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Termites are important ecosystem engineers in tropical habitats, with different feeding groups able to decompose wood, grass, litter, and soil organic matter. In most tropical regions, termite abundance and species diversity are assumed to increase with rainfall, with highest levels found in rainforests. However, in the Australian tropics, this pattern is thought to be reversed, with lower species richness and termite abundance found in rainforest than drier habitats. The potential mechanisms underlying this pattern remain unclear. We compared termite assemblages (abundance, activity, diversity, and feeding group composition) across five sites along a precipitation gradient (ranging from ∼800 to 4,000 mm annual rainfall), spanning dry and wet savanna habitats, wet sclerophyll, and lowland and upland rainforests in tropical North Queensland. Moving from dry to wet habitats, we observed dramatic decreases in termite abundance in both mounds and dead wood occupancy, with greater abundance and activity at savanna sites (low precipitation) compared with rainforest or sclerophyll sites (high precipitation). We also observed a turnover in termite species and feeding group diversity across sites that were close together, but in different habitats. Termite species and feeding group richness were highest in savanna sites, with 13 termite species from wood-, litter-, grass-, dung-, and soil-feeding groups, while only five termite species were encountered in rainforest and wet sclerophyll sites—all wood feeders. These results suggest that the Australian termite diversity anomaly may be partly driven by how specific feeding groups colonized habitats across Australia. Consequently, termites in Australian rainforests may be less important in ecosystem processes, such as carbon and nutrient cycling during decomposition, compared with termites in other tropical rainforests.

Leaf-level gas exchange data support the mechanistic understanding of plant fluxes of carbon and water. These fluxes inform our understanding of ecosystem function, are an important constraint on parameterization of terrestrial biosphere models, are necessary to understand the response of plants to global environmental change, and are integral to efforts to improve crop production. Collection of these data using gas analyzers can be both technically challenging and time consuming, and individual studies generally focus on a small range of species, restricted time periods, or limited geographic regions. The high value of these data is exemplified by the many publications that reuse and synthesize gas exchange data, however the lack of metadata and data reporting conventions make full and efficient use of these data difficult. Here we propose a reporting format for leaf-level gas exchange data and metadata to provide guidance to data contributors on how to store data in repositories to maximize their discoverability, facilitate their efficient reuse, and add value to individual datasets. For data users, the reporting format will better allow data repositories to optimize data search and extraction, and more readily integrate similar data into harmonized synthesis products. The reporting format specifies data table variable naming and unit conventions, as well as metadata characterizing experimental conditions and protocols. For common data types that were the focus of this initial version of the reporting format, i.e., survey measurements, dark respiration, carbon dioxide and light response curves, and parameters derived from those measurements, we took a further step of defining required additional data and metadata that would maximize the potential reuse of those data types. To aid data contributors and the development of data ingest tools by data repositories we provided a translation table comparing the outputs of common gas exchange instruments. Extensive consultation with data collectors, data users, instrument manufacturers, and data scientists was undertaken in order to ensure that the reporting format met community needs. The reporting format presented here is intended to form a foundation for future development that will incorporate additional data types and variables as gas exchange systems and measurement approaches advance in the future. The reporting format is published in the U.S. Department of Energy's ESS-DIVE data repository, with documentation and future development efforts being maintained in a version control system.

Elevated CO2 (eCO2) experiments provide critical information to quantify the effects of rising CO2 on vegetation1–6. Many eCO2 experiments suggest that nutrient limitations modulate the local magnitude of the eCO2 effect on plant biomass1,3,5, but the global extent of these limitations has not been empirically quantified, complicating projections of the capacity of plants to take up CO27,8. Here, we present a data-driven global quantification of the eCO2 effect on biomass based on 138 eCO2 experiments. The strength of CO2 fertilization is primarily driven by nitrogen (N) in ~65% of global vegetation and by phosphorus (P) in ~25% of global vegetation, with N- or P-limitation modulated by mycorrhizal association. Our approach suggests that CO2 levels expected by 2100 can potentially enhance plant biomass by 12 ± 3% above current values, equivalent to 59 ± 13 PgC. The global-scale response to eCO2 we derive from experiments is similar to past changes in greenness9 and biomass10 with rising CO2, suggesting that CO2 will continue to stimulate plant biomass in the future despite the constraining effect of soil nutrients. Our research reconciles conflicting evidence on CO2 fertilization across scales and provides an empirical estimate of the biomass sensitivity to eCO2 that may help to constrain climate projections.

Aim: Within C3 plants, photosynthesis is a balance between CO2 supply from the atmosphere via stomata and demand by enzymes within chloroplasts. This process is dynamic and a complex but crucial aspect of photosynthesis. We sought to understand the spatial pattern in CO2 supply–demand balance on a global scale, via analysis of stable isotopes of carbon within leaves (Δ13C), which provide an integrative record of CO2 drawdown during photosynthesis. LocationGlobal. Time period1951–2011. Major taxa studiedVascular plants. Methods: We assembled a database of leaf carbon isotope ratios containing 3,979 species–site combinations from across the globe, including 3,645 for C3 species. We examined a wide array of potential climate and soil drivers of variation in Δ13C. Results: The strongest drivers of carbon isotope discrimination at the global scale included atmospheric pressure, potential evapotranspiration and soil pH, which explained 44% of the variation in Δ13C. Addition of eight more climate and soil variables (each explaining small but highly significant amounts of variation) increased the explained variation to 60%. On top of this, the largest plant trait effect was leaf nitrogen per area, which explained 11% of Δ13C variation. Main conclusions: By considering variation in Δ13C at a considerably larger scale than previously, we were able to identify and quantify key drivers in CO2 supply–demand balance previously unacknowledged. Of special note is the key role of soil properties, with greater discrimination on low-pH and high-silt soils. Unlike other plant traits, which show typically wide variation within sets of coexisting species, the global pattern in carbon stable isotope ratios is much more conservative; there is relatively narrow variation in time-integrated CO2 concentrations at the site of carboxylation among plants in a given soil and climate.

Unravelling the complexities of transpiration can be assisted by understanding the oxygen isotope composition of transpired water vapour (δE). It is often assumed that δE is at steady state, thereby mirroring the oxygen isotope composition of source water (δsource), but this assumption has never been tested at the whole-tree scale. This study utilized the unique infrastructure of 12 whole-tree chambers enclosing Eucalyptus parramattensis E.C.Hall trees to measure δE along with concurrent temperature and gas exchange data. Six chambers tracked ambient air temperature and six were exposed to an ambient +3 °C warming treatment. Day time means for δE were within 1.2‰ of δsource (-3.3‰) but varied considerably throughout the day. Our observations show that E. parramattensis trees are seldom transpiring at isotopic steady state over a diel period, but transpiration approaches source water isotopic composition over longer time periods., (© The Author(s) 2024. Published by Oxford University Press.)

Climate change not only leads to higher air temperatures but also increases the vapour pressure deficit (VPD) of the air. Understanding the direct effect of VPD on leaf gas exchange is crucial for precise modelling of stomatal functioning. We conducted combined leaf gas exchange and online isotope discrimination measurements on four common European tree species across a VPD range of 0.8-3.6 kPa, while maintaining constant temperatures without soil water limitation. In addition to applying the standard assumption of saturated vapour pressure inside leaves (e i ), we inferred e i from oxygen isotope discrimination of CO 2 and water vapour. e i desaturated progressively with increasing VPD, consistently across species, resulting in an intercellular relative humidity as low as 0.73 ± 0.11 at the highest tested VPD. Assuming saturation of e i overestimated the extent of reductions in stomatal conductance and CO 2 mole fraction inside leaves in response to increasing VPD compared with calculations that accounted for unsaturation. In addition, a significant decrease in mesophyll conductance with increasing VPD only occurred when the unsaturation of e i was considered. We suggest that the possibility of unsaturated e i should not be overlooked in measurements related to leaf gas exchange and in stomatal models, especially at high VPD., (© 2024 The Author(s). New Phytologist © 2024 New Phytologist Foundation.)

Herbivore-plant interactions are fundamental processes shaping ecosystems, yet their study is challenged by their complex connections within broader ecosystem processes, requiring a nuanced understanding of ecosystem dynamics. This study investigated the relationship between nutrient availability and insect herbivory in the Australian Wet Tropics. Our objectives were threefold. Firstly, to understand what factors influence nutrient availability for plants and herbivores across the landscape; secondly, to investigate how trees of different species respond to nutrient availability; and thirdly, to unravel how the relationships between resources and plant chemistry affect herbivory. We established a network of 25 study sites covering important abiotic gradients, including temperature, precipitation, and geology. Employing a hierarchical modelling approach, we assessed the influence of climate and geology on resource availability for plants, primarily in the form of soil nutrients. Then, we explored the influence of the above factors on the interaction between herbivory and foliage chemistry across three widespread rainforest tree species, comparing how these relationships emerged across genera. Our findings suggest an overarching influence of climate and geology over soil chemistry, foliar nitrogen, and insect herbivory, both directly and indirectly. However, individual constituents of soil fertility showed equivocal influences on spatial patterns of foliage chemistry once site geological origin was accounted for, suggesting a questionable relationship between individual soil nutrients and foliar composition. We have demonstrated that herbivore-plant interactions are complex dynamics regulated by an intricate web of relationships spanning different biogeochemical processes. While our results provide some support to the notion that herbivory is affected by resource availability, different species growing under the same conditions can show differing responses to the same resources, highlighting the importance of identifying specific limiting factors rather than simpler proxies of resource availability., (© 2024. The Author(s).)

Modern plant physiological theory stipulates that the resistance to water movement from plants to the atmosphere is overwhelmingly dominated by stomata. This conception necessitates a corollary assumption-that the air spaces in leaves must be nearly saturated with water vapour; that is, with a relative humidity that does not decline materially below unity. As this idea became progressively engrained in scientific discourse and textbooks over the last century, observations inconsistent with this corollary assumption were occasionally reported. Yet, evidence of unsaturation gained little traction, with acceptance of the prevailing framework motivated by three considerations: (1) leaf water potentials measured by either thermocouple psychrometry or the Scholander pressure chamber are largely consistent with the framework; (2) being able to assume near saturation of intercellular air spaces was transformational to leaf gas exchange analysis; and (3) there has been no obvious mechanism to explain a variable, liquid-phase resistance in the leaf mesophyll. Here, we review the evidence that refutes the assumption of universal, near saturation of air spaces in leaves. Refining the prevailing paradigm with respect to this assumption provides opportunities for identifying and developing mechanisms for increased plant productivity in the face of increasing evaporative demand imposed by global climate change., (© 2024 The Author(s). Plant, Cell & Environment published by John Wiley & Sons Ltd.)

Measurements of stable isotope ratios in organic compounds are widely used tools for plant ecophysiological studies. However, the complexity of the processes involved in shaping hydrogen isotope values (δ 2 H) in plant carbohydrates has limited its broader application. To investigate the underlying biochemical processes responsible for 2 H fractionation among water, sugars, and cellulose in leaves, we studied the three main CO 2 fixation pathways (C 3 , C 4 , and CAM) and their response to changes in temperature and vapor pressure deficit (VPD). We show significant differences in autotrophic 2 H fractionation (ε A ) from water to sugar among the pathways and their response to changes in air temperature and VPD. The strong 2 H depleting ε A in C 3 plants is likely driven by the photosynthetic H + production within the thylakoids, a reaction that is spatially separated in C 4 and strongly reduced in CAM plants, leading to the absence of 2 H depletion in the latter two types. By contrast, we found that the heterotrophic 2 H-fractionation (ε H ) from sugar to cellulose was very similar among the three pathways and is likely driven by the plant's metabolism, rather than by isotopic exchange with leaf water. Our study offers new insights into the biochemical drivers of 2 H fractionation in plant carbohydrates., (© 2024 The Author(s). New Phytologist © 2024 New Phytologist Foundation.)

To avoid reaching lethal temperatures during periods of heat stress, plants may acclimate either their biochemical thermal tolerance or leaf morphological and physiological characteristics to reduce leaf temperature (T leaf ). While plants from warmer environments may have a greater capacity to regulate T leaf , the extent of intraspecific variation and contribution of provenance is relatively unexplored. We tested whether upland and lowland provenances of four tropical tree species grown in a common garden differed in their thermal safety margins by measuring leaf thermal traits, midday leaf-to-air temperature differences (∆T leaf ) and critical leaf temperatures defined by chlorophyll fluorescence (T crit ). Provenance variation was species- and trait-specific. Higher ∆T leaf and T crit were observed in the lowland provenance for Terminalia microcarpa, and in the upland provenance for Castanospermum australe, with no provenance effects in the other two species. Within-species covariation of T crit and ∆T leaf led to a convergence of thermal safety margins across provenances. While future studies should expand the number of provenances and species investigated, our findings suggest that lowland and upland provenances may not differ substantially in their vulnerability to heat stress, as determined by thermal safety margins, despite differences in operating temperatures and T crit ., (© 2024 The Author(s). Plant, Cell & Environment published by John Wiley & Sons Ltd.)

A fundamental assumption in plant science posits that leaf air spaces remain vapor saturated, leading to the predominant view that stomata alone control leaf water loss. This concept has been pivotal in photosynthesis and water-use efficiency research. However, recent evidence has refuted this longstanding assumption by providing evidence of unsaturation in the leaf air space of C 3 plants under relatively mild vapor pressure deficit (VPD) stress. This phenomenon represents a nonstomatal mechanism restricting water loss from the mesophyll. The potential ubiquity and physiological implications of this phenomenon, its driving mechanisms in different plant species and habitats, and its interaction with other ecological adaptations have. In this context, C 4 plants spark particular interest for their importance as crops, bundle sheath cells' unique anatomical characteristics and specialized functions, and notably higher water-use efficiency relative to C 3 plants. Here, we confirm reduced relative humidities in the substomatal cavity of the C 4 plants maize, sorghum, and proso millet down to 80% under mild VPD stress. We demonstrate the critical role of nonstomatal control in these plants, indicating that the role of the CO 2 concentration mechanism in CO 2 management at a high VPD may have been overestimated. Our findings offer a mechanistic reconciliation between discrepancies in CO 2 and VPD responses reported in C 4 species. They also reveal that nonstomatal control is integral to maintaining an advantageous microclimate of relatively higher CO 2 concentrations in the mesophyll air space of C 4 plants for carbon fixation, proving vital when these plants face VPD stress., Competing Interests: Competing interests statement:The authors declare no competing interest.

Mesophyll conductance (g m ) is a crucial plant trait that can significantly limit photosynthesis. Measurement of photosynthetic C 18 O 16 O discrimination (Δ 18 O) has proved to be the only viable means of resolving g m in both C 3 and C 4 plants. However, the currently available methods to exploit Δ 18 O for g m estimation are error prone due to their inadequacy in constraining the degree of oxygen isotope exchange (θ) during mesophyll CO 2 hydration. Here, we capitalized on experimental manipulation of leaf water isotopic dynamics to establish a novel, nonsteady state, regression-based approach for simultaneous determination of g m and θ from online Δ 18 O measurements. We demonstrated the methodological and theoretical robustness of this new Δ 18 O-g m estimation approach and showed through measurements on several C 3 and C 4 species that this approach can serve as a benchmark method against which to identify previously-unrecognized biases of the existing Δ 18 O-g m methods. Our results highlight the unique value of this nonsteady state-based approach for contributing to ongoing efforts toward quantitative understanding of mesophyll conductance for crop yield improvement and carbon cycle modeling., (© 2024 The Author(s). New Phytologist © 2024 New Phytologist Foundation.)

Elevated air temperature (T air ) and vapour pressure deficit (VPD air ) significantly influence plant functioning, yet their relative impacts are difficult to disentangle. We examined the effects of elevated T air (+6°C) and VPD air (+0.7 kPa) on the growth and physiology of six tropical tree species. Saplings were grown under well-watered conditions in climate-controlled glasshouses for 6 months under three treatments: (1) low T air and low VPD air , (2) high T air and low VPD air , and (3) high T air and high VPD air . To assess acclimation, physiological parameters were measured at a set temperature. Warm-grown plants grown under elevated VPD air had significantly reduced stomatal conductance and increased instantaneous water use efficiency compared to plants grown under low VPD air . Photosynthetic biochemistry and thermal tolerance (T crit ) were unaffected by VPD air , but elevated T air caused J max25 to decrease and T crit to increase. Sapling biomass accumulation for all species responded positively to an increase in T air , but elevated VPD air limited growth. This study shows that stomatal limitation caused by even moderate increases in VPD air can decrease productivity and growth rates in tropical species independently from T air and has important implications for modelling the impacts of climate change on tropical forests., (© 2024 The Authors. New Phytologist © 2024 New Phytologist Foundation.)

Enhanced efficiency fertilizers (EEFs) can reduce nitrogen (N) losses in temperate agriculture but are less effective in the tropics. We aimed to design a new EEF and evaluate their performance in simple-to-complex tests with tropical soils and crops. We melt-extruded urea at different loadings into biodegradable polymer matrix composites using biodegradable polyhydroxyalkanoate (PHA) or polybutylene adipate-co-terephthalate (PBAT) polymers with urea distributed throughout the pellet. These contrast with commercially coated EEF that have a polymer-coated urea core. We hypothesized that matrix fertilizers would have an intermediate N release rate compared to fast release from urea or slow release from coated EEF. Nitrogen release rates in water and sand-soil columns confirmed that the matrix fertilizer formulations had a more progressive N release than a coated EEF. A more complex picture emerged from testing sorghum [Sorghum bicolor (L.) Moench] grown to maturity in large soil pots, as the different formulations resulted in minor differences in plant N accumulation and grain production. This confirms the need to consider soil interactions, microbial processes, crop physiology, and phenology for evaluating fertilizer performance. Promisingly, crop δ 15 N signatures emerged as an integrated measure of efficacy, tracking likely N conversions and losses. The three complementary evaluations combine the advantages of standardized high-throughput screening and more resource-intensive and realistic testing in a plant-soil system. We conclude that melt-blended biodegradable polymer matrix fertilizers show promise as EEF because they can be designed toward more abiotically or more microbially driven N release by selecting biopolymer type and N loading rate., (© 2024 The Authors. Journal of Environmental Quality published by Wiley Periodicals LLC on behalf of American Society of Agronomy, Crop Science Society of America, and Soil Science Society of America.)

The strong covariation of temperature and vapour pressure deficit (VPD) in nature limits our understanding of the direct effects of temperature on leaf gas exchange. Stable isotopes in CO 2 and H 2 O vapour provide mechanistic insight into physiological and biochemical processes during leaf gas exchange. We conducted combined leaf gas exchange and online isotope discrimination measurements on four common European tree species across a leaf temperature range of 5-40°C, while maintaining a constant leaf-to-air VPD (0.8 kPa) without soil water limitation. Above the optimum temperature for photosynthesis (30°C) under the controlled environmental conditions, stomatal conductance (g s ) and net photosynthesis rate (A n ) decoupled across all tested species, with g s increasing but A n decreasing. During this decoupling, mesophyll conductance (cell wall, plasma membrane and chloroplast membrane conductance) consistently and significantly decreased among species; however, this reduction did not lead to reductions in CO 2 concentration at the chloroplast surface and stroma. We question the conventional understanding that diffusional limitations of CO 2 contribute to the reduction in photosynthesis at high temperatures. We suggest that stomata and mesophyll membranes could work strategically to facilitate transpiration cooling and CO 2 supply, thus alleviating heat stress on leaf photosynthetic function, albeit at the cost of reduced water-use efficiency., (© 2024 The Authors. New Phytologist © 2024 New Phytologist Foundation.)

A fundamental assumption when using hydrogen and oxygen stable isotopes to understand ecohydrological processes is that no isotope fractionation occurs during plant water uptake/transport/redistribution. A growing body of evidence has indicated that hydrogen isotope fractionation occurs in certain environments or for certain plant species. However, whether the plant water source hydrogen isotope offset (δ 2 H offset) is a common phenomenon and how it varies among different climates and plant functional types remains unclear. Here, we demonstrated the presence of positive, negative, and zero offsets based on extensive observations of 12 plant species of 635 paired stable isotopic compositions along a strong climate gradient within an inland river basin. Both temperature and relative humidity affected δ 2 H offsets. In cool and moist environments, temperature mainly affected δ 2 H offsets negatively due to its role in physiological activity. In warm and dry environments, relative humidity mainly affected δ 2 H offsets, likely by impacting plant leaf stomatal conductance. These δ 2 H offsets also showed substantial linkages with leaf water 18 O enrichment, an indicator of transpiration and evaporative demand. Further studies focusing on the ecophysiological and biochemical understanding of plant δ 2 H dynamics under specific environments are essential for understanding regional ecohydrological processes and for conducting paleoclimate reconstructions., (© 2024 The Authors. New Phytologist © 2024 New Phytologist Foundation.)

Woody biomass is a large carbon store in terrestrial ecosystems. In calculating biomass, tree stems are assumed to be solid structures. However, decomposer agents such as microbes and insects target stem heartwood, causing internal wood decay which is poorly quantified. We investigated internal stem damage across five sites in tropical Australia along a precipitation gradient. We estimated the amount of internal aboveground biomass damaged in living trees and measured four potential stem damage predictors: wood density, stem diameter, annual precipitation, and termite pressure (measured as termite damage in downed deadwood). Stem damage increased with increasing diameter, wood density, and termite pressure and decreased with increasing precipitation. High wood density stems sustained less damage in wet sites and more damage in dry sites, likely a result of shifting decomposer communities and their differing responses to changes in tree species and wood traits across sites. Incorporating stem damage reduced aboveground biomass estimates by > 30% in Australian savannas, compared to only 3% in rainforests. Accurate estimates of carbon storage across woody plant communities are critical for understanding the global carbon budget. Future biomass estimates should consider stem damage in concert with the effects of changes in decomposer communities and abiotic conditions., (© 2023 The Authors. New Phytologist © 2023 New Phytologist Foundation.)

Sugarcane is a vital commodity crop often grown in (sub)tropical regions which have been experiencing a recent deterioration in air quality. Unlike for other commodity crops, the risk of air pollution, specifically ozone (O 3 ), to this C 4 crop has not yet been quantified. Yet, recent work has highlighted both the potential risks of O 3 to C 4 bioenergy crops, and the emergence of O 3 exposure across the tropics as a vital factor determining global food security. Given the large extent, and planned expansion of sugarcane production in places like Brazil to meet global demand for biofuels, there is a pressing need to characterize the risk of O 3 to the industry. In this study, we sought to a) derive sugarcane O 3 dose-response functions across a range of realistic O 3 exposure and b) model the implications of this across a globally important production area. We found a significant impact of O 3 on biomass allocation (especially to leaves) and production across a range of sugarcane genotypes, including two commercially relevant varieties (e.g. CTC4, Q240). Using these data, we calculated dose-response functions for sugarcane and combined them with hourly O 3 exposure across south-central Brazil derived from the UK Earth System Model (UKESM1) to simulate the current regional impact of O 3 on sugarcane production using a dynamic global vegetation model (JULES vn 5.6). We found that between 5.6 % and 18.3 % of total crop productivity is likely lost across the region due to the direct impacts of current O 3 exposure. However, impacts depended critically on the substantial differences in O 3 susceptibility observed among sugarcane genotypes and how these were implemented in the model. Our work highlights not only the urgent need to fully elucidate the impacts of O 3 in this important bioenergetic crop, but the potential implications air quality may have upon tropical food production more generally., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2023 The Authors. Published by Elsevier B.V. All rights reserved.)

Tropospheric ozone (O3 ) is a global air pollutant that adversely affects plant growth. Whereas the impacts of O3 have previously been examined for some tropical commodity crops, no information is available for the pantropical crop, banana (Musa spp.). To address this, we exposed Australia's major banana cultivar, Williams, to a range of [O3 ] in open top chambers. In addition, we examined 46 diverse Musa lines growing in a common garden for variation in three traits that are hypothesised to shape responses to O3 : (1) leaf mass per area; (2) intrinsic water use efficiency; and (3) total antioxidant capacity. We show that O3 exposure had a significant effect on the biomass of cv. Williams, with significant reductions in both pseudostem and sucker biomass with increasing [O3 ]. This was accompanied by a significant increase in total antioxidant capacity and phenolic concentrations in older, but not younger, leaves, indicating the importance of cumulative O3 exposure. Using the observed trait diversity, we projected O3 tolerance among the 46 Musa lines growing in the common garden. Of these, cv. Williams ranked as one of the most O3 -tolerant cultivars. This suggests that other genetic lines could be even more susceptible, with implications for banana production and food security throughout the tropics.

The oxygen isotope composition (δ 18 O) of tree-ring cellulose is used to evaluate tree physiological responses to climate, but their interpretation is still limited due to the complexity of the isotope fractionation pathways. We assessed the relative contribution of seasonal needle and xylem water δ 18 O variations to the intra-annual tree-ring cellulose δ 18 O signature of larch trees at two sites with contrasting soil water availability in the Swiss Alps. We combined biweekly δ 18 O measurements of soil water, needle water, and twig xylem water with intra-annual δ 18 O measurements of tree-ring cellulose, xylogenesis analysis, and mechanistic and structural equation modeling. Intra-annual cellulose δ 18 O values resembled source water δ 18 O mean levels better than needle water δ 18 O. Large parts of the rings were formed under high proportional exchange with unenriched xylem water (p ex ). Maximum p ex values were achieved in August and imprinted on sections at 50-75% of the ring. High p ex values were associated with periods of high atmospheric evaporative demand (VPD). While VPD governed needle water δ 18 O variability, we estimated a limited Péclet effect at both sites. Due to a variable p ex , source water has a strong influence over large parts of the intra-annual tree-ring cellulose δ 18 O variations, potentially masking signals coming from needle-level processes., (© 2023 The Authors. New Phytologist © 2023 New Phytologist Foundation.)

Oxygen and hydrogen isotopes of cellulose in plant biology are commonly used to infer environmental conditions, often from time series measurements of tree rings. However, the covariation (or the lack thereof) between δ 18 O and δ 2 H in plant cellulose is still poorly understood. We compared plant water, and leaf and branch cellulose from dominant tree species across an aridity gradient in Northern Australia, to examine how δ 18 O and δ 2 H relate to each other and to mean annual precipitation (MAP). We identified a decline in covariation from xylem to leaf water, and onwards from leaf to branch wood cellulose. Covariation in leaf water isotopic enrichment (Δ) was partially preserved in leaf cellulose but not branch wood cellulose. Furthermore, whilst δ 2 H was well-correlated between leaf and branch, there was an offset in δ 18 O between organs that increased with decreasing MAP. Our findings strongly suggest that postphotosynthetic isotope exchange with water is more apparent for oxygen isotopes, whereas variable kinetic and nonequilibrium isotope effects add complexity to interpreting metabolic-induced δ 2 H patterns. Varying oxygen isotope exchange in wood and leaf cellulose must be accounted for when δ 18 O is used to reconstruct climatic scenarios. Conversely, comparing δ 2 H and δ 18 O patterns may reveal environmentally induced shifts in metabolism., (© 2023 The Authors. New Phytologist © 2023 New Phytologist Foundation.)

Atmospheric conditions are expected to become warmer and drier in the future, but little is known about how evaporative demand influences forest structure and function independently from soil moisture availability, and how fast-response variables (such as canopy water potential and stomatal conductance) may mediate longer-term changes in forest structure and function in response to climate change. We used two tropical rainforest sites with different temperatures and vapour pressure deficits (VPD), but nonlimiting soil water supply, to assess the impact of evaporative demand on ecophysiological function and forest structure. Common species between sites allowed us to test the extent to which species composition, relative abundance and intraspecific variability contributed to site-level differences. The highest VPD site had lower midday canopy water potentials, canopy conductance (g c ), annual transpiration, forest stature, and biomass, while the transpiration rate was less sensitive to changes in VPD; it also had different height-diameter allometry (accounting for 51% of the difference in biomass between sites) and higher plot-level wood density. Our findings suggest that increases in VPD, even in the absence of soil water limitation, influence fast-response variables, such as canopy water potentials and g c , potentially leading to longer-term changes in forest stature resulting in reductions in biomass., (© 2023 The Authors. New Phytologist © 2023 New Phytologist Foundation.)

Climate change is expected to increase the intensity and occurrence of drought in tropical regions, potentially affecting the phenology and physiology of tree species. Phenological activity may respond to a drying and warming environment by advancing reproductive timing and/or diminishing the production of flowers and fruits. These changes have the potential to disrupt important ecological processes, with potentially wide-ranging effects on tropical forest function. Here, we analysed the monthly flowering and fruiting phenology of a tree community (337 individuals from 30 species) over 7 years in a lowland tropical rainforest in northeastern Australia and its response to a throughfall exclusion drought experiment (TFE) that was carried out from 2016 to 2018 (3 years), excluding approximately 30% of rainfall. We further examined the ecophysiological effects of the TFE on the elemental (C:N) and stable isotope ( δ 13 C and δ 15 N) composition of leaves, and on the stable isotope composition ( δ 13 C and δ 18 O) of stem wood of four tree species. At the community level, there was no detectable effect of the TFE on flowering activity overall, but there was a significant effect recorded on fruiting and varying responses from the selected species. The reproductive phenology and physiology of the four species examined in detail were largely resistant to impacts of the TFE treatment. One canopy species in the TFE significantly increased in fruiting and flowering activity, whereas one understory species decreased significantly in both. There was a significant interaction between the TFE treatment and season on leaf C:N for two species. Stable isotope responses were also variable among species, indicating species-specific responses to the TFE. Thus, we did not observe consistent patterns in physiological and phenological changes in the tree community within the 3 years of TFE treatment examined in this study., Competing Interests: The authors have no conflict of interest to declare., (© The Author(s) 2023. Published by Oxford University Press and the Society for Experimental Biology.)

As the global climate warms, a key question is how increased leaf temperatures will affect tree physiology and the coupling between leaf and air temperatures in forests. To explore the impact of increasing temperatures on plant performance in open air, we warmed leaves in the canopy of two mature evergreen forests, a temperate Eucalyptus woodland and a tropical rainforest. The leaf heaters consistently maintained leaves at a target of 4 °C above ambient leaf temperatures. Ambient leaf temperatures (Tleaf) were mostly coupled to air temperatures (Tair), but at times, leaves could be 8-10 °C warmer than ambient air temperatures, especially in full sun. At both sites, Tleaf was warmer at higher air temperatures (Tair > 25 °C), but was cooler at lower Tair, contrary to the 'leaf homeothermy hypothesis'. Warmed leaves showed significantly lower stomatal conductance (-0.05 mol m-2 s-1 or -43% across species) and net photosynthesis (-3.91 μmol m-2 s-1 or -39%), with similar rates in leaf respiration rates at a common temperature (no acclimation). Increased canopy leaf temperatures due to future warming could reduce carbon assimilation via reduced photosynthesis in these forests, potentially weakening the land carbon sink in tropical and temperate forests., (© The Author(s) 2023. Published by Oxford University Press.)

We present a robust estimation of the CO 2 concentration at the surface of photosynthetic mesophyll cells (c w ), applicable under reasonable assumptions of assimilation distribution within the leaf. We used Capsicum annuum, Helianthus annuus and Gossypium hirsutumas model plants for our experiments. We introduce calculations to estimate c w using independent adaxial and abaxial gas exchange measurements, and accounting for the mesophyll airspace resistances. The c w was lower than adaxial and abaxial estimated intercellular CO 2 concentrations (c i ). Differences between c w and the c i of each surface were usually larger than 10 μmol mol -1 . Differences between adaxial and abaxial c i ranged from a few μmol mol -1 to almost 50 μmol mol -1 , where the largest differences were found at high air saturation deficits (ASD). Differences between adaxial and abaxial c i and the c i estimated by mixing both fluxes ranged from -30 to +20 μmol mol -1 , where the largest differences were found under high ASD or high ambient CO 2 concentrations. Accounting for c w improves the information that can be extracted from gas exchange experiments, allowing a more detailed description of the CO 2 and water vapor gradients within the leaf., (© 2023 The Authors. New Phytologist © 2023 New Phytologist Foundation.)

"Least-cost theory" posits that C 3 plants should balance rates of photosynthetic water loss and carboxylation in relation to the relative acquisition and maintenance costs of resources required for these activities. Here we investigated the dependency of photosynthetic traits on climate and soil properties using a new Australia-wide trait dataset spanning 528 species from 67 sites. We tested the hypotheses that plants on relatively cold or dry sites, or on relatively more fertile sites, would typically operate at greater CO 2 drawdown (lower ratio of leaf internal to ambient CO 2 , C i :C a ) during light-saturated photosynthesis, and at higher leaf N per area (N area ) and higher carboxylation capacity (V cmax 25 ) for a given rate of stomatal conductance to water vapour, g sw . These results would be indicative of plants having relatively higher water costs than nutrient costs. In general, our hypotheses were supported. Soil total phosphorus (P) concentration and (more weakly) soil pH exerted positive effects on the N area -g sw and V cmax 25 -g sw slopes, and negative effects on C i :C a . The P effect strengthened when the effect of climate was removed via partial regression. We observed similar trends with increasing soil cation exchange capacity and clay content, which affect soil nutrient availability, and found that soil properties explained similar amounts of variation in the focal traits as climate did. Although climate typically explained more trait variation than soil did, together they explained up to 52% of variation in the slope relationships and soil properties explained up to 30% of the variation in individual traits. Soils influenced photosynthetic traits as well as their coordination. In particular, the influence of soil P likely reflects the Australia's geologically ancient low-relief landscapes with highly leached soils. Least-cost theory provides a valuable framework for understanding trade-offs between resource costs and use in plants, including limiting soil nutrients., (© 2022 The Authors. Global Change Biology published by John Wiley & Sons Ltd.)

Deadwood is a large global carbon store with its store size partially determined by biotic decay. Microbial wood decay rates are known to respond to changing temperature and precipitation. Termites are also important decomposers in the tropics but are less well studied. An understanding of their climate sensitivities is needed to estimate climate change effects on wood carbon pools. Using data from 133 sites spanning six continents, we found that termite wood discovery and consumption were highly sensitive to temperature (with decay increasing >6.8 times per 10°C increase in temperature)-even more so than microbes. Termite decay effects were greatest in tropical seasonal forests, tropical savannas, and subtropical deserts. With tropicalization (i.e., warming shifts to tropical climates), termite wood decay will likely increase as termites access more of Earth's surface.

Tropical forests take up more carbon (C) from the atmosphere per annum by photosynthesis than any other type of vegetation. Phosphorus (P) limitations to C uptake are paramount for tropical and subtropical forests around the globe. Yet the generality of photosynthesis-P relationships underlying these limitations are in question, and hence are not represented well in terrestrial biosphere models. Here we demonstrate the dependence of photosynthesis and underlying processes on both leaf N and P concentrations. The regulation of photosynthetic capacity by P was similar across four continents. Implementing P constraints in the ORCHIDEE-CNP model, gross photosynthesis was reduced by 36% across the tropics and subtropics relative to traditional N constraints and unlimiting leaf P. Our results provide a quantitative relationship for the P dependence for photosynthesis for the front-end of global terrestrial C models that is consistent with canopy leaf measurements., (© 2022. The Author(s).)

Evidence exists that tree mortality is accelerating in some regions of the tropics 1,2 , with profound consequences for the future of the tropical carbon sink and the global anthropogenic carbon budget left to limit peak global warming below 2 °C. However, the mechanisms that may be driving such mortality changes and whether particular species are especially vulnerable remain unclear 3-8 . Here we analyse a 49-year record of tree dynamics from 24 old-growth forest plots encompassing a broad climatic gradient across the Australian moist tropics and find that annual tree mortality risk has, on average, doubled across all plots and species over the last 35 years, indicating a potential halving in life expectancy and carbon residence time. Associated losses in biomass were not offset by gains from growth and recruitment. Plots in less moist local climates presented higher average mortality risk, but local mean climate did not predict the pace of temporal increase in mortality risk. Species varied in the trajectories of their mortality risk, with the highest average risk found nearer to the upper end of the atmospheric vapour pressure deficit niches of species. A long-term increase in vapour pressure deficit was evident across the region, suggesting that thresholds involving atmospheric water stress, driven by global warming, may be a primary cause of increasing tree mortality in moist tropical forests., (© 2022. The Author(s), under exclusive licence to Springer Nature Limited.)

Stomata are orifices that connect the drier atmosphere with the interconnected network of more humid air spaces that surround the cells within a leaf. Accurate values of the humidities inside the substomatal cavity, w i , and in the air, w a , are needed to estimate stomatal conductance and the CO 2 concentration in the internal air spaces of leaves. Both are vital factors in the understanding of plant physiology and climate, ecological and crop systems. However, there is no easy way to measure w i directly. Out of necessity, w i has been taken as the saturation water vapour concentration at leaf temperature, w sat , and applied to the whole leaf intercellular air spaces. We explored the occurrence of unsaturation by examining gas exchange of leaves exposed to various magnitudes of w sat  - w a , or Δw, using a double-sided, clamp-on chamber, and estimated degrees of unsaturation from the gradient of CO 2 across the leaf that was required to sustain the rate of CO 2 assimilation through the upper surface. The relative humidity in the substomatal cavities dropped to about 97% under mild Δw and as dry as around 80% when Δw was large. Measurements of the diffusion of noble gases across the leaf indicated that there were still regions of near 100% humidity distal from the stomatal pores. We suggest that as Δw increases, the saturation edge retreats into the intercellular air spaces, accompanied by the progressive closure of mesophyll aquaporins to maintain the cytosolic water potential., (© 2022. The Author(s), under exclusive licence to Springer Nature Limited.)

We compiled hydrogen and oxygen stable isotope compositions (δ 2 H and δ 18 O) of leaf water from multiple biomes to examine variations with environmental drivers. Leaf water δ 2 H was more closely correlated with δ 2 H of xylem water or atmospheric vapour, whereas leaf water δ 18 O was more closely correlated with air relative humidity. This resulted from the larger proportional range for δ 2 H of meteoric waters relative to the extent of leaf water evaporative enrichment compared with δ 18 O. We next expressed leaf water as isotopic enrichment above xylem water (Δ 2 H and Δ 18 O) to remove the impact of xylem water isotopic variation. For Δ 2 H, leaf water still correlated with atmospheric vapour, whereas Δ 18 O showed no such correlation. This was explained by covariance between air relative humidity and the Δ 18 O of atmospheric vapour. This is consistent with a previously observed diurnal correlation between air relative humidity and the deuterium excess of atmospheric vapour across a range of ecosystems. We conclude that 2 H and 18 O in leaf water do indeed reflect the balance of environmental drivers differently; our results have implications for understanding isotopic effects associated with water cycling in terrestrial ecosystems and for inferring environmental change from isotopic biomarkers that act as proxies for leaf water., (© 2022 The Authors New Phytologist © 2022 New Phytologist Foundation.)

Predicting species-level responses to drought at the landscape scale is critical to reducing uncertainty in future terrestrial carbon and water cycle projections. We embedded a stomatal optimisation model in the Community Atmosphere Biosphere Land Exchange (CABLE) land surface model and parameterised the model for 15 canopy dominant eucalypt tree species across South-Eastern Australia (mean annual precipitation range: 344-1424 mm yr -1 ). We conducted three experiments: applying CABLE to the 2017-2019 drought; a 20% drier drought; and a 20% drier drought with a doubling of atmospheric carbon dioxide (CO 2 ). The severity of the drought was highlighted as for at least 25% of their distribution ranges, 60% of species experienced leaf water potentials beyond the water potential at which 50% of hydraulic conductivity is lost due to embolism. We identified areas of severe hydraulic stress within-species' ranges, but we also pinpointed resilience in species found in predominantly semiarid areas. The importance of the role of CO 2 in ameliorating drought stress was consistent across species. Our results represent an important advance in our capacity to forecast the resilience of individual tree species, providing an evidence base for decision-making around the resilience of restoration plantings or net-zero emission strategies., (© 2022 The Authors New Phytologist © 2022 New Phytologist Foundation.)

Tropical forests are some of the most biodiverse ecosystems in the world, yet their functioning is threatened by anthropogenic disturbances and climate change. Global actions to conserve tropical forests could be enhanced by having local knowledge on the forests' functional diversity and functional redundancy as proxies for their capacity to respond to global environmental change. Here we create estimates of plant functional diversity and redundancy across the tropics by combining a dataset of 16 morphological, chemical and photosynthetic plant traits sampled from 2,461 individual trees from 74 sites distributed across four continents together with local climate data for the past half century. Our findings suggest a strong link between climate and functional diversity and redundancy with the three trait groups responding similarly across the tropics and climate gradient. We show that drier tropical forests are overall less functionally diverse than wetter forests and that functional redundancy declines with increasing soil water and vapour pressure deficits. Areas with high functional diversity and high functional redundancy tend to better maintain ecosystem functioning, such as aboveground biomass, after extreme weather events. Our predictions suggest that the lower functional diversity and lower functional redundancy of drier tropical forests, in comparison with wetter forests, may leave them more at risk of shifting towards alternative states in face of further declines in water availability across tropical regions., (© 2022. The Author(s), under exclusive licence to Springer Nature Limited.)

In 2020, the Australian and New Zealand flux research and monitoring network, OzFlux, celebrated its 20 th anniversary by reflecting on the lessons learned through two decades of ecosystem studies on global change biology. OzFlux is a network not only for ecosystem researchers, but also for those 'next users' of the knowledge, information and data that such networks provide. Here, we focus on eight lessons across topics of climate change and variability, disturbance and resilience, drought and heat stress and synergies with remote sensing and modelling. In distilling the key lessons learned, we also identify where further research is needed to fill knowledge gaps and improve the utility and relevance of the outputs from OzFlux. Extreme climate variability across Australia and New Zealand (droughts and flooding rains) provides a natural laboratory for a global understanding of ecosystems in this time of accelerating climate change. As evidence of worsening global fire risk emerges, the natural ability of these ecosystems to recover from disturbances, such as fire and cyclones, provides lessons on adaptation and resilience to disturbance. Drought and heatwaves are common occurrences across large parts of the region and can tip an ecosystem's carbon budget from a net CO 2 sink to a net CO 2 source. Despite such responses to stress, ecosystems at OzFlux sites show their resilience to climate variability by rapidly pivoting back to a strong carbon sink upon the return of favourable conditions. Located in under-represented areas, OzFlux data have the potential for reducing uncertainties in global remote sensing products, and these data provide several opportunities to develop new theories and improve our ecosystem models. The accumulated impacts of these lessons over the last 20 years highlights the value of long-term flux observations for natural and managed systems. A future vision for OzFlux includes ongoing and newly developed synergies with ecophysiologists, ecologists, geologists, remote sensors and modellers., (© 2022 The Authors. Global Change Biology published by John Wiley & Sons Ltd.)

Background and Aims: Despite the critical role of woody tissues in determining net carbon exchange of terrestrial ecosystems, relatively little is known regarding the drivers of sapwood and bark respiration., Methods: Using one of the most comprehensive wood respiration datasets to date (82 species from Australian rainforest, savanna and temperate forest), we quantified relationships between tissue respiration rates (Rd) measured in vitro (i.e. 'respiration potential') and physical properties of bark and sapwood, and nitrogen concentration (Nmass) of leaves, sapwood and bark., Key Results: Across all sites, tissue density and thickness explained similar, and in some cases more, variation in bark and sapwood Rd than did Nmass. Higher density bark and sapwood tissues had lower Rd for a given Nmass than lower density tissues. Rd-Nmass slopes were less steep in thicker compared with thinner-barked species and less steep in sapwood than in bark. Including the interactive effects of Nmass, density and thickness significantly increased the explanatory power for bark and sapwood respiration in branches. Among these models, Nmass contributed more to explanatory power in trunks than in branches, and in sapwood than in bark. Our findings were largely consistent across sites, which varied in their climate, soils and dominant vegetation type, suggesting generality in the observed trait relationships. Compared with a global compilation of leaf, stem and root data, Australian species showed generally lower Rd and Nmass, and less steep Rd-Nmass relationships., Conclusions: To the best of our knowledge, this is the first study to report control of respiration-nitrogen relationships by physical properties of tissues, and one of few to report respiration-nitrogen relationships in bark and sapwood. Together, our findings indicate a potential path towards improving current estimates of autotrophic respiration by integrating variation across distinct plant tissues., (© The Author(s) 2022. Published by Oxford University Press on behalf of the Annals of Botany Company. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

Between late 2015 and early 2016, more than 7,000 ha of mangrove forest died along the coastline of the Gulf of Carpentaria, in northern Australia. This massive die-off was preceded by a strong 2015/2016 El Niño event, resulting in lower precipitation, a drop in sea level and higher than average temperatures in northern Australia. In this study, we investigated the role of hydraulic failure in the mortality and recovery of the dominant species, Avicennia marina , 2 years after the mortality event. We measured predawn water potential (Ψ pd ) and percent loss of stem hydraulic conductivity (PLC) in surviving individuals across a gradient of impact. We also assessed the vulnerability to drought-induced embolism (Ψ 50 ) for the species. Areas with severe canopy dieback had higher native PLC (39%) than minimally impacted areas (6%), suggesting that hydraulic recovery was ongoing. The high resistance of A. marina to water-stress-induced embolism (Ψ 50 = -9.6 MPa), indicates that severe water stress (Ψ pd < -10 MPa) would have been required to cause mortality in this species. Our data indicate that the natural gradient of water-stress enhanced the impact of El Niño, leading to hydraulic failure and mortality in A. marina growing on severely impacted (SI) zones. It is likely that lowered sea levels and less frequent inundation by seawater, combined with lower inputs of fresh water, high evaporative demand and high temperatures, led to the development of hyper-salinity and extreme water stress during the 2015/16 summer., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2022 Gauthey, Backes, Balland, Alam, Maher, Cernusak, Duke, Medlyn, Tissue and Choat.)

A better understanding of how climate affects growth in tree species is essential for improved predictions of forest dynamics under climate change. Long-term climate averages (mean climate) drive spatial variations in species' baseline growth rates, whereas deviations from these averages over time (anomalies) can create growth variation around the local baseline. However, the rarity of long-term tree census data spanning climatic gradients has so far limited our understanding of their respective role, especially in tropical systems. Furthermore, tree growth sensitivity to climate is likely to vary widely among species, and the ecological strategies underlying these differences remain poorly understood. Here, we utilize an exceptional dataset of 49 years of growth data for 509 tree species across 23 tropical rainforest plots along a climatic gradient to examine how multiannual tree growth responds to both climate means and anomalies, and how species' functional traits mediate these growth responses to climate. We show that anomalous increases in atmospheric evaporative demand and solar radiation consistently reduced tree growth. Drier forests and fast-growing species were more sensitive to water stress anomalies. In addition, species traits related to water use and photosynthesis partly explained differences in growth sensitivity to both climate means and anomalies. Our study demonstrates that both climate means and anomalies shape tree growth in tropical forests and that species traits can provide insights into understanding these demographic responses to climate change, offering a promising way forward to forecast tropical forest dynamics under different climate trajectories., (© 2021 The Authors. Global Change Biology published by John Wiley & Sons Ltd.)

Soil-leaf hydraulic conductance determines canopy-atmosphere coupling in vegetation models, but it is typically derived from ex-situ measurements of stem segments and soil samples. Using a novel approach, we derive robust in-situ estimates for whole-tree conductance (k tree ), 'functional' soil conductance (k soil ), and 'system' conductance (k system , water table to canopy), at two climatically different tropical rainforest sites. Hydraulic 'functional rooting depth', determined for each tree using profiles of soil water potential (Ψ soil ) and sap flux data, enabled a robust determination of k tree and k soil . k tree was compared across species, size classes, seasons, height above nearest drainage (HAND), two field sites, and to alternative representations of k tree ; k soil was analysed with respect to variations in site, season and HAND. k tree was lower and changed seasonally at the site with higher vapour pressure deficit (VPD) and rainfall; k tree differed little across species but scaled with tree circumference; r soil (1/k soil ) ranged from 0 in the wet season to 10× less than r tree (1/k tree ) in the dry season. VPD and not rainfall may influence plot-level k; leaf water potentials and sap flux can be used to determine k tree , k soil and k system ; Ψ soil profiles can provide mechanistic insights into ecosystem-level water fluxes., (© 2021 The Authors. New Phytologist © 2021 New Phytologist Foundation.)

Generalised dose-response curves are essential to understand how plants acclimate to atmospheric CO 2 . We carried out a meta-analysis of 630 experiments in which C 3 plants were experimentally grown at different [CO 2 ] under relatively benign conditions, and derived dose-response curves for 85 phenotypic traits. These curves were characterised by form, plasticity, consistency and reliability. Considered over a range of 200-1200 µmol mol -1 CO 2 , some traits more than doubled (e.g. area-based photosynthesis; intrinsic water-use efficiency), whereas others more than halved (area-based transpiration). At current atmospheric [CO 2 ], 64% of the total stimulation in biomass over the 200-1200 µmol mol -1 range has already been realised. We also mapped the trait responses of plants to [CO 2 ] against those we have quantified before for light intensity. For most traits, CO 2 and light responses were of similar direction. However, some traits (such as reproductive effort) only responded to light, others (such as plant height) only to [CO 2 ], and some traits (such as area-based transpiration) responded in opposite directions. This synthesis provides a comprehensive picture of plant responses to [CO 2 ] at different integration levels and offers the quantitative dose-response curves that can be used to improve global change simulation models., (© 2021 The Authors. New Phytologist © 2021 New Phytologist Foundation.)