Evaluating a simple approximation to modeling the joint evolution of self-fertilization and inbreeding depression

International audienceA comprehensive understanding of plant mating system evolution requires detailed genetic models for both the mating system and inbreeding depression, which are often intractable. A simple approximation assuming that the mating system evolves by small infrequent mutational steps has been proposed. We examine its accuracy by comparing the evolutionarily stable selfing rates it predicts to those obtained from an explicit genetic model of the selfing rate, when inbreeding depression is caused by partly recessive deleterious mutations at many loci. Both models also include pollen limitation and pollen discounting. The approximation produces reasonably accurate predictions with a low or moderate genomic mutation rate to deleterious alleles, on the order of U = 0.02 to 0.2. However, for high mutation rates, the predictions of the full genetic model differ substantially from those of the approximation, especially with nearly recessive lethal alleles. This occurs because when a modifier allele affecting the selfing rate is rare, homozygous modifiers are produced mainly by selfing, which enhances the opportunity for purging nearly recessive lethals and increases the marginal fitness of the allele modifying the selfing rate. Our results confirm that explicit genetic models of selfing rate and inbreeding depression are required to understand mating system evolution

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

International audienceWe 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, Embedded Image, 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 Embedded Image 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 Embedded Image 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 maintained by recessive lethal mutations interacting with stabilizing selection on quantitative characters in a partially self-fertilizing population

International audienceThe bimodal distribution of fitness effects of new mutations and standing genetic variation, due to early-acting strongly deleterious recessive mutations and late-acting mildly deleterious mutations, is analyzed using the Kondrashov model for lethals (K), with either the infinitesimal model for selfing (IMS) or the Gaussian allele model (GAM) for quantitative genetic variance under stabilizing selection. In the combined models (KIMS and KGAM) high genomic mutation rates to lethals and weak stabilizing selection on many characters create strong interactions between early and late inbreeding depression, by changing the distribution of lineages selfed consecutively for different numbers of generations. Alternative stable equilibria can exist at intermediate selfing rates for a given set of parameters. Evolution of quantitative genetic variance under multivariate stabilizing selection can strongly influence the purging of nearly recessive lethals, and sometimes vice versa. If the selfing rate at the purging threshold for quantitative genetic variance in IMS or GAM alone exceeds that for nearly recessive lethals in K alone, then in KIMS and KGAM stabilizing selection causes selective interference with purging of lethals, increasing the mean number of lethals compared to K; otherwise, stabilizing selection causes selective facilitation in purging of lethals, decreasing the mean number of lethals

THE EVOLUTION OF SELF-FERTILIZATION AND INBREEDING DEPRESSION UNDER POLLEN DISCOUNTING AND POLLEN LIMITATION

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

International audienceWe 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, Embedded Image, 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 Embedded Image 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 Embedded Image 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.

LANDSCAPE FEATURES ARE A BETTER CORRELATE OF WILD PLANT POLLINATION THAN AGRICULTURAL PRACTICES IN AN INTENSIVE CROPPING SYSTEM

International audienceOrganic farming is commonly associated with increased pollinator diversity and abundance, but the net effects on pollination rates are less documented. Besides, organic farms are often surrounded by more diverse landscapes than conventional farms, such that the contributions of landscape diversity vs. farming practices to pollination rates are often confounded with each other. Here, the roles of local vs. landscape scale variables on pollination rates of experimental plants are examined in agricultural landscapes. To this end, fruit set and seed production were measured in the obligate insect-pollinated Lotus corniculatus. Plants were located in pairs of neighboring organic vs. conventional farms, which were characterized by contrasting landscape structures and compositions. Fruit set, a proxy for pollinator visitation rates, was significantly related to landscape variables: fruit set was higher in farms close to a patch of semi-natural habitat, but lower in landscapes with a high cover of semi-natural habitats. Fruit set also correlated with local variables, such as habitat type, but not with farming type. Identical pollination rates in conventional and organic farms are likely due to similar diversities of habitats, crops and weeds in both farming types of the study area. These results therefore confirm that habitat diversity prevails over pesticide-free practices to explain the higher pollinator abundances usually observed in organic vs. conventional farms

Ecological specialization and rarity indices estimated for a large number of plant species in France

International audienceThe biological diversity of the Earth is being rapidly depleted due to the direct and indirect consequences of human activities. Specialist or rare species are generally thought to be more extinction prone than generalist or common species. Testing this assumption however requires that the rarity and ecological specialization of the species are quantified. Many indices have been developed to classify species as generalists vs. specialists or as rare vs. common, but large data sets are needed to calculate these indices. Here, we present a list of specialization and rarity values for more than 2800 plant species of continental France, which were computed from the large botanical and ecological dataset SOPHY. Three specialization indices were calculated using species co-occurrence data. All three indices are based on (dis)similarity among plant communities containing a focal species, quantified either as beta diversity in an additive (Fridley et al., 2007 [6]) or multiplicative (Zeleny, 2008 [15]) partitioning of diversity or as the multiple site similarity of Baselga et al. (2007) [1]. Species rarity was calculated as the inverse of a species occurrence

Assessing non-intended effects of farming practices on field margin vegetation with a functional approach

International audienceTo assess the unwanted side effects of farming practices on non-target plants, we used a nationwide survey of the vegetation of arable field margin strips. The vegetation was surveyed during two years (2013, 2014) in 430 field margins distributed over all agricultural regions of France. We used two complementary multivariate, trait-based approaches to examine how ten plant traits were related to ten environmental variables describing abiotic conditions, landscape factors, field margin management and in-field practices. Generalized additive mixed models were also developed to assess how the same environmental variables correlated with species richness, functional diversity and relative richness of agrotolerant versus hemerophobic species. Traits responded primarily to an environmental gradient of landscape diversity and field margin management. For instance, narrow field margin strips, frequent management and presence of a ditch favoured annual plants, small size at maturity and perennial plants, respectively. The second environmental gradient affecting plant traits was related to field size and intensity of in-field farming practices. On this gradient, fertilizer drift appeared to have a much stronger effect on plant trait composition of field margin strips than herbicide drift. The relationship between species richness, or functional diversity, and environment was consistent with the trait-based approach: the two former variables were negatively correlated with agriculture intensification (e.g. field size). However, this analysis also highlighted new covariates, such as a negative relationship between frequency of herbicide use and species richness. Some of the observed patterns seemed to be driven by differential responses of agrotolerant versus hemerophobic species, with the latter being more species-rich under organic than under conventional farming. Despite efforts to reduce nitrogen inputs since the 2000s, our results shows that N-fertilization still has significant non-intended effects on field margin vegetation. More generally, increasing the width of field margin strips, keeping or restoring semi-natural elements (ditches, hedges) in the field boundary, and lowering the number of management events may promote grassland plant species more typical of semi-natural habitats

Connectivity, habitat heterogeneity, and population persistence in Ranunculus nodiflorus, an endangered species in France

International audience• Here, we explore the role of habitat spatial structure in the maintenance of metapopulations of Ranunculus nodiflorus. This rare species grows in puddles that can be connected occasionally by flooded corridors. • We monitored five locations in the Fontainebleau forest, France, since 2002 and recorded the presence of corridors among puddles and evaluated their impact on puddle demography and plant fitness. • We showed that connections increased population size, by increasing both the number of puddles occupied by the species and the density of individuals within puddles, but seemed to have no direct influence on plant fitness. We found no evidence of a large persistent soil seed bank. • Natural corridors are likely to decrease the extinction probability of the populations , most probably by allowing recolonization of empty puddles after extinctions. Therefore, the preservation of corridors appears crucial for the conservation of R. nodiflorus in its natural habitat

LOSS OF GAMETOPHYTIC SELF-INCOMPATIBILITY WITH EVOLUTION OF INBREEDING DEPRESSION

International audienceGametophytic self-incompatibility (SI) in plants is a widespread mechanism preventing self-fertilization and the ensuing inbreeding depression, but it often evolves to self-compatibility. We analyze genetic mechanisms for the breakdown of gametophytic SI, incorporating a dynamic model for the evolution of inbreeding depression allowing for partial purging of nearly recessive lethal mutations by selfing, and accounting for pollen limitation and sheltered load linked to the S-locus. We consider two mechanisms for the breakdown of gametophytic SI: a nonfunctional S-allele and an unlinked modifier locus that inactivates the S-locus. We show that, under a wide range of conditions, self-compatible alleles can invade a self-incompatible population. Conditions for invasion are always less stringent for a nonfunctional S-allele than for a modifier locus. The spread of self-compatible genotypes is favored by extremely high or low selfing rates, a small number of S-alleles, and pollen limitation. Observed parameter values suggest that the maintenance of gametophytic SI is caused by a combination of high inbreeding depression in self-incompatible populations coupled with intermediate selfing rates of the self-compatible genotypes and sheltered load linked to the S-locus