Biochar dose-dependent impacts on soil bacterial and fungal diversity across the globe.

Biochar, a widely used material for soil amendment, has been found to offer numerous advantages in improving soil properties and the habitats for soil microorganisms. However, there is still a lack of global perspectives on the influence of various levels of biochar addition on soil microbial diversity and primary components. Thus, in our study, we performed a global meta-analysis of studies to determine how different doses of biochar affect soil total carbon (C), nitrogen (N), pH, alpha- and beta-diversity, and the major phyla of both bacterial and fungal communities. Our results revealed that biochar significantly increased soil pH by 4 %, soil total C and N by 68 % and 22 %, respectively, in which the positive effects increased with biochar doses. Moreover, biochar promoted soil bacterial richness and evenness by 3-8 % at the biochar concentrations of 1-5 % (w/w), while dramatically shifting bacterial beta-diversity at the doses of >2 % (w/w). Specifically, biochar exhibited significantly positive effects on bacterial phyla of Acidobacteria, Bacteroidetes, Gemmatimonadetes, and Proteobacteria, especially Deltaproteobacteria and Gammaproteobacteria, by 4-10 % depending on the concentrations. On the contrary, the bacterial phylum of Verrucomicrobia and fungal phylum of Basidiomycota showed significant negative responses to biochar by -8 % and -24 %, respectively. Therefore, our meta-analysis provides theoretical support for the development of optimized agricultural management practices by emphasizing biochar application dosing.

Intermediate soil acidification induces highest nitrous oxide emissions.

Global potent greenhouse gas nitrous oxide (N2O) emissions from soil are accelerating, with increases in the proportion of reactive nitrogen emitted as N2O, i.e., N2O emission factor (EF). Yet, the primary controls and underlying mechanisms of EFs remain unresolved. Based on two independent but complementary global syntheses, and three field studies determining effects of acidity on N2O EFs and soil denitrifying microorganisms, we show that soil pH predominantly controls N2O EFs and emissions by affecting the denitrifier community composition. Analysis of 5438 paired data points of N2O emission fluxes revealed a hump-shaped relationship between soil pH and EFs, with the highest EFs occurring in moderately acidic soils that favored N2O-producing over N2O-consuming microorganisms, and induced high N2O emissions. Our results illustrate that soil pH has a unimodal relationship with soil denitrifiers and EFs, and the net N2O emission depends on both the N2O/(N2O + N2) ratio and overall denitrification rate. These findings can inform strategies to predict and mitigate soil N2O emissions under future nitrogen input scenarios.

Monotonic trends of soil microbiomes, metagenomic and metabolomic functioning across ecosystems along water gradients in the Altai region, northwestern China.

To investigate microbial communities and their contributions to carbon and nutrient cycling along water gradients can enhance our comprehension of climate change impacts on ecosystem services. Thus, we conducted an assessment of microbial communities, metagenomic functions, and metabolomic profiles within four ecosystems, i.e., desert grassland (DG), shrub-steppe (SS), forest (FO), and marsh (MA) in the Altai region of Xinjiang, China. Our results showed that soil total carbon (TC), total nitrogen, NH4+, and NO3- increased, but pH decreased with soil water gradients. Microbial abundances and richness also increased with soil moisture except the abundances of fungi and protists being lowest in MA. A shift in microbial community composition is evident along the soil moisture gradient, with Proteobacteria, Basidiomycota, and Evosea proliferating but a decline in Actinobacteria and Cercozoa. The ß-diversity of microbiomes, metagenomic, and metabolomic functioning were correlated with soil moisture gradients and have significant associations with specific soil factors of TC, NH4+, and pH. Metagenomic functions associated with carbohydrate and DNA metabolisms, as well as phages, prophages, TE, plasmids functions diminished with moisture, whereas the genes involved in nitrogen and potassium metabolism, along with certain biological interactions and environmental information processing functions, demonstrated an augmentation. Additionally, MA harbored the most abundant metabolomics dominated by lipids and lipid-like molecules and organic oxygen compounds, except certain metabolites showing decline trends along water gradients, such as N'-Hydroxymethylnorcotinine and 5-Hydroxyenterolactone. Thus, our study suggests that future ecosystem succession facilitated by changes in rainfall patterns will significantly alter soil microbial taxa, functional potential, and metabolite fractions.

Mycorrhizae enhance reactive minerals but reduce mineral-associated carbon.

Soil organic carbon (C) is the largest active C pool of Earth's surface and is thus vital in sustaining terrestrial productivity and climate stability. Arbuscular mycorrhizal fungi (AMF) form symbioses with most terrestrial plants and critically modulate soil C dynamics. Yet, it remains unclear whether and how AMF-root associations (i.e., mycorrhizae) interact with soil minerals to affect soil C cycling. Here we showed that the presence of both roots and AMF increased soil dissolved organic C and reactive Fe minerals, as well as litter decomposition and soil CO2 emissions. However, it reduced mineral-associated C. Also, high-resolution nanoscale secondary ion mass spectrometry images showed the existence of a thin coating (0.5-1.0 µm thick) of 56 Fe16 O- (Fe minerals) on the surface of 12 C14 N- (fungal biomass), illustrating the close physical association between fungal hyphae and soil Fe minerals. In addition, AMF genera were divergently related to reactive Fe minerals, with Glomus being positively but Paraglomus and Acaulospora negatively correlated with reactive Fe minerals. Moreover, the presence of roots and AMF, particularly when combined with litter addition, enhanced the abundances of several critical soil bacterial genera that are associated with the formation of reactive minerals in soils. A conceptual framework was further proposed to illustrate how AMF-root associations impact soil C cycling in the rhizosphere. Briefly, root exudates and the inoculated AMF not only stimulated the decomposition of litter and SOC and promoted the production of CO2 emission, but also drove soil C persistence by unlocking mineral elements and promoting the formation of reactive minerals. Together, these findings provide new insights into the mechanisms that underlie the formation of reactive minerals and have significant implications for understanding and managing soil C persistence.

Soil development following glacier retreat shapes metagenomic and metabolomic functioning associated with asynchronous C and N accumulation.

Glacier retreat caused by global warming may result in the variation of soil organic carbon and nutrient cycling. Yet, the dynamic change of soil microbial functional profiles, especially C metabolism-related, with soil development following glacier retreat are still unclear. In the present study, we investigated the soil microbial communities, metagenomic functioning, and metabolomic profiles along the Hailuogou Glacier forefield representing a 120-year chronosequence. The alpha diversity indices of soil bacteria, protozoa and nifH genes showed an upward trend with increased soil ages, and the beta diversity of soil archaea, bacteria, fungi, protozoa, nifH and nirS genes were significantly correlated with soil ages, in which increasing soil C and P while decreased C/N and pH significantly contributed to the differences of soil microbial communities among the analyzed environmental variables. The metagenomic functional genes related to the metabolisms of Glycogen and Cellulosome, Iron Acquisition and Metabolism were significantly decreased with chronosequence, while the utilization of Xylose and Lactate, Potassium Metabolism, Sulfur Metabolism showing an upward trend with soil ages, in which soil C/N ratios and pH were the most influential factors. In addition, soil C and C/N ratios were also significantly correlated to metabolomic compositions, in which the complexity of the metabolite structure increased with soil ages. Our results indicate that glacier retreat may lead to the asynchronous C and N accumulation along the chronosequence, thereby affecting the metagenomic and metabolomic functioning of soil microbial communities related to C metabolisms during soil development following glacier retreat.

Moderate precipitation reduction enhances nitrogen cycling and soil nitrous oxide emissions in a semi-arid grassland.

The ongoing climate change is predicted to induce more weather extremes such as frequent drought and high-intensity precipitation events, causing more severe drying-rewetting cycles in soil. However, it remains largely unknown how these changes will affect soil nitrogen (N)-cycling microbes and the emissions of potent greenhouse gas nitrous oxide (N2 O). Utilizing a field precipitation manipulation in a semi-arid grassland on the Loess Plateau, we examined how precipitation reduction (ca. -30%) influenced soil N2 O and carbon dioxide (CO2 ) emissions in field, and in a complementary lab-incubation with simulated drying-rewetting cycles. Results obtained showed that precipitation reduction stimulated plant root turnover and N-cycling processes, enhancing soil N2 O and CO2 emissions in field, particularly after each rainfall event. Also, high-resolution isotopic analyses revealed that field soil N2 O emissions primarily originated from nitrification process. The incubation experiment further showed that in field soils under precipitation reduction, drying-rewetting stimulated N mineralization and ammonia-oxidizing bacteria in favor of genera Nitrosospira and Nitrosovibrio, increasing nitrification and N2 O emissions. These findings suggest that moderate precipitation reduction, accompanied with changes in drying-rewetting cycles under future precipitation scenarios, may enhance N cycling processes and soil N2 O emissions in semi-arid ecosystems, feeding positively back to the ongoing climate change.

Nitrogen availability mediates soil carbon cycling response to climate warming: A meta-analysis.

Global climate warming may induce a positive feedback through increasing soil carbon (C) release to the atmosphere. Although warming can affect both C input to and output from soil, direct and convincing evidence illustrating that warming induces a net change in soil C is still lacking. We synthesized the results from field warming experiments at 165 sites across the globe and found that climate warming had no significant effect on soil C stock. On average, warming significantly increased root biomass and soil respiration, but warming effects on root biomass and soil respiration strongly depended on soil nitrogen (N) availability. Under high N availability (soil C:N ratio < 15), warming had no significant effect on root biomass, but promoted the coupling between effect sizes of root biomass and soil C stock. Under relative N limitation (soil C:N ratio > 15), warming significantly enhanced root biomass. However, the enhancement of root biomass did not induce a corresponding C accumulation in soil, possibly because warming promoted microbial CO2 release that offset the increased root C input. Also, reactive N input alleviated warming-induced C loss from soil, but elevated atmospheric CO2 or precipitation increase/reduction did not. Together, our findings indicate that the relative availability of soil C to N (i.e., soil C:N ratio) critically mediates warming effects on soil C dynamics, suggesting that its incorporation into C-climate models may improve the prediction of soil C cycling under future global warming scenarios.

Arbuscular Mycorrhizae Shift Community Composition of N-Cycling Microbes and Suppress Soil N2O Emission.

Mycorrhizae are ubiquitous symbiotic associations between arbuscular mycorrhizal fungi (AMF) and terrestrial plants, in which AMF receive photosynthates from and acquire soil nutrients for their host plants. Plant uptake of soil nitrogen (N) reduces N substrate for microbial processes that generate nitrous oxide (N2O), a potent greenhouse gas. However, the underlying microbial mechanisms remain poorly understood, particularly in agroecosystems with high reactive N inputs. We examined how plant roots and AMF affect N2O emissions, N2O-producing (nirK and nirS) and N2O-consuming (nosZ) microbes under normal and high N inputs in conventional (CONV) and organically managed (OM) soils. Here, we show that high N input increased soil N2O emissions and the ratio of nirK to nirS microbes. Roots and AMF did not affect the (nirK + nirS)/nosZ ratio but significantly reduced N2O emissions and the nirK/nirS ratio. They reduced the nirK/nirS ratio by reducing nirK-Rhodobacterales but increasing nirS-Rhodocyclales in the CONV soil while decreasing nirK-Burkholderiales but increasing nirS-Rhizobiales in the OM soil. Our results indicate that plant roots and AMF reduced N2O emission directly by reducing soil N and indirectly through shifting the community composition of N2O-producing microbes in N-enriched agroecosystems, suggesting that harnessing the rhizosphere microbiome through agricultural management might offer additional potential for N2O emission mitigation.

Climate warming promotes deterministic assembly of arbuscular mycorrhizal fungal communities.

Arbuscular mycorrhizal fungi (AMF) significantly contribute to plant resource acquisition and play important roles in mediating plant interactions and soil carbon (C) dynamics. However, it remains unclear how AMF communities respond to climate change. We assessed impacts of warming and precipitation alterations (30% increase or decrease) on soil AMF communities, and examined major ecological processes shaping the AMF community assemblage in a Tibetan alpine meadow. Our results showed that warming significantly increased root biomass, and available nitrogen (N) and phosphorus (P) in soil. While precipitation alterations increased AMF abundances, they did not significantly affect the composition or diversity of AMF communities. In contrast, warming altered the composition of AMF communities and reduced their Shannon-Wiener index and Pielou's evenness. In particular, warming shifted the AMF community composition in favor of Diversisporaceae over Glomeraceae, likely through its impact on soil N and P availability. In addition, AMF communities were phylogenetically random in the unwarmed control but clustered in warming plots, implying more deterministic community assembly under climate warming. Warming enhancement of root growth, N and P availability likely reduced plant C-allocation to AMF, imposing stronger environmental filtering on AMF communities. We further proposed a conceptual framework that integrates biological and geochemical processes into a mechanistic understanding of warming and precipitation changes' effects on AMF. Taken together, these results suggest that soil AMF communities may be more sensitive to warming than expected, highlighting the need to monitor their community structure and associated functional consequences on plant communities and soil C dynamics under the future warmer climate.

Warming and elevated ozone induce tradeoffs between fine roots and mycorrhizal fungi and stimulate organic carbon decomposition.

Climate warming and elevated ozone (eO3) are important climate change components that can affect plant growth and plant-microbe interactions. However, the resulting impact on soil carbon (C) dynamics, as well as the underlying mechanisms, remains unclear. Here, we show that warming, eO3, and their combination induce tradeoffs between roots and their symbiotic arbuscular mycorrhizal fungi (AMF) and stimulate organic C decomposition in a nontilled soybean agroecosystem. While warming and eO3 reduced root biomass, tissue density, and AMF colonization, they increased specific root length and promoted decomposition of both native and newly added organic C. Also, they shifted AMF community composition in favor of the genus Paraglomus with high nutrient-absorbing hyphal surface over the genus Glomus prone to protection of soil organic C. Our findings provide deep insights into plant-microbial interactive responses to warming and eO3 and how these responses may modulate soil organic C dynamics under future climate change scenarios.

Soil acidification alters root morphology, increases root biomass but reduces root decomposition in an alpine grassland.

Soil acidification has been expanding in many areas of Asia due to increasing reactive nitrogen (N) inputs and industrial activities. While the detrimental effects of acidification on forests have been extensively studied, less attention has been paid to grasslands, particularly alpine grasslands. In a soil pH manipulation experiment in the Qinghai-Tibet Plateau, we examined the effects of soil acidification on plant roots, which account for the major part of alpine plants. After three years of manipulation, soil pH decreased from 6.0 to 4.7 with the acid-addition gradient, accompanied by significant changes in the availability of soil nitrogen, phosphorus and cations. Plant composition shifted with the soil acidity, with graminoids replacing forbs. Differing from findings in forests, soil acidification in the alpine grassland increased root biomass by increasing the fraction of coarse roots and the production of fine roots, corresponding to enhanced sedge and grass biomass, respectively. In addition, litter decomposability decreased with altered root morphological and chemical traits, and soil acidification slowed root decomposition by reducing soil microbial activity and litter quality. Our results showed that acidification effect on root dynamics in our alpine grassland was significantly different from that in forests, and supported similar results obtained in limited studies in other grassland ecosystems. These results suggest an important role of root morphology in mediating root dynamics, and imply that soil acidification may lead to transient increase in soil carbon stock as root standing biomass and undecomposed root litter. These changes may reduce nutrient cycling and further constrain ecosystem productivity in nutrient-limiting alpine systems.

Interactive global change factors mitigate soil aggregation and carbon change in a semi-arid grassland.

The ongoing global change is multi-faceted, but the interactive effects of multiple drivers on the persistence of soil carbon (C) are poorly understood. We examined the effects of warming, reactive nitrogen (N) inputs (12 g N m-2  year-1 ) and altered precipitation (+ or - 30% ambient) on soil aggregates and mineral-associated C in a 4 year manipulation experiment with a semi-arid grassland on China's Loess Plateau. Our results showed that in the absence of N inputs, precipitation additions significantly enhanced soil aggregation and promoted the coupling between aggregation and both soil fungal biomass and exchangeable Mg2+ . However, N inputs negated the promotional effects of increased precipitation, mainly through suppressing fungal growth and altering soil pH and clay-Mg2+ -OC bridging. Warming increased C content in the mineral-associated fraction, likely by increasing inputs of root-derived C, and reducing turnover of existing mineral-associated C due to suppression of fungal growth and soil respiration. Together, our results provide new insights into the potential mechanisms through which multiple global change factors control soil C persistence in arid and semi-arid grasslands. These findings suggest that the interactive effects among global change factors should be incorporated to predict the soil C dynamics under future global change scenarios.

Simulated warming enhances the responses of microbial N transformations to reactive N input in a Tibetan alpine meadow.

Alpine ecosystems worldwide are characterized with high soil organic carbon (C) and low mineral nitrogen (N). Climate warming has been predicted to stimulate microbial decomposition and N mineralization in these systems. However, experimental results are highly variable, and the underlying mechanisms remain unclear. We examined the effects of warming, N input, and their combination on soil N pools and N-cycling microbes in a field manipulation experiment. Special attention was directed to the ammonia-oxidizing bacteria and archaea, and their mediated N-cycling processes (transformation rates and N2O emissions) in the third plant growing season after the treatments were initiated. Nitrogen input (12 g m-2 y-1) alone significantly increased soil mineral N pools and plant N uptake, and stimulated the growth of AOB and N2O emissions in the late growing season. While warming (by 1.4 °C air temperature) alone did not have significant effects on most parameters, it amplified the effects of N input on soil N concentrations and AOB abundance, eliciting a chain reaction that increased nitrification potential (+83%), soil NO3--N (+200%), and N2O emissions (+412%) across the whole season. Also, N input reduced AOB diversity but increased the dominance of genus Nitrosospira within the AOB community, corresponding to the increased N2O emissions. These results showed that a small temperature increase in soil may significantly enhance N losses through NO3- leaching and N2O emissions when mineral N becomes available. These findings suggest that interactions among global change factors may predominantly affect ammonia-oxidizing microbes and their mediated N-cycling processes in alpine ecosystems under future climate change scenarios.

Combination of warming and N inputs increases the temperature sensitivity of soil N2O emission in a Tibetan alpine meadow.

Many high-elevation alpine ecosystems have been experiencing significant increases in air temperature and, to a lesser extent, nitrogen (N) deposition. These changes may affect N-cycling microbes and enhance emissions of nitrous oxide (N2O, a potent greenhouse gas) from soil. However, few studies have investigated whether and how the resulting changes in N-cycling microbes may affect the temperature sensitivity (Q10) of N2O emission and in turn feed back to N2O emissions. We conducted two incubation experiments to examine N2O emissions and their temperature sensitivities in soils that had experienced 3-yr field treatments of warming, N inputs and their combination in a Tibetan alpine meadow. Our results showed that neither N inputs nor warming alone affected the rate or Q10 of soil N2O emission, but combining the two significantly increased both parameters. Also, combined N and warming significantly increased the abundance of ammonia-oxidizing bacteria (AOB), corresponding with high soil N2O emission. In addition, N2O emission from nitrification accounted for 60-80% of total emissions in all soils, indicating that nitrifying microbes dominated the N2O production and its temperature sensitivity. Using random forest (RF) and structural equation model (SEM) analyses, we further evaluated the effects of various soil characteristics on soil N2O emissions and Q10. We identified soil moisture, pH, N mineralization and AOB abundance as the main predictors of the Q10 of N2O emissions. Together, these findings suggest that alterations in soil moisture, pH and ammonia-oxidizing bacteria induced by long-term N inputs and warming may increase temperature sensitivity of soil N2O emissions, leading to a positive climate feedback in this high-altitude alpine ecosystem.

Shifts in the Composition and Activities of Denitrifiers Dominate CO2 Stimulation of N2O Emissions.

Elevated atmospheric CO2 (eCO2) often increases soil N2O emissions, but the underlying mechanisms remain largely unknown. One hypothesis suggests that high N2O emissions may stem from increased denitrification induced by CO2 enhancement of plant carbon (C) allocation belowground. However, direct evidence illustrating linkages among N2O emissions, plant C allocation, and denitrifying microbes under eCO2 is still lacking. We examined the impact of eCO2 on plant C allocation to roots and their associated arbuscular mycorrhizal fungi and its subsequent effects on N2O emissions and denitrifying microbes in the presence of two distinct N sources, ammonium nitrogen (NH4+-N) and nitrate nitrogen (NO3--N). Our results showed that the form of the N inputs dominated the effects of eCO2 on N2O emissions: eCO2 significantly increased N2O emissions with NO3--N inputs but had no effect with NH4+-N inputs. eCO2 increased plant biomass N more with NH4+-N than with NO3--N inputs, likely reducing microbial access to available N under NH4+-N inputs and/or contributing to higher N2O emissions under NO3--N inputs. eCO2 enhanced root and mycorrhizal N uptake and also increased N2O emissions under NO3--N inputs. Further, eCO2 enhancement of N2O emissions under NO3--N inputs concurred with a shift in the soil denitrifier community composition in favor of N2O-producing (nirK- and nirS-type) over N2O-consuming (nosZ-type) denitrifiers. Together, these results indicate that eCO2 stimulated N2O emissions mainly through altering plant N preference in favor of NH4+ over NO3- and thus stimulating soil denitrifiers and their activities. These findings suggest that effective management of N sources may mitigate N2O emissions by negating the eCO2 stimulation of soil denitrifying microbes and their activities.

Contrasting Warming and Ozone Effects on Denitrifiers Dominate Soil N2O Emissions.

Nitrous oxide (N2O) in the atmosphere is a major greenhouse gas and reacts with volatile organic compounds to create ozone (an air pollutant) in the troposphere. Climate change factors such as warming and elevated ozone (eO3) affect N2O fluxes, but the direction and magnitude of these effects are uncertain and the underlying mechanisms remain unclear. We examined the impact of simulated warming (control + 3.6 °C) and eO3 (control + 45 ppb) on soil N2O fluxes in a soybean agroecosystem. Results obtained showed that warming significantly increased soil labile C, microbial biomass, and soil N mineralization, but eO3 reduced these parameters. Warming enhanced N2O-producing denitrifers ( nirS- and nirK-type), corresponding to increases in both the rate and sum of N2O emissions. In contrast, eO3 significantly reduced both N2O-producing and N2O-consuming ( nosZ-type) denitrifiers but had no impact on N2O emissions. Further, eO3 offsets the effects of warming on soil labile C, microbial biomass, and the population size of denitrifiers but still increased N2O emissions, indicating a direct effect of temperature on N2O emissions. Together, these findings suggest that warming may promote N2O production through increasing both the abundance and activities of N2O-producing microbes, positively feeding back to the ongoing climate change.

Atmospheric CO2 Enrichment and Reactive Nitrogen Inputs Interactively Stimulate Soil Cation Losses and Acidification.

Reactive N inputs (Nr) may alleviate N-limitation of plant growth and are assumed to help sustain plant responses to the rising atmospheric CO2 (eCO2). However, Nr and eCO2 may elicit a cascade reaction that alters soil chemistry and nutrient availability, shifting the limiting factors of plant growth, particularly in acidic tropical and subtropical croplands with low organic matter and low nutrient cations. Yet, few have so far examined the interactive effects of Nr and eCO2 on the dynamics of soil cation nutrients and soil acidity. We investigated the cation dynamics in the plant-soil system with exposure to eCO2 and different N sources in a subtropical, acidic agricultural soil. eCO2 and Nr, alone and interactively, increased Ca2+ and Mg2+ in soil solutions or leachates in aerobic agroecosystems. eCO2 significantly reduced soil pH, and NH4+-N inputs amplified this effect, suggesting that eCO2-induced plant preference of NH4+-N and plant growth may facilitate soil acidification. This is, to our knowledge, the first direct demonstration of eCO2 enhancement of soil acidity, although other studies have previously shown that eCO2 can increase cation release into soil solutions. Together, these findings provide new insights into the dynamics of cation nutrients and soil acidity under future climatic scenarios, highlighting the urgency for more studies on plant-soil responses to climate change in acidic tropical and subtropical ecosystems.