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Deciduomal response of the golden hamster uterus.

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Deciduomal Response of the Golden Hamster Uterus
M. J. K. HARPER
Worcester Foundation for Experimental Biology, Shmwsbury,
Massachusetts 01545
ABSTRACT
The deciduomal response to uterine trauma in pesudopregnant hamsters has been studied. Trauma was effected in one uterine horn by leaving a
sterile silk thread in the lumen, pulling a thread through the lumen, injecting oil or
air into the lumen or crushing the horn with a hemostat. At autopsy the traumatized
and untraumatized horns were weighed separately and examined histologically for a
deciduomal reaction. The day of greatest uterine sensitivity was found to be day 4 of
pseudopregnancy, and maximal deciduomal growth (mean uterine weight +- S.E. =
2582 2 279 mg) occurred by day 8. No significant further increase in uterine weight
was seen by day 9, and by day 10 separation of the deciduoma from the myometrium
had begun. Leaving a thread in the uterine lumen produced a significantly greater
uterine weight by day 8 than with any other stimulus, but good deciduomal responses
(uterine weights of 1679 2 245, 1255 156 and 1192 2 219 mg) were induced by
oil injection, pulling a thead through the uterus and crushing the uterus respectively.
A poor reponse was produced by air injection. At no particular time during day 4 of
pseudopregnancy was it possible to demonstrate a peak of uterine sensitivity using as a
traumatic stimulus either oil injection or pulling a thread through the lumen. In this
respect hamsters differ from both rats and mice, which have well defined peaks of maximal uterine sensitivity.
*
The deciduomal response of the rat
uterus to a traumatic stimulus has been extensively explored (Rothchild and Meyer,
'42; Finn and Keen, '63; Yochim and De
Feo, '62, '63; De Feo, '63a,b; von BerswordtWallrabe, Geller and Herlyn, '64 a,b; Harper, '67). The time of maximal sensitivity
of the uterus falls between 1100 and 1300
hours on day 5 of pseudopregnancy (day
1 = day of vaginal cornification) (De Feo,
'63a,b). This peak of sensitivity is related
to the release of estrogen occurring on day
4 which is necessary for induction of implantation (Canivenc and L a a r g u e , '56;
Cochrane and Meyer, '57; Psychoyos, '60,
'61; Zeilmaker, '63). In the hamster, ovarian estrogen is not required for induction
of implantation (Orsini and Meyer, '59;
Prasad, Orsini and Meyer, '60; Harper,
Prostkoff and Reeve, '66), and so this
raises the question whether a peak of
uterine sensitivity occurs in the hamster
and if so on what day. In the hamster implantation occurs during the afternoon of
day 4 of pregnancy (Orsini, '63a), and
Orsini ('63c) has reported that injection of
0.05 ml of air into the uterine lumen on
day 4 of pseudopregnancy (i.e., between 3
days 9 hours and 3 days 22 hours after
ANAT. REC., 163: 563-574.
ovulation) can induce a deciduomal reaction.
Various methods have been used for
inducing a deciduomal response in hamsters. Kent and Lytle ('60) found that
sewing threads through the uterus on the
second or third night after sterile mating
was a relatively ineffective method of inducing a deciduomal reaction unless prolactin was also given. A later report from
the same laboratory (Turnbull and Kent,
'63) indicated that using the same techniques exogenous prolactin was not in fact
essential for induction and maintenance
of a deciduomal response. Orsini ('63b)
showed that, in addition to intraluminal
injection of air, injection of histamine or
saline or insertion of threads or a needle
into the uterine lumen could induce a
deciduomal response. Intraperitoneal injection of hamsters with pyrathiazine was
an unsuccessful stimulus (Orsini, '63b)
which contrasts markedly with 100% success in rats (Shelesnyak and Kraicer, '61 ).
Blaha ('67) studied the deciduomal response to intraluminal injection of air or
oil (0.07 ml) in young and old hamsters.
The deciduomal response to these stimuli
was variable in 4 to 7-month-old hamsters.
Received Oct. 2, '68. Accep1:ed Nov. 18, '68.
563
564
M. J. K. HARPER
hematoxylin and eosin. If the uterus of an
animal failed to show a deciduomal reaction, and if the ovaries upon macroscopic and microscopic inspection did not
have normal corpora lutea, such an animd
was considered to be not pseudopregnant
'65).
and was discarded from the experimental
The Present Paper reports experiments group. The traumatized and untraumatized
in hamsters to determine (a) the time Of uterine horns were dissected out, blotted
optimal uterine sensitivity to a traumatic and weighed, and sections taken for histostimulus in pseudopregnant animals and logical examination
procedures to
(b) the deciduomal response of such Uteri those for the ovaries).
to different types of traumatic stimuli.
Only nine out 14 responded to air and five
out seven to oil injection, but six out of
six responded to a knife scratch. Intraluminal injection of oil has been very successful in producing deciduomata in both rats
(Finn and Keen, '63) and mice (Finn,
MATERIALS AND METHODS
Two hundred and forty-three mature female golden hamsters (Mesocricetus aurat u s ) purchased from a local breeder were
included in this study. Females in estrus
were placed with vasectomized males in
the evening (after 19:30 hours) and left
for at least one-half hour after first mating
was observed. The following day, on which
the postestrous discharge was observed,
was designated day 1 of pseudopregnancy.
At least five hamsters were included in
each experimental group.
On various days of pseudopregnancy a
laparotomy was performed and the left
uterine horn was traumatized in a variety
of ways: (a) A sterile silk thread attached
to a fine intestinal straight needle was
passed longitudinally through the uterine
lumen entering the uterine wall just above
the cervix and exiting just below the
uteretubal junction, the ends being left
free in the peritoneal cavity; (b) A silk
thread was passed as in (a) through the
uterine lumen but instead of being left
in situ was pulled from the lumen; (c)
Cottonseed oil (Fisher Scientific Company)
or air (Orsini, '63c) was injected into the
uterine lumen, and leakage was prevented
by gripping the uterine wall around the
needle point with fine forceps while the
needle was removed; (d) The uterus was
crushed in three places with a hemostat.
The animals were killed with chloroform
and autopsied on days 6-10 of pseudopregnancy. The ovaries were dissected free,
blotted and weighed and the corpora lutea
counted. The ovaries were then fixed in
Bouin's fluid, dehydrated, embedded in
paraplast (Fisher Scientific Company) and
sectioned at 6 p. Sections were stained with
RESULTS
Table 1 shows the effect of placing a
thread in the uterine horn on different
days of pseudopregnancy and the rate of
growth of the induced deciduoma. Traumatization on day 2 was relatively ineffective
in producing a deciduomal reaction (autopsy on day &fig. 1) and the weight of
the uterus was not significantly increased
by delaying autopsy from day 7 to day 9.
A similar but even smaller response was
seen after traumatization on day 6 of pseudopregnancy (fig. 5). Traumatization on
day 3 produced a good deciduomal response
by day 7, but the uterine weight showed
no significant increase by day 8 (fig. 2) or
9. The maximum mean weight of the
traumatized horn was significantly smaller
than that seen after traumatization on day
4 and autopsy on day 9 (the significant differences between groups were calculated
by Duncan's multiple range test-Duncan,
'55-and
are shown in table 2). There
were, however, no significant differences in
the weights of the traumatized horns of
animals autopsied on days 7 or 8 irrespective of the traumatization being done on
day 3 or 4. Traumatization of the uterus
on day 5 produced a greater weight increase and deciduomal response by day 8
(fig. 4) than did trauma on day 2 or 6, but
the response was significantly less than
those obtained by trauma performed on
day 4 (fig. 3). After traumatization of the
uterus on day 4 of pseudopregnancy only
a small deciduomal response was apparent
by day 6 (fig. 6); this had increased somewhat by day 7 (fig. 7) and reached a maximum by day 8 (fig. 3). Although greater
responses were observed if autopsy was
delayed until day 10 (fig. 8) the variabil-
565
DECIDUOMAL RESPONSE OF HAMSTER UTERUS
ity in this group made any such differences
not significant (tables 1, 2).
The highest ratio of T/U (weight of
traumatized over untraumatized uterine
horns) was produced when trauma and
autopsy were performed on days 4 and 8
respectively. There was no difference in
T/U ratio, however, between this and
either of the other groups traumatized on
day 4, and autopsied on days 7 and 9 respectively. If autopsy was delayed until
day 10, the ratio was greatly reduced owing
to the increase in weight of the untraumatized horn with the end of pseudopregnancy
and approach of estrus. In addition the
deciduoma was starting to separate from
TABLE 1
Response of the hamster uterus to intralunzinal insertion of a thread on different
days of pseudopregnancy
Day of
trauma
Day of autopsy
fi
7
2
638f52
181
(2.220.3)
695 f84
[81
(1.940.1:)
2204 k 285
[71
(11.8e1.2)
2115f182
I61
(10.5e0.9)
2065’182
161
(9.2t2.0:)
2278 -C 194
[71
(14.9f1.8)
2582e279
I111
(16.0k1.9)
2867k178
161
(13.140.9)
774 e 97
[81
(4.4e0.8)
1238f 162
I81
(6.2f 0.9)
1488’ 151
t71
(5.2t 1.1I;
415247
I61
(1.6f0.2)
4572 18
592f 98
I61
(3.4k0.6)
3
956f205
171
(5.4f1.4)
4
5
10
9
(I
6
30372508
El
(8.121.2)
[el
(1.520.11
traumatized horn (mg). Mean f S.E.
Number animals/gmup.
3 Ratio between weights of traumatized and untraumatized horns. Mean f S.E.
1 Weight of
2
TABLE 2
Significance of differences between experimental groups shown in table 1 . Means joined by vertical
lines do not differsignificantly at the 5 % level (Duncan’s multiple range test)
Day of
Trauma
Autopsy
4
4
4
4
3
3
3
5
5
4
5
2
2
2
6
6
10
9
8
7
7
8
9
9
8
6
7
9
8
7
9
8
1
Mean
weight
(mg) pf
traumatized
horn
3037
z
2278
2204
2115
2065
1488
1238
956
774
695
638
592
457
415
Significance
I
Day of
Trauma
Autopsy
4
4
4
3
3
3
4
5
8
7
9
7
8
9
10
8
6
9
7
7
8
9
8
9
‘44
11 ;
5
2
Not included in analysis because of excessive variance.
Mean
ratio
T/U
16.0
14.9
13.1
11.8
10.5
9.2
8.1
6.2
8.4
5.2
4.4
3.4
2.2
l.9
1.6
1..5
Significance
566
M.
J. K. HARPER
the underlying stroma (fig. 8). Traumatization on day 3 and autopsy on days 7, 8
or 9 produced T/U ratios not significantly
smaller than if trauma was delayed until
day 4 and autopsy performed on day 9.
Trauma on any other day irrespective of
day of autopsy resulted in consistently
smaller T/U ratios.
The results of these foregoing experiments demonstrated that day 4 of pseudopregnancy was the day when the uterus
could respond optimally to a traumatic
stimulus of a thread left in the lumen. The
next group of experiments was therefore
concerned with the uterine response to different types of traumatic stimuli applied
on this day of optimal sensitivity. Since
oil had been shown to produce a good
deciduomal response when placed in the
uterine lumen of both rats (Finn and Keen,
'63) and mice (Finn, '65), but a more
variable response in the uteri of hamsters
(Blaha, '67) this problem was reinvestigated. Table 3 shows the response of the
hamster uterus to various volumes of cottonseed oil placed in the lumen between
1300 and 1400 hours on day 4. Little response was obtained with 0.01 ml (fig. g),
some response with 0.02 ml (fig. 10) and
a maximal response with 0.04 ml (fig. 11) .
This dose was therefore used in subsequent
experiments. One disturbing fact emerged
during these experiments and this was the
highly variable response (indicated by the
larger S.E.) to the larger volumes of oil.
The deciduomal response to other types
of traumatic stimuli applied on day 4 of
pseudopregnancy was therefore studied
(table 4). None of these stimuli produced
as large a response as that seen after leaving a thread in the uterus. There was no
significant difference in mean weight of
the uterus after putting oil (0.04 ml) in
the uterus (fig. l l ) , pulling a thread
through it (fig. 12) or crushing it with a
hemostat (fig. 13). All these methods produced a significantly better response than
injecting air into the lumen (fig. 14). Consideration of the T/U ratio did not change
this situation: leaving a thread in the
uterus was the most effective stimulus and
placing air in the uterus the least effective.
There was no significant difference between the other three methods.
To investigate whether there is a particular time on day 4 of pseudopregnancy at
which the uterus exhibits a peak of sensitivity, two methods of deciduomal induction were used. Table 5 shows the results
obtained with placing oil in the uterine
TABLE 3
Response o f the hamster uterus to different volumes o f cottonseed oil inserted intraluminally
on day 4 o f pseudopregnancy: autopsy p e r f m e d on day 8
Volume
of oil
Mean weight (mg)
Mean ratio
f S.E.of uterine horns
No. of
hamsters
S.E.
Traumatized
Untraumatized
Traumatized/
Untraumatized
513 483
8484 100
1573k272
16634388
265 zk 33
2,39519
244 2 28
294rt 25
2.3C 0.8
2.540.2
7.3+- 2.0
6.341.9
ml
0.01
0.02
0.04
0.08
5
5
6
6
TABLE 4
Response o f the hamster uterus to different stimuli on day 4 o f pseudopregnancy: autopsy
performed on day 8
Stimulus
Thread left in lumen
0.04ml oil intraluminally
Thread pulled through lumen
Crushed with hemostat
0.05 ml air intraluminally
No. of
hamsters
11
11
12
6
7
Mean weight (mg)
+- S.E.of uterine horns
Mean ratio
S S.E.
Traumatized
Untraumatized
Traumatized/
Untraumatized
2582 rt 279
1679-C 245
12552 156
1192rt 219
437C 66
1705 12
2342 17
238 f 17
172511
284548
16.0k1.9
7.7f 1.4
5.6f0.8
7.4f1.6
1.640.2
567
DECIDUOMAL RESPONSE OF HAMSTER UTERUS
TABLE 5
Response of the hamster uterus to intraluminal insertion o f 0.04 ml cotton seed oil at
various times on day 4 o f pseudopregnancy: autopsy performed 01%day 8
Time of
insertion
No. of
hamsters
Mean weight ( m g )
2 S.E.of uterine horns
Traumatized
hour
900-1000
1100-1200
1300-1400
1400-1500
1600-1700
12
13
11
12
9
1103t228
1091-C 226
1679f 245
1021*219
11142 173
lumen. Although the highest mean response
was obtained by placing oil in the uterus
between 1300 and 1400 hours, there were
no significant differences between the
weight responses observed at any of these
times. The T/U ratios showed no significant
changes in relation to the time of oil insertion. Since oil, like leaving a thread in
the uterus, might be considered a continuing stimulus, a further series in which
a thread was pulled through the lumen
was done (table 6). The results of these
experiments were essentially similar to
those produced by the oil. The time of
greatest sensitivity was again apparently
between 1300 and 1400 hours on day 4
(fig. 12), but there were no signikant differences either in mean weight response or
in T/U ratio for any of the times studied
on day 4 (figs. 16, 17). Responses obtained
by pulling a thread through the uterus
late on day 3 (fig. 15) or early on day 5
of pseudopregnancy (fig. 18) were smaller
than those obtained on day 4.
DISCUSSION
In rats De Feo ('63a) observed that the
uterine response to a traumatic stimulus
reached a peak of sensitivity on day 5 of
pseudopregnancy (my dating). The responses obtained after trauma on day 4
or on day 6 were less than 31% and 13%
respectively of that obtained on day 5. It
was suggested that the phenomenon of
maximal sensitivity represented a phase in
which stromal cells were most capable of
responding to ovarian hormones. When the
stromd cells reached maximal sensitization, they apparently entered a refractory
period because of exposure to estrogen
(De Feo, '63a). These observations were
extended in a paper by Yochim and De Feo
Untraumatized
*
247 21
269t 22
2345~17
230 26
281 t 27
*
Mean ratio
+ S.E.
Tra&atized/
Untraumatized
5.1 2 1.4
5.1 & 1.6
7.7 -C 1.4
5.65 1.5
4.3 +- 0.8
('63) using cycling rats ovariectomized
and given steroid treatment. It was found
that progesterone alone caused an extended duration of Sensitivity to trauma,
but that the magnitude of the response was
a function of estrogen before trauma. In
other words, if no estrogen was given during the pre-trauma period, the sensitivity
of the uterus did not reach a sharp peak
on day 5 and only decreased gradually
during the succeeding days. If, however,
estrogen was given duriing the pre-trauma
period then there was a sharp peak of
sensitivity on day 5, and virtually no response on day 6. Pre-trauma treatment with
estrogen also caused a sharp increase in
uterine sensitivity on day 5 in rats ovariectomized on day 4 of pseudopregnancy
and given progestrone (Harper, '67).
The present experiments in the hamster
show that the period of uterine sensitivity
to placing a thread in the uterus is prolonged and somewhat similar to that seen
in ovariectomized rats given progesterone
alone (Yochim and De Feo, '63). Placing
a thread in the uterus does provide a stimulus of longer duration than the knife
scratch technique, and thus as has been
shown in the rat (De Fso, '63b) may exaggerate the duration of uterine sensitivity
to trauma. Slitting the haimster uterus with
scissors causes a large deciduomal response
five days later irrespective of whether the
trauma occurs on days 3, 4 or 5 of pseudopregnancy (De Feo, '67) but the knife
scratch technique is only maximally effective on day 4 (Atkins and De Feo, '67).
However despite this criticism, it seems
clear that the peak of uterine sensitivity in
the hamster is reached 011 day 4 of pseudopregnancy, and this agrees with earlier
findings (Orsini, '63b; Atkins and De Feo,
568
M. J. K. HARPER
TABLE 6
Response o f the hamster uterus to intraluminal insertion and removal of a thread at various
times on days 3, 4 and 5 of pseudopregnancy: autopsy performed on day 8
l i m e of
insertion
and removal
day
No. of
hamsters
Mean weight (mg)
f S.E.of uterine horns
Traumatized
Untraumatized
8622 171
1185-t171
1255f 156
892f69
453 f40
4
1
9
'
3
8
283 2 39
2 3 8 e 17
281 f31
324k24
Mean ratio
& S.E.
Traumatized/
Untraumatized
hour
3: 1600-1700
4: 900-1000
4: 1300-1400
4: 1600-1700
5: 900-1000
6
10
12
10
6
'67). De Feo ('63b) has shown that in rats
the time of maximal sensitivity on day 5
falls between 1100 and 1300 hours. No
such obvious peak of sensitivity has been
found in hamsters (Orsini, '63c and present
study - tables 5 and 6). An interesting
antithesis is provided by the steroid requirements for implantation and the uterine response to trauma in rats and hamsters.
Ovarian estrogen is not required for induction of implantation in hamsters (Orsini and Meyer, '59; Prasad et al., '60; Harper et d.,'66), and the duration of uterine
response to trauma is prolonged (Turnbull
and Kent, '63; present study): ovarian
estrogen is required for induction of implantation in rats (Canivenc and LafFargue,
'56; Cochrane and Meyer, '57; Psychoyos,
'60, '61; Zeilmaker, '63), and the duration
of uterine response to trauma is short (De
Feo, '63a; Yochim and De Feo, '63).
The growth of deciduomal tissue in the
hamster is extremely rapid, reaching a
maximum by day 8 of pseudopregnancy
in this study, although De Feo ('67) records that maximal size is not achieved
until day 9 or 10 (my dating). In four
days the uterine weight increased from less
than 200 mg to over 2500 mg, a sixteenfold
increase (table 4), whereas in rats five
days were required for the uterine horns
to increase in weight from about 200 to
2000 mg (De Feo, '63a). In the present
study no examination of the earliest stages
of decidualization was made, but Turnbull
and Kent ('63) observed decidual cells, as
early as two days after traumatization (i.e.,
about the middle of day 4). Twenty-four
hours after intraluminal injection of air on
day 4 Orsini ('63b) found discrete minute
opacities visible at the antimesometrial
surface of the endometrium. This ability of
2.1 f 0.4
5.0f 0.9
5.6f 0.8
3.720.7
1.2f 0.2
the hamster uterus to increase in weight is
not restricted to the deciduomal response,
but is also seen after estrogen treatment.
For instance, the uteri of immature rats increased in relative weight from 78 to 225
mg/100 g body weight in response to three
daily injections of 5 pg of estradiol in oil
given subcutaneously (Harper and Walpole, '67). Similar treatment of immature
hamsters caused a uterine weight increase
from 86 to 202 mg/100 g body weight,
and increasing the dose of estradiol to
50 clg/day produced a uterine weight of
401 mg/100 g body weight. Even at this
remarkable weight increase the slope of
the dose response line showed no signs of
reaching a plateau as it does in rats (Harper, '66). The hamster uterus clearly has
great growth potential given the correct
stimulus and favorable hormonal conditions. However, a massive deciduomal response was only produced by the stimulus
of a thread left in the uterus. The other
successful stimuli induced little more than
50% of this weight increase during the
same time span. The poor response to intraluminal injection of air in these studies
compared to its successful use by Orsini
('63b,c) is puzzling. Blaha ('67) reported
that the mean weight of uterine horns with
deciduomata induced by air or oil insertion
was 910 2 135mg. This figure falls almost
midway between those shown in table 4
for oil and air injection respectively. It is
not possible from the data of Blaha ('67)
to determine what weight response was obtained by air alone. Using the knife-scratch
technique on day 4 of pseudopregnancy
Atkins and De Feo ('67) recorded a mean
uterine weight five days later of 1525 -C
198 mg. This figure is comparable to those
produced in the present study by all meth-
DECIDUOMAL RESPONSE O F HAMSTER UTERUS
ods of trauma except use of air intraluminally or leaving a thread in the uterus.
In many experiments in the present
study the individual responses within a
group were very variable and this is shown
by the large standard errors of the means.
This made demonstration of signif?cant
differences between various treatments difficult. Drawing a thread through the
uterus resulted in a lower coefficient of
variation, and it was decided to use this
as a standard stimulus in future experiments concerning ovariectomy and steroid
replacement therapy. The mean weight inincrease obtained by this method was not
significantly larger than the mean weight
of the uterine horns of eight-day pregnant
hamsters (798 Ifr: 83 mg: n = 14).
De Feo (’63b) has proposed a hypothesis
to account for the development of decidua
in rats based on two variables “(a) the
duration of maximal uterine sensitivity
and (b) . . . the half-life of the inducing
stimulus.” In the hamster with a prolonged
duration of sensitivity, the use of an inducing stimulus with a long half life (i.e.,
thread left in the uterus) did produce (in
agreement with De Feo’s prediction) a massive deciduomal response. On the other
hand the use of a stimulus with a short
half life (i-e., oil in the uterus or a thread
pulled through the uterus) produced a
nodal form of deciduomal response.
ACKNOWLEDGMENTS
This investigation was supported by a
United States Public Health Service grant
HD-02597 from the National Institute of
Child Health and Human Development.
Sincere thanks for excellent technical assistance are due to Mrs. Ann S. W. Elliott.
LITERATURE CITED
Atkins, G., and V. J. De Feo 1967 Unpublished
experiments quoted by De Feo (’67).
Blaha, G. C. 1967 Effects of age, treatment,
and method of induction on deciduomata in
the golden hamster. Fert. Steril., 18: 477485.
Canivenc, R., and M. Laffargue 1956 Survie
prolong& d‘oeufs f6cond6s non implant& dans
l’ut&us de rattes castrees et injectkes de progbsterone. C. R. Acad. Sci., Paris, 242: 28572860.
Cochrane, R. L.,and R. K. Meyer 1957 Delayed
implantation in the rat induced by progesterone.
Proc. SOC.exp. Biol. Med., 96: 155-159.
569
De Feo, V. J. 1963a Temporal aspect of uterine
sensitivity in the pseudopregnant or pregnant
rat. Endocrinology, 72: 305-316.
1963b Determination of the sensitive
period for the induction of deciduomata in the
rat by different inducing procedures. Endocrinology, 73: 488497.
1967 Decidualization. In: Cellular Biology of the Uterus. Ed. by R. M. Wynn, Appleton-Century-Crofts, New York, 191-290.
Duncan, D. B. 1955 Multiple range and multiple ‘F’ tests. Biometrics, 11: 1 4 2 .
Finn, C. A. 1965 Oestrogen and the decidual
cell reaction of implantation in mice. J. Endocr.,
32: 223-229.
Finn, C. A., and P. M. Keen 1963 The induction of deciduomata in the rat. J. Embryol.
exp. Morph., 11: 673-682,.
Harper, M. J. K. 1966 unpublished experiments.
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progesterone. Acta endocr. Copenh., 54: 241248.
Harper, M. J. K., B. T. Prostkoff and R. J. Reeve
1966 Implantation and embryonic development in the ovariectoniized hamster. Acta
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implantation and mode of action in rats. J.
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Kent, G. C., Jr., and S. L. Lytle 1960 Decidual
cell responses following exogenous prolactin
in uterine-traumatized hamsters. Anat. Rec.,
138: 360-361.
Orsini, M. W. 1963a Morphological evidence on
the intrauterine career of the ovum. In: Delayed Implantation. Ed. by A. C. Enders, Chicago University Press, 155-166.
1963b Induction of deciduomata in
hamster and rat by injected air. J. Endocr.,
28: 119-121.
1963c Attemued ciecidualization in the
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PLATE
PLATE 1
EXPLANATION O F FIGURES
All uteri were photographed using bright field illumination after fivation in Bouin’s fluid
and staining with hematoxylin and eosin. Magnification in all figures x 6.
Figs. 1-5 Sections of uteri traumatized by placing a thread in the lumen on various
days of pseudopregnancy and examined on day 8, showing deciduomal response.
1
Traumatized on day 2.
2 Traumatized on day 3.
3 Traumatized on day 4.Note the optimal response to trauma on this day.
4 Traumatized on day 5.
5 Traumatized on day 6.
Figs. 6-8 Sections of uteri traumatized by placing a thread in the lumen on day 4 of
pseudopregnancy and examined on various days thereafter, showing deciduomal response.
Compare the response on days 6, 7 and 10 (see below) with that seen in figure 3 in which
trauma occurred on the same day but examination was made on day 8.
6 Examined on day 6.
7 Examined on day 7.
8 Examined on day 10.
Figs. 9-11 Sections of uteri traumatized by placing various quantities of cottonseed oil
in the lumen on day 4 of pseudopregnancy and examined on day 8, showing deciduomal response.
9 0.01 ml oil.
10 0.02 ml oil.
11 0.04mloil.
Note the best response occurred after use of 0.04 ml oil, but compare this with figure 3
(leaving a thread in the lumen) and figures 12-14 (see below).
Figs. 12-14 Sections of uteri traumatized in different ways between 1300-1400 hours on
day 4 of pseudopregnancy and examined on day 8, showing deciduomal response. The best
response was obtained by pulling a thread through the lumen, but observe similarity with
response shown in figure 11 (placing 0.04 ml oil in the lumen) and much smaller response
when compared with figure 3 (leaving a thread in the lumen).
12 Pulling a thread through the lumen.
13 Crushing the uterus with a hemostat.
14 Instillation of 0.05 ml air into the lumen.
Figs. 15-18 Sections of uteri traumatized by pulling a thread through the lumen at
various times on days 3,4 and 5 of pseudopregnancy and examined on day 8, showing deciduomal response. Compare figures 16 and 17 with figure 12 (traumatized between 1300-1400
hours on day 4). Although the best response was achieved in this group, the differences between mean weight for the group represented in figure 12 and the mean weights for the
groups represented in figures 16 and 17 were not significant.
15 Traumatized between 1600-1700 hours on day 3.
16 Traumatized between 900-1000 hours on day 4.
17 Traumatized between 1600-1700 hours on day 4.
18 Traumatized between 900-1000 hours on day 5.
572
DECIDUOMAL RESPONSE OF HAMSTER UTERUS
M. J. K. Harper
PLATE 1
573
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