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Étude De L'interaction Entre Medicago Truncatula Et Verticillium Alfalfae

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Étude De L'interaction Entre Medicago Truncatula Et Verticillium Alfalfae En vue de l'obtention du DOCTORAT DE L'UNIVERSITÉ DE TOULOUSE Délivré par : Institut National Polytechnique de Toulouse (INP Toulouse) Discipline ou spécialité : Interactions plantes-microorganismes Présentée et soutenue par : M. MAOULIDA TOUENI le lundi 17 novembre 2014 Titre : ETUDE DE L'INTERACTION ENTRE VERTICILLIUM ALFALFAE ET MEDICAGO TRUNCATULA Ecole doctorale : Sciences Ecologiques, Vétérinaires, Agronomiques et Bioingénieries (SEVAB) Unité de recherche : Ecologie Fonctionnelle (ECOLAB) Directeur(s) de Thèse : MME MARTINA RICKAUER MME CECILE BEN Rapporteurs : Mme DIANA FERNANDEZ, IRD MONTPELLIER M. PHILIPPE REIGNAULT, UNIVERSITE DU LITTORAL COTE D'OPALE Membre(s) du jury : 1 M. PHILIPPE REIGNAULT, UNIVERSITE DU LITTORAL COTE D'OPALE, Président 2 Mme CECILE BEN, INP TOULOUSE, Membre 2 Mme DIANA FERNANDEZ, IRD MONTPELLIER, Membre 2 Mme MARTINA RICKAUER, INP TOULOUSE, Membre AA.. INTROINTRODUCTIONDUCTION ............................................................................................................................................................................................................................................................................ 777 I. Les maladies des Plantes........................................................................................................ 9 1. Introduction : les maladies des plantes ............................................................................ 9 2. Les pathogènes vasculaires .............................................................................................. 13 a) Les bactéries ................................................................................................................. 13 b) Les Oomycètes .............................................................................................................. 13 c) Les champignons ......................................................................................................... 15 3. Les maladies de flétrissement vasculaires ...................................................................... 17 II. La verticilliose ...................................................................................................................... 23 1. Verticillium sp. : l’agent causal de la verticilliose .......................................................... 25 2. Cycle de la maladie .......................................................................................................... 33 3. Mécanismes de transmission et dispersion .................................................................... 35 4. Importance économique et écologique ........................................................................... 37 5. Méthodes de contrôles de la maladie .............................................................................. 39 III. Bases moléculaires de l'interaction entre Verticillium et l'hôte ................................... 39 1. Facteurs d'adaptation au xylème ............................................................................ 39 2. Effecteurs et facteurs de virulence ......................................................................... 45 3. Types d’interaction observés en réponse à Verticillium sp. ........................... 49 IV. Mécanismes de défense des plantes, notamment contre la verticilliose ...................... 51 1. Défenses préformées et induites ...................................................................................... 51 2. La perception des pathogènes ......................................................................................... 53 3. Les mécanismes de défense physico-chimiques ............................................................. 65 4. Le rôle des molécules de signalisation dans la résistance, les phytohormones ........... 75 5. Le rôle des microARN dans la régulation des gènes des mécanismes de défenses..... 89 6. Résistance contre la verticilliose ..................................................................................... 93 a) Résistance Ve1-dépendante chez la tomate ...................................................... 95 1. Structure de Ve1 et Ve2 ........................................................................................... 95 2. Signalisation de la résistance Ve1-dépendante ....................................................... 97 3. L'effecteur Ave1 ........................................................................................................ 99 b) La résistance Ve1-dépendante est une PTI ...................................................... 101 V. La verticilliose de la luzerne, modèle d'études et objectifs de la thèse .......................... 103 1. La luzerne, Medicago sativa L. ................................................................................. 103 2. La verticilliose chez Medicago sativa L. ....................................................................... 107 3. M. truncatula, plante modèle des légumineuses ........................................................... 111 4. M. truncatula est une plante hôte de souches de Verticillium isolées de luzerne ..... 115 5. Objectifs de la thèse ....................................................................................................... 119 1 B. MATMATERIELMATERIELERIEL ET METHODES ................................................................................................................................................................................................................ 123123123 I. MATÉRIEL BIOLOGIQUE ET CULTURE.................................................................. 125 1. Verticillium ...................................................................................................................... 125 2. Matériel végétal .............................................................................................................. 125 II. BIOESSAIS......................................................................................................................... 129 1. Inoculation et notation des symptômes ........................................................................ 129 2. Traitement hormonaux.................................................................................................. 131 3. Analyse statistique des données phénotypiques .......................................................... 133 a) Développement de la maladie ................................................................................... 133 b) Poids frais et secs racinaires et foliaires .................................................................... 133 1. Extraction d'ADN .......................................................................................................... 133 2. PCR ................................................................................................................................. 137 3. Quantification de l'ADN de Verticillium par qPCR ................................................... 137 4. Extraction d'ARN .......................................................................................................... 139 5. qRT-PCR pour l’étude d’expression de gènes ............................................................ 141 a) Traitement à la DNAse ............................................................................................... 141 b) Synthèse des ADN complémentaires ......................................................................... 141 c) PCR quantitative......................................................................................................... 143 I. OBSERVATIONS MICROSCOPIQUES........................................................................ 145 II. ANALYSES BIO-INFORMATIQUES ............................................................................ 147 1. Identification des gènes différentiellement exprimés .................................................. 147 2. Construction du réseau de coexpression de gènes par la méthode Weighted Gene Co- expression Network Analysis (WGCNA). ............................................................................ 149 3. Annotation fonctionnelles .............................................................................................. 149 a) Analyse des données avec le logiciel MapMan ......................................................... 149 b) Analyse des GO term ................................................................................................. 151 CC..C. RÉSURÉSULTATSLTATSRÉSULTATS .................................................................................................................................................................................................................................................................................................... 153153153 I.II.. EtuEtudede du mécanisme d'infection de M. truncatula par Verticillium alfalfae ..................... 155 1.11.. CarCaractérisationactérisation moléculaire de la souche V31V31----22 et présentation du pathosystème. ..... 155 2.22.. IntInteractioneraction entre M. truncatula et V31et V31-V31---22 en conditions hydroponiques ...................... 157 3.33.. ÉtuÉtudede comparative de la colonisation racinaire au cours d’une interaction compatible et incompatible entre M. truncatula et V. alfalfaeV. alfalfae V31V31V31-V31---2.2.2.2. ......................................................... 159 a) Sélection d’une souche monosporique exprimant la GFP. ........................................ 163 b) Étude
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