Group mating among Norway rats I. Sex differences in the pattern and neuroendocrine consequences of copulation

The copulatory pattern of groups of rats (Rattus norvegicus) was studied in the laboratory in a seminatural environment. In a given mating session, every oestrous female copulated with each male; likewise, every male copulated with each oestrous female. While individual males and females experienced similar amounts of copulation, there were dramatic sex differences in sequence and temporal pattern. Males mated in a multiple intromission pattern and had more ejaculatory series when several females were in oestrus. In contrast, females received intromissions and ejaculations in a random order, not in the sequence of a male ejaculatory series. Males copulated at shorter intervals than females did, a temporal sex difference that was determined by the pattern of female solicitations and male approaches. These sex differences are used to discuss the different units of analysis that are appropriate for male and female sexual behaviour in this species. Furthermore, the sex differences in the temporal pattern of copulation which emerged during group mating parallel the known sex differences in the temporal parameters of the neuroendocrine reflexes which mediate successful reproduction in the domestic strain.     

Copulatory behaviour of the bank voleMyodes glareolus: matings with one and two males do not make a difference

In promiscuous species in which females mate with more than one male during oestrus, males may increase their sperm expenditure or change their copulatory behaviour in response to the risk of sperm competition. I used an experimental approach to investigate the pattern of copulatory behaviour of the bank voleMyodes glareolus Schreber, 1780 depending on whether the female mated with one or two males. The work showed that the copulatory period of the bank vole lasted about 80 minutes and consisted of 4–5 ejaculatory series, with multiple intromissions preceding ejaculation. There were no significant changes in number of intromissions across the first four ejaculatory series, but I did find a relationship between number of intromissions and first ejaculation latency; also, ejaculation latencies grew shorter as the ejaculatory series proceeded. Litter size did not differ significantly between females that mated with one male and those mating with two, nor did the reproductive success of males that mated with the same female. Mating with an oestrus female appears to be advantageous for bank vole males even if they mate as the second one, and the risk of sperm competition did not trigger changes in male copulatory behaviour. The similar durations of the copulatory period and patterns of change of ejaculation latencies during copulations with one and two males point to the role of the female in temporal copulatory behaviour of the bank vole.     

Mongolian gerbil fathers avoid newborn male pups, but not newborn female pups: olfactory control of early paternal behaviour

We examined effects on the parental behaviour of male Mongolian gerbils, Meriones unguiculatus, of the sex and number of pups in a litter. Recent fathers interacting with foster litters consisting entirely of newborn males decreased the time that they spent in contact with a litter the greater the number of pups it contained. However, fathers that interacted with litters composed entirely of newborn females showed no change in the time they spent in contact with a litter as a function of its size. Fathers responded similarly to litters of 1- and 3-day-old female pups, but their responses to male pups changed from avoidance to approach as the age of males increased from 1 day to 3 days, and fathers made anosmic by intranasal administration of zinc-sulfate solution did not avoid neonatal litters. Results of a correlational study revealed that the more time males spent with newborn young during a 30-min test, the greater their latency to mate with their partners in postpartum oestrus and the shorter the duration of their mating effort during the 24 h immediately after parturition. We discuss these findings as consistent with the view that androgen-mediated olfactory stimuli produced by newborn male Mongolian gerbils make them unattractive to fathers, possibly functioning to increase the time that recent fathers mate-guard while their partners are in postpartum oestrus.     

Influence of time of mating and paced copulation on induction of pseudopregnancy in cyclic female rats

The present experiment was designed to examine whether changes occur during the course of behavioural oestrus in the sensitivity of the female to copulatory stimulation and in patterns of sexual behaviour which might influence the likelihood of luteal maintenance. Cyclic female rats were mated on the evening of pro-oestrus (21:00 h) or early on the morning of oestrus (05:00 h) and received either 5 or 10 intromissions from males under conditions which allowed or did not allow pacing of contacts with males to occur. During mating, the levels of sexual receptivity, the timing of sexual mounts from males, and pacing behaviours were measured. The occurrence of pseudopregnancy or pregnancy in each animal was determined by examining vaginal smears for 14 days after mating and by examining the uterus for the presence of fetuses at the end of the experiment. At both mating times, pacing of copulation with males increased the likelihood of prolonged luteal activity. However, females were more likely to become pseudopregnant following non-paced mating at 05:00 h than at 21:00 h the previous evening. Of those females receiving an ejaculation during mating, no difference were seen between groups in the incidence of pregnancy. This change in sensitivity to cervical-vaginal stimulation during oestrus was not associated with changes in sexual receptivity or pacing behaviour. The ability of cervical-vaginal stimulation to induce pseudopregnancy therefore increases toward the end of the period of oestrus, but the behavioural mechanisms which regulate receipt of such stimulation remain constant during that time.     

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 behaviour of male and female tammar wallabies (Macropus eugenii) at post-partum oestrus

This study describes the sexual behaviour of the adult male and female tammar wallaby from the time a female gives birth to the end of post-partum oestrus. Eleven females were continuously observed from around the time of birth until up to 3 h after the last observed mating attempt. Males began to associate with females from around the time of birth, thus females appear to be attractive to males at, but not prior to parturition. Females subsequently allowed males to mate with them from 1.3±0.8h (mean±S.D.) post-partum, although the first ejaculation was delayed until 1.8±0.8h after birth owing to intense intermale aggression directed towards any male which attempted to mate with an oestrous female. Mating chases, in which a post-partum female was pursued by several males, were observed from 1.3±0.8h post-partum. All of this mating activity usually occurred within the first 6h after birth.     

Behavioural aspects of the raccoon mating system: determinants of consortship success

We monitored raccoons Procyon lotor, in southern Texas during the 1990-1992 mating seasons to describe mating behaviour and identify factors affecting consortship success. During most of this study, raccoons were spatially aggregated, with female home ranges congregated around permanent water sources and larger home ranges of male groups encompassing each female group. Consortship success varied among males and ranged from zero to six females per male within a mating season. Individual females consorted with one to four different males during an oestrous period; however, most (62%) females consorted with only one male during their oestrus. Dominance through overt conflict appeared to influence male consortship success. During two mating seasons, one male from each group consorted with females on more days than all other males combined. Body weight of males was positively correlated with number of consortship days. As synchrony of oestrus increased, variance in number of consortship days among males decreased, and access to oestrous females increased for subordinate males. Wounding among males increased during the mating season, and was more frequent for males than for females. The mating system, as determined by consortship behaviour, appeared to shift between polygyny and promiscuity, and possibly varied annually as a result of the timing of oestrous cycles. Copyright 1999 The Association for the Study of Animal Behaviour.     

Long-term effects of accelerated or delayed sexual maturation on reproductive output in wild female house mice (Mus musculus)

The effects of acceleration and delay of puberty in female house mice on survival and reproduction were tested using 6 experimental groups: (1) control females mated at the time of first oestrus, (2) females mated at weaning, (3) females treated with male urine starting at weaning and mated at first oestrus, (4) females housed in groups and mated at first oestrus, (5) females housed alone, treated with urine from grouped females and mated at first oestrus, and (6) females housed alone and mated at 68 days of age. Females caged with males at weaning or treated with male urine and mated at puberty had lower rates of survival to 180 days of age, but did not differ in rates of fertility from mice in the other four treatments. Those females that were housed with males from weaning or treated with male urine also had smaller total numbers of litters, fewer total young, and smaller average litter sizes than did females for which the age of mating was delayed, by grouping or treatment with urine from grouped females, or by being held until age 68 days before mating. Control females mated at first oestrus generally were intermediate or did not differ from the male treatments on these dependent variables. There were no differences in the average number of female young/litter across the 6 treatments. However, females that were delayed in age of first mating had significantly more male young/litter than did females that were accelerated in their sexual development or control females.(ABSTRACT TRUNCATED AT 250 WORDS)     

Information asymmetries among males: implications for fertilization success in the thirteen-lined ground squirrel.

The classical view of mammalian mating competition focuses on combat and threat. By contrast, field studies have revealed that male mating success in some species is more strongly determined by mate location ability than by physical dominance. In thirteen-lined ground squirrels, competition in locating oestrous females is exacerbated by sperm competition that favours the first male to mate with a female. We used a female-removal experiment to identify the distinguishing characteristics of males that were the first at finding and mating with females. Each focal female was observed the day before she entered oestrus; the identities of males that made contact with her and the locations of their interactions were recorded. The following morning the females were temporarily removed, and we monitored male search behaviour in their absence. Males that arrived first were those that had spent more time with the female the previous day relative to their rivals. Time invested the day before, in turn, was highly correlated with male search persistence and familiarity with the female''s likely whereabouts. These results demonstrate that differential fertilization success can arise from information asymmetries among males: the advantaged individuals are those that have greater a priori knowledge of the reproductive state and spatial locations of prospective mates than rivals.     

Studies of the oestrous cycle, oestrus and pregnancy in the koala (Phascolarctos cinereus)

As an integral part of the development of an artificial insemination programme in the captive koala, female reproductive physiology and behaviour were studied. The oestrous cycle in non-mated and mated koalas was characterized by means of behavioural oestrus, morphology of external genitalia and changes in the peripheral plasma concentrations of oestradiol and progestogen. The mean (+/- SEM) duration of the non-mated oestrous cycle and duration of oestrus in 12 koalas was 32.9 +/- 1.1 (n = 22) and 10.3 +/- 0.9 (n = 24) days, respectively. Although the commencement of oestrous behaviour was associated with increasing or high concentrations of oestradiol, there were no consistent changes in the morphology or appearance of the clitoris, pericloacal region, pouch or mammary teats that could be used to characterize the non-mated cycle. As progestogen concentrations remained at basal values throughout the interoestrous period, non-mated cycles were considered non-luteal and presumed anovulatory. After mating of the 12 koalas, six females gave birth with a mean (+/- SEM) gestation of 34.8 +/- 0.3 days, whereas the remaining six non-parturient females returned to oestrus 49.5 +/- 1. 0 days later. After mating, oestrous behaviour ceased and the progestogen profile showed a significant increase in both pregnant and non-parturient females, indicating that a luteal phase had been induced by the physical act of mating. Progestogen concentrations throughout the luteal phase of the pregnant females were significantly higher than those of non-parturient females. Parturition was associated with a decreasing concentration of progestogen, which was increased above that of basal concentrations until 7 days post partum.     

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.     

Is Self‐Grooming by Male Prairie Voles a Predictor of Mate Choice?

Self-grooming by mammals is a form of scent dissemination in which individuals anoint themselves with salivary, anogenital, and other body odours. Self-grooming has been proposed to be a sexually selected trait favoured in reproductive competition and sexual attraction. We tested the hypothesis that females would show a mating preference for males that self-groomed more than a reproductive competitor that groomed less. In mate-choice experiments in which females had a choice of two tethered males, non pair-bonded females did not choose males based on their frequency of self-grooming. In a second experiment in which pair-bonded females in postpartum oestrus had access to their current mate and two strange males, strange males groomed significantly more than pair-bonded mates, yet attained the fewest copulations. Non pair-bonded females and pair-bonded males and females groomed significantly less often than did non pair-bonded males. Self-grooming behaviour was consistent with the sexual attraction hypothesis, but the frequency of self-grooming did not increase a male's mating success. We conclude that the frequency and time spent self grooming are not good predictors of mating success.     

Sexual and maternal behaviour at the postpartum oestrus: The role of experience in time-sharing

The postpartum sexual and maternal behaviour of Norway rats (Rattus norvegicus) was observed in a seminatural habitat. Behavioural decision-making by the female was examined by disrupting the nest and newborn offspring during mating. In experiment 1, multiparous females responded to nest disruption with an increase in maternal behaviour that was largely restricted to the post-ejaculatory intervals. The temporal and frequency measures of copulation were largely unaffected, implicating behavioural time-sharing. In experiment 2, following nest disruption primiparous females did not switch efficiently between maternal and sexual behaviour, showed poor retrieval of pups, and returned to the nest during ejaculatory series. They did not display the characteristics of behavioural time-sharing, indicating that previous experience may play a role in the ability to time-share.     

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.     

Sexual cannibalism in Nephila plumipes as a consequence of female life history strategies

The evolution of sexual cannibalism has been modelled as both an adaptive and nonadaptive female strategy. Recent evidence from several species suggests a connection between female foraging and sexual cannibalism, but the precise benefits for females have remained obscure. Here, we investigate the difference between cannibalistic and noncannibalistic female Nephila plumipes by removing the potential nutritional benefit of cannibalism. Courting and mating males that were killed by a female were immediately removed so that the female could not consume them. Nevertheless, cannibalistic females gained more mass from maturation to oviposition and produced larger first clutches than noncannibalistic females, although cannibalistic females matured at a smaller size and mass than noncannibalistic females. In juvenile instars, mass gain was generally smaller in females that moulted in a good condition but intermoult intervals were shorter. However, the time from maturity to oviposition was not shorter in females that matured in a good condition. Male behaviour did not differ according to the risk of cannibalism. We suggest that sexual cannibalism in N. plumipes is a side‐effect of an increased foraging vigour of females that matured at a smaller size and body mass. Selection pressure on males to avoid cannibalism may be weak because of limited mating opportunities.     

Effect on oestrus and ovulation of exposing creole goats to the male at three times of the year

The characteristics of the oestrous and ovarian cycles of local goats were studied over 2 years, on 96 females in March, 60 in July and 100 in November. After 3 weeks of separation between sexes, the females with inactive ovaries were identified and harnessed males were introduced. Detection of oestrus was undertaken daily and the date and rate of ovulation were checked at laparoscopies. The proportion of females with non-cyclic ovaries before the mating period varied according to the season and whether the females were nulliparous or suckling. Among the females with inactive ovaries before the mating period, 97% ovulated 2 . 8 days after the introduction of males; 68% of these initial ovulations were accompanied by oestrus and 76% resulted in short luteal phases (5 . 3 days) followed by a second ovulation accompanied by oestrus 89% of the time. The proportions of initial ovulations with oestrus and of initial ovulations followed by a short luteal phase were in linear relationship with the proportion of females with non-cyclic ovaries before the mating period. For the females with cyclic ovaries before the mating period, the distribution of first oestrus during the 8 days after the introduction of males was significantly different from the expected uniform distribution. A possible luteolytic action of teasing was suspected. The fertility of females in all groups was high (greater than 79%). Litter size was not different for non-cyclic and for cyclic females.     

Sexual behaviour and reproductive competition in male wild and domestic Norway rats

The copulatory behaviour of male wild Norway rats (Rattus norvegicus) exposed individually to females in hormone-induced oestrus is characterized by fewer mounts and intromissions per ejaculatory series, shorter ejaculation latencies, and longer ejaculatory clasps than the copulatory behaviour of domestic males. Competition between wild and domestic males tested simultaneously in pairs produced only minor changes in copulatory behaviour. The reproductive success of wild and domestic male pairs simultaneously exposed to naturally cycling females was nearly equal in both the laboratory and an outdoor enclosure. Genotype of the female had little effect on male copulatory behaviour. The copulatory behaviour of sexually naive and experienced domestic rats did not differ. However, most sexually naive wild males failed to exhibit copulatory behaviour under the test conditions employed.     

Vibratory courtship in a web-building spider: signalling quality or stimulating the female?

Courtship behaviour in spiders in the form of premating vibrations by males may function (1) as a male identity signal used for species recognition, (2) in suppression of female aggressiveness, (3) to stimulate female mating behaviour, or (4) as a quality signal used in female choice. We investigated the function of web vibration by male Stegodyphus lineatus in a series of experiments. Regardless of vibratory performance, all males mated successfully with virgin females but only 56.4% of males mated with nonvirgin females. Vibratory performance did not influence male mating success, but heavier males had a higher probability of mating with mated females. Males vibrated less often and produced fewer vibrations when introduced on the web of a mated female. Males that vibrated webs of virgin females mated faster than nonvibrating males, but there was no effect of vibration rate or body mass. There was no effect of male vibratory effort or vibration rate on female reproductive success measured as time to egg laying, clutch size, number of hatched young, number of dispersed young and offspring body mass after a single mating. Males vibrated on abandoned virgin female webs but the response decreased with increasing duration of female absence, suggesting that females produce a web-borne pheromone, which elicits male vibrating behaviour. Mated females were less receptive and not stimulated by male vibrating behaviour. We conclude that male premating vibrations in S. lineatus do not function as a male quality signal selected via female choice. Rather, the primary function of this behaviour may be to stimulate a receptive female to mate. Copyright 2003 Published by Elsevier Ltd on behalf of The Association for the Study of Animal Behaviour.      

Reinforcing stimulation of the postero-lateral hypothalamus and mating behaviour in the rat.

Male rats implanted with chronic electrodes into the postero-lateral hypothalamic site were tested for self-stimulation behaviour. Rats exhibiting steady self-stimulation behaviour were observed during mating tests with an oestrus female. During these tests the hypothalamus of male rats was stimulated. Results show that no stimulus-bound sexual behaviour was observed. Nevertheless, correlations were found between the rewarding value of the cerebral stimulation and the specific sexual components of mating behaviour. These results are interpreted with the hypothesis of a balancing effect between the reward elicited by direct stimulation of the brain and reward acquired by the presence of an oestrus female.     

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.