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The development of the suprarenal gland in the albino rat with a consideration of its possible relation to the origin of foetal movements.

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THEDEVELOPMENTOFTHESUPRARENALQLANL)
I N T H E ALBINO RAT, WITH A CONSIDERATION O F
I T S POSSIBLE RELATION TO T H E ORIGIN O F
FOETAL MOVEMENTS
DAVID S. PANKRATZ
College of Medicine, T h e University of Tennessee, and The W i s t a r Institute of
A n a t o m y and Biology
ELEVEN FIGURES
In this preliminary publication a study of the development
of the suprarenal gland is presented, from the time that the
gland first appears in embryos of higher vertebrates to the
end of foetal life. Special attention has been given to the
migration of sympatho-chromaffin cells and the development
of associated nerve fibers, since it was early noted by the
author that these processes immediately precede the appearanfie of foetal movements and may, therefore, be causally
related to the latter. In the present article attention will be
focused on the coincidence in development of the suprarenal
gland and foetal movements.
MATERIAL
Through the courtesy of the staff of The Wistar Institute
in Philadelphia, an extensive collection of serially sectioned
rat embryos was made available to the author. The material
also included some rats that Angulo ('28) had studied and
other series especially prepared by the institute f o r this
investigation.
The writer wishes here to express his appreciation and.
gratitude to the staff of The Wistar Institute of Anatomy and
Biology and especially to Dr. G . E. Coghill, who first suggested the problem and whose candid, kind, and mature
criticism has made this woik possible.
31
T H E ANATOMICAL RECORD, VOL. 49, NO. 1
32
DAVID S. PANKHATZ
OBSERVATIONS
The suprarenal glands in the albino rat appear first in the
thirteen-day-old foetus. I n the twelve-day-old foetus no
thickening of the epithelium in the dorsal wall of the coelomic
cavity can be seen (fig. 1). A slight thickening of the coelomic
epithelium medial to the cephalic portion of the mesonephric
fold marks the anlage of the suprarenal cortex. This process
is due to a proliferation of coelomic epithelium in a fold
which is continuous with the pleuroperitoiieal fold cephalically and with the genital fold caudally. This is described in
man by both Zuckerkandl ('12) and Wieman ('20). The
ABEREVIATTONS F O R A L L FIGURES
Ao., aorta
Ad.An., adrenal arilage
Ad.C., adrenal cortex
B.C., blood cells
C.C., cortical eells
C.V., central vcin
G.C., ganglion cells
L., liver
H.,mesentery
Ns.,mesonephros
M.T.,inesonepliric tubulcs
N . F . , nerve fibers
N.Npl., splanchnic nerve
P., pleuroperitoneal cushion
P.C., pleuroperitoneal cavity
P.Ep., peritoneal epithcliuin
S., stomach
Sp., spleen
Sup., suprarenal
s ~ p . L . ,suprarerial ligainent
Sy.C., sympatho-chromaffin eells (medulla)
Sy.C.R., sympatho-chromaffin ' ros4tte '
Sy.G., sympathetic ganglion
V., vertebra
rjl.
Fig. 1 Transverse section tlirongh a twelve-day-old rat foetus.
X 330.
33
SUPRARENAL-GLAND DEVELOPMENT I N T H E RAT
epithelial cells in this region become definitely larger, their
nuclei vesicular, and they appear to migrate into the mesenchyme dorsally (fig. 2). This process in the rat is much like
that described in the bird by Hays ( '14), in the pig by Whitehead ('03) and Weyman ('22), and in man by Zuckerkandl
( '12) and Fischel ( '29). Such a complex development of the
cortex as described in man by Keene and Hewer ('27) was
not observed in the rat.
Fig. 2
Transrerse section through a thirteen-day-old rat foetus.
X 330.
I n the fourteen-day and fifteen-day foetuses the thickening
of the epithelium and its dorsal migration were more apparent (figs. 3 and 4). I n these stages one could also see the
sympathetic trunk and its ganglia, the aorta, mesonephric
tubules, and stomach in their relation to the suprarenal
anlage. Growth at this time is rapid, as the figures indicate.
During the fifteenth t o the sixteenth days the cortex forms a
T H E I N A T O J I I C A L RECORD, VOL. 49, N O .
1
34
DAVID S. PAKKRATZ
marked oval mass which protrudes into the dorsal coelomic
cavity (figs. 4 and 6). This corresponds to the 25-mm. stage
of man described by Keene and Hewer ('27). At this stage
of development of the cortex, there is a 'breaking up' of the
cortex by small blood vessels, which, according to Wieman
('ZO), probably later form the central veins (fig. 4).
I n the sixteen-day f oetnses, shortly before motility was
observed (Swenson, '27), the cortex is very spongy, due to
Sy.
G. Ao.
I
I
Fig. 3
Transverse section through a fourteen-day-old r a t foetus.
X 200.
the numerous vascular spaces which later are invaded by
sympatho-chromaffin cells (fig. 6). About the sixteenth day,
small darkly staining cells migrate ventrally from the sympathetic ganglia toward the cortex. Soulie ('03) describes the
beginning of penetration of the cortex by the sympathochromaffin cells in a 15-mm. rat embryo and their reunion into
a central nucleus in a 25-mm. embryo.
Just before movements were first observed, masses of small
darkly staining cells can be seen on the medial surface of the
SUPRABENAL-GLAND DEVELOPMENT IN THE RAT
35
cortex. Sometimes one may find even a definite mass of these
cells in this position (fig. 8). At the time of first motility
small clumps of the migrating cells are found lying in the
vascular spaces of the medial part of the cortex. I n the defi-
Fig. 4
X 330.
A nearly longitudinal section through a fifteen-day-old rat foetus.
nitely motile foetuses (sixteen days) I often observed a
marked bundle of nerve fibers and ganglionic cells along their
course (figs. 6 and 6a).
Fischel ('29) says that the suprarenal gland in man remains
connected with the sympathetic chain by the processes of
36
DAVID S. PANKRATZ
these migrating cells which eventually form the medulla.
The processes on some of the large ganglion c,ells (G.C.,
fig. 6a), observed in only a few cases, appear to corroborate
his observations. Apparently there are two types of sympatho-chromaffin cells that migrate into the cortex besides the
comparatively f etv large ganglioilic cells. The small darkly
Fig.5
A section through the suprarenal cortex of a non-motile sixteen-day-
old r a t foetus, showing a mass of sympatho-chromaffin cells in a vascular space.
x
2000.
staining cells are the most numerous, especially in the early
stages of cell migration. Then there are larger and paler
cells with somewhat vesicular nuclei and cytoplasmic processes (fig. 6a). For a more detailed description, the reader
is referred to the article by Keene and Hewer ('27).
In all the rat series examined (about twenty-five series)
only a few large ganglionic cells were found in the medulla
SUPRARENAL-GLAND DEVELOPMENT I N THE BAT
37
of various ages up to birth (fig. 10). On the other hand, I
have often observed 'rosettes' of darkly staining cells in the
suprarenal gland of the actively motile rat foetuses. These
have been described in man by Keene and Hewer ( '27). Figure 9 shows such a 'rosette' with a bundle of nerve fibers
connected t o it.
P
Fig. 6 Transverse section through a motile sixteen-day-old rat foetus.
X 100.
This cell migration, which began during the sixteenth day,
continues during the following days, apparently till birth
(figs. 6, 6a, 7, and 8). I n all the foetuses of seventeen,
eighteen, and nineteen days definite nerve connections to the
gland could be seen (figs. 6,7, and 8). Mitsukuri ( '82) shows,
in figure 7, a 23-mm. rat embryo with nerve fibers passing into
the suprarenal mass. Soulie ( '03) gives a short description,
but does not show any figures f o r the rat.
38
DAVID S. PANKRATZ
I n the rat at birth a very definite nerve (N.SpZ., fig. 11)
connects the suprarenal gland with the sympathetic chain.
The medullary cells are found around and in the vascular
spaces, closely crowded against the cortical cells (fig. 11).
Fischel ('29) describes such a relation in the suprarenal in
man.
G.
Fig. 6a A section showing a higher magnification of the migrating cells of
scction shown in figurc 6. X 2000.
DISCUSSION
A survey has been made of the literature dealing with the
development of the suprarenal gland, the migration of sympatho-chromaffin cells, the first chromaffin reaction, and the
origin of foetal movements. Numerous writers, notably
Souli6 ('03) and Lutz and Case ( ' 2 5 ) , state that undoubtedly
the adrenal hormone is present and capable of functioning at
a very early age in the mammalian foetus. They also call
attention to Penger's ('12) observations which suggest that
SUPRARENAL-GLAND D E V E L O P M E N T I N T H E RAT
39
the suprarenal glands have a relatively more important r81e
in the embryo than in the adult. From the phylogenetic point
of view Gaskell ('19) points out the fact that in the lowest
vertebrates the function of the sympathetic nervous system
is largely taken over by a chromaffin system. Such facts give
credence to the view that in the embryos of higher vertebrates
Fig. 7 Transverse
(motile). X 100.
section of
a seventeen-day-and-five-hour-old rat
foetus
the suprarenal medulla may function before the sympathetic
nervous system.
Probably the most complete description of all phases of
investigation is available for the chick. Preyer ('85) was
the first investigator to study the embryonic movements of
the chick extensively. The first movements were noticed in
a five-day-old embryo and consisted in a bending of the back
from side t o side. Later, Clark and Clark ( '14), with more
refined methods, further studied the development of the
40
DAVID S . PANKRATZ
embryonic movements in the chick. Their findings corroborate Preyer's work. They describe the early stages as
consisting of passive movements of the head, tail, and extremities. They further observed that during the sixth and seventh day the bending of the body becomes more pronounced,
the tail contracts independently, the head nods, and the
Fig. 8 Transverse section of an eighteen-day-and-six-hour-oldrat foetus
(motile). X 330.
paddle-like extremities are moved inward and outward. From
the eighth to eleventh day, the movements continue to increase
in strength and variety.
In the same year, Hays ( '14) described the development of
the adrenal glands in birds, using the chick. He states that
the first evidence of any connection between the anlagen of
the prevertebral sympathetic plexuses and chromaffin substance is seen after 130 hours of incubation (5.4 days). After
SUPRARENAL-GLAND DEVELOPMENT IN THE RAT
41
168 hours (seven days) of incubation, the cells which are to
form the medulla of the gland are beginning to show some
differentiation. During the next twenty-four hours, or the
eighth day, the chromaffin cells within the glands have increased greatly in number, and most of the sympathetic cells
have disappeared from the surrounding mesenchyme. The
arrangement of the chromaffin cells undergoes a marked
change during the ninth day of incubation. At the same time
I Sy.C.R.
Fig. 9 A section showing a sympatho-chromaffin 'rosette' in an eighteen-dayand-six-hour-old rat foetus (motile). X 1500.
the chromaffin cells are found in small groups arranged
around the venous blood vessels. Furthermore, during the
period in which there is the greatest influx of cells from the
aiilagen of the prevertebral sympathetic plexuses there is also
the greatest activity in the development of the vascular systerns. Later, Lutz and Case ('25) and Okuda ('28) investigated the appearance of adrenalin in the chick by biological
and chromic reaction tests. Using the enucleated frog's eye,
Lutz and Case ('25) obtained the first positive adrenalin
reaction in seven-day-old chick embryos. Okuda ('28) used
42
DAVID S. PANKRATZ
Cannon’s intestinal method and the chromic-acid reaction.
He says:
. . . , in the chick embryo the suprarenal adrenalin appears on about
the eighth day of incubation, and chromic reaction of the gland cells
sets in a t nearly the same time. Their intimate relation which Ogata
(’16) pointed out, is thus confirmed. The adrenalin content of the
glands increases gradually with the progress of development in a n
incubator, but a more o r less marked increment is found on the
13th to 21st days.
Fig. 10 A section showing a ganglion cell in the suprarenal gland of an
eighteen day-and-four-hour-old rat foetus (motile). X 1500.
I n the chick we see that the positive tests for adrenalin
appeared a few days after the migration of the sympathochromaffin cells. On the basis of Miller’s ( ’26) and Weyman’s
( ’22) observations, possibly the actual beginning of the formation of adrenalin in the chick takes place somewhat earlier
SUPRARENAL-GLAND DEVELOPMENT I N T H E RAT
43
(fifth to sixth day) than has been described at the present.
From other investigations it appears that there is a production of adrenalin even during the later part of the sympathochromaffin cell migration.
Concerning the development of the suprarenal medulla and
the appearance of adrenalin in the mouse, &filler (’26) gives
C.
Fig. 11 Transverse section through a rat a t birth.
X 70,
a full account. According to her observations, the first
appearance of the chromaffin reaction takes place between the
latter part of fourteenth day and the first half of the fifteenth
day. “This,” she states, “is at a time when the future medullary cells are just beginning their penetration between cortical cells, leaving the sympathetic ganglion anlage, o r
44
DAVID S. PANKRATZ
sympatho-chromaffin mass, which is closely applied to the
suprarenal cortex at this time.’’ Her observations are based
on both the chromic reaction and Cannon’s intestinal method.
She also observed that the seventeen-day foetuses were highly
moti1e.l Inaba (’91) gives probably the earliest account of
the development of the suprarenal in the mouse. He gives
figures showing the final nerve connections of the gland,
Some observations have also been made on the rabbit.
According to Kolliker, the suprarenal bodies in the rabbit
appear first, on the twelfth or thirteenth day of gestation, as
masses of somewhat large round cells on each side of and
ventral to the aorta. On the fourteenth day the suprarenals
are already well marked (Mitsukuri, ’82). They consist of
a mass of cells with large nuclei, divided into indefinite cords
by blood capillaries which are already somewhat numerous.
Mitsukuri (’82) states that at this stage the connection of the
gland with the sympathetic chain, by means of a mass of cells,
is not so obvious as later on. But in the suprarenals of the
sixteen-day-old embryos great changes were observable.
Mitsukuri says that the medullary part (m., fig. 5) surrounded
for the most part by the cortical substance ( c ) can now be
clearly distinguished. I n the same figure he shows a branch
( a ,fig. 5) of the nervous mass ( n ) passing into the suprarenal and uniting with the medullary substance ( m ) . Minot
(’97) describes a similar development of the suprarenal in
the rabbit. Kohn (’03) noted the presence of the chromic
reaction as early as the period of beginning migration of the
future medullary cells into the suprarenal cortex. I n the
fourteen-day foetuses he was unable to detect any chromaffin
cells. I n fifteen-day rabbit foetuses he first observed the
chromaffin cells lying medially to the cortex; and in the
sixteen-day foetuses the reaction was very definite and the
sympatho-chromaffin cells are penetrating the cortex. This
penetration, according to Kohn (’03)’ is much more rapid in
the rabbit than in man. Soulie (’03) likewise says that about
the end of the fourteenth day one is able to see a mass of
Information on motility was given to the author by personal communication.
SUPRARENAL-GLAND DEVELOPMENT IN THE RAT
45
sympathetic ganglion cells in contact with the cortex. About
the fifteenth day the cortex receives a definite fasciculus of
fibers, but is not penetrated by the future chromaffin cells.
The first stage of motility-in the form of ventrolateral
flexion of the head and upper trunk (as described by Swenson
('27) in the rat and by Windle ('30) in the cat)-has been
first observed by the writer in fifteen-day and seventeen- to
twenty-hour rabbit foetuses.2
The observations on the cat seem to suggest a similar relation. Zuckerkandl ( 'la), speaking of the development of the
suprarenal in man and in the cat, says, "thus, for example, in
an 11mm. cat embryo the suprarenal ridge does not project
as much as in man." He further compares an 11-mm. human
embryo as being similar to a 10-mm. cat embryo in suprarenal
development. Kohn ( '03) first differentiated the sympathochromaffin cells in the ganglionated chain in a 12-mm. S.S.L.
cat embryo. Soulie ('03), in his description of the development of the suprarenal of the cat, says:
. . . . cet mas, d ' bbord homoghe, parait composit de nodules secondaires sup les embryons de 1 4 mill., et prend u n aspect nettement
reticule sur les embryons de 16 mill. C'est entre les stades 16 et de
18 mill. que commence la pitnittration des Blitments parasympathiques,
cette penetration est a peu pres achevite sur les embryons de 4 cent.
Recently, Windle ('30) has observed the first foetal movements in cat foetuses 17.5 and 18.5 mm. in length.
I n man the status of our information is in an unsettled condition. Souli6 ( '03), Keene and Hewer ( '27), and Fischel
('29) state that the cortex in man appears in a 5-to-6-mm.
embryo. Other investigators give slightly different ages.
Sonlii.5 ('03) says that the cortex develops rapidly, and in a
19-mm. embryo. According to Fischel ( 'as), the migration of
sympatho-chromaffin cells into the cortex takes place in 14to 20-mm. embryos. Keene and Hewer ('27) describe and
figure the beginning of such a migration in a 12-mm. embryo,
and show that it becomes prominent in twelve- to twenty-twoweek embryos and ceases at full term. I n the 25-mm. stage,
a
The data on these observations have not been published.
46
DAVID S. P A N B R A T Z
according to their observations, the suprarenal gland is defined and consists of true cortex, foetal cortex, and immigrating sympatho-chromaffin bundles. Jordan and Kindred ( '30)
also state that the migration begins at about the 19-mm. stage
and continues until after birth. Gradually the primordial
chromaffin cells surround the central vein, which appears at
about the 23-mm. stage, and ultimately differentiate into
characteristic chromaffin cells.
Various early investigators have failed to find any adrenalin in the suprarenal gland of human foetuses. On the other
hand, recent works show that the adrenalin begins in
relatively young foetuses. Hammar ('25) says that the
chromaffin cells appeared first in an embryo of 22.2-mm.
length, and that at that time the paraganglionic cords began
to be distinguishable. According to Keene and Hewer ( '27),
adrenalin appears at twelve weeks, and the chromaffin reaction appears at twenty-two weeks. Fenger ( '12) furthermore
says that in every case one finds adrenalin stronger relatively
in the embryo than in the adult.
The first motility in the human foetuses has been observed
as follows: Motility in the arm at 2 em. C.R. (Yanase, '07) ;
spontaneous motility in the arms and legs at 2.2 em. C.R.
(Strassmann, '00-'03) ; and oral reflex, with associated leg
movement, at 3.5 em. C.R. (Minkowski, '28). After this age,
other investigators (Bolaffio and Artom, '23-'24 ; Erbkam.
'37 ; and Krabbe, '12) have noted various types of early foetal
movements. For a more detailed discussion of these movements the reader is referred to Coghill's paper ('29). Due
to lack of material and uniform standard of age and lengths,
when dealing with human material, definite conclusions cannot be readily formulated.
The latest investigations in the vertebrates studied point
to a definite relation between the migration of sympathochromaffin cells, the chromic reaction, and the appearance of
adrenalin. Likewise, foetal movements, in the forms studied,
apparently begin at approximately the same stage of development. This striking parallelism has suggested a possible
SUPRARENAL-GLAND D E V E L O P M E N T I N THE RAT
47
relationship of cause and effect which is now being investigated in a number of different mammals.
Tabulated summary of the data and discussion
T H E MIGRATION OF
IYMPATHO-CHROMAFFIN
CELLS
Chick
Mouse
Rat
Rabbit
Cat
Man
T H E CHROMIC
1
T H E ADRENALIN
REACTION
8th day
130 hours; 5.4 days
14th-15th day
15th-16th day
15th-16th day
16-18 mm.
14-20 mm.
?
?
j ?
I
22 weeks
?
B
1 2 weeks
EARLY MOTILITY
5th day
Very motile
a t 17 days
16th day
15th-16th day
17.5 and 18.5 mm.
2-3.5 em.
CON.CLUSION
The suprarenal gland in the albino rat appears first in the
thirteen-day-old foetus. A slight thickening of the coelomic
epithelium medial to the cephalic portion of the mesonephros
marks the anlage of the suprarenal cortex. During the fifteenth to the sixteenth day the cortex forms an oval mass
which protrudes somewhat into the dorsal coelomic cavity.
About the sixteenth day, just before foetal movements begin,
cells migrate from sympathetic ganglia toward the cortex.
At the time (sixteen days or more) when foetal movements
have first been observed, a definite mass of sympathochromaffin cells is located among the medial cell cords of each
cortex. This cell migration continues during the seventeenth
and eighteenth days and possibly till birth. I n seventeen-,
eighteen-, and nineteen-day foetuses nerve fibers which connect the gland with the sympathetic chain were observed with
large ganglionic cells along their course. From the literature
on the different phases of the development of the suprarenal
gland and appearance of foetal movements in the chick,
mouse, rabbit, cat, and man, the author concludes that there
is in these animals a coincidence in development of gland and
foetal movements similar t o that which obtains in rat embryos. These facts have suggested that the beginning of
foetal movements may, perhaps, be facilitated by hormones
of the embryonic gland in some such way as adrenalin is
48
DAVID S. PANKRATZ
known to affect the threshold of skeletal muscle in adultsa hypothesis that is now being subjected to experimental
methods.
BIBLIOGRAPHY
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-4ND ARTOM 1923-1924
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~1912 a Jour. Biol. Chem., vol. 12, pp. 55-59.
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ALFRED 1929 Lehrbuch der Entwicklung des Menschen, S. 665-669.
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J. AUG. 1925 A quelle Bpoque de la vie foetale de l’homme apparaissent les premiers signes d’une activite endocrine? Upsala Lakaref.
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AND KINDRED 1930 A textbook of embryology.
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A N D E. E. HEWER 1927 Observations on the development
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KNUD 1912 Les reflexes chez le foetus. Rev. neural., T. 24, p. 434.
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R., A N D M. A. CASE 1925 The beginning of adrenal function in
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1926 The development of the epinephrine content of
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MINOT, C. S. 1897 Human embryology. New York.
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MITSUKURI,K. 1882 On the development of the suprarenal bodies in Mammalia. Q. J. M. S., vol. 22, pp. 17-29.
OGATA,T., AND A. AGATA 1916 Tokyo iji shinshi., nos. 1953, 1955, 1956.
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W. 1885 Specielle Physiologie des Embryo. Leipzig, Th. Grieben.
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STRASSMANN,
P. 1900-1903 Das Leben vor der Geburt. Samml. klin. Vort.,
n. F. (Gynak. H. 132, S. 947-96s). Leipzig.
SWENSON,
ENGELBREKT
AUGUST 1927 The development of movement of the
albino r a t before birth. Doctor’s thesis on file at the University of
Kansas.
WEYMAN,MORRIEF. 1922 Beginning and development of function in the suprarenal medulla of pig embryos. Anat. Rec., vol. 24, pp. 299-308.
WHITEHEAD,R. H. 1903 The histogenesis of the adrenal in the pig. Am. Jour.
Anat., vol. 2, pp. 349-360.
WIEMAN, H. L. 1920 Development of human suprarenal gland. Anat. Rec.,
vol. 19, pp. 268-279.
WINDLE,WILLIAM F. 1930 The earliest fetal moreriieiits in the cat correlated
with the neurofibrillar development of the spinal cord. Anat. Rec.,
rol. 45, p. 249.
YANASE,J. 1907 Beitrage zur Physiologie der peristaltischen Bewegungen des
embryonalen Darrws. Arch. f. d. ges. Physiol., Bd. 117, S. 345.
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E. 1912 I n Human Embryology, by Keibel and Mall, rol. 2,
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