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Thesis Reference Thesis Tracing back the early evolution of ParaHox genes and the ancestral neurogenic function of Gsx/Anthox2 in the developing sea anemone, Nematostella vectensis QUIQUAND, Manon Abstract The Hox/ParaHox genes are chromosomally clustered genes encoding transcription factors that regulate neurogenesis in developing bilaterians. They emerged in early eumetazoan evolution, being already expressed in cnidarians, a prebilaterian phylum where the early steps of neurogenesis can be traced back. We first performed a systematic phylogenetic reconstruction of the Hox/ParaHox families and proposed that the ProtoHox gene most likely resembled the Gsx/Pdx genes. We then made use of the sea anemone (Nematostella) cnidarian model system to investigate the developmental role of Gsx. Thanks to neuronal markers that we identified to monitor neurogenesis in developing Nematostella, we could show in loss of function assays that Gsx/Anthox2 is likely involved in nerve net formation. Moreover Gsx/Anthox2 upstream sequences drive expression in apical neurons, suggesting that they contain a neurogenic element to be identified. Our result provide advances about the genetic circuitry involved in the differentiation of neurons in early animal evolution. Reference QUIQUAND, Manon. Tracing back the early evolution of ParaHox genes and the ancestral neurogenic function of Gsx/Anthox2 in the developing sea anemone, Nematostella vectensis. Thèse de doctorat : Univ. Genève, 2009, no. Sc. 4101 URN : urn:nbn:ch:unige-46644 DOI : 10.13097/archive-ouverte/unige:4664 Available at: http://archive-ouverte.unige.ch/unige:4664 Disclaimer: layout of this document may differ from the published version. 1 / 1 UNIVERSITÉ DE GENÈVE FACULTÉ DES SCIENCES Département de Zoologie et Biologie Animale Dr. Brigitte GALLIOT Tracing back the early evolution of ParaHox genes and the ancestral neurogenic function of Gsx/Anthox2 in the developing sea anemone, Nematostella vectensis THÈSE présentée à la Faculté des sciences de l’Université de Genève pour obtenir le grade de Docteur ès sciences, mention biologie par Manon QUIQUAND de Nemours (France) Thèse N°4101 Genève Atelier d’impression ReproMail 2009 REMERCIEMENTS Ce travail a été réalisé au département de Zoologie et Biologie Animale de l’Université de Genève et financé par le Fonds National Suisse de la Recherche Scientifique et le Département de l’Instruction Publique de la République et du Canton de Genève. Introduction À l’Université Pierre et Marie Curie, entre Paris, Villefranche-sur-Mer et Montpellier, je remercie ceux qui m’ont transmis leur passion, m’ont donné l’envie de réaliser une thèse, et surtout ceux qui m’y ont aidé. Méthodes Je tiens à présenter toute ma reconnaissance à Brigitte Galliot, ma directrice de thèse, qui m’a encadrée pendant plus de cinq années et m’a donné l’opportunité de réaliser ce travail. Merci Brigitte pour votre disponibilité, vos conseils et votre soutien qui furent au rendez-vous. Je remercie les membres actuels et passés du laboratoire, ceux qui sont devenus plus que des collègues. Aussi merci aux membres des autres laboratoires du département qui se comportent souvent comme si je faisais partie du leur. Plus particulièrement merci aux Dr. Juan Montoya et Dr. Cedric Berney pour leurs conseils en phylogénie. Résultats Je remercie les membres de mon comité de thèse pour leur temps et l’intérêt qu’ils ont porté à mon manuscrit: Dr. Evelyn Houliston, Dr. Jean-Stéphane Joly et Prof. Ivan Rodriguez. Discussion Je remercie mes ami(e)s pour les moments partagés, ceux avec qui j’ai travaillé pendant cette phase de rédaction, ceux qui m’ont aidée, ceux qui ont fini par tolérer qu’une thèse à l’Université de Genève dure souvent plus de trois ans. Enfin je remercie ma famille, particulièrement ma mère et mon père, à lui je voudrais dédier cette thèse. Conclusion Belle experience !! Perspective Continuons l’aventure… ACKNOWLEDGEMENTS This work was carried out at the Department of Zoology and Animal Biology, University of Geneva. Financal support for this work was provided by le Fonds National Suisse de la Recherche Scientifique and le Département de l’Instruction Publique de la République et du Canton de Genève. I am grateful to Brigitte Galliot for giving me the oppotunity to conduct my thesis project in her laboratory. I would like to thank her for support during these years. I would like to thank all past and present members of the laboratory for their help and the stimulating environment that they create everyday : Marijana Miljkovic-Licina, Simona Chera, Renaud de Rosa, Gaspare Benenati, Yvan Wenger, Wanda Buzgariu, Luiza Ghila, Anne Forget, Josefina Lascano, Pierre Heuze, Han Petry, Lisbeth Muster, Kevin Dobretz, Virginie Voeffray ;-), Janina Karpinsky, Fadi Hamdan, Philippe Jean. Thank you also to the far or close neighbouring of the department. Thank you to my friends and my family. Futhermore, I thank the members of my thesis comittee for their time and interest : Dr. Evelyn Houliston, Dr. Jean-Stéphane Joly and Prof. Ivan Rodriguez. i INDEX REMERCIEMENTS..........................................................................................................................I ACKNOWLEDGEMENTS ...............................................................................................................I INDEX .............................................................................................................................................II RESUME..........................................................................................................................................1 SUMMARY ......................................................................................................................................5 INTRODUCTION .............................................................................................................................7 Le phylum des cnidaires..........................................................................................................8 I.1.1. Un phylum ancestral ..................................................................................................................8 I.1.2. Diversité et cycle de vie chez les cnidaires............................................................................10 I.1.3. Développement embryonnaire des cnidaires.........................................................................12 I.1.4. Organisation générale des cnidaires ......................................................................................13 I.1.4.1 Axe(s) de symétrie chez les cnidaires............................................................................13 I.1.4.2 Organisation des tissus à partir de deux feuillets: l’ectoderme, l’endoderme, (mésoderme)...................................................................................................................................13 I.1.4.3. Organisation cellulaire de l’hydre ..................................................................................14 I.1.4.4. Le système nerveux des cnidaires................................................................................15 Les neurones...............................................................................................................................................15 Les cnidocytes.............................................................................................................................................16 La transmission synaptique........................................................................................................................18 I.1.5. Nematostella vectensis : un nouveau modèle........................................................................19 I.1.5.1. Mode et cycle de vie de Nematostella..........................................................................20 I.1.5.2. Développement embryonnaire de Nematostella..........................................................21 I.1.5.3. Organisation anatomique de Nematostella...................................................................22 I.1.5.4. Système nerveux de Nematostella ...............................................................................23 I.1.6. Les cnidaires comme système modèle...................................................................................24 I.1.6.1. Les différents modèles de cnidaires utilisés.................................................................24 I.1.6.2. Avantages et inconvénients du modèle anthozoaire Nematostella.............................25 I.1.6.3. Outils génétiques disponibles chez les cnidaires.........................................................26 I.1.7. Conservation des gènes et des voies de signalisation..........................................................27 Les gènes à homéoboîte ........................................................................................................28 I.2.1. Les mutations homéotiques et les gènes à homéoboîte.......................................................28 I.2.2. La classe ANTP .......................................................................................................................29 I.2.3. Rôle des gènes Hox dans la neurogénèse.............................................................................30 I.2.3.1. Organisation des gènes Hox en complexes.................................................................30 I.2.3.2. La colinéarité spatiale et temporelle et la prévalence postérieure..............................32 I.2.3.3. Les gènes Hox fonctionnent
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