Summary Ferns and fern allies have low photosynthetic rates compared with seed plants. Their photosynthesis is thought to be limited principally by physical CO2 diffusion from the atmosphere to chloroplasts. The aim of this study was to understand the reasons for low photosynthesis in species of ferns and fern allies (Lycopodiopsida and Polypodiopsida). We performed a comprehensive assessment of the foliar gas‐exchange and mesophyll structural traits involved in photosynthetic function for 35 species of ferns and fern allies. Additionally, the leaf economics spectrum (the interrelationships between photosynthetic capacity and leaf/frond traits such as leaf dry mass per unit area or nitrogen content) was tested. Low mesophyll conductance to CO2 was the main cause for low photosynthesis in ferns and fern allies, which, in turn, was associated with thick cell walls and reduced chloroplast distribution towards intercellular mesophyll air spaces. Generally, the leaf economics spectrum in ferns follows a trend similar to that in seed plants. Nevertheless, ferns and allies had less nitrogen per unit DW than seed plants (i.e. the same slope but a different intercept) and lower photosynthesis rates per leaf mass area and per unit of nitrogen.

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Summary For land plants, water is the principal governor of growth. Photosynthetic performance is highly dependent on the stable and suitable water status of leaves, which is balanced by the water transport capacity, the water loss rate as well as the water capacitance of the plant. This review discusses the links between leaf water status and photosynthesis, specifically focussing on the coordination of CO2 and water transport within leaves, and the potential role of leaf capacitance and elasticity on CO2 and water transport.

The key role of cell walls in setting mesophyll conductance to CO2 (gm) and, consequently, photosynthesis, is reviewed. First, the theoretical properties of cell walls that can affect gm are presented. Then, we focus on cell wall thickness (Tcw) reviewing empirical evidence showing that Tcw varies strongly among species and phylogenetic groups in a way that correlates with gm and photosynthesis i.e. the thicker the mesophyll cell walls, the lower gm and photosynthesis. Potential interplays of gm, Tcw, dehydration tolerance and hydraulic properties of leaves are also discussed. Dynamic variations of Tcw in response to the environment and their implications in the regulation of photosynthesis are discussed, and recent evidence suggesting an influence of cell wall composition on gm are presented. We then propose a hypothetical mechanism for the influence of cell walls on photosynthesis, combining the effects of thickness and composition, particularly pectins. Finally, we discuss the prospects for using biotechnology for enhancing photosynthesis by means of altering cell wall-related genes.

A compromise between carbon assimilation and structure investment at the leaf level is broadly accepted, yet the relationship between net assimilation per area (A ) and leaf mass per area has been elusive. We propose bulk modulus of elasticity (ε) as a suitable parameter to reflect both leaf structure and function, and an inverse relationship between ε and A and mesophyll conductance (g ) is postulated. Using data for A , g and ε from previous studies and new measurements on a set of 20 species covering all major growth forms, a negative relationship between A or g and ε was observed. High ε was also related to low leaf capacitance and higher diffusive limitations to photosynthesis. In conclusion, ε emerges as a key trait linked with photosynthetic capacity across vascular plants, and its relationship with g suggests the existence of a common mechanistic basis, probably involving a key role of cell walls.