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The Gbeta(KlSte4p) subunit of the heterotrimeric G protein has a positive and essential role in the induction of mating in the yeast Kluyveromyces lactis.

In the yeast Saccharomyces cerevisiae the Gbetagamma dimer of the heterotrimeric G protein transduces a pheromone signal from serpentine receptor to a MAP kinase cascade that activates the mating response pathway. Haploid cells lacking the Gbeta subunit do not respond to sexual pheromone, leading to sterility. In this work we demonstrate that the beta-subunit of Kluyveromyces lactis, encoded by the KlSTE4 gene, is a component of the G protein, and that its disruption gives rise to sterile cells. However, unlike Ste4p in S. cerevisiae, its overexpression does not induce growth arrest or promote mating. It has been shown that in K. lactis, the Galpha subunit has a positive role in the mating process, hence the resulting double GalphaDelta GbetaDelta mutant was viable and sterile. Here we show that the overproduction of Gbeta subunit fails to rescue GalphaDelta mutant from sterility and that expression of a constitutive active allele of Galpha enhances transcription of the KlSTE4 gene. The mating pathway triggered by the Gbeta-subunit requires a functional KlSte12p transcription factor. Gbeta has a 10-fold higher association rate with the Galpha1 subunit involved in pheromone response than with Galpha2, the protein involved in cAMP regulation in K. lactis. Additionally, the Gbeta-subunit from K. lactis is able to interact with the Galpha-subunit from S. cerevisiae but fails to restore the mating deficiency of Scste4Delta mutant. The data presented indicate that the mating pathway of K. lactis is positively and cooperatively regulated by both the Galpha and the Gbeta subunits.

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Copulation and mate-guarding patterns in polygynous European starlings

We recorded the timing and frequency of copulation and mate-guarding behaviour during the fertile period in relation to day of first ovulation and time of day in the facultatively polygynous European starling, Sturnus vulgarisAll within-pair copulations were female-solicited. Females solicited them at a high rate during both the pre-ovulatory and ovulatory period, but there were differences in solicitation rates between monogamous, primary and secondary females. Extra-pair copulation attempts were all male-initiated and females rarely engaged in them. Attempts by males were most frequent during the ovulatory period, when the probability of extra-pair fertilization was highest. Males intensively guarded their mate during both the pre-ovulatory and ovulatory period. In contrast to most other passerines, female starlings lay their eggs not at dawn but usually between 0900 and 1100 hours, implying that they may be most fertile in the late morning. During the ovulatory period, extra-pair copulation attempts were more frequent in the late morning than in the early morning, suggesting that males try to take advantage of this putative 'insemination window'. Accordingly, there was some indication that mate guarding during the ovulatory period was more intense in the late than early morning. Females solicited within-pair copulations at a higher rate in the early morning than in the late morning, but copulation rate did not differ between early and late morning. The proportion of monogamous and polygynous males attempting extra-pair copulations did not differ significantly, but mate-guarding intensity differed significantly between female categories. Primary females were guarded less intensively than monogamous females suggesting that males trade mate guarding against mate attraction/courting additional females.

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Size-assortative mating in Drosophila malerkotliana

The relationships between wing size, locomotor activity, mating latency, courtship pattern and mating success (total number of pairs mated in a given time unit) were studied in pairs of Drosophila malerkotlianaMating latency was negatively correlated with wing length and with locomotor activity while wing length and locomotor activity were positively related. In trios of a female and a long-winged and a short-winged male, long-winged males mated more successfully than short-winged males. In trios of a male and a long-winged and a short-winged female, long-winged males mated with long-winged females, and short-winged males with short-winged females. In addition, in pairs of long-winged/short-winged flies, long-winged flies courted and mated more successfully than short-winged flies.

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Evolution of mate-choice copying: a dynamic model.

Mate-choice copying has recently been demonstrated in several species. Two, not mutually exclusive, explanations for copying have been proposed: it reduces sampling costs and/or error of mate choice. In guppies, Poecilia reticulata, and black grouse, Tetrao tetrix, young females seem most likely to copy. Therefore, copying may teach inexperienced females what attractive males look like. I developed a 2-year dynamic model, to investigate under which conditions a mate-copying strategy might first evolve. An original population of pure choosers was assumed, which was invaded by a mutant female, able to copy during her first mating season, thereby instantly improving her ability to assess male quality. Alternatively, she could either wait and learn by observing males, just as non-copiers may do, but incurring some time costs, or choose, relying on her own abilities. The degree to which copying occurred among these mutant, young, inexperienced females increased with an increasing proportion of old, experienced females in the population, and with decreasing time left until the end of the season. The model demonstrates that mate-choice copying may evolve, when young females are poor at discrimination and need to learn what high-quality males look like. Male quality proved to be unimportant for copying to evolve, as long as there are sufficient differences in quality for mate choice to be meaningful. As with previous models, time constraints are an important assumption for copying to be advantageous over non-copying. Copyright 1998 The Association for the Study of Animal Behaviour. Copyright 1998 The Association for the Study of Animal Behaviour.

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Variance in male mating success and female choice for persuasive courtship displays.

An important function of the courtship displays of male animals is to stimulate, or 'persuade', females to mate. While many studies of courtship success have shown that females choose stimulating males as partners, fewer have demonstrated that the most persuasive males in a population also enjoy the highest mating success (i.e. mate with the greatest numbers of partners). Such a relationship provides a stronger test of the hypothesis that male displays evolve by sexual selection via female choice. We tested this hypothesis in the laboratory using the North American plethodontid salamander, Desmognathus ocoee, in which females only elicit spermatophore deposition if rendered sexually responsive by male courtship. First, we determined variance among individual males in the numbers of females with which they mated across 35 encounters, in the absence of differences in partner encounter rates and direct intermale interactions. Second, we demonstrated that males with high 'historical' scores of mating success performed significantly higher frequencies of persuasive courtship displays, which provide tactile, chemical and visual stimulation to females. We conclude that sexual selection via female choice favours persuasive displays because they confer high mating success on the males that perform them at the highest frequencies. Copyright 1998 The Association for the Study of Animal Behaviour.

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Scent marks as reliable signals of the competitive ability of mates.

The quality of an individual's odour can allow potential mates to discriminate against individuals of low social class, poor health status or unsuitable genotype. Competitive scent marking provides a further mechanism which could allow mates to discriminate between individuals of apparently high quality. The presence or absence of fresh countermarks from competitors within an owner's territory or area marked by a dominant animal provides a reliable indicator of the owner's ability to defend its territory or dominate competitors. This could be used by potential mates to discriminate between individuals advertising their apparently high competitive ability through their scent-marking behaviour and odour quality. We tested this by manipulating scent marks in the neighbouring territories of wild-caught male house mice, Mus domesticus. As predicted, oestrous females used scent marks to select males apparently able to defend exclusive territories over those unable to exclude intruders. Females were more strongly attracted to the odour of owners of exclusively marked territories and showed more sexually related behaviour when interacting with these males. Furthermore, while females preferred a territory containing a better protected nest site regardless of the owner's apparent competitive ability, they still used the presence or absence of intruder countermarks when selecting a potential mate. This suggests that females use scent marks as a reliable signal of the best-quality mate among neighbouring males independently of their nest location. Since assessment depends on both the territory holder's own marks and those of competitor males, countermarking is likely to be an important mechanism of competition for mates between neighbours. Copyright 1998 The Association for the Study of Animal Behaviour

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Individual mating success, lek stability, and the neglected limitations of statistical power.

The evolution of leks (aggregations of males displaying to females) cannot be explained solely by an increasing average gain in matings for each male as group size increases. This is because the mating skew, that is, the inequality among males in mating success, is often high and may vary with lek size. Here, we show that the common observation that matings become more evenly divided as lek size increases is also insufficient to explain by itself the benefits of aggregating. The benefits to individual males are highly sensitive to the exact relationship between mating skew and lek size, and very similar relationships can lead to opposite predictions concerning individual benefits. With data on published mating success for 18 species (71 leks), we show that different species have very similar skew versus lek size relationships. With current sample sizes, however, there is insufficient statistical power to distinguish between completely different alternatives concerning individual optima of males. Copyright 1998 The Association for the Study of Animal Behaviour

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A role for pheromones in mate choice in a lekking sandfly.

There is increasing evidence to suggest that pheromone-mediated communication may play an important role in mate choice in many insects. We investigated the possible role of pheromones in mate choice in the lekking sandfly, Lutzomyia longipalpis, and explored whether males or females display any behavioural traits that predict patterns of male mating success. Pairs of virgin males were introduced into a small mating chamber and allowed to interact for approximately 10 min prior to the introduction of a single virgin female. The trial continued until the female copulated with one of the males. We recorded trials on video for later analysis. After a successful copulation, we determined the quantity of pheromone remaining in the glands of both males and measured the size of the pheromone glands and tergites. We corrected behaviours and measurements for body size (tergite width). Mated males had significantly more pheromone present in their glands after a trial than unmated males. Males differed in how much time they invested in wing fanning. Members of each pair of males spent similar amounts of time wing fanning but mated males and males with more residual pheromone spent more time wing fanning than their counterparts throughout the trial. The time spent wing fanning and the amount of residual pheromone were not correlated, however. A male's ability to win fights did not relate to his reproductive success. Finally, a female was equally likely to wing-fan directly at, or walk away from, either male in a given pair. The possible relationships between the amount of pheromone released and the residual pheromones left in the glands, and the level of male wing fanning and pheromone productivity, are discussed. Copyright 1998 The Association for the Study of Animal Behaviour.

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The role of calceoli in mate assessment and precopula guarding in Gammarus.

Gammarid crustacea exhibit precopula mate guarding in which a male will carry a potential mate beneath his ventral surface, guarding her for several days until she moults and lays her eggs. I investigated the role of the calceoli (bulb-like structures on the second antennae) in mate assessment and precopula pairing in two Gammarus species by comparing the behaviour of males with and without calceoli. Removal of the calceoli had no effect on the pattern of size-assortative mating, nor did it reduce the number of contacts made per unit time between potential mates. However, males without calceoli reduced their assessment of potential mates, were less likely to guard a suitable female and guarded for a shorter period. These data support the hypothesis that calceoli are used during contact assessment to determine a female's intermoult period and hence suitability for guarding. (c) 1998 The Association for the Study of Animal Behaviour.

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Mating ecology of the nonpollinating fig wasps of Ficus ingens.

The males of many fig wasps fight fatally for mating opportunities. The concentration of females in space has been proposed as one factor selecting for their aggressive behaviour. We studied the temporal distribution of receptive females to obtain a clearer impression of the operational sex ratio in figs. Females of nonpollinating species emerged from figs over a protracted period of time and this resulted in an extremely male-biased operational sex ratio, conducive to extreme fighting. Since there were so few receptive females at any one time, a male could defend an eclosing female. Consequently, the largest, Otitesella longicauda male in a fig had a much higher mating success than smaller males. This suggests that larger males have a larger fitness advantage than larger females and a Trivers-Willard effect could have important implications for sex allocation. Apterous and seemingly nondispersing males routinely left their figs. Such dispersal can affect both (1) sex allocation by reducing the degree of local mate competition between brothers and (2) male dimorphism by reducing the mating opportunities of males with a dispersing morphology. We show that the wingless digitata males of the Otitesella digitata species group disperse on to leaves close to their natal fig. An extremely male-biased sex ratio resulted in almost all O. longicauda females being mated. These findings suggest that the classical concept of the fig wasp mating system is too simplistic and that important assumptions of sex allocation models are violated. Copyright 1999 The Association for the Study of Animal Behaviour.

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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.

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Female mating strategy and male brood cannibalism in a sand-dwelling cardinalfish.

I investigated male and female sexual strategies of a cardinalfish, Apogon niger, which breeds in sandy areas at sporadic coral colonies. Males mouthbrood an egg mass received from one female at a time. Because of the lengthy mouthbrooding period, the interspawning interval was shorter for females than for males. Females moved between coral colonies to find mates more extensively than did males, and more frequently deserted mates after spawning. The females shortened their interspawning intervals by changing mates, especially in the late breeding season, when their mobility was highest. Their mobility was positively correlated with their disappearance rate, suggesting that mate search increases mortality. This may reduce competition between females for males, resulting in an unbiased rather than female-biased operational sex ratio. Males, on the other hand, practised filial cannibalism of entire broods, which might allow them partially to compensate for the lack of food during the mouthbrooding period. The reproductive loss entailed by filial cannibalism could be effectively offset if males remate soon afterwards. However, cannibal males took a long time to remate because few females were available. Filial cannibalism was less frequent than in a boulder-dwelling congener in which males have easy access to mates. Low mate availability may inhibit male A. niger from committing filial cannibalism. Copyright 1999 The Association for the Study of Animal Behaviour.

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Affiliative preferences are stable and predict mate choices in both sexes of Japanese quail, Coturnix japonica.

Most laboratory studies of mate choice use affiliation as a surrogate measure for actual choice of a partner for copulation. In a series of recent studies using affiliation as a dependent variable, we showed that the tendency of both male and female Japanese quail to affiliate with members of the opposite sex is profoundly affected by seeing them mate with others: female quails show an increased tendency to affiliate with males that they have seen mating; and male quails show a decreased tendency to affiliate with females that they have seen mating. In the present experiments, we examined: (1) the consistency of affiliative preferences of male and female quail for individuals of both the same and opposite sex; and (2) the validity of measures of affiliation as predictors of choice of an individual with whom to mate. We found that: (1) both male and female quail display consistency from trial to trial in their affiliative preferences for members of the opposite but not the same sex, and (2) affiliative preference is a powerful predictor of choice of a partner for copulation in both male and female quail. In Japanese quail, affiliative behaviour is consistent over time and predicts mate choice. Copyright 1999 The Association for the Study of Animal Behaviour.

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Impact of geographical origin on mating behaviour in two species of Biomphalaria (Planorbidae: Gastropoda).

Studies of inbreeding and outcrossing have traditionally concentrated on matings within populations. The influence of geographical origin on mate choice in animals from different populations has received less attention. We investigated whether planorbid snails mated preferentially within their own population or with snails from other populations. Snails from three Biomphalaria pfeifferi strains and three B. glabrata strains were allowed to mate with conspecifics in the laboratory. We recorded their matings at night using time-lapse video. When they could choose between sympatric and allopatric snails, Biomphalaria snails significantly preferred the former: snails of each population mated more often with sympatric than with allopatric snails. This tendency to avoid outcrossing may indicate that, in some species, local adaptations can be more valuable than genetic novelties. Copyright 1999 The Association for the Study of Animal Behaviour.

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Colour bands, mate choice and paternity in the bluethroat.

Studies of several bird species have shown that coloured leg bands may affect a male's success in mate attraction and/or mating competition. From a colour band experiment in the field, we have previously reported that male bluethroats, Luscinia s. svecica, with blue and orange bands (BO males) guarded their mates less intensely at the peak of female fertility, and spent more time advertising for additional mates, than males banded with non-BO colours. These responses indicated that BO males experienced less threat to their paternity than did non-BO males, possibly mediated through an increased attractiveness. Here we present paternity analyses of the broods from the field study and test whether there were differences between the two male groups in within-pair or extrapair paternity. There were no significant differences between the two groups of males in paternity, suggesting effective male protection of paternity. However, extrapair paternity was infrequent in the 2 years of the field experiment; hence, the power in detecting effects on paternity does not allow a definitive conclusion on this issue. We also conducted an aviary experiment in which females were given the choice between a BO male and a non-BO male, to test whether females had preferences for particular colour bands. Females did not associate more with BO males, as would have been expected if these males were more attractive in social mate choice. Our results suggest that the effects of colour bands on social mate choice and paternity are, at best, weak. Copyright 2000 The Association for the Study of Animal Behaviour.

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Fluctuating asymmetry, size and mating success in males of Ischnura elegans (Vander Linden) (Odonata: Coenagrionidae).

Fluctuating asymmetry (FA) is thought to be an indicator of developmental stability and negatively related to male mating success in many animal taxa. We investigated the relationships between mating success of males, body size and FA for both wing length and number of setae on the legs in the damselfly Ischnura elegans. Males were classified as mated or unmated at the time of sampling. Fluctuating asymmetry, expressed as right-left differences, showed normal distributions without evidence of directional asymmetry or antisymmetry. Univariate analyses showed a significant negative correlation between size and mating success, and significant negative correlations between FA and mating success for both characters. On the other hand, with a multivariate analysis, new to studies on FA, the effect of body size was still significant but FA did not reach significance for either character. We conclude that the multivariate analysis should be used to assess the role of the different factors affecting mating success. Copyright 2000 The Association for the Study of Animal Behaviour.

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Female mate guarding: no evidence in a socially monogamous species.

To understand the behavioural aspects of sperm competition, the costs and benefits to both sexes should be considered. However, few studies have addressed the costs to females of their social mate engaging in extrapair copulations (EPCs). Measures of female mate guarding have concentrated on female solicitation and copulation; however, females may also control access to their mate by maintaining close proximity, as is common in males. I recorded the maintenance of pair proximity behaviour of an urban population of the socially monogamous European blackbird, Turdus merula, over three breeding seasons. There was no evidence that females guarded their mates to prevent them from engaging in EPCs, nor were there any effects of the potential quality indicators of age, body size or male bill colour on the intensity of mate guarding between individuals. The study adds to a small body of literature suggesting that female mate guarding may be found in (facultatively) polygynous species, but not in socially monogamous ones. Copyright 2000 The Association for the Study of Animal Behaviour.

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Alternative mechanisms of nonindependent mate choice.

Aspects of the environment, including the social environment, can contribute to intrapopulation variation in mating preferences. One example of the effect of social environment on mate preferences is mate choice copying; however, other types of socially influenced (nonindependent) choice might exist. We develop a list of such alternatives based on possible physiological or psychological mechanisms, evaluate the evidence distinguishing one from another and clarify some controversial aspects of mate choice copying. This framework reveals many ways in which one female's mate choice can influence that of another, and suggests a broader array of hypotheses about the selective forces acting on such mechanisms. Because nonindependent choice can occur in a variety of ways, it could be more important for understanding patterns of mate choice than current theory suggests. Copyright 2000 The Association for the Study of Animal Behaviour.

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