Theory suggests that genetic conflicts drive turnovers between sex-determining mechanisms, yet these studies only apply to cases where sex allocation is independent of environment or condition. Here, we model parent-offspring conflict in the presence of condition-dependent sex allocation, where the environment has sex-specific fitness consequences. Additionally, one sex is assumed to be more costly to produce than the other, which leads offspring to favor a sex ratio less biased toward the cheaper sex in comparison to the sex ratio favored by mothers. The scope for parent-offspring conflict depends on the relative frequency of both environments: when one environment is less common than the other, parent-offspring conflict can be reduced or even entirely absent, despite a biased population sex ratio. The model shows that conflict-driven invasions of condition-independent sex factors (e.g., sex chromosomes) result either in the loss of condition-dependent sex allocation, or, interestingly, lead to stable mixtures of condition-dependent and condition-independent sex factors. The latter outcome corresponds to empirical observations in which sex chromosomes are present in organisms with environment-dependent sex determination. Finally, conflict can also favor errors in environmental perception, potentially resulting in the loss of condition-dependent sex allocation without genetic changes to sex-determining loci.
There is a growing interest in predicting the social and ecological contexts that favor the evolution of maternal effects. Most predictions focus, however, on maternal effects that affect only a single character, whereas the evolution of maternal effects is poorly understood in the presence of suites of interacting traits. To overcome this, we simulate the evolution of multivariate maternal effects (captured by the matrix M) in a fluctuating environment. We find that the rate of environmental fluctuations has a substantial effect on the properties of M: in slowly changing environments, offspring are selected to have a multivariate phenotype roughly similar to the maternal phenotype, so that M is characterized by positive dominant eigenvalues; by contrast, rapidly changing environments favor Ms with dominant eigenvalues that are negative, as offspring favor a phenotype which substantially differs from the maternal phenotype. Moreover, when fluctuating selection on one maternal character is temporally delayed relative to selection on other traits, we find a striking pattern of cross-trait maternal effects in which maternal characters influence not only the same character in offspring, but also other offspring characters. Additionally, when selection on one character contains more stochastic noise relative to selection on other traits, large cross-trait maternal effects evolve from those maternal traits that experience the smallest amounts of noise. The presence of these cross-trait maternal effects shows that individual maternal effects cannot be studied in isolation, and that their study in a multivariate context may provide important insights about the nature of past selection. Our results call for more studies that measure multivariate maternal effects in wild populations.
One key trade-off underlying life-history evolution is the one between age and size at maturity, with earlier maturation leading to greater chances of juvenile survival at the cost of reduced fecundity as an adult. Here we model the impact of limited dispersal and kin competition on the stable resolution of this trade-off. We show that if mating is at least occasionally nonlocal, then limited dispersal favors juvenile survival over adult fecundity in females, promoting earlier female maturation at the population level; at the same time, it favors adult fecundity over juvenile survival in males, promoting later male maturation. Limited dispersal and local competition can thus drive the evolution of sexual dimorphism in the timing of maturation and consequent dimorphism in body size. At the individual level, if maturation can be flexibly adjusted in response to dispersal status, then both males and females who disperse as offspring should mature earlier than those who remain on their natal patch.
An attractive way to improve our understanding of sex determination evolution is to study the underlying mechanisms in closely related species and in a phylogenetic perspective. Hymenopterans are well suited owing to the diverse sex determination mechanisms, including different types of Complementary Sex Determination (CSD) and maternal control sex determination. We investigated different types of CSD in four species within the braconid wasp genus Asobara that exhibit diverse life-history traits. Nine to thirteen generations of inbreeding were monitored for diploid male production, brood size, offspring sex ratio, and pupal mortality as indicators for CSD. In addition, simulation models were developed to compare these observations to predicted patterns for multilocus CSD with up to ten loci. The inbreeding regime did not result in diploid male production, decreased brood sizes, substantially increased offspring sex ratios nor in increased pupal mortality. The simulations further allowed us to reject CSD with up to ten loci, which is a strong refutation of the multilocus CSD model. We discuss how the absence of CSD can be reconciled with the variation in life-history traits among Asobara species, and the ramifications for the phylogenetic distribution of sex determination mechanisms in the Hymenoptera.
Female choice is a powerful selective force, driving the elaboration of conspicuous male ornaments. This process of sexual selection has profound implications for many life-history decisions, including sex allocation. For example, females with attractive partners should produce more sons, because these sons will inherit their fathers attractiveness and enjoy high mating success, thereby yielding greater fitness returns than daughters. However, previous research has overlooked the fact that there is a reciprocal feedback from life-history strategies to sexual selection. Here, using a simple mathematical model, we show that if mothers adaptively control offspring sex in relation to their partners attractiveness, sexual selection is weakened and male ornamentation declines. This weakening occurs because the ability to determine offspring sex reduces the fitness difference between females with attractive and unattractive partners. We use individual-based, evolutionary simulations to show that this result holds under more biologically realistic conditions. Sexual selection and sex allocation thus interact in a dynamic fashion: The evolution of conspicuous male ornaments favors sex-ratio adjustment, but this conditional strategy then undermines the very same process that generated it, eroding sexual selection. We predict that, all else being equal, the most elaborate sexual displays should be seen in species with little or no control over offspring sex. The feedback process we have described points to a more general evolutionary principle, in which a conditional strategy weakens directional selection on another trait by reducing fitness differences.
One basic condition of postmating sexual selection is that females mate more than once before fertilizing their ova. Knowledge of the frequency and extent of multiple mating in a given population or species is therefore important in order to fully understand the potential for sexual selection, in the form of sperm competition, sexual conflict and cryptic female choice. Surprisingly, there are only a handful of studies that have attempted to estimate the frequency of multiple mating in insects (including Drosophila) and none have made direct observations over extended periods of time. Here we use time-lapse photography to directly score matings in isolated pairs of D. melanogaster and show that multiple mating in the laboratory occurs at a high frequency but at comparable rates with wild caught females. We also find that the interval to remating rises approximately additively with each mating, indicating either an increase in female resistance or male reluctance to remate. These results suggest that the opportunity for postmating sexual selection in laboratory and natural environments are not dramatically different and that there may be a causal link between the rise in female mating resistance and the concomitant rise in the cost of mating. The method is easily executed and could be adapted to other insect models.
Natural enemies may go through genetic bottlenecks during the process of biological control introductions. Such bottlenecks are expected to be particularly detrimental in parasitoid Hymenoptera that exhibit complementary sex determination (CSD). CSD is associated with a severe form of inbreeding depression because homozygosity at one or multiple sex loci leads to the production of diploid males that are typically unviable or sterile. We observed that diploid males occur at a relatively high rate (8-13% of diploid adults) in a field population of Cotesia rubecula in Minnesota, USA, where this parasitoid was introduced for biological control of the cabbage white Pieris rapae. However, our laboratory crosses suggest two-locus CSD in a native Dutch population of C. rubecula and moderately high diploid males survival (approximately 70%), a scenario expected to produce low proportions of diploid males. We also show that courtship behavior of diploid males is similar to that of haploid males, but females mated to diploid males produce only very few daughters that are triploid. We use our laboratory data to estimate sex allele diversity in the field population of C. rubecula and discuss the possibility of a sex determination meltdown from two-locus CSD to effective single-locus CSD during or after introduction.
Intralocus sexual conflict (IASC) occurs when a trait under selection in one sex constrains the other sex from achieving its sex-specific fitness optimum. Selection pressures on body size often differ between the sexes across many species, including humans: among men individuals of average height enjoy the highest reproductive success, while shorter women have the highest reproductive success. Given its high heritability, IASC over human height is likely. Using data from sibling pairs from the Wisconsin Longitudinal Study, we present evidence for IASC over height: in shorter sibling pairs (relatively) more reproductive success (number of children) was obtained through the sister than through the brother of the sibling pair. By contrast, in average height sibling pairs most reproductive success was obtained through the brother relative to the sister. In conclusion, we show that IASC over a heritable, sexually dimorphic physical trait (human height) affects Darwinian fitness in a contemporary human population.
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