Crop-to-wild introgression in the European wild apple Malus sylvestris in Northern Britain 2 3 MARKUS RUHSAM

International audienc

Хирургическое лечение эпидермоидных кист около стволовой локализации

КИСТА ЭПИДЕРМОИДНАЯ /ХИ

New Insight into the History of Domesticated Apple: Secondary Contribution of the European Wild Apple to the Genome of Cultivated Varieties

The apple is the most common and culturally important fruit crop of temperate areas. The elucidation of its origin and domestication history is therefore of great interest. The wild Central Asian species Malus sieversii has previously been identified as the main contributor to the genome of the cultivated apple (Malus domestica), on the basis of morphological, molecular, and historical evidence. The possible contribution of other wild species present along the Silk Route running from Asia to Western Europe remains a matter of debate, particularly with respect to the contribution of the European wild apple. We used microsatellite markers and an unprecedented large sampling of five Malus species throughout Eurasia (839 accessions from China to Spain) to show that multiple species have contributed to the genetic makeup of domesticated apples. The wild European crabapple M. sylvestris, in particular, was a major secondary contributor. Bidirectional gene flow between the domesticated apple and the European crabapple resulted in the current M. domestica being genetically more closely related to this species than to its Central Asian progenitor, M. sieversii. We found no evidence of a domestication bottleneck or clonal population structure in apples, despite the use of vegetative propagation by grafting. We show that the evolution of domesticated apples occurred over a long time period and involved more than one wild species. Our results support the view that self-incompatibility, a long lifespan, and cultural practices such as selection from open-pollinated seeds have facilitated introgression from wild relatives and the maintenance of genetic variation during domestication. This combination of processes may account for the diversification of several long-lived perennial crops, yielding domestication patterns different from those observed for annual species

Diversification the genus Malus

Malgré son importance économique, culturelle et historique, l’histoire évolutive du pommier cultivé (Malus domestica) ainsi que celle de ses apparentés sauvages supposés, restaient encore très peu connues. En s’appuyant sur les nouvelles approches de génétique des populations (approximate Bayesian computation) avec l’utilisation de marqueurs microsatellites et de séquences nucléaires, cette thèse a eu pour objectif d’étudier, à différentes échelles évolutives (phylogéographie, spéciation, domestication), les mécanismes de diversification naturelle et artificielle dans le genre Malus. Mes travaux ont porté sur quatre espèces de pommiers sauvages distribuées à travers l’Eurasie (Malus orientalis (Caucase), Malus sieversii (Asie Centrale), Malus sylvestris (Europe), et Malus baccata (Sibérie)) et sur la seule espèce domestiquée du genre, Malus domestica. Cette thèse s’est articulée en quatre parties visant respectivement à inférer : (i) l’histoire de la domestication du pommier cultivé depuis son centre d’origine en Asie Centrale, (ii) l’histoire de la recolonisation post-glaciaire du pommier sauvage Européen (M. sylvestris), (iii) les histoires de spéciation entre les cinq espèces de Malus, (iv) les hybridations interspécifiques et les capacités de dispersion des trois principaux contributeurs (M. sylvestris, M. sieversii et M. orientalis) au génome du pommier cultivé. L’étude des mécanismes de diversification artificielle montre que les processus de domestication sont originaux chez cet arbre fruitier, de par l'absence de goulet d’étranglement et l’existence d’introgressions post-domestication fréquentes par une autre espèce sauvage (M. sylvestris) que l’espèce ancestrale (M. sieversii). L’étude des processus de diversification naturelle (phylogéographie, spéciation et structure des populations) révèlent de grandes tailles de populations, de forts flux de gènes et de faibles structures génétiques spatiales chez chacune des espèces. Cette thèse a aussi révélé de forts taux d’hybridations interspécifiques, en particulier de fortes introgressions des espèces de pommiers sauvages par le pommier cultivé en Europe et en Asie Centrale. Cette étude a permis l'amélioration des connaissances de la structuration des populations de pommiers sauvages ayant contribué au génome du pommier cultivé ainsi que de l’étendue des hybridations du pommier cultivé avec les espèces sauvages. Ces travaux revêtent une grande importance autant pour la conservation des pommiers sauvages, pour le maintien de leur intégrité dans des habitats fragmentés que pour l'amélioration variétale du pommier domestiqué.: Despite its economic, cultural and historical importance, few studies have investigated the evolutionary history of the domesticated apple (Malus domestica) as well as those of its wild relatives. Using new population genetic approaches (approximate Bayesian computation) with microsatellites and nuclear sequences, this thesis aimed at unravelling, at different evolutionary scales (phylogeography, speciation, domestication), the natural and artificial diversification processes at play in the Malus genus. My research focused on the four wild apple species distributed across Eurasia (Malus orientalis (Caucasus), Malus sieversii (Central Asia), Malus sylvestris (Europe), and Malus baccata (Siberia)) and on the single domesticated apple species in the genus, Malus domestica. This thesis was divided into four parts: (i) domestication history of the cultivated apple, from its origin in Central Asia to Europe, (ii) post-glacial recolonization history of the European crabapple (M. sylvestris), (iii) the history of speciation among the five Malus species, (iv) crop-to-wild gene flow and dispersal capacities of the closest wild relative species (M. sylvestris, M. sieversii and M. orientalis). By investigating artificial diversification, we evidenced unique processes of domestication in this fruit tree, with no bottleneck and with extensive post-domestication introgressions by another wild species (M. sylvestris) than the ancestral progenitor (M. sieversii). Natural diversification patterns (phylogeography, speciation and population structure) revealed large effective population sizes, high dispersal capacities and weak spatial genetic structures. This thesis also revealed high levels of interspecific hybridizations, particularly high level of crop-to-wild gene flow in Europe and Central Asia. This study extended our knowledge about population structures for wild species that contributed to the cultivated apple genome, as well as the extent of hybridization rates. This work is essential for the conservation of wild apple populations, the integrity maintenance of wild species facing fragmentation and future breeding programs concerning the domesticated apple

Diversification dans le genre Malus

: Despite its economic, cultural and historical importance, few studies have investigated the evolutionary history of the domesticated apple (Malus domestica) as well as those of its wild relatives. Using new population genetic approaches (approximate Bayesian computation) with microsatellites and nuclear sequences, this thesis aimed at unravelling, at different evolutionary scales (phylogeography, speciation, domestication), the natural and artificial diversification processes at play in the Malus genus. My research focused on the four wild apple species distributed across Eurasia (Malus orientalis (Caucasus), Malus sieversii (Central Asia), Malus sylvestris (Europe), and Malus baccata (Siberia)) and on the single domesticated apple species in the genus, Malus domestica. This thesis was divided into four parts: (i) domestication history of the cultivated apple, from its origin in Central Asia to Europe, (ii) post-glacial recolonization history of the European crabapple (M. sylvestris), (iii) the history of speciation among the five Malus species, (iv) crop-to-wild gene flow and dispersal capacities of the closest wild relative species (M. sylvestris, M. sieversii and M. orientalis). By investigating artificial diversification, we evidenced unique processes of domestication in this fruit tree, with no bottleneck and with extensive post-domestication introgressions by another wild species (M. sylvestris) than the ancestral progenitor (M. sieversii). Natural diversification patterns (phylogeography, speciation and population structure) revealed large effective population sizes, high dispersal capacities and weak spatial genetic structures. This thesis also revealed high levels of interspecific hybridizations, particularly high level of crop-to-wild gene flow in Europe and Central Asia. This study extended our knowledge about population structures for wild species that contributed to the cultivated apple genome, as well as the extent of hybridization rates. This work is essential for the conservation of wild apple populations, the integrity maintenance of wild species facing fragmentation and future breeding programs concerning the domesticated apple.Malgré son importance économique, culturelle et historique, l’histoire évolutive du pommier cultivé (Malus domestica) ainsi que celle de ses apparentés sauvages supposés, restaient encore très peu connues. En s’appuyant sur les nouvelles approches de génétique des populations (approximate Bayesian computation) avec l’utilisation de marqueurs microsatellites et de séquences nucléaires, cette thèse a eu pour objectif d’étudier, à différentes échelles évolutives (phylogéographie, spéciation, domestication), les mécanismes de diversification naturelle et artificielle dans le genre Malus. Mes travaux ont porté sur quatre espèces de pommiers sauvages distribuées à travers l’Eurasie (Malus orientalis (Caucase), Malus sieversii (Asie Centrale), Malus sylvestris (Europe), et Malus baccata (Sibérie)) et sur la seule espèce domestiquée du genre, Malus domestica. Cette thèse s’est articulée en quatre parties visant respectivement à inférer : (i) l’histoire de la domestication du pommier cultivé depuis son centre d’origine en Asie Centrale, (ii) l’histoire de la recolonisation post-glaciaire du pommier sauvage Européen (M. sylvestris), (iii) les histoires de spéciation entre les cinq espèces de Malus, (iv) les hybridations interspécifiques et les capacités de dispersion des trois principaux contributeurs (M. sylvestris, M. sieversii et M. orientalis) au génome du pommier cultivé. L’étude des mécanismes de diversification artificielle montre que les processus de domestication sont originaux chez cet arbre fruitier, de par l'absence de goulet d’étranglement et l’existence d’introgressions post-domestication fréquentes par une autre espèce sauvage (M. sylvestris) que l’espèce ancestrale (M. sieversii). L’étude des processus de diversification naturelle (phylogéographie, spéciation et structure des populations) révèlent de grandes tailles de populations, de forts flux de gènes et de faibles structures génétiques spatiales chez chacune des espèces. Cette thèse a aussi révélé de forts taux d’hybridations interspécifiques, en particulier de fortes introgressions des espèces de pommiers sauvages par le pommier cultivé en Europe et en Asie Centrale. Cette étude a permis l'amélioration des connaissances de la structuration des populations de pommiers sauvages ayant contribué au génome du pommier cultivé ainsi que de l’étendue des hybridations du pommier cultivé avec les espèces sauvages. Ces travaux revêtent une grande importance autant pour la conservation des pommiers sauvages, pour le maintien de leur intégrité dans des habitats fragmentés que pour l'amélioration variétale du pommier domestiqué

dom_ref_sylvestris_26SSR_structure

There are three files that we used for STRUCTURE analyses with the 26SSR markers that allowed us to detect hybrids and to do the following intra-species TESS analyses. Each file include wild species individuals and 40 M. domestica reference (last 40 individuals in each input file): (1) dom_ref_orientalis_26SSR_structure: the M. orientalis samples followed by the 40 M. domestica reference (2) dom_ref_sieversii_26SSR_structure: the M. sieversii samples followed by the 40 M. domestica reference (3) dom_ref_sylvestris_26SSR_structure: the M.sylvestris samples followed by the 40 M. domestica referenc

Evolutionary Genetics of Crop-Wild Complexes

Since Darwin’s time, the role of crop wild relatives (CWR), landraces, and cultivated genepools in shaping plant diversity and boosting food resources has been a major question [...

dom_ref_sieversii_26SSR_structure

There are three files that we used for STRUCTURE analyses with the 26SSR markers that allowed us to detect hybrids and to do the following intra-species TESS analyses. Each file include wild species individuals and 40 M. domestica reference (last 40 individuals in each input file): (1) dom_ref_orientalis_26SSR_structure: the M. orientalis samples followed by the 40 M. domestica reference (2) dom_ref_sieversii_26SSR_structure: the M. sieversii samples followed by the 40 M. domestica reference (3) dom_ref_sylvestris_26SSR_structure: the M.sylvestris samples followed by the 40 M. domestica referenc

Wild European apple (Malus sylvestris (L.) Mill.) population dynamics: insight from genetics and ecology in the Rhine Valley. Priorities for a future conservation programme.

The increasing fragmentation of forest habitats and the omnipresence of cultivars potentially threaten the genetic integrity of the European wild apple (Malus sylvestris (L.) Mill). However, the conservation status of this species remains unclear in Europe, other than in Belgium and the Czech Republic, where it has been declared an endangered species. The population density of M. sylvestris is higher in the forests of the upper Rhine Valley (France) than in most European forests, with an unbalanced age-structure, an overrepresentation of adults and a tendency to clump. We characterize here the ecology, age-structure and genetic diversity of wild apple populations in the Rhine Valley. We use these data to highlight links to the history of this species and to propose guidelines for future conservation strategies. In total, 255 individual wild apple trees from six forest stands (five floodplain forests and one forest growing in drier conditions) were analysed in the field, collected and genotyped on the basis of data for 15 microsatellite markers. Genetic analyses showed no escaped cultivars and few hybrids with the cultivated apple. Excluding the hybrids, the genetically "pure" populations displayed high levels of genetic diversity and a weak population structure. Age-structure and ecology studies of wild apple populations identified four categories that were not randomly distributed across the forests, reflecting the history of the Rhine forest over the last century. The Rhine wild apple populations, with their ecological strategies, high genetic diversity, and weak traces of crop-to-wild gene flow associated with the history of these floodplain forests, constitute candidate populations for inclusion in future conservation programmes for European wild apple