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• Belowground biomass was positively sensitive to early and middle droughts. • Belowground biomass was negatively sensitive to late drought. • Aboveground biomass was insensitive to all droughts. • CO 2 fluxes had the largest sensitivity to early drought. • CO 2 fluxes had the lowest sensitivity to middle drought. Quantifying the sensitivity of ecosystems to droughts, particularly with different seasonal timing, could improve our predictions of ecosystem-climate feedbacks, but few experiments have explicitly addressed seasonal timing per se effects on ecosystem sensitivity to droughts. Here, we present a seasonal timing × drought manipulation experiment to examine sensitivity (relative change in response parameters to the relative change in precipitation) of key ecosystem processes (community biomass and ecosystem CO 2 fluxes) to pulse-drought with different seasonal timing (early, middle or late) on a temperate semiarid grassland. We found belowground and total biomass were positively sensitive (i.e. ecological processes promoted by droughts and vice versa) to early and middle droughts but negatively sensitive to late drought while aboveground biomass was insensitive to all droughts. Ecosystem CO 2 fluxes had the largest negative sensitivity to early drought and smallest negative sensitivity to middle drought, although gross ecosystem production showed larger negative response to droughts than ecosystem respiration, leading to reduction in net ecosystem production, regardless of seasonal timing. Our results highlight the crucial role of seasonal drought timing in regulating sensitivity of key carbon cycle processes to droughts and suggest that droughts at plant peak stage cause the least detrimental ecological consequences. [ABSTRACT FROM AUTHOR]

• Drought reduced the soil respiration (SR), especially during the treatment. • Heat wave, alone or combined with drought, had little effect on SR. • Seasonal dynamic of SR was regulated soil moisture and microbial activity. • Inter-annual variation in SR was positively related to plant biomass and structure. Increasing frequency and magnitude of climate extremes could fundamentally affect terrestrial carbon (C) cycling. However, as the second-largest terrestrial C flux, soil respiration (SR) responses to climate extremes are not well understood. Here, we investigated the effects of drought, heat wave and drought plus heat wave on SR in a semiarid grassland during the growing season over 3 years. The results indicated that drought consistently reduced the growing-season mean SR, especially during the period of drought treatment, while heat wave, alone or when combined with drought, had little effect on SR. The decreased SR under drought at the intra-annual timescale could be primarily attributable to restriction of low soil moisture to microbial biomass, as there were no consistent changes in plant community (aboveground biomass, richness and abundance). In contrast, the inter-annual variation in SR was positively related to plant community abundance, richness and aboveground biomass in additional to soil water availability, but was not significantly affected by microbial biomass. Our study highlighted that incorporating microbial biomass in C cycling models can improve simulation of seasonal dynamics of soil respiration while incorporating plant community characteristics can benefit prediction of variation in SR across multiple years in semiarid grasslands. [ABSTRACT FROM AUTHOR]

Droughts and heat waves are increasing in magnitude and frequency, altering the carbon cycle. However, understanding of the underlying response mechanisms remains poor, especially for the combination (hot drought). We conducted a 4‐year field experiment to examine both individual and interactive effects of drought and heat wave on carbon cycling of a semiarid grassland across individual, functional group, community and ecosystem levels. Drought did not change below‐ground biomass (BGB) or above‐ground biomass (AGB) due to compensation effects between grass and non‐grass functional groups. However, consistently decreased BGB under heat waves limited such compensation effects, resulting in reduced AGB. Ecosystem CO2 fluxes were suppressed by droughts, attributed to stomatal closure‐induced reductions in leaf photosynthesis and decreased AGB of grasses, while CO2 fluxes were little affected by heat waves. Overall the hot drought produced the lowest leaf photosynthesis, AGB and ecosystem CO2 fluxes although the interactions between heat wave and drought were usually not significant. Our results highlight that the functional group compensatory effects that maintain community‐level AGB rely on feedback of root system responses, and that plant adjustments at the individual level, together with shifts in composition at the functional group level, co‐regulate ecosystem carbon sink strength under climate extremes. • It is indispensable to explore the mechanisms and impacts of climate extremes on the carbon cycle given they are increasing in magnitude and frequency.• Aboveground biomass responses to climate extremes rely on the functional group compensatory effects and feedbacks of root system responses.• Plant physiological adjustments and shifts in community composition co‐regulate ecosystem CO2 fluxes under climate extremes. [ABSTRACT FROM AUTHOR]

• Precipitation was repackaged to simulate 4-level of precipitation variability. • Extra-extreme precipitation variability scenario potentially reduced N 2 O emissions. • Effects of precipitation variability on N 2 O fluxes had high interannual variations. • Soil, microbes, functional genes, and plants explained 61% variation in N 2 O fluxes. High temporal precipitation variability, characterized by less frequent but larger-magnitude precipitation events, is increasing. However, how precipitation variability affects N 2 O emissions and the underlying mechanisms remain unclear, especially at multiple-year scales. We conducted a 3-year manipulative experiment in which the same long-term mean growing season precipitation total was repackaged into events of inversely varying magnitude and frequency to simulate four levels of precipitation variability (extra-extreme, extreme, medium, and normal) in a semiarid grassland. Cumulative N 2 O emissions was the smallest under the extra-extreme precipitation variability scenario (6 very large rainfall events), 26% less than emission under the other three treatments (10, 16, and 24 rainfall events) over the three growing seasons. However, this difference was almost entirely due to three sampling events in the third year. Plant community (biomass and biodiversity), soil abiotic properties (water, dissolved organic carbon and pH), soil microbial biomass (carbon, nitrogen and the ratio), and soil functional genes (archaeal and bacterial amoA, nirS, nirK, narG , and nosZ) explained 61% of the variation in N 2 O emissions in response to the precipitation variability. Structural equation modelling indicated that the precipitation variability had direct positive effects, and indirect negative effects via soil abiotic properties, on soil functional genes that ultimately had positive effects on N 2 O emissions. The reduction in N 2 O emission in the extra-extreme precipitation variability scenario in the third year was due to low levels of soil water content, soil pH, aboveground biomass, and microbial biomass carbon simultaneously. Our results suggest that at the multi-year timescale, semiarid grasslands may have negative feedbacks to future precipitation regimes with higher variability. [Display omitted] [ABSTRACT FROM AUTHOR]

Nitrous oxide (N2O) emissions from soil contribute to global warming and are in turn substantially affected by climate change. However, climate change impacts on N2O production across terrestrial ecosystems remain poorly understood. Here, we synthesized 46 published studies of N2O fluxes and relevant soil functional genes (SFGs, that is, archaeal amoA, bacterial amoA, nosZ, narG, nirK and nirS) to assess their responses to increased temperature, increased or decreased precipitation amounts, and prolonged drought (no change in total precipitation but increase in precipitation intervals) in terrestrial ecosystem (i.e. grasslands, forests, shrublands, tundra and croplands). Across the data set, temperature increased N2O emissions by 33%. However, the effects were highly variable across biomes, with strongest temperature responses in shrublands, variable responses in forests and negative responses in tundra. The warming methods employed also influenced the effects of temperature on N2O emissions (most effectively induced by open‐top chambers). Whole‐day or whole‐year warming treatment significantly enhanced N2O emissions, but daytime, nighttime or short‐season warming did not have significant effects. Regardless of biome, treatment method and season, increased precipitation promoted N2O emission by an average of 55%, while decreased precipitation suppressed N2O emission by 31%, predominantly driven by changes in soil moisture. The effect size of precipitation changes on nirS and nosZ showed a U‐shape relationship with soil moisture; further insight into biotic mechanisms underlying N2O emission response to climate change remain limited by data availability, underlying a need for studies that report SFG. Our findings indicate that climate change substantially affects N2O emission and highlights the urgent need to incorporate this strong feedback into most climate models for convincing projection of future climate change. [ABSTRACT FROM AUTHOR]

• N 2 O flux is sensitive and insensitive to drought and heat wave, respectively. • Soil water content is a first-order predictor of seasonal variation of N 2 O flux. • Inter-annual variations of N 2 O flux are impacted by precipitation distribution. • Archaeal amoA and nirK mainly regulates climate extremes effects on N 2 O flux. Nitrous oxide (N 2 O), the third most important greenhouse gas, contributes to the increasing frequency and severity of climate extremes. Disentangling feedbacks of climate extremes on terrestrial N 2 O emission is important for forecasting future climate changes. Here, we experimentally imposed extreme drought and heat wave events during three years in a semiarid grassland to investigate the responses of N 2 O flux. We identified that N 2 O flux suppression during droughts was mediated by soil water content (SWC), microbial biomass carbon (MBC), soil inorganic nitrogen (SIN) and dissolved organic carbon (DOC) contents, and the abundance of archaeal amoA, nirK, and narG. However, bacterial amoA, nirS, and nosZ remained stable. Upon rewetting following droughts, the SWC, SIN, DOC, archaeal amoA, nirK, narG, and resultant N 2 O fluxes recovered to the magnitude of the ambient control. In contrast, heat waves alone or in combination with drought did not impact N 2 O fluxes or the underlying physical, chemical and microbial states. Stepwise multiple linear regression suggested that SWC, DOC, and MBC were the key factors regulating immediate responses of N 2 O flux to climate extremes while the major factors regulating seasonal mean N 2 O flux in response to climate extremes were archaeal amoA abundance, nirK abundance, and MBC. Our results suggest that N 2 O fluxes were sensitive to droughts but insensitive to heat waves. Soil moisture induced changes in substrate availability, and the community size of total and functional microorganisms in soil jointly regulated N 2 O responses to climate extremes. The relative importance of regulating factors shifted at different timescales. [ABSTRACT FROM AUTHOR]

Increasing heavy rainfalls can strongly affect ecosystem nitrogen (N) cycling processes and thereby alter soil N fluxes and pools. However, the effects of heavy rainfalls on soil N fluxes and pools are poorly understood, particularly with regards to high rainfall timing under field conditions. We conducted a 3-year (2014–2016) manipulative experiment in which heavy rainfall was imposed in middle (plant peak growing stage) or late (plant senescent growth stage) growing season in a semiarid grassland of Inner Mongolia, China to explore the responses of N 2 O fluxes and soil total N contents and the underlying microbial mechanisms. Mid-season heavy rainfall promoted soil N 2 O emissions by 65% on average across the three years, attributable mainly to increases in denitrifying nirK and nirS abundances induced large denitrification at higher soil water contents. However, archaeal and bacterial amoA and narG genes did not change significantly due probably to counteracting effects of increased soil water content (positive) and soil pH (negative). Mid-season heavy rainfall led to 17% reduction in soil total N by the end of the last year of the study (2016), partly due to the enhanced accumulated N 2 O emissions over the three years. In contrast, late-season heavy rainfall did not change N 2 O emissions and soil total N contents even though soil water content, soil pH and nirK and nirS abundance were significantly increased, perhaps due to limitation by low temperature. Timing of the heavy rainfall events during the plant growing season strongly influenced their impacts on soil N fluxes and pools and heavy rainfalls in the peak stage of plant growth may potentially cause a positive feedback to global warming and exacerbate N limitation in terrestrial ecosystems. • N 2 O flux and soil total N responses to heavy rainfall depended on seasonal timing. • Mid-season heavy rainfall increased N 2 O flux by 65% due to higher denitrification. • Enhanced N 2 O emissions coincided with 17% of reduction in soil total N. • Late-season heavy rainfall did not change N 2 O emissions and soil total N contents. [ABSTRACT FROM AUTHOR]

Grazing is expected to exert a substantial influence on antibiotic resistance genes (ARGs) in grassland ecosystems. However, the precise effects of grazing on the composition of ARGs in grassland soils remain unclear. This is especially the case for grassland soils subject to long-term grazing. Here, we investigated ARGs and bacterial community composition in soils subject to long-term historic grazing (13–39 years) and corresponding ungrazed samples. Using a combination of shotgun metagenomics, amplicon analyses and associated soil physicochemical data, we provide novel insights regarding the structure of ARGs in grassland soils. Interestingly, our analysis revealed that long-term historic grazing had no impacts on the composition of ARGs in grassland soils. An average of 378 ARGs, conferring resistance to 14 major categories of antibiotics (80%), were identified in both grazing and ungrazed sites. Actinobacteria , Proteobacteria and Acidobacteria were the most prevalent predicted hosts in these soils and were also shown to harbour genetic capacity for multiple-resistant ARGs. Our results suggested that positive effects of bacterial community composition on ARGs could potentially be controlled by affecting MGEs. Soil properties had direct effects on the composition of ARGs through affecting the frequency of horizontal gene transfer among bacteria. Twelve novel ARGs were found in S. grandis steppe grasslands, indicating that different vegetation types might induce shifts in soil ARGs. Collectively, these findings suggest that soil properties, plants and microorganisms play critical roles in shaping ARG patterns in grasslands. Together, these data establish a solid baseline for understanding environmental antibiotic resistance in grasslands. Unlabelled Image • ARGs related with resistance to cephalosporin and tetracycline were dominant. • ARGs driven by factors affecting the distribution of soil microorganisms in grasslands. • Long-term historic grazing had no effect on ARGs in grassland soils. • ARGs shaped by the initial plant, soil environmental parameters and microbiomes in grassland. [ABSTRACT FROM AUTHOR]