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Doctorat De L'université De Toulouse TTHHÈÈSSEE En vue de l'obtention du DOCTORAT DE L’UNIVERSITÉ DE TOULOUSE Délivré par Université Toulouse III - Paul Sabatier Discipline ou spécialité : Neurosciences cognitives Présentée et soutenue par Aurore Avarguès-Weber Le 13 décembre 2010 Titre : Cognition visuelle chez l'abeille Apis mellifera: Catégorisation par extraction de configurations spatiales et de concepts relationnels JURY Dr. Joël Fagot (LPC; Université d'Aix-Marseille, CNRS, UMR 6146) Dr. Olivier Pascalis (LPNC; UPMF-Grenoble, CNRS, UMR 5105) Dr. Simon Thorpe (CerCo; UPS-Toulouse III, CNRS, UMR 5549) Dr. Adrian Dyer (Department of Physiology; Monash University, Melbourne) Pr. Martin Giurfa (CRCA; UPS-Toulouse III, CNRS, UMR 5169) Ecole doctorale : C.L.E.S.C.O. Unité de recherche : Centre de Recherches sur la Cognition Animale (UMR5169) Directeur(s) de Thèse : Pr. Martin Giurfa Rapporteurs : Dr. Joël Fagot; Dr. Olivier Pascalis UNIVERSITE PAUL SABATIER – TOULOUSE III U.F.R. Sciences de la Vie et de la Terre THESE en vue de l’obtention du DOCTORAT DE L’UNIVERSITE DE TOULOUSE Délivré par l’Université Paul Sabatier – Toulouse III Ecole Doctorale : C.LE.S.C.O. Discipline : Neurosciences Cognitives Présentée et soutenue publiquement Le 13 décembre 2010 par Aurore AVARGUES-WEBER « Cognition visuelle chez l'abeille Apis mellifera: Catégorisation par extraction de configurations spatiales et de concepts relationnels » Directeur de thèse : Pr. Martin Giurfa JURY Dr. Joël Fagot (LPC; Université d'Aix-Marseille, CNRS, UMR 6146) Rapporteur Dr. Olivier Pascalis (LPNC; UPMF-Grenoble, CNRS, UMR 5105) Rapporteur Dr. Simon Thorpe (CerCo; UPS-Toulouse III, CNRS, UMR 5549) Dr. Adrian Dyer (Department of Physiology; Monash University, Melbourne) Pr. Martin Giurfa (CRCA; UPS-Toulouse III, CNRS, UMR 5169) Centre de Recherches sur la Cognition Animale (CRCA) UMR 5169 – Université Paul Sabatier Toulouse III – CNRS 118 Rte de Narbonne 31062 Toulouse Cedex 9 « Tout accomplissement est une servitude, il oblige à un accomplissement plus haut. » Albert Camus Remerciements Je remercie évidemment en tout premier lieu ceux qui m’ont permis de faire cette thèse : les responsables de l’Ecole Normale Supérieure de Cachan pour s’être finalement laissé convaincre de financer une thèse dans un domaine des sciences « molles » et mon directeur Martin Giurfa pour m’avoir acceptée dans son équipe et donner les moyens de mener à bien cette thèse. Je le remercie aussi particulièrement pour m’avoir poussée à développer une rigueur scientifique associée à une recherche d’excellence dans le domaine scientifique. Je souhaite aussi le remercier pour sa compréhension et son soutien pour la conciliation de mon travail de thèse et de la maternité. Je tiens aussi à remercier ici Adrian Dyer qui a fait infiniment plus que de m’avoir donné les moyens nécessaires à la réalisation de mes expériences pendant l’été australien. Il s’est toujours montré d’une immense disponibilité pour répondre à toutes mes interrogations scientifiques et réfléchir aux divers problèmes rencontrés lors de mes expériences. Il a aussi toujours réussi à garder ma motivation au plus haut en me rappelant à quel point ce métier est passionnant. Nous sommes par ailleurs devenus de vrais amis au fil de mes séjours à Melbourne. J’en profite pour embrasser affectueusement sa femme et ses adorables fillettes rousses pour m’avoir fait partager de si bons moments familiaux et m’avoir permis de découvrir leur magnifique région en très bonne compagnie. Je voudrais aussi remercier Dave Reser, le texan célibataire, avec tout ce que ça implique, pour m’avoir ouvert son appartement lors d’un de mes séjours, son aide logistique et son soutien dans les débuts difficiles de ma grossesse. Je garde un souvenir amusé de nos échanges culturels. Retour sur Toulouse, où je souhaite remercier mille fois mes collègues de bureau sans qui ces années auraient été bien tristounettes ! On a partagé de nombreux fous rires, une grande complicité, une écoute, des conseils et un soutien sans faille. Merci donc à Edith, qui est maintenant pour moi une véritable amie, car l’on partage bien plus que des discussions de bureau. Bien sûr, on aurait toutes les deux été certainement beaucoup plus efficaces dans notre travail sans la proximité de nos bureaux, mais ça aurait été bien dommage ! Merci aussi à Théo, notre cher brésilien, qui nous a fait découvrir la véritable Caipirinha, pour l’invention de son « je désiste », expression qui prend tout son sens en thèse et pour tous nos partages de soucis… J’ai réussi à finir la thèse sans cachet, tu y arriveras aussi ;o). Et merci bien sûr à Julie et Esther, qui ont elles-aussi apporté beaucoup de vie à notre bureau. Enfin, un grand merci au ‘phénomène’ Julie (Bénard), qui est partie bien trop vite, pour m’avoir tout transmis sur l’art de dresser des abeilles et quel plaisir par la suite d’avoir une collègue qui comprenait mes galères ! Merci à tous les membres de l’équipe abeille, des services techniques et administratifs et au laboratoire en général pour leur aide et conseils aussi souvent que nécessaire. Merci à Jean marc Devaud et Christian Jost pour leur aide dans le parcours du combattant du moniteur qu’est la recherche d’heures d’enseignement pour enfin réussir à atteindre son quota. Merci aussi à tous les enseignants (en particulier, Sylvie Berkès, Kerstin Bystricky et Dominique Larrouy) avec qui j’ai collaboré pour préparer et donner mes cours, une belle bouffée d’air en dehors de ma discipline. Un grand merci à Michel Garcia, chargé de l’enseignement supérieur au rectorat, pour sa disponibilité et grande aide dans tous mes problèmes et questionnements administratifs issus de ma situation particulière de normalienne, agrégée, souhaitant continuer en post-doc et ayant bénéficié d’un congé maternité lors de ma thèse. Que d’impasses contournées grâce à vous ! Je souhaite remercier aussi particulièrement les gens du rucher-école de Pechbusque dans la campagne sud-toulousaine, et tout particulièrement Christian Pech, pour leur enthousiaste accueil et mise à disponibilité de leur ruche pour mes expériences. De bien longues journées de manips répétitives égayées par le paysage magnifique du haut de la colline, l’apparition d’un renard ou d’écureuils et le foisonnement d’insectes de toutes les tailles et de toutes les couleurs à observer. Des moments très agréables pour une naturaliste dans l’âme ! Je remercie aussi la bande des physiciens, Peyo, Stéphane, Nico, FX, Thierry, Elsa… pour tous ces bons moments partagés autour d’un bon repas le midi. Beaucoup d’échanges, de discussions, de rires, rien de tel pour se mettre dans de bonnes dispositions pour le travail de l’après-midi ! Un merci plein de tendresse et d’affection à mon mari Sébastien, pour son écoute patiente de tous mes soucis, son exemple et pour avoir comblé ma vie avec autre chose que le travail pendant ces années. Je le remercie aussi pour avoir réussi à me convaincre à tenter l’aventure d’un enfant pendant la thèse. C’est une belle victoire ! Et puis comment regretter d’avoir un bébé si sage et attachant… Merci p’tit Léon pour ces beaux moments d’émotion, de rire et de complicité lors de la rédaction de ma thèse (à côté tes chagrins dus à l’éloignement de ton père et la difficulté à tout gérer sont déjà oubliés). Mes parents ont aussi toute leur place ici pour leur soutien téléphonique sans faille à chaque fois que j’avais besoin de râler, d’être conseillée ou tout simplement de parler. Merci infiniment et particulièrement à ma maman avec qui on a battu des records de durée de conservation téléphonique. Ils ont su, ce qui s’est parfois avéré essentiel, me rappeler dans les moments difficiles pourquoi je faisais cette thèse et relativiser l’importance des problèmes rencontrés. Et pour finir, merci aux abeilles, qui se sont pliées librement et sans broncher et avec détermination à tous les casse-têtes que je leur proposais. Une petite mention spéciales aux abeilles australiennes, bien plus dociles, peut-être grâce à mon accent ‘frenchy’ lorsque je les encourageais dans leur langue natale, bien sûr ! Table des matières Remerciements ....................................................................................................................................... 3 Introduction ................................................................................................................................... 7 L’abeille Apis mellifera, un modèle invertébré classique en cognition visuelle ............................... 7 Richesse comportementale ............................................................................................................. 7 Capacités visuelles de l’abeille ......................................................................................................... 9 Protocoles d’études expérimentales .............................................................................................. 12 Performances cognitives dans le domaine de la classification visuelle .......................................... 15 Catégorisation perceptive ............................................................................................................. 15 Perception des configurations spatiales ........................................................................................ 20 Formation de concepts .................................................................................................................. 23 Objectifs ...........................................................................................................................................
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