Extended paced mating tests induces conditioned place preference without affecting sexual arousal

One way to evaluate sexual arousal is by measuring approach behavior to sexual incentive stimuli. In our case we measure approach behavior to an originally non-preferred compartment which is associated with the physiological state induced by mating. This change of preference indicative of a positive affective (reward) state can be evaluated by conditioned place preference (CPP). We have shown that the CPP induced by paced mating is mediated by opioids. The administration of opioids also induces a reward state. The present study was designed to compare the rewarding properties of paced mating and a morphine injection. One group of females was allowed to pace the sexual interaction before being placed in the non-preferred compartment. In alternate sessions they received a morphine injection before being placed in the preferred compartment. In another group of females, the treatments were reversed. Only the females placed in the originally non-preferred compartment after paced mating changed their original preference, suggesting that paced mating induces a positive affective, reward, state of higher intensity than a morphine injection of 1 mg/kg. In a second experiment we determined if females allowed to pace the sexual interaction for 1 h would still developed CPP. No change in preference was observed in the females that mated for 1 h without pacing the sexual interaction. On the other hand, females that received between 10 and 15 paced intromissions as well as females that paced the sexual interaction for 1 h developed a clear CPP. The second experiment demonstrated that pacing is rewarding even in an extended mating session in which the females received around 25 intromissions and several ejaculations. These results further demonstrate the biological relevance associated with the ability of the female to space coital stimulation received during mating. This positive affective state will contribute to increase sexual arousal the next time a rat finds an appropriate mate.     

Serum 5 alpha-androstane-3 alpha, 17 beta-diol increases in response to paced coital stimulation in cycling female rats

The cycling female rat spontaneously regulates (paces) the amount and timing of cervical-vaginal stimulation received during mating. Female pacing of coital contacts increases the ability of vaginal intromissions to induce luteal function and to abbreviate the period of behavioral estrus. In the experiments reported here, we examined whether paced mating results in alterations in serum steroid concentrations, which might contribute to these processes. In Experiment 1, estradiol (E2), progesterone (P), testosterone (T), 5 alpha-dihydrotestosterone (DHT), and 5 alpha-androstane-3 alpha, 17 beta-diol (3 alpha-androstanediol, 3 alpha-diol) were measured in sera obtained from independent groups of animals between 0 min and 12 h after mating onset. Levels of 3 alpha-diol were significantly higher at 30 min in females pacing coital stimulation (Paced) than in females receiving mounts-without-intromission (Mounts-Only) or solitary exposure to the test chamber. Serum T, E2, P, and DHT did not differ between these groups at any time. P and 3 alpha-diol levels declined significantly for all groups across the 12-h post-treatment period. In Experiment 2, serum 3 alpha-diol measured over 3h after mating was compared in females receiving Paced stimulation with females receiving temporally unregulated coital stimulation (Non-Paced), or Mounts-Only. At 20 min, serum 3 alpha-diol concentrations were significantly higher in Paced than in Non-Paced and Mounts-Only females. In Experiment 3, Paced and Non-Paced stimulation did not differentially effect the proportion of females becoming pseudopregnant/pregnant. The selective increase in serum 3 alpha-diol in females that pace matings is discussed with regard to the known inhibitory effects of 5 alpha-reduced androgens on sexual receptivity.     

Female rats develop conditioned place preferences for sex at their preferred interval

Female rats engage in approach and avoidance behaviors directed toward the male to "pace" the rate of copulation. These pacing behaviors result in a pattern of vaginocervical stimulation that triggers a neuroendocrine reflex that is important for pregnancy to result from insemination. Each female rat has a preferred pacing interval, and females develop conditioned place preferences for paced sex versus nonpaced sex. Research from this laboratory has reported that extracellular dopamine concentrations in striatum and nucleus accumbens are greater in female rats that are engaging in paced sex compared with those engaging in nonpaced sex. Furthermore, females who have males removed at their preferred intervals during a copulatory bout show extracellular dopamine concentrations comparable to females engaging in paced sex. It is unclear, however, whether they would also develop a conditioned place preference for sex under such conditions. This experiment was designed to address this question. Female rats had six exposures each to a chamber in which they engaged in nonpaced sex and a chamber in which they engaged in paced or preferred pacing interval sex. Following conditioning trials, females were tested for a conditioned place preference. The findings indicate that female rats develop conditioned place preferences for paced sex and for sex in which the male is removed at her preferred interval. This suggests that sexual behavior is reinforcing to female rats when their preferred interval is achieved, whether or not they are actively controlling the rate of copulation.     

Lesions confined to the ventromedial hypothalamus decrease the frequency of coital contacts in female rats

Ovariectomized female rats received either bilateral radiofrequency lesions of the ventromedial nucleus of the hypothalamus (VMH) or control treatments and were tested for copulatory activity following either estrogen (E) alone, or E plus progesterone (P) administration. In separate experiments the females were tested in two testing apparatuses both of which allowed the test females to control their contacts with sexually active males. One of the testing apparatuses also allowed the females to control their contacts with sexually inactive males and ovariectomized females. Females receiving VMH lesions engaged in fewer coital contacts with sexually active males than sham-operated females in the E plus P condition. Lesioned females also tended to spend less time with sexually active males than did sham operates in both the E and E plus P hormonal conditions. The VMH-lesioned females did not differ from the sham-operated females in the ability to display lordosis during the coital contacts or the frequency and duration of visits to the inactive males or ovariectomized females. The sham-operated females did have some transitory alterations in copulatory behavior in comparison to unoperated control females.     

Appetitive and Consummatory Sexual Behaviors of Female Rats in Bilevel Chambers: I. A Correlational and Factor Analysis and the Effects of Ovarian Hormones

This study investigated measures of sexual behavior displayed by female rats in bilevel chambers, the statistical relationships among the measures, and their dependency on hormone priming. Normative data from a standard 35-min test of sexual behavior were gathered from 82 fully primed sexually experienced Long-Evans females and subjected to multiple correlational and factor analyses. Several consummatory measures of copulation were related significantly, whereas appetitive level changing was statistically independent of consummatory measures. Factor analyses were conducted using orthogonal rotations of correlational matrices derived either from (a) measures of female behavior alone or (b) measures of female and male behavior together. The first analysis revealed five factors that accounted for 84% of the intersubject variance: Receptivity, Pacing, Appetitive Level Changing, Lordosis Reflex, and Solicitation. The second factor analysis with male data included revealed seven factors that accounted for 95% of the intersubject variance: Pacing, Copulatory Rate, Mount Count, Receptivity, Appetitive Level Changing, Solicitation, and Lordosis Reflex. Subsequently, subsets of these females were maintained on different steroid priming regimens (oil, low estrogen, high estrogen, high estrogen and progesterone) prior to a standard test of sexual behavior. Although the expression of all sexual behaviors required estrogen priming, appetitive level changing, solicitation, and pacing required progesterone for their full expression. Finally, appetitive level changing developed following hormone treatment alone, regardless of whether the females received access to sexually active males, inactive castrated males, or other females. Use of bilevel chambers allows complex patterns of sexual behavior to be observed in female rats and may thus facilitate the identification of neurochemical or endocrine mechanisms associated with different aspects of female sexual motivation and performance.     

Progesterone inhibits female courtship behavior in domestic canaries (Serinus canaria)

We studied copulation solicitation display (CSD) responses to playback in photostimulated female canaries given systemic injections of progesterone. Eight females received injections of 0.1 mg of progesterone dissolved in olive oil during their first breeding cycle and were untreated during their second breeding cycle; eight females received only the oil vehicle during their first breeding cycle and received no treatment during their second breeding cycle. The injections were performed every second day during 15 days, after the onset of nest building. Progesterone treatment resulted in a significant increase of plasma progesterone which in turn provoked an inhibition of females' CSDs and decreased the size of the clutch. During the first breeding cycle, progesterone-treated females had lower CSDs and egg-laying scores than did control females. During the second breeding cycle, when females received no treatment, no differences emerged between the two groups. The suppressive effect of progesterone on female sexual responses was observed as soon as 48 h after the beginning of the treatment. We propose that progesterone plays a key role in mediating the transition from active female courtship behavior to sexual refractoriness in this species. Suppressive effects of progesterone on female sexual behavior have been previously described in lizards as well as in rodents. Our data are consistent with the hypothesis of Godwin et al. (J. Godwin, V. Hartman, M. Grammer, and D. Crews, Horm. Behav. 30, 138-144, 1996) which proposed that the decrease in sexual behavior following plasma progesterone increase represents an evolutionarily conserved mechanism in the regulation of female sexual behavior.     

Hormonal and testing conditions for the induction of conditioned place preference by paced mating

The ability to control or pace the sexual interaction has important physiological and behavioral consequences for the female rat. Paced mating favors reproduction and induces a positive affective state as revealed by conditioned place preference (CPP). In the present experiment we evaluated: 1) If paced mating induces CPP in naturally cycling females; 2) If females developed a positive affective state if they paced the sexual interaction through a 1- or a 3-hole pacing chamber; 3) If females that mate with the same male without pacing the sexual interaction develop CPP. In the first experiment intact females were divided in 4 different groups; 2 paced the sexual interaction until receiving 1 or 3 ejaculations; the other 2 groups mated, without pacing the sexual interaction, until receiving 1 or 3 ejaculations. Only the group that paced the sexual interaction until receiving 3 ejaculations developed a positive affective state. In experiments 2 and 3 hormonally treated ovariectomized females were used. In experiment 2 females were allowed to pace the sexual interaction through a 1- or a 3-hole pacing chamber: A clear positive affective state was induced in both testing conditions. Finally, in experiment 3 females did not develop CPP for non-paced sex despite the fact that they mated with the same male in the conditioning sessions. These results demonstrate that the pattern of vaginocervical stimulation that the females received by engaging in approach and avoidance behaviors to pace the sexual interaction can induce a positive affective state in naturally cycling females. They also confirm the existence of a threshold of vaginocervical stimulation for paced mating to induce CPP in female rats.     

Independence of the differentiation of masculine and feminine sexual behavior in rats.

Male rats received Silastic implants of the aromatase inhibitor, 1,4,6-androstatriene-3, 17-dione (ATD), on days 2–10 of life. Controls received blank implants. There were no differences in the masculine sexual behavior of ATD and control males when they were tested as gonadally intact adults. In contrast, even without exogenous hormone treatment, nine of 14 ATD males exhibited lordosis behavior, whereas only one of 12 controls did so. In addition, during a sexual preference test in which access was provided to both a sexually receptive female and to a stud male, there was no difference in the proportions of ATD ($\text{11}\text{14}$) and control ($\text{7}\text{12}$) males that copulated with the stimulus female; however, seven of the ATD males also exhibited feminine sexual behavior including some instances of solicitation. Only one of the control males showed any lordosis behavior. In general, all animals spent more time with the stimulus female than with the stud male. At the termination of preference testing, all animals were castrated and then tested twice for feminine sexual behavior under exogenous estradiol benzoate and progesterone. All of the ATD males showed lordosis behavior with a mean lordosis quotient (LQ) of 85; and 11 of the 14 also showed solicitation behavior. Only five of 12 control males exhibited lordosis ($\text{X}\text{?}\text{LQ}\text{= 59}$) and only one showed solicitation behavior. These results indicate that the propensity of males to show feminine sexual behavior can be manipulated independently of the capacity for masculine sexual behavior. Moreover, our results suggest that the process of defeminization may occur primarily postnatally in rats since treatment during that period results in substantial increments in later feminine sexual behavior including solicitation behaviors.     

Ultrasonic Vocalizations in Rat Sexual Behavior

Ultrasonic vocalizations are very conspicuous during rat matingactivity. Two types of calls are produced by both sexes. Thefirst, brief complex calls with the main frequency centeredabout 50 kHz, occur primarily in conjunction with solicitationand mounting activity. The second type of call is the long,22 kHz whistle which is emitted mainly by the male during thepostejaculatory refractory period, but also by both male andfemale at other times during the copulatory sequence. The occurrenceof ultrasonic vocalizations is correlated with sexual motivationof rats. Males emit more 50 kHz calls before successful matingtests than before tests in which they fail to ejaculate. Furthermore,more vocalizations are emitted by the pair prior to intromissionsthan prior to mounts without intromission. Just before ejaculationthere is a large increase in the rate of calling and, at times,transition by the male to calling at 22 kHz. This latter eventmay represent physiological dearousal by the male. Followingejaculation, the male characteristically emits 22 kHz vocalizationsand exhibits a sleep-like EEG pattern. The function of the postejaculatoryvocalization may be to enforce separation between the matingpair, while at the same lime maintaining contact between thepartners. Fifty kHz calls, on the other hand, prime and facilitatesexual responsiveness of the female. Tape recorded vocalizationsof mating rats facilitate solicitation behavior of estrous femalesin the presence of castrated males, and such females also showa preference for these sounds in a "Y" maze. Deafening of femalesdoes not affect their normal pacing of copulatory contacts,but it drastically reduces their solicitation behavior. Thestudies summarized in this paper lead us to conclude that ultrasonicvocalizations play a major role in the integration of reproductiveactivity in the Norway rat, Rattus norvegicus.     

Coital stimuli controlling luteinizing hormone secretion and ovulation in the female ferret

A series of experiments focused on the masculine coital behaviors controlling pituitary luteinizing hormone (LH) secretion and reflex ovulation in the estrous female ferret. An initial experiment investigated which coital stimuli from the male are required to induce ovulation. It was found that corpus luteum formation, which served as an index of ovulation, occurred in estrous female ferrets only if the male achieved a penile intromission. Neck gripping, mounting, and pelvic thrusting behavior without intromission by the male failed to induce ovulation. A second experiment investigated the timing and magnitude of the coitus-induced LH surge associated with ovulation. Blood was obtained via jugular catheters from estrous females in various mating situations. Plasma LH concentrations were measured by a heterologous radioimmunoassay that was validated for use in the ferret. A significant surge in plasma LH occurred only when an intromission was achieved by the stud male. Plasma LH was significantly elevated 2.0 h after the introduction of the male, peak values were reached 6.0 h later, and this elevation lasted on average 5.7 hours (5/5 females). No LH rise occurred in 2/2 female ferrets in which only neck gripping, mounting, and pelvic thrusting, but no intromission, were allowed to occur. The ferret mating pattern and the resultant LH response differ from those seen in three other induced ovulators (cat, vole, and rabbit) in which the male's intromission latency and duration are much shorter than in the ferret, and in which a distinctive peak in plasma LH often occurs within 1 h after mating.     

Only Self-Paced Mating Is Rewarding in Rats of Both Sexes

When rats are mated in a traditional mating chamber (with one male and one female) in which the male dictates the pace of the copulatory sequence, males develop a reward state as evaluated by conditioned place preference (CPP). In this mating situation no reward state is induced in females. However, when female rats are able to control (pace) the rate of sexual stimulation, thereby reducing the aversive consequences associated with mating, a clear CPP is observed. In the present study the CPP paradigm was used to determine whether if the reinforced state induced by coital interactions in male rats can be maintained when females pace the sexual interaction. Adult male and female rats were mated in one of two different conditions: (1) where subjects were able to pace their coital interactions or (2) where subjects were not able to pace their sexual contacts. The results showed that when males had control over the sexual interaction they developed a clear place preference while males that mated with females that paced their coital contacts did not develop CPP. Similarly, only females that were able to pace their sexual contacts developed place preference. These results suggest that coital interactions in males, as well as in females, can induce a reward state only when they are able to control the sexual interaction. Under seminatural conditions sexual behavior in rats is highly promiscuous, they mate in groups and repeatedly change partners in the middle of copulation. This behavioral sequence allows both, male and female to control the rate of sexual interaction, assuring the induction of a reward state outlasting the actual performance of coital responses.     

Frustrated appetitive foraging behavior, stereotypic pacing, and fecal glucocorticoid levels in snow leopards (Uncia uncia) in the Zurich Zoo

This study hypothesized that permanently frustrated, appetitive-foraging behavior caused the stereotypic pacing regularly observed in captive carnivores. Using 2 adult female snow leopards (Uncia uncia), solitarily housed in the Zurich Zoo, the study tested this hypothesis experimentally with a novel feeding method: electronically controlled, time-regulated feeding boxes. The expected result of employing this active foraging device as a successful coping strategy was reduced behavioral and physiological measures of stress, compared with a control-feeding regime without feeding boxes. The study assessed this through behavioral observations and by evaluating glucocorticoid levels noninvasively from feces. Results indicated that the 2 snow leopards did not perform successful coping behavior through exercising active foraging behavior or through displaying the stereotypic pacing. The data support a possible explanation: The box-feeding method did not provide the 2 snow leopards with the external stimuli to satisfy their appetitive behavioral needs. Moreover, numerous other factors not necessarily or exclusively related to appetitive behavior could have caused and influenced the stereotypic pacing.     

Vaginocervical stimulation attenuates the sensitization of appetitive sexual behaviors by estradiol benzoate in the ovariectomized rat

The acute administration of estradiol benzoate (EB) to the ovariectomized (OVX) rat induces low levels of lordosis while sexually appetitive behaviors (e.g., hops, darts, solicitations) are absent, yet the repeated administration of EB results in a behavioral sensitization in which lordosis is potentiated and sexually appetitive behaviors are induced. We have shown that repeated copulation attenuates the sensitization of appetitive sexual behaviors. Here, we assessed which component of male stimulation during copulation is involved in the attenuation. On 8 occasions, sexually experienced OVX Long–Evans rats were treated with 10 μg EB and 48 h later assigned to one of six groups that differed in their experience on intermediates tests (2–7). One was given repeated access to a male (EB/Male), and another was placed in the copulation chamber alone (EB/Alone) on intermediate tests. Three groups were given one of three somatosensory stimuli by the experimenter: manual flank stimulation (FLS), clitoral stimulation (CLS), or vaginocervical stimulation (VCS). Finally, the control group was left undisturbed in the animal care facility (ACF). Sexual behaviors were measured on Tests 1 and 8. VCS received from the experimenter (VCS) or from the male during copulation (EB/Male) attenuated the magnitude of the sensitization of appetitive sexual behaviors compared with those that were not brought to the testing rooms (ACF), and the effect was most pronounced on sexual solicitations. These results suggest that VCS received during penile intromission inhibits the sensitization of sexually appetitive behaviors by repeated administration of EB. As such, repeated administration of EB may oppose those mechanisms that induce estrous termination, perhaps by sensitizing inhibitory processes within the ventromedial hypothalamus that typically prevent the display of sexual behaviors (i.e., by facilitating disinhibition).     

Non-intromissive mating stimuli are sufficient to enhance sexual behaviors in ovariectomized female rats

When ovariectomized/adrenalectomized female rats, injected with subthreshold doses of estradiol are given copulatory stimulation by a male rat at half hour intervals, the level of lordosis gradually increases over the course of a few hours. We tested the hypothesis that paracopulatory behaviors (behaviors that occur repetitively prior to and between mounts), also generally considered to be heavily dependent on progesterone, are enhanced by this stimulation as well. We have reported previously that the enhancement of copulatory behavior is dependent to a large extent on intromissive stimulation by the male. In the present study, mating stimulation induced high levels of paracopulatory behaviors, as well as lordosis. Surprisingly, though, and in contrast to previous findings, this increase was seen not only in rats receiving intromissive stimulation, but in those receiving non-intromissive stimulation as well. Furthermore, intromissive stimulation induced high levels of rejection behavior. In a subsequent experiment, experimenter-induced, mechanical stimulation increased only rejection behaviors, not copulatory behavior. The results collectively demonstrate that, under the conditions used in these experiments, non-intromissive stimulation is sufficient for inducing both copulatory and paracopulatory behaviors in estradiol-primed rats. However, under the conditions used in these studies, intromissive stimulation increases rejection behaviors.     

Male Syrian hamsters demonstrate a conditioned place preference for sexual behavior and female chemosensory stimuli

Sexual behavior is a natural reward for many rodent species, and it often includes chemosensory-directed components. Chemosensory stimuli themselves may also be rewarding. Conditioned place preference (CPP) is one paradigm frequently used to test the rewarding properties of a range of stimuli. Males and females of several rodent species show a CPP for sexual behavior; however, it is currently unknown whether sexual behavior can induce a CPP in male Syrian hamsters. As male Syrian hamsters are an animal model commonly used for investigation of the neurobiology of sexual behavior, understanding the rewarding components of sexual stimuli will better direct future research on brain regions and neurotransmitters involved in these behaviors. Experiment 1 tested the prediction that male hamsters show a CPP for sexual behavior. Female chemosensory stimuli are essential for the display of sexual behavior in male hamsters; however, the rewarding properties of female chemosensory stimuli contained in vaginal secretions (VS) are uncertain. Therefore, experiment 2 tested the prediction that male hamsters show a CPP for VS. This study is the first demonstration that both sexual behavior and VS induce a CPP in male hamsters. Thus, female chemosensory stimuli are a natural reward in a species that is dependent on these stimuli for reproductive fitness.     

Cuticular lipids and odors induce sex-specific behaviors in the male cricket Gryllus bimaculatus

Male crickets display sex-specific (e.g., mating and agonistic) behaviors towards conspecific individuals. One of the key signals for these behaviors is the chemical substance on the cricket body surface. In the present study, we analyzed female and male cuticular substances in behavioral assays. Antennal contact stimulation using female forewings elicited a mating behavior in males, while that using male forewings elicited an agonistic behavior in males. Thin-layer-chromatographic and other techniques analysis showed that saturated cuticular lipids were present in both female and male cuticles and that unsaturated lipids were present only in the male cuticle. Filter papers soaked with saturated or unsaturated cuticular lipids were applied to antennae of male crickets. Males showed mating behavior in response to stimulation with saturated lipids from both females and males but showed avoidance behavior in response to stimulation with male unsaturated lipids. Because cuticular lipids did not induce agonistic behavior in males, we collected odors from male crickets and found that these odors induced agonistic behavior in males. Therefore, we concluded that the key signals for mating, avoidance and agonistic behaviors of male crickets are comprised of at least three different components, saturated and unsaturated cuticular lipids and male odors, respectively.     

Previous sexual experience alters the display of paced mating behavior in female rats

The present study tested whether the display of paced mating behavior in female rats over four weekly tests is affected by sexual experience and whether test parameters, i.e., ending the test based on time or number of stimulations received, influence behavioral changes. In Experiment 1A rats with nonpaced sexual experience returned to the male more quickly overall compared to sexually naïve rats in a 30-min test of paced mating behavior. In Experiment 1B, rats received four weekly 30-min tests with one, different, male rat partner each week. Over the four tests, rats returned to the male significantly more quickly after intromissions, but significantly more slowly after ejaculations. Experiment 2A tested whether sexual experience would influence paced mating behavior in tests with a 15-intromission end criterion and the male replaced after ejaculation. Rats tested weekly under 15-intromission test conditions returned to the male significantly more quickly after intromissions, but no behavioral change was observed after ejaculations. When those same rats were given a 30-min test of paced mating behavior (Experiment 2B), they returned to the male significantly more slowly after ejaculations. Collectively, these data show that sexual experience influences the display of paced mating behavior in female rats and that the test parameters interact with sexual experience to influence the nature of the changes. Sexual experience may facilitate behaviors that promote reproductive success in female rats.     

Twenty-four hour activity budgets and patterns of behavior in captive ocelots (Leopardus pardalis)

Activity budgets of captive ocelots (Leopardus pardalis) were assessed from over 547h of observational data obtained from six ocelots; two females at the Dallas Zoo (Dallas, TX), two females at the Caldwell Zoo (Tyler, TX) and a male and female at the Fossil Rim Wildlife Center (Glen Rose, TX). Data were examined for the percentage of active behaviors exhibited during the day and nighttime hours; temporal patterns of active, pace, exploratory and marking behavior, and for significance in pacing behavior between pre- and post-feeding times. The captive cats had a bimodal pattern of active behavior similar to field studies of wild ocelots, except that the timing of the active peaks were closer to the diurnal hours for the captive cats. The captive ocelots were less active than wild ocelots, and more diurnal. Also, the captive cats exhibited stereotypic pacing. When the percentage of time of active behavior was assessed for each cat, a strong variation between individuals and institution was not seen. Pacing behavior was highest prior to the feeding times for the cats. In assessing patterns of behavior, peaks in marking and exploratory behavior in the cats did not occur at the same time as the peaks in active behavior. However, we did see institutional differences in the pattern of exploratory and marking behavior, which may have been influenced by differing management practices.     

Conditioning and sexual behavior: a review

Sexual behavior is directed by a sophisticated interplay between steroid hormone actions in the brain that give rise to sexual arousability and experience with sexual reward that gives rise to expectations of competent sexual activity, sexual desire, arousal, and performance. Sexual experience allows animals to form instrumental associations between internal or external stimuli and behaviors that lead to different sexual rewards. Furthermore, Pavlovian associations between internal and external stimuli allow animals to predict sexual outcomes. These two types of learning build upon instinctual mechanisms to create distinctive, and seemingly "automated," patterns of sexual response. This article reviews the literature on conditioning and sexual behavior with a particular emphasis on incentive sequences of sexual behavior that move animals from distal to proximal with regard to sexual stimuli during appetitive phases of behavior and ultimately result in copulatory interaction and mating during consummatory phases of behavior. Accordingly, the role of learning in sexual excitement, in behaviors that bring about the opportunity to mate, in courtship and solicitation displays, in sexual arousal and copulatory behaviors, in sexual partner preferences, and the short- and long-term influence of copulatory experience on sexual and reproductive function is examined. Although hormone actions set the stage for sexual activity by generating the ability of animals to become sexually excited and aroused, it is each animal's unique experience with sexual behavior and sexual reward that molds the strength of responses made toward sexual incentives.     

Role for prenatal estrogen in the development of masculine sexual behavior in the male ferret

Previous studies have shown that neonatal exposure to testosterone is essential for coital masculinization in male ferrets. In the present experiments, masculine sexual behavior was diminished in male ferrets by prenatal exposure to drugs which inhibited estrogenic stimulation of the brain. Similarly timed prenatal treatments with testosterone failed to masculinize the behavior of female offspring. We hypothesize that prenatal exposure of the male ferret to estrogen, derived from the neural aromatization of circulating androgen, may sensitize the developing brain to the subsequent masculinizing action of testosterone shortly after birth.