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

Éditeur(s) : HAL CCSD Oxford University Press (OUP)
Résumé : Publication Inra prise en compte dans l'analyse bibliométrique des publications scientifiques mondiales sur les Fruits, les Légumes et la Pomme de terre. Période 2000-2012. http://prodinra.inra.fr/record/256699
Background and Aims Plastic tree-shelters are increasingly used to protect tree seedlings against browsing animals and herbicide drifts. The biomass allocation in young seedlings of deciduous trees is highly disturbed by common plastic tree-shelters, resulting in poor root systems and reduced diameter growth of the trunk. The shelters have been improved by creating chimney-effect ventilation with holes drilled at the bottom, resulting in stimulated trunk diameter growth, but the root deficit has remained unchanged. An experiment was set up to elucidate the mechanisms behind the poor root growth of sheltered Prunus avium trees. Methods Tree seedlings were grown either in natural windy conditions or in tree-shelters. Mechanical wind stimuli were suppressed in ten unsheltered trees by staking. Mechanical stimuli (bending) of the stem were applied in ten sheltered trees using an original mechanical device. Key Results Sheltered trees suffered from poor root growth, but sheltered bent trees largely recovered, showing that mechano-sensing is an important mechanism governing C allocation and the shoot-root balance. The use of a few artificial mechanical stimuli increased the biomass allocation towards the roots, as did natural wind sway. It was demonstrated that there was an acclimation of plants to the imposed strain. Conclusions This study suggests that if mechanical stimuli are used to control plant growth, they should be applied at low frequency in order to be most effective. The impact on the functional equilibrium hypothesis that is used in many tree growth models is discussed. The consequence of the lack of mechanical stimuli should be incorporated in tree growth models when applied to environments protected from the wind (e.g. greenhouses, dense forests).
ISSN: 0305-7364

hal-01032056
https://hal.archives-ouvertes.fr/hal-01032056
DOI : 10.1093/aob/mcn054

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 Public Library of Science
Résumé : International audience
Domestication can influence many functional traits in plants, from overall life-history and growth form to wood density and cell wall ultrastructure. Such changes can increase fitness of the domesticate in agricultural environments but may negatively affect survival in the wild. We studied effects of domestication on stem biomechanics in manioc by comparing domesticated and ancestral wild taxa from two different regions of greater Amazonia. We compared mechanical properties, tissue organisation and wood characteristics including microfibril angles in both wild and domesticated plants, each growing in two different habitats (forest or savannah) and varying in growth form (shrub or liana). Wild taxa grew as shrubs in open savannah but as lianas in overgrown and forested habitats. Growth form plasticity was retained in domesticated manioc. However, stems of the domesticate showed brittle failure. Wild plants differed in mechanical architecture between shrub and liana phenotypes, a difference that diminished between shrubs and lianas of the domesticate. Stems of wild plants were generally stiffer, failed at higher bending stresses and were less prone to brittle fracture compared with shrub and liana phenotypes of the domesticate. Biomechanical differences between stems of wild and domesticated plants were mainly due to changes in wood density and cellulose microfibril angle rather than changes in secondary growth or tissue geometry. Domestication did not significantly modify "large-scale" trait development or growth form plasticity, since both wild and domesticated manioc can develop as shrubs or lianas. However, "finer-scale" developmental traits crucial to mechanical stability and thus ecological success of the plant were significantly modified. This profoundly influenced the likelihood of brittle failure, particularly in long climbing stems, thereby also influencing the survival of the domesticate in natural situations vulnerable to mechanical perturbation. We discuss the different selective pressures that could explain evolutionary modifications of stem biomechanical properties under domestication in manioc.
PLoS ONE

halsde-00917926
https://hal.archives-ouvertes.fr/halsde-00917926
https://hal.archives-ouvertes.fr/halsde-00917926/document
DOI : 10.1371/journal.pone.0074727

Éditeur(s) : HAL CCSD Elsevier
Résumé : International audience
Wood has a hierarchical structure involving several levels of organisation. The stiffness of wood relies on its capacity to transfer mechanical stress to its stiffest element at the lowest scale, namely crystalline cellulose. This study aims at quantifying to what extend crystalline cellulose contributes to wood stiffness depending on its moisture content. The crystal strains of cellulose were measured using X-ray diffraction on wet and dry specimens of spruce, based on a previously published methodology. The comparison between crystal strain and macroscopic strain shows that, during elastic loading, cellulose strain is lower than macroscopic strain. The means ratio of crystal/macroscopic strain amounts 0.85 for dry specimens and 0.64 for wet specimens. This strain ratio cannot be explained just by the projection effect due to the difference in orientation between cellulose microfibrils and cell wall, but results from deformation mechanisms in series with cellulose. Analysis shows that this series contribution represents a non-negligible contribution to wood compliance and is strongly moisture-dependent. This contribution amounts 9% for dry specimens and 33% for wet specimens, corresponding to a 4-fold increase in compliance for the series contribution. The origin of these strains is ascribed to mechanisms involving bending or shear strain at different scales, due to the fact that reinforcing element are neither perfectly straight nor infinitely long.
ISSN: 0266-3538

hal-01425031
https://hal.archives-ouvertes.fr/hal-01425031
https://hal.archives-ouvertes.fr/hal-01425031/document
https://hal.archives-ouvertes.fr/hal-01425031/file/Art_Almeras_al_Contribution_cellulose_moisture-dependent_elastic_behavior_2017.pdf
DOI : 10.1016/j.compscitech.2016.11.025

Éditeur(s) : HAL CCSD Springer-Verlag Berlin Heidelberg
Résumé : International audience
The main biological function of reaction wood is to act as " muscle " for trees, enabling them to control their posture. The key property to achieve this function is the development of high mechanical stress during the formation of reaction wood cells, called " maturation strains ". Actually, reaction wood formation is basically the asymmetric formation of wood around the tree circumference, with higher maturation strains on the side where reaction wood is formed than on the opposite side. This asymmetry enables stems to bend upward or to compensate for the downward bending induced by gravity. At the cross section level, the performance in this biological function is linked not only to the magnitude of this asymmetry but also to an effect of the cross-sectional size (diameter) of the stem. Eccentric growth and variations in wood mechanical stiffness are second order effects that can modify this performance. Differences in maturation strains between reaction and non-reaction woods are related to their specific cell wall structure and composition. The swelling of the cell wall matrix during maturation and the effect of microfibril angle explain the differences in maturation strains between normal and compression wood. However, this mechanism fails in explaining the high maturation shrinkage of tension wood, and several hypotheses at the molecular levels are still under debate. How trees perceive their gravitational disequilibrium is also an open question for physiologists. Integrative biomechanical modelling (from the polymer level to the cell wall, cross section and whole tree levels) enables defining key variables that explain the performance of reaction wood as a system that insures the stem motricity. Maturation strains can be precisely measured only in recently formed wood at the tree surface, but their changes during the whole tree life can also be estimated by retrospective dendrochronological analysis through structural markers of reaction wood. Lastly, wood in living trees ensures general storage, defence, vascular and skeletal functions, that ask general questions about synergies and trade-offs as the structural characteristics of reaction wood can affect all these functions.
The Biology of Reaction Wood

ISBN : 978-3-642-10813-6
hal-01452085
https://hal.archives-ouvertes.fr/hal-01452085
https://hal.archives-ouvertes.fr/hal-01452085/document
https://hal.archives-ouvertes.fr/hal-01452085/file/Chap_Gril_al_Biomechanical_action_biological_functions_2014.pdf

Éditeur(s) : HAL CCSD Elsevier
Résumé : International audience
The late Carboniferous accretionary system of the South Tianshan orogen (North-Western China) underwent complex structural and polymetamorphic evolution. Combined petrological, geochronological and microstructural analysis of (ultra)high-pressure (UHP) metabasites (eclogites and blueschists) enclosed in metapelites show a relict coarse-grained eclogitic fabric S2 surrounded by a dominant fine-grained eclogite and blueschist facies retrograde fabric S2. The S2 fabric is reworked by upright folds F3 that are responsible for a major shortening of the whole accretionary system. For both the eclogite and blueschist, peak and retrograde P–T conditions have been thermodynamically constrained at 25–26 kbar and 425–500 °C and 10–13 kbar and 500−550 °C respectively, suggesting a shared exhumation history. The garnet-whole rock-amphibole isochron in the blueschist yielded Lu–Hf and Sm–Nd ages of 326.0 ± 2.9 Ma and 318.4 ± 3.9 Ma respectively, interpreted to date the prograde to peak metamorphic assemblage. The retrograde path of the eclogite is characterized by heterogeneous omphacite recrystallization into a mylonitic fine-grained matrix and crystallization of blue amphibole. Microstructures in both pristine porphyroclastic and recrystallized fine-grained domains in the eclogite indicate a gradual evolution from constriction-dominated (L>S-type) to flattening-dominated (S>L-type) type of deformation, increase of fabric intensity reflected by gradually growing M-indexes and the development of lattice preferred orientation (LPO) typical for dislocation creep under slightly hydrated conditions. Recrystallization of the matrix in the blueschist is homogeneous, which indicates a matrix dominated channel flow during exhumation. These LPOs evolutions suggest a significant mechanical coupling with the upper plate concomitant with oroclinal bending of the Kazakh orocline. Lock up of Kazakh orocline is responsible for further stress increase resulting in horizontal shortening of South Tianshan accretionary wedge and development of D3 upright folding and steepening of the whole sequence.
ISSN: 1342-937X

hal-01481263
https://hal.archives-ouvertes.fr/hal-01481263
DOI : 10.1016/j.gr.2016.07.007

Éditeur(s) : HAL CCSD
Résumé : Several major earthquakes have struck the 2500 km long Himalayan range during the past century. However, both the maximum size of such earthquakes and the probability of occurrence of a magnitude 9 megaquake are still a mater of debate. In this framework it is of key importance to investigate the lateral segmentation of the Himalayan arc in order to deepen our understanding of the mechanisms that control such events as well as their extents. Besides the remarkably uniform tectonic set-up and the fact that most studies on the Himalayas have focused on Central Nepal, several indications of along strike changes can be gathered.This thesis brings new constraints on lateral variations of crustal structure and of deformation between western Nepal (~80°E) and eastern Bhutan (~92°E).The first part of this thesis focuses on the lateral variations of crustal structure. In the past three years we have carried out four land gravity campaigns. We have established 366 new gravity measurement points to fill the data gaps in central and western Nepal as well as in Bhutan. This new dataset, together with available seismological data, is used to constrain thermo-mechanical models of the bending of the India plate underneath the Himalayas. While the inferred crustal geometry does not exhibit major along-strike variations over central and eastern Nepal, the shorter wavelength flexure of the lithosphere in Bhutan is associated with a weaker upper mantle rheology.The second part focuses on the lateral variations of recent deformation in the Himalayas. Fifteen years of paleoseismological investigations have extended the catalogue of major Himalayan events over the last millennium. Combining these information with secular loading, we assess the spatial and temporal stress changes on the Main Himalayan Thrust along the orogen over the last nine centuries. Our calculations indicate that inter-seismic loading has now nearly overcome the Coulomb stress decrease caused by the great ~1100 medieval earthquake. Our results also point out that the 1897 Shillong plateau earthquake does not have a major influence on the stress accumulated on the Main Himalayan Thrust since then. In order to better characterize active tectonics in the Bhutan Himalayas where no studies were done so far, we carried out a morphotectonic analysis in the south-central part of the kingdom. We show that the same amount of Holocene deformation is accommodated on the frontal thrust in Bhutan as on the neighbouring portions of the Himalayan arc. We also find evidences for two major (M>8) earthquakes on this thrust in the Bhutan Himalayas during the last millennium. Our results therefore show that Bhutan cannot be considered as a seismic gap. They also challenge the interpretation of one single ~1100 medieval mega-event that would have ruptured the Main Frontal Thrust from central Nepal to eastern Bhutan.
Au cours du dernier siècle, plusieurs séismes majeurs ont affecté l'Himalaya. Cependant, la taille maximale de ces événements et la probabilité d'occurrence de méga-séismes avec des magnitudes proches de 9 sont toujours matière à débat. L'étude de la segmentation de l'arc Himalayen est donc primordiale afin de comprendre les mécanismes qui contrôlent ces séismes ainsi que leur extension spatiale. La compréhension du cycle sismique en Himalaya est aujourd'hui essentiellement basée sur des études menées au Népal central, ce qui limite notre connaissance de son fonctionnement tri-dimensionnel. Ce travail de thèse permet d'apporter de nouvelles contraintes sur les variations latérales de la déformation crustale dans la zone comprise entre l'ouest Népal (~80°E) et l'est Bhoutan (~92°E). La première partie de cette thèse est consacrée à l'étude des variations latérales de la structure lithosphérique. Quatre campagnes gravimétriques ont été réalisées entre 2010 et 2012. Elles ont permis d'étendre le réseau gravimétrique Népalais jusqu'à l'ouest Népal et d'établir le premier réseau gravimétrique Bhoutanais. Ce nouveau jeu de données, combiné avec les données sismologiques disponibles, permet de contraindre des modèles thermo-mécaniques de la flexure de la plaque Indienne sous l'Himalaya. Les résultats suggèrent qu'il n'existe pas de variations latérales majeures du comportement mécanique de la lithosphère Indienne entre le centre et l'est Népal contrairement au Bhoutan où une rigidité flexurale plus faible est nécessaire pour expliquer les données.La deuxième partie de cette thèse est consacrée à l'étude des variations latérales des déformations récentes dans le prisme Himalayen. Les études paléosismologiques menées depuis 15 ans en Himalaya ont permis d'étendre le catalogue des événements majeurs sur le dernier millénaire. En intégrant à la fois les contraintes disponibles sur ces séismes majeurs et le chargement séculaire, nous étudions les variations spatiales et temporelles de l'état de contrainte du chevauchemlent Himalayen principal le long de l'arc. Nos calculs montrent que même dans la cas où un méga-séisme se serait produit en ~1100 en Himalaya, le chargement séculaire a quasiment compensé la chute de contrainte associée. Les modèles montrent aussi que le séisme du Shillong ne joue pas un rôle majeur sur les contraintes accumulées sur le chevauchement Himalayen depuis 1897 au niveau du Bhoutan. Enfin, nous présentons les premières contraintes sur la tectonique active au Bhoutan. L'étude morphotectonique réalisée au centre du Sud-Bhoutan a permis de montrer que la majeure partie de la déformation Holocène est accommodée au niveau du front Himalayen, comme au Népal. Nous montrons aussi des évidences de ruptures de surface avec des décalages verticaux de plusieurs mètres associés à deux séismes majeurs sur le dernier millénaire. Ces évidences remettent en cause l'interprétation d'un méga-séisme en ~1100.
https://tel.archives-ouvertes.fr/tel-01002381

NNT : 2013MON20074
tel-01002381
https://tel.archives-ouvertes.fr/tel-01002381
https://tel.archives-ouvertes.fr/tel-01002381/document
https://tel.archives-ouvertes.fr/tel-01002381/file/35970_BERTHET_2013_archivage_cor.pdf

Éditeur(s) : HAL CCSD Elsevier
Résumé : We present field data from segmented normal faults having particular displacement and overlapping geometries that may be related to down-dip branching of the fault segments. Based on a 3-D numerical modeling study of computed displacement and stress fields around different geometries of down-dip branched normal fault segments, we show that the bends of fault surfaces that coalesce at depth exert a significant influence on (I) the displacement distribution on the fault segment branches and (2) the quasi-static stress fields around the relay zones. The asymmetry in the displacement profiles and low fault interaction at relay zones modeled are consistent with the fault segment geometries observed in field data. As an application, we model the geometry and interaction of the Vallo di Diano normal fault which is segmented at the Earth's surface and which produced an earthquake of magnitude 6.4 along a single fault surface at depth. Numerical modeling of the segmented fault produces an asymmetric displacement profile and low shear stress at the relay zone consistent with the profile and fault interaction inferred from the held. We conclude that asymmetry of the displacement profile and large overlap/separation ratio of the unlinked relay zone can be indicators of the presence of a continuous fault surface underneath. These geometrical attributes are therefore important to consider in the probabilistic analysis of seismic hazard along segmented normal faults.
ISSN: 0191-8141

hal-00411161
https://hal.archives-ouvertes.fr/hal-00411161
DOI : 10.1016/j.jsg.2008.07.005

Éditeur(s) : HAL CCSD Springer Verlag (Germany) Springer Verlag
Résumé : Dehydration reactions result in minerals’ replacement and a transient fluid-filled porosity. These reactions involve interface-coupled dissolution–precipitation and might therefore lead to fixed crystallographic orientation relations between reactant (protolith) and product phases (i.e. topotaxy). We investigate these two phenomena in the dehydration of a foliated antigorite (atg) serpentinite by comparing the crystallographic preferred orientation (CPO) developed by olivine (ol), orthopyroxene (opx) and chlorite (chl) during high-pressure antigorite and chlorite dehydration in piston-cylinder experiments and in natural samples recording the dehydration of antigorite (Cerro del Almirez, Betic Cordillera, Spain). Experiments were performed under undrained conditions resulting in fluid-filled porosity and in strong CPO of the prograde minerals, controlled by the pre-existing antigorite CPO in the reactant foliated serpentinite. The orientation of a ol,opx and c∗chl is parallel to c∗atg from the protolith. The Cerro del Almirez samples show similar, locally well-developed topotactic relations between orthopyroxene, chlorite and antigorite, but the product CPOs are weaker and more complex at the thin section scale. In contrast to the experiments, olivine from natural samples shows a weak correlation between b ol and the former c∗atg. We relate the strengthening of local topotactic relations and the weakening of the inherited CPO at a larger scale in natural samples to compaction and associated fluid migration. Microstructural features that might be related to compaction in the natural samples include: (1) smooth bending of the former foliation, (2) gradual crystallographic misorientation (up to 16°) of prismatic orthopyroxene due to buckling by dislocation creep, (3) inversion of enstatite to low clinoenstatite (P21/c) along lamellae and (4) brittle fracturing of prismatic orthopyroxene enclosed by plastically deformed chlorite. The coexistence of orthopyroxene buckling and clinoenstatite lamellae enables estimating the local strain rates and shear stresses generated during compaction. An lower bound for the strain rates in the order of 10−12 to 10−13 s−1 and shear stresses of 60–70 MPa are estimated based on creep data. Lower shear stresses (20–40 MPa) are retrieved using a theoretical approach. These data point to slow compaction (and fluid extraction) in nature if the system is not perturbed by external forces, with rates only marginally higher than the viscoplastic deformation of the solid matrix.
ISSN: 0010-7999

hal-01174158
https://hal.archives-ouvertes.fr/hal-01174158
DOI : 10.1007/s00410-015-1129-4

Éditeur(s) : HAL CCSD Oxford University Press (OUP): Policy B - Oxford Open Option B
Résumé : International audience
To recover verticality after disturbance, angiosperm trees produce " tension wood " allowing them to bend actively. The driving force of the tension has been shown to take place in the G-layer, a specific unlignified layer of the cell wall observed in most temperate species. However, in tropical rain forests, the G-layer is often absent and the mechanism generating the forces to reorient trees remains unclear. A study was carried out on tilted seedlings, saplings and adult Simarouba amara trees – a species known to not produce a G-layer. Microscopic observations were done on sections of normal and tension wood after staining or observed under UV light to assess the presence/absence of lignin. We showed that S. amara produces a cell-wall layer with all of the characteristics typical of G-layers, but that this G-layer can be observed only as a temporary stage of the cell-wall development because it is masked by a late lignification. Being thin and lignified, tension wood fibres cannot be distinguished from normal wood fibres in the mature wood of adult trees. These observations indicate that the mechanism generating the high tensile stress in tension wood is likely to be the same as that in species with a typical G-layer and also in species where the G-layer cannot be observed in mature cells.
ISSN: 0829-318X

hal-01229280
https://hal.archives-ouvertes.fr/hal-01229280
https://hal.archives-ouvertes.fr/hal-01229280/document
https://hal.archives-ouvertes.fr/hal-01229280/file/Roussel-Clair-2015-TreePhysiol-LateLignifGLayerSimarouba_preprintHAL.pdf
DOI : 10.1093/treephys/tpv082