Abstract
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 stress”, occurs in wood fibres during cellular maturation when their secondary cell wall thickens. In this study, the spatial and temporal stiffening kinetics of the different cell wall layers were recorded during fibre maturation on a sample of poplar tension wood using atomic force microscopy. The thickening of the different layers was also recorded. The stiffening of the CML, S1 and S2-layers was initially synchronous with the thickening of the S2-layer and continued a little after the S2-layer reached its final thickness as the G-layer began to develop. In contrast, the global stiffness of the G-layer, which initially increased with its thickening, was close to stable long before it reached its final maximum thickness. A limited radial gradient of stiffness was observed in the G-layer, but it decreased sharply on the lumen side, where the new sub-layers are deposited during cell wall thickening. Although very similar at the ultrastructural and biochemical levels, the stiffening kinetics of the poplar G-layer appears to be very different from that described in maturing bast fibres.
Competing Interest Statement
The authors have declared no competing interest.
Footnotes
The introduction has been strongly modified to give the work "a more general focus" in order to make the findings of wider interest than just those readers specialising in the specific issue of the cell wall layers of tension wood fibres. Moreover, the interest of the comparison with flax bast fibres has been explained in detail in the introduction. Finally, a conclusion has been added to introduce a more general paragraph about the main novelties of the paper for the communities of wood sciences who study wood formation and wood mechanics, beyond tension wood studies. We have also added a paragraph explaining why these new results are not sufficient to challenge the hypothesized mechanisms of maturation stress generation discussed in Almeras and Clair (2016) and what type of additional measurements should be performed to complement the current results in order to be able to challenge the existing models. All the raw AFM data and the Excel sheets have been deposited on the open repository website Zenodo: https://doi.org/10.5281/zenodo.6487575.
Abbreviations
- AFM
- Atomic force microscopy
- CML
- Compound Middle Lamella
- CCML
- Cell Corner Middle Lamella
- MFA
- Microfibril angle
- PF-QNM
- Peak-force quantitative nano-mechanics