The importance of mesophyll conductance in regulating forest ecosystem productivity during drought periods

Water availability is the most limiting factor to global plant productivity, yet photosynthetic responses to seasonal drought cycles are poorly understood, with conflicting reports on which limiting process is the most important during drought. We address the problem using a model‐data synthesis approach to look at canopy level fluxes, integrating twenty years of half hour data gathered by the FLUXNET network across six Mediterranean sites. The measured canopy level, water and carbon fluxes were used, together with an inverse canopy ecophysiological model, to estimate the bulk canopy conductance, bulk mesophyll conductance, and the canopy scale carbon pools in both the intercellular spaces and at the site of carboxylation in the chloroplasts. Thus the roles of stomatal and mesophyll conductance in the regulation of internal carbon pools and photosynthesis could be separated. A quantitative limitation analysis allowed for the relative seasonal responses of stomatal, mesophyll, and biochemical limitations to be gauged. The concentration of carbon in the chloroplast was shown to be a potentially more reliable estimator of assimilation rates than the intercellular carbon concentration. Both stomatal conductance limitations and mesophyll conductance limitations were observed to regulate the response of photosynthesis to water stress in each of the six species studied. The results suggest that mesophyll conductance could bridge the gap between conflicting reports on plant responses to soil water stress, and that the inclusion of mesophyll conductance in biosphere–atmosphere transfer models may improve their performance, in particular their ability to accurately capture the response of terrestrial vegetation productivity to drought.     

Regional atmospheric CO2 inversion reveals seasonal and geographic differences in Amazon net biome exchange

Understanding tropical rainforest carbon exchange and its response to heat and drought is critical for quantifying the effects of climate change on tropical ecosystems, including global climate–carbon feedbacks. Of particular importance for the global carbon budget is net biome exchange of CO₂ with the atmosphere (NBE), which represents nonfire carbon fluxes into and out of biomass and soils. Subannual and sub‐Basin Amazon NBE estimates have relied heavily on process‐based biosphere models, despite lack of model agreement with plot‐scale observations. We present a new analysis of airborne measurements that reveals monthly, regional‐scale (~1–8 × 10⁶ km²) NBE variations. We develop a regional atmospheric CO₂ inversion that provides the first analysis of geographic and temporal variability in Amazon biosphere–atmosphere carbon exchange and that is minimally influenced by biosphere model‐based first guesses of seasonal and annual mean fluxes. We find little evidence for a clear seasonal cycle in Amazon NBE but do find NBE sensitivity to aberrations from long‐term mean climate. In particular, we observe increased NBE (more carbon emitted to the atmosphere) associated with heat and drought in 2010, and correlations between wet season NBE and precipitation (negative correlation) and temperature (positive correlation). In the eastern Amazon, pulses of increased NBE persisted through 2011, suggesting legacy effects of 2010 heat and drought. We also identify regional differences in postdrought NBE that appear related to long‐term water availability. We examine satellite proxies and find evidence for higher gross primary productivity (GPP) during a pulse of increased carbon uptake in 2011, and lower GPP during a period of increased NBE in the 2010 dry season drought, but links between GPP and NBE changes are not conclusive. These results provide novel evidence of NBE sensitivity to short‐term temperature and moisture extremes in the Amazon, where monthly and sub‐Basin estimates have not been previously available.     

Plant water uptake modelling: added value of cross-disciplinary approaches

In recent years, research interest in plant water uptake strategies has rapidly increased in many disciplines, such as hydrology, plant ecology and ecophysiology. Quantitative modelling approaches to estimate plant water uptake and spatiotemporal dynamics have significantly advanced through different disciplines across scales. Despite this progress, major limitations, for example, predicting plant water uptake under drought or drought impact at large scales, remain. These are less attributed to limitations in process understanding, but rather to a lack of implementation of cross-disciplinary insights into plant water uptake model structure. The main goal of this review is to highlight how the four dominant model approaches, that is, Feddes approach, hydrodynamic approach, optimality and statistical approaches, can be and have been used to create interdisciplinary hybrid models enabling a holistic system understanding that, among other things, embeds plant water uptake plasticity into a broader conceptual view of soil–plant feedbacks of water, nutrient and carbon cycling, or reflects observed drought responses of plant–soil feedbacks and their dynamics under, that is, drought. Specifically, we provide examples of how integration of Bayesian and hydrodynamic approaches might overcome challenges in interpreting plant water uptake related to different travel and residence times of different plant water sources or trade-offs between root system optimization to forage for water and nutrients during different seasons and phenological stages.     

Ecosystem‐scale volatile organic compound fluxes during an extreme drought in a broadleaf temperate forest of the Missouri Ozarks (central USA)

Considerable amounts and varieties of biogenic volatile organic compounds (BVOCs) are exchanged between vegetation and the surrounding air. These BVOCs play key ecological and atmospheric roles that must be adequately represented for accurately modeling the coupled biosphere–atmosphere–climate earth system. One key uncertainty in existing models is the response of BVOC fluxes to an important global change process: drought. We describe the diurnal and seasonal variation in isoprene, monoterpene, and methanol fluxes from a temperate forest ecosystem before, during, and after an extreme 2012 drought event in the Ozark region of the central USA. BVOC fluxes were dominated by isoprene, which attained high emission rates of up to 35.4 mg m^?2 h^?1 at midday. Methanol fluxes were characterized by net deposition in the morning, changing to a net emission flux through the rest of the daylight hours. Net flux of CO₂ reached its seasonal maximum approximately a month earlier than isoprenoid fluxes, which highlights the differential response of photosynthesis and isoprenoid emissions to progressing drought conditions. Nevertheless, both processes were strongly suppressed under extreme drought, although isoprene fluxes remained relatively high compared to reported fluxes from other ecosystems. Methanol exchange was less affected by drought throughout the season, confirming the complex processes driving biogenic methanol fluxes. The fraction of daytime (7–17 h) assimilated carbon released back to the atmosphere combining the three BVOCs measured was 2% of gross primary productivity (GPP) and 4.9% of net ecosystem exchange (NEE) on average for our whole measurement campaign, while exceeding 5% of GPP and 10% of NEE just before the strongest drought phase. The megan v2.1 model correctly predicted diurnal variations in fluxes driven mainly by light and temperature, although further research is needed to address model BVOC fluxes during drought events.     

Carbon and water fluxes in ecologically vulnerable areas in China

Ecologic vulnerable areas (EVAs) are the regions where ecosystems are fragile and vulnerable to suffer from degradation with external disturbances, e.g. environmental changes and human activities (Feng et al. 2022; Wang et al. 2019). EVAs in China are widely distributed and account for more than 55% China’s land area (Ministry of Ecology and Environment of the People’s Republic of China 2008). The ecosystem in EVAs, chartered with low stability, weak resistance and high vulnerability, has been experiencing significant degradation owing to the impacts of global climate change and human activities (Bai et al. 2018; Chen et al. 2021; Yu et al. 2022). The EVAs in China are not only the most serious areas of environmental degradation, but also the most poverty-stricken regions (Wang et al. 2019). Harsh environmental condition (drought, low temperature and strong radiation) and limited resource supply (water, soil nutrients, etc.) constrain the vegetation productivity and ecosystem services of EVAs (Li et al. 2021). Climate change adds new challenges with warmer temperatures, changing rainfall regime and increasing frequency of extreme events (drought, heat wave, storms, etc.), which make it is more difficult to predict the changes of ecosystem processes and functions in future scenarios (Piao et al. 2020; Reid et al. 2014). Carbon and water fluxes are the core ecosystem processes, which is linked to diverse ecosystem services (Lian et al. 2021). Therefore, clarifying the variations and controls of ecosystem carbon and water fluxes is an effective approach to clarifying how ecosystem respond to global change in EVAs (Baldocchi 2020). As the only technique can directly measure the carbon, water and energy fluxes between vegetation and atmosphere, eddy covariance technique has been considered as a standard method for flux observations (Chen et al. 2020). By integrating long-term, eddy covariance measurements over time and space, researches are able to assess ecosystem metabolism at different time scales (hours to decades) (Forzieri et al. 2020; Han et al. 2020; Jung et al. 2017). Eddy covariance measurements also produce information on how ecosystem respond to the changes in climate, which is useful for assessing ecosystem carbon sequestration (Hu et al. 2018), water and energy balance (Forzieri et al. 2020), resource use efficiency (Liu et al. 2019) and ecosystem feedback to climate change (Huang et al. 2019; Piao et al. 2020; Yue et al. 2020). Long-term flux measurements are also vital for detecting the responses of ecosystem functions to extreme events, optimizing and validating models on regional and global scales (Baldocchi 2020). Combining with remote sensing and ecosystem modeling techniques, scientists can upscale and evaluate the functional relations between carbon and water fluxes with environmental variables at high resolution and across diverse spatial/temporal scales (Niu et al. 2017; Xia et al. 2020).     

Factors influencing carbon fixation and water use by mediterranean sclerophyll shrubs during summer drought

Summary Mediterranean sclerophyll shrubs respond to seasonal drought by adjusting the amount of leaf area exposed and by reducing gas exchange via stomatal closure mechanisms. The degree to which each of these modifications can influence plant carbon and water balances under typical mediterranean-type climate conditions is examined. Leaf area changes are assessed in the context of a canopy structure and light microclimate model. Shifts in physiological response are examined with a mechanistically-based model of C₃ leaf gas exchange that simulates progressive reduction of maximum photosynthesis and transpiration rates and increasingly strong midday stomatal closure over the course of drought. The results demonstrate that midday stomatal closure may effectively contribute to drought avoidance, increase water use efficiency, and strongly alter physiological efficiency in the conversion of intercepted light energy to photoproducts. Physiological adjustments lead to larger reductions in water use than occur when comparing leaf area index 3.5 to 1.5, extremes found for natural stands of sclerophyll shrubs in the California chaparral. Reductions in leaf area have the strongest effect on resource capture and use during non-water-stressed periods and the least effect under extreme drought conditions, while shifts in physiological response lead to large savings of water and efficient water use under extreme stress. An important model parameter termed GFAC (proportionality factor expressing the relation of conductance [g] to net photosynthesis rate) is utilized, which changes in response to the integrated water stress experimence of shrubs and alters the degree to which stomata may open for a given rate of carbon fixation. We attempt to interpret this parameter in terms of physiological mechanisms known to modify control of leaf gas exchange during drought. The analysis helps visualize means by which canopy gas exchange behavior may be coupled to physiological changes occurring in the root environment during soil drying.     

Stability of above-ground and below-ground processes to extreme drought in model grassland ecosystems: Interactions with plant species diversity and soil nitrogen availability

Extreme drought events have the potential to cause dramatic changes in ecosystem structure and function, but the controls upon ecosystem stability to drought remain poorly understood. Here we used model systems of two commonly occurring, temperate grassland communities to investigate the short-term interactive effects of a simulated 100-year summer drought event, soil nitrogen (N) availability and plant species diversity (low/high) on key ecosystem processes related to carbon (C) and N cycling. Whole ecosystem CO₂ fluxes and leaching losses were recorded during drought and post-rewetting. Litter decomposition and C/N stocks in vegetation, soil and soil microbes were assessed 4 weeks after the end of drought. Experimental drought caused strong reductions in ecosystem respiration and net ecosystem CO₂ exchange, but ecosystem fluxes recovered rapidly following rewetting irrespective of N and species diversity. As expected, root C stocks and litter decomposition were adversely affected by drought across all N and plant diversity treatments. In contrast, drought increased soil water retention, organic nutrient leaching losses and soil fertility. Drought responses of above-ground vegetation C stocks varied depending on plant diversity, with greater stability of above-ground vegetation C to drought in the high versus low diversity treatment. This positive effect of high plant diversity on above-ground vegetation C stability coincided with a decrease in the stability of microbial biomass C. Unlike species diversity, soil N availability had limited effects on the stability of ecosystem processes to extreme drought. Overall, our findings indicate that extreme drought events promote post-drought soil nutrient retention and soil fertility, with cascading effects on ecosystem C fixation rates. Data on above-ground ecosystem processes underline the importance of species diversity for grassland function in a changing environment. Furthermore, our results suggest that plant–soil interactions play a key role for the short-term stability of above-ground vegetation C storage to extreme drought events.     

Forest biogeochemistry in response to drought

Trees alter their use and allocation of nutrients in response to drought, and changes in soil nutrient cycling and trace gas flux (N₂O and CH₄) are observed when experimental drought is imposed on forests. In extreme droughts, trees are increasingly susceptible to attack by pests and pathogens, which can lead to major changes in nutrient flux to the soil. Extreme droughts often lead to more common and more intense forest fires, causing dramatic changes in the nutrient storage and loss from forest ecosystems. Changes in the future manifestation of drought will affect carbon uptake and storage in forests, leading to feedbacks to the Earth's climate system. We must improve the recognition of drought in nature, our ability to manage our forests in the face of drought, and the parameterization of drought in earth system models for improved predictions of carbon uptake and storage in the world's forests.     

A big‐microsite framework for soil carbon modeling

Soil carbon cycling processes potentially play a large role in biotic feedbacks to climate change, but little agreement exists at present on what the core of numerical soil C cycling models should look like. In contrast, most canopy models of photosynthesis and leaf gas exchange share a common ‘Farquhaur‐model’ core structure. Here, we explore why a similar core model structure for heterotrophic soil respiration remains elusive and how a pathway to that goal might be envisioned. The spatial and temporal variation in soil microsite conditions greatly complicates modeling efforts, but we believe it is possible to develop a tractable number of parameterizable equations that are organized into a coherent, modular, numerical model structure. First, we show parallels in insights gleaned from linking Arrhenius and Michaelis–Menten kinetics for both photosynthesis and soil respiration. Additional equations and layers of complexity are then added to simulate substrate supply. For soils, model modules that simulate carbon stabilization processes will be key to estimating the fraction of soil C that is accessible to enzymes. Potential modules for dynamic photosynthate input, wetting‐event inputs, freeze–thaw impacts on substrate diffusion, aggregate turnover, soluble‐C sorption, gas transport, methane respiration, and microbial dynamics are described for conceptually and numerically linking our understanding of fast‐response processes of soil gas exchange with longer‐term dynamics of soil carbon and nitrogen stocks.     

Recent research progress on soil microbial responses to drying–rewetting cycles

Climatic change, such as increases in extreme drought and rainfall events and changes in rainfall intensity and pattern, has been strongly influencing soil moisture. The climatic change impact is particularly common in arid, semi-arid and Mediterranean regions, which is causing dramatic changes in the intensity and frequency of soil drying–rewetting cycles. The soil drying–rewetting cycle is a natural phenomenon that the soil experiences drying, then wetting, and then drying and rewetting again and again. When a dry soil is being rewetted, the amount of soil microbial biomass and its activity can be sharply increasing in a short time period, and then a large amount of gaseous carbon (C) and nitrogen (N) erupts from the soil. The sudden release of gaseous C and N is caused by the stimulation of the soil microbes. Such a phenomenon is called “Birch effect”. The drying–rewetting cycles have direct and indirect effects on soil microbes, and soil microbial responses to the drying and rewetting events play an important role in the feedbacks of terrestrial ecosystems. From aspects of soil microbial biomass, microbial activities and microbial structure, we review recent advances on studies regarding microbial responses to soil drying–rewetting cycles. We interpret the microbial responses using five different types of mechanisms: (1) Microbial stress mechanism: when a soil becomes dry, microorganisms must accumulate compatible solutes such as carbohydrates and aminoacids so that the soil microbes can equilibrate with their environment in order to avoid dehydrating and being killed. When the soil is rewetted, soil microbes must dispose of those osmolytes rapidly by transforming them into carbon dioxide (CO₂), dissolved organic carbon (DOC) and nutrients in order to prevent water from being flowing into the cells. (2) Substrate supply mechanism: low soil moisture may result in the physical disruption of soil aggregates which leads to the exposure of new soil surfaces and of previously protected organic matter. When the soil is rewetted, its physical structure is further disrupted by swelling. The increased new soil surfaces and previously protected organic matter will improve the microorganism’s nutrient availability. (3) Soil hydrophobicity mechanism: soil hydrophobicity can cause the reduction of soil moisture and nutrient availability and inhibition of microbial decomposition of soil organic matter. Therefore, soil hydrophobicity is an important factor of explaining the activity of microorganism in drying and rewetting events. (4) Diffusive limitations mechanism: transportation of the soil microbe is limited in a dry soil. When soil moisture is increasing, soil microbial activity is enhanced along with the increased availability of substrate nutrients. (5) Predation mechanism: a moist soil is usually conducive to the increase of bacteria and fungi populations. In response, protozoa and nematodes also increase, leading to the fluctuation of the soil microbial community structure. On the basis of the literature review, we propose five important aspects to be considered in the future: (1) assessing soil microbes’ concrete adapting ways to the drying–rewetting cycles, (2) evaluating the microbial responses to the drying–rewetting cycles based on suitable indicators, (3) interpreting microbial responses to the drying–rewetting cycles by combining field investigation and laboratory controlling experiment, (4) investigating the microbial responses to the drying–rewetting cycles at different temporal and spatial scales.     

Integrating spatial structure and interspecific and intraspecific plant–soil feedback effects and responses into community structure

Plant–soil feedbacks have important effects on plant communities, but most theory has been derived from experiments on intraspecific plant–soil feedbacks. Much less is known about how interspecific plant–soil feedbacks affect coexistence and plant communities, due in part to experimental and analytical challenges. Here, we propose a framework for evaluating plant–soil feedbacks among multiple interacting species that incorporates 1) the average effect each species has on conspecific and heterospecific neighbors via how they modify soil biota, 2) the average response of each species to the soil modifications made by neighboring species, and 3) intraspecific feedback. We refer to this as the ‘effect–response–intraspecific’ (ERI) model. We used individual‐based models to evaluate the relative importance of intraspecific and interspecific soil feedback in determining species abundance ranks in simulated plant communities. To compare the heuristic value of the ERI model to that of an established model in which effects and responses to soil feedback are not explicitly recognized, we evaluated a ‘full‐factorial’ model in which soil feedbacks among five plant species were measured and then explicitly modeled. The ERI model indicated that the response to interspecific plant–soil feedbacks was the key factor for species’ abundance rank without spatial structure. In contrast, interspecific plant–soil feedback had no impact on species abundance with spatial structure, and intraspecific feedback became dominant. Thus, our models predict that the relative importance of intraspecific and interspecific feedbacks changes as a function of the degree of spatial structure in a system. Overall, the ERI model provides a novel and tractable framework for evaluating complex multi‐species plant–soil feedbacks.     

Feedback significantly influences the simulated effect of CO2 on seasonal evapotranspiration from two agricultural species

The direct effect of elevated carbon dioxide on evapotranspiration over a growing season was investigated by scaling up single-leaf gas exchange measurements on soybean and corn plants grown and measured at three carbon dioxide concentrations. Stomatal conductance decreased markedly with increasing carbon dioxide in these species under most conditions. Coupled soil–vegetation–atmosphere models were used to scale up these single-leaf level measurements to simulate evapotranspiration at the regional scale from planting to harvest. The coupled modelling system introduced feedbacks over the season that are not present at the measurement level, which decreased the effect of carbon dioxide on evapotranspiration. Four sets of simulations were performed to evaluate specifically the magnitude of four feedbacks; two resulting from scale, surface layer and mixed layer feedback, one resulting from soil evaporation and one resulting from the interactions of stomatal conductance and the simulated canopy microclimate (physiological feedback). The feedbacks occurring from scale were consistent with previous analytical work indicating that transpiration becomes less dependent on stomatal conductance at larger scales. Evaporation from the soil has been generally neglected in past studies on carbon dioxide effects, but was especially important in decreasing the effects of carbon dioxide on evapotranspiration and showed a seasonal dynamic. The feedback resulting from physiological responses has also received less attention than the feedbacks from scale, but was only moderately important in these simulations. We also investigated the seasonal dynamics of how the observed increase in leaf area at elevated carbon dioxide affects evapotranspiration. Considering all the feedbacks and the observed increase in leaf area at elevated carbon dioxide, the simulated decrease in evapotranspiration was not negligible but was much less than the decrease in stomatal conductance. At the regional scale and maximum complexity in our model, the simulated decrease in seasonal evapotranspiration at doubled carbon dioxide (700 μmol mol^–1) was 5.4% for soybeans and 8.6% for corn.     

Forest understory plant and soil microbial response to an experimentally induced drought and heat‐pulse event: the importance of maintaining the continuum

Drought duration and intensity are expected to increase with global climate change. How changes in water availability and temperature affect the combined plant–soil–microorganism response remains uncertain. We excavated soil monoliths from a beech (Fagus sylvatica L.) forest, thus keeping the understory plant–microbe communities intact, imposed an extreme climate event, consisting of drought and/or a single heat‐pulse event, and followed microbial community dynamics over a time period of 28 days. During the treatment, we labeled the canopy with ¹³CO₂ with the goal of (i) determining the strength of plant–microbe carbon linkages under control, drought, heat and heat–drought treatments and (ii) characterizing microbial groups that are tightly linked to the plant–soil carbon continuum based on ¹³C‐labeled PLFAs. Additionally, we used 16S rRNA sequencing of bacteria from the Ah horizon to determine the short‐term changes in the active microbial community. The treatments did not sever within‐plant transport over the experiment, and carbon sinks belowground were still active. Based on the relative distribution of labeled carbon to roots and microbial PLFAs, we determined that soil microbes appear to have a stronger carbon sink strength during environmental stress. High‐throughput sequencing of the 16S rRNA revealed multiple trajectories in microbial community shifts within the different treatments. Heat in combination with drought had a clear negative effect on microbial diversity and resulted in a distinct shift in the microbial community structure that also corresponded to the lowest level of label found in the PLFAs. Hence, the strongest changes in microbial abundances occurred in the heat–drought treatment where plants were most severely affected. Our study suggests that many of the shifts in the microbial communities that we might expect from extreme environmental stress will result from the plant–soil–microbial dynamics rather than from direct effects of drought and heat on soil microbes alone.     

Not all droughts are created equal: the impacts of interannual drought pattern and magnitude on grassland carbon cycling

Climate extremes, such as drought, may have immediate and potentially prolonged effects on carbon cycling. Grasslands store approximately one‐third of all terrestrial carbon and may become carbon sources during droughts. However, the magnitude and duration of drought‐induced disruptions to the carbon cycle, as well as the mechanisms responsible, remain poorly understood. Over the next century, global climate models predict an increase in two types of drought: chronic but subtle ‘press‐droughts’, and shorter term but extreme ‘pulse‐droughts’. Much of our current understanding of the ecological impacts of drought comes from experimental rainfall manipulations. These studies have been highly valuable, but are often short term and rarely quantify carbon feedbacks. To address this knowledge gap, we used the Community Land Model 4.0 to examine the individual and interactive effects of pulse‐ and press‐droughts on carbon cycling in a mesic grassland of the US Great Plains. A series of modeling experiments were imposed by varying drought magnitude (precipitation amount) and interannual pattern (press‐ vs. pulse‐droughts) to examine the effects on carbon storage and cycling at annual to century timescales. We present three main findings. First, a single‐year pulse‐drought had immediate and prolonged effects on carbon storage due to differential sensitivities of ecosystem respiration and gross primary production. Second, short‐term pulse‐droughts caused greater carbon loss than chronic press‐droughts when total precipitation reductions over a 20‐year period were equivalent. Third, combining pulse‐ and press‐droughts had intermediate effects on carbon loss compared to the independent drought types, except at high drought levels. Overall, these results suggest that interannual drought pattern may be as important for carbon dynamics as drought magnitude and that extreme droughts may have long‐lasting carbon feedbacks in grassland ecosystems.     

The diel imprint of leaf metabolism on the δ13 C signal of soil respiration under control and drought conditions

Recent (13) CO(2) canopy pulse chase labeling studies revealed that photosynthesis influences the carbon isotopic composition of soil respired CO(2) (δ(13) C(SR)) even on a diel timescale. However, the driving mechanisms underlying these short-term responses remain unclear, in particular under drought conditions. The gas exchange of CO(2) isotopes of canopy and soil was monitored in drought/nondrought-stressed beech (Fagus sylvatica) saplings after (13) CO(2) canopy pulse labeling. A combined canopy/soil chamber system with gas-tight separated soil and canopy compartments was coupled to a laser spectrometer measuring mixing ratios and isotopic composition of CO(2) in air at high temporal resolution. The measured δ(13) C(SR) signal was then explained and substantiated by a mechanistic carbon allocation model. Leaf metabolism had a strong imprint on diel cycles in control plants, as a result of an alternating substrate supply switching between sugar and transient starch. By contrast, diel cycles in drought-stressed plants were determined by the relative contributions of autotrophic and heterotrophic respiration throughout the day. Drought reduced the speed of the link between photosynthesis and soil respiration by a factor of c. 2.5, depending on the photosynthetic rate. Drought slows the coupling between photosynthesis and soil respiration and alters the underlying mechanism causing diel variations of δ(13) C(SR).     

Carbon balance in the tundra, boreal forest and humid tropical forest during climate change: scaling up from leaf physiology and soil carbon dynamics

Carbon exchange by the terrestrial biosphere is thought to have changed since pre-industrial times in response to increasing concentrations of atmospheric CO₂ and variations (anomalies) in inter-annual air temperatures. However, the magnitude of this response, particularly that of various ecosystem types (biomes), is uncertain. Terrestrial carbon models can be used to estimate the direction and size of the terrestrial responses expected, providing that these models have a reasonable theoretical base. We formulated a general model of ecosystem carbon fluxes by linking a process-based canopy photosynthesis model to the Rothamsted soil carbon model for biomes that are not significantly affected by water limitation. The difference between net primary production (NPP) and heterotrophic soil respiration (R_h) represents net ecosystem production (NEP). The model includes (i) multiple compartments for carbon storage in vegetation and soil organic matter, (ii) the effects of seasonal changes in environmental parameters on annual NEP, and (iii) the effects of inter-annual temperature variations on annual NEP. Past, present and projected changes in atmospheric CO₂ concentration and surface air temperature (at different latitudes) were analysed for their effects on annual NEP in tundra, boreal forest and humid tropical forest biomes. In all three biomes, annual NEP was predicted to increase with CO₂ concentration but to decrease with warming. As CO₂ concentrations and temperatures rise, the positive carbon gains through increased NPP are often outweighed by losses through increased R_h, particularly at high latitudes where global warming has been (and is expected to be) most severe. We calculated that, several times during the past 140 years, both the tundra and boreal forest biomes have switched between being carbon sources (annual NEP negative) and being carbon sinks (annual NEP positive). Most recently, significant warming at high latitudes during 1988 and 1990 caused the tundra and boreal forests to be net carbon sources. Humid tropical forests generally have been a carbon sink since 1960. These modelled responses of the various biomes are in agreement with other estimates from either field measurements or geochemical models. Under projected CO₂ and temperature increases, the tundra and boreal forests will emit increasingly more carbon to the atmosphere while the humid tropical forest will continue to store carbon. Our analyses also indicate that the relative increase in the seasonal amplitude of the accumulated NEP within a year is about 0–14% year^?1 for boreal forests and 0–23% year^?1 in the tundra between 1960 and 1990.     

Nonlinear response of ecosystem respiration to multiple levels of temperature increases

Global warming exerts profound impacts on terrestrial carbon cycles and feedback to climates. Ecosystem respiration (ER) is one of the main components of biosphere CO₂ fluxes. However, knowledge regarding how ER responds to warming is still lacking. In this study, a manipulative experiment with five simulated temperature increases (+0℃ [Control], +2.1℃ [warming 1, W1], +2.7℃ [warming 2, W2], +3.2℃ [warming 3, W3], +3.9℃ [warming 4, W4]) was conducted to investigate ER responses to warming in an alpine meadow on the Tibetan Plateau. The results showed that ER was suppressed by warming both in dry and wet years. The responses of ER to warming all followed a nonlinear pattern. The nonlinear processes can be divided into three stages, the quick‐response stage (W1), stable stage (W1–W3), and transition stage (W4). Compared with the nonlinear model, the linear model maximally overestimated the response ratios of ER to warming 2.2% and 3.2% in 2015 and 2016, respectively, and maximally underestimated the ratio 7.0% and 2.7%. The annual differences in ER responding to warming were mainly attributed to the distinct seasonal distribution of precipitation. Specially, we found that the abrupt shift response of ER to warming under W4 treatment in 2015, which might be regulated by the excitatory effect of precipitation after long‐term drought in the mid‐growing season. This study highlights the importance of the nonlinearity of warming effects on ER, which should be taken into the global‐C‐cycling models for better predicting future carbon–climate feedbacks.     

Improved temperature response functions for models of Rubisco-limited photosynthesis

Predicting the environmental responses of leaf photosynthesis is central to many models of changes in the future global carbon cycle and terrestrial biosphere. The steady-state biochemical model of C₃ photosynthesis of Farquhar et al. (Planta 149, 78–90, 1980) provides a basis for these larger scale predictions; but a weakness in the application of the model as currently parameterized is the inability to accurately predict carbon assimilation at the range of temperatures over which significant photosynthesis occurs in the natural environment. The temperature functions used in this model have been based on in vitro measurements made over a limited temperature range and require several assumptions of in vivo conditions. Since photosynthetic rates are often Rubisco-limited (ribulose, 1-5 bisphosphate carboxylase/oxygenase) under natural steady-state conditions, inaccuracies in the functions predicting Rubisco kinetic properties at different temperatures may cause significant error. In this study, transgenic tobacco containing only 10% normal levels of Rubisco were used to measure Rubisco-limited photosynthesis over a large range of CO₂ concentrations. From the responses of the rate of CO₂ assimilation at a wide range of temperatures, and CO₂ and O₂ concentrations, the temperature functions of Rubisco kinetic properties were estimated in vivo. These differed substantially from previously published functions. These new functions were then used to predict photosynthesis in lemon and found to faithfully mimic the observed pattern of temperature response. There was also a close correspondence with published C₃ photosynthesis temperature responses. The results represent an improved ability to model leaf photosynthesis over a wide range of temperatures (10–40 °C) necessary for predicting carbon uptake by terrestrial C₃ systems.     

Drought: The most important physical stress of terrestrial ecosystems

Drought is projected to become more prevalent in the future due to climate change, and its impact on the fate of terrestrial ecosystems has aroused great concern in the scientific community over the past decade. Mounting evidence suggests that drought may be the most important physical stress of terrestrial ecosystems: drought limits vegetation growth, increases wildfires, and induces tree mortality, among other impacts. Drought not only weakens the carbon sink function of terrestrial ecosystems but also may interfere directly or indirectly with biosphere–atmosphere interactions, further exacerbating climate change. This paper reviews the current evidence of the impacts of drought on terrestrial ecosystems, with particular emphasis on the ways in which drought alters the biological, biogeophysical and biogeochemical processes underlying the interaction between the biosphere and the atmosphere.     

Aboveground net primary productivity and soil respiration display different responses to precipitation changes in desert grassland

荒漠草原区地上净初级生产力和土壤呼吸对降水变化的不同响应 降水变化既影响地上植被动态,也影响地下碳循环过程,尤其以干旱半干旱生态系统对降水的响应更为敏感。然而极端降水如何影响土壤碳固存潜力仍未得出明确结果。本研究在黄土高原西部荒漠草原样地实施了为期3年的降水控制实验,该实验包含5个降水梯度(即自然降水(对照),以及在自然降水基础上的减水40%、减水20%、增水20%、增水40%)。通过对不同降水处理下植物生长指标、地上净初级生产力(ANPP)、土壤水分和土壤呼吸(Rs)进行监测,采用双侧不对称模型揭示ANPP和Rs对降水变化的响应规律;采用结构方程模型,分析降水变化下影响ANPP和Rs的直接和间接因素。研究结果表明,ANPP对极端干旱的响应比极端湿润更敏感,在干旱和湿润年份均符合负向不对称模型。ANPP的变化主要受到降水的直接影响,同时,干旱年份植物密度的变化也对ANPP产生了影响。在湿润年份,Rs对降水变化的响应也呈负向不对称性。然而,干旱年份,Rs对降水变化表现出正向不对称响应,即对降水增加响应的敏感性高于降水减少,这可能与植物生长和ANPP对增水处理的正响应增加使自氧呼吸增强,及降水事件对异氧呼吸具有较强的‘Birch效应’有关。在干旱年份Rs对极端干旱(减水40%处理)表现出饱和响应。ANPP和Rs对降水格局改变的响应模式差异表明荒漠草原区极端湿润或干旱可能降低研究区土壤碳固存的潜力。