What drives photosynthesis during desiccation? Mosses and other outliers from the photosynthesis-elasticity trade-off

In vascular plants, more rigid leaves have been linked to lower photosynthetic capacity, associated with low CO2 diffusion across the mesophyll, indirectly resulting in a trade-off between photosynthetic capacity (A(n)) and bulk modulus of elasticity (epsilon). However, we evaluated mosses, liverworts, and Chara sp., plus some lycophytes and ferns, and found that they behaved as clear outliers of the A(n)-epsilon relationship. Despite this finding, when vascular and non-vascular plants were plotted together, epsilon still linearly determined the cessation of net photosynthesis during desiccation both in species with stomata (either actively or hydro-passively regulated) and in species lacking stomata, and regardless of their leaf structure. The latter result challenges our current view of photosynthetic responses to desiccation and/or water stress. Structural features and hydric strategy are discussed as possible explanations for the deviation of these species from the A(n)-epsilon trade-off, as well as for the general linear dependency between e and the full cessation of An during desiccation.