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The frequency and intensity of droughts worldwide are challenging the conservation of soil organic carbon (SOC) pool. Microbial necromass is a key component of SOC, but how it responds to drought at specific soil depths remains largely unknown. Here, we conducted a 3-year field experiment in a forest plantation to investigate the impacts of drought intensities under three treatments (ambient control [CK], moderate drought [30% throughfall removal], and intensive drought [50% throughfall removal]) on soil microbial necromass pools (i.e., bacterial necromass carbon, fungal necromass carbon, and total microbial necromass carbon). We showed that the effects of drought on microbial necromass depended on microbial groups, soil depth, and drought intensity. While moderate drought increased total (+9.1% ± 3.3%) and fungal (+13.5% ± 4.9%) necromass carbon in the topsoil layer (0-15 cm), intensive drought reduced total (-31.6% ± 3.7%) and fungal (-43.6% ± 4.0%) necromass in the subsoil layer (15-30 cm). In contrast, both drought treatments significantly increased the BNC in the topsoil and subsoil. Our results suggested that the effects of drought on the microbial necromass of the subsoil were more pronounced than those of the topsoil. This study highlights the complex responses of microbial necromass to drought events depending on microbial community structure, drought intensity and soil depth with global implications when forecasting carbon cycling under climate change., (© 2023 John Wiley & Sons Ltd.)

Soil pH is critically important in regulating soil nutrients and thus influencing the biodiversity and ecosystem functions of terrestrial ecosystems. Despite the ongoing threat of nitrogen (N) pollution especially in the fast-developing regions, it remains unclear how increasing N deposition affects soil pH across global terrestrial ecosystems. By conducting a global meta-analysis with paired observations of soil pH under N addition and control from 634 studies spanning major types of terrestrial ecosystems, we show that soil acidification increases rapidly with N addition amount and is most severe in neutral-pH soils. Grassland soil pH decreases most strongly under high N addition while wetlands are the least acidified. By extrapolating these relationships to global mapping, we reveal that atmospheric N deposition leads to a global average soil pH decline of -0.16 in the past 40 years and regions encompassing Eastern United States, Southern Brazil, Europe, and South and East Asia are the hotspots of soil acidification under N deposition. Our results highlight that anthropogenically amplified atmospheric N deposition has profoundly altered global soil pH and chemistry. They suggest that atmospheric N deposition is a major threat to global terrestrial biodiversity and ecosystem functions., (© 2023 John Wiley & Sons Ltd.)

Soil respiration (Rs), as the second largest flux of carbon dioxide (CO 2 ) between terrestrial ecosystems and the atmosphere, is vulnerable to global nitrogen (N) enrichment. However, the global distribution of the N effects on Rs remains uncertain. Here, we compiled a new database containing 1282 observations of Rs and its heterotrophic component (Rh) in field N manipulative experiments from 317 published papers. Using this up-to-date database, we first performed a formal meta-analysis to explore the responses of Rs and Rh to N addition, and then presented a global spatially explicit quantification of the N effects using a Random Forest model. Our results showed that experimental N addition significantly increased Rs but had a minimal impact on Rh, not supporting the prevailing view that N enrichment inhibits soil microbial respiration. For the major biomes, the magnitude of N input was the main determinant of the spatial variation in Rs response, while the most important predictors for Rh response were biome specific. Based on the key predictors, global mapping visually demonstrated a positive N effect in the regions with higher anthropogenic N inputs (i.e., atmospheric N deposition and agricultural fertilization). Overall, our analysis not only provides novel insight into the N effects on soil CO 2 fluxes, but also presents a spatially explicit assessment of the N effects at the global scale, which are pivotal for understanding ecosystem carbon dynamics in future scenarios with more frequent anthropogenic activities., (© 2023 John Wiley & Sons Ltd.)

Carbon (C):nitrogen (N):phosphorus (P) stoichiometry in plants, soils, and microbial biomass influences productivity and nutrient cycling in terrestrial ecosystems. Anthropogenic inputs of P to ecosystems are increasing; however, our understanding of the impacts of P addition on terrestrial ecosystem C:N:P ratios remains elusive. By conducting a meta-analysis with 1413 paired observations from 121 publications, we showed that P addition significantly decreased plant, soil, and microbial biomass N:P and C:P ratios, but had negligible effects on C:N ratios. The reductions in N:P and C:P ratios became more evident as the P application rates and experimental duration increased. The P addition effects on terrestrial ecosystem C:N:P stoichiometry did not vary with ecosystem types or climates. Moreover, the responses of N:P and C:P ratios in soil and microbial biomass were associated with the responses of soil pH and fungi:bacteria ratios. Additionally, P additions increased net primary productivity, microbial biomass, soil respiration, N mineralization, and N nitrification, but decreased ammonium and nitrate contents. Decreases in plant N:P and C:P ratios were both negatively correlated to net primary productivity and soil respiration, but positively correlated to ammonium and nitrate contents; microbial biomass, soil respiration, ammonium contents, and nitrate contents all increased with declining soil N:P and C:P ratios. Our findings highlight that P additions could imbalance C:N:P stoichiometry and potentially impact the terrestrial ecosystem functions., (© 2022 John Wiley & Sons Ltd.)

Global environmental changes have strongly affected forest demographic rates, particularly amplified tree mortality in high latitude forests (e.g., two to five times greater mortality probability over the half-century). Although forest functional composition is critical for multitrophic biodiversity and ecosystem functioning, it remains unclear how functional composition has changed over time across large high latitude regions, which have been warming twice the rate of the globe as a whole. Using extensive spatial and long-term forest inventory data (17,107 plots monitored 1951-2016) across Canada, we found that after accounting for stand age-dependent functional shifts, functional composition shifted toward fast-growing deciduous broadleaved trees and higher drought tolerance over time. The temporal shift toward deciduous broadleaved trees was consistent across the baseline climate. However, over the study period, drought tolerance increased (or shade tolerance decreased) by 300% in colder boreal regions, while drought tolerance did not shift significantly in warmer temperate climates. A further analysis accounting for temporal changes in atmospheric CO 2 , temperature, and water availability indicated that the functional composition of colder regions shifted toward drought tolerance more rapidly with rising CO 2 than warmer regions, suggesting the greater vulnerability of boreal forests than temperate forests under ongoing global environmental changes. Future ecosystem management practices should consider spatial differences in functional responses to global environmental change, focusing on high latitude forests experiencing higher rates of warming and compositional changes., (© 2021 John Wiley & Sons Ltd.)

Soil organic carbon (SOC) is the largest carbon sink in terrestrial ecosystems and plays a critical role in mitigating climate change. Increasing reactive nitrogen (N) in ecosystems caused by anthropogenic N input substantially affects SOC dynamics. However, uncertainties remain concerning the effects of N addition on SOC in both organic and mineral soil layers over time at the global scale. Here, we analysed a large empirical data set spanning 60 years across 369 sites worldwide to explore the temporal dynamics of SOC to N addition. We found that N addition significantly increased SOC across the globe by 4.2% (2.7%-5.8%). SOC increases were amplified from short- to long-term N addition durations in both organic and mineral soil layers. The positive effects of N addition on SOC were independent of ecosystem types, mean annual temperature and precipitation. Our findings suggest that SOC increases largely resulted from the enhanced plant C input to soils coupled with reduced C loss from decomposition and amplification was associated with reduced microbial biomass and respiration under long-term N addition. Our study suggests that N addition will enhance SOC sequestration over time and contribute to future climate change mitigation., (© 2020 John Wiley & Sons Ltd.)

CO 2 fluxes from wood decomposition represent an important source of carbon from forest ecosystems to the atmosphere, which are determined by both wood traits and climate influencing the metabolic rates of decomposers. Previous studies have quantified the effects of moisture and temperature on wood decomposition, but these effects were not separated from the potential influence of wood traits. Indeed, it is not well understood how traits and climate interact to influence wood CO 2 fluxes. Here, we examined the responses of CO 2 fluxes from dead wood with different traits (angiosperm and gymnosperm) to 0%, 35%, and 70% rainfall reduction across seasonal temperature gradients. Our results showed that drought significantly decreased wood CO 2 fluxes, but its effects varied with both taxonomical group and drought intensity. Drought-induced reduction in wood CO 2 fluxes was larger in angiosperms than gymnosperms for the 35% rainfall reduction treatment, but there was no significant difference between these groups for the 70% reduction treatment. This is because wood nitrogen density and carbon quality were significantly higher in angiosperms than gymnosperms, yielding a higher moisture sensitivity of wood decomposition. These findings were demonstrated by a significant positive interaction effect between wood nitrogen and moisture on CO 2 fluxes in a structural equation model. Additionally, we ascertained that a constant temperature sensitivity of CO 2 fluxes was independent of wood traits and consistent with previous estimates for extracellular enzyme kinetics. Our results highlight the key role of wood traits in regulating drought responses of wood carbon fluxes. Given that both climate and forest management might extensively modify taxonomic compositions in the future, it is critical for carbon cycle models to account for such interactions between wood traits and climate in driving dynamics of wood decomposition., (© 2020 John Wiley & Sons Ltd.)

Carbon (C) and nitrogen (N) are the primary elements involved in the growth and development of plants. The C:N ratio is an indicator of nitrogen use efficiency (NUE) and an input parameter for some ecological and ecosystem models. However, knowledge remains limited about the convergent or divergent variation in the C:N ratios among different plant organs (e.g., leaf, branch, trunk, and root) and how evolution and environment affect the coefficient shifts. Using systematic measurements of the leaf-branch-trunk-root of 2,139 species from tropical to cold-temperate forests, we comprehensively evaluated variation in C:N ratio in different organs in different taxa and forest types. The ratios showed convergence in the direction of change but divergence in the rate of change. Plants evolved toward lower C:N ratios in the leaf and branch, with N playing a more important role than C. The C:N ratio of plant organs (except for the leaf) was constrained by phylogeny, but not strongly. Both the change of C:N during evolution and its spatial variation (lower C:N ratio at midlatitudes) help develop the adaptive growth hypothesis. That is, plants with a higher C:N ratio promote NUE under strong N-limited conditions to ensure survival priority, whereas plants with a lower C:N ratio under less N-limited environments benefit growth priority. In nature, larger proportion of species with a high C:N ratio enabled communities to inhabit more N-limited conditions. Our results provide new insights on the evolution and drivers of C:N ratio among different plant organs, as well as provide a quantitative basis to optimize land surface process models., (© 2019 John Wiley & Sons Ltd.)

The rapid global biodiversity loss has led to the decline in ecosystem function. Despite the critical importance of soil respiration (Rs) in the global carbon and nutrient cycles, how plant diversity loss affects Rs remains uncertain. Here we present a meta-analysis using 446 paired observations from 95 published studies to evaluate the effects of plant and litter mixtures on Rs and its components. We found that total Rs and heterotrophic respiration (Rh) were, on average, greater in plant mixtures than expected from those of monocultures. These mixture effects increased with increasing species richness (SR) in both plant and litter mixtures. While the positive effects of species mixtures remained similar over time for total Rs, they increased over time for Rh in plant mixtures but decreased in litter mixtures. Despite the wide range of variations in mean annual temperature, annual aridity index, and ecosystem types, the plant mixture effects on total Rs and Rh did not change geographically, except for a more pronounced increase of total Rs in species mixtures with reduced water availability. Our structural equation models suggested that the positive effects of SR and stand age on total and Rh were driven by increased plant inputs and soil microbial biomass. Our results suggest that plant diversity loss has ubiquitous negative impacts on Rs, one of the fundamental carbon-cycle processes sustaining terrestrial element cycling and ecosystem function., (© 2019 John Wiley & Sons Ltd.)

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.)

Soil respiration (Rs) is the largest terrestrial carbon (C) efflux to the atmosphere and is predicted to increase drastically through global warming. However, the responses of Rs to global warming are complicated by the fact that terrestrial plant growth and the subsequent input of plant litter to soil are also altered by ongoing climate change and human activities. Despite a number of experiments established in various ecosystems around the world, it remains a challenge to predict the magnitude and direction of changes in Rs and its temperature sensitivity (Q 10 ) due to litter alteration. We present a meta-analysis of 100 published studies to examine the responses of Rs and Q 10 to manipulated aboveground and belowground litter alterations. We found that 100% aboveground litter addition (double litter) increased Rs by 26.1% (95% confident intervals, 18.4%-33.7%), whereas 100% aboveground litter removal, root removal and litter + root removal reduced Rs by 22.8% (18.5%-27.1%), 34.1% (27.2%-40.9%) and 43.4% (36.6%-50.2%) respectively. Moreover, the effects of aboveground double litter and litter removal on Rs increased with experimental duration, but not those of root removal. Aboveground litter removal marginally increased Q 10 by 6.2% (0.2%-12.3%) because of the higher temperature sensitivity of stable C substrate than fresh litter. Estimated from the studies that simultaneously tested the responses of Rs to aboveground litter addition and removal and assuming negligible changes in root-derived Rs, "priming effect" on average accounted for 7.3% (0.6%-14.0%) of Rs and increased over time. Across the global variation in terrestrial ecosystems, the effects of aboveground litter removal, root removal, litter + root removal on Rs as well as the positive effect of litter removal on Q 10 increased with water availability. Our meta-analysis indicates that priming effects should be considered in predicting Rs to climate change-induced increases in litterfall. Our analysis also highlights the need to incorporate spatial climate gradient in projecting long-term Rs responses to litter alterations., (© 2018 John Wiley & Sons Ltd.)

Controlled experiments have shown that global changes decouple the biogeochemical cycles of carbon (C), nitrogen (N), and phosphorus (P), resulting in shifting stoichiometry that lies at the core of ecosystem functioning. However, the response of soil stoichiometry to global changes in natural ecosystems with different soil depths, vegetation types, and climate gradients remains poorly understood. Based on 2,736 observations along soil profiles of 0-150 cm depth from 1955 to 2016, we evaluated the temporal changes in soil C-N-P stoichiometry across subtropical China, where soils are P-impoverished, with diverse vegetation, soil, and parent material types and a wide range of climate gradients. We found a significant overall increase in soil total C concentration and a decrease in soil total P concentration, resulting in increasing soil C:P and N:P ratios during the past 60 years across all soil depths. Although average soil N concentration did not change, soil C:N increased in topsoil while decreasing in deeper soil. The temporal trends in soil C-N-P stoichiometry differed among vegetation, soil, parent material types, and spatial climate variations, with significantly increased C:P and N:P ratios for evergreen broadleaf forest and highly weathered Ultisols, and more pronounced temporal changes in soil C:N, N:P, and C:P ratios at low elevations. Our sensitivity analysis suggests that the temporal changes in soil stoichiometry resulted from elevated N deposition, rising atmospheric CO 2 concentration and regional warming. Our findings revealed that the responses of soil C-N-P and stoichiometry to long-term global changes have occurred across the whole soil depth in subtropical China and the magnitudes of the changes in soil stoichiometry are dependent on vegetation types, soil types, and spatial climate variations., (© 2017 John Wiley & Sons Ltd.)

Global change impacts on biogeochemical cycles have been widely studied, but our understanding of whether the responses of plant elemental composition to global change drivers differ between above- and belowground plant organs remains incomplete. We conducted a meta-analysis of 201 reports including 1,687 observations of studies that have analyzed simultaneously N and P concentrations changes in leaves and roots in the same plants in response to drought, elevated [CO 2 ], and N and P fertilization around the world, and contrasted the results within those obtained with a general database (838 reports and 14,772 observations) that analyzed the changes in N and P concentrations in leaves and/or roots of plants submitted to the commented global change drivers. At global level, elevated [CO 2 ] decreased N concentrations in leaves and roots and decreased N:P ratio in roots but no in leaves, but was not related to P concentration changes. However, the response differed among vegetation types. In temperate forests, elevated [CO 2 ] was related with lower N concentrations in leaves but not in roots, whereas in crops, the contrary patterns were observed. Elevated [CO 2 ] decreased N concentrations in leaves and roots in tundra plants, whereas not clear relationships were observed in temperate grasslands. However, when elevated [CO 2 ] and N fertilization coincided, leaves had lower N concentrations, whereas root had higher N concentrations suggesting that more nutrients will be allocated to roots to improve uptake of the soil resources not directly provided by the global change drivers. N fertilization and drought increased foliar and root N concentrations while the effects on P concentrations were less clear. The changes in N and P allocation to leaves and root, especially those occurring in opposite direction between them have the capacity to differentially affect above- and belowground ecosystem functions, such as litter mineralization and above- and belowground food webs., (© 2017 John Wiley & Sons Ltd.)