A supergene determines highly divergent male reproductive morphs in the ruff

Three strikingly different alternative male mating morphs (aggressive 'independents', semicooperative 'satellites' and female-mimic 'faeders') coexist as a balanced polymorphism in the ruff, Philomachus pugnax, a lek-breeding wading bird1, 2, 3. Major differences in body size, ornamentation, and aggressive and mating behaviors are inherited as an autosomal polymorphism4, 5. We show that development into satellites and faeders is determined by a supergene6, 7, 8 consisting of divergent alternative, dominant and non-recombining haplotypes of an inversion on chromosome 11, which contains 125 predicted genes. Independents are homozygous for the ancestral sequence. One breakpoint of the inversion disrupts the essential CENP-N gene (encoding centromere protein N), and pedigree analysis confirms the lethality of homozygosity for the inversion. We describe new differences in behavior, testis size and steroid metabolism among morphs and identify polymorphic genes within the inversion that are likely to contribute to the differences among morphs in reproductive traits

Effects of prenatal environment on phenotype are revealed by postnatal challenges : Embryonic hormone exposure, adrenocortical function, and food in seabird chicks

The interaction between prenatal environments and postnatal environments is an important source of phenotypic variability. We examined the ability of prenatal steroid exposure and postnatal energy restriction to explain adrenocortical function and fledging age in captive seabird chicks. We proposed and tested two hypotheses: (1) the strength of prenatal effects is attenuated by challenging postnatal environments (postnatal override) and (2) the strength of prenatal effects increases with the severity of postnatal challenges (postnatal reveal).We reared commonmurre (Uria aalge) chicks and measured prenatal exposure to corticosterone (CORT) and testosterone (T) from allantoic waste. Adrenocortical function was assessed after 10 d of ad lib. Feeding and then after 5 and 10 d on controlled diets. Postnatal override predicts that prenatal steroids will explain more phenotypic variation before implementation of energy restriction; postnatal reveal predicts that the contribution of prenatal steroids will increase with duration and severity of energy restriction. Energy restriction increased secretion of baseline CORT and the adrenocortical response to the standardized stressor of handling and restraint. The ability of prenatal steroids to explain baseline CORT increased with duration of energy restriction, and for day 20 free baseline CORT, there was a significant interaction between kilojoules per day and prenatal CORT levels; severity of restriction strengthened the relationship between prenatal hormone levels and postnatal hormone levels. Both maximum CORT at day 20 and fledging age were best explained by diet treatment and day 15 or day 20 baseline CORT, respectively. Overall, prenatal CORT increased fledging age and baseline secretion of CORT, while prenatal T decreased them. However, prenatal effects on adrenocortical function were apparent only under the energy restriction conditions. Thus, we found some support for the postnatal reveal hypothesis; our results suggest that some prenatal effects on phenotype may be more likely to manifest in challenging postnatal environments

Differential reproductive responses to stress reveal the role of life-history strategies within a species

Life-history strategies describe that 'slow'- in contrast to 'fast'-living species allocate resources cautiously towards reproduction to enhance survival. Recent evidence suggests that variation in strategies exists not only among species but also among populations of the same species. Here, we examined the effect of experimentally induced stress on resource allocation of breeding seabirds in two populations with contrasting life-history strategies: Slowliving Pacific and fast-living Atlantic black-legged kittiwakes. We tested the hypothesis that reproductive responses in kittiwakes under stress reflect their life-history strategies. We predicted that in response to stress, Pacific kittiwakes reduce investment in reproduction compared with Atlantic kittiwakes. We exposed chick-rearing kittiwakes to a short-term (3-day) period of increased exogenous corticosterone (CORT), a hormone that is released during food shortages. We examined changes in baseline CORT levels, parental care and effects on offspring. We found that kittiwakes from the two populations invested differently in offspring when facing stress. In response to elevated CORT, Pacific kittiwakes reduced nest attendance and deserted offspring more readily than Atlantic kittiwakes. We observed lower chick growth, a higher stress response in offspring and lower reproductive success in response to CORT implantation in Pacific kittiwakes, whereas the opposite occurred in the Atlantic. Our findings support the hypothesis that lifehistory strategies predict short-term responses of individuals to stress within a species. We conclude that behaviour and physiology under stress are consistent with trade-off priorities as predicted by life-history theory. We encourage future studies to consider the pivotal role of life-history strategies when interpreting inter-population differences of animal responses to stressful environmental events