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Droughts can affect ecosystem CO2 fluxes directly or indirectly by changing plant community composition. However, it is unknown whether shifts in plant community composition buffer or amplify the response of ecosystem CO2 fluxes to droughts with different seasonal timing, as plant phenology and physiology of the different plant functional types respond differently to droughts. To identify the interaction of drought timing and plant community composition in regulating ecosystem CO2 fluxes, we conducted a three‐year manipulative experiment in which extreme droughts occurring in the early, mid and late growing seasons were separately imposed on experimental plot communities comprising graminoids, shrubs and their combination in a semi‐arid grassland of Inner Mongolia, China. Overall, mid‐season drought caused the largest negative effects regardless of plant community composition. In addition to decreasing aboveground biomass, mid‐season drought suppressed fluxes by reducing leaf photosynthetic rate, while early‐season and late‐season drought reduced fluxes mainly by shortening growing season length. All three community compositions had consistent responses to early‐season and mid‐season droughts. However, ecosystem CO2 fluxes in the combination community were less negatively affected by late‐season drought than in either shrub or graminoid communities because the growing season length was shortened less. Synthesis. Our results highlight that it is important to account for interactions of seasonal timing and plant community composition when predicting magnitude and pathways of drought effects on ecosystem carbon cycling. [ABSTRACT FROM AUTHOR]

Extreme climatic events are likely to intensify under climate change and can have different effects on ecosystems depending on their timing and magnitude. Understanding how productivity responds to extreme precipitation patterns requires assessing stability and vulnerability during critical growing periods at the plant community level. In this study, we experimentally imposed two contrasting types of extreme precipitation patterns, including extreme drought (excluding all rainfall for 30 consecutive days) during early-, mid-, and late-stages of the growing season, and heavy rainfall (adding 14.1 mm of rainfall every day for 20 consecutive days) during mid- and late-stages of the growing season over four years (2013–2016) in a steppe community in Inner Mongolia, China. We found that extreme drought and heavy rainfall had no effect on community aboveground net primary productivity (ANPP), species richness, and dominance at any stage of the growing season. Community stability in response to extreme drought was mainly driven by compensation among species and the stability of dominant species, while the compensatory effect among species and functional groups, and the stability of dominant species contributed to the community stability in response to heavy rainfall. Overall, our findings indicate that the responses of the ecosystem to intra-seasonal contrasting extreme precipitation patterns can be driven by similar stability mechanisms and suggest that semiarid temperate steppe communities may have strong initial resistance to more frequent extreme climatic events in the future. [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]

Summary: Heavy rainfall events are expected to increase in frequency and severity in the future. However, their effects on natural ecosystems are largely unknown, in particular with different seasonal timing of the events and recurrence over multiple years.We conducted a 4 yr manipulative experiment to explore grassland response to heavy rainfall imposed in either the middle of, or late in, the growing season in Inner Mongolia, China. We measured hierarchical responses at individual, community and ecosystem levels.Surprisingly, above‐ground biomass remained stable in the face of heavy rainfall, regardless of seasonal timing, whereas heavy rainfall late in the growing season had consistent negative impacts on below‐ground and total biomass. However, such negative biomass effects were not significant for heavy rainfall in the middle of the growing season. By contrast, heavy rainfall in the middle of the growing season had greater positive effects on ecosystem CO2 exchanges, mainly reflected in the latter 2 yr of the 4 yr experiment. This two‐stage response of CO2 fluxes was regulated by increased community‐level leaf area and leaf‐level photosynthesis and interannual variability of natural precipitation.Overall, our study demonstrates that ecosystem impacts of heavy rainfall events crucially depend on the seasonal timing and multiannual recurrence. Plant physiological and morphological adjustment appeared to improve the capacity of the ecosystem to respond positively to heavy rainfall. [ABSTRACT FROM AUTHOR]

Aims: Long-duration drought can alter ecosystem plant species composition with subsequent effects on carbon cycling. We conducted a rainfall manipulation field experiment to address the question: how does drought-induced vegetation change, specifically shrub encroachment into grasslands, regulate impacts of subsequent drought on soil CO2 efflux (Rs) and its components (autotrophic and heterotrophic, Ra and Rh)? Methods: We conducted a two-year experiment in Inner Mongolia plateau, China, using constructed steppe communities including graminoids, shrubs and their mixture (graminoid + shrub) to test the effects of extreme-duration drought (60-yr return time) on Rs, Rh and Ra. Results: Our results indicated that extreme-duration drought reduced net primary production, with subsequent effects on Rs, Rh and Ra in all three vegetation communities. There was a larger relative decline in Ra (35–54%) than Rs (30–37%) and Rh (28–35%). Interestingly, we found Rs in graminoids is higher than in shrubs under extreme drought. Meanwhile, Rh declines were largest in the shrub community. Although Ra and Rh both decreased rapidly during drought treatment, Rh recovered quickly after the drought, while Ra did not, limiting the Rs recovery. Conclusions: This study suggests that plant species composition regulates several aspects of soil CO2 efflux response to climate extremes. This regulation may be limited by above- and below-ground net primary production depending on soil water availability. The results of this experiment address a critical knowledge gap in the relationship between soil respiration and plant species composition. With shrub encroachment into grasslands, total soil respiration is reduced and can partly offset the effect of reduction in productivity under drought stress. [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]

• Non-linear patterns of microbial N genes occurred along precipitation gradient. • Breaking points were clearly related to certain soil types. • Attribution of different environmental factors varied by ecosystem. Despite the importance of microorganisms in soil nitrogen (N) cycling, studies on spatial patterns of microbial N-cycling gene abundances in temperate grasslands are still lacking, whose productivity is limited by N. Here, we investigated N functional genes in soil microorganisms from 60 sites in temperate grasslands across 1661 km in Inner Mongolia, China. Abundances of all N functional genes and 16S rDNA tended to decrease from northeast to southwest, consistent with the changing tend of precipitation but contrary to that of temperature. Non-linear saturation curves dominated patterns for abundances of most N functional genes and 16S rDNA along precipitation as they increased with the rising precipitation when <288–343 mm, but remained stable after these breaking points. Interestingly, these breaking points were clearly related to certain soil types, e.g. nifH did not change with precipitation in Chernozems and Gleysols, but increased with it in other soil types. Non-linear saturation patterns were also observed for most N functional gene and 16S rDNA abundances with temperature and soil total organic carbon. In contrast, no consistent spatial pattern for ratios of N functional genes: 16S rDNA was observed, which varied greatly by different N functional genes. Moreover, decay relationships were discovered for the abundance-based matrix of N functional genes over geographic distance. From the temperate meadows to typical steppes and to temperate desert steppes, the influence of relatively long-term environmental variables increased, but that of short-term ones decreased. Overall, we revealed non-linear patterns of N functional gene abundances along precipitation with a clear relationship with soil type, and discovered that attributions of geographic distance, plant community, historical-contingency and contemporary-disturbance to N functional gene community similarity decay were ecosystem–dependent. [ABSTRACT FROM AUTHOR]

• Soil EEAs and bacterial community composition responded differently to drought. • Soil water content was most important in affecting responses. • Changes in bacterial community composition were more persistent than soil EEAs. Extreme drought can strongly impact belowground communities and biogeochemical processes, including soil microbial community composition and extracellular enzyme activities (EEAs), which are considered key agents in ecosystem carbon (C) and nutrient cycling. However, our understanding of how seasonal timing of drought during the growing season affects soil microbial communities and their activity remains notably poor. In this study, we investigated the responses of soil physicochemical properties, EEAs, and bacterial community composition to extreme-duration drought imposed in the early-, mid-, or late-stages of the growing season in a semiarid grassland ecosystem in Inner Mongolia, China. Compared with the ambient control, the activities of C-, nitrogen (N)-, and phosphorus (P)-acquisition enzymes were significantly decreased in the mid- and/or late-stages of drought. Bacterial community diversity also significantly decreased in the mid- and late-stage drought treatments. Soil water content was the most important factor explaining changes in soil EEAs and bacterial community composition. At the end of the growing season, the activities of C-, N-, and P-acquisition enzymes had mostly recovered, while the bacterial community diversity in the mid- and late-stage drought treatments was still lower than the ambient control. Overall, our study demonstrates that the effects of extreme drought on soil EEAs and bacterial community composition depend on the timing of drought. Our results highlight that understanding the effects of extreme-duration drought at different stages of the growing season may play a vital role in predicting the responses of belowground function to global changes in grassland ecosystems. [ABSTRACT FROM AUTHOR]