23)(Apertura lateralis ven- (Existence “OGG 
triculi quarti) doubted) 

24|Fastigium (See V, 42) 

25|Medulla oblongata Postoblongata Postoblongata |’85 

26|[Pars myelencephalica |METACOELIA Metacele SI 


ventriculi quarti] : 
27|Fissura mediana poster-[SULCUS DORSALIS |Dorsal sulcus [2132 


jor 
28|F. mediana anterior S. VENTRALIS Ventral sulcus 
29|Foramen caecum Recessus postpontilis |Postpontile re- |'81 
cess 
30|Pyramis [medullae ob- |PYRAMIS Pyramid "81 
longatae] 
31|Decussatio pyramidum |Decussatio pyramidum| Pyramidal decus |General 
sation 
32|Sulcus lateralis anterior|Sulcus ventrolateralis |Ventrolateral 
sulcus 
33|S. lateralis posterior |S. dorsolateralis Dorsolateral 
sulcus 
34|Oliva OLIVA Oliva 86, G. N. 
35|Corpus restiforme RESTIS Restis 89; 219 
36|Funiculus lateralis Funiculus lateralis Lateral funiculus 
37|Funiculus cuneatus Funiculus cuneatus Cuneate funi- 


culus 


WILDER, Neural Terms. 315 


VI| Myelencephalon Metencephalon Metencephal 280 
38/Tuberculum cinereum |Tuberculum cinereum /Cinereal tubercle 
39|Funiculus gracilis Funiculus gracilis Slender funiculus 
40|Clava CLAVA Clava 81, G.N. 
41|Fibrae arcuatae exter- |Fibrae arcuatae ectales|Ectal arched fi- 

nae bers 


VI. B. Sectiones Postoblongatae. 


VI| Myelencephalon Metencephalon Metencephal 80 
ING Raphe Rhaphe Rhaphe (3242) 


2|/Stratum nucleare Stratum nidale Nidal stratum 
3\Nucleus n. hypoglossi |Nidus hypoglossi Hypoglossal ni- 
dus 
4|Nucleus ambiguus 
5|Nucleus alae cinereae 
6|Tractus solitarius 
7\Nucleus tracti solitaril 
8|Tractus spinalis n. tri- 
gemini 
g|Nucleus tractus spinalis 
n. trigemini 
10|Nucleus funiculi grac- 
ilis 
11/Nucleus funiculi cu- 
neati 
12| Nuclei laterales 
13}Nucleus olivaris infe- 
rior 
14|Nucleus olivaris acces- 
sorius medialis 
15|Nucleus olivaris acces- 
sorius dorsalis 
16] Nuclei arcuati ! 
17|Fibrae arcuatae inter- |Fibrae arcuatae entales|Ental arched fi- 
nae bers 
18|Substantia reticularis 
grisea 
19|Substantia reticularis 
alba 
20|/Fasiculus longitudi- |Fasciculus longitudi- |Longitudinal 
nalis medialis nalis fasciculus 
21\Stratum interolivare 
lemnisci 
22|Decussatio lemniscor- 
um 
23|Corpus restiforme RESTIS Restis 89; 319 
24|Fasciculi corporis rest- 
iformis 
25|Fibrae cerebellodlivares 
26| Fasiculi pyramidales 
27|Fibrae arcuatae exter- |Fibrae arcuatae ectales|Ectal arched fi- 
nae bers 


NT WW 


316 JOURNAL OF GOMPARATIVE NEUROLOGY. 


VII} Medulla spinalis Myelon Myel °S1; 251 
~~ }Pars cervicalis Myelon cervicale Cervical myel 
2|Intumescentiacervicalis|Intumescentia cervi- |Cervical enlarge- 
calis ment 
3}Pars thoracalis Myelon thoracale Thoracic myel 
4|Pars lumbalis Myelon lumbale Lumbar myel 
5|Intumescentia lumbalis|Intumescentia lumbalis| Lumbar enlarge- 
ment 
6}Conus medullaris CONUS Conus 
7|Filum terminale FILUM Filum 


8|Ventriculus terminalis |RHOMBOCOELIA /Rhombocele 85 
9|Fissura mediana anter- [SULCUS VENTRAL-|Ventral sulcus 


ior Is 
1o/Sulcus medianus poster-SULCUS DORSALIS |Dorsal sulcus {#132 
ior 
11/Sulcus lateralis anterior/Sulcus ventrolateralis |Ventrolateral 
sulcus 
12/Sulcus lateralis poster- [Sulcus dorsolateralis | Dorsolateral sul- 
ior cus 
13/Sulcus intermedius an- |Sulcus intermedius ven-| Intermediate 
terior tralis ventral sulcus 
14|Funiculi medullae spin-|Columnae szve funes s, |Myelic columns |’81; 2130 
alis funiculi myeli or funes or funi- 
culi 
15 Funiculus anterior |Columna ventralis Ventral column (#132 
16} Funiculus lateralis |Columna lateralis Lateral column 
17| Funiculus posterior |Columna dorsalis Dorsal column 


Vil. B. Sectiones Myeli. 


VII| Medulla spinalis Myelon Myel °S1; 251 
1|Canalis centralis MYELOCOELIA Myelocele 85 
2|Substantia grisea cen- |ENTOCINEREA Entocinerea 89 

tralis 
3|Commissura anterior |Commissura ventralis |White ventral 
alba alba commissure ; 
4|Commissura anterior |Commissura ventralis |Gray ventral 
grisea cinerea commissure 
5|Commissura posterior |Commissura dorsalis |Dorsal commis- 
sure 
6|Columnae griseae Cornua cinerea sive {Gray cornua or |’§1; 3128 
columnae cinereae columns 
Columna anterior Cornu ventrale Ventral cornu [131 
g|Columna lateralis Cornu laterale Lateral cornu 
Columna posterior Cornu dorsale Dorsal cornu 
1o| Cervix columnae pos- 
terioris 


11, Apex col. post. 
12| Substantia gelatinosa 
[Rolandi] 
13| Nucleus dorsalis [Stil- 
lingi, Clarkii] 
14|Formatio reticularis Reticula Reticula 
15|Funiculus anterior Columna ventralis Ventral column |7130 


Vil 


Wiper, Neural Terms. S17 


16 


Medulla spinalis Myelon Myel 815 251 


Fasciculus cerebro- 
spinalis anterior 
[pyramidalis ante- 
rior] 


17| Fasciculus anterior 
proprius [Flech- 
sigi| 
18|Funiculus lateralis Columna lateralis Lateral column |?130 
19| Fasciculus cerebro- 
spinalis lateralis 
[pyramidalis later- 
alis] 
20| Fasciculus cerebello- 
spinalis 
21) Fasciculus anterolat- 
eralis superficialis 
[Gowersi] 
22| Fasciculus lateralis 
proprius [Flech- 
sigi] 
23|Funiculus posterior Columna dorsalis Dorsal column |?130 
24| Fasciculus gracilis 
[Golli] 
25| Fasciculus cuneatus 
[ Burdachi ] 
Vill Meninges 
1|Dura mater encephali }|DURA Dura (2116) 80: N. 
2|Falx cerebri FALX Falx (2124) Sq: N. 
3|Tentorium cerebelli TENTORIUM Tentorium N. 
4|Falx cerebelli FALCULA Falcula 
5|Diaphragma sellae 
6| Foramen diaphragmatis 
sellae 
7|Incisura tentorili Incisura tentoril Tentorial incis- 
ion 
8|Dura mater spinalis Dura spinalis Spinal dura 
9|Filum durae matris spi-|Filum durae Dural filum 
nalis 
Io/Cavum epidurale Cavum epidurale Epidural cavity |118 
11\Cavum subdurale Cavum subdurale Subdural cavity 
12)Arachnoidea encephali |ARACHNOIDEA Arachnoid 80; 2118 
13|Cavum subarachnoidale|Cavum subarachnoidale|Subara chnoid 
cavity 
14|Cisternae subarachnoid-|Cisternae Cisterns 
ealae 
15|Cisterna cerebello-med-|Postcisterna s. c. cere- |Postcisterna or |’89 
ullaris bellaris cerebellar cis- 
tern 
16|Cisterna fossae lateralis} Cisterna Sylviana Sylvian cistern 
cerebri (Sylvii) 
17|Cisterna chiasmatis Cisterna chiasmatis Chiasmatic cis- 
tern 
18|Cisterna interpeduncu- |Cisterna cruralis Crural cistern 2196 


laris 


318 JOURNAL OF COMPARATIVE NEUROLOGY. 


VII Meninges 


19/Cisterna venae magnae |Medicisterna Medicisterna 89 
cerebri 

20/Granulationes arach- 
noideales [Pacchi- 


oni] 

21|[Leptomeninges] Piarachnoidea Piarachnoid 89 

22|Pia mater spinalis Pia spinalis Spinal pia 

23|Ligamentum denticula-|_Ligamentum denticula-|Denticulated lig- 
tum tum ament 

24|Septum cervicale inter- 
medium 

25|Pia mater encephali PIA Pia (2117) 80; N. 

26|Tela chorioidea ventric-|-METATELA Metatela SI 
uli quarti 

27|Plexus chorioideus ven-- METAPLEXUS Metaplexus SI 
triculi quarti 

28/Tela chorioidea ventric: DIATELA Diatela Table IV 
uli tertii 

29| Plexus chorioideus ven- Diaplexus Diaplexus 
triculi tertii 

30| Plexus chorioideus ven- PARAPLEXUS Paraplexus 89 
triculi lateralis 

31)/Glomus chorioideum Glomus Glomus 

32) Acervulus Acervus Acervus 

33|[Plexus chorioideus tel-|/ PROSOPLEXUS Prosoplexus 
encephali] 

34|[Tela foraminis inter- ;|AULATELA Aulatela 
ventricularis] 

35|[Plexus chorioideus for-| f AULIPLEXUS Auliplexus "SI 
aminis interventri-| | PORTIPLEXUS Portiplexus 
cularis] 

36|[Lamina chorioidea epi--PARATELA Paratela 89 


thelialis (II, 16)] 


IX. Principal Entocranial Bloodvessels; 7242 


A, Intrinsic Encephalic Arteries. 


1|Basilaris BASILARIS Basilar General 
2\Cerebelli inferior ante- |MEDICEREBEL- Medicerebellar |’85 
rior LARIS 
3\Cerebelli inferior pos- Beata aoe Postcerebellar |’85 
terior S) 
4|Cerebelli superior PRAECEREBEL- Precerebellar 9147 
LARIS 
5|Cerebri anterior PRAECEREBRALIS |Precerebral 85 
6;Cerebri media MEDICEREBRALIS |Medicerebral 85 
7\Cerebri posterior POSTCEREBRALIS |Postcerebral 85 
8|Cerebri anterior media |TERMATICA Termatic 85 
g|Circulus arteriosus[ Wil-|Circulus Circulus 
lisi 
10 2) anterior |[PRAECOMMUNI- Precommuni- 85 
CANS cant 
11|\Communicans posterior) POSTCOMMUNI- Postcommuni- _ |’85 


CANS cant 


Witpver, Neural Terms. 319 
A. Intrinsic Encephalic Arteries. 
12|Chorioidea anterior PRAECHOR OIDEA | Prechoroid 85, 
13|Chorioidea POSTCHOROIDEA | |Postchoroid "85 
14|Perforantes anteriores |PRAECRIBRALES  |Precribral 85 
15|Perforantes posteriores |POSTCRIBRALES Postcribral 85 
16\Spinalis anterior SPINALIS VEN- Ventral spinal 
TRALIS 
17|Spinalis posterior SPINALIS DORSAL-|Dorsal spinal 
Is 
18)Ramus meningeus Ramus meningeus Meningeal ramus 
19| Vertebralis VERTEBRALIS Vertebral General 
20/Rami spinales Rami spinales Spinal rami 
B. Dural Arteries. 
1} Meningea anterior PRAEDURALIS Predural 89 
2|Meningea posterior POSTDURALIS Postdural | 
3|Meningea media MEDIDURALIS Medidural 
4|Meningea parva Parviduralis Parvidural 
5|Meningea inferior Subduralis Subdural 
C.  Sinuses. 
1|Transversus \Lateralis ' Lateral 
2|Sagittalis superior Longitudinalis Longitudinal 
3|Sagittalis inferior FALCIALIS Falcial 
4|Rectus TENTORII Tentorial 
5|Petrosus inferior Subpetrosus Subpetrosal 
6|Petrosus superior Superpetrosus Superpetrosal 
7|\Sphenoparietalis Sphenoparietalis Sphenoparietal 
8)Confluens sinuum Torcular Torcular 
D. Encephalic Veins. 
1\Cerebri superiores Supercerebrales Supercerebral |’89 
2|Cerebri media Medicerebralis Medicerebral 
3\Cerebri inferiores Subcerebrales Subcerebral 
4|Cerebelli superiores Supercerebellares Supercerebellar 
5|Cerebelli inferiores Subcerebellares Subcerebellar 
6|Cerebri internae Velares Velar 
7|Cerebri magna [Galeni]|Magna (?) 
8|Septi pellucidi Septalis Septal 
g|Terminalis 
Io) Basalis [Rosenthali] Basalis 
11|Chorioidea Choroidea Choroid 


12}Opthalmomeningea 


? 


Notr.—To save space the substantive 


(arteria, vena, stnus) are omitted. 


elements of the vasal dionyms 


320 JOURNAL OF COMPARATIVE NEUROLOGY. | 


Comments upon Table VI. 


$239. Generval.—Professor Kolliker has characterized (’96, 
814; §208) ‘‘the anatomic nomenclature coming from America 
in recent years as a complete failure.’’ Professor His has declared 
(95, 6, 7; §170) that the writer’s ‘‘ proposals tend to create 
a language entirely new and for the most part quite strange, 
and on this ground our commission cannot follow him without 


») “AS a main basis of this 


renouncing its historic principles. 
conclusion, he imputes to me either (according to the intended 
sense of ‘verlangen lauter Mononyme’’) a strong desire for 
mononyms, or a demand for them to the exclusion of all polyo- 
nyms. Since the foregoing extracts might well indicate the 
existence of a divergence, wide, radical and irreconcilable, be- 
tween the neural terminology preferred by me and that recom- 
mended by the Gesellschaft, attention is asked to the following 
statistics : 

(2) Among the (about) 540 terms on the German list 
there are about 100 concerning which I refrain from expressing 
an opinion; my doubts are indicated sometimes by blanks in 
the second column, and sometimes by interrogation points. 

(6) Among the (say) 440 remaining, the following are so 
commonly employed that I claim no especial credit for having 
adopted many of themso long ago as 1880 or 1881 :—Nervus, 
ganglion, ramus communicans, ramus anastomoticus, nervus cutan- 
eus, nervus muscularis, plexus nervorum spinalium, lobt cervebri 
(frontalis, parietals, occipitalis et temporalis), eminentia collater- 
alis, fornix, tapetum, putamen, claustrum, pulvinar, tractus opticus, 
mesencephalon, cerebellum, decussatio pyramidum. 

(c) Among the (say) 420 remaining, respecting at least 105 
(about one-fourth) there is complete, or practically complete, 
concordance between the German committee and myself; of 
these, several were adopted between 1880 and 1882, and nearly 
all prior to the report of the German committee. 


1 Ts it permissible to entertain the hypothesis that not the least operative of 
these deterrent ‘‘historic principles’? are an indifference to what is done in 
America, and an indisposition to recognize value therein ? 


WILpER, Weural Terms. 321 


(2) Among the remainder (say 315), about 25, viz., prae- 
cribrum, postcribrum, vadum, aula, porta, vipa, copula, crista, 
carina, terma, cimbia, folium, pileum, metaporus and the com- 
pounds of coelza, may be regarded as unfamiliar. 

(c) But with most of the others the differences from the 
German equivalents may be indicated by the following exam- 
ples :—Calcar (for calcar avis), callosum (for corpus callosum), 
dura (for dura mater), postcornu (for cornu posterius), praecom- 
missura (for commiéssura anterior), gyrus subfrontalis (for gyrus 
frontalis inferior), medipedunculus (for drachium pontis), habena 
(for habenula), trapezium (for corpus trapezoides), radix dorsalis 
(for radix posterior), diaplexus (for plexus choriowdeus ventricult 
tertiz) ; see Part IV. 

(f) Finally, the German list contains at least forty mono- 
nyms, while in my own list of about 440, at least 270, more 
than one half, are polyonyms; see $242, IX. 

S240. Special.in the division ‘‘Termini Neurologici 
Generales’, in the sixth line, the term TZaenza telarum of the 
German list is made equivalent to my Ripa. A similar equiva- 
lency is indicated in II, B, 15 and 23; andin III, 4. In re- 
spect to these, and also the interpretations implied in II, B, 13; 
III, 5; and VI, 19, I wish to make further observations. I was 
early (81, @) impressed with the morphologic significance of 
these marginal parts or ‘‘shore-lines’”’ of the encephalic cavities, 
but now that the foregoing list is printed I am disposed to think 
that in my recent revision sufficient attention was not paid to 
the diagrams and suggestions of Prof. His (95, 165-168). If 
his views prove to be correct I shall be pleased, since upon 
some other points I have found myself unable to agree with 
him. Upon these and upon any other features of the Table 
and of the entire paper I desire criticism and suggestions from 
all who may interested. 

II, 17, “%ssura Sylvit.—tin advocating the retention of this, 
while objecting to eponyms in general since 1880, I am cer- 
tainly open to the charge of inconsistency. The following 
points should be borne in mind:—(1) Little personality or na- 
tionality attaches to this name. (2) There are exceptions to 


322 JOURNAL OF COMPARATIVE NEUROLOGY. 


most rules (§260). (3) Even if my daszsylvian, presylvian 
and subsylvian be rejected as titles of human fissures, there are 
several fissures in animal brains that have for many years been 
known by ‘‘sylvian’” compounds. (4) The substitute proposed 
by the German committee, cerebri laterals, is rather general than 
specific in suggestion. (5) If they are right in regarding the 
fissure as collocated with the striatum (His, ’95, 170), then, 
after the fashion of Azppocampal, calcarine and collateral, the 
more appropriate term would be /zssura striatals. 

Il, 74, 75, Gyrus subcollaterahs and G. subcalcarinus.—So 
slight is the resemblance of these cortical strips to the forms in- 
dicated in the commonly accepted szwz/e names, fuszformis and 
lingualts, that I have never been able to remember their relative 
locations. It seems probable that the fissural names calcarina 
and collateralis are to persist. If so, is it not both logical and 
convenient to designate the gyres just ventrad of them by loca- 
tives indicating their positions, viz., G. subcalcarinus and G. 
subcollateralis ? 

II, 85, Sedlcus enterparietahks.—Prof. Sir William Turner or- 
iginally named the fissure zxztraparictalis, and the same form is 
employed in his last fissure paper (Jour. Anat. and Phys., Oct., 
1890). To more essential grounds for doubting the advisability 
of applying any name to this ‘‘fissure complex,’’ must be added 
the carelessness of printers and proof-readers, and even the ap- 
parent ignorance of some writers as to the distinction between 
enter and znira. 

II, B, 2, and III, B, 10, Recessus aulae.—I may err in sup- 
posing the zecessus triangularis of the German list to be identi- 
cal with the vecessus aulae described by me in 1881 (’81, @). 

II, B, 4, Parvacoca.—Even were there not adequate rea- 
sons for replacing ventriculus in all neural names by coela, para- 
coela is simply the Greek equivalent of ventriculus laterals, and 
as such has equal privileges with the German heteronym 
“« Sectenhodhle’’ which has been used heretofore and will hardly 
disappear at once. 

II, B, 18, LFxunentia occipitahs.—The name preferred by 
the German committee, Bulbus cornu posterius, is bracketed in 


Wiper, Weural Terms. 323 


their list. As I have shown (’84, a, 373, and ’89, a, 143), at 
a certain fetal stage the occipital fissure has a distinct ental cor- 
relative and is hence entitled to be ranked as a total fissure, or, 
according to the distinction proposed by the German com- 
mittee, as a fissure rather than a sulcus. But in the adult the 
general thickening of the walls commonly obliterates the orig- 
inal elevation, and the ‘‘totality” of the fissure is not apparent. 
This temporary existence of the essential character should per- 
haps constitute a fourth objection to the distinction between 
fissura and sulcus proposed by the German committee and dis- 
cussed in §$121-122. 

II, B, 37, /udustum.—This seems to have been omitted 
from the German list. Its condition in man, chimpanzee, 
monkey, cat and sheep has been discussed by Fish (’93, @). 

II, C, 1, Cortex.—I am unable to see any good reason for 
replacing this familiar and suggestive mononym by the ponder- 
ous dionym, substantea corticals. 

II, C, 14, Yapetum.—The original Latin is fapete, perhaps 
from the Greek rays. The ending e occurs with so few anatomic 
terms, e. g., ve¢e, that it is perhaps scarcely worth while to dis- 
turb the established modern usage. 

II, C, 21-23.—Why should these names of divisions in- 
clude the genitive of the major part any more than with 10-14, 
or 26-29? 

Ill, 8, 9, Postgeniculum and Praegeniculum.—These loca- 
tive names retain the essential features of the earlier polyonyms. 
The parts are strictly caudad and cephalad of one another. The 
more lateral aspect of the praegeniculum is due toa difference 
in size. 


Ill, B, 4, Horamen tnterventriculare.—This cavity, the aula 
(S211) and two portas of my list (II, B, 2, 3), connects the 
right and left ‘‘ lateral ventricles.’’ These are cavities of the 
‘‘telencephalon” (my prosencephalon), not of the diencephalon. 

III, B, 9, Praecommissura.—This is certainly prosenceph- 
alic (‘‘telencephalic’’), and to include it among diencephalic 


parts is as artificial as in the case of the ‘‘ foramen interventric- 
ulare’”’ (III, B, 4). 


324 JoURNAL OF COMPARATIVE NEUROLOGY, 


§241. IV, 21, Cvus.—The fibrous mesencephalic masses 
are mentioned ten times where the ‘‘stems”’ of the cerebellum 
or olfactory bulbs are named once; hence the greater need of a 
brief mononymic designation for the first; for the second and 
third either the longer pedunculus or the addition of a qualifier 
would prove less burdensome. 

IV, 29, /utercalatum.—This locative mononym was sug- 
gested by Spitzka in 1887; W., ’89, a, §103.' 

V, 4, Sule? cerebelli.—For most of the interfoliar crevices 
I proposed vzmula in 1889. But, as shown by Stroud, certain 
of them are deep and merit the title of sw/cus. For the minor 
crevices 77mula (or perhaps swlculus) may serve. 

V, 6, Vallis.—For the grounds of preference for the briefer 
basal word ($30) over the longer diminutive, see W. and G., 
89, 529, $75. 

V, 12, Folia [ingulae|.—For the diminutive ‘‘leaflets”’ 
on the lingula no special name seems to be needed, and if it 
were, vzucula alone would suffice. 

V, 22, Prleum.—The substitution of this mononym for 
hemisphaerium cerebeli was proposed by Dr. B. B. Stroud (’94); 
on developmental grounds he recognizes a pracpileum and fpost- 


pileum. Stroud’s ‘‘furcal sulcus”’ seems to be what is called 


‘*preclival”’ in the last edition of Quain, III, Fig. 59. Other 
features of the cerebellum described and figured by Stroud (the 
cestus and several sulci, central, culminal, tuberal, pyramidal, 
uvular and nodular) are not included in the present list. 

V, 37, Medus avis.—The depression thus designated is so 
seldom mentioned that the dionym is not burdensome. If C. 
L. Herrick had not proposed zzdulus for nucleus the former 
might be an acceptable mononym for the little ‘‘ nest.”’ 


1JIn the German list, between mesencephalon and their metencephalon (my 
epencephalon), are enumerated nine parts under the head, ‘Isthmus rhomben- 
cephali.’’ Their omission from the present table was accidental, and due to my 
reversal of the original order of enumeration of the segments. I have already 
expressed(Z197) my objection to the assignment of segmental value to this neck- 
like region, and shall consider it hereafter (3250). Lemmnzscus is included in IV, 
14and 15. Brachium conjunctivum [cerebelli| probably designates the praepe- 
dunculus (V, 40). Velum medullare antertus is the valvula (IV, 35). For gang- 
lion interpedunculare 1 prefer ganglion intercrurale, and for Nucleus trochlearis, 
Nidus trochlearis, 


Wiper, Neural Terms. 325 


V, B, 36, Stratum cinereum.—Why not (in the German 
list) stratum griseum, in accordance with the word employed 
among the ‘‘ Termini generales ?” see §36. 

VI, 1, Metepicoeha or ‘‘ ventriculus quartus.’”-—The divis- 
ion of this continuous cavity into regions corresponding with 
the two recognized segments (V, 1 and VI, 26) must be con- 
sidered upon another occasion ($250). 

VI, 25, M€edulla oblongata.—According to the German 
committee this is coextensive with their myelencephalic segment 
(my metencephalon), and the ventral portion of the segment 
next cephalad (their metencephalon, my epencephalon) is con- 
stituted by the pons. But the pons exists only in mammals ; 
hence in the other vertebrates the cerebellar ‘‘ roof’’ would be 
unsupported by a ‘‘floor;” see V, 2, preoblongata. 

VI, 30, Pyramis.—The replacement of corpus pyramidale 
and of processus clavatus (40) by pyramzs and clava respectively 
was urged by Spitzka fifteen years ago (’81, @). 

VI, B, 15, Macleus olivaris accessorius dorsals.—I\f dorsalis 
be appropriate here, why not in several other cases where pos- 
terior is employed by the German committee ? 

$242. VII, JZyelon.—Respecting the substitution of this 
mononym for the dionym medulla spinalis, which was proposed 
by Owen just half a century ago, see §51. Owen also con- 
sistently, although I think unwisely, employed the compound, 
myelencephaton, for axis cerebrospinalts. Huxley applied it to 
the last encephalic segment, and this misappropriation is 
sanctioned by the German committee. Reserving comments 
upon these points for another occasion ($250), I now claim that 
the sole justification for the use of syelencephalon is the adop- 
tion of myelon in the Owenian sense. 

VIII, 32, Acervus.—This word, signifying a heap, occurs 
in Andrews and Stoddard’s lexicon. The diminutive, acesvulus, 
is longer, needless and of modern origin. 

IX, Lloodvessels.—As admitted by me in 1884 (’84, ¢) 
and restated in §§163 and 172, absolute mononymy is unattain- 
able with large groups of organs, e. g., muscles, fissures and 
vessels. Hence, excepting with crculus (A, g) and ‘orcular 
(C, 8), with the single word adjectives in this category must be 
understood avterza, stnus or vena; and when there is any danger 
of ambiguity the substantive or its abbreviation should be 
employed, 


; (sparta jo 
- uewesi1By ue 
SI[BSiOp ‘9d {si aque ant 
wnyy fsnuod] vijueaa Binssimm0; EOL SIN Bijaooojakyw| = uojakyw| uopadwl{rTA 
wnt wnp xoqo {ey uojeyqdes) =v} eSu0] 
-zadery featjo {stmeshg}-1urvidd onessnoaq|-nsiy ‘snaodejay snxo[deja Bpayejayy| = BIaooejay|-Uaiay  |-qoisog [TA 
uinjejuap ‘snjnsd04 uojeydas wn] 
fsturiaa { e}esuo;qosrig suog epnsary snxo|didy ead vrjaooid q |-uodq -[2qa1agQ JA 
wn}e[eo wn10}U9UE 
-1ajul {snostumay {vj} |-9a} sauoijessnoap (Aoidaiey oy} uoyeygeo| wnurmes 
-snio {uinjuemsea} fsnig 'B.INSSIMIUL0D}SOg B[NATeA Ul) Bla}OSa| BIye0sosay|-Uasay  j-lupen?y) AT 
e[noiues euseryo 
‘stsXydida ‘sisfydod |: eanssrmwooipou uojeydas 
-Ay f4aqny fcunaqiojsog| { vanssrummoovidng euoqeyy snxa|deiq Ba}VIG eI[20ovIq|-Ual ImMeleYy LITT 
[ae aes See ee Se oy s (ovjaqyerrnd "(aet]a00 ele Per 
snxodeaed yeroqey | [P1978] pus |-vivd [e107 
siskyd X1U1OJ ‘uINso]]eo paar Vacoan BjazU[NeB [Vso | puv aeceee ss 
-eied {mnyepneo fejn | (stpesodura} saved) vurts} feyed fay, = Sarpnyour) |°4}8uTpnyour) jayy Surpuyoar) | Uoreydas 
-I1JU9] § B[NsUT fwintIT[eg BANSSIMMCI9BAgG|‘VIIqMIy ferusey snxa|dosoig B[9}OSO1g| vi[aodoso1g |-Uasoig wNIGs19aD/T] 
(B110}0¥j[0 sized) Sees, uojeydas| m10j08y 
uaul, { wniqimoavsg BANSSIWIWODIVIg zjaiouryy| PHP°CUIYY|-ueuryy [0 Iq[Ng IT 
: ae _,| 40d uvrvgiy ; uousog sno P quaut \uanjezsuor 
Spang sayjo aos “9 |*Ia CAUSED) Z pun uy, °9 SISNXIAT °F -uvaguay t Ayana °F | Say ce | fy 1 


“SLIJIVAVYD 1dYJO OWOS PUL S}UIWSIG ALoYy} 0} Surps0s0v spiwvg [VnoN owog Jo Surdnoay [euowu0ig ‘TTA AIAVE 


Witper, Neural Terms. 327 


Commentaries Upon Table VII. 


$243. Its purpose is two-fold :—(a) To indicate, accord- 
ing to my present information and belief, the number and con- 
stitution of the definitive encephalic segments. (4) To illus- 
trate the verbal correlations between the names of the segments 
themselves (column 2), and those of (3) their major cavities, (4) 
their membranous parietes, and (5) their vascular plexuses. 

$244. It is in some respects an amplification of the table 
on p. 409 in W. andG., ’82. It differs from that in my later 
paper (’89, @, 121) in (a) the recognition of the Rhinencephal 
and (0) the vertical arrangement of the segments. 

$245. From Schwalbe’s table (81, 397) it differs mainly 
in the absence of any attempt to indicate the relative ‘‘ values ”’ 
of the several segments upon embryologic or other grounds. 

$246. In this respect it differs also from that of His (’g5, 
162). In this latter, moreover, I have not as yet succeeded in 
recognizing consistency with (a) his other table on p. 158, (0) 
the segmental arrangement employed in the German list of neu- 
ral terms (80-87), (c) a discriminating use of terms, (d) due re- 
gard for precedent, or (¢) the facts of comparative anatomy as I 
interpret them. 

$247. Conceding the high authority of Professor His as 
to the embryology of man, I nevertheless believe it to be alto- 
gether undesirable to infer the segmental constitution of the 
vertebrate brain from the conditions presented during the devel- 
opment of the human organ. Indeed, if the embryology of 
other forms were also taken into account, the number of poten- 
tial ‘‘neuromeres’”’ would be unmanageably large, even if any 
two investigators could agree at present as to how many should 
be recognized. 

§248. While anticipating that the problems involved will 
be eventually elucidated upon the basis of all the facts concerned, 
I believe our present effort should be to agree upon a schema 
of the vertebrate brain which, while not contravening the facts 
of embryology, shall harmonize so nearly with the facts of com- 
parative anatomy as to facilitate rather than obstruct an effort 


328 JOURNAL OF COMPARATIVE NEUROLOGY. 


to describe and interpret the conditions encountered in a given 
brain. 

$249. I freely admit my ignorance or non-comprehension 
of certain points, and also that my views have varied somewhat, 
particularly as to the segmental value of the olfactory region of 
the brain. Nevertheless, I regard myself as justified in advo- 
cating the schema presented in Table VII upon the following 
grounds :—(1) For more than twenty years the general question 
has never been long out of my mind; (2) with special reference 
to it I have prepared and studied scores of brains of all classes 
and most of the orders; (3) the subject has been discussed more 
or less fully in papers by me’ upon the brains of many different 
forms; (4) papers upon other forms’ have been prepared at this 
institution ; (5) the schema has proved practically available for 
research as indicated above, and has been readily comprehended 
and remembered by even general students. 

§250. What I advocate is that there be recognized for 
the present six definitive segments of the vertebrate brain under 
the titles Rhinencephalon, Prosencephalon, Diencephalon, Mes- 
encephalon, Epencephalon* and Metencephalon. It is my in- 
tention to review the whole subject at the coming meeting of 
the Association of American Anatomists in May, 1897. 


Part VIII. Concluding Remarks. 


S251. A. Practical Suggestions.—As one of the older 
American anatomists, and as having committed at least my 
full share of terminologic errors, I venture to formulate some 
suggestions of a practical nature for the benefit of the younger 
generation. 


Wee , Bibliography, (’75, ¢3. 776, a, 6, 63/7977, 4; 781, @5. 84,14, 085) Ogee 
"87, a, 6, c; "80, @; "91, 0; ’93, a; ’96, 2d} W. &G., *82, chap. X. 
*See papers by Clark, Mrs. Gage, Fish, Humphrey, Kingsbury and Stroud. 
3Even if Osborn is correct in his interpretation of the cerebellum as 


‘* primitively ” intersegmental (’88, 57) he nevertheless admits that it ‘ second- 
arily acquires a functional importance equal to that of the other segments.” 


oS Sa. 


WILDER, WVeural Terms. 329 


$252. Caution in Publishing New Terms.—It is true that 
words needlessly introduced into anatomy have no such embar- 
rassing permanency as is conventionally assigned to synonyms 
in systematic zoology. Nevertheless, for a time at least, they 
encumber current publications and dictionaries. Hence, how- 
ever necessary and legitimate they may seem to the framer, 
neither a new term, nor an old one in a new sense, should be 
actually published without prolonged consideration, and con- 
sultation with at least four individuals representing as many 
categories of possible critics :—(a@) an investigator of the same 
general subject; (0) an experienced teacher ; .(c) an earnest stu- 
dent; (Z) a philologic expert whose admiration for the past 
has not blinded him to the needs of the present and the future. 

§253. Method of Introduction of New Terms.—As ‘‘ ur- 
gently recommended ” by the A. A. A. S. Committee on Bio- 
logical Nomenclature (§84), ‘‘ Whenever a technical word is 
used for the first time, the author should give in a special note 
(a) the Latin form, (0) the etymology, (c) the proper adopted 
form or paronym for his own language, with the adjective, etc., 
when applicable, (@) as concise and precise a definition as 
possible.”’ 

$254. Lndirect Responsibility for Latin Terms.—Even when 
the foregoing admirable rule is not followed, the validity of the 
following can hardly be questioned :—‘‘The introduction of any 
derivative, oblique case, or national paronym renders the intro- 
ducer responsible for the actual or potential Latin antecedent of 
such word in accordance with the usual rules of derivation and 
paronymy (§178). 

$255. Paronyms versus Heteronyms.—Excepting with a 
few conspicuous or particularly important parts, e. g., head, 
heart, brain, e¢c. (§48), there should be employed either the 
Latin (international) names, or the national paronyms ($46; 
Tables Il and V). Itis quite true that ‘calling a millstone 
by a Greek name does not enable us to see a whit farther into 
it’”’; yet the designation of parts of the body by terms of classic 
source, even if somewhat modified in form, enables the anato- 


330 JoURNAL oF COMPARATIVE NEUROLOGY. 


mists of other nationalities to apprehend the signification more 
readily than they might from vernacular words. 

§256. Homonyms.—As has been repeatedly observed 
(§$23, 26, 68, etc.) the context commonly averts misapprehen- 
sion as to words having two or more meanings. The proba- 
bility of confounding the mouth with a bone is scarcely greater 
than that of mistaking a mathematic fora urinary calculus. But 
when a term or phrase possibly ambiguous is first introduced in 
a given publication, and especially in the title, absolute ex- 
plicitness should be attained, no matter how many qualify- 
ing words may be required. In the title of a paper, the term 
‘“cervical follicles’ is certainly ambiguous, and while ‘‘ mental 
prominence’”’ as employed by Huxley is shown by the context 
to designate a projection in the region of the chin, in a title it 
might be readily misunderstood, particularly by a psychologist. * 

§257. Consistency.—This ranks second among the desira- 
ble attributes of all scientific writing which I have long called 
the five C’s, viz., Clearness, Consistency, Correctness, Concise- 
ness, and Completeness. The last may seldom be attained; the 
lack of the first and second is as rarely excusable.” The prac- 
tice of the virtue of Terminologic Consistency is tantamount to 
avoidance of the vice of Pecilonymy (§$34-39). 

§258.  <Avordance of Pecilonomy.—Whatever doubts a wri- 
ter may entertain as to the relative excellence, authority or 
vogue of two or more synonyms, and however he may shrink 
from committing himself to either one of them (§39), justice to 


1 The title (‘‘On the fracture system of joints, with remarks on certain 
great fractures”) of a paper just received (Bost. Soc. Nat. Hist. Proceedings, 
XX VII) might at first sight seem to concern the surgeon quite as much as the 
geologist. 


2 While never really justifiable, obscurity of style may result from conditions 
more or less difficult to avoid; let usassume that no scientific writer would delib- 
erately formulate the doctrine credited by Jules Janin to Balzac. When asked the 
meaning of a passage the novelist is reported to have replied ‘* Ceci pour le 
bourgeois”, and to have explained that an unintelligible sentence or phrase now 
and then had a good effect on the ‘‘ general reader’’, who, if the sense were al- 
ways too obvious, might flatter himself that he was equal to the writer and on a 
level with his thoughts. 


— 


ee 


| 
: 


WILDER, WVeural Terms. 331 


his readers, if not regard for their good opinion, should lead him 
to make his selection in advance and to adhere thereto through- 
out a given publication.’ 

§259.  Abbreviational Methods.—The following rules are 
recommended. 

a. The abbreviation should indicate the Latin (interna- 
tional) name. With all mononyms this will also indicate equally 
well the national paronym; but with English and German polyo- 
nyms (§47), the usual transposition of the adjective and substan- 
tive renders the recognition less easy.” 

b. Abbreviations should be formed regularly, and vowels 
excluded excepting when the initial letter is such, or when their 
absence might occasion ambiguity. 

c. In the explanation of a figure abbreviations should be 
set in alphabetic order. So natural, reasonable and just is this 
rule that its disregard can only be attributed to the selfish assump- 
tion upon the part of a writer that the time its observance would 
have cost him is of more value to the world than the time its 
non-observance costs all of his readers together, not to mention 
the ill-effects of righteous indignation. 

$260. Importance of Moderation.—As with biologic gen- 
eralizations, there are few philologic rules without exceptions. 
Yet the reformer, especially if young and enthusiastic, either 
ignorant of history or undismayed thereby, ‘‘too often imagines 


1 As stated in 355, the principle and method were adopted by me in 1880. 
At that time Henle’s works were not known tome. But in 1884 I was so im- 
pressed with his systematic employment of a single set of names that the first 
step in the collaboration toward Foster’s Medical Dictionary (258) consisted in 


photographing the ‘‘ Index’ of his ‘‘ Nervenlehre”’ and distributing copies for 
discussion, 


2 From my point of view this constitutes an argument for the conversion of 
certain polyonyms into mononyms. For example, if the dionym commissura ana 
terior be retained, the Latin and French abbreviation would bec. a., the English 


a.c.,and theGermanv.c. But of the mononym, praecommissura, pre. would 
probably serve in each case. 


332 JouRNAL oF ComMPARATIVE NEUROLOGY. 


that a principle, if right, cannot be carried too far ;’’’ (Barclay). 
In this connection may be appropriately quoted the verse from 
Horace : 


“« Est modus in vebus ; sunt certi denique fines, 
Ultra citraque nequit consistere rectum.” 


§261. B. Suggestions to American Anatomists.—Circum- 
stances have precluded the possibility of submitting either the 
manuscript or the proofs of this article to other members of the 
American committees. Hence their responsibility for its con- 
tents must be limited strictly by their official recommendation 
of certain terms or principles, and by the usages embodied in 
their individual publications. I hope they will join in whatever 
discussion of the general subject may be aroused by this article 
freely and without apprehension that opposition to my views 
will affect my personal or official relations. All I ask of them 
is the clear recognition of all the conditions. 

§262. Perhaps my own view of what the conditions really 
are may be most conveniently introduced by a commentary 
upon a varagraph in the address of the president of the Asso- 


ciation of American Anatomists a year ago. Professor Dwight 


said (’95): 

‘*German anatomists have recently adopted a report prepared 
by some of their number working in company with representatives of 
other European countries. It is for us to consider whether this one can 
be looked upon as accepted and whether it is acceptable ; whether we 
can join hands with our foreign colleagues, or whether we can devise 
an American nomenclature which shall be so much better that we can 
disregard the inconvenience of a distinct standard. We have had 
for years a committee on Anatomical Nomenclature, with Professor 
Wilder for secretary, who has given so large a part of his busy life to 
this matter. We may expect an important contribution to the matter 
in the report of this committee.” 


1 TlJustrations may be found in the record of my own terminologic progress, 
Part II, in the rigid insistence at various stages upon the indispensability of 
words in Latin form (1880-1883), and upon idionyms or terms absolutely free 
from ambiguity (prior to 1895). Cerebrocortex and cerebellocortex were products 
of the too general (and yet never sweeping) application of the principle of 
mononymy. 


ee 


Wiper, Neural Terms. 333 


$263. Dr. Dwight’s address was devoted mainly to what 
he justly characterized as ‘‘a social question of the first impor- 
tance, far transcending purely scientific discussion, vzz., the 
methods of obtaining and utilizing anatomical material.” No- 
menclature was considered briefly and almost incidentally. The 
following commentaries are designed partly to reinforce some of 
his remarks, and partly to avert possible misapprehension as to 
both what he said and what he felt obliged to omit. 

§264. Inthe first place, asa member of the committee 
on Nomenclature of the Association of American Anatomists 
since 1889 ($81), Dr. Dwight recognizes with especial clearness 
that the subject can no longer be ignored. Now that a score 
of European anatomists have given more or less attention to it 
during six years, and have expended upon it about $2500.00, 
no individual or association can hereafter treat it as insignificant. 

$265. Secondly, the approximate completeness of the 
German list of the visible parts of the entire body renders it a 
substantial basis for discussion and a starting point for further 
progress. 

§266. The two conditions just named will, as doubtless 
anticipated by Dr. Dwight, lead anatomic writers and teachers 
to pay more heed to their terminology, and to maintain at 
least a temporary consistency, z. ¢., within the limits of a single 
lecture, article or treatise. 

$267. Yet our gratification at the tardy German admission 
of the need of terminologic improvement, and our recogni- 
tion of the usefulness of the list compiled with such learning 
and industry and at such expense, should not lead us to over- 
look (a) the limitations of the German report in both intent and 
performance ; (4) the delay in its adoption by other nations ; (c) 
the qualifications of Americans for independent judgment. 

8263. Phet oie. Ni Aa 2.0¢), the Wome Anatomica 
adopted by the Anatomische Gesellschaft at Basel in 1895, is 
regarded by the Germans themselves as provisional and subject 
to modification. As stated officially (Anat. Anzeiger, Ergan- 
zungsheft, X, 161), and by Prof. His (§$2, 228) there was ap- 
pointed a standing committee of revision, which is to report 


334 JOURNAL OF COMPARATIVE NEUROLOGY. 


upon proposed changes and new terms at intervals of three 
years. 

§269. Although France and Great Britain were repre- 
sented upon the general committee, no members from those 
countries were present at the signing of the report and of the 
declaration against the efforts of the American committees, 
April 19, 1895 (Ana. Anz., Erganz., X, 162). Furthermore, as 
frankly stated by Prof. His (95, 6-8), some of the French cor- 
respondents preferred a different method of procedure, and the 
English commission had not reported at all ($170). The im- 
probability of universal and unqualified assent upon the part 
of British anatomists is indicated by the following remarks of a 
Glasgow professor (Cleland and Mackay, ’96, 3): 


‘‘ With regard to the naming of individual structures it may: be 
noted that more than one attempt has been made to impose uniformity 
of nomenclature by the arbitrary authority of an individual or com- 
mittee.? It may be doubted if any such attempt can possibly be suc- 
cessful. The ‘‘ Nomina Anatomica ” of His (’95, @) is most impor- 
tant for consultation; but the adoption of its recommendations in 
this country (Great Britain) would, in a large number of instances, in- 
volve the abandonment of good names in general use for others whose 
advantages are not obvious.” 


$270. Through its secretary the German committee de- 


clared (Krause, ’91; $158, 7) that it intended to be ‘‘conser-_ 


vative in its action.”’ Now conservatasm is notoriously difficult 
to define, and in respect to nomenclature its degrees may equal 
in number those who have opinions upon the subject. , But 


1 So far as appears in the official record (Avat. Anz. XII, Erganzungsheft, 
1896)no reference to nomenclature was made at the last meeting of the Anatom- 
ische Gesellschaft. Curiously enough, however, the title of a paper (pp. 153- 
154) by Bardeleben, who signed the antimononym declaration of the ‘‘Nomen- 
clatur Commission’ (%147),is ‘* Ueber das Praefrontale und Postfrontale des 
Menschen’. I am not disposed to cite these two words as adjectival locatives 
and as precedents for fostcava etc. (2181); but they are excellent mononymic 
adjectives used as substantives, (#115) and they do not occur in the official list 
adopted by the committee of which Bardeleben was a member. 


2No such attempt is known to me. The very notion savors of ecclesiasticism 
rather than of science. At the most, individuals have set certain fashions, more 
or less commendable and permanent, while committees have made recommend- 
ations which even their own members may disregard when their information is 
increased or their views are modified. 


eS ee ee 


WILpvER, Neural Terms. 335 


while the abolition of the vast majority of time-honored terms 
has not been even hinted at in this country, I believe many 
anatomists here and also in England have recognized earlier and 
more fully than most of the Germans the existence of two con- 
ditions (S190, F) that are essentially modern, viz., (a) the 
enormous expansion of anatomic and physiologic knowledge ; 
(b) its general diffusion among the people." 

§271. Indeed, notwithstanding the declaration of conser- 
vatism above mentioned, it is not easy for me to conceive that 
all the members of the Anatomische Gesellschaft really antici- 
pate the retention of, ¢. g., ‘‘manubrium sterni,” 


ee 
2 corpus 
sterni 


and ‘‘ processus xiphoideus ”’ for praesternum, mesoster- 


WS 


num and xiphisternum, respectively; of ‘‘squama occipitalis ”’ 
for (os) supraoccipitale ; of ‘‘arcus zygomaticus” for zygoma ; 


” ’ 


of ‘‘latissimus dorsi,’’ ‘‘ biceps brachii ’’ and ‘‘ triceps brachii” 
; Pp 


for latissimus, biceps and triceps respectively; of ‘‘ processus ver- 
miformis”’ for appendix ; of ‘‘substantia corticalis”’ for cortex ; 
of ‘‘vena cava superior’’ and ‘‘vena cava inferior’, radix an- 
terior’? and ‘‘radix posterior’, for terms not dependent for 
appropriateness upon the erect attitude of the human body. 
§272. In the declaration of the Anatomische Gesellschaft 
($147), and in the warning of its oldest member (§208), it is 
intimated that between the American and German committees 
there already exists a terminologic crevice which further ad- 
vance upon our part is likely to convert into an ‘‘impassable 
culf.”” Taken by themselves, or in connection with the pass- 


1 For nearly ten years, at Cornell University, the members of the general 
classes in physiology, candidates for first degrees in Arts and Science, and num- 
bering from 150 to 180 in each year, have individually examined, drawn and 
dissected each the brain of a sheep. At the recent meeting of the American 
Society of Naturalists, I outlined (’96,) a plan for the commencement of 
practical studies of the brain in primary schools ; this in pursuance of the 
conviction expressed seven years ago: 

‘« Aside from prejudice and lack of practical direction as to removing, pre- 
serving, and examining the organ, there is but one valid reason why every child 
of ten years should not have an accurate and somewhat extended personal ac- 
quaintance with the gross anatomy of the mammalian brain; that obstacle is 
the enormous and unmanageable accumulation of objectionable names under 
which the parts are literally buried.” W. & G., ’89, 282. 


336 JouRNAL OF CoMPARATIVE NEUROLOGY. 


ages just referred to, it seems to me that Dr. Dwight’s closing 
words convey a similar gloomy impression, and that they pre- 
sent alternatives too widely divergent. 

$273. As may be seen from Parts IV and V, with the 
single exception of the German retention of anterzor and foste- 
rior ($$132, 192), between the German committee and the 
American committees that had reported prior to the three ut- 
terances referred to in the last paragraph, the actual differences 
were simply trivial. Even the list adopted by the American 
Neurological Association contains no unfamiliar term whatever." 

$274. It must be remembered also that only neural terms 
are here referred to. As well remarked by Pye-Smith (’77, 162) 
and by His (’95, 155), encephalic nomenclature stands most in 
need of revision and offers peculiar difficulties. With the other 
regions of the body the conditions and necessities are far sim- 
pler. Hence there is no probability that any action of Ameri- 
can committees respecting anatomic nomenclature as a whole 
could eventuate in the establishment of what could be regarded 
justly as a ‘‘separate standard.” A stronger phrase for the 
hypothetic contingency could hardly be employed were the dif- 
ferences between the two sets of names comparable with the 
distinctions between the metric system and the English weights 
and measures. 

$275. The address of Dr. Dwight contained no reference 
to what has already been accomplished or proposed by Ameri- 
can organizations. At that time, of course, the action of the 
American Neurological Association had not been taken. But 
the Association of American Anatomists and the American 
Association for the Advancement of Science (§$81-85) at vari- 
ous periods between 1889 and 1892, had adopted unanimously 
the recommendations of their three committees corresponding 
with the first five sections of the report of the A. N. A. (§80). 


1The allegation of Professor His that my individual ‘* proposals tend to cre- 
ate a language entirely new and for the most part quite strange,” has already 
been met (7204). In matters non-scientific a deliberate exaggeration of like ex- 
tent would probably receive a briefer and less euphemistic characterization. 


WILpvER, WVeural Terms. 337 


§276. Although the specific terms included in these 
recommendations are few, they exemplify all the commendable 
features of the German report. Indeed, I fail to discover in 
the latter any general statement, principle, rule or suggestion 
that had not already been set forth with at least equal accuracy, 
clearness and force in the writings of British and American 
anatomists prior to 1895. 

§277. Notwithstanding the small number of individual 
terms included in the American reports, the dates of appoint- 
ment of the committees, 1885, 1889, 1891, the representative 
nature of the terms, and the comprehensiveness of the general 
recommendations, all justify deliberate and independent action 
upon the part of anatomists in this country. Hence it is grati- 
fying to see Dr. Dwight’s indication of our duty in this regard. 
He evidently advocates neither heedlessness nor a servility 
that might merit the application of the following caustic com- 
ment in an English review of an American work: 


‘¢ Our authors are merely following the lead of a certain eminent 
German anatomist, it being a fashion with American scientific writ- 
ers (except a few who prefer a sort of scientific Volapiik! ) to follow 
pretty blindly the German scientific leads in the matter of nomenclat- 
ure, and this even to the extent of bodily adopting actual German 
words into a language which can already find two or three synonyms 
for almost any word it may be desired to translate. No doubt many 
English authors are also to blame in this respect, but the fact is none 
the less to be deplored.”? Mature, Aug. 13, 1896, 341. 


$278. It seems to me that in America the present condi- 
tions are particularly favorable to deliberate thought and inde- 
pendent: conclusion upon the subject of this article. The pro- 
fessors of anatomy in some of the larger medical schools are 
young and vigorous. Few if any are rightly to be reckoned as 
‘‘old,”’ or at any rate as too old to change their minds and 


1Histologic terminology was apparently referred to here; but I imagine 
that the remark might apply equally to my series of correlated names for one of 
the encephalic segments and some of its parts, viz., metencephalon, metacoelia, 
metatela, metaplexus and metaporus; see Table VII. 


°The writer of a letter in the Waszon for Oct. 8, 1896, declares that ‘‘ there 
is a reaction setting in in America against extreme Germanization, and that it 
has not come too soon.’”’ For a comparison of the national Az/age with the in- 
Abie proton, and a citation of Aristotelian precedents for the latter, 
see $212. 


338 JOURNAL OF COMPARATIVE NEUROLOGY. 


their modes of expression when occasions arise.’ In view of 
all the circumstances, the attitude appropriate for American an- 
atomists, desirous to cOoperate yet maintaining their independ- 
ence and self-respect, is indicated in the following lines of Lu- 
cretius : 

Judicio perpende: et st tibt vera videntur, 

Dede manus: aut si falsum est, adcingere contra. 


§279. Those anatomists who are either interested already 
in the improvement of nomenclature, or whose regard for their 
successors leads them to sacrifice some present time and effort 
in their behalf, are urged to read upon the subject, to reflect, 
to confer, and to correspond freely. So intimate is the rela- 
tion between verbal expression and mental operation that, even 
when we imagine ourselves above such weakness, criticism of 
the former too often means disturbance of the latter. Hence, as 
with other matters involving individual habit and preference, an 
actual interview may sometimes be less productive of good than 
a correspondence that eliminates more completely the personal 
element and affords opportunity for reflection and for consulta- 
tion with disinterested experts.’ 


§280. Those who may entertain? a not unnatural impa- - 


tience at the apparently slow progress made in this country, 
and who may even feel mortified when comparing the two score 
terms adopted by the American Neurological Association with 


1The following incident encourages the belief that such changes of both 
opinion and custom may occur at any age. While preparing the new edition of 
his ** Anatomy’? (’89), Leidy preferred central lobe or tsland of Keil; but later, 
at the age of sixty-six, as chairman of the committee on nomenclature of the 
Association of American Anatomists, he signed the report recommending zusu/a, 


2Nearly all my letters and ‘slips’’ (278, note) from anatomists and linguists 
in this and other countries have been preserved. Always instructive and often 
encouraging, the restraining and even destructive quality of some might have 
been endured with less equanimity at a personal conference; see 371, and ’gI, 4. 


3That such sentiment, if entertained, has not been communicated to me, 
either directly or indirectly, constitutes one of the many evidences of the tolerant 
and helpful spirit that has animated American anatomists in dealing with the 
confessedly perilous qnestion as to how independent thinkers may best commu- 
nicate with their fellows. 


Ee 


Wiper, Neural Terms. 339 


the forty-five hundred recommended by the Anatomische 
Gesellschaft, may well consider : 

First, the improbability that any competent. American 
anatomist could have been diverted from his regular duties long 
enough to accomplish what was so effectively done by the sec- 
retary of the Anatomische Gesellschaft. 

Secondly, the enormous advantage afforded by the complete 
list adopted by the Gesellschaft. Many dead or dying terms 
have been disposed of, and the ‘‘ decks have been cleared”’ for 
more efficient action. 

Thirdly, whatever precipitation, vacillation, and error may 
be condoned in individuals whom volition or circumstances may 
lead to assume untenable positions, organizations legislating in 
the interest of posterity should advance so slowly as to risk 
neither recession nor even deflection. The Germans themselves 
regard their comprehensive list, as a whole, as provisional. 
The American selections (§$80, 238) constitute, we may be- 
lieve, an immortal forty. 

§281. Were neural terms to be now devised de novo, the 
hippocamp would certainly receive some less fantastic designa- 
tion, and the great cerebral commissure would be much more 
likely to be called tvabs (a beam) than corpus callosum. But both 
callosum and hippocampus are embalmed, as it were, in several 
other names and they are not sufficiently objectionable to war- 
rant their revolutionary annihilation. The best we can do is to 
effect a tolerable compromise between the imperfect conditions 
that we have inherited and the ideal conditions that we should 
like to transmit to our successors. 7 

The anatomists of to-day have an opportunity of providing 
for the future while cherishing the past; of benefiting poster- 
ity without neglecting ancestors; of lightening the burdens of 
generations to come, while recognizing the value of what was 
done by the anatomical fathers: of erecting a terminologic 
monument in which the best of what has been is cemented by 
their own labors. 


340 JouRNAL OF CoMPARATIVE NEUROLOGY. 


Part IX, Bibliography. 

In addition to papers directly referred to in the text there 
are included some in which are used the simplified terms of 
description or designation recommended by the present writer. 
A single asterisk indicates a partial, two a thorough, adoption 
of the system. Had other than neurologic papers been in- 
cluded there would appear in the second category papers by 
J. H. Comstock, S. H. Gage, G. S. Hopkins and many others. 

The names of certain periodicals are abbreviated as follows : 


A. A. A., Proceedings of the Association of American 
Anatomists. 

A, A. A. S., Proceedings of the American Association for 
the Advancement of Science, 

A. N., Transactions of the American Neurological Asso- 
ciation. 

A. P. S., Proceedings of the American Philosophical So- 
ciety of Philadelphia. 

Handbook, Reference Handbook of the Medical Sciences, 
A. H. Buck, editor, g volumes, 1884-1893. 

J. C. N., Journal of Comparative Neurology. 

FN; Deka: of Morphology. 

J. N. M. D., Journal of Nervous and Mental Disease. 


AYERS, Howarpb, ’93.—Anatomical nomenclature. Letter to 
the editor. Science, XXI, 190-191, April 7, 1893. 

BAKER, FRANK, ’84.—The rational method of teaching anatomy. 
New York Medical Record, April 19, 1884, 421-425. Six figures. 

BarKER, L.F., ’96.—Concerning neurological nomenclature. 
Johns Hopkins University Bulletin, V11, 200-201, Nov.-Dec., 1896. 
*95.—The nomenclature of nerve cells. A. A. A., 1895, 


409-44. 

Barclay, JOHN, ’03.—A new anatomical nomenclature. O., pp. 
182. Edinburg, 1803. 

*KBROWNING, W., ’89.—Vessels of the brain. Handbook, VIII, 
231-244, I pl., 3 figs., 1889. 

oa ’93.—Vessels of the brain. J/dem, IX, 138-141, 1 fig., 
1893. 

*KCLARK, TRAcy E., ’96.—Comparative anatomy of the insula. 
J. C. N., VI, 59-100, 5 plates. 


i ee a 


Wiper, Neural Terms. 341 


CLELAND, JoHN, and Mackay, J. Y., ’96.—Human anatomy, 
general and descriptive. O. pp. 833, 630 figs., London and New York, 
1896. 

Cougs, E., and SHute, D. K., ’87.—Neuro-myology. Mew 
York Medical Record, XXXII, 93-98, 121-126, July, 1887. 

*DonaALDSON, H. H., ’90.—Anatomical observations on the brain 
and sense organs of the blind deaf-mute, Laura Bridgman. Amer. 
Jour. Psychology, III, 293-342, 2 plates. Sept., 1890. 

Dwicut, T., ’95.—Our contribution to civilization and to science. 
Address as president of the Association of American Anatomists, Dec., 
1895. Proceedings of the eighth meeting, 12-15. Also in Science, 
Ill, 75-77, Jan. 17, 1896. 

’ Epincer, L., ’93.—Vorlesungen iiber den Bau der nervésen 
Centralorgane, des Menschen und der Thiere. 4th ed., O., pp. 220, 
145 figs. Leipzig, 1893. (In Table V the reference to this work is 
incorrect). 

*kFIsH, P. A., ’90.—The epithelium of the brain cavities. Amer. 
Soc. Microscopists, Proceedings, 140-144, 1 plate, 1890. 


ay ’93, @.—The indusium of the callosum. /. C. W., III, 
61-68, 1 plate. 
ot ’93, 6.—-The terminology of the nerve cell. A. A. A., 


May; 1894.) Also. Covi EN 70075: 

ao ’95 —The nervous system of Desmognathus fusca. J. M., 
X, 231-286, 4 plates. 

ek *96.—A note on the cerebral fissuration of the seal 
(Phoca vitulina). A. A. A., 1895, 61-64, 1 plate. Alsoin J. C. W., 
VI, 15-18, 1 plate, March, 1896. 

Foster, F. P., ’88-’94.—An illustrated encyclopedic medical dic- 
tionary. Being a dictionary of the technical terms used by writers on 
medicine and the collateral sciences in the Latin, English, French and 
German languages. Q., 4 vols., Mew York, 1888-1894. 

*KGAGE, Mrs. S. P., ’93.—The brain of Déemyctylus viridescens 
from larval to adult life, and comparisons with the brain of Ama and 
Petromyzon. <‘“* The Wilder Quarter-Century Book,” 1893, 259-313, 
8 plates. 

AK ’95.—Comparative morphology of the brain of the soft 
shelled turtle (Amyda mutica) and the English sparrow (Fasser domes- 
tica). Proceedings Amer. Microscopical Society, XVII, 185-228, 5 plates. 

7% —’96.—Modification of the brain during growth. Ab- 
stract of paper read before the 4. 4. A. S., Aug. 24, 1896. Amer, 
Naturalist, XXX, 836-837, October, 1896. 


342 JOURNAL OF COMPARATIVE NEUROLOGY. 


GouLp, G. M., ’96.—Concerning medical language. Address at 
meeting of Amer. Medical Editors’ Association, May, 1896. <‘‘Bor- 
derland studies,” O., pp. 380, Phil., 1896. 

HENLE, J.,’79.—Handbuch der Nervenlehre des Menschen. 
<Handbook der Systematischen Anatomie des Menschen. 2nd. ed., 
O., pp. 666, 327 figs., Braunschweig, 1879. 

*HERRICK, C, J., ’91.—Studies in the topography of the rodent 
brain. Bull. Lab. Denison Univ., VI, 26-46, 22 plates, June, 1891. 

* °93, a@.—Recent advances in the study of the nervous 
system. Kansas Acad. Sci., Trans., 1893. 

‘93, 6.—Illustrations of the surface anatomy of the brain 
of certain birds. /. C. &¥., III, 471-176,: 1/plate, \Dec., 1893: 

a3 ’94.—The cranial nerves of Amblystoma punctatum. J. 
GinN., IV, 193-207, 2 plates. ‘Dee, 2894. 

*HERRICK, C. L., ’90.—The central nervous system of rodents, 
Preliminary paper. Bull. Sct. Lab. of Dentson University, V, 35-96, 
19 plates. June 1890. 

* ’91, a —Illustrations of the architectonic of the cerebel- 
lum. /. C. W., I, 5-14, 4 plates, March, 1891. 

* 91, 6.—Topography and histology of the brain of cer- 
tain reptiles. 7. C. WV., I, 14-38, 2 plates, March, 1891. 

* 91, ¢.—Topography and histology of the brain of cer- 
tain ganoid fishes <Contributions to the comparative morphology 
of the central nervous system. /. C. V., 149-182, 4 plates, June, 
1891. 

= ’91, d.—Contributions to the morphology of the brain of 
bony fishes. /. C. WV., I, 211-245, 333-358, 3 plates, Oct. and Dec., 
1891. : 

*, *91, e.-—The commissures and histology of the teleost 
brain. Anat. Anzeiger, VI, 676-681, 3 figs., 1891. 

* ’92, a.—Notes upon the anatomy and histology of the 
prosencephalon of teleosts. American Naturalist, 112-120, 2 plates. 
Feb., 1892. 

** ’92, 6.—The cerebrum and olfactories of the opossum. 
YG. JV., 11, 1-20, 3 plates.) Bebi,) 1e92. 

* ’92, c.—Studies on the brain of some fresh-water fishes. 
J.C. N., II, 21-72,:2 plates. May, 1&92. 

* ’92, d.—Histology and physiology of the nervous ele- 
ments. /..C..4V.,:II,:137-148, Dec.,. 1892. 

* ’92, e.—Embryological notes on the brain of the snake. 
J. C. N., Il, 160-176, 2 plates. Dec., 1892. 


Wiper, Neural Terms. 343 


o 92, £—Additional notes on the teleost brain. <Anat. 
Anz., VII, 422 431, ro figs., 1892. 

*k ’92, g.—Notes upon the histology of the central nervous 
system of vertebrates. <W&. Leuckhart’s ‘‘ Festschrift.” Pp. 278-288, 
2 plates. 1892. 

* ’93, a.—Topography and histology of the brain of certain 
reptiles. /. C. M., III. 77-106, 119-140, 11 plates. June, Septem- 
ber, 1893. 

* ’93, 6.—The callosum and hippocampal region in marsu- 
pial and lower brains. /. C. W., III, 176-182, 2 plates, Dec., 1893. 

* ’93, c.—Recent progress in the comparative anatomy of 
the nervous system. Handbook, 1X, 669-696, 78 figs. 

His, W., ’80"85.—Anatomie menschlichen Embryonen. Text, 

O. I, pp. 184, 17 figs.; II, pp. 104, 67 figs; ILI, pp. 260, 156 figs.; 
Atlas, F., XIV plates. Leipzig, 1880, 82, 85. 
’9s.—Die anatomische Nomenclatur. Nomina anatomica, 
Verzeichniss der von der Anatomischen Gesellschaft auf ihrer IX. 
Versammlung in Basel angenommen Namen. Eingeleitet und im 
Einverstandniss mit dem Redactionsausschuss erlaiitert. Archiv fur 
Anatomte und Physiologie. Anat. Abth., Supplement Band, 1895. 
O, pp. 180; 27 figs., 2 plates, 1895. 
’96 —Herr Burt Wilder und die Anatomische Nomencla- 
tur. Anat. Anzeiger, XII, 446 448, Oct. 30, 1896. 

HuGuENIn, G., ’79.—Anatomie des centres nerveux. Traduit 
par Keller; annoté par Duval. O., pp. 368, 149 figs. Paris, 1879. 

*k HUMPHREY, O. D., ’94.—On the brain of the snapping turtle 
(Chelydra serpentina.) J. C. N., IV, 73-116, 3 plates. 

Huxtey, T. H.,’73.—Manual of the comparative anatomy of 
vertebrated animals. O., pp. 431, 110 figs., Mew York, 1873. 

HyrttL, JosEPH, ’80.—Onomatologia anatomica. O., pp. 626, 
Wien, 1880. 

*kK rncsBurRY, B. F., ’95.—The brain of Mecturus maculatus. /. 
C. V., V, 138-206, 3 plates. 

KoELLIKER, A., ’g2.—Nervenzellen und Nervenfasern. Diol. 
Centralblatt, XII, 33-51. Jan, 30, 1892. 

96. Handbuch der Gewebenlehre des Menchen. Sechste 
Auflage. Zweiter Band. Nervensystem des Menschen und der 
Thiere. O., pp 874, 845 figs. Letpzig, 1896. 

Krause, W., EDITOR, ’79.—Specielle u. macroscopishe Anatomie 
<Handbuch der menschlichen Anatomie, von C. F. T. Krause. O., 
pp. 1054, 570 figs. Hannover, 1879. 


344 JouRNAL OF COMPARATIVE NEUROLOGY. | 


KRavsE, W., ’9t, a2.—Myologische Nomenclatur. O., pp. 4. 
Gottingen. June, 1891. (As secretary of the committee of the 
Anatomische Gesellschaft). 

’91, 6.—On anatomical nomenclature. Lritish Assoctation 
Report, 1891, Aug., 1891. Also Jntern. Monatsch. f. Anat. u. Phys., 
1892, IX. 

*LEIDY, JOSEPH, ’89.—Human anatomy. Second edition, O., 
PP. 950, 495 figs. Philadelphia, 1889. 

*MILLs, CHARLES K., ’86.—Arrested and aberrant development 
of fissures and gyres in the brains of paranoiacs, criminals, idiots and 
negroes. Preliminary study of a Chinese brain. Presidential address. 
A. N., 1886. J. N. M. D., XIII, Sept. and Oct., 1886, 37 pages, 
2 plates. 

Minot, C. S., ’92.—Human embryology. R. O., pp. 815, 463 
figs. Mew York, 1892. : 

ONvuFROWICS, ’87.—Das balkenlose Mikrocephalgehirn Hofmann. 
Arch. f. Psychiatrie, XVIII, 305-328, 2 plates, 1887. 

*Osporn, H. F., ’83.—Preliminary observations upon the brain 
of Amphiuma. Phila. Acad. Nat. Sci , Proc., 1883, 177-186, 1 plate. 

ss -84.—Preliminary observations upon the brain of Menop- 
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plate. 

*k —’88.—A contribution to the internal structure of the 
Amphibian brain. /. JZ, II, 51-96, 3 plates. July, 1888. 

Owen, R., ’46.—Archetype and Homologies of the vertebrate 
skeleton. British Association Report, pp 169 340, 1846. 
°63.—Monograph on the aye-aye. Zoological Transactions, 
V, 33-102, 12 plates. ; 
°68.—On the anatomy of vertebrates. III, Mammals. 
O., pp. 915, 614 figs. London, 1868. 

PaRKER, T. JEFFERY, ’82.—Notes on the anatomy and develop- 
ment of Scymnus lichia. New Zealand Institute Transactions, XXV, 
222-234, 2 plates. Read Aug. 15, 1882; issued May, 1883. 

’84.—A course of instruction in zootomy. D., pp. 397, 
72 figs. London, 1884. 

* °86.—On the nomenclature of the brain and its cavities. 
Notes from the Otago University museum. JVature, XXXV, 208-210, 
2 figs., Dec. 30, 1880. 

Pye-SmituH, P. H., ’77.--Suggestions on some points of anatom- 
ical nomenclature. Journal of Anatomy and Physiology, X11, 154-175, 
Oct, 5577. 


WILDER, Neural Terms. 345 


*Ricocs, C. E., ’90.—Editor of American edition of Edinger’s 
*“Vorlesungen.” O., pp. 230, 133 figs. Philadelphia, 1890. 

SCHWALBE, G., ’81.—Lehrbuch der Neurologie. <Hoffmann’s 
Lehrbuch der Anatomie des Menschen. O., pp. 1026, 505 figs. Er- 
langen, 1881. 

*®SORENSEN, A. D., ’93, a4. —The pineal and parietal organ in 
Phrynosoma coronata. J. C. WV., III, 48-50, 1 plate, June, 1893. 

* "93, 6.—Vhe roof of the diencephalon. 7. C. W., III, 
50-52, June, 1893. © 

*SPITZKA, E. C., ’81, a.—Letter on nomenclature. Science, (ed- 
ited by J. Michel), April 9, 1881. Alsoin 7. MV. MZ. D., July, 1881, 


661-662. 
* 


’81, 6.—Further notes on the brain of the sauropsida. 
Science, May 28, 1881, 254. 

* ’81, c.—Notes on the architecture of the oblongata. WJ. 
Y. Medical Journal, Sept., 1881, pp. 10, 1 plate. 

** 84, a.—The morphological status of the cerebellum. JV. 
Y. Medical Record, XXVI, III, July 26, 1884. 

of 84, 6.—Hippocampus vs. hypocampa. WV. Y. Medical 
Record, Sept. 20, 1884, 335. 

* "84, c.—Contributions to the anatomy of the lemniscus. 
NV. Y. Medical Record, XXVI, 393-397, 421-427, 449-451, 477-481, 
16 figs. Oct. and Nov., 1884. 

5 84, d.—Mittheilung, die angebliche Abwesenheit der 
Vierhiigeltheilung bei Reptilien betreffend. Veurol. Centralblatt, 1884, 


pp. 4. 
*. 


’86.—The comparative anatomy of the pyramid tract. 
Jour. Comp. Medicine and Surgery, Jan., 1886, pp. 47, 24 figs. 

* —'88.—The oculo-motor centres and their codrdinators. 
J. N. M. D., Aug., 1888, pp. 20, 6 figs. 

* ’89, a.—Histology of the brain. Handbook, VIII, 164- 
189, 40 figs., 1889. 

* ’89, 6.—Spinal cord. Handbook, VIII, 474-484, 14 figs. 
1889. 

* ”*90..—Remarks on the brain of the seals. Amer. Natur- 
alist, XXIV, 115-122, Feb., 1890. 

*KSTOWELL. T. B., ’82.—The vagus nerve in the domestic cat. 
A. P. S., XX, 123-138, 3 plates. Read July 15,1881; printed, March, 
1882. 

sas ’85.—[Outline of practical work in vertebrate zoology.] 
This was, as I recollect it, the substance of the title. My copy is lost; 


346 JOURNAL OF COMPARATIVE NEUROLOGY. 


the busy author has none, and cannot even recall the place of publi- 
cation, probably in an educational journal. 

Solera 86, a.—The trigeminus nerve in the domestic cat. A. 
P. S., May 21, 1886, 459-478. 

eK —’86, 6.—The facial nerve in the domestic cat. 4. P. 
S., Nov. 5, 1886, 9-19, 1 fig. 

ae —’88.—The glosso-pharyngeal, the accessory and the 
hypoglossal nerves in the domestic cat. A. P. S., XXV, pp. 89-104, 
3 plates, March 2, 1888. 

eK —’gt.~ The lumbar, the sacral, and the coccygeal nerves 
in the demestic cat. /. C. WV., I, 287-314, 1 plate. 

*STRONG, OLIVER S., ’95.—The cranial nerves of amphibia. //. 
M., X, 101-203, 6 plates, 1895. 

**STROUD, B. B., ’95.—The mammalian cerebellum. Part I, 
The development of the cerebellum in man and the cat. /. C. M., 
V, 71-118, 8 plates, 1895. 

*TicHT, W. G, ’90.—[As junior author with C. L. Heriekh 
The central nervous system of rodents. Preliminary paper. ulle- 
tin of the scientific laboratories of Denison University, V, 335-396, 19 
plates, June, 1890. | 

*TURNER, C. H., ’91.—The morphology of the avian brain. /. 
C. NV., I, 39-92 ; 107-132; 265-286, 9 plates, March, June, October, 
1891. 

Waters, B. H., ’91.—Additional points on the segmentation of 
the vertebrate brain. Zoolog. Anzeiger, 1891, 141-144, May 4, 1891. - 

WERNICKE, ’81.—Lehrbuch der Gehirnkrankheiten. O., 3 vols., 
Kassel, 1881. 

WHEWELL, W., ’40.—History of the inductive sciences. 1840. 

*WRIGHT, R. Ramsay, ’?84.—On the nervous system and: sense 
organs of Amiurus. Canadian Institute Proceedings, N.S., 352-386, 
4 plates. October, 1884. 

* ’°85.—Article ‘‘The anatomy of vertebrates.” <The 
Standard Natural History, III, pp. 52, 44 figs. Boston, 1885. 

W. &G., ’82.—[WitpErR, B. G., and Gacer, S. H.|—Anatomi- 
cal technology: an introduction to human, veterinary and compara- 
tive anatomy. 1882, O., pp. 600, 120 figs., 4 plates. [Chap. X, 
comprising more than roo pages, 4 plates and 16 figures, represents an 
original investigation upon the brains of /Vecturus and the Cat. ] 
’86.—The same, second edition, revised. 

’92.—The same, third edition, from the second. 


WILDER, Neural Terms. 347 


’°89.—Anatomical terminology. Handbook. VIII, pp. 24, 
2 figs,, 1889. 

Wiper, B. G., ’71.—Intermembral homologies, the correspon- 
dence of the anterior and posterior limbs of vertebrates [1871]. Bos- 
ton, Proc. Nat. Hist. Soc., XIV, 1872, pp. 154-188, 309-339, 299- 
420; 5 figs. . 

73, 6.—The outer cerebral fissures of Mammalia (especi- 
ally of the Carnivora), and the limits of their homologies. 4. A. A. 
Se M2873, (pt. 2) app) 214-2347; inoitis. 

73, c.—Cerebral variation in domestic dogs and its bear- 
ing upon scientific phrenology. A. 4. A. S., XXII, 1873, (pt. 2), pp. 
234-249; 6 fig. 

73, d@.—Lateral asymmetry in the brains of a double hu- 
man monster. 4A. A. A. S., XXII, 1873, (pt. 2), pp. 250-251; 4 fig. 
°73, k.—The pectoral muscles of mammalia. A. JA. A. 
aL 1873% pp.s05-707- 

75, c.—Notes on the American Ganoids. IV. On the 
brains of Ama, Lepidosteus, Actpenser and Folyodon. A. A. A. S., 
XXIV, 1875, pp. 151-193 ; 3 plates. 

°76, a@.—On the brains of fishes. . Phil. Acad. Nat. Sct. 
Proceedings, XX XVIII, 51-53, April 4, 1876. 

*76, 6.—Note on the development and homologies of the 
anterior brain-mass with sharks and skates. Amer. Jour. Science, 
Series LID Xi, roz-2055 1 fig.» 1876: 

76, ¢.—On the brains of some fish-like vertebrates. A. A. 
BAd 51); ) SEXO, LO 710, ep p. 259-259- 

*77, a.—On the brain of Chimera monstrosa. Phil. Acad. 
Proc., XX1X, May 29, 1877, pp. 219-250; 1 plate. 

80, 6.—The cerebral fissures of the domestic cat (élis 
domestica). Sctence, 1, 49-51, 2 figs. 


80, d.—The foramina of Monro; some questions of ana- 
tomical history. Boston, Med. and Surg. Jour., CIII, 152-154, 1880. 
’80, e.—The foramina of Monro in man and the domestic 
cat. 4. A. A. S.; WV. Y. Medical Record, XVIII, 328, 1880. 

80, f.—Partial revision of the nomenclature of the brain. 
A. A. A. S.; N. Y. Medical Record, XVIII, 328, 1880. 

"80, g.—The crista fornicis, a part of the mammalian brain 
apparently unobserved hitherto. A. A. A. S, 1880; WV. Y. Med. 
Record, XVIII, 1880, p. 328. 

’*81, 5—A partial revision of anatomical nomenclature, 
with ‘especial reference to that of the brain. Sczence, II, 1881, pp. 


348 JOURNAL OF COMPARATIVE NEUROLOGY. 


122-126, 133-138, March. Also 7. WV. WZ. D., July, 1881, 652 661. 
81, c.—How to obtain the brain of the cat. Science, II, 
1881, pp. 158-161. 

81, d.—The brain of the cat (Felis domestica). A prelim- 
inary account of the gross anatomy. A. P. S., XIX, 1881, pp. 524- 
562; 4 plates. Read July 15; published in December. 

’83, ¢.—On the brain of a cat lacking the callosum. [4. 
A. A. S., 1879]. A. WV. A., 1883. Amer. Jour. Neurol. and Psychi- 
atry, 1883, II, pp. 491-499: 4 fig. /. V. and M. D.,N.S., VIII, 
1883, p. 62 (Abstr.); Veurol. Centralblatt, 11, 517 (Abstr.) 

—’83, d.—On the alleged homology of the carnivoral Fissura 
cructata with the primatial /. centralis. A. N. A., 1883. J. WV. M. 
De MN. S., VIIL, 1883; -pp.°62-63. 

84, a.—Methods of studying the brain. The ‘‘ Cart- 
wright Lectures ” for 1884. MV. Y. Med. Journ., XXXIX, 1884, pp. 
141-148, 177-183, 205-209, 233-237, 373-377, 457-461, 513-516, 653- 
656; XL, 113-116; 64 fig. Abstractsin VV. Y. Med. Record, XXV, 
1884, Pp. 141-143, 197-199, 225-227, 365-367, 449, 450, 545, 546. 
84, 6.—Names and synonyms. Privately printed in con- 
nection with ’84, a. O., pp. 7, Feb., 1884. 

°84, c.—On encephalic nomenclature. A. WV. A., 1884. 
JANAGM. D., 1884, pp. 18, 50; abstract: 

*84, d.—Do the cerebellum and the oblongata represent 
two encephalic segments or only one? A. A. A. S., XXXIII, 1884. 
Science, IV, 1884, p. 341; WM. Y. Med. Journ., XL, 1884, p. 324. 
"84, ¢.—On some points in anatomical nomenclature. A. 
A. A. S., XXXIII, 1884. 

"84, —The existence and dorsal circumscription of the 
porta (‘‘ Foramen Monroi’’) in the adult human brain. A. 4..A. S., 
XXXII, 1884; WV. VY. Med. Journ., XL, 1884, p. 324. 

84, g.—The relative position of the cerebrum and the 
cerebellum in the anthropoid apes. A. 4. A. S., XXXIII, 1884. 
85, a.—The use of slips in scientific correspondence. Soc. 
Natural. East. U. S., 1884. Sctence, V, 1885, p. 44. 

85, 6—Encephalic nomenclature. I. Celian terminol- 
ogy: the names of the cavities of the brain and myelon. WV. Y. Med. 
Journ., XI, pp. 325-328, 354-357, Mar. 21 and 28, 1885; 8 fig. 

’85, c.—Paronymy versus heteronymy as neuronymic prin- 
ciples. Presidential address at the 11th annual meeting of the Amer. 
Neurological Assoc., 1885. A.W. S., J. NV. M. D., XII, pp. 21. 
Abstr. in Meurolog. Centralblatt, Dec. 15, 1885. 


WILDER, Neural Terms. 349 


85, d.—Educational museums of vertebrates. Address 
(as vice-president) before the Biological section of the Amer. Assoc. 
INGV Ob SCIENCE. (Ae ALANS. | XOX S553) pp. 19. Abstract in 
Science, Sept. 11, 1885, pp. 222-224. 

’85, ¢e.-—On two little-known cerebral fissures, with sug- 
gestions as to fissural and gyral names. A. WV. A., 1885. /. WV. W. 
D., 1885, XII, pp. 3. (Abstr. in LVeurolog. Centralblatt, Dec. 15 
1885.) 


F) 


85, f.—On a seldom-described artery (4. termatica), with 
suggestions as to the names of the principal encephalic arteries. A. 
BVCA TS85:. J. Vat PD: 1885. Kil pp) 2: \(Abstractsin 7.) Y- 
Med Journ. and NV. Y. Med. Record, June 27, 1885, and in Meurolog. 
Centralblatt, Dec. 15, 1885). 

°85, g-—The names of the encephalic arteries. WV. Y. Med. 
Jour., Nov. 28, 1885. 

85, 4.—Neuronymy. WV. Y. Med. Record, Aug. 1, 1885, 


p. 139. 

*86,a.—The collocation of a suture and a fissure in the 
human’ fetus: \/.\V. 47. D.,)'1886;: XULL, pp. 6, 1 fig. ( Abstracts 
in WV. Y. Med. Record, July 31, 1886, Science, Aug. 6, 1886, and Mea- 
wcal News, Aug. 7, 1886. 

’86, 6.—Notes on the brain. /. W. AZ. D., 1886, XIII, 
pp. 4. (Abstracts as in a.) 

~86, c.—Exhibition of the medisected, alinjected head of 
amundercr. A, WV.°At, 1886. J. NVM DL XIN, p.633, (Ab- 
stracts as in @.) 

86, d.—Remarks upon a living frog which was decere- 
brized more than seven months ago. 4. WV. A., 1886. /. NV. M.D., 
XIII, p. 30. (Abstracts as in a.) 

’86, e.—The paroccipital, a newly-recognized fissural inte- 
ger of. Vi: PD Oe. pp. ins, is figs../1 886: 

’86, f—The paroccipital fissure. Letter to the editor. 
WNW Wed. Record, Oct: 2, 1886) pp. ssoi3o0: 

86, g-——Human cerebral fissures, their relations and 
names and the methods of studying them. American Naturalist, XX, 
Pp. go1-902, r plate. Oct., 1886. 

°87, a.—The dipnoan brain. (Abstract of a paper on the 
brain of Ceratodus, with remarks unon classification and the general 
morphology of the vertebrate brain, read by invitation before the 
National Academy of Sciences, April 22, 1887). American Natural- 
ist, June, 1887, XXI, 544-548, 3 figs. 


350 JOURNAL OF COMPARATIVE NEUROLOGY. 


'87, 6.—Remarks on the classification of vertebrates. 
Amer. Naturalst, XXI, 913-917, Oct. 1887; (Abstract in 4. 4. A. S., 
Proc 1887, 251.) 

’87, c.—Correction of the foregoing. Letter to the 
editor. Amer. Naturalist, X XI, 1887, 1033. 

"88, a.—The relation of the thalamus to the paracele, 
(lateral ventricle). A. M. A.,'1888, 313-320. Also J. WV. M. D., 
July, 1889, XIV, pp. 436 443, 2 figs. 

—’88, 6.—The needlessness of corpus with striatum and of 
opticus with thalamus. Letter to the editor. /. WV. M. D., Sep- 
tember, 1888. 

’*89, a.—Brain, gross or macroscopic anatomy of. Hand- 
book, VIII, pp. 58, 104 figs. 1889. 

—’89, 6.—Brain, malformations of, which are morphologi- 
cally instructive. Same, pp. 6, Io figs. 

’89, c.—Brain, removal, preservation and dissection of. 
Same, pp. 7, 5 figs. 

—’89, d.—The relation of the thalamus to the paracele, es- 
pecially in the apes. A.A. A., 1889. (See also Note on p. 317 of 
88, a.) 


’89, ¢.—The heart as the basis of an intrinsic toponymy. 
Wied. A... TS86, p25. 

’90, a.—Do the Barclayan terms cause obscurity? Letter 
to the editor. Science, April 4, 1890, p. 224. 

?90, ¢.—Remarks on the brain of Chauncey Wright, with 
commentaries upon fissural diagrams. 4. MV. A., 1890. /. WV. M.D. 
Nov. 1890, pp. 2. : 
—’go, e.—Exhibition of diagrams of the brains and medisect- 
ed heads of man and achimpanzee. A. A. A. S., 1890. Abstr. in 
Amer. Naturalist, Oct. 1, 1890, 980. 
—’go, f£.—Exhibition of diagrams illustrating the formation 
of the human Sylvian fissure. A. A. A. S., 1890. 

—’go, g.—Owen’s nomenclature of the brain, with sugges- 
tions based thereon. A. A. A., 1890. 

—’go, 4.—Macroscopic vocabulary of the brain, with syno- 
nyms and references. Presented to the Assoc. of Amer. Anatomists, 
Dec..29, O., pp. 13, 1890. 

—’g1, a.—The fundamental principles of anatomical nomen- 
clature. Zhe Medical News, 708-710, Dec. 19, 1891. 

—’g1, 6.—The morphologic importance of the membranous 


Ls lO rh ee 


Se ee a, eS 


Wivper, Neural Terms. 351 


or other thin parietes of the encephalic cavities. 7. C. WV., I, 201- 
203, 189g. 


—’g2.—American reports on anatomical nomenclature, 
188g-1890, with notes as‘secretary. O., pp. 4, 1892. 
—’93, a.—The gross or macroscopic anatomy of the brain. 
Fandbook. Pp. 99-111; 10 figs. 1893. 
93, 6.—Methods of removing, preserving, dissecting and 
drawing the brain. /dem, pp. 111-121, 2 figs. 
93, ¢.—Meninges. (The envelopes or membranes of the 
brain and spinal cord). /dem, pp. 606-616, 11 figs. 
93, d.—The metapore (Foramen of Magendie) in man 
and apes. A. WV. A. Abstract in Phil. Med. News, Oct. 14, 1893. 
—’94, a.—Exhibition of a suicide’s brain with two pistol-ball 
wounds; remarks on its fissural anomalies. /. WV. JZ. D., Dec. 1894, 
pp. 4. Alsoin A. WV. A., 1894, 102-105, and WV. Y. State Med. Soc. 
Trans., 1874, 190 194. 
—’94, c.—Problems connected with the cerebral fissures. 
A. A, A., 1894, 32-35. 

——’95, 4.—The progress of paronymy. Letter to the editor. 
Science, May 10, 1895, 515-516. 
—g5, 6.—The paroccipital fissure, should it be recognized, 
and under that name? A. A. A., 1895, pp. 69-70, 2 figs. 
—’g95, ¢.—The cerebral fissures of two philosophers. A. A. 
1895, 71-73, 1 fig. 
—’95, @.—The frog was not brainless, but decerebrized. 
Letter to the editor. Sczence, June 7, 1895, 632. 
—’95, ¢.—Meuron, neuraxis, etc. Discussion of paper of F. 
Baker, ’95, a. A. A. A., 1895, 44-45. 
—’95, 7.—Dissection of the sheep’s brain: being part IV of 
‘¢ Physiology Practicums,” with plates XVIII-XXV and an append- 
ix on the removal and preservation of the brain. 1895. 
—’96, a.—‘‘ Pontine” or ‘‘pontile”? Letter to the editor. 
Medical Record, (N. Y.) XLIX, 823, June 6, 1896. 
—’96, 6.—The names epiphysis, conarium and corpus pineale : 
correction of an error. Letter to the editor. Sczence, N. S., N., 1896. 
—’96, ¢.—A matter of terms. Letter to the editor. N. Y. 
Medical Journal, April 11, 1896, 489. 
—’g6, d.—The dorsal sack, the aulix, and the diencephalic 
Bexure, A Ve, T8060.) fe At, 2. Aug, Tego. p. 54g." 7. C. 2V.; 
VI, June, 1896, 128. 


A. 


~ 


352 JoURNAL OF COMPARATIVE NEUROLOGY. 


————’96, e.—The ectal relations of the right and left parietal 
and paroccipital fissures. A. N., 1896. /. V. AZ. D., Aug., 1896, 
543.04. C. V.,; VI, June, 1896, 229. Abstr: in WV. VY. WMedwal ieee 
ord, July 11, 1896, 61. 
—’96, f.—Some neural and descriptive terms. Circular let- 
ter. Sctence, Dec. 25, 1896, 947. Also in WV. Y. Medical Journal, 
Jan. 2, 1897; Anatomischer Anzeiger, 1897. 
—’96 g.—The desirability and the feasibility of the acquisi- 
tion of some real and accurate knowledge of the brain by precolle- 
giate scholars. Amer. Soc. Naturalists, 1896. Sczence, 1897. 


Additions and Corrections. 


There are more items under this head than there might have been had it 
always been practicable to let me see a revised proof. Ordinary errors of typo- 
graphy and punctuation are not specified. 


es line 4: For VII, read VI. 
eye tz, Lue) Aue For VI, vead VII. 

P. 230, note 1; After 84, insert ’86. 

P. 230, note 2, last line, fifth word: For many, vead neural. 

P. 253, last line: “or collosum, vead callosum. 

2147, line 7: Mor Anzeizer, read Anzeiger. 

The next three items are trivial in themselves but necessary for the main- 
tenance of the accuracy of reproduction of the original as announced on p. 268, 
note 3. 

P. 269, fifth line from bottom : Coelia has a capital initial. 
P. 269, note 1: Between Medical and News isa period. The words be- 
tween 1890 and 1889 are italicized. The y of February should be omitted. 

P. 273: The section now numbered 577 should constitute part of 176; For 
9178, read 2177. 4 

. 275, seventh line from bottom: For poslcaval, vead postcaval. 

P. 279, seventh line: For diffussion, vead diffusion. 

P. 293, Table V, 30: For 89, 153, read ’93, 196. 

2235: For certain accidental omissions from the German list, see 7241, note. 

Pp. 304-307, head-lines: For ’82, read ’81. 

P. 306, Il, B: For Corum, vead Eorum. 

2239, e, line 7: For trapezoides, vead trapezoideum. 

P, 322, II, B, 4, Paracoelia: Of course the equivalency of this word for 
ventriculus lateralis involves the acceptance of the adjectival force of the prep- 
osition, para ; 7180-190. 

P. 326, Table VII, column 8, segment VI, trapezium: That this part is 
here included in the last segment, while in Tabie VI, V, B, 20, it is left in the 
fifth, as by the German committee, exemplifies the difficulty of assigning parts 
to these two segments. 

#254, last line: The section here referred to is on p. 274, not the one wrong- 
ly numbered 4178 on p. 273. 

7268, line 1: Omit the periods between BRIN PAS 

4268, line 5: After His, read (92; ’96). 

Hat note, line 5: After 96, insert ¢. 

- 340, Title 4: This should be the second under Baker, not Barker. 


JOURNAL OF COMPARATIVE NeEvuroLoGy, Vou. VI. PLATE XIV. 


FE. C. KENYON, PHOTOGR. 


JouRNAL OF COMPARATIVE NEuROLOGY, VoL. VI. PLATE XV. 


F. C. KENYON, PHOTOGR. 


JoURNAL OF CoMPARATIVE NEuROLOGY, VoL. VI. PLATE Saves 


F. C. KENYON, PHOTOGR. 


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PiatTe XVII. 


Jeunngs OF COMPARATIVE NEuROLOGY, VoL. VI. 


F. C. KENYON, GEL. 


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EIMERARY NOTICES: 


Effects of Alcohol on the Cortical Nerve Cell.) 


The tendency apparent of late to concentrate the attention upon 
limited problems of neuro-physiology and attempt their solution by the 
various improved methods now at our command is bearing fruit in 
investigations promising to give us definite and reliable data for pathol- 
ogy and psychology. Naturally none of these problems has greater 
practical and theoretical importance than that respecting the causes and 
character of cortical degeneration. 

Dr. Berkley has sought to contribute to the solution of this prob- 
lem by minute studies of the changes in the cortical cell produced by 
the long-continued administration of alcohol to rabbits. 5 to 8 cc. of ab- 
solute alcohol were fed to the animals and the dose continued until 
their death perhaps a year after the beginning of the experiment. The 
specimens were hardened in alcohol or Miiller’s fluid. Nissl’s method 
and hematoxylin-eosin staining for cellular structures and blood-vessels 
followed alcohol hardening and an original process, the Miiller’s fluid. 
(The reviewer can but express surprise that methods known to induce 
such shrinking and alteration in the cell bodies should have been 
employed for hardening). The process referred to is as follows : 

‘«'The cerebra are treated with Miiller’s fluid until the tissue is of 
sufficient consistency to admit of fairly thin sections. The portions of 
the brain selected are then cut into pieces not more than three milli- 
meters in thickness, and the slices are immersed in a mixture of 3 per 
cent. solution of bichromate of potassium, and 1 per cent. solution of 
osmic acid, in the proportion of too parts of the former to 20 parts of 
the latter. In this mixture the slices lie from three to five days, are 
then removed from the fluid, and slightly dried on filter paper to re- 
move any superfluous bichromate, are washed for a few minutes in a 
weak solution of silver nitrate, and then go into the second or staining 
mixture, which is made by adding two drops of a ten per cent. solu- 


IBERKLEY, H, J. Studies on the Lesions produced by the Action of Cer- 
tain Poisons on the Cortical Nerve Cell.—1. Alcohol. Brain, IV, 1895. 


ii JOURNAL OF COMPARATIVE NEUROLOGY. 


tion of phosphomolybdic acid to each 60 cc. of one per cent. argentic 
nitrate solution in distilled water. The second solution is made only as 
needed and at the moment before placing the brain tissue in it.” 

This method is said to avoid the perplexing fringe of brown about 
the impregnated cells and is rather a stain than a precipitation. It is 
said to be more reliable also. 

Unaltered cells fall into several groups: arkyochromic cells, with a 
reticular arrangement of the chromophilic particles ; stichochromic cells, 
with the particles in rows ; cytochromuc cells, with large nucleus and small 
cell body ; somatochromic cells, with much protoplasm and small nucleus. 
The cortical cells are chiefly somatochromic of the stichochromic va- 
riety. 

In the normal brain the contour of the vessels is regular and the 
perivascular spaces are narrow but distinct. The nuclei of the walls 
are sharply stained. In the alcoholic brains the small capillaries are 
shrunken and irregular. ‘The nuclei are swollen and absorb more stain. 
The perivascular spaces are enlarged and are either empty or partly 
filled with hzmatoidin debris. In the somatochrome cells the nuclei 
are altered in the disposition, size, and regularity of contour of the nu- 
cleolar chromophilic particles. 

By the silver method Berkley claims to show that all the pyramidal 
cells of the cortex and many of the irregular and dngular ones have 
short rectangular or oblique projections or gemmulz given off from the 
protoplasmic processes and that these are not as Kélliker supposed ab- 
normal or artifacts. Besides these the nerve cells have upon their 
branches a small, though variable number of varicosities. 

The neuroglia cells have nodosities in their fibres which must be 
carefully differentiated. Making all possible allowances for artifacts 
and variability the author feels justified in considering a considerable 
number of the appearances in the silver preparations of alcoholic brains 
abnormal. ‘The principal lesions are diminution in size, shrinkage of a 
vast majority of the cortical cells, disappearance of the gemmule, cer- 
tain swellings of the neurodendrites, and roughening of the stronger 
processes, and to some extent of the cell body. It is a question 
whether cells undergoing degeneration are as readily impregnated as 
the normal. Our own experience would lead to the conclusion that 
active processes, whether normal or pathological, favor reactions while 
an inert condition due either to fatigue or disease predispose to rejec- 
tion of the stain. 

Tumefaction of the neurodendrite and a loss of the gemmulz go 
hand in hand during degeneration. The method employed seems to 


a 


eee 


Literary Notices. ili 


be ill adapted to bring out the lesions of the cell body, though such an 
instance as that shown in Fig. rr proves that the changes are similar to 
those ordinarily seen in alcoholic dementia where the cell body is exca- 
vated or irregularly vacuolated. It is a pity that comparison was not 
made with cells stained with- hzematoxylin-sublimate-fuchsin or some 
histological stain after complete hardening in chrom-acetic or sublimate. 

In the cerebellum similar changes were found in Purkinje’s cells 
but of a more marked character. It is certainly placed beyond ques- 
tion by this study that alcohol, in common with other irritants, pro- 
duces a very definite destructive effect on the nerve cells and in particu- 
lar upon those of the kinesodic system. Gree Vee 


Structure of Nerve Cells after Electrocution.! 


The fatiguing effect of the prolonged action of weak electric cur- 
rents on the cells of the nervous system is well known through the re- 
searches of Hodge, Mann, and Vejas. Are these structural changes 
increased by the shorter action of much stronger currents, is the ques- 
tion which Dr. Fish set before himself for solution. The first speci- 
men examined, a portion of the cervical spinal cord of a victim of an 
electrocution at 1740 volts, exhibited a pronounced vacuolation of the 
nerve cells not to be accounted for by any known conditions prior to 
the execution. A second case in all essential respects similar and with 
absolutely fresh material gave, however, negative results, there being 
no observable lesions in the nerve cells of the same region of the spinal 
cord under the same method of preparation. The latter case is re- 
garded by Dr. Fish as the typical one and is verified by the results of 
experimental electrocutions on the lower animals performed by Dr. 
Krauss of Buffalo. Dr. Fish is inclined to the hypothesis that death 
by electrocution is the result of the fixation of the cells of the central 
nervous system, that is the cells are killed instantly in practically their 
normal relations. Ci jee 


Cortical Pathology of Permanent Dementia.’ 


Dr. Berkley has advanced a theory of the pathology of dementia 
which is based on the recent advances in our knowledge of the relation 


1FISH, PIERRE A. The Action of Strong Currents of Electricity upon 
Nerve Cells. Jour. Nerv. and Mental Disease, N. S., XXI, 1, Jan., 1896; Zrans. 
Am, Microscopical Soc., X XVII, 1896. 


*BERKLEY, H. J. A Theory of the Causation of Permanent Dementia. 
Medical News, 9 Nov., 1895. 


iv JOURNAL OF COMPARATIVE NEUROLOGY. 


of nerve cells and fibres in the central nervous system, the details of 
which Dr. Berkley has himself so successfully investigated. He calls 
attention to the fact that non-medullated nerve fibres are not known to 
occur to any great extent in the cerebral cortex and expresses the belief 
that the uncovered free endings of the lateral buds or gemmules of the 
‘‘ psychic” cells of the cortex are the media of communication from 
one neuron to another. These gemmule-bearing protoplasmic processes 
are the first to suffer from the ravages of disease, the cell body degen- 
erating later and the axis cylinder being affected last of all. ‘In 
short, therefore,” to quote from a recent editorial in the Journal of the 
American Medical Association, ‘‘ the theory of Berkely as to the pathol- 
ogy of dementia is as follows: The conduction of nerve stimuli to the 
cell corpus is through the medium of the lateral gemmules of the proto- 
plasmic processes ; that these are specially liable to injury from toxic or 
morbid influences, and are the first portions of the neuron to atrophy 
and disappear in certain diseases of the brain; that with their atrophy 
and consequent loss of function we have, first, confusion and incoér- 
dination of psychic functioning, and finally with any widespread de- 
generation of the cortical elements a permanent dementia ensues. His 
conclusions have been deduced partly from examinations of human 
brains and partly from experimental investigations on animals. He 
gives with his paper a reproduction of micro-photographs, showing the 
normal primordial process of a well educated man taken from an au- 
topsy immediately after death, and of a corresponding process from a 
subject of terminal dementia, showing the atrophy and absence of the 
dendritic gemmules.” Coaiagtts 


The Functions of the Frontal Lobes.! 


The satisfactory discrimination of the functions of the frontal lobes 
from those of other cortical areas is a matter of great difficulty and the 
attempts hitherto made have produced only ambiguous and conflicting 
results. Ferrier has decided that the frontal lobes preside over atten- 
tion while also presiding over motions of the eyes and head. Munk 
and Luciani consider this region as a part of the Fiihlsphzere or senso- 
rium and the motor centre of the dorsal muscles. Wundt and Hitzig 
from a theoretical standpoint assume that it is the centre of higher psy- 
chical functions. 

The author of the paper before us reaches similar conclusions and 
offers interesting experimental evidence which if not entirely convinc- 


IBIANCHI, L. Brain, IV, 1895. 


Literary WNNotices. Vv 


ing, is at least very suggestive. One must of course be on his guard 
against attributing to the loss of brain substance phenomena which may 
be due to the stimulating or depressing effects of operation or resulting 
encephalo-meningitis. . 

The details must be sought in the original but, in general, in the 
case of monkeys after ablation of both frontal lobes, the behavior is al- 
tered, the physiognomy is stupid and less mobile, the expression is _al- 
tered and devoid of flashes of intelligence, curiosity or sociability. 
There is no evidence of affection or gratefulness. Actions of purpos- 
ive character are liable to be left incomplete. ‘There is loss of cleanli- 
ness and discrimination. ‘The disposition is fitful and cruel. 

The author denies the existence of a special centre of inhibition or 
attention but concludes that the frontal lobes are seats of codrdination 
and blending of outgoing products of the several sensory and motor 
areas of the cortex. ‘‘ The frontal lobes would thus sum up into series 
the products of the sensori-motor regions, as well as the emotive states 
which accompany all the perceptions, the fusion of which constitutes 
what has been called the psychical tone of the individual. ‘The removal 
of the frontal lobes does not so much interfere with the perceptions 
taken singly as it does disaggregate the personality, and incapacitate for 
serializing and synthesizing groups of representations.” 


“ 


(CACTI 
Cerebral Localization. 


To the surgeon whose duty it is to diagnose and treat lesions of 
the central nervous system the present state of the theory of cerebral 
localization is perplexing enough. He sees some denying the fact of 
cerebral localization zz fofo and insisting that the cortex acts as a unit; 
others who recognize the cortical areas essentially as located in the text- 
books urge that the division of these areas into sensory and motor is a 
false division, for all of the areas are really sensory, the motor zones 
exerting no control over the muscles commonly associated with them, 
but only receiving sensations of muscular and general sensations from 
those regions of the body; and yet all of the time our leaders in sur- 
gical practice are operating successfully on the basis of the old charts 
constructed on the theories of Munk and Ferrier. In view of this sit- 
uation a recent paper by Dr. C. K. Mills! offers a few timely suggest- 
ions. We quote the latter part of the paper: 

‘‘ For the practical purposes of the physician and surgeon, no 


IMitts, C. K. Cerebral Localization in the Light of recent Pathologic 
Researches. Jour. Am, Med. Assoc., XXVI, 1, 4 Jan., 1896. 


vi JOURNAL OF COMPARATIVE NEUROLOGY. 


matter what view may be taken of the nature of the processes going on 
in the cortex, it would seem best to still hold to the view of the sepa- 
rate localization of areas for the special senses, for motion, and even 
for muscular and cutaneous sensibility. Lesions of these areas produce 
phenomena of vision, audition, motion, sensation, etc., which are not 
produced when the lesions are situated outside of the special areas to 
which the functions above mentioned are assigned. The fillet radia- 
tions for cutaneous and muscular excitations, as a compact bundle, 
probably reach, or most closely approach, the outer layer of the cortex 
in the postero-parietal convolutions and in the limbic lobe. Whether 
we should regard the cells and fibers which bring about communication 
between these regions and the motor cortex as true sensory terminals or 
as simply constituting a field of conjunction, the only cortical and sub- 
cortical lesions which will produce pure and marked sensory symptoms 
will be those occurring in these areas. ‘These incoming messages,’ 
says Andriezen, ‘ which inform the brain of the movement of the limb, 
arrive (strictly speaking ) not in the pre-Rolandic but in the post-Rol- 
andic (ascending parietal) convolutions. In the pre-Rolandic or 
ascending frontal convolution, and in the adjoining posterior portions 
of the three frontal convolutions as well as the prolongation of these 
areas on the mesial (marginal) convolution, we find the last term in 
the cortical series, the finally disposed executive mechanisms.’ 

“¢ Tt would perhaps be best to define the cortical area for cutaneous 
and muscular sensations, as that part of the cerebrum where the fillet 
radiations most nearly approach the surface of the brain, before their 
final ramifications in the molecular layer, still holding to the old view 
with reference to the motor cortex. Andriezen, as already stated, 
speaks of the pyramidal and ambiguous cells as the first sensory cells 
of the cortex, because the terminals of the fillet radiations, or their ex- 
tensions, first touch the apical processes of these cells, and therefore 
these cells first receive sensory impressions from the periphery of the 
body. It would be better, following Forel and Nansen, to disregard 
entirely the subdivision into cells of sensation and motion, and take 
the broad ground that we are simply dealing with the greatest and high- 
est of sensori-motor areas, and that in the region posterior to the area 
usually recognized as motor, the last stage in the sensory process is 
reached, while in the Rolandic cortex the first stage in the motor por- 
tion of the process begins.” 

Ci aes 


Literary Notices. vii 


Motor Functions of Dorsal Spinal Nerves. 


The question proposed in this investigation is, ‘‘ Do the dorsal 
spinal nerve roots carry functional motor fibres for the splanchnic mus- 
culature, on the one hand, and on the other hand, for the musculature 
of the bladder, which is also derived from the lateral plates?’ The 
investigations of van Wyhe and Hatschek have shown that this is true 
for selachians, Amphioxus and Ammoceetes ; does it also hold for the 
higher vertebrates ? 

The dorsal roots of the spinal nerves of the frog were electrically 
excited with a Du Bois Reymond’s apparatus and the results controlled 
by means of mechanical stimuli of these roots. Such stimulation 
evoked peristaltic and and anti-peristaltic motions of the digestive tract, 
and it was demonstrated that the successive pairs of roots correspond 
to distinct, though not sharply defined successive regions of the diges- 
tive tract. This result stands in pleasant harmony with the recent work 
of Sherrington and others on the segmental distribution of the cuta- 
neous nerves. ‘The innervation is bilateral and the reaction persists after 
the cessation of the stimulus. The time intervening between the begin- 
ning of the stimulus and the first noticeable reaction varies; it is never 
less than three seconds. ‘These reactions are independent of the vagus 
or vagus centres. Study of the ventral roots showed that no motor 
fibres go to the digestive tract from the spinal cord through any of the 
ventral roots except fibres to the rectum from the sixth and seventh. 
For the details of the connections of the several roots and of the very 
interesting relations of the nerves of the rectum and bladder the orig- 
inal must be consulted. The reader will notice that these results stand 
in harmony with the anatomical discoveries of Lenhossék and others 
who have described centrifugal fibres in the dorsal roots. 

Cae. 


Spiral Fibres in the Invertebrates.’ 


Ganglion cells are described in the ventral nerve chain of Htrudo 
medicinalis which are essentially similar to those known in the sympa- 
thic system of amphibians and reptiles. The spiral fibre breaks up 
into a reticulum upon the body of the ganglion cell and the author de- 
scribes this reticulum as consisting of two parts, one extra-cellular, the 


1 Srernacu, E. and Wiener, H. Motorische Functionen hinterer Spinal- 
nervenwurzeln. Arch. f. d. ges. Physiologie, LX, p. 539. 


2 Simon, C. Sur l’existence de la cellule a fibre spirale chez les invertébrés. 
Bibliographie Anat., 111, 6, Dec., 1895. 


viii JOURNAL OF COMPARATIVE NEUROLOGY. 


other intra-cellular. The former is regarded as a reticulum of origin, 
not of termination, the individual fibres apparently arising within the 
cell and uniting to form the spiral fibres, which is considered to be a 
cellifugal prolongation of the subjacent ganglion cell. On the other 
hand the intra-cellular reticulum is composed of filaments of the same 
optical properties as those which go to make up the greater process of 
the ganglion cell and the author suggests that they are the cellipetal fibres 
which on entering the cell body spread through it and form an intra- 
cellular superficial reticulum which in some way is to come into phys- 
iological relations with the extra-cellular cellifugal reticulum. We shall 
await with interest a more full description and confirmation of the pre- 
liminary account of relations which are certainly sufficiently remark- 
able to justify a little hesitation in giving them an unqualified accept- 
ance without the most rigorous proofs. 
C.Seike 


Structure of the Thalamus.! 


Corpus Luysit, or nidus hypothalamicus of man. This receives 
fibres from the tractus opticus, which come chiefly from Meynert’s com- 
missure. Apparently Gudden’s commissure also sends fibres into the 
nidus hypothalamicus. The latter receives other fibres from the lenti- 
cular nidus which penetrate it from the lateral aspect and also from the 
tegmental bundle of the caudatum. Finally there is to be mentioned 
a commissure of the nidi hypothalamici on the dorsal side of the most 
caudal portion of the mammillaria. 

Opticus termint. In the mouse, termini of the opticus fibres were 
found in the corpus geniculatum laterale, in the thalamus itself and in 
the corpus quadrigeminum anterius. In the two first mentioned the 
termini are like those described for the lobi optici of birds, that is, with 
much branched, compact terminal tufts which lie partly in the interior 
of the corpora geniculata lateralia and of the thalamus and partly in the 
stratum zonale of the latter. In the corpus quadrigeminum anterius, 
on the other hand, no such tufts were found but endings loosely 
branched over a larger area, which spread from the second white layer 
especially into the outer grey zone. In young rabbits were found in 
the layer of the tractus opticus laterally of the thalamus large cells 
which send their nervous processes centrifugally into the tractus. It 
was impossible to determine whether these fibres pass to Gudden’s com- 


1 KOELLIKER, A. vy. Zum feineren Baue des Zwischenhirns und der Regio 
hypothalmica, Verh. Anat. Ges., IX Vers., 1895. 


Literary Notices. ix 


missure or whether they represent the centrifugal fibres found by Cajal 
in the retina. 

Coronal fibres of the thalamus. Investigation of young mammals 
shows that these fibres in the main present terminal arborizations in the 
thalamus. ‘The optic radiations, however, undoubtedly terminate in 
the cortex, as their cells of origin lie among the ends of the opticus 
fibres. 

Fasciculus thalamo-mammillarts, s. Vicg @ Azyr. In new born and 
young mice it was clearly shown that the fibres of this bundle exhibit 
terminal arborizations in the nidus dorsalis thalami. This bundle, which 
according to the researches of Gudden has nothing to do with the pil- 
lars of the fornix, must therefore take its origin in the cells of the cor- 
pus mammillare—according to Gudden in the caudo-ventral nidus. 

Pedunculus corporis mammillaris of the rabbit. The fibres of this 
bundle undoubtedly arise in the large-celled lateral nidus of the corpus 
mammillare. Of their termini nothing has hitherto been known. The 
author finds, however, that they end cephalad of the pons in the re- 
gion of the caudal end of the ganglion interpedunculare near which 
they lie, surrounding dorsally and in part also penetrating a round nidus 
already described by Gudden as lying behind the trochlearis nidus in 
the central grey, the ganglion dorsale tegmenti of K6lliker. Another part 
of these fibres ends in the central grey which surrounds this nidus. 
From this nidus and from the central grey around it there arises the 
dorsal longitudinal bundle of Schiitz which Kélliker has called the dorsal 
grey longitudinal bundle, and this may be easily followed in longitud- 
inal sections along the dorsal side of the fourth and third nidi and in a 
curved course along the floor of the third ventricle. 

Columnae fornicis of the rabbit. These tracts plainly pass only 
through the lateral part of the corpus mammillare and end crossed be- 
hind and on the dorsal side of the latter. The author has followed these 
fibres to the nidus of the oculo-motorius, the posterior commissure and 
the nidus ruber, though these relations are not definately established 
and the real termini he is not at present prepared to state. 

Ganglion habenulae of mammals. The fasciculus of Meynert 
arises in the ganglion habenulae of the opposite side in free non-med- 
ullated arborizations. The fasciculus of Meynert contains fine and 
coarse fibres, of which the latter possibly pass above the ganglion into 
the pons. In the ganglion interpedunculare arise the medullated fibres 
discovered by Ganser which pass in two bundles ventro-dorsally and 
end in the ganglion tegmenti dorsale and in the adjacent central grey. 

Stria medullaris of the rabbit. This ends for the most part in the 


x JOURNAL OF.CoMPARATIVE NEUROLOGY. 


ganglion habenulae and derives its fibres, (1) from the fornix and the 
Ammonshorn ; (2) from the basal part of the third ventricle, from the 
regio supra-optica, where the fibres in question arise in a large ganglion 
which is connected with the nidus of the basal bundle of Ganser; (3) 
from the stratum zonale of the thalamus; (4) from the interior of the 
thalamus. ‘The stria thins out from before backward and behind passes 
over into the commissure of the pineal, from which a small number of 
fibres enters into the pineal. This commissure is a union of the 
habenulae. 

Fasciculus longitudinalis dorsalis of Kélliker. The so-called dorso- 
median fasciculus ends in the mammals cephalad of the nidus III in 
ascending fibres which in part form a commissure, in part apply them- 
selves to the medullary zone on the medial side of the nidus ruber. 
A nidus of this bundle such as was found by Van Gehuchten in fishes 
cannot be demonstrated in the mammals. 

C.gpeee 
The Lateral Line System of Amphibia. 


Dr. Kingsbury has performed a service which will be appreciated 
by many students in several departments of research in mapping the 
exact distribution of the lateral line organs of many of our types of 
tailed Amphibia, as well as of some of the allied forms. Descriptions 
and figures of the following species are given, JVecturus maculatus, 
Amblystoma punctatum, Protopterus annectens, Amphiuma means, Gyrin- 
ophilus porphyriticus, Diemyctylus viridescens, Siren, Rana, Cryptobranch- 
us allegeheniensts, Lepidosiren paradoxa, as well as comparisons with 
several other types. The histology of the neuromasts ( Nervenhiigel ) 
is briefly discussed. We quote a few paragraphs. 

‘«’The Amphibia afford in certain respects peculiar opportunities 
for the study of a sensory system associated with existence in the water. 
This is due to the fact that there are here included forms purely aquatic 
-and forms as purely terrestrial in their habits of life, and yet others 
which spend a portion of their life in the water and a portion of it on 
land. In every family of the tailed Amphibia native in the United 
States the system has been found, and in five families of the tailless 
Amphibia. Since Malbranc has found the sense organs in a larval 
Pipa, and Leydig ina larva of the viviparous Salamandra atra taken 
from the oviduct of the mother, doubtless the system will be found in 


1 KinGspury, B. F, The Lateral Line System of Sense Organs in some 
Amphibia, and Comparison with the Dipnoans. Zvans. Am. Microscopical Soct- 
ety, XVII, 1896. 


Literary Notices. xi 


a more or less perfect state of development in all Amphibia at some 
period in their life-history. 

‘‘In the urodela the distribution may readily be reduced to the 
following type: Upon the body, three lines, a /ateral continuous or 
not continuous with an occipital group, though not continuous with the 
orbital lines; a ventral line extending from under the arms in the pec- 
toral region to near the hind legs; a dorsal line somewhat closely con- 
nected with the lateral at its cephalic end and seldom extending as far 
as the level of the vent. Upon the head, a series extending from be- 
hind the eye, above and below it to the snout, the swpra- and infra- 
orbital lines; a line upon the lower lip, the ova/, connected with the in- 
fra-orbital by the angular ; a line from the angle of the mouth to the 
lateral corner of the head and there meeting a diverging line upon the 
ventral side of the head, and, when this is sufficiently developed, a 
line or trend of organs upon the side of the head; these the jugular, 
gular and postorbital lines of the descriptions and figures. 

‘*Comparison with other Ichthyopsida may not be of much value ; 
however, the distribution approaches most nearly that in the Dipnoans, 
then in Elasmobranchs ; among the latter Chlamydoselachus, apparently, 
in the greater extent of the gular line, shows most resemblance to the 
Amphibia. 

‘*The significance of the arrangement into groups is apparent 
when the system is examined in the larva at different periods of devel- 
opment. Evidently, as has already been maintained by Malbranc, 
each group sprang from a single organ by repeated fission in the same 
plane. His figures and my own observations clearly show that such is 
the case, as illustrated by Fig. 45. Exactly how this takes place, how- 
ever, is unknown. Whether the sensory cells may arise from the sup- 
porting cells, or from sensory cells alone, and supported from support- 
ing cells or from ordinary epidermal cells, yet awaits solution. 

‘* Malbranc called attention to the often recurring arrangement of 
groups upon two coordinates perpendicular to each other, or nearly so 
(as in the gular line), pointing out the physical advantage in such an 
arrangement in perceiving the direction and strength of a vibration in 
the water, should such be their function. In Ichthyopsida, in which the 
sense organs are deeply sunken in canals, the pores often become many 
times divided. In Ama, Allis found that the primitive pores divide 
quite regularly in a certain plane for a number of times; these second- 
ary pores again often divide ina plane at an angle tothe first, generally 
a right angle or nearly so, reminding us of the groups in Amphibia. 
In forms, then, in which the sense organs are confined in canals, this 


xii JOURNAL OF COMPARATIVE NEUROLOGY. 


division of the pores would seem to represent a potential division of the 
sense organs, which in forms in which the sense organs are freely situ- 
ated, as in Amphibia, can be actual. 

“« Attention is called to the necessity, imposed by the life habits 
of certain urodeles, ¢. g., Diemyctylus, for the neuromasts to live over 
a period of terrestrial existence, which is accomplished by the protec- 
tion of the organs by a growth of epidermal cells. Doubtless this is 
also true for many other forms of semi-aquatic habits of life, e. ¢., 
Desmognathus. In certain other urodela, e. g., Salamandra, and | be- 
believe Plethodon (if they exist at any time), the system perishes entire- 
ly in the adult. ‘This is also the case apparently in all the Anura, 
though in Aana it persists until after both legs and arms are well devel- 
oped and the tail has begun to be absorbed. ‘There would seem, then, 
to be something other than an aquatic existence necessary for the main- 
tainance of the neuromasts, since Rana catesbiana is more purely aquat- 
ic than several of the Salamanders in which the system persists. Of 


the mode of final disappearance nothing is known.” 
C.J 


Brains of Sauropsida.' 


The large number of carefully prepared descriptive papers now 
issuing from the Cornell University laboratory of neurological research 
is an occasion of congratulation not only to the directors of this labor- 
atory but to morphologists in general. It is everywhere recognized 
that the greatest obstacle now in the way of the best morphological 
work is the lack of sufficiently full knowledge of the exact anatomical 
structure of the types under investigation. 

Studies like the one now under consideration with their detailed 
descriptions and full illustration should do much to check the prevalent 
tendency to morphological speculation by supplying such a basis of 
exact knowledge as will render possible the more satisfactory determi- 
nation of morphological fact. 

Mrs. Gage has chosen for study the soft-shelled turtle and the 
sparrow, not because they represent generalized types of Sauropsida, 
but on the contrary because they represent extremes of specialization 
of the two great divisions of this group. ‘This is based on an apprecia- 
tion of the importance of comparing through all stages of development 


1GAGE, SUSANNA PHELPS. Comparative Morphology of the Brain of the 
Soft -Shelled Turtle (Amyda mutica) and the English Sparrow (Passer domes- 
tica). Trans, Am. Microscopical Soc., XVII. 1896. 


Literary Notices. Xili 


widely different forms of brains in eadee to gain from exaggerated form 
and specialized function more light upon the truths of morphology and 
evolution, a principle which might profitably be employed more often 
than has been customary. 

The nature of the paper is such as hardly to admit of adequate 
summary. Coen 


Trophic Nervyes.' 


1. The functional influence of a nerve cell on an adjacent cell is 
always of a trophic nature, either catabolic or destructive, with aug- 
mentation of tonus [contraction], or anabolic, restorative with diminu- 
tion of tonus [relaxation]. 

2. The section of a nerve, however carefully executed, always 
provokes in that nerve an irritative process which is feeble, it is true, 
but of long duration. 

3. This irritation of degenerescence, due to the section, evokes 
variable consequences varying with the functional nature of the nerve. 
If the nerve conducts catabolic impulses, it produces in the cell with 
which it is in contact destructive phenomena, long continued and Bet 
manent; the cell atrophies. 

4. If on the contrary the cell transmits anabolic impulses [inhib- 
itory nerves, dilators], it produces in the cell with which it is in contact 
feeble but repeated and permanent plastic processes, which cause cellu- 
lar hyperplasia with karyomitosis. In this case too there is little increase 
of function. 

5. If the nerve sectioned contains both anabolic and catabolic 
fibres, we find both processes side by side. 

6. ‘The existence of trophic nerves in the sense just indicated can 
no longer be reasonably denied. 

— Revue Neurologique. 
Ampullo-oculo-motor Connections.’ 

The anatomical relations between the vestibular and the oculo- 
motor nerves so far as known are indicated in the accompanying dia- 
gram which we copy from Dr. Bonnier’s paper. ‘The diagram shows 
the direct connections between Deiter’s nidus and the internal nidus on 
the one hand and the abducens nidus on the other hand ; also the indi- 
rect connection between the former and the latter through the superior 


1 WINKLER. Les nerfs trophiques. Vandre medicale, 2 May, 1895. 


? BONNIER, PIERRE. Rapports entre l’appareil ampullaire de l’oreille interne 
et les centres oculo-moteurs. Rev. Meurol., III, 23, 15 Dec., 1895. 


xiv JouRNAL OF COMPARATIVE NEUROLOGY. 


olive. It shows further the tract of Duval and Laborde which puts 
into direct connection the abducens of one side and the nidus of origin 
of the fibres of adduction [oculo-motor fibres] of the opposite side, 
thus providing the mechanism of the conjugate movements of the eye- 
balls in the horizontal plane, and also the connection via the dorsal lon- 
gitudinal fasciculus between the sixth nidus and the fourth and third. 


Accom. distance. 
Accom. intensity. 


Adduction. 


_—— Elevation. 


Depression. 
———- Lid. 


Inferior oblique. 


_—-— Superior oblique. 


NN. Bechterew. 


NV. tnternus. 
NV. Deitters. 


Abducens. 


| Superior olive. 


tc a cee VIII nerve. 


Now the author seeks to correlate these anatomical facts with the 
results of some experimental work and a large series of clinical obser- 
vations. De Cyon first called attention to the oculo-motor disturbances 
associated with lesions of the labyrinth, a fact with which all experiment- 
ers on the labyrinth have been forcibly impressed. The most frequent 
phenomenon is nystagmus, though all disturbances may be met for they 
are all observed in the clinic. 

De Cyon was of the opinion that the directions of the oscillations 
of the eyeball were determined by the choice of the canal excited. 


Literary Notices. XV 


Breuer called attention to the inertia of the eyeball by which on a sud- 
den movement of the head the eyeball would be displaced in its orbit, 
thus interfering with the precision of the visual judgment of the actual 
angular displacement of the head. ‘The nervous connections with the 
ampullae are for the purpose of correcting this displacement of the 
eyeball by the contraction of the appropriate muscles, the latter being 
reflexly excited by the head movement as registered in the vestibular 
sense organs. It is movements in the horizontal plane which are chiefly 
to be compensated and it is a significant fact that it is the abductors 
and the adductors of the eyeball which are in most intimate connection 
with the vestibular apparatus. 

The view of Mendel that vertigo is an oculo-motor disturbance is 
combated ; on the other hand it is regarded as in every case due to an 
irritation of the connections of the eighth nerve. No doubt titubation 
can produce vertigo, but the reverse is the rule, and the fact that we 
see objects oscillating is a sort of reflex titubation of the eyeballs sec- 
ondarily induced by reason of the intimate relations existing between 
the muscles of the eyeballs and the vestibular connections. 

Upon the irritation of the labyrinth the abductor of the same side 
and the adductor of the opposite side are excited and the eye quickly 
turns toward the side of the irritation; the elasticity of the opposing 
muscles returns the eye to its normal position, but more slowly so that 
the only disturbance of sensation which results is an apparent rotation 
of the visual field in the direction opposite to the latter motion. In 
other words objects appear to move toward the side of the irritation. 
This is a fact which the author has found to be of the greatest value in 
clinical practice. Vertical nystagmus is very rare. The author has 
observed two cases of which the origin was purely labyrinthin. The 
details of several cases of ocular disturbance occasioned by both laby- 
rinthin and bulbar lesions are given and at the close the author con- 
cludes: ‘‘Ampullar disturbances can find their symptomology in all kinds 
of oculo-motor disturbances, and in the presence of the latter it should 
be remembered that after the retina itself it is the labyrinth and par- 
ticularly the ampullae which are concerned with oculo-motor functions 
as well as with equilibrium. All of the oculo-motor nidi, with the ex- 
ception perhaps of that of the oblique, which I have never seen in- 
volved, may thus be affected by reflex irradiation issuing from the 
ampullar apparatus.” 

These results of Bonnier agree also remarkably well with those ob- 
tained by Lee in his experiments upon the dog fish (Galeus Canis). 

Ce Je He 


Xvi JOURNAL OF COMPARATIVE NEUROLOGY. 


The Initial Stages of the Degeneration of Nerve Fibres.? 


The phenomena attending the degeneration of the nervous tissues 
have long been matters of the highest interest to the pathologists, and 
especially since the recent studies on the regeneration of nerve fibres 
any facts bearing on the histological processes involved in degeneration 
have an added interest. Numerous pathologists have described the 
changes observed in the structure of the nerve fibre in the various cen- 
tral and peripheral neuroses and now Dr. Klippel coGrdinates these 
morbid processes and finds that they may be resolved into a single pro- 
cess which is essentially the same in all cases. 

The first stage in the degeneration of the nerve fibre is apparently 
a remarkable hypertrophy of the nerve tube. It is not, however, a true 
hypertrophy, but a tumefaction, the first stage of a lesion essentially 
destructive. The myelin becomes hyaline and loses in the axial por- 
tion next to the axis cylinder the concentric appearance so characteris- 
tic of the normal fibre. This portion also becomes granular and stains 
more intensely than the normal myelin, though not so dark as the axis 
cylinder. The fact that this disintegration of the myelin begins ax- 
ially and not peripherally indicates that the morbid process has its ori- 
gin in the axis cylinder. And in fact simultaneously with the changes 
just described the axis cylinder also exhibits hypertrophy with a pecu- 
liar change of form. In transection it appears no longer as a circular 
dot in the centre of the sheath but as a greatly enlarged and variously 
shaped figure, a rod, a spiral, a circle or a sigmoid. In tracing a fibre 
from section to section this alters in form showing that the contour is 
flexuous and that the fibre is still in a state of degenerescence. Later, 
in the penultimate stage of the process, granular fragmentation occurs 
to be followed by the final complete resorption. 

The degenerative process of the nerve fibre may then be divided 
into three stages, the swelling and deformation of the axis cylinder with 
the figures just mentioned, the granular disintegration and fragmenta- 
tion of the axis cylinder together with the liquefaction of the myelin 
from the centre toward the periphery, and finally complete resorption. 

In diseases like general paralysis the cells of the cornua of the 
cord are affected ina manner strictly analogous. The study of the 
early stages of this process is of course attended with the difficulty aris- 
ing from the scarcity of material for histological examination taken at 
the proper stages of the disease, yet a goodly number of cases are on 


1 Kipper, M. Comment débutent les dégénérescences spinales. Arch. de 
Neurologie, 2 Serie, I, 1, Jan., 1896. 


nn EEE 


Literary WNottces. XVii 


record, sufficient to show that the process is essentially the same in both 
the acute and the chronic degenerative neuroses. ‘The chief difference 
between the acute and the chronic cases is that in the initial stages of 
the former the tumefaction is more pronounced than in the latter case. 

Though the number of these degenerative neuroses is consider- 
able, yet the lesion of the nerve fibres involved is essentially the same 
for them all. Copy ne 


Cortical Olfactory Apparatus. 


Mr. G. Elliot-Smith continues his contributions to the morphology 
of the smell centre.!_ He notes the simplicity of arrangement of these 
centres in non-placental mammals and and the similarity, already in- 
sisted on by the reviewer, to that in Sauropsida. 

The hippocampus (cornu Ammonis, subiculum and fimbria) forms 
in a typical early mammal the dorsal margin of the whole extent of the 
fissura choroidea. The ventral margin of the cerebrum in the same 
region is formed by the pyriform lobe. The prosencephalic part of the 
olfactory bulb is continued caudad as a short peduncle, which almost 
immediately divides into the ventro-mesal tuberculum olfactorium and 
a lateral pyriform lobe. The tuberculum (our post-rhinal lobe) is de- 
scribed as we have described it in the opossum and rodents. ‘‘ These 
three parts—hippocampus, pyriform and tuberculum olf.—together 
with the precommissural area [our intraventricular lobe] and ‘ septum 
lucidum’ constitute the smell centre.” ‘‘ All the rest of the cortex 
may be distinguished as ‘ pallium.’ ” 

If this distinction is adhered to of course in such cases as Perameles 
the pallium becomes greatly reduced. ‘This prepares us for Brill’s as- 
sertion that in Sauropsida the ‘‘ pallium” disappears. (Probably in 
no other branch of science than neurology is there such a felicity of 
whimsicality as that which leads authors to appropriate a word and then 
supply it with a modified connotation or alter its application and then 
permits them to use a discrepancy of their own creation to discredit 
the unfortunate coiner of the word.) 

It seems to the writer that if it could be shown that every spot on 
the superficial aspect of the cerebrum had been encroached upon by 
cells connected with the smell centres it would not be necessary to re- 
ject those areas for that reason from the category of ‘‘ cortex” or of 
“ pallium.” 

Do we not tend to undervalue the plasticity of the brain? In the 


' Anat. Anzeiger, XI, 2. 


xviii JOURNAL OF COMPARATIVE NEUROLOGY. 


tuberculum and intraventricular lobe we have an entirely different his- 
tological structure from that of the cortex. In relation with the irregu- 
larly grouped polyhedral cells of this region fibres from the bulb ter- 
minate. Here we have every morphological and physiological reason 
for assuming an infra-cortical station. All analogy requires it and all 
the histological appearances are in accord. Processes from these cells 
ascend to their cortical fields in the hippocampus. ‘The hippocampus 
is just as much a part of the cortex as the temporal lobe! 

‘The olfactory peduncle, tuberculum olf. and pyriform lobe are 
closely connected with the other hemisphere by means of the anterior 
commissure. The precommissural area (intraventr. lobe) is connected 
with the other side by a part of the hippocampal commissure, which 
Herrick calls ‘ corpus callosum.’ ” 

By means of a very strong fibre system situated in the substance 
of the tuberc. olfact. and in series with the internal capsule fibres, the 
olfactory lobe (possibly the bulb?) is intimately connected with the 
lower parts of the nervous system. In Perameles most of them enter 
the pes, a few end in the mammillary region. 

Mr. Smith, like Debiere, excludes the callosal gyrus from the 
“limbic lobe.” Interesting details respecting the fornix fibres are also 
given in the same paper. 

In the second paper by the same author! we note with gratifica- 
tion the tendency toward substantial agreement among different authors 
respecting homologies which have given so much trouble. In this re- 
spect the study of the Ornithorhynchus and lower marsupials has been 
of great help. The long neglected hippocampal commissure seems to 
be coming to its own though even Mr. Smith seems not to be aware of 
what has been more lately done in its study among Sauropsida. 

Smith finds that the cephalic part of the dorsal commissure ends 
in the intraventricular lobe but freely accords to it an independent ex- 
istence, which is a distinct gain. It is a matter of very subsidiary im- 
portance whether this cephalo-dorsal commissure of infra-placentalia is 
a homologue of the callosum. ‘The best way to show conclusively that 
it is not would be to find it present in a mammal also possessing the 
callosum. If its fibres do not pierce the cortical areas at all a strict ho- 
mology would perhaps be destroyed. Nevertheless it would be an un- 
usual method in nature for fibres to break from one external (median) 
aspect of one hemisphere and break into a corresponding aspect of the 


1 Notes upon the Morphology of the Cerebrum and its Commissures in the 
Vertebrate Series. Anat. Anz., XI, 3. 


Literary Notices. XixX 


other. Callosal fibres have developed along some pre-existing route of 
connection and then attained their present position in obedience to the 
usual laws of developmental adjustment. Smith says that the fibres 
‘‘probably belong to the lamina infra-neuroporica and supersede the 
cephalic part of the fornix commissure whose position they usurp.” 
‘‘The corresponding region of the hippocampus [in higher mammals] 
disappears and the supracallosal gyrus of Zuckerkandl is all that re- 
mains of this region in the Eutherian brain.” 

If this suggestion as to the origin of the callosal fibres could in 
any way be verified the last serious obscurity in this problem would 
seem to be solved. 

In a still more recent paper on ‘‘ Jacobson’s Organ and the Olfac- 
tory Bulb in Ornithorhynchus”! the same author corrects some very 
serious blunders in Dr. Hill’s paper in the Philos. Trans. 1893, whose 
specimen had the bulb artificially disconnected and rotated through 45 
degrees. The author finds the same olfactory fossa which we have 
called attention to in Reptilia and opossum and verifies its relation to 
Jacobson’s organ. In Platypus it is much deeper than even in the 
black snake but is on the dorso-lateral rather than mesal aspect. A 
study of the relations of the part of the olfactory bulb associated with 
Jacobson’s organ does not reveal any arrangement different from the 
rest of the bulb. It would appear that in all its connections with the 
brain the organ of Jacobson exactly resembles the olfactory apparatus 
proper and like the latter has its centres in the pyriform lobe and prob- 
ably also in the hippocampus. Gs Lane 


Fibre Connections of the Olfactory Lobe of Man.’ 


It will be recalled that recent investigators have shown by a variety 
of methods that the connections of the olfactory nerve in the olfactory 
lobe are practically the same in all vertebrates, the peripheral nerve 
forming a terminal arborization in the glomerule, there to enter into 
relations with the protoplasmic process of one or more of the mitral 
cells of the olfactory lobe whose axis cylinders effect the cortical con- 
nections. In batrachians, reptiles and birds each mitral cell gives off 
more than one protoplasmic process and thus is related to more than 
one glomerule. No mammal hitherto studied has shown this arrange- 
ment, there being but one protoplasmic process to each mitral cell. In 


1 Anat. Anz., XI, 6. 


2GEHUCHTEN, A. VAN. Le bulbe olfactif de Vhomme, Azbliog. Anatomique, 
III, 4, Aug., 1895. 


xX JouRNAL OF CoMPARATIVE NEUROLOGY. 


all lower vertebrates and in some mammals—cat, rat, mouse, rabbit— 
each glomerule receives the protoplasmic process from a single mitral 
cell, while in the dog five or six mitral cells effect connections with the 
same glomerule, thus putting each olfactory fibre into relation with a 
larger number of cortical cells. Examination of the olfactory lobe of 
the still-born child with the silver method shows the usual mammalian 
type with the following exceptions: (1) the mitral cells are more ir- 
regularly arranged than in other mammals, lying often in the glomeru- 
lar layer or even among the peripheral nerve fibres; (2) the protoplas- 
mic processes of the mitral cells occasionally branch, communicating 
with more than one glomerule as in the lower types; (3) most of the 
glomerules receive the protoplasmic process of but a single mitral cell, 
though sometimes two to four mitral cells are related to the same glom- 
erule as in the dog. CAyeHe 


The Relation of Sensory and Motor Areas of the Cortex. 


The evidence has accumulated in abundance to show that the sep- 
aration of motor and sensory areas is arbitrary and untrue to the actual 
facts. ‘The ease with which motor disturbance can be demonstrated 
may very well account for the pre-eminence given to the location of 
motor functions. Munk has shown in a series of brilliant experiments 
that even in the visual area the various portions may produce complex 
motor responses in the form of codrdinated eye movements. The same 
author has shown that the various cortical motor areas are not exclu- 
sively so but that approximately the same regions have a sensory func- 
tion also. But for our present purpose it is of importance to show that 
the same generalization holds good for the human brain. From the 
vast material at our disposal it is only necessary to call attention to the 
selected cases described by by Dr. C. L. Dana.! The analysis of these 
cases shows, as the author states, that it is the power of localization which 
is first to be disturbed in case of injury to the cortex, next tactile anzes- 
thesia then analgesia, then simple muscular anesthesia and, finally, loss 
of temperature sense. Now the higher forms of codrdinated sensations 
owe their existence as much to vestiges of earlier sense presentations as 
to the actual sense content. ‘This is especially true of localization 
and muscular sense. It appears then that one of the most constant of 
the results of cerebral injury is the impairment of the vestiges or the 
interruption of the paths connecting with the store-house of such im- 
pressions. It is a well-known fact that the extent to which vestigial 


VJourn. Nerv. and Ment. Dis., Dec., 1894. 


Literary Notices. Xxi 


impressions are made or memories are accumulated depends on the in- 
tensity of the impression or stimulus as well as on the lack of compet- 
itive impressions. ‘This may mean that the amount of irradiation of 
the stimulus is the factor. At any rate it is easily intelligible that the 
vestiges will be first disturbed and that they may in time be renewed, 
explaining the frequent return of the power of localization and codérdi- 
nation where the tactile sense is not wholly destroyed. 

The anatomical structure of the cortex is, so far as can now be gath- 
ered, conformable to the necessity imposed by the construction of the 
pathological data. Not only is each cell brought into rapport with 
many others by the neurodendrites but these connections are in both 
rank and file. For every cell that gives rise toa kinesodic fibre or 
neuraxon there are many which form its sphere of influence. It is in 
accord with the dynamic theory of nervous action to suppose that in 
the reaction between the numerous esthesodic cells su/er se and the 
other reaction between these cells and the motor initiatory cells is the 
immediate occasion of consciousness. Geir 


Is the Decorticated Dog Conscious? 


It is unfortunate that so few of the students of brain anatomy and 
physiology are also acquainted with the first principles of psychology. 
The result of this one-sided furnishing is often a deplorable inability to 
construe the results of experiment and pathology. If this were all, the 
result would have less significance but upon the false conclusions thus 
reached a superstructure is often raised, vitiated throughout by the 
same fundamental fallacy. In no instance has this lack of psychologic- 
al insight been more evident than in the various attempts to explain the 
results of total or maximal extirpation of the hemispheres. Every one 
will, of course, think at once of the celebrated instance of nearly com- 
plete removal of the hemispheres by Goltz and the subsequent reports 
by that author and Dr. Edinger. After eighteen and a half months it 
would seem that the immediate results of the operation could safely be 
considered as eliminated and the results, excluding degeneration phe- 
nomena, might be taken as those normal to the brain minus the hem- 
ispheres. Even so, the purely physiological questions are by no means 
as simple as might be supposed from current discussions. For one 
thing, the great inhibitory influence of the cerebral hemispheres being 
removed, nothing is more certain than that the infracortical centres 
would not operate in the same way that they normally do in the unin- 
jured brain. Again, the effect of the concentration of stimuli intended 
for the great terminal projection system upon one of a lower order 


Xxii JOURNAL OF COMPARATIVE NEUROLOGY. 


could hardly fail to greatly alter the reaction. In spite of these sources 
of error it is not our desire to intimate that, on the whole, the activities 
of the decorticated dog are not fairly indicative of the functions of the 
infracortical centres. Strong irritation of the skin caused the animal 
to growl and bite, he could be awaked from natural sleep by loud 
noises, the taste of food awakened the impulse to feed, etc. But when 
an author seriously claims that these facts prove that ‘‘we have un- 
doubted manifestations of the presence of every variety of sensation, 
tactile, muscular sense, sense of pain, vision, hearing, taste, and finally 
the visceral sensations of hunger and of thirst,’’! he betrays a lack of 
psychological discrimination and begs the real question. Here, as so 
generally, the content of sense is put for sensation and then the conclu- 
sion is reached that sensation is produced in the infracortical centres. 
Now it would be just as legitimate to decide that because certain pro- 
cesses of a nervous character and essential to vision are carried on in 
the eye therefore the eye is the seat of visual sensation. ‘There is no 
manner of doubt that all the preliminaries to vision including a large 
number of coérdinating reflexes are all provided for in the infracortical 
centres. It is equally certain that there is a provision for reflexes of a 
higher order—such as grow out of the relations of different senses 
inter se. All that is reported in the case of the decorticated dog may 
well belong in the categories of infraconscious codrdination. It must 
be remembered that the consciousness of a sensation is probably never 
attained until there has been a kinesodic response to it and it is not un- 
likely that it is the reflected current rather than the direct one which 
enters consciousness. It is in this way that the storage of vestiges in 
the cortex may be explained in such cases where the original stimulus 
never reached consciousness. It is absolutely necessary that the line 
between physiology and psychology should coincide with that which 
separates the conscious from the unconscious or that the distinction be 
abandoned. If sensation is selected as the unit of psychology it is ab- 
surd to speak of unconscious sensation. ‘The fact that the dog in the 
present instance acts as though conscious of a stimulus is no proof of 
such consciousness and all admit that there is a complete absence of all 
evidence of reproduced sensation or of reflection. It is true that very 
complicated sets of cyclical reflexes are produced but something very 
similar might under proper conditions be reproduced upon a corpse. 
The fact that the cortical connections are not completed until a very 
late period of the ontogeny, which is adduced in support of the idea 


1W.H. THompson. Journ. Nervous and Mental Disease, June, 1895. 


a a 


Literary Notices. XXiil 


that the dog is really conscious, in reality looks the other way. The 
presence of consciousness for some time after birth would be a great 
embarrassment to the economy of the animal. The relation between 
consciousness and educability is not denied, but it does not follow that 
the reflexes are not capable of education while the sphere of profitable 
interference of the conscious is relatively very small. 

Gee isis 


The Paraphysis.' 


This paper is mainly concerned with a summary of other and ear- 
lier papers by the same author, with, however, some new observations 
and an excellent series of photographs. He reiterates the belief that the 
paraphysis is an evagination from the cerebrum and is only secondarily 
associated with the diencephalon and that it represents, like the parie- 
tal eye, an aborted sense organ. ‘This seems to us an improbable view 
and one requiring stronger evidence than that presented by the author’s 
photographs. Yet the remarkable constancy with which this structure 
appears in the embryos of all vertebrates certainly does indicate that 
the paraphysis has now or has had an important part to play in the evo- 
lution of the vertebrate brain. Just what this part may be we cannot 
by any means regard as satisfactorily determined. 

In an earlier work the author described the parietal eye of Anguis 
as originating by a constriction of the distal end of the pineal evagina- 
tion essentially as described by Spencer and the majority of other in- 
vestigators. Béraneck, however, finds that the parietal and pineal 
evaginations have distinct origins from the roof of the diencephalon, 
and now Francotte comes over to the same ground, the error in the 
first case having arisen not from inaccuracy of observation, but from an 
anomalous condition in the embryo under under investigation. Photo- 
graphs of the normal and the abnormal brains are given. Five em- 
bryos taken from the same mother were found all to present the same 
anomaly. Whichever view of the origin of the parietal eye may 
prove to be correct, this paper will do good service in calling attention 
to a source of error and of disagreement in many another controversy 
besides this one. It is a significant fact that it was during the same year 
in which Francotte’s paper appeared that Prenant published his paper 
on accessory parietal eyes in Anguis fragilis, in which he made a sta- 


1FRANCOTTE, P. Note sur I’ceil pariétal ’épiphyse, la paraphyse et les plexus 
choroides du troiséme ventricule. Aull. 7 Acad. Royale de Belgique, 3 Series, 
XXVII, 1894. 


XXiVv JOURNAL OF COMPARATIVE NEUROLOGY. 


tistical study of the variation of these organs and got some surprising 
results. Cf. Vol. V. of this Journal, p. xv. 

Cig ae 
A New Journal. 


The first number of the ‘‘ Rrvista di Patologia nervose e mentale”’ 
appears with the new year issuing from the Clinic of Psychiatry of Flor- 
ence. ‘The directors are Dr. E. Tanzi, associated with A. Tamburini 
(Reggio Emilia) and E. Morselli (Genoa); the editors are E. Belmondo 
and E. Lugaro, together with an able board of colloborators. We are 
promised in the prospectus monthly issues making in the aggregate 
about 500 pages per year. The field to be cultivated is that of an 
“Italian Centra/blatt,” and, judging from the two numbers which have 
thus far reached us, the labors of the editors are being wisely and fruit- 
fully expended. The review department is well sustained. We append 
notices of a few of the brief original articles. 


Structure of the Cytoplasm of the Nerve Cell.! 


Nissl’s method of differential staining after alcohol hardening, 
which has given such an impulse to recent cytological work, is made 
the basis of an investigation of the relative functional value of the 
chromatic and the achromatic elements of the general protoplasm of 
the nerve cells. Dr. Lugaro comes to the conclusion that the achro- 
matic substance is the physiologically active medium, while the chro- 
matic elements are simply passive. The achromatic part is composed 
of a filar mass in the sense of Flemming, while the chromatic part 
which alone is stained by the method of Nissl represents an interfilar 
mass, and therefore the method of Nissl is absolutely negative regard- 
ing the real structure of the nerve cell. 

This conclusion is based on the mode of development of the chro- 
matic masses, their distribution in the adult cell and their relative abun- 
dance in the various processes. The fact that the chromatic substance is 
absent in the smallest ramifications of the nerve fibres, and is present in 
greater quantity and in larger masses in the larger processes of the cell 
and particularly in the cell body itself is regarded as evidence that it 
is composed either of nutritive material or of products of dissimila- 
tion. The smaller dendritic termini having a greater surface in pro- 
portion to their mass do not require any special nutritive mechanism. 


' LuGARO, E. Sul valore rispettivo della parte cromatica e della acromatica 
nel citoplasma delle cellule neryose. zvista az Patol. nerv. e ment., I, 1, Jan., 
1896. 


Literary Notices. XXV 


Dr. Lugaro has found in a study which we have noticed elsewhere in 
this number that in different functional conditions the variations of the 
quantity of the chromatic substance are very slight, while there isa much 
more constant relation between the density of the stain and the size of the 
cell. But there is another factor influencing the amount of chromatic 
substance in the cell, a ‘‘specific factor,” depending on the connections 
of the cell or the normal intensity of its discharge. ‘These points are 
illustrated by several figures of the nerve cells of both vertebrates and 
invertebrates. . Cai: 


The Pons Varolii of Man.! 


Studies on human fetuses of five to seven months and on the 
brains of young children by several of the newer methods. 

I. Substantia grisea pontis. These cells occupy the space between 
the superficial transverse fibres of the pons and the internal lemniscus, 
collecting in the interstices of the fbrae transversae pontis and the fas- 
cicles of the pyramidal tract. They are of Golgi’s first type and send 
their nervous prolongations into the middle peduncle of both the same 
and the opposite side, also into the ventral portion of the raphe. These 
fibres are regarded as the scandent fibres of the cerebellar cortex, 
whose terminal arborizations lie in connection with the Purkinje cells. 

Il. Collaterals of the pyramidal fibres. These are two kinds, di- 
rect and indirect, terminating in physiological contact with the den- 
drites of the elements of the substantia grisea pontis. 

Ill. Pedunculus medtus cerebelli. ‘The middle peduncles are com- 
posed not only of tie ascending fibres from the substantia grisea, but 
also of descending fibres from the cells of Purkinje. Part of the latter 
cross the raphe to terminate in ramifications among the elements of the 
substantia grisea of the opposite side, part effect similar connections on 
the same side, others pass via the raphe to the level of the segmentum 
pontis of the same or the opposite side. They then pass either as fibrae 
arcuatae internae or as ascending fibres of the raphe to participate in 
the formation of the fasciculus medianus. Some fibres terminate among 
the elements of the substantia reticularis grisea tegmenti pontis. To this 
tract the author gives the ponderous name, ‘‘direct tract from the cere- 
bellum to the nuclei of the tegmentum pontis.” 

IV. Internal bundle of the pes peduncult. ‘The author has estab- 
lished a direct and crossed connection between the Rolandic and fron- 


1 PusATERI, E. Sulla fine anatomia del ponte di Varolio nell’uomo. Nota 
prelim. Rzvzsta di Patol. nerv. e ment., 1, 1, Jan., 1896. 


XXVi JOURNAL OF COMPARATIVE NEUROLOGY. 


tal opercula of the cerebrum and the cerebellar cortex via the swbstan- 
tia grisea pontis. The latter has probably other important connec- 
tions as yet unknown. 

V.  Substantia reticularis grisea tegmentt pontis. ‘The cells of this 
region have large protoplasmic processes which in part join the internal 
arcuate fibres and in part cross in the raphe to the substantia reticularis 
of the opposite side, thus forming a protoplasmic commissure of this 
region. ‘The axis cylinders of these cells in part cross the raphe, in 
part join the internal arcuate fibres; others, arising nearest the raphe, 
pass dorsally to join the fasciculus medianus ; still others go to the /eg- 
mentum peduncult. : 

VI. Collaterals of the fasciculus longitudinalis postertor. These 
pass to the nucleus funicult teretis, the substantia reticularis tegmenti pon- 
tis and the nucleus reticularts tegmentt pontis of Bechterew. 

The reader will be struck with the substantial identity of many of 
these results with those of Cajal upon the brains of the dog, cat and 
rodent, as described in Bibliographie Anatomique, Dec., 1894. 

ee pas iets 


The Influence of the Cerebrum upon the Excretion of Nitrogen.! 


The experiments were performed on pigeons. ‘The cerebrum was 
removed and after complete recovery the animal was kept fasting for 
from four to six days. ‘The loss in weight from day to day and the 
amount of nitrogen contained in the urine were compared with the 
same data from unoperated specimens under the same conditions. The 
decerebrated pigeons lost weight much less rapidly than the normal, 
and the amount of nitrogen eliminated was still more conspicuously less 
in the case of the decerebrated pigeons. On the other hand, a pigeon 
which was observed on the days immediately following the operation 
showed a geater loss in weight and in amount of nitrogen excreted than 
the normal pigeons. 

The author concludes that the cerebral hemispheres, at least in 
birds, have a marked influence in regulating the metabolic processes of 
the body, acting as trophic centres to stimulate the anabolic processes. 

C. Jae 


1 BELMONDO, E. Ricerche sperimentali intorno all’influenza del cervello 
sul ricambio azotato. Révista di Patol. nerv. e ment., 1, 2, Feb., 1896. 


Literary Notices. XXVii 


Nerve Termini of the Prostate.! 


Dr. Timofeew in continuing his studies on the nerve endings of 
the sexual organs finds a peculiar structure in the outer connective tissue 
capsule of the prostate, as well as between the muscles and glands of 
that organ, and in the adjacent mucosa of the dog and cat. 


In the cut we give a diagrammatic reproduction of one of his 
figures, taken from termini in the capsule of the prostate of a dog. 
The structure consists of two fibres, both medullated and issuing from 
the same nerve trunk, one of which is much the larger and enters a 
capsule in which it terminates in a clavate or ribbon-like expansion. 
The other fibre also enters the capsule and terminates in a mass of fine 
varicose fibres which perhaps anastomose with each other, but not with 
the first fibre. Both fibres lose the myelinic sheath before entering the 
capsule. The cut will explain the relations without further description. 

Chyna 


Demonolotry of the Nineteenth Century.’ 


The volume before us is a good specimen of the modern pub- 
lishers’ art and in other respects challenges criticism. It is, moreover, 
a book destined to create a considerable amount of interest in various 
circles. Unlike most recent attempts to discuss this most perplexing 
subject it at least claims a degree of preparation on the part of writer 


1 TIMOFEEW, D. Ueber eine besondere Art von eingekapselten Nervenen- 
digungen in den mannlichen Geslechtsorganen bei Saugetieren. Axat. Anz., 
XI, 2, 22 Aug., 1895. 


2 J. L. Nevius, D.D. Demon Possession and Allied Themes. 7. A. Revel 
Co., Chicago. $1.50. 


XXVili JouRNAL oF CoMPARATIVE NEUROLOGY. 


and editor which hardly permit of an @ priort waving of its claims. It 
demands a serious hearing, if for no other reason, because it claims to 
fairly represent the calm judgment of all but an insignificant minority 
of the educated occidental missionaries at present actually living among 
the Chinese and other oriental peoples. If this is indeed the case it 
may give rise to serious reflection or even to the query whether the re- 
action of barbarism on the missionary is not as great as his influence on 
the barbarism in the opposite direction. 

We admit to a feeling of grave responsibility in dealing with such 
a work and, while we feel that scientific truth leaves us no alternative, 
it is hoped that the reviewer may be credited with no antagonism to the 
cause in the interest of which the volume was sincerely written. Even 
more, it is because the reviewer believes that the false views here pro- 
mulgated will do great injury to that very cause that he does not feel 
justified in holding his hand. 


At the outset it is freely granted that the entire honesty and credi- » 


bility of the author and his witness is assumed in all that follows. The 
author has displayed not only praiseworthy industry but considerable 
skill in the gathering of facts and discussing their significance, and 
when we are forced to add that he seemed singularly lacking in critical 
and scientific discrimination it does not follow that the value of the facts 
is invalidated. Indeed the naiveté with which facts inconsistant with 
the conclusions are set down is sufficient check in most cases. The 
author is beyond the sting of criticism and we pass to a consideration 
of the work in detail. 

In the first place we may quote the author’s synopsis of the facts 
and conclusions reached as he closes the descriptive section of his 
book. (P. 143.) It will form a convenient point of departure for our 
review of the evidence. 

‘¢7,. Certain abnormal physical and mental phenomena such as 
have been witnessed in all ages and among all nations and attributed to 
possession by demons, are of frequent occurrence in China and other 
nations and have been generally referred to the same cause. 

‘¢2, The supposed demoniac at the time of possession passes 
into an abnormal state, the character of which varies indefinitely, be- 
ing marked by depression and melancholy, or vacancy and stupidity 
amounting sometimes almost to idiocy, or it may be that he becomes 
extatic, or ferocious and malignant. 

‘¢3, During transition from the normal to the abnormal state, 
the subject is often thrown into paroxysms more or less violent, during 


Literary Notices. XX1X 


which he sometimes falls on the ground senseless, or foams at the 
mouth, presenting symptoms similar to those of epilepsy or hysteria. 

‘‘4, The intervals between these attacks vary indefinitely from 
hours to months, and during these intervals the physical and mental 
condition of the subject may be in every respect normal. ‘The dura- 
tion of the abnormal states varies from a few minutes to several days. 
The attacks are sometimes mild and sometimes violent. If frequent 
and violent the physical health suffers. 

‘‘5. During the transition period the subject often retains more 
or less of his normal consciousness. ‘The violence of the paroxysms is 
increased if the subject struggles against, and endeavors to repress the 
abnormal symptoms. When he yields to them the violence of the par- 
oxysms abates or ceases altogether. 

‘6. When normal consciousness is restored after one of these 
attacks the subject is entirely ignorant of everything which has passed 
during that state. 

“7. The most striking characteristic of these cases is that the 
subject evidences another personality and the normal personality for 
the time being is partially or wholly dormant. 

‘¢3. ‘The new personality presents traits of character utterly dif- 
ferent from those which really belong to the subject in his normal state 
and this change of character is with rare exceptions in the direction of 
moral obliquity and impurity. 

‘¢g. Many persons while ‘demon-possessed’ give evidence of 
knowledge which cannot be accounted for in ordinary ways. They 
often appear to know of the Lord Jesus Christ as a divine Person and 
show an aversion to, and fear of Him. ‘They sometimes converse in 
foreign languages of which in their normal states they are entirely 
ignorant. 

“‘to. There are often heard, in connection with ‘demon possess- 
ion’ rappings and noises in places where no physical cause for them 
can be found; and tables, chairs, crockery and the like are moved 
about without, so far as can be observed or discovered, any application 
of physical force, exactly as we are told is the case among spiritualists. 

‘11. Many cases of ‘demon possession’ have been cured by 
prayer to Christ, or in his name, some very readily, some with 
great difficulty. So far as we have been able to discover, this meth- 
od of cure has not failed in any case, however stubborn and long 
continued, in which it has been tried. And in no instance, so far as 
appears, has the malady returned if the subject has become a Christian, 
and continued to lead a Christian life.” 


SOK JOURNAL OF COMPARATIVE NEUROLOGY. 


A few words in passing upon this summary. ‘The admission of 1 
is significant and is fully borne out by the facts adduced. These so- 
called possessions are not in any material way different from phenom- 
ena with which modern pathology is dealing every day at home with 
no doubt of their pathological character. That they have been gener- 
ally referred to devils is as forcible an argument as it would be to ad- 
duce the universal belief that scrofula was due to the evil eye in a 
modern medical consultation. We might drop the matter right here 
and would be content to do so if convinced that all our readers were 
familiar with the state of scientific opinion. We have, then, not a pe- 
culiar set of phenomena but a familiar set in peculiar setting which 
alone warrants its selection for special study. Our author does not hes- 
itate to suggest (p. 182) that ‘‘ the unscientific Chinese were, so far as 
this subject is concerned, more careful observers of facts, and more 
correct in their conclusions than many who have been leaders of public 
opinion in our times ;” the direct objective being here modern patholo- 
gists. This is a remarkable statement which becomes more remarka- 
ble with every illustration of the scientific attainments of the oriental 
observer and healer. 

The symptomatic complex indicated in the following sections con- 
tains nothing novel in any way, but the strange circumstances and the 
implicit faith of the barbarians in the supernatural character of the 
manifestations very naturally produce a strong impression on the visitor. 
The author was chiefly struck by the apparent transformation of the 
personality during such attacks, yet he is not entirely ignorant of the 
fact that a change of personality is characteristic of well-known nervy- 
ous maladies. It is gravely stated that the possessed shows super- 
natural powers of speech and gaining information. ‘There is, however, 
no case given where such powers are proven. It is true that under the 
pressure of the disease the patient may seem to speak with tongues, but 
when the evidence can be thoroughly sifted such cases usually find 
their true place among familiar facts. A case in point now occurs to 
the writer. The wife of a prominent theological professor, herself a 
woman of great strength and refinement of character but not well- 
versed in the German language, awoke one morning and to her own 
great distress and the astonishment of her family was utterly unable to 
speak a word of English but made her wants known in German. Her 
husband was even less familiar than she had hitherto been with that 
language so that she was for the most part unintelligible. The good 
Doctor, in narrating the circumstances, stated that she spoke fluently, 
but admitted that neither he nor she were in a condition to gauge the 


Literary Notices. XXXi 


correctness of the language. This cultured lady had never had a mas- 
tery of the spoken tongue but suddenly, without apparent cause, lost 
the speech of her birth and entire life and seemed to have miraculously 
acquired a new faculty. . My friend, as a sensible man, did not seek to 
exorcise a demon or even send for a doctor, but employed the expect- 
ant method until the curious nervous kink straightened out of itself, 
never, so far as known, to reappear. If this circumstance, which is 
known to very few, had taken place in an ignorant community, the be- 
lief in possession would have been absolute and the sufferer, if not of 
unusually well balanced mind, would have been tormented into perma- 
nent delusions or worse. Again, in the case of the writer it has hap- 
pened while studying language and in a stage where ordinarily the act 
of composition was painfully tedious, that, say during an early morn- 
ing doze, he seemed suddenly endowed with a gift of tongues and 
rolled off sentence after sentence with the greatest ease and apparent 
accuracy. The neurological explanation of such phenomena is not 
difficult and, if it were, the appeal to demoniac possession would add 
but another and more insoluble problem. In one case cited by 
the author the subject seemed to be able to describe a distant country 
and her power was accounted miraculous, but the more than usually 
accurate reporter adds that her descriptions were only correct in the 
general outlines and not in detail and that she might have picked up 
the facts used from other sources. 

If the author had lived in this country during the last twenty years 
it is unlikely that he would have considered the resemblance of the 
oriental table-tipping performances to those which ran their course here 
as evidence in favor of their spiritual nature. 

One point requires further notice. The morbid personality is 
usually qworse than the normal. Does thisseem remarkable? We think 
it is only what should be expected. In the life of every person not ut- 
terly depraved there is a constant struggle against what are to him temp- 
tations. The even trajectory of normal life is the resultant of conflict- 
ing forces of which some are (to the subject) bad or malign. This is 
quite apart from any absolute moral system. In neurotics these anti- 
monies are doubly strong. When, then, the one set of mental forces 
are switched out of circuit the other is likely to be immanent. The 
ruling or dominant forces are just the ones to be unhorsed for the rea- 
son that they are so largely inhibitory while the other set is chiefly in- 
trenched in the elementary physical nature. 

Before passing to the evidence collected by the author let us note 
something of the setting of the facts. 


XXxXii JOURNAL OF COMPARATIVE NEUROLOGY. 


‘Tt will be observed that nearly all the incidents related are given 
on the testimony not of missionaries, but of native Christians—mostly 
native pastors.” (Introduction, p. v.) The native Christians continue 
in this belief. (P. vi, and passim.) 

‘¢ Antecedently to any knowledge of the New Testament the peo- 
ple of North China believed fully in the possession of the minds and 
bodies of men by evil spirits.” (P.iv.) The natives at once recog- 
nized the identity of the biblical possession with the phenomena about 
them. 

Our witnesses are, then, filled with a prepossession and it would 
never enter their minds to seek any other than the demoniac explana- 
tion, this ‘‘ belief being a part of that animism, or spirit worship, 
which has existed in China—as in many other countries—from the very 
beginning of history or tradition.” (P.iv.) They do not regard the 
phenomena of possession and exorcism ‘‘as anything strange or re- 
markable.” (P. 35.) 

Such are the witnesses that are regarded by the author as ‘‘ more 
careful observers and more correct in their deductions” than modern 
pathologists. We cannot expect the writer under the limitations neces- 
sary to his work to exhibit a familiarity with the recent discoveries in 
hypnotism but we must at least concern ourselves with the evident bias 
of writer and observers. 

Again, a belief in exorcism is shown to be universal. The detailed 
account of the exorcists (p. 68-71) is very full and instructive. The be- 
lief that these professional mediums, whether Confucionist or Taoist, 
have a real power to drive out demons is as implicit as that in the spir- 
itual nature of the manifestations. If the universal belief of these 
heathen is to be taken as irrefrangible evidence we must also believe in 
the supernatural power of as pitiful a set of mountebanks as ever dis- 
graced the name of man. We must believe in charms, in amulets and 
witchcraft. In other words, the learned author and by implication the 
whole body of occidental missionaries wish us to abrogate whatever 
of moral progress has been made by civilization and return to the be- 
liefs and practices of medieval superstition. Probably the author did 
not follow his argument to its legitimate conclusion but there is no logi- 
cal stopping place short of that indicated. The exorcists are supposed 
to be the special aversion of the spirits and ‘‘ never venture anywhere 
without having charms, talismans, and all kinds of abracadabras with 
them.” Armed with a black mule’s hoof and a black dog’s blood 
these practioners do, nevertheless, succeed in going about to very good 
purpose. ‘‘She spends her time going about among the villages in the 


Literary Notices. XxXxiii 


neighborhood telling fortunes, and healing diseases, and in this way 
makes a good deal of money” (p. 37). The phenomena attending 
séances are exactly such as those with which we are too well acqainted 
here. ‘‘ Tables are turned, chairs are rattled, and a general noise of 
smashing is heard.” Even Slade and his slate have their counterparts, 
‘*the pencil moving of its own accord” (p. 69). The effect of the 
mysterious is indubitable. We all recall that no less a person than Pro- 
fessor Zéllner was mystified by Slade and induced to commit himself 
to absurdities which appear in high relief since the fraud was con- 
fessed. That the oriental exorcism is associated with vulgar conjury 
and legerdermain is evident throughout these pages and the skill of 
oriental jugglers is almost past belief. That, on the other hand, there 
is a certain amount of honest belief in the powers of spirit on the part 
of the profession is also evident. A large class of semi-professional 
healers occupies the same place as the ‘‘ Christian healers” and ‘‘ faith 
healers”? of our own locality. Almost while these lines are being 
penned anew Man-Christ is touching and prescribing for all manner 
of diseases and as many as are touched are made whole. Even hand- 
kerchiefs acquire curative power by being blessed by this ignorant pea- 
sant. A thriving trade is being driven in counterfeit blessings and ad- 
vanced positions in the line at his door have sold for seven dollars. 
But it is not necessary to seek isolated cases like that of Francis Schlatter 
when any patent medicine circular will reveal a long roll of clerical at- 
testations to absurdities too patent to deceive a healthy child. The 
love of, and belief in the marvelous is too thoroughly ingrained to yield 
easily to judgment or authority. 

Prominent among the facts forming the background of these ap- 
pearances is the evidence of neurotic predispositions in the subjects 
described. 

One curious fact familiar to all physicians seems to mystify the 
author, i. e., that neurotic individuals are generally the reverse of 
feeble in appearance. They generally look younger than others of the 
same age and seem quite well to the untrained observer. Indeed it is 
one of the special inconveniences of nervous disease that it is hard for 
his friends to refer the irritable and distracted moods of the sufferer 
to a diseased condition. He gets credit for pure malignity where he 
perhaps deserves only profound sympathy, he himself frequently taking 
the same view. 

So much for the background, but we must notice that the author 
is at great pains to insist that there is no conceivable bias in favor of 
the demon theory and the resulting practice of exorcism on the side 


XXXiV JOURNAL OF COMPARATIVE NEUROLOGY. 


of the missionary. In fact, we are told that they are at first skeptical. 
How has it escaped the writer that the missionary is under the strong- 
est of all inducements to accept the current interpretation? It is, how- 
ever, a common experience that one shold be unconscious of the most 
powerful motives influencing his belief and we need not impugn the 
honesty of such an one in insisting that this motive is patent to every 
outside observer. ‘The exorcism of the natives is an integral part of 
their religion and is adopted by all sects alike. When Christianity en- 
ters the field it must at once compete with these sects in this most prac- 
tical matter. Since the function of religion is to control the spirit of 
evil, that must be the best religion which most successfully combats 
these manifestations of satanic power. ‘The author expresses surprise 
that the natives eagerly seize upon the accounts of New Testament pos- 
session and need no suggestion from the missionary to turn them to prac- 
tical account. The result is what one would expect. Grant, as we must, 
that the native doctors are often successful with their burning pills on 
the nerve plexus at the roots of the nails, and that the native conjurers 
are also frequently able to banish the disease by their incantations, it 
-would be expected that the strange message from a land beyond the 
realm of dreams should create the conditions for deliverance, nor is it 
astonishing that the complete change in mental attitude and the 
saner methods of life involved in embracing Christianity should cause, 
in many cases, a permanent cure. Even a much less significant change 
is often sufficient in practice. ‘‘ The number of those who for this 
cause have become Christians is very great.” (P. 51.) The mission- 
aries must be much less or more than human if they are uninfluenced 
at least unconsciously by this unexpected and powerful means to reach 
the otherwise inaccessible heathen. Who shall determine whether this 
is not a providential opening? Be this as it may, it would seem un- 
necessary to degrade occidental humanity to the same depths of super- 
stition and we cannot applaud the attempt. 

Fifteen cases are presented from China and we must give up our 
plan of analysing them and do so the more willingly that they present 
nothing novel or especially pertinent not already noticed. In the spirit- 
ualistic career the fortune of the experimenters often mysteriously dis- 
appears. ‘The afflicted is incidentally shown to have been a gambler 
and profligate, etc. The author explicitly denies that there is any ten- 
dency to epidemics but there is evidence, even in the few cases given, 
of contagion, i. e. that the influence of others is felt not only in pro- 
ducing but in determining the form of the attack (p. 37). The rhyth- 
mical form of expression in such cases is as old as history and the author 


—— ee ee eee eee 


Literary Notices. XXXV 


himself notices the resemblance of the utterances of these modern 
pythonesses to ‘‘ the meaningless chants heard in Buddhist temples.” 
He may not have known of the frequency with which such facility of 
versification appears in the exalted states of mania. It will even be 
recalled that the habit discoverable in Shakespeare’s characters of drop- 
ping into mongrel rhymes at the critical moments has been seriously 
defended on the ground that it is acommon tendency of overwrought 
minds to spend and distract themselves in poetical expression. 

With the second or theoretical part of the book it is difficult to 
speak patiently, perhaps unnecessary to speak at all. We are told that 
the distinguishing marks of demoniac possession are alterations in per- 
sonality. We can form only a vague guess as to what is meant by per- 
sonality, but, whatever it is, it remains certain that the vast majority of 
the insane would by this law be regarded as possessed. By a simple 
application of this conclusion it would appear that dements with morbid 
changes in the cortex (as easily detected by the microscopist as small 
pox pustules in the skin) should be treated as possessed and cured by 
exorcism. 

Cases are now well known in which an insane person has two dif- 
ferent states or ‘‘ personalities” in one of which he has one set of men- 
tal and moral attributes, and in the other, along with certain obscure 
but measureable changes in circulation and other physiological func- 
tions, displays an entirely different set of mental and moral peculiarities. 
In one case he is intelligent and speaks fluently—is crafty and excita- 
ble, while in the other he is stupid and speaks a different language. 
Nor are we wholly at a loss as to the physical cause of this double man- 
ifestation. We find, perhaps, that he is left-handed in the second state 
but right-handed in the first. We learn that he was left-handed in his boy- 
hood in Wales but learned English and the use of the right hand later, 
recelving an education in English. With abundant evidence now at 
hand that only one side of the brain is concerned with speech, and that 
the side corresponding to the educated hand, we have a right to sug- 
gest that both halves of this man’s brain have been modified, the one 
in the Welsh or left-handed, the other in the English, right-handed 
way. We cannot follow this line further, but it suggests a reasonable 
ground for the most difficult phenomena of hypnotism. ‘The author’s 
distinction between the pathological and psychical theory is arbitrary. 
We think a proper appreciation of the dignity and sphere of the soul 
would make one glad to see in the driveling lunacy of the class of phe- 
nomena here discussed matters for the alienist rather than the theolo- 
gian. It was a very true word in the author’s preface: . ‘‘ Some of the 


XXXVi JoURNAL OF COMPARATIVE NEUROLOGY. 


readers of these pages will in all probability be disappointed in finding 
the characters and doings of spirits much less interesting and credita- 
ble than they are represented in the familiar writings of Milton and 
Dante.” This in itself would be sufficient refutation of the claim to 
most candid readers, but no explanation is vouchsafed. 

We ought perhaps to accept the Chinese idea that the demons are 
simply the souls of the departed who have not been so fortunate as to 
appear in the imperial edict deifying them off-hand. 

Of the question as to the relation of the described phenomena 
with those of New Testament times we say nothing, though it forms a 
large element in the book. But we have done. ‘That the views pre- 
sented should be accepted by scientific men is impossible; that they 
should gain acceptance in the circle of religious enterprise and educa- 
tion would be very unfortunate as it would tend to perpetuate and 
‘widen a very unnecessary breach where the fullest harmony and sym- 
pathy is important. Gs ire 


The Growth of the Brain.! 


The present volume is a very worthy addition to the Contempo- 
rary Science Series, for in its 19 chapters containing 368 pages, there 
is a clear, concise, and very readable statement of the most interesting 
facts regarding the growth of the brain. 

The author gives in his first chapter an introduction to the study 
of growth and sums up the laws which seem to govern it. The grad- 
ual increase of the weight of the human body is represented by means 
of diagrams, the comparison between the male and female being well 
brought out. The next chapter gives with greater detail the relative 
increase of different parts of the body, showing the proportion between 
weight-increase and increase of stature. In the fourth chapter the 
weight of the brain and spinal cord are treated. The point is clearly 
made that in taking brain weights other things than nerve cells are 
often weighed, as for instance the membranes, pia and dura, the blood 
vessels and cavities with their fluid contents; therefore there is bound 
to be a certain discrepancy in the weights made by different observers 
if the same methods of weighing are not followed. This the author 
claims has not been done, and much of ‘the material now at hand is 
consequently of little value in making deductions. In trying to arrive 


1The Growth of the Brain, by HENry HERBERT DONALDSON, Professor of 
Neurology in the University of Chicago. Imported by Charles Scribner's Sons, 
New York, $1.25. 


Literary Notices. XXXVI 


at general conclusions Professor Donaldson believes in grouping the 
statistics, for ‘‘ age, stature, sex, bodily weight and race” are all mod- 
ifying circumstances. He concludes that there is a marked constancy 
in the percentage values. of the subdivisions of the encephalon of all 
ages, all statures, and both sexes. 

Thus after maturity the male encephalon and all its parts are lar- 
ger than the female. With increasing age there is a decrease of the 
weight of the encephalon and all its parts, and for the same age and 
sex decrease in bodily weight is accompanied by by a decrease in 
weight of the encephalon as a whole and in all its subdivisions. In 
this respect the two sexes are similar. 

In chapter V, on Increase of the Brain in Weight and Variations 
of the Cranium in Capacity, the suggestive fact is brought out that the 
greater part of the growth of the brain takes place before any of the 
formal educational processes have begun. In speaking of the weight 
of the brain of non-European races, quotations are made from Hunt’s 
observations upon weights of the brains of mulattoes and negroes, in 
which he shows that the negro brain is uniformly less than the Europ- 
ean, and in those mulattoes with a mixture of less than half of white 
blood, the result is even below the negro average, while in a mixture of 
over half white blood, the average is between that of the negro and of 
the white. In the ‘‘ Variations in Brain Weight” the author states that, 
while the heaviest brains belong to the European races and the lightest 
to the Australians, yet it is quite impossible, even in a condensed ser- 
ies, to harmonize intermediate groups with the theory that brain weight 
and culture as we measure it, are closely correlated. 

In discussing the brain weights of eminent men a number of inter- 
esting tables are given, showing that it would appear that greater brain 
weights are more frequent among eminent men even when these are 
compared with groups of men of ordinary intelligence but of large 
stature. The fact is noted however that most of the comparisons have 
been made between ‘‘ eminents” 
inals,”’ 


and ‘‘ordinaries” or even ‘‘ crim- 
while the brains of reputable and successful and professional 
men have not been weighed. ‘The insane present encephala that are 
very slightly less than the normal in absolute weight; while the differ- 
ence between the sexes is less marked than in the sane. 

Chapter VII gives a good summary of the nervous elements accord- 
ing to recent modes of interpretation. The next chapter has an inter- 
esting outline of the development of the nerve elements, a subject 
that would be of value to pedagogues. Some very important and interest- 
ing collections of data are brought together in this chapter, the author 


xxxviii © JOURNAL OF COMPARATIVE NEUROLOGY. 


stating that the enlargement of the nervous system is due, first to the 
increase in number of the neuroblasts formed, and secondly to their in- 
crease in size. He thinks that in the growth from the child to the 
adult the increase in brain weight is due to the development of the 
medullary substance about the neuron. 

In the chapter upon the ‘‘ Architecture of the Central Nervous 
System” at maturity, there are given those anatomical features of the 
brain that are familiar to most students of anatomy, but in addition 
there are a number of interesting correlations that no mere text-book 
of anatomy ever gives. 

The chapter upon ‘‘ Changes due to Growth” tells us how little 
we know upon this important subject. 

Chapter XIII discusses localization of function giving the main well 
known facts upon the subject. The ‘‘ Physiology of the Nerve Cell” 
and ‘‘ Physiological Rhythm” present some of the results of later psy- 
chological work; while the chapter on ‘‘ Fatigue” gives some of the 
observations of Hodge and Vas, in which the author is inclined to lay 
more stress upon Hodge’s work than upon the experiments of others 
that have given different results. In the ‘‘ Education of the Nerv- 
ous System” there are a number of sound generalizations. 

This book is to be heartily recommended to the student and edu- 
cated layman. While it may at times be difficult for the latter to fol- 
low all the mathematical details, yet a persistent reading will give him 
a broad grasp of many important and useful facts. 

SMITH ELY JELLIFFE, M.D. 


Recent Studies in the Forebrain of Reptiles.’ 


The latest number of this series, which comparative neurologists 
have learned to prize as much for its appreciative summaries of the 
work of others as for the well-digested results of the author’s own pa- 
tient studies, is just at hand. Amongrecent writers Dr. Edinger has 
the honorable distinction of never slurring over or consciously misrep- 
resenting the work of others. We are often obliged to seek from other 
sources the evidence of the large share the author’s own work has had 
in attaining the results accredited to others. An error is always frankly 
admitted and the genial Doctor seems to feel that a personal favor is 
conferred by the writer who is fortunate enough to set him right. It is 


1 EpINGER, Dr. L. Neue Studien iiber das Vorderhirn der Reptilien. Un- 
tersuch. ii. d. vergl, Anat. des Gehirns, 3. Frankfurt, 1896. 


Literary Notices. xo ix 


little wonder therefore that views so formed and so expressed have great 
weight with his compeers. 

It will be impossible in the space at our disposal to recapitulate the 
abundant material offered by this paper. Edinger corrects the mistake 
made in earlier papers of failing to distinguish olfactory centres of the 
first and second orders, or (to use his own nomenclature) Azechfeld and 
Ammonsrinde. ‘Yhis whole subject has been elaborately discussed by El- 
liot-Smith, Meyer and the writer. About ninety series from a wide range 
of groups were employed by Dr. Edinger in his study and the variety of 
methods was adequate to insure mutual supplementation. The nomen- 
clature used is modeled on that of the recent report of the German No- 
menclature Commission. ‘Tracts are named as far as possible by com- 
pounding their termini. Instead of using the word lobe to apply to cere- 
bral regions Edinger seeks to avoid misleading analogies by such terms 
as ‘‘ cortex medio-dorsalis.”’ Although the olfactory fossa is mentioned, 
we miss any reference to the Jacobson’s organ fibres. In the basal lobe 
three parts are distinguished as striatum, mesostriatum, and epistriatum, 
terms, however, which may be open to the charge of suggesting pre- 
mature homologies. 

In the discussion of the histology of the cortex the results are 
similar to those of Cajal. ‘The anterior mantle commissure is no longer 
as formerly homologized with the callosum but with psalterium fibres 
as a ‘‘commtssura pallit anterior.” We regret that the ambiguous terms 
anterior and posterior should be given greater currency. It is to be 
noted that in some groups the anterior commissure contains mantle 
fibres so that the ambiguity is doubled. 

The following tracts are identified with the olfactory apparatus: 
The radtatio olfactoria. This is the radix lateralis of other writers. To 
this term the author objects on the ground that it suggests a homology 
with the roots of cranial nerves. ‘The criticism is well-founded but we 
are surprised to see the discovery of the true relations attributed to Cajal. 
We had supposed that the origin of the radix fibres in the cells of the 
bulb had long been recognized. Certain it is that a good haematoxylin 
stain reveals this relation as well as the Golgi impregnation. We have 
thus demonstrated it in the Amphibia. On the next page the same 
tract seems to be formally named ¢ractus bulbo-corticales, though a part 
of the fibres end in the cortex of the lobus olfactorius and part in the 
corpus epistriatum. Thus does each part rejoice in, not one, but often 
two new names from the same source—thanks to the efforts of the Ger- 
man Nomenclature Commission. The pero of Wilder, which the au- 
thor has hitherto professed himself unable to differentiate, appears as 


xl JOURNAL OF COMPARATIVE NEUROLOGY. 


the formatio bulbaris which ‘‘ results from the union of the olfactory 
nerve fibres with the processes of the ganglion cells.” We have de- 
voted so much space to this structure and its relation to the sheath for- 
mation and method of termination that it is with some surprise that we 
note the meagerness of detail and reserve as to well authenticated 
results. 

We do not gather exactly what is meant by the lobus olfactorius— 
in figure 5, it would seem to be the pes bulbi, but the description seems 
to make it plain that the frontal protuberance of the cerebrum forming 
the support of the bulb is intended. The ¢ractus cortico-epistriaticus 
consists of fibres rising in the lobus near the termini of the radiatio. 
From the area olfactoria arises the tractus olfactorius septi which Edinger 
identifies as our radix medialis. It terminates in the cortex of the 
hippocampal region—‘‘Ammonsrinde.” The term ‘‘ area olfactoria”’ is 
applied to an enormous extent and is divided into the medulus occipital, 
tuber thenie (pyriform lobe) as well as the post-rhinal lobe, to which it 
would seem to directly apply. The various bundles connecting with the 
thalamus are all included in the theenia thalami, including ¢vactus olfac- 
to-habenularis and tractus cortico-habenularts. It seems to the reviewer 
doubtful whether we are at present justified in referring all these fibres, 
with no more to do, to the olfactory apparatus. 

If we understand correctly, Dr. Edinger accepts the suggestion 
that the epistriatum is an invaginated cortical area. It is to be regretted 
that some comparisons were not made with the basal lobe of birds as 
described by Turner, who made out similar divisions and even found 
an invaginated cortex in somewhat similar relations. 

We are glad to note that the author has so far modified his earlier 
views as to recognize the probable existence of a tract connecting the 
optic centres with the cortex, ‘‘ Sehstrahlung aus den Opticuscentren 
zur Rinde.” 

Thus the earlier position that the cortex of reptiles is wholly given 
up to the olfactory function and its associations is somewhat modified 
but appears in the following form: ‘‘ Nur lasst sich sicher bestimmen, 
dass der grdsste Teil der Reptilienrinde mit dem Riechapparat 
Zusammenhangt.” 

It is indeed a most creditable achievement to have called attention 
to the psychogenetic significance of the preponderating position among 
the cortical stations held by those of smell, but we respectfully repeat 
our statement, made on several earlier occasions, that it is not correct 
to attribute an exclusively osmatic tone to the cerebral activities of the 
Sauropsida or even the Ichthyopsida. ‘The vast complexity of cortical 


Literary Notices. xli 


histology providing, as Edinger himself recognizes, for wonderfully com- 
plicated codrdinations, even did we not know of visual tracts to the 
cortex, would suggest a greater diversity of function. We fail to un- 
derstand why a connection between the homologue of the geniculatum 
and the cortex is overlooked. 

The present work is well adapted to serve as a point of departure 
for all subsequent comparative studies of the cerebrum. 

BET 1k 


Modifications of Nerve Cells in Different Functional States.? 


A brief historical introduction serves to throw into strong relief 
the utter disagreement existing between the results of almost all of the 
investigators who have thus far studied this question. The classical 
experiments of Vas upon the cervical sympathetic ganglia were re- 
peated in 1894 by Gustav Mann (cf. the review in Vol. V of this Jour- 
nal, p. xxxil) and again by Lugaro in the paper before us. The latter 
author feels that he has removed many of the contradictions of the 
previous investigators and explained the others. He calls attention to 
the fact that the so-called resting cells of other authors are by no means 
such but that to the fatigue of normal functioning before death 
must be added the intense discharge accompanying their excision as well 
as those changes occasioned by the action of the fixing reagent between 
the time of immersion and the time of cellular death. Another source 
of error is the subjective one arising from the difficulty of estimating 
the relative sizes of the normal and fatigued cells when the sizes of both 
sets vary within wide limits. The former difficulty was met by kill- 
ing in various ways, particularly by comparing cells fixed by rapid ex- 
cision and treatment with the fixing fluid with those taken several hours 
after death from an animal killed by rapid chloroforming ; the latter diffi- 
culty, by carefully counting all of the cells from many average fields 
from each of the two sets of preparations. In each ganglion the long- 
est diameter of 1000 elements was measured. 

The results of these observations are presented in the form of a 
series of curves. Cells of ganglia taken from the body five hours after 
death when compared with normal cells (taken from the body by vivi- 
section) show a decrease in size 4.95%. Elements electrically excited 
for a time which does not exceed a half hour show an increase in size 
as compared with the normal. ‘This increase is at the maximum of 


1 Lucaro, E. Sulle Modificazioni delle Cellule nervose nei diversi Stati 
funzionali. Lo Sperimentale, XLIX, 2, Aug., 1895. 


xlii JOURNAL OF COMPARATIVE NEUROLOGY. 


6.69% with an excitation of five minutes. If the excitation is contin- 
ued longer than a half hour there is a strong diminution in size of the 
cell. After excitation of one hour the diminution is .84%, after three 
hours 11.5%, after six hours 16.53%. 

Synthetic curves based upon the measurements show that the size 
of the cell rapidly increases during moderate activity, but under pro- 
longed stimulation diminishes far below the normal ; that the size of the 
nucleus follows the same law but to a less extent and less abruptly ; 
that the size of the nucleolus increases more rapidly than the size of 
the cell and that it diminishes under prolonged excitement much more 
slowly. The author concludes that the size of the cell is acted upon 
by two tendencies, one positive, activity, the other negative, fatigue. 
Letting d represent the dimension of a cell, 7 the normal (average) size, 
a the alteration due to activity, and / that due to fatigue, d will vary as 
expressed in the following equation : 

d=—n+n(a-f). 

The tardy alterations in the nucleus are regarded as consecutive 
to those in the cytoplasm. The nuclei, contrary to Hodge and Mann, 
do not wrinkle. 

The experiments are thus summarized : 

1. The activity of the nerve cell is accompanied by a state of 
turgescence in the protoplasm of the cellular body. 

2. Fatigue causes a progressive diminution in the size of the cell- 
ular body. 

3. In moderate degrees of activity, while the cytoplasm swells, 
the nucleus suffers no modification of volume. 

4. When the activity is continuous and of long duration, the nu- 
cleus suffers modifications analogous to these of the cellular body, but 
less intense and slower. 

5. The quantity of chromatic substance in the cellular body va- 
ries always as an individual character in proportion to the size of the 
cell. Nevertheless it is probable that the first phases of cellular activ- 
ity occasion a slight increase in its amount, the last phases accompanying 
fatigue, a diminution and a more diffuse distribution. 

6. The activity of the cell occasions in the nucleolus an increase 
in volume which gradually yields to the reducing action of fatigue. 

Following these experiments are some very interesting theoretical 
observations in which attention is called to the fact that if we extend 
the knowledge acquired of the changes in form of the body of the cell 
to include the cellular processes also, assuming that the processes elon- 


Literary Notices. xiii 


gate during functional activity and shorten during fatigue, we have 
taken a long step in the direction of a knowledge of the physical basis 
of the corresponding emotions, as well as of the physiological facts as- 
sociated with exercise and fatigue. CHE 


The American Lobster.' 

Six years ago Dr. F. H. Herrick was invited by the United States 
Commissioner of Fisheries to prepare a monograph on the Biology of 
the American Lobster. <A systematic investigation was begun and in 
this final paper, which has been awaited with considerable eagerness by 
the biologists, we have a summary not only of the author’s own re- 
searches, but of practically all that is known concerning the life history 
of this species. The work makes 252 pages quarto and is accompan- 
ied by 64 full page plates, many of them colored to life, illustrating 
very fully the development and gross anatomy. ‘The details of the fine 
histology are not entered into except in a few cases. 

The practical and scientific value of the work are both very great. 
It is not common to find both of these features so well represented in 
a single monograph. 

. One of the points of greatest practical importance, bearing on the 
problems of the artificial culture of lobsters, is the chapter on The 
History of the Larval and Early Adolescent Periods. The account of 
the Embryology of the Lobster is more meager than we should have 
expected from the fact that the author has devoted so much attention 
to this subject and has already published several short papers. He 
makes no attempt to give a detailed account of the embryonic history, 
but a few notes merely on the early phases of development. 

From our standpoint the topics of especial interest centre about 
the descriptions of the habits and organs of sense of the lobster. The 
breeding habits, habits of migration and feeding and the like are quite 
fully detailed. In its native haunts it shows considerable agility in 
avoiding its enemies and cunning in the capture of its prey. The 
senses of sight and and hearing are probably far from acute, but it pos- 
sesses a keen sense of touch and of smell and probably also a sense of 
taste. It is also quite sensitive to changes of temperature, this being 
the factor in the environment which is in the main responsible for the 
periodic migrations of the lobster. 

Experiments made with various stimuli, such as electricity, heat, 
weak acetic acid, ammonia gas and clam juice, led to the conclusion that 


1HERRICK, FRANCIS HOBART. The American Lobster: A Study of its 
Habits and Development. Auzlletin U. S. Fish Commission, 1895. 


xliv JOURNAL OF COMPARATIVE NEUROLOGY. 


all or nearly all of the appendages react strongly to chemical stimuli and 
in many cases the surface of the body is capable of receiving and re- 
sponding to stimuli of various kinds. In some places the skin of the 
lobster with its shelly covering seems quite as sensitive as that of the 
frog. ‘The organs by which the stimuli are conducted through the chit- 
inous shell must be either the hair pores or the glandular ducts. The 
author decides in favor of the latter. 
Cah ie 
Pain in the Pectoral Region Sympathetically Accompanying Irritation 
of the Forearm ' 

The patient, a fireman, developed very peculiar symptoms, prob- 
ably as the result of a fall and injury of the back and elbow. Con- 
stant though unequal pain is felt in the right shoulder on the right side 
of the back at the level of the seventh dorsal spine. On the right side 
of the chest, in front, there is an area bounded laterally by a vertical 
line from the axilla to the sixth rib and mesially by a line laterad of the 
right nipple. The pains felt here have a burning or tearing character. 
Beneath the right nipple is another painful spot which only announces 
itself after exertion, as rapid walking or turning in bed. ‘The posterior 
aspect of the right arm also is subject to pains, generally induced by 
pressure over the radio-humeral articulation. The area does not in- 
clude the fingers but extends upon the dorson of the hand. Bending 
the elbow causes severe pain, as does movement of the wrist, though 
there is no swelling of the joints. 

A touch on this arm area is felt in the pectoral area, as well as in 
the arm; firm pressure over any part of the arm area, except the pain 
centre over the joint, causes severe tearing pain in the pectoral area. For 
years the patient had been in the habit of sweating profusely with the 
right side only though when at work all parts perspire alike. A fall a 
week or so before the symptoms appeared injured the elbow and back 
and there seems to have been a blow of a less violent sort on the chest. 
The author thinks this a case of sympathetic neuralgia but there is dif- 
ficulty in discovering the nervous connections necessary to satisfy the 
hppothesis. C.as en, 


Corrigendum. 


On page ix of this issue in the review of Professor K6lliker’s arti- 
cle, line 7 from the bottom of the page, insert after ‘‘ habenulae, ” 
‘¢and ends in the ganglion interpedunculare.” 


1 Munro, T. K. Brain, IV, 1895. 


LITERARY NOTICES. 


The Development of the Brain and Sense-Organs in Elasmobranchs.! 


This well-prepared paper is somewhat misrepresented by its title, 
for it is in reality, in so far as concerns the original portions, a study of 
metamerism of the brain and the origin of certain sense organs based 
on embryos of Sgualus acanthias. ‘The questions to which this paper 
addresses itself are sufficiently ambitious, being such as the following : 
(1) What was the primitive condition of the nervous system of verte- 
brates? (2) What were the number and nature of the primitive neural 
segments entering into the brain? (3) What has been, in general, the 
line of modification along which they have been converted into the brain? 
(4) What were the early steps in the differentiation of the sense-organs? 
This is certainly a program calculated to awaken high expectations and 
itis no detraction from the genuine excellence of what is offered to 
admit that the materials and their employment are undoubtedly inade- 
quate to its fulfilment. The paper does, however, give very useful 
summaries of the historical development of those phases of the sub- 
jects which are taken up. 

The earlier part of the paper is occupied with the problem of 
metamerism. ‘The author apparently agrees with McClure in mini- 
mizing the segmental value of the cranial nerves because of the uncer- 
tainty due to the degenerations of certain branches and even entire 
nerves. ‘This may appear inconsequent when he, at the same time, 
explains the segmentation of the tube as due to the cranial nerves. 
This criticism is not necessarily valid, however, for it may be true that 
the segmentation of the brain was originally due to the concentration 
of nervous matter adjacent to the point of development of the segmen- 
tal nerves and yet this segmentation might persist in an early stage of 
animals in which the subsequent development processes had been di- 
verted. In other connections, however, the author fails to avail him- 
self of this possibility. Professor Locy does not admit that the seg- 
mental divisions of the middle germ layer are primitive. He accepts 


1 Locy, Wm. A. Contribution to the structure and development of the 
vertebrate head. Journ. Morphology, X1, 3, 1895. 


xlvi JOURNAL OF COMPARATIVE NEUROLOGY. 


the writer’s suggestion that ‘‘if neuromeres once existed in the fore- 
brain they would be visible only at an early stage. . . The so-called 
fore-brain neuromeres differ from those of the medulla in involving 
only dorsal structures.” He also apparently succeeds in resolving the 
the discrepancies between Froriep’s account and that of other writers. 

Fifteen pages are devoted to original observations on metamerism 
in Acanthias and both descriptions and plates are admirably done. 
Especial praise should be awarded the plate of photographs of the very 
young stages which are particularly valuable, as also the dissections re- 
produced in plate XXVIII. The constancy of the phenomena in ex- 
tremely early stages in various groups serves to justify the present 
writer’s prediction and to furnish a new motive for a re-canvass of the 
subject. The suggestion that they are artifacts is satisfactorily disposed 
of, though it is less clear that the node-like structure sustains a positive 
and constant relation to the definitive structure of the brain. Do they 
look backward rather than forward? 

This segmentation is said to be independent of mesodermic influ- 
ence and, as proof of this, the fact is cited that it precedes the meso- 
derm segments. It should be noted, however, that if it be conceded 
that the segmentation is not due to direct mechanical influence and so 
really has morphological significance, as the author claims, then we 
must look for the cause in a state when the head was fully segmented 
and mesodermic influence then could readily be appealed to. This the 
author would doubtless agree to. 

In Sgualus he recognizes as represented in the ontogeny at least 
fourteen paired neural segments. ‘The assignment of neuromeres to 
the sense organs and nerves is as follows : 

I. Olfactory. 
ies Optics | Fore-ba 
III. Pineal sense organ ? 


IV. Oculo-motor. . . 
V. Trochlearis. \ Mid-brain, 


VI, ‘Anterior’ (cephalic?) root of V. } 
VII. Main root of V. | 
VIII. No nerve. | 
IX. Facialis. L ry: . 
X. Auditory. [ Hind (Beata, 
XI. Glossopharyngeal. i 
XII. | 
XIII. | vagus J 
XIV. 


The segments are serially homologous. In Sgualus the optic vesi- 
cles are the first rudiments of sense-organs to appear and originate be- 
fore the separation of the fore-brain. Their peculiar form has led to 


Literary Notices. xlvii 


their being long overlooked. The curious segmental sensory patches 
which were described by the author as accessory optic vesicles arise in 
in a way similar to that of the eyes. ‘They first make their appearance 
when the neural plate is broadly expanded. Four pairs of such organs, 
at least, are noted, the cephalic pair forming, as is claimed, the pineal 
body. ‘They are embryonic and transitory structures and are thought 
to point back to a multiple-eyed pre-vertebrate condition. 

A review of the literature upon the pineal seems to the author to 
lend force to the suggestion of a multiple-eyed ancestor of vertebrates. 
The results ‘‘ go to show that there are two distinct outgrowths of the 
thalamencephalon of Petromyzon, Teleosts and Lacertilia.”” The ceph- 
alic one is that developed into the pineal eye in front of the epiphysis 
in Lacertilia, and corresponds to Hill’s anterior vesicle in teleosts. 

In passing, it seems that a halt ought to be called in the unjusti- 
fiable introduction of untranslated descriptive terms from the German. 
‘¢ Zirbelpolster ” is not an English word and is not a desirable addition. 
If a popular term is needed it may be easily translated but it is much 
better to substitute a Latin equivalent. Our European friends should 
also realize that the scientific world cannot consider an organ properly 
named and presented for acceptance until a name has been proposed 
conformable to the common nomenclature of the scientific world. 

The auditory organ is briefly discussed and its relation to the lat- 
eral line system recognized. 

The paper supplies many details that have been wanting hitherto 
and has several pregnant suggestions, but its chief value is in the com- 
pleteness and apparent accuracy of illustration. 

CPL. He 


The Sensory Physiology of Actinians.! 


Recognizing the growing importance of the comparative physiolo- 
gy of the nervous system, especially among the lower invertebrates, 
Dr. Parker has undertaken a series of experiments on the common ac- 
tinian of the Atlantic coast, Metridium marginatum. In taking food, 
two kinds of responses are met, ciliary and muscular; only the latter 
shows evidence of nervous control, Numerous experiments illustrate 
the slight physiological centralization in the nervous functions of these 
organisms. Each tentacle acts like an independent organism. ‘The 
curious fact is observed that the cilia on the lips reverse their motion 


1PARKER, G. H. The Reactions of Metridium to Food and other Sub- 
stances, Bul. Museum Comp. Zool., XX1X, 2, March, 1896. 


xl viii JOURNAL OF COMPARATIVE NEUROLOGY. 


during the act of swallowing. ‘The usual stroke of these cilia is in a 
direction away from the mouth, as is shown by the motion of particles 
of carmine or other insoluble substance. But when a piece of meat or 
a drop of meat juice touches these cilia they reverse their motion after 
an appreciable latent interval. 

G.. jel 


The Histology of the Myelon.! 


The ‘* Retssner’s” or ** Dorsal Cells” of the Cord. Many years 
since, peculiar cells were noticed in the dorsal portion of the spinal cord 
of Petromyzon. Reissner described them in 1860, and, in 1877, 
Freud succeeded in tracing their axis cylinders into the dorsal root, 
though it has since been shown by Beard that they do not originally 
emerge with the sensory fibres and are of a kinesodic nature. Rohon, 
in 1885, detected similar cells in the trout and they have since been 
noticed in the embryonic stages of other Gnathostomata. According 
to Studnicka, however, these cells, though distinguished from adjacent 
elements by their great size, are not homologous with the ‘‘ colossal 
cells” of the spinal cord of Amphioxus. ‘The latter are median in po- 
sition and are sparsely distributed; their axis cylinder passes laterad and 
arches ventrad and, after crossing to the opposite side, extends toward 
the head. Some of their smaller processes were traced by Koelliker 


Horizontal Jongitudinal section of Reissner’s cells from dorsal region 
spinal cord of Ammocetes. of spinal cord of Pristiurus. 


into the dorsal roots. Respecting these Studnicka suggests that they 
may probabiy be regarded as inherited from some invertebrate prede” 
cessor. He claims that the ‘‘ medium-sized” cells of Amphioxus cor 
respond with the Reissner’s cells of higher forms. ‘These are bipolar, 


ISTuDNICKA, F. K, Ein Beitrag zur vergleichenden Histologie und His- 
togenese des Riickenmarkes. Sttzd. Aongl. bihm. Gesellsch., 1895. 


Literary Notices. xlix 


occupying a lateral position with respect to the central canal. One of 
the main processes (Studnicka calls them both axis cylinders) passes in 
a longitudinal direction, while the other emerges with the dorsal root. 
If this identification be correct Reissner’s cells occur throughout life in 
Amphioxus, Petromyzon, Protopterus, ‘Triton, and certain teleosts, 
while in the selachians and ganoids, as well as some others, they are 
embryonic and transient. 

In Lophius it will be remembered that they have a very restricted 
but specialized occurrence. The form is generally crescentic with the 
concavity mesal. One of the processes crosses to the opposite side of 
the cord and passes either toward or from the head. ‘The other or 
true axis cylinder originally passed directly ectad and innervates the 
myotome, though in some cases it associated itself with the dorsal 
root. Burckhardt’s ‘‘ Randzellen ” described in Protopterus are found 
to occur in Petromyzon also, but they have nothing to do with the 
Reissner’s cells. ‘The term selected by Studnicka, ‘‘ Hinterzellen,” 
suffers from ambiguity common to the German terms of direction. If 
we substitute ‘‘ dorsal cells” there is a possibility of confusion with 
cells of the dorsal cord so that the familiar term Reissner’s cells may 
still be employed. 

In a second paper! the same author describes the expansion of the 
canalis centralis found at the caudal extremity of the cord of Amphi- 
oxus and the Cyclostomata and differentiates it under the name ‘‘ Sinus 
terminalis” from the ‘‘ Sinus sacralis”” of acaudal vertebrates. Such 
significance as this ventricle has may be assumed to be in connection 
with the equilibrium of the cerebro-spinal fluid. 

(Go) Whig Tels 


Anatomy of the Cerebrum of Notoryctes. ” 


Although based on imperfect material and studied under adverse 
conditions, the notes upon the brain of this little-known marsupial will 
prove welcome. In respect to the brain at large the small size of the 
cerebrum and limited extent of pallium are correlated with the exposure 
of a relatively large part of the corpora quadrigemina and a very sim- 
ple condition of the cerebellum. 

The cerebrum presents an extreme simplicity of structure and 
while is has superficial resemblances to Talpa and Dasypus it is yet in 


1Ueber die terminale Partie des Riickenmarkes, 


2G, ELLIoT-SMITH. The Comparative Anatomy of the Cerebrum of No- 
toryctes typhlops. Zyrans. Roy. Soc. Australia, 1895. 


] JOURNAL OF COMPARATIVE NEUROLOGY. 


morphological plan identical with the apparently dissimilar Macropus. 
The mesal olfactory fossa is absent but it does not appear whether this 
is correlated with any peculiarity of the Jacobson’s organ. 

The pyriform lobe takes an increasing share in the formation of 
the lateral part of the hemisphere as the extent of pallium decreases. 
The region reaches the height of its development early in ontogeny and 
phylogeny, hence its surface always remains smooth. Herrick and 
Bawden are criticised for confusing of the hippocampus when they 
evidently mean the pyriform but this criticism apparently rests ona 
misapprehension. ‘The tuberculum olfactorium is of great size. Hill’s 
usage in excluding this body (the post-rhinal lobe) from the rhinenceph- 
alon is characterized as ‘‘ ridiculous” —an expression which must be 
accepted as applying to the critic’s risibilities alone and which over- 
steps as a personal reflection the limits of professional courtesy. The 
author evidently meant ‘‘ preposterous” or ‘‘ inconsistent” but this 
must depend on the morphological extension involved in the term 
rhinencephalon. 

Additional illustrations are given presumptively in favor of the 
view that the ‘‘ commissura pallii anterior” is not a callosum, but inci- 
dentally illustrating the long-questioned fact that there are two dorsal 
commissures in marsupials, which, after all, has been that chiefly in 
the minds of the writer and others who have sought to enforce the 
distinction between the hippocampal and callosal portions of the dorsal 
commissure. 

The reader may seek the original fora variety of details and 
generalizations. 

The illustrations are far from successful. (Ory BPs 


Romanes on Weismannism. ' 

This new edition of the ‘‘ Weismannism” is well printed and fur- 
nished with an excellent portrait of the author. It is one of the few 
really good cheap books which stand in so marked contrast with the ten- 
dencies of the time in book making. We have already noticed an earlier 
edition, but would again call attention to this discussion of the theories 
which Weismann has erected on the basis of his fundamental postulate 
of the non-inheritance of acquired, or somatic characters, as assuming 
a new importance in the light of the more recently published work of 


1 An Examination of Weismannism. By GEORGE JOHN ROMANES. Chicago, 
The Open Court Publishing Co., 1896, Price 35 cents. 


Literary Notices. li 


the same author in which he has discussed the truth of this postulate. 1 
The two books should be read together, if one would gain a true view 
of Mr. Romanes’ views on these important questions. Caja er: 


Germinal Selection. 2 


The address delivered by Dr. Weismann before the International 
Congress of Zoologists at Leyden, 16 Sept., 1895, is here expanded 
and translated. The fundamental difficulty urged against Weismann and 
the other so-called neo-Darwinists by their critics is the impossibility of 
accounting for many of the adaptations as we find them on the basis of 
a theory of natural selection which recognizes only indefinite variabil- 
ity. The simultaneous occurrence of the variations necessary to pro- 
duce, e. g., a case of mimicry in a butterfly, is hardly credible except 
on some hypothesis of definitely directed variability. The hypothesis 
of germinal selection assumes that when natural selection acting on 
accidental somatic variations in the manner usually described has re- 
sulted in a slight increase in the efficiency of the part in the given di- 
rection, then the determinants corresponding to that part in the germ- 
plasm of the next generation will be more vigorous than the determi- 
nants of other parts and in the struggle for food will outstrip them. 
Thus that part will in the adult be stronger than it was in the previous 
generation, and this result will be cumulative. So the germ is progress- 
ively modified in the direction set by utility. 

Though very little has yet been published on this specific phase of 
the problem, the general question of definitely directed variation is one 
to which every thoughtful biologist must have given some attention. 
The success of this attempt to explain the mechanism of the process 
will rest on the value of Dr. Weismann’s scheme of the hereditary ma- 
chinery. And in estimating this we must recognize the purely symbolic 
nature of the conceptions used by the author of this scheme, a fact 
of which, as he points out, his critics have often failed to take account. 

To one who, like the reviewer, is unwilling to admit the absolute 
non-transmissibility of somatic characters it is obvious that the argu- 
ment for germinal selection cannot be accepted exactly as here out- 


lined ; yet the principle may be a true one and is not necessarily incom- 
patible with an application of some form of the Lamarkian doctrine as 
well. (Sei fouls 

1 Darwin and After Darwin. II. Post-Darwinian Questions, Heredity and 
Utility. Chicago, The Open Court Publishing Co., 1895. 


2 WEISMANN, AuGusT. On Germinal Selection. Chicago, The Open Court 
Publishing Co., 1896. Price 25 cents. 


lii JOURNAL OF COMPARATIVE NEUROLOGY. 


The Psychology of Attention. 


This is the third revised edition of the authorized translation pub- 
lished by the Open Court Co., Chicago. There is perhaps no writer 
who has done more to popularize the idea that a neurological founda- 

.tion is essential to the full comprehension of the higher mental activi- 
ties than Professor Ribot. Of the value of this standpoint we have no 
better proof than the stimulus which has been given to recent research 
by Professor Ribot’s own works. Even those who cannot accept all of 
their conclusions are ready to acknowledge their indebtedness to them 
for many pregnant ideas. 

We can summarize the present book no better than in the author’s 
own words. ‘‘ Attention depends on emotional states ; emotional states 
are reducible to tendencies; tendencies are fundamentally movements 
(or arrested movements) and may be conscious or unconscious. Atten- 
tion, both spontaneous and voluntary is, accordingly, from its origin on, 


bound up in motor conditions.” 
Cojo 


Studies from the Yale Psychological Laboratory, 


Volumes I and III of these studies are on our table. Volume II 
has already been noticed. The first volume is concerned mainly with 
investigations in reaction time. The most extensive article is the thesis 
of Dr. Bliss on ‘‘ Reaction-time and Attention.” Throughout this 
series of many hundreds of reactions especial attention was paid to the 
subjective state of the reagent. After having demonstrated that per- 
fect darkness in the reacting room is in no way preferable to a uniform 
illumination, the reagent—in most cases Dr. Bliss himself—sat with 
pencil and paper at hand and at the close of each series of experiments 
annotated them. In many respects the most valuable part of the pa- 
per is the analysis of the reactions in the light of the data of introspec- 
tion. This principle, which has been more recently recommended and 
practiced at the Paris laboratory, is one of great importance. The 
prevailing German tendency to reduce the reacting subject to a mere 
automaton, while in many experiments useful, can never give the whole 
truth. 

Not the least valuable part of the paper by Dr. Bliss is the very 
full description of the apparatus and the details of the connections em- 
ployed. Indeed this is an important feature of nearly all of the stud- 
ies included in these three volumes. From the inauguration of the 
Yale laboratory the devising of new apparatus and laboratory conveni- 
ences has been given a prominence which puts all other workers under 


Literary Notices. liti 


obligation, an obligation which the present writer gladly acknowledges, 
as he has found many of these contrivances very useful in his own la- 
boratory. 

The leading article in the third volume is by Dr. C. E. Seashore 
on ‘‘ Measurements of Illusions and Hallucinations in Normal Life.” 
This is a paper of vital importance to every laboratory psychologist, 
for the conclusions reached have an important application to some of 
the most fundamental methods of the experimental method. By an 
elaborate series of experiments, the details of which must be sought in 
the original, Dr. Seashore has shown the truly surprising ease and _viv- 
idness with which it is possible to evoke illusions and hallucinations in 
every department of sense and that too in experienced psychologists 
and under the conditions of rigid control of the laboratory experiment. 
The hallucinations here described are evoked mainly in response to 
forced expectant attention and it is shown that this is one of the fac- 
tors which must be guarded against in all experiments on liminal differ- 
ences. Certain experiments on discrimination and many other classes 
of laboratory studies will receive cautionary hints from this paper. 

The ‘‘ Studies of Fatigue” by John H. Moore take up the effect 
of fatigue on binocular estimate of depth, the effect of fatigue on mon- 
ocular estimate of depth, the effect of fatigue on the time of monocular 
accommodation, and the effect of fatigue on the maxium rate of volun- 
tary movement. Several points of practical pedagogical interest are 
developed. For instance, several of these experiments on the estima- 
tion of depth by means of the muscles of accommodation and con- 
vergence emphasize the extreme danger to the eyes of carrying on for 
long periods of time such accommodation as must be employed in 
copying unfamiliar subjects from the blackboard to the paper or slate, as 
is so often done even in the lower grades of our schools. (Coal ini zis 


The Embryology of the Medulla of the Rabbit.' 


In this paper Dr. Dexter has followed the development of the 
medulla of the rabbit in order to make comparisons with that of man 
as worked out by His. He finds several differences, some of them 
quite unexpected. ‘The tractus solitarius is buried more deeply in the 
medulla as development progresses by the migration, as the author sup- 
poses, of scattered ectodermal elements lying on the border of the me- 
dullary wall and ‘‘ Randschlier,” and not by the formation of a rhom- 


'FRANKLIN DEXTER, M.D. A Contribution to the Morphology of the Me- 
dulla Oblongata of the Rabbit. Privately printed from the Archiv fiir Anatomie 
und Physiologie. Boston, 1896. 


liv JOURNAL OF COMPARATIVE NEUROLOGY. 


boidal lip as in the human medulla. The rhomboidal lip as described 
by His was nowhere developed. ‘The tractus solitarius is not regarded 
as wholly homologous with the oval bundle of the cord. The tractus 
solitarius is shown to receive fibers from the V, VII-VITL, Re andi 
ganglia very much as in the Amphibia Cees 


Uranism.! 


This extensive work of over 350 pages is devoted to a thorough 
exposition of the subject of unisexuality in all of its aspects. Itis a work 
of more limited scope than Krafft-Ebing’s famous ‘* Psychopathia”” and 
is written from an entirely different standpoint. It resembles that 
work, however, in the vast wealth of its historical, statistical and crit- 
ical data. It is not our purpose to attempt an abstract of the author’s 
conclusions. With many of his positions our readers are already fa- 
miliar. We need only add that we are in full accord and sympathy 
with his thesis that the misfortune of congenital sexual inversion is in 
no sense a justification for any form of sexual immorality. The subject 
is confessedly a confusing and perplexing one in all of its phases, but 
this point we think Mr. Raffalovich has quite satisfactorily established. 
And this is by no means incompatible with the further claim that the sex- 
ual invert ought not to be expected to conform in all respects to the 
mode of life of those who are normal in this respect. But if it should 
prove impossible for him to conform to this norm, it does not follow 
that he should not put the same restraint upon himself to which many 
another and perfectly normal person must often submit himself. 

Ciao 


Electricity in Electro-Therapeuties.” 


This little book is, as the title indicates, a treatise on electricity, 
nor a treatise on electro-therapeutics. ‘The authors have succeeded 
for the most part well in their attempt to so present the principles of 
electricity that any physician may comprehend them, even though he 
may have had no previous technical training. ‘The book is to be com- 
mended as just what the practitioner needs to enable him to use intelli- 
gently the electrical apparatus with which his office is, or should be, 
supplied. Coie ae 


> 


1Uranisme et Unisexualité. Etude sur différentes manifestations de ]’instinct 
sexuale. By MARC ANDKE RAFFALOVICH Bibliothéque de Criminologie. 
Lyon, 1896. 


Electricity in Electro-therapeutics. By Epwin J. Houston, Ph.D., and 
A. E. KENNELLEY, Sc.D. Mew York, The W./. Johnston Co., 1896. Price $1.00. 


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