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Université Pierre Et Marie Curie Université Pierre et Marie Curie École Doctorale Complexité du Vivant Centre Scientifique de Monaco / Département de Biologie Marine Équipe de Physiologie et Biochimie Approches physiologique et moléculaire de la calcification chez le corail rouge de Méditerranée Corallium rubrum par Carine LE GOFF Thèse de doctorat en Sciences de la vie Directrice de thèse : Sylvie Tambutté Co-encadrant : Philippe Ganot Présentée et soutenue publiquement le 14 Décembre 2016 à Monaco Devant un jury composé de : Dr. Daniel Vielzeuf Rapporteur Dr. Catherine Lorin-Nebel Rapporteur Pr. François Lallier Examinateur Dr. Stéphanie Barnay-Verdier Examinateur Dr. Clemente Capasso Examinateur Dr. Georges Carle Examinateur Dr. Sylvie Tambutté Directrice de thèse Dr. Philippe Ganot Co-encadrant À mes parents... REMERCIEMENTS Remerciements Les premières lignes de ce manuscrit sont souvent, de façon paradoxale, les dernières à être rédigées. Et lorsque l’on arrive à cette étape, qui n’est pas la plus facile, cela signifie véritablement que la thèse se termine. Je voudrais bien évidemment remercier de façon générale toutes les personnes qui ont contribué, de près ou de loin, à la réalisation de cette thèse. Ces trois années auront été une véritable aventure humaine, personnelle et professionnelle. En premier lieu, je voudrais remercier les membres de mon comité de thèse, Paola Furla, Georges Carle et Jean-Yves Sire. Ce fut un plaisir de vous présenter l’évolution de mes travaux et d’avoir pu échanger avec vous. Je tiens également à remercier les membres du jury de thèse, Daniel Vielzeuf et Catherine Lorin-Nebel pour avoir accepté d’être rapporteurs de ce travail, ainsi que François Lallier, Clemente Capasso, Stéphanie Barnay-Verdier et Georges Carle en leur qualité d’examinateurs. Je souhaite remercier le Pr. Patrick Rampal et Mme Corinne Gaziello, respectivement Président et Secrétaire Générale du Centre Scientifique de Monaco, pour m’avoir permis de réaliser cette thèse au sein de l’équipe de Physiologie et Biochimie du laboratoire. Je remercie également le Pr. Denis Allemand, Directeur du Centre Scientifique de Monaco, pour les échanges que nous avons eus, pour son enthousiasme et ses connaissances qu’il aime partager, dans le domaine de la Science et bien au-delà. Mes remerciements vont ensuite tout naturellement à mes encadrants, Sylvie Tambutté et Philippe Ganot, ainsi qu’aux autres membres de l’équipe de Physiologie et Biochimie. Merci Sylvie pour tes conseils, tes relectures, et tes corrections. Ce fût un plaisir de partager avec toi les résultats obtenus au cours de cette thèse. Le corail rouge a désormais moins de secrets pour nous. Philippe, tu auras su m’encadrer par ton côté « tyrannique» et « grand frère » (selon tes propres mots !). Egalement partenaire graphiste, malgré nos différents pour le choix des couleurs... Je te passe la main (verte ou rouge ?) pour prendre soin de ces petites colonies et boutures. Merci Didier pour tes connaissances notamment en biologie moléculaire, mais également pour les cours matinaux de Japonais et de Russe, et pour avoir essayé de parfaire ma culture musicale et cinématographique. J’espère que d’autres bretons passeront par ici pour peut-être, réussir à te faire apprécier la pluie. Merci Alex pour ton aide précieuse et tes conseils avisés lors des expériences au confocal, ce microscope qui m’a paru si difficile à maîtriser la première fois. Merci également pour ton enthousiasme permanent. Merci Eric T., pour tes connaissances en microscopie et toutes les superbes photos que tu nous fais des coraux et autres petites bêtes. Et promis, je vais m’entraîner pour te faire un kouign amann à la hauteur de celui des doctorants précédents. REMERCIEMENTS Je pense bien évidemment aussi aux deux techniciennes de l’équipe qui m’auront appris les bases pour travailler sur les coraux, et avec qui la mise au point des protocoles pour le corail rouge n’a pas été une mince affaire ! Merci également pour vos talents de cuisinières. Natacha, ce fut un plaisir de déguster tes gâteaux qui sont des réalisations incroyables à chaque fois, et Nathalie, pour m’avoir faire découvrir la cuisine Réunionnaise. Merci aux post-doc, ancienne et actuelle. Nathalie, tu auras égayé mes journées avec ta joie de vivre, ce fût un plaisir de t’avoir comme voisine de bureau et d’avoir partagé du temps ensemble au labo. Duygu, merci pour ta bonne énergie, les cours de yoga en bord de mer les lundis matins, et pour tes délicates attentions. Merci aux stagiaires pour leur bonne humeur inconditionnelle : Lisandrina, Gaëlle, Apolline, Astrid, Jonas... Je tenais également à remercier les personnes avec qui nous avons collaboré autour ce projet. Merci à Didier Aurelle et Anne Haguenauer de l’IMBE à Marseille pour les colonies de corail rouge que vous êtes allés chercher pour nous. Merci à Georges Carle de l’UMR TIRO-MATOs à Nice pour le cryobroyage des échantillons. Merci également à Claudiu Supuran de l’Université de Florence, et Clemente Capasso de l’Institute of Biosciences and BioResources de Naples, pour les échanges que l’on a eus sur les anhydrases carboniques, ainsi que pour votre accueil si chaleureux lors de notre venue à Naples. Mes remerciements vont ensuite à l’équipe d’Ecophysiologie, avec qui je n’ai pas seulement que partagé des discussions autour du corail : Christine, Renaud, Stéphanie, Eric B., Cécile, et Lucile, ainsi que les post-doc actuels, Jeroen et Vanessa. Merci aussi à Eric E. et Robert pour leur aide technique. Merci aux membres de l’équipe de Biologie Polaire actuels, avec qui c’est toujours un plaisir d’échanger : Céline, Victor, Hannah et Aymeric. Mais je pense bien évidemment aussi à Cindy, Robin et Jason, à qui je souhaite de vivre de belles aventures aux quatre coins du monde. Un grand Merci à Alexandra, notre Secrétaire scientifique, qui m’a vraiment bien aidée pendant ces trois années, et d’autant plus pendant ces dernières semaines. Merci à tous les autres membres du laboratoire du 5ème comme du 2ème. La liste serait bien trop longue pour tous vous citer mais vous faites aussi partie de cette aventure. Merci notamment à l’Administration, à Patricia, Barbara et Muriel. Un grand merci à toutes les personnes avec qui j’ai partagé de si bons moments en dehors du labo. Encore une fois, la liste serait trop longue pour tous vous citer ici. REMERCIEMENTS Leïla, un grand Merci pour beaucoup de choses. On aura vécu cette aventure en parallèle, à partager nos joies et nos stress, mais pas seulement, heureusement ! Pour les salades tomates/mozza, les pizzas, les chaï latte, pour tes nombreuses Leïlades et pour toutes nos aventures en dehors du labo, que ce soit à pied dans le Mercantour, ou au bout d’une corde sur une falaise. Nous y sommes arrivées ! Un Merci plus large à toutes les personnes du CSM qui m’auront témoigné leur soutien de quelques façons que ce soit aux moments où j’en ai eu besoin. Une petite pensée pour les trains de la région PACA, monégasques ou même Bourguignons qui m’auront menée, moi et mes compagnons de galère, tous les jours jusqu'au labo. Avec ses retards, ses suppressions et ses justifications douteuses... Je ne peux pas clôturer ces remerciements sans remercier mes ami(e)s Breton(ne)s et ma famille qui auront suivi de loin cette thèse. Merci à vous pour votre soutien, et les bouffées d’oxygène lors de mes retours, parfois trop courts. SOMMAIRE Sommaire Chapitre 1 ................................ LA BIOMINERALISATION CHEZ LES CORAUX 1 1. Généralités sur les biominéraux chez les Métazoaires ...................................................... 3 1.1. La biominéralisation dans l’échelle des temps .............................................. 3 1.2. Diversité, propriétés et fonction des biominéraux ......................................... 5 2. La biominéralisation chez les coraux ................................................................................... 8 2.1. Les coraux: des cnidaires qui (bio)minéralisent ............................................ 8 2.2. Histoire évolutive des coraux ......................................................................... 11 2.3. Processus de biominéralisation chez les coraux ............................................ 15 a) Physico-chimie de la biominéralisation b) Processus cellulaires et moléculaires de la biominéralisation 3. Problématique de l’étude de la biominéralisation .............................................................. 25 Chapitre 2 .................... LA BIOMINERALISATION CHEZ CORALLIUM RUBRUM 27 1. La biominéralisation chez les Octocoralliaires ................................................................... 29 1.1. Etat des connaissances sur la biominéralisation chez les Octocoralliaires .. 29 1.2. Diversité des structures minéralisées ............................................................ 31 2. Présentation du modèle d’étude ........................................................................................... 44 2.1. Le corail rouge : minéral, végétal ou animal ? ............................................. 44 2.2. Distribution, exploitation et protection .......................................................... 46 2.3. Classification ................................................................................................. 48 3. La biomineralisation chez Corallium rubrum...................................................................... 49 3.1. Anatomie de Corallium rubrum ..................................................................... 49 3.2. Les biominéraux chez Corallium rubrum ......................................................
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