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Summary: Contemporary climate change will push many tree species into conditions that are outside their current climate envelopes. Using the Eucalyptus genus as a model, we addressed whether species with narrower geographical distributions show constrained ability to cope with warming relative to species with wider distributions, and whether this ability differs among species from tropical and temperate climates.We grew seedlings of widely and narrowly distributed Eucalyptus species from temperate and tropical Australia in a glasshouse under two temperature regimes: the summer temperature at seed origin and +3.5°C. We measured physical traits and leaf‐level gas exchange to assess warming influences on growth rates, allocation patterns, and physiological acclimation capacity.Warming generally stimulated growth, such that higher relative growth rates early in development placed seedlings on a trajectory of greater mass accumulation. The growth enhancement under warming was larger among widely than narrowly distributed species and among temperate rather than tropical provenances. The differential growth enhancement was primarily attributable to leaf area production and adjustments of specific leaf area.Our results suggest that tree species, including those with climate envelopes that will be exceeded by contemporary climate warming, possess capacity to physiologically acclimate but may have varying ability to adjust morphology. [ABSTRACT FROM AUTHOR]

Drought‐induced groundwater decline and warming associated with climate change are primary threats to dryland riparian woodlands. We used the extreme 2012–2019 drought in southern California as a natural experiment to assess how differences in water‐use strategies and groundwater dependence may influence the drought susceptibility of dryland riparian tree species with overlapping distributions. We analyzed tree‐ring stable carbon and oxygen isotopes collected from two cottonwood species (Populus trichocarpa and P. fremontii) along the semi‐arid Santa Clara River. We also modeled tree source water δ18O composition to compare with observed source water δ18O within the floodplain to infer patterns of groundwater reliance. Our results suggest that both species functioned as facultative phreatophytes that used shallow soil moisture when available but ultimately relied on groundwater to maintain physiological function during drought. We also observed apparent species differences in water‐use strategies and groundwater dependence related to their regional distributions. P. fremontii was constrained to more arid river segments and ostensibly used a greater proportion of groundwater to satisfy higher evaporative demand. P. fremontii maintained ∆13C at pre‐drought levels up until the peak of the drought, when trees experienced a precipitous decline in ∆13C. This response pattern suggests that trees prioritized maintaining photosynthetic processes over hydraulic safety, until a critical point. In contrast, P. trichocarpa showed a more gradual and sustained reduction in ∆13C, indicating that drought conditions induced stomatal closure and higher water use efficiency. This strategy may confer drought avoidance for P. trichocarpa while increasing its susceptibility to anticipated climate warming. Key Points: Dryland riparian cottonwoods used shallow soil moisture when available but ultimately relied on groundwater for survival during droughtSpecies had different water‐use strategies related to their distribution along a gradient of increasing aridity with distance from coastPopulus trichocarpa appears more susceptible to warming, and P. fremontii more vulnerable to groundwater decline due to high water demand [ABSTRACT FROM AUTHOR]

Dryland riparian woodlands are considered to be locally buffered from droughts by shallow and stable groundwater levels. However, climate change is causing more frequent and severe drought events, accompanied by warmer temperatures, collectively threatening the persistence of these groundwater dependent ecosystems through a combination of increasing evaporative demand and decreasing groundwater supply. We conducted a dendro‐isotopic analysis of radial growth and seasonal (semi‐annual) carbon isotope discrimination (Δ13C) to investigate the response of riparian cottonwood stands to the unprecedented California‐wide drought from 2012 to 2019, along the largest remaining free‐flowing river in Southern California. Our goals were to identify principal drivers and indicators of drought stress for dryland riparian woodlands, determine their thresholds of tolerance to hydroclimatic stressors, and ultimately assess their vulnerability to climate change. Riparian trees were highly responsive to drought conditions along the river, exhibiting suppressed growth and strong stomatal closure (inferred from reduced Δ13C) during peak drought years. However, patterns of radial growth and Δ13C were quite variable among sites that differed in climatic conditions and rate of groundwater decline. We show that the rate of groundwater decline, as opposed to climate factors, was the primary driver of site differences in drought stress, and trees showed greater sensitivity to temperature at sites subjected to faster groundwater decline. Across sites, higher correlation between radial growth and Δ13C for individual trees, and higher inter‐correlation of Δ13C among trees were indicative of greater drought stress. Trees showed a threshold of tolerance to groundwater decline at 0.5 m year−1 beyond which drought stress became increasingly evident and severe. For sites that exceeded this threshold, peak physiological stress occurred when total groundwater recession exceeded ~3 m. These findings indicate that drought‐induced groundwater decline associated with more extreme droughts is a primary threat to dryland riparian woodlands and increases their susceptibility to projected warmer temperatures. [ABSTRACT FROM AUTHOR]

This article is a Commentary on Guo et al. (2023), 238: 1004–1018. [ABSTRACT FROM AUTHOR]

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]

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]

Understanding how tree growth is affected by rising temperature is a key to predicting the fate of forests in future warmer climates. Increasing temperature has direct effects on plant physiology, but there are also indirect effects of increased water limitation because evaporative demand increases with temperature in many systems. In this study, we experimentally resolved the direct and indirect effects of temperature on the response of growth and photosynthesis of the widely distributed species Eucalyptus tereticornis. We grew E. tereticornis in an array of six growth temperatures from 18 to 35.5°C, spanning the climatic distribution of the species, with two watering treatments: (a) water inputs increasing with temperature to match plant demand at all temperatures (Wincr), isolating the direct effect of temperature; and (b) water inputs constant for all temperatures, matching demand for coolest grown plants (Wconst), such that water limitation increased with growth temperature. We found that constant water inputs resulted in a reduction of temperature optima for both photosynthesis and growth by ~3°C compared to increasing water inputs. Water limitation particularly reduced the total amount of leaf area displayed at Topt and intermediate growth temperatures. The reduction in photosynthesis could be attributed to lower leaf water potential and consequent stomatal closure. The reduction in growth was a result of decreased photosynthesis, reduced total leaf area display and a reduction in specific leaf area. Water availability had no effect on the response of stem and root respiration to warming, but we observed lower leaf respiration rates under constant water inputs compared to increasing water inputs at higher growth temperatures. Overall, this study demonstrates that the indirect effect of increasing water limitation strongly modifies the potential response of tree growth to rising global temperatures. [ABSTRACT FROM AUTHOR]

Trees allocate C from sources to sinks by way of a series of processes involving carbohydrate transport and utilization. Yet these dynamics are not well characterized in trees, and it is unclear how these dynamics will respond to a warmer world. Here, we conducted a warming and pulse‐chase experiment on Eucalyptus parramattensis growing in a whole‐tree chamber system to test whether warming impacts carbon allocation by increasing the speed of carbohydrate dynamics. We pulse‐labelled large (6‐m tall) trees with 13C‐CO2 to follow recently fixed C through different organs by using compound‐specific isotope analysis of sugars. We then compared concentrations and mean residence times of individual sugars between ambient and warmed (+3°C) treatments. Trees dynamically allocated 13C‐labelled sugars throughout the aboveground‐belowground continuum. We did not, however, find a significant treatment effect on C dynamics, as sugar concentrations and mean residence times were not altered by warming. From the canopy to the root system, 13C enrichment of sugars decreased, and mean residence times increased, reflecting dilution and mixing of recent photoassimilates with older reserves along the transport pathway. Our results suggest that a locally endemic eucalypt was seemingly able to adjust its physiology to warming representative of future temperature predictions for Australia. This work combined long‐term experimental warming with 13C‐CO2 pulse‐labelling and compound‐specific isotope analysis to trace recent photoassimilates from the leaves to the roots of Eucalyptus parramattensis trees in the field. We compared the concentrations and mean residence times of individual sugars in various organs between ambient and warmed (+3°C) treatments to assess the impact of warming on whole‐tree carbon dynamics. Our results show that carbon dynamics were not altered by warming representative of future temperature predictions for Australia. [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]

Summary: The allocation of carbon (C) is an important component of tree physiology that influences growth and ecosystem C storage. Allocation is challenging to measure, and its sensitivity to environmental changes such as warming and altered water availability is uncertain.We exposed young Eucalyptus tereticornis trees to +3°C warming and elimination of summer precipitation in the field using whole‐tree chambers. We calculated C allocation terms using detailed measurements of growth and continuous whole‐crown CO2 and water exchange measurements.Trees grew from small saplings to nearly 9 m height during this 15‐month experiment. Warming accelerated growth and leaf area development, and it increased the partitioning of gross primary production (GPP) to aboveground respiration and growth while decreasing partitioning below ground. Eliminating summer precipitation reduced C gain and growth but did not impact GPP partitioning. Trees utilized deep soil water and avoided strongly negative water potentials.Warming increased growth respiration, but maintenance respiration acclimated homeostatically. The increasing growth in the warmed treatment resulted in higher rates of respiration, even with complete acclimation of maintenance respiration. Warming‐induced stimulations of tree growth likely involve increased C allocation above ground, particularly to leaf area development, whereas reduced water availability may not stimulate allocation to roots. 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]

Summary: 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. [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]

Process‐based models describing biogeochemical cycling are crucial tools to understanding long‐term nutrient dynamics, especially in the context of perturbations, such as climate and land‐use change. Such models must effectively synthesize ecological processes and properties. For example, in terrestrial ecosystems, plants are the primary source of bioavailable carbon, but turnover rates of essential nutrients are contingent on interactions between plants and soil biota. Yet, biogeochemical models have traditionally considered plant and soil communities in broad terms. The next generation of models must consider how shifts in their diversity and composition affect ecosystem processes.One promising approach to synthesize plant and soil biodiversity and their interactions into models is to consider their diversity from a functional trait perspective. Plant traits, which include heritable chemical, physical, morphological and phenological characteristics, are increasingly being used to predict ecosystem processes at a range of scales, and to interpret biodiversity–ecosystem functional relationships. There is also emerging evidence that the traits of soil microbial and faunal communities can be correlated with ecosystem functions such as decomposition, nutrient cycling, and greenhouse gas production.Here, we draw on recent advances in measuring and using traits of different biota to predict ecosystem processes, and provide a new perspective as to how biotic traits can be integrated into biogeochemical models. We first describe an explicit trait‐based model framework that operates at small scales and uses direct measurements of ecosystem properties; second, an integrated approach that operates at medium scales and includes interactions between biogeochemical cycling and soil food webs; and third, an implicit trait‐based model framework that associates soil microbial and faunal functional groups with plant functional groups, and operates at the Earth‐system level. In each of these models, we identify opportunities for inclusion of traits from all three groups to reduce model uncertainty and improve understanding of biogeochemical cycles.These model frameworks will generate improved predictive capacity of how changes in biodiversity regulate biogeochemical cycles in terrestrial ecosystems. Further, they will assist in developing a new generation of process‐based models that include plant, microbial, and faunal traits and facilitate dialogue between empirical researchers and modellers. [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]

Summary: Plant characteristics in different provenances within a single species may vary in response to climate change, which might alter soil microbial communities and ecosystem functions. We conducted a glasshouse experiment and grew seedlings of three provenances (temperate, subtropical and tropical origins) of a tree species (i.e., Eucalyptus tereticornis) at different growth temperatures (18, 21.5, 25, 28.5, 32 and 35.5°C) for 54 days. At the end of the experiment, bacterial and fungal community composition, diversity and abundance were characterized. Measured soil functions included surrogates of microbial respiration, enzyme activities and nutrient cycling. Using Permutation multivariate analysis of variance (PerMANOVA) and network analysis, we found that the identity of tree provenances regulated both structure and function of soil microbiomes. In some cases, tree provenances substantially affected the response of microbial communities to the temperature treatments. For example, we found significant interactions of temperature and tree provenance on bacterial community and relative abundances of Chloroflexi and Zygomycota, and inorganic nitrogen. Microbial abundance was altered in response to increasing temperature, but was not affected by tree provenances. Our study provides novel evidence that even a small variation in biotic components (i.e., intraspecies tree variation) can significantly influence the response of soil microbial community composition and specific soil functions to global warming. [ABSTRACT FROM AUTHOR]

Impacts of climate warming depend on the degree to which plants are constrained by adaptation to their climate-of-origin or exhibit broad climatic suitability. We grew cool-origin, central and warm-origin provenances of Eucalyptus tereticornis in an array of common temperature environments from 18 to 35.5°C to determine if this widely distributed tree species consists of geographically contrasting provenances with differentiated and narrow thermal niches, or if provenances share a common thermal niche. The temperature responses of photosynthesis, respiration, and growth were equivalent across the three provenances, reflecting a common thermal niche despite a 2,200 km geographic distance and 13°C difference in mean annual temperature at seed origin. The temperature dependence of growth was primarily mediated by changes in leaf area per unit plant mass, photosynthesis, and whole-plant respiration. Thermal acclimation of leaf, stem, and root respiration moderated the increase in respiration with temperature, but acclimation was constrained at high temperatures. We conclude that this species consists of provenances that are not differentiated in their thermal responses, thus rejecting our hypothesis of adaptation to climate-of-origin and suggesting a shared thermal niche. In addition, growth declines with warming above the temperature optima were driven by reductions in whole-plant leaf area and increased respiratory carbon losses. The impacts of climate warming will nonetheless vary across the geographic range of this and other such species, depending primarily on each provenance's climate position on the temperature response curves for photosynthesis, respiration, and growth. [ABSTRACT FROM AUTHOR]

Understanding physiological acclimation of photosynthesis and respiration is important in elucidating the metabolic performance of trees in a changing climate. Does physiological acclimation to climate warming mirror acclimation to seasonal temperature changes?, We grew Eucalyptus tereticornis trees in the field for 14 months inside 9-m tall whole-tree chambers tracking ambient air temperature ( Tair) or ambient Tair + 3°C (i.e. 'warmed'). We measured light- and CO2-saturated net photosynthesis ( Amax) and night-time dark respiration ( R) each month at 25°C to quantify acclimation. Tree growth was measured, and leaf nitrogen (N) and total nonstructural carbohydrate (TNC) concentrations were determined to investigate mechanisms of acclimation., Warming reduced Amax and R measured at 25°C compared to ambient-grown trees. Both traits also declined as mean daily Tair increased, and did so in a similar way across temperature treatments. Amax and R (at 25°C) both increased as TNC concentrations increased seasonally; these relationships appeared to arise from source-sink imbalances, suggesting potential substrate regulation of thermal acclimation., We found that photosynthesis and respiration each acclimated equivalently to experimental warming and seasonal temperature change of a similar magnitude, reflecting a common, nearly homeostatic constraint on leaf carbon exchange that will be important in governing tree responses to climate warming. [ABSTRACT FROM AUTHOR]

Given the contrasting short-term temperature dependences of gross primary production (GPP) and autotrophic respiration, the fraction of GPP respired by trees is predicted to increase with warming, providing a positive feedback to climate change. However, physiological acclimation may dampen or eliminate this response., We measured the fluxes of aboveground respiration ( Ra), GPP and their ratio ( Ra/GPP) in large, field-grown Eucalyptus tereticornis trees exposed to ambient or warmed air temperatures (+3°C). We report continuous measurements of whole-canopy CO2 exchange, direct temperature response curves of leaf and canopy respiration, leaf and branch wood respiration, and diurnal photosynthetic measurements., Warming reduced photosynthesis, whereas physiological acclimation prevented a coincident increase in Ra. Ambient and warmed trees had a common nonlinear relationship between the fraction of GPP that was respired above ground ( Ra/GPP) and the mean daily temperature. Thus, warming significantly increased Ra/GPP by moving plants to higher positions on the shared Ra/GPP vs daily temperature relationship, but this effect was modest and only notable during hot conditions., Despite the physiological acclimation of autotrophic respiration to warming, increases in temperature and the frequency of heat waves may modestly increase tree Ra/GPP, contributing to a positive feedback between climate warming and atmospheric CO2 accumulation. [ABSTRACT FROM AUTHOR]

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]

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]

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]

Forest ecosystems release large amounts of carbon to the atmosphere from fine-root respiration (Rr), but the control of this flux and its temperature sensitivity (Q10) are poorly understood. We attempted to: (1) identify the factors limiting this flux using additions of glucose and an electron transport uncoupler (carbonyl cyanide m-chlorophenylhydrazone); and (2) improve yearly estimates of Rr by directly measuring its Q10 in situ using temperature-controlled cuvettes buried around intact, attached roots. The proximal limits of Rr of loblolly pine ( Pinus taeda L.) trees exposed to free-air CO2 enrichment (FACE) and N fertilization were seasonally variable; enzyme capacity limited Rr in the winter, and a combination of substrate supply and adenylate availability limited Rr in summer months. The limiting factors of Rr were not affected by elevated CO2 or N fertilization. Elevated CO2 increased annual stand-level Rr by 34% whereas the combination of elevated CO2 and N fertilization reduced Rr by 40%. Measurements of in situ Rr with high temporal resolution detected diel patterns that were correlated with canopy photosynthesis with a lag of 1 d or less as measured by eddy covariance, indicating a dynamic link between canopy photosynthesis and root respiration. These results suggest that Rr is coupled to daily canopy photosynthesis and increases with carbon allocation below ground. [ABSTRACT FROM AUTHOR]

Carbon-use efficiency (CUE), the ratio of net primary production (NPP) to gross primary production (GPP), describes the capacity of forests to transfer carbon (C) from the atmosphere to terrestrial biomass. It is widely assumed in many landscape-scale carbon-cycling models that CUE for forests is a constant value of ∼0.5. To achieve a constant CUE, tree respiration must be a constant fraction of canopy photosynthesis. We conducted a literature survey to test the hypothesis that CUE is constant and universal among forest ecosystems. Of the 60 data points obtained from 26 papers published since 1975, more than half reported values of GPP that were not estimated independently from NPP; values of CUE calculated from independent estimates of GPP were greater than those calculated from estimates of GPP derived from NPP. The slope of the relationship between NPP and GPP for all forests was 0.53, but values of CUE varied from 0.23 to 0.83 for different forest types. CUE decreased with increasing age, and a substantial portion of the variation among forest types was caused by differences in stand age. When corrected for age the mean value of CUE was greatest for temperate deciduous forests and lowest for boreal forests. CUE also increased as the ratio of leaf mass-to-total mass increased. Contrary to the assumption of constancy, substantial variation in CUE has been reported in the literature. It may be inappropriate to assume that respiration is a constant fraction of GPP as adhering to this assumption may contribute to incorrect estimates of C cycles. A 20% error in current estimates of CUE used in landscape models (i.e. ranging from 0.4 to 0.6) could misrepresent an amount of C equal to total anthropogenic emissions of CO2 when scaled to the terrestrial biosphere. [ABSTRACT FROM AUTHOR]