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Partitioning sources of ecosystem and soil respiration (Reco and Rs) is important for understanding how climate change affects carbon cycling. Plant and microbial biomass analyses and daytime measurements of Reco and Rs were performed for 25 plots in an alpine meadow at elevation 4313 m on the Tibetan Plateau. Plant and microbial biomass were determined by harvesting method and the chloroform fumigation-extraction method, respectively. Respiration fluxes were measured by an automated CO2 flux system (LI-8100, LI-COR Biosciences, Lincoln, NE, USA). Soil respiration can be estimated by a linear or exponential relationship between Reco and aboveground plant biomass (AGB). Microbial respiration (Rm) can be estimated by a linear or exponential relationship between Rs and belowground plant biomass (BGB) or by a multiple relationship between Reco and AGB and BGB. Soil respiration (or Rm) is respiration flux when AGB (or BGB) is extrapolated to zero for the linear and exponential regression methods. Similarly, Rm is respiration flux when both AGB and BGB are zero for the multiple regression method. Our findings suggest that the exponential regression method to partition sources of Reco and Rs may be more appropriate compared to other methods for this alpine meadow of Tibet.
A warming experiment with two magnitudes was performed in an alpine meadow of Northern Tibet since late June, 2013. Open top chambers (OTCs) with two top diameters (0.60 m and 1.00 m) were used to increase soil temperature. Soil respiration (Rs) was measured during the growing season in 2013–2014. The OTCs with top diameters of 1.00 m and 0.60 m increased soil temperature by 1.30 and 3.10oC, respectively, during the whole study period, but decreased soil moisture by 0.02 and 0.05 m³ m⁻³, respectively. However, the two patters of OTCs did not affect Rs. These results implied that a higher warming did not result in a higher Rs but a greater soil drying. Therefore, a higher warming may not cause a higher soil respiration, which was most likely due to the fact that a higher warming may result in a greater soil drying.
Three types of alpine plant species,Carex montis-everestii,Quercus aquifolioidesandStipa capillacea, along an altitudinal gradient of 3005–5025 m on the Tibetan Plateau, were chosen to test the generality of the hypothesis that foliar carbon isotope composition (δ¹³C) of C3 plants increases significantly with altitude and to determine climate drivers shaping its altitudinal pattern. Temperature and relative humidity showed significantly negative correlations with altitude; however, precipitation and soil water potential remained unchanged with altitude. Foliar δ¹³C of C. montis-everestii,Q. aquifolioides,S. capillaceaalone or combined together did not significantly increase with altitude, which does not support the leading hypothesis of increased foliar δ¹³C with altitude. There was no difference in foliar δ¹³C among all three species. Multi-factor correlation analyses showed that temperature, precipitation and relative humidity alone did not affect foliar δ¹³C ofC. montis-everestiiandS. capillacea, but conferred significant effects on foliar δ¹³C of Q. aquifolioides.
The great sensitivity of the response of alpine plant community to climate change makes the identification of these responses important. In 2007, we conducted a reciprocal translocation experiment on 100 × 100 × 40 cm coherent turf and soil along an elevation gradient of 3200–3800 m on the south slope of Qilian Mountains northeast of the Qinghai-Tibetan Plateau. The aim was to understand the warming/cooling effects on the alpine ecosystem where treatments were simulated by donor elevations below/above receptors. Translocated vegetation comprised the Kobresia meadow at 3200 m, deciduous shrub meadow at 3400 m, forbs meadow at 3600 m, and sparse vegetation at 3800 m. The 5 × 5 cm grid method (50 × 50 cm, 100 grids) was used for surveying plant species absolute abundance in translocated quadrats. Results showed that species richness and Shannon-Weaver index of Kobresia meadow increased significantly (P <0.05) when translocated to 3400 m. Shannon-Weaver index of shrub meadow declined, while shrub species abundance responded slightly both to warming and cooling treatments. Both species richness and Shannon-Weaver index of forbs meadow and sparse vegetation were enhanced evidently at 3200 m and 3400 m. Four groups were identified by non-metric multidimensional scaling based on receptor elevation. Responses of the alpine plant community and the function group appeared to be specific to climate magnitude and specific to function type, respectively. Correlation indicated that climatic factors played a much more important role than soil in the response of the alpine plant community. Four vegetation types were sensitive to climate change, while Kobresia meadow behaved flexibly. Global warming would depress sedges but favor legumes and graminoids.
The effects of the growing season climatic factors (i.e., temperature, precipitation, vapor pressure and relative humidity) on the growing season maximum normalized difference vegetation index (MNDVI), which can mirror the aboveground net primary production and the vegetation maximum absorbed ability of photosynthetically active radiation, were examined during the period from 2000 to 2012 on the Tibetan Plateau. The effects of climatic factors on the MNDVI changed with vegetation types, which was probably due to the fact that the changes of climatic factors differed with the type of vegetation. There was a significant increasing trend for the spatially averaged MNDVI of the vegetation area over the entire plateau. Approximately 16 and 3% of the vegetation area demonstrated a significant MNDVI increasing and decreasing trend, respectively. The MNDVI was significantly affected by relative humidity and vapor pressure, but not affected by temperature and precipitation over the entire plateau. Our findings suggested that the environmental humidity played a predominant role in affecting the variation of MNDVI over the entire Tibetan Plateau.
Nutrients may be mobilized from senescing leaves and transported to other plant tissues, enabling plants to conserve them and reuse. Nitrogen and phosphorus are two dominant nutrients related to photosynthetic capacity and limiting plant growth. In this study, we examined the effect of canopy height on nutrient use by analyzing N and P concentrations of green and senescent leaves collected from different canopy heights of Abies georgei var. smithii (Viguie et Gaussen) Cheng growing at the alpine timberline in Sergyemla Mt. (4 350 m a.s.l), southeastern Tibetan Plateau. The results showed that N and P concentrations per unit needle mass decreased significantly with needle age, but no significant difference was found among upper, middle and lower canopy. However, area-based N and P concentrations increased with the height of canopy. The leaf level nutrient resorption and use efficiency varied in the order: upper canopy> middle canopy >lower canopy for both N and P. The higher nutrient resorption efficiency had significant relationship with leaf level nutrient use efficiency, that is, higher leaf level nutrient use efficiency was partly due to the high resorption from senescent needles. Additionally, the higher nutrient resorption was related to high current nutrient concentration. Vertical variations of leaf level nutrient use efficiency in this study reflected the strategy of alpine trees to respond to imbalance between light availability and soil nutrients.
Methane (CH4) emission was measured from an open fen on the Zoige Plateau (3500 m a.s.l.) (the eastern edge of the Qinghai- Tibetan Plateau) during two winters of 2006 and 2007. Three dominant plant stands, including Carex muliensis Hand-Mazz. (CM), Eleocharis valleculosa Ohwif. setosa (Ohwi) Kitagawa.(EV) and Kobresia tibetica Maxim (KT) were chosen to be monitored. Winter CH4 emissions were roughly estimated to be 0.94 mg CH4 m–2 h–1. High spatialtemporal variations of the emission were found in this fen (the sequence of CM> EV> KT; 0.63 and 1.24 mg CH4 m–2 h–1 for 2006 and 2007, respectively). Factors involved in the spatial-temporal variation were: 1) water table in summer determining the winter amount of “old” CH4 stored in peat; 2) ice layer determining the release of CH4; 3) plant growth determining both the quantity of CH4 stored in peat and available substrates for CH4 production in winter. However, due to the homogeneity of freezing in winters, predictive factors such as plant growth and water table in summer could contribute more to winter CH4 emission than in situ freezing conditions. Considering that plant growth and water table are also the key factors controlling the spatial-temporal variation of CH4 emission in summer, we therefore suggested that winter CH4 emission represents the “inertia” of summer CH4 emission.
The general impact of extra nitrogen on ecological stoichiometry was examined in alpine grasslands on the Tibetan Plateau. Extra nitrogen increased the ratio of nitrogen to phosphorus (N:P ratio) in leaves and aboveground parts of plants by 43.4% and 32.7%, respectively. In contrast, extra nitrogen reduced the ratio of carbon to nitrogen (C:N ratio) in leaves by 30.6%. Extra nitrogen decreased soil C:N ratio by 9.1% in alpine meadows, but increased soil C:N ratio by 3.4% in alpine steppes. Extra urea had a stronger positive impact on aboveground vegetation N:P ratio than did extra ammonium nitrate. Extra urea rather than ammonium nitrate decreased aboveground vegetation C:N ratio and soil C:N ratio. The impact of extra nitrogen on aboveground vegetation N:P ratio was positively correlated with latitude, mean annual temperature and precipitation, nitrogen application rate and accumulated amount, but negatively correlated with elevation, duration and aboveground vegetation N:P ratio of the control plots. The impact of extra nitrogen on leaves N:P ratio was positively correlated with nitrogen application rate and accumulated amount. The impact of extra nitrogen on leaves C:N ratio was positively correlated with latitude, but negatively correlated with mean annual temperature and precipitation, nitrogen application rate, accumulated amount, duration and leaves C:N ratio of the control plots. Therefore, nitrogen enrichment caused by human activities will most likely alter element balance and alpine plants from nitrogen limitation to phosphorus limitation. This effect may weaken with time, and increase with climatic warming, increased precipitation and nitrogen input rate.
Microbes remain active and play an important role in soil nitrogen (N) cycle during the winter in soil of the alpine zone. A shift from microbial N immobilization process dominant during summer to prevailing microbial mineralization process during the winter is observed. Warmer soil under deep snow cover may increase the microbial activity and rate of organic matter decomposition over the winter. Furthermore, severe shortages of dissolved carbon (C) in the winter may cause microbial mortality and lyses. Thus, C limitation on microbial growth and activity may have an important effect on winter N mineralization and even on soil N pools. However, the combined effects of additional organic C (litter inputs) and snow cover on soil N biogeochemical processes in the Tibetan Plateau remain unclear. In the current study, the in situ effects of snowpack and litter decomposition on N dynamics in the alpine zone of the Eastern Tibetan Plateau were investigated. Intact soil core incubations in three different snow regimes (0, 30 and 100 cm depth snow) in the winter were used to solve the problem by measure concentrations of mineral form of soil N. In addition, the litter bag method was used to analyze the litter decomposition over the winter. Our results indicate that the snow cover reduced the ammonium (NH₄⁺-N) content, accelerate N mineralization in soil, and did not significantly change the dissolvable organic nitrogen (DON) and microbial biomass nitrogen (MBN). Meanwhile, snowpack increased the litter N content and accelerated litter decomposition in late winter. Litter addition reduced the MBN and NH₄⁺ -N contents in soil, but increased the nitrate (NO₃⁻ -N) content and net N mineralization, suggesting that N availability to plants during the spring thaw period may be enhanced.
Artificial grassland establishment is one of the fastest and most effective ways to restore the productivity of degraded grasslands. Little is known about the effect of different types of artificial grassland establishment (i.e., single- and mixed-sowing grassland establishment with perennial grasses) on soil seed bank in degraded grassland ecosystems. Single-sowing population of a high yield species usually has a great standing biomass causing shading that may inhibit germination of seeds in soil seed bank. Thus, we hypothesized that there is higher species richness and seed density in the soil seed bank of single-sowing than mixed-sowing grasslands. Here, we investigated the soil seed bank in four-year old single-sowing and mixed-sowing and control (degraded) grasslands on the Qinghai-Tibetan Plateau. We found that the autumn seed bank of mixed-sowing grasslands had lower species richness and seed density than single-sowing grasslands, while the summer soil seed bank (persistent seed bank) showed little differences. There were differences in biomass among the three grasslands (single-sowing > mixed-sowing > control), but there was no differences in species richness of vegetation. In sum, our results of the autumn seed bank support our hypothesis. Greater above-ground biomass in single-sowing grasslands could generally cause low light availability preventing seeds in soil from germinating and support more seed output, which both may indirectly or directly result in the relatively higher species richness and seed density in the soil seed bank. Our results also suggest that artificial grasslands usually returning to native grasslands in terms of production and species composition after several years is likely due to stability of the persistent soil seed bank.
Habitat loss is one of the main factors impacting endangered wildlife in China. The Tibetan foxVulpes ferrilata Hodgson, 1842 is a characteristic species of the Tibetan Plateau. However, its habitat use is poorly known. We conducted a project (2001–2003) to better understand habitat use by the Tibetan fox in Shiqu County, Sichuan Province, China. The research site was classified into four land cover categories: grassland, grassland and shrubs, shrubs, and disturbed area (more than half of the area in a plot covered by bare soil and gravel). Their proportional share in land cover categories was estimated by line transects and used as expected land cover. Plots were also located at fox locations, fox dens, and fox feces (hereafter “fox plots”). Using Bonferroniz-statistics, land cover proportions in fox plots were compared with the expected land cover proportions. Tibetan foxes were found primarily in grassland habitat and there was evidence of use of grassland/shrub habitat and disturbed area. However, the fox were rarely observed in shrubs. Two reasons for this are: (1) the need of prey, pika, which mainly lives in open areas; (2) the requirement of security from natural enemies.
Soil phosphorus (P) availability and fractions are influenced to a large extent by land use and cover changes. Inorganic P (IP) and organic P (OP) fractions in surface soils (0–20 cm) under typical vegetation types, including subalpine coniferous forests, alpine shrubs, and alpine shrub-meadows, near the alpine timberline of the eastern Tibetan Plateau of China, were measured by a modified Hedley fraction method. The results showed that OP is the dominant soil P fraction and the main source of available P in alpine soils near the timberline. Soil organic carbon, total nitrogen, and total P contents were higher in subalpine coniferous forests than in alpine shrubs and alpine shrub-meadows. Concentrations of soil labile P (the sums of Resin–IP, NaHCO3–IP, and NaHCO3–OP) were higher in subalpine coniferous forests than in alpine shrubs and alpine shrub-meadows, an observation that may be partially ascribed to the presence of deep litter layers generated by trees. Concentrations of soil labile and moderately organic P (NaHCO3–OP and NaOH–OP) in subalpine coniferous forests were also greater than in alpine shrubs and alpine shrub-meadows. Greater amounts of soil stable OP (extracted by concentrated HCl and cHCl–OP) were accumulated in alpine shrub-meadows compared to alpine shrubs or subalpine coniferous forests. The reduced availability of OP may be attributed mainly to increasing recalcitrant soil organic matter input in alpine shrub-meadows and alpine shrubs. Concentrations of IP associated with Ca minerals and parent materials (extracted by diluted HCl and HCl–IP, and extracted by concentrated HCl and cHCl–IP, respectively) were lower in subalpine coniferous forests, indicating that coniferous forests are more likely to use recalcitrant IP than alpine shrubs and alpine shrub-meadows. In this alpine region, land cover changes from subalpine coniferous forests to alpine shrubs and alpine shrub-meadows near the alpine timberline could decrease soil P conservation, availability, and supplementation.
Seed mass is a critical lifehistory character in seed evolutionary ecology. Plant species can present responses in seed mass to environment stresses. We tested the hypotheses that seed mass was positively correlated with altitude within species. We selected four congeneric Saussurea species as study objects, and collected their seeds along altitudinal gradients (2100–4200 m a.s.l.) in the alpine area of the Qinghai-Tibetan Plateau, China. Results showed that mean seed mass of the four species were significantly affected by altitude (P <0.001). There was a general trend of an increase in seed weight with altitude among the populations of the four species. In addition, mean seed mass of four species were not significantly different, but all presented a bigger coefficients of variation within species along altitude gradients. Our results indicate selection pressure within species, with larger seeds occurring at higher altitudes.
Natural regeneration of forest depends on the light regimes of floor. Point-based methods such as fisheye photo and radiometer can not provide a full panorama of light regime of heterogeneous forest stand. Eastern Tibetan Plateau is a major forest belt characteristic of diverse forest type and topographic differentiation. Understanding the trend of changes of light regime along succession series of forest may be helpful for the management of ecosystems. Fragmented forest patches due to tectonic activity and human intervention have made this prediction difficult. We use a spatially explicit forest stand light model (tRAYci) to simulate light distribution within forest in typical subalpine forest succession series of eastern Tibetan Plateau. Due to the spatial heterogeneity of tree distribution in the subalpine area, the forest stand can be approximated with a spatially explicit model of trees. Three typical subalpine forest stands (Sabina forest (SF), Fir forest (FF) and Birch forest (BF)) are selected in the eastern Tibetan Plateau. The dominant species are sabina (Sabina saltuaria (Rehd. et Wils.) Cheng), fir (Abies faxoniana Rehd. et Wils.) and birch (Betula platyphylla Suk.) for each stand and they are spatially clumped in distribution. They represent old growth coniferous forest (SF, 330 years old), coniferous-broadleaved forest (FF, 180 ys) and pioneer broadleaved forest (BF, 40 ys). The parameters of the three-dimensional model of trees are calibrated with field measurements. The simulated values are generally consistent with observed values of radiation measured by radiometers installed in these stands and values derived from fisheye photos. Test failures may be caused by the incomplete submodel of crown as a gap free one. Light regimes in old growth and pioneer forest are much more heterogeneous than intermediate stages of forest. Light regimes of these forests are also reflected by the composition of understory herb layers.
In alpine zones, cold season processes, particularly those associated with snow accumulation and ablation, have a central role in ecosystem functioning. However, we know very little about soil carbon and nitrogen processes under the snowpack in these ecosystems, including the Tibetan Plateau. We conducted an experiment comparing three snow regimes (11 m × 1 m plots) of different snow depths and durations at an altitude of 4,100 m in the Minshan Range on the eastern Tibetan Plateau. The three snow regimes included a shallow and short duration snowpack (SS; depth <10 cm), a moderate snow depth and medium duration snowpack (MS; depth <20 cm), as well as a deep and long duration snowpack (DS; depth > 30 cm). This study explores the effects of different snow conditions on soil temperature, and further describes the sequence and timing of dissolved nutrients and microbial biomass carbon (MBC) and microbial biomass nitrogen (MBN) in soils under different snow regimes during the autumn-winter transition (i.e. November 7, 2008 – March 7, 2009). Three successive phases of temperature change were distinguished: I – initial decline – soil temperatures dropped steadily from 4°C to about 0°C at the same rate for all three snow regimes; II – moderate freezing – soil temperatures fluctuated between 0°C and –7°C under all three snow regimes; III – multiple freeze-thaw cycles took place in the SS and MS regimes, but permanent freezing occurred in the DS regime. Under moderate freezing, we found that soil temperature fluctuation was an essential factor for the transformation of soil C and N. Our results indicate that larger temperature fluctuations correlate with a greater increase in dissolved organic nitrogen (DON) content. Dissolved organic carbon (DOC) content increased markedly only under the most drastic temperature fluctuations. In contrast, MBC content increased significantly only when soil temperatures were relatively steady. Under the permanent freezing, only a large number of freeze-thaw cycles caused a significant decline of NO₃⁻–N and DOC concentrations. DON content declined markedly under permanent freezing and multiple freeze-thaw cycles. However, MBC content declined significantly only under permanent freezing. Ultimately, multiple freeze-thaw cycles resulted in the export of dissolved nutrients (organic and inorganic nitrogen) from the alpine ecosystem which had previously accumulated in the moderate freezing phase of the soil.
Field me asured N2O emissions in two years were used to parameterize and validate a process-based model, DNDC, for an alpine Kobresia humilis meadow on the Tibetan Plateau in China. Although this model failed to capture the N2O fluxes in some time periods in the spring or autumn, the modeled results showed overall a good performance in terms of simulating the seasonal variation of N2O fluxes and quantifying the annual total emissions. The relative deviation on the annual basis was about 12.4% and –15.9% for the two years, respectively. The modeled data showed that nitrification contributed about 53% of total N2O production, slightly higher than denitrification. The modeled fluxes were sensitive to soil organic content (SOC), pH, and temperature, but less sensitive to variation of precipitation, soil ammonium and nitrate contents. Further mo difications for the model were suggested to focus on the process of soil freezing and thawing as well as the crop growth sub-model that would improve the model’s performance for quantifying N2O emission from the alpine meadow.
Seed mass and light availability are important factors which affect seedling recruitment in plant populations. Effects of seed mass and light availability on seed germination and seedling recruitment in Artemisia sieversiana Willd, which is a widely distributed herbs and a common component species in alpine meadows of the Tibetan Plateau, were tested in a manipulative field experiment. Our research indicated that seed mass and light significantly affected seed germination and seedling emergence and survival and there was a significant interaction effect between seed mass and light availability on these life traits within species. Our study showed that larger seeds presented significant advantages in germination, seedling emergence and survival than smaller seeds under different light availability, especially under deep shade. Our results suggested that seedling success of Artemisia sieversiana appears to be regulated by an interactive effect of seed mass and light availability in alpine meadow of the Qinghai-Tibetan Plateau. In addition, seedlings from larger seeds have more and more stronger advantages from germination, emergence to survival stage than seedlings from smaller seed within species.
Ligularia virgaurea (Maxim.) is a typical naturally-occurring native noxious weed, widely distributed in alpine grasslands of the Tibetan Plateau, China. Three field sampling plots (30 m ´ 50 m) dominated by L. virgaurea were selected to study its population colonization mode and the relationship between sexual- and clonalrecruitment in alpine meadow of the QinghaiTibetan Plateau, NW China. Field investigations were conducted on its soil seed bank, seed rain and the individuals of new recruitment (seedlings and ramets) to study its sexual and asexual recruitment. And, 46 individuals which produced the seeds were selected randomly to study their relationship between seed production and ramet production. Results showed that there were more ramets (26.23 ramets m⁻²) and less seedlings (6.70 seedlings m⁻²) and a mean value of seed rain was close to 8.04 seeds m⁻², but the soil seed bank for this species was not found in study sites. Significantly negative correlations (r = – 0.416, P <0.001) between seed number per individual and ramet number per adult individual were found for L. vrigaurea in studied alpine grasslands. Our results revealed that clonal reproduction was the main population colonization mode. In addition, there was a significantly negative relationship between seed production and ramets for this species.
In the present study, we selected a total of 26 Salix sphaeronymphe Gorz shrubs of various sizes on a cutover in the eastern Tibetan Plateau to evaluate the effects of shrub size on richness, cover and biomass of the understorey herbaceous community, grasses and forbs, as well as litter cover and biomass. Results indicated that the richness of herbaceous community, grasses and forbs significantly increased with the increased area of both undershrub canopies and open field. However, the cover and biomass of the herbaceous community beneath the shrub canopies decreased with increasing shrub size. Grasses and forbs, as different functional groups, responded differently to the increasing area of shrubs: the cover and biomass of grasses decreased while those of forbs increased. The cover and biomass of herbaceous community, grasses and forbs in the open field did not obviously vary with area. Under the shrub canopies, the cover and mass of litter positively correlated to area, but this was not the case in the open field. Our results suggest that shrubs have a positive effect on diversity and a negative effect on the cover and productivity of the herbaceous community in forest secondary succession on the alpine cutovers, and that these effects are size–dependent. Moreover, different functional groups of herbaceous plants can respond differently to the presence of shrubs.
The aim of this study was to assess initial effects of warming on the plant growth, soil nutrient contents, microbial biomass and enzyme activities of two most widespread ecosystem types: swamp meadow (deep soil, high water content) and alpine meadow (poor soil, low water content), in the hinterland of the Qinghai-Tibet Plateau (altitude 4600–4800 m a.s.l.) The temperature of air and upper soil layer was passively enhanced using open-top chambers (OTCs) (50 cm high with 60 cm at opening and constructed of 6 mm thick translucent synthetic glass) from 2006 to 2008. The use of OTCs clearly raised temperature and decreased soil moisture. In warmed plots, monthly mean air temperature was increased by 2.10oC and 2.28oC, soil moisture of 20 cm soil layer was decreased by 2.5% and 3.9% in alpine meadow and swamp meadow, respectively. Plant biomass significantly increased by 31% in alpine meadow and 67% in swamp meadow. Warming also affected soil microbial biomass C and N at both meadows. In swamp meadow, warming caused the decrease of soil organic carbon and total nitrogen in 0–5 cm layer and an increase in 5–20 cm. While in alpine meadow, these soil parameters increased in 0–5 cm layer and decreased in 5–20 cm layer. The effects of warming on enzyme activities differed depending on the enzyme and the meadow ecosystem. In general, enzyme activities were higher in the upper soil layers (0–5 cm) than in the lower soil layers (5–20 cm). The experiment results exhibited that warming improved the soil biochemical and microbiological conditions in high- mountain meadows, at least in the short term.
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