Contrasting anatomical and biochemical controls on mesophyll conductance across plant functional types.

Mesophyll conductance (gm ) limits photosynthesis by restricting CO2 diffusion between the substomatal cavities and chloroplasts. Although it is known that gm is determined by both leaf anatomical and biochemical traits, their relative contribution across plant functional types (PFTs) is still unclear. We compiled a dataset of gm measurements and concomitant leaf traits in unstressed plants comprising 563 studies and 617 species from all major PFTs. We investigated to what extent gm limits photosynthesis across PFTs, how gm relates to structural, anatomical, biochemical, and physiological leaf properties, and whether these relationships differ among PFTs. We found that gm imposes a significant limitation to photosynthesis in all C3 PFTs, ranging from 10-30% in most herbaceous annuals to 25-50% in woody evergreens. Anatomical leaf traits explained a significant proportion of the variation in gm (R2 > 0.3) in all PFTs except annual herbs, in which gm is more strongly related to biochemical factors associated with leaf nitrogen and potassium content. Our results underline the need to elucidate mechanisms underlying the global variability of gm . We emphasise the underestimated potential of gm for improving photosynthesis in crops and identify modifications in leaf biochemistry as the most promising pathway for increasing gm in these species.

Do 2 H and 18 O in leaf water reflect environmental drivers differently?

We compiled hydrogen and oxygen stable isotope compositions (δ2 H and δ18 O) of leaf water from multiple biomes to examine variations with environmental drivers. Leaf water δ2 H was more closely correlated with δ2 H of xylem water or atmospheric vapour, whereas leaf water δ18 O was more closely correlated with air relative humidity. This resulted from the larger proportional range for δ2 H of meteoric waters relative to the extent of leaf water evaporative enrichment compared with δ18 O. We next expressed leaf water as isotopic enrichment above xylem water (Δ2 H and Δ18 O) to remove the impact of xylem water isotopic variation. For Δ2 H, leaf water still correlated with atmospheric vapour, whereas Δ18 O showed no such correlation. This was explained by covariance between air relative humidity and the Δ18 O of atmospheric vapour. This is consistent with a previously observed diurnal correlation between air relative humidity and the deuterium excess of atmospheric vapour across a range of ecosystems. We conclude that 2 H and 18 O in leaf water do indeed reflect the balance of environmental drivers differently; our results have implications for understanding isotopic effects associated with water cycling in terrestrial ecosystems and for inferring environmental change from isotopic biomarkers that act as proxies for leaf water.

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.

Growth resistance and resilience of mixed silver fir and Norway spruce forests in central Europe: Contrasting responses to mild and severe droughts.

Extreme droughts are expected to increase in frequency and severity in many regions of the world, threatening multiple ecosystem services provided by forests. Effective strategies to adapt forests to such droughts require comprehensive information on the effects and importance of the factors influencing forest resistance and resilience. We used a unique combination of inventory and dendrochronological data from a long-term (>30 years) silvicultural experiment in mixed silver fir and Norway spruce mountain forests along a temperature and precipitation gradient in southwestern Germany. We aimed at examining the mechanisms and forest stand characteristics underpinning the resistance and resilience to past mild and severe droughts. We found that (i) fir benefited from mild droughts and showed higher resistance (i.e., lower growth loss during drought) and resilience (i.e., faster return to pre-drought growth levels) than spruce to all droughts; (ii) species identity determined mild drought responses while species interactions and management-related factors strongly influenced the responses to severe droughts; (iii) intraspecific and interspecific interactions had contrasting effects on the two species, with spruce being less resistant to severe droughts when exposed to interaction with fir and beech; (iv) higher values of residual stand basal area following thinning were associated with lower resistance and resilience to severe droughts; and (v) larger trees were resilient to mild drought events but highly vulnerable to severe droughts. Our study provides an analytical approach for examining the effects of different factors on individual tree- and stand-level drought response. The forests investigated here were to a certain extent resilient to mild droughts, and even benefited from such conditions, but were strongly affected by severe droughts. Lastly, negative effects of severe droughts can be reduced through modifying species composition, tree size distribution and stand density in mixed silver fir-Norway spruce forests.

Higher than expected CO2 fertilization inferred from leaf to global observations.

Several lines of evidence point to an increase in the activity of the terrestrial biosphere over recent decades, impacting the global net land carbon sink (NLS) and its control on the growth of atmospheric carbon dioxide (ca ). Global terrestrial gross primary production (GPP)-the rate of carbon fixation by photosynthesis-is estimated to have risen by (31 ± 5)% since 1900, but the relative contributions of different putative drivers to this increase are not well known. Here we identify the rising atmospheric CO2 concentration as the dominant driver. We reconcile leaf-level and global atmospheric constraints on trends in modeled biospheric activity to reveal a global CO2 fertilization effect on photosynthesis of 30% since 1900, or 47% for a doubling of ca above the pre-industrial level. Our historic value is nearly twice as high as current estimates (17 ± 4)% that do not use the full range of available constraints. Consequently, under a future low-emission scenario, we project a land carbon sink (174 PgC, 2006-2099) that is 57 PgC larger than if a lower CO2 fertilization effect comparable with current estimates is assumed. These findings suggest a larger beneficial role of the land carbon sink in modulating future excess anthropogenic CO2 consistent with the target of the Paris Agreement to stay below 2°C warming, and underscore the importance of preserving terrestrial carbon sinks.

Validating MODIS and Sentinel-2 NDVI Products at a Temperate Deciduous Forest Site Using Two Independent Ground-Based Sensors.

Quantifying the accuracy of remote sensing products is a timely endeavor given the rapid increase in Earth observation missions. A validation site for Sentinel-2 products was hence established in central Germany. Automatic multispectral and hyperspectral sensor systems were installed in parallel with an existing eddy covariance flux tower, providing spectral information of the vegetation present at high temporal resolution. Normalized Difference Vegetation Index (NDVI) values from ground-based hyperspectral and multispectral sensors were compared with NDVI products derived from Sentinel-2A and Moderate-resolution Imaging Spectroradiometer (MODIS). The influence of different spatial and temporal resolutions was assessed. High correlations and similar phenological patterns between in situ and satellite-based NDVI time series demonstrated the reliability of satellite-based phenological metrics. Sentinel-2-derived metrics showed better agreement with in situ measurements than MODIS-derived metrics. Dynamic filtering with the best index slope extraction algorithm was nevertheless beneficial for Sentinel-2 NDVI time series despite the availability of quality information from the atmospheric correction procedure.

Oxygen isotope signatures of transpired water vapor: the role of isotopic non-steady-state transpiration under natural conditions.

The oxygen isotope signature of water is a powerful tracer of water movement from plants to the global scale. However, little is known about the short-term variability of oxygen isotopes leaving the ecosystem via transpiration, as high-frequency measurements are lacking. A laser spectrometer was coupled to a gas-exchange chamber directly estimating branch-level fluxes in order to evaluate the short-term variability of the isotopic composition of transpiration (δE ) and to investigate the role of isotopic non-steady-state transpiration under natural conditions in cork-oak trees (Quercus suber) during distinct Mediterranean seasons. The measured δ(18) O of transpiration (δE ) deviated from isotopic steady state throughout most of the day even when leaf water at the evaporating sites was near isotopic steady state. High agreement was found between estimated and modeled δE values assuming non-steady-state enrichment of leaf water. Isoforcing, that is, the influence of the transpirational δ(18) O flux on atmospheric values, deviated from steady-state calculations but daily means were similar between steady state and non-steady state. However, strong daytime isoforcing on the atmosphere implies that short-term variations in δE are likely to have consequences for large-scale applications, for example, partitioning of ecosystem fluxes or satellite-based applications.

Higher global gross primary productivity under future climate with more advanced representations of photosynthesis.

Gross primary productivity (GPP) is the key determinant of land carbon uptake, but its representation in terrestrial biosphere models (TBMs) does not reflect our latest physiological understanding. We implemented three empirically well supported but often omitted mechanisms into the TBM CABLE-POP: photosynthetic temperature acclimation, explicit mesophyll conductance, and photosynthetic optimization through redistribution of leaf nitrogen. We used the RCP8.5 climate scenario to conduct factorial model simulations characterizing the individual and combined effects of the three mechanisms on projections of GPP. Simulated global GPP increased more strongly (up to 20% by 2070-2099) in more comprehensive representations of photosynthesis compared to the model lacking the three mechanisms. The experiments revealed non-additive interactions among the mechanisms as combined effects were stronger than the sum of the individual effects. The modeled responses are explained by changes in the photosynthetic sensitivity to temperature and CO2 caused by the added mechanisms. Our results suggest that current TBMs underestimate GPP responses to future CO2 and climate conditions.

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.

The impact of soil microorganisms on the global budget of delta18O in atmospheric CO2.

Improved global estimates of terrestrial photosynthesis and respiration are critical for predicting the rate of change in atmospheric CO(2). The oxygen isotopic composition of atmospheric CO(2) can be used to estimate these fluxes because oxygen isotopic exchange between CO(2) and water creates distinct isotopic flux signatures. The enzyme carbonic anhydrase (CA) is known to accelerate this exchange in leaves, but the possibility of CA activity in soils is commonly neglected. Here, we report widespread accelerated soil CO(2) hydration. Exchange was 10-300 times faster than the uncatalyzed rate, consistent with typical population sizes for CA-containing soil microorganisms. Including accelerated soil hydration in global model simulations modifies contributions from soil and foliage to the global CO(18)O budget and eliminates persistent discrepancies existing between model and atmospheric observations. This enhanced soil hydration also increases the differences between the isotopic signatures of photosynthesis and respiration, particularly in the tropics, increasing the precision of CO(2) gross fluxes obtained by using the delta(18)O of atmospheric CO(2) by 50%.

Shifts in the Abundances of Saprotrophic and Ectomycorrhizal Fungi With Altered Leaf Litter Inputs.

Ectomycorrhizal (EcM) and saprotrophic fungi interact in the breakdown of organic matter, but the mechanisms underlying the EcM role on organic matter decomposition are not totally clear. We hypothesized that the ecological relations between EcM and saprotroph fungi are modulated by resources availability and accessibility, determining decomposition rates. We manipulated the amount of leaf litter inputs (No-Litter, Control Litter, Doubled Litter) on Trenched (root exclusion) and Non-Trenched plots (with roots) in a temperate deciduous forest of EcM-associated trees. Resultant shifts in soil fungal communities were determined by phospholipid fatty acids and DNA sequencing after 3 years, and CO2 fluxes were measured throughout this period. Different levels of leaf litter inputs generated a gradient of organic substrate availability and accessibility, altering the composition and ecological relations between EcM and saprotroph fungal communities. EcM fungi dominated at low levels of fresh organic substrates and lower organic matter quality, where short-distances exploration types seem to be better competitors, whereas saprotrophs and longer exploration types of EcM fungi tended to dominate at high levels of leaf litter inputs, where labile organic substrates were easily accessible. We were, however, not able to detect unequivocal signs of competition between these fungal groups for common resources. These results point to the relevance of substrate quality and availability as key factors determining the role of EcM and saprotroph fungi on litter and soil organic matter decay and represent a path forward on the capacity of organic matter decomposition of different exploration types of EcM fungi.

Borehole Equilibration: Testing a New Method to Monitor the Isotopic Composition of Tree Xylem Water in situ.

Forest water use has been difficult to quantify. One promising approach is to measure the isotopic composition of plant water, e.g., the transpired water vapor or xylem water. Because different water sources, e.g., groundwater versus shallow soil water, often show different isotopic signatures, isotopes can be used to investigate the depths from which plants take up their water and how this changes over time. Traditionally such measurements have relied on the extraction of wood samples, which provide limited time resolution at great expense, and risk possible artifacts. Utilizing a borehole drilled through a tree's stem, we propose a new method based on the notion that water vapor in a slow-moving airstream approaches isotopic equilibration with the much greater mass of liquid water in the xylem. We present two empirical data sets showing that the method can work in practice. We then present a theoretical model estimating equilibration times and exploring the limits at which the approach will fail. The method provides a simple, cheap, and accurate means of continuously estimating the isotopic composition of the source water for transpiration.

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'.

Altered energy partitioning across terrestrial ecosystems in the European drought year 2018.

Drought and heat events, such as the 2018 European drought, interact with the exchange of energy between the land surface and the atmosphere, potentially affecting albedo, sensible and latent heat fluxes, as well as CO2 exchange. Each of these quantities may aggravate or mitigate the drought, heat, their side effects on productivity, water scarcity and global warming. We used measurements of 56 eddy covariance sites across Europe to examine the response of fluxes to extreme drought prevailing most of the year 2018 and how the response differed across various ecosystem types (forests, grasslands, croplands and peatlands). Each component of the surface radiation and energy balance observed in 2018 was compared to available data per site during a reference period 2004-2017. Based on anomalies in precipitation and reference evapotranspiration, we classified 46 sites as drought affected. These received on average 9% more solar radiation and released 32% more sensible heat to the atmosphere compared to the mean of the reference period. In general, drought decreased net CO2 uptake by 17.8%, but did not significantly change net evapotranspiration. The response of these fluxes differed characteristically between ecosystems; in particular, the general increase in the evaporative index was strongest in peatlands and weakest in croplands. This article is part of the theme issue 'Impacts of the 2018 severe drought and heatwave in Europe: from site to continental scale'.

Water isotopes in desiccating lichens.

The stable isotopic composition of water is routinely used as a tracer to study water exchange processes in vascular plants and ecosystems. To date, no study has focussed on isotope processes in non-vascular, poikilohydric organisms such as lichens and bryophytes. To understand basic isotope exchange processes of non-vascular plants, thallus water isotopic composition was studied in various green-algal lichens exposed to desiccation. The study indicates that lichens equilibrate with the isotopic composition of surrounding water vapour. A model was developed as a proof of concept that accounts for the specific water relations of these poikilohydric organisms. The approach incorporates first their variable thallus water potential and second a compartmentation of the thallus water into two isotopically distinct but connected water pools. Moreover, the results represent first steps towards the development of poikilohydric organisms as a recorder of ambient vapour isotopic composition.

Effect of water availability on leaf water isotopic enrichment in beech seedlings shows limitations of current fractionation models.

Current models of leaf water enrichment predict that the differences between isotopic enrichment of water at the site of evaporation (Delta(e)) and mean lamina leaf water enrichment (Delta(L)) depend on transpiration rates (E), modulated by the scaled effective length (L) of water isotope movement in the leaf. However, variations in leaf parameters in response to changing environmental conditions might cause changes in the water path and thus L. We measured the diel course of Delta(L) for (18)O and (2)H in beech seedlings under well-watered and water-limited conditions. We applied evaporative enrichment models of increasing complexity to predict Delta(e) and Delta(L), and estimated L from model fits. Water-limited plants showed moderate drought stress, with lower stomatal conductance, E and stem water potential than the control. Despite having double E, the divergence between Delta(e) and Delta(L) was lower in well-watered than in water-limited plants, and thus, L should have changed to counteract differences in E. Indeed, L was about threefold higher in water-limited plants, regardless of the models used. We conclude that L changes with plant water status far beyond the variations explained by water content and other measured variables, thus limiting the use of current evaporative models under changing environmental conditions.

Robust Response of Terrestrial Plants to Rising CO2.

Human-caused CO2 emissions over the past century have caused the climate of the Earth to warm and have directly impacted on the functioning of terrestrial plants. We examine the global response of terrestrial gross primary production (GPP) to the historic change in atmospheric CO2. The GPP of the terrestrial biosphere has increased steadily, keeping pace remarkably in proportion to the rise in atmospheric CO2. Water-use efficiency, namely the ratio of CO2 uptake by photosynthesis to water loss by transpiration, has increased as a direct leaf-level effect of rising CO2. This has allowed an increase in global leaf area, which has conspired with stimulation of photosynthesis per unit leaf area to produce a maximal response of the terrestrial biosphere to rising atmospheric CO2 and contemporary climate change.

Air temperature optima of vegetation productivity across global biomes.

The global distribution of the optimum air temperature for ecosystem-level gross primary productivity ([Formula: see text]) is poorly understood, despite its importance for ecosystem carbon uptake under future warming. We provide empirical evidence for the existence of such an optimum, using measurements of in situ eddy covariance and satellite-derived proxies, and report its global distribution. [Formula: see text] is consistently lower than the physiological optimum temperature of leaf-level photosynthetic capacity, which typically exceeds 30 °C. The global average [Formula: see text] is estimated to be 23 ± 6 °C, with warmer regions having higher [Formula: see text] values than colder regions. In tropical forests in particular, [Formula: see text] is close to growing-season air temperature and is projected to fall below it under all scenarios of future climate, suggesting a limited safe operating space for these ecosystems under future warming.

Quantifying differences in water and carbon cycling between paddy and rainfed rice (Oryza sativa L.) by flux partitioning.

Agricultural crops play an important role in the global carbon and water cycle. Global climate change scenarios predict enhanced water scarcity and altered precipitation pattern in many parts of the world. Hence, a mechanistic understanding of water fluxes, productivity and water use efficiency of cultivated crops is of major importance, i.e. to adapt management practices. We compared water and carbon fluxes of paddy and rainfed rice by canopy scale gas exchange measurements, crop growth, daily evapotranspiration, transpiration and carbon flux modeling. Throughout a monsoon rice growing season, soil evaporation in paddy rice contributed strongly to evapotranspiration (96.6% to 43.3% from initial growth to fully developed canopy and amounted to 57.9% of total water losses over the growing seasons. Evaporation of rainfed rice was significantly lower (by 65% on average) particularly before canopy closure. Water use efficiency (WUE) was significantly higher in rainfed rice both from an agronomic (WUEagro, i.e. grain yield per evapotranspiration) and ecosystem (WUEeco, i.e. gross primary production per evapotranspiration) perspective. However, our results also show that higher WUE in rainfed rice comes at the expense of higher respiration losses compared to paddy rice (26% higher on average). Hence, suggestions on water management depend on the regional water availability (i.e. Mediterranean vs. Monsoon climate) and the balance between higher respiratory losses versus a potential reduction in CH4 and other greenhouse gas emissions. Our results suggest that a shift from rainfed/unsaturated soil to waterlogged paddy conditions after closure of the rice canopy might be a good compromise towards a sustainable use of water while preserving grain yield, particularly for water-limited production areas.