Bruno Clair

Relationship between growth stress, mechanical-physical properties and proportion of fibre with gelatinous layer in chestnut (Castanea sativa Mill.)

Summary A range of mechanical and physical properties were determined for 96 specimens of chestnut wood and for wood types ranging from compression to tension wood; tests included (1) growth stress, (2) longitudinal Youngs modulus in green and air-dried states (3) shrinkage in longitudinal and tangential directions. Anatomical observations permitted determination of the proportion of fibres with a gelatinous layer. The influence of these atypical fibres on macroscopic wood properties is examined and discussed. A basic model is proposed to determine their properties in theoretically isolated conditions.

Maturation Stress Generation in Poplar Tension Wood Studied by Synchrotron Radiation Microdiffraction

Tension wood is widespread in the organs of woody plants. During its formation, it generates a large tensile mechanical stress called maturation stress. Maturation stress performs essential biomechanical functions such as optimizing the mechanical resistance of the stem, performing adaptive movements, and ensuring the long-term stability of growing plants. Although various hypotheses have recently been proposed, the mechanism generating maturation stress is not yet fully understood. In order to discriminate between these hypotheses, we investigated structural changes in cellulose microfibrils along sequences of xylem cell differentiation in tension and normal wood of poplar (Populus deltoides × Populus trichocarpa ‘I45-51’). Synchrotron radiation microdiffraction was used to measure the evolution of the angle and lattice spacing of crystalline cellulose associated with the deposition of successive cell wall layers. Profiles of normal and tension wood were very similar in early development stages corresponding to the formation of the S1 layer and the outer part of the S2 layer. Subsequent layers were found with a lower microfibril angle (MFA), corresponding to the inner part of the S2 layer of normal wood (MFA approximately 10°) and the G layer of tension wood (MFA approximately 0°). In tension wood only, this steep decrease in MFA occurred together with an increase in cellulose lattice spacing. The relative increase in lattice spacing was found close to the usual value of maturation strains. Analysis showed that this increase in lattice spacing is at least partly due to mechanical stress induced in cellulose microfibrils soon after their deposition, suggesting that the G layer directly generates and supports the tensile maturation stress in poplar tension wood.

Tension wood and opposite wood in 21 tropical rain forest species. 2. Comparison of some anatomical and ultrastructural criteria

The anatomy of tension wood and opposite wood was compared in 21 tropical rain forest trees from 21 species belonging to 18 families from French Guyana. Wood specimens were taken from the upper and lower sides of naturally tilted trees. Measurement of the growth stress level ensured that the two samples were taken from wood tissues in a different mechanical state: highly tensile-stressed wood on the upper side, called tension wood and normally tensile-stressed wood on the lower side, called opposite wood. Quantitative parameters relating to fibres and vessels were measured on transverse sections of both tension and opposite wood to check if certain criteria can easily discriminate the two kinds of wood. We observed a decrease in the frequency of vessels in the tension wood in all the trees studied. Other criteria concerning shape and surface area of the vessels, fibre diameter or cell wall thickness did not reveal any general trend. At the ultrastructural level, we observed that the microfibril angle in the tension wood sample was lower than in opposite wood in all the trees except one (Licania membranacea).

Mesoporosity as a new parameter for understanding tension stress generation in trees

The mechanism for tree orientation in angiosperms is based on the production of high tensile stress on the upper side of the inclined axis. In many species, the stress level is strongly related to the presence of a peculiar layer, called the G-layer, in the fibre cell wall. The structure of the G-layer has recently been described as a hydrogel thanks to N(2) adsorption-desorption isotherms of supercritically dried samples showing a high mesoporosity (pores size from 2-50 nm). This led us to revisit the concept of the G-layer that had been, until now, only described from anatomical observation. Adsorption isotherms of both normal wood and tension wood have been measured on six tropical species. Measurements show that mesoporosity is high in tension wood with a typical thick G-layer while it is much less with a thinner G-layer, sometimes no more than normal wood. The mesoporosity of tension wood species without a G-layer is as low as in normal wood. Not depending on the amount of pores, the pore size distribution is always centred around 6-12 nm. These results suggest that, among species producing fibres with a G-layer, large structural differences of the G-layer exist between species.

Characterization of a gel in the cell wall to elucidate the paradoxical shrinkage of tension wood.

Wood behavior is characterized by high sensibility to humidity and strongly anisotropic properties. The drying shrinkage along the fibers, usually small due to the reinforcing action of cellulosic microfibrils, is surprisingly high in the so-called tension wood, produced by trees to respond to strong reorientation requirements. In this study, nitrogen adsorption-desorption isotherms of supercritically dried tension wood and normal wood show that the tension wood cell wall has a gel-like structure characterized by a pore surface more than 30 times higher than that in normal wood. Syneresis of the tension wood gel explains its paradoxical drying shrinkage. This result could help to reduce technological problems during drying. Potential applications in biomechanics and biomimetics are worth investigating, considering that, in living trees, tension wood produces tensile growth stresses 10 times higher than that of normal wood.

SHRINKAGE OF THE GELATINOUS LAYER OF POPLAR AND BEECH TENSION WOOD

Macroscopic longitudinal shrinkage of beech and poplar tension wood is higher than in normal wood. This shrinkage is the result of mechanical interactions of cell wall layers. SEM observation of cut, dried surfaces showed that longitudinal shrinkage is much greater in the gelatinous layer than in other layers. AFM topographic images of the same cells, both in water and in air-dry conditions, confirm this result. Measurements on sections indicate around 4.7% longitudinal shrinkage for the G layer.

GROWTH STRESSES ARE HIGHLY CONTROLLED BY THE AMOUNT OF G-LAYER IN POPLAR TENSION WOOD.

To determine how gelatinous fibres and gelatinous layers contribute to the magnitude of longitudinal growth stress in tension wood, anatomical measurements of gelatinous fibres were carried out on poplar tension wood (Populus I4551). It was found that (a) no gelatinous fibres were observed under a growth strain level of 0.06 to 0.08%; (b) almost 100% of the non-conductive tissues contained gelatinous fibres above a growth strain level of 0.15 to 0.19%; and (c) the area of fibres, the area of fibres with gelatinous layers per unit of tissue area, and the thickness of the gelatinous layers predominantly influenced the magnitude of growth stress

Precautions for the structural analysis of the gelatinous layer in tension wood

The gelatinous layer (G-layer) of tension wood fibres in hardwood contributes to the mechanical function of the living tree and has significant consequences on properties of solid wood. Its size, shape and structure observed by optical or electron microscopy exhibits characteristic anatomical features. However, we found that sectioning of non-embedded wood samples results in an uncontrolled swelling of the G-layer. In order to assess this artefact, the shape and thickness of the G-layer was monitored by serial sections from an embedded wood sample, from its trimmed transverse face to that located several hundreds of micrometres deep. The results revealed that the initial cutting before embedding produced a border effect responsible for the swollen nature, which is similar to sections from non-embedded material. After a conventional embedding technique was applied, a section of at least 30 micrometres below the trimming surface is required to observe an un-swollen G-layer.

On the detachment of the gelatinous layer in tension wood fiber

The detachment of the gelatinous layer (G-layer), often observed on microtome cross sections, has led some authors to believe that the G-layer cannot act as the driving force of longitudinal shrinkage in tension wood. The aim of this study was to observe the detachment of the G-layer along fibers. Green wood blocks were cut transversely into two samples. One sample was kept in water and the other was oven-dried. With one face being common to both samples, the detachment of the G-layer was studied on the same fibers. Observations were performed after blocking deformation by embedding. This revealed that the detachment of the G-layer is an effect produced by the act of cutting the transverse face of the wood block to be embedded. At distances greater than 100 µm from this primary surface of the sample, no detachment was observed. Drying shrinkage shows little or no effect on this detachment. The result seems to explain well why the detachment of the G-layer occurs during sectioning using conventional sliding microtomy. These observations prove the adhesion of the G-layer in massive wood and confirm the active role of the G-layer in tension wood properties.

Compression stress in opposite wood of angiosperms : observations in chestnut, mani and poplar

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.