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Studies on the endocrines of reptiles. I. The morphology of the pituitary gland of the lizard (Anolis carolinensis) with special reference to certain cell types

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STUDIKS ON THE ENDOCRIXES O F R E P T I L E S
I. T H E lUOlil'llOLOGY O F T l l E €'lTUITA€iY G T A ~ N DO F T l l E LlZAItD
( AKOLIS
C A R O L I N E K S I S ) W l T I I SPECIAL REFErtENCE
T O CERTAIN CELL TYPES
ETHEL G. PORIR' AND HARRY A. CIIARIPPER
V n s h u ~ g t o i eSquare Collegi, 1 - e ~Yor.1, Cnwcraity, AT( LC Yor7; Cit!,
m-o
PLBTJES (ELEVEN FIGURES)
IKTRODUCTION
Some cliaractcristics of certain members of tlic reptilian
gronp present problems of interest which fit in with the
current fields of endocrinological iiivestigations. Hypophvseal-gonadal relationships aiicl a possible pituitarv-cliromatop h o x mechanisni a r e two such problems. Thcse have been
investigated from the physiological point of view7 the former
by Hei-lant ( '33 j in l ~ a c c r t aand Anguis ; Clausen ( '35) in
snakes ; Turner ( '35) in the lizard, Eumeces laticeps ; Evans
('35 a a i d b j in Ariolis carolinensis; Mellish ('36) i n
Phryriosoma cornntum; arid Forbes ( ' 3 7 ) in the alligator
lileiriholz ( '36) used Anolis carolinensis and Parker ( ' 3 8 ) ,
Phrynosoma, to investigate llie pituital.y-clironiatop1iol.e
mcchanism. However, these in\-estigatioiis account f o r only
one aspect of the piwblcms involved.
i
as
There is anotlicr approach which has not b e ~ i considcrecl
fully as it might be, namely, the histological one. A careful
examination of the microscopic anatomy of tlie structures involved in such problcrns -\c-ould perhaps contribute niuch toward a more specific knowledge of, among other things, the
role of tlie pituitary gland in reproduction and in skin pigrnentatioii. Only two references coiicerriing the cytological
'Accepted in partial fulfillment of the reyuircnients f o r the degree of Master
of Science at S e w Y'ork University.
473
474
E. G. PORIS AND H. 4. CHARIPPER
aspects of the pituitary glancl in reptiles in relation to a
specific physiological condition are available (Viguier, ’11;
Siler, ’36). Both of these deal with the effects of thyroidectomy on pituitary cell types.
As a preliminary t o future experimental invcstigations involving changes in the glancl, it is important t o have a
thorough knowledge of the iiormal anatomy and histology
of the form to be used. Charipper (’37) stresses this from
the point of view of comparative anatomy and physiology.
He believes that “It may lead us t o a more sober attitude
in our interpretations.”
The purpose of this paper, therefore, is to describe the
normal anatomy and histology of the pituitary gland of
Anolis carolinensis as a basis for future experimental research.
The writers are indebted to Dr. G. I<. Soble of thc L4mcrican
Museum of Satural History f o r some of the animals utilized
in the initial stages of this invcstigation.
NATEIiIATd AND XETHODS
Anolis carolinensis is a species of iguaiiid lizard found in
the soutlierii United States. It is cornmoiily referred to as the
American ‘chameleon.’ The forms used in this study werc
obtained from Thiboclaux, Louisiana, at 1-arying intervals
extending over a period from April to March. The animals
survived well in the laboratory where they werc kept in a
glass terrarium with a screen top. During the summer
months they were fed on houseflies but during the remainder
of the year when flies were riot available, mealworms
(Tenebrio molitor) served as an excellent substitute. The
terrarium was sprinkled twice daily for the purpose of
supplying drinking water.
The animals used for histological study ranged from 50 nim.
to 60 mm. in length. Measurements were made from the
tip of the head to the cloaca1 slit. The entire head was
removed and trimmed off just posterior t o the pituitary
HYPOPHYSIS O F ANOLIS CAEOLlNENSIS
475
gland mhich could easily be seen through the bone. The lower
jaw was cut off and the remainder of the head immediately
dropped into fixative. IIelly ’s, Bouin’s, and Carnoy-Lebrun
were used as fixing agents. Carnoy-Lcbrun proved to be an
excellent fixative since it penetrated very rapidly inhibiting
any postmortem changes. The majority of heads were fixed
in the latter solution. As little as 6 hours fixation was found
to work well. The heads were decalcified in phloroglucin
after which the bone of the roof of the skull was removed.
The basisphenoid bone was left intact in order not to disturb
the normal position of the gland in relation to the brain and
surrounding structures. Several changes of dioxan served to
completely dehydrate and clear the heads. Cross, median
sagittal, and horizontal serial sections were made. All sections were cut at 5 p.
For a study of the general anatomical features of the gland
the slides m-erc stained in Harris’ hematoxylin and eosin.
For cellular cliffcrcntiation the Masson stain was used. In
the latter process the slides were placed into xylol and run
down the alcohols to distilled water. They were put into
Masson A (acid fuchsin) for 2 minutes, differentiated in
Masson B (phosphomolybdic acid) for 4 hours, and then
placed into Masson C (aniline blue) for 1 to 14 hours. The
slides were washed in distilled water and run quickly through
95% and absolute alcohol to xplol. They were mounted in
gum damar.
DESCRIPTION
A . Relation t o surr.ouiiding structures
The pituitary body of Anolis carolinensis lies in a slight
depression of the basisphenoid bone in the region between
the basipterygoid processes. The anterior portion of the
gland is bounded ventrally by the presphenoid cartilage which
extends forward from the basisphenoid bone. Posteriorly
this bone arches over the pituitary to completely surround
the caudal extremity of the gland and form a ledge between
it and the brain. These relationships are readily seen in a
median sagittal section through the brain arid pituitary gland
(fig. 2). An extension of the dura inater coiitiiiues into the
sella turcica covering the pituitary veiitrally and laterally
and giving of1 a partition which separates the pars anterior
i'rorn tlic p a r s intermedia except at the posterior cnd wherc
these two parts of the gland arc coniiected. The snbarachiioicl space is continuous down the stalk aiid around the pars
anterior and intermedia. giviiig iii eff'ect a rascular cavity i n
nliich the g l a d rests. Exteading from tlie lateral ventral
region of the gland on both sides are striated muscle masses
11-hich a r e directed anterolaterally and seem to he coiltinuous with tlie muscles of the eyes. They are best seen in a
cross section of the gland (fig. 4).
The pituitary is uttaclied at its anterior elid to the floor of
the diencephaloii by a relatively long thin-walled infniidihular
stalk which is directed posterjorlp and sliglitlj- ventrally. The
infnndibulai- cavi tv is bounded ventrally by the tliickciied
lamina postoptica. The cavity of tlie third ventricle is coiitinuous with the large infundibular cavity wliieh iiari*ows
down posteriorly to forin the infundibular recess [fig. 1d ) .
The cerebral carotid arteries run lateral to the p a r s interiueclia (fig. 3 ) . A large thin-walled blood sinus is present
dorsal to the gland in the caudal region (fig. 3). No blood
connections could be traced frorii iliese vessels to the pituitar) gland. Blood capilltiries, I i o w e v ~ r ,enter the gluiicl with
the coiiiiective tissue from the surrounding membranes.
B. Gross anntonzy
The pituitary of Aiiolis consists of three lob~s-thc pars
anterior, the pars intcrmcdia, arid tlic p a r s nervosa. There
is no evidence of a pars tuberalis. The p a r s anterior is the
rnobt ventral portion of the gland. It is an elongate structure
with a broad caudal elid which gradual137 decreases in width
up to a distance about one-third of tlic length from the posterior end. A t this poiiit tliere occurs a gradual iiicrease in
width toward tlie anterior extremity where the lolnc ends in
;L blunt point under the lamina postoptica. The shape is
best observed i n the miitral view shown in figure 1c. Two
miall ant criorly directed finger-like projections origiiiat e on
either side of the broad caudal end of the pars anterior. T h y
lerniiiiatc at about the level of the constriction in the anterior
lohe. The p a r s anterior is the loiigcst of the lobes, varying
i n length from Ci42.5 p iii a 50-nim. specimen to 1250 p i n a
65-nim. specimen. In cross sectional TTiews one can observe
il distinct change in shape from the aiilcrior to the posterior
cbstremities. The cranial eiid is more o r less triangular with
the broad base of the triangle dorsally situated (fig. 4). In
the constricted portion of the lobe a n elliptical shape is assnnied (fig. s), this being rciainecl almost t o the extreme p s terior end where it takes the form of an oval (fig. 3).
The p a r s anterior is attached oiilj- a l its posterior elid by
a narrow strand of cells to the caudal extremity of ihe pars
intermedia which lies directly dorsal (fig. 1 d). The p a r s
intermedia is B broad more or less ronndccl structure and is
greater i n volume than any othcr part of the gland. Yentrallp it forms a coilcavity iiito which the elliptical and oval
regions of the pars anterior fit (figs. 3 arid 5 ) . The intermedia is clccpest at about the micl-region (fig. 5). I t iiarro~17s
tlowri posteriorly to form a fla ned, broad plate of cells.
At this region there appear laterally two rather deep grooves
iyliich run a t a n angle pobteriorly and veiitrally toward the
mid-line (fig. 1 c). II is i n these grooves that tlicl c.erehr.al
carotid arteries r u n (fig. 3).
Anteriorly the pars intermedia is closely a p p l i d to the
pars nervosa. The iiiteriiiedia surrounds tlic ncrvosa laterally lout riot dorsally and reritrally except at the posterior end.
Irregular processes extend posteriorly from the dorsal p a r t
of the iiervosa and interdigitate with processes from the p a r s
iritermdia (fig. 1a ) . The infundibular cavity penetrates the
pars ncrvosa sending h o l l o ~diverticula i n t o the posterior
llrocesses. In a horioiital section through the dorsal part of
the gland, t h e system of branching cavities is quite criclent
(fig.8). In cross section the p a r s iiervosa appears t o he made
np of distinct areas separated by clorsorentral connective
478
E. G . PORIS A N D H. A. ClIARIPPER
tissue septa, each area containing within it a branch of the
infundibular cavity (fig. 4).
C. Microscopic anatomy
1. Pars a m t e k r . The cells in the anterior lobe are arranged in anastornosing and branching cell cords which run in
an anterior-posterior direction. These cords are usually two
cells in thickncss and are separated by large capillary
channels (fig. 6). I n cross section the pars anterior has a n
xiniform appearance due to the grouping of the cells about
the blood capillaries (fig. 11). I n the posterior rcgion of the
lobe there is a decided enlargement of the capillaries to form
sinnses (fig. 6). I n some regions the vascular luinina may be
seen t o be continuous with the peripheral blood sinus which
appears to surround the lobe.
The cell types differ in the anterior and posterior regions
of the gland. Anteriorly there are, bordering on the capillaries, columnar cells with a basal, spherical nucleus, and fine,
densely packed eosinophilic granules gathered on the side
between the nucleus and the apex of the cell (fig. 6). The
nucleus is vesicular in nature, and contains one eosinophilic
nucleolus. Also bordering on the capillaries are taller
columnar cells with basal vesicular nuclei and large bluestaining granules which are more diffuse than those of the
eosiiiophilic cells. Between the bases of the columnar cells are
a few round cells with an intenscly eosinophilic coarsely
granular cytoplasm and a spherical nucleus displaced t o one
side of the cell. The major proportion of the cells in this
region, however, are round cells with an extremely large
nucleus and a small amount of cytoplasm. Cell outlines are
not very distinct. A few basophilic granules are present.
Lining the blood sinuses in the posterior part of the gland
are large round to ovoid cclls which take an homogeneous
eosinophilic stain (fig. 9). The hasally situated nucleus has
an irregular spherical shape and in many cases a crescent
shape is obvious. Similarly staining cells are present between
the sinuses. In these elements the nuclcus is displaced t o one
HYPOPHYSIS O F ANOLIS CAROLINENSIS
479
side of the cytoplasm. Large round basophiles similar t o
those in the anterior region occupy the remaining area between the sinuses. They are especially abundant in the ventral portion of the lobe where they also border on the sinsues
and assume a columnar shape. A few columnar granular
eosinophiles, characterislic of tlie anterior region of the lobe,
may also be seen bordering on the sinuses.
2. Pars imntermedia. The pars intermedia in horizontal and
cross sections consists of cords of cells arranged about fine
blood capillaries. The cords run in an anteroposterior direction and make up the mass of this portion of the gland
(fig. 10). Here again, as in the pars anterior, the cavities
of the blood vessels may in places be seen to be continuous
with the lumen of the enveloping vascular sinus. The
majority of blood elements, however, enter with the connective tissue extensions from the membrane surrounding the
pars nervosa. The cells making up the cords are very tall
columnar with small basal spherical nuclei. These nuclei,
like those of the anterior lobe, are vesicular and have an
eosiiiophilic nucleolus. The cytoplasm is filled with fine and
coarse reddish purple granules. Between the bases of the
columnar cells are large, somewhat oval cells with rather
irregularly shaped nuclei (fig. 10). These cells contain a
homogeneous reddish orange cytoplasm in which may be
observed a few small clear vacuoles. Somewhat lighter
staining accumulations of an amorphous material are also
present between the bases of the cells. Vacuoles of various
sizes are present in tl!is colloid-like material (fig. 7).
3. Pars 9zeruosa. The pars nervosa consists entirely of a
rather compact fibrous tissue with a few scattered oval to
spherical nuclei. This tissue ramifies into tlie pars intermedia, not as a solid mass, however, but peiietrated by hollow
diverticula of the infundibulum. A single row of typical
epeiidymal cells lilies the cavities thus formed. The nervosa
is supplied with blood vessels from the connective tissue
surrounding the lobe. No colloid accumulations were evident in the sections studied.
480
E. G. POEIS AND H. A. CHAnTPPER
DISCUSSION
The relationship of tlie pituitary gland in Snolis to the
surrounding structures in tlie skull is similar t o that described
in other reptiles. Tilney ('11) and Stendell ('14) speak of
tlie pituitary as lying in a deep sella turcica. Siler ('36)
states that the sella burcica in which the gland of Tliamnopliis
lies is a depression in the basispherioid bone. Snolis similarly lies in a depression in the basisplieiioid bone and resembles the turtles, Chrysemys clegans and Pscuclemys picta
(Raumgartner, '16) and the garter snake, TliarnnolJliis radix
(Silcr, '36) by being completely surrounded canclallj- h y a
doi-sal extension of the hasisphenoid. Like Splienodon it is
bonucled ventrally at its anterior encl by the presphenoid
cartilage. Stadcrini ( '03) called attcntion t o two striated
mnscle masses at the sides of tlic pituitary in Goiig~lus. Gisi
( '07) noted them in Lacerta : Ranmgartiier ('16) in Sceloporus, Chrysemys, ancl k'scudcriiys ; and Xyctli and Rom
( '23) in Splienodon. The latter investigator refers t o thein
its the internal ptcrygoicl niiiscl~s. Banmgartner merely suggests that they probably represent cye muscles. The olsserratioiis on Anolis confirm this since they appear to be directly
continuous with the ocular muscles. The cerebral carotid
arteries liave bcen described as running alongside the caudal
end of the pituitary in Lacerta, Chrpsemys, Pseudemys,
Scclopoi~us,Alligator (Baurngartner, '16) and S1)llenodon
(a'ycth ancl Uow, '23). These have also been o b s e r ~ ~ ein
d
Aiiolis. No blood vessels could be traced from these vessels
into tlie gland. Baumgartner ('16) also reports that a n injected alligator showed no direct branches from thc carotids
t o the hypophysis. Baumgartner noted a large ~11011s
channel, caudodorsal t o the liypophysis in lizards and snakes,
m-hich lie believes drains the organ. h large blood sinus,
similar in position, was noted in Anolis. 'Whcther tlie organ
is draiiiccl by this clinnncl remains to be determinecl. Tn this
lizard definite capillary chaiinels 17enetrate the glaiicl from
the illvesting arachoirl membrane. This agrees with the ohservations of Siler ('36) on Thamnophis.
HPPOPHPSIS O F ANOTAIS CAROLINESSIS
481
The posteriorly directed infundibular stalk sccil in Anolis
is characteri stic of reptiles in general (de Beer, '26 ; Tilnej-,
'11; Stendell, '14; Geiiteq, '07). clc Beer describes a long
slender stalk in the tortoise, grass snake, Laeerla aiid
Sphenodon, and Stenclell speaks of a long stalked infunc3il)ular process in the Laeertidae, Varnnus, Xcincus, Chamaelcon,
Igiiana, and Basdiscus. Anolis rrsembles these forms i n regard t o the length of the stalk and simulates Lacerta ninralis
aiid L. viriclis in senclins branching diverticula from the infuiiclibular cavity into the nervosa (Stendell, '14 ; Gentes,
'07). This arrangement lias also been describcd 117Rtciiclcll
f o r Emys and by Herring ('13) f o r Tesludo.
Anolis possesses the three lobes cliaracteristic of all reptilw, nilnlel>-, the pars anterior, pars intermedia, ai1tl liars
nervosa. The veiitral position of tlie pars anterior aqrees
with the descriptions of de Beer ( %), Tilney ( ' l l ) , Sterxi
( ' O q , Baumgartiicr ( '16) and Siler ('36) on otlier relitilian
Corms. Anolis resembles other Sauropsids ( Stenclell, '14) in
having the anterior lobe only loosely connected at its 100slerior end to the pars intcrmcdia. In this respect it differs
from the Chelonians (Gentes, ' O i ) wlicre the two lobes are
joined at both anterior and posterior. ends. de Bccr ( '2G)
describes ill tlie tortoise a forward prololigation of the pa1.s
intermedia which he refers t o a s the pars tuberalis. l n
('istuda. and Slligator, Tilney ('11) speaks of an accessory
tongue-like projectioii extcncliiig forward from the anterior
extremity of the gland toward the chiasma. Baumgartncr
('16) states that the lateral buds persist to form a pars
tuheralis and corlical zone about the anterior lobe in turtles
and a tongue-like process and spiral band about tlie anteiior
lobe in alligators. Stendell ('14) states that in some forms
(Chelonia and Saurians) there is a long tong~ie-likeprocehs
projecting from l h c pars anterior.. (2 entes ( '07) remarked
that there is no formatioil in lizards which corresponds to the
lateral lobes of tlie pituitary. This observation is confirmed
in the Anolis adult hut TI-hether it exists in the embryo as has
been described f o r Lncerta muralis (deReer. '26) is not
known.
482
E. G. PORIS AND H. A. CHABIPPER
There is no evideiice of an hppophyseal cavity in Anolis.
This is not unusual in the reptilian group since the cavity
persists in some forms and disappears in others. Stendell
('14), deReer ('26), and Baumgartner ('16) describe a
residual lumen in the pars intermedia of some lizards.
Gentes ('0'7) and de Beer report the presence
Herring ( 'U),
of a cavity between the pars intermedia and the pars glandularis in Chelonians.
The cord-like arrangement of cells in the anterior lobe is
peculiar to all reptilian forms that have been described and
is also an outstanding feature of the Anolis pituitary. A
great deal of variation, however, exists in the cell types.
de Beer found very little difference between the cells apart
from being eosinophilic or basophilic. Tilney ( '11 ) described
deeply staining and faintly staining acidophiles in the turtle
as well as peripherally located basophiles. Herring ('13) reports the presence of clear non-granular cells ancl deeply
staining granular elements in the tortoise. I n the alligator
T h e y finds acidopliiles ancl lnasophiles while Baumgartner
('16) observed some cells that took eosin while others remained clear. I n Entaenia only faintly staining acidophiles
are present (Tilney, '11). Siler ('36) observed three types
of cells in Thamnophis-cells with coarse red-staining granules, cells with finer acidopliilic granules which he considers
to be chromophobes, and beta cells with fine granules. These
obviously confusing results are very likely attributable to diff erencrs in technique, thus ]nrecluding any comparison of the
work of these various investigators. Three types of cells are
also present in Anolis-cells with eosinophilic granules, cells
with basophilic granules, and non-granular chromophobes.
The anterior lobe in Anolis is extremely vascular and in
this respect agrecs with other reptiles with the exception
of Eutaenia in which no blood vessels were observed (Tilney,
'11). Anolis resembles Sphenodon in showing an increase in
the size of the vascular sinuses toward the posterior end of
the lobe. No colloid was noted in Anolis, such as has hcen
descrihed in the lnmina of the cell cords in the turtle and
IITI'OI'HPSIS
O F ANOLIS CAROLINENSIS
483
alligator (Tilney, '11); tortoise (Herring, '13) ; and intercellularly in Kmys and Cistuda (Steadell, '14).
The pars intermedia of Anolis is unusual because of its
large size, being the largest lobe in tlie gland. Stieda (1875)
fourid a small dorsal portion closely united to the lobus infundibuli and a large ventral lobe. Stendell ('14) observed a
very small '~wisehenlappeii~
in Sauropsida and Chelonia but
noted that it was fairly well dereloped in Ophidia. Herring,
('13) noted the presence of a well-marked pars intcrmedia in
Testudo and Laccrta viridis. Gentes ('07), on the other hand,
found a sinall intermedia in Lacerta viridis, L. muralis, and
Cistudo. Nost investigators noted that thc intermedia had a
poor staining reaction (de Beer, '26 ; Tilney, '11; Stendell,
'14; Gentes, '07). The well-defined, intensely staining granular cytoplasm of anolis is thus i n contrast t o what has been
noted in other forms. Coiiceriiing tlie blood supply, dc Beer
('26), Stendell ('14), and Gentes ('07) report that this lobe
is sparsely supplied with blood vessels. Tilney ('11) found
no blood vessels at all in Eutaenia, and Stendell ('14) states
that t h c intermedia of sauropsids lacks blood vessels entirely.
I n this respect too, the pars intermedia of Anolis is different
from that of other reptilcs, since it possesses a cicfinite sj-stem
of fine capillaries. On the basis of its large size, heavily granular cells, vascular supply, and colloid-like secretory material,
the pars intermedia may be interpreted as a functionally
active portion of the pituitary gland. Kleinholz ('36) has
shown that the characteristic body color cliaiiges observed
in Anolis are in part due to a pituitary liorrnone. Zondek
( '35) has isolated a chromatophoric principle, 'intermedin,'
from the pars intermedia of many forms. Further histological and cytological iiivestigation of the intermcciiate lohc
of Anolis under various conditions may offer evidence to
substantiate the conclusions of other investigators that the
pars intermedia produces an active principle which controls
color changes. It is interesting to note that Zondek ( ' 3 5 ) mas
able t o demonstrate the hormone in the walls of the third
ventricle and nowherc else, a fact which suggests the pituitary
484
E. (3. P O l l I S AND H. A. CHARIPPEP,
stalk a s a pathway f o r the distribution of this hormone. The
increased surface of the infundibular cavity i n Anolis,
brought about by the branching diverticula, might \T ell be
associated with snch a function.
Xiiolis is thus seen to resemble the other members of the
reptilian group in general anatomical features and cord-like
arrangement of cells in both pars anterior and p r s intermedia. Tlie main cliEereiice exists i n the features of the
pars intermedia which have alread? been described.
STXXARY
1. The pituitary gland of Anolis carolinensis lies i n the
hasisphenoid bone and is surrounded b-j- the dnra niater
a n d araclinoid membrane from which blood capillaries enter
the gland.
2. The short infunclibalar stalk is directed baclm-ard carrying the intermedia and nervosa posteriorl-. The anterior
lobe is ventral and is attached to the intcrmeclia only at its
posterior end. ‘Ilierc is no pars tuheralis.
3. The iiifundibular cavity is continuous with the third
1-entricle and sends liollo-cv diverticula into thc nervosa.
These cavities arc lined with a single layer of ependyina cells.
4. The pars anterior is the longest p a r t of the gland while
the p a r s intermedia is thc greatest in volume.
5. Thrce distinct cell types a r c present in llic anterior lobe
-eosiiiophilic and basophilic granular cells and pink-staining
non-granular cells.
6. The pars intermedia consists of columnar cells with
red-purple granules and oval cells with non-granular orange
cytoplasm. Colloid-like accumulations are also present.
7. There is no evidence of a residual lumen.
IIPI’OPHYSIS O F ASO1,TS CAROLIXEKFIS
485
LITERATURE CITEI)
BAEMGARTSER,
E. A. 1916 The development of the hypophgsis in Reptiles.
J. Morph., vol. 28, pp. 309-275.
DE BEER, G.. R. 1926 The comparative anaturn)., histology, and development of
the pituitary body. Oliver and Boyd, London.
CHAEIPPEA, 11. A. 1937 The morphology of the Iippopli?;sis in lowcr vertebrates,
particularly fish and Amphibin, with some notes on the cytolog.7 of the
pituitary of Carassiiis auratus (thc goldfish) and Nectiirus marulosus
(tho mudpuppy). Cold Spring Harbor Symposia on Quantitativc
Biology, vol. 5 , pp. 151-164.
CLAUSEPI’,H. J. 1935 The effects of ovariotomy and hgpophgsectoiiiy on parturition in snakes. Anat. Rec., vol. G4 (Snppl. no. I ) , p. 88.
EVAITS,L. T. 1 9 3 5 s T h e effects of antuitrin S a n d sheep pituitary extract
on thc female lizard, Aiiolis enrolinensis. Biol. BuLI., 1701. 68, pp.
355-359.
- 1936 b
The effcct of xntuitrin S on the nialc lizard, Anolis
enrolinensis. Anat. Bee., rol. 62, pp. 213-993.
FO&HES,
T. R. 1937 Studies on the reproductive svsteni of the alligator. I.
Tlic effects of prolonged injections of pituitary whole gland extract in
the iinmaturc alligator. Anat. Rec., vol. 70, pp. 113-13i.
GENTES,’L. 1907 Rrcherches sur 1’li;vpophyse et lc sac Taseulaire dcs vertbbres.
Roc. Scient. D’brch. Sta. Biol., T. 1 0 , pp. 129-281.
GIN, J. 1907 Das Gehirn von Hat.trria puiictata Dius. Rascl. Quoted froin
Maumgnrtner, 191ii.
HRRIAINT,
31. 1933 Recherches histologiqiies r t exp6riiiicritales sur les variations
cycliques du t e s t i c i i l e et des caract~&ressexuels sesondxires chcz les
reptiles. Arch. de Riol., T. 44, pp. 347-468.
IIERRIL-G,
P. T. 1913 Furtlrer ohvervations npm the comparative anntoing and
physiology of the pituitary body. Quart. J. Exp. Phys., vol. 6,
pp. 73-108.
KLEINIIOLX,
L. 11. 1936 Studies in reptilian color changes. I. A preliminary
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486
E. G. PORIS A N D H. A. CHARIPPER
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PLATE 1
EXPLBFATION O F FIGURES
1 Drawings of the pituitary gland as seen from four different views, dorsal
( A ) , lateral (B), ventral (C), and median sagittal (D). P a r s anterior is
coarsely stippled, pars intermedia finely stipplcd, and pars nervosa is solid black.
Anterior is t o the right. X 145.
2 Photomicrograph of a sagittal section through the brain and pituitary gland
showing the arching of the basisphcnoid bone over the gland, and the presphenoid cartilage extending forward from the bone. The optic ehiasma is seen
t o thc left i n front of the gland. Cxrnoy-Lebrun, Masson. X 10.
3 Photomicrograph of a cross section through the posterior portion of the
gland showing the two grooves on the sides of the pars intermedia i n which run
the cerebral carotid arteries. Dorsally situated is a large blood sinus. The pars
anterior is ventral. Carnoy-Lebrun, ;\.lasson. X 120.
4 Photomicrograph of a cross section through the anterior portion of the
gland showing the pars nervosa with four branches of the infuiidibulum. The
par? intermedia is seen lateral to t h e nervosa. Note the dark niuscle masses
under the gland. Carnoy-Lebrun. Masson., X 120.
5 Photomicrograph of a cross section through the mid-region of the gland
showing the large size of the intermedia and the cord-like arrangement of cells.
Carnoy-Lebrun, Masson. X 130.
PLATE 1
IITPOPIIYSIS 01" A U O L l S C 4 K O L I N E N S I S
E. (1. POllIh .\NU
H A
CklSI~1PPbll
487
PI, A ‘I’ b: 9
EXPI,.2Nh’I’ION
O F PIG1:RES
A11 figures arc photornicrograplis of sections f r o m glands fixed i n (‘:~rnoy-lxbrun
aiid dairied with IIassori.
li Frontal settioii thTougIi thc pars auterior sliowing tlic cords of cells
separated 1)y blood c.;ipill:irics. The voluniiiar gr:rnular cells 011 tlic rig11t arc‘
mostly eosinophiles. The dark cells o n the left are the uon-granular pink staining
r.ells eharat:t.cristic of tlir posterior end o f the lobe. >: 600.
T S~ciioizthrough t.lie pars iiitwmeilia. showing colloid-like uccuniulations
b e t ~ e e nthe h a w s of the columnar cells. X (iO0.
8 F’ront:cI section through t h e gland showing the hranching di
osa. The pars iiiterrnedia is seen closely a p p l i d t o the i i ~ r r o s : ~ .X 120.
9 C’ross section of t h c pustericr end of the pars antwior. The large dark
cclh hordcriiig 011 thc siiiuscs arc tlic same as those i n figure 6. ‘I’hc lighter
slxiriirig cells on the lcft are basupliiles. X 1400.
10 Section through the pars intermedia. showing the arrangement of cells.
Note the small dark cclls betn-ecii thc hases of the ccIluiiiiiar cells. Thrxe have
a honiogeiieous reddish orange cvtoplasm. X 600.
11 Cross scction through Ihc anterior lobe showing t h e numerous blood
sinuses. X 600.
488
IIIPOPHYSIR OF A K O L I S CAROLINBKSIS
1,
G
PORIS AhTJ H
4
PLATE 2
CII1RTPI’FPr
489
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morphology, endocrine, carolinense, anolik, references, typed, special, cells, certain, lizard, pituitary, gland, studies, reptiles
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