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.

Mediterranean evergreen oaks have to survive a long summer drought. Roots may play a relevant role under these conditions. We studied their structure and function in a mature Quercus suber L. tree in central Portugal. The root system was mapped till the lowest water table level (4.5 m depth). Xylem anatomy was analyzed in a vertical profile belowground. Sap flow was continuously monitored for 1.5 yrs in the stem and roots of this intensively studied tree (heat field deformation method) and in the stem of four trees (Granier method), in relation to environmental variables and predawn leaf water potential. The sources of water uptake were assessed by stable isotope analyses in summer. Results showed a dimorphic root system with a network of superficial roots linked to sinker roots, and a taproot diverting into tangles of deep fine roots submerged for long periods, with parenchyma aerenchyma. Transpiration was not restricted in summer due to root access to groundwater. The isotopic d 18 O signature of twig xylem water was similar to that of groundwater in the dry season. Two functional types of superficial roots were identified: shallow connected and deep connected roots. A modeling approach was built considering that each superficial root was linked to a sinker, with part of the root deep connected (between the stem and the sinker) and part shallow connected (between the sinker and topsoil). This conceptual framework simulated tree stem sap flow from root sap flow with a high efficiency (R 2 = 0.85) in four plot trees. On an annual basis, soil water and groundwater contributions were 69.5% and 30.5% of stem flow, respectively. Annual hydraulic lift and hydraulic descent were 0.9% and 37.0% of stem flow, respectively. The trees maximize the exploitation of the environmental resources by using the topsoil water during most of the year, and groundwater together with hydraulic lift (nutrient supply) in the dry summer. This study shows that a dimorphic root system, with roots reaching groundwater, is an efficient strategy of Q. suber trees to cope with seasonal drought. Knowledge of the functional behavior of Q. suber trees under shallow water table conditions may contribute to the definition of better adapted management practices and to anticipate their responses to climate change.

Mediterranean evergreen oak woodlands of southern Portugal (montados) are savannah-type ecosystems with a widely sparse tree cover, over extensive grassland. Therefore, ecosystem water fluxes derive from two quite differentiated sources: the trees and the pasture. Partitioning of fluxes according to these different sources is necessary to quantify overall ecosystem water losses as well as to improve knowledge on its functional behaviour. In southern Iberia, these woodlands are subjected to recurrent droughts. Therefore, reaction/resilience to water stress becomes an essential feature of vegetation on these ecosystems. Long-term tree transpiration was recorded for 6 years from a sample of holm oak (Quercus ilex ssp. rotundifolia) trees, using the Granier sap flow method. Ecosystem transpiration was measured by the eddy covariance technique for an 11-month period (February to December 2005), partly coincident with a drought year. Pasture transpiration was estimated as the difference between ecosystem (eddy covariance) and tree (sap flow) transpiration. Pasture transpiration stopped during the summer, when the surface soil dried up. In the other seasons, pasture transpiration showed a strong dependence on rainfall occurrence and on top soil water. Conversely, trees were able to maintain transpiration throughout the summer due to the deep root access to groundwater. Q. ilex trees showed a high resilience to both seasonal and annual drought. Tree transpiration represented more than half of ecosystem transpiration, in spite of the low tree density (30 trees ha À1) and crown cover fraction (21%). Tree evapotranspiration was dominated by transpiration (76%), and interception loss represented only 24% of overall tree evaporation.

Quercus suber and Quercus ilex trees are major elements of Mediterranean landscapes, which are threatened by the increasing water deficits related to climate change. To contribute to the understanding of the capacity of these oaks to withstand severe drought we assessed the vulnerability to xylem embolism and the anatomical traits in current-year shoots. Data were collected in mature trees at two sites, in central/coastal and southern/inland Portugal. In situ safety margins to hydraulic failure were evaluated from long-term predawn and midday leaf water potential records. Results showed that xylem vulnerability to embolism was similar in Q. ilex and Q. suber. The 50% loss in hydraulic conductivity (Ψxyl,50PLC) was observed at xylem water potentials of −2.9 and −3.2 MPa in shoots of Q. suber and Q. ilex, respectively. Values of mean vessel diameter of Q. suber shoots at both sites suggest an intra-species adaptation to the local water availability, with larger vessels at the more mesic site. In situ hydraulic safety margins observed in shoots showed that, even during the driest periods, both oaks lived comfortably above the most critical embolism thresholds. However, the hydraulic safety margins were narrower in the driest site. Results are relevant to the understanding of survival, growth, and functional behaviour of evergreen oaks in Mediterranean climates, under recurrent/seasonal drought conditions

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Abstract:Water is one of the major environmental factors limiting plant growth and survival in the Mediterranean region. Quercus suber L. woodlands occupy vast areas in the Iberian Peninsula, frequently under shallow water table conditions. The relative magnitude of soil and groundwater uptake to supply transpiration is not easy to evaluate under these circumstances. We recently developed a conceptual framework for the functioning of the root system in Q. suber that simulates well tree transpiration, based on two types of root behaviour: shallow connected and deep connected. Although this significantly improved knowledge on the functional traits of Mediterranean Q. suber, the approach has the limitation of requiring root sap flow data, which are seldom available. In this work, we present alternative methodologies to assess if trees are connected to groundwater and to estimate the soil and groundwater contributions to tree transpiration. We provide evidence on the tree unrestricted access to groundwater solely based on meteorological, stem sap flow and leaf water potential data. Using a soil mass balance approach, we estimated the yearly soil and groundwater contributions to tree transpiration: 69.7% and 30.3%, respectively. Groundwater uptake became dominant in the dry summer: 73.2% of tree transpiration. Results reproduce extremely well those derived from root modelling. Because of its simplicity both in formulation and data requirements, our approach is potentially liable to be adapted to other groundwaterdependent Mediterranean oak sites, where interactions between land use and water resources may be relevant.

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In this study we investigated the involvement of inflammatory cells in the pleural accumulation of eosinophils induced by lipopolysaccharide (LPS). Intrathoracic (i.t.) injection of LPS (250 ng/cavity) into rats induced a significant eosinophil accumulation that developed within 24 h, was maximal at 48 h, and returned to control values within 120 h. This eosinophil influx was preceded by a huge neutrophil influx within 4 h and accompanied by a mononuclear cell accumulation between 24 and 48 h. Pretreatment with an antineutrophil monoclonal antibody (RP-3, 2 ml per animal) selectively reduced the number of circulating neutrophils within 8 h but failed to alter the LPS-induced eosinophilia. Similarly, platelet depletion with an anti-rat platelet antiserum did not alter the LPS-induced eosinophil accumulation. Cyclosporine (50 mg/kg, 12 and 2 h before) partially inhibited (51%) the LPS-induced pleural eosinophilia, whereas the eosinophilia was not changed by prior degranulation of pleural mast cells with polymyxin B (10 micrograms/cavity, 24 h before). Moreover, selective depletion of T lymphocytes using an anti-Thy 1.0 monoclonal antibody significantly inhibited the eosinophilia triggered by LPS. The i.t. injection of liposomes containing dichloromethylene diphosphonate significantly reduced (65%) the number of resident macrophages after 5 days. Under this condition, the eosinophil infiltration induced by LPS was completely inhibited. Accordingly, the i.t. injection of supernatant from macrophage monolayers, obtained from the pleural cavities of LPS-injected rats, into naive recipient animals led to a twofold increase in the number of pleural eosinophils. In conclusion, our data suggest an important role for resident macrophages and T lymphocytes in the eosinophil accumulation induced by LPS.