Éditeur(s) : HAL CCSD Springer Verlag
Résumé : International audience
We explored the relationship between soil processes, estimated through soil respiration (Rsoil), and the spatial variation in forest structure, assessed through the distribution of tree size, in order to understand the determinism of spatial variations in Rsoil in a tropical forest. The influence of tree size was examined using an index (Ic) calculated for each tree as a function of (1) the trunk cross section area and (2) the distance from the measurement point. We investigated the relationships between Ic and litterfall,root mass and Rsoil, respectively. Strong significant relationships were found between Ic and both litterfall and root mass. Rsoil showed a large range of variations over the 1-ha experimental plot, from 1.5 to 12.6 gC m−2 d−1. The best relationship between Ic and Rsoil only explained 17% of the spatial variation in Rsoil. These results support the assumption that local spatial patterns in litter production and root mass depend on tree distribution in tropical forests. Our study also emphasizes the modest contribution of tree size distribution-which is mainly influenced by the presence of the biggest trees (among the large range size of the inventoried trees greater than 10 cm diameter at 1.30 m above ground level or at 0.5 m above the buttresses)-in explaining spatial variations in Rsoil.
ISSN: 0032-079X

hal-00797064
https://hal.archives-ouvertes.fr/hal-00797064

Éditeur(s) : HAL CCSD Oxford University Press (OUP): Policy B - Oxford Open Option B
Résumé :  
absent
ISSN: 0829-318X

hal-01032128
https://hal.archives-ouvertes.fr/hal-01032128
DOI : 10.1093/treephys/28.10.1513

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 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/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 Oxford University Press (OUP): Policy B - Oxford Open Option B
Résumé :  
absent
ISSN: 0829-318X

hal-01032099
https://hal.archives-ouvertes.fr/hal-01032099
DOI : 10.1093/treephys/27.11.1505

Éditeur(s) : HAL CCSD Springer Verlag
Résumé : International audience
Transverse drying shrinkage was measured at microscopic and mesoscopic levels in poplar wood characterised by an increasing growth strain (GS), from normal to tension wood. Results show that: (a) The drying shrinkage, measured as a relative thickness decrease, was significantly higher for G-layer (GL) than the other layers (OL), GL shrinkage was not significantly correlated with GS and OL shrinkage was negatively correlated with GS. (b) In gelatinous fibre (G-fibre), lumen size increased during drying and this increase was positively related with GS, but in normal wood fibre lumen size decreased during drying. These findings suggest that GL shrank outwards, so that its shrinkage feebly affected the total cell shrinkage and the mesoscopic shrinkage was controlled by the OL shrinkage which shrank inwards. (c) Measurements done on 7×7 mm² thin sections evidenced a negative correlation between transverse shrinkage and GS, significant in T direction but weak in R direction. These observations at both levels allow to discuss the contribution of GL to the mesoscopic shrinkage of tension wood.
ISSN: 0043-7719

hal-00194930
https://hal.archives-ouvertes.fr/hal-00194930
https://hal.archives-ouvertes.fr/hal-00194930/document
https://hal.archives-ouvertes.fr/hal-00194930/file/WST_FANG2006.pdf
DOI : 10.1007/s00226-007-0148-3

Éditeur(s) : HAL CCSD Walter de Gruyter
Résumé : absent
Holzforschung

hal-01031939
https://hal.archives-ouvertes.fr/hal-01031939
DOI : 10.1515/HF.2005.057

Éditeur(s) : HAL CCSD Springer Verlag (Germany)
Résumé :  
We explored the relationship between soil processes, estimated through soil respiration (Rsoil), and the spatial variation in forest structure, assessed through the distribution of tree size, in order to understand the determinism of spatial variations in Rsoil in a tropical forest. The influence of tree size was examined using an index (Ic) calculated for each tree as a function of (1) the trunk cross section area and (2) the distance from the measurement point. We investigated the relationships between Ic and litterfall, root mass and Rsoil, respectively. Strong significant relationships were found between Ic and both litterfall and root mass. Rsoil showed a large range of variations over the 1-ha experimental plot, from 1.5 to 12.6 gC m−2 d−1. The best relationship between Ic and Rsoil only explained 17% of the spatial variation in Rsoil. These results support the assumption that local spatial patterns in litter production and root mass depend on tree distribution in tropical forests. Our study also emphasizes the modest contribution of tree size distribution-which is mainly influenced by the presence of the biggest trees (among the large range size of the inventoried trees greater than 10 cm diameter at 1.30 m above ground level or at 0.5 m above the buttresses)-in explaining spatial variations in Rsoil.
Plant and Soil

hal-01032200
https://hal.archives-ouvertes.fr/hal-01032200
DOI : 10.1007/s11104-011-0848-1

Éditeur(s) : HAL CCSD Royal Society
Résumé : International audience
Trees control their posture by generating asymmetric mechanical stress around the periphery of the trunk or branches. This stress is produced in wood during the maturation of the cell wall. When the need for reaction is high, it is accompanied by strong changes in cell organization and composition called reaction wood, namely compression wood in gymnosperms and tension wood in angiosperms. The process by which stress is generated in the cell wall during its formation is not yet known, and various hypothetical mechanisms have been proposed in the literature. Here we aim at discriminating between these models. First, we summarize current knowledge about reaction wood structure, state and behaviour relevant to the understanding of maturation stress generation. Then, the mechanisms proposed in the literature are listed and discussed in order to identify which can be rejected based on their inconsistency with current knowledge at the frontier between plant science and mechanical engineering.
Journal of the Royal Society Interface

hal-01392832
https://hal.archives-ouvertes.fr/hal-01392832
https://hal.archives-ouvertes.fr/hal-01392832/document
https://hal.archives-ouvertes.fr/hal-01392832/file/AlmerasClair2016_JRSInterface_Critical%20review%20on%20the%20mechanisms%20of%20maturation%20stress%20generation%20in%20trees_HAL.pdf
DOI : 10.1098/rsif.2016.0550

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
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 Springer Verlag
Résumé : Tangential strains were measured with strain gauges at the surface of xylem and inner bark of saplings of Cryptomeria japonica D. Don. and Fagus silvatica L. during a pressurization test. The test consists in submitting the whole sapling to an artificially imposed hydrostatic pressure of increasing magnitude. The elastic response of the stems was found linear both at the surface of xylem and inner bark. A simple geometric model allows to compute radial strains in each tissue from tangential strain data. Inside inner bark, radial strains are much larger than tangential strains, because tangential strains are restrained by the core of wood. The material compliance of each tissue was computed as the ratio between the radial strain and the pressure that caused it. The material compliance of xylem is much lower than that of inner bark, but, as its thickness is much larger, its contribution to the apparent behavior of the stem is not negligible. Computation of material compliances by this pressurization test provides information about the specific behavior of each tissue in response to hydrostatic pressure. This can be used to estimate and interpret the calibration factor linking the water status of the plant to the apparent strain measured at its surface.
ISSN: 0931-1890

hal-01031900
https://hal.archives-ouvertes.fr/hal-01031900
DOI : 10.1007/s00468-006-0061-7

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 : 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 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 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 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
COST E50 Workshop: Reaction wood formation at stand, tree and cellular level

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