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Temperature is conventionally considered as the dominant factor regulating carbon fluxes of the alpine meadow on the Tibetan Plateau, while contribution from water availability is composed of large uncertainty. In this study, eddy covariance (EC) data were used to assess the relative contribution of temperature and water availability to carbon fluxes of the alpine meadow ecosystem. The results showed that soil water content (SWC) was the most important factor controlling carbon fluxes – Net Ecosystem Productivity (NEP), Gross Primary Productivity (GPP) and Ecosystem Respiration (Re). The GPP and Re increased with strengthened SWC under any temperature conditions, indicating the dominant control of water availability on carbon fluxes. In addition, water availability regulated the response size of ecosystem to temperature, and could alleviate the stress caused by low temperature. The photosynthesis capacity of alpine plants at noon was depressed by water stress rather than by high temperature. The structural equation modeling (SEM) analysis further confirmed the dominance of SWC on the carbon fluxes. This study implies that effects of climatic change on this alpine ecosystem might be more induced by changes in water pattern than increased temperature, which provides new insights into the climate controls of carbon fluxes over alpine meadow, and adds to our understanding on climate change impacts on carbon cycling on the Tibetan Plateau. [ABSTRACT FROM AUTHOR]

The effects of warming-shifted plant phenology on ecosystem carbon (C) cycling have received increasing attention in recent years. However, there is a lack of evidence and mechanistic understanding of how warming-shifted plant phenology influences ecosystem C cycling. In this study, we conducted a field experiment to investigate the effects of warming on phenology and ecosystem C exchange in a Tibetan alpine meadow during the 2014 and 2015 growing seasons. Our results indicated that warming led to later green-up in spring by aggravating water limitation but little change in autumn phenology, resulting in shortened growing season length. Interestingly, we found warming caused a seasonal shift of ecosystem C exchange. During the early summer monsoon, ecosystem C uptake was suppressed by warming due to the delay of phenological development. However, warming accelerated ecosystem C uptake and promoted ecosystem C uptake under ample water conditions during the late summer monsoon. As a result, although warming shortened the growing season length, it had no significant effects on gross primary production (GPP) and net ecosystem production (NEP). Our results will improve our understanding of the mechanisms of how warming-shifted plant phenology influences ecosystem C cycling in semiarid alpine ecosystems. [ABSTRACT FROM AUTHOR]

Highlights • Grazing could preserve the deep-layer soil moisture by reducing transpiration. • Higher productivity was maintained in grazed alpine meadow during droughts. • Younger leaves in grazed meadow could restore productivity quickly after droughts. Abstract Grazing is the primary land use practice in alpine ecosystems on the Tibetan Plateau. However, it remains unclear how grazing regulates levels of carbon and the water cycle in this ecosystem. A paired set of eddy covariance systems were set in adjacent fenced (FM) and grazed meadows (GM) to explore the grazing effects on alpine meadows. Aboveground biomass removed by grazing caused declines in grass transpiration (T), whereas the evaporation (E) was enhanced because of greater exposure to radiation, which in turn led to higher evapotranspiration (ET) in GM. However, the deep-layer soil moisture remained high because of the effects of mattic epipedon, which worked as a water-resistant layer. Therefore, the deep-layer soil moisture in GM was higher than that in FM because of decreased water consumption caused by the grazing-induced reduction in leaf area in GM. As a consequence, the deep-layer soil in GM could provide more water to help plants endure droughts. Additionally, grazing enhanced the sensitivity of productivity to soil moisture during the wet season in drought years, which could restore grassland productivity more quickly after droughts. Therefore, grazing could help this fragile ecosystem to withstand droughts to some extent and maintain higher productivity. This may provide us with an ecological approach for confronting global climate change. [ABSTRACT FROM AUTHOR]

• Atmospheric and soil droughts together greatly depressed GPP. • Atmospheric water deficits had stronger instant inhibited effects on GPP. • Drought-induced H enhancements provided positive feedback and aggravated GPP loss. Under global climate change, warmer temperatures and changing precipitation patterns will increase the relative importance of soil and atmospheric droughts in limiting productivities across different ecosystems, especially in the fragile and sensitive ecosystem on the Tibetan Plateau. Therefore, the present study examined the effects of atmospheric drought, soil drought and compound drought on alpine meadow productivity based on seven years of continuous observational data. The results demonstrated that water conditions dominated the variations in alpine meadow productivity in northern Tibet. Gross prime productivity (GPP) in the growing season was significantly inhibited during soil drought (defined as soil water content (SWC) < 0.11 m3 m−3) or atmospheric drought (defined as vapor pressure deficit (VPD) > 0.61 kPa) conditions. Pooling of the moisture data according to these thresholds revealed that soil drought occurred more frequently and accounted for 20.4% of all observed days. Atmospheric drought occurred in 2.4% of all days, and the frequency of compound drought (when SWC < 0.11 m3 m−3 and VPD > 0.61 kPa) was 8.7%. Compound drought reduced GPP by the largest extent to as much as 1.47 g C m−2 d−1. Atmospheric drought reduced GPP by 1.10 g C m−2 d−1. However, the more frequent soil drought reduced GPP by the smallest extent, with a value of 0.69 g C m−2 d−1. In spite of this, soil drought played an important role in restricting the accumulated GPP of this alpine meadow via its higher occurrence frequency and longer duration. All three types of drought significantly increased sensible heat flux (H), which provided positive feedbacks to droughts and led to further decreases in GPP. The results of the present study suggest that drought greatly threatens the alpine meadow ecosystem in northern Tibet. Therefore, more attention should be paid to water conditions under the global climate change context. [ABSTRACT FROM AUTHOR]

• Droughts enhanced the control effects of biotic factors on carbon and water fluxes. • Droughts induced changes in the driving paths of biotic and abiotic factors on RUE. • RUE responded differently to droughts in different periods in the growing season. • Droughts in the mid-growing season reduced the RUE. Drought is a threat universally faced by global terrestrial ecosystems under global climate change. The habitat of alpine ecosystems is harsh, and they necessitate to use the limited resources efficiently to survive. However, drought occurred frequently during the short growing season, which greatly affected their resource use efficiency (RUE). In this study, based on long-term flux and microclimate observations in a typical alpine meadow ecosystem in northern Tibet, the effects of drought on RUE, including water use efficiency (WUE), light use efficiency (LUE), and carbon use efficiency (CUE), were studied. The results indicate that droughts could change the driving paths of abiotic and biotic factors on carbon and water fluxes and hence affect RUE. When vegetation exhibits robust physiological activity, droughts could enhance the control effects of biotic factors. The droughts that occurred at different periods of the growing season showed divergent effects on RUE. In the early-growing season, the strong photosynthetic capacity of the nascent leaves facilitated enhancements in WUE and LUE in response to drought conditions. However, the CUE did not increase significantly due to the heightened consumption of organic matter by respiration. The mid-growing season droughts led to a sharp decrease in gross primary productivity (GPP), consequently leading to a decline in WUE, LUE and CUE. During the late-growing season, droughts together with the decomposition of chlorophyll reduced the photosynthetic capacity and led to a significant decrease in LUE. The diminished physiological activity of the senescent leaves led to insensitive responses of WUE and CUE to droughts. These findings indicate that RUE might primarily be regulated by factors affecting GPP rather than others. This study would be beneficial for assessing the responses and adaptation characteristics of alpine meadow ecosystems to future climate change and could help accurately predict their possible evolution trends. [ABSTRACT FROM AUTHOR]

• The soil water content and temperature determined the maximum photosynthesis. • Sufficient soil water elevated the optimal temperature and maximum photosynthesis. • The optimal environmental configuration for the alpine meadow was detected. • If soil water is high, higher temperature and VPD are beneficial for carbon uptake. The changes in the carbon sink function of sensitive and fragile alpine ecosystems under climate change are widely concerned. Radiation, temperature, and water resources play important roles in regulating carbon assimilation. Therefore, it is important to detect the optimal configuration of environmental factors for the photosynthesis of alpine meadow ecosystems. In this study, based on 10-year flux and corresponding environmental observations in a typical alpine meadow ecosystem, the optimal environmental configuration that could realistically occur in the natural world for photosynthesis was detected. The results indicated that the temperature and water conditions were the primary limiting factors, as radiation resources are abundant on the Tibetan Plateau. The optimal temperature for photosynthesis (TA opt) was determined to be 11.92 ℃ based on the entire observation dataset, with a maximum photosynthetic capacity (GPP max) of 0.20 mg CO 2 m−2 s−1. However, the TA opt and the corresponding GPP max could be elevated when the soil water content (SWC) was abundant. When SWC exceeded 0.26 m3 m−3, TA opt increased to 13.41 ℃, and the corresponding GPP max reached as high as 0.30 mg CO 2 m−2 s−1, which is the maximum the ecosystem could reach. Thus, the optimal environmental configuration for photosynthesis that could realistically occur in the natural world in this alpine meadow was air temperature (TA)=13.41 ℃, SWC=0.28 m3 m−3, vapor pressure deficit (VPD)=0.61 kPa, net radiation (RN)=411.53 J m−2 s−1. According to this study, it could be inferred that within a reasonable range, higher temperatures and VPD would benefit photosynthesis in this alpine meadow as long as the soil water supply is sufficient. The climate on the Tibetan Plateau is arid with high VPD and is experiencing warming due to climate change. Consequently, alpine ecosystems situated in relatively humid regions with sufficient soil water resources may exhibit greater potential for carbon sequestration than previously estimated. [ABSTRACT FROM AUTHOR]

• Water conditions dominated plant phenology and photosynthetic capacity. • Plant phenology and photosynthetic capacity co-determined to annual GPP. • Plant photosynthetic capacity dominated annual GPP. Under global climate change, climate warming and changing precipitation patterns would greatly affect plant phenology and photosynthetic capacity and hence affect the gross primary productivity (GPP) of the sensitive alpine meadow. To deeply understand the variations in GPP, it is important to clarify the joint effects of plant phenology and photosynthetic capacity. In this study, 10-year continuous flux and microclimatic data and in situ phenology period observation data were used to explore the roles of growing season length (GSL) and the seasonal maximal capacity of carbon uptake (GPP max) on annual cumulative gross primary productivity (GPP ann). The results indicated that the temperature and water conditions, especially water conditions, dominated GSL and GPP max and hence determined GPP ann. Both GSL and GPP max were dominated by annual precipitation, and the soil water content in August also showed a strong influence over them. GSL and GPP max together could explain 87% of the variation in GPP ann. Compared with GSL, GPP max had stronger effects on the interannual variability (IAV) in GPP ann. This study provides a new perspective on clarifying the proximate causes of IAV in GPP ann. The IAV in GPP ann of alpine ecosystems could be determined by these two key underlying processes, the length of the carbon uptake period and amplitude. These results are of critical importance for improving our ability to predict future changes in alpine meadow ecosystems. [ABSTRACT FROM AUTHOR]