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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.

Summary: Photosynthetic water‐use efficiency (WUE) describes the link between terrestrial carbon (C) and water cycles. Estimates of intrinsic WUE (iWUE) from gas exchange and C isotopic composition (δ13C) differ due to an internal conductance in the leaf mesophyll (gm) that is variable and seldom computed.We present the first direct estimates of whole‐tree gm, together with iWUE from whole‐tree gas exchange and δ13C of the phloem (δ13Cph). We measured gas exchange, online 13C‐discrimination, and δ13Cph monthly throughout spring, summer, and autumn in Eucalyptus tereticornis grown in large whole‐tree chambers. Six trees were grown at ambient temperatures and six at a 3°C warmer air temperature; a late‐summer drought was also imposed.Drought reduced whole‐tree gm. Warming had few direct effects, but amplified drought‐induced reductions in whole‐tree gm. Whole‐tree gm was similar to leaf gm for these same trees. iWUE estimates from δ13Cph agreed with iWUE from gas exchange, but only after incorporating gm. δ13Cph was also correlated with whole‐tree 13C‐discrimination, but offset by −2.5 ± 0.7‰, presumably due to post‐photosynthetic fractionations.We conclude that δ13Cph is a good proxy for whole‐tree iWUE, with the caveats that post‐photosynthetic fractionations and intrinsic variability of gm should be incorporated to provide reliable estimates of this trait in response to abiotic stress. [ABSTRACT FROM AUTHOR]

Summary: Thermoregulation of leaf temperature (Tleaf) may foster metabolic homeostasis in plants, but the degree to which Tleaf is moderated, and under what environmental contexts, is a topic of debate. Isotopic studies inferred the temperature of photosynthetic carbon assimilation to be a constant value of c. 20°C; by contrast, leaf biophysical theory suggests a strong dependence of Tleaf on environmental drivers. Can this apparent disparity be reconciled?We continuously measured Tleaf and whole‐crown net CO2 uptake for Eucalyptus parramattensis trees growing in field conditions in whole‐tree chambers under ambient and +3°C warming conditions, and calculated assimilation‐weighted leaf temperature (TL‐AW) across 265 d, varying in air temperature (Tair) from −1 to 45°C. We compared these data to TL‐AW derived from wood cellulose δ18O.Tleaf exhibited substantial variation driven by Tair, light intensity, and vapor pressure deficit, and Tleaf was strongly linearly correlated with Tair with a slope of c. 1.0. TL‐AW values calculated from cellulose δ18O vs crown fluxes were remarkably consistent; both varied seasonally and in response to the warming treatment, tracking variation in Tair.The leaves studied here were nearly poikilothermic, with no evidence of thermoregulation of Tleaf towards a homeostatic value. Importantly, this work supports the use of cellulose δ18O to infer TL‐AW, but does not support the concept of strong homeothermic regulation of Tleaf See also the Commentary on this article by Cavaleri, 228: 1455–1457. [ABSTRACT FROM AUTHOR]

Summary: Autotrophic respiration is a major driver of the global C cycle and may contribute a positive climate warming feedback through increased atmospheric concentrations of CO2. The extent of this feedback depends on plants' ability to acclimate respiration to maintain a constant carbon use efficiency (CUE).We quantified respiratory partitioning of gross primary production (GPP) and CUE of field‐grown trees in a long‐term warming experiment (+3°C). We delivered a 13C–CO2 pulse to whole tree crowns and chased that pulse in the respiration of leaves, whole crowns, roots, and soil. We also measured the isotopic composition of soil microbial biomass and the respiration rates of leaves and whole crowns.We documented homeostatic respiratory acclimation of foliar and whole‐crown respiration rates; the trees adjusted to experimental warming such that leaf‐level respiration rates were not increased. Experimental warming had no detectable impact on respiratory partitioning or mean residence times. Of the 13C label acquired by the trees, aboveground respiration consumed 10%, belowground respiration consumed 40%, and the remaining 50% was retained.Experimental warming of +3°C did not alter respiratory partitioning at the scale of entire trees, suggesting that complete acclimation of respiration to warming is likely to dampen a positive climate warming feedback. See also the Commentary on this article by Ryan & Asao, 222: 1167–1170. [ABSTRACT FROM AUTHOR]

Summary: The Kok and Laisk techniques can both be used to estimate light respiration Rlight. We investigated whether responses of Rlight to short‐ and long‐term changes in leaf temperature depend on the technique used to estimate Rlight.We grew Eucalyptus tereticornis in whole‐tree chambers under ambient temperature (AT) or AT + 3°C (elevated temperature, ET). We assessed dark respiration Rdark and light respiration with the Kok (RKok) and Laisk (RLaisk) 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 Rdark, RKok or RLaisk. Although the thermal sensitivities of RKok or RLaisk were similar, RKok was higher than RLaisk. We found negative values of RLaisk at the lowest measurement temperatures, indicating positive net CO2 uptake, which we propose may be related to phosphoenolpyruvate carboxylase activity. Light suppression of Rdark 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 Rlight or light suppression of Rdark between 20 and 35°C. Negative rates of RLaisk imply that this method may become less reliable at low temperatures. [ABSTRACT FROM AUTHOR]