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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...

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...

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...

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...

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...

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...

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...

To make quantitative predictions about the pollen dispersal of a plant species under different environmental conditions, it is necessary to determine its individual pollen dispersal function, i.e., the two-dimensional density function describing the probability that a pollen grain emitted in (0, 0) fertilizes an ovule in (x, y). This function will depend on biological and climate parameters. We present models for the individual dispersal function of corn. These models are based on Brownian motion with drift and integrate biological (difference of height between male and female flowers) and aerodynamic (settling velocity, wind speed, air turbulence) parameters. The models presented differ in the importance of vegetation in stopping the paths of pollen grains. The models were fitted to data from two large field experiments of corn using the color of kernels as a phenotypic marker for pollen dispersal. The resulting estimations for the parameters of the models and comparisons between models indicate that (1)... 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...

Dispersal of pollen and seeds are essential functions of plant species, with far-reaching demographic, ecological and evolutionary consequences. Interest in plant dispersal has increased with concerns about the persistence of populations and species under global change. We argue here that advances in plant dispersal ecology research will be determined by our ability to surmount challenges of spatiotemporal scales and heterogeneities and ecosystem complexity. Based on this framework, we propose a selected set of research questions, for which we suggest some specific objectives and methodological approaches. Reviewed topics include multiple vector contributions to plant dispersal, landscape-dependent dispersal patterns, long-distance dispersal events, spatiotemporal variation in dispersal, and the consequences of dispersal for plant communities, populations under climate change, and anthropogenic landscapes., This work was partly supported by CGL2009-09428 project from the Spanish Ministry of Economy and Competitiveness. more...

International audience; Reproductive strategies of closely related species distributed along successional gradients should differ as a consequence of the trade-off between competition and colonization abilities. We compared male reproductive strategies of Quercus robur and Q. petraea, two partly interfertile European oak species with different successional status. In the studied even-aged stand, trees of the late-successional species (Q. petraea) grew faster and suffered less from intertree competition than trees of the early-successional species (Q. robur). A large-scale paternity study and a spatially explicit individual-based mating model were used to estimate parameters of pollen production and dispersal as well as sexual barriers between species. Male fecundity was found to be dependent both on a tree's circumference and on its environment, particularly so for Q. petraea. Pollen dispersal was greater and more isotropic in Q. robur than in Q. petraea. Premating barriers to hybridization were strong in both species, but more so in Q. petraea than in Q. robur. Hence, predictions based on the competition-colonization trade-off are well supported, whereas the sexual barriers themselves seem to be shaped by colonization dynamics. 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; Functional statistics are commonly used to characterize spatial patterns in general and spatial genetic structures in population genetics in particular. Such functional statistics also enable the estimation of parameters of spatially explicit (and genetic) models. Recently, Approximate Bayesian Computation (ABC) has been proposed to estimate model parameters from functional statistics. However, applying ABC with functional statistics may be cumbersome because of the high dimension of the set of statistics and the dependences among them. To tackle this difficulty, we propose an ABC procedure which relies on an optimized weighted distance between observed and simulated functional statistics. We applied this procedure to a simple step model, a spatial point process characterized by its pair correlation function and a pollen dispersal model characterized by genetic differentiation as a function of distance. These applications showed how the optimized weighted distance improved estimation accuracy. In the discussion, we consider the application of the proposed ABC procedure to functional statistics characterizing non-spatial processes. 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; 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; 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...

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...

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 mδ(p)57.3 m) were found, but there was also a non-negligible immigration rate from outside the plot (m(s) = 20.5%; 71.6%m(p)77.9%). Hierarchical analyses of seedling genetic structure showed that (i) most of the genetic variation was within plots; (ii) the genetic differentiation among seedling plots was significant (F(ST) = 2.6%) while (iii) there was no effect of year-to-year seed rain variation on genetic structure. In addition, no significant effect of genetic structure on mortality was detected. The consequences of these results for the prediction of population dynamics at ecological timescales are discussed. more...

International audience; 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 maximumlikelihood approaches (e.g. Neighbourhood model) that ignore variance in male fecundity or explain it through a few covariates and (ii) indirect methods (KinDist and Two-Gener) that integrate the variance in fecundity in an ‘effective population density’. 3. The Bayesian estimates correctly considered the over-dispersion resulting from the variance infecundity, resulting in wider but more accurate confidence intervals, in particular in high-densitypopulations. The maximum likelihood methods resulted in confidence intervals with low coverageprobabilities and in widespread false-positive tests when testing the effect of covariates on malefecundity. 4. Estimated individual fecundities and estimated empirical variance in fecundity were robust tothe 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 lessinformation 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...

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...

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 (N(b) = 70 individuals, sigma(e) = 417 m) were similar to those inferred from adult genotypes (N(b) = 114 individuals, sigma(e) = 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 (sigma(c) = 211 m) was approximately 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...

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...

International audience

Background Rare long distance dispersal events impact the demography and the genetic structure of populations. When dispersal is modelled via a dispersal kernel, one possible characterisation of long-distance dispersal is given by the shape of the tail of the kernel, i.e. its type of decay. This characteristic is known to directly act on the speed and pattern of colonization, and on the spatial structure of genetic diversity during colonization. In particular, colonization waves behave differently depending on whether the kernel decreases faster or slower than an exponential (i.e. is thin-tailed vs. fat-tailed). To interpret and extend published results on the impact of long-distance dispersal on the genetic structure of populations, we examine a classification of dispersal kernels based on the shape of their tails and formally demonstrate qualitative differences among them that can influence the predicted diversity of a propagule pool sampled far from two distinct sources. Results We show that a fat-tailed kernel leads asymptotically to a diverse propagule pool containing a balanced mixing of the propagules from the two sources, whereas a thin-tailed kernel results in all propagules originating from the closest source. We further show that these results hold for biologically relevant distances under certain circumstances, and in particular if the number of propagules is large enough, as would be the case for pollen or seeds. Conclusion To understand the impact of long-distance dispersal on the structure and dynamics of a metapopulation, it might be less important to precisely estimate an average dispersal distance than to determine if the tail of the dispersal kernel is fatter or thinner than that of an exponential function. Depending solely on this characteristic, a metapopulation will behave similarly to an island model with a diverse immigrant pool or to a stepping-stone model with migrants from closest populations. Our results further help to understand why thin-tailed dispersal kernels lead to a colonization wave of constant speed, whereas fat-tailed dispersal kernels lead to a wave of increasing speed. Our results also suggest that the diversity of the pollen cloud of a mother plant should increase with increasing isolation for fat-tailed kernels, whereas it should decrease for thin-tailed kernels. more...

Background The structure and evolution of hybrid zones depend mainly on the relative importance of dispersal and local adaptation, and on the strength of assortative mating. Here, we study the influence of dispersal, temporal isolation, variability in phenotypic traits and parasite attacks on the male mating success of two parental species and hybrids by real-time pollen flow analysis. We focus on a hybrid zone population between the two closely related ash species Fraxinus excelsior L. (common ash) and F. angustifolia Vahl (narrow-leaved ash), which is composed of individuals of the two species and several hybrid types. This population is structured by flowering time: the F. excelsior individuals flower later than the F. angustifolia individuals, and the hybrid types flower in-between. Hybrids are scattered throughout the population, suggesting favorable conditions for their local adaptation. We estimate jointly the best-fitting dispersal kernel, the differences in male fecundity due to variation in phenotypic traits and level of parasite attack, and the strength of assortative mating due to differences in flowering phenology. In addition, we assess the effect of accounting for genotyping error on these estimations. Results We detected a very high pollen immigration rate and a fat-tailed dispersal kernel, counter-balanced by slight phenological assortative mating and short-distance pollen dispersal. Early intermediate flowering hybrids, which had the highest male mating success, showed optimal sex allocation and increased selfing rates. We detected asymmetry of gene flow, with early flowering trees participating more as pollen donors than late flowering trees. Conclusion This study provides striking evidence that long-distance gene flow alone is not sufficient to counter-act the effects of assortative mating and selfing. Phenological assortative mating and short-distance dispersal can create temporal and spatial structuring that appears to maintain this hybrid population. The asymmetry of gene flow, with higher fertility and increased selfing, can potentially confer a selective advantage to early flowering hybrids in the zone. In the event of climate change, hybridization may provide a means for F. angustifolia to further extend its range at the expense of F. excelsior. more...

International audience; Understanding the role of mother plants as pollen recipients in shaping mating patterns is essential for understanding the evolution of populations and in particular to predict the consequence of habitat fragmentation. Here, we investigated variation in mating patterns due to maternal phenotypic traits, phenological variance, and landscape features in Sorbus torminalis, a hermaphroditic, insect‐pollinated and low‐density, European temperate forest tree. The diversity and composition of pollen clouds received by maternal trees in S. torminalis were mainly determined by their conspecific neighborhood: isolated individuals sample more diversity through more even paternal contributions, low relatedness among paternal genes, and high rates of long‐distance pollen dispersal within their progenies. Maternal phenotypic traits related to pollinator attractiveness also had an effect, but only when competition was strong: in this case, larger mother trees with more flowers sampled more diversity. The floral architecture of S. torminalis, with multiple‐seeded fruit, strongly shaped mating patterns, with higher levels of correlated paternity among seeds belonging to the same fruit (30% full sibs) than among seeds belonging to different fruits (14% full sibs). Finally, flowering phenology affected the distribution of diversity among maternal pollen clouds, but the earliest and latest mother trees did not receive less diversity of pollen than the others. more...

Knowing the extent of gene movements from parents to offspring is essential to understand the potential of a species to adapt rapidly to a changing environment, and to design appropriate conservation strategies. In this study, we develop a nonlinear statistical model to jointly estimate the pollen dispersal kernel and the heterogeneity in fecundity among phenotypically or environmentally defined groups of males. This model uses genotype data from a sample of fruiting plants, a sample of seeds harvested on each of these plants, and all males within a circumscribed area. We apply this model to a scattered, entomophilous woody species, Sorbus torminalis (L.) Crantz, within a natural population covering more than 470 ha. We estimate a high heterogeneity in male fecundity among ecological groups, both due to phenotype (size of the trees and flowering intensity) and landscape factors (stand density within the neighbourhood). We also show that fat-tailed kernels are the most appropriate to depict the important abilities of long-distance pollen dispersal for this species. Finally, our results reveal that the spatial position of a male with respect to females affects as much its mating success as ecological determinants of male fecundity. Our study thus stresses the interest to account for the dispersal kernel when estimating heterogeneity in male fecundity, and reciprocally. more...

Gene flow from crops to wild related species has been recently under focus in risk-assessment studies of the ecological consequences of growing transgenic crops. However, experimental studies addressing this question are usually temporally or spatially limited. Indirect population-structure approaches can provide more global estimates of gene flow, but their assumptions appear inappropriate in an agricultural context. In an attempt to help the committees providing advice on the release of transgenic crops, we present a new method to estimate the quantity of genes migrating from crops to populations of related wild plants by way of pollen dispersal. This method provides an average estimate at a landscape level. Its originality is based on the measure of the inverse gene flow, i.e. gene flow from the wild plants to the crop. Such gene flow results in an observed level of impurities from wild plants in crop seeds. This level of impurity is usually known by the seed producers and, in any case, its measure is easier than a direct screen of wild populations because crop seeds are abundant and their genetic profile is known. By assuming that wild and cultivated plants have a similar individual pollen dispersal function, we infer the level of pollen-mediated gene flow from a crop to the surrounding wild populations from this observed level of impurity. We present an example for sugar beet data. Results suggest that under conditions of seed production in France (isolation distance of 1,000 m) wild beets produce high numbers of seeds fathered by cultivated plants. more...

In order to help establish a basis for the assessment of gene flow associated with the large-scale release of transgenic oilseed rape, we previously designed a method which makes it possible to retrieve the average pollen dispersal of a single plant from that of a large source plot. The ‘individual’ pollen distribution thus obtained is less dependent on the experimental design than pollen distributions usually published and could therefore be used to model the possible escape of a transgene from commercial transgenic crops. In this study we report on a field experiment set up to study the pollen dispersal from an herbicide-resistant transgenic variety of oilseed rape and to test the applicability of the method on the experimental data. Two techniques were used to determine the individual pollen dispersal, and their outcomes are compared. The results suggest that approximately half of the pollen produced by an individual plant fell within 3 m and that the probability of fertilisation afterwards decreased slowly along a negative exponential of the distance. Comparison with the global pollen distribution from the source plot indicates that pollen-dispersal distributions based on dispersal from whole plots instead of individual plants would have underestimated the proportion of pollen that was dispersed over average or long distances. more...