Among dioecious flowering plants, females and males often differ in a range of morphological, physiological, and life-history traits. This is referred to as sexual dimorphism, and understanding why it occurs is a central question in evolutionary biology. Our review documents a range of sexually dimorphic traits in angiosperm species, discusses their ecological consequences, and details the genetic and evolutionary processes that drive divergence between female and male phenotypes. We consider why sexual dimorphism in plants is generally less well developed than in many animal groups, and also the importance of sexual and natural selection in contributing to differences between the sexes. Many sexually dimorphic characters, including both vegetative and flowering traits, are associated with differences in the costs of reproduction, which are usually greater in females, particularly in longer-lived species. These differences can influence the frequency and distribution of females and males across resource gradients and within heterogeneous environments, causing niche differences and the spatial segregation of the sexes. The interplay between sex-specific adaptation and the breakdown of between-sex genetic correlations allows for the independent evolution of female and male traits, and this is influenced in some species by the presence of sex chromosomes. We conclude by providing suggestions for future work on sexual dimorphism in plants, including investigations of the ecological and genetic basis of intraspecific variation, and genetic mapping and expression studies aimed at understanding the genetic architecture of sexually dimorphic trait variation.

Heteromorphic sex chromosomes have originated independently in many species, and a common feature of their evolution is the degeneration of the Y chromosome, characterized by a loss of gene content and function. Despite being of broad significance to our understanding of sex chromosome evolution, the genetic changes that occur during the early stages of Y-chromosome degeneration are poorly understood, especially in plants. Here, we investigate sex chromosome evolution in the dioecious plant Rumex hastatulus, in which X and Y chromosomes have evolved relatively recently and occur in two distinct systems: an ancestral XX/XY system and a derived XX/XY 1 Y 2 system. This polymorphism provides a unique opportunity to investigate the effect of sex chromosome age on patterns of divergence and gene degeneration within a species. Despite recent suppression of recombination and low X-Y divergence in both systems, we find evidence that Y-linked genes have started to undergo gene loss, causing ∼28% and ∼8% hemizygosity of the ancestral and derived X chromosomes, respectively. Furthermore, genes remaining on Y chromosomes have accumulated more amino acid replacements, contain more unpreferred changes in codon use, and exhibit significantly reduced gene expression compared with their X-linked alleles, with the magnitude of these effects being greatest for older sex-linked genes. Our results provide evidence for reduced selection efficiency and ongoing Y-chromosome degeneration in a flowering plant, and indicate that Y degeneration can occur soon after recombination suppression between sex chromosomes. molecular evolution | sex linkage | dioecy S ystems of sex determination involving X and Y chromosomes have evolved multiple times in both plants and animals, with Y chromosomes having lost much of their genetic function in many species (1-3). Evidence of DNA sequence homology between X-and Y-linked gene pairs in flowering plants (4-7) and fish (8) supports the idea that sex chromosomes have evolved from autosomes and subsequently diverged following the suppression of recombination between genes involved in sex determination. Evolutionary models predict that when regions of suppressed recombination evolve on Y chromosomes, the associated reduction in the effectiveness of selection should lead to a pattern of Y-chromosome degeneration in which genes carried on the Y become impaired in function and are eventually lost (1-3). The well-studied Y chromosomes in humans and Drosophila melanogaster, for example, show clear signs of degeneration: They almost completely lack homology to the X chromosome, exhibit a highly heterochromatic chromatin structure consisting largely of repetitive and ampliconic DNA, and carry few remaining protein-coding genes (9-13).Recent genomic studies of sex chromosomes in humans, rhesus macaques, and chimpanzees (12, 13) have provided detailed information regarding the genetic structure and gene content of Y chromosomes, shedding light on the processes contributing to their deterioration. However, we still ...

Plants show a wide range of variation in mating system, ploidy level, and demographic history, allowing for unique opportunities to investigate the evolutionary and genetic factors affecting genome-wide patterns of positive and negative selection. In this review, we highlight recent progress in our understanding of the extent and nature of selection on plant genomes. We discuss differences in selection as they relate to variation in demography, recombination, mating system, and ploidy. We focus on the population genetic consequences of these factors and argue that, although variation in the magnitude of purifying selection is well documented, quantifying rates of positive selection and disentangling the relative importance of recombination, demography, and ploidy are ongoing challenges. Large-scale comparative studies that examine the relative and joint importance of these processes, combined with explicit models of population history and selection, are key and feasible goals for future work.

X and Y chromosomes differ in effective population size ( ), rates of recombination, and exposure to natural selection, all of which can affect patterns of genetic diversity. On Y chromosomes with suppressed recombination, natural selection is expected to eliminate linked neutral variation, and lower the of Y compared to X chromosomes or autosomes. However, female-biased sex ratios and high variance in male reproductive success can also reduce Y-linked , making it difficult to infer the causes of low Y-diversity. Here, we investigate the factors affecting levels of polymorphism during sex chromosome evolution in the dioecious plant (Polygonaceae). Strikingly, we find that neutral diversity for genes on the Y chromosome is, on average, 2.1% of the value for their X-linked homologs, corresponding to a chromosome-wide reduction of 93% compared to the standard neutral expectation. We demonstrate that the magnitude of this diversity loss is inconsistent with reduced male caused by neutral processes. Instead, using forward simulations and estimates of the distribution of deleterious fitness effects, we show that Y chromosome diversity loss can be explained by purifying selection acting in aggregate over a large number of genetically linked sites. Simulations also suggest that our observed level of Y-diversity is consistent with the joint action of purifying and positive selection, but only for models in which there were fewer constrained sites than we empirically estimated. Given the relatively recent origin of sex chromosomes, our results imply that Y-chromosome degeneration in the early stages may be largely driven by selective interference rather than by neutral genetic drift of silenced Y-linked genes.

The domesticated almond [Prunus dulcis (L.) Batsch] and peach [P. persica (Mill.) D. A. Webb] originated on opposite sides of Asia and were independently domesticated ∼5000 yr ago. While interfertile, they possess alternate mating systems and differ in a number of morphological and physiological traits. Here, we evaluated patterns of genome-wide diversity in both almond and peach to better understand the impacts of mating system, adaptation, and domestication on the evolution of these taxa. Almond has around seven times the genetic diversity of peach, and high genome-wide FST values support their status as separate species. We estimated a divergence time of ∼8 MYA (million years ago), coinciding with an active period of uplift in the northeast Tibetan Plateau and subsequent Asian climate change. We see no evidence of a bottleneck during domestication of either species, but identify a number of regions showing signatures of selection during domestication and a significant overlap in candidate regions between peach and almond. While we expected gene expression in fruit to overlap with candidate selected regions, instead we find enrichment for loci highly differentiated between the species, consistent with recent fossil evidence suggesting fruit divergence long preceded domestication. Taken together, this study tells us how closely related tree species evolve and are domesticated, the impact of these events on their genomes, and the utility of genomic information for long-lived species. Further exploration of this data will contribute to the genetic knowledge of these species and provide information regarding targets of selection for breeding application, and further the understanding of evolution in these species.