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
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 Springer Verlag
Résumé : A hypothetical mechanism for the generation of maturation stress in wood was tested experimentally. The hypothesis was that the maturation stress could partly originate in a physical mechanism related to daily changes in water pressure and associated diurnal strains. The matrix of lignin and hemicellulose, deposited in the cell wall during the night, would be put in compression by the effect of water tension during the next day. The cellulose framework, crystallizing during the day, would be put in tension by the decrease in tension at night and subsequent cell-wall swelling. This was tested on young saplings of sugi and beech. Half of the saplings were submitted to continuous lighting, which canceled diurnal strains. Saplings were tilted 40 degrees, and their uprighting movement was measured. The uprighting movement is directly due to the production of reaction wood and the concomitant development of large longitudinal maturation stress. It occurred in the continuously lighted plants at least as much as in control plants. We conclude that the generation of longitudinal maturation stress in tension or compression wood is not directly related to variations in water pressure and diurnal strains.
ISSN: 1435-0211

hal-01031891
https://hal.archives-ouvertes.fr/hal-01031891
DOI : 10.1007/s10086-005-0788-6

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 Wiley
Résumé : International audience
In order to progress in the understanding of mechanical stress generation, the mesoporosity of the cell wall and its changes during maturation of poplar (Populus deltoides x P. nigra) tension wood (TW) and opposite wood (OW) were measured by nitrogen adsorption–desorption. Variations in the thickness of the gelatinous layer (G-layer) were also measured to clarify whether the mesoporosity change simultaneously with the deposition of the G-layer in TW. Results show that mesoporous structures of TW and OW were very similar in early development stages before the deposition of G-layers. With the formation of the S2 layer in OW and the G-layer in TW, the mesopore volume decreased steeply before lignification. However, in TW only, the decrease in mesopore volume occurred together with the pore shape change and a progressive increase in pore size. The different patterns observed in TW revealed that pores from G-layers appear with a different shape compared to those of the compound middle lamella, and their size increases duringthe maturation process until stabilising in mature wood. This observation strongly supports the hypothesis of the swelling of the G-layer matrix during maturation as the origin of maturation stress in poplar tension wood.
ISSN: 0028-646X

hal-01081834
https://hal.archives-ouvertes.fr/hal-01081834
https://hal.archives-ouvertes.fr/hal-01081834/document
https://hal.archives-ouvertes.fr/hal-01081834/file/Chang_al_Mesoporosity_changes_New-Phyto._2015.pdf
DOI : 10.1111/nph.13126

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 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): 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 : 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 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