Summary1. Studying the effects of climate or weather extremes such as drought and heat waves on biodiversity and ecosystem functions is one of the most important facets of climate change research. In particular, primary production is amounting to the common currency in field experiments world-wide. Rarely, however, are multiple ecosystem functions measured in a single study in order to address general patterns across different categories of responses and to analyse effects of climate extremes on various ecosystem functions. 2. We set up a long-term field experiment, where we applied recurrent severe drought events annually for five consecutive years to constructed grassland communities in central Europe. The 32 response parameters studied were closely related to ecosystem functions such as primary production, nutrient cycling, carbon fixation, water regulation and community stability. 3. Surprisingly, in the face of severe drought, above-and below-ground primary production of plants remained stable across all years of the drought manipulation.4. Yet, severe drought significantly reduced below-ground performance of microbes in soil indicated by reduced soil respiration, microbial biomass and cellulose decomposition rates as well as mycorrhization rates. Furthermore, drought reduced leaf water potential, leaf gas exchange and leaf protein content, while increasing maximum uptake capacity, leaf carbon isotope signature and leaf carbohydrate content. With regard to community stability, drought induced complementary plantplant interactions and shifts in flower phenology, and decreased invasibility of plant communities and primary consumer abundance. 5. Synthesis. Our results provide the first field-based experimental evidence that climate extremes initiate plant physiological processes, which may serve to regulate ecosystem productivity. Journal of Ecology 2011Ecology , 99, 689-702 doi: 10.1111Ecology /j.1365Ecology -2745Ecology .2011 of synergisms or decoupling of biogeochemical processes, and of fundamental response dynamics to drought at the ecosystem level including potential tipping points and thresholds of regime shift. Future work is needed to elucidate the role of biodiversity and of biotic interactions in modulating ecosystem response to climate extremes.

The aim of this study was to identify the sources and depth of water uptake by 15-years old Quercus suber L. trees in southern Portugal under a Mediterranean climate, measuring d 18 O and dD in the soil-plantatmosphere continuum. Evidence for hydraulic lift was substantiated by the daily fluctuations observed in Y s at 0.4 and 1 m depth and supported by similar d 18 O values found in tree xylem sap, soil water in the rhizosphere and groundwater. From 0.25 m down to a depth of 1 m, dD trends differed according to vegetation type, showing a more depleted value in soil water collected under the evergreen trees ()47&) than under dead grasses ()35&). The hypothesis of a fractionation process occurring in the soil due to diffusion of water vapour in the dry soil is proposed to explain the more depleted soil dD signature observed under trees. Hydraulically lifted water was estimated to account for 17-81% of the water used during the following day by tree transpiration at the peak of the drought season, i.e., 0.

Container-grown seedlings of Acacia tortilis Forsk. Hayne and A. xanthophloea Benth. were watered either every other day (well watered) or every 7 days (water-stressed) for 1 year in a greenhouse. Total plant dry mass (T(dm)), carbon allocation and water relations were measured monthly. Differences in leaf area (LA) accounted for differences in T(dm) between the species, and between well-watered and water-stressed plants. Reduction in LA as a result of water stress was attributed to reduced leaf initiation, leaf growth rate and leaf size. When subjected to prolonged water stress, Acacia xanthophloea wilted more rapidly than A. tortilis and, unlike A. tortilis, lost both leaves and branches. These differences between species were attributed to differences in the allocation of carbon between leaves and roots and in the ability to adjust osmotically. Rapid recovery in A. xanthophloea following the prolonged water-stress treatment was attributed to high cell wall elasticity. Previous exposure to water stress contributed to water-stress resistance and improved recovery after stress.

This study details the physiological responses of cork oak (Quercus suber L.) to manipulated water inputs. Treatments named as dry, ambient and wet, which received 80, 100 and 120% of the annual precipitation, respectively, were applied to a Mediterranean woodland in southern Portugal. Tree ecophysiology and growth were monitored from 2003 to 2005. The impacts of the water manipulation were primarily observed in tree transpiration, especially during summer drought. Rainfall exclusion reduced the annual stand canopy transpiration by 10% over the 2-year study period, while irrigation increased it by 11%. The accumulated tree transpiration matched precipitation in spring 2004 and 2005 at the stand level, suggesting that cork oak trees rely on precipitation water sources during the peak of the growing season. However, during the summer droughts, groundwater was the main water source for trees. Despite the significant differences in soil water content and tree transpiration, no treatment effects could be detected in leaf water potential and leaf gas exchange, except for a single event after spring irrigations in the very dry year 2005. These irrigations were intentionally delayed to reduce dry spell duration during the peak of tree growing season. They resulted in an acute positive physiological response of trees from the wet treatment one week after the last irrigation event leading to a 32% raise of stem diameter increment the following months. Our results suggest that in a semi-arid environment precipitation changes in spring (amount and timing) have a stronger impact on cork oak physiology and growth than an overall change in the total annual precipitation. The extreme drought of 2005 had a negative impact on tree growth. The annual increment of tree trunk diameter in the ambient and dry treatments was reduced, while it increased for trees from the wet treatment. Water shortage also significantly reduced leaf area. The latter dropped by 10.4% in response to the extreme drought of 2005 in trees from the ambient treatment. The reduction was less pronounced in trees of the wet treatment (−7.6%), and more pronounced in trees of the dry treatment (−14.7%). Cork oak showed high resiliency to inter-annual precipitation variability. The annual accumulated tree transpiration, the minimum midday leaf water potential and the absolute amount of groundwater used Abbreviations: (A), carbon assimilation; (Amax), maximum carbon assimilation; (CV), crown volume; (DBH), diameter at breast height; (DBH inc), trunk diameter increment; (E), sap flux, tree transpiration; (E/DBH), normalized tree transpiration; (ET), potential evapotranspiration; (gs), stomatal conductance; (gsmax), maximum stomatal conductance; (Gt), whole tree specific hydraulic conductance; (LAD), leaf area density; (LAI), leaf area index; (PAR), photosynthetically active radiation; (PCA), projected crown area; (SLA), specific leaf area; (SWC), soil water content; (T), air temperature; (VPD), vapour pressure deficit; («), leaf water potential; (« md), midday leaf wat...

Studies were conducted to examine changes in soil (Ys) and plant water status during summer in a 16-year old Quercus suber plantation in southern Portugal. . Intensive measurements were conducted on five trees with mean height and DBH of 5.3 m and 11.6 cm, respectively, growing at close proximity to each other. Weather conditions and soil water potential (Ys) at the rhizosphere of each of the trees measured at 0.3 and 1 m soil depth were continuously monitored. Predawn (Ypd) and midday (Ymd) leaf water potentials were determined every month. Soil and plant samples were also collected in June and September from different locations within the study site for d 18 O isotope composition analysis. Pressure-volume (p-v) curves were constructed from plant shoots at different times during the vegetative period to determine osmotic potential at full saturation (P 100 ), water potential at turgor loss point (Y tlp ), relative water content at turgor loss point (R* tlp ) and bulk modulus of elasticity (e). Significant P < 0.05 decline in Ys occurred between May and September, the lowest value recorded being -2.0 MPa. Decline in soil moisture affected tree water status, but decline in leaf water potential varied significantly (P < 0.05) among the trees. At the end of summer drought, lowest Ypd measured was -1.7 MPa while the highest measured during this time was -0.8 MPa. Differences among trees were attributed to differences in rooting depth, as shown by regression analysis of 18 O isotopes. Radial stem growth ceased when Ys within the upper 0.3 m depth approached -1.5 MPa. The upper soil layers contributed approximately 33% of the total tree water requirement, between spring and mid summer when drought was experienced by trees. Deep soil layers however, supplied most of the water required during drought and no growth was recorded during this time. Stressed trees increased solute concentration of their tissues by a Magnitude of 0.7 MPa while bulk tissue elastic modulus increased by about 17 MPa. The study emphasizes the significance of roots as determinants of tree productivity and survival in the Mediterranean ecosystems.

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We simulated the effect of prolonged dry summer periods by lowering the water table on three manipulation plots (D(1-3)) in a minerotrophic fen in southeastern Germany in three years (2006-2008). The water table at this site was lowered by drainage and by excluding precipitation; three nonmanipulated control plots (C(1-3)) served as a reference. We found no significant differences in soil respiration (R(Soil)), gross primary production (GPP), or aboveground respiration (R(AG)) between the C(1-3) and D(1-3) plots in any of the measurement years. The water table on the control plots was naturally low, with a median water table (2006-2008) of 8 cm below the surface, and even lower during summer when respiratory activity was highest, with median values (C(1-3)) between 11 and 19 cm below the surface. If it is assumed that oxygen availability in the uppermost 10 cm was not limited by the location of the water table, manipulative lowering of the water table most likely increased oxygen availability only in deeper peat layers where we expect R(Soil) to be limited by poor substrate quality rather than anoxia. This could explain the lack of a manipulation effect. In a second approach, we estimated the influence of the water table on R(Soil) irrespective of treatment. The results showed a significant correlation between R(Soil) and water table, but with R(Soil) decreasing at lower water tables rather than increasing. We thus conclude that decomposition in the litter layer is not limited by waterlogging in summer, and deeper peat layers bear no significant decomposition potential due to poor substrate quality. Consequently, we do not expect enhanced C losses from this site due to increasing frequency of dry summers. Assimilation and respiration of aboveground vegetation were not affected by water table fluctuations between 10 and >60 cm depth, indicating the lack of stress resulting from either anoxia (high water table) or drought (low water table).