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Clair, B., Alteyrac, J., Gronvold, A., Espejo, J., Chanson, B., & Alméras, T. (2013). Patterns of longitudinal and tangential maturation stresses in Eucalyptus nitens plantation trees. Ann. Forest Sci., 70(8), 801–811.
Abstract: 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 the longitudinal direction and compressive along the 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. © 2013 # The Author(s) 2013. This article is published with open access at Springerlink.com.
Keywords: Eucalyptus nitens; G-layer; Longitudinal maturation stress; Maturation strain; Tangential maturation stress; Tension wood
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Clair, B., Arinero, R., Leveque, G., Ramonda, M., & Thibaut, B. (2003). Imaging the mechanical properties of wood cell wall layers by atomic force modulation microscopy. IAWA J., 24(3), 223–230.
Abstract: Atomic Force Microscopy in force modulation mode was used to study the elastic properties of the different fibre wall layers of the tension wood of holm oak and normal wood of boco. The method is based on the measurement of the resonance frequency of the microscope lever in contact with the sample. This frequency is related to the reduced Young modulus E* = E/(1-nu(2)) of the material, supposed to be isotropic. 'Elastic' images of the cell are obtained simultaneously with the topographic images, which allows the observation of the mechanical properties of the cells at a nanometric scale. Layers G, S-1, S-2 and ML can clearly be distinguished. By comparison with known materials an estimation of the absolute modulus is given in the range 5-20 GPa, but should be considered with caution, because the inherent anisotropy of the materials has not been taken into account.
Keywords: wood; cell wall; mechanical properties; elastic modulus; tension wood
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Clair, B., Déjardin, A., Pilate, G., & Alméras, T. (2018). Is the G-layer a tertiary cell wall? Frontiers in Plant Science, 9, 623.
Keywords: Flax; G-layer; Gelatinous layer; Maturation stress; Secondary cell wall; Tension wood; Tertiary cell wall
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Clair, B., Fournier, M., Prevost, M. F., Beauchene, J., & Bardet, S. (2003). Biomechanics of buttressed trees: Bending strains and stresses. Am. J. Bot., 90(9), 1349–1356.
Abstract: 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.
Keywords: biomechanics; buttress; Eleaocarpaceae; French Guiana; Sloanea spp.; tropical trees; wood
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Clair, B., Ghislain, B., Prunier, J., Lehnebach, R., Beauchene, J., & Alméras, T. (2019). Mechanical contribution of secondary phloem to postural control in trees: the bark side of the force. New Phytol, 221(1), 209–217.
Abstract: Summary To grow straight, plants need a motor system that controls posture by generating forces to offset gravity. This motor function in trees was long thought to be only controlled by internal forces induced in wood. Here we provide evidence that bark is involved in the generation of mechanical stresses in several tree species. Saplings of nine tropical species were grown tilted and staked in a shadehouse and the change in curvature of the stem was measured after releasing from the pole and after removing the bark. This first experiment evidenced the contribution of bark in the up-righting movement of tree stems. Combined mechanical measurements of released strains on adult trees and microstructural observations in both transverse and longitudinal/tangential plane enabled us to identify the mechanism responsible for the development of asymmetric mechanical stress in the bark of stems of these species. This mechanism does not result from cell wall maturation like in wood, or from the direct action of turgor pressure like in unlignified organs, but is the consequence of the interaction between wood radial pressure and a smartly organized trellis structure in the inner bark.
Keywords: bark; Malvaceae; maturation stress; secondary phloem; tree biomechanics
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Clair, B., Gril, J., Baba, K., Thibaut, B., & Sugiyama, J. (2005). Precautions for the structural analysis of the gelatinous layer in tension wood. IAWA J., 26(2), 189–195.
Abstract: 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. In order to assess this artefact, the shape and thickness 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 micrometres 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.
Keywords: artefact; fibre wall; gelatinous layer (G-layer); tension wood
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Clair, B., Jaouen, G., Beauchene, J., & Fournier, M. (2003). Mapping radial, tangential and longitudinal shrinkages and relation to tension wood in discs of the tropical tree Symphonia globulifera. Holzforschung, 57(6), 665–671.
Abstract: A method for measuring shrinkage resulting from drying in the three anisotropic directions is developed and tested. Measurements are performed on sawn discs, a technique which simplifies preparation and enables large numbers of measurements. Shrinkage values can be represented as a map of the disc surface. The results indicate that comparisons between shrinkage distribution and tension wood distribution on the discs show a clear relationship and can be measured with relatively high accuracy in reference to the shrinkage map. In the long term, this method could be useful in the timber industry as a means for choosing the direction in which logs are cut depending on their type of wood composition.
Keywords: drying shrinkage; tension wood; Symphonia globulifera L. f.
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Clair, B., Ruelle, J., Beauchene, J., Prevost, M. F., & Fournier, M. (2006). Tension wood and opposite wood in 21 tropical rain forest species 1. Occurrence and efficiency of the G-layer. IAWA J., 27(3), 329–338.
Abstract: 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 the G-layer indicates that the G-layer is not a key factor in the production of high tensile stressed wood.
Keywords: gelatinous layer; G-layer; French Guyana; tropical rain forest; tension wood; wood anatomy
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Clair, B., Ruelle, J., & Thibaut, B. (2003). Relationship between growth stress, mechanical-physical properties and proportion of fibre with gelatinous layer in chestnut (Castanea sativa Mill.). Holzforschung, 57(2), 189–195.
Abstract: A range of mechanical and physical properties were determined for 96 specimens of chestnut wood and for wood types ranging from compression to tension wood; tests included (1) growth stress, (2) longitudinal Young's modulus in green and air-dried states (3) shrinkage in longitudinal and tangential directions. Anatomical observations permitted determination of the proportion of fibres with a gelatinous layer. The influence of these atypical fibres on macroscopic wood properties is examined and discussed. A basic model is proposed to determine their properties in theoretically isolated conditions.
Keywords: growth stress; longitudinal Young's modulus; shrinkage; normal wood; tension wood; gelatinous layer; Castanea Sativa
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Cochard, H., Coste, S., Chanson, B., Guehl, J. M., & Nicolini, E. (2005). Hydraulic architecture correlates with bud organogenesis and primary shoot growth in beech (Fagus sylvatica). Tree Physiol., 25(12), 1545–1552.
Abstract: In beech (Fagus sylvatica L.), the number of leaf primordia preformed in the buds determines the length and the type (long versus short) of annual growth units, and thus, branch growth and architecture. We analyzed the correlation between the number of leaf primordia and the hydraulic conductance of the vascular system connected to the buds. Terminal buds of short growth units and axillary buds of long growth units on lower branches of mature trees were examined. Buds with less than four and more than five leaf primordia formed short and long growth units, respectively. Irrespective of the type of growth unit the bud was formed on, the occurrence of a large number of leaf primordia was associated with high xylem hydraulic conductance. Xylem conductance was correlated to the area of the outermost annual ring. These results suggest that organogenesis and primary growth in buds correlates with secondary growth of the growth units and thus with their hydraulic architecture. Possible causal relationships between the variables are discussed.
Keywords: development; hydraulic conductance; leaf primordia; meristem; xylem
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