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Future climate presents conflicting implications for forest biomass. We evaluate how plant hydraulic traits, elevated CO2 levels, warming, and changes in precipitation affect forest primary productivity, evapotranspiration, and the risk of hydraulic failure. We used a dynamic vegetation model with plant hydrodynamics (FATES-HYDRO) to simulate the stand-level responses to future climate changes in a wet tropical forest in Barro Colorado Island, Panama. We calibrated the model by selecting plant trait assemblages that performed well against observations. These assemblages were run with temperature and precipitation changes for two greenhouse gas emission scenarios (2086-2100: SSP2-45, SSP5-85) and two CO2 levels (contemporary, anticipated). The risk of hydraulic failure is projected to increase from a contemporary rate of 5.7% to 10.1-11.3% under future climate scenarios, and, crucially, elevated CO2 provided only slight amelioration. By contrast, elevated CO2 mitigated GPP reductions. We attribute a greater variation in hydraulic failure risk to trait assemblages than to either CO2 or climate. Our results project forests with both faster growth (through productivity increases) and higher mortality rates (through increasing rates of hydraulic failure) in the neo-tropics accompanied by certain trait plant assemblages becoming nonviable.

While traditionally considered important mainly in hypoxic roots during flooding, upregulation of fermentation pathways in plants has recently been described as an evolutionarily conserved drought survival strategy, with acetate signaling mediating reprograming of transcription and cellular carbon and energy metabolism from roots to leaves. The amount of acetate produced directly correlates with survival through potential mechanisms including defense gene activation, biosynthesis of primary and secondary metabolites, and aerobic respiration. Here, we review root ethanolic fermentation responses to hypoxia during saturated soil conditions and summarize studies highlighting acetate fermentation under aerobic conditions coupled with respiration during growth and drought responses. Recent work is discussed demonstrating long-distance transport of acetate via the transpiration stream as a respiratory substrate. While maintenance and growth respiration are often modeled separately in terrestrial models, here we propose the concept of 'Defense Respiration' fueled by acetate fermentation in which upregulation of acetate fermentation contributes acetate substrate for alternative energy production via aerobic respiration, biosynthesis of primary and secondary metabolites, and the acetylation of proteins involved in defense gene regulation. Finally, we highlight new frontiers in leaf-atmosphere emission measurements as a potential way to study acetate fermentation responses of individual leaves, branches, ecosystems, and regions.

Tropical forest dynamics play a crucial role in the global carbon, water, and energy cycles. However, realistically simulating the dynamics of competition and coexistence between different plant functional types (PFTs) in tropical forests remains a significant challenge. This study aims to improve the modeling of PFT coexistence in the Functionally Assembled Terrestrial Ecosystem Simulator (FATES), a vegetation demography model implemented in the Energy Exascale Earth System Model (E3SM) land model (ELM), ELM-FATES. Specifically, we explore (1) whether plant trait relationships established from field measurements can constrain ELM-FATES simulations and (2) whether machine learning (ML)-based surrogate models can emulate the complex ELM-FATES model and optimize parameter selections to improve PFT coexistence modeling. We conducted three ensembles of ELM-FATES experiments at a tropical forest site near Manaus, Brazil. By comparing the ensemble experiments without (Exp-CTR) and with (Exp-OBS) consideration of observed trait relationships, we found that accounting for these relationships slightly improves the simulations of water, energy, and carbon variables when compared to observations but degrades the simulation of PFT coexistence. Using ML-based surrogate models trained on Exp-CTR, we optimized the trait parameters in ELM-FATES and conducted another ensemble of experiments (Exp-ML) with these optimized parameters. The proportion of PFT coexistence experiments significantly increased from 21 % in Exp-CTR to 73 % in Exp-ML. After filtering the experiments that allow for PFT coexistence to agree with observations (within 15 % tolerance), 33 % of the Exp-ML experiments were retained, which is a significant improvement compared to the 1.4 % in Exp-CTR. Exp-ML also accurately reproduces the annual means and seasonal variations in water, energy, and carbon fluxes and the field inventory of aboveground biomass. This study represents a reproducible method that utilizes machine learning to identify parameter values that improve model fidelity against observations and PFT coexistence in vegetation demography models for diverse ecosystems. Our study also suggests the need for new mechanisms to enhance the robust simulation of coexisting plants in ELM-FATES and has significant implications for modeling the response and feedbacks of ecosystem dynamics to climate change.

Key message: Annual stem CO2 efflux increases with stem wood production rates and are inhibited by daily moisture stress. Abstract: Tropical forests cycle a large amount of CO2 between the land and atmosphere, with a substantial portion of the return flux due tree respiratory processes. However, in situ estimates of woody tissue respiratory fluxes and carbon use efficiencies (CUEW) and their dependencies on physiological processes including stem wood production (Pw) and transpiration in tropical forests remain scarce. Here, we synthesize monthly Pw and daytime stem CO2 efflux (ES) measurements over 1 year from 80 trees with variable biomass accumulation rates in the central Amazon. On average, carbon flux to woody tissues, expressed in the same stem area normalized units as ES, averaged 0.90 ± 1.2 µmol m−2 s−1 for Pw, and 0.55 ± 0.33 µmol m−2 s−1 for daytime ES. A positive linear correlation was found between stem growth rates and stem CO2 efflux, with respiratory carbon loss equivalent to 15 ± 3% of stem carbon accrual. CUEW of stems was non-linearly correlated with growth and was as high as 77–87% for a fast-growing tree. Diurnal measurements of stem CO2 efflux for three individuals showed a daytime reduction of ES by 15–50% during periods of high sap flow and transpiration. The results demonstrate that high daytime ES fluxes are associated with high CUEW during fast tree growth, reaching higher values than previously observed in the Amazon Basin (e.g., maximum CUEW up to 77–87%, versus 30–56%). The observations are consistent with the emerging view that diurnal dynamics of stem water status influences growth processes and associated respiratory metabolism.

Transpiration is often considered to be light- but not water-limited in humid tropical rainforests due to abundant soil water, even during the dry seasons. The record-breaking 2015-16 El Niño drought provided a unique opportunity to examine whether transpiration is constrained by water under severe lack of rainfall. We measured sap velocity, soil water content, and meteorological variables in an old-growth upland forest in the Central Amazon throughout the 2015-16 drought. We found a rapid decline in sap velocity (-38 ± 21%, mean ± SD.) and in its temporal variability (-88%) during the drought compared to the wet season. Such changes were accompanied by a marked decline in soil moisture and an increase in temperature and vapor pressure deficit. Sap velocity was largely limited by net radiation during the wet and normal dry seasons; however, it shifted to be primarily limited by soil moisture during the drought. The threshold in which sap velocity became dominated by soil moisture was at 0.33 m3 m-3 (around -150 kPa in soil matric potential), below which sap velocity dropped steeply. Our study provides evidence for a soil water threshold on transpiration in a moist tropical forest, suggesting a shift from light limitation to water limitation under future climate characterized by increased temperature and an increased frequency, intensity, duration and extent of extreme drought events.

Increasing severity and frequency of drought is predicted for large portions of the terrestrial biosphere, with major impacts already documented in wet tropical forests. Using a 4-year rainfall exclusion experiment in the Daintree Rainforest in northeast Australia, we examined canopy tree responses to reduced precipitation and soil water availability by quantifying seasonal changes in plant hydraulic and carbon traits for 11 tree species between control and drought treatments. Even with reduced soil volumetric water content in the upper 1 m of soil in the drought treatment, we found no significant difference between treatments for predawn and midday leaf water potential, photosynthesis, stomatal conductance, foliar stable carbon isotope composition, leaf mass per area, turgor loss point, xylem vessel anatomy, or leaf and stem nonstructural carbohydrates. While empirical measurements of aboveground traits revealed homeostatic maintenance of plant water status and traits in response to reduced soil moisture, modeled belowground dynamics revealed that trees in the drought treatment shifted the depth from which water was acquired to deeper soil layers. These findings reveal that belowground acclimation of tree water uptake depth may buffer tropical rainforests from more severe droughts that may arise in future with climate change.

Droughts in a warming climate have become more common and more extreme, making understanding forest responses to water stress increasingly pressing. Analysis of water stress in trees has long focused on water potential in xylem and leaves, which influences stomatal closure and water flow through the soil-plant-atmosphere continuum. At the same time, changes of vegetation water content (VWC) are linked to a range of tree responses, including fluxes of water and carbon, mortality, flammability, and more. Unlike water potential, which requires demanding in situ measurements, VWC can be retrieved from remote sensing measurements, particularly at microwave frequencies using radar and radiometry. Here, we highlight key frontiers through which VWC has the potential to significantly increase our understanding of forest responses to water stress. To validate remote sensing observations of VWC at landscape scale and to better relate them to data assimilation model parameters, we introduce an ecosystem-scale analog of the pressure-volume curve, the non-linear relationship between average leaf or branch water potential and water content commonly used in plant hydraulics. The sources of variability in these ecosystem-scale pressure-volume curves and their relationship to forest response to water stress are discussed. We further show to what extent diel, seasonal, and decadal dynamics of VWC reflect variations in different processes relating the tree response to water stress. VWC can also be used for inferring belowground conditions-which are difficult to impossible to observe directly. Lastly, we discuss how a dedicated geostationary spaceborne observational system for VWC, when combined with existing datasets, can capture diel and seasonal water dynamics to advance the science and applications of global forest vulnerability to future droughts.

Deep-water access is arguably the most effective, but under-studied, mechanism that plants employ to survive during drought. Vulnerability to embolism and hydraulic safety margins can predict mortality risk at given levels of dehydration, but deep-water access may delay plant dehydration. Here, we tested the role of deep-water access in enabling survival within a diverse tropical forest community in Panama using a novel data-model approach. We inversely estimated the effective rooting depth (ERD, as the average depth of water extraction), for 29 canopy species by linking diameter growth dynamics (1990-2015) to vapor pressure deficit, water potentials in the whole-soil column, and leaf hydraulic vulnerability curves. We validated ERD estimates against existing isotopic data of potential water-access depths. Across species, deeper ERD was associated with higher maximum stem hydraulic conductivity, greater vulnerability to xylem embolism, narrower safety margins, and lower mortality rates during extreme droughts over 35 years (1981-2015) among evergreen species. Species exposure to water stress declined with deeper ERD indicating that trees compensate for water stress-related mortality risk through deep-water access. The role of deep-water access in mitigating mortality of hydraulically-vulnerable trees has important implications for our predictive understanding of forest dynamics under current and future climates.

High temperatures alter the thermal sensitivities of numerous physiological and biochemical processes that impact tree growth and productivity. Foliar and root applications of methanol have been implicated in plant acclimation to high temperature via the C1 pathway. Here, we characterized temperature acclimation at 35 °C of leaf gas exchange, chlorophyll fluorescence, and extractable metabolites of potted Populus trichocarpa saplings and examined potential influences of mM concentrations of methanol added during soil watering over a two-month period. Relative to plants grown under the low growth temperature (LGT), high growth temperature (HGT) plants showed a suppression of leaf water use and carbon cycling including transpiration (E), net photosynthesis (Pn), an estimate of photorespiration (Rp), and dark respiration (Rd), attributed to reductions in stomatal conductance and direct negative effects on gas exchange and photosynthetic machinery. In contrast, HGT plants showed an upregulation of nonphotochemical quenching (NPQt), the optimum temperature for ETR, and leaf isoprene emissions at 40 °C. A large number of metabolites (867) were induced under HGT, many implicated in flavonoid biosynthesis highlighting a potentially protective role for these compounds. Methanol application did not significantly alter leaf gas exchange but slightly reduced the suppression of Rd and Rp by the high growth temperature while slightly impairing ETR, Fv′/Fm′, and qp. However, we were unable to determine if soil methanol was sufficiently taken up by the plant to have a direct effect on foliar processes. A small number of extracted leaf tissue metabolites (55 out of 10 015) showed significantly altered abundances under LGT and methanol treatments relative to water controls, and this increased in compound number (222) at the HGT. The results demonstrate the large physiological and biochemical impacts of high growth temperature on poplar seedlings and highlight the enhancement of the optimum temperature of ETR as a rapid thermal acclimation mechanism. Although no large effect on leaf physiology was observed, the results are consistent with methanol both impairing photochemistry of the light reactions via formaldehyde toxicity and stimulating photosynthesis and dark respiration through formate oxidation to CO2.

Climate change is expected to pose a global threat to forest health by intensifying extreme events like drought and insect attacks. Carbon allocation is a fundamental process that determines the adaptive responses of long-lived late-maturing organisms like trees to such stresses. However, our mechanistic understanding of how trees coordinate and set allocation priorities among different sinks (e.g., growth and storage) under severe source limitation remains limited. Using flux measurements, isotopic tracing, targeted metabolomics, and transcriptomics, we investigated how limitation of source supply influences sink activity, particularly growth and carbon storage, and their relative regulation in Norway spruce (Picea abies) clones. During photosynthetic deprivation, absolute rates of respiration, growth, and allocation to storage all decline. When trees approach neutral carbon balance, i.e., daytime net carbon gain equals nighttime carbon loss, genes encoding major enzymes of metabolic pathways remain relatively unaffected. However, under negative carbon balance, photosynthesis and growth are down-regulated while sucrose and starch biosynthesis pathways are up-regulated, indicating that trees prioritize carbon allocation to storage over growth. Moreover, trees under negative carbon balance actively increase the turnover rate of starch, lipids, and amino acids, most likely to support respiration and mitigate stress. Our study provides molecular evidence that trees faced with severe photosynthetic limitation strategically regulate storage allocation and consumption at the expense of growth. Understanding such allocation strategies is crucial for predicting how trees may respond to extreme events involving steep declines in photosynthesis, like severe drought, or defoliation by heat waves, late frost, or insect attack.

Water deficit in the atmosphere and soil are two key interactive factors that constrain transpiration and vegetation productivity. It is not clear which of these two factors is more important for the water and carbon flux response to drought stress in ecosystems. In this study, field data and numerical modeling were used to isolate their impact on evapotranspiration (ET) and gross primary productivity (GPP) at a tropical forest site in Barro Colorado Island (BCI), Panama, focusing on their response to the drought induced by the El Niño event of 2015–2016. Numerical simulations were performed using a plant hydrodynamic scheme (HYDRO) and a heuristic approach that ignores stomatal sensitivity to leaf water potential in the Energy Exascale Earth System Model (E3SM) Land Model (ELM). The sensitivity of canopy conductance (Gs) to vapor pressure deficit (VPD) obtained from eddy-covariance fluxes and measured sap flux shows that, at both ecosystem and plant scale, soil water stress is more important in limiting Gs than VPD at BCI during the El Niño event. The model simulations confirmed the importance of water stress limitation on Gs, but overestimated the VPD impact on Gs compared to that estimated from the observations. We also found that the predicted soil moisture is less sensitive to the diversity of plant hydraulic traits than ET and GPP. During the dry season at BCI, seasonal ET, especially soil evaporation at VPD > 0.42 kPa, simulated using HYDRO and ELM, were too strong and will require alternative parameterizations.

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.

Plant transpiration links physiological responses of vegetation to water supply and demand with hydrological, energy, and carbon budgets at the land-atmosphere interface. However, despite being the main land evaporative flux at the global scale, transpiration and its response to environmental drivers are currently not well constrained by observations. Here we introduce the first global compilation of whole-plant transpiration data from sap flow measurements (SAPFLUXNET, https://sapfluxnet.creaf.cat/, last access: 8 June 2021). We harmonized and quality-controlled individual datasets supplied by contributors worldwide in a semi-automatic data workflow implemented in the R programming language. Datasets include sub-daily time series of sap flow and hydrometeorological drivers for one or more growing seasons, as well as metadata on the stand characteristics, plant attributes, and technical details of the measurements. SAPFLUXNET contains 202 globally distributed datasets with sap flow time series for 2714 plants, mostly trees, of 174 species. SAPFLUXNET has a broad bioclimatic coverage, with woodland/shrubland and temperate forest biomes especially well represented (80% of the datasets). The measurements cover a wide variety of stand structural characteristics and plant sizes. The datasets encompass the period between 1995 and 2018, with 50% of the datasets being at least 3 years long. Accompanying radiation and vapour pressure deficit data are available for most of the datasets, while on-site soil water content is available for 56% of the datasets. Many datasets contain data for species that make up 90% or more of the total stand basal area, allowing the estimation of stand transpiration in diverse ecological settings. SAPFLUXNET adds to existing plant trait datasets, ecosystem flux networks, and remote sensing products to help increase our understanding of plant water use, plant responses to drought, and ecohydrological processes. SAPFLUXNET version 0.1.5 is freely available from the Zenodo repository (10.5281/zenodo.3971689; Poyatos et al., 2020a). The "sapfluxnetr"R package-designed to access, visualize, and process SAPFLUXNET data-is available from CRAN.

Leaf respiration in the dark (Rdark) and light (Rday) is poorly characterized in diverse tropical ecosystems, and little to no information exists on the degree of light suppression in common tree species within the Amazon basin, and their dependences upon plant functional traits and position within the canopy. We quantified Rdark and apparent Rday using the Kok method and measured key leaf traits in 26 tree individuals of different species distributed in three different canopy positions: canopy, lower canopy, and understory. To explore the relationships between the leaf traits we used the standardized major axis (SMA). We found that canopy trees had significantly higher rates of Rdark and Rday than trees in the understory. The difference between Rdark and Rday (the light suppression of respiration) was greatest in the understory (68 ± 9%, 95% CI) and lower canopy (49 ± 9%, 95% CI) when compared to the canopy (37 ± 10%, 95% CI). We also found that Rday was significantly and strongly correlated with Rdark (p < 0.001) for all the canopy positions. Also, leaf mass per area (LMA) and leaf Phosphorus concentration (P) had a significant relationship with Rdark (p < 0.001; p = 0.003), respectively. In addition, a significant relationship was found for LMA in the canopy and lower canopy positions (p = 0.009; p = 0.048) while P was only significant in the canopy (p = 0.044). Finally, no significant relationship was found between Rdark and nitrogen, sugars, and starch. Our results highlight the importance of including representation of the light suppression of leaf respiration in terrestrial biosphere models and also of accounting for vertical gradients within forest canopies and connections with functional traits.

Tropical forests absorb large amounts of atmospheric CO2 through photosynthesis, but high surface temperatures suppress this absorption while promoting isoprene emissions. While mechanistic isoprene emission models predict a tight coupling to photosynthetic electron transport (ETR) as a function of temperature, direct field observations of this phenomenon are lacking in the tropics and are necessary to assess the impact of a warming climate on global isoprene emissions. Here we demonstrate that in the early successional species Vismia guianensis in the central Amazon, ETR rates increased with temperature in concert with isoprene emissions, even as stomatal conductance (gs ) and net photosynthetic carbon fixation (Pn ) declined. We observed the highest temperatures of continually increasing isoprene emissions yet reported (50°C). While Pn showed an optimum value of 32.6 ± 0.4°C, isoprene emissions, ETR, and the oxidation state of PSII reaction centers (qL ) increased with leaf temperature with strong linear correlations for ETR (ƿ = 0.98) and qL (ƿ = 0.99) with leaf isoprene emissions. In contrast, other photoprotective mechanisms, such as non-photochemical quenching, were not activated at elevated temperatures. Inhibition of isoprenoid biosynthesis repressed Pn at high temperatures through a mechanism that was independent of stomatal closure. While extreme warming will decrease gs and Pn in tropical species, our observations support a thermal tolerance mechanism where the maintenance of high photosynthetic capacity under extreme warming is assisted by the simultaneous stimulation of ETR and metabolic pathways that consume the direct products of ETR including photorespiration and the biosynthesis of thermoprotective isoprenoids. Our results confirm that models which link isoprene emissions to the rate of ETR hold true in tropical species and provide necessary "ground-truthing" for simulations of the large predicted increases in tropical isoprene emissions with climate warming.

The 2015-2016 El Niño event ranks as one of the most severe on record in terms of the magnitude and extent of sea surface temperature (SST) anomalies generated in the tropical Pacific Ocean. Corresponding global impacts on the climate were expected to rival, or even surpass, those of the 1997-1998 severe El Niño event, which had SST anomalies that were similar in size. However, the 2015-2016 event failed to meet expectations for hydrologic change in many areas, including those expected to receive well above normal precipitation. To better understand how climate anomalies during an El Niño event impact soil moisture, we investigate changes in soil moisture in the humid tropics (between ±25ĝˆ ) during the three most recent super El Niño events of 1982-1983, 1997-1998 and 2015-2016, using data from the Global Land Data Assimilation System (GLDAS). First, we use in situ soil moisture observations obtained from 16 sites across five continents to validate and bias-correct estimates from GLDAS (r2Combining double low line0.54). Next, we apply a k-means cluster analysis to the soil moisture estimates during the El Niño mature phase, resulting in four groups of clustered data. The strongest and most consistent decreases in soil moisture occur in the Amazon basin and maritime southeastern Asia, while the most consistent increases occur over eastern Africa. In addition, we compare changes in soil moisture to both precipitation and evapotranspiration, which showed a lack of agreement in the direction of change between these variables and soil moisture most prominently in the southern Amazon basin, the Sahel and mainland southeastern Asia. Our results can be used to improve estimates of spatiotemporal differences in El Niño impacts on soil moisture in tropical hydrology and ecosystem models at multiple scales..

Transpiration in humid tropical forests modulates the global water cycle and is a key driver of climate regulation. Yet, our understanding of how tropical trees regulate sap flux in response to climate variability remains elusive. With a progressively warming climate, atmospheric evaporative demand [i.e., vapor pressure deficit (VPD)] will be increasingly important for plant functioning, becoming the major control of plant water use in the twenty-first century. Using measurements in 34 tree species at seven sites across a precipitation gradient in the neotropics, we determined how the maximum sap flux velocity (vmax) and the VPD threshold at which vmax is reached (VPDmax) vary with precipitation regime [mean annual precipitation (MAP); seasonal drought intensity (PDRY)] and two functional traits related to foliar and wood economics spectra [leaf mass per area (LMA); wood specific gravity (WSG)]. We show that, even though vmax is highly variable within sites, it follows a negative trend in response to increasing MAP and PDRY across sites. LMA and WSG exerted little effect on vmax and VPDmax, suggesting that these widely used functional traits provide limited explanatory power of dynamic plant responses to environmental variation within hyper-diverse forests. This study demonstrates that long-term precipitation plays an important role in the sap flux response of humid tropical forests to VPD. Our findings suggest that under higher evaporative demand, trees growing in wetter environments in humid tropical regions may be subjected to reduced water exchange with the atmosphere relative to trees growing in drier climates.

Nonstructural carbohydrates (NSCs) are essential for maintenance of plant metabolism and may be sensitive to short- and long-term climatic variation. NSC variation in moist tropical forests has rarely been studied, so regulation of NSCs in these systems is poorly understood. We measured foliar and branch NSC content in 23 tree species at three sites located across a large precipitation gradient in Panama during the 2015-2016 El Niño to examine how short- and long-term climatic variation impact carbohydrate dynamics. There was no significant difference in total NSCs as the drought progressed (leaf P = 0.32, branch P = 0.30) nor across the rainfall gradient (leaf P = 0.91, branch P = 0.96). Foliar soluble sugars decreased while starch increased over the duration of the dry period, suggesting greater partitioning of NSCs to storage than metabolism or transport as drought progressed. There was a large variation across species at all sites, but total foliar NSCs were positively correlated with leaf mass per area, whereas branch sugars were positively related to leaf temperature and negatively correlated with daily photosynthesis and wood density. The NSC homoeostasis across a wide range of conditions suggests that NSCs are an allocation priority in moist tropical forests.

Current climate change scenarios indicate warmer temperatures and the potential for more extreme droughts in the tropics, such that a mechanistic understanding of the water cycle from individual trees to landscapes is needed to adequately predict future changes in forest structure and function. In this study, we contrasted physiological responses of tropical trees during a normal dry season with the extreme dry season due to the 2015-2016 El Niño-Southern Oscillation (ENSO) event. We quantified high resolution temporal dynamics of sap velocity (Vs), stomatal conductance (gs) and leaf water potential (ΨL) of multiple canopy trees, and their correlations with leaf temperature (Tleaf) and environmental conditions [direct solar radiation, air temperature (Tair) and vapor pressure deficit (VPD)]. The experiment leveraged canopy access towers to measure adjacent trees at the ZF2 and Tapajós tropical forest research (near the cities of Manaus and Santarém). The temporal difference between the peak of gs (late morning) and the peak of VPD (early afternoon) is one of the major regulators of sap velocity hysteresis patterns. Sap velocity displayed species-specific diurnal hysteresis patterns reflected by changes in Tleaf. In the morning, Tleaf and sap velocity displayed a sigmoidal relationship. In the afternoon, stomatal conductance declined as Tleaf approached a daily peak, allowing ΨL to begin recovery, while sap velocity declined with an exponential relationship with Tleaf. In Manaus, hysteresis indices of the variables Tleaf-Tair and ΨL-Tleaf were calculated for different species and a significant difference (p < 0.01, α = 0.05) was observed when the 2015 dry season (ENSO period) was compared with the 2017 dry season ("control scenario"). In some days during the 2015 ENSO event, Tleaf approached 40°C for all studied species and the differences between Tleaf and Tair reached as high at 8°C (average difference: 1.65 ± 1.07°C). Generally, Tleaf was higher than Tair during the middle morning to early afternoon, and lower than Tair during the early morning, late afternoon and night. Our results support the hypothesis that partial stomatal closure allows for a recovery in ΨL during the afternoon period giving an observed counterclockwise hysteresis pattern between ΨL and Tleaf.

Vegetation plays an important role in regulating global carbon cycles and is a key component of the Earth system models (ESMs) that aim to project Earth's future climate. In the last decade, the vegetation component within ESMs has witnessed great progress from simple "big-leaf" approaches to demographically structured approaches, which have a better representation of plant size, canopy structure, and disturbances. These demographically structured vegetation models typically have a large number of input parameters, and sensitivity analysis is needed to quantify the impact of each parameter on the model outputs for a better understanding of model behavior. In this study, we conducted a comprehensive sensitivity analysis to diagnose the Community Land Model coupled to the Functionally Assembled Terrestrial Simulator, or CLM4.5(FATES). Specifically, we quantified the first- and second-order sensitivities of the model parameters to outputs that represent simulated growth and mortality as well as carbon fluxes and stocks for a tropical site with an extent of 1×1°. While the photosynthetic capacity parameter (Vc;max25) is found to be important for simulated carbon stocks and fluxes, we also show the importance of carbon storage and allometry parameters, which determine survival and growth strategies within the model. The parameter sensitivity changes with different sizes of trees and climate conditions. The results of this study highlight the importance of understanding the dynamics of the next generation of demographically enabled vegetation models within ESMs to improve model parameterization and structure for better model fidelity.

How plants respond physiologically to leaf warming and low water availability may determine how they will perform under future climate change. In 2015-2016, an unprecedented drought occurred across Amazonia with record-breaking high temperatures and low soil moisture, offering a unique opportunity to evaluate the performances of Amazonian trees to a severe climatic event. We quantified the responses of leaf water potential, sap velocity, whole-tree hydraulic conductance (Kwt), turgor loss and xylem embolism, during and after the 2015-2016 El Niño for five canopy-tree species. Leaf/xylem safety margins (SMs), sap velocity and Kwt showed a sharp drop during warm periods. SMs were negatively correlated with vapour pressure deficit, but had no significant relationship with soil water storage. Based on our calculations of canopy stomatal and xylem resistances, the decrease in sap velocity and Kwt was due to a combination of xylem cavitation and stomatal closure. Our results suggest that warm droughts greatly amplify the degree of trees' physiological stress and can lead to mortality. Given the extreme nature of the 2015-2016 El Niño and that temperatures are predicted to increase, this work can serve as a case study of the possible impact climate warming can have on tropical trees.This article is part of a discussion meeting issue 'The impact of the 2015/2016 El Niño on the terrestrial tropical carbon cycle: patterns, mechanisms and implications'.

Summary: Non‐structural carbohydrates (NSCs), as the labile fraction and dominant carbon currency, are essential mediators of plant adaptation to environments. However, whether and how NSC coordinates with plant economic strategy frameworks, particularly the well‐recognized leaf economics spectrums (LES) and root economics space (RES), remains unclear.We examined the relationships between NSC and key plant economics traits in leaves and fine roots across 90 alpine coniferous populations on the Tibetan Plateau, China.We observed contrasting coordination of NSC with economics traits in leaves and roots. Leaf total NSC and soluble sugar aligned with the leaf economic spectrum, conveying a trade‐off between growth and storage in leaves. However, NSC in roots was independent of the root economic spectrum, but highly coordinated with root foraging, with more starch and less sugar in forage‐efficient, thinner roots. Further, NSC‐trait coordination in leaves and roots was, respectively, driven by local temperature and precipitation.These findings highlight distinct roles of NSC in shaping the above‐ and belowground multidimensional economics trait space, and NSC‐based carbon economics provides a mechanistic understanding of how plants adapt to heterogeneous habitats and respond to environmental changes. [ABSTRACT FROM AUTHOR]

Survival rates of large trees determine forest biomass dynamics. Survival rates of small trees have been linked to mechanisms that maintain biodiversity across tropical forests. How species survival rates change with size offers insight into the links between biodiversity and ecosystem function across tropical forests. We tested patterns of size-dependent tree survival across the tropics using data from 1,781 species and over 2 million individuals to assess whether tropical forests can be characterized by size-dependent life-history survival strategies. We found that species were classifiable into four 'survival modes' that explain life-history variation that shapes carbon cycling and the relative abundance within forests. Frequently collected functional traits, such as wood density, leaf mass per area and seed mass, were not generally predictive of the survival modes of species. Mean annual temperature and cumulative water deficit predicted the proportion of biomass of survival modes, indicating important links between evolutionary strategies, climate and carbon cycling. The application of survival modes in demographic simulations predicted biomass change across forest sites. Our results reveal globally identifiable size-dependent survival strategies that differ across diverse systems in a consistent way. The abundance of survival modes and interaction with climate ultimately determine forest structure, carbon storage in biomass and future forest trajectories.

Tree mortality rates appear to be increasing in moist tropical forests (MTFs) with significant carbon cycle consequences. Here, we review the state of knowledge regarding MTF tree mortality, create a conceptual framework with testable hypotheses regarding the drivers, mechanisms and interactions that may underlie increasing MTF mortality rates, and identify the next steps for improved understanding and reduced prediction. Increasing mortality rates are associated with rising temperature and vapor pressure deficit, liana abundance, drought, wind events, fire and, possibly, CO2 fertilization-induced increases in stand thinning or acceleration of trees reaching larger, more vulnerable heights. The majority of these mortality drivers may kill trees in part through carbon starvation and hydraulic failure. The relative importance of each driver is unknown. High species diversity may buffer MTFs against large-scale mortality events, but recent and expected trends in mortality drivers give reason for concern regarding increasing mortality within MTFs. Models of tropical tree mortality are advancing the representation of hydraulics, carbon and demography, but require more empirical knowledge regarding the most common drivers and their subsequent mechanisms. We outline critical datasets and model developments required to test hypotheses regarding the underlying causes of increasing MTF mortality rates, and improve prediction of future mortality under climate change.

The fate of tropical forests under climate change is unclear as a result, in part, of the uncertainty in projected changes in precipitation and in the ability of vegetation models to capture the effects of drought-induced mortality on aboveground biomass (AGB). We evaluated the ability of a terrestrial biosphere model with demography and hydrodynamics (Ecosystem Demography, ED2-hydro) to simulate AGB and mortality of four tropical tree plant functional types (PFTs) that operate along light- and water-use axes. Model predictions were compared with observations of canopy trees at Barro Colorado Island (BCI), Panama. We then assessed the implications of eight hypothetical precipitation scenarios, including increased annual precipitation, reduced inter-annual variation, El Niño-related droughts and drier wet or dry seasons, on AGB and functional diversity of the model forest. When forced with observed meteorology, ED2-hydro predictions capture multiple BCI benchmarks. ED2-hydro predicts that AGB will be sustained under lower rainfall via shifts in the functional composition of the forest, except under the drier dry-season scenario. These results support the hypothesis that inter-annual variation in mean and seasonal precipitation promotes the coexistence of functionally diverse PFTs because of the relative differences in mortality rates. If the hydroclimate becomes chronically drier or wetter, functional evenness related to drought tolerance may decline.

Evapotranspiration (ET) is a pivotal component in catchment-scale water balance and is essential for informed watershed management. Nevertheless, uncertainties in ET observation or modeling have been hindering effective water resources management. This study addresses this gap by establishing a robust, generalized linear relationship between ET and gross primary productivity (GPP) at the catchment scale. We test the linearity of the relationships between monthly GPP and ET data at 380 near-natural catchments across various climatic and landscape conditions in the contiguous US, yielding Pearson's r≥0.6 for 97% of the 380 catchments. We then develop a regionalization strategy to parameterize this GPP-ET relationship at the catchment scale by identifying and using the linkages between the parameter values and extensively available hydroclimatic and landscape data. We demonstrate the efficacy of the proposed GPP-ET relationship and parameter regionalization strategy by their combined predictive capacity, where the predicted monthly GPP matches well with remote-sensing–based GPP product, achieving Kling-Gupta efficiency (KGE) values ≥0.5 for 92% of the catchments. In addition, we verify the relationship and its parameter regionalization at 35 AmeriFlux sites with KGE ≥0.5 for 25 sites, suggesting that the new relationship is transferable across the site, catchment, and regional scales. Our findings are valuable for improving remote-sensing–based estimation of monthly ET and diagnosing coupled water-carbon simulations in land surface and Earth system models. [ABSTRACT FROM AUTHOR]

Leaf thermoregulation has been documented in a handful of studies, but the generality and origins of this pattern are unclear. We suggest that leaf thermoregulation is widespread in both space and time, and originates from the optimization of leaf traits to maximize leaf carbon gain across and within variable environments. Here we use global data for leaf temperatures, traits and photosynthesis to evaluate predictions from a novel theory of thermoregulation that synthesizes energy budget and carbon economics theories. Our results reveal that variation in leaf temperatures and physiological performance are tightly linked to leaf traits and carbon economics. The theory, parameterized with global averaged leaf traits and microclimate, predicts a moderate level of leaf thermoregulation across a broad air temperature gradient. These predictions are supported by independent data for diverse taxa spanning a global air temperature range of ∼60 °C. Moreover, our theory predicts that net carbon assimilation can be maximized by means of a trade-off between leaf thermal stability and photosynthetic stability. This prediction is supported by globally distributed data for leaf thermal and photosynthetic traits. Our results demonstrate that the temperatures of plant tissues, and not just air, are vital to developing more accurate Earth system models.

Terrestrial disturbances are accelerating globally, but their full impact is not quantified because we lack an adequate monitoring system. Remote sensing offers a means to quantify the frequency and extent of disturbances globally. Here, we review the current application of remote sensing to this problem and offer a framework for more systematic analysis in the future. We recommend that any proposed monitoring system should not only detect disturbances, but also be able to: identify the proximate cause(s); integrate a range of spatial scales; and, ideally, incorporate process models to explain the observed patterns and predicted trends in the future. Significant remaining challenges are tied to the ecology of disturbances. To meet these challenges, more effort is required to incorporate ecological principles and understanding into the assessments of disturbance worldwide.

We related measurements of annual burned area in the southwest United States during 1984–2013 to records of climate variability. Within forests, annual burned area correlated at least as strongly with spring–summer vapour pressure deficit (VPD) as with 14 other drought-related metrics, including more complex metrics that explicitly represent fuel moisture. Particularly strong correlations with VPD arise partly because this term dictates the atmospheric moisture demand. Additionally, VPD responds to moisture supply, which is difficult to measure and model regionally due to complex micrometeorology, land cover and terrain. Thus, VPD appears to be a simple and holistic indicator of regional water balance. Coupled with the well-known positive influence of prior-year cold season precipitation on fuel availability and connectivity, VPD may be utilised for burned area forecasts and also to infer future trends, though these are subject to other complicating factors such as land cover change and management. Assuming an aggressive greenhouse gas emissions scenario, climate models predict mean spring–summer VPD will exceed the highest recorded values in the southwest in nearly 40% of years by the middle of this century. These results forewarn of continued increases in burned forest area in the southwest United States, and likely elsewhere, when fuels are not limiting.

AbstractIn 2011, exceptionally low atmospheric moisture content combined with moderately high temperatures to produce a record-high vapor pressure deficit (VPD) in the southwestern United States (SW). These conditions combined with record-low cold-season precipitation to cause widespread drought and extreme wildfires. Although interannual VPD variability is generally dominated by temperature, high VPD in 2011 was also driven by a lack of atmospheric moisture. The May–July 2011 dewpoint in the SW was 4.5 standard deviations below the long-term mean. Lack of atmospheric moisture was promoted by already very dry soils and amplified by a strong ocean-to-continent sea level pressure gradient and upper-level convergence that drove dry northerly winds and subsidence upwind of and over the SW. Subsidence drove divergence of rapid and dry surface winds over the SW, suppressing southerly moisture imports and removing moisture from already dry soils. Model projections developed for the fifth phase of the Coupled Model Intercomparison Project (CMIP5) suggest that by the 2050s warming trends will cause mean warm-season VPD to be comparable to the record-high VPD observed in 2011. CMIP5 projections also suggest increased interannual variability of VPD, independent of trends in background mean levels, as a result of increased variability of dewpoint, temperature, vapor pressure, and saturation vapor pressure. Increased variability in VPD translates to increased probability of 2011-type VPD anomalies, which would be superimposed on ever-greater background VPD levels. Although temperature will continue to be the primary driver of interannual VPD variability, 2011 served as an important reminder that atmospheric moisture content can also drive impactful VPD anomalies.