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Document 567595.Pdf Thèse présentée par Frédéric Péréfarres Pour obtenir le grade de Docteur de l’Université de La Réunion Discipline : Sciences Ecole Doctorale Interdisciplinaire E.D.I n° 0445 UMR Peuplements Végétaux et Bio-agresseurs en Milieu Tropical CIRAD – Université de La Réunion Epidémiologie et contrôle durable des Begomovirus chez la tomate Soutenance le 21 Février 2012, devant le jury composé de : POUSSIER Stéphane Professeur, Université de La Réunion Président MOURY Benoît Chercheur HDR, INRA Avignon Rapporteur MORIONES Enrique Chercheur HDR, CSIC Málaga Rapporteur REYNAUD Bernard Chercheur HDR, CIRAD Réunion Directeur de thèse Remerciements Voilà il est temps de conclure….3 années viennent de passer et cette petite aventure touche à sa fin. Faire un bilan de ces 3 années demandera certainement plus de temps et de recul que ces quelques lignes. Cette tranche de vie aura été stimulante, passionnante, stressante, fatigante par moment mais tellement enrichissante. Cette thèse n’aurait pas pu se faire toute seule donc je tiens à vous remercier pour m’avoir encouragé, soutenu, aidé et supporté quand j’avais la tête des mauvais jours. Merci à Bernard Reynaud d’être un super directeur d’unité et de faire tout son possible pour qu’on puisse avoir de bonnes conditions de travail au 3P. Je tiens à remercier Stéphane Poussier, Benoit Moury et Enrique Moriones d’avoir accepté de composer mon jury de thèse. Un grand merci bien sûr à Jacques Dintinger pour avoir monté, initié et suivi ce projet sur la résistance aux bégomovirus. Merci à Jackypédia pour ta culture, tes connaissances, ton expérience et tes conseils. Cette thèse n’aurait pas été la même sans lui, et je remercie sincèrement Jean Michel Lett pour tout ce qu’il a apporté à cette thèse. Merci pour tes conseils, ton aide précieuse (sacré NS3), ton soutien infaillible, les tartes aux fraises, les bonbons et autres sucreries. A lui aussi je lui dois beaucoup donc un grand merci à Pierrot Lefeuvre pour son exigence, sa rigueur, son œil affuté, ses jeux sur le net, les liens chronophages, les astuces au distributeur. Ca y est je n’ai plus d’excuses pour ne pas aller grimper avec vous… Merci aux participants de mes comités de thèse et autres personnes que j’ai sollicitées pendant ces 3 années pour mener à bien mon travail : Laurence Albar, Carole Caranta, André Moretti, Gaël Thébaud, Frédéric Fabre, Samuel Nibouche, Jay Scott, Grégori Bonnet. Un grand merci également aux membres ou affiliés de la Bégomo-Team, Mumu Hoareau, Magali Thierry, Alexandre de Bruyn, Hélène Delatte, Nathalie Becker, Julie Villemot, Vincent Lepitre, Martial Grondin et Didier Fontaine. Merci pour votre aide précieuse pendant ces trois années et pour la bonne ambiance qui règne dans l’équipe. La Bégomo-Team ne serait pas la même sans eux donc un grand merci à nos « Africains » Mireille, Fidèle et Innocent. Merci pour les super souvenirs d’Aussois : le ski, la raclette, le babyfoot, le Génépi…on s’en souviendra pendant longtemps ! Si je suis là c’est un peu grâce à lui aussi donc un grand merci à Pierre Yves Teycheney pour m’avoir ouvert les portes du CIRAD il y a bien longtemps maintenant. Quand tu veux on repart échantillonner à travers la Guadeloupe ! Merci aux amis de toujours Benoît, Dorian, Nicolas et Yoan. Merci d’être des mecs exceptionnels et des supers vacances passées avec vous ici ou ailleurs pendant ces 3 dernières années (Ouvéa quand tu nous tiens) Ces 3 années ont été rythmées par le travail mais pas que ça donc un grand merci à Grand Seb et Marie pour leur amitié, à Mathieu et l’autre Grand Seb pour les soirées épicuriens Ligue des Champions, au Club des 80, à Nico, à la FARRE Team, à Dr Kayser Wicker, Dr Basco-catalan (« sempre endavant »), au métronome Christian, à Miss Daphné, à Super Ja, à Ludo, à Elisabeth, à Lan. Une pensée aussi aux collègues de promo Rodolphe et Cathy et bon courage aux autres thésards du 3P Aline, Stéphanie, Toulassi, Carine, Alex, Alice. Merci à Isabelle, Lionel, Olivier, Laurent, Les Michels et l’ensemble du personnel de faire du 3P un endroit où il fait bon travailler. Petite dédicace pour les sponsors : Radio Nova, Deezer, Les Nuits de Lavige, Pascale Clark, Jean Claude Ameisen, l’After Foot, Moscato Show, l’Afrique Enchantée, Nicolas Demorand, Patrick Cohen, Jehro, Fink, Kit-Kat, Le chinois de Terre Sainte, le café philo « Chez Roger ». Merci à Stéphanie de m’avoir soutenu et encouragé ces derniers mois. Allez dans quelques temps ça sera toi. Suerte y animo !!! Enfin un grand merci à ma famille pour accepter mes choix et pour le soutien que vous m’apportez. Bienvenu à Joan, t’inquiète Tonton viendra bientôt te voir !! TABLE DES MATIERES 1. Introduction......................................................................................1 2. CHAPITRE 1 Mieux appréhender la diversité virale pour mieux la contrôler………...............................................................................34 2.1. Article 1........................................................................................35 A novel synthetic quantification standard including virus and internal report targets: application for the detection and quantification of emerging begomoviruses on tomato 2.2. Article 2........................................................................................44 Differential severity between the two main emerging strains of TYLCV 3. CHAPITRE 2 Dynamiques épidémiologiques des populations de TYLCV-Mld et TYLCV-IL à la Réunion………………………………………………….50 3.1. Article 3 Long-term viral competition monitoring: a case of epidemiological rescue 4. CHAPITRE 3 Spectre d’action des principaux gènes de résistance aux bégomovirus et caractérisation d’une résistance récessive……………………………………………………………………………………………………..…72 4.1. Article 4 Resistance to begomoviruses: phenotype and spectrum of action of a novel recessive resistance and the main resistance genes 5. CHAPITRE 4 Exploration de la résistance quantitative conférée par LA1777 aux deux souches émergentes du TYLCV……………………………………………………………………………………………………………..94 5.1 Article 5 Characterization of a quantitative resistance from Solanum habrochaites LA1777 to two emerging strains of TYLCV (TYLCV-Mld and TYLCV-IL). 6. Discussion……………………………………………………………………………………………………109 7. Références bibliographiques……………………………………………………………….121 8. Annexes…………………………………………………………………………………………………………..……141 IInnttrroodduuccttiioonn Introduction 1. Contexte général Depuis les premières descriptions du SIDA au début des années 1980, de nombreuses études ont été réalisées sur les causes et les conséquences de l'émergence ou de la ré-émergence des maladies virales. Les virus émergents sont définis comme ceux qui sont récemment apparus ou ceux dont les populations ont récemment augmenté en prévalence, en pathogénicité et/ou en répartition géographique (Holmes and Rambaut 2004 ; Jones 2009). Ce phénomène a été mis en avant ces dernières années par les épidémies virales telles que celles associées aux maladies émergentes de l’Homme (SRAS, Holmes and Rambaut 2004 ; chikungunya, Schuffenecker et al. 2006 ; grippe HIN1, Smith et al. 2009), des animaux (fièvre aphteuse, Grubman and Baxt 2004) et des plantes (mosaïque du manioc, Patil and Fauquet 2009 ; panachure jaune du riz, Fargette et al. 2006 ; enroulement foliaire et jaunissement de la tomate, Moriones et al. 2011). Même s’il est toujours difficile de prédire quel virus représente la plus grande menace d’émergence, quand et où il va survenir, des progrès importants ont été réalisés dans l’identification des facteurs écologiques et moléculaires associés à l’émergence (Fargette et al. 2006; Holmes and Rambaut 2004; Jones 2009; Moriones et al. 2011; Smith et al. 2009; Woolhouse et al. 2001). Alors que les maladies émergentes chez l’Homme et les animaux sont généralement causées par des virus à ARN, la plupart des émergences virales chez les plantes sont liées à des virus ADN du genre Begomovirus transmis par l’aleurode Bemisia tabaci (Seal et al. 2006). Le potentiel important d'évolution de ces virus à ADN, leur prévalence en augmentation permanente et l'existence de biotypes invasifs de leur insecte vecteur B. tabaci, font de ces virus un modèle idéal pour la description et la compréhension de l'émergence virale et des mécanismes sous-jacents. Les récentes avancées sur les moteurs de l’évolution des bégomovirus et les facteurs écologiques associés à leur émergence seront autant d’atouts pour la compréhension des dynamiques des populations virales et pour le développement de stratégies de contrôle. 1 Introduction 2. Les acteurs : Begomovirus et Bemisia tabaci a. La famille des Geminiviridae La famille des géminivirus présente une très grande diversité en termes d’organisation génomique, de gamme d’hôtes et d’insectes vecteurs. Sur la base de ces propriétés, les géminivirus ont été divisés en quatre genres dont le nom dérive du membre type : Begomovirus, Curtovirus, Mastrevirus et Topocuvirus. Les Mastrevirus (infectant principalement des plantes monocotylédones et transmis par cicadelle) et les Begomovirus (infectant des dicotylédones et transmis par aleurode) sont les genres les mieux caractérisés (Pour revue Rojas et al. 2005). L’espèce type du genre Begomovirus est le Bean golden mosaic virus, virus de la mosaïque dorée du haricot (Van Regenmortel et al. 2000). Les bégomovirus sont responsables de viroses sur de nombreuses cultures d’importance économique et sociale dans le monde entier et particulièrement en zones tropicales, notamment sur le manioc (Manihot esculenta), le haricot (Phaseolus vulgaris), le coton (Gossypium
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