Mating by outcrossing plants depends on the frequency and quality of the interaction between pollen vectors and individual flowers. However, the historical focus of pollination biology on individual flowers (floricentrism) cannot produce a complete understanding of the role of pollination in plant mating because mating is an aggregate process, which depends on the reproductive outcomes of all of a plant's flowers. Simultaneous display of multiple flowers in an inflorescence increases a plant's attractiveness to pollinators, which should, in general, enhance mating opportunities. However, whenever pollinators visit multiple flowers on an inflorescence, self-pollination among flowers can reduce the pollen available for export to other plants (pollen discounting) and can increase the incidence of inbreeding depression for embryos and offspring. Therefore, the size of floral displays that maximize mating frequency and quality generally balance the benefits of attractiveness against the costs of self-pollination. This balance can shift considerably if different flowers serve female and male functions at one time (sexual segregation) and flowers are arranged in inflorescences so that pollinators visit female flowers before male flowers. However, the effectiveness of sexual segregation depends on the extent to which a particular inflorescence architecture induces consistent movement patterns by pollinators. In general, the consistency of pollinator movement patterns varies with inflorescence architecture and differs between pollinator types. Such variation creates many options for the evolution of the diverse inflorescence characteristics observed within angiosperms, which can be appreciated only by moving beyond a floricentric perspective of the role of pollination in plant mating.
Sexually antagonistic (SA) genetic variation-in which alleles favored in one sex are disfavored in the other-is predicted to be common and has been documented in several animal and plant populations, yet we currently know little about its pervasiveness among species or its population genetic basis. Recent applications of genomics in studies of SA genetic variation have highlighted considerable methodological challenges to the identification and characterization of SA genes, raising questions about the feasibility of genomic approaches for inferring SA selection. The related fields of local adaptation and statistical genomics have previously dealt with similar challenges, and lessons from these disciplines can therefore help overcome current difficulties in applying genomics to study SA genetic variation. Here, we integrate theoretical and analytical concepts from local adaptation and statistical genomics research-including F ST and F IS statistics, genome-wide association studies, pedigree analyses, reciprocal transplant studies, and evolve-and-resequence experiments-to evaluate methods for identifying SA genes and genome-wide signals of SA genetic variation. We begin by developing theoretical models for between-sex F ST and F IS , including explicit null distributions for each statistic, and using them to critically evaluate putative multilocus signals of sex-specific selection in previously published datasets. We then highlight new statistics that address some of the limitations of F ST and F IS , along with applications of more direct approaches for characterizing SA genetic variation, which incorporate explicit fitness measurements. We finish by presenting practical guidelines for the validation and evolutionary analysis of candidate SA genes and discussing promising empirical systems for future work.
In hermaphrodites, pleiotropic genetic tradeoffs between female and male reproductive functions can lead to sexually antagonistic (SA) selection, where individual alleles have conflicting fitness effects on each sex function. While an extensive theory of SA selection exists for dioecious species, these results have not been generalized to hermaphrodites. We develop population genetic models of SA selection in simultaneous hermaphrodites, and evaluate effects of dominance, selection on each sex function, self-fertilization, and population size, on the maintenance of polymorphism. Under obligate outcrossing, hermaphrodite model predictions converge exactly with those of dioecious populations. Self-fertilization in hermaphrodites generates three points of divergence with dioecious theory. First, opportunities for stable polymorphism decline sharply and become less sensitive to dominance with increased selfing. Second, selfing introduces an asymmetry in the relative importance of selection through male versus female reproductive functions, expands the parameter space favorable for the evolutionary invasion of female-beneficial alleles, and restricts invasion criteria for male-beneficial alleles. Finally, contrary to models of unconditionally beneficial alleles, selfing decreases genetic hitchhiking effects of invading SA alleles, and should therefore decrease these population genetic signals of SA polymorphisms. We discuss implications of SA selection in hermaphrodites, including its potential role in the evolution of “selfing syndromes”.
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