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.

Summary• Hydraulic dysfunction in leaves determines key aspects of whole-plant responses to water stress; however, our understanding of the physiology of hydraulic dysfunction and its relationships to leaf structure and ecological strategy remains incomplete.• Here, we studied a morphologically and ecologically diverse sample of angiosperms to test whether the water potential inducing a 50% loss in leaf hydraulic conductance (P50 leaf ) is predicted by properties of leaf xylem relating to water tension-induced conduit collapse. We also assessed the relationships between P50 leaf and other traits considered to reflect drought resistance and ecological strategy.• Across species, P50 leaf was strongly correlated with a theoretical predictor of vulnerability to cell collapse in minor veins (the cubed ratio of the conduit wall thickness to the conduit lumen breadth). P50 leaf was also correlated with mesophyll traits known to be related to drought resistance, but unrelated to traits associated with carbon economy.• Our data indicate a link between the structural mechanics of leaf xylem and hydraulic function under water stress. Although it is possible that collapse may contribute directly to dysfunction, this relationship may also be a secondary product of vascular economics, suggesting that leaf xylem is dimensioned to avoid wall collapse.

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.

Efficient conduction of water inside leaves is essential for leaf function, yet the hydraulic-mediated impact of drought on gas exchange remains poorly understood. Here we examine the decline and subsequent recovery of leaf water potential (Yleaf), leaf hydraulic conductance (Kleaf), and midday transpiration (E) in four temperate woody species exposed to controlled drought conditions ranging from mild to lethal. During drought the vulnerability of Kleaf to declining Yleaf varied greatly among the species sampled. Following drought, plants were rewatered and the rate of E and Kleaf recovery was found to be strongly dependent on the severity of the drought imposed. Gas exchange recovery was strongly correlated with the relatively slow recovery of Kleaf for three of the four species, indicating conformity to a hydraulic-stomatal limitation model of plant recovery. However, there was also a shift in the sensitivity of stomata to Yleaf suggesting that the plant hormone abscisic acid may be involved in limiting the rate of stomatal reopening. The level of drought tolerance varied among the four species and was correlated with leaf hydraulic vulnerability. These results suggest that species-specific variation in hydraulic properties plays a fundamental role in steering the dynamic response of plants during recovery.

Plant hydraulic traits capture the impacts of drought stress on plant function, yet vegetation models lack sufficient information regarding trait coordination and variation with climate-of-origin across species. Here, we investigated key hydraulic and carbon economy traits of 12 woody species in Australia from a broad climatic gradient, with the aim of identifying the coordination among these traits and the role of climate in shaping cross-species trait variation. The influence of environmental variation was minimized by a common garden approach, allowing us to factor out the influence of environment on phenotypic variation across species. We found that hydraulic traits (leaf turgor loss point, stomatal sensitivity to drought [P ], xylem vulnerability to cavitation [P ], and branch capacitance [C ]) were highly coordinated across species and strongly related to rainfall and aridity in the species native distributional range. In addition, trade-offs between drought tolerance and plant growth rate were observed across species. Collectively, these results provide critical insight into the coordination among hydraulic traits in modulating drought adaptation and will significantly advance our ability to predict drought vulnerability in these dominant trees species.

Research in plant hydraulics has provided important insights into plant responses to drought and species absolute drought tolerance. However, our ability to predict when plants will die from hydraulic failure under extreme drought is limited by a lack of knowledge with regards to the dynamics of plant desiccation following stomatal closure. Thus, we develop a simple hydraulics model based on branch-level traits that incorporates key aspects of allometry, rates of water loss and resistance to embolism thresholds in order to define species differences in the time it takes plants to desiccate from stomatal closure to lethal levels of drought stress.

The ability of plants to maintain water flow through leaves under water stress-induced tension (assessed as the leaf hydraulic vulnerability; P50(leaf)) is intimately linked with survival. We examined the significance of P50(leaf) as an adaptive trait in influencing the dry-end distributional limits of cool temperate woody angiosperm species. We also examined differences in within-site variability in P50(leaf) between two high-rainfall montane rainforest sites in Tasmania and Peru, respectively. A significant relationship between P50(leaf) and the 5th percentile of mean annual rainfall across each species distribution was found in Tasmania, suggesting that P50(leaf) influences species climatic limits. Furthermore, a strong correlation between P50(leaf) and the minimum rainfall availability was found using five phylogenetically independent species pairs in wet and dry evergreen tree species, suggesting that rainfall is an important selective agent in the evolution of leaf hydraulic vulnerability. Greater within-site variability in P50(leaf) was found among dominant montane rainforest species in Tasmania than in Peru and this result is discussed within the context of differences in spatial and temporal environmental heterogeneity and parochial historical ecology.

The findings indicate that species sorting processes have influenced distributional patterns of P50leaf across the rainfall spectrum, but alternative strategies for dealing with water deficit exist within sites. The strong link to aridity suggests leaf hydraulic vulnerability may influence plant distributions under future climates.