Uncovering the critical soil moisture thresholds of plant water stress for European ecosystems.

Understanding the critical soil moisture (SM) threshold (θcrit ) of plant water stress and land surface energy partitioning is a basis to evaluate drought impacts and improve models for predicting future ecosystem condition and climate. Quantifying the θcrit across biomes and climates is challenging because observations of surface energy fluxes and SM remain sparse. Here, we used the latest database of eddy covariance measurements to estimate θcrit across Europe by evaluating evaporative fraction (EF)-SM relationships and investigating the covariance between vapor pressure deficit (VPD) and gross primary production (GPP) during SM dry-down periods. We found that the θcrit and soil matric potential threshold in Europe are 16.5% and -0.7 MPa, respectively. Surface energy partitioning characteristics varied among different vegetation types; EF in savannas had the highest sensitivities to SM in water-limited stage, and the lowest in forests. The sign of the covariance between daily VPD and GPP consistently changed from positive to negative during dry-down across all sites when EF shifted from relatively high to low values. This sign of the covariance changed after longer period of SM decline in forests than in grasslands and savannas. Estimated θcrit from the VPD-GPP covariance method match well with the EF-SM method, showing this covariance method can be used to detect the θcrit . We further found that soil texture dominates the spatial variability of θcrit while shortwave radiation and VPD are the major drivers in determining the spatial pattern of EF sensitivities. Our results highlight for the first time that the sign change of the covariance between daily VPD and GPP can be used as an indicator of how ecosystems transition from energy to SM limitation. We also characterized the corresponding θcrit and its drivers across diverse ecosystems in Europe, an essential variable to improve the representation of water stress in land surface models.

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.

Does long-term cultivation of saplings under elevated CO2 concentration influence their photosynthetic response to temperature?

BACKGROUND AND AIMS: Plants growing under elevated atmospheric CO2 concentrations often have reduced stomatal conductance and subsequently increased leaf temperature. This study therefore tested the hypothesis that under long-term elevated CO2 the temperature optima of photosynthetic processes will shift towards higher temperatures and the thermostability of the photosynthetic apparatus will increase. METHODS: The hypothesis was tested for saplings of broadleaved Fagus sylvatica and coniferous Picea abies exposed for 4-5 years to either ambient (AC; 385 µmol mol(-1)) or elevated (EC; 700 µmol mol(-1)) CO2 concentrations. Temperature response curves of photosynthetic processes were determined by gas-exchange and chlorophyll fluorescence techniques. KEY RESULTS: Initial assumptions of reduced light-saturated stomatal conductance and increased leaf temperatures for EC plants were confirmed. Temperature response curves revealed stimulation of light-saturated rates of CO2 assimilation (Amax) and a decline in photorespiration (RL) as a result of EC within a wide temperature range. However, these effects were negligible or reduced at low and high temperatures. Higher temperature optima (Topt) of Amax, Rubisco carboxylation rates (VCmax) and RL were found for EC saplings compared with AC saplings. However, the shifts in Topt of Amax were instantaneous, and disappeared when measured at identical CO2 concentrations. Higher values of Topt at elevated CO2 were attributed particularly to reduced photorespiration and prevailing limitation of photosynthesis by ribulose-1,5-bisphosphate (RuBP) regeneration. Temperature response curves of fluorescence parameters suggested a negligible effect of EC on enhancement of thermostability of photosystem II photochemistry. CONCLUSIONS: Elevated CO2 instantaneously increases temperature optima of Amax due to reduced photorespiration and limitation of photosynthesis by RuBP regeneration. However, this increase disappears when plants are exposed to identical CO2 concentrations. In addition, increased heat-stress tolerance of primary photochemistry in plants grown at elevated CO2 is unlikely. The hypothesis that long-term cultivation at elevated CO2 leads to acclimation of photosynthesis to higher temperatures is therefore rejected. Nevertheless, incorporating acclimation mechanisms into models simulating carbon flux between the atmosphere and vegetation is necessary.

Estimation of photosynthetic dynamics in forests from daily measured fluorescence and PRI data with adjustment for canopy shadow fraction.

We conducted year-long measurements of the photochemical reflectance index (PRI) and solar-induced fluorescence in the O2A oxygen band (SIFA) at a Norway spruce forest and a European beech forest to study relationships of these remote sensing variables to photosynthesis by trees in grown forest stands. Measured PRI and SIFA values were linked to changes in forest gross primary productivity (GPP) and light-use efficiency (LUE). Changes in the shadow fraction (αS) within tree crowns influenced PRI and fluorescence signals. In the spruce forest, the quantum yield of SIFA (FYSIFA) decreased around midday together with photosynthesis and GPP. Such decreases in FYSIFA were accompanied by an increase in the αS. In the beech forest, we detected an increase in FYSIFA together with a decrease in αS in the afternoon hours. The overall sensitivity of PRI to LUE was variable according to the season, presumably influenced by complex changes in photosynthetic pigments. PRI and FYSIFA showed weak correlations with canopy LUE; however, when considered together, the correlation was strengthened (R2 was 0.63 and 0.34 in spruce and beech forest, respectively). Our model predicting LUE dynamics includes a diurnal minimum of PRI and canopy αS to make allowances for seasonal changes in photosynthetic pigments and for diurnal variability of the shadow fraction in forests. The incorporation of these correcting factors allowed us to estimate LUE at R2 = 0.68 (spruce) and 0.53 (beech). The modeling equations appeared sensitive to the absorbed photosynthetically active radiation (APAR), but less sensitive to the GPP of these forests. Substituting pigments correction with introducing differential PRI (ΔPRI) into the model did not significantly improve the LUE estimation across the season. Our results show that the joint use of PRI and fluorescence improves LUE and GPP estimation accuracy in both daily and seasonal observations.

Detection and attribution of an anomaly in terrestrial photosynthesis in Europe during the COVID-19 lockdown.

Carbon dioxide (CO2) uptake by plant photosynthesis, referred to as gross primary production (GPP) at the ecosystem level, is sensitive to environmental factors, including pollutant exposure, pollutant uptake, and changes in the scattering of solar shortwave irradiance (SWin) - the energy source for photosynthesis. The 2020 spring lockdown due to COVID-19 resulted in improved air quality and atmospheric transparency, providing a unique opportunity to assess the impact of air pollutants on terrestrial ecosystem functioning. However, detecting these effects can be challenging as GPP is influenced by other meteorological drivers and management practices. Based on data collected from 44 European ecosystem-scale CO2 flux monitoring stations, we observed significant changes in spring GPP at 34 sites during 2020 compared to 2015-2019. Among these, 14 sites showed an increase in GPP associated with higher SWin, 10 sites had lower GPP linked to atmospheric and soil dryness, and seven sites were subjected to management practices. The remaining three sites exhibited varying dynamics, with one experiencing colder and rainier weather resulting in lower GPP, and two showing higher GPP associated with earlier spring melts. Analysis using the regional atmospheric chemical transport model (LOTOS-EUROS) indicated that the ozone (O3) concentration remained relatively unchanged at the research sites, making it unlikely that O3 exposure was the dominant factor driving the primary production anomaly. In contrast, SWin increased by 9.4 % at 36 sites, suggesting enhanced GPP possibly due to reduced aerosol optical depth and cloudiness. Our findings indicate that air pollution and cloudiness may weaken the terrestrial carbon sink by up to 16 %. Accurate and continuous ground-based observations are crucial for detecting and attributing subtle changes in terrestrial ecosystem functioning in response to environmental and anthropogenic drivers.

Joint optimization of land carbon uptake and albedo can help achieve moderate instantaneous and long-term cooling effects.

Both carbon dioxide uptake and albedo of the land surface affect global climate. However, climate change mitigation by increasing carbon uptake can cause a warming trade-off by decreasing albedo, with most research focusing on afforestation and its interaction with snow. Here, we present carbon uptake and albedo observations from 176 globally distributed flux stations. We demonstrate a gradual decline in maximum achievable annual albedo as carbon uptake increases, even within subgroups of non-forest and snow-free ecosystems. Based on a paired-site permutation approach, we quantify the likely impact of land use on carbon uptake and albedo. Shifting to the maximum attainable carbon uptake at each site would likely cause moderate net global warming for the first approximately 20 years, followed by a strong cooling effect. A balanced policy co-optimizing carbon uptake and albedo is possible that avoids warming on any timescale, but results in a weaker long-term cooling effect.

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.

Combining NDVI, PRI and the quantum yield of solar-induced fluorescence improves estimations of carbon fluxes in deciduous and evergreen forests.

We used automated spectroradiometers to continuously monitor changes in the optical parameters of phenological and photosynthetic traits in beech and spruce forests. We examined seasonal variations in the normalized difference vegetation index (NDVI), photochemical reflectance index (PRI), and solar-induced fluorescence in the oxygen A band (SIFA) that was estimated using a 3-FLD discrimination method from radiance data. The optical parameters tracked the activation and cessation of photosynthesis in spring and autumn. Data at photon fluxes >1200 µmol m-2 s-1 during extended noon hours were used to link the seasonal PRI and SIFA variations to the dynamics of photosynthesis. Seasonal PRI was significantly correlated with photosynthetic light-use efficiency (LUE) with R2 values of 0.66 and 0.48 for the measurements in beech and spruce forests, respectively. SIFA emissions were significantly correlated with the gross primary production (GPP) of the evergreen spruce forest (R2 = 0.47), but R2 was only 0.13 when measured in the beech forest. The correlations between the optical parameters and GPP or LUE, however, tended to be lower when using a dataset with constant NDVI. Introducing an equation combining NDVI, PRI, and the quantum yield of SIFA emission increased R2 for LUE estimation to 0.77 in the spruce forest and 0.63 in the beech forest. GPP was estimated from the parametric equation with improved accuracy reaching R2 = 0.53 and RMSE = 5.95 µmol CO2 m-2 s-1 in spruce forest and R2 = 0.58 and RMSE = 5.23 µmol CO2 m-2 s-1 in beech forest. Parametric equations were more efficient in estimating photosynthesis in datasets that consisted of an entire season's data. By combining NDVI, PRI and the quantum yield of SIFA, we could thus substantially improve estimations of carbon fluxes in diverse deciduous and evergreen canopies.

Analysis of floodplain forest sensitivity to drought.

Floodplain forests are very complex, productive ecosystems, capable of storing huge amounts of soil carbon. With the increasing occurrence of extreme events, they are today among the most threatened ecosystems. Our study's main goal was to assess the productivity of a floodplain forest located at Lanzhot in the Czech Republic from two perspectives: carbon uptake (using an eddy covariance method) and stem radius variations (using dendrometers). We aimed to determine which conditions allow for high ecosystem production and what role drought plays in reducing such production potential. Additionally, we were interested to determine the relative soil water content threshold indicating the onset and duration of this event. We hypothesized that summer drought in 2018 had the most significant negative effects on the overall annual carbon and water budgets. In contrast with our original hypothesis, we found that an exceptionally warm spring in 2018 caused a positive gross primary production (GPP) and evapotranspiration (ET) anomaly that consequently led in 2018 to the highest seasonal total GPP and ET from all of the investigated years (2015-2018). The results showed ring-porous species to be the most drought resistant. Relative soil water content threshold of approximately 0.45 was determined as indicating the onset of drought stress. This article is part of the theme issue 'Impacts of the 2018 severe drought and heatwave in Europe: from site to continental scale'.

Non-stomatal processes reduce gross primary productivity in temperate forest ecosystems during severe edaphic drought.

Severe drought events are known to cause important reductions of gross primary productivity (GPP) in forest ecosystems. However, it is still unclear whether this reduction originates from stomatal closure (Stomatal Origin Limitation) and/or non-stomatal limitations (Non-SOL). In this study, we investigated the impact of edaphic drought in 2018 on GPP and its origin (SOL, NSOL) using a dataset of 10 European forest ecosystem flux towers. In all stations where GPP reductions were observed during the drought, these were largely explained by declines in the maximum apparent canopy scale carboxylation rate VCMAX,APP (NSOL) when the soil relative extractable water content dropped below around 0.4. Concurrently, we found that the stomatal slope parameter (G1, related to SOL) of the Medlyn et al. unified optimization model linking vegetation conductance and GPP remained relatively constant. These results strengthen the increasing evidence that NSOL should be included in stomatal conductance/photosynthesis models to faithfully simulate both GPP and water fluxes in forest ecosystems during severe drought. This article is part of the theme issue 'Impacts of the 2018 severe drought and heatwave in Europe: from site to continental scale'.

Sensitivity of gross primary productivity to climatic drivers during the summer drought of 2018 in Europe.

In summer 2018, Europe experienced a record drought, but it remains unknown how the drought affected ecosystem carbon dynamics. Using observations from 34 eddy covariance sites in different biomes across Europe, we studied the sensitivity of gross primary productivity (GPP) to environmental drivers during the summer drought of 2018 versus the reference summer of 2016. We found a greater drought-induced decline of summer GPP in grasslands (-38%) than in forests (-10%), which coincided with reduced evapotranspiration and soil water content (SWC). As compared to the 'normal year' of 2016, GPP in different ecosystems exhibited more negative sensitivity to summer air temperature (Ta) but stronger positive sensitivity to SWC during summer drought in 2018, that is, a stronger reduction of GPP with soil moisture deficit. We found larger negative effects of Ta and vapour pressure deficit (VPD) but a lower positive effect of photosynthetic photon flux density on GPP in 2018 compared to 2016, which contributed to reduced summer GPP in 2018. Our results demonstrate that high temperature-induced increases in VPD and decreases in SWC aggravated drought impacts on GPP. This article is part of the theme issue 'Impacts of the 2018 severe drought and heatwave in Europe: from site to continental scale'.

Ozone flux and ozone deposition in a mountain spruce forest are modulated by sky conditions.

In order to understand the main driving factors of ozone (O3) deposition we tested the hypothesis that sky conditions (cloudy, partly cloudy, and clear sky) modulate O3 flux in forest ecosystems via stomatal regulation. The hypothesis is based on the fact that complex microclimate conditions under cloudy sky usually stimulate stomatal conductance. O3 fluxes were inferred from a concentration gradient in a mountainous Norway spruce forest in the Czech Republic (Central Europe) for years 2012-2016 and measured directly by eddy-covariance during the summer of 2017. Daily and seasonal O3 depositions were calculated separately for days with cloudy, partly cloudy, and clear sky conditions. The data show unequivocally that more O3 is taken up under cloudy and partially cloudy skies. Moreover, we found significant interactive effects of sky conditions and season on O3 flux. Though there are other mechanisms and pathways involved in the transport of O3 to the plant-soil system, the highest O3 deposition was associated to the highest stomatal conductance during partly cloudy and cloudy sky conditions in all seasons, while lower O3 ecosystem fluxes were observed under clear sky conditions despite the highest O3 concentrations at this time. These findings suggest that forests growing at sites where conditions are predominantly cloudy are expected to deposit higher extent of O3 than less-cloudy forests being thus more threatened by phytotoxic O3.

Thinning Can Reduce Losses in Carbon Use Efficiency and Carbon Stocks in Managed Forests Under Warmer Climate.

Forest carbon use efficiency (CUE, the ratio of net to gross primary productivity) represents the fraction of photosynthesis that is not used for plant respiration. Although important, it is often neglected in climate change impact analyses. Here we assess the potential impact of thinning on projected carbon cycle dynamics and implications for forest CUE and its components (i.e., gross and net primary productivity and plant respiration), as well as on forest biomass production. Using a detailed process-based forest ecosystem model forced by climate outputs of five Earth System Models under four representative climate scenarios, we investigate the sensitivity of the projected future changes in the autotrophic carbon budget of three representative European forests. We focus on changes in CUE and carbon stocks as a result of warming, rising atmospheric CO2 concentration, and forest thinning. Results show that autotrophic carbon sequestration decreases with forest development, and the decrease is faster with warming and in unthinned forests. This suggests that the combined impacts of climate change and changing CO2 concentrations lead the forests to grow faster, mature earlier, and also die younger. In addition, we show that under future climate conditions, forest thinning could mitigate the decrease in CUE, increase carbon allocation into more recalcitrant woody pools, and reduce physiological-climate-induced mortality risks. Altogether, our results show that thinning can improve the efficacy of forest-based mitigation strategies and should be carefully considered within a portfolio of mitigation options.

Protective effect of UV-A radiation during acclimation of the photosynthetic apparatus to UV-B treatment.

We examined the acclimation response of the photosynthetic apparatus of barley (Hordeum vulgare L.) to a combination of UV-A and UV-B radiation (UVAB) and to UV-B radiation alone. Our aim was to evaluate whether UV-A radiation prevents UV-B-induced damage to the photosynthetic apparatus and whether UV-A pre-acclimation is required to mitigate the negative influence of UV-B radiation. Barley plants were grown from seeds under low photosynthetically active radiation (50 µmol m(-2) s(-1)) either in the absence or presence of UV-A radiation (UVA- and UVA+ plants, respectively). After 8 days of development, plants were exposed simultaneously to UV-A and UV-B radiation for the next 6 days. Additionally, UVA- plants were exposed to UV-B radiation alone. The UVA+ plants had a higher CO2 assimilation rate near the light-saturation region (A(N)) and a higher content of both total chlorophylls (Chls) and total carotenoids than the UVA- plants. Chls content, A(N), the potential quantum yield of photosystem II (PSII) photochemistry (F(V)/F(M)), the capacity of light-induced thermal energy dissipation and the efficiency of excitation energy transfer within PSII remained the same or even increased in both UVA+ and UVA- plants after UVAB treatment. On the contrary, exposure of UVA- plants to UV-B radiation itself led to a reduction in all these characteristics. We revealed that the presence of UV-A radiation during UVAB treatment not only mitigated but completely eliminated the negative effect of UV-B radiation on the functioning of the photosynthetic apparatus and that UV-A pre-acclimation was not crucial for development of this UV-A-induced resistance against UV-B irradiation.

Impact of elevated CO2 concentration on dynamics of leaf photosynthesis in Fagus sylvatica is modulated by sky conditions.

It has been suggested that atmospheric CO2 concentration and frequency of cloud cover will increase in future. It remains unclear, however, how elevated CO2 influences photosynthesis under complex clear versus cloudy sky conditions. Accordingly, diurnal changes in photosynthetic responses among beech trees grown at ambient (AC) and doubled (EC) CO2 concentrations were studied under contrasting sky conditions. EC stimulated the daily sum of fixed CO2 and light use efficiency under clear sky. Meanwhile, both these parameters were reduced under cloudy sky as compared with AC treatment. Reduction in photosynthesis rate under cloudy sky was particularly associated with EC-stimulated, xanthophyll-dependent thermal dissipation of absorbed light energy. Under clear sky, a pronounced afternoon depression of CO2 assimilation rate was found in sun-adapted leaves under EC compared with AC conditions. This was caused in particular by stomata closure mediated by vapour pressure deficit.