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The relation of the estrogenic hormone to the formation and maintenance of corpora lutea in mature and immature rats.

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THE RELATION O F THE ESTROGENIC HORXONE TO
THE FORRIATION AND MAINTENANCE O F
CORPORA L U T E A I N MATURE AND
IMMATURE R A T S
C H A R L E S MERCKEL AiYD WARREX 0. KELSON
Dcpartincnt of Anatomy, W a y n e lJniuwsit?l College of Medicine, Detroit, ZClichigan
ONE T E S T FIGCJRE A h 9 T W O PLATES ( T E N FIGURES)
These studies were undertaken to determine whether corpora
lutea can be maintained in mature normal female rats by
injections of estrone when given in repeated small doses
a i d in a single large dose; and to determine whether estrone
exerts any luteinizing effect upon the ovaries of imiiiature
aninials.
The crucial point in these experiments lies in that phase
of the liypophyseal-gonadal relationship which is concerned
n7itli the luteinizatioxi of ovarian follicles. Evans in 1924
showed that anterior pituitary extracts produced a luteinizatioii of the ovarian follicles of both mature and immature
rats. I n 1926 Smith reported that removal of the hypophysis in
female r a t s resulted in cessation of growth and of estrous
cycles, and an atrophy of the accessory sexual organs. By
replaceinent therapy such changes could be prevented. I n the
ovaries of mature hypophysectomized animals follicles do not
develop, but such corpora lutea as a r e present a t the time of
operation may persist for an unusual length of time. The
ovaries of immature hypophysectomized rats contain no
corpora lutea, and the follicles are small aiid imniature.
The implantation of anterior pituitary tissue into immature
r a t s results in the development of a n excessive number of
follicles arid corpora lutea in the ovaries and in the occurrence
of pi-ecocious sexual maturity (Smith, '27, '30 ; Smith and
Engle, '27).
391
392
CHARLES M E R C H E L AND WARREN 0. NELSON
The physiological principles of the gonadal-hypophyseal
relationship have been summarized by Moore and Price ('30,
'32) as follows : (1) the sex hormones stimulate homologous
s e s characteristics, but have no effect upon heterologous
tissues, (2) sex hormones do not stimulate either homologous
o r heterologous gonads, (3) tlie gonadotropic secretions of the
hypophysis control the gametogenic and endocrine functions
of tlie gonads, (4) the gonadal hormones can suppress the
secretory activity of the hypophysis of either sex. Although
none of these four principles can be accepted now as entirely
correct, they have been exceedingly valuable in stimulating
studies which have furthered our understanding of the hypophyseal-gonadal relations.
Nelson ('35) found that tlie repeated daily injection of
cstrone into normal mature females produced changes which
a r e similar to the luteal effects found in pseudo-pregnancy
and are present f o r about 20 days. The ovaries of such anir i d s contain a normal number of large corpora lutea, tlic
estrous cycles a r e suppressed, the vagina is mucified, and
the lobules of the mammary gland undergo proliferation. The
hypopliysis is enlarged and its chromophilic cells become
degranulated. I f treatment is continued for a longer time
tlie corpora lutea degenerate, the effects of the corpus lnteum
hormone disappear, and the estrous reaction becomes coiitinuous. These results first reported by Scaglione ('30) liavc
been confirmed also by Hohlweg and Junkman ( ' 3 2 ) , Wolfe
('35, '36), and Sclye, Collip and Thompson ( '35). Furthermore, the same treatment in spayed and in hypophysectomizcd
animals produced a condition of sustained cstrus. This condition was reflected in the vagina, uterus, and manimarp
glands and castration changes were prevented in the pituitary
glands of the spayed animals. The ovaries of liypophysectomized r a t s which were treated with estrin showed the presence
of small, persistent corpora lutea. This picture is quite
similar to that seen in the ovaries of untreated hppophysectoniized rats.
FOR,MATION A N D R'IAINTENAIZ'CE O F CORPORA LUTEA
393
These studies indicate that estrin initiates the release of
luteinizing hormone from the hypophysis and as a consequence
maintains pre-existing corpora lutea in the ovaries of adult
rats.
It has been claimed by Hohlweg ( '34, ' 3 7 ) that the ovaries
of immature rats may be luteinized by the injection of large
doses of estrin. TVolfe ('36) showed that the injection of
estrin in young rats which had received, in addition, treatment with A.P.L. resulted in the production of larger corpora
lutea than in animals given A.P.L. only. This experiment
would indicate that estrin induces the release of luteinizing
hormone from the hypophysis.
The following sections deal with the various experiments
which have been carried out in the study of the relation of
estrogenic hormone to the development and function of the
corpus luteum in the rat.
A. E f f e c t of repeated small doses o f estrogen om adult normal
female rats
Table 1 represents data obtained from a series of female
rats treated with estrone for varying periods of time. All
injections were begun during estrus. It will be noted that all
animals given 40 R.U. daily showed an initial estrous period
of 2 to 3 days which was follo~ved,in animals permitted to
survive, by a diestrous period of 16 to 20 days. This in turn
was succeeded by a period of estrus which continued as long
as the animals were studied (fig. A, chart 2). A study of the
ovaries of animals during the long diestrous period showed
large corpora lutea. The onset of an estrous condition was
accompanied by the degeneration of these corpora lutea. Immediately after the close of the diestrous period the only
evidence of degeneration was a clumping of the lipoid drop* Grateful acknowledgment is made to Dr. Oliver Kamm of Parke, Davis &
Company and to Dr. Erwin Schwenk of t h e Schering Corporation f o r their
kindness i n furnishing the preparations of estrogenic hormone wliich were used
in these studies.
394
CHARLES MERCKEL AND W A RREN 0. NELSON
lets, but, within 7 days the corpora lutea were very small
and within 3 weeks they had undergone complete degeneration.
The gross and microscopic pictures in the uteri and vaginae
corresponded to the condition shown by the vaginal siiiears.
TABLE 1
E f f e c t of repeated doses of estrogen on mature intact females. Treatnaent zizitzated
during estrus rn all instances
__
-~_~________
RAT
NO.
R L 73
R L 74
RL 76
R L 77
R L 79
RL 80
R L 81
R L 82
R.L 83
R L 85
R L 86
R L 87
R L 89
R L 90
R L 93
R L 94
R L 97
R L 101
RL 102
RL 75
RL 78
KL 84
RL 91
R L 92
E L 95
TREATMENT
RU/day
40
40
40
40
40
40
40
40
40
40
40
40
40
40
40
40
40
40
40
20
20
20
20
20
20
Dngs
DA Y S ESTRUS
I>IMED. AFTEB
I1EG. INJECT.
11
17
17
12
10
15
8
6
6
20
22
20
21
28
27
40
38
43
67
2
3
2
2
3
3
2
2
3
2
2
3
15
14
12
17
14
17
4
15
13
4
15
4
9
I
3
2
3
2
3
9
DAYS ESTRUS
A F T E R DIEST.
PERIOD
DIit;&S
10
15
1.i
11
8
13
7
5
4
18
20
17
19
16
18
17
16
18
20
12
..
..
..
..
..
..
..
..
..
1
1
1
1
10
8
21
21
23
46
..
..
..
..
..
..
14
..
..
..
14
“,”,”;:~
Large
Large
Large
Large
Large
Large
Large
Medium
Medium
Large
Large
Large
Large
Sinall
Small
None
None
None
None
Large
None
None
Large
None
Medium
Evidence of degeiieratioii in microscopic sections.
Furthermore, niammary glands of all animals were well
developed. There would seem to be little doubt that the
administration of estrin had induced the functional and anatomical maintenance of the corpora lutea formed at the estrus
when treatment was begun, but did not induce the formation
of new corpora lutea since in no instance was an unusual
F O R M A T I O N A N D M A I N T E N A N C E O F CORPORA L U T E A
395
number of corpora present in an ovary. It is interesting t o
note that the response described in the above animals occurred
irregularly in instances where treatment was initiated during
diestrus. Thus, in only three of ten animals in which 40 R.U.
of estrone daily was injected for 20 days beginning during a
diestrous period, was there a maintenance of functional
corpora lutea. I n t.he remaining seven animals estrus was
continuous during the injection period.
I . NORMAL CYCLIC FEMALE
F-M
0
DAY
0
I
2 3 d
7
5 6 7
8
9
10 I I 12 17 14 I5 16 17 I8 I 9 20 21 2 2 29 24 25 26 27 U 29
30 71
/f l
2. INTACT MATURE ANIMALS TREATED WITH REPEATED DOSES OF ESTROGEN
:-M
D
0 I 2 3 4 5 6 7 8 9 10 I I / Z 13 /4 17 16 /7 /# ly 20 21 22 23 24 25 26 27 28 rq 70 31
3. INTACT RAT TREATED WITH ONE MASSIVE DOSE OF ESTROGEN
DAY
D
F-N
-\
0 I 2 3 4 5 6 7 8 9 I0 I / I2 I? 14 15 I6 17 /B ly W 2 / 22 27 24 25 2G 27 28 29 3031
4. SPAYED AND HYPOPHYSECTOM(ZE0 FEMALES TREATEO OAllY WITH ESTRONE
DAY
PM
DAY
0 I
2 3 4
E - ESTRUS
Fig. A
5
15 16 /7 18 3 20 21 22 23 24 25 26 27 ZS 27 3031
M - METESTRUS
D- DIESTRUS
Graphic representation of typical vaginal smear histories.
6 7 8 7 10 // I2 I3
14
- PROESTRUS
The sequence of events in the above experiments can be
reasonably explained on the basis of a release of luteinizing
hormone from the hypophysis under the influence of estrin.
This results in the maintenance of the corpora lutea and as a
consequence the production of progestin. The latter would
modify the action of the injected estrin on the vagina, uterus
and mammary glands. After about 3 weeks the corpora lutea
undergo degeneration and the modifying action of progestin is
absent. As a result the vaginal smear reveals a continuous
estrous state. The decline in function of the corpora lutea map
396
C H A R L E S M E R C K E L A N D WARREN 0. NELSON
be due to a restricted period of luteal-life (it is noteworthy
that the life of the corpora lutea of pregnancy and of lactation
is about 3 weeks) or it may be due to the exhaustion of the
supply of luteinizing hormone in the pituitary. The latter
possibility, viz., secretory exhaustion of the pituitary, is supported by the cytological picture in the hypophysis. The
basophilic cells are depleted of granules and the enlarged mitochondria and Golgi bodies have been said to be typical of cells
that have been actively engaged in the release of secretory
products.
The importance of the hypophysis in the train of events
reported in this experiment is shown by the study of the
results obtained in a series of hypophysectomized animals.
Eight female rats hypophysectomized during estrus mere injected with 40 R.U. of estrone daily f o r 20 days. The vaginal
smears showed continuous cornification during the entire
experimental period (fig. A, chart 4) and at the end of the
period corpora lutea were absent from the ovaries.
The ovaries, of course, occupy a position of importance
equal to that of the pituitary in the results obtained in this
experiment. If spayed animals are treated with estrin the
vaginal smear picture shows a continuous estrus (fig. A,
chart 4). Although estrin (affects) the hypophysis of these
animals in much the same fashion as it does normal females,
the released luteinizing hormone is without effect in the
absence of the ovaries.
B. E f e c t of a single large dose o f estrogen u p o n adult
.normal fentale rats
On the basis of the results obtained in the preceding experiment, it was of interest to observe the effect of a single injection of estrin in normal adult rats.
Ten adult normal rats were shown t o have normal estrous
cycles for 3 or 4 weeks before the initiation of treatment.
I n each case 400 to 500 R.U. of estradiol benzoate were ailministered subcutaneously in a single injection during the
period of estrus. Vaginal smears were continued and animals
FORMATION A N D MAINTENANCE OF CORPORA LUTEA
397
were sacrificed on the 7, 9, 11, 12, 13 and 14th days after
treatment. Ovaries, pituitaries, uteri, vaginae and mammary
glands were obtained for study. Three animals were continued under observation and spontaneously resumed cyclic
sexual activity 15 t o 17 days after treatment with 400 R.U.
of estrogen. These were rested about a month and then
were treated with 500 R.U. of estrogen in a single dose. Control animals were sacrificed during estrus.
The vaginal smears of intact mature animals given a single
injection of estrogen were of an estrous type during the first
2 to 4 days of treatment. Thereafter, the vaginal smears
were of the diestrous type until the fifteenth to seventeenth
days. Following this interval, in animals permitted to survivc,
the smears showed one or two cycles, then a continuous
diestrus for 9 to 12 days, and finally a resumption of cyclic
activity (fig. A, chart 3). We are unable t o expIain the occurrence of the secondary diestrous periods.
The ovaries presented no alteration in size. They all contained corpora lutea which were large or medium large and,
in contrast to corpora lutea in control ovaries, presented a
picture of functional activity. Furthermore, there were fewer
very old corpora lutea. This is in accord with Long and
Evans’ ( ’22) description of the various types of corpora lutea
which are found in the ovaries of sexually cyclic rats and
of the variations which are seen during pregnancy. I n the
latter condition, corpora lutea of pregnancy persist, those
formed previously are resorbed, and ovulation is suppressed.
S s a result only one conspicuous group of corpora lutea is
present.
The clumping of granules in the lutein cells increased with
the lapse of time after the single treatment. With incrcased
“clumping” there was a concomitant progressive nuclear
pyknosis, loss of cell outline, and shrinkage in size. The
ovarian follicles were small and few in number, although
in the rats which were kept under observation for the longer
periods the ovaries exhibited a slight increase in the number
and size of the developing follicles. The ovaries of these
398
CHABLES MEI:CICEL A N D WABBEN 0. NELSON
ariimals also showed vascular engorgement and edema
throughout their medullary regions.
The uteri of the treated animals wci'e, with one exception,
of medium size and of diestrous type. With greater closes,
the uterus acquired pseudo-pregnant characteristics. The
one rat whose estrous ,cycles were not suppi-essed exhibited
a n estrous type uterus. I n the test animals the uterine epithelium was of a low columiiar o r cuboidal type, without
vacuoles or mitoses and the glands were only slightly developed. The muscularis presented no significant deviation
from the iioriiial picture.
In all the acyclic rats, the vagina was of a diestrous,
mucifieil type, as iiiight have been expected from the vaginal
smear o n the day of sacrifice. The vaginae of the control and
cyclic rats were of an estrous type.
Proliferation of the ducts and not of the lohules and acini
was the conspicuous characteristic of the mammary glands
of animals treated with a single large dose of estrogenic
hoi-mone. The number of ducts was increased, their luiiiina
were patent, and end-huds, the forerunners of the alveoli,
were numerous. However, the connective tissue was always
precloniinant in amount over the parenchymatous tissue. I n
the cases of the two animals treated with the larger dose of
hormone and observed f o r the longest time, the iiianiniary
glands showed the best development seen in this series.
This experiment may be explained upon the same basis
a s the previous experiment, viz., the administration of estrone
to a normal adult rat during estrus induces the release of
luteinizing hormone from the hypophysis. The life and function of the corpora lutea are maintained for a longer period
than normal, but new luteinization of ovarian follicles is not
induced. The disturbance which is produced in the endocrine
system by a single dose of estrogen is not as severe as that
caused by repeated smaller doses. The secretion of follicle
stimulating hormone by the pituitary gland is only temporarily
paralyzed, and recovery from the suppression occurs before
the liheratioii of luteinizing hormone is fo1.ced to the stage
FORMATION A N D N A I N T E S A S C E O F C O R P O k 4 LUTEA
399
of exhaustion. Hypophyseal recovery occurred about 15 days
after treatment, whereas hypophyseal exhaustion did not take
place until about the twentieth day after the initiation of
treatment.
C. E f e c t o f estrogen upon immature fernale rats
Tlie finding that estrone would maintain previously formed
corpora lutea in the ovaries of adult rats suggested tlie
extension of these studies to the immature rat. This test
animal, whose ovaries are devoid of corpora lutea, is better
suited to a study of the effects of estrorie 011 the production
of new corpora lutea.
Twenty-two immature female rats 22 to 27 days of age and
with body weights ranging from 37 to 55 gni. were used in
this study. They were injected subcutaneously with single o r
repeated massive doses of estradiol-benzoate. The single
doses ranged from 200 t o 2000 R.U. Nultiple doses of 400
to 500 R.U. each, covered a total dose range of 2500 t o ,5000
R.U. Multiple doses were given a t 2-day intervals. After
intervals of 6 to 20 days following the initiation of treatiiieiit
the animals were sacrificed, and tissues were obtained f o r
study.
No corpora lutea could be found upon gross and microscopic
examination of the ovaries of either the experimental or the
control animals. However, many small follicles showing the
formation of antra were present. Witli increasing amounts
of estrin the size of the ovaries decreased, and with the largest
doses they were very small.
The vaginal smears, vaginae aiid uteri were of the estrous
type. The animals which were given repeated injections
showed a marked increase in uterine size. Frequently the
uterine lumina contained mucus and leucocytes, and were
slightly distended. The tall columnar cells were frequently
slightly vacuolated. Glands were only slightly developed.
The mammary glands of animals treated with single injections of estrone showed no stimulation. However, in rats
treated with multiple injections of estrone, there was a
400
CHARLES MERCKEL A N D WARREN 0. NELSON
stimulation of the duct system, but no proliferation of alveoli.
This development was in direct relation to increased dosage
and duration of treatment.
This experiment with immature rats fails completely to
confirm the reports of Hohlweg ('34) and of Hohlweg and
Chamorro ('37) concerning the induction, with estrone, of
TABLE
2
Immature aninulls treated with
OVARY
NO. O F
-
1
3
5
3
VAGINA
R'U.
A I1 J IA I.S
-
estrogens
I _ _ _ _ _ ~
Corpora
lutea
_.
-_____.__
Estrous
Estrous
Estrous
200
250
500
2000
UTERUS
Estrous
Estrous
Estrous
Estrous
Estrous
XAMMARY
Follicles
GLAND.
~~
Soiie
Sone
Xone
Kone
1
500
x
5
Estrous
Estrous
large
Kone
1
400 X
7
Estrous
Estrous
sl.
enlarged
Nollc
2
400 X 10
Estrous
Estrous
large
Soiie
5
Control
Estrous
Estrous
Kone
1
Control
Metestrous
Netestrous
Six large
recently
formed
Xuinerous
Sunierous
Xunierous
Sunieious
small
Numerous
siuall
Numerous
small
Iiumerous
small
Sumerous
sniall
Numerous
Not prolif.
Not prolif.
Not prolif.
S o t prolif.
XI. prolif.
F a i r prolif.
Fair prolif.
Kot prolif.
Not prolif.
small
luteinization in the ovaries of immature rats. This finding
is not surprising since it has been shown that the maintenance
of corpora lutea i n the adult female treated with estrone
was not attended by the foriliation of new corpora lutea.
DISCUSRON
Our results in adult r a t s treated with oestrone confirm the
reports of Selye, Collip and Thompson ( ' 3 5 ) , Wolfe ('35)
and Nelson ('35). The experiments in wliicli estrone was
injected into mature normal rats in repeated small doses
and single large doses demonstrate that release of luteinizing
FORMATION AND MAINTENANCE
OF CORPORA LUTEA
401
hormone from the hypophysis is stimulated and secretion of
follicle stimulating hormone is suppressed. F o r this reason,
functional corpora lutea may be maintained in the ovaries
f o r a longer time than in the normal cyclic female. Furthermore, it is note-worthy that only normal numbers of corpora
lutea were found. This indicated that new luteinization did
not occur and that only corpora lutea of the last ovulation were
affected. The subsequent production of progestin was associated with the suppression of estrous cycles and a diestrous
condition of the accessory sexual organs. This experiiiient
shows that the oestrone can indirectly stimulate the homologous gonad through the mediation of the pituitary gland, and
affirms the necessity f o r a modification of one of the major
points of the Moore and Price concept of the hypophyscalgonadal relationship.
The effect of a single large dose of estrogen was not as
marked as was the effect of repeated small doses. This was
indicated by the occurrence of estrous cycles rather than a
sustained estrus, and by a less marked degranulation of the
basophiles of the pituitary gland. I n all likelihood this was
due to the shorter period of effective action on the part of
the estrogenic hormone which was supplied by the single
treatment. As a result of this, both actions of the hormone
on the pituitary were curtailed, i.e., both the stimulus t o the
release of luteinizing hormone and the inhibition of the follicle
stimulating factor were less marked than in animals treated
continuously. Studies on the relation of excretion of the
estrogenic hormone to the recovery of the hypophysis from
its effects would be interesting.
In immature rats the formation of corpora lutea was not
induced although animals of various ages and weights and a
wide variety of dosages of hormone were employed. This is
in accord with the lack of formation of new corpora lutea
in the ovaries of adult rats under similar treatment. It should
be noted that this finding di-ffers directly with the reports
of Hohlweg and Chamorro ('37) and Hohlweg ('34) from
whose paper we quote:
402
CHARLES MERCKEL AND WARREN 0. N E L S O N
Follikelhormon in grossen Dosen hat eine ausgesprochen
luteinisierende Wirkung. So z. b. konnte schon 1TVoche nach
Injektion von 500 R.E.-12,500 internationalen Einheiten
Progynonbenzoat bei infantilen 40-50 g. schweren Rattenweibchen das auftreten von Corpora lutea beobachtet werden.
Although the rats used in his, and in the present experiments,
were comparable in weight, it should be noted that his animals
were 8 weeks old when treatment was begun, and were therefore verging on maturity; whereas our rats were but 3 to 4
weeks of age and were frankly immature. Although it was said
that no corpora lutea were found in the ovaries of their
control animals, it would seem likely that corpora lutea were
present in the ovaries of their r a ts in spite of, rather than
its a result of, the treatment.
That the ovaries of young r a t s are capable of responding
to stiniulation by gonadotropic hormone has been demonstrated by many workers. Furthermore, Wolfe ('36) lias
shown that the pituitaries of immature rats can release luteinizing hormone. Although luteinizing hormone may have been
released from the liypopliyses of our immature rats, that
possibility could not be demonstrated by our procedures.
The absence of mature follicles in the ovaries of the young
test animals would explain the lack of corpus luteum formation. The generally accepted mechanism for the development of corpora lutea includes the initial action of F S H
in the maturation of follicles a n d the subsequent action of
both P S H a n d L H in the production of corpora lutea. The
lack of either of these hormones is sufficient to prevent
luteinization. I t would seem probable that there was an
absence of available F S H in our animals. This suggestion
is supported by the fact that there was a decrease in the size
of the ovaries, a n observation which is explained on the basis
of a n inhibition of the production or release of F S H. I t
is significant in this regard that luteinization of the ovaries
could not be induccd i n either immature or mature rats by
the action of estrone. However, it should be emphasized oiice
more that the period of functional activity of previously
FORMATION AND MAIXTENANCE O F CORPORA LUTEA
403
formed corpora lutea was prolonged by the administration of
estrogenic hormone.
SUMMARY AND CONCLUSIONS
1. Daily injections of 40 R.U. of estrone into mature intact
female rats, begun during estrus, initiated a diestrous period
of about 3 weeks duration, which was followed by a vaginal
reaction of continuous estrus. These results plus those of
examination of the ovaries and pituitaries a r e explained on
the basis of a suppression of the follicle-stimulating factor
and a n increased release of luteinizing hormone. Although
new luteinization of ovarian follicles was not induced, corpora
lutea of ovulation were maintained for about 3 weeks. Following this, due, presumably, to a n exhaustion of the supply
of luteinizing hormone, the corpora lutea regressed and the
injected estrin exerted a direct estrogenic effect upon the
sex accessories.
2. A single injection of 400 R.U. of estrone into mature
intact female rats, during estrus, initiated a diestrous period
of about 2 weeks duration, which was succeeded by a return
of estrous cycles. Corpora lutea were maintained, but regressed progressively as time elapsed. New corpora lutea were
not formed. I n interpreting these effects it is supposed that
a single large dose of estrone temporarily suppresses the
follicle stimulating function of the anterior pituitary and
induced the release of some luteinizing hormone. With decline
in level of estrogenic hormone the hypophysis recovers its
cyclic control of the female reproductive system.
3. As much as 2000 R.U. of estrone injected into immature
female rats produced a n estrous response in the vaginal smear
and the accessory sex organs, but did not produce a diestrous
reaction. The ovaries were devoid of corpora lutea, follicles
were small, and with continued treatment the ovaries became
progressively smaller. This seems to indicate that the folliele
stimulating capacity of the pituitary was suppressed. Luteinizing hormone may have been released, but in the absence of
preformed corpora lutea in the ovaries of these immature
404
CHAIILES MEHCKEL AND WARREN 0. NELSOK
animals, there is no indication that it was effective in any
iiiaiiner upon the immature ovary.
L I T E R A T U R E CITED
EVANS,H. M. 1924 The function of the anterior liypopliysis. Harvey Lectures,
vol. 19, p. 212.
FOSTER,
M. A., R. C. FOSTER
AND F. L. HISAX- 1937 The interrelationship of
the pituitary sex hormones i n ovulation, corpus luteum formation and
corpus luteum secretiou in the liypophysectomized rabbit. Endocrinology, vol. 21, p. 249.
HOIIT.TVEG,
WALTER 1034 Veranderungen des Hypopli~seiivorderlappens und des
Ovariums iiach Behandlung mit grossen Dosen von Follikellior~none.
Klin. Wochnschr., vol. 13, p. 92.
HOHLTYEG,
W., AND A. C H ~ X O R R O
1937 Ueber die luteinisierende Wirkung des
Follikelhormons durch Beeinflussung der luteogenen IIypopliysenvordereappensekretion. Klin. Wochnschr., vol. 16, p. 196.
HOHLWEG,
W.,A N D K. J U N K M A N 1932 Die Hormonalnwvose Regulierung dcr
Function des Hypophysenvorderlapieus, Klin. Wochnsclir., vol. 11,
p. 321.
Lox(;, J. A., A N D H. M. EVANS 1922 The oestrous cycle in the r a t and its
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PLATES
PLATE 1
EXPLANATION OF FIGURES
1 Vagina of a n intact adult rat which received 40 R.U. oestrone daily f o r
8 days. The epithelium is mucified. X 82.
2 Vagina of an intact immature rat which received five injections of 500 R.U.
of estroiie on alternate days and was sacrificed on the tenth day. The epithelium
is prematurely cornified. X 100.
3 Ovary of a n intact adult rat treated with 40 R.U. of estrone daily for
8 days. Several large corpora lutea have been maintained. X 12.
4 Ovary of a n intact adult animal treated with a single dose of 500 R.U.
of estrone and sacrificed 11 days later. Corpora lutea persist but are diminished
in size. X 12.
5 Ovary of a n intact immature animal which received five injections of 500 R.U.
of estrone on alternate days. Corpora lutea are absent. X 12.
406
F O R M A T I O N AND M I U N T E X A N C E OF C O R P O R A LUTEA
CHAILLES M E R C K E L A N D W A R R E S 0
KELSOK
407
PLATE 1
PLATE 2
EXPLANATION OF FIGURES
6 Luteal cells from a corpus luteum of an intact adult female r a t treated
with 40 R.U. of estrone daily for 8 days. Cells are large, well shaped and well
granulated. X 384.
7 Luteal cells from a corpus luteum of an intact adult rat treated with a
Fingle dose of 300 R.U. of estrone and sacrificed 11 days later. Cells are degenerated ;
although grossly, corpora lntea are maintained. X 384.
8 Mammary gland of a n intact adult animal given one injection of 400 R.U.
of estrone, and sacrificed 14 days later. Only ducts and connective tissue are
present. X 54.
9 Mammary gland of a n intact adult animal injected daily for 10 days with
40 R.U. of estrone. Good proliferation of ducts and alveoli is present. X 54.
10 Mammary gland of a n intact immature rat which has received five injections
of 500 R.U. of estrone on alternate days. Ducts are present, there is a tendency
for end buds to form, but an alveolar system is not developed. >: 54.
408
FORNATION AND MAINTENANCE O F CORPORA LUTEA
CHAXIIES MLRCKCL A N D WARREN 0. &-ELSON
409
PLA4TE 2
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