A Comparison of Two Methods for Quantifying Soil Organic Carbon of Alpine Grasslands on the Tibetan Plateau

As CO₂ concentrations continue to rise and drive global climate change, much effort has been put into estimating soil carbon (C) stocks and dynamics over time. However, the inconsistent methods employed by researchers hamper the comparability of such works, creating a pressing need to standardize the methods for soil organic C (SOC) quantification by the various methods. Here, we collected 712 soil samples from 36 sites of alpine grasslands on the Tibetan Plateau covering different soil depths and vegetation and soil types. We used an elemental analyzer for soil total C (STC) and an inorganic carbon analyzer for soil inorganic C (SIC), and then defined the difference between STC and SIC as SOC_CNS. In addition, we employed the modified Walkley-Black (MWB) method, hereafter SOC_MWB. Our results showed that there was a strong correlation between SOC_CNS and SOC_MWB across the data set, given the application of a correction factor of 1.103. Soil depth and soil type significantly influenced on the recovery, defined as the ratio of SOC_MWB to SOC_CNS, and the recovery was closely associated with soil carbonate content and pH value as well. The differences of recovery between alpine meadow and steppe were largely driven by soil pH. In addition, statistically, a relatively strong correlation between SOC_CNS and STC was also found, suggesting that it is feasible to estimate SOC_CNS stocks through the STC data across the Tibetan grasslands. Therefore, our results suggest that in order to accurately estimate the absolute SOC stocks and its change in the Tibetan alpine grasslands, adequate correction of the modified WB measurements is essential with correct consideration of the effects of soil types, vegetation, soil pH and soil depth.     

青藏高原高寒草原近地表圈层N2O梯度及交换特征

深入了解N2O在不同生态系统土壤及大气中产生和交换特征对于全球气候变化研究具有重要意义.本研究重点探讨N2O在高寒草原近地表圈层中的产生及迁移过程机制.于2000年7月至2001年7月在青藏高原高寒草原地区从土壤1.5 m深到大气中32 m高度10个层次梯度进行N2O浓度变化的观测.结果显示,土壤和大气中N2O浓度均有明显的变化特征.大气中各个层次N2O的浓度都低于土壤中N2O浓度,此浓度差异直接导致了该地区高寒草原土壤向大气中排放N2O气体,其平均排放通量为0.05×10-4μmo1.m-2.s-1,但是在实验点上全年的观测中,N2O气体排放并没有表现出明显的季节性变化特征.土壤中N2O浓度随深度增加而不断升高,浓度最高值出现在1.5 m深处.进一步的分析表明,N2O浓度随深度递增主要是由环境因子中同样递增的土壤湿度所引起的.大气中不同梯度上N2O气体没有明显的浓度差异.近地表各个圈层中N2O浓度在季节上有非常相似的变化特征,即N2O高浓度均出现在入秋和深冬时节.除了N2O浓度变化在各个圈层之间显著相关以外,表层土壤中N2O浓度也与N2O排放变化有明显的相关关系,这表明浓度的差异是导致N2O气体排放变化的最直接因素.近地表土壤中N2O气体是土壤表层N2O气体排放的直接源泉,并且深层土壤中的N2O气体浓度高于浅层土壤,由此我们可以认定土壤中N2O气体通过微生物作用产生以后,由于浓度差异导致从深层土壤到浅层土壤的逐步扩散,最后经地表排放到大气当中去.     

Responses of Soil Microbial Communities to Experimental Warming in Alpine Grasslands on the Qinghai-Tibet Plateau

Global surface temperature is predicted to increase by at least 1.5°C by the end of this century. However, the response of soil microbial communities to global warming is still poorly understood, especially in high-elevation grasslands. We therefore conducted an experiment on three types of alpine grasslands on the Qinghai-Tibet Plateau to study the effect of experimental warming on abundance and composition of soil microbial communities at 0–10 and 10–20 cm depths. Plots were passively warmed for 3 years using open-top chambers and compared to adjacent control plots at ambient temperature. Soil microbial communities were assessed using phospholipid fatty acid (PLFA) analysis. We found that 3 years of experimental warming consistently and significantly increased microbial biomass at the 0–10 cm soil depth of alpine swamp meadow (ASM) and alpine steppe (AS) grasslands, and at both the 0–10 and 10–20 cm soil depths of alpine meadow (AM) grasslands, due primarily to the changes in soil temperature, moisture, and plant coverage. Soil microbial community composition was also significantly affected by warming at the 0–10 cm soil depth of ASM and AM and at the 10–20 cm soil depth of AM. Warming significantly decreased the ratio of fungi to bacteria and thus induced a community shift towards bacteria at the 0–10 cm soil depth of ASM and AM. While the ratio of arbuscular mycorrhizal fungi to saprotrophic fungi (AMF/SF) was significantly decreased by warming at the 0–10 cm soil depth of ASM, it was increased at the 0–10 cm soil depth of AM. These results indicate that warming had a strong influence on soil microbial communities in the studied high-elevation grasslands and that the effect was dependent on grassland type.     

青藏高原高寒草原近地表圈层N2O梯度及交换特征

深入了解N2O在不同生态系统土壤及大气中产生和交换特征对于全球气候变化研究具有重要意义。本研究重点探讨N2O在高寒草原近地表圈层中的产生及迁移过程机制。于2000年7月至2001年7月在青藏高原高寒草原地区从土壤1.5m深到大气中32m高度10个层次梯度进行N2O浓度变化的观测。结果显示,土壤和大气中N2O浓度均有明显的变化特征。大气中各个层次N2O的浓度都低于土壤中N2O浓度,此浓度差异直接导致了该地区高寒草原土壤向大气中排放N2O气体,其平均排放通量为0.05×10-4mmol.m-2.s-1,但是在实验点上全年的观测中,N2O气体排放并没有表现出明显的季节性变化特征。土壤中N2O浓度随深度增加而不断升高,浓度最高值出现在1.5m深处。进一步的分析表明,N2O浓度随深度递增主要是由环境因子中同样递增的土壤湿度所引起的。大气中不同梯度上N2O气体没有明显的浓度差异。近地表各个圈层中N2O浓度在季节上有非常相似的变化特征,即N2O高浓度均出现在入秋和深冬时节。除了N2O浓度变化在各个圈层之间显著相关以外,表层土壤中N2O浓度也与N2O排放变化有明显的相关关系,这表明浓度的差异是导致N2O气体排放变化的最直接因素。近地表土壤中N2O气体是土壤表层N2O气体排放的直接源泉,并且深层土壤中的N2O气体浓度高于浅层土壤,由此我们     

Effects of Grazing on Above- vs. Below-Ground Biomass Allocation of Alpine Grasslands on the Northern Tibetan Plateau

Biomass allocation is an essential concept for understanding above- vs. below-ground functions and for predicting the dynamics of community structure and ecosystem service under ongoing climate change. There is rare available knowledge of grazing effects on biomass allocation in multiple zonal alpine grassland types along climatic gradients across the Northern Tibetan Plateau. We collected the peak above- and below-ground biomass (AGB and BGB) values at 106 pairs of well-matched grazed vs. fenced sites during summers of 2010–2013, of which 33 pairs were subject to meadow, 52 to steppe and 21 to desert-steppe. The aboveground net primary productivity (ANPP) was represented by the peak AGB while the belowground net primary productivity (BNPP) was estimated from ANPP, the ratio of living vs. dead BGB, and the root turnover rate. Two-ways analyses of variance (ANOVA) and paired samples comparisons with t-test were applied to examine the effects of pasture managements (PMS, i.e., grazed vs. fenced) and zonal grassland types on both ANPP and BNPP. Allometric and isometric allocation hypotheses were also tested between logarithmically transformed ANPP and BNPP using standardized major axis (SMA) analyses across grazed, fenced and overall sites. In our study, a high community-dependency was observed to support the allometric biomass allocation hypothesis, in association with decreased ANPP and a decreasing-to-increasing BNPP proportions with increasing aridity across the Northern Tibetan Plateau. Grazing vs. fencing seemed to have a trivial effect on ANPP compared to the overwhelming influence of different zonal grassland types. Vegetation links above- and below-ground ecological functions through integrated meta-population adaptive strategies to the increasing severity of habitat conditions. Therefore, more detailed studies on functional diversity are essentially to achieve conservation and sustainability goals under ongoing climatic warming and intensifying human influences.     

西藏高原高寒草地群落植物多样性和地上生物量监测方法的比较研究

以西藏高原高寒草地3种植被类型(高寒草甸、高寒草原和高寒荒漠草原)为研究对象,分别采用样线法、样方法和巢式样方法进行实地调查,记录每条样线上或每个样方内出现的所有物种,并分物种刈割样方内地上部分,通过统计分析比较不同高寒草地群落植物多样性和地上生物量监测方法,以阐明西藏高原高寒草地不同植被类型的最小取样面积和最少样方数或样线长度。结果显示:(1)就物种丰富度而言,400m样线法观测到的高寒草甸、高寒草原和高寒荒漠草原的所有物种数分别占3种方法调查到的总物种数的55%、71%和50%;8m2巢式样方法调查到高寒草甸、高寒草原的总物种数占所有3种方法观测的物种总数的57.5%和57%,而8m2的巢式样方对高寒荒漠草原的调查监测到的物种数最多,其中2m2观测到的物种数就达到所有可能出现物种的83%;20个样方法监测到高寒草甸和高寒草原的物种数最多,占3种方法观测到总物种数的78%和86%,所以对物种丰富度的调查高寒草甸和高寒典型草原至少需要20个样方,高寒荒漠草原需要最小面积不少于2m2的2个样方。(2)就地上生物量而言,由地上生物量与物种数之间的变异关系得出最小样方数为7~11个,而由地上生物量的变异系数可知,在变异系数小于等于5%的前提下,高寒草甸的最小取样面积不小于0.25m2,高寒典型草原和高寒荒漠草原的最小取样面积不少于1m2。研究表明,对于生产力的监测方法而言,高寒草甸采用10个0.5m×0.5m的样方,而高寒草原和高寒荒漠草原采用10个1m×1m的样方为宜;而对于物种丰富度的监测方法而言,高寒草甸以20个0.5m×0.5m的样方和高寒草原20个1m×1m的样方为宜,高寒荒漠草原采用2个不小于2m2样方面积为宜。     

Carbon Sequestration in Two Alpine Soils on the Tibetan Plateau

Soil carbon sequestration was estimated in a conifer forest and an alpine meadow on the Tibetan Plateau using a carbon- 14 radioactive label provided by thermonuclear weapon tests （known as bomb-^14C）. Soil organic matter was physically separated into light and heavy fractions. The concentration spike of bomb-^14C occurred at a soil depth of 4 cm in both the forest soil and the alpine meadow soil. Based on the depth of the bomb-^14C spike, the carbon sequestration rate was determined to be 38.5 g C/m^2 per year for the forest soil and 27.1 g C/m^2 per year for the alpine meadow soil. Considering that more than 60% of soil organic carbon （SOC） is stored in the heavy fraction and the large area of alpine forests and meadows on the Tibetan Plateau, these alpine ecosystems might partially contribute to ＂the missing carbon sink＂.     

Carbon Sequestration in Two Alpine Soils on the Tibetan Plateau

Soil carbon sequestration was estimated in a conifer forest and an alpine meadow on the Tibetan Plateau using a carbon-14 radioactive label provided by thermonuclear weapon tests (known as bomb-¹⁴C). Soil organic matter was physically separated into light and heavy fractions. The concentration spike of bomb-¹⁴C occurred at a soil depth of 4 cm in both the forest soil and the alpine meadow soil. Based on the depth of the bomb-¹⁴C spike, the carbon sequestration rate was determined to be 38.5 g C/m² per year for the forest soil and 27.1 g C/m² per year for the alpine meadow soil. Considering that more than 60% of soil organic carbon (SOC) is stored in the heavy fraction and the large area of alpine forests and meadows on the Tibetan Plateau, these alpine ecosystems might partially contribute to "the missing carbon sink".     

Seasonal Response of Grasslands to Climate Change on the Tibetan Plateau

Background

Monitoring vegetation dynamics and their responses to climate change has been the subject of considerable research. This paper aims to detect change trends in grassland activity on the Tibetan Plateau between 1982 and 2006 and relate these to changes in climate.

Methodology/Principal Findings

Grassland activity was analyzed by evaluating remotely sensed Normalized Difference Vegetation Index (NDVI) data collected at 15-day intervals between 1982 and 2006. The timings of vegetation stages (start of green-up, beginning of the growing season, plant maturity, start of senescence and end of the growing season) were assessed using the NDVI ratio method. Mean NDVI values were determined for major vegetation stages (green-up, fast growth, maturity and senescence). All vegetation variables were linked with datasets of monthly temperature and precipitation, and correlations between variables were established using Partial Least Squares regression. Most parts of the Tibetan Plateau showed significantly increasing temperatures, as well as clear advances in late season phenological stages by several weeks. Rainfall trends and significant long-term changes in early season phenology occurred on small parts of the plateau. Vegetation activity increased significantly for all vegetation stages. Most of these changes were related to increasing temperatures during the growing season and in some cases during the previous winter. Precipitation effects appeared less pronounced. Warming thus appears to have shortened the growing season, while increasing vegetation activity.

Conclusions/Significance

Shortening of the growing season despite a longer thermally favorable period implies that vegetation on the Tibetan Plateau is unable to exploit additional thermal resources availed by climate change. Ecosystem composition may no longer be well attuned to the local temperature regime, which has changed rapidly over the past three decades. This apparent lag of the vegetation assemblage behind changes in climate should be taken into account when projecting the impacts of climate change on ecosystem processes.     

青藏高原草地双子叶植物叶片的气孔特征研究

利用光学显微镜对青藏高原29种草地双子叶植物叶片的气孔形态与数量特征进行观察及差异显著性分析,为揭示青藏高原草地双子叶植物对高原环境的适应机制及探索气孔作为辅助分类的依据奠定理论基础。结果表明:(1)青藏高原草地双子叶植物大多数种类在叶片上、下表皮均分布有气孔,气孔随机排列,气孔器多为无规则型。(2)气孔长度(SL)较小,上、下表皮的气孔平均长度分别为26.20μm与25.56μm,且气孔密度(SD)与气孔指数(SI)相对较大。(3)不同科、属、种间叶片上、下表皮的SL、SD、SI差异均极显著。(4)叶片上、下表皮的6个气孔数量特征之间具有显著相关关系。(5)上表皮的SL、SD与不同科、属、种间显著相关,下表皮除SI与物种间相关不显著外,其他指标与科、属、种间显著相关。研究认为,青藏高原草地双子叶植物独特的气孔形态与数量特征是对高寒极端环境长期适应的结果,且气孔数量特征对植物辅助分类具有重要价值。     

Soil Organic Matter Dynamics Along a Vertical Vegetation Gradient in the Gongga Mountain on the Tibetan Plateau

Our knowledge about soil organic matter (SOM) dynamics is limited although this is an important issue in the study of responses of ecosystems to global climate changes. Twelve sampling plots were set up every 200 m from 1 700 to 3 900 m along the vertical vegetation gradient along the east slope of Gongga Mountain. Samples were taken from all 12 plots for SOM content measurement, although only 5 of the 12 plots were subjected to radiocarbon measurements. A radiocarbon isotope method and a time-dependent model were used to quantify the SOM dynamics and SOM turnover rates along the vertical vegetation gradient. The results showed that the SOM turnover rate decreased and turnover time increased with soil depth for all vegetation types. The litter layer turnover rates presented a clear trend along the gradient. The litter layer turnover rates decreased with an increase in elevation, except that the litter layer turnover rate of mixed forest was higher than that of evergreen forest. Climatic factors, such as temperature and precipitation, were the main factors influencing the surface soil carbon dynamics. The turnover rates of the subsoil (including the A, B, and C horizons in the soil profiles) along the vertical gradient had no clear trends. The SOM of subalpine shrub and meadow turned over more slowly than that of the forest types in almost all soil horizons. The characteristic of short roots distributing in the upper part of the soil profile leads to different SOM dynamics of shrub and meadow compared with the forest types. Coniferous and mixed forests were susceptible to carbon loss from the young carbon pool, but their long and big roots resulted in high △^14C values of the deep soil profiles and increased the input of young carbon to the deep soil. In evergreen forest, the carbon cumulative ability from the B horizon to the C horizon was weak. The different vegetation types, together with their different modes of nutrient and carbon intake, may be the mechanism conditioning the subsoil organic matter dynamics.     

Soil Organic Matter Dynamics Along a Vertical Vegetation Gradient in the Gongga Mountain on the Tibetan Plateau

Our knowledge about soil organic matter (SOM) dynamics is limited although this is an important issue in the study of responses of ecosystems to global climate changes. Twelve sampling plots were set up every 200 m from 1 700 to 3 900 m along the vertical vegetation gradient along the east slope of Gongga Mountain. Samples were taken from all 12 plots for SOM content measurement, although only 5 of the 12plots were subjected to radiocarbon measurements. A radiocarbon isotope method and a time-dependent model were used to quantify the SOM dynamics and SOM turnover rates along the vertical vegetation gradient. The results showed that the SOM turnover rate decreased and turnover time increased with soil depth for all vegetation types. The litter layer turnover rates presented a clear trend along the gradient. The litter layer turnover rates decreased with an increase in elevation, except that the litter layer turnover rate of mixed forest was higher than that of evergreen forest. Climatic factors, such as temperature and precipitation,were the main factors influencing the surface soil carbon dynamics. The turnover rates of the subsoil (including the A, B, and C horizons in the soil profiles) along the vertical gradient had no clear trends. The SOM of subalpine shrub and meadow turned over more slowly than that of the forest types in almost all soil horizons. The characteristic of short roots distributing in the upper part of the soil profile leads to different SOM dynamics of shrub and meadow compared with the forest types. Coniferous and mixed forests were susceptible to carbon loss from the young carbon pool, but their long and big roots resulted in high △14C values of the deep soil profiles and increased the input of young carbon to the deep soil. In evergreen forest,the carbon cumulative ability from the B horizon to the C horizon was weak. The different vegetation types,together with their different modes of nutrient and carbon intake, may be the mechanism conditioning the subsoil organic matter dynamics.     

Abundance and Diversity of Glacial Bacteria on the Tibetan Plateau with Environment

The relation of glacial microorganism and their living environment is concerned but less understood for both glaciologists and microbiologists. Here we present the results about glacial snow bacteria from 3 glaciers on the Tibetan Plateau. The concentrations and diversity of bacteria collected from snow pit samples of East Rongbuk, Laohugou and Hailuogou glaciers on the Tibetan Plateau were investigated by epifluorescence microscope, denaturing gradient gel electrophoresis and Shannon-Weaver index. Concentrations and community diversity of bacteria in the East Rongbuk glacier with lower concentration and smaller size of microparticle were lower than in the Laohugou with higher concentration and larger size of microparticle. Bacterial concentration in East Rongbuk Glacier was close to that in polar regions, suggesting that Mt. Everest is a bacterial background for remote regions away from direct influence of anthropogenic sources. In addition, altitude difference was another factor for higher concentrations and community diversity of bacteria in Hailuogou and Laohugou glaciers than in East Rongbuk Glacier. The highest concentrations and community diversity of bacteria in Hailuogou Glacier were attributed to its most diverse atmospheric circulations and highest temperature among the 3 glaciers. We suggest a complicated correlation between glacial bacteria and their regional living environments.     

Variation in Natural Abundance of 15N among Plant Parts and in 15N/14N Fractionation during N2 Fixation in the Legume-Rhizobia Symbiotic System

Sixteen legumes were grown in N-free media so that N was suppliedentirely by symbiotic N₂ fixation. The plant tissues were analyzedfor natural ¹⁵N abundance (expressed as ¹⁵N per mil relativeto air N₂) with a ratio mass spectrometer. The nodules of desmodium,centro, siratro, soybean and winged bean showed high enrichmentin ¹⁵N (+9), while red clover showed slight enrichment (+2).The nodules of 9 other forage legumes (Townsville stylo, whiteclover, alsike clover, common vetch, Chinese milk vetch, senna,alfalfa, ladino clover, and hairy vetch) showed little enrichmentin ¹⁵N. In all the legumes investigated, particularly in the ureide-transportingplants such as desmodium, centro, siratro, soybean, winged beanand field bean, the ¹⁵N value of the shoots was negative (–3.2).The ¹⁵N value of the shoots in winged bean and field bean variedby about 1 depending on the Rhizobium strains used. The isotopicmass balance of 13 legumes indicated that isotopic fractionationoccurs during N₂ fixation by the legume-rhizobia symbiosis witha preference for ¹⁴N over ¹⁵N, resulting in a ¹⁵N value of –0.2to –2 in the whole plant. The results indicate that ¹⁵N/¹⁴N isotopic discrimination witha preference for the lighter atom may occur in both N₂ fixationand export of fixed N from nodules. ¹Present address: Department of Soils and Fertilizers, NationalAgriculture Research Center, Kannondai, Tsukuba, Ibaraki 305,Japan. (Received October 8, 1985; Accepted April 7, 1986)     

Effects of Grazing Regimes on Plant Traits and Soil Nutrients in an Alpine Steppe,Northern Tibetan Plateau

Understanding the impact of grazing intensity on grassland production and soil fertility is of fundamental importance for grassland conservation and management. We thus compared three types of alpine steppe management by studying vegetation traits and soil properties in response to three levels of grazing pressure: permanent grazing (M1), seasonal grazing (M2), and grazing exclusion (M3) in the alpine steppe in Xainza County, Tibetan Plateau. The results showed that community biomass allocation did not support the isometric hypothesis under different grassland management types. Plants in M1 had less aboveground biomass but more belowground biomass in the top soil layer than those in M2 and M3, which was largely due to that root/shoot ratios of dominant plants in M1 were far greater than those in M2 and M3. The interramet distance and the tiller size of the dominant clonal plants were greater in M3 than in M1 and M2, while the resprouting from rhizome buds did not differ significantly among the three greezing regimes. Both soil bulk density and soil available nitrogen in M3 were greater than in M1 at the 15–30 cm soil depth (P = 0.05). Soil organic carbon and soil total nitrogen were greater in M3 than in M1 and M2 (P = 0.05). We conclude that the isometric hypothesis is not supported in this study and fencing is a helpful grassland management in terms of plant growth and soil nutrient retention in alpine steppe. The extreme cold, scarce precipitation and short growing period may be the causation of the unique plant and soil responses to different management regimes.     

Nitrogen Transfer Between Plants: A 15N Natural Abundance Study with Crop and Weed Species

An increasing amount of evidence indicates that N can be transferred between plants. Nonetheless, a number of fundamental questions remain. A series of experiments was initiated in the field to examine N transfer between N₂-fixing soybean (Glycine max [L.] Merr.) varieties and a non-nodulating soybean, and between N₂-fixing peanut (Arachis hypogaea L.) or soybean and neighboring weed species. The experiments were conducted in soils with low N fertilities and used differences in N accumulation and/or ¹⁵N natural abundance to estimate N transfer. Mixtures of N₂-fixing and non-nod soybean indicated that substantial inter-plant N transfer occurred. Amounts were variable, ranging from negligible levels to 48% of the N found in the non-nod at maturity. Transfer did not appear to strongly penalize the N₂-fixing donor plants. But, in cases where high amounts of N were transferred, N content of donors was noticeably lowered. Differences were evident in the amount of N transferred from different N₂-fixing donor genotypes. Results of experiments with N₂-fixing crops and the weed species prickly sida (Sida spinosa L.) and sicklepod (Senna obtusifolia [L.] Irwin & Barneby) also indicated substantial N transfer occurred over a 60-day period, with amounts accounting for 30–80% of the N present in the weeds. Transfer of N, however, was generally very low in weed species that are known to be non-hosts for arbuscular mycorrhizae (yellow nutsedge, Cyperus esculentus L. and Palmer amaranth, Amaranthus palmeri [S.] Watson). The results are consistent with the view that N transfer occurs primarily through mycorrhizal hyphal networks, and they reveal that N transfer may be a contributing factor to weed problems in N₂-fixing crops in low N fertility conditions.     

Alpine Grassland Soil Organic Carbon Stock and Its Uncertainty in the Three Rivers Source Region of the Tibetan Plateau

Alpine grassland of the Tibetan Plateau is an important component of global soil organic carbon (SOC) stocks, but insufficient field observations and large spatial heterogeneity leads to great uncertainty in their estimation. In the Three Rivers Source Region (TRSR), alpine grasslands account for more than 75% of the total area. However, the regional carbon (C) stock estimate and their uncertainty have seldom been tested. Here we quantified the regional SOC stock and its uncertainty using 298 soil profiles surveyed from 35 sites across the TRSR during 2006–2008. We showed that the upper soil (0–30 cm depth) in alpine grasslands of the TRSR stores 2.03 Pg C, with a 95% confidence interval ranging from 1.25 to 2.81 Pg C. Alpine meadow soils comprised 73% (i.e. 1.48 Pg C) of the regional SOC estimate, but had the greatest uncertainty at 51%. The statistical power to detect a deviation of 10% uncertainty in grassland C stock was less than 0.50. The required sample size to detect this deviation at a power of 90% was about 6–7 times more than the number of sample sites surveyed. Comparison of our observed SOC density with the corresponding values from the dataset of Yang et al. indicates that these two datasets are comparable. The combined dataset did not reduce the uncertainty in the estimate of the regional grassland soil C stock. This result could be mainly explained by the underrepresentation of sampling sites in large areas with poor accessibility. Further research to improve the regional SOC stock estimate should optimize sampling strategy by considering the number of samples and their spatial distribution.     

Response of Alpine Plants to Nitrogen Addition on the Tibetan Plateau: A Meta-analysis

To identify the general effects of nitrogen addition on alpine plants, we used a meta-analysis approach to synthesize 599 observations from 51 studies on the Tibetan Plateau. Nitrogen addition significantly increased plant height by 19.0 %, plant biomass by 29.7 %, graminoid aboveground biomass by 89.8 %, and sedge aboveground biomass by 75.6 % but significantly decreased legume aboveground biomass by 34.5 %, forb aboveground biomass by 23.8 %, and species richness by 11.2 %. The effect of nitrogen addition on aboveground plant biomass and plant height increased with increasing the nitrogen addition rate. The effect of nitrogen addition on plant height increased with increasing mean annual precipitation but decreased with increasing mean annual temperature. Our findings suggested that the effect of nitrogen addition on alpine plants varied with plant functional types and nitrogen addition rate. In addition, climatic warming and precipitation changes may regulate the response of alpine plants to nitrogen addition on the Tibetan Plateau.