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.

• Background and Aims Stem shape in angiosperms depends on several growth traits such as elongation direction, amount and position of axillary loads, stem dimensions, wood elasticity, radial growth dynamics and active re‐orientation due to tension wood. This paper analyses the relationship between these biomechanical factors and stem shape variability. • Methods Three apricot tree varieties with contrasting stem shape were studied. Growth and bending dynamics, mechanical properties and amount of tension wood were measured on 40 1‐year‐old stems of each variety during one growth season. Formulae derived from simple biomechanical models are proposed to quantify the relationship between biomechanical factors and re‐orientation of the stems. The effect of biomechanical factors is quantified combining their mechanical sensitivity and their actual variability. • Key results Re‐orientations happened in three main periods, involving distinct biomechanical phenomena: (a) passive bending due to the increase of shoot and fruit load at the start of the season; (b) passive uprighting at the fall of fruits; (c) active uprighting due tension wood production at the end of the season. Differences between varieties mainly happened during periods (a) and (b). • Conclusions The main factors causing differences between varieties are the length/diameter and the load/cross‐sectional area ratios during period (a). Wood elasticity does not play an important role because of its low inter‐variety variability. Differences during period (b) are related to the dynamics of radial growth: varieties with early radial growth bend weakly upward because the new wood layers tend to set them in a bent position. The action of tension wood during period (c) is low when compared with passive phenomena involved in periods (a) and (b).

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.

In order to face environmental constraints, trees are able to re-orient their axes by controlling the stress level in the newly formed wood layers. Angiosperms and gymnosperms evolved into two distinct mechanisms: the former produce a wood with large tension pre-stress on the upper side of the tilted axis, while the latter produce a wood with large compression pre-stress on the lower side. In both cases, the difference between this stress level and that of the opposite side, in light tension, generates the bending of the axis. However, light values of compression were sometimes measured in the opposite side of angiosperms. By analysing old data on chestnut and mani and new data on poplar, this study shows that these values were not measurement artefacts. This reveals that generating light compression stress in opposite wood contributes to improve the performance of the re-orientation mechanism.Pour s'adapter à l'environnement, les arbres sont capables de contrôler l'état de contraintes des nouvelles couches de bois formé pour réorienter leurs axes. Angiospermes et gymnospermes ont évolué vers deux stratégies différentes : les premiers produisent un bois à forte précontrainte de tension sur la face supérieure de l'axe incliné alors que les derniers mettent en place un bois en précontrainte de compression sur la face inférieure. La différence de contrainte avec la face opposée constituée de bois en légère tension produit un même résultat, la flexion de l'axe. Pourtant, au hasard des expérimentations, des valeurs de compression ont plusieurs fois été observées dans le bois opposé des angiospermes. Cette étude reprend des donnés anciennes sur le châtaignier et le manil et les complète avec des données sur peuplier pour mettre en évidence qu'il ne s'agit par d'erreurs de mesure mais réellement d'une stratégie de ces arbres consistant à produire du bois en légère précontrainte de compression pour une meilleure efficacité du redressement