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Universite Paris-Sud UNIVERSITE PARIS-SUD ÉCOLE DOCTORALE : SCIENCES DU VÉGÉTAL Institut de Biologie des Plantes DISCIPLINE : BIOLOGIE THÈSE DE DOCTORAT Soutenance prévue le Vendredi 17/10/2014 par Linda de BONT Importance de l’homéostasie du NAD dans la productivité et la résistance aux stress chez Arabidopsis thaliana Composition du jury : Directeur de thèse : Bertrand GAKIERE Maître de Conférences (Université Paris-Sud) Rapporteurs : François BOUTEAU Maître de Conférences (Université Paris-Diderot) Yves GIBON Directeur de Recherches (INRA Bordeaux) Examinateurs : Guillaume TCHERKEZ Professeur (Université Paris-Sud) Dao-Xiu ZHOU Professeur (Université Paris-Sud) Président du jury : Anis LIMAMI Professeur (Université d’Angers) Thèse préparée au sein de l’Institut de Biologie des Plantes, Unité Mixte de Recherche CNRS – Université Paris Sud, (UMR 8618) RésuméChapitre - Abstract I Le développement et le fonctionnement harmonieux des plantes dépendent de cofacteurs comme le nicotinamide adénine dinucléotide (NAD). Outre ses rôles dans le recyclage rédox, le NAD est aussi impliqué dans des processus de signalisation cellulaire, particulièrement sollicités en situation de stress. Cela fait du NAD l’un des déterminants majeurs de l’homéostasie énergétique des plantes et donc des rendements des cultures. Par une approche de génétique inverse ciblant la L-aspartate oxydase (AO), première enzyme de la voie de biosynthèse du NAD, nous avons obtenu des plantes aux teneurs constitutivement augmentées et diminuées en nucléotides à pyridine. L’étude de ces plantes a permis d’établir que (1) le NAD stimule la croissance, le développement et la productivité des plantes, (2) coordonne les métabolismes photosynthétique, respiratoire et azoté, et (3) intervient dans les mécanismes de résistance aux stress biotiques et abiotiques. Les travaux cités ci-dessus font l’objet d’une valorisation industrielle. Mots-clef : NAD, Arabidopsis thaliana, L-aspartate oxydase, rendement, métabolisme, stress abiotique et biotique. The harmonious development and functioning of plants depend on several cofactors such as nicotinamide adenine dinucleotide. Besides its roles in redox recycling, the NAD is also involved in cellular signalling processes, which are central actors in stress situations. The NAD is thus one of the main determinants of plant energy homeostasis, and therefore of crop yield. Through a reverse genetics approach targeting the L-aspartate oxidase (AO) – the first committed enzyme of NAD biosynthesis – plants with constitutive either increased or decreased levels of pyridine nucleotides have been obtained. A further study of these plants enabled to show that the NAD (1) can improve the growth, development and productivity of plants, (2) coordinates photosynthetic, respiratory and nitrogen metabolisms, and (3) acts upon biotic and abiotic stress resistance mechanisms. This research is being valued industrially. Keywords : NAD, Arabidopsis thaliana, L-aspartate oxidase, yield, metabolism, abiotic/biotic stresses. " Venez et changez quelque chose, ne vous contentez pas de venir parler." Leymah Gbowee Prix Nobel de la Paix, 2011 Remerciements Après trois ans passés à l’Institut de Biologie des plantes, il est grand temps pour moi de remercier tous ceux qui d'une façon ou d'une autre ont contribué à sa réalisation. En premier lieu "mon chef ", Bertrand Gakière. Merci de m'avoir donné ma chance, ce qui était loin d’être gagné, il y a trois ans et demi, et de m'avoir formé. Tu m'as également laissé beaucoup (trop ?) de liberté, ce qui m'a permis de donner des directions très personnelles à ma thèse. La confiance que tu as portée en moi, ton soutien et tes encouragements ont été un facteur déterminant de ma réussite. Merci pour ton aide précieuse tout au long de ces années et plus particulièrement pendant la rédaction de ce mémoire. Travailler avec toi a vraiment été une chance aussi bien d’un point de vue scientifique que personnel. Je tiens à remercier Yves Gibon et François Bouteau de m'avoir fait l'honneur de juger ce mémoire en tant que rapporteurs. Je remercie également Anis Limami, Dao‐Xiu Zhou et Guillaume Tcherkez d'avoir accepté d'examiner mon travail de thèse. Merci à Michel Dron directeur du laboratoire, pour son accueil au sein de l’Institut de Biologie des plantes (IBP), pour sa confiance dans mon projet de thèse et d’avoir mis à disposition les outils et les moyens nécessaires au bon déroulement de ma thèse. Ce travail ne serait pas ce qu'il est sans l'aide Marie Garmier et Sophie Massot, c'est pourquoi je tiens à leur exprimer ici toute ma reconnaissance. Toutes les fois où j'ai eu la chance et le plaisir de travailler à vos cotés, vous avez toujours fait preuve de professionnalisme, de minutie et d'enthousiasme et en plus vous nous avez même pris soin de moi en amenant le petit déjeuner les matins de la toute fin de rédaction. Un grand merci à vous deux. Je tiens à remercier les membres de mon comité de thèse Anne Krapp et Juergen Kroymann pour leur très bon conseil sur le déroulement de leur thèse, leur gentillesse et leur disponibilité. Un très grand merci à Michael Hodges, chef de l’équipe Signalisation Régulation et Interactions Métaboliques. Merci pour tes encouragements et ton soutien pendant ces trois ans. Je remercie également toute les autres membres de l’équipe, Nat ‘ & Céline bien sûr, Mathieu & Younès, Guillaume, Sam & Cyril. Merci également à Jean, Françoise et Chantal, Aline, Martine et Valérie. Merci à tous pour ces bons moments passés. Un grand merci aussi à Jingfang qui va continuer de faire vivre le sujet NAD, merci pour tous les bons moments surtout lors de tes cours d’initiation au chinois. Je te souhaite plein de courage pour ta thèse. Merci énormément à Pierre de m’avoir formée quand j’étais en Master 2, Merci pour ta patience, pour tes conseils judicieux, et aussi pour ton soutien pendant ma dernière année. Mille mercis à Edouard. En plus d’être un super binôme de paillasse, tu as continué à me soutenir et m’encourager une fois parti et ce jusqu’aux corrections final du manuscrit. Merci pour ta bonne humeur, pour ton entrain et pour toutes les bières (et les camomilles) lors des « nocturnes » au labo. Un immense merci aux filles de la plateforme Métabolisme Métabolome. Avec vous toutes j’ai eu la chance d’interagir à l’IBP mais aussi en dehors. Vous avez contribué à un grand nombre de résultats de ma thèse et travailler avec vous a toujours été un grand plaisir. Merci à Françoise pour son aide lors des analyses par GC‐MS, et pour les séances de sport à l’X. Merci à Caroline pour les analyses en d’acides aminés par HPLC et pour les sorties et les footings. Merci à Florence pour les dosages d’hormones par LC‐MS et pour les sorties à la piscine et aux concerts. Merci à Valérie, pour ses aides techniques en informatique, pour son aide lors de mon déménagement et pour le sport en général. Merci à Marlène pour les analyses par EA‐IRMS et pour m’avoir motivée à faire le tri‐relais de la fac, une médaille de bronze ça ne s’oublie pas ! Un grand merci à Marina et Juergen de l’ESE, pour leur présence leur aide et leur conseil sur mon chapitre pucerons. J’en profite également pour remercier Maisera pour sa bonne humeur lors de mes passages à l’ESE. Merci à Anne Krapp de l’INRA de Versailles et à Vivi pour leur aide sur les expériences en hydroponie. Je tiens également à remercier Gabriel Cornic de l’ESE pour son aide sur la mise en place du stress hydrique et pour m’avoir initiée aux mesures de potentiels hydriques. Mes remerciements vont également à Anja Krieger‐Liszkay du CEA pour les mesures de ROS, merci pour ta patience, tes conseils et ta gentillesse. Je remercie également Sébastien de la plateforme IMAGIF pour m'avoir initié aux techniques de microscopie et Séverine de l’IBP qui m’a aidé pour l’utilisation du stéréomicroscope qui m’a donné des conseils pour la rédaction des parties abordant la microscopie. Je tiens également à remercier à tous les membres de l’IBP, surtout Jean‐Paul et Eric, je ne compte plus les fois où j’ai crevé à vélo, où les freins m’ont lâché et autres péripéties. Merci à Véronique pour son aide lors de mes derniers repiquages de plante. Merci à Gilles d’avoir fait en sorte que je puisse réaliser le stress combiné « forte lumière forte température ». Merci à Audrey et Maryelle pour leurs encouragements. Je remercie aussi Catherine, Sophie et Gilles pour les services communs, leur bonne humeur et leurs conseils. Merci également à Delphine, ma collègue du troisième étage de l’IBP, qui est désormais à Paris. Merci à la SFBV pour m’avoir octroyé la bourse « jeunes chercheur » et ainsi permettre de partir en congrès au MPMI à Rhodes et merci à Emmanuelle pour sa bonne humeur lors de ce congrès inoubliable. Durant ces trois années, j’ai eu la chance de former de près ou de loin un certain nombre de stagiaires : Du, Antoine, Kévin & Thomas, Jessica & Quentin, Clémentine et Atem. Merci à vous tous pour votre bonne humeur et pour les bons moments passés. Vous avez tous faits du très bon travail. Faites de votre mieux pour la suite ! Merci à Jacqui et Marianne, directrice et vice‐directrice de l’Ecole doctorale Sciences du Végétal 145. Merci à toutes les deux pour votre gentillesse, pour vos encouragements. Merci aussi pour votre aide pour l’organisation, le bon déroulement des PhD days et pour toutes les formalités administratives. Marianne, tu étais également ma tutrice pour le monitorat. Merci beaucoup pour tes conseils et ta présence. Un grand merci à mes collègues enseignant, Marianne, Céline Charon, Dara, Moussa et Yves qui m’ont formé lors de ma première année de monitorat.
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