The Tibetan Plateau reacts particularly sensitively to possible effects of climate change. Approximately two thirds of the total area is affected by permafrost. To get a better understanding of the role of permafrost on soil organic carbon pools and stocks, investigations were carried out including both discontinuous (site Huashixia, HUA) and continuous permafrost (site Wudaoliang, WUD). Three organic carbon fractions were isolated using density separation combined with ultrasonic dispersion: the light fractions (<1.6 g cm−3) of free particulate organic matter (FPOM) and occluded particulate organic matter (OPOM), plus a heavy fraction (>1.6 g cm−3) of mineral associated organic matter (MOM). The fractions were analyzed for C, N, and their portion of organic C. FPOM contained an average SOC content of 252 g kg−1. Higher SOC contents (320 g kg−1) were found in OPOM while MOM had the lowest SOC contents (29 g kg−1). Due to their lower density the easily decomposable fractions FPOM and OPOM contribute 27% (HUA) and 22% (WUD) to the total SOC stocks. In HUA mean SOC stocks (0–30 cm depth) account for 10.4 kg m−2, compared to 3.4 kg m−2 in WUD. 53% of the SOC is stored in the upper 10 cm in WUD, in HUA only 39%. Highest POM values of 36% occurred in profiles with high soil moisture content. SOC stocks, soil moisture and active layer thickness correlated strongly in discontinuous permafrost while no correlation between SOC stocks and active layer thickness and only a weak relation between soil moisture and SOC stocks could be found in continuous permafrost. Consequently, permafrost-affected soils in discontinuous permafrost environments are susceptible to soil moisture changes due to alterations in quantity and seasonal distribution of precipitation, increasing temperature and therefore evaporation.
Citation: Dörfer C, Kühn P, Baumann F, He J-S, Scholten T (2013) Soil Organic Carbon Pools and Stocks in Permafrost-Affected Soils on the Tibetan Plateau. PLoS ONE 8(2): e57024. https://doi.org/10.1371/journal.pone.0057024
Editor: Caroline P. Slomp, Utrecht University, Netherlands
Received: May 24, 2012; Accepted: January 21, 2013; Published: February 26, 2013
Copyright: © 2013 Dörfer et al. This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Funding: This work was supported by the Federal Ministry of Education and Research (BMBF, 03G0810A, http://www.bmbf.de/en), the German Academic Exchange Service (DAAD, http://www.daad.de) and the National Natural Science Foundation of China (31025005 and 31021001). The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.
Competing interests: The authors have declared that no competing interests exist.
The relationship between soil organic carbon (SOC) stocks and site characteristics has been well investigated in the temperate zones at local and regional scale (e.g. –), but much less studies exist about the role of SOC in high-cold alpine regions (e.g. –). Studies in Arctic regions have shown that permafrost-influenced alpine ecosystems are highly sensitive to global climate change . Prevailing low temperatures and permanently low turn-over rates result in large SOC stocks, providing a great emission potential for greenhouse gases such as CO2 and CH4 –. Thus, an estimate of SOC stocks in their extent and distribution is essential to predict feedback of SOC on global climate change , , . Furthermore, differentiation between carbon pools is necessary because various SOM fractions show large differences in their turn-over rates according to their mineral binding –.
The Tibetan Plateau is a particularly sensitive area in terms of possible effects of global climate change. It is the largest and highest plateau on earth and covers an area of 2.5 million square kilometers with an average altitude of more than 4000 m a.s.l. comprising about a quarter of China’s mainland . On the Tibetan Plateau seasonally frozen soils are widespread . Approximately two thirds of the total area is affected by permafrost . Many studies focus on recent changes of the permafrost conditions on the Tibetan Plateau, attesting increasing permafrost temperatures, active layer thickness and rising degradation (e.g. –. Due to relatively high permafrost temperatures just below the freeze-thaw point, the so-called warm permafrost is more sensitive to global warming than the cold permafrost of higher latitudes . The permafrost distribution is closely related to the characteristics of the land surface such as slope, exposure, vegetation distribution and snow cover. In this study permafrost is classified after the Chinese Permafrost Classification which differs from the classification of the International Permafrost Association (IPA). 50–90% of permafrost is required for classifying discontinuous permafrost after IPA Permafrost Classification; 30–70% is required after Chinese Permafrost Classification .
Recent soil ecological research mainly focuses on soil temperature as the main driving force for ecosystem processes (e.g. , –). Baumann et al.  showed that nutrient availability is a limiting factor for plant growth as well, which in turn is controlled by soil moisture. Furthermore studies show soil moisture is the dominant parameter regarding to the spatial variation of SOC contents  and soil CO2 efflux  on the landscape scale in permafrost influenced ecosystems. Zhao et al.  found a negative relationship between vegetation cover/biomass and active layer thickness in alpine meadow ecosystems. Permafrost favors the development of alpine meadow ecosystems, protecting in turn permafrost by their dense vegetation cover from degradation , .
According to Wang et al.  about 33.52 Pg SOC are stored in grassland soils of the Tibetan Plateau down to a depth of 70 cm. Alpine meadow and alpine steppe soils have a share of 23.2 Pg, which represents 2.5% of the global soil carbon pool . Alpine meadows make up 38.2% of total grassland soil carbon in Chinese grassland soils . Wang et al.  estimated that the degradation of grassland is resulting in a loss of 57% of SOC in heavy fractions (HF) and 84% in light fractions (LF) from alpine meadow soils in Dari County (Qinghai Province). From 1986 to 2000 land cover changes have led to a loss of 1.8 Gg SOC and a mass loss of 65% in the LF in the upper 30 cm .
Major objectives of this study are (1) to investigate SOC stocks and their affiliation to pools by density fractionation and (2) to examine interactions of SOC, soil moisture, and active layer thickness in permafrost-affected soils on the Tibetan Plateau. For comparing stocks and processes in soils affected by continuous and discontinuous permafrost adequately, sites with similar rainfall under varying temperatures were selected.
Materials and Methods
The study sites are located on the northeastern Tibetan Plateau, Qinghai Province, China and were investigated in May/June 2009 and 2010 (Fig. 1). Site HUA is situated near the settlement of Huashixia in Maduo County in the Yellow River catchment area, 4300 m a.s.l. The area is affected by the subtropical East Asian Monsoon, which transports air masses with high water vapor content from the lowlands to the Tibetan Plateau through the meridional flow furrows , leading to relatively high rainfall. The nearest climate station at Maduo shows a Mean Annual Air Temperature (MAAT) of −4.1°C and a Mean Annual Precipitation (MAP) of 326 mm . The catchment area of the Yellow River (Huang He) is characterized by discontinuous, unstable permafrost. The soils freeze to a depth of 2–3 m, while the upper limit of the permafrost lies in 4–7 m depth; so-called taliks have developed . This vertical disconnection of the permafrost is widespread near the study site. The site is influenced by severe summer grazing with yak (Bos grunniens) and sheep (Ovis aries) and a temporary settlement by nomads.
Site WUD is located in the headwaters of the Yangtze River, in the middle between Wudaoliang and Tuotuohe next to the Qinghai-Tibet Highway. Precipitation decreases from SE to NW on the Plateau, but due to its location on the Fenghuo Shan mountains, site WUD is characterized by higher precipitation and lower temperatures as the surrounding area (MAP: 348 mm, MAAT: −5.75°C, 4801 m a.s.l., ). The high elevation and the low influence of the South Asian Monsoon are responsible for the widespread continuous permafrost in that area  which is relatively poor in ice . This also results in a shorter vegetation growth period compared to HUA. During the field campaign no grazing occurred. Nevertheless, the area is used only as late summer pasture due to extreme climate conditions. Generally, grazing is less intensive compared to HUA.
The soils in both study areas are developed in sandy loess, mixed with coarser material derived from frost weathering processes. Poorly developed soils at site HUA are classified as gleyic Fluvisols, haplic Regosols and mollic Cryosols, whereas cambic Cryosols are common in WUD . Alpine Kobresia meadow ecosystems are the most common vegetation types on the plateau  occurring at elevations ranging from 3200 to 5200 m a.s.l. . At the study sites, particularly Kobresia pygmaea and K. humilis are widespread. Plant composition is similar at both sites, differing along the altitudinal gradient according to water supply. Strongly rooted, partly felty topsoils are common .
During May and June in 2009, 11 soil profiles were sampled (HUA: 6, WUD: 5). All soil profiles were arranged along an elevation gradient and affected by permafrost with an active layer thickness less than 100 cm at both sites. At each site two soil profiles in footslope position, two soil profiles in lower mid-slope position and two (HUA) and one (WUD) soil profiles in upper mid-slope position were sampled. We set a high value on the comparability of both study sites in topographic position of the soil profiles, inclination of the slopes and vegetative cover. Soil sampling was split into three parts: horizon-wise sampling for pedogenesis and soil chemical analyses using soil pits, schematic sampling conducted by drilling at four depth-increments (0–5, 5–10, 10–20 and 20–30 cm, four replicates each) for C analysis and volumetric sampling at the same depths for bulk density and gravimetric water content determination (three replicates each). Detailed description of soil profiles and pedogenic implications will be published elsewhere.
All soil samples were air-dried and sieved to <2 mm. The pH of the samples was measured in deionized water and in 0.01 M CaCl2 solution at a solution:soil ratio of 2.5∶1 with a glass electrode (Sentix 81, WTW, pH 340). CaCO3 was analyzed gasvolumetrically on ground subsamples (Calcimeter, Eijkelkamp). Total C and N in bulk soil samples and density fractions were determined on ground subsamples by heat combustion with a CNS analyzer (Vario EL III, Elementar GmbH, Hanau, Germany). SOC in bulk soils and density fractions was calculated as the difference between soil total and soil inorganic carbon. Water content was determined by gravimetric water content analysis, corrected by the skeleton content (>2 mm).
Density fractionation was carried out using sodium polytungstate, following the procedures of Grünewald et al. . It is generally accepted that the density of OM is <1.5 g cm−3. After Golchin et al.  a density fractionation at 1.6 g cm−3 separates light organic fractions from mineral dominated heavy fractions under the assumption, that most mineral particles contain less than 20% OM . Three fractions were isolated: the light fractions free particulate organic matter (FPOM) and occluded particulate organic matter (OPOM) with a density <1.6 g cm−3, plus a heavy fraction of mineral associated organic matter (MOM) with a density of >1.6 g cm−3. FPOM was separated by floatation after gently shaking in a sodium polytungstate solution (soil:solution ratio 1∶5) and centrifugation at 4,500 rpm for 20 min, followed by vacuum filtration. OPOM was separated after ultrasonic dispersion (58 J ml−1) and centrifugation at 4,500 rpm for 15 min. Calibration of the ultrasonic output energy was carried out according to Roscoe et al. . The remains (MOM) were washed three times to remove the salt. Due to soil inhomogeneity fractionation was carried out twice per sample. The dried and ground fractions were analyzed for C, N and SOC. The density solution was recycled after Six et al. . SOC stocks for bulk soils and individual fractions were calculated down to a depth of 30 cm, according to Ohtsuka et al. :where M is the soil layer thickness, is the bulk density (g cm−3) of the soil, SOC is the soil organic carbon content (Mass%) and S is the skeleton content (Mass%).
Soil Organic Carbon Content and SOC/N Ratios
The SOC content of bulk soil decreased with increasing depth at both sites (Fig. 2). In HUA (51 g kg−1) significantly higher mean values were reached than in WUD (19 g kg−1). Highest contents occurred in the OPOM fractions (320 g kg−1), FPOM followed with 252 g kg−1, while they were lowest in the MOM fractions (29 g kg−1). The SOC contents in the FPOM decreased with depth, while there were increasing SOC values in the OPOM fractions. The SOC contents of the MOM fraction follow the same depth gradient as SOC in bulk soils at both sites. Significantly higher SOC contents were found in this fraction on site HUA, showing a large variation. The mean recovery of SOC after density fractionation was 95%.
SOC/N ratios of the bulk soils were relatively similar at both sites and decreased with depth (from 12 to 9) – except in extremely moist soils in HUA, where the SOC/N-ratios remained constant (13). The SOC/N ratios of the three fractions were remarkably different. Highest ratios were found in FPOM with a mean ranging from 15–36, followed by OPOM with 14–29. Generally, lower SOC/N ratios occurred in MOM fractions (3–12). We observed slightly rising SOC/N ratios with depth for the POM fractions in all soils, whereas ratios varied less with depth for the MOM fractions.
Soil Organic Carbon Stocks of Bulk Soil and Fractions
Total SOC stocks in HUA ranged from 1.9 kg m−2 to 19.3 kg m−2 up to 30 cm depth with a mean of 10.4 kg m−2 (Tab. 1). Significantly lower stocks were found in WUD ranging from 2.5 kg m−2 to 5.0 kg m−2 (mean: 3.4 kg m−2).
In line with the higher SOC contents, soils in HUA (0.41 kg m−2) showed twice as high stocks as in WUD in the top 5 cm (Tab. 1). Fractional SOC stocks in particular depths were highest in the MOM fraction. In WUD, stocks decreased with depth for all three fractions whereas increasing SOC stocks in the OPOM fraction in 20–30 cm depth were evident compared to 10–20 cm depth at HUA.
Variations of the SOC stocks were much higher in HUA than in WUD, especially in the light POM fractions (Tab. 1).The POM fractions in HUA and WUD contributed 27% and 22% to the SOC stocks with 8% in the OPOM fractions at both sites (Fig. 3). Comprising the different depth levels, 53% of SOC is stored in the upper 10 cm in WUD. In HUA only 39% is stored in the upper 10 cm and the portion of FPOM on SOC stocks remained constant with depth, while the portion of OPOM increased. The portion of FPOM on SOC stocks decreased slightly, whereas OPOM stocks increased with depth in WUD. The portion of MOM stock remained relatively constant at 77%, thus slightly higher than in HUA.
Correlations between Soil Organic Carbon Stocks, Active Layer Thickness and Soil Moisture
To assess the influence of soil hydrological properties on SOC stocks, active layer thickness and soil moisture content were taken into account. Since we did not reach the maximum active layer depth in May/June 2009, we used active layer and corresponding soil moisture data from August/September 2011. A significant correlation between SOC stocks and soil moisture can be confirmed for both study sites (Fig. 4). Soil moisture in HUA varies from 7 to almost 56 Vol. % with a high correlation of R2 = 0.74 between SOC stocks and soil moisture. The range in WUD is smaller with values between 11 and 50 Vol. % soil moisture and a weak correlation (R2 = 0.05).
An inverse correlation can be observed for active layer thickness and soil moisture (Fig. 5). The mean thaw depth at both locations is similar (HUA: 97 cm, WUD: 99 cm). Contrarily, the range differs distinctly, with a variation coefficient of 22.0 in HUA and 15.8 in WUD. The interrelation between active layer thickness and SOC stocks is positive for site HUA with R2 = 0.77. For site WUD, similar significant correlations could not be detected.
Soil Organic Carbon Content
SOC contents decreased with soil depth at both locations in all soil profiles. At site HUA, however, much higher overall values were found than in WUD. The more humid, partly water logging conditions inhibit microbial decomposition processes  leading in combination with a more dense vegetation cover to a larger accumulation of organic matter.
Concerning the relative amount of OM fractions, our findings are in line with the results of Golchin et al. , who found higher SOC contents in the OPOM than in the FPOM and MOM fractions as well. Wang et al.  isolated a light fraction (LF) and a heavy fraction (HF) using a 1.8 g cm−3 density solution. In LF SOC contents of 290 g kg−1 (0–10 cm) and 260 g kg−1 (10–20 cm), for HF 30 g kg−1 (0–10 cm) and 40 g kg−1 (10–20 cm) were found. The comparably slightly lower SOC contents in the LF may reflect the influence of the MOM fraction. However, the comparison is difficult, because different density ranges for the fractionation were used .
SOC in FPOM and MOM decreased with depth, whereas the contents of the OPOM fractions increased slightly at both sites. Larger aggregates composed of coarse textured OPOM with a lower degree of decomposition were present in the depth increment 0–10 cm. A reduction of particle size with depth was clearly observable during the fractionation process. Even though the SOC content was relatively small compared with the POM fractions, the portion of total SOC stored in OPOM is large (Fig. 2).
OC/N ratios of bulk soil decreased with depth indicating a higher age and grade of humification in the subsoil . SOC/N ratios in water-saturated soils in HUA remained relatively stable with depth indicating inhibited decomposition processes. The highest SOC/N ratios with depth were found in the FPOM fraction followed by OPOM and MOM. Comparable results were reported by Grünewald et al.  and John et al. , who also observed decreasing SOC/N ratios with depth from FPOM>OPOM>MOM for all soils, indicating an increasing degree of OM degradation and humification. Golchin et al.  observed higher ratios for OPOM than for FPOM, whereas Kölbl and Kögel-Knabner  found no differences between both fractions.
OC/N ratios in FPOM showed little variations with depth, while the portion of carbon content decreases. Further, we observed a decrease in particle size in the POM fractions during the fractionation that may also contribute to a shift in the SOC/N ratio with depth. The very low SOC/N ratios in MOM fractions suggest a generally larger contribution of microbial biomass and, hence, stronger microbial decomposition of plant debris than for POM fractions. The contribution of inorganic N to the total N content – resulting in a very low SOC/N ratio – cannot be excluded.
Soil and Fractional Organic Carbon Stocks
Our results on SOC stocks are in line with other published data on alpine meadow soils on the Tibetan Plateau , , ,  as well as on tundra soils in Siberia  at a depth of 0–30 cm (Tab. 2). In HUA mean SOC stocks of 10.4 kg m−2 were found, in WUD comparably low 3.4 kg m−2 (0–30 cm). Due to small-scale differences in substrate, soil moisture and hence vegetation cover, carbon stocks differ more in HUA than in WUD. Highest stocks occurred in water saturated profiles at HUA (19.3 kg m−3) whereas lowest stocks were found in dry profiles with a lower vegetation density in WUD (2.7 kg m−3). Our results show that high soil moistures combined with low soil temperatures (due to the isolating effect of dense vegetation) lead to an increased accumulation of soil organic matter  and therefore higher SOC stocks than in drier soil profiles. SOC stocks decreased with depth, especially in WUD (Tab. 1).
The comparison of fractional SOC stocks with the results of other published research is challenging, as different density ranges and fractionation methods are used and the number of studies on the Tibetan Plateau is limited. Compared to other grassland ecosystems like steppe soils in Ukraine and Kazakhstan  and grassland soils in Saxony-Anhalt, Germany , in this study significantly higher portions of FPOM and OPOM on SOC stocks were found. About 18% of the total contents in HUA and 14% in WUD were contributed by the FPOM fraction. The OPOM portions were the same at both sites (8%) and the MOM fractions contributed 74 and 78% to the total SOC stocks (Fig. 3). Due to lower litter production in WUD, lower fraction masses and SOC contents led to lower portions of POM on total SOC stocks (Fig. 3). At site HUA the share of FPOM remained relatively constant with depth, while the stocks in WUD were decreasing. The share of OPOM increased at both sites with increasing depth. The increase in OPOM stocks was linked also to the increasing SOC content of the fractions with increasing depth.
The limitation of water caused a lower turnover of organic matter at very dry profile sites in HUA and WUD. This resulted in a relatively high share of FPOM on SOC with small portions of OPOM similar to desert soils investigated by Kadono et al. . The largest FPOM and OPOM shares were found in water-saturated profiles at site HUA with a contribution of 39% of the POM fractions to the SOC stocks since further degradation of SOM is strongly inhibited there.
Our results are comparable to studies in Siberia  and the Tibetan Plateau , . Wang et al.  isolated a light and heavy fraction (1.85 g cm−3 density solution) without distinction between FPOM and OPOM. In this case only trends can be compared with our results. The LF contained 7 kg m−2 and thus about 37–44% of SOC from 0–30 cm depth. Altogether, SOC stocks comprised 9.81 kg m−2 at a depth of 30 cm. Wang et al.  isolated LF and HF with a density range of 1.8 g cm−3. They found a SOC stock of 7.5 kg m−2 at a depth of 0–20 cm, with 0.8 kg m−2 in LF and 2.8 kg m−2 in HF in the upper 10 cm. 0.4 kg m−2 in LF and 3.5 kg m−2 in HF were contained in 10–20 cm depth.
Correlation between Organic Carbon Stocks, Active Layer Thickness and Soil Moisture
Similar mean active layer thicknesses are evident at both sites, but significantly larger variations were observed in HUA. These spatial dynamics are related to small scale changes in substrate, bulk density, soil moisture values  and hence in vegetation coverage leading to larger active layer thicknesses. Patches of dense vegetation have an isolating effect, protecting permafrost from thawing ,  resulting in shallower active layer depths.
The detected thawing depths of 55–124 cm in HUA and 84–122 cm in WUD correspond to the range of the maximum thawing depths of 80–150 cm near WUD in September published by Wang et al. . Yang et al.  and Baumann et al.  showed, that soil moisture affects significantly extension and distribution of carbon stocks on the Tibetan Plateau, as we found as well. Soil moisture, active layer thickness and carbon stocks correlated strongly at HUA (Fig. 4 and 5). As a consequence of the moist to water-saturated conditions and the dense vegetation cover, a higher amount of organic matter is accumulated in HUA compared to WUD, where a lower litter input due to the shorter growing season and drier conditions accelerating mineralization, are prominent. In WUD we found no correlation between soil moisture and SOC stocks as well as between SOC stocks and active layer thickness.
In this paper we investigated the interactions of SOC stocks and the proportion of light and heavy SOC fractions with soil moisture and active layer thickness in permafrost-affected soils on the Tibetan Plateau. Furthermore, the affiliation of SOC stocks into different SOC pools was examined. The research sites are located in both continuous (WUD) and discontinuous (HUA) permafrost areas.
SOC stocks, soil moisture, and active layer thickness correlated strongly in discontinuous permafrost, whereas no correlation between SOC stocks and active layer thickness and only a weak relation between SOC stocks and soil moisture could be detected for continuous permafrost. Organic carbon contents and SOC stocks were remarkably lower under continuous permafrost conditions. We conclude that drier soil conditions and a shorter vegetation period compared to areas with discontinuous permafrost account for this. Moreover, these soils contain higher portions of easily decomposable POM fractions.
Although the POM fractions comprise only a small portion of the organic carbon mass balance, they contribute a large proportion on SOC stocks due to their high SOC contents. These results show that different POM fractions play specific roles under the scope of climate change: light POM fractions have short turnover rates and are particularly vulnerable to increasing temperatures in terms of potential CO2 and CH4 emission from soils.
We thank Yang Xiaoxia and Mi Zhaorong (NWIPB, CAS, China) for supporting the field campaign and Christian Wolf, Kathrin Drechsel and Sabine Flaiz (University of Tuebingen, Germany) for their assistance during laboratory analyses.
Conceived and designed the experiments: CD FB. Performed the experiments: CD FB. Analyzed the data: CD. Wrote the paper: CD PK FB JSH TS.
- 1. Jobbágy EG, Jackson RB (2000) The vertical distribution of soil organic carbon and its relation to climate and vegetation. Ecological Applications 10 (2): 423–436.
- 2. Bockheim JG, Hinkel KM, Nelson FE (2003) Predicting Carbon Storage in Tundra Soils of Arctic Alaska. Soil Sci. Soc. Am. J 67 (3): 948–950 Available: https://www.crops.org/publications/sssaj/abstracts/67/3/948.
- 3. Callesen I, Listi J, Raulund-Rasmussen K, Olsson MT, Tau-Strand L, et al. (2003) Soil carbon stores in Nordic well-drained forest soils - relationships with climate and texture class. Global Change Biology 9: 358–370.
- 4. Post WM, Emanuel WR, Zinke PJ, Stangenberger AG (1982) Soil carbon pools and world life zones. Nature 298: 156–159.
- 5. Wang G, Qian J, Cheng G, Lai Y (2002) Soil organic carbon pool of grassland soils on the Qinghai-Tibetan Plateau and its global implication. The Science of the Total Environment 291: 207–217.
- 6. Baumann F, He J, Schmidt K, Kühn P, Scholten T (2009) Pedogenesis, permafrost, and soil moisture as controlling factors for soil nitrogen and carbon contents across the Tibetan Plateau. Global Change Biology 15 (12): 3001–3017 Available: http://dx.doi.org/10.1111/j.1365-2486.2009.01953.x.
- 7. Ohtsuka T, Hirota M, Zhang X, Shimono A, Senga Y, et al. (2008) Soil organic carbon pools in alpine to nival zones along an altitudinal gradient (4400–5300 m) on the Tibetan Plateau. Polar Science 2 (4): 277–285.
- 8. Zhang Y, Tang Y, Jiang J, Yang Y (2007) Characterizing the dynamics of soil organic carbon in grasslands on the Qinghai-Tibetan Plateau. Science in China Series D: Earth Sciences 50 (1): 113–120.
- 9. Luo T, Li W, Zhu H (2002) Estimated biomass and productivity of natural vegetation on the Tibetan plateau. Ecological Applications 12 (4): 980–997 .
- 10. Kirschbaum MUF (1995) The temperature dependence of soil organic matter decomposition, and the effect of global warming on soil organic C storage. Soil Biology & Biochemistry 27 (6): 753–760.
- 11. Wagner D, Liebner S (2009) Global Warming and Carbon Dynamics in Permafrost Soils: Methane Production and Oxidation. In: Margesin R, editor. Permafrost Soils. Berlin/Heidelberg: Springer. 219–236.
- 12. Tarnocai C, Canadell JG, Schuur EAG, Kuhry P, Mazhitova G, et al. (2009) Soil organic carbon pools in the northern circumpolar permafrost region. Global Biogeochem. Cycles 23 (2): GB2023 Available: http://dx.doi.org/10.1029/2008GB003327.
- 13. Grosse G, Romanovsky V, Jorgenson T, Anthony KW, Brown J, et al. (2011) Vulnerability and Feedbacks of Permafrost to Climate Change. Eos Trans. AGU 92 (9): 73–74 .
- 14. Schuur EAG, Bockheim J, Canadell JG, Euskirchen E, Field CB, et al. (2008) Vulnerability of Permafrost Carbon to Climate Change: Implications for the Global Carbon Cycle. BioScience 58 (8): 701–714.
- 15. Torn MS, Trumbore SE, Chadwick OA, Vitousek PM, Hendricks DM (1997) Mineral control of soil organic carbon storage and turnover. Nature 389 (6647): 170–173 Available: http://dx.doi.org/10.1038/38260.
- 16. Trumbore S (2006) Carbon respired by terrestrial ecosystems – recent progress and challenges. Global Change Biology 12 (2): 141–153.
- 17. Lützow M, Kögel-Knabner I, Ekschmitt K, Flessa H, Guggenberger G, et al. (2007) SOM fractionation methods: Relevance to functional pools and to stabilization mechanisms. Soil Biology and Biochemistry 39: 2183–2207.
- 18. Du Zheng, Zhang Q, Wu S (2000) Mountain geoecology and sustainable development of the Tibetan Plateau. Dordrecht/Boston: Kluwer Academic Publishers.
- 19. Yang Y, Fang J, Tang Y, Ji C, Zheng C, et al. (2008) Storage, patterns and controls of soil organic carbon in the Tibetan grasslands. Global Change Biology 14: 1592–1599.
- 20. Zhao L, Cheng G, Li S, Zhao X, Wang S (2000) Thawing and freezing processes of active layer in Wudaoliang region of Tibetan Plateau. Chinese Science Bulletin 45: 2181–2187.
- 21. Kang SXY, You Q, Flügel W, Pepin N, Yao T (2010) Review of climate and cryospheric change in the Tibetan Plateau. Environ. Res. Lett. 5: 1–8.
- 22. Jin H, Zhao L, Wang S, Jin R (2006) Thermal regimes and degradation modes of permafrost along the Qinghai-Tibet Highway. Science in China Series D: Earth Sciences 49 (11): 1170–1183.
- 23. Yang M, Wang S, Yao T, Gou X, Lu A, et al. (2004) Desertification and its relationship with permafrost degradation in Qinghai-Xizang (Tibet) plateau. Cold Regions Science and Technology 39: 47–53.
- 24. Cheng G, Wu T (2007) Responses of permafrost to climate change and their environmental significance, Qinghai-Tibet Plateau. J. Geophys. Res 112: F02S03 .
- 25. Nan Z, Li S, Cheng G (2005) Prediction of permafrost distribution on the Qinghai-Tibet Plateau in the next 50 and 100 years. Science in China Series D: Earth Sciences 48 (6): 797–804.
- 26. Yang M, Nelson FE, Shiklomanov NI, Guo D, Wan G (2010) Permafrost degradation and its environmental effects on the Tibetan Plateau: A review of recent research. Earth-Science Reviews 103 (1–2): 31–44 Available: http://www.sciencedirect.com/science/article/pii/S0012825210000826.
- 27. Wang B, French HM (1995) Permafrost on the Tibet Plateau, China. Quaternary Science Reviews 14: 255–274.
- 28. Zhang T (2005) Historical Overview of Permafrost Studies in China. Physical Geography 26 (4): 279–298 Available: http://dx.doi.org/10.2747/0272-3618.104.22.1689.
- 29. Davidson EA, Janssens IA (2006) Temperature sensitivity of soil carbon decomposition and feedbacks to climate change. Nature 440 (7081): 165–173 Available: http://dx.doi.org/10.1038/nature04514.
- 30. Saito M, Kato T, Tang Y (2009) Temperature controls ecosystem CO2 exchange of an alpine meadow on the northeastern Tibetan Plateau. Global Change Biology 15: 221–228.
- 31. Kirschbaum MUF (2006) The temperature dependence of organic-matter decomposition – still a topic of debate. Soil Biology & Biochemistry 38: 2510–2518.
- 32. Geng Y, Wang Z, Liang C, Fang J, Baumann F, et al. (2012) Soil Respiration in Tibetan Alpine Grasslands: Belowground Biomass and Soil Moisture, but Not Soil Temperature, Best Explain the Large-Scale Patterns. PLoS ONE 7 (4): e34968 .
- 33. Wang WY, Wang QJ, Lu ZY (2009) Soil organic carbon and nitrogen content of density fractions and effect of meadow degradation to soil carbon and nitrogen of fractions in alpine Kobresia meadow. Science in China Series D: Earth Sciences 52 (5): 660–668 Available: http://dx.doi.org/10.1007/s11430-009-0056-5.
- 34. Geng Y, Wang Z, Liang C, Fang J, Baumann F, et al.. (2012) Effect of geographical range size on plant functional traits and the relationships between plant, soil and climate in Chinese grasslands. Global Ecology and Biogeography: no. Available: http://dx.doi.org/10.1111/j.1466-8238.2011.00692.x.
- 35. Ni J (2002) Carbon storage in grasslands of China. Journal of Arid Environments 50: 205–218.
- 36. Wang G, Li Y, Hu H, Wang Y (2008) Synergistic effect of vegetation and air temperature changes on soil water content in alpine frost meadow soil in the permafrost region of Qinghai-Tibet. Hydrological Processes 22: 3310–3320.
- 37. Weischet W, Endlicher W (2000) Regionale Klimatologie Teil 2. Stuttgart: Schweizerbart’sche Verlagsbuchhandlung.
- 38. Zhou H, Zhao X, Tang Y, Gu S, Zhou L (2005) Alpine grassland degradation and its control in the source regions of the Yangtze and the Yellow Rivers, China. Grassland Science 51: 191–203.
- 39. Jin H, Li S, Cheng G, Shaoling W, Li X (2000) Permafrost and climatic change in China. Global and Planetary Change 26: 387–404.
- 40. Wang G, Wang Y, Li Y, Cheng H (2007) Influences of alpine ecosystem responses to climatic change on soil properties on the Qinghai-Tibet Plateau, China. Catena 70 (3): 506–514.
- 41. IUSS Working Group WRB. 2007. World Reference Base for Soil Resources 2006, first update 2007. World Soil Resources Reports No. 103. FAO, Rome.
- 42. Kato T, Tang Y, Gu S, Cui X, Hirota M, et al. (2004) Carbon dioxide exchange between the atmosphere and an alpine meadow ecosystem on the Qinghai–Tibetan Plateau, China. Agricultural and forest meteorology 124 (1–2): 121–134 Available: http://www.sciencedirect.com/science/article/pii/S0168192304000024.
- 43. Kaiser K, Miehe G, Barthelmes A (2008) Turf-bearing topsoils on the central Tibetan Plateau, China: Pedology, botany, geochronology. Catena 73 (3): 300–311.
- 44. Grünewald G, Kaiser K, Jahn R, Guggenberger G (2006) Organic matter stabilization in young calcareous soils as revealed by density fractionation and analysis of lignin-derived constituents. Organic Geochemistry 37: 1573–1589.
- 45. Golchin A, Oades JM, Skemstad J, Clarke P (1994) Study of free and occluded organic matter in soils by 13C CP/MAS NMR spectroscopy and scanning electron microscopy. Australian Journal of Soil Research (32): 285–309.
- 46. Christensen B (1992) Physical fractionation of soil and organic matter in primary particle size and density separates. Advances in Soil Science (20): 1–90.
- 47. Roscoe R, Buurman P, Velthorst EJ (2000) Disruption of soil aggregates by varied amounts of ultrasonic energy in fractionation of organic matter of a clay Latosol: carbon, nitrogen and d13C distribution in particle-size fractions. European Journal of Soil Science 51: 445–454.
- 48. Six J, Schultz PA, Jastrow JD, Merckx R (1999) Recycling of sodium polytungstate used in soil organic matter studies. Soil Biology & Biochemistry 31: 1193–1196.
- 49. Wagner D, Kobabe S, Liebner S (2009) Bacterial community structure and carbon turnover in permafrost-affected soils of the Lena Delta, northeastern Siberia. Canadian Journal of Microbiology 55 (1): 73–83 Available: http://dx.doi.org/10.1139/W08-121.
- 50. Golchin A, Oades JM, Skjemstad J, Clarke P (1995) Structural and dynamic properties of soil organic-matter as reflected by 13C natural-abundance, pyrolysis mass-spectrometry and solid-state 13C NMR-spectroscopy in density fractions of an oxisol under forest and pasture. Australian Journal of Soil Research 33: 59–76 Available: http://dx.doi.org/10.1071/SR9950059.
- 51. Callesen I, Raulund-Rasmussen K, Westman C, Tau-Strand L (2007) Nitrogen pools and C:N ratios in well-drained Nordic forest soils related to climate and soil texture. Boreal Environment Research 12 (6): 681–692.
- 52. John B, Yamashita T, Ludwig B, Flessa H (2005) Storage of organic carbon in aggregate and density fractions of silty soils under different types of land use. Geoderma 128: 63–79.
- 53. Kölbl A, Kögel-Knabner I (2004) Content and composition of free and occluded particulate organic matter in a differently textured arable Cambisol as revealed by solid-state 13C NMR spectroscopy. Journal of Plant Nutrition and Soil Science 167 (1): 45–53 .
- 54. Wang G, Li Y, Wang Y, Wu Q (2008) Effects of permafrost thawing on vegetation and soil carbon losses on the Qinghai-Tibet Plateau, China. Geoderma 143: 143–152.
- 55. Yang YH, Fang JY, Guo DL, Ji CJ, Ma WH (2010) Vertical patterns of soil carbon, nitrogen and carbon: nitrogen stoichiometry in Tibetan grasslands. Biogeosciences Discuss. 7: 1–24.
- 56. Uhlirova E, Santruckova H, Davidov SP (2007) Quality and potential biodegradability of soil organic matter preserved in permafrost of Siberian tussock tundra. Soil Biology & Biochemistry 39: 1978–1989.
- 57. Wang G, Wang Y, Qian J, Wu Q (2006) Land cover change and its impact on soil C and N in two watersheds in the center of the Qinghai-Tibetan Plateau. Mountain Research and Development 26 (2): 153–162.
- 58. Kadono A, Funakawa S, Kosaki T (2008) Factors controlling mineralization of soil organic matter in the Eurasian steppe. Soil Biology and Biochemistry 40 (4): 947–955 Available: http://www.sciencedirect.com/science/article/pii/S0038071707004567.
- 59. Gundelwein A, Müller-Lupp T, Sommerkorn M, Haupt ETK, Pfeiffer E, et al. (2007) Carbon in tundra soils in the Lake Labaz region of arctic Siberia. European Journal of Soil Science 58 (5): 1164–1174 Available: http://dx.doi.org/10.1111/j.1365-2389.2007.00908.x.