Female tail wagging enhances sexual performance in male goats

Preference testing has shown that sexually experienced male goats choose females that are tail wagging, a behavior that may function as both attractivity and proceptivity, over those that are not. We hypothesized that exposure to females expressing high rates of tail wagging would arouse males, increasing sexual performance. Tail wagging rate could be manipulated because we have shown previously that flutamide treatment increases the frequency of tail wagging in estrous goats. Sexually experienced males observed different stimuli for 10 min before a 20 min sexual performance test (SPT). The stimuli were an empty pen (MT), or groups of three females that were all estrous (E), non-estrous (NE), estrous + flutamide (E_F) or non-estrous + flutamide (NE_F). During the stimulus observation period, tail wagging was recorded. During SPT, frequencies and latencies of sexual behaviors were recorded. E_F females displayed the most tail wagging. Viewing E_F females before SPT increased the number of ejaculations attained by males and decreased the latencies to first and second ejaculation, as well as the inter-ejaculatory interval. Viewing estrous females (E and E_F) before SPT decreased the latency to first mount, as compared to non-estrous females (NE and NE_F). We conclude that male goats are sexually aroused by tail wagging. This study and previous work demonstrate that tail wagging functions as both attractivity and proceptivity in goats.     

The relationship of male-male mounting to mate choice and sexual performance in male dairy goats

Rearing of male farm animals in unisexual groups has been implicated as a factor contributing to the failure of many males to breed as adults. The present study examines the relationship of male-male mounting in yearling dairy goats to subsequent mate preferences and sexual performance.Twenty-four sexually inexperienced male dairy goats, representing the Alpine, LaMancha, Saanen and Toggenburg breeds, were observed for male-male mounting in their home enclosure and then tested for mate choice and sexual performance when exposed to male and female (estrous and diestrous) stimulus animals. Their sexual behavior was compared with 7 adult goats with previous breeding experience.In the mate choice-sexual performance tests, 4 sexually inexperienced goats (17%) were sexually inactive, 6 (25%) mounted both male and female stimulus animals and 14 (58%) mounted only the female stimuli. Mate choice and sexual performance of the 20 sexually active males was not related to the number of male-male mounts initiated or the number of different males mounted in their home enclosure. However, the goats that received the greatest number of mounts in their home pen tended to be bisexual (would mount both male and female stimulus animals) in the mate choice tests. Males that were sexually inactive in mate choice-sexual performance tests repeatedly mounted the same male during home pen observations. Except for ejaculation frequency, the sexual performance of the sexually naive and experienced goats was similar. Goats of the Saanen breed were favored recipients of mounts from other males. There was no relationship between the number of male-male mounts performed and received.It was hypothesized that the reproductive failure of many male farm animals reared in all-male groups may be more closely related to the formation of specific sexual attachments to other males rather than the frequency with which they exhibit homosexual behaviors.     

Effects of sexual stimulation on the sexual performance of rams

The sexual performance of bulls and male goats is improved if they are allowed to view the hetero-sexual behavior of other males as a prelude to mating. The purpose of the following study was to determine whether sexual stimulation enhances the sexual performance of rams. In Experiment 1, 11 sexually experienced ram lambs ( 9 months of age) and 18 sexually experienced yearling and 2-year-old rams were individually exposed to 4 unrestrained, hormone-induced estrous ewes for 60 min after viewing the courtship and mounting behaviors of a male conspecific for 20 min (two tests) and in the absence of stimulator animals (two tests). In contrast to the results with bulls and bucks, the rams were hardly influenced by the sexual stimulation treatment. Latencies for first mount and first ejaculation were shorter for sexually stimulated ram lambs; otherwise, treatment differences were negligible.

A similar follow-up experiment was administered to 12 mature rams using restrained females in the sexual performance tests. Again, treatment differences were minor.

It was concluded that sexual stimulation does not functionally enhance the sexual performance of rams. Species differences in response to sexual stimulation are discussed in terms of female sexual behaviors that may result in a selective (competitive) advantage to males that are stimulated to locate and mate with females early in the estrous period.     

Copulatory behavior of sexually inexperienced male voles paired with proestrous females

The copulatory behavior in sexually inexperienced male voles aged 9-13 weeks old was observed under dim red illumination. Proestrous females were used as stimulus voles in copulatory behavior tests. A 30-min test session was recorded. All males showed at least one ejaculation within 30 min. In addition, the pattern of copulatory behavior in male voles was compared to that reported previously in male hamsters. The frequencies of mount, intromission and ejaculation in male voles were similar to that in male hamsters but there were significant differences in the latencies of mounting, intromission, ejaculation and post-ejaculation between the two species.     

Homosexual behavior in male goats is more frequent during breeding season and in bucks isolated from females

Male homosexual behavior is observed in many ruminant species; seasonality and isolation from females may affect the display of male–male sexual behavior. Our objectives were to determine whether: 1) homosexual behavior has a seasonal pattern in male goats (Capra hircus); 2) isolation from female goats influences the frequency of the display of homosexual behavior; and 3) the frequency of homosexual behaviors is related to testosterone concentration. Bucks were housed in a pen adjacent to another pen with 3 estrous goats (FC group) or in complete isolation from females (IF group). Homosexual behaviors and testosterone concentrations were recorded in October, December, February and May. Frequencies of penile display and flehmen increased in May and were observed more frequently in the IF than in the FC group. Ano-genital sniffing was more frequent in February and May and was more frequent in IF than in FC bucks. Lateral approaches were more frequent in May, in which period this was more frequently observed in IF bucks. Mount attempts and mounts were only observed in IF bucks. Testosterone concentrations increased from December to February and decreased in May, although they were still greater than in October and December. In May IF bucks had greater testosterone concentrations than FC bucks. Testosterone concentrations were only related to the number of lateral approaches in IF bucks in December. In conclusion, homosexual behavior was more frequent during the late breeding season than during the early and the non-breeding seasons, and in males isolated from females than in males housed near estrous females. Testosterone concentrations appear to be unrelated to homosexual behavior.     

Neurological effects of aromatase deficiency in the mouse

In the brain, the conversion from androgen into estrogen is an important process for the differentiation of the brain function in male rodents. The aromatase is expressed in some nucleus of the brain. To assess the functional significance of the aromatase gene in development and activation of sex-specific behavior, we analyzed behavioral phenotypes of the aromatase knockout (ArKO) male mice. ArKO males obviously decreased their fertility and showed deficits in male sexual behavior including mount, intromission and ejaculation. Noncontact penile erection was not significantly affected by defect of the aromatase gene. A reduction of aggressive behavior against male intruders was also observed in ArKO males, while they tend to exhibit aggression toward estrous females during male copulatory tests. Moreover, the infanticide toward the pups was observed in the ArKO males, whereas characteristic parental behavior, but not infanticide was observed in wild-type males. These results indicate that aromatase gene expression is a critical step not only for motivational and consummatory aspects of male sexual behavior, but also for aggressive and parental behaviors in male mice.     

The relationship of male-male mounting to the sexual preferences of young rams

The purpose of this study was to determine the relationship between the frequency of male-male mounting of young rams in all-male groups and their subsequent preferences for female vs. male sexual partners. Mount interactions were monitored in the home pen of 29 rams reared in 1984 and 25 rams reared in 1985. At ∼ 8–10 months of age, and prior to copulatory experience, individual rams were simultaneously exposed to two unfamiliar estrous ewes and two unfamiliar rams. The four stimulus animals were restrained and cloth covers were placed over their perinea to prevent copulations. The Binomial Test was used to verify sexual preferences.Of the 44 rams that were sexually active in the sexual performance tests, 30 (68.2%) met the criterion for female preference, 4 (9.1%) preferred males and 10 (22.7%) exhibited no preference (i.e., were bisexual in orientation). There was little relationship between the frequency of male-male mounting by rams in all-male rearing groups and their subsequent sexual preferences. The majority of young rams prefer to mount ewes, even prior to copulatory experience, but in the absence of females will often mount other males.     

Parity of female goats does not influence their estrous and ovulatory responses to the male effect

The objective of the present study was to determine whether parity is a factor that influences the estrous and ovulatory responses of female goats when they are stimulated by males that show increased sexual activity. To stimulate sexual activity, four adult male goats were subjected to photoperiodic treatment for 2.5 months comprising long days, with the treatment commencing on 1 November. On 14 April at 1900 h, a group of multiparous females (n = 21) and a group of 16 months-old nulliparous females (n = 19) were exposed to four bucks (two per group) for 15 days. Throughout the study period, the estrous behavior of these female goats was detected twice on a daily basis. Ovulations of the female goats were determined by ecography on days 7 and 18 after exposure to males. The sexual behavior of males was recorded twice every day from 0800 to 0900 h and from 1730 to 1830 h during the first 4 days after introduction in the pen of females. The total cumulative proportion of multiparous females that had ovulations (100%) and displayed estrous behavior (100%) during the 15 days of exposure to males did not differ (P > 0.05) from that of nulliparous females (100% and 95%, respectively). The interval between introduction of males and onset of estrous behavior did not differ (P > 0.05) between multiparous (1.9 ± 0.1 days) and nulliparous (1.7 ± 0.2 days) females. The proportion of females displaying a short estrous cycle was greater (P < 0.05) in multiparous (13/21, 62%) than in nulliparous (5/19, 26%) females. Duration of these shorter than typical estrous cycles did not differ (P > 0.05) between groups (multiparous: 5.2 ± 0.3 days, nulliparous: 4.5 ± 0.1 days). The number of anogenital sniffings was greater (P < 0.001) in males exposed to nulliparous than in those exposed to multiparous females. In contrast, the number of mounting attempts was greater (P < 0.01) in males that were introduced to multiparous than in those that were introduced to nulliparous does. The number of flehmen, nudging, self-marking with urine, and mounts was not different (P > 0.05) between males that were in contact with multiparous and nulliparous females. These results indicate that regardless of parity, female goats respond to male introduction if they are stimulated by males that were previously exposed to artificial long days to increase their sexual behavior.     

Lack of sexual experience does not reduce the responses of LH, estrus or fertility in anestrous goats exposed to sexually active males

We investigated whether LH secretion, estrous behavior and fertility would differ between sexually inexperienced and experienced anestrous goats exposed to the males. Male goats were rendered sexually active during the reproductive rest season by exposure to 2.5 months of artificial long days. Two groups of anovulatory sexually inexperienced and sexually experienced does were exposed to males during 15 days (n = 20 per group). LH pulsatility was determined every 15 min from 4 h before to 8 h after introducing males (Day 0). Estrous behavior was recorded twice daily. Pregnancy rates were determined on Day 50. Fertility was determined at parturition. Male sexual behavior was registered on days 1 and 2 during 1 h. Before introducing the males, the number of LH pulses did not differ between groups. After introduction of the males, all females increased their LH pulsatility, but the number of pulses did not differ between sexually inexperienced and experienced goats. The proportion of females displaying estrous behavior with a high pregnancy rate and fertility did not differ between inexperienced and experienced goats. The sexual behavior of the males did not differ significantly between those interacting with sexually inexperienced or experienced goats. We conclude that goats can show substantial endocrine and reproductive responses to males, even in the absence of previous sexual experience, when sexually active bucks are used.     

A contextual definition of male sexual arousal

Sexual arousal is a construct without a widely shared definition. Historically, sexual arousal has usually referred to a central physiological state, but there has been much less agreement on its relation to motivation, emotion, and - for males - penile erection and ejaculation. Many behavioral and physiological measures have been used as operational definitions of sexual arousal, but the relation of the measure to arousal is often assumed rather than tested. For men, penile erection in the presence of erotic stimuli has been considered the most reliable and valid indicator of sexual arousal. The adoption of analogous criteria is recommended for research on other male mammals in order to establish a minimal basis for inferring that they are sexually aroused. That is, sexual arousal should be inferred only when penile erection is observed in a sexual context. A sexual context is provisionally defined as an environment that tends in most reproductively active males of the species to provoke further sexual stimulation, e.g., copulation or self-stimulation to ejaculation. Erection occurring outside of a sexual context, as during REM sleep or from injection of drugs, is not grounds for inferring arousal. Conversely, males engaging in behavior directed toward estrous females may be sexually motivated, but in the absence of erection, the males should not be assumed to be sexually aroused. Implications of other erection-context interactions are also considered. Adoption of these more conservative criteria for inferring sexual arousal may promote greater precision in identifying the physiological systems mediating this hypothetical construct.     

Sexual behavior in brown capuchins ( Cebus apella )

We observed sexual behavior patterns in two captive groups of Cebus apella.We obtained data on intersexual solicitations and intersexual competition. The length of proceptivity cycles by females averaged 18.8 ± 1.2 days; this corresponds with published reports of physiological measures of ovulatory cycles. Females initiated sexual interactions with males significantly more than males initiated interactions with females, and female solicitations increased in frequency over the duration of displayed periods of proceptivity. In contrast, male solicitations of females were infrequent and did not change significantly throughout female proceptive phases. Each female solicited the same male throughout a given proceptive phase. We observed no evidence of male-male or female-female competition. Males seem to respond to female sexual behavior directed toward them, rather than initiating sexual interactions themselves. Even so, male sexual interactions with females followed only a small proportion of solicitations.     

Sexual behavior in brown capuchins (Cebus apella)

We observed sexual behavior patterns in two captive groups of Cebus apella.We obtained data on intersexual solicitations and intersexual competition. The length of proceptivity cycles by females averaged 18.8 ± 1.2 days; this corresponds with published reports of physiological measures of ovulatory cycles. Females initiated sexual interactions with males significantly more than males initiated interactions with females, and female solicitations increased in frequency over the duration of displayed periods of proceptivity. In contrast, male solicitations of females were infrequent and did not change significantly throughout female proceptive phases. Each female solicited the same male throughout a given proceptive phase. We observed no evidence of male-male or female-female competition. Males seem to respond to female sexual behavior directed toward them, rather than initiating sexual interactions themselves. Even so, male sexual interactions with females followed only a small proportion of solicitations.     

Potential contribution of prenatal estrogens to the sexual differentiation of mate preferences in mice

The neural mechanisms controlling sexual behavior are sexually differentiated by perinatal actions of gonadal hormones. We recently observed using female mice deficient in alpha-fetoprotein (AFP-KO) and which lack the protective actions of AFP against maternal estrogens, that exposure to prenatal estrogens completely defeminized their potential to show lordosis behavior in adulthood. Therefore, we determined here whether mate preferences were also affected in female AFP-KO mice. We observed a robust preference for an estrous female over an intact male in female AFP-KO mice, which were ovariectomized in adulthood and subsequently treated with estradiol and progesterone, whereas similarly treated WT females preferred the intact male over the estrous female. Gonadally intact WT males preferred the estrous female over the male, but only when visual cues were blocked by placing stimulus animals behind opaque partitions. Furthermore, when given the choice between an intact male and a castrated male, WT females preferred the intact male, whereas AFP-KO females showed no preference. Finally when given the choice between an estrous female and an ovariectomized female, WT males preferred the estrous female whereas AFP-KO females preferred the ovariectomized female or showed no preference depending on whether they could see the stimulus animals or not. Taken together, when AFP-KO females are tested under estrous conditions, they do not show any male-directed preferences, indicating a reduced sexual motivation to seek out the male in these females. However, they do not completely resemble males in their mate preferences suggesting that the male-typical pattern of mate preferences is not solely organized by prenatal estrogens.     

Importance of the signals provided by the buck for the success of the male effect in goats

Under temperate and subtropical latitudes, ewes and goats display a reproductive seasonal pattern and their sexual activity during the anestrous period can be stimulated and synchronized by the introduction of males in the group, which is called the "male effect". The response of females to the male effect in the middle of the anestrous season is weak or absent. This failure may be due to the inability of the female to respond to males, as a result of a refractoriness of the female to the male stimulus. But, it may also be due to a low quality stimulus provided by the male which is, as the females, in seasonal rest. We tested this latter hypothesis in seasonally inactive goats kept under subtropical conditions by comparing the use of males with their sexual behavior stimulated or not by photoperiodic treatments. Treated males were able to induce estrous activity of females during the whole anestrus season. We have also determined that previous separation of the males and continuous contact during teasing are not absolute requisites when active bucks are used. While odor from the male and its sexual behavior play a primary role in inducing ovulation, vocalizations appear to facilitate the display of the does' estrous. It remains to be determined to which extent these conclusions apply under temperate latitudes and with more seasonal breeds.     

Male reproductive condition is the limiting factor of efficiency in the male effect during seasonal anestrus in female goats

Two experiments were conducted to determine whether the failure of males to induce sexual activity in goats during seasonal anestrus is due to unresponsiveness of females to male stimulus or insufficient stimulation from males. In the first study, one group of males (sexually inactive, SI; n = 4) was kept under natural photoperiod while the other (sexually active, SA; n = 4) was subjected to 2.5 mo of long days (16L:8D) and received 2 s.c. implants of melatonin. Two mo later, 2 different flocks of anovulatory goats previously separated from bucks were exposed to either SI (n = 34) or SA (n = 40) bucks. Progesterone assays and estrous behavior were used to determine ovarian and behavioral responses of the females to teasing. Of the goats exposed to SI males, only 2 ovulated, and none showed estrous behavior during the 35 days of the study. In contrast, all females (40 of 40) in contact with SA males ovulated and showed at least one estrous behavior during the first 11 days following male introduction (P < 0.001). Overall, 38 of 40 females stimulated with SA bucks were diagnosed pregnant at Day 35, according to progesterone assay (versus 0 in SI-treated group: P < 0.001). To control for a possible difference of responsiveness between flocks, the experiment was repeated 1 yr later using a single flock of goats divided into 2 groups. Again, over the first 14 days, 1 of 33 goats showed estrous behavior in the SI-treated group versus 27 of 33 in the SA-treated group (P < 0.001). Therefore, treating bucks with long days and melatonin increased their teasing capacity to induce sexual activity in females during anestrus. These results indicate that the absence of response to teasing at this time of the year is not due to female unresponsiveness, but to insufficient stimulation from the male.     

Stimulus conditions influencing self-enurination,genital grooming and flehmen in male goats

The stimulus conditions which influence the frequencies of self-enurination, genital grooming and flehmen by domestic male goats (Capra hircus L.) were studied in two experiments in which the sexual arousal and performance of the subjects were varied either naturally or artificially.Sexually aroused males exhibit more frequent self-enurination, genital grooming and flehmen than sexually inactive males. In natural mating, self-enurination and genital grooming are exhibited mainly when sexual behaviors are naturally inhibited, as during the post-ejaculatory refractory period. Frequencies of self-enurination and genital grooming can be artificially increased by enhancing sexual arousal in situations that preclude physical contact with females, such as when observing other males copulate or when restrained in close proximity to a female immediately following ejaculation.     

Mating Tactics in Response to Costs Incurred by Mating with Multiple Males in Wild Female Japanese Macaques

We investigated the costs of mating with multiple males in terms of feeding time, traveling distances, sexual proceptivity, and male aggression, for wild female (Macaca fuscata yakui) on Yakushima Island, Japan. We analyzed all-day focal sampling data from 7 females during the mating season (Sept.-Nov. 1996). On days when estrous females copulated with multiple males, they decreased their feeding time to half that of anestrous days, traveled longer distances, showed more proceptive sexual behaviors and received more aggression from subordinate males than on days when they copulated with only the 1st-ranking male. On days when females copulated with only the 1st-ranking male, they showed no difference in feeding time with that of anestrous days, and expended less effort than the above mating pattern because of short traveling distances, diminished sexual proceptivity and a lower frequency of aggression received. The results suggest that the costs of estrous vary according to female sexual proceptivity and the number and social status of mating partners. Female Japanese macaques exhibit a mixed mating strategy over prolonged estrous periods, which may provide females with opportunities to maximize the benefits of copulating with multiple males and to minimize the costs of estrus by mating with only the 1st-ranking male. During an estrous cycle, females may be adjusting efforts for reproduction and survival; i.e., mating vs. feeding.     

Dose-response and time-response relationships between progesterone and the display of patterns of receptive and proceptive behavior in the female rat

The relationship between administration of progesterone and the display of patterns of receptive (response to the male) and preceptive (female initiated) sexual behavior was examined in ovariectomized, estrogen-primed female rats in a “restrained male” test situation. It was found that the degree of receptivity and proceptivity displayed was directly proportional to progesterone dose and time from progesterone injection (up to 4.5 hr). Higher progesterone doses and longer period of time from progesterone injection (up to 4.5 hr) were both associated with shorter latencies to return to the male following intromission and ejaculation. Receptivity could be induced with estrogen alone but progesterone was required for the display of proceptivity and higher doses of progesterone were needed to effect increases in proceptivity relative to receptivity. Proceptive behavior also occurred in a narrower time range than did receptive behavior. Receptivity alone is characterized as the lowest degree, and receptivity plus proceptivity as the highest degree, of expression of the total behavior pattern of the estrous female rat. Receptivity and proceptivity together constitute a continuum of estrous responsiveness. Increasing the progesterone dose from 0 to 200 μg, and increasing the latency from progesterone injection from 0 to 4.5 hr, were associated with increasing degree of expression of the total behavioral continuum.     

Sexual arousal,the coolidge effect and dominance in the rat (rattus norvegicus)

Male rats were ranked in dominance orders, were allowed to copulate to exhaustion and then were tested for sexual rearousal in five different treatment conditions. Novel females rearoused 60 per cent of the males to ejaculation. Five-minute encounters with dominant or subordinate males stimulated 33 per cent to ejaculation with a familiar female. Fifteen per cent ejaculated with the familiar female after a 5 min rest and 15 per cent after an encounter with an unknown male. The degree of rearousal was not related to fighting or to mounting during male-male encounters, nor did the dominant males show a higher degree of sexual potency.     

Male effect in seasonally anovulatory lactating goats depends on the presence of sexually active bucks,but not estrous females

A study was conducted in subtropical northern Mexico (26 degrees N) to determine whether the presence of estrous females can improve the response of seasonally anovulatory goats to the introduction of bucks in the group. The induction of estrous activity was studied in three groups of anovulatory lactating goats during seasonal anestrus. These females were of the Mexican Creole breed. In the control group (sexually inactive (SI), n = 20), two control (SI) bucks exposed to normal seasonal daylength variations were used. In the second group (SI + E, n = 20 + 3), two control males were also used, but in addition, three females of the group were in estrus at the time of male introduction. In the third group (sexually active, SA + E, n = 19 + 4), anovulatory females were exposed to two bucks made sexually active by exposure to 2.5 months of long days (16L:8D) followed by two subcutaneous 18 mg melatonin implants, and four estrous females were also present when introducing the bucks. In all groups, males were introduced on 15 March and estrous detection was conducted twice daily for 15 days. The sexual activity of the bucks was observed from 08:00 to 10:00 h during the first five days of exposure to females. More females displayed estrous behavior in the first 15 days following the introduction of the males in the SA + E group (18/19) as compared with the SI or SI + E groups (2/20 and 0/20, respectively; P < 0.001). No difference was observed between the two latter groups. Thirteen females of SA + E group showed a second estrus between days 6 and 11 (short estrous cycle duration: 5.4 +/- 0.4 days). By contrast, in the SI group none showed a second estrus. The sexual behavior of the males in the SA + E group was greater as compared with that of the males in SI and SI + E groups (over 80% of the total sexual activity recorded in the three groups; P < 0.001). By contrast, no differences were found between SI and SI + E males. These results indicate that the presence of estrous females alone at the time of buck introduction is not sufficient to induce an adequate stimulation of seasonally inactive males. The use of sexually active bucks is necessary to induce reproductive activity in anovulatory females, whereas preparation of the bucks with long days followed by melatonin implants allows them to gain such a capacity.