Éditeur(s) : HAL CCSD
Résumé :  
Wood samples were taken from the upper and lower sides of 21 naturally tilted trees from 18 families of angiosperms in the tropical rain forest in French Guyana. The measurement of growth stresses 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 lower tensile stressed wood on the lower side, called opposite wood. Eight species had tension wood fibres with a distinct gelatinous layer (G-layer). The distribution of gelatinous fibres varied from species to species. One of the species, Casearia javitensis (Flacourtiaceae), showed a peculiar multilayered secondary wall in its reaction wood. Comparison between the stress level and the occurrence of G-layer indicates that the G-layer is not a key factor in the production of high tensile stressed wood.
IAWA Journal

hal-01032051
https://hal.archives-ouvertes.fr/hal-01032051

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
Résumé : Wood samples were taken from the upper and lower sides of 21 naturally tilted trees from 18 families of angiosperms in the tropical rainforest in French Guyana. The measurement of growth stresses 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 lower tensile stressed wood on the lower side, called opposite wood. Eight species had tension wood fibres with a distinct gelatinous layer (G-layer). The distribution of gelatinous fibres varied from species to species. One of the species, Casearia javitensis (Flacourtiaceae), showed a peculiar multilayered secondary wall in its reaction wood. Comparison between the stress level and the occurrence of the G-layer indicates that the G-layer is not a key factor in the production of high tensile stressed wood.
IAWA Journal

hal-00112579
https://hal.archives-ouvertes.fr/hal-00112579
https://hal.archives-ouvertes.fr/hal-00112579/document
https://hal.archives-ouvertes.fr/hal-00112579/file/IAWA2006_ClairRuelleBeauchenePrevostFournier_ccsd.pdf
IRD : PAR00001136

Éditeur(s) : HAL CCSD
Résumé :  
The gelatinous layer (G-layer) of tension wood fibres in hardwood contributes to the mechanical function of the living tree and has significant consequences on properties of solid wood. Its size, shape and structure observed by optical or electron microscopy exhibits characteristic anatomical features. However, we found that sectioning of non-embedded wood samples results in an uncontrolled swelling of the G-layer was monitored by serial sections, from an embedded wood sample, from its trimmed transverse face to that located several hundreds of micrometers deep. The results revealed that the initial cutting before embedding produced a border effect responsible for the swollen nature, which is similar to sections from non-embedded material. After a conventional embedding technique was applied, a section of at least 30 micrometres below the trimming surface is required to observe an un-swollen G-layer.
IAWA Journal

hal-01032054
https://hal.archives-ouvertes.fr/hal-01032054

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 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 Springer Verlag
Résumé : Increasing interest in understanding tension stress generation in tension wood with fibres having a gelatinous layer (G-layer) has focused attention on the specific role of this layer. To distinguish its contribution from those of other wall layers, the G-layer of wood sections was isolated by ultrasonication. The aim of this study was to assess the effect of sectioning and of the ultrasonic treatment on the mesoporosity of tension wood using nitrogen adsorption-desorption analysis. The results showed that the process of isolating the G-layer using ultrasonication strongly affects its mesoporosity. Most damage was found to occur during sectioning rather than as a result of the 15-min ultrasonic treatment.
ISSN: 0043-7719

hal-01601127
https://hal.archives-ouvertes.fr/hal-01601127
DOI : 10.1007/s00226-017-0890-0
PRODINRA : 401888

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
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 Springer-Verlag Berlin Heidelberg
Résumé : International audience
Reaction wood produces very peculiar maturation stresses at the tree periphery i.e. compressive stress or very high tensile stress, for compression and tension wood respectively, as compared to moderately high tensile stress for normal wood. This means that both its mechanical state and its mechanical and physical properties differ from normal wood. Compression wood shows big differences from normal wood in conifers, for all physical and mechanical properties: higher density and axial crushing strength (MOR) but lower modulus of elasticity (MOE), far higher axial (longitudinal) shrinkage but lower radial and tangential shrinkage, sometimes even lower than the axial shrinkage. For tension wood things are less simple and can vary a lot from hardwood species to species. Globally there are no systematic differences in density and transverse shrinkage, MOE tends to be a little higher while MOR is slightly lower. However, axial shrinkage is much higher for tension wood with a gelatinous layer (G layer) than normal wood due to the specific gel like organization of cellulose microfibrils in the G layer. For tension wood without a G layer(which is rather frequent) axial shrinkage in the first drying operation is around two times higher than in normal wood mainly because of a kind of hygro-thermal release of locked in maturation stress. Overall the very high tensile stress and stored elastic energy in tension wood lead to problemsin wood processing (end splitting and board warping), which is far less the case for compression wood. But due to the large difference in properties relative to normal wood, compression wood occurrence is always a big problem for the in service behaviour of timber, which is seldom the case for tension wood.
The biology of reaction wood

ISBN : 978-3-642-10813-6
hal-01135273
https://hal.archives-ouvertes.fr/hal-01135273

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