Influence of male endocrine condition on the choice of social contact in pair-living Mongolian gerbils (Meriones unguiculatus)

In two previous studies influence of gonadal hormones on male—female interaction has been investigated. Interaction between pair-living cagemates and non-cagemates was compared. In the first study the animals met in direct paired encounters, and in the second study the tested animals had the choice of wiremesh-restricted contact with the cagemate or a non-cagemate. The present choice test involves direct contact with the animal chosen in a series of consecutive trials. The two incentive animals were again non-cagemate and the cagemate of the experimental animal. The males tested were intact or castrated. The females were tested in induced oestrus. The females tested with intact males showes a higher choice of the cagemate. This was also the case in castrated males. Intact males and females tested with castrated males showed no significant difference in their choice. Differences in the results and testing conditions of the present and the previous choice test allowing restricted contact are discussed.     

Influence of gonadal hormones and social housing conditions on agonistic,copulatory, and related socio-sexual behaviour in the Mongolian gerbil (Meriones unguiculatus)

The Mongolian gerbil is colonial, but its social organization within its colonies is not known. Pair formation has been suggested by breeders and has been observed under semi-natural conditions. Family units remain stable under such conditions. In the solitary-living hamster (Cricetus auratus), agonistic behaviour decreases in male-female encounters during female heat. This report concerns the influence of female and male gonadal hormones on agonistic behaviour in gerbils studied in cagemate and non-cagemate encounters. Further, the occurrence of mating, marking, and drumming was studied in the same situations. No effects were found of either the female or the male endocrine condition on the low amount of agonistic behaviour between cagemates. A reduction in the high amount of agonistic responses in encounters between non-cagemates was indicated. It was related to the castrated-male and induced-oestrous female conditions. Copulatory behaviour was not influenced by social maintenance conditions. Marking and drumming were most frequent in females in oestrous and intact males. The occurences in the females were influenced by the individual's endocrine condition, but not by the opponent's. Intact males in encounters with non-cagemates marked more frequently when the females were in anoestrous. Drumming in the intact males increased in tests with oestrous versus anoestrous females but occurred more frequently in encounters with non-cagemates than in those with cagemates. The behaviour of castrated males was not influenced by the endocrine or by social housing conditions. The limited variation in agonistic behaviour relative to the endocrine conditions may reflect a non-solitary social organization.     

The hormonal response of female European Stonechats to a territorial intrusion: the role of the male partner

In many bird species, the female participates in defending a pair's breeding territory, however, the endocrine control mechanism of female aggressive behavior is largely unknown. The general statement that androgens are involved in the regulation of aggressive behavior is based on studies conducted only in males. Here, we tested whether paired female stonechats show a hormonal response to a simulated male territorial intruder. Since in males of territorial bird species androgen levels usually increase following a male-male encounter, we measured androgen-levels before and after a simulated male intrusion. In addition, we measured estradiol, the main gonadal hormone in females, and corticosterone, a stress hormone. The results show that a male intruder does not affect any of the measured hormones in females. In a second experiment, we also tested whether the endocrine state of the male partner affects the hormonal response of females to a male intruder by comparing the hormonal response of females paired with pharmacologically castrated males and females paired with control males. Females paired with pharmacologically castrated males had lower corticosterone levels both before and after the intrusion than females paired with control males. Additionally, in both groups, female corticosterone levels were increased following a male intrusion. We suggest that the differences found between females paired with pharmacologically castrated males and females paired with control males are due to differences in intra-pair interactions.     

Attractivity of male and female rats after early endocrine manipulation

Approach behavior toward males and approach behavior toward females were studied in the male rat. Adult male rats approached neonatally castrated male rats more than they approached intact males. Neonatally androgenized female rats and estrous female rats were equally approached. A preferential approach toward the androgenized female was evident when the alternative incentive was a proestrous female. These adult patterns of approach behavior appeared on Day 50 after birth. It is concluded that early endocrine manipulation influences the attractivity of the incentives.     

Partner preference of intact and ovariectomized female gray short-tailed opossums (Monodelphis domestica)

Intact, ovariectomized and ovariectomized estradiol (E)-treated female gray short-tailed opossums were placed in a test situation in which they could choose between an intact and a castrated male. Intact females chose to visit intact males first and visited them more frequently and spent more time with intact than with castrated males. Ovariectomized (OVX) females did not show this preference for visiting intact males over castrates. When compared to OVX females with blank implants, OVX females with E implants spent less time with castrated males. Like intact females, OVX and OVX-E-treated females preferred to stay in close proximity to but not actually in the cage of intact rather than castrated males. To our knowledge, this is the first experimental study of partner preference and its relationship to hormonal condition in a female marsupial.     

Why Do African Elephants (Loxodonta africana) Simulate Oestrus? An Analysis of Longitudinal Data

Female African elephants signal oestrus via chemicals in their urine, but they also exhibit characteristic changes to their posture, gait and behaviour when sexually receptive. Free-ranging females visually signal receptivity by holding their heads and tails high, walking with an exaggerated gait, and displaying increased tactile behaviour towards males. Parous females occasionally exhibit these visual signals at times when they are thought not to be cycling and without attracting interest from musth males. Using demographic and behavioural records spanning a continuous 28-year period, we investigated the occurrence of this “simulated” oestrus behaviour. We show that parous females in the Amboseli elephant population do simulate receptive oestrus behaviours, and this false oestrus occurs disproportionately in the presence of naïve female kin who are observed coming into oestrus for the first time. We compare several alternative hypotheses for the occurrence of this simulation: 1) false oestrus has no functional purpose (e.g., it merely results from abnormal hormonal changes); 2) false oestrus increases the reproductive success of the simulating female, by inducing sexual receptivity; and 3) false oestrus increases the inclusive fitness of the simulating female, either by increasing the access of related females to suitable males, or by encouraging appropriate oestrus behaviours from female relatives who are not responding correctly to males. Although the observed data do not fully conform to the predictions of any of these hypotheses, we rule out the first two, and tentatively suggest that parous females most likely exhibit false oestrus behaviours in order to demonstrate to naïve relatives at whom to direct their behaviour.     

Sexual motivation suppresses paternal behaviour of male gerbils during their mates' postpartum oestrus

Adult male Mongolian gerbils, Meriones unguiculatus, avoid contact with their young on the day that the young are born. However, on succeeding days, fathers spend nearly as much time in contact with their offspring as do mothers. We undertook a series of studies to investigate the causes of the day-to-day change in male parental behaviour. In experiment 1, we tested males' response to pups before, after and on the day of their mates' parturition, and found that males were more parental both before and after the day of birth of their young than on that day. In experiment 2, we compared the parental behaviour of males paired either with intact or with ovariectomized dams (which do not come into postpartum oestrus) and found that males that were mated to intact females were less attentive to pups on the day of their birth than were the males mated to ovariectomized females. In experiment 3, we compared the parental responses of castrated and intact males to newborn pups and found that castrated males were more parental than intact males. In experiment 4, we compared the parental responses of males that were exposed to postpartum oestrous females but prevented from mating for 24 h. Extending the period of male sexual arousal to 24 h inhibited paternal responsiveness to neonates for 24 h. We interpret these results as being consistent with the hypothesis that on the day of birth of a litter, a male's parental behaviour is inhibited by the motivation to mate during his partner's postpartum oestrus.     

Sexual and social behavior of adult saddle-back tamarins (Saguinus fuscicollis), castrated as neonates

The behavior of four adult males castrated as neonates was compared with that of three neonatally sham-castrated males and two untreated males serving as controls. Two males castrated as sexually experienced adults served as additional controls. All males lived in permanent male-female pairs. The interactions of neonatally castrated, sham-castrated, and intact control males with strange conspecifics were tested in a situation analogous to territorial defense. In addition, the interactions of all males with their female pair mates were tested. Neonatally castrated males showed less injurious aggression against strangers and interacted with them less frequently than control males. There were significant differences between both groups of males in some of the agonistic behaviors recorded, but not in all. In pair tests, neonatally castrated males failed to show any coordinated male copulatory behavior but control males did. Males castrated as adults showed rates of mounting and thrusting similar to those of intact controls. Neonatally castrated males tended to interact with their females on a lower level than neonatally sham-castrated and untreated control males. How ever, most differences between both groups in the levels of pair mate interactions were not statistically significant. It is concluded that castration during neonatal life prevents the display of male copulatory behavior in adulthood and selectively affects the display of a number of other behaviors in adulthood. It is not possible at this point to decide whether these results are due to hormonal deficiencies on an organizational or activational level or both. However, data from studies currently in progress suggest that organizational processes might be responsible for part of the effect.     

Sexual hormones: effects on cardiac and mitochondrial activity after ischemia-reperfusion in adult rats. Gender difference

In this work we studied the influence of sex hormones on heart and mitochondrial functions, from adult castrated female and male, and intact rats. Castration was performed at their third week of life and on the fourth month animals were subjected to heart ischemia and reperfusion. Electrocardiogram and blood pressure recordings were made, cytokines levels were measured, histopathological studies were performed and thiobarbituric acid reactive species were determined. At the mitochondrial level respiratory control, transmembranal potential and calcium management were determined; Western blot of some mitochondrial components was also performed. Alterations in cardiac function were worst in intact males and castrated females as compared with those found in intact females and castrated males, cytokine levels were modulated also by hormonal status. Regarding mitochondria, in those obtained from hearts from castrated females without ischemia-reperfusion, all evaluated parameters were similar to those observed in mitochondria after ischemia-reperfusion. The results show hormonal influences on the heart at functional and mitochondrial levels.     

Effect of female sexual displays on the endocrine physiology and behaviour of male white-crowned sparrows, Zonotrichia leucophrys

In photoperiodic birds, endocrine responses to behavioural interactions between males and females may be involved in temporally "fine-tuning" the onset of reproduction to yearly variations in the environment. This study examined the endocrine and behavioural responses of male White-crowned sparrows ( Zonotrichia leucophrys ) to changes in the endocrine state of the female, as signalled by changes in her behaviour. Males on different photoperiodic regimes were paired with oestrogen-treated, sexually receptive females. Males exposed to gonadostimulatory long days mounted and copulated with oestrogen-treated females even before gonadal development was complete. These males had higher plasma levels of testosterone and luteinizing hormone and maintained enlarged testes longer than control males paired with untreated, nonreceptive females. Males maintained on nonstimulatory short days also mounted oestrogen-treated females; however, testes of these males remained nonfunctional and their plasma levels of testosterone and luteinizing hormone were basal. Thus, reproductive function of photostimulated males is profoundly affected by changes in the endocrine state and behaviour of the female. However, male sexual behaviours are expressed in response to visual and auditory stimuli from the female regardless of male hormonal condition or photoperiodic treatment.     

Dynamics of male residence and female oestrus during a breeding season of blue monkeys in the Kakamega Forest, Kenya

Blue monkeys (Cercopithecus mitis stuhlmanni) are seasonal breeders with a dynamic mating system in which typical one‐male social units are regularly disrupted and replaced by multi‐male ones. The number of males in the group is correlated with the number of oestrous females. We used observations of male presence and female oestrus on individual days during a 6‐month period to assess whether the presence of multiple males in a group stimulates female oestrus or whether oestrous females attract multiple males to the group. We confirmed prior observations with our finding that the number of males in a group was significantly correlated (r_s = 0.435, P < 0.0001) with the number of oestrous females across 126 observation days. A transition matrix did not show an obvious relationship between day‐to‐day changes in the numbers of oestrous females and males. However, cross‐correlation analysis provided stronger support for the idea that the number of oestrous females attracts males to a group than for the idea that the influx of strange males stimulates oestrus in female blue monkeys. Autocorrelation analysis showed that while female oestrus appeared to show a high degree of synchrony, as expected in a seasonal breeder, there was no evidence that the number of males accompanying a group of females influenced the likelihood of other males joining or leaving the group. Overall, our results confirm that female oestrous behaviour stimulates changes in male residence patterns. However, other observations suggest that changes in male residence may also stimulate female oestrus in some circumstances.     

Postisolation aggression and olfactory cues.

Aggressive maternal behavior was tested by placing intruders in the home cages of 210 individually housed lactating mice 3-8 days postpartum. The intruders were of 6 types: castrated males and females, intact males and females, mothers of 3-8 day old pups, and mothers of older pups. All fights were initiated by the resident lactating female. There were significantly fewer attacks on castrated males than on intact males (p less than .001), and fewer attacks on castrated females than on intact males and females (p less than .001, p less than .01, respectively). There were no attacks on mothers of 3-8 day old pups. Attacks on older mothers were significantly more frequent than on younger mothers (p less than .01). The results indicate that maternal aggression in albino mice is strongly influenced by the hormonal and reproductive state of the intruder.     

Olfactory preferences, scent marking, and "proceptivity" in female hamsters

Two appetitive sexual behaviors of female hamsters, attraction to odors of males and vaginal scent marking, were investigated and the correlations of each to reproductive states were determined. Both estrous and diestrous females were more attracted to the odors of intact males than to the odors of females or castrated males; this differential attraction was more pronounced for estrous females. Lactating females, but not pregnant females, were also preferentially attracted to odors of intact males. Thus, odors alone provided sufficient information to allow sexual discrimination by females, and estrous, diestrous, and lactating females were preferentially attracted to odors of intact males whereas pregnant females were not. Reproductive condition had no influence on responses to odors of castrated males or diestrous females. Vaginal scent marking was strongly influenced by reproductive state: Marking frequency was low in estrous, pregnant, and early lactation females, but high in late lactation and diestrous females, suggesting a function of advertisement of approaching receptivity. Thus sexual preferences based on olfactory cues and vaginal scent-marking frequency differed in their relationships to reproductive conditions. It is suggested that in other species such relationships between appetitive sexual behaviors and reproductive condition will depend on the species social organization, ecological niche, and taxonomic group, and consequently that the concept of preceptive behaviors should be broadened to include all appetitive sexual behaviors of females, not just those that are correlated with the hormonal state characterizing estrus.     

Activation of estrus by pheromones in a marsupial: stimulus control and endocrine factors

Little is known about the effects of social factors or about the hormonal control of social factors on reproduction in marsupials. In previous studies on the gray opossum, a small, Brazilian didelphid, the presence of males was found to activate reproduction in females. In this study, the effects of pheromones produced by intact males, castrated males, and intact females on estrus in female gray opossums was examined. It was found that females housed alone rarely showed spontaneous periods of estrus, that exposure to pheromonal cues provided by intact males reliably induced estrus in 75-100% of the experimental females, and that pheromonal cues provided by castrated males or intact, stimulus females induced estrus in 25% of the experimental females. These findings are discussed with respect to the ecology, social behavior, and reproductive characteristics of the didelphid marsupials.     

Sexual behaviour and its olfactory control in the desert woodrat (Neotoma lepida lepida)

In studies 1 and 2 nine pairs of desert woodrats (Neotoma lepida lepida) were observed for copulatory behaviour when the female was in a state of naturally occurring oestrus (study 1) and following ovariectomy and oestrogen and progesterone replacement (study 2). Males and females respond in a similar way under conditions of natural and hormone-induced oestrus. Males show a consummatory pattern involving multiple mounts and ejaculations, with ejaculations occurring after single intromissions. Females show the lordosis reflex accompanied by hop-and-dart and ear-wiggling responses. In addition, both sexes show appetitive precopulatory behaviours; the male emits an audible rasping vocalization as he trails and mounts the female, following a period of intense sniffing of the female's anogenital region. The female also frequently approaches and sniffs the male. In study 3, the role of female odours in the sexual behaviour of the male was examined in eight of the nine pairs used in studies 1 and 2. This was done by applying to the anogenital region of ovariectomized females a combination of urine and vaginal secretions taken from familiar and unfamiliar, and oestrogen-primed or non-oestrogen-primed females. The results show that odours from oestrogen-primed females are not sufficient to elicit male sexual behaviour, if the female is not sexually active. In study 4 the eight males were tested for their preferences for urine and vaginal secretion odours taken from females in different reproductive states and applied to cotton swabs. These males spent more time sniffing unfamiliar oestrous odours than unfamiliar non-oestrous odours and more time sniffing oestrous odours from a familiar female over those taken from an unfamiliar female.In study 5, 12 sexually active males were tested with oestrogen-primed females before and after either olfactory bulb removal or sham-surgery. Bulbectomized animals ceased copulating with females although females showed precopulatory approaches.Taken together, these studies suggest that normal sexual behaviour in the male woodrat requires that the female both possess the attractive odours (of oestrus) and that she engage in appetitive precopulatory behaviour.     

Social influences on plasma testosterone levels in male talapoin monkeys

Three social groups of laboratory-housed talapoin monkeys (Miopithecus talapoin) consisting of four adult males and four or five adult females, were observed over a 4-year period. All females were ovariectomized and given estradiol implants at intervals to render them sexually attractive; except for two castrated males with testosterone implants, all males were intact. Sexual and aggressive interactions were recorded, and testosterone levels were measured in plasma taken from males twice weekly. In each group males formed a linear dominance order, defined in terms of the direction of aggression between animals. The hormonal responses of intact males were monitored with respect to the presence of attractive females, access to these females, and transfer from the social group to isolation. In all groups the behavioral and endocrine responses of males to these treatments were rank related. In some instances, rising in rank was associated with elevated testosterone and falling in rank with decreased testosterone; these hormonal changes were associated with changes in sexual and aggressive interactions. The effects of sexual and aggressive behavior on plasma testosterone titers are discussed.     

Individual differences in aggressiveness of female hamsters: response to intact and castrated males and to females

The aggressive behaviour of female hamsters was studied while they were housed in large enclosures with males and in brief tests with males or females. Some females are not aggressive with any male, whereas others are very aggressive toward all males in both testing conditions. Females that are not aggressive toward intact males may be very aggressive toward castrated males or females. When the animals are housed together for long periods of time, males dominate only if they are much heavier. Male dominance takes a relatively long time to establish and often there is an equivocal period characterized by reversals of dominance. Female dominance is rapidly established. Unless the male is much heavier, the female determines the presence or absence of agonistic behaviour.     

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.     

Hormonal and experiential control of female-male mounting in the female rat

Mounting behavior in the female rat has been studied extensively in same-sex interactions, but not in the heterosexual dyad. The present study examined the display of female mounting of castrated noncopulating male rats (FMM). Ovariectomized (OVX) sexually naive female rats (N = 80) were given either estrogen (E) + progesterone (P), E + oil (O), P + O, or O + O treatment for five tests with castrated male rats. FMMs were observed in both the E + P and E + O females. The influence of the ovarian cycle on FMM was also investigated. Vaginal smears from sexually naive females (N = 16) were taken daily for 12 days immediately after testing with castrated males. FMM frequency was greatest during Proestrus. Finally, OVX females (N = 30) treated with E + P were given either 0, 1, 10 multi-ejaculatory heterosexual experiences with intact, sexually experienced males, prior to tests with intact, copulating or castrated, noncopulating males for five tests. Sexually naive females displayed a greater number of mounts relative to the sexually experienced females when tested with castrated, noncopulating males. In contrast, very few FMMs were observed in females of any group tested with intact, copulating males. These data suggest that FMM occurs naturally in rats as a "super-solicitational" behavior that is modified by hormone treatment and prior heterosexual experience.     

Expression of male-typical behavior in adult female pseudohermaphroditic rhesus: Comparisons with normal males and neonatally gonadectomized males and females

Two types of pseudohermaphroditic female rhesus produced by exposure to either testosterone propionate (TP) or dihydrotestosterone propionate (DHTP) prior to birth were ovariectomized postpuberally and evaluated for the display of male-typical sexual behavior in response to exogenous TP in adulthood (2 mg/kg/day for 12 weeks). Their performance in standardized tests with estrogenized female partners was compared to that of neonatally gonadectomized males and females identically tested and treated with exogenous TP as adults. In addition intact adult males not given exogenous TP were tested with the same estrogenized female partners. There were no reliable differences between the two types of pseudohermaphrodites on any measure of behavior shown during the tests. Accordingly results were combined. Reliable behavioral changes induced by the TP given in adulthood were limited to increases in purse-lip responses, the induced increases were similar in pseudohermaphrodites and castrated males, and increases were reliably greater in these two groups of subjects than in females. Pseudohermaphrodites and castrated males did not differ reliably from intact males in performance of purse-lip gestures during TP treatment. In the performance of mounting, however, pseudohermaphrodites and castrated males remained consistently below the standard of the intact males. The estrogenized female partners displayed proceptive responses most frequently to the intact males and least frequently to the females. Their proceptive responses with castrated males resembled their performance with intact males, but with pseudohermaphrodites their proceptive responses more closely resembled their performance with females. Receptive behavior of the female partners was displayed most frequently to intact males, at intermediate levels to castrated males, and least often to pseudohermaphrodites. Results are completely consistent with the notion that androgens in high concentrations before birth alter mechanisms related to the later display of masculine behavior. These alterations in behavioral mechanisms are of such a nature that the display of male-typical behavior induced by androgens in adulthood is more pronounced and more frequent than it would have been otherwise. The alterations in masculine behavior observed in pseudohermaphroditic rhesus are not different in kind or scope than those reported extensively for lower mammals.(ABSTRACT TRUNCATED AT 400 WORDS)