Community assembly influences plant trait economic spectra and functional trade-offs at ecosystem scales.

Plant functional traits hold the potential to greatly improve the understanding and prediction of climate impacts on ecosystems and carbon cycle feedback to climate change. Traits are commonly used to place species along a global conservative-acquisitive trade-off, yet how and if functional traits and conservative-acquisitive trade-offs scale up to mediate community and ecosystem fluxes is largely unknown. Here, we combine functional trait datasets and multibiome datasets of forest water and carbon fluxes at the species, community, and ecosystem-levels to quantify the scaling of the tradeoff between maximum flux and sensitivity to vapor pressure deficit. We find a strong conservative-acquisitive trade-off at the species scale, which weakens modestly at the community scale and largely disappears at the ecosystem scale. Functional traits, particularly plant water transport (hydraulic) traits, are strongly associated with the key dimensions of the conservative-acquisitive trade-off at community and ecosystem scales, highlighting that trait composition appears to influence community and ecosystem flux dynamics. Our findings provide a foundation for improving carbon cycle models by revealing i) that plant hydraulic traits are most strongly associated with community- and ecosystem scale flux dynamics and ii) community assembly dynamics likely need to be considered explicitly, as they give rise to ecosystem-level flux dynamics that differ substantially from trade-offs identified at the species-level.

A new empirical framework to quantify the hydraulic effects of soil and atmospheric drivers on plant water status.

Metrics to quantify regulation of plant water status at the daily as opposed to the seasonal scale do not presently exist. This gap is significant since plants are hypothesised to regulate their water potential not only with respect to slowly changing soil drought but also with respect to faster changes in air vapour pressure deficit (VPD), a variable whose importance for plant physiology is expected to grow because of higher temperatures in the coming decades. We present a metric, the stringency of water potential regulation, that can be employed at the daily scale and quantifies the effects exerted on plants by the separate and combined effect of soil and atmospheric drought. We test our theory using datasets from two experiments where air temperature and VPD were experimentally manipulated. In contrast to existing metrics based on soil drought that can only be applied at the seasonal scale, our metric successfully detects the impact of atmospheric warming on the regulation of plant water status. We show that the thermodynamic effect of VPD on plant water status can be isolated and compared against that exerted by soil drought and the covariation between VPD and soil drought. Furthermore, in three of three cases, VPD accounted for more than 5 MPa of potential effect on leaf water potential. We explore the significance of our findings in the context of potential future applications of this metric from plant to ecosystem scale.

The role of height-driven constraints and compensations on tree vulnerability to drought.

Frequent observations of higher mortality in larger trees than in smaller ones during droughts have sparked an increasing interest in size-dependent drought-induced mortality. However, the underlying physiological mechanisms are not well understood, with height-associated hydraulic constraints often being implied as the potential mechanism driving increased drought vulnerability. We performed a quantitative synthesis on how key traits that drive plant water and carbon economy change with tree height within species and assessed the implications that the different constraints and compensations may have on the interacting mechanisms (hydraulic failure, carbon starvation and/or biotic-agent attacks) affecting tree vulnerability to drought. While xylem tension increases with tree height, taller trees present a range of structural and functional adjustments, including more efficient water use and transport and greater water uptake and storage capacity, that mitigate the path-length-associated drop in water potential. These adaptations allow taller trees to withstand episodic water stress. Conclusive evidence for height-dependent increased vulnerability to hydraulic failure and carbon starvation, and their coupling to defence mechanisms and pest and pathogen dynamics, is still lacking. Further research is needed, particularly at the intraspecific level, to ascertain the specific conditions and thresholds above which height hinders tree survival under drought.

Daytime stomatal regulation in mature temperate trees prioritizes stem rehydration at night.

Trees remain sufficiently hydrated during drought by closing stomata and reducing canopy conductance (Gc ) in response to variations in atmospheric water demand and soil water availability. Thresholds that control the reduction of Gc are proposed to optimize hydraulic safety against carbon assimilation efficiency. However, the link between Gc and the ability of stem tissues to rehydrate at night remains unclear. We investigated whether species-specific Gc responses aim to prevent branch embolisms, or enable night-time stem rehydration, which is critical for turgor-dependent growth. For this, we used a unique combination of concurrent dendrometer, sap flow and leaf water potential measurements and collected branch-vulnerability curves of six common European tree species. Species-specific Gc reduction was weakly related to the water potentials at which 50% of branch xylem conductivity is lost (P50 ). Instead, we found a stronger relationship with stem rehydration. Species with a stronger Gc control were less effective at refilling stem-water storage as the soil dries, which appeared related to their xylem architecture. Our findings highlight the importance of stem rehydration for water-use regulation in mature trees, which likely relates to the maintenance of adequate stem turgor. We thus conclude that stem rehydration must complement the widely accepted safety-efficiency stomatal control paradigm.

Simple Models Outperform More Complex Big-Leaf Models of Daily Transpiration in Forested Biomes.

Transpiration makes up the bulk of total evaporation in forested environments yet remains challenging to predict at landscape-to-global scales. We harnessed independent estimates of daily transpiration derived from co-located sap flow and eddy-covariance measurement systems and applied the triple collocation technique to evaluate predictions from big leaf models requiring no calibration. In total, four models in 608 unique configurations were evaluated at 21 forested sites spanning a wide diversity of biophysical attributes and environmental backgrounds. We found that simpler models that neither explicitly represented aerodynamic forcing nor canopy conductance achieved higher accuracy and signal-to-noise levels when optimally configured (rRMSE = 20%; R 2 = 0.89). Irrespective of model type, optimal configurations were those making use of key plant functional type dependent parameters, daily LAI, and constraints based on atmospheric moisture demand over soil moisture supply. Our findings have implications for more informed water resource management based on hydrological modeling and remote sensing.

Climate and functional traits jointly mediate tree water-use strategies.

Tree water use is central to plant function and ecosystem fluxes. However, it is still unknown how organ-level water-relations traits are coordinated to determine whole-tree water-use strategies in response to drought, and whether this coordination depends on climate. Here we used a global sap flow database (SAPFLUXNET) to study the response of water use, in terms of whole-tree canopy conductance (G), to vapour pressure deficit (VPD) and to soil water content (SWC) for 142 tree species. We investigated the individual and coordinated effect of six water-relations traits (vulnerability to embolism, Huber value, hydraulic conductivity, turgor-loss point, rooting depth and leaf size) on water-use parameters, also accounting for the effect of tree height and climate (mean annual precipitation, MAP). Reference G and its sensitivity to VPD were tightly coordinated with water-relations traits rather than with MAP. Species with efficient xylem transport had higher canopy conductance but also higher sensitivity to VPD. Moreover, we found that angiosperms had higher reference G and higher sensitivity to VPD than did gymnosperms. Our results highlight the need to consider trait integration and reveal the complications and challenges of defining a single, whole-plant resource use spectrum ranging from 'acquisitive' to 'conservative'.

Towards a statistically robust determination of minimum water potential and hydraulic risk in plants.

Minimum water potential (Ψmin ) is a key variable for characterizing dehydration tolerance and hydraulic safety margins (HSMs) in plants. Ψmin is usually estimated as the absolute minimum tissue Ψ experienced by a species, but this is problematic because sample extremes are affected by sample size and the underlying probability distribution. We compare alternative approaches to estimate Ψmin and assess the corresponding uncertainties and biases; propose statistically robust estimation methods based on extreme value theory (EVT); and assess the implications of our results for the characterization of hydraulic risk. Our results show that current estimates of Ψmin and HSMs are biased, as they are strongly affected by sample size. Because sampling effort is generally higher for species living in dry environments, the differences in current Ψmin estimates between these species and those living under milder conditions are partly artefactual. When this bias is corrected using EVT methods, resulting HSMs tend to increase substantially with resistance to embolism across species. Although data availability and representativeness remain the main challenges for proper determination of Ψmin , a closer look at Ψ distributions and the use of statistically robust methods to estimate Ψmin opens new ground for characterizing plant hydraulic risks.

Spatiotemporal variability and origin of CO2 and CH4 tree stem fluxes in an upland forest.

The exchange of multiple greenhouse gases (i.e., CO2 and CH4 ) between tree stems and the atmosphere represents a knowledge gap in the global carbon cycle. Stem CO2 and CH4 fluxes vary across time and space and are unclear, which are their individual or shared drivers. Here we measured CO2 and CH4 fluxes at different stem heights combining manual (biweekly; n = 678) and automated (hourly; n > 38,000) measurements in a temperate upland forest. All trees showed CO2 and CH4 emissions despite 20% of measurements showing net CH4 uptake. Stem CO2 fluxes presented clear seasonal trends from manual and automated measurements. Only automated measurements captured the high temporal variability of stem CH4 fluxes revealing clear seasonal trends. Despite that temporal integration, the limited number of automated chambers made stand-level mean CH4 fluxes sensitive to "hot spots," resulting in mean fluxes with high uncertainty. Manual measurements provided better integration of spatial variability, but their lack of temporal variability integration hindered the detection of temporal trends and stand-level mean fluxes. These results highlight the potential bias of previous studies of stem CH4 fluxes solely based on manual or automated measurements. Stem height, temperature, and soil moisture only explained 7% and 11% of the stem CH4 flux variability compared to 42% and 81% for CO2 (manual and automated measurements, respectively). This large unexplained variability, in combination with high CH4 concentrations in the trees' heartwood, suggests that stem CH4 fluxes might be more influenced by gas transport and diffusivity through the wood than by drivers of respiratory CO2 flux, which has crucial implications for developing process-based ecosystem models. We postulate that CH4 is likely originated within tree stems because of lack of a consistent vertical pattern in CH4 fluxes, evidence of CH4 production in wood incubations, and low CH4 concentration in the soil profile but high concentrations within the trees' heartwood.

Statistical upscaling of ecosystem CO2 fluxes across the terrestrial tundra and boreal domain: Regional patterns and uncertainties.

The regional variability in tundra and boreal carbon dioxide (CO2 ) fluxes can be high, complicating efforts to quantify sink-source patterns across the entire region. Statistical models are increasingly used to predict (i.e., upscale) CO2 fluxes across large spatial domains, but the reliability of different modeling techniques, each with different specifications and assumptions, has not been assessed in detail. Here, we compile eddy covariance and chamber measurements of annual and growing season CO2 fluxes of gross primary productivity (GPP), ecosystem respiration (ER), and net ecosystem exchange (NEE) during 1990-2015 from 148 terrestrial high-latitude (i.e., tundra and boreal) sites to analyze the spatial patterns and drivers of CO2 fluxes and test the accuracy and uncertainty of different statistical models. CO2 fluxes were upscaled at relatively high spatial resolution (1 km2 ) across the high-latitude region using five commonly used statistical models and their ensemble, that is, the median of all five models, using climatic, vegetation, and soil predictors. We found the performance of machine learning and ensemble predictions to outperform traditional regression methods. We also found the predictive performance of NEE-focused models to be low, relative to models predicting GPP and ER. Our data compilation and ensemble predictions showed that CO2 sink strength was larger in the boreal biome (observed and predicted average annual NEE -46 and -29 g C m-2  yr-1 , respectively) compared to tundra (average annual NEE +10 and -2 g C m-2  yr-1 ). This pattern was associated with large spatial variability, reflecting local heterogeneity in soil organic carbon stocks, climate, and vegetation productivity. The terrestrial ecosystem CO2 budget, estimated using the annual NEE ensemble prediction, suggests the high-latitude region was on average an annual CO2 sink during 1990-2015, although uncertainty remains high.

Ecosystem transpiration and evaporation: Insights from three water flux partitioning methods across FLUXNET sites.

We apply and compare three widely applicable methods for estimating ecosystem transpiration (T) from eddy covariance (EC) data across 251 FLUXNET sites globally. All three methods are based on the coupled water and carbon relationship, but they differ in assumptions and parameterizations. Intercomparison of the three daily T estimates shows high correlation among methods (R between .89 and .94), but a spread in magnitudes of T/ET (evapotranspiration) from 45% to 77%. When compared at six sites with concurrent EC and sap flow measurements, all three EC-based T estimates show higher correlation to sap flow-based T than EC-based ET. The partitioning methods show expected tendencies of T/ET increasing with dryness (vapor pressure deficit and days since rain) and with leaf area index (LAI). Analysis of 140 sites with high-quality estimates for at least two continuous years shows that T/ET variability was 1.6 times higher across sites than across years. Spatial variability of T/ET was primarily driven by vegetation and soil characteristics (e.g., crop or grass designation, minimum annual LAI, soil coarse fragment volume) rather than climatic variables such as mean/standard deviation of temperature or precipitation. Overall, T and T/ET patterns are plausible and qualitatively consistent among the different water flux partitioning methods implying a significant advance made for estimating and understanding T globally, while the magnitudes remain uncertain. Our results represent the first extensive EC data-based estimates of ecosystem T permitting a data-driven perspective on the role of plants' water use for global water and carbon cycling in a changing climate.

Quantification of uncertainties in conifer sap flow measured with the thermal dissipation method.

Trees play a key role in the global hydrological cycle and measurements performed with the thermal dissipation method (TDM) have been crucial in providing whole-tree water-use estimates. Yet, different data processing to calculate whole-tree water use encapsulates uncertainties that have not been systematically assessed. We quantified uncertainties in conifer sap flux density (Fd ) and stand water use caused by commonly applied methods for deriving zero-flow conditions, dampening and sensor calibration. Their contribution has been assessed using a stem segment calibration experiment and 4 yr of TDM measurements in Picea abies and Larix decidua growing in contrasting environments. Uncertainties were then projected on TDM data from different conifers across the northern hemisphere. Commonly applied methods mostly underestimated absolute Fd . Lacking a site- and species-specific calibrations reduced our stand water-use measurements by 37% and induced uncertainty in northern hemisphere Fd . Additionally, although the interdaily variability was maintained, disregarding dampening and/or applying zero-flow conditions that ignored night-time water use reduced the correlation between environment and Fd . The presented ensemble of calibration curves and proposed dampening correction, together with the systematic quantification of data-processing uncertainties, provide crucial steps in improving whole-tree water-use estimates across spatial and temporal scales.

Stand dynamics modulate water cycling and mortality risk in droughted tropical forest.

Transpiration from the Amazon rainforest generates an essential water source at a global and local scale. However, changes in rainforest function with climate change can disrupt this process, causing significant reductions in precipitation across Amazonia, and potentially at a global scale. We report the only study of forest transpiration following a long-term (>10 year) experimental drought treatment in Amazonian forest. After 15 years of receiving half the normal rainfall, drought-related tree mortality caused total forest transpiration to decrease by 30%. However, the surviving droughted trees maintained or increased transpiration because of reduced competition for water and increased light availability, which is consistent with increased growth rates. Consequently, the amount of water supplied as rainfall reaching the soil and directly recycled as transpiration increased to 100%. This value was 25% greater than for adjacent nondroughted forest. If these drought conditions were accompanied by a modest increase in temperature (e.g., 1.5°C), water demand would exceed supply, making the forest more prone to increased tree mortality.

Strong resilience of soil respiration components to drought-induced die-off resulting in forest secondary succession.

How forests cope with drought-induced perturbations and how the dependence of soil respiration on environmental and biological drivers is affected in a warming and drying context are becoming key questions. The aims of this study were to determine whether drought-induced die-off and forest succession were reflected in soil respiration and its components and to determine the influence of climate on the soil respiration components. We used the mesh exclusion method to study seasonal variations in soil respiration (R S) and its components: heterotrophic (R H) and autotrophic (R A) [further split into fine root (R R) and mycorrhizal respiration (R M)] in a mixed Mediterranean forest where Scots pine (Pinus sylvestris L.) is undergoing a drought-induced die-off and is being replaced by holm oak (Quercus ilex L.). Drought-induced pine die-off was not reflected in R S nor in its components, which denotes a high functional resilience of the plant and soil system to pine die-off. However, the succession from Scots pine to holm oak resulted in a reduction of R H and thus in an important decrease of total respiration (R S was 36 % lower in holm oaks than in non-defoliated pines). Furthermore, R S and all its components were strongly regulated by soil water content-and-temperature interaction. Since Scots pine die-off and Quercus species colonization seems to be widely occurring at the driest limit of the Scots pine distribution, the functional resilience of the soil system over die-off and the decrease of R S from Scots pine to holm oak could have direct consequences for the C balance of these ecosystems.

Balancing the risks of hydraulic failure and carbon starvation: a twig scale analysis in declining Scots pine.

Understanding physiological processes involved in drought-induced mortality is important for predicting the future of forests and for modelling the carbon and water cycles. Recent research has highlighted the variable risks of carbon starvation and hydraulic failure in drought-exposed trees. However, little is known about the specific responses of leaves and supporting twigs, despite their critical role in balancing carbon acquisition and water loss. Comparing healthy (non-defoliated) and unhealthy (defoliated) Scots pine at the same site, we measured the physiological variables involved in regulating carbon and water resources. Defoliated trees showed different responses to summer drought compared with non-defoliated trees. Defoliated trees maintained gas exchange while non-defoliated trees reduced photosynthesis and transpiration during the drought period. At the branch scale, very few differences were observed in non-structural carbohydrate concentrations between health classes. However, defoliated trees tended to have lower water potentials and smaller hydraulic safety margins. While non-defoliated trees showed a typical response to drought for an isohydric species, the physiology appears to be driven in defoliated trees by the need to maintain carbon resources in twigs. These responses put defoliated trees at higher risk of branch hydraulic failure and help explain the interaction between carbon starvation and hydraulic failure in dying trees.

A new look at water transport regulation in plants.

Plant function requires effective mechanisms to regulate water transport at a variety of scales. Here, we develop a new theoretical framework describing plant responses to drying soil, based on the relationship between midday and predawn leaf water potentials. The intercept of the relationship (Λ) characterizes the maximum transpiration rate per unit of hydraulic transport capacity, whereas the slope (σ) measures the relative sensitivity of the transpiration rate and plant hydraulic conductance to declining water availability. This framework was applied to a newly compiled global database of leaf water potentials to estimate the values of Λ and σ for 102 plant species. Our results show that our characterization of drought responses is largely consistent within species, and that the parameters Λ and σ show meaningful associations with climate across species. Parameter σ was ≤1 in most species, indicating a tight coordination between the gas and liquid phases of water transport, in which canopy transpiration tended to decline faster than hydraulic conductance during drought, thus reducing the pressure drop through the plant. The quantitative framework presented here offers a new way of characterizing water transport regulation in plants that can be used to assess their vulnerability to drought under current and future climatic conditions.

Urinary phthalate/DINCH metabolites associations with kisspeptin and reproductive hormones in teenagers: A cross-sectional study from the HBM4EU aligned studies.

BACKGROUND: Exposure to phthalate/DINCH metabolites can induce human reproductive toxicity, however, their endocrine-disrupting mechanisms are not fully elucidated. OBJECTIVE: To investigate the association between concentrations of phthalate/DINCH metabolites, serum kisspeptin, and reproductive hormones among European teenagers from three of the HBM4EU Aligned Studies. METHODS: In 733 Belgian (FLEHS IV study), Slovak (PCB cohort follow-up), and Spanish (BEA study) teenagers, ten phthalate and two DINCH metabolites were measured in urine by high-performance liquid chromatography-tandem mass spectrometry. Serum kisspeptin (kiss54) protein, follicle-stimulating hormone (FSH), total testosterone (TT), estradiol (E2), and sex hormone-binding globulin (SHBG) levels were measured by immunosorbent assays. Free Androgen Index (FAI) was calculated as a proxy of free testosterone. Adjusted sex-stratified linear regression models for individual studies, mixed effect models (LME) accounting for random effects for pooled studies, and g-computation and Bayesian kernel machine regression (BKMR) models for the phthalate/DINCH mixture were performed. RESULTS: The LME suggested that each IQR increase in ln-transformed levels of several phthalates was associated with lower kisspeptin [MnBP: %change (95%CI): -2.8 (-4.2;-0.4); MEHP: -1.4 (-3.4,0.2)] and higher FSH [∑DINP: 11.8 (-0.6;25.1)] levels in females from pooled studies. G-computation showed that the phthalates/DINCH mixture was associated with lower kisspeptin [-4.28 (-8.07;-0.34)] and higher FSH [22.13 (0.5;48.4)] also in females; BKMR showed similar although non-significant pattern. In males, higher phthalates metabolites [MEHP: -12.22 (-21.09;-1.18); oxo-MEHP: -12.73 (-22.34;-1.93)] were associated with lower TT and FAI, although higher DINCH [OH-MINCH: 16.31 (6.23;27.35), cx-MINCH: 16.80 (7.03;27.46), ∑DINCH: 17.37 (7.26;29.74)] were associated with higher TT levels. No mixture associations were found in males. CONCLUSION: We observed sex-specific associations between urinary concentrations of phthalate/DINCH metabolites and the panel of selected effect biomarkers (kisspeptin and reproductive hormones). This suggests that exposure to phthalates would be associated with changes in kisspeptin levels, which would affect the HPG axis and thus influence reproductive health. However, further research is needed, particularly for phthalate replacements such as DINCH.

Drought-induced defoliation and long periods of near-zero gas exchange play a key role in accentuating metabolic decline of Scots pine.

Drought-induced defoliation has recently been associated with the depletion of carbon reserves and increased mortality risk in Scots pine (Pinus sylvestris). We hypothesize that defoliated individuals are more sensitive to drought, implying that potentially higher gas exchange (per unit of leaf area) during wet periods may not compensate for their reduced photosynthetic area. We measured sap flow, needle water potentials and whole-tree hydraulic conductance to analyse the drought responses of co-occurring defoliated and nondefoliated Scots pines in northeast Spain during typical (2010) and extreme (2011) drought conditions. Defoliated Scots pines showed higher sap flow per unit leaf area during spring, but were more sensitive to summer drought, relative to nondefoliated pines. This pattern was associated with a steeper decline in soil-to-leaf hydraulic conductance with drought and an enhanced sensitivity of canopy conductance to soil water availability. Near-homeostasis in midday water potentials was observed across years and defoliation classes, with minimum values of -2.5 MPa. Enhanced sensitivity to drought and prolonged periods of near-zero gas exchange were consistent with low levels of carbohydrate reserves in defoliated trees. Our results support the critical links between defoliation, water and carbon availability, and their key roles in determining tree survival and recovery under drought.

Networking the forest infrastructure towards near real-time monitoring - A white paper.

Forests account for nearly 90 % of the world's terrestrial biomass in the form of carbon and they support 80 % of the global biodiversity. To understand the underlying forest dynamics, we need a long-term but also relatively high-frequency, networked monitoring system, as traditionally used in meteorology or hydrology. While there are numerous existing forest monitoring sites, particularly in temperate regions, the resulting data streams are rarely connected and do not provide information promptly, which hampers real-time assessments of forest responses to extreme climate events. The technology to build a better global forest monitoring network now exists. This white paper addresses the key structural components needed to achieve a novel meta-network. We propose to complement - rather than replace or unify - the existing heterogeneous infrastructure with standardized, quality-assured linking methods and interacting data processing centers to create an integrated forest monitoring network. These automated (research topic-dependent) linking methods in atmosphere, biosphere, and pedosphere play a key role in scaling site-specific results and processing them in a timely manner. To ensure broad participation from existing monitoring sites and to establish new sites, these linking methods must be as informative, reliable, affordable, and maintainable as possible, and should be supplemented by near real-time remote sensing data. The proposed novel meta-network will enable the detection of emergent patterns that would not be visible from isolated analyses of individual sites. In addition, the near real-time availability of data will facilitate predictions of current forest conditions (nowcasts), which are urgently needed for research and decision making in the face of rapid climate change. We call for international and interdisciplinary efforts in this direction.

A deep learning-based hybrid model of global terrestrial evaporation.

Terrestrial evaporation (E) is a key climatic variable that is controlled by a plethora of environmental factors. The constraints that modulate the evaporation from plant leaves (or transpiration, Et) are particularly complex, yet are often assumed to interact linearly in global models due to our limited knowledge based on local studies. Here, we train deep learning algorithms using eddy covariance and sap flow data together with satellite observations, aiming to model transpiration stress (St), i.e., the reduction of Et from its theoretical maximum. Then, we embed the new St formulation within a process-based model of E to yield a global hybrid E model. In this hybrid model, the St formulation is bidirectionally coupled to the host model at daily timescales. Comparisons against in situ data and satellite-based proxies demonstrate an enhanced ability to estimate St and E globally. The proposed framework may be extended to improve the estimation of E in Earth System Models and enhance our understanding of this crucial climatic variable.