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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]

Fine roots are a key component of carbon and nutrient dynamics in forest ecosystems. Rising atmospheric [CO2] (eCO2) is likely to alter the production and activity of fine roots, with important consequences for forest carbon storage. Yet empirical evidence of the role of eCO2 in driving root dynamics is limited, particularly for grassy woodlands, an ecosystem type of global importance.We sampled fine roots across seasons over a 2‐year period to examine the effects of eCO2 on their biomass, production, turnover and functional traits in a native mature grassy Eucalyptus woodland in eastern Australia (EucFACE).Fine root biomass, production and turnover varied greatly through time, increasing as soil water content declined. Despite a lack of consistent effects of eCO2 on fine root biomass, production or turnover across the 2‐year sampling period, we found enhanced production pulses under eCO2 between 10‐ and 30‐cm soil depth. In addition, eCO2 led to greater carbon and phosphorus concentrations in fine roots and increased root diameter, but no detectable effects on other morphological traits.Synthesis. We found minor quantitative effects of eCO2 on fine root biomass dynamics that were largely driven by temporal variations in soil water availability. Our results suggest that in this mature grassy woodland, and perhaps also in other similar forested ecosystem types, eCO2 effects are small and transient. This also implies a limited ability of these systems to mitigate climate change through below‐ground mechanisms. A free Plain Language Summary can be found within the Supporting Information of this article. [ABSTRACT FROM AUTHOR]

The triose phosphate utilization (TPU) rate has been identified as one of the processes that can limit terrestrial plant photosynthesis. However, we lack a robust quantitative assessment of TPU limitation of photosynthesis at the global scale. As a result, TPU, and its potential limitation of photosynthesis, is poorly represented in terrestrial biosphere models (TBMs). In this study, we utilized a global data set of photosynthetic CO2 response curves representing 141 species from tropical rainforests to Arctic tundra. We quantified TPU by fitting the standard biochemical model of C3 photosynthesis to measured photosynthetic CO2 response curves and characterized its instantaneous temperature response. Our results demonstrate that TPU does not limit leaf photosynthesis at the current ambient atmospheric CO2 concentration. Furthermore, our results showed that the light‐saturated photosynthetic rates of plants growing in cold environments are not more often limited by TPU than those of plants growing in warmer environments. In addition, our study showed that the instantaneous temperature response of TPU is distinct from temperature response of the maximum rate of Rubisco carboxylation. The new formulations of the temperature response of TPU derived in this study may prove useful in quantifying the biochemical limits to terrestrial plant photosynthesis and improve the representation of plant photosynthesis in TBMs. We demonstrate that triose phosphate utilization rate does not limit leaf photosynthesis at the current ambient atmospheric CO2 concentration. Our results do not support the hypothesis that the light‐saturated photosynthetic rates of plants growing in cold environments are more often limited by triose phosphate utilization rate than those of plants growing in warmer environments. In addition, we showed that instantaneous temperature responses of triose phosphate utilization rate are distinct from temperature responses of the maximum rate of Rubisco carboxylation. [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]

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

Recent experimental evidence suggests that during heat extremes, wooded ecosystems may decouple photosynthesis and transpiration: reducing photosynthesis to near zero but increasing transpiration into the boundary layer. This feedback may act to dampen, rather than amplify, heat extremes in wooded ecosystems. We examined eddy-covariance databases (OzFlux and FLUXNET2015) to identify whether there was field-based evidence to support these experimental findings. We focused on two types of heat extremes: (i) the three days leading up to a temperature extreme, defined as including a daily maximum temperature > 37 °C (similar to the widely used TXx metric) and (ii) heatwaves, defined as three or more consecutive days above 35 °C. When focussing on (i), we found some evidence of reduced photosynthesis and sustained or increased latent heat fluxes in seven Australian evergreen wooded flux sites. However, when considering the role of vapour pressure deficit and focusing on (ii), we were unable to conclusively disentangle the decoupling between photosynthesis and latent heat flux from the effect of increasing vapour pressure deficit. Outside of Australia, the Tier-1 FLUXNET2015 database provided limited scope to tackle this issue as it does not sample sufficient high temperature events with which to probe the physiological response of trees to extreme heat. Thus, further work is required to determine whether this photosynthetic decoupling occurs widely, ideally by matching experimental species with those found at eddy-covariance towers sites. Such measurements would allow this decoupling mechanism to be probed experimentally and at the ecosystem scale. Transpiration during heatwaves remains a key issue to resolve, as no land surface model includes a decoupling mechanism, and any potential dampening of the land-atmosphere amplification is thus not included in climate model projections. [ABSTRACT FROM AUTHOR]

The rate of CO2 diffusion from soils to the atmosphere (soil CO2 efflux, soil respiration; Rsoil) reflects the integrated activity of roots and microbes and is among the largest fluxes of the terrestrial global C cycle. Most experiments have demonstrated that Rsoil increases by 20-35% following the exposure of an ecosystem to an atmosphere enriched in CO2 (i.e., eCO2), but such experiments have largely been performed in young and N-limited ecosystems. Here, we exposed a mature and phosphorus-limited eucalypt woodland to eCO2 and measured Rsoil across three full years with a combination of manual surveys and automated measurements. We also implemented an empirical model describing the dependence of Rsoil on volumetric soil water content (θ) and soil temperature (Tsoil) to produce annual Rsoil flux estimates. Rsoil varied strongly with Tsoil, θ, and precipitation in complex and interacting ways. The realized long-term (weeks to years) temperature dependence (Q10) of Rsoil increased from ~ 1.6 at low θ up to ~ 3 at high θ. Additionally, Rsoil responded strongly and rapidly to precipitation events in a manner that depended on the conditions of θ and Tsoil at the beginning of the rain event; Rsoil increased by up to 300% within 30 min when rain fell on dry soil that had not experience rain in the preceding week, but Rsoil was rapidly reduced by up to 70% when rain fell on wet soil, leading to flooding. Repeated measures analysis of Rsoil observations over 3 years indicated no significant change in response to CO2 enrichment (P = 0.7), and elevated CO2 did not alter the dependence of Rsoil on Tsoil or θ. However, eCO2 increased Rsoil observations by ~ 10% under some constrained and moderate environmental conditions. Annual Rsoil flux sums estimated with an empirical model were ~ 7% higher in eCO2 plots than in aCO2 plots, but this difference was not statistically significant. The lack of a large eCO2 effect on Rsoil is consistent with recent evidence that aboveground net primary production was not stimulated by eCO2 in this ecosystem. The C budget of this mature woodland may be less affected by eCO2 than the young N-limited ecosystems that have been studied previously. [ABSTRACT FROM AUTHOR]