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Titre Du Document Présentée et soutenue publiquement le 20 avril 2016 par : Auteur du mémoire Titre du mémoire, v.1.10 Thèse En vue de l’obtention du grade de Docteur de l’Université de Guyane Discipline : Sciences et Technologies Spécialité : Sciences forestières et du bois Présentée en vue d’être soutenue publiquement le 17 octobre 2017 par : BARBARA GHISLAIN Diversité anatomique et efficience du bois de tension des arbres de forêt tropi- cale humide. JURY Pr, NCB Naturalis, Leiden Pieter BAAS Rapporteur (NL) Pr, AgroParisTech LERFoB, Mériem FOURNIER Rapporteur Nancy DR, CNRS AMAP, Montpel- Nick ROWE Examinateur lier CR, CNRS LMGC, Montpel- Tancrède ALMÉRAS Examinateur lier Jacques BEAUCHÊNE Ch, Cirad EcoFoG, Kourou Examinateur Bruno CLAIR DR, CNRS EcoFoG, Kourou Directeur de Thèse Ecole Doctorale n°587 : Diversités, Santé et développement en Amazonie REMERCIEMENTS Si l’arbre est capable de cacher la forêt, alors un thésard cache certainement une foule. Par ces quelques lignes, je souhaite remercier chaleureusement les nom- breuses personnes qui m’ont apporté leur aide, qu’elle soit de nature scientifique, logistique ou simplement humaine. Tout d’abord merci à Bruno Clair, pour m’avoir orienté au cours de cette grande aventure. Merci de m’avoir transmis un grand nombre de connaissances ainsi que la motivation d’aller toujours plus loin. Merci à Tancrède Alméras de m’avoir accompagné dans l’aventure de l’efficience et d’avoir clarifié de nombreux concepts en étant un excellent professeur de méca- nique. Merci à Jacques Beauchêne pour son grand savoir en anatomie des bois guyanais et sa bienveillance légendaire. Merci à Pieter Baas, Meriem Fournier et Nick Rowe d’avoir donné une plus grande envergure à mon travail de thèse lors de la soutenance. Vient ensuite une foule de personnes, impossible à lister dans un ordre précis. A tous les membres de l’ANR StressInTrees, merci pour les réunions et les confé- rences passées ensemble. Merci aux membres du CEBA pour les conférences qui agrandissent notre savoir sur le Bassin Amazonien. Merci également au personnel de l’école doctorale de l’Université de Guyane pour la diversité des formations. Je remercie Olivier Marnette pour la formidable formation « Experimentarium » ! Merci aux botanistes Julien Engel et Pascal Petronelli d’avoir identifié de nombreux troncs et plantules. Merci à Eric Nicolini pour la collaboration « Salicaceae ». Merci à Jean-Yves Goret, Benoît Burban et Onoefe Ngwete de m’avoir amené à quelques sorties « récolte de graines ». Merci à Sabrina Coste pour son savoir en serre et en manip’ d’écophysiologie .Un tout grand merci à Jocelyn Cazal pour l’énorme travail de rénovation et d’entretien de la serre, ainsi qu’à Audin Patient. Merci à l’ensemble du labo écophysio pour ses divers dépannages. Merci à Soepe Koese d’avoir patiem- ment scié et assemblé 63 systèmes d’inclinaisons de pots et scié 500 tuteurs. Merci aux stagiaires qui ont débroussaillé le terrain, Jean-Romain Roussel et Raïssa Romain. Un énorme merci à Aurélie Cuvelier pour son travail de césure et les nombreux terrains passés ensemble. Cela vaut aussi pour Grégory Faure et son formidable travail de césure en serre. Merci également aux stagiaires Nicolas Gen- tis et Camille Gérard, ainsi que tous ceux passés sur des périodes plus courtes. Un grand merci à tous les étudiants du module FTH qui nous ont aidés de 2014 à 2017. Merci à Jonathan Prunier pour son aide précieuse depuis son arrivée au labo bois. Merci à Patrick Heuret d’apprécier les poissons d’avril douteux. Un clin d’œil à tous les jeunes chercheurs du labo bois pour les coups de main et l’ambiance du labo bois: 3 Hélène Morel, Julie Bossu, Romain Lehnebach, Clémence Ogéron, Claudia Flora, Bach Nguyen et Sébastien Levionnois. Merci à Eliane Louisianna pour son aide sur Laetia. Merci à Valérie Troispoux pour sa touche de sécurité dans le labo. Merci à Frédéric Petitclerc de continuer d’améliorer le colorospeed, indispensable outil pour les coupes à haut débit. Merci à Annie Koutouan et Anthony Chassaing pour l’administratif, Josie Santini pour les aspects financiers et Carole Legrand pour l’aide à la documentation. De manière générale je voudrais remercier tous les membres de l’UMR EcoFoG, qui font de cette unité un cadre de travail à la fois dynamique et très agréable. Merci à Takabe-sensei, Arata-sensei et Awano-sensei, ainsi qu’aux autres membres du Tree Cell Laboratory à l’Université de Kyoto, au Japon, de m’avoir accueillie au sein de leur équipe pendant 3 mois. Enfin, merci à toutes les personnes dans les coulisses qui font de cette expérience un équilibre. Un énorme merci à Delphine, Damien et petit Nolan pour tous les moments ma- giques passés entre voisins. Merci à la joyeuse bande du volley pour l’ambiance. Merci à l’ensemble des Compagnons pour les bons moments. Merci à tous les co- pains pour les sorties carbets. Merci à ma famille d’être telle qu’elle est. Au risque de me répéter : Vous savez ce que je sais, non ? Le meilleur pour la fin : Stéphane, ta contribution essentielle serait trop longue à expliquer. Alors simplement : merci d’être au top ! 4 RESUME Le bois de tension est un tissu développé par les angiospermes afin de redresser ou de maintenir leur position verticale. Ce tissu génère de fortes contraintes de tension capables de courber un tronc vers le haut. Dans la littérature, l’anatomie du bois de tension est caractérisée par la présence d’une couche gélatineuse (couche G) non lignifiée dans la paroi des fibres. Cette thèse vise à étudier la diversité anatomique du bois de tension ainsi que les mécanismes de génération des contraintes de ten- sion associés et les éventuelles variations de l’efficience du redressement, afin de comprendre comment les arbres se redressent. Elle s’appuie sur des observations anatomiques sur 291 espèces tropicales, sur des estimations de contraintes de ma- turation sur arbres adultes ainsi que sur une nouvelle méthode d’estimation des contraintes de maturation sur des individus juvéniles artificiellement inclinés et tuteurés en serre. Nos résultats montrent que la couche G est présente dans la ma- jorité des espèces, bien qu’elle soit majoritairement masquée par de la lignine. Dans un faible nombre d’espèces, la couche G est absente du bois de tension. Dans ces es- pèces, le mécanisme de génération des contraintes de tension implique l’interaction du bois avec l’écorce. Bien que la contrainte de tension soit générée dans des com- partiments distincts (le bois et/ou l’écorce), ces deux mécanismes de génération des contraintes de tension ont une efficience de redressement similaire dans le stade juvénile. Les résultats de cette étude ouvrent de nouvelles perspectives de re- cherche, notamment sur le rôle fonctionnel de la lignine dans la couche G. Abstract Tension wood is a tissue developed by angiosperms to upright or maintain their ver- tical position. This tissue generates a strong tensile stress able to curve the stem upwards. In the literature, tension wood anatomy is characterized by the presence of an unlignified gelatinous layer (G-layer) in the fibre cell wall. The aims of this thesis are to study the anatomical diversity of tension wood as well as the asso- ciated mechanisms of tensile stress generation and eventual variations of uprigh- ting efficiency in order to understand how trees upright. This thesis relies on ana- tomical observations of 291 tropical species, on estimations of maturation strain on adult trees and on estimations of maturation strain on artificially tilted young trees tied to a pole in a greenhouse. Our results show that the G-layer is present in the majority of the species, although it is mainly hidden by lignin. The G-layer is absent in tension wood of a few number of species. In these species, the mechanism of ten- sile stress generation involves the interaction of wood and bark. Although tensile stress is generated in separated tissues (wood and/or bark), these two mechanism of tensile stress generation show a similar efficiency of uprighting in juvenile trees. Results of this study open new prospects of research, including on functional role of lignin in the G-layer. 5 DIVERSITE ANATOMIQUE ET EFFICIENCE DU BOIS DE TENSION DES ARBRES DE FORET TROPICALE HUMIDE TABLE DES MATIERES Remerciements .......................................................................................... 3 Résumé ...................................................................................................... 5 Avant-propos général ................................................................................ 9 Contexte de l’étude ............................................................................................................... 9 Questionnement et objectif ..................................................................................................10 Organisation du manuscrit de thèse ...................................................................................10 Chapitre 1: : Introduction générale ...................................................... 12 1.1. Avant-propos .............................................................................................................12 1.2. Caractéristiques de la forêt guyanaise et importance de la biomécanique ..............12 1.2.1. Pédologie ............................................................................................................................ 12 1.2.2. Caractéristique des peuplements et richesse spécifique ................................................ 13 1.2.3. Climat................................................................................................................................
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