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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.

We investigate the effects of an isolated meso-γ-scale surface heterogeneity for roughness and albedo on the atmospheric boundary-layer (ABL) height, with a case study at a semi-arid forest surrounded by sparse shrubland (forest area: 28km2, forest length in the main wind direction: 7 km). Doppler lidar and ceilometer measurements at this semi-arid forest show an increase in the ABL height over the forest compared with the shrubland on four out of eight days. The differences in the ABL height between shrubland and forest are explained for all days with a model that assumes a linear growth of the internal boundary layer of the forest through the convective ABL upwind of the forest followed by a square-root growth into the stable free atmosphere. For the environmental conditions that existed during our measurements, the increase in ABL height due to large sensible heat fluxes from the forest (600Wm-2 in summer) is subdued by stable stratification in the free atmosphere above the ABL, or reduced by high wind speeds in the mixed layer.

For the past decade, observations of carbonyl sulfide (OCS or COS) have been investigated as a proxy for carbon uptake by plants. OCS is destroyed by enzymes that interact with CO2 during photosynthesis, namely carbonic anhydrase (CA) and RuBisCO, where CA is the more important one. The majority of sources of OCS to the atmosphere are geographically separated from this large plant sink, whereas the sources and sinks of CO2 are co-located in ecosystems. The drawdown of OCS can therefore be related to the uptake of CO2 without the added complication of co-located emissions comparable in magnitude. Here we review the state of our understanding of the global OCS cycle and its applications to ecosystem carbon cycle science. OCS uptake is correlated well to plant carbon uptake, especially at the regional scale. OCS can be used in conjunction with other independent measures of ecosystem function, like solar-induced fluorescence and carbon and water isotope studies. More work needs to be done to generate global coverage for OCS observations and to link this powerful atmospheric tracer to systems where fundamental questions concerning the carbon and water cycle remain.

Abstract. For the past decade, observations of carbonyl sulfide (OCS or COS) have been investigated as a proxy for carbon uptake by plants. OCS is destroyed by enzymes that interact with CO2 during photosynthesis, namely carbonic anhydrase (CA) and RuBisCO, where CA is the more important. The majority of sources of OCS to the atmosphere are geographically separated from this large plant sink, whereas the sources and sinks of CO2 are co-located in ecosystems. The drawdown of OCS can therefore be related to the uptake of CO2 without the added complication of co-located emissions comparable in magnitude. Here we review the state of our understanding of the global OCS cycle and its applications to ecosystem carbon cycle science. OCS uptake is correlated well to plant carbon uptake, especially at the regional scale. OCS can be used in conjunction with other independent measures of ecosystem function, like solar-induced fluorescence and carbon and water isotope studies. More work needs to be done to generate global coverage for OCS observations and to link this powerful atmospheric tracer to systems where fundamental questions concerning the carbon and water cycle remain.

Current climate models disagree on how much carbon dioxide land ecosystems take up for photosynthesis. Tracking the stronger carbonyl sulfide signal could help.

Drought stress causes multiple feedback responses in plants. These responses span from stomata closure and enzymatic downregulation of photosynthetic activity to structural adjustments of xylem biomass and leaf area. Some of these processes are not easily reversible and may persist long after the stress has ended. Despite a multitude of hydraulic model approaches, simulation models still widely lack an integrative mechanistic description of how this sequence of physiological to structural tree responses may be realized that is also simple enough to be generally applicable. Here, we suggest an integrative, sequential approach to simulate drought stress responses. First, decreasing plant water potential triggers stomatal closure alongside a downregulation of photosynthetic performance, thereby effectively slowing down further desiccation. A second protective mechanism is introduced by increasing the soil–root resistance, represented by a disconnection of fine roots after a threshold soil water potential has been reached. Further decreases in plant water potential due to residual transpiration and loss of internal stem water storage consistently lead to a loss of hydraulic functioning, which is reflected in sapwood loss and foliage senescence. This new model functionality has been used to investigate the responses of tree hydraulics, carbon uptake, and transpiration to soil and atmospheric drought in an extremely dry Aleppo pine (Pinus halepensis Mill.) plantation. Using the hypothesis of a sequential triggering of stress-mitigating responses, the model was able to reflect carbon uptake and transpiration patterns under varying soil water supply and atmospheric demand conditions – especially during summer – and respond realistically regarding medium-term responses, such as leaf and sapwood senescence. We could show that the observed avoidance strategy was only achieved when the model accounted for very early photosynthesis downregulation, and the relatively high measured plant water potentials were well reproduced with a root–soil disconnection strategy that started before major xylem conductance losses occurred. Residual canopy conductance was found to be pivotal in explaining dehydration and transpiration patterns during summer, but it also disclosed the fact that explaining the water balance in the driest periods requires water supply from stem water and deep soil layers. In agreement with the high drought resistance observed at the site, our model indicated little loss of hydraulic functioning in Aleppo pine, despite the intensive seasonal summer drought. [ABSTRACT FROM AUTHOR]

Abstract Background Net primary productivity (NPP) in forests plays an important role in the global carbon cycle. However, it is not well known about the increase rate of China’s forest NPP, and there are different opinions about the key factors controlling the variability of forest NPP. Methods This paper established a statistics-based multiple regression model to estimate forest NPP, using the observed NPP, meteorological and remote sensing data in five major forest ecosystems. The fluctuation values of NPP and environment variables were extracted to identify the key variables influencing the variation of forest NPP by correlation analysis. Results The long-term trends and annual fluctuations of forest NPP between 2000 and 2018 were examined. The results showed a significant increase in forest NPP for all five forest ecosystems, with an average rise of 5.2 gC·m− 2·year− 1 over China. Over 90% of the forest area had an increasing NPP range of 0–161 gC·m− 2·year− 1. Forest NPP had an interannual fluctuation of 50–269 gC·m− 2·year− 1 for the five major forest ecosystems. The evergreen broadleaf forest had the largest fluctuation. The variability in forest NPP was caused mainly by variations in precipitation, then by temperature fluctuations. Conclusions All five forest ecosystems in China exhibited a significant increasing NPP along with annual fluctuations evidently during 2000–2018. The variations in China’s forest NPP were controlled mainly by changes in precipitation.

Carbonyl sulfide (COS) is an atmospheric trace gas that participates in some key reactions of the carbon cycle and thus holds great promise for studies of carbon cycle processes. Global monitoring networks and atmospheric sampling programs provide concurrent data on COS and CO2 concentrations in the free troposphere and atmospheric boundary layer over vegetated areas. Here we present a modeling framework for interpreting these data and illustrate what COS measurements might tell us about carbon cycle processes. We implemented mechanistic and empirical descriptions of leaf and soil COS uptake into a global carbon cycle model (SiB 3) to obtain new estimates of the COS land flux. We then introduced these revised boundary conditions to an atmospheric transport model (Parameterized Chemical Transport Model) to simulate the variations in the concentration of COS and CO2 in the global atmosphere. To balance the threefold increase in the global vegetation sink relative to the previous baseline estimate, we propose a new ocean COS source. Using a simple inversion approach, we optimized the latitudinal distribution of this ocean source and found that it is concentrated in the tropics. The new model is capable of reproducing the seasonal variation in atmospheric concentration at most background atmospheric sites. The model also reproduces the observed large vertical gradients in COS between the boundary layer and free troposphere. Using a simulation experiment, we demonstrate that comparing drawdown of CO 2 with COS could provide additional constraints on differential responses of photosynthesis and respiration to environmental forcing. The separation of these two distinct processes is essential to understand the carbon cycle components for improved prediction of future responses of the terrestrial biosphere to changing environmental conditions. Key PointsCarbonyl sulfide can help falsify carbon cycle modelsCarbonyl sulfide can aid separation of NPP into GPP and RespThe oceanic COS source is probably much larger than currently thought ©2013. American Geophysical Union. All Rights Reserved.

The ability of plants to adjust to the adverse effects of climate change is important for their survival and for their contribution to the global carbon cycle. This is particularly true in the Mediterranean region, which is among the regions that are most vulnerable to climate change. Here, we carried out a 2-year comparative ecophysiological study of ecosystem function in two similar Eastern Mediterranean forests of the same tree species (Pinus halepensis Mill.) under mild (Sani, Greece) and extreme (Yatir, Israel) climatic conditions. The partial effects of key environmental variables, including radiation, vapor pressure deficit, air temperature and soil moisture (R g, D , T and soil water content (SWC), respectively), on the ecosystems' CO2 and water vapor fluxes were estimated using generalized additive models (GAMs). The results showed a large adjustment between sites in the seasonal patterns of both carbon and water fluxes and in the time and duration of the optimal period (defined here as the time when fluxes were within 85% of the seasonal maximum). The GAM analysis indicated that the main factor influencing the seasonal patterns was SWC, while T and D had significant but milder effects. During the respective optimal periods, the two ecosystems showed strong similarities in the fluxes' responses to the measured environmental variables, indicating similarity in their underlying physiological characteristics. The results indicate that Aleppo pine forests have a strong phenotypic adjustment potential to cope with increasing environmental stresses. This, in turn, will help their survival and their continued contribution to the terrestrial carbon sink in the face of climate change in this region. [ABSTRACT FROM AUTHOR]

Climate change is often associated with increasing vapour pressure deficit (VPD) and changes in soil moisture (SM). While atmospheric and soil drying often co‐occur, their differential effects on plant functioning and productivity remain uncertain. We investigated the divergent effects and underlying mechanisms of soil and atmospheric drought based on continuous, in situ measurements of branch gas exchange with automated chambers in a mature semiarid Aleppo pine forest. We investigated the response of control trees exposed to combined soil‒atmospheric drought (low SM, high VPD) during the rainless Mediterranean summer and that of trees experimentally unconstrained by soil dryness (high SM; using supplementary dry season water supply) but subjected to atmospheric drought (high VPD). During the seasonal dry period, branch conductance (gbr), transpiration rate (E) and net photosynthesis (Anet) decreased in low‐SM trees but greatly increased in high‐SM trees. The response of E and gbr to the massive rise in VPD (to 7 kPa) was negative in low‐SM trees and positive in high‐SM trees. These observations were consistent with predictions based on a simple plant hydraulic model showing the importance of plant water potential in the gbr and E response to VPD. These results demonstrate that avoiding drought on the supply side (SM) and relying on plant hydraulic regulation constrains the effects of atmospheric drought (VPD) as a stressor on canopy gas exchange in mature pine trees under field conditions. Summary statement: Under high evaporative demand conditions, mature Aleppo pine trees demonstrated increased transpiration and photosynthesis in response to supplementary irrigation. Stomatal conductance exhibited hypo‐sensitivity to increasing water demand (high vapour pressure deficit) but was sensitive to a decrease in water supply, expressed as soil water content or plant water potential. Our findings highlight that water supply, rather than water demand, is the primary limiting factor for transpiration in Aleppo pine trees. [ABSTRACT FROM AUTHOR]

The 18O content of atmospheric O2 is an important tracer for past changes in the biosphere. Its quantitative use depends on knowledge of the discrimination against 18O associated with the various O2 consumption processes. Here we evaluated, for the first time, the in situ 18O discrimination associated with soil respiration in natural ecosystems. The discrimination was estimated from the measured [O2] and δ18O of O2 in the soil-air. The discriminations that were found are 10.1 ± 1.5‰, 17.8 ± 1.0‰, and 22.5 ± 3.6‰, for tropical, temperate, and boreal forests, respectively, 17.9 ± 2.5‰ for Mediterranean woodland, and 15.4 ± 1.6‰ for tropical shrub land. Current understanding of the isotopic composition of atmospheric O2 is based on the assumption that the magnitude of the fractionation in soil respiration is identical to that of dark respiration through the cytochrome pathway alone (∼18‰). The discrimination we found in the tropical sites is significantly lower, and is explained by slow diffusion in soil aggregates and root tissues that limits the O2 concentration in the consumption sites. The high discrimination in the boreal sites may be the result of high engagement of the alternative oxidase pathway (AOX), which has high discrimination associated with it (∼27‰). The intermediate discrimination (∼18‰) in the temperate and Mediterranean sites can be explained by the opposing effects of AOX and diffusion limitation that cancel out. Since soil respiration is a major component of the global oxygen uptake, the contribution of large variations in the discrimination, observed here, to the global Dole Effect should be considered in global scale studies.

The modulation of the leaf energy budget components to maintain optimal leaf temperature are fundamental aspects of plant functioning and survival. Better understanding these aspects becomes increasingly important under a drying and warming climate when cooling through evapotranspiration (E) is suppressed. Combining novel measurements and theoretical estimates, we obtained unusually comprehensive twig‐scale leaf energy budgets under extreme field conditions in droughted (suppressed E) and non‐droughted (enhanced E) plots of a semi‐arid pine forest. Under the same high mid‐summer radiative load, leaf cooling shifted from relying on nearly equal contributions of sensible (H) and latent (LE) energy fluxes in non‐droughted trees to relying almost exclusively on H in droughted ones, with no change in leaf temperature. Relying on our detailed leaf energy budget, we could demonstrate that this is due to a 2× reduction in leaf aerodynamic resistance. This capability for LE‐to‐H shift in leaves of mature Aleppo pine trees under droughted field conditions without increasing leaf temperature is likely a critical factor in the resilience and relatively high productivity of this important Mediterranean tree species under drying conditions. Summary statement: Relying on detailed leaf energy budgets in mature pines, we demonstrate that leaves greatly enhance their sensible heat flux to prevent overheating under drought, which must rely on changes in leaf aerodynamic resistance. This may be a critical factor in the resilience of trees to drying conditions. [ABSTRACT FROM AUTHOR]

Drought stress is imposing multiple feedback responses in plants. These responses span from stomata closure and enzymatic downregulation of photosynthetic activity to structural adjustments in leaf area. Some of these processes are not easily reversible and may persist long after the stress ended. Unfortunately, simulation models widely lack an integrative mechanistic description on how this sequence of tree physiological to structural responses occur. Here, we suggest an integrative approach to simulate drought stress responses. Firstly, a decreasing plant water potential triggers stomatal closure alongside a downregulation of photosynthetic performance. This is followed by a disconnection of roots and soil and the reliance on internal stem water storage or water uptake from deep soil layers. Consistently, loss in hydraulic functioning is reflected in sapwood loss of functionality and foliage senescence. This new model functionality has been used to investigate responses of tree hydraulics, carbon uptake and transpiration to soil- and atmospheric drought in an extremely dry Aleppo pine (Pinus halepensis L.) plantation. Using the hypothesis of a sequential triggering of stress-mitigating responses, the model was able to reflect the carbon uptake and transpiration patterns under varying soil water supply and atmospheric demand - especially during summer - and responded realistically regarding medium-term responses such as leaf and sapwood senescence. In agreement with the high drought resistance observed at the site our model indicated little loss of hydraulic functioning in Aleppo pine, despite the intensive seasonal summer drought. [ABSTRACT FROM AUTHOR]