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Différenciation Sociale Et Sa Régulation Au Sein D'un Clone : Cas De La UNIVERSITE PARIS XIII Ecole Doctorale de Galilée THESE Présentée pour obtenir le grade de DOCTEUR DE L’UNIVERSITE PARIS XIII Discipline : Ethologie Par Emmanuel LECOUTEY Différenciation sociale et sa régulation au sein d’un clone : cas de la fourmi parthénogénétique Cerapachys biroi Soutenue le 11 Décembre 2009 devant le jury composé de : D. FRESNEAU Professeur Président (Université Paris XIII) J. L. DENEUBOURG Professeur Rapporteur (Université Libre de Bruxelles) C. PEETERS Directeur de Recherche CNRS Rapporteur (Université Paris VI) R. JEANSON Chargé de Recherche CNRS Examinateur (Université Toulouse III) N. CHÂLINE Maître de Conférences Examinateur (Université Paris XIII) (Codirecteur de thèse) P. JAISSON Professeur Examinateur (Université Paris XIII) (Codirecteur de thèse) Laboratoire d'Ethologie Expérimentale et Comparée (EA 4443) Remerciements Ces cinq années passées au LEEC m’ont offert l’opportunité de rencontrer une flopée de personnalités, venues d’horizons divers et variés, que seul le petit monde de l’éthologie pouvait rassembler. La liste est longue, je tâcherai de ne pas avoir la mémoire courte. Je tiens, en tout premier lieu, à remercier Pierre Jaisson et Claude Baudoin. Sans cette porte qu’ils m’ont ouverte sur le DEA de Biologie du Comportement, l’éthologie ne serait restée qu’un doux rêve auquel j’étais sur le point de renoncer. J’exprime une profonde reconnaissance à Pierre Jaisson pour la confiance qu’il m’a toujours témoignée. S’il n’avait pas entrepris les démarches nécessaires à l’obtention de mon allocation de recherche, tout ce travail n’aurait jamais vu le jour… Un grand merci aussi pour avoir défendu ma candidature au monitorat. Enseigner en faculté de Médecine fut alors une expérience particulière mais pour le moins enrichissante!!! Pierre m’a également donné la chance de partir à Okinawa pour aller y récolter mes chères Cerapachys biroi. A cette générosité s’ajoute enfin l’extrême liberté qu’il m’a laissée tout au long de mon DEA et de ma thèse. Ce parti pris m’a permis de mener librement mon projet scientifique, sans jamais aucune pression. Merci pour tout ! Mon entrée dans la recherche fût des plus agréables. Fabien y est pour beaucoup. Dès mon arrivée, il m’a pris sous son aile pour me former efficacement sur C. biroi. Avant son départ et depuis le pays du soleil levant, Fabien a toujours été de précieux conseils et d’une aide incommensurable dans la rédaction de mon rapport et de mon premier article. Toute ma thèse n’est que le fruit de son joli travail et d’une réflexion qu’il avait préalablement ébauché. Par-dessus tout, je garde un magnifique souvenir de ces deux mois où il m’a accueilli, à bras ouverts, à Okinawa. Les récoltes, la plongée, la découverte de cette île et de ses habitants ont été rendues inoubliables en sa compagnie !!! A très vite sur Taiwan p’tit père !! Comment aussi remercier Nico qui, tout au long de cette thèse, m’a spontanément encadré ? Toujours enthousiaste et disponible, son intelligence, sa rigueur scientifique et son esprit critique exacerbé ont toujours considérablement enrichi ce travail. Merci également à l’ensemble des occupants du LEEC qui de part leur disponibilité, leur sympathie, et leur humour font régner une chaleureuse ambiance au sein de ce labo perdu dans l’univers parisien. Je pense notamment à : Monique et son dévouement quotidien, Alain qui revient de loin, Jean-Luc pour ses conseils statistiques, Patrick et ses têtes à claques, Christophe et sa pomme, Dominique et sa lose éternelle, Stéphane pour son engagement de chaque instant, la mignonne Simone, Renée et sa discrétion, Chantal pour ses tartes poire- chocolat, Fabrice pour la chromato et Catherine pour les joies des commandes ! Difficile d’oublier Paul et Marie-Claire qui se sont occupés respectivement de façon permanente et intermittente de mes élevages d’Aphaenogaster et de Tetramorium. Merci à vous deux ! J’ai une pensée sincère envers tous les scientifiques passionnés avec lesquels j’ai pu collaborer au cours de ce travail et/ou qui m’ont gentiment accueilli dans leur laboratoire. Un grand merci donc à Abraham Hefetz, Alain Lenoir, Kazuki Tsuji et Guénaël Cabanes. 2 J’aimerai bien sûr vivement remercier Jean-Louis Deneubourg, Christian Peeters et Raphaël Jeanson qui m’ont fait l’honneur et le plaisir d’examiner ce travail. Je salue enfin mon cher Dominique qui, à ma plus grande joie, a accepté de présider ce jury. J’ai cette fâcheuse tendance à ne dévoiler mon affection et mon amitié que par des actes, des attentions. Il est donc des choses difficiles à dire pour un roux qui n’a jamais pris l’habitude de s’épancher. Cependant, une thèse c’est avant tout la chance incroyable de partager plus que des réflexions scientifiques. Pour toutes ces rencontres, ces beaux moments partagés, qui embellissent ou compliquent une existence, mais aussi pour ces périodes difficiles où l’on se serre les coudes et où l’on ose se livrer enfin, je remercie du fond du cœur : Ronara, pour incarner tout ce que l’être humain a de plus beau et généreux… Olivier, pour m’avoir écouté et, avec qui, parler de la vie est un régal ! Nico, pour les joies et les peines traversées ensemble… Anaïs, parce que je sais que tu sais que je sais… Fabos, pour ce qu’on a déjà vécu et ce qu’on vivra encore… Pierre, avec qui la simple rédaction d’un article se transforme en expédition ! Gilles, pour nos discussions, ton humour, et ta générosité ! Rum, pour notre gourmandise commune et ces beaux jours de récolte passés avec Olivier. Alain, pour avoir gentiment relu et corrigé mon manuscrit. Pour m’avoir secouru à chacune de mes galères informatiques ! Mais aussi pour m’avoir fait partager tes voyages tout au long de cette thèse… Paul, pour m’avoir ouvert ta coquille, même éphémèrement… dElfe, pour ce soutien mutuel au cours de la rédaction ! Pour Okinawa, ta magie et ton univers… Aurélia, pour avoir écouté, en boucle, Michael Jackson tout au long de ma rédaction ! Nanie, Marie et Max pour ses 5 ans de colocation mémorables ! Je me permets également de remercier Vio, Tibo, J.C., Steve, Daphné, Clovis, Léa, Damien et Cédric qui ont tous chacun contribué à égayer la vie du 6e étage au cours de ces quelques années. Je ne peux conclure ces interminables remerciements sans évoquer ceux qui me portent depuis tout ce temps. Je pense évidemment à mes amis normands et parisiens d’adoption qui sauront tous se reconnaître chacun dans leur petit coin ! Un grand merci également à Ninon pour avoir enduré mon absence durant la quasi-totalité de cet été 2009 passé à rédiger. Merci pour ton soutien et ton réconfort quotidien ! Enfin je ne remercierai jamais assez mes parents, mes sœurs Solène & Fanny, mes grands- parents ainsi que Pierre & Gisèle pour la confiance aveugle et l’Amour que vous m’avez toujours témoignés. Les mots manquent pour dire ce que vous tous représentez à mes yeux… 3 à Pierrot 4 TABLE DES MATIERES Introduction - Différenciation sociale : du gène au comportement................... 8 1) Variation génétique et division du travail ....................................................................... 14 1.1) Le déterminisme génétique reine/ouvrière................................................................ 15 1.2) Les spécialisations entre reines................................................................................. 16 1.3) Les spécialisations entre ouvrières............................................................................ 17 2) La différenciation pré-imaginale...................................................................................... 20 2.1) Influence de la reine au stade œuf............................................................................. 22 2.2) Influence de la température....................................................................................... 23 2.3) Le signal alimentaire................................................................................................. 24 2.4) Inhibition royale des larves par contrôle du comportement des ouvrières ............... 26 2.5) Contrôle phéromonal direct du déterminisme larvaire ............................................. 27 3) La différenciation imaginale ............................................................................................. 33 3.1) Le polyéthisme d’âge................................................................................................ 33 3.2) Le rôle de l’expérience.............................................................................................. 34 3.3) Les interactions sociales ........................................................................................... 36 3.4) Les modèles proximaux de l’organisation du travail................................................ 40 4) Objectifs de la thèse ........................................................................................................... 44 Chapitre 1 - Présentation du modèle biologique.................................................... 46 1) Présentation phylogénétique............................................................................................. 47 2) Distribution géographique et habitat............................................................................... 49 3) Récolte et élevage au laboratoire...................................................................................... 49 3.1) Bilan des récoltes ...................................................................................................... 49 3.2) Conditions d’élevage ...............................................................................................
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