• Sciences du Vivant/Sciences agricoles/Sylviculture, foresterie
  
• Sciences de l'ingénieur/Mécanique/Mécanique des matériaux
  
• Physique/Mécanique/Biomécanique
  
• Physique/Mécanique/Mécanique des matériaux
  
• Physique/Physique/Biophysique
  

Abstract Trees can generate large mechanical stresses at the stem periphery to control the orientation of their axes. This key factor in the biomechanical design of trees, named “maturation stress”, occurs in wood fibres during cellular maturation when their secondary cell wall thickens. In this study, the spatial and temporal stiffening kinetics of the different cell wall layers were recorded during fibre maturation on a sample of poplar tension wood using atomic force microscopy. The thickening of the different layers was also recorded. The stiffening of the CML, S 1 and S 2 -layers was initially synchronous with the thickening of the S 2 layer and continued a little after the S 2 -layer reached its final thickness as the G-layer begins to develop. In contrast, the global stiffness of the G-layer, which initially increased with its thickening, was almost stable long before it reached its final maximum thickness. A limited radial gradient of stiffness was observed in the G-layer, but it decreased sharply on the lumen side, where the new sub-layers are deposited during cell wall thickening. Although very similar at the ultrastructural and biochemical levels, the stiffening kinetics of the poplar G-layer appears to be very different from that described in maturing bast fibres. Highlight New insights into the changes in mechanical properties within the cell wall of poplar tension wood fibres during maturation have been obtained using atomic force microscopy.

Trees can modify their wood structure in response to changes in mechanical, hydraulic and storage demands during their life-cycles. Thus, examining radial variations in wood traits is important to expand our knowledge of tree functioning and species ecological strategies. Yet, several aspects of radial changes in wood functional traits are still poorly understood, especially in angiosperm trees from tropical humid forests. Here, we examined radial shifts in wood traits in trunks of tropical forest species and explored their potential ecological implications. We first examined radial variations in wood specific gravity (WSG). Then, we asked what anatomical traits drove radial variations in WSG, and whether WSG, vessel fraction and specific hydraulic conductivity vary independently from each other along the radius gradients. We measured WSG and eight wood anatomical traits, at different radial positions along the trunks, in 19 tree species with contrasting shade-tolerance from a lowland tropical forest in eastern Amazonia. Most species had significant radials shifts in WSG. Positive radial gradients in WSG (i.e., increments from pith to bark) were common among shade-intolerant species and were explained by different combinations of fiber and parenchyma traits, while negative radial shifts in WSG (e.g., decreases towards the bark) were present in shade-tolerants, but were generally weakly related to anatomical traits. We also found that, in general, WSG was unrelated to vessel fraction and specific hydraulic conductivity in any radial position. This study illustrates the contrasting radial variations in wood functional traits that occur in tree species from a humid lowland tropical forest. In particular, our results provide valuable insights into the anatomical traits driving WSG variations during tree development. These insights are important to expand our knowledge on tree ecological strategies by providing evidence on how wood allocation varies as trees grow, which in turn can be useful in studying trait-demography associations, and in estimating tree above-ground biomass.

Abstract The leaf size-stem size spectrum is one of the main dimensions of plant ecological strategies. Yet the anatomical, mechanical, and hydraulic implications of small vs. large shoots are still poorly understood. We investigated 42 tropical rainforest tree species in French Guiana, with a wide range of leaf areas at the shoot level. We quantified the scaling of hydraulic and mechanical constraints with shoot size estimated as the water potential difference ΔΨ and the bending angle ΔΦ, respectively. We investigated how anatomical tissue area, flexural stiffness and xylem vascular architecture affect such scaling by deviating (or not) from theoretical isometry with shoot size variation. Vessel diameter and conductive path length were found to be allometrically related to shoot size, thereby explaining the independence between ΔΨ and shoot size. Leaf mass per area, stem length, and the modulus of elasticity were allometrically related with shoot size, explaining the independence between ΔΦ and shoot size. Our study also shows that the maintenance of both water supply and mechanical stability across the shoot size range are not in conflict.

Recent works revealed that bark is able to produce mechanical stress to control the orientation of young tilted stems. Here we report how the potential performance of this function changes with stem size in six Amazonian species with contrasted bark anatomy. The potential performance of the mechanism depends both on the magnitude of bark stress and the relative thickness of the bark. We measured bark longitudinal residual strain and density, and the allometric relationship between bark thickness and stem radius over a gradient of tree sizes. Constant tensile stress was found in species that rely on bark for the control of stem orientation in young stages. Other species had increasing compres-sive stress, associated with increasing density attributed to the development of sclereids. Compressive stress was also associated with low relative bark thickness. The relative thickness of bark decreased with size in all species, suggesting that a reorientation mechanism based on bark progressively performs less well as the tree grows. However, greater relative thickness was observed in species with more tensile stress, thereby evidencing that this reduction in performance is mitigated in species that rely on bark for reorientation.

Angiosperm trees produce tension wood to actively control their vertical position. Tension wood has often been characterised by the presence of an unlignified inner fibre wall layer called the G-layer. Using this definition, previous reports indicate that only one-third of all tree species have tension wood with G-layers. Here we aim to (i) describe the large diversity of tension wood anatomy in tropical tree species, taking advantage of the recent understanding of tension wood anatomy and (ii) explore any link between this diversity and other ecological traits of the species. We sampled tension wood and normal wood in 432 trees from 242 species in French Guiana. The samples were observed using safranin and astra blue staining combined with optical microscopy. Species were assigned to four anatomical groups depending on the presence/absence of G-layers, and their degree of lignification. The groups were analysed for functional traits including wood density and light preferences. Eighty-six% of the species had G-layers in their tension wood which was lignified in most species, with various patterns of lignification. Only a few species did not have G-layers. We found significantly more species with lignified G-layers among shade-tolerant and shade-demanding species as well as species with a high wood density. Our results bring up-to-date the incidence of species with/without G-layers in the tropical lowland forest where lignified G-layers are the most common anatomy of tension wood. Species without G-layers may share a common mechanism with the bark motor taking over the wood motor. We discuss the functional role of lignin in the G-layer.