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Couplage Entre Introduction Et Réparation Des Cassures Double Brin Pendant Les Réarrangements Programmés Du Génome Chez Paramecium Tetraurelia

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Couplage Entre Introduction Et Réparation Des Cassures Double Brin Pendant Les Réarrangements Programmés Du Génome Chez Paramecium Tetraurelia Centre de génétique moléculaire Bâtiment 26 1 avenue de la Terrasse 91190 Gif sur Yvette Thèse de doctorat de l’université Paris Sud XI Ecole doctorale : Gènes, Génomes, Cellules Présentée par Antoine MARMIGNON Couplage entre introduction et réparation des cassures double brin pendant les réarrangements programmés du génome chez Paramecium tetraurelia. Soutenance le 27 septembre 2013 Devant le jury composé de : Président du jury : Jean Cohen Rapporteur : Jean Baptiste Charbonnier Rapporteure : Gaelle Legube Examinatrice : Ariane Gratias-Weill Directrice de thèse : Mireille Bétermier 1 2 Remerciements On m’avait dit que les remerciements s’écrivaient en dernier, tout à la fin, et que c’était très facile, le tout étant de ne pas provoquer de crise diplomatique en oubliant quelqu’un. Rien de tel pour nous mettre la pression, à moi et ma mémoire de poisson rouge. J’ai déjà du mal à retenir les dates d’anniversaire de ma famille alors… pour tout ceux que je vais oublier, car il y en aura probablement, je m’excuse par avance et vous remercie mille fois également. En premier lieu, je tiens à remercier les membres de mon jury qui ont accepté de juger mon travail de thèse. Certains ont eu besoin d’être convaincu. Pour obtenir leur accord, je me suis engagé à ne pas terminer la soutenance par un paint-ball endiablé ou un jeu de rôle moyenâgeux. Je tiendrais promesse. Je souhaite également remercier les organismes qui ont financé ma recherche, le ministère d’abord, puis la fondation ARC. L’idéal serait que la fondation ARC reste dans ces bonnes dispositions et porte un regard bienveillant sur mes futures demandes d’argent, ca faciliterait grandement la poursuite de ma carrière dans un futur très très proche. S’il est une personne que je n’oublierai pas dans ces remerciements, c’est bien Mireille… Mais revenons un peu dans le temps d’abord. Et intéressons nous à moi, étudiant de M2, il y a quelques années. Je dois faire un séminaire bibliographique dans le cadre de ma formation à l’université. Manque de bol (ou plutôt coup de bol) le sujet choisi dans la liste est trop plébiscité et comme je suis bonne pate, j’accepte un autre sujet, délaissé jusqu’ici. Cela 3 concerne les réarrangements programmés du génome et les marques épigénétiques chez les ciliés, encadré par Sandra Duharcourt (C’est ainsi que j’ai fait mes premiers pas dans le monde des ciliés. Et non ce n’est pas encore Mireille, mais merci Sandra pour m’avoir permis de mettre le pied dans la porte). Je trouve le système fascinant. Plus tard je devrai trouver un sujet de thèse, à l’occasion d’une discussion avec Jean-Luc Ferat au deuxième étage du CGM, je parle de ce que j’ai fait pendant le semestre, et il m’annonce qu’au rez de chaussée du bâtiment, Mireille Bétermier travaille sur les réarrangements programmés du génome de la paramécie (Merci Jean-Luc pour le tuyau). Je descends donc au rez de chaussée, passe la tête par la porte de la pièce 38 et dit que je cherche un stage avec une possibilité de thèse après. Mireille, que je rencontre pour la première fois, me répond que là tout de suite elle est occupée mais me propose de revenir le lendemain même heure pour en discuter. Ce fut le début de mon aventure dans la paramécie et je ne regrette pas. Etre encadré par Mireille, c’est avoir une chef dynamique, patiente, toujours disponible, investie, rigoureuse, et profondément humaine (La liste est plus longue mais j’arrive déjà au bout de la première page…). C’est certainement grâce à elle et à la bonne ambiance qui règne dans le laboratoire, grâce à tous ses membres, que venir au labo le matin, et même la nuit, et le week-end (oui oui) pour pouponner des paramécies ne fut jamais une corvée. Donc Mireille, Aurélie, Céline, Jacek, Emeline, Nathalie, Julien, Vinciane, et toutes les personnes qui sont passées au labo ou qui font ou ont fait partie de l’aile bisounours pendant ces quatre années, merci beaucoup. Le monde de la paramécie est une petite communauté, riche en interactions, humaines et scientifiques, où il est agréable de travailler. Ainsi je remercie également tous les « gens de Paris » pour leur retours, leurs idées, leurs conseils et leur présence. 4 Puisque je parle d’environnement, intéressons nous au CGM, le Centre de Génétique Moléculaire pour ceux à qui cet acronyme ne signifie rien. Je ne peux pas faire une liste exhaustive de tous les gens que j’y ai croisés et que je tiens à remercier, mais rien n’aurait été possible sans chacune de ses parties, les dames de la laverie toujours souriantes ; le secrétariat qui gère beaucoup mieux l’administratif que moi, heureusement qu’ils sont tous là ; l’atelier et le magasin pour gérer les urgences ; l’ensemble des scientifiques qui font du bâtiment un incubateur à idées ; l’association étudiante, qui permet de tenir le coup dans l’adversité ; et la direction qui gère tout ca, si ma présence au Conseil de Laboratoire m’a appris quelquechose, c’est que diriger un institut de recherche n’a rien d’une partie de plaisir, je suis bien content de ne pas être en charge. Bref, à tous ces gens merci. A tous mes amis hors du monde de la biologie, Charles, Matthieu, Louis, Hadrien, Jean-Yves, Jonathan, Jérôme, Sarah, Charlotte, Blandine, pour m’avoir aidé à tenir le coup (oui oui je sais j’ai dit que venir au labo n’était pas une corvée, mais je n’ai jamais dit qu’être thésard était facile), qui me changent les idées, m’emmènent voir des navets au cinéma pour déconnecter les neurones (pas que, je vais aussi voir des choses plus intellectuelles), me permettent d’avoir une vie en dehors du labo, je tiens à vous dire merci, merci beaucoup. A ma famille, je sais que le cursus universitaire n’a pas toujours semblé à mes parents la voie idéale pour me garantir une vie longue et prospère. Mais c’est grâce à eux que j’ai développé ma curiosité de tout, mon goût pour la science et les questions existentielles qui m’ont fait m’épanouir dans la pratique de la recherche. JE NE REMERCIE PAS, les paramécies asynchrones qui m’ont valu de nombreuses nuits de veille pour rien au laboratoire, à les couver d’un regard implorant pour qu’elles aient 5 la bonne volonté de démarrer les processus sexuels toutes en même temps. De toute façon, celles qui n’ont pas obéi ont été bien punies… je les ai javellisées sans remords (C’est le coté obscur de la Science). JE NE REMERCIE PAS, ma voiture. Si l’avoir m’a permis de continuer à pratiquer l’équitation dans mon club préféré tout au long de ma thèse, la tentation de l’utiliser abusivement a été trop forte et se ressent sur mon tour de taille (On pourrait croire que c’est de la mauvaise foi et que je devrais ne m’en prendre qu’à moi ; on aurait raison). Ca y est l’inspiration me fait défaut… Encore une fois, à tous ceux que j’oublie, ce n’est pas que vous n’avez pas compté ; blâmez ma mémoire défaillante, et recevez tout de même tous mes remerciements. 6 Table des matières REMERCIEMENTS ......................................................................................................................... 3 TABLE DES MATIERES .................................................................................................................... 7 LISTE DES FIGURES........................................................................................................................ 9 LISTE DES ABREVIATIONS ............................................................................................................. 12 INTRODUCTION .............................................................................................................. 15 LES CASSURES DE L’ADN ............................................................................................................. 17 REPARATION DES CASSURES DOUBLE BRIN DE L’ADN ........................................................................ 19 La recombinaison homologue : ........................................................................................ 21 Les principaux acteurs de la recombinaison homologue ......................................................... 21 Mécanisme programmé utilisant la recombinaison homologue : la méiose ........................... 25 Les voies apparentées à la recombinaison homologue ........................................................... 27 Synthesis Dependent Strand Annealing (SDSA) ................................................................... 27 Single Strand Annealing (SSA) .............................................................................................. 27 La voie de Non Homologous End Joining : ........................................................................ 31 Les principaux acteurs du NHEJ ................................................................................................ 31 Mécanisme programmé utilisant la voie du NHEJ : la recombinaison V(D)J ........................... 39 Les voies apparentées au NHEJ ................................................................................................ 41 Quelle voie choisir pour réparer une cassure double brin de l’ADN? ............................... 43 Choisir de ne pas réparer .................................................................................................. 47 Le cas des télomères ................................................................................................................ 47 LA PARAMECIE COMME ORGANISME MODELE. ................................................................................. 49 La paramécie, sa vie. .......................................................................................................
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