Enhanced spring warming in a Mediterranean mountain by atmospheric circulation.

We analyzed trends of air temperature across the Cévennes National Park in Southern France, a mid-altitude coastal mountain experiencing a rapid spread of forests at the expense of rangelands and submitted to Mediterranean Sea influences and so, impacted by local and regional processes of climate change. Since 1980, April to June warming trend reached a maximum temperature increase of + 0.124 °C year-1 and uniform whatever the altitude. Minimum temperature increased by + 0.058 °C year-1 at 500 m altitude and + 0.089 °C year-1 at 1500 m. Concomitantly, forest cover is increasing by + 0.51% year-1. Using an intrinsic biophysical mechanism model, we demonstrated that, at monthly scale, the forest surface is 1.7-3.1 °C cooler than that of nearby grasslands. As a result, the decrease in albedo corresponding to the conversion from grasslands to dense forests, translates into a cooling of maximum air temperatures of 0.023 °C year-1 which contributes to slow down the warming rate enhancement. Spring warming trends co-varied with negative WeMO phases associated with a low in the Gulf of Cádiz and an anticyclone in Central Europe. An east to west pressure gradient increases atmospheric humidity leading to a strong water vapor feedback, enhancing the forcing of thermal long wave radiations and hence the rise in temperature.

How reliable are methods to assess xylem vulnerability to cavitation? The issue of 'open vessel' artifact in oaks.

Three methods are in widespread use to build vulnerability curves (VCs) to cavitation. The bench drying (BD) method is considered as a reference because embolism and xylem pressure are measured on large branches dehydrating in the air, in conditions similar to what happens in nature. Two other methods of embolism induction have been increasingly used. While the Cavitron (CA) uses centrifugal force to induce embolism, in the air injection (AI) method embolism is induced by forcing pressurized air to enter a stem segment. Recent studies have suggested that the AI and CA methods are inappropriate in long-vesselled species because they produce a very high-threshold xylem pressure for embolism (e.g., P50) compared with what is expected from (i) their ecophysiology in the field (native embolism, water potential and stomatal response to xylem pressure) and (ii) the P50 obtained with the BD method. However, other authors have argued that the CA and AI methods may be valid because they produce VCs similar to the BD method. In order to clarify this issue, we assessed VCs with the three above-mentioned methods on the long-vesselled Quercus ilex L. We showed that the BD VC yielded threshold xylem pressure for embolism consistent with in situ measurements of native embolism, minimal water potential and stomatal conductance. We therefore concluded that the BD method provides a reliable estimate of the VC for this species. The CA method produced a very high P50 (i.e., less negative) compared with the BD method, which is consistent with an artifact related to the vessel length. The VCs obtained with the AI method were highly variable, producing P50 ranging from -2 to -8.2 MPa. This wide variability was more related to differences in base diameter among samples than to differences in the length of samples. We concluded that this method is probably subject to an artifact linked to the distribution of vessel lengths within the sample. Overall, our results indicate that the CA and the AI should be used with extreme caution on long-vesselled species. Our results also highlight that several criteria may be helpful to assess the validity of a VC.

Mast seeding under increasing drought: results from a long-term data set and from a rainfall exclusion experiment.

Mast seeding, the synchronous, highly variable seed production among years, is very common in tree species, but there is no consensus about its main causes and the main environmental factors affecting it. In this study, we first analyze a long-term data set on reproductive and vegetative growth of Quercus ilex in a mediterranean woodland in order to identify the main environmental drivers of interannual variation in flower and seed production and contrast the impact of climate vs. adaptive factors as main causes of masting. Second, we conducted an experiment of rainfall exclusion to evaluate the effects of an increasing drought (simulating predictions of global change models) on both reproductive processes. The annual seed crop was always affected by environmental factors related to the precipitation pattern, these abiotic factors disrupting the fruiting process at different periods of time. Seed production was strongly dependent upon water availability for the plant at initial (spring) and advanced (summer) stages of the acorn maturation cycle, whereas the final step of seed development was negatively affected by the frequency of torrential-rain events. We also found clear evidence that seed masting in the study species is not only regulated by selective endogenous rhythms, but is mainly a physiological response to the variable environment. Our results from the rainfall exclusion experiment corroborated the conclusions obtained from the 26-year fruiting record and demonstrated that the high interannual variation in seed crop was mainly determined by the success in seed development rather than by the flowering effort. Under a global change scenario, it could be expected that the drier conditions predicted by climate models reinforce the negative effects of summer drought on seed production, leading to negative consequences for tree recruitment and forest dynamics.

Summer and winter sensitivity of leaves and xylem to minimum freezing temperatures: a comparison of co-occurring Mediterranean oaks that differ in leaf lifespan.

Freezing sensitivity of leaves and xylem was examined in four co-occurring Mediterranean oaks (Quercus spp.) grown in a common garden to determine whether freezing responses of leaves and xylem were coordinated and could be predicted by leaf lifespan. Freezing-induced embolism and loss of photosynthetic function were measured after overnight exposure to a range of subzero temperatures in both summer and winter. Both measures were found to be dependent on minimum freezing temperature and were correlated with leaf lifespan and vessel diameter. The dependence of xylem embolism on minimum freezing temperature may result from the decline in water potential with ice temperature that influences the redistribution of water during freezing and leads to an increase in xylem tension. Winter acclimatization had a relatively small effect on the vulnerability to freezing-induced embolism, although leaf photosynthetic function showed a strong acclimatization response, particularly in the two evergreen species. Quercus ilex, the species with the longest leaf lifespan and narrowest vessel diameters, showed the highest freezing tolerance. This helps explain its ability to inhabit a broad range throughout the Mediterranean region. By contrast, the inability of the deciduous oaks to maintain photosynthetic and vascular function throughout the winter indicates a competitive disadvantage that may prevent them from expanding their ranges.

Europe-wide reduction in primary productivity caused by the heat and drought in 2003.

Future climate warming is expected to enhance plant growth in temperate ecosystems and to increase carbon sequestration. But although severe regional heatwaves may become more frequent in a changing climate, their impact on terrestrial carbon cycling is unclear. Here we report measurements of ecosystem carbon dioxide fluxes, remotely sensed radiation absorbed by plants, and country-level crop yields taken during the European heatwave in 2003. We use a terrestrial biosphere simulation model to assess continental-scale changes in primary productivity during 2003, and their consequences for the net carbon balance. We estimate a 30 per cent reduction in gross primary productivity over Europe, which resulted in a strong anomalous net source of carbon dioxide (0.5 Pg C yr(-1)) to the atmosphere and reversed the effect of four years of net ecosystem carbon sequestration. Our results suggest that productivity reduction in eastern and western Europe can be explained by rainfall deficit and extreme summer heat, respectively. We also find that ecosystem respiration decreased together with gross primary productivity, rather than accelerating with the temperature rise. Model results, corroborated by historical records of crop yields, suggest that such a reduction in Europe's primary productivity is unprecedented during the last century. An increase in future drought events could turn temperate ecosystems into carbon sources, contributing to positive carbon-climate feedbacks already anticipated in the tropics and at high latitudes.

Effect of elevated CO2 on monoterpene emission of young Quercus ilex trees and its relation to structural and ecophysiological parameters.

We investigated growth, leaf monoterpene emission, gas exchange, leaf structure and leaf chemical composition of 1-year-old Quercus ilex L. seedlings grown in ambient (350 microl l(-1)) and elevated (700 microl l(-1)) CO2 concentrations ([CO2]). Monoterpene emission and gas exchange were determined at constant temperature and irradiance (25 degrees C and 1000 micromol m(-2) s(-1) of photosynthetically active radiation) at an assay [CO2] of 350 or 700 microl l(-1). Measurements were made on intact shoots after the end of the growing season between mid-October and mid-February. On average, plants grown in elevated [CO2] had significantly increased foliage biomass (about 50%). Leaves in the elevated [CO2] treatment were significantly thicker and had significantly higher concentrations of cellulose and lignin and significantly lower concentrations of nitrogen and minerals than leaves in the ambient [CO2] treatment. Leaf dry matter density and leaf concentrations of starch, soluble sugars, lipids and hemi-cellulose were not significantly affected by growth in elevated [CO2]. Monoterpene emissions of seedlings were significantly increased by elevated [CO2] but were insensitive to short-term changes in assay [CO2]. On average, plants grown in elevated [CO2] had 1.8-fold higher monoterpene emissions irrespective of the assay [CO2]. Conversely, assay [CO2] rapidly affected photosynthetic rate, but there was no apparent long-term acclimation of photosynthesis to growth in elevated [CO2]. Regardless of growth [CO2], photosynthetic rates of all plants almost doubled when the assay [CO2] was switched from 350 to 700 microl l(-1). At the same assay [CO2], mean photosynthetic rates of seedlings in the two growth CO2 treatments were similar. The percentage of assimilated carbon lost as monoterpenes was not significantly altered by CO2 enrichment. Leaf emission rates were correlated with leaf thickness, leaf concentrations of cellulose, lignin and nitrogen, and total plant leaf area. In all plants, monoterpene emissions strongly declined during the winter independently of CO2 treatment. The results are discussed in the context of the acquisition and allocation of resources by Q. ilex seedlings and evaluated in terms of emission predictions.

Within-tree variation in transpiration in isolated evergreen oak trees: evidence in support of the pipe model theory.

Within-tree variation in sap flow density (SFD) was measured in two isolated evergreen oak (Quercus ilex L.) trees growing in an oak savannah (dehesa) in southwest Spain. Sap flow was estimated by the constant heating method. Three sensors were installed in the trunk of each tree in three orientations: northeast (NE), northwest (NW) and south (S). Sap flow density was monitored continuously from May 18 to September 27, 1993. Daily values of SFD ranged between 500 and 4500 mm3 mm-2 day-1. There were significant differences in SFD between orientations; SFD was higher in the NE and NW orientations than in the S orientation. These differences were noted on both a daily and seasonal time scale, and were less pronounced on cloudy days and at the end of the drought period, when SFD was relatively low. Our results support the idea that branches of trees can be viewed as a collection of small independent plants.

Internal remobilization of carbohydrates, lipids, nitrogen and phosphorus in the Mediterranean evergreen oak Quercus ilex.

Remobilization of internal resources is an important mechanism enabling plants to be partly independent of external nutrient availability. We assessed resource remobilization during the growing period in woody and foliar tissues of leafy branches of mature evergreen Mediterranean oak (Quercus ilex L.) at three field sites. We compared nonstructural carbohydrates, lipids, nitrogen and phosphorus pools in leaves and stems before bud burst (March) and at the end of the growing period (July). We also experimentally defoliated leafy branches to determine the storage function of old leaves. Changes in pools of carbon compounds in leaves and stems during spring and in response to defoliation indicated that foliar and woody tissues could provide carbon to support shoot growth. Independently of stem age, soluble sugar and lipid pools decreased significantly during spring. Changes in leaf pools between March and July involved all compounds measured except starch and were accompanied by a 5% decrease in mean leaf biomass. During the same period, 15% of the nitrogen and 25% of the phosphorus were removed from leaves. In contrast, woody tissues did not remobilize nitrogen or phosphorus. Our results support earlier hypotheses that leaves of evergreen species have a primary role in resource remobilization.

Between-tree variations in leaf δ13C of Quercus pubescens and Quercus ilex among Mediterranean habitats with different water availability.

In this study, sun leaf carbon isotope composition (δ13C) of two co-occurring woody Mediterranean species (Quercus pubescens Willd., a deciduous oak, and Q. ilex L., an evergreen one) was investigated on four sites with different water availability. The total range of δ13C values was 4.4 and 3.1‰ for Q. pubescens and Q. ilex respectively. The intra-site variability was about 3‰. Total mean per species was equal. There were significant differences among sites, but at each site means of δ13C were not significantly different between species. A simple physiological model predicts no difference in intrinsic water-use efficiency (WUEi) between evergreen and deciduous oaks. The relationship between site means of δ13C and water parameters suggests that there is a leaf functional adjustment with respect to available water resource. No correlation was found between δ13C and the contents of any mass-based biochemical constituent. Nevertheless there was a significant correlation between δ13C and leaf mass per area of Q. ilex. For both species, there is also a positive correlation between leaf δ13C and individual crown area, i.e. a structural characteristic at tree level. Causal relations between δ13C and plant-environment interactions are discussed.

Water balance and pattern of root water uptake by a Quercus coccifera L. evergreen srub.

The water balance of a Quercus coccifera evergreen scrub was studied over 7 consecutive years. This scrub grows on hard limestone. Soil water content was measured with a neutron meter. Calibration curves were calculated from (1) the thermal neutron macroscopic cross-sections of soil (<2-mm fraction) and rock samples, and (2) the profile of wet bulk density measured with a subsurface gamma-ray gauge. The annual and seasonal patterns of actual evapotranspiration and of deep drainage were calculated using field-measured drainage characteristics. The soil water content data were used to compute water uptake rates and pattern for the root zone over a 4-month drying period. The 906 mm of mean annual precipitation yielded 603 mm of actual evapotranspiration (AET) and 296 mm of drainage. No drainage occured with precipitation less than 578 mm. The average AET values for the months from April to September were 57, 74, 89, 96, 70, and 42 mm respectively. It was found that Quercus coccifera consumed considerable quantities of water from the soil-rock complex. Roots could extract 270 mm of water in the first 470 cm of soil. The results showed a gradual downward shift of the zone of maximum root water uptake as the soil dried.