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The allocation of carbon (C) is an important component of tree physiology that influences growth and ecosystem C storage. Allocation is challenging to measure, and its sensitivity to environmental changes such as warming and altered water availability is uncertain. We exposed young Eucalyptus tereticornis trees to +3°C warming and elimination of summer precipitation in the field using whole-tree chambers. We calculated C allocation terms using detailed measurements of growth and continuous whole-crown CO 2 and water exchange measurements. Trees grew from small saplings to nearly 9 m height during this 15-month experiment. Warming accelerated growth and leaf area development, and it increased the partitioning of gross primary production (GPP) to aboveground respiration and growth while decreasing partitioning below ground. Eliminating summer precipitation reduced C gain and growth but did not impact GPP partitioning. Trees utilized deep soil water and avoided strongly negative water potentials. Warming increased growth respiration, but maintenance respiration acclimated homeostatically. The increasing growth in the warmed treatment resulted in higher rates of respiration, even with complete acclimation of maintenance respiration. Warming-induced stimulations of tree growth likely involve increased C allocation above ground, particularly to leaf area development, whereas reduced water availability may not stimulate allocation to roots., (© 2018 The Authors. New Phytologist © 2018 New Phytologist Trust.)

Understanding forest tree responses to climate warming and heatwaves is important for predicting changes in tree species diversity, forest C uptake, and vegetation-climate interactions. Yet, tree species differences in heatwave tolerance and their plasticity to growth temperature remain poorly understood. In this study, populations of four Eucalyptus species, two with large range sizes and two with comparatively small range sizes, were grown under two temperature treatments (cool and warm) before being exposed to an equivalent experimental heatwave. We tested whether the species with large and small range sizes differed in heatwave tolerance, and whether trees grown under warmer temperatures were more tolerant of heatwave conditions than trees grown under cooler temperatures. Visible heatwave damage was more common and severe in the species with small rather than large range sizes. In general, species that showed less tissue damage maintained higher stomatal conductance, lower leaf temperatures, larger increases in isoprene emissions, and less photosynthetic inhibition than species that showed more damage. Species exhibiting more severe visible damage had larger increases in heat shock proteins (HSPs) and respiratory thermotolerance (T max ). Thus, across species, increases in HSPs and T max were positively correlated, but inversely related to increases in isoprene emissions. Integration of leaf gas-exchange, isoprene emissions, proteomics, and respiratory thermotolerance measurements provided new insight into mechanisms underlying variability in tree species heatwave tolerance. Importantly, warm-grown seedlings were, surprisingly, more susceptible to heatwave damage than cool-grown seedlings, which could be associated with reduced enzyme concentrations in leaves. We conclude that species with restricted range sizes, along with trees growing under climate warming, may be more vulnerable to heatwaves of the future., (© 2019 John Wiley & Sons Ltd.)

The Kok and Laisk techniques can both be used to estimate light respiration R light . We investigated whether responses of R light to short- and long-term changes in leaf temperature depend on the technique used to estimate R light . We grew Eucalyptus tereticornis in whole-tree chambers under ambient temperature (AT) or AT + 3°C (elevated temperature, ET). We assessed dark respiration R dark and light respiration with the Kok (R Kok ) and Laisk (R Laisk ) methods at four temperatures to determine the degree of light suppression of respiration using both methods in AT and ET trees. The ET treatment had little impact on R dark , R Kok or R Laisk . Although the thermal sensitivities of R Kok or R Laisk were similar, R Kok was higher than R Laisk . We found negative values of R Laisk at the lowest measurement temperatures, indicating positive net CO 2 uptake, which we propose may be related to phosphoenolpyruvate carboxylase activity. Light suppression of R dark decreased with increasing leaf temperature, but the degree of suppression depended on the method used. The Kok and Laisk methods do not generate the same estimates of R light or light suppression of R dark between 20 and 35°C. Negative rates of R Laisk imply that this method may become less reliable at low temperatures., (© 2018 The Authors. New Phytologist © 2018 New Phytologist Trust.)

The temperature response of photosynthesis is one of the key factors determining predicted responses to warming in global vegetation models (GVMs). The response may vary geographically, owing to genetic adaptation to climate, and temporally, as a result of acclimation to changes in ambient temperature. Our goal was to develop a robust quantitative global model representing acclimation and adaptation of photosynthetic temperature responses. We quantified and modelled key mechanisms responsible for photosynthetic temperature acclimation and adaptation using a global dataset of photosynthetic CO 2 response curves, including data from 141 C 3 species from tropical rainforest to Arctic tundra. We separated temperature acclimation and adaptation processes by considering seasonal and common-garden datasets, respectively. The observed global variation in the temperature optimum of photosynthesis was primarily explained by biochemical limitations to photosynthesis, rather than stomatal conductance or respiration. We found acclimation to growth temperature to be a stronger driver of this variation than adaptation to temperature at climate of origin. We developed a summary model to represent photosynthetic temperature responses and showed that it predicted the observed global variation in optimal temperatures with high accuracy. This novel algorithm should enable improved prediction of the function of global ecosystems in a warming climate., (© 2018 The Authors. New Phytologist © 2018 New Phytologist Trust.)

Short-term acclimation and long-term adaptation represent two ways in which forest trees can respond to changes in temperature. Yet, the relative contribution of thermal acclimation and adaptation to tree physiological responses to temperature remains poorly understood. Here, we grew two cool-origin and two warm-origin populations of a widespread broad-leaved evergreen tree species (Corymbia calophylla (Lindl.) K.D.Hill & L.A.S.Johnson) from a Mediterranean climate in southwestern Australia under two growth temperatures representative of the cool- and warm-edge of the species distribution. The populations selected from each thermal environment represented both high and low precipitation sites. We measured the short-term temperature response of leaf photosynthesis (A) and dark respiration (R), and attributed observed variation to acclimation, adaptation or the combination of both. We observed limited variation in the temperature optimum (Topt) of A between temperature treatments or among populations, suggesting little plasticity or genetic differentiation in the Topt of A. Yet, other aspects of the temperature response of A and R were dependent upon population and growth temperature. Under cooler growth temperatures, the population from the coolest, wettest environment had the lowest A (at 25 °C) among all four populations, but exhibited the highest A (at 25 °C) under warmer growth temperatures. Populations varied in R (at 20 °C) and the temperature sensitivity of R (i.e., Q10 or activation energy) under cool, but not warm growth temperatures. However, populations showed similar yet lower R (at 20 °C) and no differences in the temperature sensitivity of R under warmer growth temperatures. We conclude that C. calophylla populations from contrasting climates vary in physiological acclimation to temperature, which might influence how this ecologically important tree species and the forests of southwestern Australia respond to climate change., (© The Author 2017. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.)

The ability of plants to maintain an intact water transport system in leaves under drought conditions is intimately linked to survival and can been be seen as adaptive in shaping species climatic limits. Large differences in leaf hydraulic vulnerability to drought are known among species from contrasting climates, yet whether this trait varies among populations within a single species and, furthermore, whether it is altered by changes in growth conditions, remain unclear. We examined intraspecific variation in both leaf water transport capacity (Kleaf) and leaf hydraulic vulnerability to drought (P50leaf) among eight populations of Corymbia calophylla (R. Br.) K.D. Hill & L.A.S. Johnson (Myrtaceae) from both cool and warm climatic regions grown reciprocally under two temperature treatments representing the cool and warm edge of the species distribution. Kleaf did not vary between cool and warm-climate populations, nor was it affected by variable growth temperature. In contrast, population origin and growth temperature independently altered P50leaf. Using data pooled across growth temperatures, cool-climate populations showed significantly higher leaf hydraulic vulnerability (P50leaf = -3.55 ± 0.18 MPa) than warm-climate populations (P50leaf = -3.78 ± 0.08 MPa). Across populations, P50leaf decreased as population home-climate temperature increased, but was unrelated to rainfall and aridity. For populations from both cool and warm climatic regions, P50leaf was lower under the warmer growth conditions. These results provide evidence of trait plasticity in leaf hydraulic vulnerability to drought in response to variable growth temperature. Furthermore, they suggest that climate, and in particular temperature, may be a strong selective force in shaping intraspecific variation in leaf hydraulic vulnerability to drought., (© The Author 2017. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.)

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 (Tair ) or ambient Tair  + 3°C (i.e. 'warmed'). We measured light- and CO2 -saturated net photosynthesis (Amax ) 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 Amax and R measured at 25°C compared to ambient-grown trees. Both traits also declined as mean daily Tair increased, and did so in a similar way across temperature treatments. Amax 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., (© 2016 The Authors. New Phytologist © 2016 New Phytologist Trust.)

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 CO2 and H2O, 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. [ABSTRACT FROM AUTHOR]

As rapid climate warming creates a mismatch between forest trees and their home environment, the ability of trees to cope with warming depends on their capacity to physiologically adjust to higher temperatures. In widespread species, individual trees in cooler home climates are hypothesized to more successfully acclimate to warming than their counterparts in warmer climates that may approach thermal limits. We tested this prediction with a climate-shift experiment in widely distributed Eucalyptus tereticornis and E. grandis using provenances originating along a ~2500 km latitudinal transect (15.5-38.0°S) in eastern Australia. We grew 21 provenances in conditions approximating summer temperatures at seed origin and warmed temperatures (+3.5 °C) using a series of climate-controlled glasshouse bays. The effects of +3.5 °C warming strongly depended on home climate. Cool-origin provenances responded to warming through an increase in photosynthetic capacity and total leaf area, leading to enhanced growth of 20-60%. Warm-origin provenances, however, responded to warming through a reduction in photosynthetic capacity and total leaf area, leading to reduced growth of approximately 10%. These results suggest that there is predictable intraspecific variation in the capacity of trees to respond to warming; cool-origin taxa are likely to benefit from warming, while warm-origin taxa may be negatively affected. [ABSTRACT FROM AUTHOR]