-The extreme drought event that occurred in Western Europe during 2003 highlighted the need to understand the key processes that may allow trees and stands to overcome such severe water shortages. We therefore reviewed the current knowledge available about such processes. First, impact of drought on exchanges at soil-root and canopy-atmosphere interfaces are presented and illustrated with examples from water and CO 2 flux measurements. The decline in transpiration and water uptake and in net carbon assimilation due to stomatal closure has been quantified and modelled. The resulting models were used to compute water balance at stand level basing on the 2003 climate in nine European forest sites from the CARBOEUROPE network. Estimates of soil water deficit were produced and provided a quantitative index of soil water shortage and therefore of the intensity of drought stress experienced by trees during summer 2003. In a second section, we review the irreversible damage that could be imposed on water transfer within trees and particularly within xylem. A special attention was paid to the inter-specific variability of these properties among a wide range of tree species. The inter-specific diversity of hydraulic and stomatal responses to soil water deficit is also discussed as it might reflect a large diversity in traits potentially related to drought tolerance. Finally, tree decline and mortality due to recurrent or extreme drought events are discussed on the basis of a literature review and recent decline studies. The potential involvement of hydraulic dysfunctions or of deficits in carbon storage as causes for the observed long term (several years) decline of tree growth and development and for the onset of tree dieback is discussed. As an example, the starch content in stem tissues recorded at the end of the 2003's summer was used to predict crown conditions of oak trees during the following spring: low starch contents were correlated with large twig and branch decline in the crown of trees. drought / water balance / time lag effect / hydraulic properties / dieback Résumé -Arbres et peuplements forestiers tempérés soumis à sécheresse : une revue des réponses écophysiologiques, des processus d'adaptation et des conséquences à long terme. La sécheresse exceptionnelle de 2003 a été l'occasion de faire le point de nos connaissances sur les mécanismes écophysiologiques permettant aux arbres de traverser un tel évènement climatique extrême. L'analyse a été conduite à l'échelle de l'arbre et du peuplement, tandis que l'intensité de la sécheresse a été quantifiée à l'aide d'un calcul de bilan hydrique sur neuf sites forestiers européens contrastés du réseau CARBOEUROPE. Le rôle du couvert dans les échanges avec l'atmosphère est rappelé puis intégré dans l'analyse des réductions de bilan d'eau et de carbone en 2003 dus à la régulation stomatique. Les caractéristiques du complexe sol-racine, important à la fois pour l'accès à la ressource et à l'efficience de son absorption, constituent un des premiers traits d'ada...

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Leaf gas exchange rates, predawn Ψ and daily minimum Ψ leaf water potentials were measured during a wet-to-dry season transition in pioneer (Jacaranda copaia, Goupia glabra andCarapa guianensis) and late stage rainforest tree species (Dicorynia guianensis andEperua falcata) growing in common conditions in artificial stands in French Guiana. Carbon isotope discrimination (Δ) was assessed by measuring the stable carbon isotope composition of the cellulose fraction of wood cores. The Δ values were 2.7‰ higher in the pioneer species than in the late stage species. The calculated time integratedC values derived from the Δ values averaged 281 μmol mol in the pioneers and 240 μmol mol in the late stage species. The corresponding time-integrated values of intrinsinc water-use efficiency [ratio CO assimilation rate (A)/leaf conductance (g)] ranged from 37 to 47 mmol mol in the pioneers and the values were 64 and 74 mmol mol for the two late stage species. The high Δ values were associated-at least inJ. copaia-with high maximumg values and with high plant intrinsinc specific hydraulic conductance [C≔g/(Ψ-Ψ], which could reflect a high competitive ability for water and nutrient uptake in the absence of soil drought in the pioneers. A further clear discriminating trait of the pioneer species was the very sensitive stomatal response to drought in the soil, which might be associated with a high vulnerability to cavitation in these species. From a methodological point of view, the results show the relevance of Δ for distinguishing ecophysiological functional types among rainforest trees.

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.

Mediterranean tree species experience unpredictable climate environments and severe summer droughts and they may be impaired by the trend of decline in precipitation projected as a consequence of global climate change. The response of Quercus ilex to drought was studied by measuring hydraulic traits of trees growing in a mature forest subjected to partial throughfall exclusion for 6 years. We measured hydraulic conductivity, xylem vulnerability to embolism, and anatomical features in branches and roots. Xylem vulnerability to embolism was higher in the dry treatment than in the control treatment, P₅₀ of branches was on average -3.88 +/- 0.80 MPa for the control treatment compared with -3.41 +/- 0.80 MPa for the dry treatment, but the difference was not statistically significant. A similar difference between treatments was observed for roots, which exhibited lower P₅₀ values. This change of xylem vulnerability to embolism was not linked to modification of the hydraulic conductivity or vessel anatomy, which remained unaffected by the throughfall exclusion treatment. The xylem density of branches was lower in the dry treatment. The hydraulic conductivity was correlated with the mean vessel diameter of xylem, but the P₅₀ was not. The main response of trees from the dry treatment to reduced water availability appeared to be a reduction in the transpiring leaf area, which resulted in significantly increased leaf-specific conductivity.

Xylem vulnerability to cavitation is a key parameter in understanding drought resistance of trees. We determined the xylem water pressure causing 50% loss of hydraulic conductivity (P(50)), a proxy of vulnerability to cavitation, and we evaluated the variability of this trait at tree and population levels for Fagus sylvatica. We checked for the effects of light on vulnerability to cavitation of stem segments together with a time series variation of P(50). Full sunlight-exposed stem segments were less vulnerable to cavitation than shade-exposed ones. We found no clear seasonal change of P(50), suggesting that this trait was designed for a restricted period. P(50) varied for populations settled along a latitudinal gradient, but not for those sampled along an altitudinal gradient. Moreover, mountainside exposure seemed to play a major role in the vulnerability to cavitation of beech populations, as we observed the differences along north-facing sides but not on south-facing sides. Unexpectedly, both north-facing mountainside and northern populations appeared less vulnerable than those grown on the southern mountainside or in the South of France. These results on beech populations were discussed with respect to the results at within-tree level.

Controlled, localized heating was applied to the trunks of young Aleppo pines (Pinushalepensis Mill.) during the winter. Three levels of burning were applied to the circumference of the trunks (zero, partial, or total), and the effects on tree physiology and growth were monitored during the 7 months following heating. Effects of the heat treatment on tree physiology were assessed by measuring predawn water potential, sap flux density, microvariations in trunk diameter, electrical resistance of the cambium, and gas exchanges from the needles (photosynthesis and stomatal conductance). Morphological response measurements included height and radial growth of the trees and morphological characteristics of the needles. The amount of chlorotic foliage was also measured. Aleppo pine was highly resistant to trunk injury, surviving when 80% of the trunk circumference was destroyed by fire. The behaviour of partially burnt pines was not significantly different from that of controls, for all variables studied. Trees whose trunks were totally burnt died from 1 to 5 months after treatment. Sap flux density of totally burnt trees was already significantly lower than that of other trees in the first week following the treatment. A decrease in net photosynthesis and stomatal conductance in the still surviving totally burnt trees was also recorded during the first measurements made in July. Treatment differences in predawn water potential, microvariations in trunk diameter, and electrical resistance of the cambium only appeared a few days before death of the totally burnt trees.

We studied hydraulic traits of young plants of the Mediterranean cedar species Cedrus atlantica (Endl.) G. Manetti ex Carrière (Luberon, France), C. brevifolia (Hook. f.) Henry (Cyprus), C. libani A. Rich (Hadeth El Jebbe, Lebanon) and C. libani (Armut Alani, Turkey). With an optimum water supply, no major differences were observed among species or provenances in either stem hydraulic conductivity (Ks) or leaf specific conductivity (Kl) measured on the main shoot. A moderate soil drought applied for 10 weeks induced marked acclimation through a reduction in Ks, particularly in the Lebanese provenance of C. libani, and a decrease in tracheid lumen size in all species. Cedrus atlantica, which had the smallest tracheids, was the species most vulnerable to embolism: a 50% loss in hydraulic conductivity (PsiPLC50) occurred at a water potential of -4.4 MPa in the well-watered treatment, and at -6.0 MPa in the moderate drought treatment. In the other species, PsiPLC50 was unaffected by moderate soil drought, and only declined sharply at water potentials between -6.4 and -7.5 MPa in both irrigation treatments. During severe drought, Ks of twigs and stomatal conductance (g(s)) were measured simultaneously as leaf water potential declined. For all species, lower vulnerability to embolism based on loss of Ks was recorded on current-year twigs. The threshold for stomatal closure (10% of maximum g(s)) was reached at a predawn water potential (Psi(pd)) of -2.5 MPa in C. atlantica (Luberon) and at -3.1 MPa in C. libani (Lebanon), whereas the other provenance and species had intermediate Psi(pd) values. Cedrus brevifolia, with a Psi(pd) (-3.0 MPa) close to that of C. libani (Lebanon), had the highest stomatal conductance of the study species. The importance of a margin of safety between water potential causing stomatal closure and that causing xylem embolism induction is discussed.