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C:\Users\Ludo\Desktop\THESE Ludovic Sawelew Compilee Pour THESE DE DOCTORAT D’AIX-MARSEILLE UNIVERSITE Ecole Doctorale Sciences de La Vie et de la Santé Présentée par Ludovic SAWELEW Pour obtenir le grade de Docteur d’Aix-Marseille Université Spécialité : biologie végétale La palytoxine chez les Zoantharia (Cnidaria) : biodiversité des organismes producteurs et activité anticancéreuse de la toxine Soutenue à huis clos le 2 octobre 2017 devant le jury composé de : Mme Patricia BONIN, Directrice de Recherche, CNRS Présidente du jury M. Denis ALLEMAND, Professeur, Centre Scientifique de Monaco Rapporteur M. Mohamed MEHIRI, Maître de Conférence, Univ. Nice Sophia Antipolis Rapporteur M. Jean LORQUIN, Ingénieur de Recherche, IRD Directeur de thèse Institut Méditerranéen d’Océanologie (MIO) Coral Biome Laboratoire de Biochimie Structurale & Bio-Procédés Grand Luminy Technopôle Institut de Recherche pour le Développement (IRD) Zone Entreprises Biotech, UMR235-IRD/7294-CNRS/AMU/Univ de Toulon Case 922, 163 Av de Luminy, Aix-Marseille Université, Polytech, Case 925, 13288 Marseille cedex 9 163 Av de Luminy, 13288 Marseille cedex 9 Remerciements Premièrement je tiens à exprimer ma gratitude à M. Yvan PEREZ sans qui je n’aurais jamais eu l’occasion de réaliser cette thèse. Lorsque j’ai décroché une bourse de mobilité européenne en 2013 pour effectuer un stage professionnalisant post master, tu m’as accueilli pendant 6 mois à l’IMBE et tu as su me transmettre une partie de ton immense passion pour l’étude et la culture des coraux. Après m’avoir présenté l’équipe de Coral Biome, nous avons envisagé la réalisation d’un travail à plus long terme et c’est ainsi, qu’un an et demi, et beaucoup de démarches administratives plus tard, j'ai enfin pu commencer ma thèse dont je n’osais même plus rêver. Encore merci à toi ! Je tiens à remercier M. Denis ALLEMAND et M. Mohamed MEHIRI pour me faire l’honneur d’être les rapporteurs de cette thèse ainsi que Mme Patricia BONIN pour avoir accepté de juger mon travail. J’exprime toute ma gratitude à M. Jean LORQUIN, mon directeur de thèse, pour m’avoir encadré durant la plus grande partie de cette aventure. Merci pour tes encouragements et ton soutien sans faille, tes corrections de rédaction qui ont ajouté une plus-value à ce manuscrit et pour tout ce que tu as pu m’apprendre en chimie (HPLC, spectrométrie de masse, réactions multiples et variées, etc.) Merci également à Coral Biome, le partenaire industriel de cette thèse et plus particulièrement à M. Frédéric GAULT pour m’avoir accueilli dans son entreprise durant 3 années. Je remercie également Mme Sandrine COURTES et Mme Carole VALENTI pour la relecture du manuscrit, leurs conseils et leur aide sur les expérimentations, notamment la spectrométrie de masse et l’imagerie, et pour m’avoir formé aux bases et à la rigueur du travail en entreprise. Merci aussi à M. Mathieu FERRY et Mme Andréa MERIOT pour la relecture des parties de ce manuscrit portant sur les coraux, pour toutes les connaissances et l’aide qu’ils m’ont apporté sur la culture des coraux ainsi que pour leur bonne humeur au quotidien. Au cours des travaux réalisés pendant cette thèse, j’ai eu l’occasion de collaborer avec plusieurs équipes de recherche qui m’ont fait découvrir différents domaines passionnants : Je tiens d’abord à remercier M. Daniel LAFITTE et M. Claude VILLARD pour m’avoir ouvert les locaux de la Plateforme Protéomique et Innovation Technologique Timone (PIT2) où j’ai passé de très bons moments à faire de l’imagerie et de la spectrométrie de masse MALDI-ToF. Un merci tout particulier à M. Christopher NUCCIO pour son aide précieuse, ses explications et ses conseils au cours de ces manips ainsi que pour m’avoir tenu au courant de tout ce qui se passait à l’OM. Merci à M. Christos PANAGIOTOPOULOS et M. Vincent FAUVELLE de l’Institut Méditerranéen d'Océanologie (MIO) pour leur accueil chaleureux, leur aide et la mise à disposition du matériel LC-QToF nécessaire aux analyses par spectrométrie de masse et à la fragmentation. Merci à Mme Elsa CASTELLANI et M. Cédric MATTHEWS de l’Institut du Biologie de Développement de Marseille pour m’avoir formé et conseillé lors de diverses expérimentations d’imagerie par microscope. Je remercie également M. Olivier ZELPHATI pour m’avoir autorisé à entrer régulièrement dans les locaux d’OZ Bioscience pour interagir avec son équipe. Un merci tout particulier à Mme Flavie SICARD et Mme Elodie BERTOSIO pour leurs échanges riches et variés notamment sur les cultures cellulaires ainsi que pour leur aide et leurs conseils sur la microscopie à épifluorescence et les dosages Bradford. Enfin je remercie Mme Christine FERRIER-PAGES du Centre Scientifique de Monaco de m’avoir envoyé une souche de Symbiodinium spp. ainsi que du milieu de culture ASP-8A et pour ses conseils sur la culture des Symbiodinium spp. Je remercie également M. Craig FAULDS pour la relecture des résumés anglais de cette thèse. I Sommaire Remerciements ........................................................................................................................................................ I Abréviations ....................................................................................................................................................... VII Résumé confidentiel ............................................................................................................................................. IX Introduction, objectifs et contexte du travail de thèse ..................................................................................... 1 1. La palytoxine ...................................................................................................................................................... 1 1.1. Une origine légendaire ................................................................................................................................... 1 1.2. La découverte de la palytoxine ...................................................................................................................... 2 1.3. Structures et propriétés physico-chimiques .................................................................................................... 3 1.3.1. Solubilité, pouvoir rotatoire et stabilité ............................................................................................... 3 1.3.2. Stratégies d’identification ................................................................................................................... 3 1.3.3. Structure chimique .............................................................................................................................. 5 1.3.4. Spectroscopie UV ............................................................................................................................... 6 1.3.5. Identification de la palytoxine et de ses analogues par spectrométrie de masse .................................. 7 1.4. Sources de palytoxine et d’analogues ............................................................................................................ 9 1.4.1. Organismes producteurs ...................................................................................................................... 9 1.4.2. Effet des facteurs environnementaux sur la production de palytoxine .............................................. 13 1.5. Aspects toxicologiques de la palytoxine ...................................................................................................... 13 1.5.1. Toxicité et symptômes chez les mammifères .................................................................................... 13 1.6. Mécanisme d’action de la palytoxine : la pompe sodium-potassium (NaK) ................................................ 18 1.6.1. Rôle physiologique de la NaK .......................................................................................................... 18 1.6.2. Structure de la NaK ........................................................................................................................... 20 1.6.3. Mécanisme de fixation de la palytoxine sur la NaK .......................................................................... 21 1.7. Méthodes de détection biologiques de la palytoxine et de ses analogues ..................................................... 22 1.7.1. Tests biologiques sur des rongeurs .................................................................................................... 23 1.7.2. Tests de cytotoxicité ......................................................................................................................... 23 1.7.3. Tests hémolytiques............................................................................................................................ 24 1.7.4. Tests immunologiques ...................................................................................................................... 25 2. Les genres Palythoa et Zoanthus ...................................................................................................................... 26 2.1. Position taxinomique ................................................................................................................................... 26 2.1.1. L’embranchement Cnidaria ............................................................................................................... 26 2.2. L’ordre Zoantharia ....................................................................................................................................... 26 2.2.1. Répartition géographique
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