Tancrède Alméras

Integrative biomechanics for tree ecology: beyond wood density and strength

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, oversimplified 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 stiffness, 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.

Critical review on the mechanisms of maturation stress generation in trees.

Trees control their posture by generating asymmetric mechanical stress around the periphery of the trunk or branches. This stress is produced in wood during the maturation of the cell wall. When the need for reaction is high, it is accompanied by strong changes in cell organization and composition called reaction wood, namely compression wood in gymnosperms and tension wood in angiosperms. The process by which stress is generated in the cell wall during its formation is not yet known, and various hypothetical mechanisms have been proposed in the literature. Here we aim at discriminating between these models. First, we summarize current knowledge about reaction wood structure, state and behaviour relevant to the understanding of maturation stress generation. Then, the mechanisms proposed in the literature are listed and discussed in order to identify which can be rejected based on their inconsistency with current knowledge at the frontier between plant science and mechanical engineering.

Does spatial distribution of tree size account for spatial variation in soil respiration in a tropical forest

We explored the relationship between soil processes, estimated through soil respiration (Rsoil), and the spatial variation in forest structure, assessed through the distribution of tree size, in order to understand the determinism of spatial variations in Rsoil in a tropical forest. The influence of tree size was examined using an index (Ic) calculated for each tree as a function of (1) the trunk cross section area and (2) the distance from the measurement point. We investigated the relationships between Ic and litterfall, root mass and Rsoil, respectively. Strong significant relationships were found between Ic and both litterfall and root mass. Rsoil showed a large range of variations over the 1-ha experimental plot, from 1.5 to 12.6xa0gC m−2 d−1. The best relationship between Ic and Rsoil only explained 17% of the spatial variation in Rsoil. These results support the assumption that local spatial patterns in litter production and root mass depend on tree distribution in tropical forests. Our study also emphasizes the modest contribution of tree size distribution–which is mainly influenced by the presence of the biggest trees (among the large range size of the inventoried trees greater than 10xa0cm diameter at 1.30xa0m above ground level or at 0.5xa0m above the buttresses)–in explaining spatial variations in Rsoil.

Mesoporosity changes from cambium to mature tension wood: a new step toward the understanding of maturation stress generation in trees

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 × 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 during the 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.

Patterns of longitudinal and tangential maturation stresses in Eucalyptus nitens plantation trees

ContextTree 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.AimsWe aimed at evaluating patterns of maturation stress on eucalypt plantation trees and their relation with growth, with a focus on tangential stress evaluation.MethodsReleased maturation strains along longitudinal and tangential directions were measured around the circumference of 29 Eucalyptus nitens trees, including both straight and leaning trees.ResultsMost 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.ConclusionThe 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.

Representativeness of wood biomechanical properties measured after storage in different conditions

Obtaining representative values of green wood properties is essential for studies investigating the biomechanical aspects of tree development and ecology. Here, we compare the biomechanical properties of wood stored in various conditions between their collection in the field and their measurement. The study was performed on a large sample of wood specimens from different tropical species and different location in the trees, representing a wide diversity in wood structures. Elastic and viscoelastic properties are measured on green wood, and measured again after storage in different conditions: immersion in cold water during various durations, storage in an ethanol solution with or without washing in water, and air drying with or without rehydration. The systematic and random errors induced by these storage methods are quantified. Storage in cold water is the best way to preserve wood native properties. Soaking in ethanol is a fair alternative regarding elastic properties, but induces a significant change in viscoelastic properties. Air drying causes important, and partly irreversible, changes in mechanical properties. However, regarding elastic properties, this change is a systematic bias so that the air-dried elastic modulus provides a good basis for comparative studies of green wood stiffness.

Is the G-Layer a Tertiary Cell Wall?

HAL is a multi-disciplinary open access archive for the deposit and dissemination of scientific research documents, whether they are published or not. The documents may come from teaching and research institutions in France or abroad, or from public or private research centers. L’archive ouverte pluridisciplinaire HAL, est destinée au dépôt et à la diffusion de documents scientifiques de niveau recherche, publiés ou non, émanant des établissements d’enseignement et de recherche français ou étrangers, des laboratoires publics ou privés. Is the G-Layer a Tertiary Cell Wall? Bruno Clair, Annabelle Dejardin, Gilles Pilate, Tancrède Alméras

Quantifying the motor power of trees

Key MessageWood maturation strains can be estimated from the change in curvature that occurs when a stem grown staked in tilted position is released from the stake.AbstractTrees have a motor system to enable upright growth in the field of gravity. This motor function is taken on by reaction wood, a special kind of wood that typically develops in leaning axes and generates mechanical force during its formation, curving up the stem and counteracting the effect of gravity or other mechanical disturbances. Quantifying the mechanical stress induced in wood during maturation is essential to many areas of research ranging from tree architecture to functional genomics. Here, we present a new method for quantifying wood maturation stress. It consists of tilting a tree, tying it to a stake, letting it grow in tilted position, and recording the change in stem curvature that occurs when the stem is released from the stake. A mechanical model is developed to make explicit the link between the change in curvature, maturation strain and morphological traits of the stem section. A parametric study is conducted to analyse how different parameters influence the change in curvature. This method is applied to the estimation of maturation strain in two different datasets. Results show that the method is able to detect genotypic variations in motor power expression. As predicted by the model, we observe that the change in stem curvature is correlated to stem diameter and diameter growth. In contrast, wood maturation strain is independent from these dimensional effects, and is suitable as an intrinsic parameter characterising the magnitude of the plant’s gravitropic reaction.

Modelling, Evaluation and Biomechanical Consequences of Growth Stress Profiles Inside Tree Stems

The diameter growth of trees occurs by the progressive deposition of new wood layers at the stem periphery. These wood layers are submitted to at least two kinds of mechanical loads: maturation stress induced in wood during its formation, and the effect of the increasing self-weight. Interaction between growth and these loads causes mechanical stress with a particular distribution within the stem, called growth stresses. Growth stresses have technical consequences, such as cracks and deformations of lumber occurring during sawing, and biological consequences through their effect on stem strength. The first model for computing the field of stress inside a growing stem was set long ago by Kubler. Here, we extend these analytical formulations to cases with heterogeneous wood properties, eccentricity and bending stresses. Simulated profiles show reasonable agreement with measured profiles of released strains in logs. The particular shape of these profiles has consequences on stem bending strength. During bending in response to transient loads such as wind, most of the load is supported by outer parts of a stem cross section. The tensile maturation stress at this level increases the bending strength of the stem by delaying compression failure. Compressive stress in reaction to this tension does not reduce the bending strength because it is located near the centre of the stem and thus not loaded during bending, except if growth is strongly eccentric. Permanent bending stresses are concentrated at the mid-radius of the section, so that they do not cumulate with above-mentioned sources of stress. This smart distribution of stresses makes it possible that the stem is stronger than the wood it is made of, and that a growing stem can bend considerably more than its non-growing beam equivalent without breaking.

Mechanical contribution of secondary phloem to postural control in trees: the bark side of the force

To grow straight, plants need a motor systemxa0that 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.