Photosynthetic limitation of several representative subalpine species in the Catalan Pyrenees in summer.

Information on the photosynthetic process and its limitations is essential in order to predict both the capacity of species to adapt to conditions associated with climate change and the likely changes in plant communities. Considering that high-mountain species are especially sensitive, three species representative of subalpine forests of the Central Catalan Pyrenees: mountain pine (Pinus uncinata Mill.), birch (Betula pendula Roth) and rhododendron (Rhododendron ferrugineum L.) were studied under conditions associated with climate change, such as low precipitation, elevated atmospheric [CO2 ] and high solar irradiation incident at Earth's surface, in order to detect any photosynthetic limitations. Short-term high [CO2 ] increased photosynthesis rates (A) and water use efficiency (WUE), especially in birch and mountain pine, whereas stomatal conductance (gs ) was not altered in either species. Birch showed photosynthesis limitation through stomatal closure related to low rainfall, which induced photoinhibition and early foliar senescence. Rhododendron was especially affected by high irradiance, showing early photosynthetic saturation in low light, highest chlorophyll content, lowest gas exchange rates and least photoprotection. Mountain pine had the highest A, photosynthetic capacity (Amax ) and light-saturated rates of net CO2 assimilation (Asat ), which were maintained under reduced precipitation. Furthermore, maximum quantum yield (Fv /Fm ), thermal energy dissipation, PRI and SIPI radiometric index, and ascorbate content indicated improved photoprotection with respect to the other two species. However, maximum velocity of carboxylation of RuBisco (Vcmax ) indicated that N availability would be the main photosynthetic limitation in this species.

Antioxidant and photoprotective responses to elevated CO(2) and heat stress during holm oak regeneration by resprouting, evaluated with NIRS (near-infrared reflectance spectroscopy).

Photosynthetic, photoprotective and antioxidant responses during high temperature stress were determined in leaves of evergreen holm oak (Quercus ilex L.), the main species in Mediterranean forests, during resprouting under elevated CO(2) (750 µl·l(-1) ). Leaf chemicals, chloroplast pigments and non-enzymatic antioxidants were quantified in a single measurement using NIRS (near-infrared spectroscopy), a rapid and suitable method for ecophysiological purposes. Resprouts from plants grown under elevated CO(2) (RE) showed photosynthetic down-regulation, higher starch content and lower stomatal conductance, but similar stomatal density, than plants grown under current CO(2) concentrations (350 µl·l(-1) ) (RA). The photosynthetic sink reduction and need for more antioxidants and photoprotection in RE were reflected in an increased concentration of ascorbate (Asc) and phenolic compounds and in the contribution of the xanthophyll (Z/VAZ) and lutein epoxide cycles to excess energy dissipation as heat, and also reflected in chlorophyll fluorescence measurements. CO(2) assimilation parameters were stable from 25 to 35 °C in RE and RA, declining thereafter in RA in spite of a 2.3 °C lower leaf temperature. RE showed a more marked decline in photorespiration above 35 °C and less sensitive stomatal responses to high temperature stress than RA. During heat stress, RE had higher Asc, Z/VAZ and phenolics content, together with delayed enhancement of chloroplast lipophilic antioxidant compounds (carotenes and tocopherols). The high contribution of photoprotective systems and high temperature tolerance in resprouts developed under elevated CO(2) would mitigate the effect of photosynthesis acclimation during the regeneration of Q. ilex plants under climate change.

Photosynthesis and photoprotection in Quercus ilex resprouts after fire.

Plants that resprout after fires often have higher rates of photosynthesis than before a fire. To elucidate the mechanism of this response, we studied gas exchange and chlorophyll fluorescence in Quercus ilex L. plants growing on control (unburned) sites and on sites that had been burned the preceding summer. In early July, photosynthetic rates and stomatal conductance were similar in plants on unburned and burned plots, and in young and old foliage within unburned plots. At this time, photochemical efficiency of photosystem II (PSII), nonphotochemical quenching of chlorophyll fluorescence (NPQ), and the de-epoxidation of violaxanthin to zeaxanthin were also similar among leaves of different ages and treatments. In late July, photosynthetic rates and stomatal conductances were much greater in resprouts on the burned areas than in unburned plants. From early to late July, unburned plants showed an increase in NPQ and the de-epoxidation of violaxanthin to zeaxanthin, indicating increased photoprotection as a result of enhanced nonradiative dissipation of excess light energy. Plants on the burned plots did not show these changes. Leaves of all ages and treatments showed no substantial reduction in potential quantum yield of PSII (F(v)/F(m)) at midday or predawn, indicating that there was little or no photoinhibition. Leaf nitrogen and soluble protein contents varied with leaf age during July, but did not vary between treatments. We conclude that the primary effect of burning is an increase in water availability to resprouting plants that eliminates the need for photoprotection, at least in the short term. The decrease in photosynthetic rates of unburned leaves in late July was the result of reduced stomatal conductance. We suggest that lowered stomatal conductance is the primary limiting factor in Q. ilex leaves, governing the regulation of carboxylation activity and energy dissipation processes.