To grow straight, plants need a motor system that 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.

International audienceTension wood, a tissue developed by angiosperm trees to actively recover their verticality, has long been defined by the presence of an unlignified cellulosic inner layer in the cell wall of fibres, called the G-layer. Although it was known that some species have no G-layer, the definition was appropriate since it enabled easy detection of tension wood zones using various staining techniques for either cellulose or lignin. For several years now, irrespective of its anatomical structure, tension wood has been defined by its high mechanical internal tensile stress. This definition enables screening of the diversity of cell walls in tension wood fibres. Recent results obtained in tropical species with tension wood with a delay in the lignification of the G-layer opened our eyes to the effective presence of large amounts of lignin in the G-layer of some species. This led us to review older literature mentioning the presence of lignin deposits in the G-layer and give them credit. Advances in the knowledge of tension wood fibres allow us to reconsider some previous classifications of the diversity in the organisation of the fibre walls of the tension wood

Trees 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.

Salicaceae have been enlarged to include a majority of the species formerly placed in the polyphyletic tropical Flacourtiaceae. Several studies have reported a peculiar and infrequently formed multilayered structure of tension wood in four of the tropical genera. Tension wood is a tissue produced by trees to restore their vertical orientation and most studies have focused on trees developing tension wood by means of cellulose‐rich, gelatinous fibres, as in Populus and Salix (Salicaceae s.s.). This study aims to determine if the multilayered structure of tension wood is an anatomical characteristic common in other Salicaceae and, if so, how its distribution correlates to phylogenetic relationships. Therefore, we studied the tension wood of 14 genera of Salicaceae and two genera of Achariaceae, one genus of Goupiaceae and one genus of Lacistemataceae, families closely related to Salicaceae or formerly placed in Flacourtiaceae. Opposite wood and tension wood were compared with light microscopy and three‐dimensional laser scanning confocal microscopy. The results indicate that a multilayered structure of tension wood is common in the family except in Salix, Populus and one of their closest relatives, Idesia polycarpa. We suggest that tension wood may be a useful anatomical character in understanding phylogenetic relationships in Salicaceae. Further investigation is still needed on the tension wood of several other putatively close relatives of Salix and Populus, in particular Bennettiodendron, Macrohasseltia and Itoa.

Acer pseudoplatanus is a worldwide-distributed tree which contains toxins, among them hypoglycin A (HGA). This toxin is known to be responsible for poisoning in various species, including humans, equids, Père David’s deer and two-humped camels. We hypothesized that any herbivore pasturing with A. pseudoplatanus in their vicinity may be at risk for HGA poisoning. To test this hypothesis, we surveyed the HGA exposure from A. pseudoplatanus in species not yet described as being at risk. Animals in zoological parks were the major focus, as they are at high probability to be exposed to A. pseudoplatanus in enclosures. We also searched for a toxic metabolite of HGA (i.e., methylenecyclopropylacetyl-carnitine; MCPA-carnitine) in blood and an alteration of the acylcarnitines profile in HGA-positive animals to document the potential risk of declaring clinical signs. We describe for the first instance cases of HGA poisoning in Bovidae. Two gnus (Connochaetes taurinus taurinus) exposed to A. pseudoplatanus in their enclosure presented severe clinical signs, serum HGA and MCPA-carnitine and a marked modification of the acylcarnitines profile. In this study, even though all herbivores were exposed to A. pseudoplatanus, proximal fermenters species seemed less susceptible to HGA poisoning. Therefore, a ruminal transformation of HGA is hypothesized. Additionally, we suggest a gradual alteration of the fatty acid metabolism in case of HGA poisoning and thus the existence of subclinical cases.

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.

& Key message Gravitropic movements in angiosperm woody stems are achieved through the action of bark and/or wood motor, depending on the bark and wood fibre anatomy (with trellis structure or not; with G-layers or not). Bark motor is as efficient as wood motor to recover from tilting in young trees of 21 tropical species. & Context Angiosperm trees produce tension wood to control their orientation through changes in stem curvature. Tension wood is classified into 3 anatomical groups: with unlignified G-layer, with lignified G-layer and without G-layer. & Aims This study aimed at assessing whether this anatomical diversity reflects a diversity in efficiency of gravitropic movement. & Methods The study was conducted on tropical seedling from the three anatomical groups. Seedlings were staked and grown tilted. At the end of the experiment, changes in curvature when releasing the stem from the stake and when removing bark were measured. Three parameters were computed to compare the global efficiency of gravitropism (stem gravitropic efficiency) and the specific efficiency of motor mechanism based on wood (maturation strain of tension wood) and bark (standardized debarking curvature). & Results The maturation strain of tension wood was similar between species with unlignified and lignified G-layer. Species without G-layer exhibited low maturation strain and large debarking curvature, showing they rely on bark for gravitropism. Bark and wood achieved similar motor efficiency. & Conclusion Lignin does not affect the generation of tensile stress in the G-layer. Bark can be as efficient as wood as a motor of gravitropic movements.

In spring, grazing equids may suffer from a severe rhabdomyolysis syndrome named atypical myopathy. This environmental intoxication results from ingestion of toxins contained in Acer pseudoplatanus seedlings. The aim of this study was to investigate the effectiveness of herbicide spraying and mowing to reduce the toxic pressure of sycamore seedlings. In a first experiment, the efficacy of three herbicides to eradicate seedlings was compared to mowing. In a second experiment, the influence of the mowing timing on pasture productivity was determined. In both experiments, sycamore seedling counting, grass height and botanical composition were determined. In experiment 2, the final harvest biomass and its nutritional value were also determined. Herbicides and mowing both reduced the number of seedlings, which nevertheless disappear naturally over time without intervention (i.e., in control areas). As opposed to mowing, herbicide spraying altered the composition of the pasture flora. Both sprayed and mowed seedlings remain toxic until full decomposition. Early mowing (i.e., early April) did not affect the harvest yield. Late mowing (i.e., end of April) reduced the harvest yield but its nutritional value fitted the horses’ need. In conclusion, mowing is the best strategy to reduce the risk of Acer pseudoplatanus poisoning in grazing equids while preserving ecological sustainability and nutritional value of pastures.