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Almeras, T., Thibaut, A., & Gril, J. (2005). Effect of circumferential heterogeneity of wood maturation strain, modulus of elasticity and radial growth on the regulation of stem orientation in trees. Trees-Struct. Funct., 19(4), 457–467.
Abstract: Active mechanisms of re-orientation are necessary to maintain the verticality of tree stems. They are achieved through the production of reaction wood, associated with circumferential variations of three factors related to cambial activity: maturation strain, longitudinal modulus of elasticity (MOE) and eccentric growth. These factors were measured on 17 mature trees from different botanical families and geographical locations. Various patterns of circumferential variation of these factors were identified. A biomechanical analysis based on beam theory was performed to quantify the individual impact of each factor. The main factor of re-orientation is the circumferential variation of maturation strains. However, this factor alone explains only 57% of the re-orientations. Other factors also have an effect through their interaction with maturation strains. Eccentric growth is generally associated with heterogeneity of maturation strains, and has an important complementary role, by increasing the width of wood with high maturation strain. Without this factor, the efficiency of re-orientations would be reduced by 31% for angiosperms and 26% for gymnosperms. In the case of angiosperms, MOE is often larger in tension wood than in normal wood. Without these variations, the efficiency of re-orientations would be reduced by 13%. In the case of gymnosperm trees, MOE of compression wood is lower than that of normal wood, so that re-orientation efficiency would be increased by 24% without this factor of variations.
Keywords: biomechanics; reaction wood; maturation strain; Young's modulus; eccentricity
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Almeras, T., Derycke, M., Jaouen, G., Beauchene, J., & Fournier, M. (2009). Functional diversity in gravitropic reaction among tropical seedlings in relation to ecological and developmental traits. J. Exp. Bot., 60(15), 4397–4410.
Abstract: Gravitropism is necessary for plants to control the orientation of their axes while they grow in height. In woody plants, stem re-orientations are costly because they are achieved through diameter growth. The functional diversity of gravitropism was studied to check if the mechanisms involved and their efficiency may contribute to the differentiation of height growth strategies between forest tree species at the seedling stage. Seedlings of eight tropical species were grown tilted in a greenhouse, and their up-righting movement and diameter growth were measured over three months. Morphological, anatomical, and biomechanical traits were measured at the end of the survey. Curvature analysis was used to analyse the up-righting response along the stems. Variations in stem curvature depend on diameter growth, size effects, the increase in self-weight, and the efficiency of the gravitropic reaction. A biomechanical model was used to separate these contributions. Results showed that (i) gravitropic movements were based on a common mechanism associated to similar dynamic patterns, (ii) clear differences in efficiency (defined as the change in curvature achieved during an elementary diameter increment for a given stem diameter) existed between species, (iii) the equilibrium angle of the stem and the anatomical characters associated with the efficiency of the reaction also differed between species, and (iv) the differences in gravitropic reaction were related to the light requirements: heliophilic species, compared to more shade-tolerant species, had a larger efficiency and an equilibrium angle closer to vertical. This suggests that traits determining the gravitropic reaction are related to the strategy of light interception and may contribute to the differentiation of ecological strategies promoting the maintenance of biodiversity in tropical rainforests.
Keywords: Biomechanics; French Guiana; functional diversity; gravitropism; reaction wood; tropical rainforest
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Fournier, M., Dlouhá, J., Jaouen, G., & Almeras, T. (2013). Integrative biomechanics for tree ecology: Beyond wood density and strength. J. Exp. Bot., 64(15), 4793–4815.
Abstract: Functional ecology has long considered the support function as important, but its biomechanical complexity is only just being elucidated. We show here that it can be described on the basis of four biomechanical traits, two safety traits against winds and self-buckling, and two motricity traits involved in sustaining an upright position, tropic motion velocity (MV) and posture control (PC). All these traits are integrated at the tree scale, combining tree size and shape together with wood properties. The assumption of trait constancy has been used to derive allometric scaling laws, but it was more recently found that observing their variations among environments and functional groups, or during ontogeny, provides more insights into adaptive syndromes of tree shape and wood properties. However, oversimpli-fed expressions have often been used, possibly concealing key adaptive drivers. An extreme case of oversimplification is the use of wood basic density as a proxy for safety. Actually, as wood density is involved in stiffiness, loads, and construction costs, the impact of its variations on safety is non-trivial. Moreover, other wood features, especially the microfibril angle (MFA), are also involved. Furthermore, wood is not only stiff and strong, but it also acts as a motor for MV and PC. The relevant wood trait for this is maturation strain asymmetry. Maturation strains vary with cell-wall characteristics such as MFA, rather than with wood density. Finally, the need for further studies about the ecological relevance of branching patterns, motricity traits, and growth responses to mechanical loads is discussed. © The Author 2013.
Keywords: Biomechanics; Ecological strategy; Gravitropism; Shape; Size; Trees; Wood
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Jaouen, G., Almeras, T., Coutand, C., & Fournier, M. (2007). How to determine sapling buckling risk with only a few measurements. Am. J. Bot., 94(10), 1583–1593.
Abstract: Tree buckling risk (actual height/critical buckling height) is an important biomechanical trait of plant growth strategies, and one that contributes to species coexistence. To estimate the diversity of this trait among wide samples, a method that minimizes damage to the plants is necessary. On the basis of the rarely used, complete version of Greenhill's model (1881, Proceedings of the Cambridge Philosophical Society 4(2): 65-73), we precisely measured all the necessary parameters on a sample of 236 saplings of 16 species. Then, using sensitivity (variance) analysis, regressions between successive models for risk factors and species ranks and the use of these models on samples of self- and nonself-supporting saplings, we tested different degrees of simplification up to the most simple and widely used formula that assumes that the tree is a cylindrical homogeneous pole. The size factor had the greatest effect on buckling risk, followed by the form factor and the modulus of elasticity of the wood. Therefore, estimates of buckling risk must consider not only the wood properties but especially the form factor. Finally, we proposed a simple but accurate method of assessing tree buckling risk that is applicable to a wide range of samples and that requires mostly nondestructive measurements.
Keywords: biomechanics; critical buckling height; French Guiana; risk factor; sapling; stem form; tropical rain forest; trunk volume
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Almeras, T., & Gril, J. (2007). Mechanical analysis of the strains generated by water tension in plant stems. Part 1: stress transmission from the water to the cell walls. Tree Physiol., 27(11), 1505–1516.
Abstract: Plant tissues shrink and swell in response to changes in water pressure. These strains can be easily measured, e.g., at the surface of tree stems, to obtain indirect information about plant water status and other physiological parameters. We developed a mechanical model to clarify how water pressure is transmitted to cell walls and causes shrinkage of plant tissues, particularly in the case of thick-walled cells such as wood fibers. Our analysis shows that the stress inside the fiber cell walls is lower than the water tension. The difference is accounted for by a stress transmission factor that depends on two main effects. The first effect is the dilution of the stress through the cell wall, because water acts at the lumen border and is transmitted to the cuter border of the cell, which has a larger circumference. The second effect is the partial conversion of radial stress into tangential stress. Both effects are quantified as functions of parameters of the cell wall structure and its mechanical properties.
Keywords: biomechanics; cell mechanics; diurnal strains; mechanical model; multilayer cylinder; stress transtnissionjactor
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Almeras, T. (2008). Mechanical analysis of the strains generated by water tension in plant stems. Part II: strains in wood and bark and apparent compliance. Tree Physiol., 28(10), 1513–1523.
Abstract: Tree steins shrink in diameter during the day and swell during the night in response to changes in water tension in the xylem. Stein shrinkage can easily be measured in a nondestructive way, to derive continuous information about tree water status. The relationship between the strain and the change in water tension can be evaluated by empirical calibrations, or can be related to the structure of the plant. A mechanical analysis was performed to make this relationship explicit. The stem is modeled as a cylinder made of multiple layers of tissues, including heartwood, sapwood, and inner and outer bark. The effect of changes in water tension on the apparent strain at the surface of a tissue is quantified as a function of parameters defining stem anatomy and the mechanical properties of the tissues. Various possible applications in the context of tree physiology are suggested.
Keywords: biomechanics; calibration; diurnal strains; mechanical model; multilayer cylinder; water potential
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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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Thomas, H. J. D., Bjorkman, A. D., Myers-Smith, I. H., Elmendorf, S. C., Kattge, J., Diaz, S., et al. (2020). Global plant trait relationships extend to the climatic extremes of the tundra biome. Nat. Commun., 11(1351).
Abstract: The majority of variation in six traits critical to the growth, survival and reproduction of plant species is thought to be organised along just two dimensions, corresponding to strategies of plant size and resource acquisition. However, it is unknown whether global plant trait relationships extend to climatic extremes, and if these interspecific relationships are confounded by trait variation within species. We test whether trait relationships extend to the cold extremes of life on Earth using the largest database of tundra plant traits yet compiled. We show that tundra plants demonstrate remarkably similar resource economic traits, but not size traits, compared to global distributions, and exhibit the same two dimensions of trait variation. Three quarters of trait variation occurs among species, mirroring global estimates of interspecific trait variation. Plant trait relationships are thus generalizable to the edge of global trait-space, informing prediction of plant community change in a warming world. © 2020, Crown.
Keywords: biome; climate change; extreme event; global change; growth; interspecific interaction; plant community; tundra; article; plant community; prediction; tundra; warming; classification; climate; ecosystem; genetics; plant; plant development; Climate; Ecosystem; Plant Development; Plants; Tundra
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Schwalm, C. R., Williams, C. A., Schaefer, K., Arneth, A., Bonal, D., Buchmann, N., et al. (2010). Assimilation exceeds respiration sensitivity to drought: A FLUXNET synthesis. Glob. Change Biol., 16(2), 657–670.
Abstract: The intensification of the hydrological cycle, with an observed and modeled increase in drought incidence and severity, underscores the need to quantify drought effects on carbon cycling and the terrestrial sink. FLUXNET, a global network of eddy covariance towers, provides dense data streams of meteorological data, and through flux partitioning and gap filling algorithms, estimates of net ecosystem productivity (F-NEP), gross ecosystem productivity (P), and ecosystem respiration (R). We analyzed the functional relationship of these three carbon fluxes relative to evaporative fraction (EF), an index of drought and site water status, using monthly data records from 238 micrometeorological tower sites distributed globally across 11 biomes. The analysis was based on relative anomalies of both EF and carbon fluxes and focused on drought episodes by biome and climatic season. Globally P was approximate to 50% more sensitive to a drought event than R. Network-wide drought-induced decreases in carbon flux averaged -16.6 and -9.3 g C m-2 month-1 for P and R, i.e., drought events induced a net decline in the terrestrial sink. However, in evergreen forests and wetlands drought was coincident with an increase in P or R during parts of the growing season. The most robust relationships between carbon flux and EF occurred during climatic spring for F-NEP and in climatic summer for P and R. Upscaling flux sensitivities to a global map showed that spatial patterns for all three carbon fluxes were linked to the distribution of croplands. Agricultural areas exhibited the highest sensitivity whereas the tropical region had minimal sensitivity to drought. Combining gridded flux sensitivities with their uncertainties and the spatial grid of FLUXNET revealed that a more robust quantification of carbon flux response to drought requires additional towers in all biomes of Africa and Asia as well as in the cropland, shrubland, savannah, and wetland biomes globally.
Keywords: biome; carbon cycling; drought; eddy covariance; evaporative fraction; FLUXNET; synthesis
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Grau, O., Peñuelas, J., Ferry, B., Freycon, V., Blanc, L., Desprez, M., et al. (2017). Nutrient-cycling mechanisms other than the direct absorption from soil may control forest structure and dynamics in poor Amazonian soils. Sci. Rep., 7, 45017.
Abstract: Tropical forests store large amounts of biomass despite they generally grow in nutrient-poor soils, suggesting that the role of soil characteristics in the structure and dynamics of tropical forests is complex. We used data for >34 000 trees from several permanent plots in French Guiana to investigate if soil characteristics could predict the structure (tree diameter, density and aboveground biomass), and dynamics (growth, mortality, aboveground wood productivity) of nutrient-poor tropical forests. Most variables did not covary with site-level changes in soil nutrient content, indicating that nutrient-cycling mechanisms other than the direct absorption from soil (e.g. the nutrient uptake from litter, the resorption, or the storage of nutrients in the biomass), may strongly control forest structure and dynamics. Ecosystem-level adaptations to low soil nutrient availability and long-term low levels of disturbance may help to account for the lower productivity and higher accumulation of biomass in nutrient-poor forests compared to nutrient-richer forests.
Keywords: biomass; forest structure; French Guiana; mortality; nutrient availability; nutrient content; nutrient cycling; nutrient uptake; productivity; soil; storage; tropical rain forest
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