HORMONES AND BEHAVIOR 3, 307-320 (1972)

Mounting Behavior in the Female Rat During the

Estrous Cycle, After Ovariectomy, and After Estrogen or
Testosterone Administration

PER SODERSTEN

Department of Psychology, University of Gbteborg

Sweden

Female rats displaying regular 4-day estrous cycles were tested for
male sexual behavior with a receptive stimulus female during five consec-
utive estrous cycles, after ovariectomy, and after estrogen or testosterone
treatment. Such behaviors as mounts (mount with pelvic thrusts), climbings
(mount without pelvic thrusts), and sniffing at the genital area of the
stimulus female varied systematically during the estrous cycle, being at
minimal values when the rats were in heat. Intromission patterns (mount
with pelvic thrusts accompanied by a final deep thrust and by genital
grooming) were also displayed by most females but at relatively low rates.
Ovariectomy decreased, but did not abolish masculine behavior. Estrogen in
different doses (2, 10, and 50 pg/kg) stimulated the display of intromission
patterns and mounts and depressed the frequency of climbings. The effects
of various doses of testosterone (100, 500, and 2500 pg/kg) were similar to
those of estrogen, i.e., testosterone stimulated the display of intromission
patterns and mounts and depressed the frequency of climbings. No
systematic dose-response relationships between amount of injected hormone
and sniffing frequency or latencies to the first intromission pattern or
mount could be detected. The results suggest that ovarian hormones may be
involved in the regulation of male sexual behavior in the female rat.

Early investigators thought that male sexual behavior is only rarely

displayed by female rats (Moore, 1919; Long and Evans, 1922; Hemmingsen,
1933). In 1938 and 1942 Beach demonstrated that female rats are normally
capable of showing such male sexual behaviors as mounts with pelvic thrusts
(mount) and mounts with pelvic thrusts plus a final deep thrust, i.e., the
behavior pattern which is accompanied by intromission in the male rat
(intromission pattern). These experiments by Beach demonstrated that the
females do show masculine copulatory behaviors, and subsequently Beach and
Rasquin (1942) reported that female rats frequently display such behaviors.
Of 20 females 15 mounted estrous females and three of these females also
exhibited intromission patterns. In the first experiments by Beach (1938)

307

Copyright @ 1972 by Academic Press, Inc.

All rights of reproduction in any form reserved.
308 SdDERSTEN

1942a) those females that would mount a receptive stimulus female were
themselves in behavioral estrus, and the study by Beach and Rasquin (1942)
was designed to investigate the possible relationship between ovarian hormones
and masculine sexual behavior in the female. However, it was found that the
females mounted throughout the estrous cycle and only a slight decrement
when the females themselves were receptive was noted. Further, ovariec-
tomized females mounted as much as did intacts, and the combined treatment
with estrogen and progesterone did not augment mounting behavior in these
females. Beach and Rasquin (1942; see also Beach, 1968) accordingly
suggested that mounting behavior in the female rat may be independent of
ovarian hormones. However, Pfaff (1970) later showed that estrogen
stimulates mounting behavior in spayed female rats, and in pilot studies in this
laboratory we observed that ovariectomized females only infrequently mount
estrous females. The present experiment was, therefore, designed to reinvesti-
gate the possible influence of ovarian hormones upon male behavior in the
female rat. Female rats were tested with receptive females for the display of
masculine sexual behaviors throughout five consecutive estrous cycles, they
were then ovariectomized and again tested for male behavior. Ovariectomy
decreased mounting behavior and estrogen in different doses was injected in
an attempt to reveal any dose-response relationship between amount of
injected estrogen and male behavior.
From the studies by Ball (1937, 1940), Beach (1942b), and Koster
(1943) it is well known that androgen stimulates masculine copulatory
behavior in spayed female rats. In the present experiment female rats were
treated with different doses of testosterone propionate (TP) to see whether TP
stimulates mounting behavior in the female rat in a dose-dependent fashion.

METHODS

Subjects

The subjects taking part in this study were 31 female rats born in this
laboratory. They were 118-128 days old and weighed approximately 250 g at
the beginning of the experiment.

Procedure

Prior to being used as experimental subjects all rats had displayed at

least three consecutive 4-day estrous cycles. Ovarian cyclicity was established
by the manual stimulation technique described by Zucker (1967).
Two days before an expected day of behavioral estrus each female was
placed in an individual semicircular (radius 25 cm, depth 50 cm, height 30 cm)
 HORMONESANDMOUNTINGINFEMALERATS 309

cage with a sawdust-covered floor and a Plexiglas front. The females lived in
these cages throughout the experiment in an air-conditioned colony room with
a lo/14 hr dark-light cycle (dark 1230-2300) and had free access to food and
water.
Starting on the day when the females were in heat and daily throughout
five subsequent estrous cycles all females were presented with a stimulus
female in estrus and male sexual behaviors were recorded. The stimulus
females were 4-day cycling animals of the same age as the test rats.
After these tests all females were ovariectomized and allowed a 7-day
recovery period. They were thereafter tested four times with receptive
stimulus females for masculine behaviors every fifth day.
The rats were then randomly divided into three groups and injected
daily with estradiol benzoate (EB) in different doses. One group (N=lO)
received 2 pglkg, another group (N=lO) 10 Mg/kg, and a third group (N=l 1)
SOpglkg EB. The hormone was dissolved in oleum arachidis and injected in a
volume of 0.5 ml/kg SC. This hormone treatment was given for 1.5 days and
the rats were tested with estrous females for mounting behavior every third
day.
After 15 days of EB treatment hormone injections were withdrawn and
the rats were tested every fifth day for male behavior until, after three tests,
the behavior was back to castration levels. The rats were then randomly
divided into three groups and injected daily with testosterone propionate (TP)
as follows: one group (N=lO) was injected with 100 @g/kg, another group
(N=lO) with 500 /&kg, and a third group (N=l 1) with 2500/.&kg. The
hormone was dissolved in oleum arachidis and injected in a volume of
0.5 ml/kg SC. As in the EB experiment TP treatment lasted 15 days and the
animals were tested for masculine behavior every third day.

Behavioral Testing Procedure

Tests for mounting behavior were started at 1600-1630 hr (i.e., 3.5-4 hr

after lights off) each day. At this time all rats that were expected to be in
heat were so invariably. An estrous (natural heat) female was introduced and
remained with the test animal for 15 min. A 25-W red light bulb was used as
illumination. All rats were tested daily during five consecutive estrous cycles,
and the cycle days were arbitrarily called Day 1 (day when the animals were
in heat), 2, 3, and 4. Any Day 1 female was manually stimulated after the
test to ascertain that she was in heat. In most cases this was not necessary,
since when a rat was in heat, she was mostly mounted by the stimulus female,
and in response to these mounts she would show lordosis.
The following behaviors were recorded: Intromission pattern: mount
with pelvic thrusts followed by a deep Enal thrust. This behavior pattern is in
most cases followed by genital licking. Mount: mount with pelvic thrusts.
310 So DERSTEN

Climbing: mount without pelvic thrusts. This behavior corresponds to what

Beach and Rasquin (1942) called sexual clasp. Sniffing: each time a female
investigated and sniffed the genital region of the stimulus female was
recorded. Latencies to the first mount and intromission pattern, mount
latency and intromission pattern latency, respectively, were also taken. Each
time an experimental female was mounted by a stimulus female was also
counted.
Two females together will also show other behaviors such as mutual
grooming and climbings from the side. These behaviors were not studied. An
attempt was also made to record aggressive behavior as measured by the
adoption of the fighting posture. This behavior was very infrequently seen and
will, therefore, not be considered further in this paper.

RESULTS

Mounting Behavior During the Estrous Qcle

Intromission patterns were observed in all but seven of the females. This
behavior was, however, displayed at relatively low frequencies (Fig. 1 A). A
maximum of 13 intromission patterns was recorded in one animal. Overall
mean frequencies for the respective cycle days were calculated (histograms to
the right in Fig. 1). No variation in the frequency of intromission patterns
dependent upon on any special day of the estrous cycle could be detected
(Table 1).
Mounts were very frequently recorded (Fig. 1 B). All females with the
exception of only two mounted their stimulus partner. As many as 41 mounts
were recorded in one animal. It is clear from Fig. 1 B that on cycle Day 1
(i.e., when the females were in heat) mounting behavior was at minimal
values. On Day 2 mounts increased, on Day 3 there was again a reduction,

TABLE 1
Probability Values for Cycle Day Comparisons of Overall Mean Frequencies
of Intromission Patterns, Mounts, Climbings, and Sniffing for Tests
During the Estrous Cycle. Comparisons Were Performed with the Wilcoxon t-Test

Comparisons of cycle days

l-2 l-3 l-4 2-3 2-4 3-4

Intromission patterns NS NS NS NS NS NS
Mounts <O.OOl <O.OOl <O.OOl <O.OOl NS <O.Ol
Climbings <O.OOl <O.OOl <O.OOl NS <O.Ol <O.OOl
Sniffing <O.Ol <O.OOl <O.OOl NS NS NS
 HORMONES AND MOUNTING IN FEMALE RATS 311

“is &

123456789101112 14 16 18 20 1234
,5 t 8 MOUNTS

123456789101112 14 16 18 20 1234

; 2 3 4

D SNIFFING
101

1234

Fig. 1. Mean frequencies of intromission patterns (A), mounts (B), climbings (C),
and sniffing (D) during the estrous cycle. The numbers on the abscissa refer to the 20
successive daily tests, black dots indicate days of the cycle when the rats were in heat.
The histograms to the right in the figure are overall mean frequencies for Days 1, 2, 3,
and 4 of the estrous cycle.

and finally on Day 4 mount frequency increased. The associated statistical

signifkances for overall mount frequencies are seen in Table 1.
High climbing frequencies were also seen (Fig. 1 C). All rats showed
climbings. A maximum of 25 climbings was recorded for one subject. On
cycle Day 1 climbing was much reduced, on Days 2 and 3 climbing frequency
increased, and on Day 4 there was a reduction. (For statistical significances
see Table 1.)
All rats also sniffed at the genital area of the stimulus female. A
maximum of 28 such sniffings was once observed for one individual. When the
rats were in heat they sniffed significantly less than on other cycle days (Fig.
1 D). No other systematic variation in sniffing frequency during the estrous
cycle than this reduction at heat could be detected (Table 1).
312 SdDBKSTEN

Median response latencies to the first intromission pattern and mount

were calculated. If the rat responded fewer than three out of the possible five
times on one particular day of the estrous cycle, the value for this day would
be > 15 min. The overall median for intromission patterns were based on tests
only when intromission patterns occurred, because intromission patterns were
seen in less than 5% of the tests. All tests were included in the calculation ot
the median mount latency, because the females displayed mounts in most
tests. The latency values in Table 2 should be interpreted with these
methodological difficulties in mind. There was no systematic variation in
intromission pattern latency dependent upon the stage of the estrous cycle,
and only a tendency for a prolongation of the mount latency on the day of
heat was observed (Table 2). The mean number of times a female was
mounted by the stimulus female was also calculated (Table 2). The rats were
mounted throughout the estrous cycle, and, not surprisingly, most frequently
when in heat.

Mounting Behavior Afier Ovariectomy

Starting 7 days after ovariectomy the rats were tested for masculine
behavior four times with S-day intervals. During these tests intromission
patterns were absent (Table 3). Only at the first test was one subject observed
to perform two intromission patterns. Mounting was displayed by 19 of the
animals. The frequency of this behavior was, however, reduced far below
values seen during ovarian cyclicity (Fig. 2 A and Table 3). It should be
pointed out, though, that as many as 12 mounts were seen in one subject at
the third test.
Interestingly climbing frequency increased after ovariectomy (Fig. 2 B
and Table 3), and sniffing, finally, was reduced (Fig. 2 C and Table 3).
Latency values during ovariectomy tests would be of questionable

TABLE 2
Median Latencies to the First Intromission Pattern and Mount, and Mean Times
Mounted by Stimulus Female During the Estrous Cycle. Latency to First Intromission
Pattern Is Based on Tests When Intromission Patterns Were Seen, Mount
Latencies Are Based on all Tests. The Numbers in ParenthesesAre the Number of Cases
Upon Which the Medians Are Based. The Days of the Estrous Cycle Were Compared
with the Friedman Two-Way Analysis of Variance

Day of estrous cycle

1 2 3 4 P
..- - _-.- - - -. - -.. - -..- -. --- - -.-.
Median intromission pattern latency (min) 5.2 (12) 4.2 (10) 5.7 (8) 4.0 (11) NS
Median mount latency (min) 4.6 (21) 2.6 (24) 3.8 (22) 3.6 (21) NS
Mean timesmountedby stimulusfemale 4.3 0.5 0.5 0.5 <O.OOl
 HORMONES AND MOUNTING IN FEMALE RATS 313

A MOUNTS

B CLIMEINGS

IO

:A
, C SNIFFING

5
4

II-.-
3
2

1 2 3 4
I
I 2 3 4

Fig. 2. Mean frequencies of mounts (A), climbings (B), and sniffing (C) by
ovariectomized females during four consecutive tests separated by 5 days.

TABLE 3
Overall Mean Frequencies of Intromission Patterns, Mounts, Climbings, and
Sniffing During Tests with Ovaries in situ and After Ovariectomy.
Comparisons Were Performed with the Wilcoxon t Test

Cycle tests Ovariectomy tests P

Intromission patterns 0.9 0.0 <O.OOl

Mounts 8.5 1.2 <O.OOl
Climbings 6.0 9.5 <O.OOl
Sniffing 7.0 5.2 <O.OOl

informative value, since such values would be based on very few cases. The
rats were occasionally mounted by the stimulus females, however, at very low
rates.

Mom ting Behavior During EB Injection Tests

After the ovariectomy tests the rats were injected with 2, 10, and
50 E.tg/kgEB for 15 days and tested for mounting behavior every third day.
Figure 3 A shows that EB in all doses used stimulated intromission patterns
above castration levels. 4 of 10 in the 2pg/kg group, 7 of 10 in the IO-pg/kg
group, and 7 of 11 in the 50-@g/kg group displayed intromission patterns. The
overall mean frequencies were not significantly different (histogram to the
right in Fig. 3 A).
Most rats also exhibited mounts (Fig. 3 B). The proportions of animals
displaying mounts were 10 of 10, 9 of 10, and 11 of 11 in the 2, 10, and
314 SeDERSTEN

A INTROMISSION
PATTERNS

B MOUNTS

,_

1 2 3 4 5

C CLIMBINGS

I .
I 2 3 4 5

0 SNIFFING

1. . .
1 1 3 4 5 2 10 50

Fig. 3. Mean frequencies of intromission patterns (A), mounts (B), climbings (C),
and sniffing (D) during estrogen injection tests. The rats were injected daily with 2 pg/kg
(black dots), 10 wg/kg (triangles), or 50 &kg (open circles) of estradiol benzoate for 15
days and tested every third day. The histograms to the right in the figure are overall
mean frequencies for the respective estrogen doses.
 HORMONES AND MOUNTING IN FEMALE RATS 315

50pgjkg EB groups, respectively. Although there was a tendency for the

animals that were injected with the higher EB doses (10 or 50yg/kg) to
display more mounts than the rats treated with the lower dose (2 pg/kg), the
overall mean mount frequencies did not differ significantly (Fig. 3 B,
histograms to the right in the figure).
From these observations it is apparent that EB stimulates both mount
and intromission patterns, even as low a dose as 2 pg/kg facilitates the display
of these behaviors in spayed rats. Also EB depressesclimbing frequency (Fig.
3 C compared to Fig. 2 B). This depression was found to be dose-dependent
(P< 0.02 for overall means, Kruskall-Wallis one way analysis of variance, see
histogram to the right in Fig. 3 C). Rats injected with 2 or 10 pg/kg EB
displayed significantly more climbings than rats treated with 50,ug/kg
(P < 0.05 for both comparisons, Mann-Whitney U test). The difference
between the 2- and the lOyg/kg groups, however, did not reach statistical
significance. Thus, the more EB is injected into an ovariectomized female the
less it will climb, and, to some extent, the more it will mount an estrous
stimulus female. It should be noted that all females in all EB groups climbed,
the high EB doses did not abolish climbing.
Sniffing frequency remained approximately at the same levels as those
seen during the ovariectomy tests (Fig. 3 D and Fig. 2 C). No systematic
dose-response relationship could be revealed.
Response latencies were calculated as in the tests during the estrous
cycle. Since for intromission patterns the number of cases on which the values
are based are quite small and variable, the data in Table 4 should be
interpreted cautiously. None of the differences are statistically significant.
The rats in the different EB groups were also mounted by stimulus
females. Overall means were 0.6, 0.4, and 1.1, respectively, for the 2-, lo-,
and 50yglkg groups. It is worthwhile to point out that these values are
considerably below the number of times the rats were mounted when in heat
during the estrous cycle (Table 2). When mounted, though, the rats in any EB
group would show lordosis.

TABLE 4
Median ResponseLatenciesDuring EB Tests. The Ratios in Parentheses Are the
Numbers of Cases Upon Which Each Median Is Based

Mount latency (min) Intromission pattern latency (min)

EB dose (m/kg)
2 2.3 (6/10) 7.8 (l/10)
10 1.8 (g/10) 8.4 (6/10)
50 1.4 (9/10) 6.4 (3/11)
 316 S6 DERSTEN

Mounting Behavior During TP Injection Tests

All rats were tested three times every fifth day after the termination of
EB treatment. At the end of this period mounting behavior was back to
castrational levels. The rats were then randomly divided into three groups and
treated with TP in doses 100, 500, and 2500ng/kg for 1.5 days. They were
tested for masculine behavior every third day. The lowest TP dose resulted in

12 34 5 100 I2500
vm ---
c. CLIMBINGS

IC

5
a, ‘“I 5

L
e:,b
1 2 3 4 5 loo
500
Em

10 1D SNIFFING 10 1

Fig. 4. Mean frequencies of intromission patterns (A), mounts (B), climbings (C),
and sniffing (D) during testosterone injection tests. The rats were injected daily with
100 &kg (open circles), 500 Ecg/kg (triangles), or 2500 &kg (black dots) of testosterone
propionate for 15 days and tested every third day. The histograms to the right in the
figure are overall mean frequencies for the respective testosterone doses.
 HORMONESANDMOUNTINGIN FEMALE RATS 317
very few intromission patterns, only one of the animals in this group showed
intromission patterns. Seven of ten and 8 of 11 of the animals in the 500- and
2500~pg/kg TP groups displayed intromission patterns. The behavior was,
however, observed at relatively low frequencies (Fig. 4 A, histogram to the
right in the figure). A Kruskall-Wallis one-way analysis of variance revealed
that the difference in overall frequencies was significant at the 0.02 level, and
between group comparisons showed that the 500- and 2500+g/kg TP groups
exhibited significantly more intromission patterns than did the 100~@g/kgTP
group (I’< 0.05 for comparisons, Mann-Whitney U test), whereas there was no
difference between these two groups.
All TP doses also stimulated mounts (Fig. 4 B). The proportions of
animals displaying mounts were 10 of 10 (100 /.&kg), 10 of 10 (500 pg/kg),
and 10 of 11 (2500&kg). In the lowest TP group, however, mount
frequency was below the frequencies seen in the two higher groups (P< 0.01
for overall mean frequencies, Kruskall-Wallis one-way analysis of variance, see
histogram to the right in Fig. 4 B). Between-group comparisons revealed that
both the 500- and 2500-@g/kg TP groups showed statistically more mounts
than did the 100~rig/kg TP group (I’< 0.02 for both comparisons,
Mann-Whitney U test), whereas these groups did not differ in mount
frequency.
These data substantiate the findings of earlier research (Ball, 1937,
1940; Beach, 1942b, and Koster, 1943) that TP stimulates both mounts and
intromission patterns in spayed rats. A low dose (IOOpgIkg) of TP stimulates
male behavior only slightly, increasing the dose five times (500pglkg) results
in a marked stimulation of this behavior, but a further increment of the TP
dose (2500pg/kg) results only in slightly more masculine behavior.
The data for climbing is the mirror image of the mounting data (Fig.
4 C). lOOpg/kg TP was not effective in depressing climbing frequency,
500 pg/kg and 2500 /&kg strongly depressed the display of this behavior. The
difference in overall mean frequency was highly significant (I’< 0.001,
Kruskall-Wallis one-way analysis of variance), and between-group comparisons
revealed P values of 0.05 (lOO-500pg/kg and 500-2500 &kg), and 0.002
(lOO-2500&kg). Climbing was displayed by all rats however, but during the
last test 1 of 10 in the 500~&kg group and 6 of 11 in the 2500+g/kg group did
not climb. Thus, the more TP a spayed rat receives the less it will climb, and, to
a certain degree, the more it will mount.
As in the EB experiment no systematic dose-response relationship in
sniffing frequency could be detected (Fig. 4 D). Response latencies were
calculated as in the above experiments (Table 5). There was a tendency for
mount latency to become progressively shorter the higher the injected TP
dose, but, due to great individual variation, the difference was not statistically
significant.
Only rarely did the stimulus females mount a TP-treated rat. Once,
318 SODERSTEN

TABLE 5
Median Response Latencies During TP Tests. The Ratios in Parentheses Are the
Number of Cases Upon Which Each Median is Based

Mount latency (min) Intromission patterns latency (min)

TP dose Wkd
100 4.8 (9/10) 5.6 (l/10)
500 1.3 (lO/lO) 3.1 (2/10)
2500 .5 (10/l 1) 6.8 (5111)

during the last test, was one rat (500ng/kg TP group) mounted seven times,
and this rat showed lordosis twice in response to these mounts. At the same
test, this individual rat was observed to mount the stimulus female 21 times.

DISCUSSION

The results of these experiments agree with those of Beach and Rasquin
(1942) that masculine sexual behaviors are commonly displayed by female
rats. Beach and Rasquin (1942), however, found that female rats mount with
equal frequencies throughout the estrous cycle. In the present experiment a
clear reduction of male behavior on days when the females were in heat was
consistently found throughout five consecutive estrous cycles. This finding is
not necessarily divergent from the earlier findings by Beach and Rasquin
(1942), since these workers also reported a decrement of male behavior on
days when the females were in heat. On a day of heat a female will, of
course, also show lordosis, and at this time the animals were found to be
mounted by their stimulus partners significantly more than on other days of
the estrous cycle. It might, therefore, be that this increase in mounts by the
stimulus female makes the probability that the female herself will mount
lower than on other days, when stimulus females are less likely to mount, this
same probability is higher. When the female rat displays lordosis ovarian
progesterone secretion is at its highest values (Feder, Resko, and Coy, 1968;
Hashimoto, Henricks, Anderson, and Melampy, 1968; Ushida, Kadowaki, and
Miyake, 1969) and one other possible explanation for the reduction of male
behavior seen at the time of behavioral estrus would be that ovarian
progesterone per se inhibits the display of masculine behavior in the female
rat. In this present report a significant reduction of male behavior at the
middle of the estrous cycle was also found, and at this time, when lordosis is
not displayed, there is evidence suggesting that ovarian progesterone secretion
again increases (Feder et aE., 1968; Hashimoto et al., 1968; Ushida et al.,
1969). It, therefore, appears that the intensity of mounting behavior in the
female rat is inversely related to ovarian progesterone secretion. Further,
 HORMONES AND MOUNTING IN FEMALE RATS 319

experiments now in progress in this laboratory, show that during pregnancy

and pseudopregnancy, when plasma levels of progesterone are high, mounting
behavior is very much reduced.
The significant drop in climbing frequency on the fourth day of the
estrous cycle may be related to the increase of’ estrogen secretion from the
ovaries at this time (Hori, Ide, and Miyake, 1968; Yoshinaga, Hawkins, and
Stocker, 1969; Brown-Grant, Exley, and Naftolin, 1970), since it was
demonstrated that estrogen injections depress the display of climbing.
Simultaneously with this decrease in climbing behavior an increase in mount-
ing frequency was observed, which may be related to the increase of plasma
estrogen concentration, since it was also demonstrated that estrogen stimulates
mounting behavior in spayed females. However, on Day 2 of the estrous cycle,
mounting behavior was also at high rates, and at this time ovarian estrogen
secretion is low (Hori et al., 1968; Yoshinaga et al., 1969; Brown-Grant et al.,
1970). Thus, the possible importance of estrogen in the regulation of
masculine sexual behavior under physiological conditions in the female rat
remains to be determine.
Beach and Rasquin (1942) reported that ovariectomy does not decrease
the frequency of mounting behavior in female rats. The present results are in
direct contrast to these findings. It was found that ovariectomized females
mounted estrous stimulus females at very low rates. Unfortunately, we have
no other explanation for this discrepancy than to refer to procedural
differences. Beach and Rasquin (1942), in an attempt to get measures not
only of male, but also of female behavior in their rats, used vasectomized
males, which were present during parts of the tests, and this might, as Beach
and Rasquin (1942) pointed out, have resulted in pseudopregnancies, and
hence progesterone secretion, which in turn may have reduced mounting in
their females. However, the present results agree with those of Beach and
Rasquin (1942) that ovariectomized females do show mounting behavior, and
this suggests the involvement of some controlling factor beside the ovaries in
the regulation of male behavior in female rats.
Injecting spayed females with estrogen or testosterone resulted not only
in a enhancement of mounts and intromission patterns, but also in a
depression of climbing. This is in agreement with Pfaff (1970), who also
reported that estrogen and testosterone stimulate mounting behavior in
castrated female rats.
Though the present findings do not tell what hormones are involved in
the control of male sexual behavior in the female rat, they suggest the
involvement of ovarian hormones.
ACKNOWLEDGMENTS

This work was supported by grants from the Swedish Council for Social Research
&W/71 p) and RiksbankensJubileumsfond (68/50:1) to ProfessorKnut Larsson. We
thank Organon, the Netherlands, for generous supply of hormones. The skillful technical
 320 SdDERSTEN

assistance of Mr. Nils Carlsson, Mrs. Margit Barscay, and Mr. Miclos Barscay is gratefully
acknowledged

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