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Elevated testosterone reduces choosiness in female dark-eyed juncos (Junco hyemalis): evidence for a hormonal constraint on sexual selection?

Because testosterone (T) often mediates the expression of attractive displays and ornaments, in the absence of constraints sexual selection should lead to an evolutionary increase in male T levels. One candidate constraint would be a genetic correlation between the sexes that leads to a correlated response in females. If increased T in females were to have deleterious effects on mate choice, the effect of sexual selection on male T would be weakened. Using female dark-eyed juncos (Junco hyemalis), we tested whether experimentally enhancing female T would lead to a decrease in discrimination between two classes of males, one treated with T (T-males) and one control (C-males). The two female treatments (T-implanted and C-females) spent equal amounts of time with both classes of males, but T-treated females failed to show a preference for either male treatment, whereas C-females showed a significant preference, albeit in an unexpected direction (for C-males). T-females were less discriminating than C-females, irrespective of the direction of their preference. To our knowledge, this is the first study to show that circulating hormones can alter female choosiness without reducing sexual motivation. Our results suggest that hormonal correlations between the sexes have the potential to constrain sexual selection on males.

Analysis of Variance↗

Female sexual preferences differ in Mus spicilegus and Mus musculus domesticus: the role of familiarization and sexual experience.

Mating systems correspond to particular ecological conditions and result from proximate interactions between individuals. We compared the mating preferences of female mice of two species: the house mouse, Mus musculus domesticus, and the mound-builder mouse, Mus spicilegus. Because of differences in their habitat, we expected to observe differences in their sexual preferences. We studied female preferences for a familiar or an unfamiliar male and the occurrence of copulation with the unfamiliar male, during two states of female sexual activity: (1) the postpartum oestrus of paired females, to evaluate the stability of their sexual partnership; and (2) the oestrus of females familiarized with a male, to study the mechanisms underlying their sexual preferences. In the polygamous house mouse, postpartum oestrous females did not show a clear preference between their familiar male and the unfamiliar one. Moreover, oestrous females, familiarized with a male (without sexual interactions), preferred an unfamiliar male and copulated with him. In contrast, postpartum oestrous females and oestrous females of M. spicilegus preferred their familiar male and rarely copulated with the unfamiliar male. This study indicates a strong pair bond in established breeding pairs in M. spicilegus and shows that this bond can be established by familiarization, which is not the case in M. m. domesticus. Our study suggests the existence of monogamous traits in M. spicilegus in contrast to the polygamous M. m. domesticus. (c) 1998 The Association for the Study of Animal Behaviour.

Journal Article↗

Infusions of 5alpha-pregnan-3alpha-ol-20-one (3alpha,5alpha-THP) to the ventral tegmental area, but not the substantia nigra, enhance exploratory, anti-anxiety, social and sexual behaviours and concomitantly increase 3alpha,5alpha-THP concentrations in the hippocampus, diencephalon and cortex of ovariectomised oestrogen-primed rats.

Sequential actions of 17beta-oestradiol (E2) and progesterone (P4) in the ventromedial hypothalamus (VMH) and ventral tegmental area (VTA) mediate sexual behaviour of female rodents. In the presence of appropriate environmental stimuli, E2 and P4 can facilitate initiation of sex behaviour (i.e. lordosis), in part through classic actions at intracellular progestin receptors in the VMH. However, the effects of P4 in the VTA to modulate lordosis involve its metabolite, 5alpha-pregnan-3alpha-ol-20-one (3alpha,5alpha-THP), which can have paracrine effects in the brain to reduce anxiety and stress. We investigated the effects of 3alpha,5alpha-THP infusions to the VTA, and a control site, the substantia nigra (SN), on exploratory, anti-anxiety, social and sexual behaviours (socio-sexual behaviours) and hormone levels in the midbrain and other regions (hippocampus, diencephalon and cortex) that may mediate these functions. Ovariectomised, rats were E2-primed (10 microg, s.c.) at 0 h and were infused with beta-cyclodextrin vehicle or 3alpha,5alpha-THP to the VTA or SN 44-48 h later. Ten minutes after infusions, rats were tested in the open field, plus maze, partner preference, social interaction and paced mating tasks, or served as nontested controls. Infusions of 3alpha,5alpha-THP to the VTA, but not the SN, increased central entries in the open field, open arm time in the plus maze, time spent in proximity to a male, duration of social interaction, incidence and intensity of lordosis, pacing, proceptivity, and anti-conflict behaviour. 3Alpha,5alpha-THP, but not vehicle, infusions to the VTA (but not the SN) also increased 3alpha,5alpha-THP levels in the midbrain, as well as the hippocampus, diencephalon and cortex. Behavioural testing increased levels of the precursor of 3alpha,5alpha-THP precursor, dihydroprogesterone (DHP). Thus, infusions of 3alpha,5alpha-THP to the VTA enhance socio-sexual behaviours and increase 3alpha,5alpha-THP levels in the hippocampus, diencephalon and cortex, and behavioural testing increases DHP levels in brain areas involved in modulating socio-sexual behaviours.

Animals↗

Stress hormone is implicated in satellite-caller associations and sexual selection in the Great Plains toad.

The effects of androgens on male-typical traits suggest that variation among males in circulating levels can play a major role in sexual selection. We examined whether variation in vocal attractiveness is attributable to differences in androgen levels among Great Plains toads (Bufo cognatus). We found that noncalling "satellite" males practicing an alternative mating tactic were more likely to associate with males producing long calls. However, callers with satellites did not have higher androgen levels than callers without satellites. Rather, callers with satellites had significantly lower corticosterone (CORT) levels than callers without satellites. A CORT manipulation experiment suggested that differences in calls for males with and without satellites were related to differences in CORT levels. Furthermore, there was a negative correlation between CORT level and call duration within most nights of chorus activity. However, the correlation was weak for the pooled data (across all nights), suggesting that local environmental and/or social factors also affect call duration. Last, we show that females preferred broadcast calls of longer duration, characteristic of males with satellites and low CORT. These results imply that satellites optimize their reproductive success by associating with males producing long calls. However, this association should negatively affect the fitness of attractive callers.

Analysis of Variance↗

Mating propensities and variations in enzyme activities in long-term cage populations of Drosophila melanogaster.

Environment-dependent reproductive isolation was established between cage populations (Bs) of Drosophila melanogaster originated from a Greek natural population (summer 1973) and maintained for about five years under different diets (poor-rich). The detected deviation from random mating involved no homogametic or heterogametic preference but rather, a significantly increased activity of males from populations maintained on the rich food medium. This observation indicates that the male parental investment is not negligible and under certain conditions sexual isolation can be a function not only of female behavior but also of male behavior. Differences also were found in various enzyme activities on the inter- and intra-population levels. Given those observations as well as the observed different behavioral patterns of Bs and Cs-Ds populations 19, a preliminary attempt was made to associate adaptive evolution with differences in enzyme activities. The differences in enzyme activities between populations reared on different media are not due to allozymic differences. It also was shown that in some populations environmental effects do not always elicit differences in enzyme activity. It was concluded, therefore, that the observed variations were the result of environmental effects interacting with modifier genes.

Animals↗

Mate choice and imprinting in birds studied by cross-fostering in the wild.

Sexual-selection theories generally assume that mating preferences are heritable traits. However, there is substantial evidence that the rearing environment may be important for the development of mating preferences, indicating that they may be learnt, or modified by experience. The relative importance of such sexual imprinting across species remains largely unexplored. Here, we report results of a large-scale cross-fostering experiment in the wild in which nestling birds were raised by parents of a different species. We show that resulting sexual imprinting may have a negative effect on pairing success in one species (the great tit, Parus major), but not in two other species (the blue tit, P. caeruleus and the pied flycatcher, Ficedula hypoleuca). A remarkable variation thus seems to exist, even between species that are congeneric and have similar breeding ecologies. The cross-fostering resulted in heterospecific pairings between the two tit species (female blue tit breeding with male great tit), which has never, to our knowledge, been previously documented. However, the chicks fledging from these nests were all blue tit.

Animals↗

Evaluation of Dorset, Finnsheep, Romanov, Texel, and Montadale breeds of sheep: I. Effects of ram breed on productivity of ewes of two crossbred populations.

Effects of Dorset, Finnsheep, Romanov, Texel, and Montadale breeds for performance as sires were estimated in the initial phase of a comprehensive evaluation of these breeds as contributors to sheep crossbreeding systems. Objectives were to evaluate the effects of ram breed, ewe breed, season of mating, and two-way interactions. Rams from the five breeds were single-sire-mated with ewes from two breed types to produce lambs over a 3-yr period. Ewes were assigned to one of three distinct 35-d mating seasons initiated each year in August, October, and December. A different sample of six rams per breed was used each year across all three seasons, and each ram was penned with ewes of both breeds. Traits evaluated and number of ewe records were conception rate and litter weaning weight per ewe exposed (n = 3,261) and number born, litter birth weight, average birth weight, number weaned, and litter weaning weight per ewe lambing (n = 2,751). Ram breed and ewe breed interacted (P < .01) for conception rate and litter weaning weight per ewe exposed, implicating mating preferences, particularly of Romanov rams. In mixed groups of ewes exposed to Romanov rams, conception rate was 12.7% lower and litter weight weaned was 8.4 kg lower in the ewe breed presumably less preferred for mating by the rams. On a per ewe exposed basis, Romanov-sired litters produced either the largest or the smallest values for litter weaning weight, depending on the breed of ewe. Effects of ram breed on number born and litter birth weight interacted (P < .05) with season of mating. The largest litters within each ram breed were associated with the October mating season. Montadale and Romanov rams sired larger and heavier litters from August matings than from December matings, whereas the opposite was true for Dorset-sired litters. Texel- and Finnsheep-sired litters were similar in size and weight from August and December matings. Breed of ram differences affected per ewe lambing productivity measurements (P < .01). Differences between ram breeds for ewe productivity were noted, with increased number born and improved survival of crossbred progeny to weaning for Romanov-sired litters. These results may have implications for using these ram breeds as sires in different crossbreeding systems. Structured mating systems or the creation of new composite populations involving these breeds could be used to match the resources, environment, and market of specific production situations.

Animals↗

The competing countermarks hypothesis: reliable assessment of competitive ability by potential mates.

Scent marking on top of (overmarking), or in the vicinity of, a scent mark already present is commonly termed countermarking. Scent marks and countermarks provide a continuous record of competitive challenges between conspecifics, thus providing a reliable advertisement of an individual's ability to dominate or defend an area to other competitors and potential mates. To test the hypothesis that females should prefer males that countermark competing scent marks in their territory over those whose own marks are partially countermarked by a competing male, we manipulated scent marks in the territories of neighbouring male house mice (captive-bred Mus domesticus). As predicted, oestrous females were more strongly attracted to approach territory owners that countermarked the scent mark challenges of competitors than those that had been countermarked, and females themselves deposited more scent marks near the scents of these males. To investigate whether female mice use scent age, overlap or intrinsic qualitative or quantitative differences between scent marks and countermarks to make this discrimination, we redeposited male scent marks artificially as marks and partially overlapping countermarks, with or without a 24-h age difference between them. While the intrinsic quality or quantity of countermarks did not affect discrimination, an age difference between the original mark and subsequent countermark was important for consistent discrimination. The ultimate function of such competitive scent signalling thus may be to provide potential mates with a reliable indicator of the competitive ability of individuals advertising their high status. Since scent marks remain in the environment and are continuously available to challenge and investigation, they may provide a particularly effective and reliable means of dominance advertisement. Copyright 1999 The Association for the Study of Animal Behaviour.

Journal Article↗

Sexual experience and conditioned place preference in male rats.

We have previously shown that sexual behavior induces a reward state, as evaluated by conditioned place preference (CPP), only when males or females are able to control the rate of sexual interaction. In the present experiment, we evaluated if male rats that are repeatedly tested in a situation in which they are not able to control the sexual interaction eventually develop CPP. Three groups of sexually naïve male rats were used. One group never mated. A second group was tested once a week for 10 consecutive weeks in a chamber in which they controlled the rate of the sexual interaction. The third group was mated for the same number of weeks in a chamber in which the female, but not the male, controlled mating. The three groups were then tested for CPP. Only the group able to control the sexual interaction developed CPP. The group that had no control over the rate of the sexual interaction did not develop CPP even after 10 tests in which they consistently displayed sexual behavior. These results suggest that an estrous female and/or sexual behavior are powerful incentives that maintain mating even if the rewarding properties of the incentive are reduced.

Animals↗

Evolution of moth sex pheromones via ancestral genes.

Mate finding in most moth species involves long-distance signaling via female-emitted sex pheromones. There is a great diversity of pheromone structures used throughout the Lepidoptera, even among closely related species. The conundrum is how signal divergence has occurred. With strong normalizing selection pressure on blend composition and response preferences, it is improbable that shifts to pheromones of diverse structures occur through adaptive changes in small steps. Here, we present data supporting the hypothesis that a major shift in the pheromone of an Ostrinia species occurred by activation of a nonfunctional desaturase gene transcript present in the pheromone gland. We also demonstrate the existence of rare males that respond to the new pheromone blend. Their presence would allow for asymmetric tracking of male response to the new blend and, thus, evolution of an Ostrinia species with structurally different sex pheromone components.

Amino Acid Sequence↗

Effects of a kaolin-based particle film on obliquebanded leafroller (Lepidoptera: Tortricidae).

Studies were conducted in 1997 to evaluate the effects of the kaolin-based particle film formulation M96-018 on adults, eggs, and larvae of the obliquebanded leafroller, Choristoneura rosaceana (Harris). Particle film treatments significantly reduced female longevity, mating success, and number of egg masses oviposited compared with moths on untreated apple leaves in sleeve-cage and screen-cage tests. No differences in mating success or oviposition were caused by the application rates and coverage density of M96-018 on foliage. Females avoided ovipositing on particle film-treated leaves in choice tests. Larval hatch was not affected by topical application or residual exposure to M96-018. Larval weight gain and pupal weight were significantly reduced and larval mortality increased in no-choice feeding tests with M96-018. In choice tests, larvae preferred to feed on untreated leaf surfaces. The negative effects on larval development and survivorship on M96-018-treated foliage did not differ across a fourfold difference in spray application rate. A significant reduction in the number of infested shoots was found in orchard trials when M96-018 was applied before bud break in late March compared with untreated trees. No reductions in larval densities were found compared with an untreated control following prebloom and postbloom applications.

Animals↗

Mate location mechanism and phase-related mate preferences in solitarius desert locust, Schistocerca gregaria.

Mate location responses of male and female solitary-reared locusts that had either experienced no crowding or that had been crowded for varying periods were studied in a flatbed wind tunnel. Two hypotheses were explored: that both sexes of this phase of the locust participate in locating the other by using a combination of chemical and visual signals, and that individuals that experience some crowding (i.e., undergo varying levels of phase shift) can compete effectively with their solitary counterparts in mate location and mating. Our results confirm that both male and female solitarious locusts actively participate in mate location, although the former is the more aggressive partner. The responses of the insects are stronger when a visual cue is provided with the olfactory signal. Crowding of solitary-reared adults enhances their responsiveness to the other sex in the absence and presence of the visual cue. This phenomenon may constitute one of several mechanisms that are involved in recruiting solitary individuals into gregarizing groups and facilitating the spread of gregarious characters across a reproductively active solitarious population.

Animals↗

No apparent indirect genetic benefits to female red flour beetles preferring males with intense copulatory courtship.

Whether females gain indirect genetic benefits through mate choice is a controversial issue since this requires additive genetic variance in the preferred male traits. Condition dependence could maintain the necessary genetic variance by linking the expression of male traits to the supposedly large number of genes affecting condition. Copulating males of the red flour beetle Tribolium castaneum rub their legs along the female elytra. Females favor males with vigorous rubbing through cryptic female choice. We measured the repeatability and heritability of this behavior and assessed its potential use as indicator of viability and condition. We found genetic variance in larval to adult survival and in the rate of leg rubbing in males. However, the rate of leg rubbing was not related to offspring survival or condition dependent. The genetic variance in leg rubbing was mostly non-additive with very low narrow sense heritability. Therefore, we failed to document any indirect genetic benefits to choosy females through viability of their offspring or attractiveness of their sons.

Animals↗

The effect of Wolbachia versus genetic incompatibilities on reinforcement and speciation.

Wolbachia is a widespread group of intracellular bacteria commonly found in arthropods. In many insect species, Wolbachia induce a cytoplasmic mating incompatibility (CI). If different Wolbachia infections occur in the same host species, bidirectional CI is often induced. Bidirectional CI acts as a postzygotic isolation mechanism if parapatric host populations are infected with different Wolbachia strains. Therefore, it has been suggested that Wolbachia could promote speciation in their hosts. In this article we investigate theoretically whether Wolbachia-induced bidirectional CI selects for premating isolation and therefore reinforces genetic divergence between parapatric host populations. To achieve this we combined models for Wolbachia dynamics with a well-studied reinforcement model. This new model allows us to compare the effect of bidirectional CI on the evolution of female mating preferences with a situation in which postzygotic isolation is caused by nuclear genetic incompatibilities (NI). We distinguish between nuclear incompatibilities caused by two loci with epistatic interactions, and a single locus with incompatibility among heterozygotes in the diploid phase. Our main findings are: (1) bidirectional CI and single locus NI select for premating isolation with a higher speed and for a wider parameter range than epistatic NI; (2) under certain parameter values, runaway sexual selection leads to the increase of an introduced female preference allele and fixation of its preferred male trait allele in both populations, whereas under others it leads to divergence in the two populations in preference and trait alleles; and (3) bidirectional CI and single locus NI can stably persist up to migration rates that are two times higher than seen for epistatic NI. The latter finding is important because the speed with which mutants at the preference locus spread increases exponentially with the migration rate. In summary, our results show that bidirectional CI selects for rapid premating isolation and so generally support the view that Wolbachia can promote speciation in their hosts.

Animals↗

Why do multiple traits determine mating success? Differential use in female choice and male competition in a water boatman.

Mating success is often determined by multiple traits, but why this occurs is largely unknown. Much attention has been paid to female preferences for multiple traits, but surprisingly few researchers have addressed the possibility that multiple traits are important because they serve different functions in female choice and male-male competition. Differential trait function could result from a conflict of interest between the sexes or from constraints forcing the sexes to pay attention to different traits. I show that traits determined at distinct life-history stages differ in their importance in female choice and male-male competition in a water boatman Sigara falleni. Juvenile conditions determined body and foreleg pala size and were the main determinants of mating success under female choice, whereas adult conditions determined body mass and influenced mating success when male competition was included. This differential use of condition-dependent traits under the two selection regimes appeared to arise partly from a conflict between the sexes, since the two selection forces (female choice and male competition) conflict for selection on pala size, and partly from constraints, as females appeared unable to assess adult condition.

Age Factors↗

Sexual behaviour of rams: male orientation and its endocrine correlates.

The components of heterosexual behaviour in rams are reviewed as a basis for understanding partner preference behaviour. A small percentage of rams will not mate with oestrous females and if given a choice will display courtship behaviour towards another ram in preference to a female. Some of the endocrine profiles of these male-oriented rams differ from those of heterosexual controls. These differences include reduced serum concentrations of testosterone, oestradiol and oestrone, reduced capacity to produce testosterone in vitro, and reduced capacity to aromatize androgens in the preoptic-anterior hypothalamus of the brain. Our observation that aromatase activity is significantly lower in the preoptic-anterior hypothalamic area of male-oriented rams than in female-oriented rams may indicate an important neurochemical link to sexual behaviour that should be investigated. The defect in steroid hormone production by the adult testes of the male-oriented ram may represent a defect that can be traced to the fetal testes. If this contention is correct, partner preference behaviour of rams may also be traceable to fetal development and represent a phenomenon of sexual differentiation.

Animals↗

Chronic prenatal exposure to carbamazepine and perinatal outcomes of C3H/He mice.

OBJECTIVES: This study was undertaken to determine a daily therapeutic dose of carbamazepine and to measure its effect on reproductive performance and perinatal outcomes of mice. STUDY DESIGN: Adult C3H/He mice were given carbamazepine in rodent chow in either a 0.25% or a 1.0% mixture. Comparisons between doses included nongravid weight change, plasma drug steady-state concentrations, and response to a maximal electroshock seizure test. The strain was then fed either the preferred dose of carbamazepine or a placebo 1 week before starting to mate and throughout gestation to compare reproductive performance and offspring early development. RESULTS: Mice who ate the 0.25% carbamazepine mixture displayed no evoked seizure activity and, in contrast to the 1.0% mixture, did not lose weight. This daily dose of 542+/-35 mg/kg produced a trough steady-state plasma concentration that was consistent with a protective threshold in humans. Differences from placebo controls were not statistically significant for the number of cycles necessary to conceive or for the duration of gestation. The litter size, survival rates, birth weights, weight gain, and onset of eye openings and teeth eruptions of the pups were not statistically significant between the two groups. CONCLUSION: Long-term prenatal exposure to a subtoxic yet therapeutic dose of carbamazepine did not impair reproductive performance or early growth and development of exposed mice offspring.

Animals↗

Intraspecific sexual selection on a speciation trait, male coloration, in the Lake Victoria cichlid Pundamilia nyererei.

The haplochromine cichlids of Lake Victoria constitute a classical example of explosive speciation. Extensive intra- and interspecific variation in male nuptial coloration and female mating preferences, in the absence of postzygotic isolation between species, has inspired the hypothesis that sexual selection has been a driving force in the origin of this species flock. This hypothesis rests on the premise that the phenotypic traits that underlie behavioural reproductive isolation between sister species diverged under sexual selection within a species. We test this premise in a Lake Victoria cichlid, by using laboratory experiments and field observations. We report that a male colour trait, which has previously been shown to be important for behavioural reproductive isolation between this species and a close relative, is under directional sexual selection by female mate choice within this species. This is consistent with the hypothesis that female choice has driven the divergence in male coloration between the two species. We also find that male territoriality is vital for male reproductive success and that multiple mating by females is common.

Animals↗