Reductions in mesophyll conductance under drought stress are influenced by increases in cell wall chelator-soluble pectin content and denser microfibril alignment in cotton

Plants commonly undergo leaf morphoanatomy and composition modifications to cope with drought stress, and these tend to reduce mesophyll conductance to CO2 diffusion (gm), a key limitation to photosynthesis. The cell wall appears to play a crucial role in this reduction, yet the specific effect of cell wall component on gm and the underlying regulatory mechanisms of cell wall thickness (Tcw) variation are not well understood. In this study, we subjected cotton plants to varying levels of water deficit to investigate the impact of leaf cell wall component and the arrangement patterns of microfibrils within cell walls on Tcw and leaf gas exchange. Drought stress resulted in a significant thickening of cell walls and a decrease in gm. Concurrently, drought stress increased the content of chelator-soluble pectin and cellulose while reducing hemicellulose content. The alignment of cellulose microfibrils became more parallel and their diameter increased under drought conditions, suggesting a decrease in cell wall effective porosity which coincides with the observed reduction in gm. This research demonstrates that reduced gm typically observed under drought stress is related not only to thickened cell walls, but also to ultra-anatomical and compositional variations. Specifically, increases in cellulose content, diameter, and a highly aligned arrangement of cellulose microfibrils collectively contributed to an increase in Tcw, which, together with increases in chelator-soluble pectin content, resulted in an increased cell wall resistance to CO2 diffusion.