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Recently, the Qinghai–Tibetan Plateau has experienced significant warming. Climate warming is expected to have profound effects on plant community productivity and composition, which can drive ecosystem structure and function. To explore effects of warming on plant community productivity and composition, we conducted a warming experiment using open top chambers (OTCs) from 2012 to 2014 in alpine meadow and alpine steppe habitat on the central Qinghai–Tibetan Plateau. We measured aboveground net primary productivity (ANPP), community composition and species diversity under ambient and two levels of artificially warmed conditions across three years. Our results showed that warming significantly stimulated plant growth in the alpine meadow, but reduced growth on the alpine steppe. The increase of ANPP in alpine meadow was a result of an increase of plant height under warming. Warming-induced drought conditions were primarily responsible for the observed decrease of ANPP in an alpine steppe. Plant community composition and species diversity were not influenced by warming in alpine meadow. Alternatively, in alpine steppe, cover of graminoids and forbs significantly declined while legumes substantially increased under warming, subsequently resulting in rapid species losses. Changes in soil moisture were responsible for observed changes in graminoids and legumes in the alpine steppe. Overall, experimental results demonstrated that warming had a positive impact on plant community structure and function in alpine meadow and had a negative impact on these characteristics in an alpine steppe. This work highlights the important role of soil moisture for regulating plant productivity and community composition response to warming in the alpine steppe. In particular, the deep-rooted, drought resistant plants may increase in a warmer future in the central Qinghai–Tibetan Plateau. These changes may reduce habitat quality for the local community of grazers because many of the species that increased are also unpalatable to grazers. [ABSTRACT FROM AUTHOR]

To analyze CO2 fluxes under conditions of climate change in an alpine meadow on the central Qinghai–Tibetan Plateau, we simulated the effect of warming using open top chambers (OTCs) from 2012 to 2014. The OTCs increased soil temperature by 1.62°C (P < 0.05), but decreased soil moisture (1.38%, P < 0.05) during the experiments. The response of ecosystem CO2 fluxes to warming was variable, and dependent on the year. Under conditions of warming, mean gross ecosystem productivity (GEP) during the growing season increased significantly in 2012 and 2014 (P < 0.05); however, ecosystem respiration (ER) increased substantially only in 2012 (P < 0.05). The net ecosystem CO2 exchange (NEE) increased marginally in 2012 (P = 0.056), did not change in 2013(P > 0.05), and increased significantly in 2014 (P = 0.034) under conditions of warming. The GEP was more sensitive to climate variations than was the ER, resulting in a large increase in net carbon uptake under warming in the alpine meadow. Under warming, the 3-year averages of GEP, ER, and NEE increased by 19.6%, 15.1%, and 21.1%, respectively. The seasonal dynamic patterns of GEP and NEE, but not ER, were significantly impacted by warming. Aboveground biomass, particularly the graminoid biomass increased significantly under conditions of warming. Soil moisture, soil temperature, and aboveground biomass were the main factors that affected the variation of the ecosystem CO2 fluxes. The effect of warming on inter- and intra-annual patterns of ecosystem CO2 fluxes and the mechanism of different sensitivities in GEP and ER to warming, require further researched. [ABSTRACT FROM AUTHOR]

Rare species can switch between abundant and rare, respond to various external disturbances, enhance community resilience, and reduce the lag in ecosystem recovery. However, the impact of N enrichment on microbial community dynamic changes in rare species during the growing season is still poorly understood. To clarify which taxa contribute most to the community variation under N enrichment and whether the responses of these taxa vary at different periods of the growing season, we set up a 6-years multi-level field N addition experiment in an alpine meadow on the Tibetan Plateau. The variation in soil bacterial community diversity, composition, species interactions, and assembly process of different abundance profiles were investigated in the early, mid, and late-growing seasons. Approximately 80% of bacteria were classified as rare taxa (including permanently and conditionally rare taxa). The influence of N enrichment on the bacterial community was seasonally dependent. Bacterial community diversity and composition dissimilarity significantly increased only in the early growing season, especially in permanently rare taxa and conditionally rare taxa, but remained stable in the mid-and late-growing seasons. Seasonal fluctuations led to a shift in community composition, and conditionally rare taxa were the primary component of seasonal variation in community composition. Simultaneously, microbial network complexity and stability gradually increased over time, as evidenced by decreased modularity and vulnerability and increased robustness, clustering coefficient, connectivity, and keystone nodes. The assembly of conditionally rare taxa showed a significant difference under N addition, presented as decreased drift and increased dispersal limitation, which are closely related to the composition of the entire bacterial community. Our findings emphasize that the conditionally rare taxa decisive contribute to changes in the bacterial community under N enrichment, but seasonal fluctuations can make bacterial communities resistant to the influence of N enrichment. [ABSTRACT FROM AUTHOR]

[Display omitted] Structural information of grassland changes on the Tibetan Plateau is essential for understanding alterations in critical ecosystem functioning and their underlying drivers that may reflect environmental changes. However, such information at the regional scale is still lacking due to methodological limitations. Beyond remote sensing indicators only recognizing vegetation productivity, we utilized multivariate data fusion and deep learning to characterize formation-based plant community structure in alpine grasslands at the regional scale of the Tibetan Plateau for the first time and compared it with the earlier version of Vegetation Map of China for historical changes. Over the past 40 years, we revealed that (1) the proportion of alpine meadows in alpine grasslands increased from 50% to 69%, well-reflecting the warming and wetting trend; (2) dominances of Kobresia pygmaea and Stipa purpurea formations in alpine meadows and steppes were strengthened to 76% and 92%, respectively; (3) the climate factor mainly drove the distribution of Stipa purpurea formation, but not the recent distribution of Kobresia pygmaea formation that was likely shaped by human activities. Therefore, the underlying mechanisms of grassland changes over the past 40 years were considered to be formation dependent. Overall, the first exploration for structural information of plant community changes in this study not only provides a new perspective to understand drivers of grassland changes and their spatial heterogeneity at the regional scale of the Tibetan Plateau, but also innovates large-scale vegetation study paradigm. [ABSTRACT FROM AUTHOR]

• The relative abundance of soil rare biosphere was stable under climate changes. • The specific soil microbial taxa strongly related to soil rare biosphere. • The rare biosphere strongly explained variations of soil available nitrogen. Ecological roles of "rare biosphere" (RB) were rarely discussed in alpine grassland ecosystems. We explored the roles of RB and specific taxa of soil bacteria and fungi in regulating soil nutrients and aboveground biomass of an alpine meadow and alpine steppe under warming and enhanced rainfall scenarios on the Qinghai-Tibetan plateau. In the context of alpine meadow and alpine steppe, the relative abundance of bacterial RB was ∼40% in the bacterial community, and it was <10% for fungal RB in the fungal community. The relative abundance of the RB of soil bacteria and fungi were not changed by warming or enhanced rainfall. All specific bacterial taxa and over 90% specific fungal taxa belonged to the RB. The RB, including the most specific taxa, explained many more variations in the contents of soil NO 3 -N and NH 4 -N than the high abundant microbial species under warming and enhanced rainfall conditions. The aboveground biomass of the alpine meadow and steppe were strongly affected by the contents of soil NO 3 -N, available phosphorus and total nitrogen under the condition of warming and enhanced rainfall. Our findings suggest that the robust soil microbial RB plays an important role in regulating the content of soil available nitrogen, which could profoundly affect aboveground plant biomass of alpine grassland ecosystems under climate change conditions. [ABSTRACT FROM AUTHOR]

Phenology is an important indicator of plant responses to environmental changes and is closely correlated with biomass production. However, how changes in phenological events affect plant biomass production when exposed to changing temperature and precipitation remain unclear.We conducted a 4‐year manipulative experiment of warming and precipitation addition to explore phenology–biomass interactions under climate change in a dry alpine meadow on the central Qinghai–Tibetan Plateau from 2015 to 2018.In dry and warm years, warming delayed phenology and precipitation addition advanced them. Warming decreased the biomass of Kobresia pygmaea in 2018 and the biomass of Poa pratensis in 2015, 2017 and 2018. However, precipitation addition significantly increased the biomass of Poa pratensis and Potentilla multifida in most of the experimental years. Phenological changes regulated the responses of biomass to treatments. Specifically, delay of green up of P. pratensis and delay of withering of K. pygmaea induced by warming can increase biomass production, but it can be offset by the direct negative effects of warming on biomass.Synthesis. Here we show how warming‐induced drought tend to decrease the biomass production of graminoids and the negative effects of warming on the biomass of P. pratensis and K. pygmaea were partially offset by green up postponement and withering postponement respectively. Our results highlights phenology is a crucial regulator for biomass production under climate change. Hence, both direct and indirect effects of warming and precipitation addition on phenology and biomass cannot be ignored when predicting biomass responses to climate change. [ABSTRACT FROM AUTHOR]

Atmospheric concentrations of nitrous oxide (N 2 O) have increased over the last 150 years due to human activities. Soils are important source of N 2 O where its production is largely regulated by biological processes. Nitrogen (N) deposition can alter the processes of autotrophic nitrification and denitrification, thus affecting the rate and direction of the soil N 2 O exchange with the atmosphere. It is obvious that atmospheric N deposition has influenced a dry alpine meadow in the Qinghai–Tibetan Plateau (QTP). Therefore, we investigated the effects of N deposition on the annual N 2 O emissions in an alpine meadow of the QTP. The N 2 O flux was measured for 2 years using static chambers and gas chromatography methods at four treatments (N0, background level; N7, add 7 kg N ha −1  yr −1 ; N20, add 20 kg N ha −1  yr −1 ; N40, add 40 kg N ha −1  yr −1 ). We found that high N deposition increased the N 2 O flux, not only in the growing season but also in winter and the spring thaw period. The average annual N 2 O fluxes at N0 and N40 plots were 3.1 and 6.1 μg m −2  h −1 , respectively. Compared with N0, the average annual N 2 O fluxes increased at N7, N20, and N40 plots by 13.7%, 47.6%, and 98.7%, respectively, the N 2 O fluxes in N40 plots increased by 113.6%, 41.6%, and 78.7% during the growing season, winter and spring thaw period, respectively. The emission of N 2 O during the growing season, winter and spring thaw period accounted for 63.2%, 29.9% and 6.9% of annual total emission of N 2 O, respectively. The N 2 O flux correlated significantly with air temperature, soil temperature, soil water content, total nitrogen, total organic carbon, and soil NH 4 + –N content in the alpine meadow. [ABSTRACT FROM AUTHOR]

• Alpine meadow had higher CUE and lower WUE than cultivated grassland under warming. • Warming decreased biomass of cultivated grassland in the later of the experiment. • Warming increased ET, did not affect GEP, and decreased WUE in alpine meadow. • Warming decreased NEE, did not alter ER, and decrease CUE in cultivated grassland. Carbon and water use efficiency (CUE and WUE) are crucial indicators of grassland ecosystem responses to climate change. Grass planting is one of the dominant methods of grassland restoration and agricultural production. However, how warming and reclamation impact grassland CUE and WUE remain unclear. Here we examined how warming affected carbon and water processes in natural alpine meadow and cultivated grassland, including gross ecosystem production (GEP), net ecosystem CO 2 exchange (NEE), evapotranspiration (ET), CUE and WUE. We deployed a seven-year warming experiment using open top chambers in alpine meadow and cultivated grassland on the Tibetan Plateau. Our results showed that the alpine meadow had a higher CUE and a lower WUE compared to the cultivated grassland under warming. Warming increased biomass production of the cultivated grassland in the beginning of the experiment but decreased it later in the experiment. Warming increased ET, did not affect GEP, and finally decreased WUE in the natural alpine meadow. In the cultivated grassland, warming decreased NEE, did not alter ER, and resulted the decrease in CUE. Our results highlight that reclamation decreases CUE and warming decreased WUE of alpine meadow. Although grass planting can increase biomass, the negative impacts of warming on carbon and water processes of cultivated grassland should be considered when reclaiming alpine meadow under warmer future. [ABSTRACT FROM AUTHOR]

Alpine grasslands on the Qinghai-Tibetan Plateau are sensitive and vulnerable to climate change and human activities. Climate warming and overgrazing have already caused degradation in a large fraction of alpine grasslands on this plateau. However, it remains unclear how human activities (mainly livestock grazing) regulates vegetation dynamics under climate change. Here, alpine grassland productivity (substituted with the normalized difference vegetation index, NDVI) is hypothesized to vary in a nonlinear trajectory to follow climate fluctuations and human disturbances. With generalized additive mixed modelling (GAMM) and residual-trend (RESTREND) analysis together, both magnitude and direction of climatic (in terms of temperature, precipitation, and radiation) and anthropogenic impacts on NDVI variation were examined across alpine meadows, steppes, and desert-steppes on the Qinghai-Tibetan Plateau. The results revealed that accelerating warming and greening, respectively, took place in 76.2% and 78.8% of alpine grasslands on the Qinghai-Tibetan Plateau. The relative importance of temperature, precipitation, and radiation impacts was comparable, between 20.4% and 24.8%, and combined to explain 66.2% of NDVI variance at the pixel scale. The human influence was strengthening and weakening, respectively, in 15.5% and 14.3% of grassland pixels, being slightly larger than any sole climatic variable across the entire plateau. Anthropogenic and climatic factors can be in opposite ways to affect alpine grasslands, even within the same grassland type, likely regulated by plant community assembly and species functional traits. Therefore, the underlying mechanisms of how plant functional diversity regulates nonlinear ecosystem response to climatic and anthropogenic stresses should be carefully explored in the future. • Generalized additive model revealed nonlinear climatic and anthropogenic impacts on ecosystems. • Climate change dominantly drove grassland productivity changes on the Qinghai-Tibetan Plateau. • Human impacts on grassland productivity on the Qinghai-Tibetan Plateau weakened in past decades. [ABSTRACT FROM AUTHOR]

• The effect of grazing regime on functional genes were different at distinct grazing period. • Grazing regime affected the temporal variation of soil microbial functional genes. • Grazing rather than exclosure promote C fixation and decomposition and N cycling. • There existed trade-offs between plant and soil nutrients along the grazing time. • Continuous grazing promoted CH 4 metabolism. Microbial functional genes can reflect the nutrient cycle activities in soil, because they encode various enzymes involved in the material cycle. Therefore, the gene abundance variation can reveal the impact of interference on the material cycle of grassland. To study the cycle of carbon (C), nitrogen (N) and phosphorus (P) between plant and soil in grassland under different grazing regimes, we investigated the soil microbial functional genes related to C, N, and P cycling by high-throughput quantitative PCR and 16S rRNA-based Illumina sequencing analysis under grazing exclusion (GE), rotational grazing (RG), and continuous grazing (CG) in alpine meadow of the Qinghai-Tibetan Plateau, where climate is characterized by little rain and low temperature, and grassland is very sensitive to grazing. The results showed at the early grazing period, C fixtion (rbcL , korA , and frdA) and lignin degradation (abfA , xylA , exg , lig , exoPG , chiA , and glx) processes were slower under GE; CH 4 metabolism (mcrA) was faster under CG; RG and CG improved the denitrification process (narG); RG slowed down organic-P mineralization (phoD). At the late grazing period, C fixation (accA and frdA) and degradation (mnp , apu , and amyA) processes were slower under GE; CH 4 metabolism (pmoA and mxa) was faster under CG; RG and CG improved the ammonia-oxidizing (amoA2), nitrification (hao), and denitrification (nirS3 and nirK1) processes. The majority of the genes involved in C, N, and P cycling decreased, the C, N and P content in plant leaf decreased, while that of soil increased from early to late grazing period. No matter grazing or not, there were negative relationships between genes and soil nutrients, and positive relationships between genes and plant nutrients, implying a trade-off between plant nutrients and soil nutrients along the grazing time. The genes responsible for regulating C and N cycling were increased under grazing, implying that reasonable grazing is beneficial to the nutrients cycling of grassland. [ABSTRACT FROM AUTHOR]

• Responses of green up to warming were different among years. • Water addition did not advance community phenology except in a dry and warm year. • Warming increased annual variability of plant green up and flowering. • Flowering of K. pygmaea advanced under warming plus water addition in all three years. • Soil moisture plays a critical role in determining phenological response to warming. Temperature and precipitation are primary regulators of plant phenology. However, our knowledge of how these factors might interact to affect plant phenology is incomplete. The Qinghai-Tibetan Plateau, a cold and high region, has experienced no consistent changes in spring phenology, despite a significant warming trend. We conducted a manipulative experiment of warming and precipitation addition in an alpine meadow on the Qinghai-Tibetan Plateau in 2015 (cold and wet), 2016 (warm and dry) and 2017 (mild and very wet). We found that warming increased annual variability of plant spring phenology. Warming delayed green up of all monitored species in 2016, advanced green up of early flowering species in 2015, and did not alter green up in 2017. For example, green up of the shallow rooted Kobresia pygmaea advanced 8 (± 2) days in 2015 and was delayed by 23 (± 3) days in a dry year (2016) under warming compared with control. Early spring precipitation addition can offset the delaying effects of warming in a dry year on the Qinghai-Tibetan Plateau. Under warming plus precipitation addition, community average green up advanced compared to control plots in 2015 and 2016, and community average flowering advanced for all three years. In 2016, flowering of K. pygmaea (an early flowering species) advanced under warming plus precipitation addition compared to control while flowering of other species did not change. Our results highlight that annual variation of soil moisture condition plays a critical role in determining the magnitude and direction of spring phenology response to warming. We provide insights in how plant spring phenology might change in a warmer future in the presence or absence of precipitation increase. [ABSTRACT FROM AUTHOR]