Gender specialisation and stigma height dimorphism in Mediterranean Lithodora fruticosa (Boraginaceae).

Dimorphism in style height has evolved repeatedly in flowering plants, with some individuals having short and others long styles; in the case of distylous species, stigma position varies reciprocally with that of the anthers. Distyly can be associated with divergence in the functional gender between long- and short-styled individuals, but gender divergence has rarely been investigated in species with a simple stigma height polymorphism in the absence of reciprocal dimorphism in anther position. To evaluate the relation between stigma height polymorphism and gender, I measured the dimensions of floral morphology and seed production for the two morphs of a large population of the Iberian species Lithodora fruticosa (Boraginaceae). Results confirm the existence of a stigma height polymorphism in L. fruticosa, with long- and short-styled individuals at a 1:1 ratio in the studied population. Long-styled individuals produced substantially more seeds than did short-styled individuals, pointing to strong divergence in functional gender between the two morphs. The results of this study are puzzling in light of recent work that suggests that L. fruticosa has a multi-allelic self-incompatibility system. I discuss the significance of gender divergence in L. fruticosa and evaluate hypotheses that might explain it

Inbreeding depression is high in a self-incompatible perennial herb population but absent in a self-compatible population showing mixed mating.

High inbreeding depression is thought to be one of the major factors preventing evolutionary transitions in hermaphroditic plants from self-incompatibility (SI) and outcrossing toward self-compatibility (SC) and selfing. However, when selfing does evolve, inbreeding depression can be quickly purged, allowing the evolution of complete self-fertilization. In contrast, populations that show intermediate selfing rates (a mixed-mating system) typically show levels of inbreeding depression similar to those in outcrossing species, suggesting that selection against inbreeding might be responsible for preventing the transition toward complete self-fertilization. By implication, crosses among populations should reveal patterns of heterosis for mixed-mating populations that are similar to those expected for outcrossing populations. Using hand-pollination crosses, we compared levels of inbreeding depression and heterosis between populations of Linaria cavanillesii (Plantaginaceae), a perennial herb showing contrasting mating systems. The SI population showed high inbreeding depression, whereas the SC population displaying mixed mating showed no inbreeding depression. In contrast, we found that heterosis based on between-population crosses was similar for SI and SC populations. Our results are consistent with the rapid purging of inbreeding depression in the derived SC population, despite the persistence of mixed mating. However, the maintenance of outcrossing after a transition to SC is inconsistent with the prediction that populations that have purged their inbreeding depression should evolve toward complete selfing, suggesting that the transition to SC in L. cavanillesii has been recent. SC in L. cavanillesii thus exemplifies a situation in which the mating system is likely not at an equilibrium with inbreeding depression

No difference in plasticity between different ploidy levels in the Mediterranean herb Mercurialis annua.

Increased phenotypic plasticity for a number of plant traits has been suggested as a possible reason for the success and spread of polyploids. One such trait is a plant's sex allocation (or gender), which influences its reproductive success directly as a function of the potentially heterogeneous mating prospects in the population. However, it is unknown how polyploidy per se might affect plasticity in a plant's sex allocation. Although there have been numerous comparisons between diploid and (usually) tetraploid taxa, we know very little about how elevated ploidy above the diploid level might affect plasticity. Here, we ask whether different ploidy levels > 2x express different plasticity in the ruderal plant Mercurialis annua. We grew tetraploid and hexaploid hermaphrodites under different levels of nutrient availability and compared their reaction norms for growth (above-ground biomass, SLA) and reproductive traits (reproductive effort, phenotypic gender). Overall, we found that an increase in ploidy level from 4x to 6x in M. annua is associated with an increase in the relative biomass allocated to seeds, measured as female reproductive effort. However, our study provides no support for the idea that increasing ploidy level increases the ability to express different phenotypes in response to changes in the environment

Kin discrimination allows plants to modify investment towards pollinator attraction.

Pollinators tend to be preferentially attracted to large floral displays that may comprise more than one plant in a patch. Attracting pollinators thus not only benefits individuals investing in advertising, but also other plants in a patch through a 'magnet' effect. Accordingly, there could be an indirect fitness advantage to greater investment in costly floral displays by plants in kin-structured groups than when in groups of unrelated individuals. Here, we seek evidence for this strategy by manipulating relatedness in groups of the plant Moricandia moricandioides, an insect-pollinated herb that typically grows in patches. As predicted, individuals growing with kin, particularly at high density, produced larger floral displays than those growing with non-kin. Investment in attracting pollinators was thus moulded by the presence and relatedness of neighbours, exemplifying the importance of kin recognition in the evolution of plant reproductive strategies

High rates of evolution preceded shifts to sex-biased gene expression in Leucadendron, the most sexually dimorphic angiosperms.

Differences between males and females are usually more subtle in dioecious plants than animals, but strong sexual dimorphism has evolved convergently in the South African Cape plant genus Leucadendron. Such sexual dimorphism in leaf size is expected largely to be due to differential gene expression between the sexes. We compared patterns of gene expression in leaves among 10 Leucadendron species across the genus. Surprisingly, we found no positive association between sexual dimorphism in morphology and the number or the percentage of sex-biased genes (SBGs). Sex bias in most SBGs evolved recently and was species specific. We compared rates of evolutionary change in expression for genes that were sex biased in one species but unbiased in others and found that SBGs evolved faster in expression than unbiased genes. This greater rate of expression evolution of SBGs, also documented in animals, might suggest the possible role of sexual selection in the evolution of gene expression. However, our comparative analysis clearly indicates that the more rapid rate of expression evolution of SBGs predated the origin of bias, and shifts towards bias were depleted in signatures of adaptation. Our results are thus more consistent with the view that sex bias is simply freer to evolve in genes less subject to constraints in expression level

The divergence history of the perennial plant Linaria cavanillesii confirms a recent loss of self-incompatibility.

Many angiosperms prevent inbreeding through a self-incompatibility (SI) system, but the loss of SI has been frequent in their evolutionary history. The loss of SI may often lead to an increase in the selfing rate, with the purging of inbreeding depression and the ultimate evolution of a selfing syndrome, where plants have smaller flowers with reduced pollen and nectar production. In this study, we used approximate Bayesian computation (ABC) to estimate the timing of divergence between populations of the plant Linaria cavanillesii that differ in SI status and in which SI is associated with low inbreeding depression but not with a transition to full selfing or a selfing syndrome. Our analysis suggests that the mixed-mating self-compatible (SC) population may have begun to diverge from the SI populations around 2810 generation ago, a period perhaps too short for the evolution of a selfing syndrome. We conjecture that the SC population of L. cavanillesii is at an intermediate stage of transition between outcrossing and selfing

Development and characterization of microsatellite markers for diploid populations of the wind-pollinated herb Mercurialis annua.

Mercurialis annua is a wind-pollinated annual plant that has long been used as a model for the study of ploidy and sexual-systems evolution. However, no molecular markers are yet available for genetic studies of its diploid populations. Here, we develop and characterize a set of eight polymorphic microsatellite markers for diploid dioecious M. annua. Following an SSR-enrichment protocol, 13 microsatellite markers were proposed, eight of which yielded successful amplification and polymorphism. We screened the eight microsatellite loci in 100 individuals. The number of alleles per marker ranged from 6 to 12, and observed heterozygosity ranged from 0.57 to 0.76. To estimate potential allele scoring errors, these individuals' offspring were genotyped for the same loci, and error rates were estimated from parentage analyses. Error rates ranged from 0 to 6.8%. Cross-amplification tests were performed for congeneric M. huetti and M. canariensis, with successful amplification for seven and six of the eight loci, respectively. The novel microsatellite markers proposed here will be crucial for a multitude of genetic studies of M. annua and further establish its importance as a model species for addressing ecological and population genetic questions

Exogenous selection shapes germination behaviour and seedling traits of populations at different altitudes in a Senecio hybrid zone.

BACKGROUND AND AIMS: The Senecio hybrid zone on Mt Etna, Sicily, is characterized by steep altitudinal clines in quantitative traits and genetic variation. Such clines are thought to be maintained by a combination of 'endogenous' selection arising from genetic incompatibilities and environment-dependent 'exogenous' selection leading to local adaptation. Here, the hypothesis was tested that local adaptation to the altitudinal temperature gradient contributes to maintaining divergence between the parental species, S. chrysanthemifolius and S. aethnensis. METHODS: Intra- and inter-population crosses were performed between five populations from across the hybrid zone and the germination and early seedling growth of the progeny were assessed. KEY RESULTS: Seedlings from higher-altitude populations germinated better under low temperatures (9-13 °C) than those from lower altitude populations. Seedlings from higher-altitude populations had lower survival rates under warm conditions (25/15 °C) than those from lower altitude populations, but also attained greater biomass. There was no altitudinal variation in growth or survival under cold conditions (15/5 °C). Population-level plasticity increased with altitude. Germination, growth and survival of natural hybrids and experimentally generated F(1)s generally exceeded the worse-performing parent. CONCLUSIONS: Limited evidence was found for endogenous selection against hybrids but relatively clear evidence was found for divergence in seed and seedling traits, which is probably adaptive. The combination of low-temperature germination and faster growth in warm conditions might enable high-altitude S. aethnensis to maximize its growth during a shorter growing season, while the slower growth of S. chrysanthemifolius may be an adaptation to drought stress at low altitudes. This study indicates that temperature gradients are likely to be an important environmental factor generating and maintaining adaptive divergence across the Senecio hybrid zone on Mt Etna