Éditeur(s) : HAL CCSD
Résumé : International audience
The anatomy of tension wood and opposite wood was compared in 21 tropical rain forest trees from 21 species belonging to 18 families from French Guyana. Wood specimens were taken from the upper and lower sides of naturally tilted trees. Measurement of the growth stress level ensured that the two samples were taken from wood tissues in a different mechanical state: highly tensile-stressed wood on the upper side, called tension wood and normally tensile-stressed wood on the lower side, called opposite wood. Quantitative parameters relating to fibres and vessels were measured on transverse sections of both tension and opposite wood to check if certain criteria can easily discriminate the two kinds of wood. We observed a decrease in the frequency of vessels in the tension wood in all the trees studied. Other criteria concerning shape and surface area of the vessels, fibre diameter or cell wall thickness did not reveal any general trend. At the ultrastructural level, we observed that the microfibril angle in the tension wood sample was lower than in opposite wood in all the trees except one (Licania membranacea).
IAWA Journal

hal-00194915
https://hal.archives-ouvertes.fr/hal-00194915
https://hal.archives-ouvertes.fr/hal-00194915/document
https://hal.archives-ouvertes.fr/hal-00194915/file/IAWA2006_RuelleClairBeauchenePrevostFournier_ccsd.pdf
IRD : PAR00001321

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
Résumé : International audience
Tension wood, a tissue developed by angiosperm trees to actively recover their verticality, has long been defined by the presence of an unlignified cellulosic inner layer in the cell wall of fibres, called the G-layer. Although it was known that some species have no G-layer, the definition was appropriate since it enabled easy detection of tension wood zones using various staining techniques for either cellulose or lignin. For several years now, irrespective of its anatomical structure, tension wood has been defined by its high mechanical internal tensile stress. This definition enables screening of the diversity of cell walls in tension wood fibres. Recent results obtained in tropical species with tension wood with a delay in the lignification of the G-layer opened our eyes to the effective presence of large amounts of lignin in the G-layer of some species. This led us to review older literature mentioning the presence of lignin deposits in the G-layer and give them credit. Advances in the knowledge of tension wood fibres allow us to reconsider some previous classifications of the diversity in the organisation of the fibre walls of the tension wood.
ISSN: 0928-1541

hal-01636686
https://hal.archives-ouvertes.fr/hal-01636686
https://hal.archives-ouvertes.fr/hal-01636686/document
https://hal.archives-ouvertes.fr/hal-01636686/file/2017_Ghislain-Clair_IAWAJ_DiversityOrganisationLignificationTensionWoodFibreWalls-AReview.pdf
DOI : 10.1163/22941932-20170170

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 :  
The anatomy of tension wood and opposite wood was compared in 21 tropical rain forest trees from 21 species belonging to 18 families from French Guyana. Wood secimens were taken from the upper and lower sides of naturally titled trees. Measurement of the growth stress level ensured that the two samples were taken from wood tissues in a different mechanical state: highly tensile-stressed wood on the upper side, called tension wood and normally tensile-stressed wood on the lower side, called opposite wood. Quantitative parameters relating to fibres and vessels were measured on transverse sections of both tension and opposite wood to check if certain criteria can easily discriminate the two kinds of wood. We observed a decrease in the frequency of vessels in the tension wood in all the trees studied. Other criteria concerning shape and surface area of the vessels, fibre diameter or cell wall thickness did not reveal any general trend.At the ultrastructural level, we observed that the microfibril angle in the tension wood sample was lower than in opposite wood in all the trees except one (Licania membranacea).
IAWA Journal

hal-01032149
https://hal.archives-ouvertes.fr/hal-01032149

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 Springer Verlag
Résumé :  
Growth strains were measured in situ in nine trees of three species from a French Guiana tropical rainforest in a clearly active verticality restoration process. The aim was to detect tension wood within the samples. Wood specimens were cut in the vicinity of the growth strain measurements in order to determine the microfibril angle and some mechanical and physical properties. As suspected, tensile growth strain was much higher in tension wood zones, as shown by the slightly higher longitudinal modulus of elasticity. Conversely, tension wood showed reduced compression strength. Longitudinal shrinkage was much higher in tension wood than in opposite wood. Clear relationships between the microfibril angle and longitudinal properties were noted in comparison (i) with those observed in gymnosperm compression wood and (ii) with expected relationships from the organization of wood fibres cell wall structure.
ISSN: 0043-7719

hal-01032114
https://hal.archives-ouvertes.fr/hal-01032114
DOI : 10.1007/s00226-010-0323-9

Éditeur(s) : HAL CCSD Springer Verlag
Résumé : International audience
Tension wood of Laetia procera (Poepp.) Eichl. (Flacourtiaceae), a neo-tropical forest species, shows a peculiar secondary wall structure, with an alternance of thick and thin layers, while opposite wood of this species has a typical secondary wall structure (S1 + S2 + S3). Samples for the study of microstructural properties were collected upon the estimation of growth stresses in the living tree, in order to analyze the correlation of the former with the latter. Investigation using optical microscopy, scanning electron microscopy and UV microspectrophotometry allowed the description of the anatomy, ultra-structure and chemistry of this peculiar polylaminate secondary wall. In the thick layers, cellulose microfibril angle is very low (i.e., microfibril orientation is close to fibre axis) and cellulose microfibrils are well organized and parallel to each other. In the thin layers, microfibrils (only observable in the inner layer) are less organized and are oriented with a large angle relative to the axis of the cell. Thick layers are lightly lignified although thin layers show a higher content of lignin, close to that of opposite wood secondary wall. The more the wood was under tensile stress, the less the secondary wall was lignified, and lower the syringyl on guaiacyl lignin units' ratio was. The innermost layer of the secondary wall looks like a typical S3 layer with large microfibril angle and lignin occurrence. The interest of this kind of structure for the understanding of stress generation is discussed.
ISSN: 0931-1890

hal-00194923
https://hal.archives-ouvertes.fr/hal-00194923
https://hal.archives-ouvertes.fr/hal-00194923/document
https://hal.archives-ouvertes.fr/hal-00194923/file/Trees_RuelleYoshidaClairThibaut_submitted2006.pdf
DOI : 10.1007/s00468-007-0128-0

Éditeur(s) : HAL CCSD Oxford University Press (OUP): Policy B - Oxford Open Option A
Résumé :  
The mechanism for tree orientation in angiosperms is based on the production of high tensile stress on the upper side of the inclined axis. In many species, the stress level is strongly related to the presence of a peculiar layer, called the G-layer, in the fibre cell wall. The structure of the G-layer has recently been described as a hydrogel thanks to N2 adsorption-desorption isotherms of supercritically dried samples showing a high mesoporosity (pores size from 2-50 nm). This led us to revisit the concept of the G-layer that had been, until now, only described from anatomical observation. Adsorption isotherms of both normal wood and tension wood have been measured on six tropical species. Measurements show that mesoporosity is high in tension wood with a typical thick G-layer while it is much less with a thinner G-layer, sometimes no more than normal wood. The mesoporosity of tension wood species without a G-layer is as low as in normal wood. Not depending on the amount of pores, the pore size distribution is always centred around 6-12 nm. These results suggest that, among species producing fibres with a G-layer, large structural differences of the G-layer exist between species.
Journal of Experimental Botany

hal-01032161
https://hal.archives-ouvertes.fr/hal-01032161
DOI : 10.1093/jxb/erp133

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 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 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