Ontogenetic variations in leaf traits of the homoblastic species dipterocarpus alatus roxb ex g don under two light conditions at cat tien national park, vietnam

UNIVERSITY OF MONTPELLIER II UNIVERSITY OF SCIENCE IN HO CHI MINH CITY Doctoral School SIBAGHE Speciality Evolution, Ecology, Genetic Resources, Paleontology ONTOGENETIC VARIATIONS IN LEAF TRAITS OF THE HOMOBLASTIC SPECIES DIPTEROCARPUS ALATUS ROXB EX G DON UNDER TWO LIGHT CONDITIONS AT CAT TIEN NATIONAL PARK, VIETNAM Joint Thesis submitted in fulfillment of the requirements for the degree of Doctor of Sciences By Anh Tuan DANG LE Presented on 21 December 2012 Before the jury MEYER-BERTHAUD Brigitte Directrice of Research Examinator CNRS UMR AMAP Montpellier COCHARD Hervé Director of Research Reporter INRA UMR PIAF Clermont-Ferrand SANOJA Elio Professor Reporter UNED, Puerto Ordaz, Venezuela EDELIN Claude Senior Researcher Co-director CNRS UMR AMAP Montpellier LE CONG Kiet Emeritus Professor University of Sciences, Ho Chi Minh City Co-director ONTOGENETIC VARIATIONS IN LEAF TRAITS OF THE HOMOBLASTIC SPECIES DIPTEROCARPUS ALATUS UNDER TWO LIGHT CONDITIONS AT CAT TIEN NATIONAL PARK, VIETNAM ABSTRACT When explaining variations in leaf parameters, analysis is based solely on environmental factors will lead to significant errors if the plant shows substantial ontogenetic variations in leaf properties We evaluated variations in 27 morphoanatomical leaf traits of Dipterocarpus alatus over six architectural development stages and between axes at a given development stage under two different light conditions An architectural analysis was conducted to distinguish precisely and objectively axis categories and developmental stages of D alatus Leaves were collected on the most recent complete growth unit on the top of the trunk and the outermost part of the middle-crown branch from trees growing under two different light conditions Twenty two leaf traits were measured and calculated using ImageJ on images of leaf tracings and cross-sections Stomatal density was calculated on nail varnish impression of the leaf lower surface Four chlorophyll traits were determined ANOVA and Tukey post-hoc tests were used to determine differences between development stages and axes The lmg relative contribution was calculated and compared to determine ontogenetic stage, axis category or light intensity, which explained more the variations in leaf traits Leaf traits differed strongly during ontogeny for all four axes and under both light conditions The axis effect was displayed at stages B, C, D and F but was insignificant at stage E Tree ontogeny explained more variations in leaf traits than light intensity Axis category explained more variations in leaf morphology but not in leaf anatomy than tree ontogeny Strong and significant intraspecific variations (ontogeny and axes) may influence the interspecific variations, and thus challenge the validity of the mean value of leaf traits between species Tree ontogeny contributed more than light intensity in explanation of the variability in leaf traits both on the trunk and second axis order strongly recommends that studies on the responses of leaf anatomy to the environment might need to correct for an ontogeny effect Key words: Dipterocarpus alatus; architecture; ontogeny; intraspecific; leaf morphology; leaf anatomy, homoblastic, light intensity VARIATIONS ONTOGENETIQUES DES TRAITS FOLIAIRES DE L'ESPÈCE HOMOBLASTIQUE DIPTEROCARPUS ALATUS SOUS DEUX CONDITIONS LUMINEUSES AU PARC NATIONAL DE CAT TIEN, VIETNAM RÉSUMÉ En expliquant les variations des paramètres feuilles, l'analyse est fondée uniquement sur des facteurs environnementaux conduire des erreurs importantes si la plante montre des variations ontogénétiques substantielles des propriétés des feuilles Nous avons évalué les variations des 27 traits morpho-anatomiques des feuilles de Dipterocarpus alatus durant six stades de développement architectural et entre les axes un stade de développement donné sous deux conditions lumineuses différentes Une analyse architecturale a été réalisée pour distinguer précisément et objectivement des catégories d'axes et stades de développement de D alatus Les feuilles ont été collectées sur l'unité de croissance plus récente et complète sur la partie supérieure du tronc et sur la partie plus externe de la branche du milieu couronne des arbres qui poussent sous deux différentes conditions lumineuses Vingtdeux traits foliaires ont été mesurées et calculées sur des images de tracés feuilles et de sections transversaux l'aide de logiciel ImageJ Densité stomatal a été calculée sur l'impression de vernis ongles de la surface inférieure des feuilles Quatre traits de chlorophylle ont été déterminés Les tests d’ANOVA et Tukey ont été utilisés pour déterminer les différences entre les stades de développement et des axes La contribution relative lmg a été calculée et comparées pour déterminer le stade ontogénétique, catégorie d’axe ou l'intensité lumineuse, ce qui est expliqué plus les variations de traits foliaire Les traits foliaires diffère fortement durant l'ontogenèse pour les quatre axes et sous des conditions lumineuses L'effet de l'axe a été présentée au stade B, C, D et F, mais a été insignifiante au stade E Ontogenèse de l’arbre expliqué plus de variations des traits foliaire que l'intensité lumineuse Catégorie d’axe expliqué plus de variations dans la morphologie des feuilles, mais pas dans l'anatomie des feuilles que l’ontogenèse de l’arbre Fortes et significatives variations intra-spécifiques (l'ontogenèse et axes) peuvent influencer les variations inter-spécifiques, et donc contestent la validité de la valeur moyenne des traits foliaires entre les espèces Ontogenèse de l’arbre contribué plus que l'intensité lumineuse dans l'explication de la variabilité des traits foliaire la fois sur le tronc et deuxième l’ordre l’axe recommande fortement que les études sur les réponses de la morphologie et l'anatomie des feuilles l'environnement doivent corriger l'effet de l'ontogenèse Mots clés: Dipterocarpus alatus, architecture, ontogénèse; intraspécifique; la morphologie des feuilles; l'anatomie des feuilles, homoblastique, l’intensité lumineuse This thesis was granted by Agence Universitaire de la Francophonie (AUF), programme Sud Experts Plantes (SEP), University of Montpellier (UM2), UMR-AMAP, France and Vietnam International Education Development, programme 322 (VIED), Vietnam PREFACE The purpose of this thesis is to study the variations in leaf morphological and anatomical traits during ontogeny and between axes at a given development stage of the homoblastic species, Dipterocarpus alatus that grows under two contrasting light conditions at Cat Tien national park, Vietnam This manuscript was represented under article format in six chapters The first chapter introduced the basic structure of leaf morphology and anatomy, their applications in plant taxonomy, floristic, evolution and ecology The general variations of leaf traits during plant development, within different positions in a plant and due to environment were also mentioned Finally, the remained problems and objectives were noted In the second chapter, the general methods used in this thesis were described in details Within the next three chapters, the 3rd, 4th and 5th, variations in leaf morphology and anatomy during tree ontogeny and among different axes (orders and categories) were evaluated for trees that grow under low light and high light Ontogenetic changes in leaf morphological traits of the trunk were found in chapter three Variations in leaf morphology during ontogeny of four axes and variations in leaf morphological traits between axes at a given development stage were reported in chapter The same analysis was performed for leaf anatomy in chapter Chapter gives a general discussion on how leaf morphology and anatomy changes during ontogeny of an axis and between axes at a given development stage This last chapter also provides the interpretation of variations in leaf traits in the viewpoint of adaptation and morphogenetic movement In brief, both leaf morphological and anatomical traits changed significantly during tree ontogeny and between different axes at a given development stage The most important result was that tree ontogeny explained much more the variability in leaf traits than light intensity did, and this was found on the trunk and second order axis under both low light and high light Our results give a strong argument that studies on the responses of leaf traits to the environment might need to correct for the ontogeny effect Montpellier, France November 2012 Anh Tuan DANG LE For the circle of life, Buddha told Goethe: “Everything is born, grows, gets old, has sickness and finally disappears” Goethe would praise: “What a beautiful metamorphosis” - Be inspired - ACKNOWLEDGEMENT To be here, I am grateful to: Teachings of Buddha, Yoga and Mediation for my peacefulness, Ms VO Kim Tien for her heart to show me the first step of research, Prof LE CONG Kiet for his wisdom to refine me on the stair of research, Dr EDELIN Claude and Ms Martine for their great enthusiasm and patience to enlarge my view of humanity and to extend my vision of research, Dr MILLET Jérôme for his generosity to be a teacher and a friend, Staffs in Cat Tien National Park for their help to facilitate my field trip, Dr LE MOGUÉDEC Gilles for his statistical lessons at the first step, Dr MUNOZ Franỗois and Prof NIMON Kim for their help on statistical issues, Dr ROWE Nick, Dr REY Hervé and Dr HEINZ Christine for their useful comments to the manuscript, Ms BROHARD Yannick for her great sympathy to find any document I need, Ms DE GUILLEN Marie-Laure and Ms LOPEZ Joelle for their rapid solution to any administrative difficulties, Dr AUCLAIR Daniel , Dr CARAGLIO Yves, Dr BARTHELEMY Daniel, Dr COUTERON Pierre, Dr MEYER-BERTHAUD Brigitte, Dr LEBRUN Michel, Ms TRAN Thi Phuong Giang and Mr TRUONG Quang The for their help to facilitate administration process, Tristan, Mathieu, Trung, Mai and Hang for their great friendship, All AMAPiens for 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architecture; ontogeny; intraspecific; leaf morphology; leaf anatomy, homoblastic, light intensity VARIATIONS ONTOGENETIQUES DES TRAITS FOLIAIRE DE L'ESPÈCE HOMOBLASTIC DIPTEROCARPUS ALATUS SOUS DEUX CONDITIONS LUMINEUSES AU PARC NATIONAL DE CAT TIEN, VIETNAM RÉSUMÉ En expliquant les variations des paramètres feuilles, l'analyse est fondée uniquement sur des facteurs environnementaux conduire des erreurs importantes si la plante montre des variations ontogénétiques substantielles des propriétés des feuilles Nous avons évalué les variations des 27 traits morpho-anatomiques des feuilles de Dipterocarpus alatus durant six stades de développement architectural et entre les axes un stade de développement donné sous deux conditions lumineuses différentes Une analyse architecturale a été réalisée pour distinguer précisément et objectivement des catégories d'axes et stades de développement de D alatus Les feuilles ont été collectées sur l'unité de croissance plus récente et complète sur la partie supérieure du tronc et sur la partie plus externe de la branche du milieu couronne des arbres qui poussent sous deux différentes conditions lumineuses Vingt-deux traits foliaires ont été mesurées et calculées sur des images de tracés feuilles et de sections transversaux l'aide de logiciel ImageJ Densité stomatal a été calculée sur l'impression de vernis ongles de la surface inférieure des feuilles Quatre traits de chlorophylle ont été déterminés Les tests d’ANOVA et Tukey ont été utilisés pour déterminer les différences entre les stades de développement et des axes La contribution relative lmg a été calculée et comparées pour déterminer le stade ontogénétique, catégorie d’axe ou l'intensité lumineuse, ce qui est expliqué plus les variations de traits foliaire Les traits foliaire diffère fortement durant l'ontogenèse pour les quatre axes et sous des conditions lumineuses L'effet de l'axe a été présentée au stade B, C, D et F, mais a été insignifiante au stade E Ontogenèse de l’arbre expliqué plus de variations des traits foliaire que l'intensité lumineuse Catégorie d’axe expliqué plus de variations dans la morphologie des feuilles, mais pas dans l'anatomie des feuilles que l’ontogenèse de l’arbre Fortes et significatives variations intra-spécifiques (l'ontogenèse et axes) peuvent influencer les variations inter-spécifiques, et donc contestent la validité de la valeur moyenne des traits foliaires entre les espèces Ontogenèse de l’arbre contribué plus que l'intensité lumineuse dans l'explication de la variabilité des traits foliaire la fois sur le tronc et deuxième l’ordre l’axe recommande fortement que les études sur les réponses de la morphologie et l'anatomie des feuilles l'environnement doivent corriger l'effet de l'ontogenèse Mots clés: Dipterocarpus alatus, architecture, ontogénèse; intraspécifique; la morphologie des feuilles; l'anatomie des feuilles, homoblastique, l’intensité lumineuse This thesis was granted by Agence Universitaire de la Francophonie (AUF), program Sud Experts Plantes (SEP), University of Montpellier (UM2), UMR-AMAP, France and Vietnam International Education Development, programme 322 (VIED), Vietnam ... ONTOGENETIC VARIATIONS IN LEAF TRAITS OF THE HOMOBLASTIC SPECIES DIPTEROCARPUS ALATUS UNDER TWO LIGHT CONDITIONS AT CAT TIEN NATIONAL PARK, VIETNAM ABSTRACT When explaining variations in leaf. .. during ontogeny and between axes at a given development stage of the homoblastic species, Dipterocarpus alatus that grows under two contrasting light conditions at Cat Tien national park, Vietnam. .. was calculated and compared to determine ontogenetic stage, axis category or light intensity, which explained more the variations in leaf traits Leaf traits differed strongly during ontogeny for