Widespread and complex drought effects on vegetation physiology inferred from space.

The response of vegetation physiology to drought at large spatial scales is poorly understood due to a lack of direct observations. Here, we study vegetation drought responses related to photosynthesis, evaporation, and vegetation water content using remotely sensed data, and we isolate physiological responses using a machine learning technique. We find that vegetation functional decreases are largely driven by the downregulation of vegetation physiology such as stomatal conductance and light use efficiency, with the strongest downregulation in water-limited regions. Vegetation physiological decreases in wet regions also result in a discrepancy between functional and structural changes under severe drought. We find similar patterns of physiological drought response using simulations from a soil-plant-atmosphere continuum model coupled with a radiative transfer model. Observation-derived vegetation physiological responses to drought across space are mainly controlled by aridity and additionally modulated by abnormal hydro-meteorological conditions and vegetation types. Hence, isolating and quantifying vegetation physiological responses to drought enables a better understanding of ecosystem biogeochemical and biophysical feedback in modulating climate change.

Leaf-level coordination principles propagate to the ecosystem scale.

Fundamental axes of variation in plant traits result from trade-offs between costs and benefits of resource-use strategies at the leaf scale. However, it is unclear whether similar trade-offs propagate to the ecosystem level. Here, we test whether trait correlation patterns predicted by three well-known leaf- and plant-level coordination theories - the leaf economics spectrum, the global spectrum of plant form and function, and the least-cost hypothesis - are also observed between community mean traits and ecosystem processes. We combined ecosystem functional properties from FLUXNET sites, vegetation properties, and community mean plant traits into three corresponding principal component analyses. We find that the leaf economics spectrum (90 sites), the global spectrum of plant form and function (89 sites), and the least-cost hypothesis (82 sites) all propagate at the ecosystem level. However, we also find evidence of additional scale-emergent properties. Evaluating the coordination of ecosystem functional properties may aid the development of more realistic global dynamic vegetation models with critical empirical data, reducing the uncertainty of climate change projections.

Microbial carbon use efficiency promotes global soil carbon storage.

Soils store more carbon than other terrestrial ecosystems1,2. How soil organic carbon (SOC) forms and persists remains uncertain1,3, which makes it challenging to understand how it will respond to climatic change3,4. It has been suggested that soil microorganisms play an important role in SOC formation, preservation and loss5-7. Although microorganisms affect the accumulation and loss of soil organic matter through many pathways4,6,8-11, microbial carbon use efficiency (CUE) is an integrative metric that can capture the balance of these processes12,13. Although CUE has the potential to act as a predictor of variation in SOC storage, the role of CUE in SOC persistence remains unresolved7,14,15. Here we examine the relationship between CUE and the preservation of SOC, and interactions with climate, vegetation and edaphic properties, using a combination of global-scale datasets, a microbial-process explicit model, data assimilation, deep learning and meta-analysis. We find that CUE is at least four times as important as other evaluated factors, such as carbon input, decomposition or vertical transport, in determining SOC storage and its spatial variation across the globe. In addition, CUE shows a positive correlation with SOC content. Our findings point to microbial CUE as a major determinant of global SOC storage. Understanding the microbial processes underlying CUE and their environmental dependence may help the prediction of SOC feedback to a changing climate.

Emergence of the physiological effects of elevated CO2 on land-atmosphere exchange of carbon and water.

Elevated atmospheric CO2 (eCO2 ) influences the carbon assimilation rate and stomatal conductance of plants, thereby affecting the global cycles of carbon and water. Yet, the detection of these physiological effects of eCO2 in observational data remains challenging, because natural variations and confounding factors (e.g., warming) can overshadow the eCO2 effects in observational data of real-world ecosystems. In this study, we aim at developing a method to detect the emergence of the physiological CO2 effects on various variables related to carbon and water fluxes. We mimic the observational setting in ecosystems using a comprehensive process-based land surface model QUINCY to simulate the leaf-level effects of increasing atmospheric CO2 concentrations and their century-long propagation through the terrestrial carbon and water cycles across different climate regimes and biomes. We then develop a statistical method based on the signal-to-noise ratio to detect the emergence of the eCO2 effects. The eCO2 effect on gross primary productivity (GPP) emerges at relatively low CO2 increase (∆[CO2 ] ~ 20 ppm) where the leaf area index is relatively high. Compared to GPP, the eCO2 effect causing reduced transpiration water flux (normalized to leaf area) emerges only at relatively high CO2 increase (∆[CO2 ] >> 40 ppm), due to the high sensitivity to climate variability and thus lower signal-to-noise ratio. In general, the response to eCO2 is detectable earlier for variables related to the carbon cycle than the water cycle, when plant productivity is not limited by climatic constraints, and stronger in forest-dominated rather than in grass-dominated ecosystems. Our results provide a step toward when and where we expect to detect physiological CO2 effects in in-situ flux measurements, how to detect them and encourage future efforts to improve the understanding and quantification of these effects in observations of terrestrial carbon and water dynamics.

Observation-based assessment of secondary water effects on seasonal vegetation decay across Africa.

Local studies and modeling experiments suggest that shallow groundwater and lateral redistribution of soil moisture, together with soil properties, can be highly important secondary water sources for vegetation in water-limited ecosystems. However, there is a lack of observation-based studies of these terrain-associated secondary water effects on vegetation over large spatial domains. Here, we quantify the role of terrain properties on the spatial variations of dry season vegetation decay rate across Africa obtained from geostationary satellite acquisitions to assess the large-scale relevance of secondary water effects. We use machine learning based attribution to identify where and under which conditions terrain properties related to topography, water table depth, and soil hydraulic properties influence the rate of vegetation decay. Over the study domain, the machine learning model attributes about one-third of the spatial variations of vegetation decay rates to terrain properties, which is roughly equally split between direct terrain effects and interaction effects with climate and vegetation variables. The importance of secondary water effects increases with increasing topographic variability, shallower groundwater levels, and the propensity to capillary rise given by soil properties. In regions with favorable terrain properties, more than 60% of the variations in the decay rate of vegetation are attributed to terrain properties, highlighting the importance of secondary water effects on vegetation in Africa. Our findings provide an empirical assessment of the importance of local-scale secondary water effects on vegetation over Africa and help to improve hydrological and vegetation models for the challenge of bridging processes across spatial scales.

Research priorities for global food security under extreme events.

Extreme events, such as those caused by climate change, economic or geopolitical shocks, and pest or disease epidemics, threaten global food security. The complexity of causation, as well as the myriad ways that an event, or a sequence of events, creates cascading and systemic impacts, poses significant challenges to food systems research and policy alike. To identify priority food security risks and research opportunities, we asked experts from a range of fields and geographies to describe key threats to global food security over the next two decades and to suggest key research questions and gaps on this topic. Here, we present a prioritization of threats to global food security from extreme events, as well as emerging research questions that highlight the conceptual and practical challenges that exist in designing, adopting, and governing resilient food systems. We hope that these findings help in directing research funding and resources toward food system transformations needed to help society tackle major food system risks and food insecurity under extreme events.

Widespread increasing vegetation sensitivity to soil moisture.

Global vegetation and associated ecosystem services critically depend on soil moisture availability which has decreased in many regions during the last three decades. While spatial patterns of vegetation sensitivity to global soil water have been recently investigated, long-term changes in vegetation sensitivity to soil water availability are still unclear. Here we assess global vegetation sensitivity to soil moisture during 1982-2017 by applying explainable machine learning with observation-based leaf area index (LAI) and hydro-climate anomaly data. We show that LAI sensitivity to soil moisture significantly increases in many semi-arid and arid regions. LAI sensitivity trends are associated with multiple hydro-climate and ecological variables, and strongest increasing trends occur in the most water-sensitive regions which additionally experience declining precipitation. State-of-the-art land surface models do not reproduce this increasing sensitivity as they misrepresent water-sensitive regions and sensitivity strength. Our sensitivity results imply an increasing ecosystem vulnerability to water availability which can lead to exacerbated reductions in vegetation carbon uptake under future intensified drought, consequently amplifying climate change.

Characterizing the Response of Vegetation Cover to Water Limitation in Africa Using Geostationary Satellites.

Hydrological interactions between vegetation, soil, and topography are complex, and heterogeneous in semi-arid landscapes. This along with data scarcity poses challenges for large-scale modeling of vegetation-water interactions. Here, we exploit metrics derived from daily Meteosat data over Africa at ca. 5 km spatial resolution for ecohydrological analysis. Their spatial patterns are based on Fractional Vegetation Cover (FVC) time series and emphasize limiting conditions of the seasonal wet to dry transition: the minimum and maximum FVC of temporal record, the FVC decay rate and the FVC integral over the decay period. We investigate the relevance of these metrics for large scale ecohydrological studies by assessing their co-variation with soil moisture, and with topographic, soil, and vegetation factors. Consistent with our initial hypothesis, FVC minimum and maximum increase with soil moisture, while the FVC integral and decay rate peak at intermediate soil moisture. We find evidence for the relevance of topographic moisture variations in arid regions, which, counter-intuitively, is detectable in the maximum but not in the minimum FVC. We find no clear evidence for wide-spread occurrence of the "inverse texture effect" on FVC. The FVC integral over the decay period correlates with independent data sets of plant water storage capacity or rooting depth while correlations increase with aridity. In arid regions, the FVC decay rate decreases with canopy height and tree cover fraction as expected for ecosystems with a more conservative water-use strategy. Thus, our observation-based products have large potential for better understanding complex vegetation-water interactions from regional to continental scales.

Heatwave breaks down the linearity between sun-induced fluorescence and gross primary production.

Sun-induced fluorescence in the far-red region (SIF) is increasingly used as a remote and proximal-sensing tool capable of tracking vegetation gross primary production (GPP). However, the use of SIF to probe changes in GPP is challenged during extreme climatic events, such as heatwaves. Here, we examined how the 2018 European heatwave (HW) affected the GPP-SIF relationship in evergreen broadleaved trees with a relatively invariant canopy structure. To do so, we combined canopy-scale SIF measurements, GPP estimated from an eddy covariance tower, and active pulse amplitude modulation fluorescence. The HW caused an inversion of the photosynthesis-fluorescence relationship at both the canopy and leaf scales. The highly nonlinear relationship was strongly shaped by nonphotochemical quenching (NPQ), that is, a dissipation mechanism to protect from the adverse effects of high light intensity. During the extreme heat stress, plants experienced a saturation of NPQ, causing a change in the allocation of energy dissipation pathways towards SIF. Our results show the complex modulation of the NPQ-SIF-GPP relationship at an extreme level of heat stress, which is not completely represented in state-of-the-art coupled radiative transfer and photosynthesis models.

Climatic and soil factors explain the two-dimensional spectrum of global plant trait variation.

Plant functional traits can predict community assembly and ecosystem functioning and are thus widely used in global models of vegetation dynamics and land-climate feedbacks. Still, we lack a global understanding of how land and climate affect plant traits. A previous global analysis of six traits observed two main axes of variation: (1) size variation at the organ and plant level and (2) leaf economics balancing leaf persistence against plant growth potential. The orthogonality of these two axes suggests they are differently influenced by environmental drivers. We find that these axes persist in a global dataset of 17 traits across more than 20,000 species. We find a dominant joint effect of climate and soil on trait variation. Additional independent climate effects are also observed across most traits, whereas independent soil effects are almost exclusively observed for economics traits. Variation in size traits correlates well with a latitudinal gradient related to water or energy limitation. In contrast, variation in economics traits is better explained by interactions of climate with soil fertility. These findings have the potential to improve our understanding of biodiversity patterns and our predictions of climate change impacts on biogeochemical cycles.

Soil moisture-atmosphere feedback dominates land carbon uptake variability.

Year-to-year changes in carbon uptake by terrestrial ecosystems have an essential role in determining atmospheric carbon dioxide concentrations1. It remains uncertain to what extent temperature and water availability can explain these variations at the global scale2-5. Here we use factorial climate model simulations6 and show that variability in soil moisture drives 90 per cent of the inter-annual variability in global land carbon uptake, mainly through its impact on photosynthesis. We find that most of this ecosystem response occurs indirectly as soil moisture-atmosphere feedback amplifies temperature and humidity anomalies and enhances the direct effects of soil water stress. The strength of this feedback mechanism explains why coupled climate models indicate that soil moisture has a dominant role4, which is not readily apparent from land surface model simulations and observational analyses2,5. These findings highlight the need to account for feedback between soil and atmospheric dryness when estimating the response of the carbon cycle to climatic change globally5,7, as well as when conducting field-scale investigations of the response of the ecosystem to droughts8,9. Our results show that most of the global variability in modelled land carbon uptake is driven by temperature and vapour pressure deficit effects that are controlled by soil moisture.

A unified vegetation index for quantifying the terrestrial biosphere.

Empirical vegetation indices derived from spectral reflectance data are widely used in remote sensing of the biosphere, as they represent robust proxies for canopy structure, leaf pigment content, and, subsequently, plant photosynthetic potential. Here, we generalize the broad family of commonly used vegetation indices by exploiting all higher-order relations between the spectral channels involved. This results in a higher sensitivity to vegetation biophysical and physiological parameters. The presented nonlinear generalization of the celebrated normalized difference vegetation index (NDVI) consistently improves accuracy in monitoring key parameters, such as leaf area index, gross primary productivity, and sun-induced chlorophyll fluorescence. Results suggest that the statistical approach maximally exploits the spectral information and addresses long-standing problems in satellite Earth Observation of the terrestrial biosphere. The nonlinear NDVI will allow more accurate measures of terrestrial carbon source/sink dynamics and potentials for stabilizing atmospheric CO2 and mitigating global climate change.

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.

Asymmetric responses of ecosystem productivity to rainfall anomalies vary inversely with mean annual rainfall over the conterminous United States.

The CONterminous United States (CONUS) presents a large range of climate conditions and biomes where terrestrial primary productivity and its inter-annual variability are controlled regionally by rainfall and/or temperature. Here, the response of ecosystem productivity to those climate variables was investigated across different biomes from 2010 to 2018 using three climate datasets of precipitation, air temperature or drought severity, combined with several proxies of ecosystem productivity: a remote sensing product of aboveground biomass, an net primary productivity (NPP) remote sensing product, an NPP model-based product and four gross primary productivity products. We used an asymmetry index (AI) where positive AI indicates a greater increase of ecosystem productivity in wet years compared to the decline in dry years, and negative AI indicates a greater decline of ecosystem productivity in dry years compared to the increase in wet years. We found consistent spatial patterns of AI across the CONUS for the different products, with negative asymmetries over the Great Plains and positive asymmetries over the southwestern CONUS. Shrubs and, to a lesser extent, evergreen forests show a persistent positive asymmetry, whilst (natural) grasslands appear to have transitioned from positive to negative anomalies during the last decade. The general tendency of dominant negative asymmetry response for ecosystem productivity across the CONUS appears to be influenced by the negative asymmetry of precipitation anomalies. AI was found to be a function of mean rainfall: more positive AIs were found in dry areas where plants are adapted to drought and take advantage of rainfall pulses, and more negative AIs were found in wet areas, with a threshold delineating the two regimes corresponding to a mean annual rainfall of 200-400 mm/year.

Drought and heatwave impacts on semi-arid ecosystems' carbon fluxes along a precipitation gradient.

The inter-annual variability (IAV) of the terrestrial carbon cycle is tightly linked to the variability of semi-arid ecosystems. Thus, it is of utmost importance to understand what the main meteorological drivers for the IAV of such ecosystems are, and how they respond to extreme events such as droughts and heatwaves. To shed light onto these questions, we analyse the IAV of carbon fluxes, its relation with meteorological variables, and the impact of compound drought and heatwave on the carbon cycle of two similar ecosystems, along a precipitation gradient. A four-year long dataset from 2016 to 2019 was used for the FLUXNET sites ES-LMa and ES-Abr, located in central (39°56'25″ N 5°46'28″ W) and southeastern (38°42'6″ N 6°47'9″ W) Spain. We analyse the physiological impact of compound drought and heatwave on the dominant tree species, Quercus ilex. Our results show that the gross primary productivity of the wetter ecosystem was less sensitive to changes in soil water content, compared to the dryer site. Still, the wetter ecosystem was a source of CO2 each year, owing to large ecosystem respiration during summer; while the dry site turned into a CO2 sink during wet years. Overall, the impact of the summertime compound event on annual CO2 fluxes was marginal at both sites, compared to drought events during spring or autumn. This highlights that drought timing is crucial to determine the annual carbon fluxes in these semi-arid ecosystems. This article is part of the theme issue 'Impacts of the 2018 severe drought and heatwave in Europe: from site to continental scale'.

The FLUXNET2015 dataset and the ONEFlux processing pipeline for eddy covariance data.

The FLUXNET2015 dataset provides ecosystem-scale data on CO2, water, and energy exchange between the biosphere and the atmosphere, and other meteorological and biological measurements, from 212 sites around the globe (over 1500 site-years, up to and including year 2014). These sites, independently managed and operated, voluntarily contributed their data to create global datasets. Data were quality controlled and processed using uniform methods, to improve consistency and intercomparability across sites. The dataset is already being used in a number of applications, including ecophysiology studies, remote sensing studies, and development of ecosystem and Earth system models. FLUXNET2015 includes derived-data products, such as gap-filled time series, ecosystem respiration and photosynthetic uptake estimates, estimation of uncertainties, and metadata about the measurements, presented for the first time in this paper. In addition, 206 of these sites are for the first time distributed under a Creative Commons (CC-BY 4.0) license. This paper details this enhanced dataset and the processing methods, now made available as open-source codes, making the dataset more accessible, transparent, and reproducible.

Partitioning net carbon dioxide fluxes into photosynthesis and respiration using neural networks.

The eddy covariance (EC) technique is used to measure the net ecosystem exchange (NEE) of CO2 between ecosystems and the atmosphere, offering a unique opportunity to study ecosystem responses to climate change. NEE is the difference between the total CO2 release due to all respiration processes (RECO), and the gross carbon uptake by photosynthesis (GPP). These two gross CO2 fluxes are derived from EC measurements by applying partitioning methods that rely on physiologically based functional relationships with a limited number of environmental drivers. However, the partitioning methods applied in the global FLUXNET network of EC observations do not account for the multiple co-acting factors that modulate GPP and RECO flux dynamics. To overcome this limitation, we developed a hybrid data-driven approach based on combined neural networks (NNC-part ). NNC-part incorporates process knowledge by introducing a photosynthetic response based on the light-use efficiency (LUE) concept, and uses a comprehensive dataset of soil and micrometeorological variables as fluxes drivers. We applied the method to 36 sites from the FLUXNET2015 dataset and found a high consistency in the results with those derived from other standard partitioning methods for both GPP (R2  > .94) and RECO (R2  > .8). High consistency was also found for (a) the diurnal and seasonal patterns of fluxes and (b) the ecosystem functional responses. NNC-part performed more realistic than the traditional methods for predicting additional patterns of gross CO2 fluxes, such as: (a) the GPP response to VPD, (b) direct effects of air temperature on GPP dynamics, (c) hysteresis in the diel cycle of gross CO2 fluxes, (d) the sensitivity of LUE to the diffuse to direct radiation ratio, and (e) the post rain respiration pulse after a long dry period. In conclusion, NNC-part is a valid data-driven approach to provide GPP and RECO estimates and complementary to the existing partitioning methods.