Journal of Experimental Psychology: Copyright 1986 by the American Psychological Association, Inc.

Animal Behavior Processes 8097-7403/86/S00.75

1986, Vol. 12, No. 4,403-411

Pavlovian Conditioning of Sexual Arousal: Parametric and

Background Manipulations

Edward Zamble, John B. Mitchell, and Helen Findlay

Queen's University at Kingston, Kingston, Ontario, Canada

Recent studies have shown that a Pavlovian conditioned stimulus (CS) for unconsummated sexual
arousal can increase the rate of copulation in the rat. This report includes five experiments examin-
ing the effects of parametric manipulations on the conditioned arousal response. Results show that
between six and nine trials are necessary for reliable conditioning, but extinction is somewhat slower
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than acquisition. The function for the CS-US (unconditioned stimulus) interval is quadratic, with a
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minimum of several minutes required for effective conditioning. In the first three experiments, it
appeared that background cues were conditioned as well as the designated CSs. and this was tested
explicitly in the last two studies. In one, the effect of background cues was shown by training and
testing in different situations; in the second, background cues were shown to be subject to latent
inhibition. These results demonstrate the influence of Pavlovian learning in sexual behavior and help
to provide the basis for an animal model of the acquisition of deviant sexual arousal in humans.

Sexual behavior of an adult can be influenced by early experi-
ence, inappropriate sexual imprinting (Lorenz, 1965), or rear-
ing conditions (Meaney & Stewart, 1979). Similarly, a number
of observations and anecdotes have accumulated indicating that
learning in mature organisms can also affect sexual responding.
For example, Beach and Fowler (1959), and Dewsbury (1969)
both reported that repeated testing in the same arena led to an
increase in the rate of copulation that may have been the result
of some conditioned sexual arousal related to the apparatus.
Exposure to a mating arena has also been reported to elicit hor-
monal changes similar to those from exposure to an estrous fe-
male rat (Kamel, Mock, Wright, & Frankel, 1975). Although
these studies implicate a Pavlovian mechanism, they were not
designed to examine conditioning effects, and so the conclu-
sions that can be drawn from them are limited. Until recently,
no unequivocal demonstrations of the classical conditioning of
sexual arousal could be found in the literature, especially for
mammalian species.
However, Graham and Desjardins (1980) have reported re-
sults consistent with those of Kamel et al. (1975). They found
that an olfactory conditional stimulus (CS) paired with expo-
sure to a sexually receptive female became as effective as the
female in eliciting increases in luteinizing hormone and testos-
terone levels. Unfortunately, they did not include any behav-
ioral tests to measure the effect of this conditioned endocrine
response on copulation itself, although their data indicate that
at least a concomitant of mating is conditionable.
Some recent experiments from our laboratory have provided
behavioral evidence for the conditioning of sexual arousal by
This research was supported by a grant from the National Research
Council of Canada. John Mitchell is now at Concordia University, and
Helen Findlay is at Smith Klein and French Ltd., Canada.
Correspondence concerning this article should be addressed to Ed-
ward Zamble, Department of Psychology, Queen's University, Kings-
ton, Ontario, Canada K7L 3N6.
excitatory Pavlovian processes (Zamble, Hadad, & Mitchell,
1985; Zamble, Hadad, Mitchell, & Cutmore, 1985). Uncon-
summated sexual arousal functioned as an effective uncondi-
tioned response (UCR) in classical conditioning, just as it had
been found to function as an effective reinforcer in instrumental
learning (Sheffield, Wulff, & Backer, 1951; Whalen, 1961). A
CS paired with this response gained the power to accelerate cop-
ulation, indicating the development of a conditioned arousal re-
sponse.
Although these studies showed the effect of a Pavlovian pro-
cess in sexual responding, the conditioning parameters used
were unusual, for example, the interval between the condi-
tioned and unconditioned stimulus (US) was 10 min. To define
the conditioned arousal effect better, we investigated the results
of some variations in the parameters of the conditioning situa-
tion. We varied the number of acquisition trials and the CS-US
interval to delineate the basic parameters of this paradigm and
investigated the extinction of conditioned sexual arousal. Fi-
nally, familiarity with the cues involved in conditioning was
manipulated by both changing the testing environment and by
using the procedure of latent inhibition. The change in copula-
tion rate should vary with these manipulations similarly to
other conditioned responses (CRs), thus providing further evi-
dence that it is indeed a learned response and not the result of
nonspecific factors.
Experiment 1
The first parameter manipulated was the number of condi-
tioning trials necessary to establish an effect of a CS on sexual
arousal.
Any attempt to relate a conditioned sexual response to what
might occur during naturalistic conditions requires that the CR
be established with relatively few trials. One of the problems
with previous studies claiming to have found evidence for con-
ditioning of sexual arousal with human subjects is that a great
many pairings were needed to show any effects at all. For exam-

403
 404 E. ZAMBLE, J. MITCHELL, AND H. FINDLAY
pie, Rachman (1966) reported using from 24 to over 60 trials
before any CR was visible. It is unlikely that this could occur
outside of a highly contrived laboratory situation.
Although in previous laboratory work we have found signifi-
cant conditioning after 8 or 10 CS-US pairings, the minimum
number of trials required and the development of the CR have
not been investigated. This manipulation should allow a learn-
ing curve to be defined, as would be expected for any CR.
As before, the dependent measure was the time from the first
intromission to ejaculation, ejaculation latency (EL). This is an
index of the rate or intensity of copulation, because the number
of intromissions necessary to reach ejaculation are stereotypi-
cal of rat copulation and relatively inflexible (Adler, 1978; Sachs
& Barfield, 1976). EL was the most sensitive measure of condi-
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tioning in our previous experiments (Zamble, Hadad, Mitchell,
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& Cutmore, 198S), and it is generally accepted as a good mea-
sure of sexual arousal (Sachs & Barfield, 1976).
Method
Subjects. The subjects of this experiment were 40 naive male Long-
Evans hooded rats (Biobreeders Ltd., Canada) selected from an original
population of 56 animals as described below. These animals were sexu-
ally mature but had no sexual experience prior to the experimental situ-
ation. They were individually housed in wire cages with free access to
food and water. The animal room had a reversed light-dark cycle, with
lights off between 8 a.m. and 8 p.m.
The experiment also used 60 female rats of the same strain, which
had been ovariectomized at least 1 month prior to the experiment.
Housing and maintenance were the same as for the males.
Apparatus. Mating trials were conducted in arenas measuring 60
cm wide X 60 cm deep X 50 cm high. Three sides and the floor were
plywood painted flat black, and the front was of removable Plexiglas.
The arenas had no tops and were located in a dimly illuminated room.
For training trials a set of conditioning arenas was used. They were
similar to the mating arenas, differing only in having a wire mesh screen,
which divided the interiors in half. They were also equipped with
weighted tops to prevent animals from climbing over the divider.
For CS presentations the males were placed in white plastic tubs, 28
cm long X 18 cm wide X 13 cm high, with metal grid tops and bottoms
covered with wood chips.
Procedure. The female rats were made sexually receptive by subcu-
taneous injections of 0.1 mg estradiol benzoate (Sigma Chemical Co.,
St. Louis, Missouri) dissolved in 0.1 mL peanut oil, and 1 mg progester-
one (Sigma Chemical Co., St. Louis, Missouri) in 0.1 mL peanut oil
(Bermant, 1964). Trials were conducted 3 to 6 hr after the second (pro-
gesterone) injection to ensure that sexual receptivity had time to de-
velop. The females were maintained on a 5-day injection schedule to
approximate normal cycling and to ensure that some were available for
trials on each day. A single female was normally paired with two consec-
utive males, chosen randomly with the restriction that each male had
an equal number of first and second accesses during training and testing.
Before the start of conditioning, a pretraining test was conducted to
eliminate noncopulators. The male was carried from the home room,
immediately placed into a mating arena with a receptive female, and
they were allowed to copulate until the male ejaculated. When he had
disengaged himself completely from the female and was engaged in post-
ejaculatory grooming, the male was removed from the arena and re-
turned to his home cage. If the male did not achieve an intromission
within 20 min or ejaculation with 30 min, the trial was terminated and
he was eliminated from the experiment. This single test was the extent
of the males' copulatory experience before training began.
Each intromission was recorded, along with the time of the first intro-
mission and the time of ejaculation. In all tests the occurrence of ejacu-
lation was verified by visual inspection of the female for the presence of
a semen plug. The arenas were cleaned with a mild soap and dried after
each trial to minimize the buildup of odors. All trials were conducted
between 10:30 a.m. and 6 p.m.
After the completion of the pretests, five experimental groups of 8
subjects each were formed by stratified random assignment so that EL
scores from the pretests were roughly similar across groups. The groups
differed in the number of conditioning trials to be given, with 3,6,9, 12
and 15 trials, respectively.
On each conditioning trial the male was carried from the home room
to a holding room in his home cage, and then placed into a plastic hold-
ing tub for a 10-min period (CS). This stimulus complex had been used
successfully in previous experiments; it was originally chosen to present
the subject with a variety of cues in several modalities.
Following exposure to the CS, the male was carried to an adjacent
room and placed into one side of a conditioning arena. On the other
side of the arena was a receptive female rat (US), separated from the
male by the wire divider. Thus, the male could see, hear, smell and—to
a limited extent—touch the female, but copulation was not possible.
The animals were left in the arena for 15 min, after which the male was
placed in his home cage and returned to the home room. Subjects were
given single daily conditioning trials on 3 out of every 4 days, until the
requisite number of trials for each group had occurred.
After conditioning there were two copulation test trials. The first was
a CS test trial, on which the male was given the CS period and then
placed in a mating arena with unrestricted access to a female. They
were allowed to copulate until the male reached ejaculation. Thus, this
pastiest was like the pretest except for the presence of the CS, and it
measured the changes in copulation which occurred as the result of
conditioning.
Four days after the CS test, all subjects were given a final test trial
in which the female was presented without the CS. This trial was thus
identical to the pretest.
Results and Discussion
The mean ejaculation latency (EL) values from the various
groups are presented in Figure 1. It can be seen that posttest
scores were only slightly lower than those in the pretest for the
three- and six-trial groups, but with more training, the mean
EL scores for the posttests dropped. In addition, it can be seen
-! 13.0
I 12.0
5 M.O
U
10.0
-J
z 9.0
0
§ 8.0
.J
a 7.0
u
^
s S.O
2
2 5.0
3 6 9 12 15
CONDITIONING TRIALS
Figure 1. Mean ejaculation latencies for each test for each
of the five groups in Experiment 1.
 CONDITIONING OF SEXUAL AROUSAL
that the values for the CS and no-CS tests were very similar for
all groups.
EL scores were transformed logarithmically before statistical
analysis because latency measures typically show a skewed dis-
tribution, and this transformation helps to normalize the distri-
bution (Myers, 1979). The changes apparent across tests were
assessed in an analysis of variance using test (pre-test, CS, and
no-CS tests) as a wi thin-subjects factor and the number of train-
ing trials (groups) as a between-subjects factor.
The analysis revealed a significant effect of tests, F(2, 70) =
23.04, p < .001. This difference across tests was examined with
planned comparisons using the Bonferroni t statistic (Myers,
1979). The differences between the pretest and the CS test were
not close to significance for the three- and six-trial groups (ps >
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.1, for both), whereas the pretest and CS test did differ signifi-
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cantly for each of the other groups (ps < .05). The CS test and
no-CS test did not differ significantly for any of the groups.
A similar analysis was performed on the latency to the first
intromission. There was a decrease in latency from the pretest
to the posttests, resulting in a significant effect of test, F(2,70) -
23.9 l , p < .001. In multiple comparisons we found that for all
groups the CS test was different from the pretest (ps < .05), and
that the CS and no-CS tests did not differ significantly (ps >
.2). Thus, the intromission latency data did not discriminate
between groups. This is consonant with the results of previous
experiments where intromission latency always decreased over
trials but showed no evidence of conditioning (Zamble, Hadad,
Mitchell, & Cutmore, 1985).
The number of intromissions to ejaculation was also ana-
lyzed, and there were no significant differences on any compari-
sons (p > .2). This is in agreement with other literature which
has argued that the number of intromissions to ejaculation is
species typical and resistant to change by most manipulations
(Sachs & Barfield, 1976). The consistency in the number of in-
tromissions indicates that the differences in EL are due to an
increase in the rate of copulation.
Thus, the decreased EL values with nine or more trials indi-
cate an effect of conditioning training; with nine or more pair-
ings of the CS and a receptive female, the CS gained an arousal-
eliciting capacity that was reflected in an increased rate of copu-
lation. Previous studies (Zamble, Hadad, Mitchell, & Cutmore,
1985) have shown significant changes after eight training trials,
so the minimum number to show consistent effects is probably
seven or eight pairings with the present CS and US.
At the same time, the absence of any differences between the
two posttests was unexpected. Whether the posttest was pre-
ceded by the programmed CS or was unsignaled did not seem to
have any effect on copulation. Although the differences between
groups argue for conditioning, the absence of within-groups
differences on the posttests leaves some uncertainty about the
results.
There are several possible explanations for this result. One is
that conditioning did not occur and that the gains apparent on
the posttests were produced by nonassociative factors. However,
other data from our laboratory indicate that with the same CS
and US and using comparable numbers of trials (8-10) EL re-
mains stable if the CS and US are explicitly unpaired (Zamble,
Hadad, Mitchell, & Cutmore, 1985). Thus, it is difficult to ar-
gue that nonassociative or experiential factors alone could ac-
405
count for the EL changes that happen here after 9 or more
paired-conditioning trials.
A more likely possibility is that conditioning did occur but
that conditioned sexual arousal developed not only to the pro-
grammed CS but also to a variety of background cues (removal
from the animal room, transport, the arena, the experimenter,
etc.). Training that was sufficient to establish the programmed
CS was also sufficient to establish various background cues as
CSs which controlled conditioned arousal. This will be exam-
ined further in subsequent experiments; if the background cues
in this experiment were conditioned along with the pro-
grammed CS, then they should respond to parametric manipu-
lations in a manner similar to the CS.
Experiment 2
In the first experiment we traced the development of the CR
across training trials and showed that it is established relatively
quickly. If an effect on sexual arousal is established by classical
conditioning, then it should also be possible to extinguish it
with CS-only presentations. In this experiment we examined
the persistence of the conditioned arousal response under ex-
tinction training.
The establishment of an extinction curve is not only impor-
tant to strengthen the case for conditioning, but it is also rele-
vant to the anomalous aspects of the results of the previous ex-
periment. If we are correct that a CR developed for both the
experimenter-designated CS and background cues, then similar
extinction curves should be found for both.
Method
Subjects. Forty-eight male Long-Evans rats were used, after pretest
selection from an original population of 60 animals. As in Experiment
1,24 ovariectomized female rats were also used.
Apparatus. The mating and conditioning arenas described in Ex-
periment 1 were used, as were the plastic holding tubs.
Procedure. The females were treated with hormone injections as in
Experiment I, and males were given a single copulation pretest as de-
scribed earlier.
All males selected for the experiment were given 10 conditioning tri-
als; on each trial, a 10-min CS was followed by 15-min exposure to a
receptive female, as in the preceding experiment. From earlier results it
was assumed that this would yield a strong CR for all groups. The male
rats were then divided into six groups of 8 subjects by a stratified ran-
domization procedure as before. These groups varied according to
treatment in the next stage, in which extinction trials were given.
An extinction trial consisted of a 10-min placement in the CS holding
cage, followed by a 15-min placement in a conditioning arena with no
female (UCS) present. Thus, the only difference from a conditioning
trial was the absence of the female.
Four of the six groups received extinction trials, with 5, 10, 20, and
30 trials, respectively. The fifth group was given no extinction training
and was tested immediately after conditioning, that is, on the day follow-
ing the last conditioning trial. The last group was a control for the delay
between conditioning and extinction tests: It received no extinction
training but was not tested until the 30-trial extinction group had com-
pleted its trials.
When their allotted training was completed, subjects were given two
copulation test trials, as in the previous experiment. As before, one test
was preceded by the CS, but in the other test the CS was omitted. Two
animals, both in the 30-trial extinction group, failed to meet the crite-
 406 E. ZAMBLE, J. MITCHELL, AND H. FINDLAY
rion of ejaculation within 30 min on one of the tests; they were given a
(successful) replacement trial on the next day, and these scores were
used.
Results and Discussion
Initial comparisons were performed between the two nonex-
tinction groups to establish that conditioning had occurred and
that the delay itself did not attenuate the conditioning. Mean
ejaculatory latency decreased substantially from pretest to post-
tests for both groups; however, as in the preceding experiment
the means for the CS tests were not different from those for the
no-CS tests.
An analysis of variance was performed to verify the apparent
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changes after conditioning, comparing the two control groups
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across the tests. This indicated an overall effect for tests, f\2,
28) = 7.83, p < .01. Orthogonal comparisons showed a signifi-
cant difference between the pretest and posttests, F(l, 2&) =
11.90, p < .01, but the CS test versus no-CS test comparison
did not quite reach significance, F(l,2&) = 3.97, .05 < p < .10.
At the same time, there was no suggestion of an overall between-
groups difference (p > .5).
Thus, it appears that the changes expected with conditioning
did occur and that the delay between training and testing for the
extinction groups did not have any effect on test scores. Given
this, we considered the effects of the extinction trials. Mean
scores for the immediately tested nonextinction group and for
the four extinction training groups are shown in Figure 2.
It can be seen that there was a substantial difference between
pretests and posttests after conditioning trials and that this
difference did not disappear after a few extinction trials. How-
ever, after 30 extinction trials, EL scores returned virtually to
the pretest baselines. An overall analysis of variance was per-
formed on the EL scores. The overall between-groups term ap-
proached significance, ^4,43) = 2. I I , p< .10, and the within-
groups term (tests) was statistically significant, f\2, 86) = 9.52,
p < .001. As in Experiment 1, comparisons were performed
within each group using the Bonferroni t statistic. The pretest
and CS test differed significantly for the 0, 5, and 10 extinction
0 5 10 20 30
EXTINCTION TRIALS
Figure 2. Mean ejaculation latencies for each test for the immediately
tested nonextinction group and the four extinction groups in Experi-
ment 2.
trial groups (ps < .05). However, the differences for the 20 and
30 extinction trial groups did not approach significance (ps >
.1), showing the loss of conditioning that is apparent from in-
spection of the means. As in Experiment 1, the CS and no-CS
tests did not differ significantly for any group.
There were no interesting changes in intromission latency;
posttest scores tended to be lower than pretest values, although
the differences were statistically significant only for the 0 extinc-
tion trial group (CS test), and the 20 extinction trial group (no-
CS test). The number of intromissions to ejaculation was con-
sistent across tests and groups.
Thus, this experiment demonstrates that extinction of condi-
tioned sexual arousal does occur, that is, the facilitation of copu-
lation gained by pairing the CS with a receptive female is lost
after extended experience with the CS alone.
One can also see that the pattern of difference across the
number of extinction trials was very similar for both posttests.
Whether the test was preceded by the programmed CS or by the
background cues alone, the rates of copulation were virtually
identical within each group. Although it is not convincing evi-
dence, this supports the argument that conditioned sexual
arousal was evoked by a complex of stimuli including both the
CS and background cues. During extinction, both types of cues
were weakened in a parallel fashion.
Experiment 3
The final parametric manipulation was that of the CS-US
interval. A brief survey of the conditioning literature reveals
that the optimal CS-US interval varies widely, depending on
the response system being conditioned (Mackintosh, 1974).
There is some indication that at least one of the unconditioned
response systems involved in the present conditioning process
is the slow-acting hypothalamic-pituitary-gonadal axis (Gra-
ham & Desjardins, 1980; Kamel et al., 1975); this is consistent
with the long (10 min) CS duration found to be effective in all
of our previous experiments. As with other response systems
and conditioning paradigms, CS-US intervals that are either
too short or too long should result in poorer conditioning.
Therefore, we included a range of intervals centering on the val-
ues known to produce effective conditioning.
Method
Subjects. From an original pool of 54 rats, 40 males like those in
the previous experiments were selected and used. Females were used as
before.
Apparatus. The arenas and plastic holding tubs were the same as in
Experiments 1 and 2.
Procedure. After selection by pretesting, the males were assigned to
one of five experimental groups that differed according to the length of
the CS-US interval (and also the total CS duration) during condition-
ing. On training trials, after transport to the holding room each male
was placed in the CS tub for either 2,4, 8, 16, or 32 min, according to
group. Males were then given exposure to a female for 15 min in the
conditioning arena, as in previous experiments.
There was a total of 10 training trials for all groups, followed by CS
and no-CS test trials. Exposure to the CS during the CS-posttest was of
the same duration as during training for each group. Testing and other
details of the procedure were the same as in the previous two experi-
ments.
 CONDITIONING OF SEXUAL AROUSAL
CS-TEST
• HO-CS TEST
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CS-US INTERVAL
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tmlnutts)
Figure 3. Mean ejaculation latencies for each test for
each of the five groups in Experiment 3.
Results and Discussion
The mean ejaculatory latency for each group may be seen in
Figure 3. There is an apparent U-shaped function between the
CS-US interval and scores on the posttests, with the shortest
and longest intervals being relatively ineffective in producing
ejaculatory latencies shorter than the pretest levels.
This pattern was confirmed by an analysis of variance. There
was a significant effect of tests, F(2, 70) = 49.24, p < .001, and
a significant Group X Test interaction, F(&, 70) = 3.0, p < .01.
The comparisons between tests revealed that the pretest and CS
test differed significantly for the 4-, 8-, and 16-min groups (ps <
.05), but not for the 2- or 32-min groups (ps > . 1). The no-CS
test followed a similar pattern except that for the 32-min group
the no-CS test also differed statistically from the pretest. The
reason for this is not clear.
In the analysis of intromission latency, we found a significant
effect of tests, (F(2, 70) = 80.71, p < .001; for all groups the
posttests were significantly lower than the pretests (ps < .05).
The analysis of the number of intromissions to ejaculation re-
vealed no significant differences (p > . 1). Because in the first
three experiments and in our other work (Zamble, Hadad,
Mitchell, & Cutmore, 1985) intromission latency has been
found to be relatively insensitive to conditioning, it will not be
reported for the subsequent experiments. Similarly, because the
number of intromissions is stable, as would be predicted from
the literature, it will not be reported further.
The present results of manipulating the CS-US interval
again fit the expectations of a Pavlovian model. Compared with
most other examples of classical conditioning, the effective in-
tervals, approximately 4 min to 16 min, are long.
Taken together, Experiments 1-3 provide strong evidence
that the conditioned arousal response conforms to a condition-
ing paradigm. They define a learning curve, an extinction
curve, and a quadratic CS-US interval function. However, in
these experiments we also failed to find any differential effect of
the designated CS in the posttests.
The differences between pretest and posttests, as well as the
differences between groups, evidence conditioning. Yet the lack
407
of differences between CS and no-CS posttests was unexpected,
and this presents a problem. As explained above, we expect that
cues other than the experimenter-programmed CS were acting
as predictive stimuli for the subjects and that those cues were
responsible for the increased arousal shown in the no-CS tests.
Although this explanation is consistent with the results of Ex-
periments 1-3, any evidence for it so far is indirect. The follow-
ing experiments were designed to provide some direct tests.
Experiment 4
The previous experiments have indicated that arousal may
easily become conditioned to background cues present during
conditioning trials. There were many stimuli in the experimen-
tal situation other than those which we explicitly manipulated
as the designated CS, and some of them were systematically pre-
dictive of the US. In particular, cues associated with removal
from the home room and transportation to the arenas were reg-
ularly part of the complex of cues that predicted exposure to a
female, and local cues in the holding rooms and arenas were
also available. We hypothesize that these cues became capable
of eliciting arousal, just as the designated CS did. This would
account for the lack of differences between the CS and no-CS
tests, because at least some of the original complex of cues were
available during both test trials.
This experiment was a direct test for the development of con-
ditioning to background cues. For some animals, transportation
and arena cues were minimized by conducting the conditioning
trials in their home cages. During testing, the importance of the
remaining background cues was evaluated by testing some of
the subjects in locations different from those in which they had
been trained.
Method
Subjects. As in the preceding experiments, a total of 40 male rats
was used after selection from an original pool of 70 by pretests. In addi-
tion, 24 female rats were used.
Apparatus. The arenas used were the same as those in Experiments
1-3. For some groups, conditioning was carried out in wire mesh alumi-
num cages (90 cm long X 20 cm deep X 20 cm high) in the animals'
home room; these cages were like those in which the subjects were ordi-
narily housed except they were twice the normal size. For conditioning
trials they were divided across the center with wire mesh barriers. Sim-
ilar cages without the dividers were used for test trials when appropriate.
The CS presentation was the same as in Experiments 1-3, except for
the differences in location discussed below.
Procedure. As before, males were selected from the original pool by
means of a copulation pretest and then assigned to one of four experi-
mental groups.
On conditioning trials, two of the groups (home/home and home/
arena) were removed from their home cages and placed into the CS
holding tub in the home room; after the 10-min CS period they were
placed into the oversized conditioning cages (still in the home room)
with a receptive female on the other side of the wire barrier. Subjects in
the other two groups (arena/arena and arena/home) were placed into
the CS tub and then transported to a holding room for the CS period,
after which they were placed into the conditioning arenas, again with a
receptive female on the other side of the wire mesh divider. In either
case, males were returned to their home cages after a 10-min exposure
to the female. Thus, the groups differed primarily in the background
cues that preceded exposure to the female.
 408 E. ZAMBLE, J. MITCHELL, AND H. FTNDLAY
After 10 conditioning trials, copulation tests were given. Two of the
groups (arena/arena and home/home) were tested in the environments
in which they had been trained, the arena and home cage, respectively.
Animals in the other two groups (arena/home and home/arena) were
tested in the alternate environment so that the background cues that
they experienced during the tests did not correspond to those during
conditioning trials.
There were two test trials, with one preceded by a 10-min CS expo-
sure and the other unsignaled. The order of the tests was counterbal-
anced.
Results
There was significant heterogeneity of variance between
groups (p < .01) probably because of the increased variances
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in the groups that had changed background conditions during
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testing. Therefore, individual analyses of variance were calcu-
lated for each group, and multiple comparisons were performed
where appropriate to test for within-groups differences.
Mean ejaculatory latency values for each group and test can
be seen in Figure 4. Group Arena/Arena, comparable to the
groups in Experiments 1-3, showed changes from the original
(pretest) scores during both the CS and no-CS tests. In contrast,
Group Home/Home, trained in the familiar environment of the
home room, showed a substantial CS effect but no change with
the background cues alone on the no-CS test. There appears to
have been little facilitation on either test for Groups Home/
Arena and Arena/Home, both of which received novel com-
plexes of stimuli during testing.
The pattern apparent visually was substantiated by the indi-
vidual group analyses. Considering Group Arena/Arena first,
the overall effect of tests was reliable, F(2, 16) = 12.66, p < .01.
Multiple comparisons showed that the mean value on the CS
test was reliably lower than for the no-CS test (p < .05), and the
mean value for the no-CS test was, in turn, lower than the pre-
Figure 4. Mean ejaculation latencies for each test for the various groups
in Experiment 4. (Each group is designated by the places where training
and testing occurred, either in arenas [A] or in subjects' home cages
[HC]. Thus, Groups A/HC and HC/A were tested in environments in
which they had not had conditioning trials.)
test (p < .01). In Group Home/Home the overall within-groups
term was also significant (p < .05), as was the comparison be-
tween the CS test and the no-CS test (p < .05); however, the
comparison between the no-CS test and the pretest showed no
indication of a difference (p > .5). For the two groups that
changed environments between training and testing, the overall
terms for tests did not approach significance (p > .2), and the
other comparisons were not calculated.
Discussion
The present results demonstrate the role of background and
general environmental cues in the control of conditioned sexual
arousal. When subjects were conditioned in a novel room in a
novel apparatus (arenas), the situational background cues ac-
quired the power to evoke a CR and thus to affect the rate of
copulation. As a result, the additional effect of the experimen-
ter-designated CS was relatively small, although it was still sig-
nificant in this experiment.
In contrast, when conditioning occurred in the familiar envi-
ronment of the animals' home room, the background cues were
ineffective, although the designated CS was still a reliable signal
for exposure to the female; as a result, a much clearer discrimi-
nation was apparent. This explanation, of course, assumes that
the cues of the home environment had been subject to latent
inhibition, as will be tested in the next experiment.
The results for the two groups with cues changed from train-
ing to testing again show the power of the background stimuli.
Even though the designated CS was unchanged for subjects in
the home/arena and arena/home groups, it had little effect, indi-
cating that it was seen by subjects as only one part of the com-
plex which was effecting the CR. It appears that the various
situational cues were relatively much more salient than our des-
ignated CS in the effective CS complex. Only when we made
some effort to eliminate the effects of the background, as in
Group Home/Home, did our designated CS become a sole
strong cue for conditioned arousal.
Experiment 5
The previous experiment showed that changing environmen-
tal cues between training and testing disrupts the conditioned
facilitation of copulation and thus indicates the importance of
background cues in the learning situation. However, our expla-
nation of the results depends on the occurrence of latent inhibi-
tion to the background cues in the group which was both
trained and tested in the home room. In this experiment we
tested for the development of latent inhibition directly, by giving
varying numbers of preexposures to background stimuli before
conditioning (Lubow, 1973). The results should support the in-
ferences made about the previous study and should also provide
another demonstration of the importance of background cues
in conditioned sexual arousal.
The pretest environment was changed so that the rats would
have no experience with the background cues in the experimen-
tal situation prior to the preexposure trials. This was achieved
by using the animals' home cages for the pretest. Thus, any in-
terference from a pairing of copulation and the background
stimuli was effectively removed. In addition, some of the ani-
 CONDITIONING OF SEXUAL AROUSAL 409

mals were allowed copulatory experience before training began, SEXUALLY INEXPERIENCED
in order to reduce the novelty of the UCS and to investigate the
effects of sexual experience on the development both of latent
inhibition and of conditioned sexual arousal generally.

Method
Subjects. Forty-eight male rats, selected from an original pool of 64
animals, were used. Twenty-eight female rats were also used. The male
rats were housed singly in double-sized colony cages; otherwise, condi-
tions were the same as before.
Apparatus. Mating and conditioning arenas were the same as those
used previously.
The CS environment was a cage 24.5 cm long X 20 cm deep X 20 cm
high, inside a sound-attenuating chamber 76.5 cm long X 46 cm deep X o
1- SEXUALLY EXPERIENCED
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51 cm high. To the front and center of the chamber, 7 cm above the top
< 12
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of the cage, there was a 7.5-W light bulb. Power reached the light 1^ '1 t
through a GEM Flasher, causing it to flash every 5 s. => 10
Procedure. The pretest used to eliminate noncopulators was con- O

i
ducted as in previous experiments except that it was performed by in- < 8
troducing a female into the home cage of the male. LU
Males that copulated successfully on the pretest were assigned to one 6 _T_
T
_L
of six groups, with 8 rats in each group. The groups differed according
2 4
to the number of preexposures to the background that they received
^ y,
and as to whether or not they were allowed further sexual experience
before the classical conditioning procedure, as explained below.
Three groups (sexually inexperienced) were allowed no further copu-
latory experience until the posttests and were given either 0, 15, or 30
preexposure trials. On a preexposure trial the male was carried to the
5
2 -

0
i^
15
//
30

conditioning arena in his home cage and placed into one side of the PREEXPOSURES
^

conditioning arena (without a female present) for 15 min. This exposed

the male to most of the stimuli involved in conditioning trials, that is,
handling, transportation cues, arena cues, and other cues in the condi-
tioning room.
The three other groups were given sexual experience prior to the stim-
ulus preexposures. Males in these groups had a female rat introduced
into their home cages on each of 5 different days. They were allowed
to copulate, and the female was removed after the male ejaculated. To
minimize the importance of time cues, these trials were given at random
times between 11 a.m. and 6 p.m. If a male failed to ejaculate within 30
min on any occasion, the female was removed and a replacement trial
was given the next day. If a male failed to copulate on a total of three
experience trials or on two consecutive trials, he was eliminated from
the experiment. The choice of five copulation experiences (or six includ-
ing the pretest) followed from the finding that the male rat's sexual per-
formance reaches asymptote after about six opportunities to copulate
to ejaculation (Sachs & Barfield, 1976; Zamble, Hadad, & Mitchell,
1985). Thus, for these three groups performance on the posttests should
not exhibit any further gains after conditioning owing to practice alone.
After experience trials, the preexposure trials for the sexually experi-
enced subjects were conducted the same as for the inexperienced males,
with the three groups receiving either 0, 15 or 30 preexposures, respec-
tively.
After sexual experience and preexposure training, all subjects were
given 10 conditioning trials. These were conducted as previously, with
a 10-min placement in the CS box followed by a 15-min placement
in the conditioning arena, with noncopulatory exposure to a receptive
female. Following completion of the conditioning trials, the male rats
were given two copulation test trials, a CS test and a no-CS test, as in
previous experiments. The order of the tests was counterbalanced.
Results
During the sexual experience trials it was necessary to give
four replacement trials. These failures to copulate to ejacula-
I 1 PRETEST
[ [ C S TEST
£^CS TEST
Figure 5. Mean ejaculation latencies for each test for each of the groups
in Experiment 5. (The groups with sexual experience before condition-
ing are shown in the bottom panel, and those without experience trials
are above.)
tion within 30 min were distributed among 3 animals, one from
each of the three experience groups. The rat that failed to copu-
late on two occasions was not eliminated from the experiment
because the failures were not on successive trials.
As expected, there was a considerable overall decrease in ejac-
ulatory latency from the pretest to the posttests, as shown in
Figure 5. The decrease in the CS test values was roughly equiva-
lent for all groups, with no differences apparent from either ex-
perience or the number of preexposures. However, on the no-
CS tests the amount of change from the pretest levels was visibly
reduced for the 30-preexposure groups, regardless of the level
of sexual experience. Thus, extensive preexposure to the back-
ground cues lessened the postcondition! tig effect of those cues
on copulation.
An analysis of variance was computed, comparing pre-CS,
CS, and no-CS tests within groups and both the level of sexual
experience and number of preexposures between groups. The
differences that are visually apparent were confirmed by a sig-
nificant difference for tests, f\2, 84) = 93.34, p< .001, and by
 410 E. ZAMBLE, J. MITCHELL, AND H. FINDLAY
a Test X Preexposure interaction which was close to statistical
significance, F(4, 84) - 2.34, p < .07. Interestingly, the effect
of sexual experience and the several terms involving it did not
approach significance (ps < .2). Multiple comparisons, calcu-
lated using the Bonferroni t statistic, showed that for both the
sexually inexperienced and the sexually experienced groups
that received 30 preexposures the Cs test and no-CS test differed
significantly (ps < .05). None of the other groups showed any
reliable differences between the posttests. Thus, 30 preexpo-
sures to the background cues reduced the ability of the back-
ground cues alone to increase arousal, although the power of
the experimenter-designated CS was left intact.
Discussion
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This document is copyrighted by the American Psychological Association or one of its allied publishers.
These results again demonstrate the differential control of
classically conditioned sexual arousal by the programmed CS
and the complex of background stimuli. When exposure to a
receptive female was signaled, copulation was facilitated. When
the signal was omitted on a test trial, the only available cues
were those of the background. If the background cues had been
relatively novel during training, they were effective in evoking
arousal and facilitating copulation. However, if the background
cues had been encountered extensively in the past with no im-
portant consequences, that is, if they had been subject to latent
inhibition, their postconditioning effect was reduced.
However, the no-CS test performance did not return to pre-
test levels even in the groups showing latent inhibition. Such a
return to pretest levels has been found with other experiments
(see Experiments 2 and 4, above). This suggests that despite the
preexposures the background cues still gained some control
over sexual arousal. In other experiments in our laboratory as
many as 60 preexposures to the background stimuli have been
used, and the resulting values were similar to those from the 30-
preexposure group reported here. Thus, the amount of stimulus
experience necessary to block conditioning entirely may be very
large and certainly more than with other responses investigated
previously. As few as 10 CS preexposures retard acquisition of
a leg flexion CR in sheep and goats (Lubow & Moore, 1959),
and significant latent inhibition is often reported with 16-20
preexposures of the to-be-conditioned stimulus (Lubow, 1973).
In the present case, no effects were seen with 15 or fewer preex-
posures. The number of preexposures that are necessary to at-
tentuate conditioning to the background may be added to other
indications of the robustness of this type of conditioning.
Finally, it is perhaps surprising that the effect of prior sexual
experience was not evident here. The sexually experienced
groups had a marginally lower overall copulation time and
slightly lower within-group variances, but neither of these ap-
proached statistical significance. At the most, sexual experience
may have attenuated the effect of preexposures, as the differ-
ences between CS and no-CS tests were slightly less in experi-
enced subjects than in the inexperienced group.
General Discussion
These experiments demonstrate again the classical condition-
ing of sexual arousal as measured by facilitation of copulation
in the male rat. The CR, which we have called "conditioned
sexual arousal" affects the rate or intensity of copulation. In
no instance was there a conditioning effect on the number of
intromissions required to reach ejaculation. Therefore, we con-
clude that the changes we saw were owing to an increase in the
rate of intromissions and not to a reduction in the total amount
of copulatory behavior. Because the speed and intensity of the
consummatory response have been widely used as measures of
motivation, we can argue that the present conditioned response
reflects an increase in the level of sexual arousal. Until data
using more direct measures of arousal are available, describing
the CR as conditioned sexual arousal seems to be warranted.
The present results show that we are dealing with a robust
example of Pavlovian conditioning. The conditioned response
develops rapidly over a relatively small number of trials and oc-
curs to a wide variety of external stimuli as CSs, including un-
controlled background cues. Finally, very extensive preexpo-
sure training is required to make a stimulus ineffective as a sub-
sequent CS.
Graham and Desjardin (1980) have provided evidence that
the endocrine system can be conditioned; conditioning can in-
crease luteinizing hormone and testosterone levels. The present
experiments, along with others which we have reported (Zam-
ble, Hadad, Mitchell, & Cutmore, 1985), provide evidence that
a similar conditioning procedure can establish behavioral sex-
ual arousal as a CR. Thus, Pavlovian conditioning can influence
both the endocrine system and copulatory behavior, although at
present there are no data indicating how or if these conditioning
effects interact.
In the last experiment we found that additional copulatory
experience prior to conditioning had little effect on the power
of the CS. Thus, subjects do not need to be sexually naive for
conditioning to occur. This adds to the established generality of
the effect as an example of Pavlovian control of a vital response
system. It is also important for any attempt to apply our results
to naturalistic settings or to use them as a model for the acquisi-
tion of deviant sexual arousal in human beings.
One area in which the results reported here appear to differ
from our previously published findings (Zamble, Hadad,
Mitchell, & Cutmore, 1985) is in the conditioning to back-
ground cues. The effects of background were not visible before
because experimental designs compared paired-conditioning
groups with unpaired controls. In the first three experiments
reported here, control procedures were based on within-sub-
jects comparisons because of the large, reliable individual
differences found with rat copulatory behavior and the desir-
ability of using subjects as their own controls whenever possible
(Dewsbury, 1969). Fortunately, the parametric variations in the
effects made it clear that conditioning was present, despite the
lack of differences between CS and no-CS tests, and Experi-
ments 4 and 5 clarify this interpretation by demonstrating con-
trol of conditioned arousal by background cues. In the end, we
have another demonstration of the power of the conditioned
arousal paradigm: When the appropriate US and conditioning
parameters are employed, the problem is not how to generate
the effect but rather how to maintain an acceptable level of stim-
ulus control.
In summary, we can say that the conditioning of sexual
arousal in the rat is both powerful and simple to establish. There
are obvious differences between the copulatory patterns of rats
 CONDITIONING OF SEXUAL AROUSAL 411

and humans, as well as differences in the presence and likely
influence of various controlling factors. However, with the assis-
tance of an animal model it should be possible to investigate the
effects of learning on human sexual behavior more productively
than in the past.
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