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We dedicate this manuscript to I.O. who passed away before the completion of this study, which she initiated. She inspired and is still inspiring our research in many ways.; International audience; Sexual dimorphism in plants may emerge as a result of sex-specific selection on traits enhancing access to nutritive resources and/or to sexual partners. Here we investigated sex-specific differences in selection of sexually dimorphic traits and in the spatial distribution of effective fecundity (our fitness proxy) in a highly dimorphic dioecious wind-pollinated shrub, Leucadendron rubrum. In particular, we tested for the effect of density on male and female effective fecundity. We used spatial and genotypic data of parent and offspring cohorts to jointly estimate individual male and female effective fecundity on the one hand and pollen and seed dispersal kernels on the other hand. This methodology was adapted to the case of dioecious species. Explicitly modelling dispersal avoids the confounding effects of heterogeneous spatial distribution of mates and sampled seedlings on the estimation of effective fecundity. We also estimated selection gradients on plant traits while modelling sex-specific spatial autocorrelation in fecundity. Males exhibited spatial autocorrelation in effective fecundity at a smaller scale than females. A higher local density of plants was associated with lower effective fecundity in males but was not related to female effective fecundity. These results suggest sex-specific sensitivities to environmental heterogeneity in L. rubrum. Despite these sexual differences, we found directional selection for wider canopies and smaller leaves in both sexes, and no sexually antagonistic selection on strongly dimorphic traits in L. rubrum. Many empirical studies in animals similarly failed to detect sexually antagonistic selection in species expressing strong sexual dimorphism, and we discuss reasons explaining this common pattern. more...

International audience; Most existing forests are subjected to natural and human-mediated selection pressures, which have increased due to climate change and the increasing needs of human societies for wood, fibre and fuel resources. It remains largely unknown how these pressures trigger evolutionary changes. We address this issue here for temperate European oaks (Quercus petraeaandQ. robur), which grow in mixed stands, under even-aged management regimes. We screened numerous functional traits for univariate selection gradients and for expected and observed genetic changes over two successive generations. In both species, growth, leaf morphology and physiology, and defence-related traits displayed significant selection gradients and predicted shifts, whereas phenology, water metabolism, structure and resilience-related traits did not. However, the direction of the selection response and the potential for adaptive evolution differed between the two species.Quercus petraeahad a much larger phenotypic and genetic variance of fitness thanQ. robur. This difference raises concerns about the adaptive response ofQ. roburto contemporary selection pressures. Our investigations suggest thatQ. roburwill probably decline steadily, particularly in mixed stands withQ. petraea, consistent with the contrasting demographic dynamics of the two species. more...

Understanding the ecological and evolutionary processes occurring during species range shifts is important in the current context of global change. Here, we investigate the interplay between recent expansion, gene flow and genetic drift, and their consequences for genetic diversity and structure at landscape and local scales in European beech (Fagus sylvatica L.) On Mont Ventoux, South-Eastern France, we located beech forest refugia at the time of the most recent population minimum, ~150 years ago, and sampled 71 populations (2042 trees) in both refugia and expanding populations over an area of 15,000 ha. We inferred patterns of gene flow and genetic structure using 12 microsatellite markers. We identified six plots as originating from planting, rather than natural establishment, mostly from local genetic material. Comparing genetic diversity and structure in refugia versus recent populations did not support the existence of founder effects: heterozygosity (He = 0.667) and allelic richness (Ar = 4.298) were similar, and FST was low (0.031 overall). Still, significant spatial evidence of colonization was detected, with He increasing along the expansion front, while genetic differentiation from the entire pool (βWT) decreased. Isolation by distance was found in refugia but not in recently expanding populations. Our study indicates that beech capacities for colonization and gene flow were sufficient to preserve genetic diversity despite recent forest contraction and expansion. Because beech has long distance pollen and seed dispersal, these results illustrate a 'best case scenario' for the maintenance of high genetic diversity and adaptive potential under climate-change-related range change. more...

National audience; Recent interest for microgeographic adaptation, i.e. adaptation at spatial scales compatible with substantial amount of gene dispersal, offers to reconsider the scale at which evolution occurs (Richardson et al. 2014). Whether gene flow is constraining or facilitating local adaptation at this fine spatial scale remains an unresolved question. Too important gene flow would overwhelm the effects of natural selection and decrease local adaptation along environmental gradients. Conversely, gene flow, and particularly long-distance dispersal events, could play a major role in resupplying the genetic variation of populations and favouring the spread of beneficial alleles (Kremer et al. 2012). Hence, the high dispersal capacities of trees are often assumed to be the main process maintaining the large levels of genetic variation measured in their natural populations. However, evidences for microgeographic adaptation and the quantitative assessment of the impact of gene flow on adaptive genetic variation are still limited in most temperate trees. Here, we sampled 60 open-pollinated families of European beech (Fagus sylvatica L.) from three natural plots, spreading along a short elevation gradient (∼1.5 km) at the warm margin of this species distribution. We analysed the phenotypic and genotypic data of ∼2,300 seedlings grown in a common garden. We focused on 11 potentially adaptive traits with significant heritabilities (Gauzere et al. 2016) and tested for signature of local selection on quantitative trait differentiation. We then identified the offspring likely originating from local or distant pollen immigration events and quantified the role of gene flow in increasing locally the additive variance of traits under selection. We found a significant signal of adaptive differentiation among plots separated by less than one kilometre, with local selection acting on growth and phenological traits. We found that trees in the plots at high elevation, experiencing the lowest temperature conditions, flushed earlier and had a higher height and diameter growth in our common garden than trees from the plot at low elevation. Beech populations originating from higher longitude or elevation have also been shown to be genetically earlier in provenance tests, suggesting that these populations evolved phenological traits promoting a longer vegetation period. At this southern margin of the species, the reduced allocation to stem growth at the low elevation plot is likely an adaptive response to drought, which has previously been described by comparing marginal vs central beech populations. Consistently with theoretical expectations, our results suggest a beneficial effect of pollen dispersal by increasing the genetic diversity for these locally differentiated traits. These effects were quantitatively high, with more than twice higher genetic variance for immigrant than local offspring, although with large standard errors around estimates. Our results highlight that local selection is an important evolutionary force in natural tree populations, and provide a strong evidence that adaptive genetic differentiation can occur despite high gene flow. For the two genetically differentiated traits, our analyses suggested a beneficial effect of pollen dispersal by increasing genetic diversity after one episode of reproduction. The findings suggest that conservation and management interventions to facilitate movement of gametes along short ecological gradients would boost genetic diversity of individual tree populations, and thereby enhance their adaptive potential. more...

Where and when alien organisms are successfully introduced are central questions to elucidate biotic and abiotic conditions favorable to the introduction, establishment and spread of invasive species. We propose a modelling framework to analyze multiple introductions by several invasive genotypes or genetic variants, in competition with a resident population, when observations provide knowledge on the relative proportions of each variant at some dates and places. This framework is based on a mechanistic-statistical model coupling a reaction-diffusion model with a probabilistic observation model. We apply it to a spatio-temporal dataset reporting the relative proportions of five genetic variants of watermelon mosaic virus (WMV, genus Potyvirus, family Potyviridae) in infections of commercial cucurbit fields. Despite the parsimonious nature of the model, it succeeds in fitting the data well and provides an estimation of the dates and places of successful introduction of each emerging variant as well as a reconstruction of the dynamics of each variant since its introduction. more...

Forest fragmentation may affect mating and pollen dispersal patterns through conversion of continuous forests into small, spatially isolated remnant patches and individual trees in an anthropogenic landscape. We investigated reproductive investment and success, pollen dispersal, mating system, and genetic diversity and spatial structure of Qualea grandiflora trees in two environmental contexts: a continuous natural Cerrado area and isolated individuals on roadsides. Roadside trees produced more flowers and more fruit than Cerrado trees. Pollen dispersal kernels were fat-tailed in both contexts, indicating long-distance dispersal, but in Cerrado the mean pollen dispersal distance (524.7 m) and the effective number of pollen donors per mother-tree (Nep = 12.7) were higher than for roadside trees (60.9 m, Nep = 4.6). The levels and structure of genetic diversity, outcrossing rates ( $$t_{m}$$ > 0.98), and mating among relatives ( $$t_{m} - t_{s}$$ < 0.1) were similar in both environmental contexts. Allelic richness and number of private alleles were similar between the two environments. The fixation index was significantly lower in adults (minimum of 0.08) than in offspring (minimum of 0.23) in both contexts, suggesting selection against inbred individuals between offspring and adult stage. Our results indicate that the spatial isolation of roadside trees, by increasing the number of flowers produced, decreased pollinator movements, thereby reducing effective pollen flow and the number of pollen donors. All these results suggest that roadside trees can be used for harvesting seeds for recovery plans, and that these trees are a biological legacy, and reservoir of Q. grandiflora genetic diversity, from the original Cerrado forest. more...

International audience; Sexual selection may contribute to the evolution of plant sexual dimorphism by favoring architectural traits in males that improve pollen dispersal to mates. In both sexes, larger individuals may be favored by allowing the allocation of more resources to gamete production (a ‘budget’ effect of size). In wind‐pollinated plants, large size may also benefit males by allowing the liberation pollen from a greater height, fostering its dispersal (a ‘direct’ effect of size). To assess these effects and their implications for trait selection, we measured selection on plant morphology in both males and females of the wind‐pollinated dioecious herb Mercurialis annua in two separate experimental common gardens at contrasting density. In both gardens, selection strongly favored males that disperse their pollen further. Selection for pollen production was observed in the high‐density garden only and was weak. In addition, male morphologies associated with increased mean pollen dispersal differed between the two gardens, as elongated branches were favored in the high‐density garden while shorter plants with longer inflorescence stalks were favored in the low‐density garden. Larger females were selected in both gardens. Our results point to the importance of both a direct effect of selection on male traits that affect pollen dispersal, and, to a lesser extent, a budget effect of selection on pollen production. more...

International audience; Cedar is an emblematic Mediterranean forest tree. Genetic research activities on Mediterranean Cedrus species are progressively developed in relation to conservation, restoration, and exploitation plans of these forest tree species. However, currently available molecular biology tools supporting genetic research in these fields are still scarce and have a limited genetic resolution potential. We developed a new set of 12 nuclear microsatellite markers (nSSRs) on Cedrus atlantica that we combined with four previously developed ones in three multiplexes. We checked their monogenic inheritance in controlled crosses. We used a collection of 131 samples from six populations of C. atlantica to estimate null allele frequencies and probability of identity and to characterize the structure of genetic diversity in the fragmented distribution range of this species. We also tested the transferability of the markers to another set of 36 samples from the other Mediterranean Cedrus species and performed a phylogenetic analysis. The three multiplexes reached a high level of resolution potential that we used to evaluate sampling quality. Null allele frequency estimates showed no specific pattern across populations or across species and did not affect the results of biogeographic and phylogenic analyses. Our results reveal a very clear geographical genetic structure within C. atlantica, and the phylogenic tree matched previous analyses based on other markers. Our results confirm the potential interest of these nSSR multiplexes for genotyping in Cedrus. more...

International audience; Understanding how invasive species establish and spread is vital for developing effective management strategies for invaded areas and identifying new areas where the risk of invasion is highest. We investigated the explanatory power of dispersal histories reconstructed based on local-scale wind data and a regionalscale wind-dispersed particle trajectory model for the invasive seed chalcid wasp Megastigmus schimitscheki (Hymenoptera: Torymidae) in France. The explanatory power was tested by: (1) survival analysis of empirical data on M. schimitscheki presence, absence and year of arrival at 52 stands of the wasp’s obligate hosts, Cedrus (true cedar trees); and (2) Approximate Bayesian analysis of M. schimitscheki genetic data using a coalescence model. The Bayesian demographic modeling and traditional population genetic analysis suggested that initial invasion across the range was the result of long-distance dispersal from the longest established sites. The survival analyses of the windborne expansion patterns derived from a particle dispersal model indicated that there was an informative correlation between the M. schimitscheki presence/absence data from the annual surveys and the scenarios based on regional-scale wind data. These three very different analyses produced highly congruent results supporting our proposal that wind is the most probable vector for passive long-distance dispersal of this invasive seed wasp. This result confirms that long-distance dispersal from introduction areas is a likely driver of secondary expansion of alien invasive species. Based on our results, management programs for this and other windborne invasive species may consider (1) focusing effort at the longest established sites and (2) monitoring outlying populations remains critically important due to their influence on rates of spread. We also suggest that there is a distinct need for new analysis methods that have the capacity to combine empirical spatiotemporal field data, genetic data, and environmental data to investigate dispersal and invasion more...

Dispersal ecology is a broad topical discipline that tackles important conceptual and applied issues, such as the study of the ability of plants to transmit their propagules across fragmented and managed landscapes. The relevance of dispersal for plant populations is threefold because: (i) dispersal processes scale from genes and individuals that disperse (or produce propagules to be dispersed) to population dynamics and both local and regional distribution patterns; (ii) by dispersing propagules or individuals and the genes they carry, dispersal inherently links demographic and genetic dynamics across the landscape; and (iii) dispersal elicits key ecological and evolutionary processes that sustain biodiversity, such as species assembly in species-rich communities. The steady improvement of tracking devices and molecular tools that trace the movement or infer provenance of organisms and the pressing need to address conservation issues have expanded the disciplinary boundaries of dispersal ecology. The discussion on the main advances on and challenges for dispersal ecology was the main motivation for the organization of a thematic topic session entitled Dispersal processes driving plant movement: challenges for understanding and predicting range shifts in a changing world at the Annual Meeting of the British Ecological Society (2015, Edinburgh). This session brought together researchers with different types of expertise and interests on dispersal processes and their contributions are now included in this special feature together with a few additional articles. Synthesis. Overall, this special feature illustrates that dispersal ecology spans a broad range of research topics by integrating eight contributions that cover key aspects of this discipline: from conceptual and methodological advances to the study of the ecological and evolutionary outcomes. more...

1. Hybridization is a key evolutionary process with major consequences for conservation and speciation. However, sexual barriers interact with the local context to determine hybridization rates in a way that is still poorly explored. For instance, in the context of an expanding or introduced plant population, where a few individuals are isolated in populations dominated by heterospecific individuals, what is the hybridization potential? 2. To obtain baseline predictions on hybridization rate between two species differing in abundance and to evaluate the effect of pollen limitation, we first used a mean-field model. We then explored a spatially explicit individual-based mating model relying on pollen dispersal kernels to predict hybridization rates in a mixed-species stand, under different scenarios for (i) the strength of sexual barriers; (ii) the spatial distribution of individuals within the site; (iii) the variation in individual fecundity; and (iv) the magnitude, shape and asymmetry of pollen dispersal kernels. 3. Pollen limitation was shown to have the potential to greatly increase hybridization rates. Similarly, fine-scale variation in species composition can result in elevated hybridization rates compared to mean-field predictions, especially with strong sexual barriers. However, species clustering in combination with reduced pollen dispersal has the opposite effect, protecting from hybridization. 4. Synthesis. Our simulation results show that variation of the pollen pool composition at the scale of individuals or stigmas due to spatial configurations or pollen limitation can substantially modify hybridization rates. We explain this by the disproportionate effect of some pollen environments on average hybridization rates. This suggests thoroughly evaluating individual behaviour in terms of hybridization, especially for rare or patchy species. more...

Understanding the determinants of phenotypic variation is critical to evaluate the ability of traits to evolve in a changing environment. In trees, the genetic component of the phenotypic variance is most often estimated based on maternal progeny tests. However, the lack of knowledge about the paternal relatedness hampers the accurate estimation of additive genetic and maternal effects. Here, we investigate how different methods accounting for paternal relatedness allow the estimation of heritability and maternal determinants of adaptive traits in a natural population of Fagus sylvatica L., presenting non-random mating. Twelve potentially adaptive functional traits were measured in 60 maternal families in a nursery. We genotyped a subset of offspring and of all the potentially reproductive adults in the population at 13 microsatellite markers to infer paternal relationships and to estimate average relatedness within and between maternal families. This relatedness information was then used in family and animal models to estimate the components of phenotypic variance. All the studied traits displayed significant genetic variance and moderate heritability. Maternal effects were detected for the diameter increment, stem volume and bud burst. Comparison of family and animal models showed that unbalanced mating system led to only slight departures from maternal family assumptions in the progeny trial. However, neglecting the significant maternal effects led to an overestimation of the heritability. Overall, we highlighted the usefulness of relatedness pattern analyses using polymorphic molecular markers to accurately analyse tree sibling designs. more...

International audience; Inter-individual variation in fecundities has major consequences on population evolutionary potential, through genetic drift and selection. Using two spatially explicit mating models that analyze the genotypes of seeds and seedlings, we investigated the variation of male and female fecundities within and among three European beech (Fagus sylvatica) stands situated along an altitudinal gradient. Female and male individual fecundity distributions were both skewed in this monoecious species, and we found a higher variance in female as compared to male fecundities. Both female and male fecundities increased with tree size and decreased with density and competition in the neighbourhood, the details of these effects suggesting sex-specific strategies to deal with the impact of limited resource on fecundity. The studied populations were functionally male-biased. Among-individual variations in functional gender were not driven by tree size but by density and competition in the neighbourhood. Femaleness decreased under limited resource availability, an expected consequence of the higher cost of female reproduction. Considering the variation of gene flow and genetic drift across elevation, our results suggest that the adaptive potential could be enhanced by low genetic drift at low elevation, and by high pollen-mediated gene flow at high elevation. Finally, this study predicts a more efficient response to selection for traits related to male versus female fitness, for a given selection intensity. more...

Range expansion and contraction has occurred in the history of most species and can seriously impact patterns of genetic diversity. Historical data about range change are rare and generally appropriate for studies at large scales, whereas the individual pollen and seed dispersal events that form the basis of geneflow and colonization generally occur at a local scale. In this study, we investigated range change in Fagus sylvatica on Mont Ventoux, France, using historical data from 1838 to the present and approximate Bayesian computation (ABC) analyses of genetic data. From the historical data, we identified a population minimum in 1845 and located remnant populations at least 200 years old. The ABC analysis selected a demographic scenario with three populations, corresponding to two remnant populations and one area of recent expansion. It also identified expansion from a smaller ancestral population but did not find that this expansion followed a population bottleneck, as suggested by the historical data. Despite a strong support to the selected scenario for our data set, the ABC approach showed a low power to discriminate among scenarios on average and a low ability to accurately estimate effective population sizes and divergence dates, probably due to the temporal scale of the study. This study provides an unusual opportunity to test ABC analysis in a system with a well-documented demographic history and identify discrepancies between the results of historical, classical population genetic and ABC analyses. The results also provide valuable insights into genetic processes at work at a fine spatial and temporal scale in range change and colonization. more...

Negative frequency dependent selection (NFDS) is supposed to be the main force controlling allele evolution at the gametophytic self-incompatibility locus (S-locus) in strictly outcrossing species. Genetic drift also influences S-allele evolution. In perennial sessile organisms, evolution of allelic frequencies over two generations is mainly shaped by individual fecundities and spatial processes. Using wild cherry populations between two successive generations, we tested whether S-alleles evolved following NFDS qualitative and quantitative predictions. We showed that allelic variation was negatively correlated with parental allelic frequency as expected under NFDS. However, NFDS predictions in finite population failed to predict more than half S-allele quantitative evolution. We developed a spatially explicit mating model that included the S-locus. We studied the effects of self-incompatibility and local drift within populations due to pollen dispersal in spatially distributed individuals, and variation in male fecundity on male mating success and allelic frequency evolution. Male mating success was negatively related to male allelic frequency as expected under NFDS. Spatial genetic structure combined with self-incompatibility resulted in higher effective pollen dispersal. Limited pollen dispersal in structured distributions of individuals and genotypes and unequal pollen production significantly contributed to S-allele frequency evolution by creating local drift effects strong enough to counteract the NFDS effect on some alleles. more...

Trees' long lifespan, long-distance dispersal abilities and high year-to-year variability in fecundity are thought to have pervasive consequences for the demographic and genetic structure of recruited seedlings. However, we still lack experimental studies quantifying the respective roles of spatial processes such as restricted seed and pollen dispersal and temporal processes such as mast seeding on patterns of regeneration. Dynamics of European beech (Fagus sylvatica) seedling recruitment was monitored in three plots from 2004 to 2006. Six polymorphic microsatellite genetic markers were used to characterize seedlings and their potential parents in a 7.2-ha stand. These seedlings were shown to result from 12 years of recruitment, with one predominant year of seedling recruitment in 2002 and several years without significant recruitment. Using a spatially explicit mating model based on parentage assignment, short average dispersal distances for seed (δ(s) = 10.9 m) and pollen (43.7 m < δ(p) more...

Summary 1. Characterizing fine scale mating patterns in plant populations makes it possible to investigate genetic drift, gene flow and selection gradients at a contemporary time scale. Molecular markers are valuable tools for this type of analysis, and numerous statistical methods have been developed to make the best of the information they can provide. In particular, we recently proposed a Bayesian approach based on a paternity analysis wherein we estimate jointly the variance in male fecundity and the pollen dispersal kernel. 2. Here, we use simulated data sets to investigate the accuracy of the Bayesian approach compared to (i) classical maximum likelihood approaches (e.g. Neighbourhood model) that ignore variance in male fecundity or explain it through a few covariates and (ii) indirect methods (KinDist and TwoGener) that integrate the variance in fecundity in an ‘effective population density’. 3. The Bayesian estimates correctly considered the over-dispersion resulting from the variance in fecundity, resulting in wider but more accurate confidence intervals, in particular in high-density populations. The maximum likelihood methods resulted in confidence intervals with low coverage probabilities and in widespread false-positive tests when testing the effect of covariates on male fecundity. 4. Estimated individual fecundities and estimated empirical variance in fecundity were robust to the distribution assumed for the individual random fecundities (log-normal or Gamma). In contrast, the theoretical variance estimate critically depended on the assumed distribution. 5. The indirect methods provided much more variable estimators, as expected because they use less information about pollen sources and consider the molecular information only through genetic structure indices. 6. Disentangling the fecundity from the spatial effects in paternity analyses is necessary when studying selection in natura and or when addressing the effects of spatial distribution on effective gene flow. The Bayesian approach studied here successfully accounts for the variance in fecundity when a large fraction of it is not explained by the studied covariates. The Mixed Effect Mating Model computer program introduced here is devoted to its implementation. more...

Summary 1. Interest in pollen-borne gene dispersal has grown with the cultivation of genetically modified plants. To date, both experimental data and models of oilseed rape (OSR) Brassica napus pollen movement at the landscape scale do not clearly differentiate between wind- and insect-mediated dispersal. Estimations of insect-mediated gene dispersal would be valuable for managing potential escapes of transgenes. 2. To quantify the intensity and spatial scale of pollen dispersal by insect pollinators in an agricultural landscape, bait points made of flowering male-sterile OSR that attract OSR pollinators were located at six distances (10–1100 m) from the closest OSR fields and feral populations. Flower-visiting insects were caught by net on these male-sterile flowers and were rubbed onto the stigmas of male-sterile OSR plants grown in a pollen-proof greenhouse to do a manual pollination of their flowers. In this way we were able to assess the insects’ OSR pollen load and seed production at each of the six distances. 3. A large diversity of insects carried OSR pollen and contributed to seed production, but not pollen beetles Meligethes aeneus. Logistic regression analyses of the seed-set success from the manual pollination demonstrated that seed set significantly increased with the proximity of OSR fields, the size of the pollinating insect, and the main daily temperature. Seed set was not affected by the pollinating insect’s order or genus. Seed set, both observed and predicted by the model, was above zero for flowers pollinated with large bees caught at >1100 m from the nearest OSR field. 4. Synthesis and applications. Our study provides clear evidence that a large variety of insect species can transfer viable pollen between OSR plants over considerable distances. However, only 39AE4% of the insects caught on male-sterile flowers carried OSR pollen. Our results provide valuable data to improve models of pollen dispersal for entomophilous crops at the landscape scale. These models are essential to help land-managers reduce pollen-borne gene dispersal from genetically modified plants to wild and cultivated relatives. more...

The comparison between estimates of historical gene flow, using variance in allelic frequencies, and estimates of contemporary gene flow, using parentage assignment, is expected to provide insights into ecological and evolutionary processes at work within and among populations. Genetic variation at microsatellite loci was used to quantify genetic structure in two wind pollinated, gravity and animal-dispersed tree species (Fagus sylvatica L. and Fagus crenata Blum.) and to derive historical estimates of gene flow. The gene dispersal distances estimated assuming effective population density to be 1/4 of the observed density were ∼77 m in European beech and ∼40 m in Japanese beech. Parentage analyses and a neighbourhood model approach were used to estimate contemporary patterns of seed and pollen dispersal. Our results suggest restricted seed dispersal abilities in both European beech (δs = 10.5 m) and Japanese beech (δs = 12.4 m), with an exponential shaped seed dispersal kernel. A non-negligible rate of seed immigration (ms = 27%) was detected in European beech sites but not in Japanese beech site. Pollen dispersal within studied sites also appeared limited (δp = 41.63 m in European beech and δp = 79.4 m in Japanese beech), despite high rate of pollen immigration (mp = 68% in European beech and mp = 40% m in Japanese beech). Interestingly, contemporary and historical estimates of gene flow were within the same order of magnitude (a few tens of meters). more...

International audience; There is an increasing recognition that long distance dispersal (LDD) plays a key role in establishing spatial genetic structure during colonization. Recent works, focused on short distance dispersal, demonstrated that a neutral mutation arising at the colonization front can either 'surf' with the wave front and reach high frequencies or stay near its place of origin at low frequencies. Here, we examine how LDD, and more generally the shape of the dispersal kernel, modifies this phenomenon and how the width of the colonization corridor affects the fate of the mutation. We demonstrate that when LDD events are more frequent, the 'surfing phenomenon' is less frequent, probably because any alleles can get far ahead from the colonization front and preclude the invasion by others alleles, thus leading to an attenuation of the diversity loss. We also demonstrate that the width of the colonization corridor influences the fate of the mutation, wide spaces decreasing the probability of invasion. Overall, the genetic structure of diversity resulted not only from LDD but also particularly from the shape of the dispersal kernel more...

Interindividual variance of male reproductive success (MRS) contributes to genetic drift, which in turn interacts with selection and migration to determine the short-term response of populations to rapid changes in their environment. Individual relative MRS can be estimated through paternity analysis and can be further dissected into fecundity and spatial components. Existing methods to achieve this decomposition either rely on the strong assumption of a random distribution of pollen donors (TwoGener) or estimate only the part of the variance of male fecundity that is explained by few covariates. We developed here a method to estimate jointly the whole variance of male fecundity and the pollen dispersal curve from the genotypic information of sampled seeds and their putative fathers and geographical information of all individuals in the study area. We modelled the relative individual fecundities as a log-normally distributed random effect. We used a Bayesian approach, well suited to the hierarchical nature of the model, to estimate these fecundities. When applied to Sorbus torminalis, the estimated variance of male fecundity corresponded to an effective density of trees 13 times lower than the observed density (d(obs)/d(ep ) approximately 13). This value is between the value (approximately 2) estimated with a classical mating model including three covariates (neighbourhood density, diameter, flowering intensity) that affect fecundity and the value (approximately 30) estimated with TwoGener. The estimated dispersal kernel was close to previous results. This approach allows fine monitoring of ongoing genetic drift in natural populations, and quantitative dissection of the processes contributing to drift, including human actions. more...

The comparison between historical estimates of gene flow, using variance in allelic frequencies, and contemporary estimates of gene flow, using parentage assignment, is expected to provide insights into ecological and evolutionary processes at work within and among populations. Genetic variation at six microsatellite loci was used to quantify genetic structure in the insect-pollinated, animal-dispersed, low-density tree Sorbus torminalis L. Crantz, and to derive historical estimates of gene flow. The neighbourhood size and root-mean-squared dispersal distance inferred from seedling genotypes (Nb = 70 individuals, σe = 417 m) were similar to those inferred from adult genotypes (Nb = 114 individuals, σe = 472 m). We also used parentage analyses and a neighbourhood model approach after an evaluation of the statistical properties of this method on simulated data. From our data, we estimated even contributions of seed- and pollen-mediated dispersal to the genetic composition of established seedlings, with both fat-tailed pollen and seed dispersal kernels, and slightly higher mean distance of pollen dispersal (248 m) as compared to seed dispersal (135 m). The resulting contemporary estimate of gene dispersal distance (σc = 211 m) was ∼twofold smaller than the historical estimates. Besides different assumptions and statistical nuances of both approaches, this discrepancy is likely to reflect a recent restriction in the scale of gene flow which requires manager's attention in a context of increasing forest fragmentation. more...

The genetic consequences of range expansions have generally been investigated at wide geographical and temporal scales, long after the colonization event. A unique ecological system enabled us to both monitor the colonization dynamics and decipher the genetic footprints of expansion over a very short time period. Each year an epidemic of the poplar rust (Melampsora larici‐populina) expands clonally and linearly along the Durance River, in the Alps. The colonization dynamics observed in 2004 showed two phases with different genetic outcomes. Upstream, fast colonization maintained high genetic diversity. Downstream, the colonization wave progressively faltered, diversity eroded, and differentiation increased, as expected under recurrent founder events. In line with the high dispersal abilities of rust pathogens, we provide evidence for leapfrog dispersal of clones. Our results thus emphasize the importance of colonization dynamics in shaping spatial genetic structure in the face of high gene flow. more...

Summary 1 Evaluation of our ability to predict cross-pollination rates (CPR) at long distance using dispersal functions estimated in experimental set-ups differing by either their scale or their environmental conditions is crucial to establish appropriate management rules following the release of genetically modified (GM) crops. 2 From 1998 to 2004 we measured oilseed rape CPR in commercial fields for 44 donor-recipient couples separated by 220 up to 2000 m, from up to three different pollen donor cultivars in two French oilseed rape production areas. We followed the same sampling and screening designs and tested the effect of region, year, size, cultivar and distance on the observed CPR. 3 We then compared observed CPR to predictions from six empirical pollen dispersal models based on dispersal kernels that were fitted previously at the local and landscape scales. These predictions allowed us to test the possibility to extrapolate and up-scale dispersal kernels. 4 The observed CPR varied from 0% to 0·092%. They were higher in Champagne-Ardennes, where they depended negatively on distance, than in Bourgogne where they did not depend on distance. CPR also differed among years, being nil for the last 3 successive years, partly because of different environmental conditions and detection issues. CPR depended further on the source cultivars due to differences in pollen production. Dispersal kernels fitted at the local scale lead to systematic and huge underestimation of CPR, when observed. The power-law kernel fitted at the landscape scale under-estimated CPR by two orders of magnitude above 400 m but followed the rate of decrease of the observed pollination with distance. 5 Synthesis and applications. Caution should be taken when extrapolating and up-scaling dispersal kernels and models because predictions may differ greatly from observations. Models should rely at least upon dispersal kernels estimated from landscape-scale data obtained in different regions. Models should also integrate several phenomena at the agro-ecosystem scale including the dispersal by insects and become more mechanistic to account for variations observed among years, environments and distance to borders. more...

Understanding patterns of pollen movement at the landscape scale is important for establishing management rules following the release of genetically modified (GM) crops. We use here a mating model adapted to cultivated species to estimate dispersal kernels from the genotypes of the progenies of male-sterile plants positioned at different sampling sites within a 10 x 10-km oilseed rape production area. Half of the pollen clouds sampled by the male-sterile plants originated from uncharacterized pollen sources that could consist of both large volunteer and feral populations, and fields within and outside the study area. The geometric dispersal kernel was the most appropriate to predict pollen movement in the study area. It predicted a much larger proportion of long-distance pollination than previously fitted dispersal kernels. This best-fitting mating model underestimated the level of differentiation among pollen clouds but could predict its spatial structure. The estimation method was validated on simulated genotypic data, and proved to provide good estimates of both the shape of the dispersal kernel and the rate and composition of pollen issued from uncharacterized pollen sources. The best dispersal kernel fitted here, the geometric kernel, should now be integrated into models that aim at predicting gene flow at the landscape level, in particular between GM and non-GM crops. more...

Summary 1. Debate continues regarding the ecological impacts of genetically modified (GM) crops and their coexistence with non-GM crops in Europe. In this debate, quantitative predictions of gene dispersal by pollen are necessary, and as a result numerous plot-to-plot gene flow experiments have been performed with various crops. However, plot-to-plot cross-pollination rates (CPR) depend on spatial configuration of plots, implying that (i) they are difficult to compare among experiments and (ii) functions directly fitted on CPR data are inappropriate for predictions in other spatial contexts. 2. Modelling pollen dispersal via an individual dispersal function (IDF) circumvents these problems by accounting for spatial designs. We detail for oilseed rape how this approach can be used to both estimate an IDF from field data and predict CPR between two neighbouring fields of various sizes and shapes. Predictions were used to investigate the sensitivity of CPR to the family of IDF, the uncertainty in parameter estimates and the effects of field dimensions and isolation distances. 3. We fitted a range of families of IDF, including several types of tails, on previously published data. The best IDF was a fat-tailed power-law function, meaning frequent long-distance dispersal. 4. The choice of IDF appeared crucial when predicting CPR between fields, occasionally being even more important than the distance between fields. Width of the source field and depth of the recipient field were next in importance. When approximated CPR were calculated without considering field dimensions, using distance between field centres gave better performance than field margins. 5. Synthesis and applications. This study demonstrates the value of IDF for quantitative predictions of pollen flow in variable spatial configurations. A spatially explicit model of agro-ecosystems used to define management rules for the commercial release of GM crops in Europe already employs IDF but underestimates long-distance dispersal for oilseed rape. These new parameter estimates will refine the performance of these models. Moreover, the detailed guidelines for estimating an IDF should encourage such statistical analysis of other dispersal data, enabling comparisons of dispersal data obtained for different environments and species and providing new IDF for management models. more...

Estimating the frequency of long-distance pollination is important in cultivated species, particularly to assess the risk of gene transfer following the release of genetically modified crops. For this purpose, we estimated the diversity and origin of fertilizing pollen in a 10 x 10 km French oilseed rape production area. First, the cultivar grown in each field was identified through surveys to farmers and using microsatellite markers. Examination of the seed set in fields indicated high rates of seed contamination (8.7%) and pollination from other sources (5%). Then, male-sterile plants were scattered over the study area and their seed genotyped using the same markers. Most pollination was local: 65% of the seeds had a compatible sire in the closest field, i.e. at 50 or 300 m depending on site, but the nearest compatible field was found more than 1000 m away for 13% of the seeds. To assess the diversity of fertilizing pollen, each seed was assigned to the nearest putative siring cultivar. The observed diversity of pollen was then compared to that predicted by simulations using three empirical dispersal models with increasing proportion of long-distance pollination. The diversity was sensitive to the dispersal kernel used in the simulations, fatter-tailed functions predicting higher diversities. The dispersal kernel that was more consistent with our data predicted more long-distance dispersal than the exponential function. more...

International audience; Abstract Key message With a retrospective growth analysis over 20 years, we showed that silviculture (in particular thinning) ofCedrus atlantica(Manetti) could to some extent mitigate the effects of drought episodes. Severe reductions of density resulted in faster recovery of radial growth after severe drought episodes. Context Climate change is expected to increase the frequency and the intensity of drought in southern Europe. Silviculture could help in mitigating the effects of water deficit on forests. Aim The aim of this study is to assess the effect of thinning intensity on the climate-growth relationship of different crown classes in Cedrus atlantica (Manetti). Material and methods A 20-year annual survey was conducted on plots managed under contrasted thinning regimes in a site of the southern French Alps. Using a linear mixed modeling approach, we evaluated the modulation of the growth response to annual climate by thinning (climate × thinning interaction effect) during the experiment. The effect of thinning on the growth sensitivity to annual climate and the effect of thinning on the growth response to pointer years were also quantified. Results The highest intensity of thinning significantly changed the climate-growth relationship of the two most dominant crown classes. Heavy thinning reduced the impact of negative pointer years during a period of 4–5 years and improved the post-drought recovery for a decade. Heavily thinned plots and dominant crown classes had significantly higher relative growth sensitivities to annual climate over the studied period. Conclusions Forest management has the potential to reduce the effect of water deficit on the growth of Atlas cedar stands. We provide keystones for future silvicultural guidelines, in the context of a larger introduction of C. atlantica in the French territory. more...

Having a precise knowledge of the dispersal ability of a population in a heterogeneous environment is of critical importance in agroecology and conservation biology as it can provide management tools to limit the effects of pests or to increase the survival of endangered species. In this paper, we propose a mechanistic-statistical method to estimate space-dependent diffusion parameters of spatially-explicit models based on stochastic differential equations, using genetic data. Dividing the total population into subpopulations corresponding to different habitat patches with known allele frequencies, the expected proportions of individuals from each subpopulation at each position is computed by solving a system of reaction-diffusion equations. Modelling the capture and genotyping of the individuals with a statistical approach, we derive a numerically tractable formula for the likelihood function associated with the diffusion parameters. In a simulated environment made of three types of regions, each associated with a different diffusion coefficient, we successfully estimate the diffusion parameters with a maximum-likelihood approach. Although higher genetic differentiation among subpopulations leads to more accurate estimations, once a certain level of differentiation has been reached, the finite size of the genotyped population becomes the limiting factor for accurate estimation. more...

Understanding precisely how plants disperse their seeds and pollen in their neighbourhood is a central question for both ecologists and evolutionary biologists because seed and pollen dispersal governs both the rate of spread of an expanding population and gene flow within and among populations. The concept of a ‘dispersal kernel’ has become extremely popular in dispersal ecology as a tool that summarizes how dispersal distributes individuals and genes in space and at a given scale. In this issue of Molecular Ecology, the study by Moran & Clark (2011) (M&C in the following) shows how genotypic and spatial data of established seedlings can be analysed in a Bayesian framework to estimate jointly the pollen and seed dispersal kernels and finally derive a parentage analysis from a full-probability approach. This approach applied to red oak shows important dispersal of seeds (138 m on average) and pollen (178 m on average). For seeds, this estimate contrasts with previous results from inverse modelling on seed trap data (9.3 m). This research gathers several methodological advances made in recent years in two research communities and could become a cornerstone for dispersal ecology. more...

Conditions of persistence or extinction of a metapopulation of a colonizing annual species are studied in a heterogeneous landscape, a mixture of two elementary landscapes. An elementary landscape is a landscape whose age-structure is described by only one transition matrix, giving the probability for a site to be disturbed, or to follow the process of succession. We first provide an analytical study of the range of dispersal rates that allow metapopulation persistence in an elementary landscape. Second, conditions for metapopulation persistence in a heterogeneous landscape are derived from results obtained in each elementary landscape. Three cases are distinguished. If the two ranges of dispersal rates defined in each elementary landscape overlap, the metapopulation persists in any mixture of the elementary landscapes. If these two dispersal rates ranges are non-overlapping, either the metapopulation goes extinct for some values of the proportion of the elementary landscapes, or two discontinuous ranges of dispersal rates allow the metapopulation persistence. The consequences of these results are discussed in terms of landscape management. In particular, it is shown that under some conditions, a rapid change in environment (from one elementary landscape to another one) might less often lead to metapopulation extinction than a slower change. Copyright 1999 Academic Press. more...

International audience; Studies addressing the variation of mating system between plant populations rarely account for the variability of these parameters between individuals within populations, although this variability is often non-negligible. Here, we propose a new direct method based on paternity analyses (Mixed Effect Mating Model) to estimate individual migration (m(i)) and selfing rates (s(i)) together with the pollen dispersal kernel. Using this method and the KINDIST approach, we investigated the variation of mating system parameters within and between three populations of Fagus sylvatica along an elevational gradient. Among the mother trees, s(i) varied from 0% to 48%, m(i) varied from 12% to 86% and the effective number of pollen donors (Nep(i)) varied from 2 to 364. The mating patterns differed along the gradient, the top population showing higher m and lower s, and a trend to higher Nep than the bottom populations. The phenological lag shaped long-distance pollen flow both within population (by increasing m(i) at mother-tree level) and between populations (by increasing m at high elevation). Rather than the mate density, the canopy density was detected as a major mating system determinant within population; it acted as a barrier to pollen flow, decreasing the proportion of long-distance pollen flow and increasing s(i). Overall, the effects of ecological factors on mating system were not the same within vs. between populations across the gradient, and these factors also differed from those traditionally found to shape variation at range-wide scale, highlighting the interest of multiscale approaches. more...

International audience; Widespread ecosystem change has led to declines in species world-wide. The loss of pollinators in particular constitutes a problem for ecosystem function and crop production. Understanding how landscape change affects pollinator movement, effective pollen flow, and plant and pollinator survival is therefore a global priority. In this study we investigated patterns of effective pollen flow, using wild cherry tree (Prunus avium) progeny arrays, to address two questions in three case studies: Do land-use types present different resistances to pollinator movement? Which pollinator travel path best explains the pollination data (straight lines, weighted straight lines, least cost paths or pair-wise resistance)? Trees and progeny arrays were genotyped and effective pollen flow and pollinator movement were estimated using the spatially explicit mating model. We found that pollinators did modify their travel paths in response to land-use type and arrangement, but the travel path that best described pollinator movement and the resistance rank of the land uses depended on the type and size of land-use patches and the landscape context. We propose a novel theoretical framework rooted in behavioural ecology, the resource model, for interpreting pollinator behaviour in heterogeneous landscapes. We conclude by discussing the importance and practicality of conservation and management strategies in which native and non-native land-use types together provide functional habitat and support ecosystem services across economic landscapes. more...

International audience; In plant quantitative genetic studies conducted ex situ, the large number of seeds produced per individual has promoted the use of open-pollinated progeny tests. In subsequent analyses, seeds collected on the same mother-plant are assumed to be half-sibs. The consequences of the departure from half-sib assumption in progeny tests have been investigated since the 1960s using simulation approaches and, more recently, using molecular-based experimental approaches. This review aims to synthesize the results and conclusions of these simulation and empirical studies. We focus on tree species, where controlled crosses are difficult to carry out experimentally and departures from half-sib assumptions occur frequently in natural populations. First, the average level of relatedness expected within maternal progeny for many tree populations is higher than that of half-sibs. This is the consequence of non-random mating resulting from the small number of effective pollen donors per female, unequal male reproductive success and/or selfing. As result, estimates of genetic variance and heritability for quantitative traits may be upward biased. Alternatively, inbreeding depression, dominance effects and the heterogeneity of the male gamete pool among females are often neglected, which may lead to underestimation of the heritability of traits. A correction based on the mean genetic relatedness between offspring and the relatedness between parents is often used to compensate those biases. However, such correction cannot accurately adjust the estimates in situations where variable levels of genetic relatedness among families, dominance effects or inbreeding depression exist within the progeny. An alternative and promising approach is the use of the "animal model" approach, which optimizes the use of molecular data and paternal information to estimate heritability more accurately. more...

International audience; A major challenge in population ecology of long-lived plants is to disentangle the intricate effects of seed dispersal and seedlings mortality on spatial and genetic structure of recruited seedlings. A common way to address this issue is to compare dispersal patterns among seeds (i.e. basic dispersal) and established seedlings (i.e. effective dispersal). High density-dependent mortality near adults is expected to shift the mode of the effective dispersal distance away from the mother-plant relative to the mode of the basic dispersal distance. Here, we proposed a new approach to address this issue by using seedlings cohorts of different age and by combining parentage analyses to retrieve effective seed dispersal and Spatially Explicit Mating Models (SEMM) to estimate basic seed dispersal. We applied this approach in two seedling cohorts at each of two plots of Fagus sylvatica L., relying on adult and seedling genotypes at 10 microsatellite loci. Basic seed dispersal was primarily local, with mean distances of 12-25 m (young cohort) and 22-25 m (old cohort). The effective seed dispersal distance increased between young and old cohorts from 33 m to 48 m at site I and from 23 m to 33 m at site 2. The hypothesis of density-dependent mortality was supported at site 2 but not at site 1 possibly because of different ecological conditions among sites. Our results shed light on the interplay between seed dispersal and density-dependent mortality in F. sylvatica. Moreover, the new approach proposed here alleviates general problems associated to synchronic studies comparing spatial patterns across different-age cohorts at a single point in time. more...

International audience; 1. The seed dispersal kernel is a major determinant of spatial population dynamics and spatial distribution of genetic diversity. Among the main methods to estimate it, inverse modelling (IM) and gene shadow model (GSM) rely on seed counts in traps, whereas competing source model (CSM) and spatially explicit mating models (SEMMs) rely on compositions of seed pools. Moreover, GSM, CSM and SEMM exploit genetic information from molecular markers, whereas IM only exploits seed counts ignoring seed origins. These methods were also applied to established seedlings. 2. In the presence of post-dispersal density-dependent mortality (DDM), the effective dispersal kernel, describing the spatial distribution of established seedlings relatively to the seed source, is notoriously different from the basic dispersal kernel, describing the spatial distribution of seed deposition sites relatively to the source. Using simulated data sets, we investigated whether IM, GSM, CSM and SEMM applied to established seedlings estimate the basic or the effective dispersal kernel. In our simulations, DDM resulted in a shift of the mean basic dispersal distance (10m) towards substantially higher effective mean dispersal distances (15m and 208m). 3. We demonstrated that CSM and SEMM estimate the basic seed dispersal kernel, independently from the presence of post-dispersal mortality. By contrast, GSM estimates the effective dispersal kernel. IM failed to provide satisfactory estimates in the presence of DDM in our sampling design. Besides, for all methods, seed migration was inflated in the presence of DDM, due to lower mortality among randomly distributed immigrants relatively to local seedlings. 4. It could seem intuitive that estimates based on seedlings or seeds provide effective or basic dispersal kernels respectively. Our results showed that it is not true for estimates obtained with CSM or SEMM because they rely on the composition of seed/seedling pools and not seed/seedling counts such as IM or GSM. This has important consequences for life stage studies where the discordance of dispersal kernels estimated from different cohorts is used to investigate post-dispersal density-dependent mortality. more...

International audience; Natural hybridization is attracting much interest in modern speciation and conservation biology studies, but the underlying mechanisms remain poorly understood. In particular, it is unclear why environmental changes often increase hybridization rates. To study this question, we surveyed mating events in a mixed oak stand and developed a spatially explicit individual-based hybridization model. This model, where hybridization is frequency-dependent, pollen is nonlimiting and which allows immigrant pollen to compete with local pollen, takes into account species-specific pollen dispersal and sexual barriers to hybridization. The consequences of pollen limitation on hybridization were studied using another simple model. The results indicate that environmental changes could increase hybridization rates through two distinct mechanisms. First, by disrupting the spatial organization of communities, they should decrease the proportion of conspecific pollen available for mating, thus increasing hybridization rates. Second, by decreasing the density of conspecifics, they should increase pollen limitation and thus hybridization rates, as a consequence of chance pollination predominating over deterministic pollen competition. Altogether, our results point to a need for considering hybridization events at the appropriate level of organization and provide new insights into why hybridization rates generally increase in disturbed environments. more...

International audience; Variation among individuals in reproductive success is advocated as a major process driving evolution of sexual polymorphisms in plants, such as gynodioecy where females and hermaphrodites coexist. In gynodioecious Beta vulgaris ssp. maritima, sex determination involves cytoplasmic male sterility genes (CMS) and nuclear restorers of male fertility. Both restored CMS and non-CMS hermaphrodites co-occur. Genotype-specific differences in male fitness are theoretically expected to explain the maintenance of cytonuclear polymorphism. * Using genotypic information on seedlings and flowering plants within two metapopulations, we investigated whether male fecundity was influenced by ecological, phenotypic and genetic factors, while taking into account the shape and scale of pollen dispersal. * Along with spatially restricted pollen flow, we showed that male fecundity was affected by flowering synchrony, investment in reproduction, pollen production and cytoplasmic identity of potential fathers. Siring success of non-CMS hermaphrodites was higher than restored CMS hermaphrodites. However, the magnitude of difference in fecundity depended on the likelihood of carrying restorer alleles for non-CMS hermaphrodites. Our results suggest the occurrence of a cost of silent restorers, a condition supported by scarce empirical evidence, but theoretically required to maintain a stable sexual polymorphism in gynodioecious species. more...

International audience; Most mathematical studies on expanding populations have fo-cused on the rate of range expansion of a population. However, the genetic consequences of population expansion remain an under-studied body of theory. Describing an expanding population as a traveling wave solution derived from a classical reaction-diffusion model, we analyze the spatio-temporal evolution of its genetic structure. We show that the presence of an Allee effect (i.e., a lower per capita growth rate at low densities) drastically modifies genetic diversity, both in the colonization front and behind it. With an Allee effect (i.e., pushed colonization waves), all of the genetic diversity of a population is conserved in the colonization front. In the absence of an Allee effect (i.e., pulled waves), only the furthest forward members of the initial population persist in the coloniza-tion front, indicating a strong erosion of the diversity in this population. These results counteract commonly held notions that the Allee effect generally has adverse consequences. Our study contributes new knowledge to the surfing phenomenon in continuous models without random genetic drift. It also provides insight into the dynamics of traveling wave solutions and leads to a new interpretation of the mathematical notions of pulled and pushed waves. R more...

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International audience; Inbreeding depression is a key factor affecting the persistence of natural populations, particularly when they are fragmented. In species with mixed mating systems, inbreeding depression can be estimated at the population level by regressing the average progeny fitness by the selfing rate of their mothers. We applied this method using simulated populations to investigate how population genetic parameters can affect the detection power of inbreeding depression. We simulated individual selfing rates and genetic loads from which we computed fitness values. The regression method yielded high statistical power, inbreeding depression being detected as significant (5 % level) in 92 % of the simulations. High individual variation in selfing rate and high mean genetic load led to better detection of inbreeding depression while high among-individual variation in genetic load made it more difficult to detect inbreeding depression. For a constant sampling effort, increasing the number of progenies while decreasing the number of individuals per progeny enhanced the detection power of inbreeding depression. We discuss the implication of among-mother variability of genetic load and selfing rate on inbreeding depression studies. more...

Forest trees are the dominant species in many parts of the world and predicting how they might respond to climate change is a vital global concern. Trees are capable of long-distance gene flow, which can promote adaptive evolution in novel environments by increasing genetic variation for fitness. It is unclear, however, if this can compensate for maladaptive effects of gene flow and for the long-generation times of trees. We critically review data on the extent of long-distance gene flow and summarise theory that allows us to predict evolutionary responses of trees to climate change. Estimates of long-distance gene flow based both on direct observations and on genetic methods provide evidence that genes can move over spatial scales larger than habitat shifts predicted under climate change within one generation. Both theoretical and empirical data suggest that the positive effects of gene flow on adaptation may dominate in many instances. The balance of positive to negative consequences of gene flow may, however, differ for leading edge, core and rear sections of forest distributions. We propose future experimental and theoretical research that would better integrate dispersal biology with evolutionary quantitative genetics and improve predictions of tree responses to climate change. © 2012 Blackwell Publishing Ltd/CNRS. more...

Estimation of demographic parameters is important for understanding the functioning of natural populations and the underlying ecological and evolutionary processes that may impact their dynamics. Here, we used sibship assignment methods to shed light on the local dynamics of codling moth females in eight orchards in a 90-ha domain near Valence, France. Based on full-sib inference among 1,063 genotyped moths, we estimated (1) the effective number of females that had offspring, (2) their fertility and (3) the distribution of their oviposition sites within and among orchards. The average number of females in all the orchards increased between the first (~130) and the second (~235) annual generations. The average fertilities of the females were similar at each generation according to the host plant considered (apple, pear, or walnut), but differed between commercial (~10) and non-treated (~25) apple orchards. Females mainly clustered their eggs on contiguous trees along orchard borders, but they also occasionally dispersed their eggs among different orchards independently of the cultivated host plants or the inter-orchard distances (up to 698 m) during the second annual generation. The mean distance between two oviposition sites was 30 m. Sibship estimates of both the effective number of females and the inter-orchard migration rates (~5%) were in agreement with the observed genetic differentiation among the eight orchards (0.006 more...

International audience; Diffusion is one of the most frequently used assumptions to explain dispersal. Diffusion models and in particular reaction–diffusion equations usually lead to solutions moving at constant speeds, too slow compared to observations. As early as 1899, Reid had found that the rate of spread of tree species migrating to northern environments at the beginning of the Holocene was too fast to be explained by diffusive dispersal. Rapid spreading is generally explained using long distance dispersal events, modelled through integro-differential equations (IDEs) with exponentially unbounded (EU) kernels, i.e. decaying slower than any exponential. We show here that classical reaction–diffusion models of the Fisher–Kolmogorov–Petrovsky–Piskunov type can produce patterns of colonisation very similar to those of IDEs, if the initial population is EU at the beginning of the considered colonisation event. Many similarities between reaction–diffusion models with EU initial data and IDEs with EU kernels are found; in particular comparable accelerating rates of spread and flattening of the solutions. There was previously no systematic mathematical theory for such reaction–diffusion models with EU initial data. Yet, EU initial data can easily be understood as consequences of colonisation–retraction events and lead to fast spreading and accelerating rates of spread without the long distance hypothesis. more...

We investigated the inter-individual variation of phenotypic plasticity and its evolution across three generations within an expanding forest. Plasticity was assessed in situ from dendrochronological data as the response of radial growth to summer rainfall. A linear mixed model was used to account for spatial effects (environment and stand structure), temporal factors (stand dynamics) and the variation with age. Beyond these effects, our results reveal a significant inter-individual variance of growth and plasticity within each generation. We also show that the mean values and variances of growth and plasticity changed significantly across generations, with different patterns for both traits. The possible environmental and genetic drivers of these changes are discussed. Contrasting with the trade-off between stress tolerance and plasticity generally observed among populations, we detected a positive covariance at the individual level, which does not support the cost of plasticity hypothesis in this case. more...

International audience; Pollen-mediated gene flow has important implications for biodiversity conservation and for breeders and farmers’ activities. In sugar beet production fields, a few sugar beet bolters can produce pollen as well as be fertilized by wild and weed beet. Since the crop, the wild beets, and the weed beets are the same species and intercross freely, the question of pollen flow is an important issue to determine the potential dispersal of transgenes from field to field and to wild habitats. We report here an experiment to describe pollen dispersal from a small herbicide-resistant sugar beet source towards male sterile target plants located along radiating lines up to 1,200 m away. Individual dispersal functions were inferred from statistical analyses and compared. Pollen limitation, as expected in root-production fields, was confirmed at all the distances from the pollen source. The number of resistant seeds produced by bait plants best fitted a fat-tailed probability distribution curve of pollen grains (power–law) dependent on the distance from the pollen source. A literature survey confirmed that power–law function could fit in most cases. The b coefficient was lower than 2. The number of fertilized flowers by background (herbicide-susceptible) pollen grains was uniform across the whole field. Airborne pollen had a fertilization impact equivalent to that of one adjacent bolter. The individual dispersal function from different pollen sources can be integrated to provide the pollen cloud composition for a given target plant, thus allowing modeling of gene flow in a field, inter-fields in a small region, and also in seed-production area. Long-distance pollen flow is not negligible and could play an important role in rapid transgene dispersal from crop to wild and weed beets in the landscape. The removing of any bolting, herbicide-resistant sugar beet should be compulsory to prevent the occurrence of herbicide-resistant weed beet, thus preventing gene flow to wild populations and preserving the sustainable utility of the resistant varieties. Whether such a goal is attainable remains an open question and certainly would be worth a large scale experimental study. more...

International audience; Frequency-dependent selection is a major force determining the evolutionary dynamics of alleles at the self-incompatibility locus (S-locus) in flowering plants. We introduce a general method using numerical simulations to test several alternative models of frequency-dependent selection on S-locus data from sporophytic systems, taking into account both genetic drift and observed patterns of dominance interactions among S-locus haplotypes (S-haplotypes). Using a molecular typing method, we estimated S-haplotype frequencies in a sample of 322 adult plants and of 245 offspring obtained from seeds sampled on 22 maternal plants, collected in a single population of Arabidopsis halleri (Brassicaceae).We found eight different S-haplotypes and characterized their dominance interactions by controlled pollinations. We then compared the likelihood of different models of frequency-dependent selection: we found that the observed haplotype frequencies and observed frequency changes in one generation best fitted a model with (1) the observed dominance interactions and (2) no pollen limitation. Overall, our population genetic models of frequency-dependent selection, including patterns of dominance interactions among S-haplotypes and genetic drift, can reliably predict polymorphism at the S-locus. We discuss how these approaches allow detecting additional processes influencing the evolutionary dynamics of the S-locus, such as purifying selection on linked loci. more...

New concerns about crop coexistence in agricultural landscapes are being expressed in reaction to the prospect of introducing transgenic crops into European cropping systems: these include meeting current consumer demand for non-GM products, respecting threshold levels required for organic farming labels as well as keeping food cultures separated from those destined for the pharmaceutical and energy industries. To address these concerns in the case of maize crops, we have chosen a modelling approach. Our aim was to simulate cross-pollination in the case of existing agricultural landscapes, taking into account the effect of climate and cropping techniques in order to forecast gene escape from genetically modified maize to non-GM maize. The resulting spatially explicit model, MAPOD (Matricial Approach to POllen Dispersal), is presented in this paper. A preliminary evaluation is also provided. Pollen exchanges between GM and non-GM maize crops are simulated and influencing factors such as field sizes and shapes, distribution of GM and non-GM fields in the agricultural landscape as well as flowering dates and dynamics are integrated. Model parameter values were either derived from existing models of pollen dispersal or estimated from experimental field studies. The preliminary evaluation of MAPOD was carried out by comparing simulation results with data from two French and one American gene flow field trials. MAPOD was found to provide good average predictive values. Examples of output data illustrate the capacity of the model to simulate a wide range of agricultural contexts. These simulation results provide a basis for designing coexistence rules and monitoring procedure set-up. more...

http://www3.interscience.wiley.com/cgi-bin/fulltext/120696197/HTMLSTART; International audience; It is well established that pollen-mediated gene flow among natural plant populations depends on a complex interaction between the spatial distribution of pollen sources and the short- and long-distance components of pollen dispersal. Despite this knowledge, spatial isolation strategies proposed in Europe to ensure the harvest purity of conventional crops are based on distance from the nearest genetically modified (GM) crop and on empirical data from two-plot experiments. Here, we investigate the circumstances under which the multiplicity of pollen sources over the landscape should be considered in strategies to contain GM crops. We simulated pollen dispersal over eighty 6 × 6 km simulated landscapes differing in field characteristics and in amount of GM and conventional maize. Pollen dispersal was modelled either via a Normal Inverse Gaussian (NIG, currently used for European coexistence studies) or a bivariate Student (2Dt) kernel. These kernels differ in their amount of short- and long-distance dispersal. We used linear models to analyse the impact of local and landscape variables on impurity rates (i.e. proportion of seeds sired by pollen from a transgenic crop) in conventional fields and quantified their increase due to dispersal from other than the closest GM crops. The average impurity rate over a landscape increased linearly with the proportion of GM maize over that landscape. The increase was twice as fast using the NIG kernel and was governed by the short-distance dispersal component. Variation in impurity rates largely depended on the distance to the closest GM crop and the size of the receptor field. However, impurity rates were generally underestimated when only dispersal from the closest GM field was considered. Synthesis and applications. Distance to the closest GM crop had most impact on impurity rates in conventional fields. However, impurity rates also depended on intermediate- to long-distance dispersal from distant GM crops. Therefore, isolation distances as currently defined will probably not allow long-term coexistence of GM and conventional crops, especially as the proportion of GM crops grown increases. We suggest strategies to account for this impact of long-distance dispersal. more...

Mixed mating, where a single tree progeny results from a mixture of selfing and outcrossing, is widespread in conifers and could be an evolutionary advantage at ecological margins when mating partners become scarce. This study analyzes how the mating system responds to bioclimate and density variations. We surveyed published data on the mating system of Abies, Picea and Pinus species when information on bioclimate and stand density was available. Our survey revealed that Mediterranean species demonstrate a lower selfing rate than other species and that the proportion of selfed versus outcrossed progeny is not fixed within species. The highest variability in mating types within populations was found when stand density was the most variable. To show how density affects the proportion of selfed versus outcrossed progeny, we used isozymes to genotype single tree seeds from a marginal Abies alba forest in Mediterranean France (Mont Ventoux) where low-to high-density stands are found. We then tested the adaptive potential of the different high and low density progenies by sowing them under controlled nursery conditions and measuring germination rate and seedling survival after 4 yr under 3 different water regimes. Although the mean value of outcrossing rate was typical for mixed mating conifers (tm = 0.85), individual outcrossing rates varied from 0.05 to 0.99 and were strongly correlated with stand type and density (tm from 0.87 in high-density to 0.43 in low-density marginal stands). Significantly fewer seeds from the low density marginal stand germinated (32% vs. 53% in the high density mature stand), although seedlings from the marginal stand had a better 4-yr survival (81%) than seedlings from the high-density mature stand (63%) when the three water regimes (from least to most stressful) were averaged. Trees from low density stands may be at a selective disadvantage because they produce more selfed progeny than trees from high-density stands. However, selfed seeds may be purged early on (at the embryo stage) and the remaining seeds tend to produce seedlings with better fitness. Variability of the selfing rate might thus be an adaptive strategy for species in highly contrasted environments where selection of both reproductive assurance and avoidance of inbreeding depression may be density dependent. more...