THE C E R E B E L L U M O F hfYXINOIDS AND P E T R O -

AIYZONTS INCLUDING D E V E L O P N E N T A L
S T A G E S I N THE L A R I P R E P S
0 . LARSELL
Anato?nical Lnbmatory, U n i t e l a i t y of Orep71, Uedical School, Portland

THIRTY FIGURES

INTRODUCTION

The hagfishes and the lampreys sliow such great differences

in development of the cerebellar region that common features
a r e difficult to recognize. Interpretations of the hagfish brain
have ranged from no cerebellum present to the view that tlie
region now recognized as midbrain represents the cerebellum.
The latter viewpoint was advanced especially by Joliannes
Miiller (1838), Sanders (1894), Worthington ( ’05) and Holm-
gren (’19), while Anders Retzius (l822), Holm (’01) and
Edinger (’06) considered it a s midbrain. Gustaf Retzius
(1893) states (p. 56) that the medulla oblongata of hlysine
has no trace of a cerebellum, but describes (1). 61) tentatively
a s “Hinterhirn” the part of the brain which Conel ( ’29, ’31)
and Jansen (’30) have shown to be midbrain. Retzius, how-
ever, does not refer to it as cerebellum. The more recent
studies of Conel and of Jansen, above cited, have clarified the
difficult problems of the morphology of the mysinoicl brain and
prepared the way for a restudy of tlie cerebellar region.
Conel (’29, p. 378) stated that a “small cerebellum is
probably present a t the cephalic end of tlie rhornheiicephalon”
in embryos of Bddlostorna stouti, but that further study was
necessary before he could be positive a s to its presence or
absence. Subsequently (’31, p. 403) he stated, “If there is a

395
396 0. LARSELL

cerebellum in Bdellostoma, it is in the anteromedial part of

the roof of the rhombencephalon which forms the posterior
wall of the plica rhombomesencephalica. ” He recognized
fibers, in some stages of the embryos, which could be inter-
preted as a cerebellar commissure, and also a ridge of cells in
the blase of the plica rhombencephalica connected by these
fibers, but concluded (p. 421) that “the evidence is too scant to
permit one to say definitely that this ridge of cells constitutes
a cerebellum.”
Jansen (’30, pp. 494-496) has reviewed the conflicting
interpretations of the region which he and Conel have shown
is midbrain. Holmgren (’19) has described a commissure
cerebellaris which Jansen held (p. 443) is the commissure
posterior tecti, although admitting that it may contain a few
fibers of a rudimentary cerebellar commissure connecting
the 2 auricular regions. Jansen was unable, however, to es-
tablish such connections.
A t the Cleveland meeting of the America11 Association of
Anatomists in April, 1946, the present writer described a
commissure between the acousticolateral areas of adult Bdel-
lostoma, with cells along its lateral extent 011 each side, as rep-
resenting the incipient cerebellum. The description, published
in abstract (Larsell, ’46) was illustrated with photomicro-
graphs of Cajal preparations. I n a paper which probably
was in press when the above mentioned paper was read Holm-
gren (’46) described a cerebellar commissure in 2 embryos of
Myxine glutinosa. I n the larger embryo nerve cells surround
the commissure. Holmgren states that there is “strong evi-
dence i n favor of accepting the presence of a not insignificant
cerebellar portion of the ‘midbrain’ in front of the cerebellar
cornmi~sui-e,’’ describing the cerebellar commissure as con-
taining a bundle of fibers which enters the midbrain and splits
up in it. This point will be discussed further in the descriptive
portion of this paper.
All students of the petromyzont brain have recognized a
cerebellum and Johnston ( ’02) established the foundations
of a clear interpretation of this organ, in part, in his study
 CEREBELLUM O F CYCLOSTOMES 397

of the brain of the brook lamprey. Clark ('06) and Pearsoii

( '36) have described the cerebellar connections in several
species of petromyzonts.
In the present paper the cerebellar region of adult Bdello-
stoma is described from material prepared by the method of
Ramcin y Cajal, and a restudy of the petroinyzont cerebellum,
including the principal features of its deveiopmeat, is pre-
sented, based on the large collection of material indicated
below.
MATERIAL AND METHODS

Many years ago the late Dr. Howard Ayers presented the author
with a number of brains of aclult Bdellostoma stonti (Lockington)
prepared by the method of Ram& y Cajal. These subsequently were
sectioned, forming the following :
3 series cut in the horizontal plane.
3 series cut in the sagittal plane.
4 series cut in the transverse plane.
The petromyzont material studied consisth of the following series :
Adult brains.
Entosphenus tridentatus (Gairdner) :
1 series, iron-hematoxylin, horizontal.
1 series, iron-hematoxylin, transverse.
1series, Cajal, sagittal.
1 series, Cajal, transverse.
1 series, Cajal, horizontal.
Entosphenus appendix : 1 series, pyridine-silver, horizontal, IIuher
Collection €120.
Ichthyomyzon unicolor : 1 series, toluidine blue, transverse, Huber
Collection H4.
Ichthyomyzon concolor : 1 series, iron-hematoxylin, transverse, C. J.
Herrick series.
Petromyzon niarinus unicolor :
1 series, pyridine-silver, horizontal, Huber Collection H18.
1 series, pyridine-silver, sagittal, Huber Collection H17.
1 series, toluidine blne, transverse, IIuber Collection H2.
Lampetra wilderi (J. B. Johnston series) :
4 series, Golgi, horizontal.
5 series, Golgi, transverse.
2 series, Golgi, saKitta1.
398 0. LARSELL

Animocoete brains, Entosphenus tridentatus :

7 mm, 1 series, iron-hernatoxylin, transverse.
7 nim, 2 series, borax-carmine and Lyons blue, transverse.
7 nim, 1 series, borax-carmine a n d Lyons blue, sagittal.
42 inm, 1 series, protargol, transverse.
50 mm, 1 series, iron-hematoxylin, sagittal.
55 nim, 1 series, borax-carmine and Lyons blue, transverse.
53 iiini, 1 series, borax-carmine and Lyons blue, sagittal.
63 mni, 1 series, protargol, transverse.
65 mm, 1 series, borax-carmine and Lyons blue, sagittal.
71 nim, 1 series, protargol, transverse.
120 mm, 1 serivs, protargol, transverse.
I25 nim, 1 series, protargol, horizontal.
127 nim, 1 series, protargol, sagittal.
133 m m , 1 series, hematosylin and eosin, transverse.
133 mm, 1 series, borax-carmine (celloidin), transverse.
147 mm, 1 series, protargol, transverse.
160 nim, 1 series, protargol, transrcrse.
The Golgi series above indicated were prepared by the late Dr. J. E.
.Johnston and were the basis of his paper entitled, “ T h r Brain of
PPtromyzon,” published i n 1902. Thej- were given to me by Dr. A. T.
Rasniussen a number of years ago. The series indicated “ H u b e r
Collection” were kindly placed a t my disposal by Ih. Elizabeth C.
Crosby. They had been used by Dr. A. A. Pearson in his study of
the petroinyzont cerebellum arid acousticolateral area. The series
of Ichthyornyzon concolor has been loaned me by Dr. C. Judson Her-
rick. It served as tlie hasis of the paper o n the brain of Ichthyoinyzon
published by Herrick and Obenchain in 1913. The series of Ento-
sphenns tridentatus, larval and adult, have been prepared by the author
a? tlie study progressed. The serial sections have been supplemented
by dissections of the adnlt brains of Entospheiins tridentatus and
Bdellostoma.
To Doctor Crosby, Doctor Herrick and Doctor Ra.;nius~enI express
my appreciation for the material they have made available to me and
my thanks also are due Dr. R. L. Bolin, of tlie Hopkins Marine Station,
for heads of Bdcllostonia.

DESCKIPTIVE
BDELLOSTOX A

Ayers aiicl TTrortliiiigton ( ’08) described the acousticolateral

area in Bdellostoiiia under the name “acusticus nucleus” a i d
stated that it receives 5 sets of fibers, namely, “those of tlie
 CEREBELLUM O F CTCLOSTOMES 399

r. acusticus utrioulus, of r. acusticus saccularis, of n. lateralis

posterior and 2 sets f o r n. lateralis anterior.” These investi-
gators found 2 classes of cells, namely, large and small, the
latter preponderating and being scattered throughout the
nucleus. The small cells include both spindle-shaped and multi-
polar types. The large cells also are scattered throughout the
nucleus. Many of those lying on or near its borders send their
processes into adjacent parts of the brain, some into the gen-
eral cutaneous nucleus, while cells of the latter nucleus also
send processes into the nucleus acusticus.
Among the fiber tracts, Ayers and Worthington described a
small “tractus acustico-cerebellaris,” arising “near the
cephalic end of the acusticum dlong the mesial border, near
the mid dorsal-ventral plane. It runs caudo-mesad close t o
the mesial surface for about 1.2 mm; then it turns mesad
into the cerebellum, runs slightly dorso-mesad, and crosses
to the other side. After crossing its fibers separate and turn
cephalad. They run dorso-latero-cephalad in small bundles for
varying distances, and are eventually distributed in the
cerebellum. ” I n Cajal series of Bdellostoma I have found
fibers corresponding t o this description, but in view of the
demonstration by Cone1 ( ’29, ’31) and Jansen ( ’30) that the
portion of the myxinoid brain which Ayers and Worthington
considered a s cerebellum, is in fact the midbrain, these fibers
must be regarded a s a n acousticolateral-mesencephalic tract.
It seems likely that the cerebellar commissural fibers described
by Holmgren ( ’46) in Myxine embryos as ending in the mid-
brain represent the same tract.
Jansen’s description of the acousticolateral area of adult
Ifyxine is in essential agreement with Ayers and Worthington,
but no mention is made of the “acustico-cerebellar ” tract.
Possibly Jansen’s “loose bundle of fine fibers” (Jansen, ’30,
p. 463) which often cross the midplane dorsal to the ventricle
in the region caudal to the isthmic fissure constitutes this tract,
in part.
The brain of Bdellostoma does not show the cerebellum ex-
ternally (fig. 1). It differs but slightly from the brain of
 ABBREVIATIOR’S
aq.cer., cerebral aqueduct.
ax., axon.
bLv., blood jessel.
cb., cerebellum.
cb.v.pl., cerebellovelar plate.
co.ac.lat., acousticolateral commissure.
eo.ac.lat.p.nuc.dors., dorsal nucleus com-
ponent of acousticolateral eommis-
sure.
co.p.t., posterior tectal commissure.
C O . ~ . ,velar commissure.
co.ven., ventral commissure.
cr.cb., cerebellar crest.
dec.r.IV, dccussation of I V t h root.
dien., diencephalon.
ep., ependyma.
f.isth., fovea isthmi.
fi.int.arc., internal areuate fibers.
fi.r.VII.l.l.v., fiber of ventral lateral line
V I I t h root.
gl., glomeruli.
gn.V, ganglion of Vth nerve.
gn.VII1, ganglion of V I I I t h nerve.
gn.max.mand., niaxillo-mandibular gan-
glion.
gn. oph.prof., deep ophthalmic ganglion.
gn.sac., saccular ganglion.
gn.utr., utricular ganglion.
hab., habenula.
lo.ac.lat., acousticolateral lobe or area.
lo.olf., olfactory lobe.
lo.opt., optic lobe.
M.c., Miiller cell.
rndobl., mednlla oblongata.
mes., mesencephalon.
n.1, olfactory nerve.
n.11, optic nerve.
n.IV, trochlear nerve.
nuc.IV, nucleus of I V t h nerve.
nue.V.m., motor nucleus of Vth nerve.
nuc.V.s., sensory Vth nucleus.
nuc.VII1.m. ant., anterior motor V I I I t h
nucleus.
n.sp.l., first spinal nerve.
nb., neuroblast.
nuc.eb., nucleus cerebelli.
nue.dors., dorsal nucleus of acoustico-
lateral lobe.
nuc.med., medial iinrleus of acoustico-
lateral lobc.
400
nue.ven., ventral nucleus of acoustico-
lateral lobe.
ot., otocyst.
p.g.c., primitive granule cell.
p.P.e., primitive Purkinje cell.
p.st., pineal stalk.
pl.ch., chorioid plexus.
pli.ene., plica encephali.
r.IV, root of I V t h nerve.
r.V, root of Vth nerve.
r..V ds., descending root of Vth nerve.
r.Vm., motor root of Vth nerve.
r.V s., sensory root of Vth nerve.
r.VI1, root of V I I t h nerve.
r.VII.l.l., lateral line root of V I I t h
nerve.
r.VII.l.l.v., ventral root of lateral line
V I I t h nerve.
r.VII.l.ld., dorsal root of lateral line
V I I t h nerve.
r.VII1, root of VIIItli nerve.
r.VIII.as., ascending root of V I I I t h
nerve.
r.ant.VII1, anterior root of V I I I t h
nerve.
r.post.VII1, posterior root of VIIItli
nerve.
r.IXfX, roots of glossopharyngeus and
vagus nerves.
r.X, roots of vagus nerve.
r.X.1.1., lateral line root of vagus nervr.
nuc.hab.d., right Einbenular nucleus.
nuc.hab.si., left liahenular nucleus.
sp.c., spinal cord.
str.gr.eb., cerebellar gray layer.
str.mol., molecular layer.
t., taenia of fourth ventricle.
tr.VIII.as., n scending VIIItli tract.
tr.VIII.m.ant., anterior motor V I I I t h
tract.
t.ch., tela chorioiden.
tel., teleneephalon.
tr.cb.teg., eerebello-teemental tract.
tr.sp.cb., spino-cerebellar tract.
tr.sp.t., spino-tectal tract.
tr.t.v.+t.cb., teeto-velar and terto-crre-
bellar tract.
tr.trig.cb., triRemino-cerebellar tract.
v.4, fourth ventricle.
vx., ventricular canal.
 CEREBELLUM O F CYCLOSTOMES 401

Myxine, a s figured by Gustaf Retzius (1893), but is somewhat

more elongated and the mesenceplialoii is smaller. The posi-
tion of the acousticolateral commissure, more fully described
below, which marks the beginning of cerebellar structure is

Fig. 1 Dorsal view of the I:rain of Bdtllostoma stouti. X 7.5.

shown in figure 2, i n the midsagittal plane, while fignre 3

shows this commissure in relation to the acousticolateral areas
or lobes and to other parts of the medulla oblongata a h well
as to the midbrain, a s seen in transverse section. Figures 4
and 5 show the commissure more clearly and also show a
402 0. LARSELL

medial extension of small cells from the acousticolateral area.

of either side. These cells lie between the bundles of large
arcuate fibers and the acousticolateral commissure, approach-
ing the midplane but not reaching it. Many of the cells are
spindle-shaped but, lacking Golgi material, I cannot state
with certainty that the others a r e multipolar. All have the
same appearance a s those of the acousticolateral area and
they also correspond in size. A thin nexus of cells is contin-
uous with the scattered cells of the periventricular gray, to
which the connecting cells correspond in size.
Beginning at the medial margins of the main mass of the
acousticolateral area on each side, a bundle of very fine fibers
extends medially, arches over the ventricular canal and passes

Fig. 2 Outline of midsagittal section of the brain of Rdellostoma. Method of

Ram6n y Cajal. Edinger-Lcitz projection apparatus. X 14.5.

ventrolaterally, to be lost among the cells of the opposite

acousticolateral area. The bundle is augmented by fibers from
the medial extension of cells ahove described. The latter fibers
form a small, separate bundle which, however, intermingles
with the larger one above the ventricular canal (fig. 5). The
combined commissnral bundle then passes through the junc-
tion of the midbrain with the thin roof of the ventricular
canal. In the midsagittal region the bundle lies just above
the ependymal layer before the latter continues caudally as
the roof of the ventricular canal (fig. 6A). More laterally it
is seen, both i n sagittal and transverse sections, to pass into
the acousticolateral area (figs. 5 and 6B). It is somewhat
intermingled with the posterior tectal commissure but its
fibers a r e finer and more brownish in color. Also they arch
 CEREBELLUM O F CY CLOSTOMES 403

Fig. 3 Photomicrograph of transverse section of brain of Bdellostoma a t plane

marked a-b in figure 2. Method of Ram& y Cajal.

Fig. 4 Photomicrograph of medial region of same section, a t higher magnifica

tion, showing acousticolateral commissure. Method of Ram6n y Cajal.
404 0. LARSELL

laterally and d o w n w a d , a s described, instead of laterally and

upward as do the fibers of the posterior tectal commissure.
It is possible that some pass into the posterior tectal coin-
Inissure, but close study of the sect'ions indicates that the
brownish fibers all pass veritrolateral1~- into the acousti-
colateral area. There is thus formed a dorsal commissure be-
tween the acousticolateral lobes of the 2 sides which is the fore-
runner of the fibrous connection between the auricles of uro-
deles (Herrick, '14, '24; Larsell, '20, '31, '32) which we
have called the lateral commissure. As seen in the haqfish, this

CO.ac 1at lo ac 1a.t

Pig. 5 The same region as figure 4, showing the acoustieolateral and posterior
tectal commissures, and tlie median extensions of cells from the acousticolateral
areas i n greater detail. Camera lucida. X 51.

coniniissure and the cells extending medially along its course

constitute the first intimation of a cerebellar structure in
the vertebrate phylogenetic series. The fibers undoubtedly
arise from cells of the acousticolateral lobe and tlie medial
extension of cells above described.
There is no real cerebellar tissue in Bdellostoma, a s Cone1
states, but the medial extensions of small cells of the acousti-
colateral lobes represent the beginning of its cellular elenients.
As these cells a r e drawl farther medially along the fibrous
bridge they form the basic cellular layer of the ccrebellum
 CEREBELLUM O F CYCLOSTOMES 405

into which larger cells later migrate or develop among the

small cells. As illustrated in figures 4 and 5, a large cell is
present on each side in the cellular layer. Only 1 pair of such
cells was found in the entire region, both being shown in the
single section illustrated in the figures. The main dendrites
as well as the axons a r e directed lateralward. These cells proh-
ably represent reticular elements rather than Purkinje cells.

B
Fig. 6 Sagittal section through lower caudal region of midbrain of Rdellostoma,
showing acousticolateral and posterior tectal commissures. Method of Ramdii y
Cajal. X 260.
A. Section through midsagittal plane.
B. Section through same series lateral to A.

According to Ayers and Worthington ( '08) the utricular

division of the V I I I t h nerve has a large general cutaneous com-
ponent of fibers that innervate the lining membrane of the
ear, in addition to fibers that pass to the serisorv epithelium.
The Vth and V I I t h nerves also have 2 or more sets of fnnc-
tionally distinct fibers, according to these authors. The acous-
ticolateral area, into which VIIth and VIIIth fibers enter
directly, thus may receive not only lateral line and vestibular
fibers through these nerves but also cutaneous fibers. The
acousticolateral area also has numerous connections with the
general cutaneous nuclei by individual cells and in other maps,
according t o these investigators. It is, therefore, a center
for coordination of lateral line, vestibular, cutaneous and pos-
406 0. LARSELL

sibly other impulses in the hagfish. The commissural fibers,

connecting the 2 sides, accordingly must carry impulses co-
ordinated from these several primary types of stimuli from
one side of the brain to the other.

PETROMYZONTS

The relation of the acousticolateral lobes to the adult

cerebellum in Lampetra has been described by Johnston ( '02) ,
while Pearson ('36) has provided a description of the aeousti-
colateral lobes and the connections of the cerebellum in sev-
eral species of lampreys. In the present account some of the
principal stages of the development of these regions in am-
mocoete larvae of Entosphenus tridentatus, and the cerebellum
of the adult Pacific lamprey will be described, together with a
number of details gleaned from a restudy of the Gol,'01 ma-
terial of Lampetra wilderi and of pyridine-silver and toluidine
blue series of Entosphenus appendix, Petromyzon marinus
and 2 species of Ichthyomyzon.
The acousticolateral lobes and cerebellum of adult Ento-
sphenus tridentatus a r e shown in figure 7, together with their
relations to the other divisions of the brain and the cranial
nerves visible from the dorsal view. The fourth ventricle is
much broader rostrally than in Petromyzon marinus, as il-
lustrated in figure 135 of Kappers, Huber, and Crosby ( '36).
Figure 753 in Parker and Haswell (1897), representing the
brain of the same species, has greater resemblance to Ento-
sphenns, but in both figures the Petromyzon brain is repre-
sented a s more elongated than i n Entosphenus. The cerebellar
region of the latter forms a nearly transverse band over the
rostra1 p a r t of the ventricle, dorsal and slightly caudal to the
entrance of the Vth nerve into the medulla oblongata. The
I V t h nerves emerge, i n part, from the dorsolateral surface
of the medial extension of the acousticolateral lobes which will
be designated cerebellum in this description. This apparently
led Clark ('06) to call the IVth nerve the cerebellar root of
the trigerninus. The relations of the IVth nerve to the cere-
 CEREBELLUM O F CYCLOSTOMES 407

bellar region in the lampreys have been very puzzling, but a

study of the Entosphenus ammocoete stages has made clear
the reasons not only for the apparent attachment to the
cerebellum but also for the dorsal position of this motor nerve.
This aspect of the present investigation is presented in a
separat.e paper (Larsell, ’47) t o which frequent reference will
be made.

Fig. 7 Dorsal view of brain of Entosphenus tridentatus (Gsirdner). X 7.5.

The acousticoZateraE lobe

Johnston ( ’02) designated the part of the medulla obloiigata
in which the lateral liiie and vestibular roots are distributed
as the “lobus acusticus,” while Kappers in various publica-
tions has called it the area statica. Herrick and others have
used the term acousticolateral area. It appears to me that the
408 0. L A R S E LL

iiaine acousticolateral lobe is preferable since the brain part

represented is a 3 dimensional structure of considerable size.
The principal steps in its development, especially with relation
to formation of the cerebellum and related structures, will
be presented from selected stages of the ammocoete series
above listed.
The youngest Entosphenns embryos available, having a total
length of 7 mm and still possessing considerable yolk, show a
neural tube (fig. 8). The mantle layer is composed of closely

Fig. 8 Transverse seetioils through brain of larval Entosphenus tridentatus of

7 mm total length. The right side of the figures is more rostral than the left side.
Iron-hematoxylin stain. X 205.
A. Section at rostralmost level of medulla oblongata.
B. Sectiol~a t le\el of rostral p a r t of otoeyst, on left side.

crowded nuclei among which a few early iieuroblasts are

recognizable. A thin inargiiial layer is present. Roof plate and
floor plate have the appearance typical of the early neural tube
stage in other vertebrates. In the acousticolateral region,
which a t this stage is very short rostrocaudally, a few VIIIth
root fibers a r e present and fibers froin the VIIth and Xth
lateral line nerves may be followed for a short distance. In
the mantle layer there is a slight lateral swelling of the alar
plate. A few of the crowded iieuroblasts already have sliort
processes extendin? vciitrolateraily, while in the dorsomedial
 CEREBELLUM O F CYCLOSTOMES 409

region some show short processes extending dorsomedially

toward the roof plate.
I n the next stage available, an ammocoete of 42-mm total
length, the acousticolateral lobe forms a broad area at the level
of entrance of the VIIIth and anterior lateral line roots (fig.
9). The 3 nuclei, designated by Johnston ( '02) as dorsal nu-
cleus or lateral line lobe, dorsomedian nucleus, and ventro-
lateral nucleus, but which I shall call dorsal, medial and ventral
nuclei, respectively, following the usage of Kappers ( '20)

Fig. 9 Transverse sections through brain of Entosphenus ammocoete of 42 mm

total length. Iron-hematoxylin stain. X 97.
A. Section a t level of acousticolateral commissure.
B. Section at level of entrance of lateral line V I I t h and V I I I t h nerve roots.

and Pearson ('36), are present in an early stage of differen-

tiation. The root fibers of the above named nerves enter the
nuclei, as illustrated in the left side of figure 9B. The fourth
ventricle extends laterally above the acousticolateral lobe,
which is thick at this level, as shown in the figure, but rostrally
the medial wall of the acousticolat4ral lobe rapidly assumes a
nearly vertical position again, as shown in figure 9A. As it
continues forward the lobe becomes progressively narrower
until just caudal to the isthmus it has the appearance shown
410 0. LAGSELL

in figure 9A, right side. The fourth ventricle a t this level has
again assumed the form of a vertical slit with but little widen-
ing. As compared with the adult brain the acousticolateral
lobes in larval stages are greatly elongated. Their tips ap-
proach each other and are connected together by a commissure
composed of a compact group of fibers from the dorsal lateral
line VIIth tract and a more scattered bundle from the ventral
lateral line VIIth tract. Vestibular and lateral line Xtli root

Fig. 10 Transverse sections through braill of Entosphenus ammocoete of 55

mm total length. Borax-carmine and Lyons blue stain. X 102.5.
A. Seetion at level of acousticolateral commissure.
B. Section a t level of entrance of lateral line V I I t h and V I I I t h nerre roots.

fibers do not appear to reach the commissure at this stage but

a few fibers from the medial nucleus may do so. A small bulbo-
and spino-acousticolateral tract is present ventral to the
descending Vth root fibers. It reaches the rostra1 part of the
acousticolateral lobe but there is no sign of a bulbo- or spino-
cerebellar component of tlie cerebellar commissural mass at
this stage.
According t o Kappers, Huber and Crosby ('36) the decus-
sating bundles in adult petromyzonts include the lateral line
 CEIIEBELLUM O F CYCLOSTOMES 411

Xth tract. Johnston ('03, p. 19) thought the fibers of this tract
end in the medial nucleus without running forward as far as
the V I I I t h fibers. Pearson ( '36, p. 216) states that a few fibers
appear to reach the cerebellum. I n the adult material at my
disposal lateral line Xth fibers a r e difficult to follow in the
rostral p a r t of the acousticolateral lobe. F o r some distance
rostral t o its entrance the posterior lateral line tract forms
a compact buiidle of large myelinated fibers which can easily
be seen. The bundle breaks up, however, in its course along the

Fig. 11 Horizontal section through rostral p a r t of medulla oblongata of Ento-

sphenus ammocoete 64 mm long. Iron-liematoxylin stain. X 102.5.

medial nucleus so that rostrally the fibers become less nu-

merous and more scattered. Some of the ammocoete stages,
however, show that this tract reaches the cerebellar region and
decussates.
The relations of the commissural components from the an-
terior and posterior lateral line root fibers, together with addi-
tions from the dorsal nucleus and from the medial nucleus, a r e
shown in figure 11, prepared from a horizontal series of a
64-mm ammocoete. It is evident from the figure that the
medulla oblongata is greatly elongated between the region of
the V I I I t h and lateral line roots, on the one hand, and the
commissural region, on the other. J u s t rostral to the level of
412 0. LARSELL

the VIIIth and lateral line VIIth roots the ventricle has
begun to broaden between the lobes but it tapers rostrally and
caudally from this point, as shown in the figure. The section
represented was so cut that the right (lower) side of the figure
is slightly ventral to the left side. It shows the 3 chief com-
ponents of the commissure a t this stage, namely, the dorsal
lateral line VIIth bundle, the ventral lateral line VIIth bundle,
and the lateral line Xth bundle. Fibers arising from the medial
nucleus accompany the root fibers. The rostral end of the

Fig. 1 2 Transverse sections through brain of Entosphenus ammocoete 120 mm

long. Protargol stain. X 82.
A. Section a t level of acoustieolateral comrnissure.
B. Section slightly rostral to entrance of lateral line V I I t h and V I I I t h roots.

dorsal nucleus is the starting point for a compact bundle

which also reaches the commissure where it intermingles with
the dorsal tract fibers. I n the 42-mm ammocoete, a s in later
stages, the dorsoventral level of the commissure is the same as
that of the 3 main lateral line tracts which form it.
The rostral part of the medulla oblongata forms a deep
and narrow trough which is closed above a t its rostral tip
by the commissure just described and caudally by the conver-
gence of the lateral walls at the calamus scriptorius. I n the
 CEREBELLUM O F CYC!LOSTOMES 413

region of entrance of the lateral line VIIth and the VIIIth

roots the upper part of the ventricle extends laterally over
the dorsal surface of the acousticolateral lobe (fig. SB), but
farther rostrally its walls gradually assume a nearly dorso-
ventral position (fig. SA). By the 55-mm stage (fig. 10) the
ventricle has widened somewhat rostrally but the breadth be-
tween the taeniae at the level of entrance of the VIIth and
VIIItli roots has become relatively less than in the 42-mm
stage due to the growth of the acousticolateral lobes, as
described below. I n the 120-mm ammocoete the taenial mar-
gins of the acousticolateral lobes have tilted mediallp so that
the dorsal part of the ventricle has narrowed, but it is wide
between the ventralmost part of the acousticolatcral lobes arid
the underlying motor column at the VIIth and VIIIth root
levels, tapering rostrally from this region until at the level
of the commissure it is again a narrow canal whose lateral
walls a r e almost vertical save that dorsally they have a slight
medial inclination (fig. 12). The decussation of fibers across
the roof of the ventricle is still very short from side to side.

Thc cerehellovelcrr plate

The chorioid roof of the fourth ventricle lies well above
the dorsal margins of the medulla oblongata, to which it is at-
tached on each side, and it continues rostrally into the cho-
rioidea of the midbrain region (fig. 14). I n the isthmus region,
just caudal to the fovea isthmi but above it, the chorioidea
loops downward (fig. 13A). The lower margin of the caudal
p a r t of the chorioidal loop is attached to a thickened portion,
with which the aconsticolateral lobes are continuous on either
side, which will be termed the cerebellovelar plate. The rostra1
part of the loop is attached to the margin of the optocoele
(figs. 13R and 14). The cerebellovelar plate develops into the
medial, fibrous portion of the cerebellum and into the rudi-
mentary anterior medullary velum, but these parts do not
become sharply differentiated from each other even in adult
petromyzonts, as the further description will show.
414 0. LARSELL

The developing optic lobe and the cerebellovelar plate, as seen

in sagittal section, form a V-shaped mass with commissural
fibers in its deepest p a r t and with the acousticolateral com-
niissure in the dorsalmost p a r t where it continues, laterally
arid caudally, with the acousticolateral lobes. The rostra1 por-
tion of the V becomes the roof of tlie caudal part of the mid-
brain. As shown in figure 13A, there is proliferation of cells

r-
f. 6th
Mcer

-C

Fig. 13 A. Midsngittal section through ccrebellovelar plate and midbrain of

Entosphenus ammoeoete 65 mm long. Borax-carmine and Lyons blue stain. X 164.
B. Sagittal section, slightly t o one side of midplane, showing cerebellovelar
plate and midbrain of Entosphcnus aminocoete 127 nim long. Protargol stain.
X 104.

above the ependymal layer in the ventral part of both seg-

ments, thus thickening them. Less proliferation of cells at the
apex of the V apparently makes this the region of folding,
as the chorioidea is raised upward with growth of the brain,
so that the loop of tela pulls on the segments. Evidently the
position of tlie fold is partly determined also by the commis-
sure-like bundles of fibers, including the posterior tectal com-
 CEREBELLUM O F CYCLOSTOMES 41.5

missure arid IVth root fibers (Larsell, '47), which are present
a t this point.. The commissure between the acousticolateral
lobes remains short from side to side.
Apparently because of the increasingly high walls of the
ventricle on each side the cerebellovelar plate becomes strongly
tilted upward from the isthmus region as growth continues
(figs. 13 and 14). I n the 65-mm and 127-mm ammocoetes this
is well shown in figure 13, the rostra1 part of the plate having
a much more ventral position than the caudal. This condition
prevails until the ventricle broadens rostrally and the acousti-
colateral lobes are spread farther. apart, their dorsal margins

Fig. 14 Photomicrograph of sagittal section through brain of Entosphenus

ammocoete 1 2 7 mm long. Protargol stain.

assuming as a result the relatively more ventral position char-

acterisitic of the adult (figs. 7 and 15-18). At the 127-mm
stage the IVth nerve decussatioii is well shown in the velar
part of the cerebellovelar plate (fig. 13B) and IVth nucleus
cells are found well dorsally near the acousticolateral fibers
which cross from side to side. The ventral part of the plate
is swollen in the midplane, while the dorsal part is relatively
thin arid contains only commissural fibers. I n the adult,
save for increased thickness due to the more numerous fibers
and greater volume because of enlarged myelin sheaths, ap-
proximately the same relative thickness and relations of the
2 parts of the plate prevails (fig. 27). This figure, prepared
from the midsagittal section of an adult Entosphenus brain
416 0. LARSELL

fixed in situ in Bouin's fluid and subsequently embedded in

situ by the celloidin method, shows the normal relations of
tlie parts of the cerebellovelar plate much more closely than
figure 26, prepared from the midsagittal section of a pyridine-
silver series of Petromyzon. The dorsal part belongs to the
cerebellum but the ventral part is anterior medullary velum,
despite the fact that it tilts sharply upward and is bounded
from the midbrain by a deep sulcus. The sharp dorsal tilt
is no doubt due t o the dorsal anchorage afforded during de-
velopment by the commissural fibers which pass between the
rostral ends of the high lateral walls of the acousticolateral
lobes and by the tela chorioidea, a s shown in figures 13 and 14.
The ventrolateral margins of the plate, on the other hand, are
anchored to the mesencephalic and bulbar parts of the con-
stricted isthmus. The IVtli nerve and its nucleus should riot
be regarded as lying in the cerebellum but in the velar part
of the cerebellar plate. The dorsocaudal part of the plate
contains merely a mass of commissural fibers. In adult lam-
preys, a s well as in the larvae, it must be regarded as more
closely related to the acousticolateral lobes than it is to defini-
tive cerebellum.
As the midbrain increases in size it crowds backward
against the rostral end of the medulla oblongata. This me-
chanical factor, together with continued widening of the ros-
tral part of the fourth ventricle, produces a medial bending
of the rostral portion of the acousticolateral lobe from each
side which gradually changes the form of this region from an
elongated trough with high lateral walls, in ammocoete stages
(figs. 9-14), to the adult condition (figs. 7 and 15-19) in which
the rostral ends of the acousticolateral lobes continue from
either side almost t o the midplane and form an arch over the
rostral end of the fourth ventricle. The medial part of the
arch is entirely fibrous save for cells of the IVth nucleus, some
of which have reached this part of the cerebellovelar plate by
migration (Larsell, '47). Developmentally, therefore, the
petromyzont cerebellum is a continuation, deflected medially,
of the rostral ends of the acousticolateral lobes. It differs,
 CEREBELLUM O F CYCLOSTOMES 417

however, from the main p a r t of these lobes in including only

the medial nucleus of each lobe, in some of its fiber connections,
and in a slight modification of some of its cells. The fiber
conne,ctions, nuclei and cells a r e described below. The close
relationship of the cerebellum to the acousticolateral lobe in
petromyzonts, pointed out by many authors, thus is further
emphasized by its developmental history.

T h e acousticolateral commissure
The commissural connection between the rostra1 ends of the
acousticolateral lobes in larval stages of Entosphenus consists
at first entirely of lateral line root fibers, to which VIIIth
root fibers a r e soon added. Therefore it seems desirable to
designate it the acousticolateral commissure rather than to
employ the more general term cerebellar commissure, as used

Fig. 15 Photomicrograph of one side of medulla oblongata of adult Entosphenus

tridentatus at level of entrance of lateral line V I I t h and V I I I t h roots. Iroii-
lienixtoxylin stain. Transverse series.
418 0. LARSELL

by Pearson ('36) in petromyzonts, or lateral comniissure a s

it was called by Herrick ( '14) and myself ('20, '31, '32) in
urodeles, although it is essentially the same as the lateral
commissure of amphibians. I n ammocoetes decussating fibers
are recognizable in the 42-inm stage, as already stated, which
cross through the dorsal part of the cerehellovelar plate (fiy.
9A) as continuations of the bundles of the lateral line roots
of the VIIth and Xth nerves. Vestibular root fibers become
scattered among them, especially in the ventral lateral line
VIIth and the lateral line Xth part of the commissure. They
terminate in the gray layer, as Johnston ( '02) found, although
Tretjakoff ('09) described thein a s ending in the molecular
layer. The fibers of the dorsal lateral line VIIth tract are
coarser than the majority of those of the ventral tract arid
form a compact bundle, in the commissural niass, located
fartliest dorsally and caudally in the cerebellovelar plate (figs.
CIA, IOA, 12A, 13 and 14). The ventral lateral line VIIth and
the posterior lateral line portions of the commissure are more
diffuse. I n the adult a numbei* of coarse ventral lateral line
VTIth fibers are scattered through the medial nucleus, some
of them ieeaching the cerebellum.
The nerve root-fiber elements of the commissure are aug-
iriented by fibers arising from the dorsal and medial acousti-
colatcral nuclei (fig. 11) as already stated. In earlier stages
such fibers are not iiunierous but by the 120-mm stage they
constitute a considerable element of the commissure. Pearson
('36), in adult lampreys, groups with his bulbo-cerebellar
system the fibers arising in the acousticolateral nuclei, which
were described also by Tretjakoff ( '09) and Kappers ( '20).
Because they undoubtedly are closely related, functionally, to
the VIIIth and lateral live root fiber components of the acous-
ticolateral commissure I shall include them in this commis-
bure, reserving the tern1 bulbo-cerebellar for fibers arising in
the sensory Vth and other nuclei, a s described below.
I n adult Entosplienus trideiitatus and Lampetra, especially,
Cajal and Golgi series show fibers from cells in the medial tip
of the cerebellar gray which give rise to comniissural fibers
 CEREBELLUM O F CYCLOSTOMES 419

(fig. 22). Pyridine silver series of adult Petromyzon marinus

and Entosphenus appendix show a bundle of relatively small
fibers issuing from the rostral tip of the dorsal nucleus and
continuing forward into the commissure. Such a bundle is
shown also in the 64-mm ammocoete (fig. 11). These may rep-
resent a gathering together of diffuse fibers passing through
the dorsal nucleus but the indications a r e that most of them

Fig. 16 Photomicrograph of one side of medulla oblongata of adult Entosphenus

rostral to entrance of lateral line V I I t h and V I I I t h roots, showing acousticolateral
lobe. Iron-hematoxylin stain. Transverse series.

arise within it. Another component of the commissural mass

crossing the midplane arises a s a small but compact bundle
of fine fibers from the periventricular gray ventral to the
cerebellar p a r t of the medial acousticolateral nucleus. It
appears to correspond to the ventral element of the acoustico-
lateral commissure in Bdellostoma. The large ventral lateral
line VITth fibers found in the adult cerebellum (fig. 22) do not
decussate save as they subdivide into finer terminal branches.
420 0. LARSELL

Johnston ('02) divides the commissural system in adult

Lampetra into a thick dorsal and a thin ventral commissure.
Pearson ('36) includes 4 components in his cerebellar com-
missure, namely: (1)complex primary lateral line and VIIIth
root fibers, together with some bulbo-cerebellar fibers that
enter the cerebellum by way of the cerebellar crests; (2)
fibers of the bulbo- and spino-cerebellar tracts which enter in
front of the cerebellar crests; (3) fibers that establish rela-
tions between the optic tectum and the cerebellum; and (4)
I V t h root fibers. H e states that complete decussations of
these systems do not occur, with the exception of the motor
root of the I V t h nerve. Since the I V th roots and IVth nucleus
belong to the velar part of the cerebellovelar plate (Larsell,
'47), the IVth roots and their decussation, which also is but
partial, may be removed from the cerebellar commissural
complex although more or less intermingled with it because of
incomplete differentiation of velum and cerebellum.
The more dorsal part of the remaining portion of the com-
missural complex I sliall designate the acousticolateral com-
missure, as already described, and the more ventrorostral
p a r t as the bulbo- and spino-cerebellar commissure. The fibers
between midbrain and cerebellum, as described by Pearson
('36), save the tecto-cerebellar tract, appears to me to be
related to the velar p a r t of the cerebellovelar plate arid to
belong, so far as they decussate, to the velar commissure
rather than to the cominissural complex of the cerebellum.

T h e acoztsticolutcral nuclei and the cerebellum

The nuclei of the acousticolateral lobes have begun to appear
in the 42-mm ammocoete (fig. 9 ) as lateral extensions of cells
from the periventricular gray, no doubt drawn by the neuro-
biotactic influence of the lateral line and vestibular roots as
Pearson ('36) suggests. A t this stage the cells are scattered
and not numerous, without sharp demarcation between the 3
more definitive nuclei of later ammocoete stages arid adult
lampreys. I n the 55-mm ammocoete the cells have increased
 CEREBELLUM O F CYCLOSTOMES 421

in number and a rough grouping into dorsal, medial and ven-

tral nuclei is apparent (fig. 10B) in tlie region of entrance of
tlie lateral line VIIth and the V I I I t h roots. Only the medial
nucleus, however, extends to the rostra1 part of the acousti-
colateral lobe from which the cerebellum develops (figs. 10A,
11, 12A). The acousticolateral lobe, in the 42-mm ammocoete,
is somewhat flattened dorsorentrallp in the region where the

Fig. 1 T Photomicrograph of medulla oblougata and caudal p a r t of cerebellum

of adult Entosphenus. Iron-liematoxylin stain. Trausverse series.

lateral line V I I t h and V I I I t h roots enter. It is relatively thick

between the fourth ventricle and the external surface. The
attachment of the tela chorioidea is lateral (fig. 9 ) . Rostrally
the lobe tapers toward tlie acousticolateral conimissural re-
gion, a s map be seen by comparison of the left and right sides
of figure 9A, the right side of which represents the more ros-
tral p a r t of the section whieli was cut somewhat obliquely
422 0. LARSELL

t o the transverse plane. The tapering also is shown in figure

11. As the nuclear masses increase in size by continued addi-
tions of cells and Iierve fibers, the transverse axis of the lobe
rotates upward and medially in such fashion that the taenia
is carried from the lateral position it occupies in the 42-mm
larva to a dorsal position by the 55-mm stage (fig. 10B) and
to a dorsomedial position in the 120-mm arnmocoete (fig. 12R).

Fig. 18 Photomicrograph of rostra1 part of cerebellum and medulla oblongata

in adult Entosphenus. Iron-hematoxylin stain. Transverse series.

As seen in transverse sections the lobe increases greatly in its

ciorsoventral nieasurements (dorsomedial to ventrolatera1 in
the 120-mm stage) but becomes relatively thinner from ven-
tricular surface to external surface. In the last named stage
the acousticolateral area s11ows distinct dorsal, medial and
ventral nuclei, with the fiber coiinections of each essentially
as described by Pearson ( '36) in the adult petromyzont. The
 CEREBELIJUM O F CYCLOSTOMES 423

fourth ventricle extends laterally under the aconsticolateral

lobe which has tilted medially, as described, a t the VIItli and
VTIIth root levels. Fa r t h e r rostrally the ventricular surface
gradually assumes a dorsoventral direction again (fig. l 2 A ) .
Tlie medial nucleus extends to the rostral tip of the lobe on
each side, but as the tips turn medially to join the cerebello-
velar plate the medial nuclei of the 2 sides do not fuse.
The dorsal nucleus continues relatively mnch farther ros-
trally in adult Entospherius tridentatus than in ammocoete
stages but i t takes no p a r t in formation of the cerebellum
save through tlie fibers that pass from it to the commissure
aiid the rudimentary molecular layer, described below. The
ventral nucleus, a s described by Pearson ( '36), remains con-
fined to the region of entrance of the VIIIth roots, not extend-
ing f a r beyond the most rostral of the entering fibers.

Tlie cerebellar gray substmice

Johnston ( '02) described the cellular part of the cerebellum
a s composed of small cells similar to those in his dorsomedial
nucleus, i.e., the medial nucleus of the present description, but
with larger cells intermingled. He refers to it as the granular
layer. Because of the presence of IVth nucleus cells in and
near the midplane, more coiispicuous in the adult than i i i
larval stages, there appears to be continuity of the gray
layer of the cerebellum from 1 side to tlie other. The cells of
the cerebellar gray proper do not reach the midplane in the
ammocoetes, as pointed out above, o r in adult lampreys,
as Pearson ('36, p. 214) has stated, but the medial tips of the
gray masses in each lateral half of the organ are joined b)-
connnissural fibers (figs. 17, 22, 24). Pearson also describes
the cerebellar gray as not continuous with tlie gray of the
midbrain. This is certainly true laterally, as his figure 4
well sliows, but more medially there is a n upward extension of
the tegmental gray into the base of the cerebellar region, while
still more medially there is continuity of cells in the deep part
of the cerebellovelar plate with the gray of the midbrain (fig.
424 0. LARSELL

13B). This also is shown in figure 351 of Kappers, Huller and

Crosby ( '36), representing adult Petromyzon. In the iron-
hematoxylin series of adult Entosphenus tridentatus there is
something of a hiatus between the deeper mass of gray, which
I regard as pertaining to the velum, and the more dorsally
situated mass of cerebellar gray proper, with only scattered
cells between the 2 masses. Pearson describes a fusion of the

Fig. 19 Photomicrograph tliroiigh anterior mrdullary reliim and medulla ob-

longata a t level of entrance of the trigeminal roots in adult Entosphenus. Iron-
hrmatoxylin stain. Transverse series.

cephalic portion of the niedial nucleus of the acousticolateral

area and the correspoiidiiig periventricular gray, the combined
mass continuing into the cerebellum as his deep cerebellar
gray. I can corroborate this description but see little need for
dividing the cerebellar gray into superficial and deep portions
since the 2 a r e continuous farther medially, while the separa-
tion seen in many of the sections is due to fiber bundles rather
than to a continuous sheet of fibers.
 CEREBELLUM O F CYCLOSTOMES 425

The gray substance of the cerebellum proper, therefore, is

continuous laterocaudally with the medial nuclei of the acous-
ticolateral lobes and represents the medially deflected rostra1
portion of these nuclei, as its development makes clear, to-
gether with additions from the periventricular grisea. While
the cerebellar part of the medial nucleus undoubtedly is the
precursor of the granular layer of the cerebellum of higher
vertebrates, in petromyzonts it is more closely related to the
acousticolateral lobe, functionally and structurally, than to the
definitive granular layer. Some functional modification has
been brought about by the entrance into the cerebellum of the
lobo-cerebellar and tecto-cerebellar tracts described below,
as well a s by the absence in the cerebellum of the cellular inter-
connections between the medial nucleus with the dorsal and
ventral nuclei which are characteristic of the aconsticolateral
lobe proper. However, lateral line and vestibular fibers con-
stitute its chief afferent supply, supplemented by spino- and
bulbo-cerebellar fibers, much as in the acousticolateral lobe.

T h e cell t y p e s in acousticolateral lobe and

cerebellum
By the 120-mm stage of the larvae many of the cells of the
grisea have enlarged and show dendritic processes a s well a s
axons. Johnston ('02) has described the cells of the acousti-
colateral lobe and the cerebellum from Golgi series of adult
Lampetra which I have restudied rather exhaustively. I can
confirm his conclusions and add some details of observation.
In the acousticolateral lobe are found cells of several sizes.
Many, if not all of the small cells in the dorsal and medial
nuclei send their axons into the cerebellar crest (figs. 20 and
21). Their dendrites spread out within the nuclei, as shown
in the figures. Entirely similar cells are found in the gray
substance of the cerebellum (figs. 22-24). There can be no
doubt that they are primitive granule cells. Their axons pass
into the superficial fibrous layer to augment similar fibers
from the cerebellar crest. Other medial nucleus cells of inter-
426 0. LARSELL

mediate size give off dendrites both into the cerebellar crest
and into the dorsal nucleus (fig. 20A) a t levels caudal to the
cerebellum. Still others appear to have their dendrites spread
wholly within the medial or ventral nuclei. All give rise
to arcuate fibers which for the most part pass ventromedially.

Fig. 20 Transverse sections through acousticolateral lobe of Lampetra wilderi.

Golgi method.
A. Section through acousticolateral lobe a t level of entrance of lateral line
V I I t h and V I I I roots, showing txpes of cells in the dorsal, medial and ventral
nuclei. X 52.
B. Section at same level on opposite side of brain. X 82.

Still larger cells, with bodies nearer tlic ventricular surface,

give off dendrites to 2 or all 3 of the nuclei (fig. 2 0 s ) .
Within the cerebellum a r e found large cells with wide-
spreading dendrites, some of which branch among the com-
missural fibers (figs. 22-25), others among the granules and
many reach the cerebellar continuation of the cerebellar crest,
 CEREBELLUM O F CYCLOSTOMES 427

as described below. Many of the dendritic processes have

types of telodendrites very similar to those shown by medium
and large cells in the acousticolateral lobe. As shown in
figures 20 and 22, the dendritic tips break up into small twigs,
b,oth in the acousticolateral lobe and in the cerebellum, not
only in the crista cerebellaris and its continuation as tlie
rudimentary molecular layer of the cerebellum but also in the
cellular layer of the latter and the dorsal and medial nuclei
of the acousticolateral lobe. I n addition, many of the dendrites
in the cerebellum have short processes a t intervals before tlie
terminal tuft is reached (fig. 22). Thehe cells undoubtedly are
precursors of Purkinje cells, a s Johnston ( '02) regarded them,
but in the lampreys they are even more closely related to the
larger cells of the acousticolateral lobe.

The cerebellar crest and the molecular layer

The cerebellar crest in petromyzonts (figs. 9, 12, 15, 16, 20,
21) is a relatively thin zone of fibers extending from a level
caudal to the entrance of the lateral line and vestibular roots
into the cerebellum, a s described by Pearson ( '36). While it
contains fibers of passage of lateral line, vestibular and other
systems, it should be regarded primarily a s a zone of neuropil.
I n the younger ammocoetes a small zone devoid of stained
fibers, lying dorsal to the entering lateral line and VIIIth
root fibers and lateral to the ascending lateral line tracts,
represents its location (fig. 9). By the 120-mm stage the area
is well marked off from the acousticolateral nuclei, medially,
but only a few fine fibers are stained in the most favorable
preparations. Rostrally, as seen in transverse and sagittal
sections of ammocoete stages (figs. 9,12 and 13), the cerebellar
crest is much reduced in area, being absent in the cerebellar
midplane region in the 65-mm ammocoete (fig. 1 3 A ) . As
the cerebellum develops and increases in size in successive
stages the rostral part of the cerebellar crest grows forward,
eventually forming a layer in the dorsal part of the rostral
face of the cerebellum which crosses the midplane in later
428 0. LAliSELL

Fig. 2 1 Section :it level of entranee of V I I I t h root showing small cells in dorsal
nucleus whose axons pass into the cerebellar erest. Lampetra. Transverse section.
Golgi method. X 102.5.

Fig. 2 2 Section of cerebrlluni showing cell types. Lanipetra. Golgi method.

x 102.5.
 CEREBELLUM OF CYCLOSTOMES 429

stage ammocoetes and the adult (figs. 1 6 2 3 and 25-27) and

forms what is usually called the molecular layer of the cere-
bellum. From the phylogenetic viewpoint the term cerebellar
crest is misleading, implying as it does a continuation of
cerebellum onto the medulla oblongata. Since it is a part of
the acousticolateral lobe before the cerebellum has formed
another name would be preferable. Pearson’s term “stratum
fibrosum et mo!eculare” is better because it is a t least descrip-
tive of structure. However, “cerebellar crest” is so embedded
in the literature that I shall continue to use it.
Golgi series of adult Lampetra show the bulbar part of the
cereb,ellar crest to be composed largely of fine fibers given off
chiefly by the small cells of the dorsal and medial nuclei (figs.
20, 2 l ) , the fibers from the dorsal nucleus preponderating in
the material examined. Larger fibers of lateral line origin
also a r e found. Dendritic processes from larger cells in both
dorsal and medial nuclei extend into the neuropil formed by
the fine fibers. The tips of many of the dendrites break up
into short, knobbed terminal tufts (fig. 20A). S o f a r as ob-
served few, if any, cells of the ventral nucleus give off den-
drites to the cerebellar crest although dendritic processes
which pass into the neighboring medial nucleus end among the
fibers in the latter. Many of these fibers arise from small cells
similar to those which give rise to cerebellar crest fibers (figs.
20 and 21). The axoiis of the larger cells become iiicluclecl
in the arcuate fiber system. Some of the widespreading
dendrites of the relatively large cells in the cerebellum, above
described, terminate in the cerebellar continuation of the
cerebellar crest, while others end in the cerebellar part of the
medial nucleus (figs. 22-24).
The cerebellar continuation of the crista cerebellaris forms
the molecular layer of the cerebellum, as all writers on the
snbject a r e agreed. While there is much decussation of the
fine fibers in the molecular layer, it should be regarded as a
zone of neuropil, rather than a commissure. Because of the
entrance of fiber tracts that a r e not represented in the acous-
ticolateral lobes, into both layers of the cerebellum, as de-
430 0. LARSELL

scribed below, the molecular laper receives a somewhat dif-

ferent complex of stimuli tliaii does the cerebellar crest.
These fiber tracts are the lobo-cerebellar tract and the tecto-
cerebellar tract. To them, as differentiating factors, should
be added a larger, or a t least a more coilcentrated represeiita-
tioii of trigeminal and spinal fillers, as compared with the

Fig. 23 Sagittal sections showing cell types in cerebellum. Lampetra. Golgi

method. X 102.5.

Fig. 24 Horizontal section showing primitive Purkinje cells, a primitive granule

cell and fibers of the V I I l t h root in the ncousticolnteral commissure. Lampetra.
Golgi method. X 65.
 CEREBELLUM O F CYCLOSTOMES 431

acousticolateral lobes. Also the V I I Ith root fibers to the cere-

bellum should be included since they come into relation with
a different complex of other impulses than is true in the ventral
acousticolateral nucleus and other p arts of the acousticolateral
lobe which a r e reached by vestibular fibers. Therefore, while
structurally identical with the cerebellar crest, the molecular
layer has begun to differentiate functionally from it.

Afferent fiber tracts

The acousticolateral coiiiiections and commissure have been
described as a group because they form the beginning of the
cerebellum in lamprey larvae as well as in the adult hagfish.
Other connections enter in, as the amniocoetes develop, which
now will be considered.
Fine fibers from the sensory Vth root, which apparently
represent ascending rami of the main root fibers, enter the
lateral portion of the cerebellar base in the 120-mm ammocoete
and later stages but I have been unable to find any evidence of
such fibers in the cerebellar commissural complex. Tretjakoff
( '09) described trigemino-cerebellar fibers in the ammocoete
and sometimes is incorrectly cited as describing a trigeminal
commissure in the cerebellum. According to Tretjalzoff's de-
scription (p. 672), which I can corroborate in Entosphenus
larvae, a small proportion of the finer trigeminal fibers divide
into 2 branches on entering the brain, as already stated, the
rostral rami forming a small cerebellar tract. This is a com-
pact bundle for a short distance only, the fibers spreading out
between other systems, a s the cerebellum is approached, and
becoming lost among them. Some, however, can be followed
into the cerebellum. Johnston ( '02) found trigemino-cere-
bellar fibers in Lampetra and Pearsoii ('36) also describes
them. They a r e shown in adult Entosphenus tridentatus (fig.
28), the description above given for them in ammocoetes ap-
plying also to the adult in which, however, they form a larger
bundle. A series of small fascicles arise from the rostral
p a r t of the trigeminal nucleus and constitute a secondary
432 0. LARSELL

Fig. 25 Oblique longitudinal section showing cells in cerebellar gray and the
spino-cerebellar tract. Lamprtra. Golgi method. X 102.5.

Fig. 06 Midsagittal section through cerebellum and anterior medullary velum

of Prtromyzon marinus. I’gridine-silver method. X 130.
 CEREBELLUM O F CYCLOSTOMES 433

trigemino-cerebellar tract. These also appear to be entirely

homolateral although it is possible that some of them, as well
a s fibers from the direct trigeminal tract, may decussate in
the loose aggregation of fibers just caudal to the spino-cere-
bellar tract shown in figure 26.
The spilzo-cerebellar tract. In the ammocoete series the
bulbo- and spino-cerebellar tract is shown a s a small bundle
(figs. 9, 12) throughout most of the length of the medulla
oblongata lying ventral to the spinal Vth tract in the position
occupied by the spino-cerebellar and spino-tectal tracts of
adult lampreys. Fibers pass from it into the acousticolateral
lobe. I n a sagittal series of the 127 mm stage, well stained
with protargol, collaterals from the spino-tectal tract pass into
the cerebellar region, as Herrick (’14) found in larval Ambly-
stoma. I n the ammocoete they are given off at right angles
to the spino-tectal fibers in the region rostra1 to the anterior
motor VIIIth tract and the sensory Vth roots and they enter
the grisea in the base of the cerebellar region. Farther caudally
similar fibers arch dorsorostrally toward the cerebellum, ac-
companied by a few fibers which are not collaterals. These
more caudal fibers *become closely related in position to the
trigemino-cerebellar fibers. They interdigitate to some extent
with Vth root bundles. Some decussate a t this staqe, a small
fascicle of about a dozen fibers being present near the midplane
of the cerebellovelar plate (fig. 13B) in the position occupied
by the spino-cerebellar tract in the adult lamprey. They may
be followed across the midsagittal plane.
Pearson ( ’36) describes a spino-acoustico-latero-cerebellar
system” as a n accumulation of fibers at the lower end of the
medulla oblongata, ventral to the descending Vth root and its
nucleus, which continues into the lateral funiculus of the spinal
cord. These fibers and his further description of this system
I can corroborate in adult lampreys. As the tract is followed
rostrally it separates into an acousticolateral component and
a cerebellar bundle. When the Vth roots are approached the
spino-cerebellar tract divides into small fascicles which curve
dorsally, interdigitating between the Vth root bundles, and
434 0. LARSELL

enter the base of the cerebellum. The relations are very simi-
lar to those found in urodeles (Larsell, '20, '31, '32). This
tract is undoubtedly the spino-cerebellar tract, corresponding
to the ventral spino-cerebellar tract of higher vertebrates. It
is accompanied by the spino-tectal tract a s f a r as the rostra1
level of the Vth roots, a t which point the spino-cerebellar tract
turns more dorsalward to enter the cerebellum, while the
spino-tectal tract curves more gradually upward into the mid-
brain (fig. 25). I n the Golgi series of adult Lampetra from
which the figure was drawn there are suggestions of collaterals
from the spino-tectal fibers passing into the spino-cerebellar
tract but the heavy impregnation of the sections made it im-
possible to follow the apparent branches very f a r with any
certainty. A spino-cerebellar tract was thought present in
Petromyzon by Clark ('06) and Addison ('29). Clark was
unable to trace it to the spinal cord and Addison did not
describe it. The spino-cerebellar fibers decussate in Petro-
myzoii, as Pearson ('36) states, and also in adult Entosphenus
tridentatus. In the Golgi series of Lampetra the decussation
is not evident but it probably is present in this genus also.
hly observations indicate, also in accord. with Pearson, that
the tract is derived from the spinal cord, with accessions from
the bulb.
The cerebellar comnissurc a s the term has been used by
Herrick ('14, and subsequent papers) and myself ('20, and
subsequent papers) is thus represented, in part, in petro-
myzonts by the decussating spino-cerebellar tracts in the lower
cerebellar portion of the cerebellovelar plate, in the midplane
region, and extending laterally and ventrocaudally through
the cereloellum from this zone t o tlie medulla oblongata. This
component of tlie commissural complex of the cerebellum must
be regarded a s distinct, functionally as well a s in its origin,
from the acousticolateral commissure above described. I n
urodeles (Herrick, '14, '24; Larsell, '20, '31, '32) the spino-
and trigemino-cerebellar decussation, as compared with the
acousticolateral commissure, is predominant. Also in the
salamanders and in higher vertebrates the spino-cerebellar
 C ER EB ELLU M O F CYCLOSTOMES 435

mes.

Fig. 27 Midsagittal section through cerebellum and anterior medullary velum

of Entosphenus tridentatus. Celloidin series, after fixation in Bouin ’s fluid. Iron-
hematoxylin stain. X 130.

Fig. 28 Transverse section showing anterior motor V I I I t h nucleus and the ad-
jacent structures in adult Entosphenus tridentatus. Iron-hematoxylin stain.
X 102.5.
436 0. LARSELL

and t rigemino-cerebellar decussations, which a r e closely re-

lated in position a s well as in probable function, constitute the
bridge along which the corpus cerebelli is formed. I n petro-
myzonts, however, tlie corpus cerebelli does not exist save as
it is foresliadowcd by the cells in tlie rostra1 part of the
medulla oblongata and ventral to the medial nucleus aniong
which terminate the spinal, bulbar and trigeminal tracts above
described, with some of tlie spinal fibers decussating.
T h e tecto-cerebellar tract (fig. 2 6 ) was described by Clark
('06) and by Pearson ('36) in adult petromyzonts. It is well
shown in protargol stained series of larval Entosplienus (fig.
13B). Many of its fibers terminate in the IVth nucleus but
others continue into cerebellar territory. F o r this reason I
have divided this fiber bundle into tecto-velar and tecto-
cerebellar tracts. The fibers appear to cross in the cerebello-
velar plate.
T h e cerebello-tecfal fibers, which Pearson describes as de-
cussating in the cerehellar commissure, appear to me to be
p a r t of the posterior tectal commissure, passing through the
velar part of the cerebellovelar plate.
T h e i)~esencephalo-cerebellartract of Pearson is easily seen
in later larval and adult stages. I t arises in the tegmental part
of the midbrain and is distributed to the region below the
cerebellum which appears to be more closely allied to reticular
formation than to cerebellum. The bundle gives the impression
of being related to the tecto-reticular tract of higher forms.
T h e lobo-cerebellar tract was described by Johnston ( '02)
in Lamptera as passing from the inferior lobes of the hypo-
thalamus to the cerebellum, p a r t of the fibers crossing in tlie
postoptic commissure. I can confirm this in ammocoete stages
as well as in adult Cajal ancl iron-hematoxylin series of Ento-
splieiius and in pyridine-silver series of Petromyzon. The
fibers a r e scattered and the cells of origin cannot be deter-
mined, the direction of conduction in tlie tract being assumed
from a degeneration esperiment 011 the corresponding tract
in fishes reported by Wallenberg ( '07). The lobo-cerebellar
tract is intermingled with the anterior fibers of the cerehello-
 CEREBELLUM O F CYCLOSTOMES 437

motor system, as described by Johnston ('02) and Pearsori

( '36). It ends in the basal p a r t of the cerebellum.

Deep nuclei in. cerebellar regiovz

T h e anterior niotor V I I I t h wucleus. At the rostra1 end of
the ascending V I I I t h tract, just dorsomedial to the entrance
of the sensory Vth root, is located a group of large cells which
Kappers ( '20, p. 640) has called the nucleus octavo-motorius
anterior. The large axons of these cells pass rostroventrally
and medially a s the anterior motor VIIIth tract. This tract
reaches the region of the I I I r d nucleus and decussates in the
ansulate commissure, but collaterals from its fibers pass to
the I I I r d nucleus in ammocoete stages (Larsell, '47). Pearson
('36) describes some of its fibers a s ending in the I I I r d nucleus
in adult lampreys. I n adult Entosphenus tridentatus large
and numerous dendritic processes from many of the cells
(fig. 28) pass dorsomedially into the cerebellum, ending among
the fibers of the acousticolateral commissure and among the
cells of the medial nucleus. Golgi sections of Lampetra show
similar dendrites (fig. 29). In these sections, and especially
in protargol sections of ammocoete stages of Entosphenus,
dendrites may be seen which extend caudally into the ascend-
ing VTIIth tract which also has its terminus in the nucleus.
The anterior motor VIIIth nucleus thus receives impulses
from both vestibular and lateral line systems, and probably
also more or less correlated impulses from the cerebellar part
of the acousticolateral lobe. As Kappers, Huber and Crosby
('36, p. '709) point out, this nucleus is subcerebellar, cor-
responding in its position to the nuclei of the vestikular and
lateral line nerves.
T h e rtucleus cerebelli. Kappers, Huber and Crosby ('36)
suggest, a s had Kappers ( ' Z O ) , that the cerebellar nuclei have
their origin in relation to the region of the anterior motor
VIIIth nucleus. With this statement I am in agreement, with
the reservation that the nucleus octavo-motorius anterior itself
not be included. Cajal sections of adult Entosphenus show
438 0. LARSELL

/ Cb.

Fig. 29 Cells of the anterior motor V I I I t h nucleus, with dendrites extending

dorsomedially into the lateral p a r t of the acousticolateral commissure. Lampetra.
Golgi method. X 130.

Fig. 30 Rostra1 p a r t of medulla oblongata showing anterior motor V I I I t h

nucleus and scattered cells of primitive nucleus cerebelli. Entosphenus tridentatus,
adult. Method of Ramdn y Cajal. Oblique horizontal series. X 205.
 CEREBELLUM O F CYCLOSTOMES 439

numerous cells of much smaller size located dorsally and

dorsomedially to the anterior motor VIIIth nucleus (fig. 30)
many of whose axons collect into a fairly compact tract that
passes ventrorostrally, as shown in the figure.
The afferent connections of the cells of the incipient nucleus
cerebelli probably a r e the trigemino-cerebellar fibers and the
bulbo- and spino-cerebellar tracts which reach this region
of the zone belonging more or less in common to the deep part
of the cerebellum and the rostral end of the medulla oblongata.
Whether o r not any axons of the primitive Purkinje cells above
described end in this region could not be ascertained. Such
cells give off axons into the internal arcuate system, a s above
stated and a s shown in figure 22, but they do not form fascicles
that can be followed in the lampreys. Other cells of the same
general group give off axons which pass ventrally or ventro-
caudally into the medulla oblongata a s scattered fibers or
small fascicles. The entire group of smaller cells in the lateral
portion of the cerebellum corresponds, in a more diffuse
manner, to the nucleus cerebelli of amphibians (Larsell, '23).
The anterior motor VIIIth nucleus appears somewhat related,
pet because of its special connections it should not be regarded
as a cerebellar nucleus but as belonging to the acousticolateral
and eye-muscle systems, plus whatever connections it makes
through the ansulate commissure.

Eferelzt fibers
The efferent fibers of the cerebellum form a fan radiating
from the base of the cerebellar region into the motor regions
of the midbrain and medulla oblongata. The more rostral fibers
a r e interniingled with lobo-cerebellar fibers, while the caudal
ones a r e continuous with similar appearing fibers from the
acousticolateral area. Nost of them, rostral, intermediate, and
caudal, no doubt a r e axons of the primitive Purkinje cells, as
shown in figure 22, and correspond to the dorsal arcuate fibers
from the related cells of the acousticolateral lobes, forming
both crossed and uncrossed connections with motor elements.
440 0. LARSELL

Those that cross, ventral to the floor of the fourth ventricle,

form a decussation that appears to cori*espoiid, in a diffuse
manner, to the “Haubeiikommissur” of the auricles of fishes,
demonstrated by degeneration methods by Wallenberg ( ’07).
This author ( ’05) previously had demonstrated a flocculo-teg-
mental conimissure in the rabbit, with which he homologized
the Hauberikommissur of fishes, and he regarded such a
ventral conimissure a s present in all vertebrate groups. Dow
(’36) has demonstrated by the hlarclii method that direct
fibers from the flocculus reach the vestibular nuclei in the rat
aiid cat. There thus is evidence that the primitive pattern of
established connections between the primordial Purkinje cells
in the vestibulolateral p a r t of thd lamprey cerebellum, which
corresponds to the auricles of fishes aiid of which the mam-
malian flocculus represents a niodification, is maintained
throughon t tlie vertebra t c series.
Some of the fibers of the radiating mass appear to arise from
smaller cells in the deep part of the cerebellar region which
corresponds to the emineiitia subcerebellaris. Rostrally these
constitute a definite cerebello-tegmental tract.
T h e ccr~beZlo-teg.ilienta1t r a c t is made up of the fibers above
described as constituting a ventrorostrally directed bundle
from the incipient nucleus cerebclli. It lies dorsal and rostra1
to the anterior motor VITItli tract for a short distance and
then fans out into the tegmentum of the midbrain so that the
term cei.ebello-niesenceplialic tract would, l)erhaps, be more
appropriate. Pearson ( ’36), however, has used this name for
another tract. While consisting chieflr of axons from. cells of
the eminentia subcerebellaris, it may also include axoiis of
primitive Purkinje cells. It undoubtedly represents the begin-
ning of the brachium conjunctivum of higher vertebrates.
SUMMARY AND CONC1,TTSIONS

1. I n Bdellostoiiia a coinniissure composed of very fine fibers

extends between the ventromeclial border of the acoustico-
lateral areas, nrching dorsal to the ventricular canal and pass-
ing through the midbrain at the margin of attachment of tlie
 CEREBELLUM O F CYCLOSTOMES 441

tela chorioidea of that, part of the ventricular canal which

corresponds to the fourth ventricle of higher vertebrates. I t is
joined by a similar bundle of fine fibers arising from a medial
extension of small cells from the acousticolateral area on
either side. The 2 fascicles become blended together as they
approach the midplane so that they form one commissural
bundle, the acousticolateral commissure, in which the 2
fascicles are not distinguishable. The commissure lies just
caudal to the posterior tectal commissure in the region of the
midplane of the mesencephalon, but is differentiated from the
latter by the lighter stain and smaller size of its fibers. Farther
laterally, on each side, it passes veiitrolaterally into the
acousticolateral area, while the posterior tectal commissure
arches dorsally and rostrolaterally in the midbrain.
2. The medial extension of cells from the acousticolateral
area is the forerunner of the gray substance of the cerebellum.
The cells are small bipolar and multipolar elements. Only 2
large cells, one on either side and symmetrically placed, were
found in the cerebellar region. They are regarded as reticular
elements rather than a s related to Purkinje cells.
3. The cerebellum of Bdellostoma, therefore, consists of a
small acousticolateral commissure accompanied by a medial
extension of cells from the acousticolateral area. The fibers
decussate but the cell masses stop short of the midsagittal
plane.
4. The acousticolateral lobes in ainmocoete stages of Ento-
sphenus form the walls of a deep and narrow trough, the
rostral end of which is the cerebellovelar plate. The acoustico-
lateral commissure, formed by lateral line VIIth, lateral line
Xtli and vestibular tracts, passes through the dorsal part of
this plate a s a short commissure at the same dorsoventral level
as the tracts of xvhich it represents a decussation. This forms
the fibrous, medial part of the cerebellum. By the adult stage
the rostral ends of the acousticolateral lobes have been bent
medially, forming the lateral and massive portion of the
cerebellum.
442 0. LARSELL

5. The medial nuclei of the acoustjcolateral lobes extend

farther rostrally, as the lobes develop, than do the dorsal or
ventral nuclei and turn medially with the bending of the rostral
ends of the acousticolateral lobes. Together with peri-
ventricular gray they form the gray substance of the cere-
bellum, but they do not fuse across the midplane. The
cerebellar gray is composed of primitive granule cells and
forerunners of Purkinje cells. The latter, however, are more
closely related to the larger cells of the acousticolateral lobes
than to definitive Purkinje cells, while the granule cells have
their counterparts in the dorsal and medial nuclei of the
acousticolateral lobes.
6. The gray layer of the cerebellum, while a continuation of
the medial nuclei of the acousticolateral lobes, differs func-
tionally from the latter because of differences in connections.
A slight modification of some of its cells from related cells in
the acousticolateral lobe appears to be an expression of the
functional divergence.
7. The cerebellar crest develops in ammocoetes a s a zone
of neuropil on the dorsolateral margin of the acousticolateral
lobe. It continues into the cerebellum as the molecular layer
of the latter. While similar structurally to the cerebellar crest
the molecular layer must differ in the types of stimuli it cor-
relates because of the differences in afferent connections be-
tween cerebellum and acousticolateral lobes above mentioned.
8. The fiber tract connections of the cerebellum are similar
to those of the acousticolateral lobes in that both regions re-
ceive lateral line, trigeminal and spinal fibers. They differ
in the important features that the cerebellar region receives
lobo-cerebellar and tecto-cerebellar fibers, a larger concentra-
tion of spinal and trigeminal fibers, and in the decussation of
the acousticolateral and spino-cerebellar tracts as the com-
niissures above named. The latter feature is probably the
most important in laying the foundation of the cerebellum as
a n organ of the whole body, rather than a unilateral part of
the nervous system. The entrance of the tracts from more
rostral parts of the brain makes it a coordinating center for
 C E R EB ELLU M O F CYCLOSTOMES 443

optic and presumably olfactory stimuli, in relation to the

cutaneous, lateral line, vestibular and probable propioceptive
stimuli i t receives from its spinal and bulbar connections.
9. The efferent fibers of the cerebellum a r e derived in part
from small cells in the zone between the lateral part of the
cerebellum and the medulla oblongata and in part from the
primitive Purkinje cells. The small cells collectively consti-
tute the forerunner of the nucleus cerebelli and give off axons
which radiate vent roros trally, ventrally and vent rocaudally,
many of the ventrorostral fibers being aggregated into a cere-
bello-tegmental tract in Entosphenus. The asons of most of
the primitive Purkinje cells appear to enter the system of
arcuate fibers but some probably end among the small cells
of the incipient nucleus cerebelli.
10. The anterior motor VIIIth nucleus is not a part of the
cerebellum, although i t sends dendrites into the acoustico-
lateral commissure and the lateral portion of the cerebellar
gray. It also receives vestibular impulses through the ascend-
ing V I I I t h tract. There are, however, no connections between
it and the cerebellar crest or the molecular layer of the cere-
bellum. It probably serves reflexes that are relatively simple
as compared with those made possible throuyh the more
numerous connections and more elaborate structure of the
cerebellum.
11. The cerebellum of petromyzonts corresponds to the
auricles of fishes and urodeles. The corpus cerebelli is fore-
shadowed by the decussation of the spino-cerebellar tract, the
incipient nucleus cerebelli and the cerebello-tegmental tract,
hut i t is buried beneath the dominant acousticolateral part of
the cerebellum.
LITERATURE CITED
ADDISON,W. H. F. 1929 The phylogeny of the afferent cerehellar pathways.
The Cerebellum. Publ. Assoc. for Rrsrarrh in Nero. and Ment. Diseases,
6: 329-333. The Willianis and Wilkins Conipany, Baltimore.
AYERS,HOWARD A., A N D .JIJLIA WORTHINCTON1908 The finer nn:itomy of the
brain of Rdellostoma domhcyi. I. The acustiro-lateral system. Am. J.
Anat., 8: 1-16.
444 0. LARSELL

CLARK,W. B. 1906 The cerebellum of Petromyzon fluviatilis. J. Anat., 4 0 :

318-325.
CONEL,J. L. 1929 The development of the brain of Bellostoma stouti. I. Ex-
ternal growth changes. J. Comp. Neur., 4 7 : 343-403.
1931 The development of the brain of Bdellostoma stouti. 11. I n -
ternal growth changes. J. Comp. Neur., 52: 365-499.
DoW, R. S. 1936 The fiber connections of the posterior parts of the cerebellum
in the r a t and cat. J. Comp. Neur., 63: 527-548.
EDINGER, L. 1906 Uber das Gehirn von Myxine glutinosa. Physik. Abhandl. der
konig. preuss. Akad. der Wissensch., Berlin. (Cited from Conel, 1931.)
HERRIPK, C. JUDSON 1914 The medulla oblongata of larval Amblystoma. J.
Comp. Neur., 6 4 : 343-427.
1924 Origin and evolution of the cerebellum. Arch. Neurol. and
Psychiat., 11: 621-652.
HOLM,J. 1901 The finer anatomy of the nervous system of Myxine glutinosa.
Morph. Jahrb., 29 : 365.
HOLMGREN, NILS 1919 Zur Anatomie des Gehirns von Myxine. Kungl. Svenska
Vetenskapakad. Handl., 60 : 1-96.
1946 On two embryos of Myxine glutinosa. Aeta Zool., 27: 2-90.
JANSEN, J. 1930 The brain of Myxine glutinosa. J. Comp. Neur., 4 9 : 359-507.
JOHNSTON, J. B. 1902 The brain of Petromyzon. J. Comp. Neur., 22: 1-86.
KAPPERS, C. U. ARIENS 1920-1921 Die vergleichende Anatomie des Nerven-
systems der Wirbeltiere nnd des Mensehen. De Erven F. Bohn, Haarlem.
KAPPERS, C. U. ARIENS, G. CARL HIJBER AND ELIZABETH C. CROSBY 1936 The
comparative anatomy of the nervous system of vertebrates, including
man. The Maemillan Company, New York.
LARSELL, 0. 1920 The cerebellum of Amblystoma. J. Comp. Neur., 31 : 259-282.
1923 The cerebellum of the frog. J. Comp. Keur., 3 6 : 89-112.
1931 The cerebellum of Triturus torosus. J. Comp. Neur., 53: 1-54.
1932 The development of the cerebellum in Smblystoma. J. Comp.
Neur., 54: 357-435.
1946 The Cerebellum of cyclostomes. Anat. Rec., 9 4 : 34.
1947 The nucleus of the I V t h nerve in petromyzonts. J. Comp.
Neur., 86 : 447-466.
MULLER,JOHANNES 1938 Die vergleichende Anatomie der Wirbeltiere. Abhandl.
der konigl. Akad. der Wissensch., Berlin. (Cited from Conel, 1931.)
PARKER, T. JEFFERY, AND WILLIAMA. HASWELL1897 A text-book of zoology,
2 : 123. The Macmillan Company, New York.
PEARSON, ANTHONYA. 1936 The aeustico-lateral centers and the cerebellum,
with fiber cohnections, of fishes. J. Comp. Neur., 65: 201-294.
RETZIUS, ANDERS 1882 Ridrag till Ader- och Nerfsystemets Anatomie hos
Myxine glntinosa. Kungl. Rrenska Vetenskapsekad. Handl. (Cited from
Conel, 1931.)
RETZIIjS, GrJsTaF 1893 Das Gehirn und dxs Auge von Myxine. Rial. Untersuch.,
N. F., 5: 55-68.
SANDERS, A. 1894 Researches in the nervous system of Myxine plutinosa. London.
TRETJAKOFF, D. 1909 Das Nervensvstcm von Ammocoetes. 11. Gehirn. Arch. f.
mikros. Anat., 7 4 : 636-779.
 CEREBELLUM O F CYCLOSTOMES 445

A. 1905 Sekundare Bahnen aus dem frontalen sensiblen Trigc-

WALLENBERG,
minuskerne des Kaninchens. Anat. Anz., 26: 145-155.
1907 Beitrage zur Kenntnis des Gehirns der Teleostier und
Selachier. Anat. Anz., 31 I 369-399.
WORTHINGTON,
JULIA 1905 The descriptive anatomy of the brain and cranial
nerves of Bdellostoma dombeyi. Quart. J. Micros. Sei., 49: 137-181.