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The leaf economics spectrum1,2 and the global spectrum of plant forms and functions3 revealed fundamental axes of variation in plant traits, which represent different ecological strategies that are shaped by the evolutionary development of plant species2. Ecosystem functions depend on environmental conditions and the traits of species that comprise the ecological communities4. However, the axes of variation of ecosystem functions are largely unknown, which limits our understanding of how ecosystems respond as a whole to anthropogenic drivers, climate and environmental variability4,5. Here we derive a set of ecosystem functions6 from a dataset of surface gas exchange measurements across major terrestrial biomes. We find that most of the variability within ecosystem functions (71.8%) is captured by three key axes. The first axis reflects maximum ecosystem productivity and is mostly explained by vegetation structure. The second axis reflects ecosystem water-use strategies and is jointly explained by variation in vegetation height and climate. The third axis, which represents ecosystem carbon-use efficiency, features a gradient related to aridity, and is explained primarily by variation in vegetation structure. We show that two state-of-the-art land surface models reproduce the first and most important axis of ecosystem functions. However, the models tend to simulate more strongly correlated functions than those observed, which limits their ability to accurately predict the full range of responses to environmental changes in carbon, water and energy cycling in terrestrial ecosystems7,8., Three key axes of variation of ecosystem functional changes and their underlying causes are identified from a dataset of surface gas exchange measurements across major terrestrial biomes and climate zones.

13 Pág. Departamento de Medio Ambiente y Agronomía​ (INIA), Remote sensing capabilities to monitor evergreen broadleaved vegetation are limited by the low temporal variability in the greenness signal. With canopy greenness computed from digital repeat photography (PhenoCam), we investigated how canopy greenness related to seasonal changes in leaf age and traits as well as variation of trees’ water fluxes (characterized by sap flow and canopy conductance). The results showed that sprouting leaves are mainly responsible for the rapid increase in canopy green chromatic coordinate (GCC) in spring. We found statistically significantly differences in leaf traits and spectral properties among leaves of different leaf ages. Specifically, mean GCC of young leaves was 0.385 ± 0.010 (mean ± SD), while for mature and old leaves was 0.369 ± 0.003, and 0.376 ± 0.004, respectively. Thus, the temporal dynamics of canopy GCC can be explained by changes in leaf spectral properties and leaf age. Sap flow and canopy conductance are both well explained by a combination of environmental drivers and greenness (96% and 87% of the variance explained, respectively). In particular, air temperature and vapor pressure deficit (VPD) explained most of sap flow and canopy conductance variance, respectively. Besides, GCC is an important explanatory variable for variation of canopy conductance may because GCC can represent the leaf ontogeny information. We conclude that PhenoCam GCC can be used to identify the leaf flushing for evergreen broadleaved trees, which carries important information about leaf ontogeny and traits. Thus, it can be helpful for better estimating canopy conductance which constraints water fluxes., The authors acknowledge the Alexander von Humboldt Foundation for supporting this research with the Max Planck Prize to Markus Reichstein. Yunpeng Luo and Mirco Migliavacca gratefully acknowledge the financial support from the China Scholarship Council. ADR acknowledges support for the PhenoCam network from the National Science Foundation ( DEB- 1702697 ). Javier Pacheco-Labrador and Mirco Migliavacca acknowledge the German Aerospace Center (DLR) project OBEF-Accross2 “The Potential of Earth Observations to Capture Patterns of Biodiversity” (Contract No. 50EE1912). The research also received funding from the European Union's Horizon 2020 research and innovation programme under the Marie Sklodowska-Curie grant agreement No. 721995 and Ministerio de Economíay Competitividad through FLUXPEC CGL2012-34383 and SynerTGE CGL2015-G9095-R (MINECO/FEDER, UE) projects.

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.

Quantifying global photosynthesis remains a challenge due to a lack of accurate remote sensing proxies. Solar-induced chlorophyll fluorescence (SIF) has been shown to be a good indicator of photosynthetic activity across various spatial scales. However, a global and spatially challenging estimate of terrestrial gross primary production (GPP) based on satellite SIF remains unresolved due to the confounding effects of species-specific physical and physiological traits and external factors, such as canopy structure or photosynthetic pathway (C-3 or C-4). Here we analyze an ensemble of far-red SIF data from OCO-2 satellite and ground observations at multiple sites, using the spectral invariant theory to reduce the effects of canopy structure and to retrieve a structure-corrected total canopy SIF emission (SIFtotal). We find that the relationships between observed canopy-leaving SIF and ecosystem GPP vary significantly among biomes. In contrast, the relationships between SIFtotal and GPP converge around two unique models, one for C-3 and one for C-4 plants. We show that the two single empirical models can be used to globally scale satellite SIF observations to terrestrial GPP. We obtain an independent estimate of global terrestrial GPP of 129.56 +/- 6.54 PgC/year for the 2015-2017 period, which is consistent with the state-of-the-art data- and process-oriented models. The new GPP product shows improved sensitivity to previously undetected 'hotspots' of productivity, being able to resolve the double-peak in GPP due to rotational cropping systems. We suggest that the direct scheme to estimate GPP presented here, which is based on satellite SIF, may open up new possibilities to resolve the dynamics of global terrestrial GPP across space and time.

Remote Sensing (RS) has traditionally provided estimates of key biophysical properties controlling light interaction with the canopy (e.g., chlorophyll content (Cab) or leaf area index (LAI)). However, recent and upcoming developments in hyperspectral RS are expected to lead to a new generation of products such as vegetation functional traits that control leaf carbon and water gas exchange. This information is pivotal to improve our understanding and capability to predict biosphere-atmosphere fluxes at global scale. Yet, the retrieval of key functional traits such as maximum carboxylation rate (Vcmax) or the Ball-Berry stomatal sensitivity parameter (m) remains challenging, as they only have a weak and indirect influence on optical reflectance factors. Recently, the assimilation of different observations in coupled soil-vegetation-atmosphere transfer (SVAT) and radiative transfer models (RTM) is allowing Vcmax and m estimates; notably using the Soil Canopy Observation of Photosynthesis and Energy fluxes (SCOPE) model. In this work we assess the potential of airborne and satellite emulated hyperspectral imagery jointly with eddy covariance (EC) data for the retrieval of functional traits. Specifically, we made use of time series of gross primary production (GPP) and thermal irradiance measured with net radiometers, together with 17 hyperspectral airborne images. The potential of satellite-borne sensors was tested with emulated EnMAP imagery from the airborne data. EnMAP was selected because of the availability of the emulator, and because is one of the foreseen hyperspectral satellite missions expected to contribute to a new generation of RS products. We estimated ecosystem functional traits by inverting the senSCOPE model, a novel version of SCOPE adapted to represent partly senescent canopies. The experiment takes place in a Mediterranean tree-grass ecosystem subject of a large scale manipulation experiment with nitrogen and nitrogen plus phosphorus, monitored by three EC towers. Parameter estimates and predicted fluxes were evaluated using both ground observations and pattern-oriented model evaluation approach. The method developed in this study provided robust estimates of functional and biophysical parameters for both airborne and synthetic EnMAP datasets. Cab and Vcmax estimates followed observed relationships with leaf nitrogen concentration; whereas m and predicted underlying water use efficiency showed expected relationships with discrimination of 13C isotope in leaves. Results prove that the inversion of coupled RTM-SVAT models against a combination of hyperspectral imagery (e.g., EnMAP), and time series of GPP and thermal irradiance provides reliable estimates of key functional parameters of vegetation that are robust to several sources of uncertainty. The forthcoming satellite hyperspectral missions combined with ecosystem station networks (e.g. Integrated Carbon Observation System (ICOS), NEON, FLUXNET, etc…), offers unique possibilities to characterize the spatiotemporal distribution of functional parameters relevant for terrestrial biosphere modeling.

Summary Plant biomass allocation may be optimized to acquire and conserve resources. How trade‐offs in the allocation of tropical tree seedlings depend on different stressors remains poorly understood. Here we test whether above‐ and below‐ground traits of tropical tree seedlings could explain observed occurrence along gradients of resources (light, water) and defoliation (fire, herbivory).We grew 24 tree species occurring in five African vegetation types, varying from dry savanna to moist forest, in a glasshouse for 6 months, and measured traits associated with biomass allocation.Classification based on above‐ground traits resulted in clusters representing savanna and forest species, with low and high shoot investment, respectively. Classification based on root traits resulted in four clusters representing dry savanna, humid savanna, dry forest and moist forest, characterized by a deep mean rooting depth, root starch investment, high specific root length in deeper soil layers, and high specific root length in the top soil layer, respectively.In conclusion, tree seedlings in this study show root trait syndromes, which vary along gradients of resources and defoliation: seedlings from dry areas invest in deep roots, seedlings from shaded environments optimize shoot investment, and seedlings experiencing frequent defoliation store resources in the roots.

The most recent efforts to provide remote sensing (RS) estimates of plant function rely on the combination of Radiative Transfer Models (RTM) and Soil-Vegetation-Atmosphere Transfer (SVAT) models, such as the SoilCanopy Observation Photosynthesis and Energy fluxes (SCOPE) model. In this work we used ground spectroradiometric and chamber-based CO2 flux measurements in a nutrient manipulated Mediterranean grassland in order to: 1) develop a multiple-constraint inversion approach of SCOPE able to retrieve vegetation biochemical, structural as well as key functional traits, such as chlorophyll concentration (Cab), leaf area index (LAI), maximum carboxylation rate (Vcmax) and the Ball-Berry sensitivity parameter (m); and 2) compare the potential of the of gross primary production (GPP) and sun-induced fluorescence (SIF), together with up-welling Thermal Infrared (TIR) radiance and optical reflectance factors (RF), to estimate such parameters. The performance of the proposed inversion method as well as of the different sets of constraints was assessed with contemporary measurements of water and heat fluxes and leaf nitrogen content, using pattern-oriented model evaluation. The multiple-constraint inversion approach proposed together with the combination of optical RF and diel GPP and TIR data provided reliable estimates of parameters, and improved predicted water and heat fluxes. The addition of SIF to this scheme slightly improved the estimation of m. Parameter estimates were coherent with the variability imposed by the fertilization and the seasonality of the grassland. Results revealed that fertilization had an impact on Vcmax, while no significant differences were found for m. The combination of RF, SIF and diel TIR data weakly constrained functional traits. Approaches not including GPP failed to estimate LAI; however GPP overestimated Cab in the dry period. These problems might be related to the presence of high fractions of senescent leaves in the grassland. The proposed inversion approach together with pattern-oriented model evaluation open new perspectives for the retrieval of plant functional traits relevant for land surface models, and can be utilized at various research sites where hyperspectral remote sensing imagery and eddy covariance flux measurements are simultaneously taken, JPL, MM, and MR acknowledge the EnMAP project MoReDEHESHyReS “Modelling Responses of Dehesas with Hyperspectral Remote Sensing” (Contract No. 50EE1621, German Aerospace Center (DLR) and the German Federal Ministry of Economic Affairs and Energy). DM, MM and MR received funding from the European Union’s Horizon 2020 research and innovation programme via the TRUSTEE project under the Marie Skłodowska-Curie grant agreement No. 721995. Authors acknowledge the Alexander von Humboldt Foundation for supporting this research with the Max-Planck Prize to Markus Reichstein; SynerTGE “Landsat-8 + Sentinel-2: exploring sensor synergies for monitoring and modelling key vegetation biophysical variables in tree-grass ecosystems” (CGL2015-69095-R, Spanish Ministry of Science, Innovation and Universities); and FLUχPEC “Monitoring changes in water and carbon fluxes from remote and proximal sensing in Mediterranean ‘dehesa’ ecosystem” (CGL2012- 34383, Spanish Ministry of Economy and Competitiveness). Authors are very thankful to Dr. Karl Segl and Prof. Dr. Luis Guanter for their support with the EnMAP end-to-end simulator; as well as the MPI-BGC Freiland Group and especially Martin Hertel as well as Ramón LópezJiménez (CEAM) for technical assistance. We are grateful to all the colleagues from MPI-BGC, University of Extremadura, University of Milano-Bicocca, SpecLab-CSIC, INIA and CEAM which have collaborated in any of the field and laboratory works. We acknowledge the Majadas de Tiétar city council for its support.

Sun-Induced fluorescence at 760 nm (F760) is increasingly being used to predict gross primary production (GPP) through light use efficiency (LUE) modeling, even though the mechanistic processes that link the two are not well understood. We analyzed the effect of nitrogen (N) and phosphorous (P) availability on the processes that link GPP and F760 in a Mediterranean grassland manipulated with nutrient addition. To do so, we used a combination of process-based modeling with Soil-Canopy Observation of Photosynthesis and Energy (SCOPE), and statistical analyses such as path modeling. With this study, we uncover the mechanisms that link the fertilization-driven changes in canopy nitrogen concentration (N%) to the observed changes in F760 and GPP. N addition changed plant community structure and increased canopy chlorophyll content, which jointly led to changes in photosynthetic active radiation (APAR), ultimately affecting both GPP and F760. Changes in the abundance of graminoids, (%graminoids) driven by N addition led to changes in structural properties of the canopy such as leaf angle distribution, that ultimately influenced observed F760 by controlling the escape probability of F760 (Fesc). In particular, we found a change in GPP¿F760 relationship between the first and the second year of the experiment that was largely driven by the effect of plant type composition on Fesc, whose best predictor is %graminoids. The P addition led to a statistically significant increase on light use efficiency of fluorescence emission (LUEf), in particular in plots also with N addition, consistent with leaf level studies. The N addition induced changes in the biophysical properties of the canopy that led to a trade-off between surface temperature (Ts), which decreased, and F760 at leaf scale (F760leaf,fw), which increased. We found that Ts is an important predictor of the light use efficiency of photosynthesis, indicating the importance of Ts in LUE modeling approaches to predict GPP, The project received funding from the European Union’s Horizon 2020 research and innovation program under the Marie Sklodowska-Curie grant agreement No. 721995. The authors acknowledge the Alexander von Humboldt Foundation for supporting this research with the Max-Planck Prize to Markus Reichstein, and the EUFAR TA project DEHESHyrE (EU FP7 Program), the EnMAP project “MoReDEHESHyReS” (Contract No. 50EE1621, German Aerospace Center (DLR) and German Federal Ministry of Economic Affairs and Energy), SynerTGE (CGL2015-69095-R, MINECO/FEDER, UE) and FLUχPEC (CGL2012-34383, Spanish Ministry of Economy and Competitiveness). This work was supported by a research grant (18968) from VILLUM FONDEN

Vegetation transpiration (T) is the process of plant water loss through the stomata on the leaf surface and plays a key role in the energy and water balance of the land surface, especially with dense vegetation cover. To date, however, estimation of ecosystem-scale T is still rather uncertain mainly due to errors in modeling canopy resistance or conductance. Considering the intrinsic link between photosynthesis and chlorophyll fluorescence, the recent available remote sensing of solar-induced chlorophyll fluorescence (SIF) provides a valuable opportunity to estimate plants T at large scales. In this study, we demonstrate how remote sensing of SIF relates to canopy stomatal conductance and transpiration at diurnal and seasonal scales with continuous ground measurements of SIF at three flux sites in forest, cropland and grassland ecosystems. The results show that both ground and spaceborne SIF observations are good indicators of canopy conductance at both diurnal and seasonal scales (R2 = 0.57 and 0.74 for forest, R2 = 0.62 and 0.80 for cropland, R2 = 0.52 and 0.63 for grassland, respectively). Then, empirical SIF-based canopy conductance models are employed to estimate hourly and daily transpiration. We evaluate our ecosystem T estimations against latent heat fluxes measured by eddy covariance systems with more satisfactory results for forest (R2 = 0.57 and 0.71), and cropland (R2 = 0.77 and 0.83) than for grassland (R2 = 0.13 and 0.22) at hourly and daily time scales. Our results suggest the potential of remotely-sensed SIF for estimating canopy conductance and plant transpiration, but a more mechanistic understanding is needed for their link.

Sun induced fluorescence at 760 nm (F 760 ) has shown to provide a valid approach to quantify gross primary production (GPP) at various scales, however the relationship between GPP and F760 is influenced by the escape probability of fluorescence (Fesc), a variable which is not still fully understood. Combining radiative transfer modelling approaches, by means of the SCOPE model, and a data driven methodology based on variable selection methods we identify the predictors of Fesc, focusing on the effect of functional and structural traits. We show that Fesc is mainly predicted by structural variables such as fraction of grasses and near infrared reflectance. Building on the analysis of the predictor of Fesc, LUE p and LUE f we present a semi-empirical model formulation based only on optical data that significantly improves the GPP prediction.

Omega‐6 (n‐6) arachidonic acid (AA) and its pro‐inflammatory metabolites, including prostaglandin E2 (PGE (2)), are known to promote tumorigenesis. Delta‐6 desaturase (D6D) is the rate‐limiting enzyme for converting n‐6 linoleic acid (LA) to AA. Our objective was to determine if AA synthesis, specifically D6D activity, and PGE (2) levels are increased in cancerous breast tissue, and whether these variables differ between estrogen receptor positive (ER+) and negative (ER−) breast cancers. Gas chromatography was performed on surgical breast tissue samples collected from 69 women with breast cancer. Fifty‐four had ER+ breast cancer, and 15 had ER− breast cancer. Liquid chromatography‐mass spectrometry was used to determine PGE (2) levels. Lipid analysis revealed higher levels of LA metabolites (C18:3 n‐6, C20:3 n‐6, and AA) in cancerous tissue than in adjacent noncancerous tissue (P