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Analyses Génétiques Et Moléculaires Du Locus Skr Impliqué Dans L’Aptitude Du Blé (Triticum Aestivum L.) Au Croisement Avec Le Seigle (Secale Cereale L.) Walid Alfares

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Analyses Génétiques Et Moléculaires Du Locus Skr Impliqué Dans L’Aptitude Du Blé (Triticum Aestivum L.) Au Croisement Avec Le Seigle (Secale Cereale L.) Walid Alfares Analyses génétiques et moléculaires du locus SKr impliqué dans l’aptitude du blé (Triticum aestivum L.) au croisement avec le seigle (Secale cereale L.) Walid Alfares To cite this version: Walid Alfares. Analyses génétiques et moléculaires du locus SKr impliqué dans l’aptitude du blé (Triticum aestivum L.) au croisement avec le seigle (Secale cereale L.). Génétique des plantes. Uni- versité Blaise Pascal - Clermont-Ferrand II, 2009. Français. NNT : 2009CLF21984. tel-00724743 HAL Id: tel-00724743 https://tel.archives-ouvertes.fr/tel-00724743 Submitted on 22 Aug 2012 HAL is a multi-disciplinary open access L’archive ouverte pluridisciplinaire HAL, est archive for the deposit and dissemination of sci- destinée au dépôt et à la diffusion de documents entific research documents, whether they are pub- scientifiques de niveau recherche, publiés ou non, lished or not. The documents may come from émanant des établissements d’enseignement et de teaching and research institutions in France or recherche français ou étrangers, des laboratoires abroad, or from public or private research centers. publics ou privés. Université BLAISE PASCAL Université d’AUVERGNE N° D.U. 1984 Année 2009 ECOLE DOCTORALE DES SCIENCES DE LA VIE ET DE LA SANTE N° d’ordre 513 Thèse Présentée à l’Université Blaise Pascal Pour l’obtention du grade de Docteur d’Université (Spécialité : Physiologie et Génétique Moléculaire Végétale) Soutenue le 4 décembre 2009 Walid ALFARES Analyses génétiques et moléculaires du locus SKr impliqué dans l'aptitude du blé (Triticum aestivum L.) au croisement avec le seigle (Secale cereale L.) JURY Président Said MOUZEYAR PR, Université Blaise Pascal Rapporteurs Pierre DEVAUX DR, Florimond Desprez Alain GHESQUIERE DR, IRD Examinateurs Nabila YAHIAOUI CR, CIRAD Catherine FEUILLET DR, INRA Pierre SOURDILLE CR, INRA Michel BERNARD DR, INRA UMR1095 INRA/UBP Génétique, Diversité et Ecophysiologie des Céréales Résumé La plupart de variétés élites de blé tendre ne peuvent pas être croisées avec des espèces apparentées ce qui restreint considérablement la base génétique qui peut être utilisée pour l'introgression de nouveaux allèles dans les programmes de sélection. L'inhibition de l'hybridation entre le blé et les espèces apparentées (e.g. seigle, orge) est génétiquement contrôlée. Un certain nombre de QTLs ont été identifiés à ce jour, y compris les gènes Kr1 sur le chromosome 5BL et SKr, un QTL majeur identifié au laboratoire en 1998 sur le bras court du chromosome 5B, tous deux impliqués dans l'inhibition du croisement entre le blé tendre et le seigle. Dans cette étude, nous avons utilisé une population recombinante SSD provenant d'un croisement entre la variété Courtot non croisable et la lignée MP98 croisable pour caractériser l'effet majeur dominant de SKr. Le gène a ensuite été cartographié génétiquement sur la partie distale du chromosome 5BS à proximité du locus GSP (Grain Softness Protein) dont l'homéologue sur le chromosome 5D est impliqué dans la dureté du grain (locus Ha). Les relations de colinéarité avec l'orge et le riz ont été utilisées pour saturer la région de SKr par de nouveaux marqueurs et établir des relations orthologues avec une région de 54 kb sur le chromosome 12L de riz. Au total, 6 marqueurs moléculaires ont été cartographiés dans un intervalle génétique de 0,3 cM, et 400 kb de contigs physiques de BAC ont été établis des deux cotés du gène afin de jeter les bases du clonage positionnel de SKr. De nouvelles populations de grands effectifs ont été développées pour la localisation précise du gène SKr sur les cartes génétiques et physiques. 223 individus d'une population HIF (SSD254.14) ont été testés pour leur aptitude au croisement avec le seigle et génotypés avec les marqueurs proches du gène pour confirmer les données obtenues dans la population de départ. Les résultats montrent que SKr est localisé dans une région hautement recombinante et que les relations entre distances génétiques et distances physiques sont favorables aux dernières étapes de clonage positionnel du gène. Enfin, deux marqueurs SSR complètement liés au gène SKr ont été utilisés pour évaluer une collection de descendances de blé aptes au croisement avec le seigle originaires d'un programme de sélection de triticale primaire. Les résultats confirment l'effet majeur de SKr sur l’aptitude au croisement et l'utilité des deux marqueurs pour introgresser l'aptitude au croisement interspécifique dans des variétés élites de blé tendre. 4 Abstract Although wheat can be crossed with a wide range of related species, most adapted bread wheat varieties are non-crossable because of the failure of pollen to fertilize the ovary, greatly restricting the germplasm that can be used for alien introgression. Inhibition to crossability between wheat (Triticum aestivum L.) and related species such as rye (Secale cereale L.) is genetically controlled. A number of QTL have been identified to date, including Kr1 on chromosome 5BL and SKr, a strong QTL mapped previously in our group at the distal end of chromosome 5BS. In this work, we have used 50 recombinant SSD individuals originating from a cross between the poorly crossable cultivar Courtot and the crossable line MP98 to first characterize the effect of SKr. Two classes of crossability were observed with a 1:1 segregation as expected for a single major gene in a SSD population after 6 generations of selfing, thereby confirming the major dominant effect of SKr on crossability. Additional SSR markers were mapped at the proximal end of the gene on 5BS and new markers were developed from the colinearity with barley and rice. High disruption of colinearity between wheat and rice in the telomeric region of 5BS did not allow to use rice in the first place but the use of a high-density barley EST map allowed us to indentify a new marker at the distal end of SKr and establish a syntenic relationship with rice chromosome 12L in the region. New wheat SSR markers were also derived from a BAC sequence carrying the GSP locus that was identify in the vicinity of the SKr locus. In total, 6 additional markers were mapped at the SKr locus and 400 kb of physical contig were established on both sides of the gene after screening the Chinese Spring BAC library with flanking markers. Sequence comparisons showed that the region spanning SKr in wheat corresponds to a 54 kb region carrying 6 genes on rice chromosome 12L. New high-resolution mapping populations have been developped to achieve the map-based cloning of the SKr gene. Two SSR markers completely linked to SKr were further used to evaluate a collection of crossable wheat progenies originating from a primary triticale program developed in the institute. High association was observed between the crossability and the alleles from the crossable lines at the SKr locus. This confirmed the major effect of SKr on crossability in wheat and provided very useful markers for the efficient introgression of Skr alleles in elite varieties to increase genetic diversity in wheat and triticale breeding programs. 5 6 Remerciements Ce travail a été effectué au sein de l’UMR 1095 INRA-Université Blaise Pascal Génétique, Diversité et Ecophysiologie des Céréales. Je tiens à remercier Gilles Charmet pour m’avoir accueilli au sein de l’unité. Je tiens à remercier également l’Université d’Alep – Syrie qui m’a attribuée la bourse de la thèse dans le cadre de l’accord de la collaboration scientifique Franco-syrien (groupe 7), et je suis très reconnaissant à M. Kinan Darkazanli qui a encadré mes études supérieures du coté syrien. Je voudrais exprimer mes sincères remerciements aux membres du jury, qui ont accepté d’évaluer mon travail de thèse : M. Said Mouzeyar, de l’UBP, qui a accepté de présider le jury de cette thèse. M. Pierre Devaux de Florimond Desprez et M. Alain Ghesquiere de l’IRD qui ont bien voulu être les rapporteurs de ce manuscrit. Mme. Nabila Yahiaoui du CIRAD en tant qu’examinateur de mon mémoire de thèse. Je souhaite exprimer toute ma gratitude à Catherine Feuillet, ma directrice de thèse, qui m’a accueilli dans son équipe et m’a proposé un sujet riche et passionnant. Je la remercie très sincèrement pour l’encadrement scientifique apporté au cours de ces années de thèse. Elle a toujours été disponible, à l’écoute de mes nombreuses questions, et s’est toujours intéressée à l’avancée de mes travaux. Grâce à ses grandes qualités humaines elle m’a aidé à passer tous les caps difficiles en m’encourageant lorsque j’en avais besoin. Encore merci pour l’aide apportée à la rédaction en français de ce rapport. Je voudrais remercier tout particulièrement Michel Bernard, mon co-directeur de thèse, pour son suivi et sa disponibilité. Malgré son départ à la retraite, il est resté à l’écoute de mes nombreuses questions. Merci pour ses conseils, ses qualités humaines et pour le temps qu’il a consacré aux relectures et aux précieuses corrections. Je voudrais exprimer mes profonds respects à Pierre Sourdille qui a répondu avec gentillesse à toutes mes questions durant ces années. Nos discussions scientifiques m’ont apporté une aide précieuse. Merci pour l’aide aux rédactions de l’article et de mon rapport. Encore merci pour avoir accepté d’examiner mon mémoire de thèse. Je remercie Georges Gay et Alain Loussert pour leur aide précieuse lors des cultures en serre nécessaires au développement des populations. Le test du phénotypage a occasionné un grand nombre de castrations et pollinisations. Je dois beaucoup à Georges qui n’a pas hésité à m’apprendre la technique et enrichir mes connaissances par des discussions sur les croisements interspécifiques. Merci à Ludo, Stéphane et Marie Reine qui ont aussi participé au phénotypage 2009.
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