Fabien Halkett

French National Institute for Agricultural Research, Lutetia Parisorum, Île-de-France, France

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Publications (24)117.18 Total impact

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    ABSTRACT: The genetic structure of ectomycorrhizal (ECM) fungal populations results from both vegetative and sexual propagation. In this study, we have analysed the spatial genetic structure of Tuber melanosporum populations, a heterothallic ascomycete that produces edible fruit bodies. Ectomycorrhizas from oaks and hazels from two orchards were mapped and genotyped using simple sequence repeat markers and the mating type locus. The distribution of the two T. melanosporum mating types was also monitored in the soil. In one orchard, the genetic profiles of the ascocarps were compared with those of the underlying mycorrhizas. A pronounced spatial genetic structure was found. The maximum genet sizes were 2.35 and 4.70 m in the two orchards, with most manifesting a size < 1 m. Few genets persisted throughout two seasons. A nonrandom distribution pattern of the T. melanosporum was observed, resulting in field patches colonized by genets that shared the same mating types. Our findings suggest that competition occurs between genets and provide basic information on T. melanosporum propagation patterns that are relevant for the management of productive truffle orchards.
    New Phytologist 04/2013; · 6.74 Impact Factor
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    ABSTRACT: Alder decline caused by Phytophthora alni has been one of the most important diseases of natural ecosystems in Europe during the last 20 years. The emergence of Phytophthora alni subsp. alni (Paa)-the pathogen responsible for the epidemic-is linked to an interspecific hybridization event between two parental species: Phytophthora alni subsp. multiformis (Pam) and Phytophthora alni subsp. uniformis (Pau). One of the parental species, Pau, has been isolated in several European countries and recently in North America. The objective of this work was to assess the level of genetic diversity, the population genetic structure, and the putative reproduction mode and mating system of Pau. Five new polymorphic microsatellite markers were used to contrast both geographical populations. The study comprised 71 isolates of Pau collected from 8 European countries and 10 locations in North America. Our results revealed strong differences between continental populations (Fst=0.88; Rst=0.74), with no evidence for gene flow. European isolates showed extremely low genetic diversity compared to the North American collection. Selfing appears to be the predominant mating system in both continental collections. The results suggest that the European Pau population is most likely alien and derives from the introduction of a few individuals, while the North American population probably is an indigenous population.
    Phytopathology 01/2013; 103:190-199. · 2.97 Impact Factor
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    ABSTRACT: Fungal plant pathogens, especially rust fungi (Pucciniales), are well known for their complex life cycles, which include phases of sexual and asexual reproduction. The effect of asexual multiplication on population genetic diversity has been investigated in the poplar rust fungus Melampsora larici-populina using a nested hierarchical sampling scheme. Four hierarchical levels were considered: leaf, twig, tree and site. Both cultivated and wild poplar stands were sampled at two time points at the start and end of rust epidemics. A total of 641 fungal isolates was analysed using nine microsatellite markers. This study revealed that the genetic signature of asexual multiplication in the wild poplar stand was seen only at lower hierarchical levels (leaf and twig). Moreover, we observed an erosion of clonal structure through time, with an increase in both gene and genotypic diversity. New genotypes contributed to host infection over time, which demonstrates the importance of allo-infection in the epidemic process in this host-pathogen system. Compared with the wild stands, the nearly lack of detection of clonal structure in the cultivated stands reflects the higher infection level on cultivated poplars. More generally, this genetic analysis illustrates the utility of population genetics approach for elucidating the proportion of asexual reproduction in the multiplication of isolates during an epidemic, and for proper quantification of asexual dispersal in plant pathogens.
    Molecular Ecology 09/2012; 21(20):4996-5008. · 6.28 Impact Factor
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    ABSTRACT: The advent of molecular epidemiology has greatly improved our ability to identify the population sources and track the pathogen movement. Yet the wide spatial and temporal scales usually considered are useful only to infer historical migration pathways. In this study, Bayesian genetic assignments and a landscape epidemiology approach were combined to unravel genetic origin and annual spread during a single epidemic of a plant pathogen: the poplar rust fungus Melampsora larici-populina. The study focused on a particular area-the Durance River valley-which enabled inoculum sources to be identified and channelled spread of the epidemic along a one-dimensional corridor. Spatio-temporal monitoring of disease showed that the epidemic began in the upstream part of the valley and spread out downstream. Using genetic assignment tests, individuals collected at the end of the epidemic were sorted into two genetic groups; very few hybrids were detected, although individuals from both groups coexisted locally downstream in the valley. The epidemic originated from two genetically distinct inoculum sources. Individuals of each group then dispersed southwards along the Durance River and became mixed in poplar riparian stands. These two genetic groups were found previously at a wider spatial scale and proved to result from distinct evolutionary histories on either wild or cultivated poplars. This study showed that the two groups can mix during an epidemic but do not hybridize because they then reproduce asexually. In general, the methods employed here could be useful for elucidating the genetic origin and retracing the colonization history and migration pathways of recent epidemics.
    Molecular Ecology 04/2012; 21(10):2383-98. · 6.28 Impact Factor
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    ABSTRACT: Wild and cultivated plants represent very different habitats for pathogens, especially when cultivated plants bear qualitative resistance genes. Here, we investigated to what extent the population genetic structure of a plant pathogenic fungus collected on its wild host can be impacted by the deployment of resistant cultivars. We studied one of the main poplar diseases, poplar rust, caused by the fungus Melampsora larici-populina. A thousand and fifty individuals sampled from several locations in France were phenotyped for their virulence profile (ability to infect or not the most deployed resistant cultivar 'Beaupré'), and a subset of these was genotyped using 25 microsatellite markers. Bayesian assignment tests on genetic data clustered the 476 genotyped individuals into three genetic groups. Group 1 gathered most virulent individuals and displayed evidence for selection and drastic demographic changes resulting from breakdown of the poplar cultivar 'Beaupré'. Group 2 comprised individuals corresponding to ancestral populations of M. larici-populina naturally occurring in the native range. Group 3 displayed the hallmarks of strict asexual reproduction, which has never previously been demonstrated in this species. We discuss how poplar cultivation has influenced the spatial and genetic structure of this plant pathogenic fungus, and has led to the spread of virulence alleles (gene swamping) in M. larici-populina populations evolving on the wild host.
    Molecular Ecology 07/2011; 20(13):2739-55. · 6.28 Impact Factor
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    ABSTRACT: Summary1. Biological invasions are a major consequence of globalization and pose a significant threat to biodiversity. Because only a small fraction of introduced species become invasive, identification of those species most likely to become invasive after introduction is highly desirable to focus management efforts. The predictive potential of species-specific traits has been much investigated in plants and animals. However, despite the importance of fungi as a biological group and the potentially severe effects of pathogenic fungi on agrosystems and natural ecosystems, the specific identification of traits correlated with the invasion success of fungi has not been attempted previously.2. We addressed this question by constructing an ad hoc data set including invasive and non-invasive species of forest pathogenic fungi introduced into Europe. Data were analysed with a machine learning method based on classification trees (Random Forest). The performance of the classification rule based on species traits was compared with that of several random decision rules, and the principal trait predictors associated with invasive species were identified.3. Invasion success was more accurately predicted by the classification rule including biological traits than by random decision rules. The effect of species traits was maintained when confounding variables linked to residence time and habitat availability were included. The selected traits were unlikely to be affected by a phylogenetic bias as invasive and non-invasive species were evenly distributed in fungal clades.4. The species-level predictors identified as useful for distinguishing between invasive and non-invasive species were traits related to long-distance dispersal, sexual reproduction (in addition to asexual reproduction), spore shape and size, number of cells in spores, optimal temperature for growth and parasitic specialization (host range and infected organs).5. Synthesis and applications. This study demonstrates that some species-level traits are predictors of invasion success for forest pathogenic fungi in Europe. These traits could be used to refine current pest risk assessment (PRA) schemes. Our results suggest that current schemes, which are mostly based on sequential questionnaires, could be improved by taking into account trait interactions or combinations. More generally, our results confirm the interest of machine learning methods, such as Random Forest, for species classification in ecology.
    Journal of Applied Ecology 06/2011; 48(6):1381 - 1390. · 4.74 Impact Factor
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    ABSTRACT: Landscape genetics, which combines population genetics, landscape ecology and spatial statistics, has emerged recently as a new discipline that can be used to assess how landscape features or environmental variables can influence gene flow and spatial genetic variation. We applied this approach to the invasive plant pathogenic fungus Mycosphaerella fijiensis, which causes black leaf streak disease of banana. Around 880 isolates were sampled within a 50 × 50 km area located in a fragmented banana production zone in Cameroon that includes several potential physical barriers to gene flow. Two clustering algorithms and a new F(ST) -based procedure were applied to define the number of genetic entities and their spatial domain without a priori assumptions. Two populations were clearly delineated, and the genetic discontinuity appeared sharp but asymmetric. Interestingly, no landscape features matched this genetic discontinuity, and no isolation by distance (IBD) was found within populations. Our results suggest that the genetic structure observed in this production area reflects the recent history of M. fijiensis expansion in Cameroon rather than resulting from contemporary gene flow. Finally, we discuss the influence of the suspected high effective population size for such an organism on (i) the absence of an IBD signal, (ii) the characterization of contemporary gene-flow events through assignation methods of analysis and (iii) the evolution of the genetic discontinuity detected in this study.
    Molecular Ecology 03/2011; 20(8):1661-74. · 6.28 Impact Factor
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    ABSTRACT: In some diseases-in particular, tree root infection-stages of infection and inoculum production level and timing are not readily observable because of uncertainty or time lags in symptom appearance. Here, we pose a criterion, based on relative hazard of disease symptoms, to discriminate between healthy and asymptomatic infected individuals. We design a statistical procedure to estimate the criterion for a 6-year survey of alder decline along a northeastern French river. Individual tree symptom hazard was modeled with Cox's regression model, taking estimation of local infection pressure as a risk factor. From an inoculum production experiment, we thereafter assessed the inoculum production level of target trees, including symptomatic and asymptomatic trees ranked according to their symptoms hazard. Using receiver operating characteristic methods, we first evaluated the criterion performance and determined the discrimination threshold to sort out asymptomatic individuals into healthy and infected. Then, we highlighted the fact that the infected asymptomatic trees were among the major inoculum producers whereas severely declining and dead trees were found to be poor inoculum sources.
    Phytopathology 11/2010; 100(11):1262-9. · 2.97 Impact Factor
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    ABSTRACT: Dispersal processes of fungal plant pathogens can be inferred from analysis of spatial genetic structures resulting from recent range expansion. The relative importance of long-distance dispersal (LDD) events vs. gradual dispersal in shaping population structures depends on the geographical scale considered. The fungus Mycosphaerella fijiensis, pathogenic on banana, is an example of a recent worldwide epidemic. Founder effects in this species were detected at both global and continental scale, suggesting stochastic spread of the disease through LDD events. In this study, we analysed the structure of M. fijiensis populations in two recently (∼1979-1980) colonized areas in Costa Rica and Cameroon. Isolates collected in 10-15 sites distributed along a ∼250- to 300- km-long transect in each country were analysed using 19 microsatellite markers. We detected low-to-moderate genetic differentiation among populations in both countries and isolation by distance in Cameroon. Combined with historical data, these observations suggest continuous range expansion at the scale of banana-production area through gradual dispersal of spores. However, both countries displayed specific additional signatures of colonization: a sharp discontinuity in gene frequencies was observed along the Cameroon transect, while the Costa Rican populations seemed not yet to have reached genetic equilibrium. These differences in the genetic characteristics of M. fijiensis populations in two recently colonized areas are discussed in the light of historical data on disease spread and ecological data on landscape features.
    Molecular Ecology 09/2010; 19(18):3909-23. · 6.28 Impact Factor
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    ABSTRACT: In aphids, reproductive mode is generally assumed to be selected for by winter climate. Sexual lineages produce frost-resistant eggs, conferring an advantage in regions with cold winters, while asexual lineages predominate in regions with mild winters. However, habitat and resource heterogeneities are known to exert a strong influence on sex maintenance and might modulate the effect of climate on aphid reproductive strategies. We carried out a hierarchical sampling in northern France to investigate whether reproductive mode variation of the aphid Rhopalosiphum padi is driven by winter climate conditions, by habitat and resource heterogeneities represented by a range of host plants or by both factors. We confirmed the coexistence in R. padi populations of two genetic clusters associated with distinct reproductive strategies. Asexual lineages predominated, whatever the surveyed year and location. However, we detected a between-year variation in the local contribution of both clusters, presumably associated with preceding winter severity. No evidence for host-driven niche differentiation was found in the field on six Poaceae among sexual and asexual lineages. Two dominant multilocus genotypes ( approximately 70% of the sample), having persisted over a 10-year period, were equally abundant on different plant species and locations, indicating their large ecological tolerance. Our results fit theoretical predictions of the influence of winter climate on the balance between sexual and asexual lineages. They also highlight the importance of current agricultural practices which seem to favour a small number of asexual generalist genotypes and their migration across large areas of monotonous environments.
    Molecular Ecology 07/2009; 18(14):3050-61. · 6.28 Impact Factor
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    ABSTRACT: Life-history theory postulates that evolution is constrained by trade-offs (i.e., negative genetic correlations) among traits that contribute to fitness. However, in organisms with complex life cycles, trade-offs may drastically differ between phases, putatively leading to different evolutionary trajectories. Here, we tested this possibility by examining changes in life-history traits in an aphid species that alternates asexual and sexual reproduction in its life cycle. The quantitative genetics of reproductive and dispersal traits was studied in 23 lineages (genotypes) of the bird cherry-oat aphid Rhopalosiphum padi, during both the sexual and asexual phases, which were induced experimentally under specific environmental conditions. We found large and significant heritabilities (broad-sense) for all traits and several negative genetic correlations between traits (trade-offs), which are related to reproduction (i.e., numbers of the various sexual or asexual morphs) or dispersal (i.e., numbers of winged or wingless morphs). These results suggest that R. padi exhibits lineage specialization both in reproductive and dispersal strategies. In addition, we found important differences in the structure of genetic variance-covariance matrices (G) between phases. These differences were due to two large, negative genetic correlations detected during the asexual phase only: (1) between fecundity and age at maturity and (2) between the production of wingless and winged parthenogenetic females. We propose that this differential expression in genetic architecture results from a reallocation scheme during the asexual phase, when sexual morphs are not produced. We also found significant G x E interaction and nonsignificant genetic correlations across phases, indicating that genotypes could respond independently to selection in each phase. Our results reveal a rather unique situation in which the same population and even the same genotypes express different genetic (co)variation under different environmental conditions, driven by optimal resource allocation criteria.
    Evolution 05/2009; 63(9):2402-12. · 4.86 Impact Factor
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    ABSTRACT: Aggressiveness, the quantitative component of pathogenicity, and its role in the adaptation of plant pathogens are still insufficiently investigated. Using mainly examples of biotrophic and necrotrophic fungal pathogens of cereals and Phytophthora infestans on potato, the empirical knowledge on the nature of aggressiveness components and their evolution in response to host and environment is reviewed. Means of measuring aggressiveness components are considered, as well as the sources of environmental variance in these traits. The adaptive potential of aggressiveness components is evaluated by reviewing evidence for their heritability, as well as for constraints on their evolution, including differential interactions between host and pathogen genotypes and trade-offs between components of pathogenicity. Adaptations of pathogen aggressiveness components to host and environment are analysed, showing that: (i) selection for aggressiveness in pathogen populations can be mediated by climatic parameters; (ii) global population changes or remarkable population structures may be explained by variation in aggressiveness; and (iii) selection for quantitative traits can influence pathogen evolution in agricultural pathosystems and can result in differential adaptation to host cultivars, sometimes leading to erosion of quantitative resistance. Possible links with concepts in evolutionary ecology are suggested.
    Plant Pathology 01/2009; 58:409-424. · 2.73 Impact Factor
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    ABSTRACT: Forty-one microsatellite markers were developed for the poplar rust fungus Melampsora larici-populina. The genome sequence of M. larici-populina strain 98AG31 was screened in silico for microsatellite loci using Magellan software. Thirty-nine loci, evaluated on a sexual population of 35 M. larici-populina isolates sampled on larch trees in the French Alps, were identified to be highly polymorphic. These markers revealed two to 34 alleles, with an average of 7.23 alleles per locus. The estimated heterozygosity ranged from 0.029 to 0.958 across the 41 microsatellite loci. These markers will be highly useful for population genetics studies of M. larici-populina.
    Conservation Genetics Resources 01/2009; 1(1):21-25. · 0.71 Impact Factor
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    ABSTRACT: Many organisms considered as strictly clonal may in fact experience some rare events of sexual reproduction with their sexual relatives. However, the rate of sexual-asexual gene flow has rarely been assessed mainly because its evaluation is difficult to achieve in the field. In the cyclically parthenogenetic aphid Rhopalosiphum padi, two main sets of lineages, differing in their investment in sexual reproduction and in their genetic attributes, co-exist even at a very fine scale: the 'sexual' lineages which have a full commitment to the sexual reproduction, and the 'facultatively asexual' lineages, which allocate investment in the sexual and parthenogenetic reproduction. This system offers a unique opportunity to tackle the genetic interactions between two contrasting reproductive modes. Here, we provide evidence that gene flow occurred between sexual and facultatively asexual lineages of R. padi. We carefully examined the shuffling in phenotypic and genotypic variation following a sexual reproduction event that took place in the field. Combining genotypic data and phenotypic measurements showed that this gene mixing led to the production of a wide array of reproductive modes, including strictly asexual lineages. Finally, we discuss the central role played by facultatively asexual lineages on the maintenance of reproductive mode variation.
    Molecular Ecology 07/2008; 17(12):2998-3007. · 6.28 Impact Factor
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    ABSTRACT: Changes in the genetic structure and genotypic variation of the aphid Sitobion avenae collected from cereal crops in northern France were examined by analysing variation at five microsatellite loci across several years and seasons. Little regional and temporal differentiation was detected, as shown by very low FST among populations. Repeated genotypes, significant heterozygote deficits, positive FIS values and frequent linkage disequilibria were found in nearly all samples, suggesting an overall pattern of reproductive mode variation in S. avenae populations. In addition, samples from Brittany (Bretagne) showed greater signs of asexual reproduction than those from the north of France, indicating a trend toward increasing sexuality northward. These patterns of reproductive variation in S. avenae are consistent with theoretical models of selection of aphid reproductive modes by climate. Contrasting with little changes in allelic frequencies, genotypic composition varied substantially in time and, to a lesser extent, in space. An important part of changes in genotypic arrays was due to the variation in frequency distribution of common genotypes, i.e. those that were found at several instances in the samples. Genotypic composition was also shown to vary according to climate, as genotypic diversity in spring was significantly correlated with the severity of the previous winter and autumn. We propose that the genetic homogeneity among S. avenae populations shown here across large temporal and spatial scales is the result of two forces: (i) migration conferred by high dispersal capabilities, and (ii) selection over millions of hectares of cereals (mostly wheat) bred from a narrow genetic base.
    Bulletin of Entomological Research 05/2008; 98(2):159-68. · 1.99 Impact Factor
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    ABSTRACT: Dispersal has a great impact on the genetic structure of populations, but remains difficult to estimate by direct measures. In particular, gradual and stochastic dispersal are often difficult to assess and to distinguish, although they have different evolutionary consequences. Plant pathogens, especially rust fungi, are suspected to display both dispersal modes, though on different spatial scales. In this study, we inferred dispersal capacities of the poplar rust fungus Melampsora larici-populina by examining the genetic diversity and structure of 13 populations from eight European and two overseas countries in the Northern hemisphere. M. larici-populina was sampled from both cultivated hybrid poplars and on the wild host, Populus nigra. The populations were analyzed with 11 microsatellite and 8 virulence markers. Although isolates displayed different virulence profiles according to the host plant, neutral markers revealed little population differentiation with respect to the type of host. This suggests an absence of reproductive isolation between populations sampled from cultivated and wild poplars. Conversely, studying the relationship between geographic and genetic structure allowed us to distinguish between isolation by distance (IBD) patterns and long distance dispersal (LDD) events. The European populations exhibited a significant IBD pattern, suggesting a regular and gradual dispersal of the pathogen over this spatial scale. Nonetheless, the genetic differentiation between these populations was low, suggesting an important gene flow on a continental scale. The two overseas populations from Iceland and Canada were shown to result from rare LDD events, and exhibited signatures of strong founder effects. Furthermore, the high genetic differentiation between both populations suggested that these two recent introductions were independent. This study illustrated how the proper use of population genetics methods can enable contrasted dispersal modes to be revealed.
    Infection Genetics and Evolution 04/2008; 8(5):577-587. · 2.77 Impact Factor
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    ABSTRACT: The geographic range of plant pests can be modified by the use of glasshouses. Bemisia tabaci, originating from warm to hot climates, has been shown to be a complex of distinct genetic groups with very limited gene flow. The genetic structure of this pest was studied in glasshouses in southern France, a region beyond the northern limit of its open-field development area in Europe. Seven microsatellite loci were scored in 22 populations sampled from various regions over 3 years. Two genetic groups were distinguished using a Bayesian clustering method and were assigned to the so-called biotypes B and Q using the gene sequence of cytochrome oxidase 1 (CO1). All but one population corresponded to biotype Q, even though only biotype B was previously reported. Despite the enclosed environment of glasshouses and their expected isolation due to low outdoor survival during the winter, only limited differentiation among biotype Q glasshouses was observed. A single sample site was notable for a decrease in expected heterozygosity and the mean number of alleles over the years. The lack of spatial genetic structure among biotype Q populations was indicative of a recent colonization event combined with large dispersal at all spatial scales. This migration pattern of biotype Q populations was further supported by additional CO1 sequences, since individuals from France, Asia and America exhibited 100% nucleotide identity. The evolution of genetic diversity observed in glasshouses in France is part of the worldwide invasion of biotype Q, which is discussed in light of human activities.
    Heredity 04/2008; 100(3):316-25. · 4.11 Impact Factor
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    ABSTRACT: Clonal reproduction in Puccinia triticina, the cause of wheat leaf rust, has long been hypothesized but has never been demonstrated. Using a population genetics approach and microsatellite markers, we analysed genetic diversity of this fungus at each level of genome organisation. Sampling included isolates from two field populations growing on two cultivars carrying specific resistance genes, completed with isolates representing the main pathotypes identified from a national survey. For the two cultivars, populations differentiated according to the distribution of their genotypes and pathotypes. There was a high proportion of repeated genotypes, combined with a significant linkage disequilibrium and a strong negative value for FIS. These three factors, especially heterozygote excess, strongly support the hypothesis of a high rate of clonal reproduction. Each pathotype matched a unique multilocus genotype, except for a few isolates, which were taken to be mutants of the dominant genotype. We discussed the strong relationship between pathotypes and genotypes as the consequence of clonal reproduction combined with a strong selection exerted by host cultivars. (c) 2007 Elsevier Inc. All rights reserved.
    Fungal Genetics and Biology 07/2007; 44(6):474-483. · 3.26 Impact Factor
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    ABSTRACT: Cases of coexisting sexual and asexual relatives are puzzling, as evolutionary theory predicts that competition for the same ecological niches should lead to the exclusion of one or the other population. In the cyclically parthenogenetic aphid, Rhopalosiphum padi, sexual and facultative asexual lineages are admixed in space at the time of sexual reproduction. We investigated how the interaction of reproductive mode and environment can lead to temporal niche differentiation. We demonstrated theoretically that differential sensitivity of sexual and facultatively asexual aphids to an environmental parameter (mating host suitability) shapes the two strategies: whereas the sexual lineages switch earlier to the production of sexual forms, the facultative asexual lineages delay and spread out their investment in sexual reproduction. This predicted pattern of niche specialization is in agreement with the temporal structure revealed in natura by demographic and genetic data. We propose that partial loss of sex by one pool of aphids and subsequent reduction in gene flow between lineages may favour temporal specialization through disruptive selection.
    Journal of Evolutionary Biology 06/2006; 19(3):809-15. · 3.48 Impact Factor
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    ABSTRACT: Many clonal organisms experience occasional events of sexual recombination, with profound consequences for their population dynamics and evolutionary trajectories. With the recent development of polymorphic genetic markers and new statistical methods, we now have an unprecedented ability to detect recombination in organisms that are thought to reproduce strictly, or essentially asexually. However, it is not always obvious which methodology to apply. Consequently, biologists might decide how to analyse their data without clear guidelines. Here, we discuss the available methods, focusing on those best suited when working with limited genetic information, such as a few genetic markers or DNA sequences. We conclude by commenting on the prospects offered by some recent conceptual advances and the access to high throughput technologies in an increasing number of model organisms.
    Trends in Ecology & Evolution 05/2005; 20(4):194-201. · 15.39 Impact Factor

Publication Stats

507 Citations
117.18 Total Impact Points

Institutions

  • 2004–2013
    • French National Institute for Agricultural Research
      • Interactions Arbres/Micro-organismes (IAM)
      Lutetia Parisorum, Île-de-France, France
  • 2008–2012
    • University of Bordeaux
      Burdeos, Aquitaine, France
  • 2006
    • La Trobe University
      • Department of Genetics
      Melbourne, Victoria, Australia
    • Cirad - La recherche agronomique pour le développement
      Montpelhièr, Languedoc-Roussillon, France