Emmanuelle Porcher

Plant mating systems in a changing world

There is increasing evidence that human disturbance can negatively impact plant-pollinator interactions such as outcross pollination. We present a meta-analysis of 22 studies involving 27 plant species showing a significant reduction in the proportion of seeds outcrossed in response to anthropogenic habitat modifications. We discuss the evolutionary consequences of disturbance on plant mating systems, and in particular whether reproductive assurance through selfing effectively compensates for reduced outcrossing. The extent to which disturbance reduces pollinator versus mate availability could generate diverse selective forces on reproductive traits. Investigating how anthropogenic change influences plant mating will lead to new opportunities for better understanding of how mating systems evolve, as well as of the ecological and evolutionary consequences of human activities and how to mitigate them.

Correlations among Fertility Components Can Maintain Mixed Mating in Plants

Classical models studying the evolution of self‐fertilization in plants conclude that only complete selfing and complete outcrossing are evolutionarily stable. In contrast with this prediction, 42% of seed‐plant species are reported to have rates of self‐fertilization between 0.2 and 0.8. We propose that many previous models fail to predict intermediate selfing rates because they do not allow for functional relationships among three components of reproductive fitness: self‐fertilized ovules, outcrossed ovules, and ovules sired by successful pollen export. Because the optimal design for fertility components may differ, conflicts among the alternative pathways to fitness are possible, and the greatest fertility may be achieved with some self‐fertilization. Here we develop and analyze a model to predict optimal selfing rates that includes a range of possible relationships among the three components of reproductive fitness, as well as the effects of evolving inbreeding depression caused by deleterious mutations and of selection on total seed number. We demonstrate that intermediate selfing is optimal for a wide variety of relationships among fitness components and that inbreeding depression is not a good predictor of selfing‐rate evolution. Functional relationships subsume the myriad effects of individual plant traits and thus offer a more general and simpler perspective on mating system evolution.

Evaluation of floristic diversity in urban areas as a basis for habitat management

ABSTRACT Questions: How can floristic diversity be evaluated in conservation plans to identify sites of highest interest for biodiversity? What are the mechanisms influencing the distribution of species in human-dominated environments? What are the best criteria to identify sites where active urban management is most likely to enhance floristic diversity? Location: The Hauts-de-Seine district bordering Paris, France. Methods: We described the floristic diversity in one of the most urbanized French districts through the inventory of ca. 1000 sites located in 23 habitats. We built a new index of floristic interest (IFI), integrating information on richness, indigeneity, typicality and rarity of species, to identify sites and habitats of highest interest for conservation. Finally, we explored the relationship between site IFI and land use patterns (LUP). Results: We observed a total of 626 vascular plant species. Habitats with highest IFI were typically situated in semi-natural environments or environments with moderate human impact. We also showed that neighbouring (urban) structures had a significant influence on the floristic interest of sites: for example, the presence of collective dwellings around a site had a strong negative impact on IFI. Conclusions: Our approach can be used to optimize management in urban zones; we illustrate such possibilities by defining a ‘Site Potential Value’, which was then compared with the observed IFI, to identify areas (e.g. river banks) where better management could improve the districts biodiversity. Nomenclature: Kerguélen (2003).

Constraints imposed by pollinator behaviour on the ecology and evolution of plant mating systems

Most flowering plants rely on pollinators for their reproduction. Plant‐pollinator interactions, although mutualistic, involve an inherent conflict of interest between both partners and may constrain plant mating systems at multiple levels: the immediate ecological plant selfing rates, their distribution in and contribution to pollination networks, and their evolution. Here, we review experimental evidence that pollinator behaviour influences plant selfing rates in pairs of interacting species, and that plants can modify pollinator behaviour through plastic and evolutionary changes in floral traits. We also examine how theoretical studies include pollinators, implicitly or explicitly, to investigate the role of their foraging behaviour in plant mating system evolution. In doing so, we call for more evolutionary models combining ecological and genetic factors, and additional experimental data, particularly to describe pollinator foraging behaviour. Finally, we show that recent developments in ecological network theory help clarify the impact of community‐level interactions on plant selfing rates and their evolution and suggest new research avenues to expand the study of mating systems of animal‐pollinated plant species to the level of the plant‐pollinator networks.

EXPERIMENTAL DEMONSTRATION OF A CAUSAL RELATIONSHIP BETWEEN HETEROGENEITY OF SELECTION AND GENETIC DIFFERENTIATION IN QUANTITATIVE TRAITS

Abstract Comparisons of estimates of genetic differentiation at molecular markers (FST) and at quantitative traits (QST) are a means of inferring the level and heterogeneity of selection in natural populations. However, such comparisons are questionable because they require that the influence of drift and selection on QST be detectable over possible background influences of environmental or nonadditive genetic effects on QST‐values. Here we test this using an experimental evolution approach in metapopulations of Arabidopsis thaliana experiencing different levels of drift and selection heterogeneity. We estimated the intensity and heterogeneity of selection on morphological and phe‐nological traits via selection differentials. We demonstrate that QST‐values increased with increasing selection heterogeneity when genetic drift was limited. The effect of selection on QST was thus detectable despite significant genotype‐by‐environment interactions that most probably biased the estimates of genetic differentiation. Although they cannot be used as a direct validation of the conclusions of prior studies, our results strongly support both the relevance of QST as an estimator of genetic differentiation and the role of local selection in shaping the genetic differentiation of natural populations.

The genetic consequences of fluctuating inbreeding depression and the evolution of plant selfing rates.

The magnitude of inbreeding depression, a central parameter in the evolution of plant mating systems, can vary depending on environmental conditions. However, the underlying genetic mechanisms causing environmental fluctuations in inbreeding depression, and the consequences of this variation for the evolution of self‐fertilization, have been little studied. Here, we consider temporal fluctuations of the selection coefficient in an explicit genetic model of inbreeding depression. We show that substantial variance in inbreeding depression can be generated at equilibrium by fluctuating selection, although the simulated variance tends to be lower than has been measured in experimental studies. Our simulations also reveal that purging of deleterious mutations does not depend on the variance in their selection coefficient. Finally, an evolutionary analysis shows that, in contrast to previous theoretical approaches, intermediate selfing rates are never evolutionarily stable when the variation in inbreeding depression is due to fluctuations in the selection coefficient on deleterious mutations.

On the use of parataxonomy in biodiversity monitoring: a case study on wild flora

Monitoring programs that assess species-richness and turnover are now regarded as essential to document biodiversity loss worldwide. Implementation of such programs is impeded by a general decrease in the number of skilled naturalists. Here we studied how morphotypes, instead of species, might be used by unskilled participants (referred to as “volunteers”) to survey common plant communities. Our main questions were: (1) Can morphotypes be used as a robust estimator of species-richness (α-diversity) and assemblage turnover (β-diversity)? and (2) What is the robustness (reproducibility and repeatability) of such methods? Double inventories were performed on 150 plots in arable field margins, one by a non-expert using morphotypes, the other by a taxonomist using species. To test the robustness of morphotype identification among participants, 20 additional plots were surveyed by eight volunteers using the same protocol. We showed that (1) the number of morphotypes identified by unskilled volunteers in a plot was always strongly correlated with species-richness. (2) Morphotypes were sensitive to differences among habitats but were less accurate than species to detect these differences. (3) Morphotype identification varied significantly within and between volunteers. Due to this lack of repeatability and reproducibility, parataxonomy cannot be considered a good surrogate for taxonomy. Nevertheless, assuming that morphotypes are identified with standardized methods, and that results are used only to evaluate gross species-richness but not species turnover, parataxonomy might be a valuable tool for rapid biodiversity assessment of common wild flora.

Maintenance of Quantitative Genetic Variance Under Partial Self-Fertilization, with Implications for Evolution of Selfing

We analyze two models of the maintenance of quantitative genetic variance in a mixed-mating system of self-fertilization and outcrossing. In both models purely additive genetic variance is maintained by mutation and recombination under stabilizing selection on the phenotype of one or more quantitative characters. The Gaussian allele model (GAM) involves a finite number of unlinked loci in an infinitely large population, with a normal distribution of allelic effects at each locus within lineages selfed for τ consecutive generations since their last outcross. The infinitesimal model for partial selfing (IMS) involves an infinite number of loci in a large but finite population, with a normal distribution of breeding values in lineages of selfing age τ. In both models a stable equilibrium genetic variance exists, the outcrossed equilibrium, nearly equal to that under random mating, for all selfing rates, r, up to critical value, r^, the purging threshold, which approximately equals the mean fitness under random mating relative to that under complete selfing. In the GAM a second stable equilibrium, the purged equilibrium, exists for any positive selfing rate, with genetic variance less than or equal to that under pure selfing; as r increases above r^ the outcrossed equilibrium collapses sharply to the purged equilibrium genetic variance. In the IMS a single stable equilibrium genetic variance exists at each selfing rate; as r increases above r^ the equilibrium genetic variance drops sharply and then declines gradually to that maintained under complete selfing. The implications for evolution of selfing rates, and for adaptive evolution and persistence of predominantly selfing species, provide a theoretical basis for the classical view of Stebbins that predominant selfing constitutes an “evolutionary dead end.”

Inbreeding depression under mixed outcrossing, self-fertilization and sib-mating

BackgroundBiparental inbreeding, mating between two relatives, occurs at a low frequency in many natural plant populations, which also often have substantial rates of self-fertilization. Although biparental inbreeding is likely to influence the dynamics of inbreeding depression and the evolution of selfing rates, it has received limited theoretical attention in comparison to selfing. The only previous model suggested that biparental inbreeding can favour the maintenance of stable intermediate selfing rates, but made unrealistic assumptions about the genetic basis of inbreeding depression. Here we extend a genetic model of inbreeding depression, describing nearly recessive lethal mutations at a very large number of loci, to incorporate sib-mating. We also include a constant component of inbreeding depression modelling the effects of mildly deleterious, nearly additive alleles. We analyze how observed rates of sib-mating influence the mean number of heterozygous lethals alleles and inbreeding depression in a population reproducing by a mixture of self-fertilization, sib-mating and outcrossing. We finally use the ensuing relationship between equilibrium inbreeding depression and population selfing rate to infer the evolutionarily stable selfing rates expected under such a mixed mating system.ResultsWe show that for a given rate of inbreeding, sib-mating is more efficient at purging inbreeding depression than selfing, because homozygosity of lethals increases more gradually through sib-mating than through selfing. Because sib-mating promotes the purging of inbreeding depression and the evolution of selfing, our genetic model of inbreeding depression also predicts that sib-mating is unlikely to maintain stable intermediate selfing rates.ConclusionsOur results imply that even low rates of sib-mating affect plant mating system evolution, by facilitating the evolution of selfing via more efficient purging of inbreeding depression. Alternative mechanisms, such as pollination ecology, are necessary to explain stable mixed selfing and outcrossing.

Biogeographic comparisons of herbivore attack, growth and impact of Japanese knotweed between Japan and France

To shed light on the process of how exotic species become invasive, it is necessary to study them both in their native and non-native ranges. Our intent was to measure differences in herbivory, plant growth and the impact on other species in Fallopia japonica in its native and non-native ranges. We performed a cross-range full descriptive, field study in Japan (native range) and France (non-native range). We assessed DNA ploidy levels, the presence of phytophagous enemies, the amount of leaf damage, several growth parameters and the co-occurrence of Fallopia japonica with other plant species of herbaceous communities. Invasive Fallopia japonica plants were all octoploid, a ploidy level we did not encounter in the native range, where plants were all tetraploid. Octoploids in France harboured far less phytophagous enemies, suffered much lower levels of herbivory, grew larger and had a much stronger impact on plant communities than tetraploid conspecifics in the native range in Japan. Our data confirm that Fallopia japonica performs better-plant vigour and dominance in the her-baceous community-in its non-native than its native range. Because we could not find octoploids in the native range, we cannot separate the effects of differences in ploidy from other biogeographic factors. To go further, common garden experiments would now be needed to disentangle the proper role of each factor, taking into account the ploidy levels of plants in their native and non-native ranges. Synthesis. As the process by which invasive plants successfully invade ecosystems in their non-native range is probably multifactorial in most cases, examining several components-plant growth, herbivory load, impact on recipient systems-of plant invasions through biogeographic comparisons is important. Our study contributes towards filling this gap in the research, and it is hoped that this method will spread in invasion ecology, making such an approach more common.