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Id | Title * | Authors * | Abstract * ▲ | Picture * | Thematic fields * | Recommender | Reviewers | Submission date | |
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08 Jul 2022
![]() Mechanical characterisation of the developing cell wall layers of tension wood fibres by Atomic Force MicroscopyOlivier Arnould, Marie Capron, Michel Ramonda, Françoise Laurans, Tancrède Alméras, Gilles Pilate, Bruno Clair https://doi.org/10.1101/2021.09.23.461481“Touching the Void”Recommended by Barry Gardiner based on reviews by Meriem Fournier and Félix HartmannUnderstanding the processes allowing trees to orientate their stems and branches requires an understanding of the mechanical properties of individual cells. As the cells are formed, maturation stresses are created that can lead to the reorientation of the tree. But measuring the properties within the different wood cells produced in normal wood, tension wood or compression wood requires measurements at very fine spatial resolution and the wood cells must remain in-situ so that the cell mechanical characteristics are preserved. In the article of Arnould et al (2022), measurements of the mechanical characteristics of poplar tension wood were measured in comparison to normal wood at different distances from the cambium and therefore different levels of maturation. The work required incredible care to embed the wood in resin, to cut the wood with extremely sharp microtone blades in order to minimize artefacts in the measurements, and then ultra-careful atomic force microscope (AFM) measurements across cell walls from the edge of the lumen to the middle lamella at extremely high spatial resolution. The result is a detailed picture of the kinetic development and maturation of tension wood cells in a tree. The measurements showed that the G-layer reaches close to its final stiffness long before its final thickness, and this is different from the maturation kinetics of other cell wall layers where thickening and stiffening are generally synchronous. Finally, although the G-layer in poplar tension wood fibres and in flax phloem fibres are in many ways very similar there are clear differences in the kinetics of their development and maturation. The detailed information presented in this paper can therefore help to clarify the different hypothetical mechanisms proposed to explain excess stress generation in the tension wood of trees and help move us towards a full understanding of how the “muscles” of trees work. References Arnould O, Capron M, Ramonda M, Laurans F, Alméras T, Pilate G, Clair B (2022) Mechanical characterisation of the developing cell wall layers of tension wood fibres by Atomic Force Microscopy. bioRxiv, 2021.09.23.461481, ver. 4 peer-reviewed and recommended by Peer Community in Forest and Wood Science. https://doi.org/10.1101/2021.09.23.461481 | Mechanical characterisation of the developing cell wall layers of tension wood fibres by Atomic Force Microscopy | Olivier Arnould, Marie Capron, Michel Ramonda, Françoise Laurans, Tancrède Alméras, Gilles Pilate, Bruno Clair | <p style="text-align: justify;">Trees generate mechanical stresses at periphery of stem and branches to improve their strength and to control the orientation of their axes. This key factor in the biomechanical design of trees, named “maturation st... | ![]() | Biology of wood in living trees | Barry Gardiner | 2022-01-11 17:49:47 | View | |
24 Jan 2025
![]() The diversity of radial variations of wood properties in European beech reveals the plastic nature of juvenile woodALMERAS Tancrède, JULLIEN Delphine, LIU Shengquan, LOUP Caroline, GRIL Joseph, THIBAUT Bernard https://hal.science/hal-04133248Recommendation of an interesting analysis of ontogenic and adaptive variations in local wood properties in European Beech (Fagus sylvatica L.)Recommended by Erwin DreyerThe mechanics of wood as a material for construction, furniture, pulp and other uses have been addressed in a very large number of papers and are a well-established field for both research and technical applications (for example among many others, see Pöhler et al., 2006 for beechwood). In addition to such approaches that derive from material sciences, further developments based on similar physical concepts addressed the questions raised by the biomechanics of the standing and the growing tree which requires some degree of postural control and sensing of specific signals (gravity, movements…; see for instance Fournier et al. 2013; Dlouha et al., 2025). Within this field of research, the question of the correlation of wood anatomy (diameter of xylem tracheids or vessels, fibre content and angles, vessel wall thickness… ) and biomechanical properties is of prime importance, and specific responses of wood and bark components have been identified over the last decades. In particular the occurrence of reaction wood generates local strains and contributes to the postural control (Ruelle, 2014). In this preprint, Almeras et al. address a complementary question related to the properties of juvenile wood in trees. During the first years of the growth of young trees, the annual tree rings display quite specific properties (large tree rings, less dense wood, …) that gradually change with age and dimensions of the trees until reaching a range of values typical for adult trees. During the first years, the interannual changes might follow an ontogenetic trajectory mainly related to age (and dimensions) while in later stages, they appear to be strongly controlled by environment (wind, soil fertility, site index, irradiance, water availability, ...). All these changes result in radial profiles along tree rings (from the pith to the bark) of three main features that govern the biomechanical properties of wood, namely the width of the annual tree ring, the local specific gravity (wood density), and the specific modulus which contributes with density to the local modulus of elasticity (Fournier et al. 2013). Such gradients of local wood properties within stems have been analysed and synthesised in the last years (Lachenbruch et al. 2011, Meinzer et al. 2014). Here, the authors address the question of local variations of such properties within tree stems as a function of the distance to the pith (inversely related to the age of the trees when the ring was formed) in a broadleaved species, European Beech (Fagus sylvatica L.). They checked whether ring width, specific gravity and specific modulus display systematic trends from pith to bark across tree stems, and whether these trends enable the detection of a general ontogenic (age-related) effect with very similar patterns in juvenile wood of different individuals, or whether adaptive factors (modulated by the environment and by the mechanical constraints induced by the postural control of growth) dominate already in juvenile wood, like it does at later stages. Such questions were already analysed in the wood of some coniferous species (softwood with tracheids), but less frequently in hardwood species (angiosperms, like Beech with its diffuse porous wood anatomy). Before starting the analysis of age-related tree ring properties in juvenile wood, the authors addressed the potential impact of duraminisation, which affects the oldest tree rings in the inner wood (that is those formed during the juvenile growth stages). Duraminisation results from local deposition of a number of secondary metabolites and results in the build-up of heartwood; in the case of beech however, reddish heartwood is less present than in other species (Knoke, 2003). Almeras et al showed here that the occurrence of reddish wood did only marginally affect the mechanical properties and contributed only marginally to the observed variations among trees The very solid experimental design enabled the authors to clearly assign a fraction of the observed variation in the three parameters to (i) the site where trees had grown, (ii) to the individuals within these sites and (iii) to the position of the ring within the stem. The intraindividual component of the variation was much larger than the former. However, the observed asymmetry in the patterns of ring properties in juvenile wood, and the large variability in these patterns among trees led the authors conclude that the ontogenic juvenility effects, visible in ring width were largely dominated by other effects influenced by the local environment. In this respect, the results differ from those that were recorded earlier with Pinus taeda L. in a plantation (i.e., trees of the same age and homogenous spatial distribution, Bendtsen and Senft, 1986). The recommended version of the preprint is very original as it shows how local (radial) variations of biomechanical wood properties can be addressed in a systematic way. This lead to novel approaches that share light on the processes governing wood formation in trees. The first version of the preprint was submitted over a year ago. The recommended version differs in many respects from the initial one. The two rounds of reviews with external reviewers, and the additional one with the recommender resulted in an in-depth reorganisation of the statistical analysis and of the demonstration. This took some time, but shows also the benefits that may be gained during an open peer review process like the one developed by the Peer Community in…. References Bendtsen BA, Senft J. 1986. Mechanical and anatomical properties in individual growth rings of plantation-grown eastern cottonwood and loblolly pine. Wood Fiber Sci 18: 23-38. Dlouhá J, Moulia B, Fournier, M et al. 2025 Beyond the perception of wind only as a meteorological hazard: importance of mechanobiology for biomass allocation, forest ecology and management. Ann For Sci 82, 1. https://doi.org/10.1186/s13595-024-01271-6 Fournier M, Dlouha J, Jaouen G, Alméras T. 2013. Integrative biomechanics for tree ecology: beyond wood density and strength. J Exp Bot, 64, 4793-4815. https://doi.org/10.1093/jxb/ert279 Knoke T. 2003 Predicting red heartwood formation in beech trees (Fagus sylvatica L.)? Ecol. Model. 169, 289-312. https://doi.org/10.1016/S0304-3800(03)00276-X Lachenbruch, B., Moore, J.R., Evans, R. (2011). Radial Variation in wood structure and function in woodypPlants, and hypotheses for its occurrence. In: Meinzer, F., Lachenbruch, B., Dawson, T. (eds) Size- and age-related changes in tree structure and function. Tree Physiology, vol 4. Springer, Dordrecht. https://doi.org/10.1007/978-94-007-1242-3_5 Longuetaud F, Mothe F, Santenoise P, Diop N, Dlouha J, Fournier M, Deleuze C. 2017. Patterns of within-stem variations in wood specific gravity and water content for five temperate tree species. Ann For Sci 74:64. https://doi.org/10.1007/s13595-017-0657-7 Meinzer, F., Lachenbruch, B., Dawson, T. (eds). 2014. Size- and age-related changes in tree structure and function. Tree Physiology, vol 4. Springer, Dordrecht. https://doi.org/10.1007/978-94-007-1242-3_5 Pöhler E, Klingner R, Künniger T. 2006. Beech (Fagus sylvatica L.) - Technological properties, adhesion behaviour and colour stability with and without coatings of the red heartwood. Ann For Sci 63: 129-137. https://doi.org/10.1051/forest:2005105 Ruelle J. 2014. Morphology, anatomy and ultrastructure of reaction wood. In: Gardiner B, Barnett J, Saranpää P, Gril J (eds) The Biology of Reaction Wood. Springer Series in Wood Science. Springer, Berlin, Heidelberg. https://doi.org/10.1007/978-3-642-10814-3_2 Cite the recommended preprint: Almeras Tancrède, Jullien Delphine, Liu Shengquan, Loup Caroline, Gril Joseph, Thibaut Bernard (2025) The diversity of radial variations of wood properties in European beech reveals the plastic nature of juvenile wood. HAL, ver.6 peer-reviewed and recommended by PCI Forest and Wood Sciences https://hal.science/hal-04133248 | The diversity of radial variations of wood properties in European beech reveals the plastic nature of juvenile wood | ALMERAS Tancrède, JULLIEN Delphine, LIU Shengquan, LOUP Caroline, GRIL Joseph, THIBAUT Bernard | <p>The long -term (as opposed to short-term intra-ring) radial variation of wood properties in European beech (Fagus sylvatica L.) from pith to bark are largest in the young ages of the tree (internal core). This so-called juvenility reflects both... | ![]() | Wood properties and wood products | Erwin Dreyer | 2023-07-01 10:42:48 | View | |
15 Mar 2021
![]() Variability in seeds’ physicochemical characteristics, germination and seedling growth within and between two French Populus nigra L. populationsMarlène Lefebvre, Marc Villar, Nathalie Boizot, Armelle Delile, Benjamin Dimouro, Anne-Marie Lomelech, Caroline Teyssier https://doi.org/10.48550/arXiv.2008.05744Black poplar (Populus nigra L.) seed physiology: an important issue for the in situ conservation of this riparian speciesRecommended by Erwin DreyerSeed physiology, which was a little forgotten in forest ecology since several decades, needs be revitalized as a research area given the many open questions about population dynamics and demography in rapidly changing environments (and not only for endangered species). Indeed, seed physiology was long mobilized mainly to optimize seed conservation and germination for the production of plant material in a range of tree species used for afforestation/plantation. In the case of back poplar (which by the way is the male genitor of the multiple hybrid Populus x euramerica poplar cultivars), the focus is rather on the conservation ecology of this riparian species, and mainly on in situ conservation (Lefèvre et al, 1998). Indeed, the protection of populations of Populus nigra L requires an improved understanding of the ecology of this species with a focus on reproduction. Indeed, black poplar seeds need to be rapidly disseminated, to germinate as soon as the conditions are favourable (with rather small time windows) and establish seedlings with access to water in the rather harsh environment of mobile and sandy river banks submitted to alternating periods of flooding and of severe water deficits during low river flows in summer (Imbert and Lefèvre, 2003; Corenblit et al, 2014; Tinschert et al, 2020). This process is therefore central to the propagation/maintenance of these populations that are threatened by the destruction of river banks and by introgression by either genes from the widespread “Italica” cultivar of black poplar, of from other poplar species leading to a variety of natural hybrids (Smulders et al, 2008). Many questions remain open about seeds of black poplar (Michalak et al, 2015). One of the most intriguing one is to what extent seed properties and physiology differ within and among local populations from different river catchments. This question was addressed in this preprint by Lefebvre et al. (2021) that provides a very detailed and comparative analysis of two populations from central and southern France, each represented by 10 half sib families (i.e., seed collected separately from 10 adult individuals after open pollination). Investigated properties were mainly seed biomass, anatomy, germination rate, root growth, lipid and sugar contents, protein content (with identification of some major protein families). The within populations variability was indeed quite large, but nevertheless there were significant differences between the two populations in several traits, like seed weight, lipid content, and starch content. Storage proteins differed among families, but only slightly between the two populations. However, the main conclusion was that intrinsic qualities of the seeds were not critical for early stage establishment in the two populations, despite some significant differences in mean seed biomass, in lipid and in soluble sugars contents. The preprint nicely analyses these differences, brings a large set of new observations about the seed physiology of Populus nigra. The referees found the data produced during this research quite important and original. This is why, despite the fact that the number of tested groups of populations remains rather small and the link with seedling establishment remains rather weak, this study is an important contribution to conservation ecology. This research (and that of many other groups) needs be further developed with an emphasis on inter and intra population variation and on demogenetics of forest tree species. References Corenblit D., Steiger J., González, E et al. (2014), The biogeomorphological life cycle of poplars during the fluvial biogeomorphological succession: a special focus on Populus nigra L.. Earth Surf. Process. Landforms, 39: 546-563. doi: https://doi.org/10.1002/esp.3515 Imbert E. and Lefèvre F. (2003) Dispersal and geneflow of Populus nigra (Salicaceae) along a dynamic river system. Journal of Ecology 91: 447-456. doi: https://doi.org/10.1046/j.1365-2745.2003.00772.x Lefebvre M., Villar M., Boizot N., Delile A., Dimouro B., Lomelech A.-M. and Teyssier, C. (2021) Variability in seeds’ physicochemical characteristics, germination and seedling growth within and between two French Populus nigra populations. arXiv, 2008.05744, ver 3 peer-reviewed and recommended by Peer community in Forest and Wood Sciences. https://arxiv.org/abs/2008.05744 Lefèvre F., Légionnet A., de Vries S. and Turok J. (1998) Strategies for the conservation of a pioneer tree species, Populus nigra L., in Europe. Genetics, Selection, Evolution 30, S181-196. doi: https://doi.org/10.1186/1297-9686-30-S1-S181 Michalak M., Plitta B.P., Tylkowski T. et al. (2015) Desiccation tolerance and cryopreservation of seeds of black poplar (Populus nigra L.), a disappearing tree species in Europe. European Journal of Forest Research 134, 53–60. doi: https://doi.org/10.1007/s10342-014-0832-4 Smulder M.J.M., Beringen R., Volosyanchuk R. et al. (2008) Natural hybridisation between Populus nigra L. and P. x canadensis Moench. Hybrid offspring competes for niches along the Rhine river in the Netherlands. Tree Genetics & Genomes 4, 663–675. doi: https://doi.org/10.1007/s11295-008-0141-5 Tinschert E., Egger G., Wendelgass J. et al. (2020) Alternate reproductive strategies of Populus nigra influence diversity, structure and successional processes within riparian woodlands along the Allier River, France. Journal of Hydro-environment research 30, 100-108. doi: https://doi.org/10.1016/j.jher.2020.03.004 | Variability in seeds’ physicochemical characteristics, germination and seedling growth within and between two French Populus nigra L. populations | Marlène Lefebvre, Marc Villar, Nathalie Boizot, Armelle Delile, Benjamin Dimouro, Anne-Marie Lomelech, Caroline Teyssier | <p>To improve understanding of the physiology, ecology and regeneration requirements of black poplar (Populus nigra), a severely endangered tree species in Europe, we analysed the biochemical composition of seeds from 20 families derived from open... | ![]() | Tree biology and physiology | Erwin Dreyer | 2020-08-14 09:59:17 | View |
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