Rôle Du Système De Sécrétion De Type III SPI-1 Et Des Mégaenzymes

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Rôle Du Système De Sécrétion De Type III SPI-1 Et Des Mégaenzymes UNIVERSITE MONTPELLIER II SCIENCES ET TECHNIQUES DU LANGUEDOC THESE Pour obtenir le grade de DOCTEUR DE L’UNIVERSITE DE MONTPELLIER II Ecole doctorale : SIBAGHE Discipline : Biologie Intégrative des Plantes Présentée et soutenue publiquement par Mélanie MARGUERETTAZ Le 7 décembre 2010 Rôle du système de sécrétion de type III SPI-1 et des mégaenzymes NRPS dans le cycle de vie de Xanthomonas albilineans, l’agent causal de l'échaudure des feuilles de la canne à sucre JURY : M. Guy CONDEMINE (Directeur de Recherche, CNRS Lyon), Rapporteur. M. Alain GIVAUDAN (Directeur de Recherche, INRA Montpellier), Président du jury. *,ime Marie-Agnès JACQUES (Chargé de Recherche, INRA Angers), Rapporteur. Mme Monique ROYER (Directeur de Recherche, CIRAD Montpellier), Directeur de thèse. Remerciements Je tiens à remercier Monique ROYER qui a encadré ce travail. Elle a toujours été de bons conseils et disponible. Mes remerciements vont également à Marie-Agnès Jacques et Guy Condemine pour avoir accepté d'évaluer ce travail, ainsi qu'à Alain Givaudan pour m'avoir suivie au cours de cette thèse. Je remercie la région Languedoc-Roussillon et le Cirad pour avoir co-financé cette thèse. Merci aux membres de l'équipe 3 : Philippe, Isabelle, Stéphane et Sandrine, pour m'avoir accueillie durant ces 3 ans, ainsi que pour leurs conseils, leur aide et leur soutien. Et Philippe on est d'accord : s'il n'y a pas d'albicidine ce ne sont pas de vraies lignes blanches!! Et merci à Jérôme pour son travail sur le Spy One et sa bonne humeur au labo. ) Une dédicace spéciale pour le groupe des "Tueurs de canne" avec lesquels j'ai passé de lonnnngues journées. Marie-Jo et Sandrine, heureusement que vous avez été là pour couler ces milliers de boîtes! Vous êtes définitivement les reines de la logistique. Marc, bien que tu sois parti loin, j ’ai énormément apprécié nos discussions philosophiques dans la serre souvent suivies de tes disparitions mystérieuses. Et enfin Damien, qui en venant renforcer le groupe, n'a pas hésité une seconde à faire couler la sève des pauvres cannes pour la science. A mes formidables collègues de bureau! Il faut bien avouer que j'ai vraiment eu de la chance. Philippe et Laurence, nos discussions mi-scientifiques, mi-potin me manquent encore...et merci Bernard pour tes dictées corsées. Heureusement que le Potager est là pour prolonger ces bons moments. Audrey et Pierre-o, c'est vrai on a formé un bureau de choc, c'était trop court! Quoiqu'il en soit je vous remercie tous les quatre pour votre soutien au cours de cette thèse (et les pauses sandwichs). Audrey, prépare ta poupée, la séance d’exorcisme va commencer ! ! Petite pensée pour mon frère de galère qui se reconnaîtra...We did it! Ainsi que pour Mauricio et ses conseils avisés. 3 Je tiens à remercier mes collègues de BGPI pour la bonne humeur. Amandine et Joan pour les petites pauses entre deux manips, Emmanuel pour ses histoires plus drôles les unes que les autres. Et tous les autres trop nombreux à citer. Merci, merci, merci à tous mes amis qui sont toujours là pour moi. Cora ! La vache ça va bientôt faire 15 ans qu'on se connaît, faudra qu'on pense à se faire engager dans le même labo un jour! Vraiment merci d'être là même depuis toutes ces années. Merci à toi aussi Marion! La Rebelle de médecine! La fétarde de tous les instants! Ma psy aussi... Enfin merci à mes deux réunionnais préférés Willy et Tony. Je terminerai en remerciant ma famille, ça y est c'est fini! Vous m'avez soutenu toutes ces années et c'est grâce à vous aussi, et surtout, que j'en suis là. Enfin Vincent, même si tout n'a pas été rose, tu es toujours là pour m'épauler, merci. Cette thèse a été sponsorisée par RedBull, Picard et Rivière du Mât. 4 Table des matières Liste des figures...................................................................................................................10 Liste des tableaux................................................................................................................12 Abréviations......................................................................................................................... 13 Objectifs de la thèse........................................................................................................... 15 Chapitre 1. Synthèse bibliographique............................................................................21 1 Les bases moléculaires des interactions plante-bactérie....................................... 23 1.1 Les mécanismes de défense chez les plantes............................................................ 23 1.1.1 Les défenses préformées................................................................................23 1.1.2 Les défenses inductibles................................................................................. 24 1.1.2.1 La reconnaissance de l'agent pathogène.......................................24 1.1.2.2 Les réactions de défense suite à la reconnaissance d ’un microorganisme............................................................................................... 24 1.1.2.3 Le modèle en zigzag..........................................................................26 1.2 Les principaux facteurs de pathogénie chez les bactéries...................................... 26 1.2.1 Les enzymes de dégradation de la paroi cellulaire......................................27 1.2.2 Les exopolysaccharides.................................................................................. 27 1.2.3 Les structures impliquées dans la mobilité..................................................29 1.2.3.1 Le flagelle...........................................................................................29 1.2.3.2 Le pilus de type IV.............................................................................30 1.2.4 Les adhésines non fibrillaires........................................................................ 31 1.2.5 Les systèmes de captation du fer...................................................................32 1.2.6 La signalisation cellulaire..............................................................................33 1.2.6.1 Gènes rpf............................................................................................33 1.2.6.2 Gènes pig............................................................................................35 1.2.7 Les pompes d’efflux........................................................................................35 1.2.8 Les systèmes de sécrétion de protéines........................................................ 37 1.2.8.1 Le système de sécrétion de type 1....................................................39 1.2.8.2 Le système de sécrétion de type II...................................................40 1.2.8.3 Le système de sécrétion de type III .................................................42 1.2.8.4 Le système de sécrétion de type IV..................................................44 1.2.8.5 Le système de sécrétion de type V...................................................46 1.2.8.6 Le système de sécrétion de type VI................................................. 48 5 1.2.9 Les effecteurs de virulence ou d’avirulence sécrétés par les bactéries Gram négatif phytopathogènes..........................................................................................49 1.2.9.1 Les effecteurs de type III.................................................................. 49 1.2.9.2 Un exemple d ’effecteur d ’avirulence sécrété par le système de sécrétion de type 1 ...........................................................................................51 1.2.9.3 Les effecteurs sécrétés par des pompes d ’efflux...........................52 2 Modes de transmission et de dissémination des bactéries phytopathogènes.... 53 2.1 Les bactéries phytopathogènes dans l'environnement..............................................53 2.1.1 Le sol.................................................................................................................53 2.1.2 L'eau..................................................................................................................53 2.1.3 Les repousses et les plantes adventices........................................................ 55 2.2 La colonisation épiphyte................................................................................................ 55 2.3 La transmission par le biais d'insectes........................................................................57 3 La colonisation du xylème par les bactéries phytopathogènes............................ 59 3.1 Morphologie du xylème.................................................................................................. 59 3.2 Composition de la sève brute.........................................................................................61 3.3 Les bactéries colonisatrices du xylème........................................................................61 3.3.1 Xanthomonas oryzae pv. oryzae................................................................... 61 3.3.2Xanthomonas campestris pv. campestris..................................................... 62 3.3.3 Ralstonia solanacearum................................................................................. 63 3.3.4 Xanthomonas axonopodis pv. vasculorum...................................................64 3.3.5 Xylella fastidiosa............................................................................................. 65 3.3.6Leifsoniaxyli subsp. x y li...............................................................................
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