Severe droughts have caused widespread tree mortality across many forest biomes with profound effects on the function of ecosystems and carbon balance. Climate change is expected to intensify regional-scale droughts, focusing attention on the physiological basis of drought-induced tree mortality. Recent work has shown that catastrophic failure of the plant hydraulic system is a principal mechanism involved in extensive crown death and tree mortality during drought, but the multi-dimensional response of trees to desiccation is complex. Here we focus on the current understanding of tree hydraulic performance under drought, the identification of physiological thresholds that precipitate mortality and the mechanisms of recovery after drought. Building on this, we discuss the potential application of hydraulic thresholds to process-based models that predict mortality.

Contents Summary I. Introduction II. Comparison of various definitions and measurement techniques of minimum conductance III. Cuticular conductance IV. Contribution of stomata V. Environmental and ecological variation in minimum conductance VI. Use of minimum conductance in models VII. Conclusions Acknowledgements References SUMMARY: When the rate of photosynthesis is greatly diminished, such as during severe drought, extreme temperature or low light, it seems advantageous for plants to close stomata and completely halt water loss. However, water loss continues through the cuticle and incompletely closed stomata, together constituting the leaf minimum conductance (g ). In this review, we critically evaluate the sources of variation in g , quantitatively compare various methods for its estimation, and illustrate the role of g in models of leaf gas exchange. A literature compilation of g as measured by the weight loss of detached leaves is presented, which shows much variation in this trait, which is not clearly related to species groups, climate of origin or leaf type. Much evidence points to the idea that g is highly responsive to the growing conditions of the plant, including soil water availability, temperature and air humidity - as we further demonstrate with two case studies. We pay special attention to the role of the minimum conductance in the Ball-Berry model of stomatal conductance, and caution against the usual regression-based method for its estimation. The synthesis presented here provides guidelines for the use of g in ecosystem models, and points to clear research gaps for this drought tolerance trait.

Heatwaves are likely to increase in frequency and intensity with climate change, which may impair tree function and forest C uptake. However, we have little information regarding the impact of extreme heatwaves on the physiological performance of large trees in the field. Here, we grew Eucalyptus parramattensis trees for 1 year with experimental warming (+3°C) in a field setting, until they were greater than 6 m tall. We withheld irrigation for 1 month to dry the surface soils and then implemented an extreme heatwave treatment of 4 consecutive days with air temperatures exceeding 43°C, while monitoring whole-canopy exchange of CO and H O, leaf temperatures, leaf thermal tolerance, and leaf and branch hydraulic status. The heatwave reduced midday canopy photosynthesis to near zero but transpiration persisted, maintaining canopy cooling. A standard photosynthetic model was unable to capture the observed decoupling between photosynthesis and transpiration at high temperatures, suggesting that climate models may underestimate a moderating feedback of vegetation on heatwave intensity. The heatwave also triggered a rapid increase in leaf thermal tolerance, such that leaf temperatures observed during the heatwave were maintained within the thermal limits of leaf function. All responses were equivalent for trees with a prior history of ambient and warmed (+3°C) temperatures, indicating that climate warming conferred no added tolerance of heatwaves expected in the future. This coordinated physiological response utilizing latent cooling and adjustment of thermal thresholds has implications for tree tolerance of future climate extremes as well as model predictions of future heatwave intensity at landscape and global scales.

Water stress (WS) slows growth and photosynthesis (A(n)), but most knowledge comes from short-time studies that do not account for longer term acclimation processes that are especially relevant in tree species. Using two Eucalyptus species that contrast in drought tolerance, we induced moderate and severe water deficits by withholding water until stomatal conductance (g(sw)) decreased to two pre-defined values for 24 d, WS was maintained at the target g(sw) for 29 d and then plants were re-watered. Additionally, we developed new equations to simulate the effect on mesophyll conductance (g(m)) of accounting for the resistance to refixation of CO(2). The diffusive limitations to CO(2), dominated by the stomata, were the most important constraints to A(n). Full recovery of A(n) was reached after re-watering, characterized by quick recovery of gm and even higher biochemical capacity, in contrast to the slower recovery of g(sw). The acclimation to long-term WS led to decreased mesophyll and biochemical limitations, in contrast to studies in which stress was imposed more rapidly. Finally, we provide evidence that higher gm under WS contributes to higher intrinsic water-use efficiency (iWUE) and reduces the leaf oxidative stress, highlighting the importance of gm as a target for breeding/genetic engineering.

The aim of this study was to provide new insights into how intraspecific variability in the response of key functional traits to drought dictates the interplay between gas-exchange parameters and the hydraulic architecture of European beech (Fagus sylvatica L.). Considering the relationships between hydraulic and leaf functional traits, we tested whether local adaptation to water stress occurs in this species. To address these objectives, we conducted a glasshouse experiment in which 2-year-old saplings from six beech populations were subjected to different watering treatments. These populations encompassed central and marginal areas of the range, with variation in macro- and microclimatic water availability. The results highlight subtle but significant differences among populations in their functional response to drought. Interpopulation differences in hydraulic traits suggest that vulnerability to cavitation is higher in populations with higher sensitivity to drought. However, there was no clear relationship between variables related to hydraulic efficiency, such as xylem-specific hydraulic conductivity or stomatal conductance, and those that reflect resistance to xylem cavitation (i.e., Ψ(12), the water potential corresponding to a 12% loss of stem hydraulic conductivity). The results suggest that while a trade-off between photosynthetic capacity at the leaf level and hydraulic function of xylem could be established across populations, it functions independently of the compromise between safety and efficiency of the hydraulic system with regard to water use at the interpopulation level.

The ability of P. canariensis to inhabit a wide range of ecosystems seemed to be associated with high phenotypic plasticity and some degree of local adaptations of xylem and leaf traits. Resistance to cavitation conferred adaptive potential for this species to adapt successfully to xeric conditions.

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Dutch elm disease (DED) is a vascular wilt disease that causes the occlusion and cavitation of xylem vessels. Therefore, it is hypothesized that those elms that are less vulnerable to cavitation by drought might be more resistant to DED. To test this hypothesis, the relationship between xylem vulnerability to cavitation and susceptibility to DED was examined in progenies of crosses between susceptible and resistant individuals of Ulmus minor. Hydraulic conductivity and xylem vulnerability curves were evaluated and anatomical features such as vessel size, length and grouping were measured. Next, elms were inoculated with Ophiostoma novo-ulmi, the cause of DED, and pre-dawn and midday water potentials, stomatal conductance and wilting percentages were assessed. Progenies of R 9 R crosses showed significantly lower mean wilting percentages (30-50%) than the progeny of S 9 S crosses (75%). Fifty percent conductivity loss was reached at c. À1 MPa, pointing out a high vulnerability of this species to drought-induced cavitation. Crown wilting percentage as a result of inoculation and xylem vulnerability to cavitation by water stress did not show any significant correlation. Nevertheless, significant differences in theoretical hydraulic conductivity and vessel size parameters (diameter, length and size distributions) were found among the tested progenies. Susceptible trees had significantly wider and longer vessels. Xylem structure of resistant elms seems to restrict pathogen spread rather than prevent cavitation.