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Understanding how canopy-scale photosynthesis responds to temperature is of paramount importance for realistic prediction of the likely impact of climate change on forest growth. The effects of temperature on leaf-scale photosynthesis have been extensively documented but data demonstrating the temperature response of canopy-scale photosynthesis are relatively rare, and the mechanisms that determine the response are not well quantified. Here, we compared leaf- and canopy-scale photosynthesis responses to temperature measured in a whole-tree chamber experiment and tested mechanisms that could explain the difference between leaf and crown scale temperature optima for photosynthesis. We hypothesised that 1) there is a large contribution of non-light saturated leaves to total crown photosynthesis; 2) photosynthetic component processes vary vertically through the canopy following the gradient in incident light; and 3) seasonal temperature acclimation of photosynthetic biochemistry has a significant role in determining the overall temperature response of canopy photosynthesis. We tested these hypotheses using three models of canopy radiation interception and photosynthesis parameterized with leaf-level physiological data and estimates of canopy leaf area. Our results identified the influence of non-light saturated leaves as a key determinant of the lower temperature optimum of canopy photosynthesis, which reduced the temperature optimum of canopy photosynthesis by 6-8 °C compared to the leaf scale. Further, we demonstrate the importance of accounting for within-canopy variation and seasonal temperature acclimation of photosynthetic biochemistry in determining the magnitude of canopy photosynthesis. Overall, our study identifies key processes that need to be incorporated in terrestrial biosphere models to accurately predict temperature responses of whole-tree photosynthesis., (© The Author(s) 2024. Published by Oxford University Press.)

The importance of phosphorus (P) in regulating ecosystem responses to climate change has fostered P-cycle implementation in land surface models, but their CO 2 effects predictions have not been evaluated against measurements. Here, we perform a data-driven model evaluation where simulations of eight widely used P-enabled models were confronted with observations from a long-term free-air CO 2 enrichment experiment in a mature, P-limited Eucalyptus forest. We show that most models predicted the correct sign and magnitude of the CO 2 effect on ecosystem carbon (C) sequestration, but they generally overestimated the effects on plant C uptake and growth. We identify leaf-to-canopy scaling of photosynthesis, plant tissue stoichiometry, plant belowground C allocation, and the subsequent consequences for plant-microbial interaction as key areas in which models of ecosystem C-P interaction can be improved. Together, this data-model intercomparison reveals data-driven insights into the performance and functionality of P-enabled models and adds to the existing evidence that the global CO 2 -driven carbon sink is overestimated by models.

The capacity for terrestrial ecosystems to sequester additional carbon (C) with rising CO 2 concentrations depends on soil nutrient availability 1,2 . Previous evidence suggested that mature forests growing on phosphorus (P)-deprived soils had limited capacity to sequester extra biomass under elevated CO 2 (refs. 3-6 ), but uncertainty about ecosystem P cycling and its CO 2 response represents a crucial bottleneck for mechanistic prediction of the land C sink under climate change 7 . Here, by compiling the first comprehensive P budget for a P-limited mature forest exposed to elevated CO 2 , we show a high likelihood that P captured by soil microorganisms constrains ecosystem P recycling and availability for plant uptake. Trees used P efficiently, but microbial pre-emption of mineralized soil P seemed to limit the capacity of trees for increased P uptake and assimilation under elevated CO 2 and, therefore, their capacity to sequester extra C. Plant strategies to stimulate microbial P cycling and plant P uptake, such as increasing rhizosphere C release to soil, will probably be necessary for P-limited forests to increase C capture into new biomass. Our results identify the key mechanisms by which P availability limits CO 2 fertilization of tree growth and will guide the development of Earth system models to predict future long-term C storage., (© 2024. Crown.)

The critical temperature beyond which photosynthetic machinery in tropical trees begins to fail averages approximately 46.7 °C (T crit ) 1 . However, it remains unclear whether leaf temperatures experienced by tropical vegetation approach this threshold or soon will under climate change. Here we found that pantropical canopy temperatures independently triangulated from individual leaf thermocouples, pyrgeometers and remote sensing (ECOSTRESS) have midday peak temperatures of approximately 34 °C during dry periods, with a long high-temperature tail that can exceed 40 °C. Leaf thermocouple data from multiple sites across the tropics suggest that even within pixels of moderate temperatures, upper canopy leaves exceed T crit 0.01% of the time. Furthermore, upper canopy leaf warming experiments (+2, 3 and 4 °C in Brazil, Puerto Rico and Australia, respectively) increased leaf temperatures non-linearly, with peak leaf temperatures exceeding T crit 1.3% of the time (11% for more than 43.5 °C, and 0.3% for more than 49.9 °C). Using an empirical model incorporating these dynamics (validated with warming experiment data), we found that tropical forests can withstand up to a 3.9 ± 0.5 °C increase in air temperatures before a potential tipping point in metabolic function, but remaining uncertainty in the plasticity and range of T crit in tropical trees and the effect of leaf death on tree death could drastically change this prediction. The 4.0 °C estimate is within the 'worst-case scenario' (representative concentration pathway (RCP) 8.5) of climate change predictions 2 for tropical forests and therefore it is still within our power to decide (for example, by not taking the RCP 6.0 or 8.5 route) the fate of these critical realms of carbon, water and biodiversity 3,4 ., (© 2023. The Author(s), under exclusive licence to Springer Nature Limited.)

As the global climate warms, a key question is how increased leaf temperatures will affect tree physiology and the coupling between leaf and air temperatures in forests. To explore the impact of increasing temperatures on plant performance in open air, we warmed leaves in the canopy of two mature evergreen forests, a temperate Eucalyptus woodland and a tropical rainforest. The leaf heaters consistently maintained leaves at a target of 4 °C above ambient leaf temperatures. Ambient leaf temperatures (Tleaf) were mostly coupled to air temperatures (Tair), but at times, leaves could be 8-10 °C warmer than ambient air temperatures, especially in full sun. At both sites, Tleaf was warmer at higher air temperatures (Tair > 25 °C), but was cooler at lower Tair, contrary to the 'leaf homeothermy hypothesis'. Warmed leaves showed significantly lower stomatal conductance (-0.05 mol m-2 s-1 or -43% across species) and net photosynthesis (-3.91 μmol m-2 s-1 or -39%), with similar rates in leaf respiration rates at a common temperature (no acclimation). Increased canopy leaf temperatures due to future warming could reduce carbon assimilation via reduced photosynthesis in these forests, potentially weakening the land carbon sink in tropical and temperate forests., (© The Author(s) 2023. Published by Oxford University Press.)

Optimal stomatal theory predicts that stomata operate to maximise photosynthesis (A net ) and minimise transpirational water loss to achieve optimal intrinsic water-use efficiency (iWUE). We tested whether this theory can predict stomatal responses to elevated atmospheric CO 2 (eCO 2 ), and whether it can capture differences in responsiveness among woody plant functional types (PFTs). We conducted a meta-analysis of tree studies of the effect of eCO 2 on iWUE and its components A net and stomatal conductance (g s ). We compared three PFTs, using the unified stomatal optimisation (USO) model to account for confounding effects of leaf-air vapour pressure difference (D). We expected smaller g s , but greater A net , responses to eCO 2 in gymnosperms compared with angiosperm PFTs. We found that iWUE increased in proportion to increasing eCO 2 in all PFTs, and that increases in A net had stronger effects than reductions in g s . The USO model correctly captured stomatal behaviour with eCO 2 across most datasets. The chief difference among PFTs was a lower stomatal slope parameter (g 1 ) for the gymnosperm, compared with angiosperm, species. Land surface models can use the USO model to describe stomatal behaviour under changing atmospheric CO 2 conditions., (© 2022 The Authors. New Phytologist © 2022 New Phytologist Foundation.)

There is a pressing need to better understand ecosystem resilience to droughts and heatwaves. Eco-evolutionary optimization approaches have been proposed as means to build this understanding in land surface models and improve their predictive capability, but competing approaches are yet to be tested together. Here, we coupled approaches that optimize canopy gas exchange and leaf nitrogen investment, respectively, extending both approaches to account for hydraulic impairment. We assessed model predictions using observations from a native Eucalyptus woodland that experienced repeated droughts and heatwaves between 2013 and 2020, whilst exposed to an elevated [CO 2 ] treatment. Our combined approaches improved predictions of transpiration and enhanced the simulated magnitude of the CO 2 fertilization effect on gross primary productivity. The competing approaches also worked consistently along axes of change in soil moisture, leaf area, and [CO 2 ]. Despite predictions of a significant percentage loss of hydraulic conductivity due to embolism (PLC) in 2013, 2014, 2016, and 2017 (99th percentile PLC > 45%), simulated hydraulic legacy effects were small and short-lived (2 months). Our analysis suggests that leaf shedding and/or suppressed foliage growth formed a strategy to mitigate drought risk. Accounting for foliage responses to water availability has the potential to improve model predictions of ecosystem resilience., (© 2022 The Authors. Plant, Cell & Environment published by John Wiley & Sons Ltd.)

Tropical forests take up more carbon (C) from the atmosphere per annum by photosynthesis than any other type of vegetation. Phosphorus (P) limitations to C uptake are paramount for tropical and subtropical forests around the globe. Yet the generality of photosynthesis-P relationships underlying these limitations are in question, and hence are not represented well in terrestrial biosphere models. Here we demonstrate the dependence of photosynthesis and underlying processes on both leaf N and P concentrations. The regulation of photosynthetic capacity by P was similar across four continents. Implementing P constraints in the ORCHIDEE-CNP model, gross photosynthesis was reduced by 36% across the tropics and subtropics relative to traditional N constraints and unlimiting leaf P. Our results provide a quantitative relationship for the P dependence for photosynthesis for the front-end of global terrestrial C models that is consistent with canopy leaf measurements., (© 2022. The Author(s).)

While trees can acclimate to warming, there is concern that tropical rainforest species may be less able to acclimate because they have adapted to a relatively stable thermal environment. Here we tested whether the physiological adjustments to warming differed among Australian tropical, subtropical and warm-temperate rainforest trees. Photosynthesis and respiration temperature responses were quantified in six Australian rainforest seedlings of tropical, subtropical and warm-temperate climates grown across four growth temperatures in a glasshouse. Temperature-response models were fitted to identify mechanisms underpinning the response to warming. Tropical and subtropical species had higher temperature optima for photosynthesis (T optA ) than temperate species. There was acclimation of T optA to warmer growth temperatures. The rate of acclimation (0.35-0.78°C °C -1 ) was higher in tropical and subtropical than in warm-temperate trees and attributed to differences in underlying biochemical parameters, particularly increased temperature optima of V cmax25 and J max25 . The temperature sensitivity of respiration (Q 10 ) was 24% lower in tropical and subtropical compared with warm-temperate species. Overall, tropical and subtropical species had a similar capacity to acclimate to changes in growth temperature as warm-temperate species, despite being grown at higher temperatures. Quantifying the physiological acclimation in rainforests can improve accuracy of future climate predictions and assess their potential vulnerability to warming., (© 2022 The Authors. New Phytologist © 2022 New Phytologist Foundation.)

Evergreen species are widespread across the globe, representing two major plant functional forms in terrestrial models. We reviewed and analysed the responses of photosynthesis and respiration to warming in 101 evergreen species from boreal to tropical biomes. Summertime temperatures affected both latitudinal gas exchange rates and the degree of responsiveness to experimental warming. The decrease in net photosynthesis at 25°C (A net25 ) was larger with warming in tropical climates than cooler ones. Respiration at 25°C (R 25 ) was reduced by 14% in response to warming across species and biomes. Gymnosperms were more sensitive to greater amounts of warming than broadleaved evergreens, with A net25 and R 25 reduced c. 30-40% with > 10°C warming. While standardised rates of carboxylation (V cmax25 ) and electron transport (J max25 ) adjusted to warming, the magnitude of this adjustment was not related to warming amount (range 0.6-16°C). The temperature optimum of photosynthesis (T optA ) increased on average 0.34°C per °C warming. The combination of more constrained acclimation of photosynthesis and increasing respiration rates with warming could possibly result in a reduced carbon sink in future warmer climates. The predictable patterns of thermal acclimation across biomes provide a strong basis to improve modelling predictions of the future terrestrial carbon sink with warming., (© 2022 The Authors New Phytologist © 2022 New Phytologist Foundation.)

Current carbon cycle models attribute rising atmospheric CO2 as the major driver of the increased terrestrial carbon sink, but with substantial uncertainties. The photosynthetic response of trees to elevated atmospheric CO2 is a necessary step, but not the only one, for sustaining the terrestrial carbon uptake, but can vary diurnally, seasonally and with duration of CO2 exposure. Hence, we sought to quantify the photosynthetic response of the canopy-dominant species, Quercus robur, in a mature deciduous forest to elevated CO2 (eCO2) (+150 μmol mol-1 CO2) over the first 3 years of a long-term free air CO2 enrichment facility at the Birmingham Institute of Forest Research in central England (BIFoR FACE). Over 3000 measurements of leaf gas exchange and related biochemical parameters were conducted in the upper canopy to assess the diurnal and seasonal responses of photosynthesis during the 2nd and 3rd year of eCO2 exposure. Measurements of photosynthetic capacity via biochemical parameters, derived from CO2 response curves, (Vcmax and Jmax) together with leaf nitrogen concentrations from the pre-treatment year to the 3rd year of eCO2 exposure, were examined. We hypothesized an initial enhancement in light-saturated net photosynthetic rates (Asat) with CO2 enrichment of ≈37% based on theory but also expected photosynthetic capacity would fall over the duration of the study. Over the 3-year period, Asat of upper-canopy leaves was 33 ± 8% higher (mean and standard error) in trees grown in eCO2 compared with ambient CO2 (aCO2), and photosynthetic enhancement decreased with decreasing light. There were no significant effects of CO2 treatment on Vcmax or Jmax, nor leaf nitrogen. Our results suggest that mature Q. robur may exhibit a sustained, positive response to eCO2 without photosynthetic downregulation, suggesting that, with adequate nutrients, there will be sustained enhancement in C assimilated by these mature trees. Further research will be required to understand the location and role of the additionally assimilated carbon., (© The Author(s) 2021. Published by Oxford University Press.)

We introduce the AusTraits database - a compilation of values of plant traits for taxa in the Australian flora (hereafter AusTraits). AusTraits synthesises data on 448 traits across 28,640 taxa from field campaigns, published literature, taxonomic monographs, and individual taxon descriptions. Traits vary in scope from physiological measures of performance (e.g. photosynthetic gas exchange, water-use efficiency) to morphological attributes (e.g. leaf area, seed mass, plant height) which link to aspects of ecological variation. AusTraits contains curated and harmonised individual- and species-level measurements coupled to, where available, contextual information on site properties and experimental conditions. This article provides information on version 3.0.2 of AusTraits which contains data for 997,808 trait-by-taxon combinations. We envision AusTraits as an ongoing collaborative initiative for easily archiving and sharing trait data, which also provides a template for other national or regional initiatives globally to fill persistent gaps in trait knowledge., (© 2021. The Author(s).)

Phosphorus (P) is an essential macro-nutrient required for plant metabolism and growth. Low P availability could potentially limit plant responses to elevated carbon dioxide (eCO 2 ), but consensus has yet to be reached on the extent of this limitation. Here, based on data from experiments that manipulated both CO 2 and P for young individuals of woody and non-woody species, we present a meta-analysis of P limitation impacts on plant growth, physiological, and morphological response to eCO 2 . We show that low P availability attenuated plant photosynthetic response to eCO 2 by approximately one-quarter, leading to a reduced, but still positive photosynthetic response to eCO 2 compared to those under high P availability. Furthermore, low P limited plant aboveground, belowground, and total biomass responses to eCO 2 , by 14.7%, 14.3%, and 12.4%, respectively, equivalent to an approximate halving of the eCO 2 responses observed under high P availability. In comparison, low P availability did not significantly alter the eCO 2 -induced changes in plant tissue nutrient concentration, suggesting tissue nutrient flexibility is an important mechanism allowing biomass response to eCO 2 under low P availability. Low P significantly reduced the eCO 2 -induced increase in leaf area by 14.3%, mirroring the aboveground biomass response, but low P did not affect the eCO 2 -induced increase in root length. Woody plants exhibited stronger attenuation effect of low P on aboveground biomass response to eCO 2 than non-woody plants, while plants with different mycorrhizal associations showed similar responses to low P and eCO 2 interaction. This meta-analysis highlights crucial data gaps in capturing plant responses to eCO 2 and low P availability. Field-based experiments with longer-term exposure of both CO 2 and P manipulations are critically needed to provide ecosystem-scale understanding. Taken together, our results provide a quantitative baseline to constrain model-based hypotheses of plant responses to eCO 2 under P limitation, thereby improving projections of future global change impacts., (© 2020 John Wiley & Sons Ltd.)

Leaves are exposed to different light conditions according to their canopy position, resulting in structural and anatomical differences with consequences for carbon uptake. While these structure-function relationships have been thoroughly explored in dense forest canopies, such gradients may be diminished in open canopies, and they are often ignored in ecosystem models. We tested within-canopy differences in photosynthetic properties and structural traits in leaves in a mature Eucalyptus tereticornis canopy exposed to long-term elevated CO2 for up to three years. We explored these traits in relation to anatomical variation and diffusive processes for CO2 (i.e., stomatal conductance, gs and mesophyll conductance, gm) in both upper and lower portions of the canopy receiving ambient and elevated CO2. While shade resulted in 13% lower leaf mass per area ratio (MA) in lower versus upper canopy leaves, there was no relationship between leaf Nmass and canopy gap fraction. Both maximum carboxylation capacity (Vcmax) and maximum electron transport (Jmax) were ~ 18% lower in shaded leaves and were also reduced by ~ 22% with leaf aging. In mature leaves, we found no canopy differences for gm or gs, despite anatomical differences in MA, leaf thickness and mean mesophyll thickness between canopy positions. There was a positive relationship between net photosynthesis and gm or gs in mature leaves. Mesophyll conductance was negatively correlated with mean parenchyma length, suggesting that long palisade cells may contribute to a longer CO2 diffusional pathway and more resistance to CO2 transfer to chloroplasts. Few other relationships between gm and anatomical variables were found in mature leaves, which may be due to the open crown of Eucalyptus. Consideration of shade effects and leaf-age dependent responses to photosynthetic capacity and mesophyll conductance are critical to improve canopy photosynthesis models and will improve understanding of long-term responses to elevated CO2 in tree canopies., (© The Author(s) 2020. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permission@oup.com.)

Plant respiration can acclimate to changing environmental conditions and vary between species as well as biome types, although belowground respiration responses to ongoing climate warming are not well understood. Understanding the thermal acclimation capacity of root respiration (Rroot) in relation to increasing temperatures is therefore critical in elucidating a key uncertainty in plant function in response to warming. However, the degree of temperature acclimation of Rroot in rainforest trees and how root chemical and morphological traits are related to acclimation is unknown. Here we investigated the extent to which respiration of fine roots (≤2 mm) of four tropical and four warm-temperate rainforest tree seedlings differed in response to warmer growth temperatures (control and +6 °C), including temperature sensitivity (Q10) and the degree of acclimation of Rroot. Regardless of biome type, we found no consistent pattern in the short-term temperature responses of Rroot to elevated growth temperature: a significant reduction in the temperature response of Rroot to +6 °C treatment was only observed for a tropical species, Cryptocarya mackinnoniana, whereas the other seven species had either some stimulation or no alteration. Across species, Rroot was positively correlated with root tissue nitrogen concentration (mg g-1), while Q10 was positively correlated with root tissue density (g cm-3). Warming increased root tissue density by 20.8% but did not alter root nitrogen across species. We conclude that thermal acclimation capacity of Rroot to warming is species-specific and suggest that root tissue density is a useful predictor of Rroot and its thermal responses in rainforest tree seedlings., (© The Author(s) 2020. Published by Oxford University Press. All rights reserved. For permissions, please email: journals.permission@oup.com.)

Atmospheric carbon dioxide enrichment (eCO 2 ) can enhance plant carbon uptake and growth 1-5 , thereby providing an important negative feedback to climate change by slowing the rate of increase of the atmospheric CO 2 concentration 6 . Although evidence gathered from young aggrading forests has generally indicated a strong CO 2 fertilization effect on biomass growth 3-5 , it is unclear whether mature forests respond to eCO 2 in a similar way. In mature trees and forest stands 7-10 , photosynthetic uptake has been found to increase under eCO 2 without any apparent accompanying growth response, leaving the fate of additional carbon fixed under eCO 2 unclear 4,5,7-11 . Here using data from the first ecosystem-scale Free-Air CO 2 Enrichment (FACE) experiment in a mature forest, we constructed a comprehensive ecosystem carbon budget to track the fate of carbon as the forest responded to four years of eCO 2 exposure. We show that, although the eCO 2 treatment of +150 parts per million (+38 per cent) above ambient levels induced a 12 per cent (+247 grams of carbon per square metre per year) increase in carbon uptake through gross primary production, this additional carbon uptake did not lead to increased carbon sequestration at the ecosystem level. Instead, the majority of the extra carbon was emitted back into the atmosphere via several respiratory fluxes, with increased soil respiration alone accounting for half of the total uptake surplus. Our results call into question the predominant thinking that the capacity of forests to act as carbon sinks will be generally enhanced under eCO 2 , and challenge the efficacy of climate mitigation strategies that rely on ubiquitous CO 2 fertilization as a driver of increased carbon sinks in global forests.

Vapour pressure deficit (D) is projected to increase in the future as temperature rises. In response to increased D, stomatal conductance (gs) and photosynthesis (A) are reduced, which may result in significant reductions in terrestrial carbon, water and energy fluxes. It is thus important for gas exchange models to capture the observed responses of gs and A with increasing D. We tested a series of coupled A-gs models against leaf gas exchange measurements from the Cumberland Plain Woodland (Australia), where D regularly exceeds 2 kPa and can reach 8 kPa in summer. Two commonly used A-gs models were not able to capture the observed decrease in A and gs with increasing D at the leaf scale. To explain this decrease in A and gs, two alternative hypotheses were tested: hydraulic limitation (i.e., plants reduce gs and/or A due to insufficient water supply) and non-stomatal limitation (i.e., downregulation of photosynthetic capacity). We found that the model that incorporated a non-stomatal limitation captured the observations with high fidelity and required the fewest number of parameters. Whilst the model incorporating hydraulic limitation captured the observed A and gs, it did so via a physical mechanism that is incorrect. We then incorporated a non-stomatal limitation into the stand model, MAESPA, to examine its impact on canopy transpiration and gross primary production. Accounting for a non-stomatal limitation reduced the predicted transpiration by ~19%, improving the correspondence with sap flow measurements, and gross primary production by ~14%. Given the projected global increases in D associated with future warming, these findings suggest that models may need to incorporate non-stomatal limitation to accurately simulate A and gs in the future with high D. Further data on non-stomatal limitation at high D should be a priority, in order to determine the generality of our results and develop a widely applicable model., (© The Author(s) 2019. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

To quantify stem respiration (R S ) under elevated CO 2 (eCO 2 ), stem CO 2 efflux (E A ) and CO 2 flux through the xylem (F T ) should be accounted for, because part of respired CO 2 is transported upwards with the sap solution. However, previous studies have used E A as a proxy of R S , which could lead to equivocal conclusions. Here, to test the effect of eCO 2 on R S , both E A and F T were measured in a free-air CO 2 enrichment experiment located in a mature Eucalyptus native forest. Drought stress substantially reduced E A and R S , which were unaffected by eCO 2 , likely as a consequence of its neutral effect on stem growth in this phosphorus-limited site. However, xylem CO 2 concentration measured near the stem base was higher under eCO 2 , and decreased along the stem resulting in a negative contribution of F T to R S , whereas the contribution of F T to R S under ambient CO 2 was positive. Negative F T indicates net efflux of CO 2 respired below the monitored stem segment, likely coming from the roots. Our results highlight the role of nutrient availability on the dependency of R S on eCO 2 and suggest stimulated root respiration under eCO 2 that may shift vertical gradients in xylem [CO 2 ] confounding the interpretation of E A measurements., (© 2019 John Wiley & Sons Ltd.)

Elevated CO 2 affects C cycling processes which in turn can influence the nitrogen (N) and phosphorus (P) concentrations of plant tissues. Given differences in how N and P are used by plants, we asked if their stoichiometry in leaves and wood was maintained or altered in a long-term elevated CO 2 experiment in a mature Eucalyptus forest on a low P soil (EucFACE). We measured N and P concentrations in green leaves at different ages at the top of mature trees across 6 years including 5 years in elevated CO 2 . N and P concentrations in green and senesced leaves and wood were determined to evaluate both spatial and temporal variation of leaf N and P concentrations, including the N and P retranslocation in leaves and wood. Leaf P concentrations were 32% lower in old mature leaves compared to newly flushed leaves with no effect of elevated CO 2 on leaf P. By contrast, elevated CO 2 significantly decreased leaf N concentrations in newly flushed leaves but this effect disappeared as leaves matured. As such, newly flushed leaves had 9% lower N:P ratios in elevated CO 2 and N:P ratios were not different in mature green leaves (CO 2 by Age effect, P = 0.02). Over time, leaf N and P concentrations in the upper canopy slightly declined in both CO 2 treatments compared to before the start of the experiment. P retranslocation in leaves was 50%, almost double that of N retranslocation (29%), indicating that this site was P-limited and that P retranslocation was an important mechanism in this ecosystem to retain P in plants. As P-limited trees tend to store relatively more N than P, we found an increased N:P ratio in sapwood in response to elevated CO 2 ( P < 0.01), implying N accumulation in live wood. The flexible stoichiometric ratios we observed can have important implications for how plants adjust to variable environmental conditions including climate change. Hence, variable nutrient stoichiometry should be accounted for in large-scale Earth Systems models invoking biogeochemical processes.

Rising atmospheric CO 2 concentrations is expected to stimulate photosynthesis and carbohydrate production, while inhibiting photorespiration. By contrast, nitrogen (N) concentrations in leaves generally tend to decline under elevated CO 2 (eCO 2 ), which may reduce the magnitude of photosynthetic enhancement. We tested two hypotheses as to why leaf N is reduced under eCO 2 : (a) A "dilution effect" caused by increased concentration of leaf carbohydrates; and (b) inhibited nitrate assimilation caused by reduced supply of reductant from photorespiration under eCO 2 . This second hypothesis is fully tested in the field for the first time here, using tall trees of a mature Eucalyptus forest exposed to Free-Air CO 2 Enrichment (EucFACE) for five years. Fully expanded young and mature leaves were both measured for net photosynthesis, photorespiration, total leaf N, nitrate ( N O 3 - ) concentrations, carbohydrates and N O 3 - reductase activity to test these hypotheses. Foliar N concentrations declined by 8% under eCO 2 in new leaves, while the N O 3 - fraction and total carbohydrate concentrations remained unchanged by CO 2 treatment for either new or mature leaves. Photorespiration decreased 31% under eCO 2 supplying less reductant, and in situ N O 3 - reductase activity was concurrently reduced (-34%) in eCO 2 , especially in new leaves during summer periods. Hence, N O 3 - assimilation was inhibited in leaves of E. tereticornis and the evidence did not support a significant dilution effect as a contributor to the observed reductions in leaf N concentration. This finding suggests that the reduction of N O 3 - reductase activity due to lower photorespiration in eCO 2 can contribute to understanding how eCO 2 -induced photosynthetic enhancement may be lower than previously expected. We suggest that large-scale vegetation models simulating effects of eCO 2 on N biogeochemistry include both mechanisms, especially where N O 3 - is major N source to the dominant vegetation and where leaf flushing and emergence occur in temperatures that promote high photorespiration rates., (© 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.)

Climate is an important factor limiting tree distributions and adaptation to different thermal environments may influence how tree populations respond to climate warming. Given the current rate of warming, it has been hypothesized that tree populations in warmer, more thermally stable climates may have limited capacity to respond physiologically to warming compared to populations from cooler, more seasonal climates. We determined in a controlled environment how several provenances of widely distributed Eucalyptus tereticornis and E. grandis adjusted their photosynthetic capacity to +3.5°C warming along their native distribution range (~16-38°S) and whether climate of seed origin of the provenances influenced their response to different growth temperatures. We also tested how temperature optima (T opt ) of photosynthesis and J max responded to higher growth temperatures. Our results showed increased photosynthesis rates at a standardized temperature with warming in temperate provenances, while rates in tropical provenances were reduced by about 40% compared to their temperate counterparts. Temperature optima of photosynthesis increased as provenances were exposed to warmer growth temperatures. Both species had ~30% reduced photosynthetic capacity in tropical and subtropical provenances related to reduced leaf nitrogen and leaf Rubisco content compared to temperate provenances. Tropical provenances operated closer to their thermal optimum and came within 3% of the T opt of J max during the daily temperature maxima. Hence, further warming may negatively affect C uptake and tree growth in warmer climates, whereas eucalypts in cooler climates may benefit from moderate warming., (© 2018 John Wiley & Sons Ltd.)

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 2 and H 2 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., (© 2018 John Wiley & Sons Ltd.)

Contents 986 I. 987 II. 987 III. 988 IV. 991 V. 992 VI. 995 VII. 997 VIII. 998 References 998 SUMMARY: It has been 75 yr since leaf respiratory metabolism in the light (day respiration) was identified as a low-flux metabolic pathway that accompanies photosynthesis. In principle, it provides carbon backbones for nitrogen assimilation and evolves CO 2 and thus impacts on plant carbon and nitrogen balances. However, for a long time, uncertainties have remained as to whether techniques used to measure day respiratory efflux were valid and whether day respiration responded to environmental gaseous conditions. In the past few years, significant advances have been made using carbon isotopes, 'omics' analyses and surveys of respiration rates in mesocosms or ecosystems. There is substantial evidence that day respiration should be viewed as a highly dynamic metabolic pathway that interacts with photosynthesis and photorespiration and responds to atmospheric CO 2 mole fraction. The view of leaf day respiration as a constant and/or negligible parameter of net carbon exchange is now outdated and it should now be regarded as a central actor of plant carbon-use efficiency., (© 2017 The Authors. New Phytologist © 2017 New Phytologist Trust.)

Elevated atmospheric CO 2 (eCO 2 ) is expected to reduce the impacts of drought and increase photosynthetic rates via two key mechanisms: first, through decreased stomatal conductance (g s ) and increased soil water content (V SWC ) and second, through increased leaf internal CO 2 (C i ) and decreased stomatal limitations (S lim ). It is unclear if such findings from temperate grassland studies similarly pertain to warmer ecosystems with periodic water deficits. We tested these mechanisms in three important C 3 herbaceous species in a periodically dry Eucalyptus woodland and investigated how eCO 2 -induced photosynthetic enhancement varied with seasonal water availability, over a 3 year period. Leaf photosynthesis increased by 10%-50% with a 150 μmol mol -1 increase in atmospheric CO 2 across seasons. This eCO 2 -induced increase in photosynthesis was a function of seasonal water availability, given by recent precipitation and mean daily V SWC . The highest photosynthetic enhancement by eCO 2 (>30%) was observed during the most water-limited period, for example, with V SWC <0.07 in this sandy surface soil. Under eCO 2 there was neither a significant decrease in g s in the three herbaceous species, nor increases in V SWC , indicating no "water-savings effect" of eCO 2 . Periods of low V SWC showed lower g s (less than ≈ 0.12 mol m -2  s -1 ), higher relative S lim (>30%) and decreased C i under the ambient CO 2 concentration (aCO 2 ), with leaf photosynthesis strongly carboxylation-limited. The alleviation of S lim by eCO 2 was facilitated by increasing C i , thus yielding a larger photosynthetic enhancement during dry periods. We demonstrated that water availability, but not eCO 2 , controls g s and hence the magnitude of photosynthetic enhancement in the understory herbaceous plants. Thus, eCO 2 has the potential to alter vegetation functioning in a periodically dry woodland understory through changes in stomatal limitation to photosynthesis, not by the "water-savings effect" usually invoked in grasslands., (© 2017 John Wiley & Sons Ltd.)

Quantifying the adjustments of leaf respiration in response to seasonal temperature variation and climate warming is crucial because carbon loss from vegetation is a large but uncertain part of the global carbon cycle. We grew fast-growing Eucalyptus globulus Labill. trees exposed to +3 °C warming and elevated CO2 in 10-m tall whole-tree chambers and measured the temperature responses of leaf mitochondrial respiration, both in light (RLight) and in darkness (RDark), over a 20-40 °C temperature range and during two different seasons. RLight was assessed using the Laisk method. Respiration rates measured at a standard temperature (25 °C - R25) were higher in warm-grown trees and in the warm season, related to higher total leaf nitrogen (N) investment with higher temperatures (both experimental and seasonal), indicating that leaf N concentrations modulated the respiratory capacity to changes in temperature. Once differences in leaf N were accounted for, there were no differences in R25 but the Q10 (i.e., short-term temperature sensitivity) was higher in late summer compared with early spring. The variation in RLight between experimental treatments and seasons was positively correlated with carboxylation capacity and photorespiration. RLight was less responsive to short-term changes in temperature than RDark, as shown by a lower Q10 in RLight compared with RDark. The overall light inhibition of R was ∼40%. Our results highlight the dynamic nature of leaf respiration to temperature variation and that the responses of RLight do not simply mirror those of RDark. Therefore, it is important not to assume that RLight is the same as RDark in ecosystem models, as doing so may lead to large errors in predicting plant CO2 release and productivity., (© The Author 2017. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.)

Nitrogen (N) and phosphorus (P) have key roles in leaf metabolism, resulting in a strong coupling of chemical composition traits to metabolic rates in field-based studies. However, in such studies, it is difficult to disentangle the effects of nutrient supply per se on trait-trait relationships. Our study assessed how high and low N (5 mM and 0.4 mM, respectively) and P (1 mM and 2 μM, respectively) supply in 37 species from six plant functional types (PTFs) affected photosynthesis (A) and respiration (R) (in darkness and light) in a controlled environment. Low P supply increased scaling exponents (slopes) of area-based log-log A-N or R-N relationships when N supply was not limiting, whereas there was no P effect under low N supply. By contrast, scaling exponents of A-P and R-P relationships were altered by P and N supply. Neither R : A nor light inhibition of leaf R was affected by nutrient supply. Light inhibition was 26% across nutrient treatments; herbaceous species exhibited a lower degree of light inhibition than woody species. Because N and P supply modulates leaf trait-trait relationships, the next generation of terrestrial biosphere models may need to consider how limitations in N and P availability affect trait-trait relationships when predicting carbon exchange., (© 2017 The Authors. New Phytologist © 2017 New Phytologist Trust.)

Leaf-level photosynthetic processes and their environmental dependencies are critical for estimating CO2 uptake from the atmosphere. These estimates use biochemical-based models of photosynthesis that require accurate Rubisco kinetics. We investigated the effects of canopy position, elevated atmospheric CO2 [eC; ambient CO2 (aC)+240 ppm] and elevated air temperature (eT; ambient temperature (aT)+3 °C) on Rubisco content and activity together with the relationship between leaf N and Vcmax (maximal Rubisco carboxylation rate) of 7 m tall, soil-grown Eucalyptus globulus trees. The kinetics of E. globulus and tobacco Rubisco at 25 °C were similar. In vitro estimates of Vcmax derived from measures of E. globulus Rubisco content and kinetics were consistent, although slightly lower, than the in vivo rates extrapolated from gas exchange. In E. globulus, the fraction of N invested in Rubisco was substantially lower than for crop species and varied with treatments. Photosynthetic acclimation of E. globulus leaves to eC was underpinned by reduced leaf N and Rubisco contents; the opposite occurred in response to eT coinciding with growth resumption in spring. Our findings highlight the adaptive capacity of this key forest species to allocate leaf N flexibly to Rubisco and other photosynthetic proteins across differing canopy positions in response to future, warmer and elevated [CO2] climates., (© The Author 2017. Published by Oxford University Press on behalf of the Society for Experimental Biology.)

The response of terrestrial ecosystems to rising atmospheric CO2 concentration (Ca ), particularly under nutrient-limited conditions, is a major uncertainty in Earth System models. The Eucalyptus Free-Air CO2 Enrichment (EucFACE) experiment, recently established in a nutrient- and water-limited woodland presents a unique opportunity to address this uncertainty, but can best do so if key model uncertainties have been identified in advance. We applied seven vegetation models, which have previously been comprehensively assessed against earlier forest FACE experiments, to simulate a priori possible outcomes from EucFACE. Our goals were to provide quantitative projections against which to evaluate data as they are collected, and to identify key measurements that should be made in the experiment to allow discrimination among alternative model assumptions in a postexperiment model intercomparison. Simulated responses of annual net primary productivity (NPP) to elevated Ca ranged from 0.5 to 25% across models. The simulated reduction of NPP during a low-rainfall year also varied widely, from 24 to 70%. Key processes where assumptions caused disagreement among models included nutrient limitations to growth; feedbacks to nutrient uptake; autotrophic respiration; and the impact of low soil moisture availability on plant processes. Knowledge of the causes of variation among models is now guiding data collection in the experiment, with the expectation that the experimental data can optimally inform future model improvements., (© 2016 John Wiley & Sons Ltd.)

Simulations of photosynthesis by terrestrial biosphere models typically need a specification of the maximum carboxylation rate (Vcmax ). Estimating this parameter using A-Ci curves (net photosynthesis, A, vs intercellular CO2 concentration, Ci ) is laborious, which limits availability of Vcmax data. However, many multispecies field datasets include net photosynthetic rate at saturating irradiance and at ambient atmospheric CO2 concentration (Asat ) measurements, from which Vcmax can be extracted using a 'one-point method'. We used a global dataset of A-Ci curves (564 species from 46 field sites, covering a range of plant functional types) to test the validity of an alternative approach to estimate Vcmax from Asat via this 'one-point method'. If leaf respiration during the day (Rday ) is known exactly, Vcmax can be estimated with an r(2) value of 0.98 and a root-mean-squared error (RMSE) of 8.19 μmol m(-2) s(-1) . However, Rday typically must be estimated. Estimating Rday as 1.5% of Vcmax, we found that Vcmax could be estimated with an r(2) of 0.95 and an RMSE of 17.1 μmol m(-2) s(-1) . The one-point method provides a robust means to expand current databases of field-measured Vcmax , giving new potential to improve vegetation models and quantify the environmental drivers of Vcmax variation., (© 2015 The Authors. New Phytologist © 2015 New Phytologist Trust.)

Canopy leaf area, quantified by the leaf area index (L), is a crucial driver of forest productivity, water use and energy balance. Because L responds to environmental drivers, it can represent an important feedback to climate change, but its responses to rising atmospheric [CO2] and water availability of forests have been poorly quantified. We studied canopy leaf area dynamics for 28 months in a native evergreen Eucalyptus woodland exposed to free-air CO2 enrichment (the EucFACE experiment), in a subtropical climate where water limitation is common. We hypothesized that, because of expected stimulation of productivity and water-use efficiency, L should increase with elevated [CO2]. We estimated L from diffuse canopy transmittance, and measured monthly leaf litter production. Contrary to expectation, L did not respond to elevated [CO2]. We found that L varied between 1.10 and 2.20 across the study period. The dynamics of L showed a quick increase after heavy rainfall and a steady decrease during periods of low rainfall. Leaf litter production was correlated to changes in L, both during periods of decreasing L (when no leaf growth occurred) and during periods of increasing L (active shedding of old foliage when new leaf growth occurred). Leaf lifespan, estimated from mean L and total annual litter production, was up to 2 months longer under elevated [CO2] (1.18 vs. 1.01 years; P = 0.05). Our main finding that L was not responsive to elevated CO2 is consistent with other forest FACE studies, but contrasts with the positive response of L commonly predicted by many ecosystem models., (© 2015 John Wiley & Sons Ltd.)

Projections of future climate are highly sensitive to uncertainties regarding carbon (C) uptake and storage by terrestrial ecosystems. The Eucalyptus Free-Air CO2 Enrichment (EucFACE) experiment was established to study the effects of elevated atmospheric CO2 concentrations (eCO2 ) on a native mature eucalypt woodland with low fertility soils in southeast Australia. In contrast to other FACE experiments, the concentration of CO2 at EucFACE was increased gradually in steps above ambient (+0, 30, 60, 90, 120, and 150 ppm CO2 above ambient of ~400 ppm), with each step lasting approximately 5 weeks. This provided a unique opportunity to study the short-term (weeks to months) response of C cycle flux components to eCO2 across a range of CO2 concentrations in an intact ecosystem. Soil CO2 efflux (i.e., soil respiration or Rsoil ) increased in response to initial enrichment (e.g., +30 and +60 ppm CO2 ) but did not continue to increase as the CO2 enrichment was stepped up to higher concentrations. Light-saturated photosynthesis of canopy leaves (Asat ) also showed similar stimulation by elevated CO2 at +60 ppm as at +150 ppm CO2 . The lack of significant effects of eCO2 on soil moisture, microbial biomass, or activity suggests that the increase in Rsoil likely reflected increased root and rhizosphere respiration rather than increased microbial decomposition of soil organic matter. This rapid increase in Rsoil suggests that under eCO2, additional photosynthate was produced, transported belowground, and respired. The consequences of this increased belowground activity and whether it is sustained through time in mature ecosystems under eCO2 are a priority for future research., (© 2015 John Wiley & Sons Ltd.)

Arguments based on the biochemistry of photosynthesis predict a positive interaction between elevated atmospheric [CO2] and temperature on photosynthesis as well as growth. In contrast, few long-term studies on trees find greater stimulation of photosynthesis in response to elevated [CO2] at warmer compared with cooler temperatures. To test for CO2 × temperature interactions on leaf photosynthesis and whole-plant growth, we planted Eucalyptus globulus Labill. in climate-controlled chambers in the field at the Hawkesbury Forest Experiment research site, and investigated how photosynthetic enhancement changed across a range of seasonal temperatures. Trees were grown in a complete two-way factorial design with two CO2 concentrations (ambient and ambient + 240 ppm) and two temperatures (ambient and ambient + 3 °C) for 15 months until they reached ∼10 m height, after which they were harvested for biomass. There was significant enhancement of photosynthesis and growth with elevated [CO2], with the photosynthetic stimulation varying with season, but there was no significant effect of warming. Photosynthetic enhancement was higher in summer (+46% at 28 °C) than in winter (+14% at 20 °C). Photosynthetic enhancement as a function of leaf temperature was consistent with theoretical expectations, but was strongly mediated by the intercellular [CO2]/ambient [CO2] (Ci/Ca) ratio across seasons. Total tree biomass after 15 months was 66% larger in elevated CO2 (P = 0.017) with no significant warming effect detected. The fraction of biomass in coarse roots was reduced in warmed trees compared with ambient temperature controls, but there was no evidence of changed biomass allocation patterns in elevated CO2. We conclude that there are strong and consistent elevated CO2 effects on photosynthesis and biomass of E. globulus. It is crucial to consider stomatal conductance under a range of conditions to appraise the interactive effect of [CO2] and temperature on photosynthetic enhancement and subsequent implications for tree growth and forest productivity in future climates., (© The Author 2015. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.)

Differential species responses to atmospheric CO 2 concentration (Ca) could lead to quantitative changes in competition among species and community composition, with flow-on effects for ecosystem function. However, there has been little theoretical analysis of how elevated Ca (eC a) will affect plant competition, or how composition of plant communities might change. Such theoretical analysis is needed for developing testable hypotheses to frame experimental research. Here, we investigated theoretically how plant competition might change under eC a by implementing two alternative competition theories, resource use theory and resource capture theory, in a plant carbon and nitrogen cycling model. The model makes several novel predictions for the impact of eC a on plant community composition. Using resource use theory, the model predicts that eC a is unlikely to change species dominance in competition, but is likely to increase coexistence among species. Using resource capture theory, the model predicts that eC a may increase community evenness. Collectively, both theories suggest that eC a will favor coexistence and hence that species diversity should increase with eC a. Our theoretical analysis leads to a novel hypothesis for the impact of eC a on plant community composition. This hypothesis has potential to help guide the design and interpretation of eC a experiments.

Leaf photosynthetic CO2 responses can provide insight into how major nutrients, such as phosphorus (P), constrain leaf CO2 assimilation rates (Anet). However, triose-phosphate limitations are rarely employed in the classic photosynthesis model and it is uncertain as to what extent these limitations occur in field situations. In contrast to predictions from biochemical theory of photosynthesis, we found consistent evidence in the field of lower Anet in high [CO2] and low [O2 ] than at ambient [O2 ]. For 10 species of trees and shrubs across a range of soil P availability in Australia, none of them showed a positive response of Anet at saturating [CO2] (i.e. Amax) to 2 kPa O2. Three species showed >20% reductions in Amax in low [O2], a phenomenon potentially explained by orthophosphate (Pi) savings during photorespiration. These species, with largest photosynthetic capacity and Pi  > 2 mmol P m(-2), rely the most on additional Pi made available from photorespiration rather than species growing in P-impoverished soils. The results suggest that rarely used adjustments to a biochemical photosynthesis model are useful for predicting Amax and give insight into the biochemical limitations of photosynthesis rates at a range of leaf P concentrations. Phosphate limitations to photosynthetic capacity are likely more common in the field than previously considered., (© 2014 John Wiley & Sons Ltds.)

Leaf dark respiration (Rdark ) is an important yet poorly quantified component of the global carbon cycle. Given this, we analyzed a new global database of Rdark and associated leaf traits. Data for 899 species were compiled from 100 sites (from the Arctic to the tropics). Several woody and nonwoody plant functional types (PFTs) were represented. Mixed-effects models were used to disentangle sources of variation in Rdark . Area-based Rdark at the prevailing average daily growth temperature (T) of each site increased only twofold from the Arctic to the tropics, despite a 20°C increase in growing T (8-28°C). By contrast, Rdark at a standard T (25°C, Rdark (25) ) was threefold higher in the Arctic than in the tropics, and twofold higher at arid than at mesic sites. Species and PFTs at cold sites exhibited higher Rdark (25) at a given photosynthetic capacity (Vcmax (25) ) or leaf nitrogen concentration ([N]) than species at warmer sites. Rdark (25) values at any given Vcmax (25) or [N] were higher in herbs than in woody plants. The results highlight variation in Rdark among species and across global gradients in T and aridity. In addition to their ecological significance, the results provide a framework for improving representation of Rdark in terrestrial biosphere models (TBMs) and associated land-surface components of Earth system models (ESMs)., (© 2015 The Authors. New Phytologist © 2015 New Phytologist Trust.)

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., (© 2014 John Wiley & Sons Ltd.)

Climate change is resulting in increasing atmospheric [CO2], rising growth temperature (T), and greater frequency/severity of drought, with each factor having the potential to alter the respiratory metabolism of leaves. Here, the effects of elevated atmospheric [CO2], sustained warming, and drought on leaf dark respiration (R(dark)), and the short-term T response of R(dark) were examined in Eucalyptus globulus. Comparisons were made using seedlings grown under different [CO2], T, and drought treatments. Using high resolution T-response curves of R(dark) measured over the 15-65 °C range, it was found that elevated [CO2], elevated growth T, and drought had little effect on rates of R(dark) measured at T <35 °C and that there was no interactive effect of [CO2], growth T, and drought on T response of R(dark). However, drought increased R(dark) at high leaf T typical of heatwave events (35-45 °C), and increased the measuring T at which maximal rates of R(dark) occurred (Tmax) by 8 °C (from 52 °C in well-watered plants to 60 °C in drought-treated plants). Leaf starch and soluble sugars decreased under drought and elevated growth T, respectively, but no effect was found under elevated [CO2]. Elevated [CO2] increased the Q 10 of R(dark) (i.e. proportional rise in R(dark) per 10 °C) over the 15-35 °C range, while drought increased Q 10 values between 35 °C and 45 °C. Collectively, the study highlights the dynamic nature of the T dependence of R dark in plants experiencing future climate change scenarios, particularly with respect to drought and elevated [CO2]., (© The Author 2014. Published by Oxford University Press on behalf of the Society for Experimental Biology.)

We explored the impact of canopy position on leaf respiration (R) and associated traits in tree and shrub species growing in a lowland tropical rainforest in Far North Queensland, Australia. The range of traits quantified included: leaf R in darkness (RD) and in the light (RL; estimated using the Kok method); the temperature (T)-sensitivity of RD; light-saturated photosynthesis (Asat); leaf dry mass per unit area (LMA); and concentrations of leaf nitrogen (N), phosphorus (P), soluble sugars and starch. We found that LMA, and area-based N, P, sugars and starch concentrations were all higher in sun-exposed/upper canopy leaves, compared with their shaded/lower canopy and deep-shade/understory counterparts; similarly, area-based rates of RD, RL and Asat (at 28 °C) were all higher in the upper canopy leaves, indicating higher metabolic capacity in the upper canopy. The extent to which light inhibited R did not differ significantly between upper and lower canopy leaves, with the overall average inhibition being 32% across both canopy levels. Log-log RD-Asat relationships differed between upper and lower canopy leaves, with upper canopy leaves exhibiting higher rates of RD for a given Asat (both on an area and mass basis), as well as higher mass-based rates of RD for a given [N] and [P]. Over the 25-45 °C range, the T-sensitivity of RD was similar in upper and lower canopy leaves, with both canopy positions exhibiting Q10 values near 2.0 (i.e., doubling for every 10 °C rise in T) and Tmax values near 60 °C (i.e., T where RD reached maximal values). Thus, while rates of RD at 28 °C decreased with increasing depth in the canopy, the T-dependence of RD remained constant; these findings have important implications for vegetation-climate models that seek to predict carbon fluxes between tropical lowland rainforests and the atmosphere., (© The Author 2014. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.)

Eucalyptus species are grown widely outside of their native ranges in plantations on all vegetated continents of the world. We predicted that such a plantation species would show high potential for acclimation of photosynthetic traits across a wide range of growth conditions, including elevated [CO2] and climate warming. To test this prediction, we planted temperate Eucalyptus globulus Labill. seedlings in climate-controlled chambers in the field located >700 km closer to the equator than the nearest natural occurrence of this species. Trees were grown in a complete factorial combination of elevated CO2 concentration (eC; ambient [CO2] +240 ppm) and air warming treatments (eT; ambient +3 °C) for 15 months until they reached ca. 10 m height. There was little acclimation of photosynthetic capacity to eC and hence the CO2-induced photosynthetic enhancement was large (ca. 50%) in this treatment during summer. The warming treatment significantly increased rates of both carboxylation capacity (V(cmax)) and electron transport (Jmax) (measured at a common temperature of 25 °C) during winter, but decreased them significantly by 20-30% in summer. The photosynthetic CO2 compensation point in the absence of dark respiration (Γ*) was relatively less sensitive to temperature in this temperate eucalypt species than for warm-season tobacco. The temperature optima for photosynthesis and Jmax significantly changed by about 6 °C between winter and summer, but without further adjustment from early to late summer. These results suggest that there is an upper limit for the photosynthetic capacity of E. globulus ssp. globulus outside its native range to acclimate to growth temperatures above 25 °C. Limitations to temperature acclimation of photosynthesis in summer may be one factor that defines climate zones where E. globulus plantation productivity can be sustained under anticipated global environmental change., (© 2013 John Wiley & Sons Ltd.)

Predicted responses of transpiration to elevated atmospheric CO2 concentration (eCO2 ) are highly variable amongst process-based models. To better understand and constrain this variability amongst models, we conducted an intercomparison of 11 ecosystem models applied to data from two forest free-air CO2 enrichment (FACE) experiments at Duke University and Oak Ridge National Laboratory. We analysed model structures to identify the key underlying assumptions causing differences in model predictions of transpiration and canopy water use efficiency. We then compared the models against data to identify model assumptions that are incorrect or are large sources of uncertainty. We found that model-to-model and model-to-observations differences resulted from four key sets of assumptions, namely (i) the nature of the stomatal response to elevated CO2 (coupling between photosynthesis and stomata was supported by the data); (ii) the roles of the leaf and atmospheric boundary layer (models which assumed multiple conductance terms in series predicted more decoupled fluxes than observed at the broadleaf site); (iii) the treatment of canopy interception (large intermodel variability, 2-15%); and (iv) the impact of soil moisture stress (process uncertainty in how models limit carbon and water fluxes during moisture stress). Overall, model predictions of the CO2 effect on WUE were reasonable (intermodel μ = approximately 28% ± 10%) compared to the observations (μ = approximately 30% ± 13%) at the well-coupled coniferous site (Duke), but poor (intermodel μ = approximately 24% ± 6%; observations μ = approximately 38% ± 7%) at the broadleaf site (Oak Ridge). The study yields a framework for analysing and interpreting model predictions of transpiration responses to eCO2 , and highlights key improvements to these types of models., (© 2013 Blackwell Publishing Ltd.)

We investigated whether the degree of light inhibition of leaf respiration (R) differs among large Eucalyptus saligna grown in whole-tree chambers and exposed to present and future atmospheric [CO(2) ] and summer drought. Associated with month-to-month changes in temperature were concomitant changes in R in the light (R(light) ) and darkness (R(dark) ), with both processes being more temperature dependent in well-watered trees than under drought. Overall rates of R(light) and R(dark) were not significantly affected by [CO(2) ]. By contrast, overall rates of R(dark) (averaged across both [CO(2) ]) were ca. 25% lower under drought than in well-watered trees. During summer, the degree of light inhibition of leaf R was greater in droughted (ca. 80% inhibition) than well-watered trees (ca. 50% inhibition). Notwithstanding these treatment differences, an overall positive relationship was observed between R(light) and R(dark) when data from all months/treatments were combined (R(2)  = 0.8). Variations in R(light) were also positively correlated with rates of Rubisco activity and nitrogen concentration. Light inhibition resulted in a marked decrease in the proportion of light-saturated photosynthesis respired (i.e. reduced R/A(sat) ). Collectively, these results highlight the need to account for light inhibition when assessing impacts of global change drivers on the carbon economy of tree canopies., (© 2011 Blackwell Publishing Ltd.)

To investigate whether long-term elevated carbon dioxide concentration ([CO(2)]) causes declines in photosynthetic enhancement and leaf nitrogen (N) owing to limited soil fertility, we measured photosynthesis, carboxylation capacity and area-based leaf nitrogen concentration (N(a)) in Pinus taeda L. growing in a long-term free-air CO(2) enrichment (FACE) facility at an N-limited site. We also determined how maximum rates of carboxylation (V(cmax)) and electron transport (J(max)) varied with N(a) under elevated [CO(2)]. In trees exposed to elevated [CO(2)] for 5 to 9 years, the slope of the relationship between leaf photosynthetic capacity (A(net-Ca)) and N(a) was significantly reduced by 37% in 1-year-old needles, whereas it was unaffected in current-year needles. The slope of the relationships of both V(cmax) and J(max) with N(a) decreased in 1-year-old needles after up to 9 years of growth in elevated [CO(2)], which was accompanied by a 15% reduction in N allocation to the carboxylating enzyme. Nitrogen fertilization (110 kg N ha(-1)) in the ninth year of exposure to elevated [CO(2)] restored the slopes of the relationships of V(cmax) and J(max) with N(a) to those of control trees (i.e., in ambient [CO(2)]). The J(max):V(cmax) ratio was unaffected by either [CO(2)] or N fertilization. Changes in the apparent allocation of N to photosynthetic components may be an important adjustment in pines exposed to elevated [CO(2)] on low-fertility sites. We conclude that fundamental relationships between photosynthesis or its component processes with N(a) may be altered in aging pine needles after more than 5 years of exposure to elevated atmospheric [CO(2)].

Few studies have examined the effects of elevated CO2 concentration ([CO2]) on the physiology of intact forest canopies, despite the need to understand how leaf-level responses can be aggregated to assess effects on whole-canopy functioning. We examined the long-term effects of elevated [CO2] (ambient + 200 ppm CO2) on two age classes of needles in the upper and lower canopy of Pinus taeda L. during the second through sixth year of exposure to elevated [CO2] in free-air (free-air CO2 enrichment (FACE)) in North Carolina, USA. Strong photosynthetic enhancement in response to elevated [CO2] (e.g., +60% across age classes and canopy locations) was observed across the years. This stimulation was 33% greater for current-year needles than for 1-year-old needles in the fifth and sixth years of treatment. Although photosynthetic stimulation in response to elevated [CO2] was maintained through the sixth year of exposure, we found evidence of concurrent down-regulation of Rubisco and electron transport capacity in the upper-canopy sunlit leaves. The lower canopy showed no evidence of down-regulation. The upper canopy down-regulated carboxylation capacity (Vcmax) and electron transport capacity (Jmax) by about 17-20% in 1-year-old needles; however, this response was significant across sampling years only for Jmax in 1-year-old needles (P < 0.02). A reduction in leaf photosynthetic capacity in aging conifer needles at the canopy top could have important consequences for canopy carbon balance and global carbon sinks because 1-year-old sunlit needles contribute a major proportion of the annual carbon balance of these conifers. Our finding of a significant interaction between canopy position and CO2 treatment on the biochemical capacity for CO2 assimilation suggests that it is important to take canopy position and needle aging into account because morphologically and physiologically distinct leaves could respond differently to elevated [CO2].