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Plant responses to elevated atmospheric carbon dioxide (eCO₂) have been hypothesized as a key mechanism that may ameliorate the impact of future drought. Yet, despite decades of experiments, the question of whether eCO₂ reduces plant water use, yielding ‘water savings’ that can be used to maintain plant function during periods of water stress, remains unresolved. In this Viewpoint, we identify the experimental challenges and limitations to our understanding of plant responses to drought under eCO₂. In particular, we argue that future studies need to move beyond exploring whether eCO₂ played ‘a role’ or ‘no role’ in responses to drought, but instead more carefully consider the timescales and conditions that would induce an influence. We also argue that considering emergent differences in soil water content may be an insufficient means of assessing the impact of eCO₂. We identify eCO₂ impact during severe drought (e.g. to the point of mortality), interactions with future changes in vapour pressure deficit and uncertainty about changes in leaf area as key gaps in our current understanding. New insights into CO₂ × drought interactions are essential to better constrain model theory that governs future climate model projections of land–atmosphere interactions during periods of water stress.

Plant survival during drought requires adequate hydration in living tissues and carbohydrate reserves for maintenance and recovery. We hypothesized that tree growth and hydraulic strategy determines the intensity and duration of the ‘physiological drought’, thereby affecting the relative contributions of loss of hydraulic function and carbohydrate depletion during mortality. We compared patterns in growth rate, water relations, gas exchange and carbohydrate dynamics in three tree species subjected to prolonged drought. Two Eucalyptus species (E. globulus, E. smithii) exhibited high growth rates and water-use resulting in rapid declines in water status and hydraulic conductance. In contrast, conservative growth and water relations in Pinus radiata resulted in longer periods of negative carbon balance and significant depletion of stored carbohydrates in all organs. The ongoing demand for carbohydrates from sustained respiration highlighted the role that duration of drought plays in facilitating carbohydrate consumption. Two drought strategies were revealed, differentiated by plant regulation of water status: plants maximized gas exchange, but were exposed to low water potentials and rapid hydraulic dysfunction; and tight regulation of gas exchange at the cost of carbohydrate depletion. These findings provide evidence for a relationship between hydraulic regulation of water status and carbohydrate depletion during terminal drought.

Summary statement: Mesophyll conductance ( g m ) was negatively correlated with wheat leaf age but was positively correlated with the surface area of chloroplasts exposed to intercellular airspaces ( S c ). The rate of decline in photosynthetic rate and g m as leaves aged was slower for water-stressed than well-watered plants. Upon rewatering, the degree of recovery from water-stress depended on the age of the leaves, with the strongest recovery for mature leaves, rather than young or old leaves. Diffusion of CO 2 from the intercellular airspaces to the site of Rubisco within C 3 plant chloroplasts ( g m ) governs photosynthetic CO 2 assimilation ( A ). However, variation in g m in response to environmental stress during leaf development remains poorly understood. Age-dependent changes in leaf ultrastructure and potential impacts on g m , A , and stomatal conductance to CO 2 ( g sc ) were investigated for wheat ( Triticum aestivum L.) in well-watered and water-stressed plants, and after recovery by re-watering of droughted plants. Significant reductions in A and g m were found as leaves aged. The oldest plants (15 days and 22 days) in water-stressed conditions showed higher A and gm compared to irrigated plants. The rate of decline in A and g m as leaves aged was slower for water-stressed compared to well-watered plants. When droughted plants were rewatered, the degree of recovery depended on the age of the leaves, but only for g m . The surface area of chloroplasts exposed to intercellular airspaces ( S c ) and the size of individual chloroplasts declined as leaves aged, resulting in a positive correlation between g m and S c . Leaf age significantly affected cell wall thickness ( t cw ), which was higher in old leaves compared to mature/young leaves. Greater knowledge of leaf anatomical traits associated with g m partially explained changes in physiology with leaf age and plant water status, which in turn should create more possibilities for improving photosynthesis using breeding/biotechnological strategies.

1077 I. 1078 II. 1079 III. 1080 IV. 1081 V. 1084 VI. 1087 VII. 1088 1089 References 1089 SUMMARY: The rate of CO₂ assimilation by plants is directly influenced by the concentration of CO₂ in the atmosphere, cₐ. As an environmental variable, cₐ also has a unique global and historic significance. Although relatively stable and uniform in the short term, global cₐ has varied substantially on the timescale of thousands to millions of years, and currently is increasing at seemingly an unprecedented rate. This may exert profound impacts on both climate and plant function. Here we utilise extensive datasets and models to develop an integrated, multi‐scale assessment of the impact of changing cₐ on plant carbon dioxide uptake and water use. We find that, overall, the sensitivity of plants to rising or falling cₐ is qualitatively similar across all scales considered. It is characterised by an adaptive feedback response that tends to maintain 1 − cᵢ/cₐ, the relative gradient for CO₂ diffusion into the leaf, relatively constant. This is achieved through predictable adjustments to stomatal anatomy and chloroplast biochemistry. Importantly, the long‐term response to changing cₐ can be described by simple equations rooted in the formulation of more commonly studied short‐term responses.

The iso/anisohydric continuum describes how plants regulate leaf water potential and is commonly used to classify species drought response strategies. However, drought response strategies comprise more than just this continuum, incorporating a suite of stomatal and hydraulic traits. Using a common garden experiment, we compared and contrasted four metrics commonly used to describe water use strategy during drought in 10 eucalyptus species comprising four major ecosystems in eastern Australia. We examined the degree to which these metrics were aligned with key stomatal and hydraulic traits related to plant water use and drought tolerance. Species rankings of water use strategy were inconsistent across four metrics. A newer metric (Hydroscape) was strongly linked to various plant traits, including the leaf turgor loss (TLP), water potential at stomatal closure (Pgs90), leaf and stem hydraulic vulnerability to embolism (PL50 and Px50), safety margin of hydraulic segmentation (HSMHS), maximum stomatal conductance (gsmax) and Huber value (HV). In addition, Hydroscape was correlated with climatic variables representing the water availability at the seed source site. Along the continuum of water regulation strategy, species with narrow Hydroscapes tended to occupy mesic regions and exhibit high TLP, PL50 and Px50 values and narrow HSMHS. High gsmax recorded in species with broad hydroscapes was also associated with high HV. Despite a fourfold difference in Hydroscape area, all species closed their stomata prior to the onset of hydraulic dysfunction, suggesting a common stomatal response across species that minimizes embolism risk during drought. Hydroscape area is useful in bridging stomatal regulation, hydraulic architecture and species drought tolerance, thus providing insight into species water use strategies. A plain language summary is available for this article. Plain Language Summary

Background and aims Extreme climate events, including flooding and prolonged drought, may establish long-lasting (legacy) effects on soil abiotic and biotic properties, potentially influencing soil N-cycling, microbial communities, and plant productivity. Nitrogen (N) fertilizer often stimulates plant growth, but it remains unknown whether N addition can alleviate the impact of legacy drought or waterlogging events on crops. Our aim was to assess the interactive effects of legacy extreme climate events and N-addition on these processes. Methods Using cotton as a model system, soils previously exposed to waterlogging and prolonged drought were used to examine potential legacy impacts of extreme climate on soil N process rates, abundance and structure of associated microbial communities, and cotton growth and productivity under different levels of N fertilizer application (0, 100, 200 and 300 kg N/ha). Results The deleterious legacy effects of prolonged drought on plant productivity were due to negative impacts on microbial abundance and community structure, and soil nutrient availability, thereby negatively influencing the rate of nitrification, and consequently plant available N. The legacy impacts of prolonged drought persisted throughout the experiment despite fertiliser applications of up to 300 kg of N/ha. The only observed legacy impacts of waterlogging were low NO3− levels in soils without N-addition and shifts in the abundance and structure of the N2O-reducing community. Conclusions There were strong legacy impacts of prolonged drought, but minor legacy impacts of waterlogging, on soils and crop yields which could not be fully counteracted by the high rates of N fertilizer application. This study provides critical knowledge contributing to the development of adaptation and soil N management strategies to minimize the loss of farm productivity, within the context of increased frequencies and intensities of extreme weather events.

Detrimental impacts of drought on crop yield have tripled in the last 50 years with climate models predicting that the frequency of such droughts will intensify in the future. Silicon (Si) accumulation, especially in Poaceae crops such as wheat ( Triticum aestivum L.), may alleviate the adverse impacts of drought. We have very limited information, however, about whether Si supplementation could alleviate the impacts of drought under field conditions and no studies have specifically manipulated rainfall. Using field–based rain exclusion shelters, we determined whether Si supplementation (equivalent to 39, 78 and 117 kg ha -1 ) affected T. aestivum growth, elemental chemistry [Si, carbon (C) and nitrogen (N)], physiology (rates of photosynthesis, transpiration, stomatal conductance, and water use efficiency) and yield (grain production) under ambient and drought (50% of ambient) rainfall scenarios. Averaged across Si treatments, drought reduced shoot mass by 21% and grain production by 18%. Si supplementation increased shoot mass by up to 43% and 73% in ambient and drought water treatments, respectively, and restored grain production in droughted plants to levels comparable with plants supplied with ambient rainfall. Si supplementation increased leaf-level water use efficiency by 32–74%, depending on Si supplementation rates. Water supply and Si supplementation did not alter concentrations of C and N, but Si supplementation increased shoot C content by 39% and 83% under ambient and drought conditions, respectively. This equates to an increase from 6.4 to 8.9 tonnes C ha -1 and from 4.03 to 7.35 tonnes C ha -1 under ambient and drought conditions, respectively. We conclude that Si supplementation ameliorated the negative impacts of drought on T. aestivum growth and grain yield, potentially through its beneficial impacts on water use efficiency. Moreover, the beneficial impacts of Si on plant growth and C storage may render Si supplementation a useful tool for both drought mitigation and C sequestration.

Understanding physiological acclimation of photosynthesis and respiration is important in elucidating the metabolic performance of trees in a changing climate. Does physiological acclimation to climate warming mirror acclimation to seasonal temperature changes? We grew Eucalyptus tereticornis trees in the field for 14 months inside 9-m tall whole-tree chambers tracking ambient air temperature (T air) or ambient T air + 3°C (i.e. ‘warmed’). We measured light- and CO2-saturated net photosynthesis (A max) and night-time dark respiration (R) each month at 25°C to quantify acclimation. Tree growth was measured, and leaf nitrogen (N) and total nonstructural carbohydrate (TNC) concentrations were determined to investigate mechanisms of acclimation. Warming reduced A max and R measured at 25°C compared to ambient-grown trees. Both traits also declined as mean daily T air increased, and did so in a similar way across temperature treatments. A max and R (at 25°C) both increased as TNC concentrations increased seasonally; these relationships appeared to arise from source–sink imbalances, suggesting potential substrate regulation of thermal acclimation. We found that photosynthesis and respiration each acclimated equivalently to experimental warming and seasonal temperature change of a similar magnitude, reflecting a common, nearly homeostatic constraint on leaf carbon exchange that will be important in governing tree responses to climate warming.

Understanding the physiological and biochemical responses of tree seedlings under extreme drought stress, along with recovery during rewatering, and potential intra-species differences, will allow us to more accurately predict forest responses under future climate change. Here, we selected seedlings from four provenances (AH (Anhui), JX (Jiangxi), HN (Hunan) and GX (Guangxi)) of Schima superba and carried out a simulated drought-rewatering experiment in a field-based rain-out shelter. Seedlings were progressively dried until they reached 50% and 88% loss of xylem hydraulic conductivity (PLC) (i.e. P 50 and P 88 ), respectively, before they were rehydrated and maintained at field capacity for 30 days. Leaf photosynthesis ( A sat ), water status, activity of superoxide dismutase (SOD), and proline (Pro) concentration were monitored and their associations were determined. Increasing drought significantly reduced A sat , relative water content (RWC) and SOD activity in all provenances, and Pro concentration was increased to improve water retention; all four provenances exhibited similar response patterns, associated with similar leaf ultrastructure at pre-drought. Upon rewatering, physiological and biochemical traits were restored to well-watered control values in P 50 -stressed seedlings. In P 88 -stressed seedlings, Pro was restored to control values, while SOD was not fully recovered. The recovery pattern differed partially among provenances. There was a progression of recovery following watering, with RWC firstly recovered, followed by SOD and Pro, and then A sat , but with significant associations among these traits. Collectively, the intra-specific differences of S. superba seedlings in recovery of physiology and biochemistry following rewatering highlight the need to consider variations within a given tree species coping with future more frequent drought stress.

Climate change is testing the resilience of forests worldwide pushing physiological tolerance to climatic extremes. Plant functional traits have been shown to be adapted to climate and have evolved patterns of trait correlations (similar patterns of distribution) and coordinations (mechanistic trade‐off). We predicted that traits would differentiate between populations associated with climatic gradients, suggestive of adaptive variation, and correlated traits would adapt to future climate scenarios in similar ways. We measured genetically determined trait variation and described patterns of correlation for seven traits: photochemical reflectance index (PRI), normalized difference vegetation index (NDVI), leaf size (LS), specific leaf area (SLA), δ13C (integrated water‐use efficiency, WUE), nitrogen concentration (NCONC), and wood density (WD). All measures were conducted in an experimental plantation on 960 trees sourced from 12 populations of a key forest canopy species in southwestern Australia. Significant differences were found between populations for all traits. Narrow‐sense heritability was significant for five traits (0.15–0.21), indicating that natural selection can drive differentiation; however, SLA (0.08) and PRI (0.11) were not significantly heritable. Generalized additive models predicted trait values across the landscape for current and future climatic conditions (>90% variance). The percent change differed markedly among traits between current and future predictions (differing as little as 1.5% (δ13C) or as much as 30% (PRI)). Some trait correlations were predicted to break down in the future (SLA:NCONC, δ13C:PRI, and NCONC:WD). Synthesis: Our results suggest that traits have contrasting genotypic patterns and will be subjected to different climate selection pressures, which may lower the working optimum for functional traits. Further, traits are independently associated with different climate factors, indicating that some trait correlations may be disrupted in the future. Genetic constraints and trait correlations may limit the ability for functional traits to adapt to climate change. We tested whether functional traits of an important tree species in southwestern Australia were genetically determined and able to undergo selection. Our results suggest that some traits are heritable (Water Use Efficiency) while others are not (SLA), indicating that in the future, under different climates, selection will differentially effect traits. Therefore, it is likely that current trait combinations will change with climate.