Section: Forest & Wood Sciences
Topic:
Plant biology,
Agricultural sciences,
Physiology
Tree growth forces and wood properties
Corresponding author(s): Thibaut, Bernard (bernard.thibaut@umontpellier.fr); Gril, Joseph (joseph.gril@cnrs.fr)
10.24072/pcjournal.48 - Peer Community Journal, Volume 1 (2021), article no. e46.
Get full text PDF Peer reviewed and recommended by PCILiving wood in the tree performs a muscular action by generating forces at the sapwood periphery and residual strains in dead sapwood fibres. Dissymmetric force generation around the tree trunk is the motor system allowing movement, posture control and tree reshaping after accidents. Rather young trees are able to restore the verticality of their trunk after accidental rotation of the soil-root system due to wind or landslide, leading to typically curved stems shape. The very high dissymmetry of forces for the motor action is associated with the occurrence of reaction wood on one side of the inclined stem during many successive years. A method to reconstitute this biomechanical history from observations after tree felling, on either green or dry wood, is discussed. A selection of 17 trees from 15 different species (13 different families), tropical and temperate, hardwoods and softwoods, were selected and peripheral residual strains were measured in situ before felling, on 8 positions for each stem. Matched wooden rods were sawn and measured for their mechanical and physical properties in the green and dry states, allowing the estimation of tree growth stress, i.e., the force created by the living wood. It was possible to build easy-to-use conversion coefficients between the growth stress indicator (GSI), measured in situ by the single hole method, and growth strain and growth stress with the knowledge of basic density and green longitudinal elastic modulus. Maturation strain, specific modulus (as a proxy of micro-fibril angle) and longitudinal shrinkage are properties independent from basic density, whose variation among species was very large. For the whole range of compression wood, normal wood and tension wood, strong relationships between these 3 properties were observed, but together no single model, based on cell-wall microfibril angle only, could be defined. Growth forces are the product of 4 parameters: ring width, basic density, basic specific modulus and maturation strain, all of them being the result of wood formation. Thanks to the wide range of wood types and species, simple and highly significant formulas were obtained for the relationship between basic and dry density, green and dry longitudinal modulus of elasticity, basic and dry specific modulus. To estimate ring width in the green state from values in dry state, radial shrinkage needs to be measured afterwards on dry specimens. Maturation strains is less accurately linked to late measurements on dry wood, but longitudinal shrinkage offers a rather good solution for an estimation provided that the wood type (softwood, hardwood with-G layer, hardwood without G-Layer) is known.
Type: Research article
Thibaut, Bernard 1; Gril, Joseph 2, 3
@article{10_24072_pcjournal_48, author = {Thibaut, Bernard and Gril, Joseph}, title = {Tree growth forces and wood properties}, journal = {Peer Community Journal}, eid = {e46}, publisher = {Peer Community In}, volume = {1}, year = {2021}, doi = {10.24072/pcjournal.48}, url = {https://peercommunityjournal.org/articles/10.24072/pcjournal.48/} }
Thibaut, Bernard; Gril, Joseph. Tree growth forces and wood properties. Peer Community Journal, Volume 1 (2021), article no. e46. doi : 10.24072/pcjournal.48. https://peercommunityjournal.org/articles/10.24072/pcjournal.48/
PCI peer reviews and recommendation, and links to data, scripts, code and supplementary information: 10.24072/pci.forestwoodsci.100006
Conflict of interest of the recommender and peer reviewers:
The recommender in charge of the evaluation of the article and the reviewers declared that they have no conflict of interest (as defined in the code of conduct of PCI) with the authors or with the content of the article.
[1] Modelling anisotropic maturation strains in wood in relation to fibre boundary conditions, microstructure and maturation kinetics, Holzforschung, Volume 59 (2005) no. 3, pp. 347-353 | DOI
[2] Effect of circumferential heterogeneity of wood maturation strain, modulus of elasticity and radial growth on the regulation of stem orientation in trees, Trees, Volume 19 (2005) no. 4, pp. 457-467 | DOI
[3] Functional diversity in gravitropic reaction among tropical seedlings in relation to ecological and developmental traits, Journal of Experimental Botany, Volume 60 (2009) no. 15, pp. 4397-4410 | DOI
[4] Critical review on the mechanisms of maturation stress generation in trees, Journal of The Royal Society Interface, Volume 13 (2016) no. 122 | DOI
[5] Modelling, Evaluation and Biomechanical Consequences of Growth Stress Profiles Inside Tree Stems, Plant Biomechanics, Springer International Publishing, Cham, 2018, pp. 21-48 | DOI
[6] Quantifying the motor power of trees, Trees, Volume 32 (2018) no. 3, pp. 689-702 | DOI
[7] Application of a new method for the growth stress measurement for Pinus Caribea. IUFRO P5-01, Properties and utilisation of tropical woods, Manaus, Brasil, 19-23/11/1984., 1994
[8] Growth Stresses and Strains in Trees, Springer Series in Wood Science, Springer Berlin Heidelberg, Berlin, Heidelberg, 1986 | DOI
[9] Structure, composition chimique et retraits de maturation du bois chez les clones d'Eucalyptus, Annales des Sciences Forestières, Volume 52 (1995) no. 2, pp. 157-172 | DOI
[10] Précontraintes de croissance et propriétés mécano-physiques de clones d’Eucalyptus (Pointe Noire, Congo) : hétérogénéités, corrélations et interprétations histologiques. Thèse en sciences du bois, Université Bordeaux 1 (In French). https://www.theses.fr/1994BOR10521 (1994)
[11] A general theory for the origin of growth stresses in reaction wood: how trees stay upright, IAWA Journal, Volume 22 (2001) no. 3, pp. 205-212 | DOI
[12] Plant Architecture: A Dynamic, Multilevel and Comprehensive Approach to Plant Form, Structure and Ontogeny, Annals of Botany, Volume 99 (2007) no. 3, pp. 375-407 | DOI
[13] Occurrence and relevance of growth stresses in Beech (Fagus sylvatica L.) in Central Europe, Final Report of FAIR-project CT 98-3606, Coordinator G. Becker, Institut für Forstbenutzung und forstliche Arbeitwissenschaft, Albert-Ludwigs Universität, Freiburg, Germany, 323 p., 2001
[14] Module dynamique et frottement intérieur dans le bois : mesures sur poutres flottantes en vibrations naturelles. Thèse de Doctorat en Sciences du Bois, Institut National Polytechnique de Lorraine, Nancy. https://www.theses.fr/1989NAN10387, 1989
[15] Prediction of Coefficients of Thermal Expansion for Unidirectional Composites, Journal of Composite Materials, Volume 23 (1988) no. 4, pp. 370-388 | DOI
[16] Tree growth stresses ? Part V: Evidence of an origin in differentiation and lignification, Wood Science and Technology, Volume 6 (1972) no. 4, pp. 251-262 | DOI
[17] Natural vibration analysis of clear wooden beams: a theoretical review, Wood Science and Technology, Volume 36 (2002) no. 4, pp. 347-365 | DOI
[18] Changes in viscoelastic vibrational properties between compression and normal wood: roles of microfibril angle and of lignin, Holzforschung, Volume 67 (2013) no. 1, pp. 75-85 | DOI
[19] Cellulose microfibril angles and cell-wall polymers in different wood types of Pinus radiata, Cellulose, Volume 19 (2012) no. 4, pp. 1385-1404 | DOI
[20] A theory of the shrinkage of wood, Wood Science and Technology, Volume 6 (1972) no. 4, pp. 284-292 | DOI
[21] The anisotropic elasticity of the plant cell wall, Wood Science and Technology, Volume 2 (1968) no. 4, pp. 268-278 | DOI
[22] Mesoporosity as a new parameter for understanding tension stress generation in trees, Journal of Experimental Botany, Volume 60 (2009) no. 11, pp. 3023-3030 | DOI
[23] Mesoporosity changes from cambium to mature tension wood: a new step toward the understanding of maturation stress generation in trees, New Phytologist, Volume 205 (2015) no. 3, pp. 1277-1287 | DOI
[24] Tension Wood and Opposite Wood in 21 Tropical Rain Forest Species, IAWA Journal, Volume 27 (2006) no. 3, pp. 329-338 | DOI
[25] Patterns of longitudinal and tangential maturation stresses in Eucalyptus nitens plantation trees, Annals of Forest Science, Volume 70 (2013) no. 8, pp. 801-811 | DOI
[26] Physical and Mechanical Properties of Reaction Wood, The Biology of Reaction Wood, Springer Berlin Heidelberg, Berlin, Heidelberg, 2014, pp. 171-200 | DOI
[27] The Gravitropic Response of Poplar Trunks: Key Roles of Prestressed Wood Regulation and the Relative Kinetics of Cambial Growth versus Wood Maturation, Plant Physiology, Volume 144 (2007) no. 2, pp. 1166-1180 | DOI
[28] Elasticity and microfibrillar angle in the wood of Sitka spruce, Proceedings of the Royal Society of London. Series B. Biological Sciences, Volume 166 (1966) no. 1004, pp. 245-272 | DOI
[29] Cell Wall Polymers in Reaction Wood, The Biology of Reaction Wood, Springer Berlin Heidelberg, Berlin, Heidelberg, 2014, pp. 37-106 | DOI
[30] Plant anatomy. Fourth edition, Pergamon press., 1990
[31] Growth Stresses are Highly Controlled by the Amount of G-Layer in Poplar Tension Wood, IAWA Journal, Volume 29 (2008) no. 3, pp. 237-246 | DOI
[32] Mesures des déformations résiduelles de croissance à la surface des arbres, en relation avec leur morphologie. Observations sur différentes espèces, Annales des Sciences Forestières, Volume 51 (1994) no. 3, pp. 249-266 | DOI
[33] Mécanique de l'arbre sur pied : modélisation d'une structure en croissance soumise à des chargements permanents et évolutifs. 1. Analyse des contraintes de support, Annales des Sciences Forestières, Volume 48 (1991) no. 5, pp. 513-525 | DOI
[34] Mécanique de l'arbre sur pied : modélisation d'une structure en croissance soumise à des chargements permanents et évolutifs. 2. Analyse tridimensionnelle des contraintes de maturation, cas du feuillu standard, Annales des Sciences Forestières, Volume 48 (1991) no. 5, pp. 527-546 | DOI
[35] Integrative biomechanics for tree ecology: beyond wood density and strength, Journal of Experimental Botany, Volume 64 (2013) no. 15, pp. 4793-4815 | DOI
[36] Biomechanical Action and Biological Functions, The Biology of Reaction Wood, Springer Berlin Heidelberg, Berlin, Heidelberg, 2014, pp. 139-169 | DOI
[37] The Biology of Reaction Wood, Springer Series in Wood Science, Springer Berlin Heidelberg, Berlin, Heidelberg, 2014 | DOI
[38] Diversity in the organisation and lignification of tension wood fibre walls – A review, IAWA Journal, Volume 38 (2017) no. 2, pp. 245-265 | DOI
[39] Diversity of anatomical structure of tension wood among 242 tropical tree species, IAWA Journal, Volume 40 (2019) no. 4, pp. 765-784 | DOI
[40] Structures or Why things don’t fall down, Springer US, Boston, MA, 1978 | DOI
[41] Plant ‘muscles’: fibers with a tertiary cell wall, New Phytologist, Volume 218 (2018) no. 1, pp. 66-72 | DOI
[42] Tree growth stress and related problems, Journal of Wood Science, Volume 63 (2017) no. 5, pp. 411-432 | DOI
[43] Growth stress distribution in leaning trunks of Cryptomeria japonica, Tree Physiology, Volume 21 (2001) no. 4, pp. 261-266 | DOI
[44] Relationship between tree morphology and growth stress in mature European beech stands, Annals of Forest Science, Volume 70 (2013) no. 2, pp. 133-142 | DOI
[45] Thermal expansion behaviour of unidirectional carbon-fibre-reinforced copper-matrix composites, Composites Part A: Applied Science and Manufacturing, Volume 29 (1998) no. 12, pp. 1563-1567 | DOI
[46] Strength and related properties of woods grown in the United States. USDA technical bulletin N° 479, Washington D.C., 1935
[47] Effects of stand density and seedlot on three wood properties of young radiata pine grown at a dry-land site in New Zealand, New Zealand Journal of Forestry Science, Volume 45 (2015) no. 1 | DOI
[48] The power and control of gravitropic movements in plants: a biomechanical and systems biology view, Journal of Experimental Botany, Volume 60 (2009) no. 2, pp. 461-486 | DOI
[49] Posture control in land plants: growth, position sensing, proprioception, balance, and elasticity, Journal of Experimental Botany, Volume 70 (2019) no. 14, pp. 3467-3494 | DOI
[50] Quantitative chemical indicators to assess the gradation of compression wood, Holzforschung, Volume 63 (2009) no. 4 | DOI
[51] Growth stresses in tension wood: role of microfibrils and lignification, Annales des Sciences Forestières, Volume 51 (1994) no. 3, pp. 291-300 | DOI
[52] Comparative anatomy normal wood / reaction wood and observation of structure / properties relationships (in French). MS Thesis, Université de Nancy., 2003
[53] Growth stresses and cellulose structural parameters in tension and normal wood from three tropical rainforest angiosperm species, BioResources, Volume 2 (2007) no. 2, pp. 235-251 | DOI
[54] Variations in physical and mechanical properties between tension and opposite wood from three tropical rainforest species, Wood Science and Technology, Volume 45 (2011) no. 2, pp. 339-357 | DOI
[55] Morphology, Anatomy and Ultrastructure of Reaction Wood, The Biology of Reaction Wood, Springer Berlin Heidelberg, Berlin, Heidelberg, 2014, pp. 13-35 | DOI
[56] The evolution process of the growth stress in the tree. The surface stress on the tree, Mokuzai Gakkaishi (Journal of the Japanese Wood Research Association), Volume 29 (1978)
[57] Measuring microfibrillar angles using light microscopy, Wood and fiber science, Volume 17 (1985)
[58] Assessment of Fiber Orientation on the Mechanical Properties of PA6/Cellulose Composite, Applied Sciences, Volume 10 (2020) no. 16 | DOI
[59] Generation process of growth stresses in cell walls: Relation between longitudinal released strain and chemical composition, Wood Science and Technology, Volume 27 (1993) no. 4 | DOI
[60] Valorisation du bois de châtaignier : prévoir et réduire les risques de roulure à la production, Rapport final contrat “Agriculture demain” MRT 91.G.0491, 12.95. (In French), 1995
[61] Three-dimensional printing, muscles, and skeleton: mechanical functions of living wood, Journal of Experimental Botany, Volume 70 (2019) no. 14, pp. 3453-3466 | DOI
[62] Compression Wood in Gymnosperms, Springer Berlin Heidelberg, Berlin, Heidelberg, 1986 | DOI
[63] The Molecular Mechanisms of Reaction Wood Induction, The Biology of Reaction Wood, Springer Berlin Heidelberg, Berlin, Heidelberg, 2014, pp. 107-138 | DOI
[64] Relations entre Contraintes de Croissance Longitudinales et Bois de Tension, dans le Hêtre(Fagus sylvaticaL.), Holzforschung, Volume 29 (1975) no. 6, pp. 217-223 | DOI
[65] New formula and conversion factor to compute tree species basic wood density from a global wood technology database, American Journal of Botany, Volume 105 | DOI
[66] Influence of tree morphology, genetics, and initial stand density on outerwood modulus of elasticity of 17-year-old Pinus radiata, Forest Ecology and Management, Volume 244 (2007) no. 1-3, pp. 86-92 | DOI
[67] Shrinkage and elasticity of normal and compression wood in conifers, Mokuzai Gakkaishi (Journal of the Japanese Wood Research Association), Volume 42 (1996)
[68] Modelling the influence of environment and stand characteristics on basic density and modulus of elasticity for young Pinus radiata and Cupressus lusitanica, Forest Ecology and Management, Volume 255 (2008) no. 3-4, pp. 1023-1033 | DOI
[69] Growth stress generation and microfibril angle in reaction wood In: Microfibril angle in wood. Butterfield BG (ed) (1998), pp. 225-239
[70] A model of anisotropic swelling and shrinking process of wood, Wood Science and Technology, Volume 35 (2001) no. 1-2, pp. 167-181 | DOI
[71] Errata: Wood Science and Technology 35: 167-181 and Wood Science and Technology 35: 269-282, Wood Science and Technology, Volume 35 (2001) no. 4, p. 377-377 | DOI
[72] Measurement methods for longitudinal surface strain in trees: a review, Australian Forestry, Volume 68 (2005) no. 1, pp. 34-43 | DOI
[73] Tension wood and growth stress induced by artificial inclination in textitLiriodendron tulipifera Linn. and Prunus spachiana Kitamura f. ascendens Kitamura, Annals of Forest Science, Volume 57 (2000) no. 8, pp. 739-746 | DOI
[74] Techniques for Measuring Growth Stress on the Xylem Surface Using Strain and Dial Gauges, Holzforschung, Volume 56 (2002) no. 5, pp. 461-467 | DOI
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