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Sexual equality in zebra finch song preference: evidence for a dissociation between song recognition and production learning.

Song in oscine birds is a culturally inherited mating signal and sexually dimorphic. From differences in song production learning, sex differences in song recognition learning have been inferred but rarely put to a stringent test. In zebra finches, Taeniopygia guttata, females never sing and the species has one of the greatest neuroanatomical differences in song-related brain nuclei reported for songbirds. Preference tests with sibling groups for which exposure to song had been identical during the sensitive phase for song learning in males, revealed equally strong influence of the tutor's song (here the father) on males' and females' adult song preferences. Both sexes significantly preferred the father's over unfamiliar song when having free control over exposure to playbacks via an operant task. The sibling comparisons suggest that this preference developed independently of the song's absolute quality: variation between siblings was as great as between nests. The results show that early exposure has an equally strong influence on males' and females' song preferences despite the sexual asymmetry in song production learning. This suggests that the trajectory for song recognition learning is independent of the one for song production learning.

Acoustic Stimulation↗

Kin preference in infant Macaca nemestrina.

The ability to recoginize conspecifics is a prerequisite for many types of social behaviour, including, for example, parent-offspring relation, mate selection and recognition, territorial defence and dominance coalitions. This ability is of special importance to Hamilton's kin selection hypothesis, which predicts that an individual's behaviour towards a conspecific will depend on the degree of genetic relatedness between them. Although recognition depends on previous experience between individuals in some species, this does not precluded the possibility that recognition could occur in its absence. For example, juveniles who disperse before nonlittermate siblings are born or adult males who do not participate in rearing their young might benefit from recognition abilities that are independent of prior association between the individuals. Here we show that young pigtail macaques prefer to interact with a related over an unrelated monkey in a laboratory test. Because subjects were separated from their dams at birth and reared apart from all other relatives, results suggest that kin recognition can occur in the absence of prior association with relatives.

Age Factors↗

Mutual mate choice in sticklebacks: attractive males choose big females, which lay big eggs.

Brighter red three-spined stickleback, Gasterosteus aculeatus, males have been shown to be preferred by females in the laboratory but in the field, these males did not receive more eggs. Instead, they had heavier eggs in their brood. We investigated the hypothesis that sexual selection for red coloration in male sticklebacks acts through mate choice by preferred males, who can afford to be choosy, for high-quality females which lay heavier eggs. We assume here that heavier eggs provide a direct fitness advantage. In simultaneous choice tests males were presented with two females differing in size. The number of zigzags directed to and the time spent orienting to each female were measured. After the test the females laid eggs, which we counted and weighed. Bigger (i.e. longer and heavier) females laid significantly more and heavier eggs than smaller females. For all 23 males pooled together, the preferred female was the bigger of the two in 17 cases, laid more eggs in 18 cases, but laid heavier eggs in only 13 cases. When bright and dull males were analysed separately, we found that bright but not dull males spent more time oriented to the bigger female, and to the female that laid more eggs. Females preferred by bright males tended to lay heavier eggs than nonpreferred females, although this result was not quite significant. We conclude that in nature this preference for bigger females results in brighter males receiving on average heavier eggs. Assuming higher survival of bigger offspring, we propose that this can explain how brightness can be sexually selected in spite of brighter males not receiving more eggs. Copyright 1998 The Association for the Study of Animal Behaviour.

Journal Article↗

Effects of developmental environment on signal-preference coupling in a Hawaiian cricket.

Previous work has suggested that developmental temperature influences expression of the adult male calling song of the cricket, a sexually selected mate recognition signal. The role of developmental temperature in shaping female preference functions, and thus its influence on signal-preference coupling has not been investigated. In this study, the effects of developmental temperature are examined in both males and females of the Hawaiian cricket, Laupala cerasina, to determine the degree of signal-preference matching between male song and female preference due to developmental environment. We found that rearing females in different temperature environments affected adult female acoustic preference functions in the same direction as male calling song, thereby influencing variation in adult reproductive behavior in such a way that male and female components remain coordinated. However, we further demonstrate that for male song, the effect of the rearing environment is not permanent but appears only to exert influence for a period of days. This mid-term temperature effect is distinct from the effect of short-term ambient temperature, which influences song in a matter of minutes and has been well documented. Signal-preference coordination, and sexual selection pressures due to mismatching within natural populations, likely will be influenced by nymphal developmental environments of males and females, as well as by adult singing and preference environments.

Animal Communication↗

DNA fingerprinting reveals female preference for male parental care in Savannah Sparrows.

According to sexual selection theory, females choose mates to ensure access to high quality resources, male parental care, or good genes. This last hypothesis has been hotly debated on both theoretical and empirical grounds. In contrast, female preference for male parental care has received less attention, primarily because the potential benefits of paternal effort seem obvious. The fitness relations are less clear in double-brood species, however, because females can base mating decisions on their prior experience with male parental care. Here, the extent of male parental care delivered to first-brood offspring may indicate male genetic quality and/or be the target of female manipulation via her subsequent mating fidelity. In Savannah Sparrows (Passerculus sandwichensis), multi-locus DNA fingerprinting of 203 adults and young revealed substantial female infidelity in first and second broods: overall, 24 of 80 first-brood young (30%) and 13 of 80 second-brood young (16.3%) were the product of extra-pair fertilizations. Among 12 females altering fidelity between broods, absolute male feeding rate to first-brood offspring was a strong, positive predictor of change in female fidelity. Because the extent of male parental care reflects a male's viability, the data support a 'good-genes' interpretation.

Analysis of Variance↗

Production objectives and management strategies of livestock keepers in south-east Kenya: implications for a breeding programme.

A survey of pastoralist and agropastoralist households in south-east Kenya was conducted to determine their production objectives and management strategies in order to optimize and extend a breeding programme for indigenous small East African Shorthorn Zebu cattle. The reasons for keeping cattle and the breed/trait preferences identified reflect the multiple objectives of the livestock keepers, with both adaptive traits and productive/reproductive traits rated as important. Although the Maasai and Kamba zebu (M&KZ) breeds were ranked highly with regard to adaptive traits, the population is considered to have been in decline over recent years. In order to promote the conservation and sustainable use of the M&KZ cattle, the formation of an open nucleus breeding scheme is recommended. In particular, such a scheme would be able to address several existing constraints (e.g. individual herds are very small and communal use of pastures/water makes controlled mating difficult). Such interventions would require the full participation of the livestock keepers, as well as ensuring that a holistic approach to species and breed attributes is taken into account in setting breeding goals, such that the full array of contributions that livestock make to livelihoods and the genetic characteristics related to these contributions are fully incorporated into the programme.

Adult↗

Germ-line transmission of a disrupted beta 2-microglobulin gene produced by homologous recombination in embryonic stem cells.

Major histocompatibility complex (MHC) class I molecules are integral membrane proteins present on virtually all vertebrate cells and consist of a heterodimer between the highly polymorphic alpha-chain and the beta 2-microglobulin (beta 2-m) protein of relative molecular mass 12,000 (ref. 1). These cell-surface molecules play a pivotal part in the recognition of antigens, the cytotoxic response of T cells, and the induction of self tolerance. It is possible, however, that the function of MHC class I molecules is not restricted to the immune system, but extends to a wide variety of biological reactions including cell-cell interactions. For example, MHC class I molecules seem to be associated with various cell-surface proteins, including the receptors for insulin, epidermal growth factor, luteinizing hormone and the beta-adrenergic receptor. In mice, class I molecules are secreted in the urine and act as highly specific olfactory cues which influence mating preference. The beta 2-m protein has also been identified as the smaller component of the Fc receptor in neonatal intestinal cells, and it has been suggested that the protein induces collagenase in fibroblasts. Cells lacking beta 2-m are deficient in the expression of MHC class I molecules, indicating that the association with beta 2-m is crucial for the transport of MHC class I molecules to the cell surface. The most direct means of unravelling the many biological functions of beta 2-m is to create a mutant mouse with a defective beta 2-m gene. We have now used the technique of homologous recombination to disrupt the beta 2-m gene. We report here that introduction of a targeting vector into embryonic stem cells resulted in beta 2-m gene disruption with high frequency. Chimaeric mice derived from blastocysts injected with mutant embryonic stem cell clones transmit the mutant allele to their offspring.

Animals↗

Response of female Cydia molesta (Lepidoptera: Tortricidae) to plant derived volatiles.

Peach shoot volatiles were attractive to mated female oriental fruit moth, Cydia molesta (Busck), in a dual choice arena. No preference was observed between leaf odours from the principle host plant, peach, and the secondary host plant, apple. Twenty-two compounds were identified in headspace volatiles of peach shoots using gas chromatography-mass spectrometry. Green leaf volatiles accounted for more than 50% of the total emitted volatiles. A bioassay-assisted fractionation using different sorbent polymers indicated an attractant effect of compounds with a chain length of 6-8 carbon atoms. The major compounds of this fraction were tested either singly or in combinations for behavioural response of females. Significant bioactivity was found for a three-component mixture of (Z)-3-hexen-1-yl acetate, (Z)-3-hexen-1-ol and benzaldehyde in a 4:1:1 ratio. This synthetic mixture elicited a similar attractant effect as the full natural blend from peach shoots as well as the bioactive fraction.

Animals↗

Strategic mating with common preferences.

We present a two-sided search model in which individuals from two groups (males and females, employers and workers) would like to form a long-term relationship with a highly ranked individual of the other group, but are limited to individuals who they randomly encounter and to those who also accept them. This article extends the research program, begun in Alpern and Reyniers [1999. J. Theor. Biol. 198, 71-88], of providing a game theoretic analysis for the Kalick-Hamilton [1986. J. Personality Soc. Psychol. 51, 673-682] mating model in which a cohort of males and females of various 'fitness' or 'attractiveness' levels are randomly paired in successive periods and mate if they accept each other. Their model compared two acceptance rules chosen to represent homotypic (similarity) preferences and common (or 'type') preferences. Our earlier paper modeled the first kind by assuming that if a level x male mates with a level y female, both get utility -|x-y|, whereas this paper models the second kind by giving the male utility y and the female utility x. Our model can also be seen as a continuous generalization of the discrete fitness-level game of Johnstone [1997. Behav. Ecol. Sociobiol. 40, 51-59]. We establish the existence of equilibrium strategy pairs, give examples of multiple equilibria, and conditions guaranteeing uniqueness. In all equilibria individuals become less choosy over time, with high fitness individuals pairing off with each other first, leaving the rest to pair off later. This route to assortative mating was suggested by Parker [1983. Mate Choice, Cambridge University Press, Cambridge, pp. 141-164]. If the initial fitness distributions have atoms, then mixed strategy equilibria may also occur. If these distributions are unknown, there are equilibria in which only individuals in the same fitness band are mated, as in the steady-state model of MacNamara and Collins [1990. J. Appl. Prob. 28, 815-827] for the job search problem.

Animals↗

Negative genetic correlation between male sexual attractiveness and survival.

Indirect selection of female mating preferences may result from a genetic association between male attractiveness and offspring fitness. The offspring of attractive males may have enhanced growth, fecundity, viability or attractiveness. However, the extent to which attractive males bear genes that reduce other fitness components has remained unexplored. Here I show that sexual attractiveness in male guppies (Poecilia reticulata) is heritable and genetically correlated with ornamentation. Like ornamentation, attractiveness may be substantially Y-linked. The benefit of mating with attractive males, and thus having attractive sons, is opposed by strong negative genetic correlation between attractiveness and both offspring survival and the number of sons maturing. Such correlations suggest either antagonistic pleiotropy between attractiveness and survival or linkage disequilibrium between attractive and deleterious alleles. The presence of many colour pattern genes on or near the non-recombining section of the Y chromosome may facilitate the accumulation of deleterious mutations by genetic hitchhiking. These findings show that genes enhancing sexual attractiveness may be associated with pleiotropic costs or heavy mutational loads.

Animals↗

Pair bonding and "the widow effect" in female prairie voles.

We conducted field and laboratory experiments with the well-studied monogamous prairie vole, Microtus ochrogaster, to distinguish among three hypotheses for the failure of females that lose their mates to bond with a new male ("the widow effect"). The reproductive value hypothesis predicts that males prefer young to older females because they potentially have a longer reproductive lifespan. The mate rejection hypothesis predicts that females will prevent repairing by aggressively deterring males that might harm their current offspring. The misdirected paternal care hypothesis assumes that females will mate during postpartum estrus and thus will be pregnant and/or nursing young throughout the breeding season; males will avoid pairing with older females to avoid providing care to unrelated offspring and/or because of a delay to the next breeding opportunity. Males associated and bred more with older than young females, allowing us to reject the reproductive value hypothesis. Our results were consistent with the male rejection hypothesis in that females were aggressive toward unfamiliar males. Our results were most consistent with the misdirected paternal care hypothesis in that once females started breeding, they continued to become pregnant and nurse young throughout the study period. Thus, our findings suggest that the potential of misdirected paternal care and delayed mating opportunity in conjunction with the aggressive behavior of females toward unfamiliar males are likely explanations for the lack of repairing for widow females.

Animals↗

Prenatal blockade of androgen receptors reduces the number of intromissions needed to induce conditioned place preference after paced mating in female rats.

Estrous female rats pace their coital contacts to control the rate of cervical/vaginal stimulation. Females that receive at least 10 paced intromissions develop a reward state, evaluated by conditioned place preference (CPP), whereas females that do not pace their coital contacts do not develop CPP. We asked if the blockade of androgen receptors could modify the number of intromissions needed during paced mating to develop CPP. Pregnant females received daily injections of the androgen receptor antagonist flutamide from day 12 of pregnancy until pups were born. When adults, females exposed prenatally to flutamide or vehicle were ovariectomized and hormonally primed to evaluate if paced and non-paced mating induced CPP. The prenatal flutamide treatment did not affect the capacity of females to develop CPP to a systemic morphine injection. Flutamide-treated females that paced their sexual contacts developed CPP with fewer intromissions than control females. No effect on conditioning was observed when females were not allowed to pace their sexual contacts. The results suggest the existence of a threshold of cervical/vaginal stimulation that correlates with the induction of a reward state and that this threshold can be reduced by prenatal blockade of androgen receptors.

Androgen Antagonists↗

Delayed selfing in relation to the availability of a mating partner in the cestode Schistocephalus solidus.

A hermaphroditic individual that prefers to outbreed but that has the potential of selfing faces a dilemma: in the absence of a partner, should it wait for one to arrive or should it produce offspring by selfing? Recent theory on this question suggests that the evolutionary solution is to find an optimal delay of reproduction that balances the potential benefit of outcrossing and the cost of delaying the onset of reproduction. Assuming that resources retained from breeding can be reallocated to future reproduction, isolated individuals, compared with individuals with available mates, are predicted to delay their age at first reproduction to wait for future outcrossing. Here, I present empirical support for this idea with experimental data from the hermaphroditic cestode Schistocephalus solidus. I show that individuals breeding alone delay their reproduction and initially produce their eggs at a slow rate relative to cestodes breeding in pairs. This delay is partly compensated for by a later higher egg production, although singly breeding cestodes still pay a cost of overall lower egg production.

Animals↗

Comparison of three methods for age-grading of female neotropical phlebotomine sandflies.

Changes in appearance of the accessory glands (AG), ovaries (OV) and the absence or presence of crease marks at the edges of the genital atrium armature (GA) were compared as methods to age-grade female laboratory-bred (Lutzomyia migonei) and wild-caught (Lu.youngi, Lu.spinicrassa, Lu.ovallesi) Neotropical sandflies. Dissections at different physiological stages revealed that all parous and high proportions of nulliparous (4/38) unmated, 12/12 mated) females showed similar residual AG secretions. Hence AG appearance (nullipars with AG translucent, versus yellowish opaque granulated AG previously regarded as signs of parousness) is unreliable for age-grading. Readily recognizable changes in both GA (98% creased) and the OV (100% yellowish with loose ovarioles) indicated that these organs could be employed to discriminate between nulliparous and parous females. Despite finding one misdiagnosed female (a nullipar having creased GA indicative of parity), the GA method of age-grading is recommended because females may be examined satisfactorily at any stage of the gonotrophic cycle. Moreover, the GA characteristics of pickled or dry preserved sandflies as well as fresh slide-mounted specimens can be determined easily. Among sixty-five wild-caught Lutzomyia females dissected (mostly Lu.spinicrassa and Lu.youngi), the proportions of specimens yielding preparations suitable for examination were 97% for GA, 89% for OV, but only 58% for AG, which reinforces GA as the preferred character for age-grading.

Aging↗

An invasion of cheats; the evolution of worthless nuptial gifts.

Nuptial gifts are food items or inedible tokens that are transferred to females during courtship or copulation . Tokens are of no direct value to females, and it is unknown why females require such worthless gifts as a precondition of mating. One hypothesis is that token giving arose in species that gave nutritious gifts and males exploited female preferences for nutritional gifts by substituting more easily obtainable but worthless items. An invasion of such behavior would require that females accept the substitute gift and copulate for a period of time similar to that with genuine gifts. We show that both these prerequisites are met in the dance fly Rhamphomyia sulcata, in which females normally accept a nutritious gift. We removed the gift from copulating pairs and replaced it with either a large or small prey item or inedible token. We found that although pairs copulated longest with a large genuine gift, the tokens resulted in copula durations equivalent to those with a small genuine gift. We also observed that males that returned to the lek with tokens re-paired successfully. These findings suggest that female behavior in genuine gift-giving species is susceptible to the invasion of male cheating on reproductive investment.

Animals↗

Female meadow voles have a preferred mating pattern predicted by photoperiod, which influences fertility.

In a previous study, Meek and Lee found that female meadow voles mated within three distinct time periods after pairing with a male, and fertility was influenced by the time of mating and photoperiod. In the present studies, we tested the hypothesis that different patterns of mating would correlate with high and low fertility. We found two very different patterns of mating were correlated with high fertility. Females housed in long daylengths (long day; 14 h light/day) and mating within 48 h of pairing with a male, received few mounts, intromissions, and intromissions per ejaculatory series, with very short interintromission intervals. All long-day females utilizing this mating pattern ovulated, and 86-100% produced litters. We suggest this pattern of mating is associated with spontaneous estrus. In contrast to long-day females mating within 48 h, females housed in short daylengths (short day; 10 h light/day) and mating between 14-48 h after pairing with a male, received significantly more short mounts and intromissions, with longer interintromission intervals. These short day females all ovulated and 86% produced litters. We suggest that this pattern of mating is associated with an induced estrus. Short-day females that did not exhibit this pattern of mating (66%) produced far fewer litters. We discuss two mechanisms by which fertility may be inhibited in most short-day females.

Animals↗

Naloxone inhibits mating and conditioned place preference for an estrous female in male rats soon after castration.

Three experiments were conducted to assess the role of endogenous opioids in controlling mating behavior and sexual reward in the male rat. In Experiment 1 SC administration of naloxone (0.5, 1.0, 5.0, or 10.0 mg/kg) significantly reduced mounting and ejaculation in male rats tested 14, but not 7 days, after castration. In Experiment 2 naloxone (5.0 mg/kg) administered SC to gonadally intact males, which had ejaculated repeatedly with one female until they were sexually sated, significantly inhibited the resumption of mating after the reintroduction of a female partner. One interpretation of these results is that naloxone attenuated the reward experienced by castrated and sexually sated males in the presence of an estrous female, thereby disrupting males' coital performance. This hypothesis was tested in Experiment 3 using a conditioned place preference paradigm in which males copulated with an estrous female in an initially "non-preferred" (white) compartment, whereas on alternate days they remained alone in an initially "preferred" (black) compartment. After 10 such conditioning sessions, males were either castrated or sham-operated. They later were given free access to both compartments in the absence of an estrous female. Seven days after conditioning and surgery, sham-operated, naloxone-injected males and both groups of castrates spent significantly less time than sham-operated, saline-injected controls in the initially "non-preferred" compartment. Fourteen days after conditioning and surgery castrated, naloxone-treated males spent significantly less time in the "non-preferred" compartment than males in the other three groups. Endogenous opioids may play an important role in the interpretation by males of the incentive motivational stimuli which emanate from an estrous female.

Animals↗

The role of juvenile hormone in immune function and pheromone production trade-offs: a test of the immunocompetence handicap principle.

The immunocompetence handicap hypothesis postulates that secondary sexual traits are honest signals of mate quality because the hormones (e.g. testosterone) needed to develop secondary sexual traits have immunosuppressive effects. The best support for predictions arising from the immunocompetence handicap hypothesis so far comes from studies of insects, although they lack male-specific hormones such as testosterone. In our previous studies, we found that female mealworm beetles prefer pheromones of immunocompetent males. Here, we tested how juvenile hormone (JH) affects male investment in secondary sexual characteristics and immune functions in the mealworm beetle, Tenebrio molitor. We injected male mealworm beetles with JH (type III) and found that injection increased the attractiveness of male pheromones but simultaneously suppressed immune functions (phenoloxidase activity and encapsulation). Our results suggest that JH, which is involved in the control of reproduction and morphogenesis, also plays a central role in the regulation of a trade-off between the immune system and sexual advertisement in insects. Thus, the results reflect a general mechanism by which the immunocompetence handicap hypothesis may work in insects.

Animals↗