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Populations from different climates often show unique growth responses to temperature, reflecting temperature adaptation. Yet, whether populations from different climates differ in physiological temperature acclimation remains unclear. Here, we test whether populations from differing thermal environments exhibit different growth responses to temperature and differences in temperature acclimation of leaf respiration. We grew tropical and subtropical populations of two mangrove species (Avicennia germinans and Rhizophora mangle) under ambient and experimentally warmed conditions in a common garden at the species’ northern range limit. We quantified growth and temperature responses of leaf respiration (R) at seven time points over ~10 months. Warming increased productivity of tropical populations more than subtropical populations, reflecting a higher temperature optimum for growth. In both species, R measured at 25 °C declined as seasonal temperatures increased, demonstrating thermal acclimation. Contrary to our expectations, acclimation of R was consistent across populations and temperature treatments. However, populations differed in adjusting the temperature sensitivity of R (Q10) to seasonal temperatures. Following a freeze event, tropical Avicennia showed greater freeze damage than subtropical Avicennia, while both Rhizophora populations appeared equally susceptible. We found evidence of temperature adaptation at the whole-plant scale but little evidence for population differences in thermal acclimation of leaf physiology. Studies that examine potential costs and benefits of thermal acclimation in an evolutionary context may provide new insights into limits of thermal acclimation.

Sustaining grassland production in a changing climate requires an understanding of plant adaptation strategies, including trait plasticity under warmer and drier conditions. However, our knowledge to date disproportionately relies on aboveground responses, despite the importance of belowground traits in maintaining aboveground growth, especially in grazed systems. We subjected a perennial pasture grass, Festuca arundinacea, to year-round warming (+3 °C) and cool-season drought (60% rainfall reduction) in a factorial field experiment to test the hypotheses that: (i) drought and warming increase carbon allocation belowground and shift root traits towards greater resource acquisition and (ii) increased belowground carbon reserves support post-drought aboveground recovery. Drought and warming reduced plant production and biomass allocation belowground. Drought increased specific root length and reduced root diameter in warmed plots but increased root starch concentrations under ambient temperature. Higher diameter and soluble sugar concentrations of roots and starch storage in crowns explained aboveground production under climate extremes. However, the lack of association between post-drought aboveground biomass and belowground carbon and nitrogen reserves contrasted with our predictions. These findings demonstrate that root trait plasticity and belowground carbon reserves play a key role in aboveground production during climate stress, helping predict pasture responses and inform management decisions under future climates.

Rising atmospheric [CO2 ] (Ca ) generally enhances tree growth if nutrients are not limiting. However, reduced water availability and elevated evaporative demand may offset such fertilization. Trees with access to deep soil water may be able to mitigate such stresses and respond more positively to Ca . Here, we sought to evaluate how increased vapor pressure deficit and reduced precipitation are likely to modify the impact of elevated Ca (eCa ) on tree productivity in an Australian Eucalyptus saligna Sm. plantation with access to deep soil water. We parameterized a forest growth simulation model (GOTILWA+) using data from two field experiments on E. saligna: a 2-year whole-tree chamber experiment with factorial Ca (ambient =380, elevated =620 μmol mol-1 ) and watering treatments, and a 10-year stand-scale irrigation experiment. Model evaluation showed that GOTILWA+ can capture the responses of canopy C uptake to (1) rising vapor pressure deficit (D) under both Ca treatments; (2) alterations in tree water uptake from shallow and deep soil layers during soil dry-down; and (3) the impact of irrigation on tree growth. Simulations suggest that increasing Ca up to 700 μmol mol-1 alone would result in a 33% increase in annual gross primary production (GPP) and a 62% increase in biomass over 10 years. However, a combined 48% increase in D and a 20% reduction in precipitation would halve these values. Our simulations identify high D conditions as a key limiting factor for GPP. They also suggest that rising Ca will compensate for increasing aridity limitations in E. saligna trees with access to deep soil water under non-nutrient limiting conditions, thereby reducing the negative impacts of global warming upon this eucalypt species. Simulation models not accounting for water sources available to deep-rooting trees are likely to overestimate aridity impacts on forest productivity and C stocks.

Cities have been described as 'heat islands' and 'dry islands' due to hotter, drier air in urban areas, relative to the surrounding landscape. As climate change intensifies, the health of urban trees will be increasingly impacted. Here, we posed the question: Is it possible to predict urban tree species mortality using (1) species climate envelopes and (2) plant functional traits? To answer these, we tracked patterns of crown dieback and recovery for 23 common urban tree and shrub species in Sydney, Australia during the record-breaking austral 2019-2020 summer. We identified 10 heat-tolerant species including five native and five exotic species, which represent climate-resilient options for urban plantings that are likely to continue to thrive for decades. Thirteen species were considered vulnerable to adverse conditions due to their mortality, poor health leading to tree removal, and/or extensive crown dieback. Crown dieback increased with increasing precipitation of the driest month of species climate of origin, suggesting that species from dry climates may be better suited for urban forests in future climates. We effectively grouped species according to their drought strategy (i.e., tolerance versus avoidance) using a simple trait-based framework that was directly linked with species mortality. The seven most climate-vulnerable species used a drought-avoidance strategy, having low wood density and high turgor loss points along with large, thin leaves with low heat tolerance. Overall, plant functional traits were better than species climate envelopes at explaining crown dieback. Recovery after stress required two mild, wet years for most species, resulting in prolonged loss of cooling benefits as well as economic losses due to replacement of dead/damaged trees. Hotter, longer, and more frequent heatwaves will require selection of more climate-resilient species in urban forests, and our results suggest that future research should focus on plant thermal traits to improve prediction models and species selection.

Ecosystems integrity and services are threatened by anthropogenic global changes. Mitigating and adapting to these changes require knowledge of ecosystem functioning in the expected novel environments, informed in large part through experimentation and modelling. This paper describes 13 advanced controlled environment facilities for experimental ecosystem studies, herein termed ecotrons, open to the international community. Ecotrons enable simulation of a wide range of natural environmental conditions in replicated and independent experimental units while measuring various ecosystem processes. This capacity to realistically control ecosystem environments is used to emulate a variety of climatic scenarios and soil conditions, in natural sunlight or through broad‐spectrum lighting. The use of large ecosystem samples, intact or reconstructed, minimizes border effects and increases biological and physical complexity. Measurements of concentrations of greenhouse trace gases as well as their net exchange between the ecosystem and the atmosphere are performed in most ecotrons, often quasi continuously. The flow of matter is often tracked with the use of stable isotope tracers of carbon and other elements. Equipment is available for measurements of soil water status as well as root and canopy growth. The experiments ran so far emphasize the diversity of the hosted research. Half of them concern global changes, often with a manipulation of more than one driver. About a quarter deal with the impact of biodiversity loss on ecosystem functioning and one quarter with ecosystem or plant physiology. We discuss how the methodology for environmental simulation and process measurements, especially in soil, can be improved and stress the need to establish stronger links with modelling in future projects. These developments will enable further improvements in mechanistic understanding and predictive capacity of ecotron research which will play, in complementarity with field experimentation and monitoring, a crucial role in exploring the ecosystem consequences of environmental changes., Experimentation and modelling are necessary to predict ecosystem functioning under future environments and to develop mitigating and adapting strategies. This paper describes 13 advanced controlled environment facilities, called ecotrons, open to the international community. An ecotron comprises a set of replicated enclosures designed to host ecosystems samples and enable realistic simulations of above‐ and belowground environmental conditions, while simultaneously and automatically measuring ecosystem processes. The characteristics of these infrastructures are given as well as examples of collaborative projects hosted so far.

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 δ18 O. 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 δ18 O 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 δ18 O to infer TL-AW , but does not support the concept of strong homeothermic regulation of Tleaf.

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

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.

Carbon allocation determines plant growth, fitness and reproductive success. However, climate warming and drought impacts on carbon allocation patterns in grasses are not well known, particularly following grazing or clipping. A widespread C

Tree mortality during global-change-type drought is usually attributed to xylem dysfunction, but as climate change increases the frequency of extreme heat events, it is necessary to better understand the interactive role of heat stress. We hypothesized that some drought-stressed plants paradoxically open stomata in heatwaves to prevent leaves from critically overheating. We experimentally imposed heat (>40°C) and drought stress onto 20 broadleaf evergreen tree/shrub species in a glasshouse study. Most well-watered plants avoided lethal overheating, but drought exacerbated thermal damage during heatwaves. Thermal safety margins (TSM) quantifying the difference between leaf surface temperature and leaf critical temperature, where photosynthesis is disrupted, identified species vulnerability to heatwaves. Several mechanisms contributed to high heat tolerance and avoidance of damaging leaf temperatures-small leaf size, low leaf osmotic potential, high leaf mass per area (i.e., thick, dense leaves), high transpirational capacity, and access to water. Water-stressed plants had smaller TSM, greater crown dieback, and a fundamentally different stomatal heatwave response relative to well-watered plants. On average, well-watered plants closed stomata and decreased stomatal conductance (gs ) during the heatwave, but droughted plants did not. Plant species with low gs , either due to isohydric stomatal behavior under water deficit or inherently low transpirational capacity, opened stomata and increased gs under high temperatures. The current paradigm maintains that stomata close before hydraulic thresholds are surpassed, but our results suggest that isohydric species may dramatically increase gs (over sixfold increases) even past their leaf turgor loss point. By actively increasing water loss at high temperatures, plants can be driven toward mortality thresholds more rapidly than has been previously recognized. The inclusion of TSM and responses to heat stress could improve our ability to predict the vulnerability of different tree species to future droughts.

While trees can acclimate to warming, there is concern that tropical rainforest species may be less able to acclimate because they have adapted to a relatively stable thermal environment. Here we tested whether the physiological adjustments to warming differed among Australian tropical, subtropical and warm-temperate rainforest trees. Photosynthesis and respiration temperature responses were quantified in six Australian rainforest seedlings of tropical, subtropical and warm-temperate climates grown across four growth temperatures in a glasshouse. Temperature-response models were fitted to identify mechanisms underpinning the response to warming. Tropical and subtropical species had higher temperature optima for photosynthesis (T

Shifts in the timing, intensity and/or frequency of climate extremes, such as severe drought and heatwaves, can generate sustained shifts in ecosystem function with important ecological and economic impacts for rangelands and managed pastures. The Pastures and Climate Extremes experiment (PACE) in Southeast Australia was designed to investigate the impacts of a severe winter/spring drought (60% rainfall reduction) and, for a subset of species, a factorial combination of drought and elevated temperature (ambient +3°C) on pasture productivity. The experiment included nine common pasture and Australian rangeland species from three plant functional groups (C3 grasses, C4 grasses and legumes) planted in monoculture. Winter/spring drought resulted in productivity declines of 45% on average and up to 74% for the most affected species (Digitaria eriantha) during the 6-month treatment period, with eight of the nine species exhibiting significant yield reductions. Despite considerable variation in species’ sensitivity to drought, C4 grasses were more strongly affected by this treatment than C3 grasses or legumes. Warming also had negative effects on cool-season productivity, associated at least partially with exceedance of optimum growth temperatures in spring and indirect effects on soil water content. The combination of winter/spring drought and year-round warming resulted in the greatest yield reductions. We identified responses that were either additive (Festuca), or less-than-additive (Medicago), where warming reduced the magnitude of drought effects. Results from this study highlight the sensitivity of diverse pasture species to increases in winter and spring drought severity similar to those predicted for this region, and that anticipated benefits of cool-season warming are unlikely to be realized. Overall, the substantial negative impacts on productivity suggest that future, warmer, drier climates will result in shortfalls in cool-season forage availability, with profound implications for the livestock industry and natural grazer communities.

Heatwaves are becoming more frequent with climate warming and can impact tree growth and reproduction. Eucalyptus parramattensis can cope with an extreme heatwave in the field via transpiratory cooling and enhanced leaf thermal tolerance that protected foliar tissues from photo-inhibition and photo-oxidation during natural midday irradiance. Here, we explored whether changes in foliar carotenoids and/or the xanthophyll cycle state can facilitate leaf acclimation to long-term warming and/or an extreme heatwave event. We found that leaves had similar carotenoid levels when grown for one year under ambient and experimental long-term warming (+ 3 °C) conditions in whole tree chambers. Exposure to a 4-day heatwave (> 43 °C) significantly altered the xanthophyll de-epoxidation state of carotenoids revealing one mechanism by which trees could minimise foliar photo-oxidative damage. The levels of zeaxanthin were significantly higher in both young and old leaves during the heatwave, revealing that violaxanthin de-epoxidation and perhaps de novo zeaxanthin synthesis contributed to enhancement of the xanthophyll cycle state. In a future climate of long-term warming and increased heatwave events, leaves of E. parramattensis will be able to utilise biochemical strategies to alter the xanthophyll cycle state and cope with extreme temperatures under natural solar irradiation.

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.

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 13 C-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 13 C 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.

It is easy to measure annual growth of a tree stem. It is hard to measure its daily growth. The reason for this difficulty is the microscopic scale and the need to separate processes that simultaneously result in reversible and irreversible stem expansion. Here we present a model that separates reversible from irreversible cell expansion. Our model is novel, because it explains reversible expansion as consequence of longitudinally and, importantly, radially transmitted changes of hydraulic and osmotic pressures in xylem and bark. To capture and quantify these changes, we manipulated daily stem growth by applying a phloem girdle to stems of 9-m tall trees. The model was informed by measurements of radial movement in stem tissues and sap flow before and after and positions below and above the girdle. Additional measurements of whole-crown fluxes of H2O and CO2, leaf water potentials, non-structural carbohydrates and respiration were used to document the physiological impacts of girdling. This work sheds new light on the role of radial transport processes underpinning daily growth of tree stems. The model helps explain diel patterns of stem growth in trees.

Urban forests (i.e. all vegetation present in urban areas), provide environmental and socio-economic benefits to more than half of the global population. Projected climate change threatens these benefits to society. Here, we assess vulnerability to climate change of 16,006 plant species present in the urban forests of 1,010 cities within 93 countries, using three vulnerability metrics: exposure, safety margin and risk. Exposure expresses the magnitude of projected changes in climate in a given area, safety margin measures species' sensitivity to climate change, and risk is the difference between exposure and safety margin. We identified 9,676 (60.5%) and 8,344 (52.1%) species exceeding their current climatic tolerance (i.e. safety margin) for mean annual temperature (MAT) and annual precipitation (AP), respectively. By 2050, 13,479 (84.2%) and 9,960 (62.2%) species are predicted to be at risk from projected changes in MAT and AP, respectively, with risk increasing in cities at lower latitudes. Our results can aid evaluation of the impacts of climate change on urban forests and identify the species most at risk. Considering future climates when selecting species for urban plantings will enhance the long-term societal benefits provided by urban forests, including their contribution to mitigating the magnitude and impacts of climate change.

summary The effects of soil nitrogen availability and chronic ozone stress on carbon and nutrient economy were investigated in loblolly pine (Pinus. taeda L.) and yellow-poplar (Liriodendron tulipifera L.). One-year-old seedlings were planted individually in pots in forest soil of low (58 μg g−1), medium (96 μg g−1) or high (172 μg g−1) initial concentrations of soluble nitrogen. The seedlings were exposed to ozone in open-top field chambers at sub-ambient (charcoal-filtered air), ambient, and elevated (ambient + 60 nl 1−1 O3) (32, 56, 108 nl 1−1 O3, 1 h seasonal mean, respectively) levels for 18 weeks. At final harvest loblolly pine dry matter increased by 50% at the highest soil K level relative to the low with the largest gains in new needle biomass. Elevated ozone reduced the biomass of current-year needles by 20% in plants grown at the highest N level. Higher soil N supply increased the concentration of nitrogen in needles, stimulated current-year needle photosynthesis and increased needle and whole-plant water-use efficiencies. Ozone treatment had no significant effect on photosynthesis or water-use efficiency in either species, although ozone exposure tended to reduce- stomatal conductance in loblolly pine. The low N treatment increased the proportion of dry matter allocated to fine roots in yellow-poplar, but whole-plant dry weight had not responded to N fertilization at the final harvest, suggesting other limitations on growth. Ozone exposure increased leaf abscission and doubled leaf turnover m yellow-poplar. Although yellow-poplar was highly sensitive to ozone-induced leaf abscission, final whole-plant dry weights were not affected. The indeterminate growth habit of yellow-poplar permitted compensatory leaf growth which may have ameliorated effects of chronic ozone stress on biomass gain. Ozone exposure also decreased shoot weight more than root weight, resulting in higher root:leaf ratios in loblolly pine and a similar trend m higher fine roor:leaf ratios in yellow-poplar. Greater proportional allocation of carbon to roots in response to nutrient deficiency may preclude an increased allocation to shoots often observed in response to air pollution stress. Interspecific differences in growth response to chronic ozone and nutrient stress may be influenced by differences in leaf growth habit.

SUMMARY Hybrid poplar (Populus tristis Fisch. ×P. balsamifera L., cv. Tristis) and sugar maple (Acer saccharum Marsh.) seedlings were grown under contrasting light and ozone treatments to investigate the role of the light environment in their response to chronic ozone stress. In consecutive growth chamber experiments, cuttings of shade-intolerant poplar and 3-yr-old seedlings of shade-tolerant sugar maple were grown in pots for 6 and 10 wk, respectively, under shaded, low light irradiance (c. 2.5 mol m−2 d−1 PPFD or 7% of full sunlight) and six-fold greater irradiance (c. 16.6 mol m−2 d−1 PPFD or 45% of full sunlight) in combination with low (< 10 nl 1−1) and elevated levels of ozone (c. 99–115 nl 1−1). In unshaded poplar plants, ozone exposure reduced root dry mass by 33% at final harvest, while shaded plants had no such response. By comparison, sugar maple root dry mass was reduced by ozone in shaded plants by 10%, but was unaffected by ozone in unshaded plants. In poplar, leaf area: plant dry mass ratios were unaffected by ozone, whereas in sugar maple ozone-exposed plants had a 24% lower leaf area: plant dry mass ratio in the shaded treatment. In shade-grown sugar maple, ozone doubled dark respiration rates of leaves, but in unshaded seedlings ozone had no effect on respiration. In comparison, in poplar plants ozone exposure resulted in greater increases in dark respiration under unshaded than shaded conditions. In unshaded plants, ozone treatment resulted in lower in situ net photosynthesis in poplar, but not in sugar maple. Overall, shade-grown sugar maple appeared more sensitive to ozone stress than unshaded plants in terms of lower leaf area: plant dry mass ratio and root growth and higher leaf respiration. In poplar on the other hand, root growth, leaf respiration and photosynthesis were more affected by ozone in unshaded than in shaded plants. These findings suggest that shade-grown sugar maple and unshaded poplar may experience greater reductions in carbon gain and growth under elevated levels of ozone than plants under the opposite light conditions.

Understanding seasonal and diurnal dynamics of ecosystem respiration (Reco) in forests is challenging, because Reco can only be measured directly during night-time by eddy-covariance flux towers. Reco is the sum of soil respiration (Rsoil) and above-ground respiration (in theory, RAG = Reco − Rsoil). Rsoil can be measured day and night and can provide a check of consistency on Reco, as the difference in magnitude and time dynamic between Reco and Rsoil should be explained by RAG. We assessed the temporal patterns and climatic drivers of Rsoil and Reco in a mature eucalypt woodland, using continuous measurements (only at night for Reco) at half-hourly resolution over 4 years (2014–2017). Our data showed large seasonal and diurnal (overnight) variation of Reco, while Rsoil had a low diurnal amplitude and their difference (Reco − Rsoil, or RAG) had a low seasonal amplitude. This result implies at first glance that seasonal variation of Reco was mainly influenced by Rsoil while its diurnal variation was mainly influenced by RAG. However, our analysis suggests that the night-time Reco decline cannot realistically be explained by a decline of RAG. Chamber measurements of autotrophic components at half-hourly time resolution are needed to quantify how much of the Reco decline overnight is due to declines in leaf or stem respiration, and how much is due to missing storage or advection, which may create a systematic bias in Reco measurements. Our findings emphasize the need for reconciling bottom-up (via components measured with chambers) and direct estimates of Reco (via eddy-covariance method).

Shifts in the timing, intensity and/or frequency of climate extremes, such as severe drought and heatwaves, can generate sustained shifts in ecosystem function with important ecological and economic impacts for rangelands and managed pastures. The Pastures and Climate Extremes experiment (PACE) in Southeast Australia was designed to investigate the impacts of a severe winter/spring drought (60% rainfall reduction) and, for a subset of species, a factorial combination of drought and elevated temperature (ambient +3 °C) on pasture productivity. The experiment included nine common pasture and Australian rangeland species from three plant functional groups (C3 grasses, C4 grasses and legumes) planted in monoculture. Winter/spring drought resulted in productivity declines of 45% on average and up to 74% for the most affected species (Digitaria eriantha) during the 6-month treatment period, with eight of the nine species exhibiting significant yield reductions. Despite considerable variation in species’ sensitivity to drought, C4 grasses were more strongly affected by this treatment than C3 grasses or legumes. Warming also had negative effects on cool-season productivity, associated at least partially with exceedance of optimum growth temperatures in spring and indirect effects on soil water content. The combination of winter/spring drought and year-round warming resulted in the greatest yield reductions. We identified responses that were either additive such that there was only as significant warming effect under drought (Festuca), or less-than-additive, where there was no drought effect under warming (Medicago), compared to ambient plots. Results from this study highlight the sensitivity of diverse pasture species to increases in winter and spring drought severity similar to those predicted for this region, and that anticipated benefits of cool-season warming are unlikely to be realised. Overall, the substantial negative impacts on productivity suggest that future, warmer, drier climates will result in shortfalls in cool-season forage availability, with profound implications for the livestock industry and natural grazer communities.

Rising atmospheric [CO

Globally, soils store two to three times as much carbon as currently resides in the atmosphere, and it is critical to understand how soil greenhouse gas (GHG) emissions and uptake will respond to ongoing climate change. In particular, the soil‐to‐atmosphere CO2 flux, commonly though imprecisely termed soil respiration (R S), is one of the largest carbon fluxes in the Earth system. An increasing number of high‐frequency R S measurements (typically, from an automated system with hourly sampling) have been made over the last two decades; an increasing number of methane measurements are being made with such systems as well. Such high frequency data are an invaluable resource for understanding GHG fluxes, but lack a central database or repository. Here we describe the lightweight, open‐source COSORE (COntinuous SOil REspiration) database and software, that focuses on automated, continuous and long‐term GHG flux datasets, and is intended to serve as a community resource for earth sciences, climate change syntheses and model evaluation. Contributed datasets are mapped to a single, consistent standard, with metadata on contributors, geographic location, measurement conditions and ancillary data. The design emphasizes the importance of reproducibility, scientific transparency and open access to data. While being oriented towards continuously measured R S, the database design accommodates other soil‐atmosphere measurements (e.g. ecosystem respiration, chamber‐measured net ecosystem exchange, methane fluxes) as well as experimental treatments (heterotrophic only, etc.). We give brief examples of the types of analyses possible using this new community resource and describe its accompanying R software package., Here we describe the lightweight, open source COSORE (COntinuous SOil REspiration) database and software. COSORE focuses on automated, continuous and long‐term greenhouse gas flux datasets, and is intended to serve as a community resource for earth sciences, climate change syntheses and model evaluation.

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 (δ13 C) 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 δ13 C of the phloem (δ13 Cph ). We measured gas exchange, online 13 C-discrimination, and δ13 Cph 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 δ13 Cph agreed with iWUE from gas exchange, but only after incorporating gm . δ13 Cph was also correlated with whole-tree 13 C-discrimination, but offset by -2.5 ± 0.7‰, presumably due to post-photosynthetic fractionations. We conclude that δ13 Cph 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.

International audience; Short-term temperature response curves of leaf dark respiration (R-T) provide insights into a critical process that influences plant net carbon exchange. This includes how respiratory traits acclimate to sustained changes in the environment. Our study analysed 860 high-resolutionR-T(10-70 degrees C range) curves for: (a) 62 evergreen species measured in two contrasting seasons across several field sites/biomes; and (b) 21 species (subset of those sampled in the field) grown in glasshouses at 20 degrees C : 15 degrees C, 25 degrees C : 20 degrees C and 30 degrees C : 25 degrees C, day : night. In the field, across all sites/seasons, variations inR(25)(measured at 25 degrees C) and the leafTwhereRreached its maximum (T-max) were explained by growthT(mean air-Tof 30-d before measurement), solar irradiance and vapour pressure deficit, with growthThaving the strongest influence.R(25)decreased andT(max)increased with rising growthTacross all sites and seasons with the single exception of winter at the cool-temperate rainforest site where irradiance was low. The glasshouse study confirmed thatR(25)andT(max)thermally acclimated. Collectively, the results suggest: (1) thermal acclimation of leafRis common in most biomes; and (2) the highTthreshold of respiration dynamically adjusts upward when plants are challenged with warmer and hotter climates.

Predicting future air quality in Australian cities dominated by eucalypt emissions requires an understanding of their emission potentials in a warmer climate. Here we measure the temperature response in isoprene emissions from saplings of four different Eucalyptus species grown under current and future average summertime temperature conditions. The future conditions represent a 2050 climate under Representative Concentration Pathway 8.5, with average daytime temperatures of 294.5 K. Ramping the temperature from 293 to 328 K resulted in these eucalypts emitting isoprene at temperatures 4–9 K higher than the default maximum emission temperature in the Model of Emissions of Gases and Aerosols from Nature (MEGAN). New basal emission rate measurements were obtained at the standard conditions of 303 K leaf temperature and 1000 µmol m−2 s−1 photosynthetically active radiation and converted into landscape emission factors. We applied the eucalypt temperature responses and emission factors to Australian trees within MEGAN and ran the CSIRO Chemical Transport Model for three summertime campaigns in Australia. Compared to the default model, the new temperature responses resulted in less isoprene emission in the morning and more during hot afternoons, improving the statistical fit of modelled to observed ambient isoprene. Compared to current conditions, an additional 2 ppb of isoprene is predicted in 2050, causing hourly increases up to 21 ppb of ozone and 24-hourly increases of 0.4 µg m−3 of aerosol in Sydney. A 550 ppm CO2 atmosphere in 2050 mitigates these peak Sydney ozone mixing ratios by 4 ppb. Nevertheless, these forecasted increases in ozone are up to one-fifth of the hourly Australian air quality limit, suggesting that anthropogenic NOx should be further reduced to maintain healthy air quality in future.

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.

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.

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.

Extreme climate conditions, including more frequent and prolonged heatwaves, are increasing in many regions throughout the world. Urban trees can aid in mitigating the adverse impacts of climate extremes; however, their capacity to do so is limited by species differences in ability to maintain function and retain leaves to provide shade under extreme temperatures. To assess tree vulnerability to heatwaves, we used a common garden experiment in Western Sydney, Australia, which included four widely-planted tree species; two native evergreen and two exotic deciduous species. Data were collected during 2019 and 2020, the two hottest summers on local record with maximum air temperatures reaching 45 °C. We monitored plant physiological status as stomatal conductance (gS) and midday leaf water potential (Ψmid). We determined heat tolerance by measuring the leaf critical temperature (Tcrit) for photosynthesis and leaf turgor loss point (πtlp) as thresholds for loss of function and calculated the thermal safety margin. Plant performance was assessed by measuring tree growth and leaf damage after heatwaves. Species responded dynamically with gS decreasing as air temperature and vapor pressure deficit increased during heatwaves. Exotic species had higher gS, Tcrit and πtlp than native species. Leaf damage under heatwave conditions was overall lower in native species. Our results highlight the impact of heatwaves on urban trees and the value of physiological metrics to inform tree species selection to create more resilient urban environments.

The CO2 efflux from tree stem surfaces to atmosphere (RS) is an important component in the carbon (C) balance of forest ecosystems. Rising atmospheric carbon dioxide concentrations [CO2] are expected to stimulate RS, because of greater C assimilation and carbohydrate supply to stems under rising [CO2]. Growth respiration (Rg) and maintenance respiration (Rm) during the warm growing season may respond differently to rising [CO2] due to different metabolic demands. To test the effect of elevated [CO2] (eCO2, ambient +150 ppm) on RS, we examined RS in mature Eucalyptus trees on a monthly basis for an entire year during the seventh year of exposure to eCO2. RS varied seasonally and mirrored seasonal variation in temperature. While RS was not significantly increased under eCO2 compared to ambient CO2 (aCO2), its temperature sensitivity was significantly decreased (Q10 of 1.92 for aCO2 and 1.56 for eCO2). The estimated annual Rg accounted for approximately 7–8% of annual total RS, 419 ± 103 g C m−2 yr-1, indicating that Rm contributes substantially to total RS in this mature woodland. Monthly mean RS was correlated with monthly mean soil temperature, soil moisture and monthly stem growth rate in this dry year, but soil moisture levels may have been insufficient to observe the impacts of eCO2 on stem growth in this droughted and phosphorous limited site. Our results highlight that eCO2 tends to increase Rm at low temperatures during the non-growing season, thus decreasing the temperature sensitivity of RS, despite a neutral effect of eCO2 on RS rates on a yearly basis.

Drought strongly influences terrestrial C cycling via its effects on plant H2O and CO2 exchange. However, the treatment of photosynthetic physiology under drought by many ecosystem and earth system models remains poorly constrained by data. We measured the drought response of four tree species and evaluated alternative model formulations for drought effects on photosynthesis (A). We implemented a series of soil drying and rewetting events (i.e. multiple droughts) with four contrasting tree species in large pots (75 L) placed in the field under rainout shelters. We measured leaf-level gas exchange, predawn and midday leaf water potential (Ψpd and Ψmd), and leaf isotopic composition (δ13C) and calculated discrimination relative to the atmosphere (Δ). We then evaluated eight modeling frameworks that simulate the effects of drought in different ways. With moderate reductions in volumetric soil water content (θ), all species reduced stomatal conductance (gs), leading to an equivalent increase in water use efficiency across species inferred from both leaf gas exchange and Δ, despite a small reduction in photosynthetic capacity. With severe reductions in θ, all species strongly reduced gs along with a coincident reduction in photosynthetic capacity, illustrating the joint importance of stomatal and non-stomatal limitations of photosynthesis under strong drought conditions. Simple empirical models as well as complex mechanistic model formulations were equally successful at capturing the measured variation in A and gs, as long as the predictor variables were available from direct measurements (θ, Ψpd, and Ψmd). However, models based on leaf water potential face an additional challenge, as we found that Ψpd was substantially different from Ψsoil predicted by standard approaches based on θ. Modeling frameworks that combine gas exchange and hydraulic traits have the advantage of mechanistic realism, but sacrificed parsimony without an improvement in predictive power in this comparison. Model choice depends on the desired balance between simple empiricism and mechanistic realism. We suggest that empirical models implementing stomatal and non-stomatal limitations based on θ are highly predictive simple models. Mechanistic models that incorporate hydraulic traits have excellent potential, but several challenges currently limit their widespread implementation.

The potential of the vegetation to sequester C is determined by the balance between assimilation and respiration. Respiration is under environmental and substrate-driven control, but the circadian clock might also contribute. To assess circadian control on night-time dark respiration (RD) and on light enhanced dark respiration (LEDR) - the latter providing information on the metabolic reorganization in the leaf during light-dark transitions - we performed experiments in macrocosms hosting canopies of bean and cotton. Under constant darkness (plus constant air temperature and air humidity), we tested whether circadian regulation of RD scaled from leaf to canopy respiration. Under constant light (plus constant air temperature and air humidity), we assessed the potential for leaf-level circadian regulation of LEDR. There was a clear circadian oscillation of leaf-level RD in both species and circadian patterns scaled to the canopy. LEDR was under circadian control in cotton, but not in bean indicating species-specific controls. The circadian rhythm of LEDR in cotton might indicate variable suppression of the normal cyclic function of the tricarboxylic-acid-cycle in the light. Since circadian regulation is assumed to act as an adaptive memory to adjust plant metabolism based on environmental conditions from previous days, circadian control of RD may help to explain temporal variability of ecosystem respiration. This study benefited from the CNRS human and technical resources allocated to the ECOTRONS Research Infrastructures as well as from the state allocation ‘Investissement d'Avenir’ AnaEE-France ANR-11-INBS-0001, ExpeER Transnational Access program, Ramón y Cajal fellowships (RYC-2012-10970 to VRD and RYC-2008-02050 to JPF), the Erasmus Mundus Master Course MEDfOR, internal grants from UWS-HIE to VRD and ZALF to AG and Juan de la Cierva-fellowships (IJCI-2014-21393 to JGA). We remain indebted to E. Gerardeau, D. Dessauw, J. Jean, P. Prudent (Aïda CIRAD), J.-J. Drevon, C. Pernot (Eco&Sol INRA), B. Buatois, A. Rocheteau (CEFE CNRS), A. Pra, A. Mokhtar and the full Ecotron team, in particular C. Escape, for outstanding technical assistance.

Short-term acclimation and long-term adaptation represent two ways in which forest trees can respond to changes in temperature. Yet, the relative contribution of thermal acclimation and adaptation to tree physiological responses to temperature remains poorly understood. Here, we grew two cool-origin and two warm-origin populations of a widespread broad-leaved evergreen tree species (Corymbia calophylla (Lindl.) K.D.Hill & L.A.S.Johnson) from a Mediterranean climate in southwestern Australia under two growth temperatures representative of the cool- and warm-edge of the species distribution. The populations selected from each thermal environment represented both high and low precipitation sites. We measured the short-term temperature response of leaf photosynthesis (A) and dark respiration (R), and attributed observed variation to acclimation, adaptation or the combination of both. We observed limited variation in the temperature optimum (Topt) of A between temperature treatments or among populations, suggesting little plasticity or genetic differentiation in the Topt of A. Yet, other aspects of the temperature response of A and R were dependent upon population and growth temperature. Under cooler growth temperatures, the population from the coolest, wettest environment had the lowest A (at 25 °C) among all four populations, but exhibited the highest A (at 25 °C) under warmer growth temperatures. Populations varied in R (at 20 °C) and the temperature sensitivity of R (i.e., Q10 or activation energy) under cool, but not warm growth temperatures. However, populations showed similar yet lower R (at 20 °C) and no differences in the temperature sensitivity of R under warmer growth temperatures. We conclude that C. calophylla populations from contrasting climates vary in physiological acclimation to temperature, which might influence how this ecologically important tree species and the forests of southwestern Australia respond to climate change.

Rhizosphere processes are integral to carbon sequestration by terrestrial ecosystems in response to rising concentrations of atmospheric CO2. Yet, the nature and magnitude of rhizosphere responses to elevated CO2, particularly in nutrient and water-limited forest ecosystems, remain poorly understood. We investigated rhizosphere responses (enzyme activities and nutrient availability) to atmospheric CO2 enrichment (ambient +150 μmol CO2 mol−1) in a phosphorus-limited mature eucalypt woodland in south-eastern Australia (the EucFACE experiment). Following 17 months of treatment, the activity of rhizosphere soil exoenzymes related to starch and cellulose degradation decreased between 0 and 10 cm and increased from 10 to 30 cm depth under elevated CO2. This response was concurrent with increases in nitrogen and phosphorus availability and smaller C:P nutrient ratios in rhizosphere soil under elevated CO2. This nutrient-poor eucalypt woodland exhibited rhizosphere responses to atmospheric CO2 enrichment that increased nutrient availability in rhizosphere soil and suggest accelerated rates of soil organic matter decomposition, both of which may, in turn, promote plant growth under elevated CO2 concentrations.

Atmospheric carbon dioxide enrichment (eCO2) can enhance plant carbon uptake and growth1–5, thereby providing an important negative feedback to climate change by slowing the rate of increase of the atmospheric CO2 concentration6. Although evidence gathered from young aggrading forests has generally indicated a strong CO2 fertilization effect on biomass growth3–5, it is unclear whether mature forests respond to eCO2 in a similar way. In mature trees and forest stands7–10, photosynthetic uptake has been found to increase under eCO2 without any apparent accompanying growth response, leaving the fate of additional carbon fixed under eCO2 unclear4,5,7–11. Here using data from the first ecosystem-scale Free-Air CO2 Enrichment (FACE) experiment in a mature forest, we constructed a comprehensive ecosystem carbon budget to track the fate of carbon as the forest responded to four years of eCO2 exposure. We show that, although the eCO2 treatment of +150 parts per million (+38 per cent) above ambient levels induced a 12 per cent (+247 grams of carbon per square metre per year) increase in carbon uptake through gross primary production, this additional carbon uptake did not lead to increased carbon sequestration at the ecosystem level. Instead, the majority of the extra carbon was emitted back into the atmosphere via several respiratory fluxes, with increased soil respiration alone accounting for half of the total uptake surplus. Our results call into question the predominant thinking that the capacity of forests to act as carbon sinks will be generally enhanced under eCO2, and challenge the efficacy of climate mitigation strategies that rely on ubiquitous CO2 fertilization as a driver of increased carbon sinks in global forests. Carbon dioxide enrichment of a mature forest resulted in the emission of the excess carbon back into the atmosphere via enhanced ecosystem respiration, suggesting that mature forests may be limited in their capacity to mitigate climate change.