Éditeur(s) : HAL CCSD Elsevier
Résumé : International audience
Studies on tree biomechanical design usually focus on stem stiffness, resistance to breakage or uprooting, and elastic stability. Here we consider another biomechanical constraint related to the interaction between growth and gravity. Because stems are slender structures and are never perfectly symmetric, the increase in tree mass always causes bending movements. Given the current mechanical design of trees, integration of these movements over time would ultimately lead to a weeping habit unless some gravitropic correction occurs. This correction is achieved by asymmetric internal forces induced during the maturation of new wood. The long-term stability of a growing stem therefore depends on how the gravitropic correction that is generated by diameter growth balances the disturbance due to increasing self weight. General mechanical formulations based on beam theory are proposed to model these phenomena. The rates of disturbance and correction associated with a growth increment are deduced and expressed as a function of elementary traits of stem morphology, cross-section anatomy and wood properties. Evaluation of these traits using previously published data shows that the balance between the correction and the disturbance strongly depends on the efficiency of the gravitropic correction, which depends on the asymmetry of wood maturation strain, eccentric growth, and gradients in wood stiffness. By combining disturbance and correction rates, the gravitropic performance indicates the dynamics of stem bending during growth. It depends on stem biomechanical traits and dimensions. By analyzing dimensional effects, we show that the necessity for gravitropic correction might constrain stem allometric growth in the long-term. This constraint is compared to the requirement for elastic stability, showing that gravitropic performance limits the increase in height of tilted stem and branches. The performance of this function may thus limit the slenderness and lean of stems, and therefore the ability of the tree to capture light in a heterogeneous environment.
ISSN: 0022-5193

hal-00534316
https://hal.archives-ouvertes.fr/hal-00534316
DOI : 10.1016/j.jtbi.2008.10.011

Éditeur(s) : HAL CCSD Oxford University Press (OUP)
Résumé : International audience
Gravitropism is necessary for plants to control the orientation of their axes while they grow in height. In woody plants, stem re-orientations are costly because they are achieved through diameter growth. The functional diversity of gravitropism was studied to check if the mechanisms involved and their efficiency may contribute to the differentiation of height growth strategies between forest tree species at the seedling stage. Seedlings of eight tropical species were grown tilted in a greenhouse, and their up-righting movement and diameter growth were measured during three months. Morphological, anatomical and biomechanical traits were measured at the end of the survey. Curvature analysis was used to analyse the up-righting response along the stems. Variations in stem curvature depend on diameter growth, size effects, the increase in self-weight, and the efficiency of the gravitropic reaction. A biomechanical model was used to separate these contributions. Results showed that (1) gravitropic movements were based on a common mechanism associated to similar dynamic patterns, (2) clear differences in efficiency (defined as the change in curvature achieved during an elementary diameter increment for a given stem diameter) existed between species, (3) the equilibrium angle of the stem and the anatomical characters associated to the efficiency of the reaction also differed between species, (4) the differences in gravitropic reaction were related to the light requirements: heliophilic species, compared to more shade-tolerant species, had a larger efficiency and an equilibrium angle closer to vertical. This suggests that traits determining the gravitropic reaction are related to the strategy of light interception and may contribute to the differentiation of ecological strategies promoting the maintenance of biodiversity in tropical rain forests.
ISSN: 0022-0957

hal-00797023
https://hal.archives-ouvertes.fr/hal-00797023
https://hal.archives-ouvertes.fr/hal-00797023/document
https://hal.archives-ouvertes.fr/hal-00797023/file/Functional_diversity_gravitropic_Almeras_al.pdf

Éditeur(s) : HAL CCSD
Résumé : Gravitropism is a major process in vertical growth as it allows the positioning of plant in the gravity field and thus, for trees, it takes part in the light foraging strategies. Without any gravitropism, trees would adopt a weeping habit or fall on the ground due to the increasing with growth supported weight, especially at the advance regeneration stage (sapling stage) in the understorey, where tree are very slender and poorly stiff. Most of the gravitropic studies are made by physiologists in order to understand the process and thus are associated to complex laboratory methods (genetic engineering, anatomical studies, accurate 2D ou 3D measurements of form changes with time). In order to study the ecological relevance and the specific diversity of gravitropic performances in natural forests, we developed a framework that uses different kinds of datas. Synchronical analysis has been done i) at the population level in large sample of trees and ii) at the tree level from destructive measurement of the dissymmetry maturation strains known to be the main motor of the posture control reaction of trees. In both cases, biomechanical models allows to estimate some aspects of the dynamic gravitropic process. Diachronical analysis has been done in natural conditions on permanent plots, or in a more traditional way, by observing in greenhouse experiments the righting movements of plants artificially tilted. In the first case, the problem is the high variability of natural disturbances and the slowness of growth and movements in the dark understorey. The second choice allows a measurement of a capacity (i.e. the ability of the species to react in quite extreme conditions of verticality disturbance but good for growth light and nutrient conditions) that is more easy to analyse, but is maybe not so relevant for estimating the ecological strategy (some species could have developed a high efficience of gravitropic reaction in natural conditions by avoiding the mechanical disturbance or adapting the reaction to slow growth and changing environments in natural conditions). All approaches use the same basic biomechanical modelling, i) to analyse simultaneously the reaction and the disturbance (due to the weight increase) during growth, or ii) to combine different parameters (geometry, growth, weight, maturation strains) involved in shape changes and posture control. They give a classification of the species according to their gravitropic performances. Results on 15 species are discussed, with relation to shade-tolerance or avoidance.
6th Plant Biomechanics Conference
Cayenne, France

hal-00565956
https://hal.archives-ouvertes.fr/hal-00565956

Éditeur(s) : HAL CCSD
Résumé : The functioning of ecosystems is based on the functional diversity, which is the complementto the species diversity. In the tropical rain forest, the saplings grow in the dense and stratifiedunderstorey. This thesis deals with the diversity of realisation of the support function by thesaplings in this context. The hypotheses are that the race through higher and so brighter levelsconstrains the mechanical security of the saplings with necessary biomass saving and need ofreorientation (gravitropism) and acclimation (mecanoperception and response) processes andthat diversity is thus structured by the species light-demand and affinity for the risk. Using awide sample of 16 species, at the Paracou station, relevant functional traits - risk of not-selfsupportingand gravitropic performance - are defined through adapted modellings whichcombine trunk shape, biomass distribution and wood properties. The traits values varybetween species and also with ontogeny and saplings environment. These variations are dueto elementary traits (shape, biomass, wood) with strong geometrical effects. The less selfsupportingspecies (low mechanical security) have the highest gravitropic performances andare also the most light-demanders. A species, Tachigali melinonii, distinguishes itself bystrong slendernesses and a frequent not-self-supporting habit letting suppose a strongadaptability to its mechanical environment. An experiment (growth response to staking),however, did not provide the expected results whereas the species with lower risks, Dicoryniaguianensis, reacted more.
Le fonctionnement des écosystèmes repose sur la diversité fonctionnelle, complément de ladiversité spécifique. En forêt tropicale humide, les jeunes arbres croissent en sous-bois denseet stratifié. Cette thèse porte sur la diversité de réalisation de la fonction biomécanique desoutien. Les hypothèses sont que la "course" vers les strates plus éclairées contraint lasécurité mécanique des jeunes arbres, avec de nécessaires économies de matière et recours àdes processus de réparation (gravitropisme) et d'acclimatation (mécanoperception et réponse),et que la diversité est donc structurée par le besoin en lumière et l'affinité pour le risque desespèces. Avec un échantillon de 16 espèces sur le dispositif de Paracou, des traits fonctionnelspertinents - risque de non-autoportance et performance gravitropique - sont définis au traversde modélisations adaptées qui combinent forme des troncs, répartition des masses etpropriétés du bois. Ces traits varient entre espèces mais aussi avec le développement etl'environnement des arbres. Ces variations sont dues aux variations des traits élémentaires(forme, masses, bois) avec de forts effets géométriques. Les espèces les moins autoportantes(faible sécurité) ont les plus fortes performances gravitropiques. Ce sont aussi les plusdemandeuses de lumière. Une espèce Tachigali melinonii se distingue par de fortsélancements et une fréquente non-autoportance laissant supposer une adaptabilité forte àl'environnement mécanique. Une expérimentation (réponse de croissance au tuteurage) n'acependant pas abouti aux résultats supposés puisque c'est au contraire l'espèce à faible risque,Dicorynia guianensis, qui s'est montrée plus réactive.
https://tel.archives-ouvertes.fr/tel-00272100

tel-00272100
https://tel.archives-ouvertes.fr/tel-00272100
https://tel.archives-ouvertes.fr/tel-00272100/document
https://tel.archives-ouvertes.fr/tel-00272100/file/TheseJaouen.pdf

Éditeur(s) : HAL CCSD
Résumé :   Diffusion du document : INRA Antilles-Guyane, UMR ECOFOG, 97387 Kourou, Guyane (France) Diplôme : Master Recherche
il s'agit d'un type de produit dont les métadonnées ne correspondent pas aux métadonnées attendues dans les autres types de produit : DISSERTATION
The gravitropic reaction is a fundamental phenomenon in the life of the young trees in under-storey of a tropical rain forest. It enables them to preserve a growth in height even after a fall of remains or a wind fallen wood had leaning them. This reaction passes mainly by the differentiation of reaction wood and the offsetting of the radial growth. The gravitropic reaction is also under the control of various parameters such as the diameter of the stems, the hardness of wood, or masses they carried. The tropical rain forest contains a great specific diversity and ecological strategies. In this work we seek to know if this diversity is reflected in the strategies of recovery of the species in particular on a level of efficiency of reaction. We compared eight species of tropical rain forest with various ecological behaviours. We studied their changes in form in the course of time, as to their growth. We showed that there are indeed interspecific differences in the gravitropic reaction. Some behaviours are specific to some species. We also showed that for the majority of the species, the deformation energy is strongly correlated with the relative growth of the species.
Le redressement gravitropique est un phénomène fondamental dans la vie des jeunes arbres en sous-bois de forêt tropicale. Il leur permet de conserver une croissance en hauteur même après avoir subi la chute de débris ou même un chablis ayant conduit à leur inclinaison. Ce redressement passe principalement par la différentiation de bois de réaction et l’excentrement de la croissance radiale. Le redressement gravitropique est également sous le contrôle de différents paramètres tels que le diamètre des tiges, la dureté du bois, ou les masse portées. La forêt tropicale humide contient une grande diversité spécifique et de stratégies écologiques. Dans ce travail nous cherchons à savoir si cette diversité se reflète dans les stratégies de redressement des espèces notamment au niveau de l’efficience de la réaction. Nous avons comparé huit espèces de forêt tropicale humide avec différents comportements écologiques. Nous avons étudié leurs changements de forme au cours du temps, ainsi que leur croissance. Nous avons réussi à montrer qu’il existe effectivement des différences interspécifiques de redressement. Des comportements particuliers à certaines espèces ont pu être dégagés. Nous avons également montré que pour la majorité des espèces, l’énergie de déformation est fortement corrélée à la croissance relative de l’espèce.
https://hal.archives-ouvertes.fr/hal-01189172

hal-01189172
https://hal.archives-ouvertes.fr/hal-01189172
PRODINRA : 11396

Éditeur(s) : HAL CCSD
Résumé : For their survival, plants need to grow in height to access light resource while ensuring their mechanical stability. Growth in height amplifies the mechanical constraints due to wind and gravity. Nevertheless these constraints represent morphogenetic signals that plants respond to by modifying their growth. There are two regulatory processes of growth by mechanical signals: thigmomorphogenesis and gravitropism. Thigmomorphogenesis expresses by a reduction of growth in height and an increase of growth in girth under the effect of mechanical signals. This is a way for plants to grow with controlling slenderness of their axes to avoid buckling (beating down). In some cases (storm, sudden exposure to a new mechanical environment) this process is not sufficient to prevent buckling, gravitropism then acts to restore axes orientation in case of no lethal buckling. With measuring several traits, we have demonstrated that these processes have a fundamental ecological role in the control of tree habit and the vertical structuration of ecosystems. As studies of climate changes seem to predict a higher frequency of storms it is important to study the diversity of their efficiency.
Colloque Adaptation au Changement Climatique de l'Agriculture et des Ecosystèmes

hal-00797377
https://hal.archives-ouvertes.fr/hal-00797377

Éditeur(s) : HAL CCSD
Résumé :   Diplôme : DEA
il s'agit d'un type de produit dont les métadonnées ne correspondent pas aux métadonnées attendues dans les autres types de produit : DISSERTATION
La lumière est le principal facteur limitant de la croissance en forêt tropicale humide. Malgré 1% du rayonnement incendie disponible en sous-bois, les jeunes arbres doivent rejoindre la canopée pour assurer la régénération du couvert. La compétition pour l'accès à la lumière est forte, et leur stabilité mécanique est souvent perturbée. La gravité étant un facteur omniprésent , tout déséquilibre entraîne des flexions de l'arbre et peut-être dramatique. Les arbres peuvent cependant palier ces déséquilibres en réorientant leurs axes par gravitropisme. Les "moteurs" des réorientations dans les axes lignifiés, sont la croissance radiale, la différentiation de bois de réaction et souvent, une croissance radiale excentrée.La capacité au redressement dépend aussi des dimensions des axes. Ces différents paramètres modulent donc la réponse gravitropique. Les réorientations dépendent de la croissance et donc également des ressources en lumière. Ce travail avait pour buts de quantifier l'impact de la croissance radiale sur le redressement d'une espèce tropicale et de mettre en évidence d'éventuelles différences d'efficience, de réorientation des arbres en fonction de la disponibilité en lumière. La réponse gravitropique de jeunes Symphonia globulifera artificiellement inclinés a été étudiée sous deux niveaux de lumière (1% et 25%°). L'importance de la croissance radiale dans la réaction gravitropique a été quantifiée par une analyse cinématique du redressement et l'utilisation d'un modèle biomécanique simple. Ce travail montre que même à 1% de lumière, les arbres ont une réponse gravitropique. L'efficience de réaction est d'ailleurs plus forte qu'à 25% de lumière. Cette réaction semble donc être une fonction prédominante pour les plantes en sous-bois. Nous avons également montré que le redressement est fortement corrélé à la vitesse de croissance radiale. Les données obtenues complètent celles déjà acquises sur des espèces tempérées. Elles permettront des comparaisons interspécifiques du processus gravitropique et serviront à valider des modèles biomécaniques plus complexes que celui utilisé dans cette étude.
https://hal.archives-ouvertes.fr/hal-01189227

hal-01189227
https://hal.archives-ouvertes.fr/hal-01189227
PRODINRA : 16967

Éditeur(s) : HAL CCSD Oxford University Press (OUP)
Résumé : International audience
Functional ecology has long considered the support function as important, but its biomechanical complexity is only just being elucidated. We show here that it can be described on the basis of four biomechanical traits, two safety traits against winds (SW) and self-buckling (SB), and two motricity traits involved in sustaining an upright position, tropic motion velocity (MV) and posture control (PC). All these traits are integrated at the tree scale, combining tree size and shape together with wood properties. The assumption of trait constancy has been used to derive allometric scaling laws, but it was more recently found that observing their variations among environments and functional groups, or during ontogeny, provides more insights into adaptive syndromes of tree shape and wood properties. However, over-simplified expressions have often been used, possibly concealing key adaptive drivers. An extreme case of over-simplification is the use of wood basic density as a proxy for safety. Actually, since wood density is involved in stiffness, loads and construction costs, the impact of its variations on safety is non-trivial. Moreover, other wood features, especially the microfibril angle (MFA), are also involved. Furthermore, wood is not only stiff and strong, but it also acts as a motor for MV and PC. The relevant wood trait for that is maturation strain asymmetry. Maturation strains vary with cell wall characteristics such as MFA, rather than with wood density. Finally, the need for further studies about the ecological relevance of branching patterns, motricity traits and growth responses to mechanical loads is discussed.
ISSN: 0022-0957

hal-00909904
https://hal.archives-ouvertes.fr/hal-00909904
https://hal.archives-ouvertes.fr/hal-00909904/document
https://hal.archives-ouvertes.fr/hal-00909904/file/Integrative_biomechanics_for_tree_Fournier_al.pdf
DOI : 10.1093/jxb/ert279

Éditeur(s) : HAL CCSD
COST E50 Workshop: Reaction wood formation at stand, tree and cellular level

hal-00797326
https://hal.archives-ouvertes.fr/hal-00797326

Éditeur(s) : HAL CCSD Oxford University Press (OUP): Policy B - Oxford Open Option A
Résumé :  
absent
Journal of Experimental Botany

hal-01032195
https://hal.archives-ouvertes.fr/hal-01032195
DOI : 10.1093/jxb/erp276

Éditeur(s) : HAL CCSD
Résumé : International audience
The main biological function of reaction wood is to act as “muscle” for trees, enabling them to control their posture and right stems when they are permanently disturbed from their set-point inclination by gravity or winds. The key property to achieve this function is the development of high mechanical stress during the formation of reaction wood cells, called “maturation strains”. Moreover, this function is basically performed through the asymmetric formation of wood (in quantity or quality) around the tree circumference. This asymmetry enables stems to bend upward or to compensate for the downward bending induced by gravity, as an asymmetric force associated to maturation strains creates a bending moment (i.e. one side of the stem “pulls” the other one). We will discuss the traits (cross sectional size and shape, wood features as G fibers or microfibril angle, eccentric growth even in usual or extreme cases as buttresses, total biomass growth and height of center of mass, stem lean and set-point angle) involved to understand how the posture control works. We will give simple formulas to estimate a “muscle efficiency” from biomechanical modeling. We will give practical methods to estimate all the involved traits, especially maturation strains from direct and indirect measurements, in peripheral functional wood or by retrospective dendrochronological analysis. Then, we will illustrate how tree stem tropisms and especially gravitropism, widely studied by physiologists, are key processes in natural ecosystems, through practical examples from tropical rainforest tree communities and regeneration of European mixed beech forests. Using the previous definition of postural control efficiency, we will show that the cross section size is rapidly a limiting factor, opposing the extreme strategies of flexible small stems, which bend easily but also right themselves quickly, to stiff big stems. Lastly, as wood in living trees ensures general storage, defence, vascular and skeletal functions, we will ask general questions about synergies and trade-offs between posture control and other functions.
Botany 2014 : new frontiers in botany
Boise, United States

hal-01627031
https://hal.archives-ouvertes.fr/hal-01627031