Female-female mounting among goats stimulates sexual performance in males

The hypothesis that female-female mounting is proceptivity in goats, in that male goats are aroused by the visual cues of this mounting behavior, was tested. Once a week, male goats were randomly selected and placed in a test pen in which they were allowed to observe one of six selected social or sexual stimulus conditions. The stimulus conditions were one familiar male with two estrous females (MEE); three estrous females that displayed female-female mounting (E(m)); three estrous females that did not mount (E(nm)); three non-estrous females (N(E)); three familiar males (M); and no animals in the pen (Empty). After 10 min, the stimulus animals were removed, and an estrous female was placed in the test pen with the male for a 20-min sexual performance test. During sexual performance tests, the frequencies and latencies of all sexual behaviors were recorded. This procedure was repeated so all males (n = 6) were tested once each test day, and all the stimulus conditions were presented each test day. This was repeated weekly until all males had been exposed to each stimulus condition. Viewing mounting behavior, whether male-female or female-female, increased the total number of sexual behaviors displayed, increased ejaculation frequency, and decreased latency to first mount and ejaculation, post-ejaculatory interval, and the interval between ejaculations. We conclude that male goats are aroused by the visual cues of mounting behavior, and that female-female mounting is proceptivity in goats.     

Divergent roles for estrogens and androgens in the expression of female goat sexual behavior

We tested the hypothesis that the activation of the androgen receptor (AR) is required for full expression of female goat sexual behavior. Once a week for 6 weeks, ovariectomized (OVX) females were given priming doses of progesterone 72 and 48 h before behavioral observation. Estradiol (E(2); 100 microg), testosterone (T; 100 mg), or sesame oil was supplied 14 h before behavioral testing. Six goats received the AR antagonist flutamide (9 mg/kg sc) 8 h before and 4 h after steroid injection. Six goats received the carrier only. After 3 weeks, flutamide and carrier treatments were switched so that all females received all treatments. Treatments with E(2) and T were equally effective in eliciting estrus-typical behaviors (sniffing, courting, leg kicks, mount attempts by males, bouts of thrusting by males, ejaculations, and flehman responses) compared to treatment with oil. Flutamide treatment enhanced proceptive behaviors in E(2)-treated females compared to other treatment groups; this was most likely via enhanced tail wagging. Moreover, compared to goats given T + carrier, T + flutamide significantly reduced receptivity in females. The results of this experiment implicate the AR as an important facilitator of some aspects of the female goat sexual behavior. However, the results of this experiment do not show whether androgens influence estrous behaviors alone or in some combination with estrogen.     

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.     

Food Deprivation and Restriction During late Gestation Affect the Sexual Behavior of Postpartum Female Meadow Voles,Microtus pennsylvanicus

In many species of small mammals, including meadow voles, Microtus pennsylvanicus, females come into postpartum estrus (PPE) within 12–24 h of giving birth, allowing them to mate and become pregnant while raising the current litter. PPE females show increases in attractivity, proceptivity, and receptivity, the three components of sexual behavior, relative to females not in PPE. Several studies have shown that food deprivation and restriction reduce attractivity, proceptivity, and receptivity of females not in PPE. We tested the hypothesis that food deprivation and restriction during late gestation cause deficits and decrease the attractivity, proceptivity, and receptivity of females when they enter PPE. Our data support the hypothesis. On day 1 of lactation, females that were food deprived and food restricted produced scent marks that were significantly less attractive as those produced by control PPE females. Food deprivation but not food restriction caused females to no longer display significant preferences for the scent marks of males over those of females (proceptivity). Food deprivation and food restriction were sufficient to induce females to become significantly less sexually receptive than control females. Eleven of 12 control PPE females mated, 4 of 12 food‐restricted females mated, and 3 of 12 food‐deprived females mated. Dams facing food deprivation or restriction during late gestation may have to balance the benefits of mating during PPE with the increased costs associated with getting pregnant while they are lactating.     

Sex differences in DHEA and estradiol during development in a wild songbird: Jugular versus brachial plasma

Sexual differentiation of the brain has traditionally been thought to be driven by gonadal hormones, particularly testosterone (T). Recent studies in songbirds and other species have indicated that non-gonadal sex steroids may also be important. For example, dehydroepiandrosterone (DHEA) - a sex steroid precursor that can be synthesized in the adrenal glands and/or brain - can be converted into active sex steroids, such as 17β-estradiol (E₂), within the brain. Here, we examine plasma DHEA and E₂ levels in wild developing European starlings (Sturnus vulgaris), from hatch (P0) to fledging (P20). Blood samples were collected from either the brachial vein (n = 143) or the jugular vein (n = 129). In songbirds, jugular plasma is enriched with neurally-synthesized steroids and, therefore, jugular plasma is an indirect measure of the neural steroidal milieu. Interestingly, brachial DHEA levels were higher in males than females at P4. In contrast, jugular DHEA levels were higher in females than males at P0 and P10. Brachial E₂ levels were higher in males than females at P6. Surprisingly, jugular E₂ levels were not high and showed no sex differences. Also, we calculated the difference between brachial and jugular steroid levels. At several ages, jugular steroid levels were lower than brachial levels, particularly in males, suggesting greater neural metabolism of circulating DHEA and E₂ in males than females. At a few ages, jugular steroid levels were higher than brachial levels, suggesting neural secretion of DHEA or E₂ into the general circulation. Taken together, these data suggest that DHEA may play a role in brain sexual differentiation in songbirds.     

Dexamethasone suppression of sexual behavior in the ovariectomized mare

The influence of steroids of adrenal cortical origin on estrous behavior in the ovariectomized mare was evaluated by adrenal suppression via dexamethasone (DEX) administration in two experiments. In Experiment I, 12 mares (six DEX, six control) were tested for sexual behavior in harem groups (two DEX and two control mares plus one stallion per group) for 9 consecutive days. In Experiment II, estradiol (E₂) was given to a group of DEX-treated mares as an additional control. Twelve mares (four DEX, four DEX + E₂, and four control) were tested in harem groups (one DEX, one DEX + E₂, and one control mare plus one stallion per group) for 10 days. All DEX mares showed a clear suppression of sexual response compared to control or DEX + E₂ mares, indicating that the estrous behavior seen in ovariectomized mares may be due to steroids from the adrenal cortex. The control and DEX + E₂ mares were similar in all measures of proceptivity. Despite being more receptive, as indicated by fewer negative responses, the DEX + E₂ mares received fewer intromissions and ejaculations than did the control animals. The ability of estradiol to induce estrous behavior in the dexamethasone-suppressed mare notwithstanding, other adrenal steroids, e.g., androgens, may be involved in estrous behavior in the untreated, ovariectomized mare.     

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.     

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.     

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.     

Relationships between rapid changes in local aromatase activity and estradiol concentrations in male and female quail brain

Estradiol-17β (E₂) synthesized in the brain plays a critical role in the activation of sexual behavior in many vertebrate species. Because E₂ concentrations depend on aromatization of testosterone, changes in aromatase enzymatic activity (AA) are often utilized as a proxy to describe E₂ concentrations. Utilizing two types of stimuli (sexual interactions and acute restraint stress) that have been demonstrated to reliably alter AA within minutes in opposite directions (sexual interactions = decrease, stress = increase), we tested in Japanese quail whether rapid changes in AA are paralleled by changes in E₂ concentrations in discrete brain areas. In males, E₂ in the pooled medial preoptic nucleus/medial portion of the bed nucleus of the stria terminalis (POM/BST) positively correlated with AA following sexual interactions. However, following acute stress, E₂ decreased significantly (approximately 2-fold) in the male POM/BST despite a significant increase in AA. In females, AA positively correlated with E₂ in both the POM/BST and mediobasal hypothalamus supporting a role for local, as opposed to ovarian, production regulating brain E₂ concentrations. In addition, correlations of individual E₂ in POM/BST and measurements of female sexual behavior suggested a role for local E₂ synthesis in female receptivity. These data demonstrate that local E₂ in the male brain changes in response to stimuli on a time course suggestive of potential non-genomic effects on brain and behavior. Overall, this study highlights the complex mechanisms regulating local E₂ concentrations including rapid stimulus-driven changes in production and stress-induced changes in catabolism.     

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.     

Expression of adult female patterns of sexual behavior by male, female, and pseudohermaphroditic female rhesus monkeys

Gonadally intact pseudohermaphroditic female and normal female and neonatally castrated male rhesus monkeys were given estrogen treatment as adults and evaluated for attractivity, proceptivity, and receptivity during tests with a tethered stud male. Pseudohermaphrodites were produced by injecting their mothers during pregnancy with either testosterone propionate (TP) or dihydrotestosterone propionate (DHTP). Castrated males had reliably lower attractivity than normal females on all indicator responses shown by the tethered males. Additionally, castrated males showed reliably fewer proceptive responses on 4 of 5 measures than normal females. Receptivity could not be assessed in this situation for castrated males, because tethered males never contacted them unless the castrated males were displaying presentation. No reliable differences were observed between pseudohermaphrodites produced by prenatal treatments with TP or DHTP. Pseudohermaphrodites generally showed reliably less attractivity and proceptivity than normal females and reliably more of these traits than castrated males. Attractivity scores for pseudohermaphrodites were not different from those for normal females until proximity to the tethered male was established. Receptivity was not different in pseudohermaphrodites compared with normal females. Results indicate prenatal androgenization and its developmental sequelae lead to a defeminization in adulthood which, in this testing situation, was principally manifested by a deficiency in the performance of proceptive behaviors. Additionally, defeminization in rhesus monkeys, unlike that demonstrated in rodents, does not depend upon actions of an aromatizable androgen.     

Strong sexual selection in males against a mutation load that reduces offspring production in seed beetles

Theory predicts that sexual reproduction can increase population viability relative to asexual reproduction by allowing sexual selection in males to remove deleterious mutations from the population without large demographic costs. This requires that selection acts more strongly in males than females and that mutations affecting male reproductive success have pleiotropic effects on population productivity, but empirical support for these assumptions is mixed. We used the seed beetle Callosobruchus maculatus to implement a three‐generation breeding design where we induced mutations via ionizing radiation (IR) in the F₀ generation and measured mutational effects (relative to nonirradiated controls) on an estimate of population productivity in the F₁ and effects on sex‐specific competitive lifetime reproductive success (LRS) in the F₂. Regardless of whether mutations were induced via F₀ males or females, they had strong negative effects on male LRS, but a nonsignificant influence on female LRS, suggesting that selection is more efficient in removing deleterious alleles in males. Moreover, mutations had seemingly shared effects on population productivity and competitive LRS in both sexes. Thus, our results lend support to the hypothesis that strong sexual selection on males can act to remove the mutation load on population viability, thereby offering a benefit to sexual reproduction.     

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.     

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.     

Functional significance of behavioral,morphological, and endocrine correlates across the ovarian cycle in semifree ranging female Tonkean macaques

The role of sexual displays in mating strategies and their reliability in indicating the time of ovulation has given rise to multiple explanations in nonhuman primates. In order to discriminate among hypotheses, socio-sexual behaviors were recorded in a semifree ranging group of Tonkean macaques (Macaca tonkeana), together with sexual skin swelling volumes and measurements of urinary concentrations of estrone conjugates and pregnanediol glucuronide. A clear preovulatory peak of urinary estrogen levels occurred 2 days before a defined rise in pregnanediol glucuronide concentrations, indicating that both hormones pointed out the female's ovulatory period. The concept of estrus rightly could be applied to female Tonkean macaques since fluctuations in estrogen levels correlated with cyclic changes in genital swelling sizes and rates of female behavioral attractivity and proceptivity. Males proved to be capable of recognizing the optimal conception period as judged from the occurrence of maximal rates of following behavior, serial matings, and ejaculations during the peri-ovulatory phase. During this time, males succeeded in maintaining exclusive and enduring associations with females. However, consortships occurred precociously, with males starting to affiliate with females, follow, and mount them 1 week before the presumed time of ovulation. These long-lasting consortships appear to be a consequence of the female extended follicular phase. This presumably sexually selected character allowed females to extend conspicuous sexual displays: genital swelling and utterance of an estrous call, which might attract males' attention and arouse them. With regard to female mating tactics, the combination of reliably indicating the time of ovulation to the male and durable periods of competitor exclusion led to reject explanations assuming manipulation about paternity or long-lasting intermale competition incitement in Tonkean macaques. Competition for mates between females also turned out to be an irrelevant factor as it was very low in the species. We conclude that the main function of sexual displays is to herald the approach of ovulation toward available mates. Am. J. Primatol. 46:285–309, 1998. © 1998 Wiley-Liss, Inc.     

Male aggression and the sexual behavior of Japanese monkeys

The factors that regulate male aggressive behavior towards females during the mating season and their relationship to the sexual behavior of Japanese monkeys were investigated. Observations were made on the Shiga A troop in Nagano Prefecture for 132 days during both the non-mating and mating seasons in three successive years. As a result, chasing towards females was observed in males older than 4 years. The frequencies of this type of behavior increased in the mating season. The chasing rank of the individual males did not correlate with the male dominance rank but with the tree-shaking rank. Chasing was directed not only at estrous females but also at non-estrous females. Pairs of both sexes in which chasing was observed, tended to have sexual interactions. Both hormonal and social factors should be considered in the regulation of chasing. The role of aggression in the formation of new male-female bonds in discussed.     

Attractivity of male and female rats which are hormonally manipulated during early development and in adulthood

Attractivity is one aspect of female sexuality relevant for the understanding of male-female sexual interactions. In a previous study, it was shown that intact males were equally attracted to early androgenized, gonadally intact females as to normally developed, estrous females. The present study was designed to investigate in what way hormones given in adulthood might influence attractivity of early androgenized females in adulthood. Specifically, we compared the attractivity of neonatally androgenized females (NeoTP) to the attractivity of normally developed females (NeoOIL), neonatally castrated males (NeoCASTR), and neonatally sham-castrated males (NeoSHAM) when different groups received either OIL, estradiol benzoate (EB) or testosterone propionate (TP) in adulthood. The male's preference to stay in the vicinity of one incentive in favor of the other was taken as an index of attractivity. The results show that, under the present hormonal conditions, NeoTP-females are generally less attractive than NeoOIL-females, more attractive than NeoSHAM-males, and equally attractive as NeoCASTR-males. TP-treated androgenized females were found to be equally attractive as TP-treated NeoSHAM-males. It is concluded that, relative to normally developed females, androgenized females become less attractive when the endogenous secretion of sex steroids is artifically controlled by gonadectomy and/or by administration of fixed amounts of sex steroids.     

Dynamics of social and energetic stress in wild female chimpanzees

Stress hormone measurements can reinforce and refine hypotheses about the costs of particular contexts or behaviors in wild animals. For social species, this is complicated because potential stressors may come from the physical environment, social environment, or some combination of both, while the stress response itself is generalized. Here, we present a multivariate examination of urinary cortisol dynamics over 6 years in the lives of wild female chimpanzees in the Kanyawara community of Kibale National Park, Uganda. We hypothesized that chimpanzee socioecology provides strong indications of both energetic and social stress to females, but that the salience of these stressors might vary over a female's life history in accordance with their changing reproductive costs and social interactions. Using linear mixed models, we found that urinary cortisol levels increased significantly with age but were also elevated in young immigrants to the community. Across reproductive states, cycling, non-estrous females had relatively low cortisol compared to lactating, estrous, or pregnant females. Aggression from males led to higher cortisol levels among estrous females, frequent targets of aggressive sexual coercion. In contrast, energetic stress was most salient to lactating females, who experienced higher cortisol during months of low fruit consumption. Low dominance rank was associated with increased cortisol, particularly during the energetically demanding period of lactation. The effects of female conflict were felt widely, even among those who were the primary aggressors, providing further evidence that long-term resource competition, while apparently muted, exerts a far-reaching impact on the lives of chimpanzee females.     

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.