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 (AN) 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 AN 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 AN) 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 AN with LMA across this species. The results revealed that gm was the most limiting factor for AN 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 (Tcw), chloroplast thickness (Tchl) and chloroplast exposed surface area facing intercellular air spaces (Sc/S). A positive scaling of gm and AN with Vc,max was also found, associated with an increased nitrogen content per area. A strong correlation of maximum photosynthetic electron transport (Jmax) with AN 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 within-species increases found in AN 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 Vc,max.

Cell-level anatomical characteristics explain high mesophyll conductance and photosynthetic capacity in sclerophyllous Mediterranean oaks.

Leaf mass per area (LMA) has been suggested to negatively affect the mesophyll conductance to CO2 (gm ), which is the most limiting factor for area-based photosynthesis (AN ) in many Mediterranean sclerophyll species. However, despite their high LMA, these species have similar AN to plants from other biomes. Variations in other leaf anatomical traits, such as mesophyll and chloroplast surface area exposed to intercellular air space (Sm /S and Sc /S), may offset the restrictions imposed by high LMA in gm and AN in these species. Seven sclerophyllous Mediterranean oaks from Europe/North Africa and North America with contrasting LMA were compared in terms of morphological, anatomical and photosynthetic traits. Mediterranean oaks showed specific differences in AN that go beyond the common morphological leaf traits reported for these species (reduced leaf area and thick leaves). These variations resulted mainly from the differences in gm , the most limiting factor for carbon assimilation in these species. Species with higher AN showed increased Sc /S, which implies increased gm without changes in stomatal conductance. The occurrence of this anatomical adaptation at the cell level allowed evergreen oaks to reach AN values comparable to congeneric deciduous species despite their higher LMA.

Leaf morphological and physiological adaptations of a deciduous oak (Quercus faginea Lam.) to the Mediterranean climate: a comparison with a closely related temperate species (Quercus robur L.).

'White oaks'--one of the main groups of the genus Quercus L.--are represented in western Eurasia by the 'roburoid oaks', a deciduous and closely related genetic group that should have an Arcto-Tertiary origin under temperate-nemoral climates. Nowadays, roburoid oak species such as Quercus robur L. are still present in these temperate climates in Europe, but others are also present in southern Europe under Mediterranean-type climates, such as Quercus faginea Lam. We hypothesize the existence of a coordinated functional response at the whole-shoot scale in Q. faginea under Mediterranean conditions to adapt to more xeric habitats. The results reveal a clear morphological and physiological segregation between Q. robur and Q. faginea, which constitute two very contrasting functional types in response to climate dryness. The most outstanding divergence between the two species is the reduction in transpiring area in Q. faginea, which is the main trait imposed by the water deficit in Mediterranean-type climates. The reduction in leaf area ratio in Q. faginea should have a negative effect on carbon gain that is partially counteracted by a higher inherent photosynthetic ability of Q. faginea when compared with Q. robur, as a consequence of higher mesophyll conductance, higher maximum velocity of carboxylation and much higher stomatal conductance (gs). The extremely high gs of Q. faginea counteracts the expected reduction in gs imposed by the stomatal sensitivity to vapor pressure deficit, allowing this species to diminish water losses maintaining high net CO2 assimilation values along the vegetative period under nonlimiting soil water potential values. In conclusion, the present study demonstrates that Q. faginea can be regarded as an example of adaptation of a deciduous oak to Mediterranean-type climates.

Leaf functional plasticity decreases the water consumption without further consequences for carbon uptake in Quercus coccifera L. under Mediterranean conditions.

The accumulation of epicuticular waxes over stomata in Quercus coccifera L. contributes to a severe reduction in maximum stomatal conductance (g s,max) under Mediterranean (MED) conditions. However, this phenomenon was not observed in this species under temperate (TEM) conditions, which could lead to differences in the ability to assimilate CO2 between the sites. We hypothesise that the overall importance of such a reduction in gs,max on photosynthesis is modulated by other factors affecting carbon gain, mainly mesophyll conductance to CO2 (g m), through a plastic response to changes in environmental conditions (i.e., vapour pressure deficit, VPD, and mean daily quantum flux density, Q int). The results reveal that leaves grown at the TEM site did not show an increased ability for net CO2 assimilation (A N), mainly due to an equal gm at both sites. This fact is explained by a trade-off between an increased conductance of the gas phase (g ias) and a reduced conductance of the liquid phase (g liq) at the TEM site compared with the MED site. In spite of the reduction in gs,max at the MED site, transpiration (E) did not diminish during midsummer to the levels of the TEM site due to a higher VPD found at the MED site, yielding a higher water use efficiency (AN/E) at the TEM site. Moreover, photosynthetic nitrogen use efficiency was also higher at the TEM site, indicating these leaves can reach similar values of AN with lower nitrogen investment that those at the MED site. These results suggest that Q. coccifera does not always use the main resources (water and nutrients) at leaf level as efficiently as possible. Moreover, the different patterns of resource use (in particular N), together with the functional plasticity, cannot overcome the morpho-functional constraints that limit photosynthetic activity, even under potentially favourable conditions.