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Drained wetlands are thought to be carbon (C) source hotspots, and rewetting is advocated to restore C storage in drained wetlands for climate change mitigation. However, current assessments of wetland C balance mainly focus on vertical fluxes between the land and atmosphere, frequently neglecting lateral carbon fluxes and land-use effects. Here, we conduct a global synthesis of 893 annual net ecosystem C balance (NECB) measures that include net ecosystem exchange of CO 2 , along with C input via manure fertilization, and C removal through biomass harvest or hydrological exports of dissolved organic and inorganic carbon, across wetlands of different status and land uses. We find that elevating water table substantially reduces net ecosystem C losses, with the annual NECB decreasing from 2579 (95% interval: 1976 to 3214) kg C ha -1  year -1 in drained wetlands to -422 (-658 to -176) kg C ha -1  year -1 in natural wetlands, and to -934 (-1532 to -399) kg C ha -1  year -1 in rewetted wetlands globally. Climate, land-use history, and time since water table changes introduce variabilities, with drainage for (sub)tropical agriculture or forestry uses showing high annual C losses, while the net C losses from drained wetlands can continue to affect soil C pools for several decades. Rewetting all types of drained wetlands is needed, particularly for those formerly agriculture-used (sub)tropical wetlands where net ecosystem C losses can be largely reduced. Our findings suggest that elevating water table is an important initiative to reduce C losses in degraded wetlands, which could contribute to policy decisions for managing wetlands to enhance their C sequestration., (© 2024 John Wiley & Sons Ltd.)

Determination of tipping points in nitrogen (N) isotope (δ 15 N) natural abundance, especially soil δ 15 N, with increasing aridity, is critical for estimating N-cycling dynamics and N limitation in terrestrial ecosystems. However, whether there are linear or nonlinear responses of soil δ 15 N to increases in aridity and if these responses correspond well with soil N cycling remains largely unknown. In this study, we investigated soil δ 15 N and soil N-cycling characteristics in both topsoil and subsoil layers along a drought gradient across a 3000-km transect of drylands on the Qinghai-Tibetan Plateau. We found that the effect of increasing aridity on soil δ 15 N values shifted from negative to positive with thresholds at aridity index (AI) = 0.27 and 0.29 for the topsoil and subsoil, respectively, although soil N pools and N transformation rates linearly decreased with increasing aridity in both soil layers. Furthermore, we identified markedly different correlations between soil δ 15 N and soil N-cycling traits above and below the AI thresholds (0.27 and 0.29 for topsoil and subsoil, respectively). Specifically, in wetter regions, soil δ 15 N positively correlated with most soil N-cycling traits, suggesting that high soil δ 15 N may result from the "openness" of soil N cycling. Conversely, in drier regions, soil δ 15 N showed insignificant relationships with soil N-cycling traits and correlated well with factors, such as soil-available phosphorus and foliage δ 15 N, demonstrating that pathways other than typical soil N cycling may dominate soil δ 15 N under drier conditions. Overall, these results highlight that different ecosystem N-cycling processes may drive soil δ 15 N along the aridity gradient, broadening our understanding of N cycling as indicated by soil δ 15 N under changing drought regimes. The aridity threshold of soil δ 15 N should be considered in terrestrial N-cycling models when incorporating 15 N isotope signals to predict N cycling and availability under climatic dryness., (© 2024 John Wiley & Sons Ltd.)

Extreme drought is found to cause a threshold response in photosynthesis in ecosystem level. However, the mechanisms behind this phenomenon are not well understood, highlighting the importance of revealing the drought thresholds for multiple leaf-level photosynthetic processes. Thus, we conducted a long-term experiment involving precipitation reduction and nitrogen (N) addition. Moreover, an extreme drought event occurred within the experimental period. We found the presence of drought thresholds for multiple leaf-level photosynthetic processes, with the leaf light-saturated carbon assimilation rate (A sat ) displaying the highest threshold (10.76 v/v%) and the maximum rate of carboxylation by Rubisco (Vc max ) showing the lowest threshold (5.38 v/v%). Beyond the drought thresholds, the sensitivities of leaf-level photosynthetic processes to soil water content could be greater. Moreover, N addition lowered the drought thresholds of A sat and stomatal conductance (g s ), but had no effect on that of Vc max . Among species, plants with higher leaf K concentration traits had a lower drought threshold of A sat . Overall, this study highlights that leaf photosynthesis may be suppressed abruptly as soil water content surpasses the drought threshold. However, N enrichment helps to improve the resistance via delaying drought threshold response. These new findings have important implications for understanding the nonlinearity of ecosystem productivity response and early warning management in the scenario of combined extreme drought events and continuous N deposition., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2023 Elsevier B.V. All rights reserved.)

Grassland roots are fundamental to obtain the most limiting soil water and nitrogen (N) resources. However, this natural pattern could be significantly changed by recent co-occurrence of N deposition and extreme precipitations, likely with complex interactions on grassland root production and respiration. Despite this nonlinearity, we still know little about how extreme precipitation change nonlinearly regulates the responses of root respiration to N enrichment. Here, we conducted a 6-year experiment of N addition in an alpine meadow, coincidently experiencing extreme precipitations among experimental years. Our results demonstrated that root respiration showed divergent responses to N addition along with extreme precipitation changes among years. Under normal rainfall year, root respiration was significantly stimulated by N addition, whereas it was depressed under high or low water. Moreover, we revealed that both root biomass and traits (i.e. specific root length) were critical mechanisms in affecting root respiration response, but their relative importance changed with water condition. For example, specific root length and specific root respiration were more dominant than root biomass in determining root respiration response under low water, or vice versa. Overall, this study comprehensively reveals the nonlinearity of root respiration responses to the interactions of N enrichment and extreme water change. These new findings help to reconcile previously conflicting results that obtain in a specific episode of water gradient, with important implications for understanding grassland belowground carbon dynamics in facing combined N deposition and extreme precipitation events., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2023 Elsevier B.V. All rights reserved.)

Climate change is increasing the frequency and severity of short-term (~1 y) drought events-the most common duration of drought-globally. Yet the impact of this intensification of drought on ecosystem functioning remains poorly resolved. This is due in part to the widely disparate approaches ecologists have employed to study drought, variation in the severity and duration of drought studied, and differences among ecosystems in vegetation, edaphic and climatic attributes that can mediate drought impacts. To overcome these problems and better identify the factors that modulate drought responses, we used a coordinated distributed experiment to quantify the impact of short-term drought on grassland and shrubland ecosystems. With a standardized approach, we imposed ~a single year of drought at 100 sites on six continents. Here we show that loss of a foundational ecosystem function-aboveground net primary production (ANPP)-was 60% greater at sites that experienced statistically extreme drought (1-in-100-y event) vs. those sites where drought was nominal (historically more common) in magnitude (35% vs. 21%, respectively). This reduction in a key carbon cycle process with a single year of extreme drought greatly exceeds previously reported losses for grasslands and shrublands. Our global experiment also revealed high variability in drought response but that relative reductions in ANPP were greater in drier ecosystems and those with fewer plant species. Overall, our results demonstrate with unprecedented rigor that the global impacts of projected increases in drought severity have been significantly underestimated and that drier and less diverse sites are likely to be most vulnerable to extreme drought., Competing Interests: Competing interests statement:The authors declare no competing interest.

Ecosystem water use efficiency (WUE) is a coupled index of carbon (gross ecosystem productivity, GEP) and water fluxes (transpiration, Tr or evapotranspiration, ET), reflecting how ecosystem uses water efficiently to increase its carbon uptake. Though ecosystem WUE is generally considered to decrease with increasing precipitation levels, it remains elusive whether and how it nonlinearly responds to extreme water changes. Here, we performed a 5-year precipitation halving experiment in an alpine meadow, combined with extremely interannual precipitation fluctuations, to create a large range of soil water variations. Our results showed that WUE Tr and WUE ET consistently showed a quadratic pattern in response to soil water. Such quadratic patterns were steadily held at different stages of growing seasons, with minor changes in the optimal water thresholds (25.0-28.4 %). Below the water threshold, more soil water stimulated GEP but reduced Tr and ET by lowering soil temperature, resulting in a positive response of ecosystem WUE to soil water. Above the threshold, soil water stimulated GEP less than Tr (ET), leading to a negative response of ecosystem WUE to soil water. However, biological processes, including plant cover and belowground biomass as well as vertical root biomass distribution, had less effect on ecosystem WUE. Overall, this work is among the first to reveal the nonlinearity and optimal water thresholds of ecosystem WUE across a broad range of soil water, suggesting that future extreme precipitation events will more frequently surpass the water threshold and differently change the coupling relationships of carbon and water fluxes in alpine grasslands., Competing Interests: Declaration of competing interest The authors declare that they have no competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2023 Elsevier B.V. All rights reserved.)

Climate warming, often accompanied by extreme drought events, could have profound effects on both plant community structure and ecosystem functioning. However, how warming interacts with extreme drought to affect community- and ecosystem-level stability remains a largely open question. Using data from a manipulative experiment with three warming treatments in an alpine meadow that experienced one extreme drought event, we investigated how warming modulates resistance and recovery of community structural and ecosystem functional stability in facing with extreme drought. We found warming decreased resistance and recovery of aboveground net primary productivity (ANPP) and structural resistance but increased resistance and recovery of belowground net primary productivity (BNPP), overall net primary productivity (NPP), and structural recovery. The findings highlight the importance of jointly considering above- and belowground processes when evaluating ecosystem stability under global warming and extreme climate events. The stability of dominant species, rather than species richness and species asynchrony, was identified as a key predictor of ecosystem functional resistance and recovery, except for BNPP recovery. In addition, structural resistance of common species contributed strongly to the resistance changes in BNPP and NPP. Importantly, community structural resistance and recovery dominated the resistance and recovery of BNPP and NPP, but not for ANPP, suggesting the different mechanisms underlie the maintenance of stability of above- versus belowground productivity. This study is among the first to explain that warming modulates ecosystem stability in the face of extreme drought and lay stress on the need to investigate ecological stability at the community level for a more mechanistic understanding of ecosystem stability in response to climate extremes., (© 2023 The Ecological Society of America.)

Multiple global change drivers typically co-occur in terrestrial ecosystems, usually with complex interactions on ecosystem carbon fluxes. However, how they interactively impact terrestrial carbon sinks remains unknown. Here, we synthesized 82 field experiments that studied the individual and pairwise effects among nitrogen addition (N), increased precipitation (IP), elevated CO 2 (eCO 2 ) and warming, with direct measurements of net ecosystem productivity (NEP), gross ecosystem productivity (GEP) and ecosystem respiration (ER). We found that synergistic interactions mostly occurred between pairs of global change drivers on carbon fluxes. Moreover, these interactions varied with treatment magnitude, experimental duration and background precipitation. Specifically, the synergistic effect of N × IP became stronger with experimental precipitation magnitude and background rainfall. With an increasing N addition rate, N and eCO 2 had weaker interactive effects on NEP. Warming and IP were more synergic to enhance NEP with higher levels of warming magnitude. However, the interactive effects of N × eCO 2 on ER decreased over the experimental duration. Overall, this study provides new insights into the context-dependent occurrence of interactions among multiple global change drivers on ecosystem carbon sinks. These new findings are valuable to validate land C-cycle models with complex global change interactions and advance the next generations of future experimental design., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2023 Elsevier B.V. All rights reserved.)

Climate change leads to increasing temperature and more extreme hot and drought events. Ecosystem capability to cope with climate warming depends on vegetation's adjusting pace with temperature change. How environmental stresses impair such a vegetation pace has not been carefully investigated. Here we show that dryness substantially dampens vegetation pace in warm regions to adjust the optimal temperature of gross primary production (GPP) ( T opt GPP ) in response to change in temperature over space and time. T opt GPP spatially converges to an increase of 1.01°C (95% CI: 0.97, 1.05) per 1°C increase in the yearly maximum temperature (T max ) across humid or cold sites worldwide (37 o S-79 o N) but only 0.59°C (95% CI: 0.46, 0.74) per 1°C increase in T max across dry and warm sites. T opt GPP temporally changes by 0.81°C (95% CI: 0.75, 0.87) per 1°C interannual variation in T max at humid or cold sites and 0.42°C (95% CI: 0.17, 0.66) at dry and warm sites. Regardless of the water limitation, the maximum GPP (GPP max ) similarly increases by 0.23 g C m -2 day -1 per 1°C increase in T opt GPP in either humid or dry areas. Our results indicate that the future climate warming likely stimulates vegetation productivity more substantially in humid than water-limited regions., (© 2023 John Wiley & Sons Ltd.)

Extreme climate events, such as severe droughts and heavy rainfall, have profound impacts on the sustainable provision of ecosystem functions and services. However, how N enrichment interacts with discrete extreme climate events to affect ecosystem functions is largely unknown. Here, we investigated the responses of the temporal stability (i.e., resistance, recovery, and resilience) of aboveground net primary productivity (ANPP) in an alpine meadow to extreme dry and wet events under six N addition treatments (0, 2, 4, 8, 16, 32 g N m -2  year -1 ). We found that N addition had contrasting effects on the responses of ANPP to the extreme dry versus wet events, which resulted in no overall significant effects on ANPP stability across 2015-2019. Specifically, high N addition rates reduced the stability, resistance, and resilience of ANPP in response to extreme drought, whereas medium N addition rates increased ANPP stability and recovery in response to the extreme wet event. The main mechanisms underlying the response of ANPP to extreme drought and wet events were discrepant. Species richness, asynchrony, and dominant species resistance contributed most to the reduction of ANPP resistance to extreme drought, while species asynchrony and dominant and common species resilience contributed most to the decrease of ANPP resilience from extreme drought with N enrichment. The ANPP recovery from the extreme wet event was mainly explained by dominant and common species recovery. Our results provide strong evidence that N deposition mediates ecosystem stability in response to extreme dry and wet events in different ways and modulates the provisioning of grassland ecosystem functions under increasing extreme climate events., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2023 Elsevier B.V. All rights reserved.)

Carbon-use efficiency (CUE) has been widely used as a constant value in many earth system models to simulate how assimilated C is partitioned in ecosystems, to estimate ecosystem C budgets, and investigate C feedbacks to climate warming. Although correlative relationships from previous studies indicated that CUE could vary with temperature, and relying on a fixed CUE value could cause large uncertainty in model projections, however, due to the lack of manipulative experiment, it remains unclear how CUE at the plant (CUEp) and ecosystem (CUEe) levels respond to warming. Based on a 7-year manipulative warming experiment in an alpine meadow ecosystem on the Qinghai-Tibet Plateau, we quantitatively distinguished various C flux components of CUE, including gross ecosystem productivity, net primary productivity, net ecosystem productivity, ecosystem respiration, plant autotrophic respiration, and microbial heterotrophic respiration and explored how CUE at different levels responded to climate warming. We found large variations in both CUEp (0.60 to 0.77) and CUEe (from 0.38 to 0.59). The warming effect on CUEp was positively correlated with ambient soil water content (SWC) and the warming effect on CUEe was negatively correlated with ambient soil temperature (ST), but was positively correlated with warming-induced changes in ST. We also found that the direction and magnitude of the warming effects on different CUE components scaled differently with changes in the background environment, which explained the variation in CUE's warming response under environmental changes. Our new insights have important implications for reducing modelling uncertainty of ecosystem C budgets and improving our ability to predict ecosystem C-climate feedbacks under climate warming., Competing Interests: Declaration of competing interest The authors declare that they have no competing interests., (Copyright © 2023 Elsevier B.V. All rights reserved.)

Heavy grazing generally reduces grassland biomass, further decreasing its carbon sink. Grassland carbon sink is determined by both plant biomass and carbon sink per unit biomass (specific carbon sink). This specific carbon sink could reflect grassland adaptative response, because plants generally tend to adaptively enhance the functioning of their remaining biomass after grazing (i.e. higher leaf nitrogen content). Though we know well about the regulation of grassland biomass on carbon sink, little attention is paid to the role of specific carbon sink. Thus, we conducted a 14-year grazing experiment in a desert grassland. Ecosystem carbon fluxes, including net ecosystem CO 2 exchange (NEE), gross ecosystem productivity (GEP) and ecosystem respiration (ER), were measured frequently during five consecutive growing seasons with contrasting precipitation events. We found that heavy grazing reduced NEE more in drier (-94.0 %) than wetter (-33.9 %) years. However, grazing did not reduce community biomass much more in drier (-70.4 %) than wetter years (-66.0 %). These meant a positive response of specific NEE (NEE per unit biomass) to grazing in wetter years. This positive response of specific NEE was mainly caused by a higher biomass ratio of other species versus perennial grasses with greater leaf nitrogen content and specific leaf area in wetter years. In addition, we also detected a shift of grazing effects on specific NEE from positive in wetter years to negative in drier years. Overall, this study is among the first to reveal the adaptive response of grassland specific carbon sink to experimental grazing in plant trait view. The stimulation response of specific carbon sink can partially compensate for the loss of grassland carbon storage under grazing. These new findings highlight the role of grassland adaptive response in decelerating climate warming., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2023 Elsevier B.V. All rights reserved.)

Elucidating the effects underlying soil organic carbon (SOC) variation is imperative for ascertaining the potential drivers of mitigating climate change. However, the drivers of variations in various SOC fractions (e.g., macroaggregate C, microaggregate C, and silt and clay C) at different soil depths remain poorly understood. Here, we investigated the effects and relative contributions of climatic, plant, edaphic, and microbial factors on soil aggregate C between the topsoil (0-10 cm) and subsoil (20-30 cm) across alpine grasslands on the Tibetan Plateau. Results showed that the C content of macroaggregates, microaggregates, and silt and clay fractions in the topsoil was 128.6 %, 49.6 %, and 242.4 % higher than that in the subsoil, respectively. Overall, plant properties were the most determinants controlling soil macroaggregate, microaggregate, and silt + clay associated C for both two soil depths, accounting for 32.2 %, 37.4 %, and 38.8 % of the variation, respectively, followed by edaphic, microbial, and climatic factors. The aggregate C of both soil depths was significantly related with the climatic, plant, edaphic, and microbial factors, but the relative importance of these determinants was soil-depth dependent. Specifically, the effects of plant root biomass and microbial (e.g., microbial biomass carbon and fungal diversity index) factors on each aggregate C weakened with soil depth, but the importance of edaphic factors (e.g., clay content, pH, and bulk density) strengthened with soil depth, except for the weakened effect of bulk density on the microaggregate C. And the effects of climatic factor (e.g., mean annual precipitation) on macroaggregate and microaggregate C increased with soil depth. Our results highlight differential drivers and their impacts on soil aggregate C between the topsoil and subsoil, which benefits biogeochemical models for more accurately forecasting soil C dynamics and its feedbacks to environmental changes., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2023 Elsevier B.V. All rights reserved.)

Eutrophication generally promotes but destabilizes grassland productivity. Under eutrophication, plants tend to decrease biomass allocation to roots but increase aboveground allocation and light limitation, likely affecting community stability. However, it remains unclear to understand how shifting plant biomass allocation and light limitation regulate grassland stability in response to eutrophication. Here, using a 5-yr multiple nutrient addition experiment in an alpine meadow, we explored the role of changes in plant biomass allocation and light limitation on its community stability under eutrophication as well as traditionally established mechanisms (i.e., plant Shannon diversity, species asynchrony and grass subcommunity stability). Our results showed that nitrogen (N) addition, rather than phosphorus (P) or potassium (K) addition, significantly reduced the temporal stability of the alpine meadow. In accordance with previous studies, we found that N addition decreased plant Shannon diversity, species asynchrony and grass subcommunity stability, further destabilizing meadow community productivity. In addition, we also found the decrease in biomass allocation to belowground by N addition, further weakening its community stability. Moreover, this shifts in plant biomass allocation from below- to aboveground, intensifying plant light limitation. Further, the light limitation reduced plant species asynchrony, which finally weakened its community stability. Overall, in addition to traditionally established mechanisms, this study highlights the role of plant biomass allocation shifting from belowground to aboveground in determining grassland community stability. These "unseen" mechanisms might improve our understanding of grassland stability in the context of ongoing eutrophication., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2022 Elsevier B.V. All rights reserved.)

Plant and microbial diversity are key to determine ecosystem functioning. Despite the well-known role of local-scale α diversity in affecting vegetation biomass, the effects of community heterogeneity (β diversity) of plants and soil microbes on above- and belowground biomass (AGB and BGB) across contrasting environments still remain unclear. Here, we conducted a dryness-gradient transect survey over 3000 km across grasslands on the Tibetan Plateau. We found that plant β diversity was more dominant than α diversity in maintaining higher levels of AGB, while soil fungal β diversity was the key driver in enhancing BGB. However, these positive effects of plant and microbial β diversity on AGB and BGB were strongly weakened by increasing climatic dryness, mainly because higher soil available phosphorus caused by increasing dryness reduced both plant and soil fungal β diversities. Overall, these new findings highlight the critical role of above- and belowground β diversity in sustaining grassland biomass, raising our awareness to the ecological risks of large-scale biotic homogenization under future climate change., (© 2022 John Wiley & Sons Ltd.)

Epiphytic microbes on the surfaces of leaves and roots can bring substantial benefits or damages to their plant hosts. Although various factors have been proposed for shaping the epiphytic microbial composition, the contributions of environment factors, endogenous microbial taxa interactions, host plant traits, and their interactive effects are poorly understood. Here, we conducted a field investigation along a precipitation gradient and collected leaf and root surface microbes of two alpine plant species for 16S rRNA sequencing. We found that epiphytic bacterial community composition significantly changed along the precipitation gradient through ordination analyses and permutational multivariate analysis of variance. Beneficial bacterial taxa from Caulobacteraceae, Sphingomonadaceae, Comamonadaceae and Rhizobiales were enriched in the high precipitation zones. The stress-tolerant Hymenobacteraceae, Micrococcaceae, and Geodermatophilaceae occurred more frequently in the phyllosphere, and the Thermoleophilia, Thermomicrobiales and Bacillales were enriched in the rhizosphere at the drier sites. Mean annual precipitation was the most important factor regulating the epiphytic bacterial community composition. The direct effect of climate on bacterial community composition was higher in the phyllosphere than in the rhizosphere where joint effects of climate, plant traits and soil properties predominated. Distinct leaf trichome cover and plant height clearly explained the host effect on the phyllosphere bacterial community composition while belowground traits did not explain the host effect well on the rhizosphere bacterial community composition. We detected a significant role of bacterial taxa interactions in shaping microbial communities, where greater negative taxa interactions led to lesser composition changes. Structural equation modeling showed that environmental factors and bacterial interactions substantially contributed to the variation in epiphytic community composition, followed by host plant traits. This study advances our understanding of complex factors affecting alpine epiphytic community assembly and further confirms the role of biotic interactions., Competing Interests: Declaration of competing interest No conflict of interest exists in the submission of this manuscript, and the manuscript was approved by all authors for publication., (Copyright © 2022 Elsevier B.V. All rights reserved.)

Aims: Leaf chlorophyll (Chl) is a fundamental component and good proxy for plant photosynthesis. However, we know little about the large-scale patterns of leaf Chl and the relative roles of current environment changes vs. plant evolution in driving leaf Chl variations., Locations: The east to west grassland transect of the Tibetan Plateau., Methods: We performed a grassland transect over 1,600 km across the Tibetan Plateau, measuring leaf Chl among 677 site-species., Results: Leaf Chl showed a significantly spatial pattern across the grasslands in the Tibetan Plateau, decreasing with latitude but increasing with longitude. Along with environmental gradient, leaf Chl decreased with photosynthetically active radiation (PAR), but increased with water availability and soil nitrogen availability. Furthermore, leaf Chl also showed significant differences among functional groups (C 4 > C 3 species; legumes < non-legume species), but no difference between annual and perennial species. However, we surprisingly found that plant evolution played a dominant role in shaping leaf Chl variations when comparing the sum and individual effects of all the environmental factors above. Moreover, we revealed that leaf Chl non-linearly decreased with plant evolutionary divergence time. This well-matches the non-linearly increasing trend in PAR or decreasing trend in temperature during the geological time-scale uplift of the Tibetan Plateau., Main Conclusion: This study highlights the dominant role of plant evolution in determining leaf Chl variations across the Tibetan Plateau. Given the fundamental role of Chl for photosynthesis, these results provide new insights into reconsidering photosynthesis capacity in alpine plants and the carbon cycle in an evolutionary view., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2022 He, Li, Zhang, Wang, Zhu, Li, Chen, Pan, Shen, Wang, Li and Tian.)

Ecological restoration is essential to reverse land degradation worldwide. Most studies have assessed the restoration of ecosystem functions individually, as opposed to a holistic view. Here we developed a network-based ecosystem multifunctionality (EMF) framework to identify key functions in evaluating EMF restoration. Through synthesizing 293 restoration studies (2900 observations) following cropland abandonment, we found that individual soil functions played different roles in determining the restoration of belowground EMF. Soil carbon, total nitrogen, and phosphatase were key functions to predict the recovery of belowground EMF. On average, abandoned cropland recovered ~19% of EMF during 18 years. The restoration of EMF became larger with longer recovery time and higher humidity index, but lower with increasing soil depth and initial soil carbon. Overall, this study presents a network-based EMF framework, effectively helping to evaluate the success of ecosystem restoration and identify the key functions., (© 2022 The Ecological Society of America.)

The denitrification process profoundly affects soil nitrogen (N) availability and generates its byproduct, nitrous oxide, as a potent greenhouse gas. There are large uncertainties in predicting global denitrification because its controlling factors remain elusive. In this study, we compiled 4301 observations of denitrification rates across a variety of terrestrial ecosystems from 214 papers published in the literature. The averaged denitrification rate was 3516.3 ± 91.1 µg N kg -1  soil day -1 . The highest denitrification rate was 4242.3 ± 152.3 µg N kg -1  soil day -1 under humid subtropical climates, and the lowest was 965.8 ± 150.4 µg N kg -1 under dry climates. The denitrification rate increased with temperature, precipitation, soil carbon and N contents, as well as microbial biomass carbon and N, but decreased with soil clay contents. The variables related to soil N contents (e.g., nitrate, ammonium, and total N) explained the variation of denitrification more than climatic and edaphic variables (e.g., mean annual temperature (MAT), soil moisture, soil pH, and clay content) according to structural equation models. Soil microbial biomass carbon, which was influenced by soil nitrate, ammonium, and total N, also strongly influenced denitrification at a global scale. Collectively, soil N contents, microbial biomass, pH, texture, moisture, and MAT accounted for 60% of the variation in global denitrification rates. The findings suggest that soil N contents and microbial biomass are strong predictors of denitrification at the global scale., (© 2021 John Wiley & Sons Ltd.)

Forest thinning is a major forest management practice worldwide and may lead to profound alterations in the fluxes of soil greenhouse gases (GHGs). However, the global patterns and underlying mechanisms of soil GHG fluxes in response to forest thinning remain poorly understood. Here, we conducted a global meta-analysis of 106 studies to assess the effects of forest thinning on soil GHG fluxes and the underpinning mechanisms. The results showed that forest thinning significantly increased soil CO 2 emission (mean lnRR: 0.07, 95% CI: 0.03-0.11), N 2 O emission (mean lnRR: 0.39, 95% CI: 0.16-0.61) and decreased CH 4 uptake (mean Hedges' d: 0.98, 95% CI: 0.32-1.64). Furthermore, the negative response of soil CH 4 uptake was amplified by thinning intensity, and the positive response of soil N 2 O emission decreased with recovery time after thinning. The response of soil CO 2 emission was mainly correlated with changes in fine root biomass and soil nitrogen content, and the response of soil CH 4 uptake was related to the changes in soil moisture and litterfall. Moreover, the response of soil N 2 O emission was associated with changes in soil temperature and soil nitrate nitrogen content. Thinning also increased the total balance of the three greenhouse gas fluxes in combination, which decreased with recovery time. Our findings highlight that thinning significantly increases soil GHG emissions, which is crucial to understanding and predicting ecosystem-climate feedbacks in managed forests., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2021 Elsevier B.V. All rights reserved.)

Biodiversity drives ecosystem functioning across grassland ecosystems. However, few studies have examined how grazing intensity affects ecosystem multifunctionality (EMF) via its effects on plant diversity and soil microbial diversity in dry grasslands. We conducted a 12-year experiment manipulating sheep grazing intensity in a desert steppe of northern China. Through measuring plant species diversity, soil microbial diversity (bacteria diversity) and multiple ecosystem functions (i.e., aboveground net primary productivity, belowground biomass of plant community, temporal stability of ANPP, soil organic matter, moisture, available nitrogen and phosphorus, ecosystem respiration and gross ecosystem productivity), we aimed to understand how grazing intensity affected EMF via changing the diversity of plants and microbes. Our results showed that increasing grazing intensity significantly reduced EMF and most individual ecosystem functions, as well as the diversity of plants and microbes, while EMF and most individual functions were positively related to plant diversity and soil microbial diversity under all grazing intensities. In particular, soil microbial diversity in shallow soil layers (0-5 cm depth) had stronger positive correlations with plant diversity and EMF than in deeper soil layers. Furthermore, structural equation modeling (SEM) showed that grazing reduced EMF mainly via reducing plant diversity, rather than by reducing soil microbial diversity. Thus, plant diversity played a more important role in mediating the response of EMF to grazing disturbance. This study highlights the critical role of above- and belowground diversity in mediating the response of EMF to grazing intensity, which has important implications for biodiversity conservation and sustainability in arid grasslands., Competing Interests: Declaration of competing interest The authors declare that they have no conflict of interest., (Copyright © 2021 Elsevier B.V. All rights reserved.)

Whether and how warming alters functional traits of absorptive plant roots remains to be answered across the globe. Tackling this question is crucial to better understanding terrestrial responses to climate change as fine-root traits drive many ecosystem processes. We carried out a detailed synthesis of fine-root trait responses to experimental warming by performing a meta-analysis of 964 paired observations from 177 publications. Warming increased fine-root biomass, production, respiration and nitrogen concentration as well as decreased root carbon : nitrogen ratio and nonstructural carbohydrates. Warming effects on fine-root biomass decreased with greater warming magnitude, especially in short-term experiments. Furthermore, the positive effect of warming on fine-root biomass was strongest in deeper soil horizons and in colder and drier regions. Total fine-root length, morphology, mortality, life span and turnover were unresponsive to warming. Our results highlight the significant changes in fine-root traits in response to warming as well as the importance of warming magnitude and duration in understanding fine-root responses. These changes have strong implications for global soil carbon stocks in a warmer world associated with increased root-derived carbon inputs into deeper soil horizons and increases in fine-root respiration., (© 2021 The Authors New Phytologist © 2021 New Phytologist Foundation.)

It is well documented that warming can accelerate greenhouse gas (GHG) emissions, further inducing a positive feedback and reinforcing future climate warming. However, how different kinds of GHGs respond to various warming magnitudes remains largely unclear, especially in the cold regions that are more sensitive to climate warming. Here, we concurrently measured carbon dioxide (CO 2 ), methane (CH 4 ), and nitrous oxide (N 2 O) fluxes and their total balance in an alpine meadow in response to three levels of warming (ambient, +1.5 °C, +3.0 °C). We found warming-induced increases in CH 4 uptake, decreases in N 2 O emissions and increases in CO 2 emissions at the annual basis. Expressed as CO 2 -equivalents with a global warming potential of 100 years (GWP100), the enhancement of CH 4 uptake and reduction of N 2 O emissions offset only 9% of the warming-induced increase in CO 2 emissions for 1.5 °C warming, and only 7% for 3.0 °C warming. CO 2 emissions were strongly stimulated, leading to a significantly positive feedback to climate system, for 3.0 °C warming but less for 1.5 °C warming. The warming with 3.0 °C altered the total GHG balance mainly by stimulating CO 2 emissions in the non-growing season due to warmer soil temperatures, longer unfrozen period, and increased soil water content. The findings provide an empirical evidence that warming beyond global 2 °C target can trigger a positive GHG-climate feedback and highlight the contribution from non-growing season to this positive feedback loop in cold ecosystems., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2021 Elsevier B.V. All rights reserved.)

Nitrogen immobilization usually leads to nitrogen retention in soil and, thus, influences soil nitrogen supply for plant growth. Understanding soil nitrogen immobilization is important for predicting soil nitrogen cycling under anthropogenic activities and climate changes. However, the global patterns and drivers of soil nitrogen immobilization remain unclear. We synthesized 1350 observations of gross soil nitrogen immobilization rate (NIR) from 97 articles to identify patterns and drivers of NIR. The global mean NIR was 8.77 ± 1.01 mg N kg -1  soil day -1 . It was 5.55 ± 0.41 mg N kg -1  soil day -1 in croplands, 15.74 ± 3.02 mg N kg -1  soil day -1 in wetlands, and 15.26 ± 2.98 mg N kg -1  soil day -1 in forests. The NIR increased with mean annual temperature, precipitation, soil moisture, soil organic carbon, total nitrogen, dissolved organic nitrogen, ammonium, nitrate, phosphorus, and microbial biomass carbon. But it decreased with soil pH. The results of structural equation models showed that soil microbial biomass carbon was a pivotal driver of NIR, because temperature, total soil nitrogen, and soil pH mostly indirectly influenced NIR via changing soil microbial biomass. Moreover, microbial biomass carbon accounted for most of the variations in NIR among all direct relationships. Furthermore, the efficiency of transforming the immobilized nitrogen to microbial biomass nitrogen was lower in croplands than in natural ecosystems (i.e., forests, grasslands, and wetlands). These findings suggested that soil nitrogen retention may decrease under the land use change from forests or wetlands to croplands, but NIR was expected to increase due to increased microbial biomass under global warming. The identified patterns and drivers of soil nitrogen immobilization in this study are crucial to project the changes in soil nitrogen retention., (© 2021 John Wiley & Sons Ltd.)

Soil nitrification, an important pathway of nitrogen transformation in ecosystems, produces soil nitrate that influences net primary productivity, while the by-product of nitrification, nitrous oxide, is a significant greenhouse gas. Although there have been many studies addressing the microbiology, physiology, and impacting environment factors of soil nitrification at local scales, there are very few studies on soil nitrification rate over large scales. We conducted a global synthesis on the patterns and controlling factors of soil nitrification rate normalized at 25°C by compiling 3,140 observations from 186 published articles across terrestrial ecosystems. Soil nitrification rate tended to decrease with increasing latitude, especially in the Northern Hemisphere, and varied largely with ecosystem types. The soil nitrification rate significantly increased with mean annual temperature (MAT), soil nitrogen content, microbial biomass carbon and nitrogen, soil ammonium, and soil pH, but decreased with soil carbon:nitrogen and carbon:nitrogen of microbial biomass. The total soil nitrogen content contributed the most to the variations of global soil nitrification rate (total coefficient = 0.29) in structural equation models. The microbial biomass nitrogen (MBN; total coefficient = 0.19) was nearly of equivalent importance relative to MAT (total coefficient = 0.25) and soil pH (total coefficient = 0.24) in determining soil nitrification rate, while soil nitrogen and pH influenced soil nitrification via changing soil MBN. Moreover, the emission of soil nitrous oxide was positively related to soil nitrification rate at a global scale. This synthesis will advance our current understanding on the mechanisms underlying large-scale variations of soil nitrification and benefit the biogeochemical models in simulating global nitrogen cycling., (© 2020 John Wiley & Sons Ltd.)

Soil enzymes play critical roles in the decomposition of organic matter and determine the availability of soil nutrients, however, there are significant uncertainties in regard to how enzymatic responses to global warming. To reveal the general response patterns and controlling factors of various extracellular enzyme activities (EEA), we collected data from 78 peer-reviewed papers to investigate the responses of extracellular enzyme activities (EEA), including β-1,4-glucosidase (BG), β-d-cellobiosidase (CBH), β-1,4-xylosidase (XYL), leucine amino peptidase (LAP), N-acetyl-glucosaminidase (NAG), urease (URE), phosphatase (PHO), peroxidase (PER), phenol oxidase (POX), and polyphenol oxidase (PPO), to experimental warming. Our results showed that warming treatments increased soil temperature by 1.9 °C on average. The oxidative EEA, calculated as the sum of PER, POX and PPO, was on average stimulated by 9.4% under warming. However, the responses of C acquisition EEA (the sum of BG, CBH and XYL), N acquisition EEA (the sum of LAP, NAG and URE), and P acquisition EEA to warming had large variations across studies. The warming effects on C, N, P acquisition EEA and oxidative EEA tended to increase with soil warming magnitude and duration as well as the mean annual temperature. The response of C acquisition EEA to warming was positively correlated with fungal biomass, while that of P acquisition EEA had positive relationships with fungi: bacteria ratios. The response of oxidative EEA was negatively correlated with the abundance of gram-positive bacterial biomass. Our results suggested that warming consistently stimulated oxidative EEA, but had diverse effects on hydrolytic EEA, which were dependent on the warming magnitude or duration, or environmental factors. The observed relationships between changes in microbial traits and extracellular enzymes suggested that microbial compositions drive changes in enzyme decomposition under warming. Thus, incorporation of microbial modification in biogeochemistry models is essential to better predict ecosystem carbon and nutrient dynamics., (Copyright © 2018 Elsevier B.V. All rights reserved.)

It is a central ecological goal to explore the effects of global change factors on soil microbial communities. The vast functional gene repertoire of soil microbial communities is composed of both core and accessory genes, which may be governed by distinct drivers. This intuitive hypothesis, however, remains largely unexplored. We conducted a 5-year nitrogen and water addition experiment in the Eurasian steppe and quantified microbial gene diversity via shotgun metagenomics. Nitrogen addition led to an 11-fold increase in the abundance (based on quantitative PCR [qPCR]) of ammonia-oxidizing bacteria, which have mainly core community genes and few accessory community genes. Thus, nitrogen addition substantially increased the relative abundance of many core genes at the whole-community level. Water addition stimulated both plant diversity and microbial respiration; however, increased carbon/energy resources from plants did not counteract increased respiration, so soil carbon/energy resources became more limited. Thus, water addition selected for microorganisms with genes responsible for degrading recalcitrant soil organic matter. Accordingly, many other microorganisms without these genes (but likely with other accessory community genes due to relatively stable average microbial genome size) were selected against, leading to the decrease in the diversity of accessory community genes. In summary, nitrogen addition primarily affected core community genes through nitrogen-cycling processes, and water addition primarily regulated accessory community genes through carbon-cycling processes. Although both gene components may significantly respond as the intensity of nitrogen/water addition increases, our results demonstrated how these common global change factors distinctly impact each component. IMPORTANCE Our results demonstrated increased ecosystem nitrogen and water content as the primary drivers of the core and accessory components of soil microbial community functional diversity, respectively. Our findings suggested that more attention should be paid to certain components of community functional diversity under specific global change conditions. Our findings also indicated that microbial communities have adapted to nitrogen addition by strengthening the function of ammonia oxidization to deplete the excess nitrogen, thus maintaining ecosystem homeostasis. Because community gene richness is primarily determined by the presence/absence of accessory community genes, our findings further implied that strategies such as maintaining the amount of soil organic matter could be adopted to effectively improve the functional gene diversity of soil microbial communities subject to global change factors., (Copyright © 2019 Zhang et al.)

Atmospheric nitrogen (N) deposition and phosphorus (P) addition both can change soil bacterial and fungal community structure with a consequent impact on ecosystem functions. However, which factor plays an important role in regulating responses of bacterial and fungal community to N and P enrichments remains unclear. We conducted a manipulative experiment to simulate N and P inputs (10 g N · m -2 · yr -1 NH 4 NO 3 or 10 g P · m -2 · yr -1 NaH 2 PO 4 ) and compared their effects on soil bacterial and fungal species richness and community composition. The results showed that the addition of N significantly increased NH 4 + and Al 3+ by 99.6% and 57.4%, respectively, and consequently led to a decline in soil pH from 4.18 to 3.75 after a 5-year treatment. P addition increased Al 3+ and available P by 27.0% and 10-fold, respectively, but had no effect on soil pH. N addition significantly decreased bacterial species richness and Shannon index and resulted in a substantial shift of bacterial community composition, whereas P addition did not. Neither N nor P addition changed fungal species richness, Shannon index, and fungal community composition. A structural equation model showed that the shift in bacterial community composition was related to an increase in soil acid cations. The principal component scores of soil nutrients showed a significantly positive relationship with fungal community composition. Our results suggest that N and P additions affect soil bacterial and fungal communities in different ways in subtropical forest. These findings highlight how the diversity of microbial communities of subtropical forest soil will depend on future scenarios of anthropogenic N deposition and P enrichment, with a particular sensitivity of bacterial community to N addition., Competing Interests: The authors declare that they have no competing interests.

It has been well established by field experiments that warming stimulates either net ecosystem carbon uptake or release, leading to negative or positive carbon cycle-climate change feedback, respectively. This variation in carbon-climate feedback has been partially attributed to water availability. However, it remains unclear under what conditions water availability enhances or weakens carbon-climate feedback or even changes its direction. Combining a field experiment with a global synthesis, we show that warming stimulates net carbon uptake (negative feedback) under wet conditions, but depresses it (positive feedback) under very dry conditions. This switch in carbon-climate feedback direction arises mainly from scaling effects of warming-induced decreases in soil water content on net ecosystem productivity. This water scaling of warming effects offers generalizable mechanisms not only to help explain varying magnitudes and directions of observed carbon-climate feedback but also to improve model prediction of ecosystem carbon dynamics in response to climate change.

The effects of grazing on methane (CH 4 ) budgets are important for understanding the balance of greenhouse gas emissions and removals in grassland ecosystems. However, the CH 4 budgets of grazing systems, that is simultaneously considering CH 4 uptake by grassland soils and emissions from ruminant enteric fermentation, livestock folds and animal feces, are poorly investigated, particularly for Chinese grasslands, and thus, remained unclear currently. Here, a synthesis of 43 individual studies was carried out to assess the grazing season/annual CH 4 budgets and their responses to grazing in grassland ecosystems of China. The results showed that heavy grazing (HG) significantly decreased, while light grazing (LG) and moderate grazing (MG) had no significant effects soil CH 4 uptake, as compared to un-grazing sites. Grazing has shifted Chinese grasslands from a sink to source for atmospheric CH 4 , and the grazing season/annual CH 4 budgets increased with increasing grazing intensity, while the offset of CH 4 uptake by grassland soils to total CH 4 emissions from sheep, sheepfolds and feces were exponentially decreased with increasing grazing intensity. Moreover, the herbage biomass (HBM), organic matter intake (OMI) and live weight gain (LWG) were decreased while CH 4 emission intensities (i.e., CH 4 emission per HBM, OMI, and LWG) were linearly increased with increasing grazing intensity. Our results demonstrate that mediating grazing intensity, e.g., from HG to LG, could yield the optimal balance between maintaining productive grasslands and meanwhile mitigating CH 4 emissions. This study could help for building strategies with implications for grassland management in China with similar CH 4 emission problems., (Copyright © 2018 Elsevier B.V. All rights reserved.)

Grazing degrades worldwide grasslands and possibly suppresses soil greenhouse gas (GHG: CO 2 , CH 4 and N 2 O) fluxes. However, the global patterns of these three gas fluxes in response to grazing and the general mechanisms remain poorly understood. Here, we performed a meta-analysis of 63 independent grazing studies that measured soil GHG fluxes across global grasslands. Our results revealed that light and moderate grazing had no significant effect on soil CH 4 uptake, N 2 O and CO 2 emission, but heavy grazing consistently reduced them. The magnitudes of their responses to grazing were regulated by grazing duration and precipitation. In comparison with CO 2 emission, soil CH 4 uptake and N 2 O emission were reduced much more under heavier grazing, longer grazing duration or less precipitation. The decrease in soil CO 2 emission was possibly caused by grazing-induced reduction in root biomass and soil moisture, while the decline in soil CH 4 uptake and N 2 O emission was due to decreased soil moisture and substrate availability. Overall, this study provides the first large-scale evaluation on three main soil GHG fluxes in response to grazing, highlighting grazing inhibition of GHG emission but at the cost of plant productivity and soil fertility. We call for future efforts to identify an appropriate grazing intensity that is optimal to balance these complicated impacts., (Copyright © 2018 Elsevier B.V. All rights reserved.)

Soil net nitrogen mineralization rate (N min ), which is critical for soil nitrogen availability and plant growth, is thought to be primarily controlled by climate and soil physical and/or chemical properties. However, the role of microbes on regulating soil N min has not been evaluated on the global scale. By compiling 1565 observational data points of potential net N min from 198 published studies across terrestrial ecosystems, we found that N min significantly increased with soil microbial biomass, total nitrogen, and mean annual precipitation, but decreased with soil pH. The variation of N min was ascribed predominantly to soil microbial biomass on global and biome scales. Mean annual precipitation, soil pH, and total soil nitrogen significantly influenced N min through soil microbes. The structural equation models (SEM) showed that soil substrates were the main factors controlling N min when microbial biomass was excluded. Microbe became the primary driver when it was included in SEM analysis. SEM with soil microbial biomass improved the N min prediction by 19% in comparison with that devoid of soil microbial biomass. The changes in N min contributed the most to global soil NH 4 + -N variations in contrast to climate and soil properties. This study reveals the complex interactions of climate, soil properties, and microbes on N min and highlights the importance of soil microbial biomass in determining N min and nitrogen availability across the globe. The findings necessitate accurate representation of microbes in Earth system models to better predict nitrogen cycle under global change., (© 2018 John Wiley & Sons Ltd.)

Transgenic Bacillus thuringiensis (Bt) crops have been widely planted, and the resulting environmental risks have attracted extensive attention. To foresee the impacts of Bt crops on soil quality, it is essential to understand how Bt crops alter the soil enzymatic activities and what the important influencing factors are. We compiled data from 41 published papers that studied soil enzymatic activities with Bt crops and their non-Bt counterparts. The results showed that dehydrogenase and urease significantly increased, but neutral phosphatase significantly decreased under Bt crop cultivations without Bt residues incorporation. The activities of dehydrogenase, β-glucosidase, urease, nitrate reductase, alkaline phosphatase, and aryl sulfatase significantly increased under Bt crop cultivation with Bt residues incorporation. The response ratios of other enzymes were not significantly changed. Generally, the response ratios of soil enzymes were greater with Bt residues incorporation than those of Bt crop cultivations without Bt residues incorporation. Further, the response ratios of soil enzymes varied with Bt crop types and growth periods. It was the strongest under Bt cotton among Bt crops, and the significant responses usually appeared in the middle growth stages. The responses of soil enzymes ascribed more to the properties of Bt crops than to soil properties across sites. Given - significant responses of some soil enzymes to Bt crops, we recommended that soil environmental risks should be carefully evaluated over the transgenic crops., (Copyright © 2018 Elsevier B.V. All rights reserved.)

Coastal wetlands are considered as a significant sink of global carbon due to their tremendous organic carbon storage. Coastal CO2 and CH4 flux rates play an important role in regulating atmospheric CO2 and CH4 concentrations. However, the relative contributions of vegetation, soil properties, and spatial structure on dry-season ecosystem carbon (C) rates (net ecosystem CO2 exchange, NEE; ecosystem respiration, ER; gross ecosystem productivity, GEP; and CH4) remain unclear at a regional scale. Here, we compared dry-season ecosystem C rates, plant, and soil properties across three vegetation types from 13 locations at a regional scale in the Yellow River Delta (YRD). The results showed that the Phragmites australis stand had the greatest NEE (-1365.4 μmol m-2 s-1), ER (660.2 μmol m-2 s-1), GEP (-2025.5 μmol m-2 s-1) and acted as a CH4 source (0.27 μmol m-2 s-1), whereas the Suaeda heteroptera and Tamarix chinensis stands uptook CH4 (-0.02 to -0.12 μmol m-2 s-1). Stepwise multiple regression analysis demonstrated that plant biomass was the main factor explaining all of the investigated carbon rates (GEP, ER, NEE, and CH4); while soil organic carbon was shown to be the most important for explaining the variability in the processes of carbon release to the atmosphere, i.e., ER and CH4. Variation partitioning results showed that vegetation and soil properties played equally important roles in shaping the pattern of C rates in the YRD. These results provide a better understanding of the link between ecosystem C rates and environmental drivers, and provide a framework to predict regional-scale ecosystem C fluxes under future climate change., Competing Interests: The authors have declared that no competing interests exist.

Because of increases in atmospheric nitrogen (N) deposition worldwide, nutrient imbalances and phosphorus (P) limitations in soil are aggravated, with the result that P fertilizer applications to terrestrial ecosystems worldwide may increase. Nitrification and denitrification in soil are major sources of nitrous oxide emissions, especially in soils treated with fertilizers. However, few researchers have studied how forest soils respond to nutrient additions, so we are not sure how the potential nitrification and denitrification activities (PNA and PDA, respectively) and microbial communities involved in these processes might respond when N and P are added to temperate and subtropical forest soils. We investigated how the PNA, PDA, the abundances and community compositions of nitrifiers and denitrifiers, and environmental properties, including soil pH, soil total and dissolved organic carbon, total and available N and phosphorus P, changed when N and/or P were added to subtropical and temperate forest soils. We quantified the abundance, and analyzed the composition, of functional marker genes of nitrifiers (ammonia-oxidizing bacteria and archaea amoA) and denitrifiers (nirK and nirS) using quantitative PCR and sequencing, respectively. We found that the PNA and PDA in the subtropical soil increased when P was added and PNA in the temperate forest soil increased when either N or P was added. The PNA and PDA were positively correlated with the abundance of ammonia-oxidizing bacteria and nirK-denitrifiers, respectively, in the subtropical forest soil but were not correlated with changes in corresponding community compositions in either of the forest soils. The soil total N to total P ratio explained most of the variabilities in the PNA and PDA in the subtropical forest soils, and the soil exchangeable ammonium concentrations and pH were the main controls on the PNA and PDA, respectively, in the temperate forest soils. Our results indicate that soil environmental conditions have more influence on variations in the PNA and PDA in forest soils fertilized with N and P than the corresponding microbial properties., (Copyright © 2018 Elsevier B.V. All rights reserved.)

Plant stoichiometric coupling among all elements is fundamental to maintaining growth-related ecosystem functions. However, our understanding of nutrient balance in response to global changes remains greatly limited to plant carbon : nitrogen : phosphorus (C : N : P) coupling. Here we evaluated nine element stoichiometric variations with one meta-analysis of 112 global change experiments conducted across global terrestrial ecosystems and one synthesis over 1900 species observations along natural environment gradients across China. We found that experimentally increased soil N and P respectively enhanced plant N : potassium (K), N : calcium (Ca) and N : magnesium (Mg), and P : K, P : Ca and P : Mg, and natural increases in soil N and P resulted in qualitatively similar responses. The ratios of N and P to base cations decreased both under experimental warming and with naturally increasing temperature. With decreasing precipitation, these ratios increased in experiments but decreased under natural environments. Based on these results, we propose a new stoichiometric framework in which all plant element contents and their coupling are not only affected by soil nutrient availability, but also by plant nutrient demand to maintain diverse functions under climate change. This study offers new insights into understanding plant stoichiometric variations across a full set of mineral elements under global changes., (© 2018 The Authors. New Phytologist © 2018 New Phytologist Trust.)

Defoliation is widely used for grassland management. Our understanding of how grass species adjust their regrowth and regain balance after defoliation remains limited. In the present study, we examined the regrowth processes of two dominant species after defoliation in grasslands in Inner Mongolia. Our results showed that the aboveground biomass and total biomass of both species significantly decreased and did not completely recover to the control level after 30 days of regrowth. The leaf mass ratio of Leymus chinensis reached the control level at 15 days, but that of Stipa grandis did not recover to the control level. The root mass ratio of these species reached the same levels as that of the control plants within 10 days after defoliation. As indicated by the dynamics of water-soluble carbohydrates (WSCs), protein, and biomass-based shoot: root ratios, both species regained balances of WSCs and protein between above- and below-ground organs at day 10 after defoliation; however, the biomass regained balance 15 days after defoliation. We deduced that the biomass-based shoot:root ratio was regulated by the WSCs and protein concentrations. In conclusion, following defoliation, both grass species first restore their nutrient-based balance between above- and below-ground parts and then regain biomass balance.

Atmospheric nitrogen (N) deposition and soil acidification both can largely change soil microbial activity and root growth with a consequent impact on soil respiration (R s ). However, it remains unclear which one, N enrichment or soil acidification, plays more important role in impacting soil respiration. We conducted a manipulative experiment to simulate N enrichment (10gm -2 yr -1 NH 4 NO 3 ) and soil acidity (0.552molH + m -2 yr -1 sulfuric acid) and compared their effects on R s and its components in a subtropical forest. The results showed that soil pH was reduced by 0.4 similarly under N addition or acid addition after 3years' treatment. Acid addition decreased autotrophic respiration (R a ) by 22-35% and heterotrophic respiration (R h ) by 22-23%, resulting in a reduction of R s by 22-26% in the two years. N addition reduced R a , R h , R s less than acid addition did. The reductions of R s and its components were attributed to increase of soil acid cations and reduction of cellulose degrading enzymes activity. N addition and soil acidification significantly enhanced fungal to bacterial ratio. All the cellulose degrading enzymes were reduced more by soil acidity (43-50%) than N addition (30-39%). The principal component scores of degrading enzymes activity showed significantly positive relationships with R h . Structural equation model showed that soil acidification played more important role than N enrichment in changing R s and its components. We therefore suggest that soil acidification is an important mechanism underlying soil respiration changes, and should be incorporated into biogeochemical models to improve the prediction of ecosystem C cycling in the future scenarios of anthropogenic N deposition and acid enrichment., (Copyright © 2017 Elsevier B.V. All rights reserved.)

Understanding the impacts of biodiversity loss on ecosystem functioning and services has been a central issue in ecology. Experiments in synthetic communities suggest that biodiversity loss may erode a set of ecosystem functions, but studies in natural communities indicate that the effects of biodiversity loss are usually weak and that multiple functions can be sustained by relatively few species. Yet, the mechanisms by which natural ecosystems are able to maintain multiple functions in the face of diversity loss remain poorly understood. With a long-term and large-scale removal experiment in the Inner Mongolian grassland, here we showed that losses of plant functional groups (PFGs) can reduce multiple ecosystem functions, including biomass production, soil NO 3 -N use, net ecosystem carbon exchange, gross ecosystem productivity, and ecosystem respiration, but the magnitudes of these effects depended largely on which PFGs were removed. Removing the two dominant PFGs (perennial rhizomatous grasses and perennial bunchgrasses) simultaneously resulted in dramatic declines in all examined functions, but such declines were circumvented when either dominant PFG was present. We identify the major mechanism for this as a compensation effect by which each dominant PFG can mitigate the losses of others. This study provides evidence that compensation ensuing from PFG losses can mitigate their negative consequence, and thus natural communities may be more resilient to biodiversity loss than currently thought if the remaining PFGs have strong compensation capabilities. On the other hand, ecosystems without well-developed compensatory functional diversity may be much more vulnerable to biodiversity loss., (© 2016 by the Ecological Society of America.)

Nitrogen (N) deposition is impacting the services that ecosystems provide to humanity. However, the mechanisms determining impacts on the N cycle are not fully understood. To explore the mechanistic underpinnings of N impacts on N cycle processes, we reviewed and synthesised recent progress in ecosystem N research through empirical studies, conceptual analysis and model simulations. Experimental and observational studies have revealed that the stimulation of plant N uptake and soil retention generally diminishes as N loading increases, while dissolved and gaseous losses of N occur at low N availability but increase exponentially and become the dominant fate of N at high loading rates. The original N saturation hypothesis emphasises sequential N saturation from plant uptake to soil retention before N losses occur. However, biogeochemical models that simulate simultaneous competition for soil N substrates by multiple processes match the observed patterns of N losses better than models based on sequential competition. To enable better prediction of terrestrial N cycle responses to N loading, we recommend that future research identifies the response functions of different N processes to substrate availability using manipulative experiments, and incorporates the measured N saturation response functions into conceptual, theoretical and quantitative analyses., (© 2016 John Wiley & Sons Ltd/CNRS.)

Bunchgrasses are one of the most important plant functional groups in grassland ecosystems. Reproductive allocation (RA) for a bunchgrass is a hierarchical process; however, how bunchgrasses adjust their RAs along hierarchical levels in response to nutrient addition has never been addressed. Here, utilizing an 11-year nutrient addition experiment, we examined the patterns and variations in RA of Agropyron cristatum at the individual, tiller and spike levels. We evaluated the reproductive allometric relationship at each level by type II regression analysis to determine size-dependent and size-independent effects on plant RA variations. Our results indicate that the proportion of reproductive individuals in A. cristatum increased significantly after 11 years of nutrient addition. Adjustments in RA in A. cristatum were mainly occurred at the individual and tiller levels but not at the spike level. A size-dependent effect was a dominant mechanism underlying the changes in plant RA at both individual and tiller levels. Likewise, the distribution of plant size was markedly changed with large individuals increasing after nutrient addition. Tiller-level RA may be a limiting factor for the adjustment of RA in A. cristatum. To the best of our knowledge, this study is the first to examine plant responses in terms of reproductive allocation and allometry to nutrient enrichment within a bunchgrass population from a hierarchical view. Our findings have important implications for understanding the mechanisms underlying bunchgrass responses in RA to future eutrophication due to human activities. In addition, we developed a hierarchical analysis method for disentangling the mechanisms that lead to variation in RA for perennial bunchgrasses.