In the Mediterranean evergreen oak woodlands of southern Portugal, the main tree species are Quercus ilex ssp. rotundifolia Lam. (holm oak) and Quercus suber L. (cork oak). We studied a savannah-type woodland where these species coexist, with the aim of better understanding the mechanisms of tree adaptation to seasonal drought. In both species, seasonal variations in transpiration and predawn leaf water potential showed a maximum in spring followed by a decline through the rainless summer and a recovery with autumn rainfall. Although the observed decrease in predawn leaf water potential in summer indicates soil water depletion, trees maintained transpiration rates above 0.7 mm day−1 during the summer drought. By that time, more than 70% of the transpired water was being taken from groundwater sources. The daily fluctuations in soil water content suggest that some root uptake of groundwater was mediated through the upper soil layers by hydraulic lift. During the dry season, Q. ilex maintained higher predawn leaf water potentials, canopy conductances and transpiration rates than Q. suber. The higher water status of Q. ilex was likely associated with their deeper root systems compared with Q. suber. Whole-tree hydraulic conductance and minimum midday leaf water potential were lower in Q. ilex, indicating that Q. ilex was more tolerant to drought than Q. suber. Overall, Q. ilex seemed to have more effective drought avoidance and drought tolerance mechanisms than Q. suber.
Responses of leaf water relations and photosynthesis to summer drought and autumn rewetting were studied in two evergreen Mediterranean oak species, Quercus ilex spp. rotundifolia and Quercus suber. The predawn leaf water potential (ΨlPD), stomatal conductance (gs) and photosynthetic rate (A) at ambient conditions were measured seasonally over a 3-year period. We also measured the photosynthetic response to light and to intercellular CO2 (A/PPFD and A/Ci response curves) under water stress (summer) and after recovery due to autumn rainfall. Photosynthetic parameters, Vcmax, Jmax and triose phosphate utilization (TPU) rate, were estimated using the Farquhar model. RuBisCo activity, leaf chlorophyll, leaf nitrogen concentration and leaf carbohydrate concentration were also measured. All measurements were performed in the spring leaves of the current year. In both species, the predawn leaf water potential, stomatal conductance and photosynthetic rate peaked in spring, progressively declined throughout the summer and recovered upon autumn rainfall. During the drought period, Q. ilex maintained a higher predawn leaf water potential and stomatal conductance than Q. suber. During this period, we found that photosynthesis was not only limited by stomatal closure, but was also downregulated as a consequence of a decrease in the maximum carboxylation rate (Vcmax) and the light-saturated rate of photosynthetic electron transport (Jmax) in both species. The Vcmax and Jmax increased after the first autumnal rains and this increase was related to RuBisCo activity, leaf nitrogen concentration and chlorophyll concentration. In addition, an increase in the TPU rate and in soluble leaf sugar concentration was observed in this period. The results obtained indicate a high resilience of the photosynthetic apparatus to summer drought as well as good recovery in the following autumn rains of these evergreen oak species.
The effects of growth at elevated CO2 on the response to high temperatures in terms of carbon assimilation (net photosynthesis, stomatal conductance, amount and activity of Rubisco, and concentrations of total soluble sugars and starch) and of photochemistry (for example, the efficiency of excitation energy captured by open photosystem II reaction centres) were studied in cork oak (Quercus suber L.). Plants grown in elevated CO2 (700 ppm) showed a down-regulation of photosynthesis and had lower amounts and activity of Rubisco than plants grown at ambient CO2 (350 ppm), after 14 months in the greenhouse. At that time plants were subjected to a heat-shock treatment (4 h at 45°C in a chamber with 80% relative humidity and 800–1000 µmol m−2 s−1 photon flux density). Growth in a CO2-enriched atmosphere seems to protect cork oak leaves from the short-term effects of high temperature. Elevated CO2 plants had positive net carbon uptake rates during the heat shock treatment whereas plants grown at ambient CO2 showed negative rates. Moreover, recovery was faster in high CO2-grown plants which, after 30 min at 25°C, exhibited higher net carbon uptake rates and lower decreases in photosynthetic capacity (Amax as well as in the efficiency of excitation energy captured by open photosystem II reaction centres (FvJFm than plants grown at ambient CO2. The stomata of elevated CO2 plants were also less responsive when exposed to high temperature.