Some histological considerations of the aortic paraganglionic bodies (of Zuckerkandl) of the guinea pig

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Content SOME HISTOLOGICAL CONSIDERATIONS OF
THE AORTIC PARAGANGLIONIC BODIES
(OP ZUCKERKANDL) OF THE GUINEA PIG
A Thesis
Presented to
The Faculty of the Department of Zoology
University of Southern California
In Partial Fulfillment
of the Requirements for the Degree
Master of Arts
by
Wilbur L. Brandt
August, 1942
UMI Number: EP67151
All rights reserved
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OissGftation Publishing
UMI EP67151
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This thesis^ written by
........... %llbiir JLiK .Brafi&t............
under the direction of Æ L s . . Faculty Committee,
and approved by all its members, has been
presented to and accepted by the Council on
Graduate Study and Research in partial fulfill
ment of the requirements for the degree of
.Master
Secretary
Dnt. AURUSt,1942
Dean
Faculty Committee
Chairman
ACKNOWLEDGMENTS
The author wishes to express his gratitude and thanks
to Dr. P. M. Baldv/in, chairman of the Thesis Committee, for
his advice and help in the preparation of the Thesis.
Thanks are also expressed to Drs. B. M. Harrison and W. G.
Campbell, members of the Thesis Committee, for their
assistance in reading and correcting the manuscript.
I am grateful to Mrs. Willard Kleiter for her help in
typing and proof-reading the manuscript.
Thanks to Mr. 0. B. Wiswell and Mr. Willard Kleiter
for their help in taking and printing the photomicrographs
and diagrams.
A special thanks to Mrs. Wilbur L. Brandt, without
whose inspiration, help and persistence the Thesis might
never have been written.
W.L.B.
TABLE OP CONTENTS
CHAPTER PAGE
I. Introduction 1
II. Review of the Literature 5
III. Materials and Methods 10
IV. Observations 15
V• Summary 40
Bibliography
ILLUSTRATIONS
PLATE PAGE
I. Diagram showing the Chromaffin
System of Mammals
II. Diagram showing the Location of 8
the Paraganglionic Bodies in Man
III.-A and B Photomicrograph showing a lY
Myelinated Sympathetic Nerve
IV. Photomicrograph showing the Hilus 21
of the Paraganglionic Body
V. Photomicrograph showing a Secondary 24
Lymph Nodule
VI. Photomicrograph showing an Artery 26
in the Paraganglionic Body
VII. Photomicrograph showing a Blood 28
Sinus
VIII. Photomicrograph shov/ing a tubule 31
which has assumed the Function of
a Blood Vessel
IX.-A and B Photomicrograph showing a 34
Tubule
X. Photomicrograph showing a "Corpuscle" 36
XI. Photomicrograph shov/ing an Artery, 39
Vein and Blood Sinus
CHAPTER I
Introduction
Though chromaffin cells occur throughout the body
of animals, vertebrate and invertebrate alike, and have
been described by many v/orkers, their physiological signi
ficance and varying anatomical topography are not very well
understood. As the name implies, these cells are particu
larly responsive to staining with chromic acid, osmic acid,
and certain chrome salts. When so treated, the cells take
on a yellowish, dark-brovm, or black coloration, which, in
the opinion of such workers as Maximow and Bloom (1938),
Morris (1933), Cowdry (1938), is caused by the supposed pre
sence of epinephrin (adrenalin) in the cells.
In invertebrates, chromaffin cells are distributed
throughout the body being found in every organ. As we pro
ceed up the scale to the higher type animals, the chromaffin
tissues and cells become more and more localized until in
mammals they appear as definite masses occurring in definite
anatomical positions. (Plate I)
In mammals, definite masses of chromaffin cells are
closely associated with the sympathetic ganglia, occur in the
cervical, abdominal, and coccygeal regions ofthe mammalian
body, the best known aggregation of chromaffin cells being
found in the medulla of the suprarenal gland.
PLATE I - Diagram showing the Constituents
of the Chromaffin System in Mammals*
(Morris’ - Human Anatomy)
Carotid body
Sup. cerv. ganglion
Inf. cerv. ganglion
Stellate ganglion
Sympathetic ganglion
Sympathetic trunk
Chromaffin groups in sympathetic ganglion
Accessory cortical body
Suprarenal cortex
Suprarenal medulla
0
O"
O"
Abdominal chromafSn cell-groups
oo
Aortic or lum bar paraganglion
Accessory cortical body
PLATE I
4
The present problem evolved from some preliminary in
vestigation carried on by a member of the Physiology depart
ment. The purposes of this investigation are: (1) to study
the histological structure of the aortic paraganglionic
bodies; and (2) from the histological study, draw such con
clusions as seem reasonable as to their possible physiological
function.
CHAPTER II
Review of Literature
Literature on the aortic paraganglionic bodies is
very limited.. There is also considerable disagreement
among the authors concerning the appropriate name. G-errish
(1902), Piersol (1918), and Jackson (1933) give the name of
"aortic paraganglionic bodies” (of Zuckerkandl) describing
them as a group of chromaffin bodies which have developed
from the sympathetic nodules. In man, these bodies attain
a large size, which may represent a fusion of several
chromaffin bodies into an elongated cylinderical mass over
one centimeter in length in the young. They lie retroperi-
toneally, ventrad to the dorsal aorta, at the level of the
inferior mesenteric artery. Their size and extension is
extremely variable. In the human, the bodies are said to
undergo partial retrogression after the state of puberty
has been reached.
Cunningham (1903) and Gray (1930) give substantially
the same account of the bodies but refer to them as the
”aortic bodies”. Schmidt and Comroe (1940) apply the name
”aortic bodies” to a group of chromaffin bodies in the cer
vical region. Baldwin, Irons and Wiswell (1942) in an un
published paper, given before the meeting of the Western
Division of the American Academy for the Advancement of
Science, on the anatomical considerations of the bodies,
attaches the name, "aortic paraganglionic bodies'* •
After having done considerable work with these bodies,
the name "aortic paraganglionic bodies*' or "lumbar paragang
lia” seems to give a more accurate description of them and
thus it will De used throughout the paper.
During man* s fetal development, the aortic or lumbar
paraganglionic bodies arise from the sympathetic nodules,
and are considered by most workers to be of ectodermal origin.
Arey (1937) says that the bodies are recognizable toward the
end of the second month of embryonic life at the root of the
inferior mesenteric artery. At birth they attain a length
of about one centimeter and are composed of cords of chrom
affin cells intermingled v/ith strands of connective tissue,
the whole being surrounded by a connective tissue capsule.
Arey further says that after birth the chromaffin bodies de
generate but do not entirely disappear (Plate II).
Cowdry (1938) applies the name ”aortic bodies” to
chromaffin bodies in the cervical region associated with the
carotid bodies. This coincides vfith the report of Schmidt
and Comroe. Cowdry says nothing specific about the structure
of the aortic paraganglionic, or lumbar, bodies, but says
that extra adrenal chromaffin tissues, paraganglia, aortic
and carotid bodies, come into play when there is an increased
PLATE II- Original Diagram showing the
Location of the Aortic Paraganglionic
Bodies in Man.
8
-Aorta
-Inf. Vena
Cava
— Left Kidney
Left Renal
Vein
Spermatic
Vein
Sympathetic
Trunk
Aortic paraganglionic,
or lumbar, body
Inf. Mesenteric
Artery
Left Common
Iliac Artery
=-Plexus Aorticus
PLATE II
9
demand and tension on the medulla of the suprarenal gland by-
taking up the shock and secrete epinephrin to obviate the
necessity of hypertrophy of the true medulla.
Bremer (1936) and Maximow and Bloom (1938) give much
the same account of paraganglia as Cowdry, but they do not
mention the aortic bodies.
Literature on the physiology of the aortic paragang
lionic bodies is lacking. However, much is written about
the sympathetic nervous system and its constituents and much
about chromaffin tissue generally.
All of the above investigations apply to man. To the
author's knowledge, no literature exists which reports any
investigation on the guinea pig.
CHAPTER III
Materials and Methods ♦
Twenty-three pairs of aortic paraganglionic bodies
of the guinea pig were used in this investigation. About
one half, or eleven pairs, of the bodies were obtained by-
autopsy from animals that had been killed by ether asphyx
iation. The remainder, twelve pairs, were removed from the
animals by surgical operation under an ether anesthetic. In
each instance the bodies were placed in a killing and fixing
reagent immediately after removal from the animal. These re
agents were Bouin* s solution, and ten per cent formalin.
Bouin’s solution did not prove to be very satisfactory as it
contained a saturated solution of picric acid. The picric
acid was so impregnated in the tissues that several washings
(eight or ten) in seventy per cent alcohol, with lithi-um car
bonate added, failed to remove it. This later impaired the
staining qualities*of the tissues. For this reason, it was
decided to discard Benin's solution.
Ten per cent formalin had its disadvantage, too.
Formalin left the tissue so white that it was very hard to
distinguish the tissue from the surrounding paraffin, used
as the imbedding material. Just enough of a one per cent
picric @.cid solution was added to the ten per cent formalin
to give the formalin a pale yellow color. This would color
11
the tissue sufficiently to make it easily distinguishable in
the paraffin block. Yet the picric acid was easily washed
out of the tissue with two or three v/ashings in seventy per
cent alconol, with lithium carbonate added. Three or four
drops of lithium carbonate was found to be sufficient to re
move the picric acid.
TO demonstrate the chromaffin cells, the killing and
fixing solution of Wiesel's method, as suggested by Carlton
(1926) was used. This solution contains 10 c.c. of a five
per cent solution of potassium dichrornate, 20 c.c. of ten
per cent formalin, and 20 c.c. ofdistilled water.
In all of the killing and fixing reagents, Bouin's
formalin, or Wiesel's, the tissues were left for two to four
days.
The chromaffin reaction, which was expected to have
been relatively strong, was very weak. It was thought that
the formalin might have reduced the chromaffin reaction,
which is a reaction betv/een the epinephrin in the cell and
the potassium dichromate in the killing and fixing reagent.
The killing and fixing procedure was modified so that
the tissue was treated v/ith a ten per cent aqueous solution
of potassium dichromate and normal (0.7 per cent) saline
solution. This solution was heated to 37 degrees centigrade
and the tissue placed in it immediately after removal from
12
the animal* These tissues and solution v/ere placed in an
incubator at 57 degrees centigrade for tv/o hours. The
tissues were then placed in ten per cent formalin where they
were left for two to four days.
The chromaffin reaction was not sufficiently improved
by this procedure. Extensive research on the subject soon
revealed that drugs and anesthetics such as morphine, ether,
etc., reduced the chromaffin reaction to almost nothing.
The guinea pigs used in this investigation were either killed
by ether asphyxiation, or were operated on while under an
ether anesthetic.
The ages of guinea pigs used during this investigation
extended from three months to over two years and their
weights ranged from 150 to 400 grams. Both sexes were in
cluded.
Two guinea pigs were killed by injecting a little
air into the blood stream of the animal, causing an embolism.
The aortic paraganglionic bodies from these animals were
placed in the saline potassium dichromate mixture and the
potassium dichromate and formalin mixture. In each case the
chromaffin reaction was very strong proving that the ether
was the cause of reducing the chromaffin reaction, not the
formalin as first supposed.
Following is a brief summary of the reagents used and
13
the length of treatment in each.
1. Killing and fixing reagent — two to four days.
2. Fifty per cent dioxane (several changes) -- twenty-
four to thirty-six hours.
5. Absolute dioxane (eight to ten changes) -- one hour
each.
4. Dioxane-paraffin (one half and one half mixture) --
two to three hours at fifty-five to sixty degrees centigrade.
5. Pure paraffin (two baths) -- three hours each#
6. A third bath in a mixture of nine parts, by v/eight,
of paraffin to one part, by weight, of bayberry wax -- one
hour.
7. Imbed in a separate mixture of the same ingred
ients as No. 6.
The sections were cut eight to ten microns thick.
Both cross sections and longitudinal sections were cut in
serial, and mounted on slides in the usual manner.
The sections were stained with Delafield* s haemotox-
ylin and counter-stained v/ith alcoholic eosin, Mallory's
Triple stain according to the procedure given by Harrison
and Richins; Heidenhain's Azan stain, and Hansen's Chrom
Alum haematoxylin counter-stained with alcoholic eosin accord
ing to the procedure given by Bensley and Bensley; Bodian
silver reduction (protargol) stain according to the procedure
14
given by Bodian in the Anatomical Record.
The sections were mounted with clarite, a sixty per
cent solution in Toluene or Xylene.
CHAPTER IV
Observations
The aortic paraganglionic bodies (of Zuckerkandl)
of the guinea pig are small, about 5 mm., elongated bodies.
Their color ranges from a glistening v/hite in the young
animals, from birth to three or four months ; to an egg-shell
white in young adult animals, four months to a year and one
half; to a grayish-white in animals over a year and one half.
The bodies are located retroperitoneally on either
side of the dorsal aorta at the bifurcation of the aorta
into the right and left common iliac arteries. They are not
found in the same anatomical position in every animal. In
the younger animals, the bodies are found just below the
origin of the inferior mesenteric artery. In older animals,
the bodies are usually found just below the bifurcation of
the aorta along the side of the common iliac arteries. In
every case, however, the bodies are attached to the ventral
fascia of the psoas muscles.
The blood supply is very abundant. The arteries arise
from minute branches off the right or left iliac artery. The
veins which drain the right body lead into the inferior vena
cava, while the veins which drain the left body lead into the
ovarian or spermatic vein. The medulla is highly vascular.
Plates III-A and B, Plate VI, Plate VII and Plate XI show
PLATS III-B. Photomicrograph showing
the Myelinated Sympathetic Nerve, x900
SN - Sympathetic Nerve
Vn - Venule
Ar - Arterioles
PLATE III-A, Photomicrograph showing
the Myelinated Sympathetic Nerve# x400
SN - Sympathetic Nerve
Vn - Venule
Ar - Arterioles
17
PLATE III -B
W
iitf
m
PLATE III -A
18
arterioles, venules and blood sinuses. Nearly every section
of the medulla will show one or all of these in it.
The sympathetic nerve supply comes from the abdominal
aortic plexus. The nerve enters the body as a single branch
from the plexus, penetrating to the medulla of the body be
fore it divides. Plate III-A shows this nerve just as it is
dividing. The nerve is surrounded by prominently nucleated
lymphocytes in contrast to the loose connective tissue
surrounding the arteriole and venule also shown in the section,
Under somewhat higher magnification (Plate III-B) the nerve
becomes much more conspicuous and the axis cylinders and the
myelin sheathes become visible. The arteries, veins and
nerves enter or leave through a hilus near the anterior pole
ofthe body (Plate IV).
In the adult guinea pig the right and left aortic par
aganglionic bodies are not the same size. The left is
usually slightly larger. The left body usually measures
five to seven millimeters, v/hile the right only four to six.
The width is usually two millimeters for each.
Histologically, the bodies are unique. They resemble
a lymph node in that they possess a cortex and a medulla,
secondary lymph nodules, reticular connective tissue forming
the framework, and the predominating cells being lymphocytes.
A fibrous connective tissue capsule surrounds the body and
19
becomes the trabeculae as it penetrates the body. The para
ganglionic bodies resemble the hypophysis cerebri in that
alpha cells (eosinophils) are found in large numbers in the
medulla. The resemblance to the thymus gland is noted by
the presence of a corpuscle-like structure resembling the
thymic corpuscle (Hassall's). (Plate XI). They differ from
the thymic corpuscle, however, by having a lumen, which is
not found in Hassall's corpuscle of the thymus. They are
found exclusively in the center portion of the medulla, and
range in size from fifty to eighty micra. These corpuscle
like structures in each case are surrounded by large numbers
of eosinophils. In one or two instances, a fibrinous mater
ial was found in the lumen, but in the majority of cases, the
lumen was clear.
After studying the sections, the idea suggests itself
that these bodies seem to represent a fusion of some of the
cells of the lymph glands, pituitary and thymus, v/hich, dur
ing their development, were not included in them, but somehow
became detached, and all these cells fused together to form
the paraganglionic bodies. Of course, such an origin could
have no scientific basis, as no tissue, or organ, develops
as an accumulation of cast off cells of other tissues. It
does, however, illustrate the complexity of the cellular
structure of the paraganglionic bodies. The cells resembling
PLATE IV. Photomicrograph showing the
tiilus of the Paraganglionic Body. xBO
H - Hilus
V - Vein
A - Artery
PLW - Primary Lymph nodule
Ar - Arteriole
21
%
V > ‘
PLATE IV
22
the lymph glands, lymphocytes, constitute the greatest maj
ority of the cells. Next, in equal proportions, are the
eosinophils and the red blood cells. In lesser numbers are
the cuboidal epithelium and the erescent-shaped cells.
The paraganglionic bodies appear to be some type of
lymph gland because of the close similarity in structure
to them. The cortex is composed mainly of large lymphocytes.
These lymphocytes, when stained v/ith a haemotoxylin (Dela-
fieldds), are very dark because of the perponderance of
nuclear material. Some of the lymphocytes have become more
closely compact giving the appearance of primary lymph
nodules. However, the light spot, or "germinal center",
usually called the secondary lymph nodules, is absent
(Plate V). In one of the largest sections, there appear to
be approximately six primary lymph nodules in the cortex.
The medulla of the paraganglionic bodies bear a
slight resemblance to that of the lymph glands. The arter
ies, which enter through the hilus, go to the center of the
body where they break up into small arterioles and then
into blood sinuses. Numerous phagocytic cells line the walls
of the arterioles and veins. These phagocytic cells, how
ever, are not found in the blood sinuses.
The medulla consists of a framework of reticular con
nective tissue with lymphocytes filling its meshes. The oval
PLATE V, Photomicrograph showing the
Primary Lymph Nodules in the Cortex. xBO
PLN - Primary Lymph Nodules
24
4
JELÜ
PLATE V
PLATE VI, Photomicrograph showing an
Artery in the Paraganglionic Body. x900
A - Artery
V - Vein
RBG - Red Blood Cells
Ar - Arteriole
Vn - Venule
26
K
PLATE VI
PLATE VII. Photomicrograph showing a Vein
and a Blood Sinus in the Paraganglionic Body,
x400
BS - Blood Sinus
V - Vein
RBG - Red Blood Cells
28
r-.
# - ^
&
V
¥,
PLATE VII
29
nuclei of the reticular cells are easily seen but the anastom
osing processes are mostly invisible because of the over-
lyinglymphocytes.
The lymphocytes in the medulla differ slightly from
these in the cortex. The difference is mainly in the amount
of nuclear material present. The nuclear material in the
lymphocytes of the medulla is not as much as in the cortex,
thereby revealing a small amount of cytoplasm.
Three main types of tubules were numerous throughout
the medulla. First, there were tubules composed of true
cuboidal epithelial cells. These cells had a dense nucleus,
surrounded by an almost transparent cytoplasm. In some
tubules there seemed to be a secretion of some colloidal
material in the lumen. In others, the lumen was clear. The
cuboidal cells, which formed the tubule, were surrounded by
a nucleated basement membrane, which in turn was surrounded
by connective tissue and lymphocytes.
Second, there were tubules lined v/ith a low type cu
boidal cells (Plate VIII). These tubules seemed to have
assumed the function of blood vessels as there were red blood
cells in the lumen of the tubule. This tubule could not be
mistaken for an artery lined with phagocytic cells because
the elastic membrane, adventitia, and muscular layer, all
typical constituents of arteries, were lacking. Neither
PLATE VIII, Photomicrograph showing a
Tubule which has assumed the function
of a Blood Vessel. x970
BS - Blood Sinus
CE - Cuboidal Epithelium
RBC - Red Blood Cells
BM - Basement Membrane
T - Tubule
31
PLATE VIII
32
could they have been mistaken for a vein as the adventitia,
internal elastic membrane, and muscular layer, typical con
stituents of a vein, were lacking.
Third, there are tubules composed of low columnar,
or high cuboidal, cells. These cells had a dense nucleus
surrounded by an almost transparent cytoplasm. A basement
membrane surrounded the cells. This membrane, in turn, was
surrounded by connective tissue and lymphocytes. There was
no evidence of a secretion of any kind in the lumen, it be
ing perfectly clear (Plate IX-A and B).
A structure which resembled a thymic corpuscle
(Hassall*s corpuscle) was found in the medulla (Plate X).
The cells seemed to have formed concentric layers around the
lumen. The nuclei of the cells had become completely oblit
erated. Within the lumen there appeared to be some type of
secretion which contained connective tissue that resembled
fibers of some body fluid.
Surrounding the three types of tubules, described
above, are from one to forty or fifty eosinophilic cells.
There appeared to be two types of these cells. The first
type had a cytoplasm of red granules with two distinct, un
connected nuclei. This could very easily have been inter
preted as an eosinophilic myelocyte in mitosis. The second
type was an eosinophil leucocyte with a red granular
PLATES IX-A and B. Photomicrograph showing
a Tubule in the Paraganglionic Body. x400
CE - Cuboidal Epithelium
L - Lumen
BM - Basement Membrane
34
PLATE IX -A
CE
PLATE IX -B
PLATE X. Photomicrograph showing a Corpuscle'
like Structure in the Paraganglionic Body.
x900
Eo - Eosinophil
CSC - Crescent-shaped Cells
L - Lumen
36
fiSfi
L
o
PLATE X
37
cytoplasm, and a double, or bllobed, nucleus connected by an
isthmus.
The medulla gave the appearance of being a haemolymph
gland in that red blood cells were found, not only in the
sinuses but distributed in great numbers throughout the
medulla.
There was no stain found that was sufficient in itself.
Heindenhain*s Azan stain gave the best differentiation of the
various cell types, thus it was used most frequently. It was
from slides stained with this that most of the observations
of this investigation were made.
The hilus, through which all of the arteries, veins
and nerves enter or leave the body, is not distinguishable
by gross anatomical study, out is quite evident when the
sections are studied. This hilus was not mentioned by any
author in discussion of the aortic paraganglionic bodies.
PLATE XI. Photomicrograph showing an
Arteriole, Venule, and a Blood Sinus. x400
Ar - Arteriole
BS - Blood Sinus
Vn - Venule
39
I
PLATE XI
CHAPTER V
Summary
The lack of previous study on the aortic paraganglionic
bodies leaves a wide field open for investigation. This in
vestigation is confined to a histological consideration of
the bodies. It is hoped that this study v/ill provide a step
ping stone for future work on the bodies in the fields of
physiology, bichemistry, and bacteriology.
The investigation of the cell types revealed that the
bodies are composed of lymphocytes, eosinophils, cuboidal
epithelial cells, acidophils, phagocytic cells, and red blood
cells. There are probably other types present, but more
selective stains will De necessary before they can be demon
strated.
The most significant feature in the histological re
lationship of the cells is that the tubules of the cuboidal
epithelium and the eosinophils are singular in their relation
ship, since in every instance observed, the eosinophils v/ere
grouped around these tubules. Such a relationship would seem
to indicate that the body might be a site of leukopoiesis in
the case of infections by great numbers of bacteria. Such a
theory would have to be proved bactériologieally, or physiol
ogically.
The similarity in structure of the paraganglionic
41
bodies and the lymph glands make it apparent that these bodies
are some type of lymphatic body. The large numbers of phago
cytic cells in the arterioles and venules would seem to fur
ther indicate that the oodles were, in some way, connected
with the immunity of the animal to bacterial infection.
Such a supposition has been further substantiated when sev
eral of the animals whose paraganglionic bodies were removed
developed bacterial infections of the streptococcus type.
The control animals, however, whose body cavities were merely
opened and closed, did not in any instance develop an in
fection.
Since three types of tubules were present, it is quite
possible that they were developmental stages in the production
of the corpuscle-like structure. This development might
start with the high cuboidal cells, to the true cuboidal cells,
to the lovv cuboidal cells, thence to the corpuscle-like
structure, which in turn, degenerates and gives rise to the
fibrinous material present in the lumen.
Like the thymic corpuscle, this corpuscle may be re
lated in some way with the reproductive organs, especially
of the testes, since in operated male animals degenerative
changes take place whicn are visible on gross anatomical ex
amination of the testes. The testes are about two-tnirds
normal size, soft, and have changed color from a glistening
42
white to a light grayish white*
Upon sectioning and examining the testes of the oper
ated animals, it was found that spermatogenesis was halted at
the stage of the secondary spermatocyte. Examination of the
testes in animals where the body cavity was merely opened
and closed, or when one body only was removed, showed no sig
nificant change from the normal.
The possibility of some physiological function found
further corrobaration, when after the removal of one body,
the other body increased in size, therefore increasing its
function to maintain proper balance. There appeared to be
no significant change in the histological structure except
that each had increased its size in proportion to the increase
of the body.
If the body is connected with the haemolymph glands,
experimental bleeding of the animal should produce an in
crease in the haemopocitlc activity in response to the
bodies’ demands. There were large numbers of red blood cells
distributed throughout the medulla.
Most workers claim that there is partial retrogression
after the state of puberty has been reached. This would tend
to make the aortic, or lumbar, paraganglionic bodies primar
ily bodies of fetal and early childhood. Nothing discovered
in this investigation bears this out. In fact, the reverse
43
has been found to be true. In newly born and very young
animals, the bodies are less than five millimeters long.
By the time the animal reaches the age of six months, they
have attained their normal size of five to seven millimeters
and remain so until the death of the animal.
This study has been an attempt to establish some
basis for further study, especially in the fields of hist
ology, physiology, bacteriology, bichemistry, and embryol
ogy. Each of these fields is as yet uninvestigated. It
is hoped that this study may strike a spark in the minds of
future investigators, and eventually a complete study in
all the aforementioned fields will establish its function.
BIBLIOGRAPHY
Bibliography
Arey, L. B.
1957. Developmental Anatomy, 3rd, ed,
W, B. Saunders Co, Philla, pp, 443
Baker, John R,
1933. Cytologlcal Technique.
Methuen and Co, Ltd, London, pp. 64-74
Bensley, R, R, and S, H, Bensley
1938. Handbook of Histological and Cytologlcal
Technique, Univ. of Chicago Press,
pp. 73-102
Bodian, David
1936, Reduced Silver Method (Protargol)
Anatomical Record, 65, no, 1, pp. 84-97
Bremer, J, L,
1936, A Textbook of Histology, 5th, ed,
P. Blakiston*s Son and Co. Phila, pp. 355
Carlton, H, M.
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Asset Metadata

Creator Brandt, W. L. (author)

Core Title Some histological considerations of the aortic paraganglionic bodies (of Zuckerkandl) of the guinea pig

Contributor Digitized by ProQuest (provenance)

Degree Master of Arts

Publisher University of Southern California (original), University of Southern California. Libraries (digital)

Tag Biological Sciences,OAI-PMH Harvest

Format application/pdf (imt)

Language English

Permanent Link (DOI) https://doi.org/10.25549/usctheses-c39-273493

Unique identifier UC11314884

Identifier EP67151.pdf (filename),usctheses-c39-273493 (legacy record id)

Legacy Identifier EP67151.pdf

Dmrecord 273493

Document Type Thesis

Format application/pdf (imt)

Rights Brandt, W. L.

Type texts

Source University of Southern California (contributing entity), University of Southern California Dissertations and Theses (collection)

Access Conditions The author retains rights to his/her dissertation, thesis or other graduate work according to U.S. copyright law. Electronic access is being provided by the USC Libraries in agreement with the au...

Repository Name University of Southern California Digital Library

Repository Location USC Digital Library, University of Southern California, University Park Campus, Los Angeles, California 90089, USA

Linked assets

University of Southern California Dissertations and Theses