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Mating in Heliothis virescens: transfer of juvenile hormone during copulation by male to female and stimulation of biosynthesis of endogenous juvenile hormone.

Studies were undertaken to determine whether adult males of Heliothis virescens transfer juvenile hormone (JH) to females during copulation, and an in vitro radiochemical assay was used to determine whether mating causes an allatotropic effect, i.e., stimulation of JH biosynthesis by corpora allata (CA). In vitro, CA from 3-day-old mated females synthesized and released approximately 2.5 times total JH as that of CA from comparably aged virgin females. Of the homologues, JH II exhibited significant increase in mated females; JH I also increased but not significantly. JH III remained similar to that of virgin females. This is the first demonstration of an allatotropic effect of mating in moths. In contrast to the female, CA of virgin males did not produce any JH, but accessory sex glands (ASG) in 3-day-old males synthesized small amounts of JH. Immediately after adult emergence, male ASG contained approximately 1.5 ng JH I and II, which increased by 12 h after emergence and remained at this high level up to 54 h after emergence. JH III was barely detected in ASG. JH in ASG of mated male immediately after uncoupling was depleted almost completely, and 24 h later recovered to levels comparable to that of 54-h-old virgin male. Virgin female bursa copulatrix did not contain any JH, but mated female bursa, immediately after uncoupling, had JH at levels comparable to that observed in virgin male ASG. By 6 h after uncoupling, JH levels decreased dramatically in mated female bursa. These data suggest the transfer of JH to females by the male.

Animals↗

Male mating strategies and reproductive constraints in a group of wild tufted capuchin monkeys (Cebus apella nigritus).

Tufted capuchin monkeys (Cebus apella) provide an extreme example of active female sexual solicitation of males. In spite of being targeted by females for sex, males may delay copulation for hours or days. Data were collected on the sexual interactions in one wild capuchin group at the Estação Biológica de Caratinga in Brazil from September 1996 to August 1997. All successful conceptions during this year occurred in the dry season, yet sexual behavior was observed during 9 months of the year. This study tested whether male sexual response to female proceptivity was seasonally-mediated. Male consortship participation, solicitation of females, latency to copulation, and copulation frequency were compared between fertile and nonconceptive females. Seasonal patterns in copulation interference, mating style, and alternative mating strategies were also examined. Thirty-two copulations were observed. The alpha male was solicited for significantly more consortship days per female, but his mating success, in terms of copulation frequency, did not differ from that of two other adult males in the group. In the dry season, when the females were fertile, the males showed increased contest competition for mates, a higher frequency of alternative mating strategies against copulation interference, and increased monitoring of the females' condition. However, contrary to expectations, the alpha male's latency to copulation was significantly longer in the fertile season than in the nonconceptive months, and no males were observed to mate more than one time per day, even at the conceptive peak. Male mating strategies were affected by both season and rank, and there was evidence for reproductive constraints on males throughout the year. Limited male ejaculatory capacity and male choice in the timing of copulations within female proceptive phases may both be important factors in driving the sexual dynamics of this species.

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Sexual selection in Japanese macaques I: female mate choice or male sexual coercion?

Mate choice is defined as a behavioural pattern in one sex which increases the probability of fertile matings with certain members of the opposite sex. The chosen sex may use force, however, to coerce matings from reluctant members of the choosing sex. In a confined group of Japanese macaques, Macaca fuscata fuscatahoused at the Primate Research Institute of Kyoto University, Japan, behavioural and endocrine data were combined to assess the validity of several mate-choice behaviours and the potential influence of sexual coercion. Preliminary evidence was found for both female mate choice of males and male sexual coercion of females. Both female proximity maintenance towards males and male aggression towards females were correlated with an increase in fertile matings. Most male aggression appeared to be the by-product of increased time spent in proximity due to female proximity maintenance, but some male aggression appeared to be sexual coercion. These results are interpreted in light of recent research on male sexual coercion, and a tentative model of mate choice for this species is proposed.1997The Association for the Study of Animal Behaviour

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Mate-choice copying in Japanese quail, Coturnix coturnix japonica

We performed four experiments to examine effects on the mate choices of female Japanese quail, Coturnix coturnix japonica, of observing a male mate with another female. Each experiment was conducted in three phases: (1) a pre-test during which subject females were allowed to choose between two males with which to affiliate; (2) an observation phase, in which subject females either watched or did not watch the male they had spent less time near during the pre-test (their 'non-preferred' male) copulate with a 'model' female; and (3) a post-test when subject females again chose between non-preferred and preferred males. Only females that had watched their non-preferred male mate with a model female during the observation phase spent significantly more time affiliating with him during the post-test than they had during the pre-test. Watching mating did not change females' criteria for choosing males, and non-preferred males that had mated recently were no more attractive to females than were non-preferred males that had not done so, unless subject females actually observed the mating take place. The results were consistent with the hypothesis that female quail copy one another's mate choices. Copyright 1998 The Association for the Study of Animal Behaviour Copyright 1998 The Association for the Study of Animal Behaviour.

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Non-genetic benefits of mate choice: fecundity enhancement and sexy sons.

I compared reproductive success (lifetime number of fertilized eggs) as a function of mate choice among females of the stink bug, Nezara viridula L. (Hemiptera: Pentatomidae). 'Choosing' (C) females were placed with one of two males on alternate days. CI females chose between inexperienced males while CR females chose between males previously rejected by CI females. 'Non-choosing' (N) females were placed with the same male every day. Non-choosing NI, NR and NA females encountered, respectively, inexperienced males, previously rejected males, or previously accepted and mated males. Reproductive success was highest for CI females, showing direct selection on mate preferences. Reproductive success did not differ between CR and NR females, indicating that male quality, not the act of choosing a mate, affects fitness. CI females preferred males with longer antennae and their fecundity (lifetime number of eggs) was correlated with male antenna length, consistent with antenna length as an indicator of male ability to transfer nutritive sperm produced in paired harlequin lobes of the testes. Harlequin lobes were smaller in rejected than chosen males. In second-generation mate choice trials, sons of NR females competed well against sons of NA females but not against sons of CI females. This suggests that non-genetic paternal contributions that decline with prior mating account for the attractiveness of sons because sons of CI and NA females shared the same fathers. Sons experiencing mating success came from larger eggs and egg size was greatest for CI females, perhaps as a consequence of paternal nutritional contributions. Copyright 1998 The Association for the Study of Animal Behaviour. Copyright 1998 The Association for the Study of Animal Behaviour.

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Male potential reproductive rate influences mate choice in a bushcricket.

Male bushcrickets, Requena verticalis, provide their mates with a proteinaceous nuptial gift, which functions both as paternal investment in offspring and to ensure sperm transfer. When nutrients are limited, males have a lower potential reproductive rate than females, so the operational sex ratio becomes female biased. Males are then expected to discriminate more in their choice of mate for two reasons: (1) the relatively higher female potential reproductive rate should reduce the costs of rejecting a female; (2) multiple mating by females should increase variance in female mate quality, because of first male sperm precedence. Males are known to discriminate against old females, which are more likely to have mated previously. Our objective in this study was to partition experimentally the influences of male potential reproductive rate (manipulated by diet) and variance in female quality (estimated by age) and to assess their relative influence on the level of mate choice of male bushcrickets. The potential reproductive rate of males had a great impact on their choosiness, whereas variance in female age had no effect. These results support recent theoretical models that predict costs, rather than benefits, will primarily influence the level of mate choice. Copyright 1998 The Association for the Study of Animal Behaviour. Copyright 1998 The Association for the Study of Animal Behaviour.

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Assortative mating and female clutch investment in black grouse.

Variation in female behaviour has only recently received attention in studies of sexual selection. It has been suggested that females may invest differentially in their offspring in relation to the quality of their mate. This may lead to females that mate with high-quality and/or attractive males laying larger clutches. Females may also differ in their ability to choose between males. For example, females in good physical condition may make better choices. If physical condition and clutch size are positively correlated, this hypothesis could also produce a relationship between male attractiveness and female clutch size. We found, in lekking black grouse, Tetrao tetrix, that females mated to the highest ranked males laid the largest clutches. Furthermore we found, regardless of female age, a positive relationship between a measure of female condition and male rank but not between female condition and her clutch size. In addition, females in good condition visited a larger number of different male territories, and old females produced the largest clutches. Our results suggest two mechanisms to explain our findings. First, females in good physical condition tend to mate with the top males, suggesting an assortative mating pattern. Second, females mating with the highest ranked males lay larger clutches as a consequence of their choice. In general, our result calls for caution in evaluating studies that look at the consequences of mate choice. It may be that differences in female quality produce effects that may be wrongly interpreted as male quality effects. (c) 1998 The Association for the Study of Animal Behaviour.

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The cost of peripheral males in a brook trout mating system.

A focus on the reproductive contributions of males displaying alternative life histories has neglected the role of size-dependent peripheral males in salmonine mating systems. We documented mating behaviour of brook trout Salvelinus fontinalis, including observations of spawning, over two breeding seasons to determine the mating costs of peripheral males to dominant males (kleptogamy) and females (egg cannibalism). For males and females, the mating costs of peripheral males were substantial because more than half (56%) of all observed brook trout spawnings involved peripheral males. Males that paired with large females experienced a greater incidence of kleptogamy due to increased numbers of peripheral males present. Large males face a conflict when mating in that they prefer to spawn with large females; however, these same females attract numerous males against which the dominant male cannot defend. From paternity studies, we estimated that males that had peripheral males participate in spawning may fertilize, on average, equal numbers of eggs compared to males spawning solely with a smaller female. Females that paired with relatively smaller males had significantly more eggs eaten by peripheral males than females that paired with relatively larger males. Latency to spawn by females increased when paired with a relatively small male, and resulted in females obtaining a larger spawning partner. The observed patterns of size-assortative mating, kleptogamy and cannibalism are discussed in relation to mate choice for this population of brook trout. Copyright 1999 The Association for the Study of Animal Behaviour.

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Cumulative long-term investment in vocalization and mating success of fallow bucks, Dama dama.

We carried out behavioural observations to investigate the function of long-term investment in vocal display by fallow bucks during the breeding season. The measures of long-term investment used were the date of initiation of vocal activity, the number of days vocal during the breeding season, and the proportion of time spent vocalizing. We analysed data from 3 years (1993-1995) to assess the relationship between the date of initiation of vocal activity, and the number of days vocal, and age, dominance rank and mating success. Observations from a sample of focal males in 1996 were used to determine the effect of the proportion of time vocal during the breeding season on dominance rank and mating success. The majority of socially immature males (</=3 years old) did not vocalize; among socially mature males (>/= 4 years old), dominance rank was more important than age in explaining variation in vocal activity. The onset of vocal activity by fallow bucks was not a direct consequence of the presence of mating opportunities since the first males became vocal more than 3 weeks before any matings occurred. Long-term investment in vocal activity did not alter the dominance relationships that had been established between males before they became vocal. When we considered all mature males from 3 years of observations, the majority of matings were achieved by those that had high rank, initiated vocal activity early during the breeding season and remained vocal on most days. For the 1-year sample of mature males, the factor most highly correlated with mating success was the proportion of time that males spent vocalizing during the rut. Thus we have shown a strong relationship between the time invested in vocal display by fallow bucks and their mating success. Copyright 1999 The Association for the Study of Animal Behaviour.

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Parasites promote mating success: the case of a midge and a mite.

I tested the prediction that the hydracharinid mite Unionicola ypsilophora reduces mating success in the chironomid midge Paratrichocladius rufiventris. Males of the midge form mating swarms through which females fly to emerge after a few minutes with a mate. This mating system is believed to depend upon the male capturing a mate after aerial competition between males. Thus aerobatic ability is expected to determine success and a large ectoparasite should impair aerial performance. The proportion of infested males in swarms (ca. 4%) was less than that in mated pairs (ca. 15%). Infestation thus improved the expected mating success of the male midge. This finding is counterintuitive and may be a chance effect of no adaptive value to host or parasite. Alternatively, it may be an adaptive manipulation of the host by the parasite. This study provides evidence for the latter explanation. Copyright 1999 The Association for the Study of Animal Behaviour.

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Female-female aggression and female mate choice on black grouse leks.

We studied female-female aggression in relation to female mate choice in black grouse, Tetrao tetrix, in central Finland, in 1994-1998. Aggression occurred on average every other minute when there was more than one female on a territory, and aggressive behaviour was most prominent when several females attended the lek. Interactions tended to be proportionally most frequent on the territories of the highest-ranking males, although not significantly so. Females that were chased by other females did not mate with lower-ranking males than their aggressors did. Furthermore, chased females were only rarely (6% of cases) forced to move off the territory by agonistic interactions and copulations were disrupted by other females even less often (3% of cases). The choice of a mating territory did not depend on the outcome of aggression even though the aggressors were more likely to mate on the territory where aggression occurred than elsewhere. There was a marginally significant tendency for aggressors to mate earlier in the season. Females placed themselves further away from other females on the territory when soliciting a copulation than just before aggression. Our results suggest that aggression between females does not effectively constrain female choice in black grouse. Its function may be to aid females to secure undisturbed mating opportunities for themselves rather than to prevent others from mating with a particular male. Copyright 2000 The Association for the Study of Animal Behaviour.

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Differences between the sexes in direction and duration of response to seeing a potential sex partner mate with another.

We have shown previously that: (1) female Japanese quail, Coturnix japonica, increase and male Japanese quail decrease their tendencies to affiliate with potential sex partners after seeing them mate, and (2) in both sexes of quail, affiliative preferences and choice of a sex partner are highly correlated. Here we predict that because effects of a prior male's sperm on a second male's probability of fertilizing a female are relatively brief, a male's avoidance of whichever member of a pair of females he has seen mating should be transitory. Conversely, because female quail seek high-quality males as mates and quality is a relatively permanent characteristic, females' preferences between males should remain constant over time. We found, as predicted, that the durations of effects on affiliation of seeing a potential sex partner mate differed in male and female quail. Forty-eight hours after male quail saw a female mate, they no longer avoided her, whereas 48 h after female quail saw a male mate, his attractiveness remained enhanced. We conclude by suggesting that both the direction and the duration of responses of male and female Japanese quail to seeing a member of the other sex mating enhance the fitness of members of each sex. Copyright 2000 The Association for the Study of Animal Behaviour.

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Mate fidelity and breeding site tenacity in a monogamous sandpiper, the black turnstone.

We examined the relationship between mate fidelity and breeding site tenacity during a 5-year study of the black turnstone, Arenaria melanocephala, a socially monogamous sandpiper breeding in subArctic Alaska. We tested the predictions of several hypotheses regarding the incidence of divorce and the benefits of fidelity to mate and breeding site. Interannual return rates to the breeding grounds (88% for males, 79% for females) were among the highest yet recorded for any scolopacid sandpiper, and 88% of returning birds nested on their previous year's territory. The annual divorce rate was only 11%, and mate fidelity was significantly linked to fidelity to territory but independent of sex and year. Males arrived in spring significantly earlier than their mates and interannual fidelity was influenced by the relative timing of arrival of pair members. Reunited pairs had significantly higher fledging success than new pairs formed after death or divorce. The incidence of divorce was unrelated to reproductive success the previous year, although birds nested significantly further away after failure than after a successful nesting attempt. Sightings of marked individuals suggested that members of pairs do not winter together, and breeding site tenacity provides a mechanism through which pair members can reunite. We reject the 'incompatibility' hypothesis for divorce in turnstones, and our data contradict predictions of the 'better option' hypothesis. Alternatively, we propose the 'bet-hedging' hypothesis to explain the occurrence of divorce, which transpires when an individual pairs with a new mate to avoid the cost of waiting for a previous mate to return. Such costs can include remaining unmated, if the former mate has died, or experiencing lower reproductive success because of delayed breeding. Copyright 2000 The Association for the Study of Animal Behaviour.

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Only self-paced mating is rewarding in rats of both sexes.

When rats are mated in a traditional mating chamber (with one male and one female) in which the male dictates the pace of the copulatory sequence, males develop a reward state as evaluated by conditioned place preference (CPP). In this mating situation no reward state is induced in females. However, when female rats are able to control (pace) the rate of sexual stimulation, thereby reducing the aversive consequences associated with mating, a clear CPP is observed. In the present study the CPP paradigm was used to determine whether if the reinforced state induced by coital interactions in male rats can be maintained when females pace the sexual interaction. Adult male and female rats were mated in one of two different conditions: (1) where subjects were able to pace their coital interactions or (2) where subjects were not able to pace their sexual contacts. The results showed that when males had control over the sexual interaction they developed a clear place preference while males that mated with females that paced their coital contacts did not develop CPP. Similarly, only females that were able to pace their sexual contacts developed place preference. These results suggest that coital interactions in males, as well as in females, can induce a reward state only when they are able to control the sexual interaction. Under seminatural conditions sexual behavior in rats is highly promiscuous, they mate in groups and repeatedly change partners in the middle of copulation. This behavioral sequence allows both, male and female to control the rate of sexual interaction, assuring the induction of a reward state outlasting the actual performance of coital responses.

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Temporal and spatial quantitation of nesting and mating behaviors among mice housed in a semi-natural environment.

Enhanced behavioral complexity may be observed when animals are tested in naturalistic environments and engaging in nonforced social interactions. For each of six experimental runs, different groups of five adult Swiss-Webster mice (four ovariectomized females and a single male) were maintained under 12h dark:12h light in a 122 x 122 x 30.5-cm open box containing six peripheral "nestboxes." On Day 1, females were released into the box first and their nesting behavior was observed. Two days later, each female was injected with estradiol benzoate and the male was introduced into the environment. Females were injected with progesterone (P) 48 h later and the animals were observed for an additional 14 h. Behaviors were recorded with a video camera suspended over the apparatus. Mating occurred only post-P and males always mated preferentially with certain females. The amount of nesting behavior per female on Day 1 correlated significantly with the number of times each female was mated by the male (r = 0.57, P < 0.005). In all but one run, the male ejaculated with the female who performed the most nesting behavior. While 63% of mating was in the open, 56% of nestbox matings resulted in female postmating darting to alternate nestboxes; in 19% of these cases, the female quickly returned to the mating nestbox and was mated there again. Direct approaches by females to the male and behaviors which affect pacing were observed. These behaviors have not been reported previously for mice and may provide additional endpoints for the exploration of hormonal and genetic influences on reproductive behaviors.

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Prenatal stress alters reproductive responses of rats in behavioral estrus and paced mating of hormone-primed rats.

Adult female offspring of dams exposed to gestational stress (prenatal stress, PNS) may show altered reproductive behavior, exploration in novel environments, and/or social interactions than do their non-PNS counterparts. These behavioral differences may be more readily observed in a seminatural, paced mating paradigm, in which females have greater control of their sexual contacts, than in a standard mating situation. Adult offspring of dams exposed to restraint and lights for 45 min on Gestational Days 14-20 (PNS) were compared with those not subjected to stress (non-PNS, control condition). The motor, reproductive, and sociosexual behaviors of hormone-primed (Experiment 1) or cycling adult offspring in behavioral estrus (Experiment 2) were examined following 20 min of restraint stress under bright lights (postnatal stress). Hormone-primed PNS rats displayed less motor behavior in a novel arena than did non-PNS rats. In a standard mating test, hormone-primed PNS females tended to be more aggressive toward the male than were non-PNS rats. In a seminatural mating situation, hormone-primed PNS females showed increased avoidance behavior, such as longer latencies to the initial intromission, greater return latencies following mounts and intromissions, and more exiting subsequent to mounts and intromissions, than did non-PNS rats. PNS rats in behavioral estrus had decreased incidence and intensity of lordosis, and fewer solicitation behaviors, in both standard or paced mating situations, in which latency to and number of mounts were also increased. Thus, hormone-primed PNS rats exposed to restraint showed more avoidance behaviors in paced mating situations, while cycling PNS rats in behavioral estrus had greater disruption of reproductive responses in standard or paced mating paradigms than did non-PNS control rats.

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Structure-activity relationships in corpora allata of the cockroach Diploptera punctata: roles of mating and the ovary.

Morphometric studies were made on corpora allata of the cockroach Diploptera punctata from animals in which increasing gland size is not coupled to hormone synthesis (ovariectomized mated females; last-instar larvae) and in which gland size is coupled to hormone synthesis (normal mated and virgin females; penultimate-instar larvae). Cell number, gland volume, and juvenile hormone synthesis were measured. From electron micrographs, nuclear, cytoplasmic, and extracellular volumes; and cell membrane area were calculated; and fine structure described. Low-activity glands of ovariectomized mated females resembled high-activity glands from mated females in high cell number, large overall and cytoplasmic volume, and low nuclear-cytoplasmic ratio; they differed in having organelles typical of low-activity glands, mitochondria with dense matrices and large whorls of smooth endoplasmic reticulum. Inactive last-instar larval glands resembled mated ovariectomized female glands in increased cell number and organelles characteristic of inactive glands; however, their nuclear-cytoplasmic volume ratio was much higher. Penultimate larval glands with high activity per cell resembled active glands of normal mated females. Ovariectomy did not change morphometric parameters of virgin female glands; thus mating results in increase in size of adult female glands whereas the growing ovary is needed for changes in mitochondria and endoplasmic reticulum associated with high juvenile hormone synthesis.

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Analysis of truncated distributions: mating speed in Drosophila melanogaster.

The distribution of mating speeds in wild-type Drosophila melanogaster is shown to be log normal. The analysis of mating speeds by methods for truncated distributions is validated, and unbiased estimates of the mean mating speed, the variance of mating speed, and the proportion of flies capable of ever mating are produced. In pair matings, not all pairs are capable of copulating, given even a 7-day mating period.

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