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 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 Springer Verlag (Germany)
Résumé : International audience
*Context: Tree orientation is controlled by asymmetric mechanical stresses set during wood maturation. The magnitude of maturation stress differs between longitudinal and tangential directions, and between normal and tension woods. * Aims: We aimed at evaluating patterns of maturation stress on eucalypt plantation trees and their relation with growth, with a focus on tangential stress evaluation. * Methods: Released maturation strains along longitudinal and tangential directions were measured around the circumference of 29 Eucalyptus nitens trees, including both straight and leaning trees. * Results: Most trees produced asymmetric patterns of longitudinal maturation strain but more than half of the maturation strain variability occurred between trees. Many trees produced high longitudinal tensile stress all around their circumference. High longitudinal tensile stress was not systematically associated with the presence of gelatinous layer. The average magnitude of released longitudinal maturation strain was found negatively correlated to the growth rate. A methodology is proposed to ensure reliable evaluation of released maturation strain in both longitudinal and tangential directions. Tangential strain evaluated with this method was lower than previously reported. * Conclusion: The stress was always tensile along longitudinal direction and compressive along tangential direction, and their respective magnitude was positively correlated. This correlation does not result from a Poisson effect but may be related to the mechanism of maturation stress generation.
Annals of Forest Science

hal-00913611
https://hal.archives-ouvertes.fr/hal-00913611
https://hal.archives-ouvertes.fr/hal-00913611/document
DOI : 10.1007/s13595-013-0318-4

Éditeur(s) : HAL CCSD Springer Verlag/EDP Sciences
Résumé : International audience
In the dense tropical rainforest understorey, saplings exhibit different growth strategies aiming at reaching light levels better fitting their ecology. Investing mainly in height growth, at the expense of their width, a lot are close to mechanical instability. Tachigali melinonii, a long living heliophilic tree species, is frequently observed to be extremely slender and supported by neighbours. Such observations suggest an active growth control through the perception of mechanical environment. * Mechanical environment or light availability, which one is the most influent on growth and slenderness (H/D)? To test this question, we recorded growth of control and staked saplings of two species with contrasting habits and ecology: T. melinonii, and Dicorynia guianensis, along a natural light gradient. * Dicorynia, the more stable, responded more clearly to the staking treatment, showing slenderness increase when light is available, whereas for Tachigali, only light availability governed growth. * For Tachigali, growth allocation is mainly governed by light availability and ontogeny, whereas Dicorynia is probably similar to the average tree strategy, using the thigmomorphogenetic physiological process to control its stability.
ISSN: 1286-4560

hal-00534019
https://hal.archives-ouvertes.fr/hal-00534019
DOI : 10.1051/forest/2009104

Éditeur(s) : HAL CCSD Springer Verlag/EDP Sciences
Résumé : International audience
Context and aim : This study aimed to examine the effect of the tension wood G‐layer on the viscoelastic properties of wood. Methods : Tension wood and opposite wood samples were obtained from six French Guianese tropical rainforest species (Sextonia rubra, Ocotea guyanensis, Inga alba, Tachigali melinoni, Iyranthera sagotiana and Virola michelii); the tension wood of the former three of these species had a Glayer, whilst the tension wood from the latter three had no Glayer. Tensile dynamic mechanical analysis (DMA) was performed on green never dried wood samples in the longitudinal direction with samples submerged in a water bath at a temperature (30°C) and frequency (1 Hz) representative of the conditions experienced by wood within a living tree. Then, DMA was repeated with samples conditioned to an air-dried state. Finally, samples were oven-dried to measure longitudinal shrinkage. Results : Tension wood did not always have a higher longitudinal storage (elastic) modulus than opposite wood from the same tree regardless of the presence or absence of a G‐layer. For the species containing a G‐layer, tension wood had a higher damping coefficient and experienced a greater longitudinal shrinkage upon drying than opposite wood from the same species. No difference was found in damping coefficients between tension wood and opposite wood for the species that had no G‐layer. Conclusion : It is proposed that the different molecular composition of the G-layer matrix has an influence on the viscoelasticity of wood, even if a biomechanical gain is not yet clear. This study shows that rheological properties and longitudinal shrinkage can be used to detect the presence of a G‐layer in tension wood.
ISSN: 1286-4560

hal-00677460
https://hal.archives-ouvertes.fr/hal-00677460
https://hal.archives-ouvertes.fr/hal-00677460/document
https://hal.archives-ouvertes.fr/hal-00677460/file/Clair2.pdf
DOI : 10.1007/s13595-011-0164-1

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 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
Résumé : International audience
Trees can control their shape and resist gravity thanks to their ability to produce wood under tensile stress. This stress is known to be produced during the maturation of wood fibres but the mechanism of its generation remains unclear. This study focuses on the formation of the secondary wall in tension wood produced in artificially tilted poplar saplings. Thickness of secondary wall layer (SL) and gelatinous layer (GL) were measured from cambium to mature wood in several trees sampled at different times after tilting. Measurements on wood fibres produced before tilting show the progressive increase of secondary wall thickness during the growing season. After the tilting date, SL thickness decreased markedly from normal wood to tension wood while the total thickness increased compared to normal wood, with the development of a thick GL. However, even after GL formation, SL thickness continues to increase during the growing season. GL thickening was observed to be faster than SL thickening. The development of the unlignified GL is proposed to be a low cost, efficient strategy for a fast generation of tensile stress in broadleaved trees.
IAWA journal / International Association of Wood Anatomists.

hal-01229277
https://hal.archives-ouvertes.fr/hal-01229277
https://hal.archives-ouvertes.fr/hal-01229277/document
https://hal.archives-ouvertes.fr/hal-01229277/file/AbediniClair-Iawa-TensionCellWallThickening_preprint_HAL.pdf
DOI : 10.1163/22941932-00000084

Éditeur(s) : HAL CCSD Botanical Society of America
Résumé :  
Tree buckling risk (actual height/critical buckling height) is an important biomechanical trait of plant growth strategies, and one that contributes to species coexistence. To estimate the diversity of this trait among wide samples, a method that minimizes damage to the plants is necessary. On the basis of the rarely used, complete version of Greenhill's model (1881, Proceedings of the Cambridge Philosophical Society 4(2): 65-73), we precisely measured all the necessary parameters on a sample of 236 saplings of 16 species. Then, using sensitivity (variance) analysis, regressions between successive models for risk factors and species ranks and the use of these models on samples of self- and nonself-supporting saplings, we tested different degrees of simplification up to the most simple and widely used formula that assumes that the tree is a cylindrical homogeneous pole. The size factor had the greatest effect on buckling risk, followed by the form factor and the modulus of elasticity of the wood. Therefore, estimates of buckling risk must consider not only the wood properties but especially the form factor. Finally, we proposed a simple but accurate method of assessing tree buckling risk that is applicable to a wide range of samples and that requires mostly nondestructive measurements.
ISSN: 0002-9122

hal-01032059
https://hal.archives-ouvertes.fr/hal-01032059

Éditeur(s) : HAL CCSD
Résumé :  
The main aspects of wood mechanics are approached through the role played by wood as the building material of the tree. The main concepts used by nature: progressive setting up of tree weight and architecture, multifunctional role of wood and adaptability to hazard and long-term evolution, are clarified together with their consequence on the multilayered, anisotropic nature of wood. The technological choices of the plant world as the cellular structure, the use of composite with oriented fibres and hydro-carbonated polymers, as well as the systematic prestressing of every new layer, bear obvious consequences on the type of mechanical behaviour of wood. The local control of the level of prestressing through minor modifications of the cell-wall biosynthesis allows the tree to adapt to conditions of light exploration that will evolve according to time and risk occurrence. This analysis of wood genesis process permits in return to improve our understanding of the mechanical behaviour of the material in relation with the parameters of the microstructure.
Comptes Rendus de l'Académie des Sciences Série 2

hal-01032151
https://hal.archives-ouvertes.fr/hal-01032151
DOI : 10.1016/S1620-7742(01)01380-0

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
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 Springer Verlag
Résumé : Active mechanisms of re-orientation are necessary to maintain the verticality of tree stems. They are achieved through the production of reaction wood, associated with circumferential variations of three factors related to cambial activity: maturation strain, longitudinal modulus of elasticity (MOE) and eccentric growth. These factors were measured on 17 mature trees from different botanical families and geographical locations. Various patterns of circumferential variation of these factors were identified. A biomechanical analysis based on beam theory was performed to quantify the individual impact of each factor. The main factor of re-orientation is the circumferential variation of maturation strains. However, this factor alone explains only 57% of the re-orientations. Other factors also have an effect through their interaction with maturation strains. Eccentric growth is generally associated with heterogeneity of maturation strains, and has an important complementary role, by increasing the width of wood with high maturation strain. Without this factor, the efficiency of re-orientations would be reduced by 31% for angiosperms and 26% for gymnosperms. In the case of angiosperms, MOE is often larger in tension wood than in normal wood. Without these variations, the efficiency of re-orientations would be reduced by 13%. In the case of gymnosperm trees, MOE of compression wood is lower than that of normal wood, so that re-orientation efficiency would be increased by 24% without this factor of variations.
ISSN: 0931-1890

hal-01031825
https://hal.archives-ouvertes.fr/hal-01031825
DOI : 10.1007/s00468-005-0407-6

Éditeur(s) : HAL CCSD Springer Verlag/EDP Sciences
Résumé : International audience
Background : This study examines the radial trend in wood stiffness of tropical rainforest trees. The objective was to determine if the type of growing environment (exposed plantation or dense primary forest) would have an effect on this radial trend. Methods : The axial elastic modulus of wood samples, representing a pith to bark cross-section, of six trees from several French Guianese species (two of Eperua falcata, one of Eperua grandiflora, two of Carapa procera and one of Symphonia gloubulifera) was measured using a dynamic "forced vibration" method. Results : Primary forest trees were observed to have a decrease in wood stiffness from pith to bark, whereas plantation trees, from the same genus or species, displayed a corresponding increase in wood stiffness. Juvenile wood stiffness appears to vary depending on the environment in which the tree had grown. Conclusion : We suggest that the growth strategy of primary forest trees is to produce wood resistant to selfbuckling so that the height of the canopy may be obtained with the maximum of efficiency. In contrast, the growth strategy of the trees growing in an exposed plantation is to produce low-stiffness wood, important to provide flexibility in wind. Further experiments to study the behaviour of more species, with more individuals per species, growing across a range of physical environments, are required.
ISSN: 1286-4560

hal-00644335
https://hal.archives-ouvertes.fr/hal-00644335
https://hal.archives-ouvertes.fr/hal-00644335/document
https://hal.archives-ouvertes.fr/hal-00644335/file/McLean_al.pdf
DOI : 10.1007/s13595-011-0085-z

Éditeur(s) : HAL CCSD Oxford University Press (OUP)
Résumé : International audience
Background and Aims: In rain forests, sapling survival is highly dependent on the regulation of trunk slenderness (height/diameter ratio): shade-intolerant species have to grow in height as fast as possible to reach the canopy but also have to withstand mechanical loadings (wind and their own weight) to avoid buckling. Recent studies suggest that mechanosensing is essential to control tree dimensions and stability-related morphogenesis. Differences in species slenderness have been observed among rainforest trees; the present study thus investigates whether species with different slenderness and growth habits exhibit differences in mechanosensitivity. Methods: Recent studies have led to a model of mechanosensing (sum-of-strains model) that predicts a quantitative relationship between the applied sum of longitudinal strains and the plant's responses in the case of a single bending. Saplings of five different neotropical species (Eperua falcata, E. grandiflora, Tachigali melinonii, Symphonia globulifera and Bauhinia guianensis) were subjected to a regimen of controlled mechanical loading phases (bending) alternating with still phases over a period of 2 months. Mechanical loading was controlled in terms of strains and the five species were subjected to the same range of sum of strains. The application of the sum-of-strain model led to a dose-response curve for each species. Dose-response curves were then compared between tested species. Key Results: The model of mechanosensing (sum-of-strain model) applied in the case of multiple bending as long as the bending frequency was low. A comparison of dose-response curves for each species demonstrated differences in the stimulus threshold, suggesting two groups of responses among the species. Interestingly, the liana species B. guianensis exhibited a higher threshold than other Leguminosae species tested. Conclusions: This study provides a conceptual framework to study variability in plant mechanosensing and demonstrated interspecific variability in mechanosensing.
ISSN: 0305-7364

halsde-00454145
https://hal.archives-ouvertes.fr/halsde-00454145
DOI : 10.1093/aob/mcp286

Éditeur(s) : HAL CCSD Botanical Society of America
Résumé :  
International audience
The different hypotheses about buttress function and formation mainly involve mechanical theory. Forces were applied to two trees of Sloanea spp., a tropical genus that develops typical thin buttresses, and the three-dimensional strains were measured at different parts of the trunk base. Risks of failure were greater on the buttress sides, where shear and tangential stresses are greater, not on the ridges, in spite of high longitudinal (parallel to the grain) stresses. A simple beam model, computed from the second moment of area of digitized cross sections, is consistent with longitudinal strain variations but cannot predict accurately variations with height. Patterns of longitudinal strain variation along ridges are very different in the two individuals, owing to a pronounced lateral curvature in one specimen. The constant stress hypothesis is discussed based on these results. Without chronological data during the development of the tree, it cannot be proved that buttress formation is activated by stress or strain.
ISSN: 0002-9122

hal-01032155
https://hal.archives-ouvertes.fr/hal-01032155
https://hal.archives-ouvertes.fr/hal-01032155/document
https://hal.archives-ouvertes.fr/hal-01032155/file/Biomechanics_buttressed_trees_Clair_al_2003.pdf
DOI : 10.3732/ajb.90.9.1349

Éditeur(s) : HAL CCSD
Résumé : International audience
A key element of the biomechanical design of trees is their ability to generate largemechanical stresses in wood at the stem periphery. This function is necessary for the tree tocontrol the orientation of its axes, and therefore to grow in height, maintain its branches at anoptimal angle or achieve adaptive reorientations (Fournier et al., 2013). This “maturationstress” appears in wood fibres during their cellular maturation when their secondary cell wallis thickening. In hardwoods, the stress asymmetry is generated by the formation of specificfibres with a very high tensile growth stress on the upper side of the inclined axis. Theresulting tension wood has microstructural features highly different from the normal wood.Tension wood in almost all temperate species has a specific gelatinous cell wall layer, calledG-layer, that is not lignified and highly mesoporous (Chang et al., 2015). It has been shownrecently, at the macroscopic scale, that cellulose tension appears during maturation and issynchronous with the development of this specific layer (Clair et al., 2011). Tropical species,like simarouba, are able to generate lignified tension wood fibres. The mechanisms of stressgeneration in tension wood fibres are not yet well known. The aim of our study is tounderstand these mechanisms and to know the spatial and temporal kinetics of the differentcell wall layers stiffening during maturation, in parallel with tensile stress generation.Contact-Resonance Atomic Force Microscopy, CR-AFM, was used here to address thisquestion on embedded samples (Arnould and Arinero, 2015) using dual resonance frequencytracking (DRFT, Rodriguez et al., 2007). Change in the contact modulus between, and within,each layer of the cell wall has been measured on several radial lines of developing fibres atdifferent steps. In order to understand the obtained results, and to estimate the sensitivity ofthe AFM indentation-like technique to the different cell wall components stiffness, an elasticanisotropic indentation model was used (Vlassak et al., 2003; Jäger et al., 2011). Finally,these mechanical measurements have been compared to topochemical data obtained on thesame fibres.
8th Plant Biomechanics International Conference
Nagoya, Japan

hal-01436113
https://hal.archives-ouvertes.fr/hal-01436113

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

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