A Juvenile Hormone analogue enhances homosexual behaviour in female‐deprived males of the migratory locust

In laboratory colonies of crowded migratory locusts Locusta migratoria (L.), homosexual behaviour (i.e. males mounting other males) is commonly observed. Female‐deprived males of Locusta migratoria migratorioides R. & F. mount each other in a characteristic mating position, often forming a group of several insects. When allatectomized males are placed together with intact, female‐deprived males, the former are usually mounted by the latter, demonstrating some degree of control involving the corpora allata over homosexual behaviour. This may be related to the positive effects of Juvenile Hormone (JH) or Juvenile Hormone analogues (JHAs) in enhancing male sexual behaviour, as is shown in some other insects. In the present study, a potent JHA (i.e. pyriproxyfen) is injected into a group of young, crowded L. m. migratorioides males, and an equivalent group of control males is maintained in a separate cage. Both groups are deprived of females. Three times a week, during 2‐h observation periods, JHA‐injected and control males are placed together in the absence of females, and homosexual mountings, per group and per individual, are recorded every 10 min. Observations are performed for 10 weeks. Analysis of these data, including the time spent in mounting behaviour, the percentage of individuals within a group involved in the behaviour during the observation periods and the identity of both partners, reveals that the JHA‐treated males show a more intense homosexual mating behaviour than control males in all quantified parameters. This is the first report of the enhancement of homosexual behaviour by an endocrine factor in insects.     

Contextual determinants of female-female mounting in laboratory rats

Female-female mounting is widespread among mammalian species, but little is known about the proximal function of this behaviour. While such mounting is often regarded as a 'masculine' trait, its widespread occurrence may indicate that it serves specific functions within the context of female-female social behaviour. We valuated female mounting behaviour in Long-Evans rats in standard observation chambers and in a seminatural enclosure. Under these conditions, we examined a number of potential factors that might influence mounting, including the oestrous cycle, social hierarchy, familiarity and male presence. The female's mounting was not influenced by her own oestrous cycle, but did vary with the oestrous cycle of the stimulus female. Socially dominant females mounted significantly more than subordinate females, and mounting by the dominant female was most frequent when the subordinate female was sexually receptive. Females mounted (and fought with) unfamiliar females significantly more than they did with familiar cagemates. Female-female mounting was dramatically reduced when males were present. Further testing showed that female mounting did not affect the induction of the progestational state of pregnancy, suggesting that female mounting does not function as a pseudomale behaviour that can substitute for genital stimulation provided by the male. Based on these data, female mounting does not appear to function as a sexual behaviour per se, but may serve as a form of female social behaviour related to maintenance of the female's social status within female groups. In this regard, the results of this study suggest that female mounting is part of the normal female's complex behavioural repertoire and does not necessarily reflect masculinization of some underlying neural substrate. Copyright 1999 The Association for the Study of Animal Behaviour.     

Gonadal hormones and sexual behavior in groups of adult talapoin monkeys (Miopithecus talapoin).

Three heterosexual groups of six to eight monkeys were studied; all females were ovariectomized, whereas males were either intact or castrated. Aggressive hierarchies were evident in all groups, with females generally outranking males. When females were treated with estradiol, all males looked more frequently at the latters' sexual skin swellings, but only one male who was both dominant and intact copulated with them. Thus, either castration or low rank resulted in decreased levels of sexual behavior in male talapoins. The sexual behavior of dominant castrated males was restored by testosterone therapy, whereas subordinate castrates never copulated, even after large doses of testosterone, though penile erections and ejaculatory reflex (during masturbation) were restored. Following removal of a dominant male, the sexual behavior of the next male in rank was restored, provided he was not castrated and untreated. In contrast to males, female talapoins showed no consistent correlation between their rank and sexual activity. Estradiol therapy was without overall effect upon the frequency of female mounting behavior, though some females mounted and presented to one another more often. Estradiol treatment also caused females to present to males more frequently, but only to those that were sexually active (i.e., who mounted females).     

Mounting Interactions between Female Japanese Macaques: Testing the Influence of Dominance and Aggression

The goal of the work reported here was to determine whether female Japanese macaques (Macaca fuscata) participated in same-sex mounting interactions during homosexual consortships to communicate about asymmetries in their dominance relationships and to reduce aggression. Focal data were collected during 21 homosexual consortships involving 14 females living in a captive, mixed-sex group of 37 individuals. We identified eight types of mounts, one solicitation used specifically to request to mount (hands-on-hindquarters solicitation), two solicitations used specifically to request to be mounted (hindquarter and back presentations), and one behavior employed to facilitate mounts-in-progress (clasping). We tested whether dominant consort partners (1) mounted more and (2) requested to mount more than their subordinate partners and whether subordinate consort partners (1) requested to be mounted more and (2) facilitated mounts-in-progress more than their dominant partners. Finally, we examined whether mounting was temporally linked to the onset of aggressive interactions between consort partners and whether it functioned to defuse incipient aggression. None of these predictions was supported. All types of mounts, mount solicitations, and clasping occurred bi-directionally within consort dyads. Mutual sexual attraction and gratification provided the proximate motivation for these mounting interactions and, in turn, for the formation and maintenance of their homosexual consortships.     

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.     

The reproductive behavior of golden monkeys in captivity (Rhinopithecus roxellana roxellana)

The reproductive behavior of two captive harem breeding groups of golden monkeys was observed over one mating season and compared to non-seasonal reproductive behavior. Observation of over two thousand sexual interactions suggests that 1) females were responsible for 95% of all sexual solicitations (proceptive behavior), of which the typical pattern was prostration, while males were responsible for only 5%; 2) 17% of female solicitations were interrupted by another female; 3) 52% of female solicitations resulted in mounting by males; 4) the frequencies of solicitations and receipt of mounts and ejaculations varied greatly among females; 5) the frequency of ejaculations, the ratio of ejaculations to mountings, and ejaculation patterns were different between two males, and 6) the golden monkey is a seasonal breeding animal.     

野生川金丝猴一个全雄青年猴群的同性爬背行为

本研究采用行为取样方法,首次对秦岭野生川金丝猴全雄青年群中的同性爬跨行为进行报道。研究中我们共观察到21 次同性爬背行为,其平均持续时间为5.53 ± 3.11s;雄性同性爬背发生前的行为中以玩耍行为(47.62% )和3 种不同姿势的邀配模仿行为(42.85% )为主;而同性爬背行为发生后则主要为彼此间的相互理毛行为(47.62% )和玩耍行为(23.81% )。对比先前婴幼猴时期的相关研究结果,青年猴同性爬背行为前邀配模仿资势的多样化在一定程度上体现了野生状态下雄性川金丝猴个体青少年阶段的性行为发育进程。考虑到川金丝猴全雄群的社会结构,青年猴同性爬背行为具有巩固社群稳定和加强个体关系的功能。此外,研究中观察到76.19% 的同性爬背行为发生在昼间休息期之后的第一个小时内,这可能是“睡眠与觉醒循环”对性激素的反调节作用所致。     

Social dominance and female reproductive behaviour in rhesus monkeys (Macaca mulatta)

The relationship between dominance rank and female sexual behaviour was examined in rheusus monkeys (Macaca mulatta) living in a social group. High-ranking females engaged in copulatory series as frequently as lower-ranking females. Furthermore, lower-ranking females copulated with as many available males as did high-ranking females. Social rank did appear to influence the pattern of sexual activity exhibited, in that copulatory series were more often initiated by the higher-ranking animal of the mating pair. Copulatory series involving high-ranking females were characterized by more mounts by males and were longer in duration. Higher-ranking animals interfered more often with copulatory series involving other animals, but such interference was not effective in preventing completion of the series. These data indicate that any reproductive advantage conferred to high-ranking females is not the result of sexual competition in social-living rhesus monkeys.     

人工饲养长江江豚的同性性行为和自慰行为

雌雄性之间的性行为对鲸类的生存和繁衍有着极其重要的意义。然而不具有繁殖功能的同性性行为和自慰行为在多种鲸类中存在。因此对鲸类非繁殖性性行为研究,将有助于深入认识该物种性行为特点和规律,并解决室内饲养可能存在的问题以及促进可控性人工繁殖的成功。本研究以人工饲养的7头长江江豚为研究对象,定义并描述了14种性行为。其中,同性之间的性行为定义为同性性行为,雌雄个体单独的性行为定义为自慰行为。雄性同性性行为发现于所有定义的性行为中,但雌性同性性行为仅在7种性行为中观察到。同性性行为（3633次）占总性行为（5563次）的65.31%,自慰行为（254次）占4.57%。同性性行为中,雄性之间的同性性行为（3438次,94.63%）远多于雌性同性性行为（195次,5.37%）。两头有父子关系的江豚（阿福和淘淘）之间的雄性同性性行为最多（1921次,55.88%）,雄性江豚之间的同性性行为可能存在结盟竞争关系或教育练习作用。两头青少年雌豚（F7和F9）之间的同性性行为最活跃（117次,60.00%）,雌性江豚之间的同性性行为很可能具有结伴交流的功能。     

Courtship behaviour in Japanese macaques during heterosexual and homosexual consortships

Female Japanese macaques (Macaca fuscata) in the Arashiyama population near Kyoto, Japan, are unusual, in that they exhibit what many would consider to be male-typical sexual characteristics. Specifically, they mount other females within the context of temporary, but exclusive, sexual relationships (i.e., homosexual consortships) and they sometimes exhibit a preference for female sexual partners, even when given the choice of a sexually motivated male alternative. In this study, we examined whether female Japanese macaques also exhibited male-typical patterns of courtship behaviour during homosexual consortships. Data were collected on courtship behaviour during heterosexual and homosexual consortships in free-ranging Japanese macaques from the Arashiyama (Japan) population. We analyzed the occurrence of 12 different courtship behaviours during 3374 heterosexual inter-mount intervals and 1412 homosexual inter-mount intervals. Sex differences between heterosexually consorting males and females existed for only two of the 12 courtship behaviours we investigated: inclined-back presentations and sexual vocalizations. Dominant and subordinate homosexually consorting females were sex-typical in their expression of inclined-back presentations and sexual vocalizations. Consequently, facultative same-sex sexual partner preference, mounting and consortships do not co-occur with male-typical patterns of courtship behaviour in female Japanese macaques.     

Mating behavior ofXylotrechus pyrrhoderus Bates (Coleoptera: Cerambycidae) V. Female mounting behavior

Female mounting behavior was observed in the grape borerXylotrechus pyrrhoderus Bates (Coleoptera: Cerambycidae) under laboratory conditions. This behavior was directed towards either females (i.e., female homosexual mounting) or males. The female homosexual mounting was strongly elicited in females which had previously been allowed to fly. The frequency of this behavior was not related to the experience of mating. The investigation of the daily temporal pattern in female homosexual mounting activity indicated that the behavior most frequently occurred between 12∶00 and 16∶00, i.e., when females were searching for the pheromone-releasing males. Oviposition was also observed during this time interval. A detailed analysis revealed female mounting behaviors which are characteristic of male mounting behaviors, such as chasing conspecifics and synchronism of licking and abdominal bending. On the other hand, the frequency of male homosexual mounting was lower than that of females. Taken together with cases of other species previously reported, the expression of mounting behavior in females may be related to the evolution of the reproductive system, in which females approach their mates.     

Effects of ovariectomy and estradiol replacement therapy upon the sexual and aggressive behavior of the greater galago (Galago crassicaudatus crassicaudatus)

The effects of ovariectomy and estradiol treatment upon sexual and aggressive behavior were studied in a prosimian primate, the greater galago. Ovariectomized galagos were sexually unreceptive and frequently aggressive, but retained their sexual attractiveness to males. When females were treated with estradiol monobenzoate, however, their aggression and refusals of males' mounting attempts decreased markedly. Although males mounted these females, they usually failed to copulate, possibly because the females did not perform certain postural adjustments which assist males to intromit. Estradiol benzoate alone, even in large doses, does not fully restore patterns of mating behavior in ovariectomized female greater galagos. These observations on a prosimian primate are in striking contrast to the results of similar work on Old World monkeys and chimpanzees.     

Female American mink, Mustela vison, mate multiply in a free-choice environment

Like most solitary carnivores, the home ranges of male American mink overlap with those of several females, but each female typically shares its range with only a single male. Nevertheless, female mink produce multiply sired litters. Unusually among mammals, the physiological characteristics of female American mink make them highly suited to multiple paternity, suggesting that polyandry in the species may result from female behaviour rather than from coercion by males. In a free-choice experiment on captive-bred animals, all female mink mated multiply, with seven of the eight mating with all three available partners. Since females sometimes resisted copulations, multiple mating in this species may result from their behaviour rather than male coercion. Females visited males nonrandomly, but visiting patterns did not predict mating patterns, suggesting disparate social and mating preferences. There was no evidence for precopulatory female choice, since the allocation of copulations among available males did not deviate significantly from the expected distribution. Females mated multiply only around the predicted times of ovulation, contrary to the expectation that they should be more selective at times of peak fertility. Larger males copulated for longer in total: copulation duration is probably under male control, suggesting that larger males could influence mating duration, perhaps increasing their share of paternity by greater sperm transfer. Our observations suggest that in wild populations, mating patterns could be influenced by female preference for multiple partners.     

Mate choice by wild and mass‐reared females of the Mediterranean fruit fly

The sterile insect technique (SIT) is used to control Mediterranean fruit fly, Ceratitis capitata (Wiedemann), but its effectiveness is limited by low sexual competitiveness of mass‐reared males. This study investigated whether wild and mass‐reared [from a temperature sensitive lethal (tsl) genetic sexing strain] females display similar mate preferences and thus exert similar selective forces on the evolution of male courtship behaviour. Wild females preferred wild males over tsl males, whereas tsl females mated indiscriminately. The probability that mounting resulted in copulation was related to the duration of pre‐mount courtship for wild females, and wild males performed longer courtships than tsl males. Copulation occurred independently of courtship duration in tsl females. Counter to the aim of the SIT, female choice by tsl females appears to promote the evolution of male behaviour disfavoured by wild females.     

Intrasexual mounting in the beetle Diaprepes abbreviatus (L.).

The weevil Diaprepes abbreviatus shows three kinds of same-sex mountings: males mount other unpaired males, males mount males already engaged in copulation and females mount other females. Four hypotheses were evaluated in order to explain same-sex matings by males: (i) female mimicry by inferior males, (ii) dominance of larger males which affects the behaviour of small males, (iii) sperm transfer in which smaller males gain some reproductive success by 'hitchhiking' their sperm with the sperm of larger males, and (iv) poor sex recognition. Data from mate choice and sperm competition experiments rejected the female mimicry, dominance and sperm transfer hypotheses and supported the poor sex recognition hypothesis. We tested three hypotheses in order to explain female mounting behaviour: (i) females mimic male behaviour in order to reduce sexual harassment by males, (ii) females mount other females in order to appear larger and thereby attract more and larger males for mating, and (iii) female mimicry of males. The results of our mate choice experiments suggested that the female mimicry of males hypothesis best explains the observed female mounting behaviour. This result is also consistent with the poor sex recognition hypothesis which is the most likely explanation for male and female intrasexual mating behaviour in many insect species.     

Social relationships between adult females and the alpha male in wild tufted capuchin monkeys

Primates are notable for the widespread presence of long-term female-male associations which go beyond the mating context. However, little attention has been given to the factors that affect within-species variation in female-male relationships, especially among New World primates. Although detailed accounts of heterosexual relationships in Cebus species are scarce, a few studies have suggested the occurrence of strong associations between adult females and high-ranking males. This study explores affiliative relationships between females and the alpha male during the nonbreeding season in wild tufted capuchin monkeys (Cebus apella nigritus). Affiliative relationships were explored through female-male patterns of spatial proximity and grooming. By adopting a social network approach, we analyzed: (1) whether the alpha male is the preferred male partner for females and, (2) whether variation (if any) in female-alpha male affiliation can be explained through both female individual characteristics and social network metrics. Our results showed that alpha males were the favorite male partner for adult females in the proximity networks, but this did not hold true in the grooming networks. In addition, female-alpha male interaction patterns showed considerable variation, with only some females being strongly associated with the alpha male. Our results suggest that such a variation can be explained by female dominance rank, level of centrality (the quantity and intensity of spatial connection with other females) and prestige (the quantity of grooming received by other females) in female-female social networks. Taken together, these findings highlight two aspects of female-alpha male relationships in tufted capuchin monkeys: the alpha male represents the most socially integrated male in the group, and females with high dominance ranks and high centrality in both proximity and grooming networks show stronger relationships with the alpha male.     

Application of paternity discrimination by DNA polymorphism to the analysis of the social behavior of primates

Paternity in many primate species cannot be established reliably on behavioral grounds. For instance, Japanese macaques (Macaca fuscata) have a multi-male group structure and promiscuous mating patterns. On the other hand, accurate evaluation of male reproductive success is needed to analyze primate behavior. DNA finger-printing techniques were applied to 2 captive groups of Japanese macaques in the Primate Research Institute of Kyoto University for identification of paternity. Also, mating behavior of a captive group was observed in order to compare the reproductive success of each male with that expected on the basis of his observed mating activity. The number of offspring of full adult males was not related to their social rank although the number of copulations with ejaculation was highly correlated with their social rank. Monitoring of the female sexual cycles from the plasma profiles of gonadotropins and ovarian hormones suggested that males could not choose females at days of ovulation. The results of two-free-ranging wild troops, like those of the captive groups, indicated that high-ranking males could not monopolize the paternity of offspring. The results of paternity discrimination in Japanese macaques were compared with results from patas monkeys (Erythrocebus patas) in a discussion of social structure and male reproductive success. Some aspects of polymorphism detection techniques are also discussed.     

Nonconceptive Sexual Behavior in Bonobos and Capuchins

Sexual behavior by infecundable females, and by same-sex and adult-immature dyads, occurs in wild and captive bonobos (Pan paniscus). Proposed functions of these behaviors, in social primates generally, include practice, paternity confusion, exchange, and communication as well as appeasement. We used this framework to interpret and to compare observations of sexual behavior in a captive bonobo group and a wild white-faced capuchin (Cebus capucinus) group. In both species, (a) sexual behavior was no more frequent in cycling females than in pregnant or lactating females and (b) same-sex and adult-immature dyads engaged in as much mounting or genitogenital contact as adult heterosexual dyads did. The species differed in that (a) bonobos engaged in sexual behavior 65 times as frequently as capuchins, (b) only bonobos engaged in sexual contact other than ventrodorsal mounting during focal observation, and (c) bonobo sexual contact was concentrated most heavily in socially tense situations in adult female–female dyads, whereas capuchin sexual contact was concentrated most heavily in socially tense situations in adult male–male dyads. These data and published literature indicate that (a) practice sex occurs in both species, (b) paternity confusion may be a current function of C. capucinus nonconceptive sex, (c) exchange sex remains undemonstrated in capuchins, and (d) communication sex is more important to members of the transferring sex—female bonobos and male capuchins—than to members of the philopatric sex.     

Behaviorial effects of a synthetic mixture of aliphatic acids in rhesus monkeys (Macaca mulatta).

The stimulation of sexual behavior by a synthetic mixture of volatile aliphatic acids (acetic, propanoic, methylpropanoic, butanoic, methylbutanoic, methylpentanoic) was studied in male rhesus monkeys. Twelve intact adult males and 12 long-ovariectomized adult females were used in 24 paired combinations (541 tests each of 1 hr). A mixture of authentic acids similar to that found in the vaginal secretions of estrogenized females was applied to the sexual skin area of ovariectomized females immediately before tests with males. There was marked between-pair variability during the application of both control and test substances. However, using rigorous behavioral criteria, there was a well-marked stimulation either of male mounting attempts or of ejaculations in 12 of 24 pairs involving 9 of 12 males. Three males responded with both female partners, three responded with neither female partner, and six responded with one partner only. In the responding pairs, there were highly significant increases in mounting attempts and ejaculations, an effect that could be attributed only to treatment. We conclude, therefore, that these aliphatic acids (copulins), which act via olfactory pathways, have communicatory significance in rhesus monkeys.     

Female-male mounting in Japanese macaques: the proximate role of sexual reward

Female Japanese macaques (Macaca fuscata), in certain populations, routinely engage in female-male mounting. In this study, we documented patterns of vulvar, perineal and anal (VPA) stimulation during female-male mounting in Japanese macaques. During approximately 45% of the female-male mounts analyzed, two thirds of female mounters engaged in VPA stimulation. Given the VPA region's primary role in mediating sexual response, the results of this research provide direct evidence bearing on the sexual nature of a substantial proportion of female-male mounts in Japanese macaques.