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Les Hormones Thyroïdiennes, Leurs Récepteurs Et L'évolution De La

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Les Hormones Thyroïdiennes, Leurs Récepteurs Et L'évolution De La Les hormones thyroïdiennes, leurs récepteurs et l’évolution de la métamorphose chez les Chordés. Mathilde Paris To cite this version: Mathilde Paris. Les hormones thyroïdiennes, leurs récepteurs et l’évolution de la métamorphose chez les Chordés.. Biochimie [q-bio.BM]. Ecole normale supérieure de lyon - ENS LYON, 2008. Français. tel-00366311 HAL Id: tel-00366311 https://tel.archives-ouvertes.fr/tel-00366311 Submitted on 6 Mar 2009 HAL is a multi-disciplinary open access L’archive ouverte pluridisciplinaire HAL, est archive for the deposit and dissemination of sci- destinée au dépôt et à la diffusion de documents entific research documents, whether they are pub- scientifiques de niveau recherche, publiés ou non, lished or not. The documents may come from émanant des établissements d’enseignement et de teaching and research institutions in France or recherche français ou étrangers, des laboratoires abroad, or from public or private research centers. publics ou privés. THÈSE Présentée devant L’ECOLE NORMALE SUPÉRIEURE DE LYON pour l’obtention du DIPLÔME DE DOCTORAT soutenue le 18 décembre 2008 par MATHILDE PARIS THYROIDHORMONES,THEIRRECEPTORS AND THE EVOLUTION OF METAMORPHOSIS IN CHORDATES Jury: Vincent LAUDET Directeur de thèse Béatrice DESVERGNE Rapportrice Detlev ARENDT Rapporteur Philippe JANVIER Examinateur Nicholas HOLLAND Examinateur Frédéric FLAMANT Examinateur Hector ESCRIVA Membre invité THYROIDHORMONES,THEIRRECEPTORS ANDTHEEVOLUTIONOFMETAMORPHOSIS INCHORDATES mathilde paris Doctorate of Life Science 18 December 2008 ABSTRACT In an attempt to understand how the regulation of development evolves, particular attention has been put on transcription factors, which regulate gene expression during development. Among transcription factors, nuclear hormone receptors (NRs) have a peculiar status linked to their ligand-dependent activity. However, their role in the evolution of development is still poorly understood. In this context, I studied during my thesis the evolution of the estrogen receptor (ER) and of the thyroid hormone receptor (TR) in chordates (including vertebrates, urochordates like sea squirt and cephalochordates like amphioxus), by focusing on the amphioxus species Branchiostoma floridae. Combined molecular and bioinformatic approaches allowed me to show that the amphioxus ER does not bind estrogen, suggesting that estrogen binding appeared in the vertebrate lineage. In parallel, a large part of my thesis was dedicated to understand how metamorphosis evolved in the chordate lineage, as a paradigm of the evolution of a developmental process. Although most chordates metamorphose, the morphological changes during larva-to-adult transitions vary extensively from one species to another. Does the molecular determinism of metamorphosis in this group reflect this morphological diversity? In the well-studied vertebrates, metamorphosis is triggered by thyroid hormones (THs) binding to their receptor TR, member of the NR superfamily. In order to get better insight into the evolution of the molecular determinism of metamorphosis in chordates, I focused on the most basal chordate amphioxus. Combined biochemical and phylogenetic approaches allowed me to establish that amphioxus produces various THs through metabolic pathways homologous to vertebrate ones. Then I showed that TH-dependent TR activation is essential for metamorphosis induction in amphioxus, like in vertebrates, with the slight difference that the active TH is not T3, the classical vertebrate TH, but possibly its derivative TRIAC. Consequently the homology of metamorphosis in chordates is revealed by the conservation of its triggering mechanism. This suggests that the evolution of metamorphosis in chordates is marked by the conservation of the couple TH/TR whereas other parts of the regulatory network may change to underlie the morphological diversity observed nowadays. 3 RÉSUMÉ Un des principaux objectifs de ma thèse a été de comprendre comment les pro- cessus du développement évoluent. Dans ce contexte, les facteurs de transcription sont importants car ils régulent l’expression génique au cours du développement. Parmi les facteurs de transcription, les récepteurs nucléaires (RNs) ont un statut particulier car leur activité est régulée par un ligand. Le rôle qu’ils jouent dans l’évolution du développement et plus généralement leur évolution (leur fonction, leur capacité à lier un ligand...) sont mal compris. Durant ma thèse, je me suis intéressée à l’évolution des RNs chez les chordés (comprenant les vertébrés, les urochordés tels que les tuniciés et les céphalochordés tels que l’amphioxus) en étudiant deux RNs particuliers : le récepteurs aux oestrogènes (ER) et le récepteur aux hormones thyroïdiennes (TR). Les études que j’ai menées ont été réalisées en particulier chez l’amphioxus Branchiostoma floridae. J’ai montré que l’ER d’amphioxus ne lie pas les oestrogènes, suggérant que cette capacité est apparue chez les vertébrés. La majeure partie de ma thèse a été dédiée à l’étude de l’évolution de la métamorphose chez les chordés. Bien que la plupart des chordés métamorphosent, les modifications morphologiques caractéristiques du passage d’une forme larvaire à une forme juvénile sont très variables d’une espèce à l’autre. Cette variabilité morphologique se reflète-telle dans le déterminisme moléculaire de la métamorphose ? Ce dernier est encore mal connu en dehors des modèles vertébrés classiques chez qui la métamorphose est induite par la fixation des hormones thyroïdiennes (HTs) sur TR. Afin de combler ce manque, je me suis intéressée au protochordé amphioxus. Par des approches bioinformatiques et biochimiques, j’ai établi que l’amphioxus produit des HTs par une voie métabolique homologue à celle des vertébrés. J’ai alors montré que les HTs régulent la métamorphose de l’amphioxus, en modulant l’activité de TR, comme chez les vertébrés. Une différence notable est la nature de l’HT active : T3 chez les vertébrés contre TRIAC, un dérivé de T3, chez l’amphioxus. J’ai ainsi proposé que l’homologie de la métamorphose chez les chordés est soutenue par la conservation du déterminisme moléculaire de ce processus du développement (le couple HT/TR) alors que les autre parties de la voie de régulation ont été moins conservées au cours de l’évolution et expliquent la diversité morphologique que l’on observe de nos jours. 4 PUBLICATIONS Holland N. D., Paris M., and Koop D. (2009) The club-shaped gland of am- phioxus: export of secretion to the pharynx in pre-metamorphic larvae and apopto- sis during metamorphosis. Acta Zoologica 89, doi: 10.1111/j.1463–6395.(2008)00379.x. Tocchini-Valentini G. D., Rochel N., Escriva H., Germain P., Peluso-Iltis C., Paris M., Sanglier-Cianferani S., Dorsselaer A. V., Moras D., and Laudet V. (2008) Structural and functional insights into the ligand binding domain of a non- duplicated RXR from the invertebrate chordate amphioxus. J Biol Chem 284, 1938- 1948. Paris M., Brunet F., Markov G., Schubert M., and Laudet V. (2008) The amphioxus genome enlightens the evolution of the thyroid hormone signaling pathway. Dev Genes Evol 218, 667-680. Paris M., and Laudet V. (2008) The history of a developmental stage: Metamor- phosis in chordates. Genesis 46, 657-672. Schubert M., Brunet F., Paris M., Bertrand S., Benoit G., and Laudet V. (2008) Nuclear hormone receptor signaling in amphioxus. Dev Genes Evol 218, 651-665. Paris M., Pettersson K., Schubert M., Bertrand S., Pongratz I., Escriva H., and Laudet V. (2008) An amphioxus orthologue of the estrogen receptor that does not bind estradiol: insights into estrogen receptor evolution. BMC Evol Biol, 8, 219. Paris M., Escriva H., Schubert M., Brunet F., Brtko J., Ciesielski F., Roecklin D., Vivat-Hannah V., Jamin E. L., Cravedi J. P., Scanlan T. S., Renaud J. P., Holland N. D., and Laudet V. (2008). Amphioxus postembryonic development reveals the homology of chordate metamorphosis. Curr Biol, 18, 825–30. Markov G. V., Paris M., Bertrand S., and Laudet V. (2008) The evolution of the ligand / receptor couple: a long road from comparative endocrinology to compara- 5 tive genomics. Molecular and Cellular Endocrinology, 293, 5–16. Pagès M., Calvignac S., Klein C., Paris M., Hugues S., and Hänni C. (2008) Combined analysis of fourteen nuclear genes refines the Ursidae phylogeny. Mol Phylogenet Evol, 47, 73-83. Fuentes M., Benito E., Bertrand S., Paris M., Mignardot A., Godoy L., Jimenez- Delgado S., Oliveri D., Candiani S., Hirsinger E., D’Aniello S., Pascual-Anaya J., Maeso I., Pestarino M., Vernier P., Nicolas J. F., Schubert M., Laudet V., Geneviere A. M., Albalat R., Garcia Fernandez J., Holland N. D., and Escriva H. (2007) Insights into spawning behavior and development of the european amphioxus (Branchios- toma lanceolatum). J Exp Zool B Mol Dev Evol, 308, 484-493. Bertrand S., Camasses A., Paris M., Holland N.D., and Escriva H. (2006) Phylo- genetic analysis of Amphioxus genes of the proprotein convertase family, including aPC6C, a marker of epithelial fusions during embryology. Int J Biol Sci 2, 125-132. Hassani Z., François J.-C., Alfama G., Dubois G. M., Paris M., Giovannangeli C., and Demeneix B. A. (2007) A hybrid CMV-H1 construct improves efficiency of PEI-delivered shRNA in the mouse brain. Nucleic Acids Res, 35, e65. Brunet F. G., Crollius H. R., Paris M., Aury J. M., Gibert P., Jaillon O., Laudet V., and Robinson-Rechavi M. (2006) Gene loss and evolutionary rates following whole-genome duplication in teleost fishes. Mol Biol Evol, 23, 1808–16. 6 REMERCIEMENTS Je voudrais tout d’abord remercier les membres de mon jury de thèse Béatrice Desvergne, Detlev Arendt, Philippe Janvier, Nicholas Holland et Frédéric Flamant pour avoir accepté de juger mon travail, pour leur indulgence et leurs commentaires précieux. Je voudrais remercier Vincent Laudet de m’avoir accueillie dans son laboratoire (pendant 5 ans et demi tout de même). On n’a pas assez discuté, on n’a pas toujours été d’accord mais on a toujours fini par un bon compromis. Je rends hommage à son ouverture d’esprit et à sa porte toujours ouverte. Vincent m’a donné un passionnant sujet de thèse, qui a métamorphosé ma vision du développement tardif chez les chordés (non pas que j’en avais une avant la thèse...).
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