Evolvability and trait function predict phenotypic divergence of plant populations.

Understanding the causes and limits of population divergence in phenotypic traits is a fundamental aim of evolutionary biology, with the potential to yield predictions of adaptation to environmental change. Reciprocal transplant experiments and the evaluation of optimality models suggest that local adaptation is common but not universal, and some studies suggest that trait divergence is highly constrained by genetic variances and covariances of complex phenotypes. We analyze a large database of population divergence in plants and evaluate whether evolutionary divergence scales positively with standing genetic variation within populations (evolvability), as expected if genetic constraints are evolutionarily important.

Sex-specific natural selection on SNPs in .

Selection that acts in a sex-specific manner causes the evolution of sexual dimorphism. Sex-specific phenotypic selection has been demonstrated in many taxa and can be in the same direction in the two sexes (differing only in magnitude), limited to one sex, or in opposing directions (antagonistic). Attempts to detect the signal of sex-specific selection from genomic data have confronted numerous difficulties.

Parasitic manipulation or by-product of infection: an experimental approach using trematode-infected snails.

Natural selection should favour parasite genotypes that manipulate hosts in ways that enhance parasite fitness. However, it is also possible that the effects of infection are not adaptive. Here we experimentally examined the phenotypic effects of infection in a snail-trematode system.

Observational evidence of herbivore-specific associational effects between neighboring conspecifics in natural, dimorphic populations of .

Associational effects-in which the vulnerability of a plant to herbivores is influenced by its neighbors-have been widely implicated in mediating plant-herbivore interactions. Studies of associational effects typically focus on interspecific interactions or pest-crop dynamics. However, associational effects may also be important for species with intraspecific variation in defensive traits.

Herbivore-mediated negative frequency-dependent selection underlies a trichome dimorphism in nature.

Negative frequency-dependent selection (NFDS) has been shown to maintain polymorphism in a diverse array of traits. The action of NFDS has been confirmed through modeling, experimental approaches, and genetic analyses. In this study, we investigated NFDS in the wild using morph-frequency changes spanning a 20-year period from over 30 dimorphic populations of .

Rapid reversal of a potentially constraining genetic covariance between leaf and flower traits in .

Genetic covariance between two traits generates correlated responses to selection, and may either enhance or constrain adaptation. exhibits potentially constraining genetic covariance between specific leaf area (SLA) and flower number in males. Flower number is likely to increase via fecundity selection but the correlated increase in SLA increases mortality, and SLA is under selection to decrease in dry habitats.

Water availability drives population divergence and sex-specific responses in a dioecious plant.

Premise: Water availability is an important abiotic factor, resulting in differences between plant species growing in xeric and mesic habitats. Species with populations occurring in both habitat types allow examination of whether water availability has acted as a selective force at the intraspecific level. Investigating responses to water availability with a dioecious species allows determination of whether males and females, which often have different physiologies and life histories, respond differently.

The X chromosome is necessary for ovule production in Silene latifolia.

Sex chromosomes stop recombining and accumulate differences over time. In particular, genes on the chromosome restricted to the heterogametic sex degenerate and become non-functional. Here, we investigated whether or not the degeneration of a plant Y chromosome was sufficient to cause ovules containing a Y to fail to develop, thereby eliminating the possibility of YY individuals.

Pollen competition is the mechanism underlying a variety of evolutionary phenomena in dioecious plants.

It has long been known that more pollen grains often arrive on stigmas than there are ovules to fertilize, resulting in pollen competition. Moreover, this competition among pollen grains (gametophytes) depends, in part, on their extensive haploid gene expression. Here I review how this leads to a variety of phenomena in dioecious plants of interest to evolutionary biologists.

Periodic, Parasite-Mediated Selection For and Against Sex.

Asexual lineages should rapidly replace sexual populations. Why sex then? The Red Queen hypothesis proposes that parasite-mediated selection against common host genotypes could counteract the per capita birth rate advantage of asexuals. Under the Red Queen hypothesis, fluctuations in parasite-mediated selection can drive fluctuations in the asexual population, leading to the coexistence of sexual and asexual reproduction.

The two-fold cost of sex: experimental evidence from a natural system.

Over four decades ago, John Maynard Smith showed that a mutation causing asexual reproduction should rapidly spread in a dioecious sexual population. His reasoning was that the per-capita birth rate of an asexual population would exceed that of a sexual population, because asexual females do not invest in sons. Hence, there is a cost of sexual reproduction that Maynard Smith called the "cost of males.

Divergence in style length and pollen size leads to a postmating-prezygotic reproductive barrier among populations of Silene latifolia.

A central tenet of speciation research is the need to identify reproductive isolating barriers. One approach to this line of research is to identify the phenotypes that lead to reproductive isolation. Several studies on flowering plants have shown that differences in style length contribute to reproductive isolation between species, leading us to consider whether style length could act as a reproductive barrier among populations of a single species.

Lineages of Silene nutans developed rapid, strong, asymmetric postzygotic reproductive isolation in allopatry.

Reproductive isolation can rise either as a consequence of genomic divergence in allopatry or as a byproduct of divergent selection in parapatry. To determine whether reproductive isolation in gynodioecious Silene nutans results from allopatric divergence or from ecological adaptation following secondary contact, we investigated the pattern of postzygotic reproductive isolation and hybridization in natural populations using two phylogeographic lineages, western (W1) and eastern (E1). Experimental crosses between the lineages identified strong, asymmetric postzygotic isolation between the W1 and the E1 lineages, independent of geographic overlap.

Evolution: Selfing Takes Species Down Stebbins's Blind Alley.

Shifts from outcrossing to selfing have occurred thousands of times across the tree of life. By reducing the size of the gene pool, selfing should limit adaptive potential. A refreshing empirical experiment with snails supports this long-standing hypothesis.

Male-female genotype interactions maintain variation in traits important for sexual interactions and reproductive isolation.

Prezygotic reproductive isolation can evolve quickly when sexual selection drives divergence in traits important for sexual interactions between populations. It has been hypothesized that standing variation for male/female traits and preferences facilitates this rapid evolution and that variation in these traits is maintained by male-female genotype interactions in which specific female genotypes prefer specific male traits. This hypothesis can also explain patterns of speciation when ecological divergence is lacking, but this remains untested because it requires information about sexual interactions in ancestral lineages.

Haldane's Rule: Genetic Bases and Their Empirical Support.

There are few patterns in evolution that are as rigidly held as Haldane's rule (HR), which states, "When in the first generation between hybrids between 2 species, 1 sex is absent, rare, or sterile, that sex is always the heterogametic sex." Yet despite considerable attention for almost a century, questions persist as to how many independent examples exist and what is (are) the underlying genetic cause(s). Here, we review recent evidence extending HR to plants, where previously it has only been documented in animals.

Experimental evolution: Assortative mating and sexual selection, independent of local adaptation, lead to reproductive isolation in the nematode Caenorhabditis remanei.

Using experimental evolution, we investigated the contributions of ecological divergence, sexual selection, and genetic drift to the evolution of reproductive isolation in Caenorhabditis remanei. The nematodes were reared on two different environments for 100 generations. They were assayed for fitness on both environments after 30, 64, and 100 generations, and hybrid fitness were analyzed after 64 and 100 generations.

Differences in style length confer prezygotic isolation between two dioecious species of Silene in sympatry.

One fundamental signature of reinforcement is elevated prezygotic reproductive isolation between related species in sympatry relative to allopatry. However, this alone is inadequate evidence for reinforcement, as traits conferring reproductive isolation can occur as a by-product of other forces. We conducted crosses between Silene latifolia and S.

Commentary: When does understanding phenotypic evolution require identification of the underlying genes?

Adaptive evolution is fundamentally a genetic process. Over the past three decades, characterizing the genes underlying adaptive phenotypic change has revealed many important aspects of evolutionary change. At the same time, natural selection is often fundamentally an ecological process that can often be studied without identifying the genes underlying the variation on which it acts.

Genetic architecture of isolation between two species of Silene with sex chromosomes and Haldane's rule.

Examination of the genetic architecture of hybrid breakdown can provide insight into the genetic mechanisms of commonly observed isolating phenomena such as Haldane's rule. We used line-cross analysis to dissect the genetic architecture of divergence between two plant species that exhibit Haldane's rule for male sterility and rarity, Silene latifolia and Silene diclinis. We made 15 types of crosses, including reciprocal F1, F2, backcrosses, and later-generation crosses, grew the seeds to flowering, and measured the number of viable ovules, proportion of viable pollen, and sex ratio.

On the importance of balancing selection in plants.

Balancing selection refers to a variety of selective regimes that maintain advantageous genetic diversity within populations. We review the history of the ideas regarding the types of selection that maintain such polymorphism in flowering plants, notably heterozygote advantage, negative frequency-dependent selection, and spatial heterogeneity. One shared feature of these mechanisms is that whether an allele is beneficial or detrimental is conditional on its frequency in the population.

Multiple paternity in the freshwater snail, Potamopyrgus antipodarum.

Mating multiply may incur costs, such as exposure to predators and to sexually transmitted diseases. Nevertheless, it may be favored, in spite of these costs, as a way to increase the genetic diversity of offspring through fertilization by multiple males. Here, we tested for multiple paternity in a freshwater snail (Potamopyrgus antipodarum), which is host to several species of sterilizing trematode worms.