Coordinated modifications in mesophyll conductance, photosynthetic potentials and leaf nitrogen contribute to explain the large variation in foliage net assimilation rates across Quercus ilex provenances

Leaf dry mass per unit area (LMA) has been suggested to negatively affect the mesophyll conductance to CO2 (gm), the most limiting factor for photosynthesis per unit leaf area (A(N)) in many evergreens. Several anatomical traits (i.e., greater leaf thickness and thicker cell walls) constraining gm could explain the negative scaling of gm and A(N) with LMA across species. However, the Mediterranean sclerophyll Quercus ilex L. shows a major within-species variation in functional traits (greater LMA associated with higher nitrogen content and A(N)) that might contrast the worldwide trends. The objective of this study was to elucidate the existence of variations in other leaf anatomical parameters determining gm and/or biochemical traits improving the capacity of carboxylation (Vc, max) that could modulate the relationship of A(N) with LMA across this species. The results revealed that gm was the most limiting factor for A(N) in all the studied Q. ilex provenances from Spain and Italy. The within-species differences in gm can be partly attributed to the variation in several leaf anatomical traits, mainly cell-wall thickness (T-cw), chloroplast thickness (T-chl) and chloroplast exposed surface area facing intercellular air spaces (Sc/S). A positive scaling of gm and A(N) with V-c, max was also found, associated with an increased nitrogen content per area. A strong correlation of maximum photosynthetic electron transport (J(max)) with A(N) further indicated a coordination between the carboxylase activity and the electron transport chain. In conclusion, we have confirmed the strong ecotypic variation in the photosynthetic performance of individual provenances of Q. ilex. Thus, the withinspecies increases found in A(N) for Q. ilex with increasing foliage robustness can be explained by a synergistic effect among anatomical (at the subcellular and cellular level) and biochemical traits, which markedly improved gm and V-c, max.