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Biodiversité, Biochimie Et Pharmacologie Des Peptides De Venins De Fourmis

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Biodiversité, Biochimie Et Pharmacologie Des Peptides De Venins De Fourmis UNIVERSITE DES ANTILLES ET DE LA GUYANE Faculté des Sciences Exactes et Naturelles École doctorale pluridisciplinaire Thèse pour le Doctorat en Sciences de la vie Axel TOUCHARD Biodiversité, biochimie et pharmacologie des peptides de venins de fourmis Sous la direction de Jérôme ORIVEL et Pierre ESCOUBAS Soutenue le 18 Mars 2015 à l’UMR EcoFoG (Kourou) N : [0000AGUY0000] Jury : Alain LENOIR , Professeur émérite, IRBI, Université de Tours Examinateur Sébastien DUTERTRE, CR CNRS, IBMM, Université de Montpellier Examinateur Jean ROSSIER, Professeur, CPN, Hôpital Sainte Anne Examinateur Alain MILON, Professeur, IPBS, Université de Toulouse Rapporteur Mohamed AMICHE, CR CNRS, IMRB U955 INSERM, Université Paris Est Créteil Rapporteur Jérôme ORIVEL, DR CNRS, UMR EcoFoG Directeur 2 3 —Remerciements— Remerciements En premier lieu je tiens à remercier Pierre Escoubas, co-directeur de cette thèse et directeur de mon stage de Master 2, pour la confiance qu’il a su m’accorder, pour son soutien et ses conseils. Ses connaissances sur les venins, les toxines peptidiques et la spectrométrie de masse furent indispensables pour l’élaboration de cette thèse. Un grand merci à mon directeur de thèse, Jérôme Orivel. Il a su me faire profiter de son savoir myrmécologique et a en plus, apporté des critiques constructives pour me faire aller de l’avant et orienter mes recherches dans la bonne direction. Toute ma reconnaissance à Alain Dejean, sans qui ce travail n’aurait pas été pareil et qui m’a fait profiter de sa passion pour les insectes sociaux. Alain était le porteur du projet FEDER BI-Appli (115/SGAR-DE/2011/052274) qui a financé le travail effectué au cours de cette thèse. Un grand merci également à Andrea Yockey-Dejean pour ses nombreuses corrections d’anglais dans les articles scientifiques. J’adresse mes sincères remerciements à Alain Milon et Mohamed Amiche pour avoir accepté d’être les rapporteurs de ce travail. Merci également à Alain Lenoir, Jean Rossier et Sébastien Dutertre d’avoir accepté de participer à l’évaluation de cette thèse. Certaines parties de ce travail ont été effectuées en collaboration avec des laboratoires australiens. Un grand merci notamment à Graham Nicholson, Samira Aili, Jennifer Koh et Glenn King pour l’accueil chaleureux dans leurs laboratoires de Sydney et de Brisbane ainsi que leur disponibilité. 4 —Remerciements— J’adresse un remerciement particulier à Christophe Duplais et le labo SNA, je remercie également les personnes que j’ai pu côtoyer à VenomeTech; Fab, Math, Seb, Mélo, M-J, Perrine et Wela ainsi qu’a EcoFoG; Audrey, Caro, Alex Salas, Olivier, Stan, Céline, Mélanie, Roro, Bender (Juju), Romain G, Kaka, Leeloo, Élo, Niklas, pour leur bonne humeur et toutes ces bonnes soirées passées ensemble autour d’un (ou plusieurs) Ti-punch. Merci également à tous mes colocs de Parcouri et de la Maison Bleue et plus particulièrement Mélaine, Gus, Bourget, Pierrot, Bennie et Gigi. La vie à Kourou aurait été bien différente sans eux. Un grand merci à tous mes autres potes kourouciens; Marie-Mélina, Giselle, Niko Ko, Sophie, Isa, Claire, Nathan, Costo, Colette et tous les étudiants EFT et stagiaires du campus. Enfin, je remercie ma famille (mes parents, mon frère Matt et mes grands parents) pour m’avoir toujours soutenu tout au long de mes études. J’ai une pensée particulière pour mon grand père Claude à qui je dédie cette thèse. 5 —Sommaire— Sommaire Avant propos ………………………………………………………………………………………………………… 7 Introduction générale …………………………………………………….………………………………….… 10 1. Sources de produits naturels pour la recherche pharmaceutique ……..….….. 10 2. Les animaux venimeux…………………………………………………….……………………..…. 14 3. Les fourmis ……………………………………………………………….…………………..…….…... 20 4. Les venins de fourmis ……………………………………………………………………..………… 27 5. Les peptides de venins de fourmis ………………………………………………….….…….. 31 5.1. Les peptides linéaires ………………………………………………………………….……. 32 5.2. Les peptides dimériques ………………………………………………………………….. 34 5.3. Les peptides ICK ………………………………………………………………………..…….. 36 6. Objectifs généraux …………………………………………………………………………………… 39 Chapitre 1. Biodiversité ………………………………………………………………………………….…….. 43 1. Taxonomie par spectrométrie de masse ……………………………………………..….. 45 2. Application aux venins de fourmis ………………………………………………………..….. 46 Article 2 ………………………………………………………………………………………………………….. 49 Chapitre 2. Écologie …………………………………………………………………………………..………… 82 1. Venin et polyéthisme ................................................................................... 84 Article 3 ……………………………………………………………………………………..…………..…….. 87 2. Spécialisation trophique ………………………………………………………………………….. 106 3. Spécialisation défensive ……………………………………………………..…………………... 114 Article 4 ………………………………………………………………………………………..…………..….. 119 Chapitre 3. Biochimie ………………………………………………………………….………………………. 145 1. Diversité structurale des peptides …………………………………….……………………… 147 Article 5 …………………………………………………………………………………….……………..……. 149 2. Caractérisation de peptides originaux ………………………………………….………….. 182 Chapitre 4. Pharmacologie ………………………………………………………………………….………. 185 1. Neurotoxines ………………………………………………………………………………….……….. 188 Article 6 ………………………………………………………………………………………………….…..…. 191 2. Nocitoxines ……………………………………………………………………………………….…….. 217 Discussion générale ……………………………………………………………….……………………………… 220 Conclusion et perspectives ………………………………………………………………………………….. 227 Bibliographie …………………………………………………………………………………………………….…. 230 6 —Sommaire— Annexe 1— Article 1 ……………………………………………………………………………………….….. 258 Annexe 2— Protocole électrophysiologie …………………………………………………..…….. 291 Annexe 3— Abstract ……………………………………………………………………………………….…. 292 7 —Avant-propos— Avant-propos Les venins sont des armes chimiques très sophistiquées, composées de plusieurs centaines de molécules bioactives qui ont été présélectionnées et pré-optimisées par la nature. Parmi ces composés chimiques, les peptides sont les toxines majoritaires qui possèdent une formidable diversité structurale et pharmacologique. De plus ces toxines peptidiques sont souvent structurées par des ponts disulfure leur conférant une remarquable stabilité chimique ainsi qu’une grande résistance aux dégradations enzymatiques. Ces peptides bioactifs sont promis à un grand avenir et pourraient trouver de nombreuses applications dans le domaine des biotechnologies et notamment pour l’industrie pharmaceutique. En effet les peptides présentent plusieurs avantages par rapport aux petites molécules qui constituent les médicaments traditionnels. Le principal atout des peptides est qu’ils offrent souvent une plus grande efficacité et surtout une plus haute sélectivité que les petites molécules. De surcroit, leur nature protéique leur apporte un temps de demi-vie très court bien que cela puisse également être une contrainte. De plus les produits de dégradation des peptides sont peu toxiques car ce sont des acides aminés. Tout cela offre l’espoir d’en dériver des agents thérapeutiques efficaces possédant des activités très ciblées et avec moins d’effets secondaires [King, 2011; Lewis and Garcia, 2003; Uhlig et al., 2014]. D’ailleurs plusieurs peptides issus de venins ont déjà permis le développement de médicaments et de nombreux autres sont actuellement en essais cliniques et précliniques. Le Prialt®, dérivé d’une conotoxine et le Captopril® issu d’un venin de serpent sont deux exemples remarquables de médicaments provenant de venin [Smith and Vane, 2003; Staats et al., 2004]. 8 —Avant-propos— La composition biochimique des venins est strictement spécifique et donc l’ensemble des venins représentent une source presque inépuisable de peptides bioactifs. La grande biodiversité des organismes venimeux va de pair avec une grande diversité de peptides aux pharmacologies variées. L’ensemble de ces venins constitue une importante librairie de peptides bioactifs exploitable pour la recherche de molécules d’intérêt. Ainsi l’exploration de l’ensemble de la diversité taxonomique des organismes venimeux est la clé pour la découverte de nouvelles molécules. Dans cette optique, l’étude des venins de certains groupes d’animaux inexploités à ce jour, pourrait permettre la découverte de nombreuses molécules originales aux propriétés pharmacologiques uniques. Parmi les venins d’animaux, les venins de fourmis restent parmi les moins étudiés en raison de plusieurs désavantages. Le problème majeur dans l'étude des venins de fourmis est la petite taille des animaux, posant un défi majeur pour l’obtention d’une quantité de venin exploitable. Pour la reproductibilité des études biochimiques et pharmacologiques, il est primordial de pouvoir identifier de façon indiscutable les espèces étudiées mais l’identification des espèces de fourmis reste l’apanage de quelques rares spécialistes. Enfin, au contraire des venins d’araignées ou de cônes, l’absence d’une base de données spécifique aux toxines peptidiques provenant des venins de fourmis est aussi un frein à l’exploration des ces peptides. Malgré cela, les fourmis (Formicidae) constituent probablement la famille la plus facilement exploitable parmi les hyménoptères car elles offrent plusieurs avantages. Ce sont des insectes sociaux qui vivent en colonie allant de quelques individus à plusieurs millions. En plus de cela, la taxonomie des fourmis est probablement l’une des mieux connues de tous les hyménoptères et plusieurs bases de données taxonomiques existent [AntWeb, 2014b]. L’immense biodiversité des fourmis laisse envisager une grande diversité biochimique des toxines. En parallèle les avancées technologiques en termes de biochimie analytique
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