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The effect of estrin and male hormone injected separately and simultaneously upon the smooth muscle and epithelium of the seminal vesicle in the albino rat.

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THE E F F E C T O F ESTRIN AND MALE HORMONE
I N J E C T E D SEPARATELY AND SIMULTANEOUSLY
UPON THE SMOOTH MUSCLE AND EPITHELIUM
O F T H E SEMINAL VESICLE I N THE ALBINO RAT
MILTON D. OVERIIOLSER AND WARREN 0. NELSON
Department of Anatomy, Uniuersity of Missouri, Columbia
T W O PLATES (EIGHTEEN FIGURES)
INTRODUCTION
Freud (’33) reported that estrin was a stimulator of smooth
muscle growth in the seminal vesicles of castrated immature
rats. The animals were castrated at 3 to 5 weeks of age
and hormone injections started 3 days later. Subcutaneous
injections were made twice a day for 4 days. On the day
following the last injections the animals were killed, both
seminal vesicles cut out in one piece and weighed.
Freud’s principal findings may be summarized as follows :
The optimum growth of the seminal vesicles was not obtained
by the male hormone alone; the effect of this hormone upon
smooth muscle tissue being only moderate, its main point of
attack being on the epithelium. Estrin caused growth of
the smooth muscle and either did not affect the epithelium
or induced degeneration. The combined effect of follicular
and male hormones upon the seminal vesicles was a mutual
reinforcement of each other’s actions. F o r this phenomenon
the term ‘pace-making’ was suggested.
In a preliminary publication we (Overholser and Nelson,
’34) reported that estrin stimulated the growth of smooth
muscle in the seminal vesicles of adult castrated rats.
247
THE ANATOMICAL RECORD, VOL. 62, NO. S
248
MILTON D. OVERHOLSER AND WARREN 0. NELSON
MATERIAL ,4ND METHODS
Adult male albino rats subjected to long standing and
recent castration and normal adult animals were injected with
20 rat units of estrin daily for 15 days. I n two animals the
injection period was 30 days, one of the animals receiving
40 r a t units daily.
The effect of estrin and male hormone given separately and
together (two levels of male hormone) was determined in
immature castrated rats. Estrin was also given to normal
immature animals. The injection period was 12 days, the
daily dose of estrin being 20 rat units and of male hormone
3 bird units (first level) and 2 bird units (second level). All
injections of hormones were made subcutaneously in oil
solution.
Control material consisted of uninjected castrated adult
and immature animals, and normal animals sacrificed at the
beginning and end of the injection period of the experimental
animals.
In some cases one seminal vesicle was removed, measured
and weighed before injections and compared with the remaining seminal vesicle a t the end of the injection period.
I n other cases the size and weight of both seminal vesicles
were taken at the end of the injection periods and compared
with other groups receiving different treatment and with the
control vesicles.
Both seminal vesicles were removed in one piece with a
small bridge of prostatic tissue and immediately measured
and weighed. The vesicles were then fixed in Bouin’s fluid,
cut at 10 p and stained with Nallorp’s triple stain for histological examination and comparison. Prostates and testes
were stained with Harris’ hemotoxylin after fixation in
Bouin’s fluid.
249
EFFECT OF SEX HORMONES ON SEMINAL VESICLE
PRESENTBTION OF DATA
1. Effect of estrirt
0%
serni.lzal vesides of long time castrated
awimaEs
Four rats were castrated at 2 months of age. Three months
later one seminal vesicle was removed from three of the animals, an unsuccessful attempt being made to secure a seminal
vesicle from the fourth. The seminal vesicles were all at an
extremely infantile level, measuring 3 x 1 mm.
The animals were then injected daily with 20 r a t units of
estrin (Thee1in)l for 15 days. At the end of' the injection
period it was found that the remaining seminal vesicles had
more than doubled in size (table 1).
TABLE 1
Effect of 80 rat units of estrin daily for 15 days on size of seminal vesicles in
animals castrated for 9 months
ANIMAL
SIZlC OF 8EEMINAL
VESICLES REMOVED
(BEFORE INJECTIONS
SIZE O F SEMINAL
VESICLE ABTEF*
TINJECTIONS
BODY WEIGHT AT
STAET OB
INJECTIONS
BODY WEIQHT
AFTER
INJEOTIONS
mm.
mm.
urn.
m.
214
3.0 X 1
217
3.0 X 1
2.5 X 1
6 X 3.5
6 X 3.5
7 x 3.5
6 X 3.0
6.25 X 3.37
160
160
160
157
159.2
155
155
142
141
148.2
Average
.. .. .. . .
I
2.83
x
1
Figure 1is a section of the seminal vesicle from animal 217
removed before the estrin injections and figure 2 is the second
seminal vesicle after the estrin injections. The marked
growth of the smooth muscle is very striking, and occurred in
all four of the animals. I n animals 214 and 216 there appeared to be a slight stimulation of the epithelium in addition
to the muscular growth. I n these two animals the epithelium
seemed t o be slightly higher as a result of the eshrin treatment, but no mitotic figures were f ound.
Three animals were castrated at 1month of age and allowed
t o go without treatment of any kind for 10 months. Two of
The Theclin used in these experiments was supplied through the kindness of
Dr. Oliver Kamm of Parke, Davis & Co.
250
MILTON D. OVERHOLSER AND WARIXEN 0. XELSON
the animals were then injected with estrin for 30 days, 40
and 20 rat units daily, respectively. The third animal served
as a control. At the end of the injection period all three
animals were sacrificed and the seminal vesicles measured
and sectioned (table 2). The seminal vesicles of the two injected animals were larger than those of the control. There
was marked growth of the smooth muscle with no effect on
the epithelium in the seminal vesicles of both injected animals.
Figure 3 shows the seminal vesicle from the control animal
205 and in figure 4 is seen the seminal vesicle from animal 204
which received 20 rat units of estrin daily for 30 days. The
increase in smooth muscle is very striking and occurred in a
slightly greater degree in animal 203 which received 40 rat
units of estrin daily for 30 days.
TABLE 2
E f e c t of cst& injections for 30 days on sin0 of seminal vesicles of rats
castrated for 10 month.7
AIVIXAL
DAILY DOSE OF ESTRIN
FOR 30 DAYS
R.U.
203
40
204
205
20 R.U.
Control
SIZE O F SEXINAL VESICLE
AFTER IXJECTIONS
BODY WEIGHT AFTER
INJECTIONS
mm.
Urn.
x
295
330
332
9
5.5
8 X 5.0
5 x 3.2
Effect of estrin. on. seminal vesicles of adult castrated
aNimals
Fifteen adult rats were castrated and one seminal vesicle
removed from each. Ten of the animals were then injected
daily with 20 rat units of estrin for 15 days. At the end of
2.
the injection period all fifteen animals were sacrificed and
the remaining seminal vesicles measured and sectioned.
Table 3 gives the data of the injected animals and table 4
of the controls.
In the ten injected animals castration changes occurred in
the epithelium, secretion practically disappeared and the
vesicles decreased in size. In seven of the ten animals a
noticeable increase in the thickness of the smooth muscle of
EFFECT OF SEX HORMONES
251
ON SEMINAL VESICLE
the seminal vesicle occurred. This can be seen by comparing
figures 5 and 6. I n the five uninjected animals the usual
castration changes occurred (fig.7), and no decrease or increase in the thickness of the musculature could be detected.
TABLE 3
Effect of 20 rat units of estrin daily for 15 days on size of seminal vesicles in
castrated adult animals
ANIMAL
55
56
57
58
59
60
61
62
63
64
Average
SIZE OF SEMINAL
VESICbE ZEMOVED
AT CASTRATIOY
RODY WEIQHT A T
BTABT O F
MJECTIONS
SIZE OF SXMINAD
VSSICl23 ABT'EB
ISJECPIONS
BODY WEIGHT
AFTER
INJECTIONS
mm.
mm.
Bm.
m.
16 x 6.0
14 X 5.0
15 X 6.0
17 X 8.0
12 x 4.5
18 X 8.0
16 X 8.0
14 X 7.0
17 X 8.0
19 x 9.0
15.8 X 6.95
15 X 5.0
11 x 4.5
13 X 5.0
14 X 4.0
9 x 3.5
12 X 6.0
10 X 6.0
10 X 4.0
13 X 5.0
13 X 7.0
12.0 X 5.0
165
170
150
165
160
170
180
160
185
185
169.0
162
162
152
162
153
155
156
150
165
174
159.1
-
T.4BLE 4
Control castrated adult
animals
'
ANIMAL
SIZE O F S E X I N A L VESICLE
W T E R 15 DAYS CASTBATION
mm.
gm.
gm.
50
51
52
53
54
10 X 4.0
11 X 5.0
10 x 3.5
12 X 4.0
11X 4.0
160
170
150
190
185
160
190
151
187
162
Average
10.8 X 4.41
171
170
BODY WEIQHT AT
CABTRATION
~
BODY WEIGHT AFTER
15 DAYS CABTRATION
It was noted, however, that the castration changes in the
epithelium were more marked than those in the animals that
received estrin injections. The difference is not great but
was noted independently by two observers and occurred in
all ten injected animals.
252
MILTON
n.
OVERHOLSER
3. E f f e c t of estrin 0%
AND WARREN 0. NELSON
seminal vesicles of iiorrnal adult alzimals
One seminal vesicle was removed from five normal adult
rats. The animals were then injected daily with 20 rat units
of estrin for 15 days and then the remaining seminal vesicles
measured and sectioned (table 5).
I n all five animals there was a noticeable increase in the
amount of the smooth muscle in the seminal vesicle and a
marked suppression of the epithelium and secretion following the estrin injections (fig. 8). The marked atrophy of the
epithelium as an indirect result of the estrin is clearly shown.
TABLE 5
Ereat of 20 rat units of estrin daily for 15 days
on
size of seminal vesicles in
normd adult rats
SIZE OF SEMINAI,
VISICLE REMOVED
BEIWRE INJECTIONS
ALNlMAL
mm.
12 x 7
15 X 9
15 X 7
10 x 7
12 X 6
65
66
67
68
69
Average
4.
SIZE OF SEMINAL
VESICLES AFTER
INJECTIONS
I
12.8 X 7.2
mm.
12 X
15 X
14 X
14 X
12 X
1
5.0
5.0
5.5
4.5
5.0
13.4 X 5
RODY WEIGHT A T
I
~
cm.
147
175
155
180
160
163.4
~
~
BaDy
AFTEE I S J l C T I O N B
~
~
N
s
gm.
140
162
165
171
145
'
i
156.6
Comparisoi$ of the effects of male hormone a i d estrin on.
the seminal vesicles of immature castrated animals
Ten immature rats were castrated at 36 days of age and
injections started the following day. Five of these animals
received daily injections of 3 bird units of male hormone2 for
12 days, and five received 20 rat units of estrin ( A m n i ~ t i n ) ~
daily for 12 days. The animals were sacrificed the day after
the last injection. The data from these two series are shown
in table 6. Table 7 gives seminal vesicles sizes and weights
Speeial measures were taken t o free the male hormone used in these experiments
from estrin. We are indebted to Dr. T. F. Gallagher of The University of
Chicago for this material.
aDr. J. A. Morrell of E. R. Squibb and Sons very kindly furnished us with the
Amniotin employed in this work.
253
EFFECT O F SEX HORMONES ON SEMINAL VESICLE
of sacrificed immature rats and uninjected castrated and
normal animals sacrificed at same ages as those receiving
injections.
It is seen that the estrin injections caused no growth of
the seminal vesicle (table 6 B ; fig. 9) when compared with the
normal controls sacrificed a t the beginning of the injection
period (table 7 A) and with the uninjected normal controls
(table 7 C). However, when compared with the uninjected
TABLE 6
Comparivon of the effects of male hormone and estrin on the size and weight o f
the seminal vesicles i n immature catrated rats. Animals injected
daily for 12 days
1 1
OASTRATIOIN
BODY WEIUHT
C*S-TION
AT
1
j
BODY WEIGHT
Ih’lECTIoJS
AFTER
1
SIZE O F SEMINAL
v
~
~
1
WEIUHT OB
B O T ~9.~MINAL
VESICLES
~
~ AFTEB
~
INJECTIONS
T
~
~
A. Three bird units of male hormone daily
days
126
127
128
129
142
36
36
36
36
36
Average
130
131
137
143
144
36
36
36
36
36
Average
gm.
m.
35
45
45
43
32
40
___---
57
72
72
68
54
64.6
mm.
8.0 x
8.5 X
9.0 X
8.0 X
-8.0 X
8.3 X
mg.
32.6
38.2
32.2
26.4
27.2
31.3
3.5
3.5
4.0
4.0
3.5
3.7
I
_
_
_
_
:i
I
38
33
29
37.6
1I
1
1
56
64
57
48
35
52
j
6.0 X 2.0
6.5 X 2.0
6.0 X 2.5
5.5 x 2.0
4.5 x 2.0
5.7 x 2.1
i
8.4
9.0
8.2
7.3
6.4
7.86
castrate controls (table 7 B ) it is seen that the estrin injections have caused a slight growth of the seminal vesicles due
to stimulation of smooth muscle growth as the epithelium was
unaffected.
I n the animals receiving the male hormone (table 6 A ; fig.
10) there was a normal increase in the size and weight of the
seminal vesicles when compared with the uninjected normal
controls (table 7 C).
254
MILTON D. OVERHOLSER
AND WARREN 0. NELSON
TABLE
7
Control anim1.s
A . Sacrificed immature normal animals, 39 to 99 days old
I
AGE
ANIMAL
WEIQRT OF
days
79
80
81
82
83
123
124
125
146
147
148
VESICLES
mm.
gna.
31
31
31
34
34
32
30
29
39
39
---39
Average
1
BOTH SEMINAL
31ZE OF SBHIINAL VESICLE
BODY WEIQHT
35
39
33
40
43
54
54
50
52
43
31
43.0
1
3.5
3.7
3.5
3.7
3.5
5.0 X
5.0 X
4.5 x
5.7 X
5.0 X
5.0 X
4.37 X
I
I
1
nag.
x 2.0
x 2.0
x 2.0
x 2.2
x 2.0
AQE AT
BODY WBIQRT
AGE AT
BODY WEIQRT
CASTRATION AT' CASTRATION SACRIFICE AT RACRIIrICE
7.3
8.0
7.2
10.3
10.3
14.0
10.1
8.1
13.3
9.3
9.4___
I
.
_
I
9.75
2.5
2.0
2.0
2.5
2.2
2.2
2.14
'IZE OF
B
~~~~~~
I
.Bm.
135
136
141
36
36
36
Average
43
31
29
1
days
50
50
50
34.3
gm.
53
41
40
44.6
mm.
mg.
X
X
X
X
8.3
5.4
7.O
6.9
5.0
3.5
4.5
4.33
1.7
1.7
1.5
1.63
SIZE OF
ANIYAb
AQE
dapr
153
39
154
39
155
39
39
156
________
Average
o
-
BODY WEIURT
____
m.
38
36
36
35
S:gt&~~&~~~~~ "v",",:,",","
aarrs
53
53
53
53
gm.
67
67
67
65
66.5
mn.
6.5 x 3.0
7.5 X 3.0
8.0 X 3.0
7.5 x 3.5
7.37 X 3.12
WBIQHT OF
BOTI~,",",fZA&
mg.
18.3
31.0
33.3
40.0
30.65
_
_
_
_
I
_
36.2
I
I
~
255
EFFECT OF SEX HORMONES ON SEMINAL VESICLE
5.
Efect of combinatiolz of irtjectiolzs of male hormone alzd
estriw on the seminal vesicles of immature
castrated n n i n d s
Nine immature rats were castrated, seven being 36 days old
and two 39 days of age. Five of the animals received daily
injections of 20 rat units of estrin and 3 bird units of male
hormone and four were given daily injections of 20 rat units
TABLE 8
Erect of combination of injections of male hormone a d estrin on the size and
weight of the seminal vesicles of immature castrated rats. A n i d s
injected daily f o r 18 days
A. Twenty rat units of estrin and 3 birds units of male hormone daily
132
133
134
145
138
139
140
151
159
1
1
1
day8
m.
36
36
36
36
36
Average
40
43
38
34
27
36.4
36
36
39
39 Average
33
31
41
36
35.2
mm.
gm.
59
64
47
45
- 45
52
49
42
59
65
53.7
mg.
11.0 X
9.5 X
10.0 X
9.5 X
10.5 X
5.0
4.0
5.0
4.0
4.5
61.3
57.4
66.0
54.4
67.4
10.0 x
9.5 X
10.0 x
8.5 X
9.5 X
4.5
4.0
4.5
4.0
4.25
66.2
55.1
44.4
36.0
50.42
_-
of estrin and 2 bird units of male hormone. The animals
were injeated for 12 days and sacrificed the day after the
last injection (table 8).
By comparing table 8 with table 6 it is seen that by injecting estrin and male hormone together the growth of the senima1 vesicle was much greater than when male hormone alone
was injected. I n the former animals the seminal vesicle
weights are almost twice that of those receiving only male
hormone. As the dose of male hormone was the same in each
256
MILTON D. OVERHOLSER AND WARREN 0. NELSON
group of animals and as estrin alone caused no growth of
the seminal vesicles (table 6 B) the result appears to be
due to a synergistic action of estrin and male hormone. I n
table 8 B it is seen that when the male hormone dosage was
reduced to 2 bird units the size and weight of the seminal
vesicles were less than those receiving 3 bird units, but still
greater than those receiving 3 bird units of male hormone
without the estrin.
A section of the seminal vesicle from an animal receiving
simultaneous injections of estrin and male hormone is shown
in figure 11. By comparing figure 11with the seminal vesicles
from the animals receiving estrin alone (fig. 9) and male
hormone alone (fig. 10) the striking synergistic action of the
two hormones administered together is clearly shown.
A study of the epithelium of the injected animals and controls reveals further differences. I n the normal controls
sacrificed at 29 to 39 days of age the seminal vesicles showed
a low columnar epithelium with no secretion granules (fig.
12). Seminal vesicles of normal controls at 53 days have
numerous secretion granules, many of which have halos (fig.
14). In our experience secretion granules appear in seminal
vesicles weighing about 25 mg. a t the age of about 45 days.
F o r a full account of the relation of the secretion granules
to seminal vesicle function the reader is referred to the paper
by Moore, Hughes and Gallagher ('30). In the uninjected
castrated controls sacrificed a t 50 days of age (fig. 13) the
epithelium is quite comparable to that of normal controls
sacrificed at 39 days (fig. 12). Castration has prevented the
normal developmental changes but has not resulted in degenerative changes since appreciable amounts of male hormone apparently are not secreted in rats of this age. Estrin
did not stimulate the epithelium in the injected castrates
(fig. 15)' the seminal vesicles of these animals resembling
those of the 39-day controls. The seminal vesicles of the
castrates receiving male hormone alone (fig. 16) presented a
picture approaching that of the 53-day normal controls (fig.
14). The epithelium is medium columnar with many mitoses.
257
EFFECT OF SEX HORMONES ON SEMINAL VESICLE
Secretion granules are fairly numerous, but few have halos.
I n animals receiving simultaneous injections of male hormone
and estrin the epithelium shows considerable stimulation (fig.
17). The lobules are greatly extended and the cells are verging on high columnar. Mitoses and secretion granules with
halos are numerous. There seems to be every indication of a
synergistic action of estrin and male hormone upon the
seminal vesicle epithelium.
TABLE 9
Effect of injeations of eslrin on the size and weight of the seminal ve&cZes of
immature normal rats. Anzi.mal8 hjected daily with
20 rat units of estrin for 12 dugs
ANIMAL
1 1
AGEAT
START OF
INSECTIONS
I
BODYWEIGHT
AT 8l'ART OF
INJPCTIONS
BODY WEIGHT
APPER
INJECTIONS
W R I G H T OF BOTH
s
~
s
~
lNJECT1oNS
- _____days
149
150
152
157
158
39
39
39
gm.
45
41
45
35
m.
79
61
79
68
70
71.4
A b VESICLES
&S E X~I NAFTER
~ & ~ ~
INJECTIONS
-I___
mm.
8.5 X 4.0
7.0 X 3.3
8.0 X 4.5
6.5 X 3.5
8
.5 X 3.5
___.__
7.7 X 3.76
mg.
36.3
20.2
44.0
25.0
33.3
31.76
E f e c t of estrirt on the seminal vesicles of qzormal inmature
animals
Daily injections of 20 rat units of estrin were begun in five
normal animals at 39 days of age and continued for 12 days,
6.
the animals being sacrificed the day after the last injection
(table 9). By comparing table 9 with the normal uninjected
control animals (table 7 C) it is seen that the estrin injections
have caused no increase in the size or weight of the seminal
vesicles over the normal growth.
The seminal vesicle epithelium of the estrin injected normal
animals (fig. 18) is only slightly lower than that of uninjected normal controls (fig. 14). It is apparent that estrin
has not suppressed the epithelium in these immature animals
to the same degree that it did in the adult series.
258
MILTON D. OVERHOLSER AND WABREN 0. NELSON
DISCUSSION
The experiments in which estrin was injected in adult animals of long standing castration definitely show that growth
of smooth muscle of the seminal vesicle was stimulated (figs.
2, 4). There was apparently a slight stimulation of the epithelium in two of the animals.
The results of estrin injections in adult animals recently
castrated are less striking. I n seven of the ten animals a
slight increase in the thickness of the muscle was noticed as
compared with the vesicles removed before the injections
(fig. 6). It was also observed that usual castration changes
in the epithelium were not quite as marked as in the controls
(compare fig. 6 with fig. 7). Moore and Price ('32) reported
that estrin injections in ten adult castrated rats resulted in
slightly less degenerate epithelium in two animals as compared with uninjected castrated rats. They stated that there
is the barest suggestion that estrin delays the appearance of
castration changes in the epithelium of the seminal vesicle
and prostate but concluded that the difference is so slight it
was to be judged insignificant. I n the present experiment the
difference occurred in all ten of the animals and was noticed
independently by two observers. It may be due to a general
nutritional factor caused by a richer blood supply to the
seminal vesicles as a result of the estrin injections.
Estrin injections in the normal adult animals produced a
slight noticeable increase in the thickness of the muscle and
a marked suppression of the epithelium and secretion (fig.8).
The estrin indirectly suppressed the epithelium by suppressing the gonad stimulating hormone of the anterior pituitary
which resulted in a lessened secretion of the male hormone
from the testis and therefore a degeneration of the epithelium
of the seminal vesicle. This mechanism was shown by Moore
and Price ('32). We have studied the prostates and testes
of our injected animals, and although we are not reporting
the details of that study it should be stated that the findings
have confirmed those of Moore and Price.
EFFECT O F SEX HORMONES ON SEMINdL VESICLE
259
I n immature, castrated animals estrin caused a slight stimulation of smooth muscle growth. when compared with the
uninjected castrate controls. This does not completely correspond with the results of Freud ('33) who obtained marked
smooth muscle growth in similar experiments. There was no
stimulation of the epithelium as a result of the estrin (fig. 15).
Although areas of degenerating and sloughing epithelium
were present we attach no significance to this as it also occurred in the normal controls and in the uninjected castrate
controls. This is in opposition to the statement of David,
Freud and de Jongh ( '34) that estrin damages the epithelium
of the seminal vesicles in castrated animals. I n fact our
experiments on long time castrated animals give an indication
that estrin has a slight stimulatory effect on the seminal
vesicle epithelium.
Three bird units of male hormone daily in the immature
castrates maintained normal growth of the seminal vesicles as
to size and weight when compared with the normal control
animals (tables 6 A and 7 C) . The histological picture was, in
general, the same as that of the normal control. I n both cases
areas of sloughing epithelium were found.
I n the animals receiving simultaneous injections of male
hormone and estrin at the same dosage as given separately,
a very striking synergistic effect was obtained (table 8; compart With table 6). A second group received the same dosage
of estrin but 2 bird units instead of 3 bird units of male hormone. I n the first group the average weight of the seminal
vesicles was practically double those of the animals receiving
only the male hormone. The average size was also increased
from 8.3 x 3.7 mm. to 10.1 x 4.5 mm. I n the second group
the same striking increase in weight and size occurred, although the increase was somewhat less due to the fact that
only 2 bird units of male hormone were given.
The histological picture (figs. 11, 17) of the estrin-male
hormone injected animals showed that the epithelium had been
stimulated to a greater extent than when male hormone was
given alone, The muscle was also distinctly increased. These
260
MILTON D. OVERHOLSER AND WARREN 0. NELSON
findings therefore confirm Freud’s contention that the simultaneous administration of estrin and male hormone results in
a mutual reinforcement of each other’s actions. We do not
consider his term ‘pace-making’ for this phenomenon a
desirable one.
The stimulation of smooth muscle growth obtained in these
experiments we believe is due mainly to an hypertrophy of
the fibers. Hyperplasia may have occurred but we have been
unable to discern it in our preparations.
I n the injection of estrin in immature normal animals (table
9) the results were negative. The size and weight of the
seminal vesicles were the same as in the normal controls
(table 7 C). The histological picture (fig. 18) was the same
as that in the normal control (fig. 14). a r e a s of sloughing
epithelium were encountered but we attach no significance to
this as the same thing was seen in the controls and even in
those receiving male hormone.
I n the present regard it is of interest to note that castrated
adult male rats injected with about three times the amount of
male hormone (15 B.U. daily) required to maintain an essentially normal picture in the epithelium have seminal
vesicles which weigh considerably less than control glands.
Such control tissue is represented by seminal vesicles removed a t the time of castration. Although we do not have
sufficient data on this subject to warrant conclusions it seems
probable that failure to maintain normal weight of the
seminal vesicles by male hormone alone may be corrected by
the simultaneous administration of estrin. This synergistic
effect would appear to be mainly related to the extra-epithelial tissues, but the epithelium too, may be stimulated by the
addition of estrin. Hansen (’33) has reported that 7 bird
units of male hormone daily will maintain a normal epithelial
picture except that the cells fail to maintain their normal
height. We have observed the same condition in males receiving 10 bird units daily. The results reported in this
paper indicate that this failure to maintain an entirely normal
epithelium may be due to the lack of normal amounts of
estrin acting in conjunction with male hormone.
EFFECT OF SEX HORMONES ON SEMINAL VESICLE
261
We believe that the conception of estrin acting only on
female structures or homologues is no longer tenable and that
instead in estrin we are dealing with a principle which exerts
a more widespread influence than has been commonly supposed. There appears to be at least one factor which presents an obstacle to the analysis of the exact parts played by
the male hormone and estrin in their relation to the male
sex-accessories. This factor is the possibility that there is
a conversion of one hormone into the other within the body.
Support of this concept is found in the recent work of Ruzicka,
Goldberg and Briingger ( '34 a, b). Their work has demonstrated that the two hormones seem to be structurally very
similar.
SUMMARY AND CONCLTJSIONS
1. Daily injections of 20 rat units of estrin for 15 and 30
days in adult rats (5 and 11 months old) castrated at 1 o r 2
months of age caused a definite growth of the smooth muscle
of the seminal vesicle.
2. Daily injections of 20 rat units of estrin for 15 days in
adult rats recently castrated caused a slight stimulation of
smooth muscle growth and a questionable delay in the typical
castration changes in the epithelium of the seminal vesicle.
3. Daily injections of 20 rat units of estrin for 15 days in
normal adult rats caused a slight stimulation of smooth
muscle growth and an indirect suppression of the epithelium
in the seminal vesicle.
4. Daily injections of 20 rat units of estrin for 12 days in
immature castrated rats resulted in a slight stimulation of
smooth muscle growth in the seminal vesicle when compared
with uninjected castrated controls.
5 . Daily injections of 3 bird units of male hormone f o r 12
days in immature castrated rats maintained the normal
growth of smooth muscle and epithelium of the seminal
vesicle.
6. Simultaneous daily injections of 20 rat units of estrin
and 3 bird units of male hormone for 12 days resulted in a
doubling of the normal weight increase of the seminal vesicle.
262
MILTON D. OVERHOLSER AND WARREN 0. NELSON
T,here was stimulation of smooth muscle growth and epithelial
differentiation beyond that obtained in rats receiving the hormones separately. This is apparently due to a synergistic
action of the two hormones.
7. Daily injections of 20 r a t units of estrin for 12 days in
normal immature animals had no apparent effect on the
smooth muscle or epithelium of the seminal vesicles.
It is concluded that estrin stimulates smooth muscle
hypertrophy in the r a t seminal vesicle. There is evidence
also that it has a slight direct stimulating action on the
seminal vesicle epithelium in castrated animals. Estrin and
male hormone act synergistically upon both the smooth
muscle and epithelium when they are administered simultaneously.
LITERATURE CTTED
DAVID,K., J. FREUD
AND S. E. DE JONGH 1934 Conditions of hypertrophy of
seminal vesicles i n rats. 11. The effect of derivatives of oestrone
(menformon). Bioehem. J., vol. 25, pp. 1360-1367.
FREUD,
D. 1933 Conditions of hypertrophy of the seminal vesicles in rats.
Biochem. J., vol. 27, pp. 1438-1450.
HANSEN,J. R. 1933 Rat seminal vesicles and prostate glands a s quantivative
indicators of testicular hormones. Endocrinol., vof. 17, pp. 163-179.
MOORE,
C. R., W. HUGHESA h 3 T. F. GAUAGRER 1930 Rat semiual vesicle
eytology as a testis-hormone indicator and the prevention of castration changes by testis-extract injection. Am. 5. Anat., vol. 45,
pp. 109-135.
MOORE,C. R., AND D. PRICE1932 Gonad hormone functions, and the reciprocal
influence between gonads and hypophysis with its bearing on the
problem of sex-hormone antagonism. Am. J. Anat., vol. 50, pp. 13-71.
RUZICKA,L., M. W. GOLDRERG
AND H. BRENGGER
1934a Zur Kenntnis der
Sexualhormone. I. o b e r die Gewinnung von 3-Crhlor- und 3 0x7atioallocholanon- (17). Synthese einer Verbindung von den Eigensehaften des Testikelhormons. Helvetica Chimica Aeta, vol. 17, pp.
1389-1394.
RUZICXA,
L.,&f. w. GOLDBERG., JULES MEPER, H. BRCNGGEP. AKD E. EICHENRERGER
1934 b Z u r Kenntnis der Sexualhormone. IT. Uber die Synthese des
Testikelhormons (Androsteron) und Stereoisomerer desselben durch
Abbau hydrierter Sterine. HeIvetiea Chimica Acta, vol. 17, pp. 13951406.
OVERHOI,SER,
M. D., AND W. 0. NELSON1934 The effect of estrin administration
on the seminal vesicle musculature of castrated rats. Anat. Rec.
(supplement), vol. 58, pp. 31-32.
PLATES
PLATE 1
EXPLANATION
or
FIGURES
1 Sciiiiiiill r ~ s k l efin111 :idult int 217 ~cmorrd3 niniiths a f t e r castration. X 12.
3 Scwnirl scminal vesicle from rat 217 after jiijectioiis of 20 rat mrits of
wtiiii daily f o r 1.5 days. Marked growth o f the smooth niirsclc wit11 no effect ozr
t h e epithelium. x 12.
3 Seminal vesicle of ndiilt rat 205 rnrrorrd 11 moiiths after castration. x 13.
4 Reiiiiiixl wsiclr of a d u l t Tat 204 rciiiorcd 11 inoiitlia after castration aild
folloniiig iirjcctiniis of 20 rat iiiiitq of rstrin daily f o r 30 4ajs. Marked g r o n t h
of tlrr smooth i i i ~ ~ cwith
l r iio cffcct oil the ~pitlielinin. X 12.
5 Semin:il veqicle reiiio~edfrom iioriiia! adult rat 57 at time of castration.
x 12.
6 Seroiitl scwiir:rl vesicle Ternox e& froin rat 5; fifteen days after castrntioii
duriiig hirh tinic 20 i n t uiiits of pstrin n e r c iujcctrtl daily. Noticeable increase
iii thr smooth niusc.lc nitlr restiatioii iliaiigrs hi the epithelium. X 1 2 .
7 Sciniiinl rcsiclc rcriioled f i o m :idtilt rat 51 fiftceii da) s after castration.
Castration changes i n cpithcliuin niorc iiiarked than in estrin iii jeetcd animal.
x
12.
8 Srmiiial vesicle reinovd froni normal ,adult rat E.6 after iiijections of 20 rat
units of estriii daily f o r 15 days. Xoticcublr increase in the snrooth iniiscle
with marked snpprwsioii of the cpitliplium.
X 13.
EFFECT O F SEX H0IG;ZIONES ON SEMINATJ VESICI.BJ
MITITON D. OVERHOLSER AND W-4R.lIGN 0. NRLSOh-
265
PLATE 2
EXPLANATION OF FIGIJRES
9 Scmiiial vesicle from immature castrated rat 131 after iujections of 20 rat
tlllits of estriii daily f o r 12 clays. No effect on epitlielium but sljght stimulation
of the sliiooth muscle when compared with uninjeeted castrated coiitrol animals.
x 20.
1 0 Seminal vesicle from immature castrated rat 227 a f t e r injections of 3 bird
imitr of male hoimoue daily f o r 1 2 days. Eormal growth of muscle and epithelium. X 20.
11 Seminal vesicle from immature castrated rat 145 a f t e r simultaiieous daily
injections of 20 r a t uiiits of estrin and 3 bird units of male liornioiie for 1 2 days.
Excessive grortlr of smooth muscle aiid epithelium due to synergistic action of
the two hormoues. X 20.
1 2 Epithelium from seminal vesicle of imiiiaturc normal animal sacrificed
a t heginning of injection period m euperimrntal animals. Low columnar epithelium with no secretion granules. X 785.
13 Epithelium froin seminal vesirle of uniiijected castrated control animal.
LONeoluninar cpitlielinm with no secretion grannles. x 7x5.
14 Epithelium from seminal vesicle of uiiinjected iiormal control animal.
Colunuiar epitlieliuin with iiumeroiis secretion graunles many oC which have halop.
X 785.
13 Epitheliuni from seminal vesicle showu in figure 9. Jmmature castrated
aiiiiiial a f t e r estrin injections. No stimulation of epithelium. >< 785.
16 Epithcliurn from seminal vesicle shomn in figure 10. Immature castrated
aiiimnl after injections of niale hormone. Seeretiou granules fairly iiunierous
with a few halos. X 785.
17 Epithelium froiii seminal vesicle shovvn in figure 11. Iminaturc castratccl
animal after aimu1t:iucous injections of male hormone and estrin, showing the
synergistic action of the t w o horniones on the epitlirluim. High columnar cells
with many secretion granules and halos. >< 785.
18 Epithelium from 8enimal vesicle of immature normal :ininial after injectionr
of estrin, showing normal epithelium. x 7Aj.
266
E F F E C T O F SEX IIORMONES OK SEhllNAL VI~CSTCJAR
XILTON D. OVEBHOLSEE AN1) WARREN 0. S B L S O N
THE ANATOXICAT. RBCORI), VOL.
62,
XO.
3
PLATE 2
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