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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.

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.

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 CO2 response curves, including data from 141 C3 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.

Earth system models (ESMs) use photosynthetic capacity, indexed by the maximum Rubisco carboxylation rate (Vcmax ), to simulate carbon assimilation and typically rely on empirical estimates, including an assumed dependence on leaf nitrogen determined from soil fertility. In contrast, new theory, based on biochemical coordination and co-optimization of carboxylation and water costs for photosynthesis, suggests that optimal Vcmax can be predicted from climate alone, irrespective of soil fertility. Here, we develop this theory and find it captures 64% of observed variability in a global, field-measured Vcmax dataset for C3 plants. Soil fertility indices explained substantially less variation (32%). These results indicate that environmentally regulated biophysical constraints and light availability are the first-order drivers of global photosynthetic capacity. Through acclimation and adaptation, plants efficiently utilize resources at the leaf level, thus maximizing potential resource use for growth and reproduction. Our theory offers a robust strategy for dynamically predicting photosynthetic capacity in ESMs.

The capacity for terrestrial ecosystems to sequester additional carbon (C) with rising CO2 concentrations depends on soil nutrient availability1,2. Previous evidence suggested that mature forests growing on phosphorus (P)-deprived soils had limited capacity to sequester extra biomass under elevated CO2 (refs. 3–6), but uncertainty about ecosystem P cycling and its CO2 response represents a crucial bottleneck for mechanistic prediction of the land C sink under climate change7. Here, by compiling the first comprehensive P budget for a P-limited mature forest exposed to elevated CO2, 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 CO2 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 CO2 fertilization of tree growth and will guide the development of Earth system models to predict future long-term C storage.Microbial pre-emption of mineralized soil P limits the capacity of trees for increased P uptake and assimilation under elevated CO2 and therefore restricts their capacity to sequester extra C. [ABSTRACT FROM AUTHOR]

Summary: Optimal stomatal theory predicts that stomata operate to maximise photosynthesis (Anet) 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 CO2 (eCO2), 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 eCO2 on iWUE and its components Anet and stomatal conductance (gs). 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 gs, but greater Anet, responses to eCO2 in gymnosperms compared with angiosperm PFTs.We found that iWUE increased in proportion to increasing eCO2 in all PFTs, and that increases in Anet had stronger effects than reductions in gs. The USO model correctly captured stomatal behaviour with eCO2 across most datasets. The chief difference among PFTs was a lower stomatal slope parameter (g1) for the gymnosperm, compared with angiosperm, species.Land surface models can use the USO model to describe stomatal behaviour under changing atmospheric CO2 conditions. [ABSTRACT FROM AUTHOR]

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 [CO2] treatment. Our combined approaches improved predictions of transpiration and enhanced the simulated magnitude of the CO2 fertilization effect on gross primary productivity. The competing approaches also worked consistently along axes of change in soil moisture, leaf area, and [CO2]. 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. [ABSTRACT FROM AUTHOR]

Summary: 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 (ToptA) than temperate species. There was acclimation of ToptA 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 Vcmax25 and Jmax25. The temperature sensitivity of respiration (Q10) 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. [ABSTRACT FROM AUTHOR]

Summary: 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 (Anet25) was larger with warming in tropical climates than cooler ones. Respiration at 25°C (R25) 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 Anet25 and R25 reduced c. 30–40% with > 10°C warming. While standardised rates of carboxylation (Vcmax25) and electron transport (Jmax25) adjusted to warming, the magnitude of this adjustment was not related to warming amount (range 0.6–16°C). The temperature optimum of photosynthesis (ToptA) 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. [ABSTRACT FROM AUTHOR]

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 (eCO2), but consensus has yet to be reached on the extent of this limitation. Here, based on data from experiments that manipulated both CO2 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 eCO2. We show that low P availability attenuated plant photosynthetic response to eCO2 by approximately one‐quarter, leading to a reduced, but still positive photosynthetic response to eCO2 compared to those under high P availability. Furthermore, low P limited plant aboveground, belowground, and total biomass responses to eCO2, by 14.7%, 14.3%, and 12.4%, respectively, equivalent to an approximate halving of the eCO2 responses observed under high P availability. In comparison, low P availability did not significantly alter the eCO2‐induced changes in plant tissue nutrient concentration, suggesting tissue nutrient flexibility is an important mechanism allowing biomass response to eCO2 under low P availability. Low P significantly reduced the eCO2‐induced increase in leaf area by 14.3%, mirroring the aboveground biomass response, but low P did not affect the eCO2‐induced increase in root length. Woody plants exhibited stronger attenuation effect of low P on aboveground biomass response to eCO2 than non‐woody plants, while plants with different mycorrhizal associations showed similar responses to low P and eCO2 interaction. This meta‐analysis highlights crucial data gaps in capturing plant responses to eCO2 and low P availability. Field‐based experiments with longer‐term exposure of both CO2 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 eCO2 under P limitation, thereby improving projections of future global change impacts. [ABSTRACT FROM AUTHOR]

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 (R root) 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 R root 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 (Q 10) and the degree of acclimation of R root. Regardless of biome type, we found no consistent pattern in the short-term temperature responses of R root to elevated growth temperature: a significant reduction in the temperature response of R root 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, R root was positively correlated with root tissue nitrogen concentration (mg g−1), while Q 10 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 R root to warming is species-specific and suggest that root tissue density is a useful predictor of R root and its thermal responses in rainforest tree seedlings. [ABSTRACT FROM AUTHOR]

The response of mature forest ecosystems to a rising atmospheric carbon dioxide concentration (Ca) is a major uncertainty in projecting the future trajectory of the Earth's climate. Although leaf-level net photosynthesis is typically stimulated by exposure to elevated Ca (e Ca), it is unclear how this stimulation translates into carbon cycle responses at the ecosystem scale. Here we estimate a key component of the carbon cycle, the gross primary productivity (GPP), of a mature native eucalypt forest exposed to free-air CO2 enrichment (the EucFACE experiment). In this experiment, light-saturated leaf photosynthesis increased by 19 % in response to a 38 % increase in Ca. We used the process-based forest canopy model, MAESPA, to upscale these leaf-level measurements of photosynthesis with canopy structure to estimate the GPP and its response to e Ca. We assessed the direct impact of e Ca , as well as the indirect effect of photosynthetic acclimation to e Ca and variability among treatment plots using different model scenarios. At the canopy scale, MAESPA estimated a GPP of 1574 g C m -2 yr -1 under ambient conditions across 4 years and a direct increase in the GPP of + 11 % in response to e Ca. The smaller canopy-scale response simulated by the model, as compared with the leaf-level response, could be attributed to the prevalence of RuBP regeneration limitation of leaf photosynthesis within the canopy. Photosynthetic acclimation reduced this estimated response to 10 %. After taking the baseline variability in the leaf area index across plots in account, we estimated a field GPP response to e Ca of 6 % with a 95 % confidence interval (- 2 %, 14 %). These findings highlight that the GPP response of mature forests to e Ca is likely to be considerably lower than the response of light-saturated leaf photosynthesis. Our results provide an important context for interpreting the e Ca responses of other components of the ecosystem carbon cycle. [ABSTRACT FROM AUTHOR]

To quantify stem respiration (RS) under elevated CO2 (eCO2), stem CO2 efflux (EA) and CO2 flux through the xylem (FT) should be accounted for, because part of respired CO2 is transported upwards with the sap solution. However, previous studies have used EA as a proxy of RS, which could lead to equivocal conclusions. Here, to test the effect of eCO2 on RS, both EA and FT were measured in a free‐air CO2 enrichment experiment located in a mature Eucalyptus native forest. Drought stress substantially reduced EA and RS, which were unaffected by eCO2, likely as a consequence of its neutral effect on stem growth in this phosphorus‐limited site. However, xylem CO2 concentration measured near the stem base was higher under eCO2, and decreased along the stem resulting in a negative contribution of FT to RS, whereas the contribution of FT to RS under ambient CO2 was positive. Negative FT indicates net efflux of CO2 respired below the monitored stem segment, likely coming from the roots. Our results highlight the role of nutrient availability on the dependency of RS on eCO2 and suggest stimulated root respiration under eCO2 that may shift vertical gradients in xylem [CO2] confounding the interpretation of EA measurements. In phosphorus‐limited sites, stem respiratory metabolism will remain unaffected by elevated CO2 as long as stem growth does not respond to it. Stimulated carbon allocation belowground and greater root respiration under elevated CO2 may shift vertical gradients in xylem [CO2] confounding the interpretation of stem CO2 efflux measurements. [ABSTRACT FROM AUTHOR]

The response of mature forest ecosystems to rising atmospheric carbon dioxide concentration (Ca) is a major uncertainty in projecting the future trajectory of the Earth's climate. Although leaf-level net photosynthesis is typically stimulated by exposure to elevated Ca (eCa), it is unclear how this stimulation translates into carbon cycle responses at whole-ecosystem scale. Here we estimate a key component of the carbon cycle, the gross primary productivity (GPP), of a mature native Eucalypt forest exposed to Free Air CO2 Enrichment (the EucFACE experiment). In this experiment, light-saturated leaf photosynthesis increased by 19 % in response to a 38 % increase in Ca. We used the process-based forest canopy model, MAESPA, to upscale these leaf-level measurements of photosynthesis with canopy structure to estimate Gross Primary Production (GPP) and its response to eCa. We assessed the direct impact of eCa, as well as the indirect effect of photosynthetic acclimation to eCa and variability among treatment plots via different model scenarios. At the canopy scale, MAESPA estimated a GPP of 1574 g C m-2 yr-1 under ambient conditions across four years and a direct increase in GPP of +11 % in response to eCa. The smaller canopy-scale response simulated by the model, as compared to the leaf-level response, could be attributed to the prevalence of RuBP-regeneration limitation of leaf photosynthesis within the canopy. Photosynthetic acclimation reduced this estimated response to 10 %. Considering variability in leaf area index across plots, we estimated a mean GPP response to eCa of 6 % with a 95 % CI of (-2 %, 14 %). These findings highlight that the GPP response of mature forests to eCa is likely to be considerably lower than the response of light-saturated leaf photosynthesis. Our results provide an important context for interpreting eCa responses of other components of the ecosystem carbon cycle. [ABSTRACT FROM AUTHOR]

Elevated CO2 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 CO2 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 CO2. 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 CO2 on leaf P. By contrast, elevated CO2 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 CO2 and N:P ratios were not different in mature green leaves (CO2 by Age effect, P = 0.02). Over time, leaf N and P concentrations in the upper canopy slightly declined in both CO2 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 CO2 (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. [ABSTRACT FROM AUTHOR]

Rising atmospheric CO2 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 CO2 (eCO2), which may reduce the magnitude of photosynthetic enhancement. We tested two hypotheses as to why leaf N is reduced under eCO2: (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 eCO2. 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 CO2 Enrichment (EucFACE) for five years. Fully expanded young and mature leaves were both measured for net photosynthesis, photorespiration, total leaf N, nitrate (NO3-) concentrations, carbohydrates and NO3- reductase activity to test these hypotheses. Foliar N concentrations declined by 8% under eCO2 in new leaves, while the NO3- fraction and total carbohydrate concentrations remained unchanged by CO2 treatment for either new or mature leaves. Photorespiration decreased 31% under eCO2 supplying less reductant, and in situ NO3- reductase activity was concurrently reduced (−34%) in eCO2, especially in new leaves during summer periods. Hence, NO3- 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 NO3- reductase activity due to lower photorespiration in eCO2 can contribute to understanding how eCO2‐induced photosynthetic enhancement may be lower than previously expected. We suggest that large‐scale vegetation models simulating effects of eCO2 on N biogeochemistry include both mechanisms, especially where NO3- is major N source to the dominant vegetation and where leaf flushing and emergence occur in temperatures that promote high photorespiration rates. Nearly, every experiment in elevated CO2 (eCO2) reported a decline in leaf N concentration, and a turndown in photorespiration has been suggested as one explanation. When increased atmospheric CO2 concentration enhances photosynthetic activity, carbon assimilation and carbohydrate production are favoured but photorespiration is inhibited. Photorespiration supplies reductant, nicotinamide adenine dinucleotide hydride (NADH), for the first step of nitrate (NO3-) assimilation. When photorespiration is inhibited under eCO2, malate export is reduced, and thus, availability of NADH also declines. This results in lower conversion of NO3- into nitrite (NO2−) by enzyme nitrate reductase (NR), a likely cause for declining leaf N concentrations in mature Eucalyptus trees in free‐air (FACE), especially for recently produced leaves. [ABSTRACT FROM AUTHOR]

Abstract: 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 (Topt) of photosynthesis and Jmax 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 Topt of Jmax 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. [ABSTRACT FROM AUTHOR]

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]

Contents986I.987II.987III.988IV.991V.992VI.995VII.997VIII.998References998 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 CO2 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 CO2 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. [ABSTRACT FROM AUTHOR]

Elevated atmospheric CO2 ( eCO2) is expected to reduce the impacts of drought and increase photosynthetic rates via two key mechanisms: first, through decreased stomatal conductance (gs) and increased soil water content ( VSWC) and second, through increased leaf internal CO2 (Ci) and decreased stomatal limitations (Slim). 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 C3 herbaceous species in a periodically dry Eucalyptus woodland and investigated how eCO2-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 CO2 across seasons. This eCO2-induced increase in photosynthesis was a function of seasonal water availability, given by recent precipitation and mean daily VSWC. The highest photosynthetic enhancement by eCO2 (>30%) was observed during the most water-limited period, for example, with VSWC <0.07 in this sandy surface soil. Under eCO2 there was neither a significant decrease in gs in the three herbaceous species, nor increases in VSWC, indicating no 'water-savings effect' of eCO2. Periods of low VSWC showed lower gs (less than ≈ 0.12 mol m−2 s−1), higher relative Slim (>30%) and decreased Ci under the ambient CO2 concentration ( aCO2), with leaf photosynthesis strongly carboxylation-limited. The alleviation of Slim by eCO2 was facilitated by increasing Ci, thus yielding a larger photosynthetic enhancement during dry periods. We demonstrated that water availability, but not eCO2, controls gs and hence the magnitude of photosynthetic enhancement in the understory herbaceous plants. Thus, eCO2 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. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Equatorial species may have less capacity to adjust to warmer temperatures because they have adapted to stable thermal conditions year-round compared to species from cooler climates (higher latitudes) where temperature fluctuates strongly among the seasons. Given that temperature responses of plants to warming are fundamental in any environment, how are plant function and metabolism linked to seasonal variations and climate? Understanding how temperature dependence of physiological processes is related to climate variation is critical to predict how species will adjust to warmer temperatures, what their thermal tolerances are, and ultimately lead to understanding how future species distributions may change. Plant carbon metabolism and climate change: elevated CO 2 and temperature impacts on photosynthesis, photorespiration and respiration. [Extracted from the article]

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. [ABSTRACT FROM AUTHOR]

Rising levels of atmospheric CO2 concentration ( Ca) and simultaneous climate change profoundly affect plant physiological performance while challenging our ability to estimate vegetation-atmosphere fluxes. To predict rates of water and carbon exchange between vegetation and the atmosphere, we require a formulation for stomatal conductance ( gs) that captures the multidimensional response of stomata to changing environmental conditions. The unified stomatal optimization (USO) theory provides a formulation for gs with the ability to predict the response of gs to novel environmental conditions such as elevated Ca (e Ca), warmer temperatures and/or changing water availability., We tested for the effect of e Ca and seasonally varying climate on stomatal behaviour, as defined by the USO theory, during the first year of free-air CO2 enrichment in a native eucalypt woodland (the EucFACE experiment). We hypothesized that under e Ca, gs would decrease and photosynthesis ( Anet) would increase, but fundamental stomatal behaviour described in the USO model would remain unchanged. We also predicted that the USO slope parameter g1 would increase with temperature and water availability. Over 20 months, we performed quarterly gas exchange campaigns encompassing a wide range of temperatures and water availabilities. We measured gs, Anet and leaf water potential (Ψ) at mid-morning, midday and pre-dawn (Ψ only) under ambient and e Ca and prevailing climatic conditions, at the tree tops (20 m height)., We found that e Ca induced a 20% reduction in stomatal conductance under non-limiting water availability, enhanced mid-morning Anet by 24% in three out of five measurement campaigns and had no significant effect on Ψ. The parameter g1 was conserved under e Ca, weakly increased with temperature and did not respond to increasing water availability., Our results suggest that under e Ca and variable rainfall, mature eucalypt trees exhibit a conservative water-use strategy, but this strategy may be modified by growth temperature. We show that the USO theory successfully predicts coupling of carbon uptake and water loss in future atmospheric conditions in a native woodland and thus could be incorporated into ecosystem-scale and global vegetation models. [ABSTRACT FROM AUTHOR]

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 r2 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 r2 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. [ABSTRACT FROM AUTHOR]

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 Euc FACE 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. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Differential species responses to atmospheric CO2 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 ( eCa) 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 eCa 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 eCa on plant community composition. Using resource use theory, the model predicts that eCa is unlikely to change species dominance in competition, but is likely to increase coexistence among species. Using resource capture theory, the model predicts that eCa may increase community evenness. Collectively, both theories suggest that eCa will favor coexistence and hence that species diversity should increase with eCa. Our theoretical analysis leads to a novel hypothesis for the impact of eCa on plant community composition. This hypothesis has potential to help guide the design and interpretation of eCa experiments. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

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, Rdark25) 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 Rdark25 at a given photosynthetic capacity ( Vcmax25) or leaf nitrogen concentration ([N]) than species at warmer sites. Rdark25 values at any given Vcmax25 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). [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]

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 ( Vcmax) 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. [ABSTRACT FROM AUTHOR]

Atmospheric carbon dioxide concentration ( Ca) has a direct and measurable effect on plant growth. However, it does not affect all plant species equally, which could lead to shifts in competitive dominance of species in ecosystems., We used a dynamic plant carbon-nitrogen model to systematically examine how species traits affect the long-term Ca responsiveness of C3 plants when growing as established monocultures in the field. The model was tested against responses of 7 C3 herbaceous species growing in a free-air Ca enrichment ( FACE) experiment ( Bio CON) in Minnesota, USA., Model simulations showed that several species traits affected the Ca response strongly, giving rise to a number of testable hypotheses about interspecific differences in responsiveness to Ca. The largest responses to rising Ca were obtained for species with low carbon-use efficiency (net primary production: gross primary production ratio), low foliar carbon allocation, low stomatal conductance, low instantaneous photosynthetic nitrogen use efficiency and low specific leaf area., In general, our model predicted that, for established plants growing in resource-limited field conditions, species with slow growth rates would be most responsive to elevated Ca. This prediction was supported by data from the Bio CON experiment., Our model also predicts that, for young plants growing in non-resource-limited conditions, species with high growth rates will be most responsive to elevated Ca. This difference in species ranking under different resource availabilities is largely explained by the indirect effects of Ca on leaf area. Leaf-area feedbacks favour fast-growing species the most during leaf-area expansion, but following stand maturation they favour slow-growing species the most., These results imply that species that respond strongly to elevated Ca in short-term (non-resource-limited) glasshouse experiments are unlikely to also be the most responsive in resource-limited field conditions, and therefore that we cannot directly extrapolate from glasshouse experiments to predict which species will be most responsive to elevated Ca in the long term. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

SUMMARY 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 [CO2] and summer drought. Associated with month-to-month changes in temperature were concomitant changes in R in the light ( Rlight) and darkness ( Rdark), with both processes being more temperature dependent in well-watered trees than under drought. Overall rates of Rlight and Rdark were not significantly affected by [CO2]. By contrast, overall rates of Rdark (averaged across both [CO2]) 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 Rlight and Rdark when data from all months/treatments were combined (R2 = 0.8). Variations in Rlight 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/ Asat). 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. [ABSTRACT FROM AUTHOR]

Rising atmospheric concentrations of CO2 ( Ca) can reduce stomatal conductance and transpiration rate in trees, but the magnitude of this effect varies considerably among experiments. The theory of optimal stomatal behaviour predicts that the ratio of photosynthesis to transpiration (instantaneous transpiration efficiency, ITE) should increase in proportion to Ca. We hypothesized that plants regulate stomatal conductance optimally in response to rising Ca. We tested this hypothesis with data from young Eucalyptus saligna Sm. trees grown in 12 climate-controlled whole-tree chambers for 2 years at ambient and elevated Ca. Elevated Ca was ambient + 240 ppm, 60% higher than ambient Ca. Leaf-scale gas exchange was measured throughout the second year of the study and leaf-scale ITE increased by 60% under elevated Ca, as predicted. Values of leaf-scale ITE depended strongly on vapour pressure deficit ( D) in both CO2 treatments. Whole-canopy CO2 and H2O fluxes were also monitored continuously for each chamber throughout the second year. There were small differences in D between Ca treatments, which had important effects on values of canopy-scale ITE. However, when Ca treatments were compared at the same D, canopy-scale ITE was consistently increased by 60%, again as predicted. Importantly, leaf and canopy-scale ITE were not significantly different, indicating that ITE was not scale-dependent. Observed changes in transpiration rate could be explained on the basis that ITE increased in proportion to Ca. The effect of elevated Ca on photosynthesis increased with rising D. At high D, Ca had a large effect on photosynthesis and a small effect on transpiration rate. At low D, in contrast, there was a small effect of Ca on photosynthesis, but a much larger effect on transpiration rate. If shown to be a general response, the proportionality of ITE with Ca will allow us to predict the effects of Ca on transpiration rate. [ABSTRACT FROM AUTHOR]

Leaf responses to elevated atmospheric CO2 concentration (Ca) are central to models of forest CO2 exchange with the atmosphere and constrain the magnitude of the future carbon sink. Estimating the magnitude of primary productivity enhancement of forests in elevated Ca requires an understanding of how photosynthesis is regulated by diffusional and biochemical components and up-scaled to entire canopies. To test the sensitivity of leaf photosynthesis and stomatal conductance to elevated Ca in time and space, we compiled a comprehensive dataset measured over 10 years for a temperate pine forest of P inus taeda, but also including deciduous species, primarily L iquidambar styraciflua. We combined over one thousand controlled-response curves of photosynthesis as a function of environmental drivers (light, air Ca and temperature) measured at canopy heights up to 20 m over 11 years (1996-2006) to generate parameterizations for leaf-scale models for the Duke free-air CO2 enrichment ( FACE) experiment. The enhancement of leaf net photosynthesis ( Anet) in P . taeda by elevated Ca of +200 μmol mol−1 was 67% for current-year needles in the upper crown in summer conditions over 10 years. Photosynthetic enhancement of P . taeda at the leaf-scale increased by two-fold from the driest to wettest growing seasons. Current-year pine foliage Anet was sensitive to temporal variation, whereas previous-year foliage Anet was less responsive and overall showed less enhancement (+30%). Photosynthetic downregulation in overwintering upper canopy pine needles was small at average leaf N ( Narea), but statistically significant. In contrast, co-dominant and subcanopy L . styraciflua trees showed Anet enhancement of 62% and no Anet- Narea adjustments. Various understory deciduous tree species showed an average Anet enhancement of 42%. Differences in photosynthetic responses between overwintering pine needles and subcanopy deciduous leaves suggest that increased Ca has the potential to enhance the mixed-species composition of planted pine stands and, by extension, naturally regenerating pine-dominated stands. [ABSTRACT FROM AUTHOR]

Models of vegetation function are widely used to predict the effects of climate change on carbon, water and nutrient cycles of terrestrial ecosystems, and their feedbacks to climate. Stomatal conductance, the process that governs plant water use and carbon uptake, is fundamental to such models. In this paper, we reconcile two long-standing theories of stomatal conductance. The empirical approach, which is most commonly used in vegetation models, is phenomenological, based on experimental observations of stomatal behaviour in response to environmental conditions. The optimal approach is based on the theoretical argument that stomata should act to minimize the amount of water used per unit carbon gained. We reconcile these two approaches by showing that the theory of optimal stomatal conductance can be used to derive a model of stomatal conductance that is closely analogous to the empirical models. Consequently, we obtain a unified stomatal model which has a similar form to existing empirical models, but which now provides a theoretical interpretation for model parameter values. The key model parameter, g, is predicted to increase with growth temperature and with the marginal water cost of carbon gain. The new model is fitted to a range of datasets ranging from tropical to boreal trees. The parameter g is shown to vary with growth temperature, as predicted, and also with plant functional type. The model is shown to correctly capture responses of stomatal conductance to changing atmospheric CO, and thus can be used to test for stomatal acclimation to elevated CO. The reconciliation of the optimal and empirical approaches to modelling stomatal conductance is important for global change biology because it provides a simple theoretical framework for analyzing, and simulating, the coupling between carbon and water cycles under environmental change. [ABSTRACT FROM AUTHOR]