Peili Shi

Nitrogen and carbon source-sink relationships in trees at the Himalayan treelines compared with lower elevations.

No single hypothesis or theory has been widely accepted for explaining the functional mechanism of global alpine/arctic treeline formation. The present study tested whether the alpine treeline is determined by (1) the needle nitrogen content associated with photosynthesis (carbon gain); (2) a sufficient source-sink ratio of carbon; or (3) a sufficient C-N ratio. Nitrogen does not limit the growth and development of trees studied at the Himalayan treelines. Levels of non-structural carbohydrates (NSC) in trees were species-specific and site-dependent; therefore, the treeline cases studied did not show consistent evidence of source/carbon limitation or sink/growth limitation in treeline trees. However, results of the combined three treelines showed that the treeline trees may suffer from a winter carbon shortage. The source capacity and the sink capacity of a tree influence its tissue NSC concentrations and the carbon balance; therefore, we suggest that the persistence and development of treeline trees in a harsh alpine environment may require a minimum level of the total NSC concentration, a sufficiently high sugar:starch ratio, and a balanced carbon source-sink relationship.

Changes in individual plant traits and biomass allocation in alpine meadow with elevation variation on the Qinghai-Tibetan Plateau

Plant traits and individual plant biomass allocation of 57 perennial herbaceous species, belonging to three common functional groups (forbs, grasses and sedges) at subalpine (3700 m ASL), alpine (4300 m ASL) and subnival (⩾5000 m ASL) sites were examined to test the hypothesis that at high altitudes, plants reduce the proportion of aboveground parts and allocate more biomass to belowground parts, especially storage organs, as altitude increases, so as to geminate and resist environmental stress. However, results indicate that some divergence in biomass allocation exists among organs. With increasing altitude, the mean fractions of total biomass allocated to aboveground parts decreased. The mean fractions of total biomass allocation to storage organs at the subalpine site (7%±2% S.E.) were distinct from those at the alpine (23%±6%) and subnival (21%±6%) sites, while the proportions of green leaves at all altitudes remained almost constant. At 4300 m and 5000 m, the mean fractions of flower stems decreased by 45% and 41%, respectively, while fine roots increased by 86% and 102%, respectively. Specific leaf areas and leaf areas of forbs and grasses deceased with rising elevation, while sedges showed opposite trends. For all three functional groups, leaf area ratio and leaf area root mass ratio decreased, while fine root biomass increased at higher altitudes. Biomass allocation patterns of alpine plants were characterized by a reduction in aboveground reproductive organs and enlargement of fine roots, while the proportion of leaves remained stable. It was beneficial for high altitude plants to compensate carbon gain and nutrient uptake under low temperature and limited nutrients by stabilizing biomass investment to photosynthetic structures and increasing the absorption surface area of fine roots. In contrast to forbs and grasses that had high mycorrhizal infection, sedges had higher single leaf area and more root fraction, especially fine roots.

Effects of Grazing Exclusion on Plant Productivity and Soil Carbon, Nitrogen Storage in Alpine Meadows in Northern Tibet, China

Grazing exclusion is widely adopted in restoring degraded alpine grasslands on the Qinghai-Tibetan Plateau. However, its effectiveness remains poorly understood. In this study, we investigated the effects of grazing exclusion on plant productivity, species diversity and soil organic carbon (SOC) and soil total nitrogen (STN) storage along a transect spanning from east to west of alpine meadows in northern Tibet, China. After six years of grazing exclusion, plant cover, aboveground biomass (AGB), belowground biomass (BGB), SOC and STN were increased, but species diversity indices declined. The enhancement of AGB and SOC caused by grazing exclusion was correlated positively with mean annual precipitation (MAP). Grazing exclusion led to remarkable biomass increase of sedge species, especially Kobresia pygmaea, whereas decrease of biomass in forbs and no obvious change in grass, leguminous and noxious species. Root biomass was concentrated in the near surface layer (10 cm) after grazing exclusion. The effects of grazing exclusion on SOC storage were confined to shallow soil layer in sites with lower MAP. It is indicated that grazing exclusion is an effective measure to increase forage production and enhance soil carbon sequestration in the studied region. The effect is more efficient in sites with higher precipitation. However, the results revealed a tradeoff between vegetation restoration and ecological biodiversity. Therefore, carbon pools recover more quickly than plant biodiversity in the alpine meadows. We suggest that grazing exclusion should be combined with other measures to reconcile grassland restoration and biodiversity conservation.

Experimental warming does not enhance gross primary production and above-ground biomass in the alpine meadow of Tibet

Abstract In order to understand the response of gross primary production (GPP) and above-ground biomass (AGB) to warming, a field warming experiment using open-top chambers was conducted in an alpine meadow at three elevations (i.e., 4313, 4513, and 4693 m) on the Northern Tibetan Plateau in May 2010. We calculated GPP from the moderate-resolution imaging spectroradiometer algorithm and AGB using the surface measured data in 2012. Average GPP and AGB at elevation 4313 m was significantly decreased by experimental warming, whereas the declines at elevations 4513 and 4693 m were not statistically significant across all sampling dates. The negative effects of experimental warming on GPP and AGB may be related to experimental warming-induced soil drying. The different responses of GPP and AGB to experimental warming among the three alpine meadow sites could be dependent on climate conditions. Our findings suggested that experimental warming did not enhance GPP and AGB in the alpine meadow, and its effects differed among alpine meadows on the Tibetan Plateau.

Calibration of MODIS-based gross primary production over an alpine meadow on the Tibetan Plateau

Moderate-resolution imaging spectroradiometer (MODIS) gross primary production (GPP) was compared with estimated GPP (GPP_EC) from eddy covariance measurements over an alpine meadow on the Tibetan Plateau in 2005–2007. The MODIS GPP (GPP_MOD17A2) with a bias of −0.38 g C m−2 d−1 (i.e., about −40.58% of the mean of the GPP_EC) strongly underestimated the GPP_EC for the alpine meadow. The MODIS GPP was recalibrated using measured surface meteorological data, including photosynthetically active radiation (PAR), daily minimum air temperature (Tamin) and daytime mean vapor pressure deficit (VPD), revised fractional photosynthetically active radiation (FPAR), and the revised maximum light use efficiency (LUEmax) of 0.81 g C MJ−1 (compared with the default value of 0.68 g C MJ−1 for grassland in the MODIS GPP algorithm) for the alpine meadow. The MODIS-based FPAR was about 14.70% larger than the surface-estimated FPAR using surface-measured leaf area index (LAI) data. Additionally, the temporal resolution of surface-measured LAI data was relatively low. Therefore, the linear relationship between surface-measured LAI and MODIS-based LAI was established (R2 > 0.80, P < 0.001). Then the revised MODIS LAI datasets were used to calculate the revised FPAR. The revised LUEmax was optimized from the MOD17A2 algorithm using daily surface measurements, including LAI, PAR, VPD, Tamin and GPP_EC. The calibrated MOD17A2 algorithm could explain 88% of GPP_EC variance for the alpine meadow. The bias between GPP_MOD17A2 and calculated GPP from the MOD17A2 algorithm using surface-measured PAR, Tamin, and VPD, MODIS-based FPAR, and the default LUEmax of 0.68 g C MJ−1 was −0.17 g C m−2 d−1 (i.e., about −17.60% of the mean of the GPP_EC). The underestimation of LUEmax caused a 13.78% underestimation of GPP. In contrast, the overestimation of FPAR resulted in a 7.17% overestimation of GPP. The net effect of meteorology data and FPAR resulted in a 13.84% underestimation of GPP. These results showed that MODIS-based meteorology data, FPAR, and LUEmax for the alpine meadow needed to be adjusted.

Response of Soil Respiration to Grazing in an Alpine Meadow at Three Elevations in Tibet

Alpine meadows are one major type of pastureland on the Tibetan Plateau. However, few studies have evaluated the response of soil respiration (R s) to grazing along an elevation gradient in an alpine meadow on the Tibetan Plateau. Here three fenced enclosures were established in an alpine meadow at three elevations (i.e., 4313 m, 4513 m, and 4693 m) in July 2008. We measured R s inside and outside the three fenced enclosures in July–September, 2010-2011. Topsoil (0–20 cm) samples were gathered in July, August, and September, 2011. There were no significant differences for R s, dissolved organic C (DOC), and belowground root biomass (BGB) between the grazed and ungrazed soils. Soil respiration was positively correlated with soil organic C (SOC), microbial biomass (MBC), DOC, and BGB. In addition, both R s and BGB increased with total N (TN), the ratio of SOC to TN, ammonium N (NH4 +-N), and the ratio of NH4 +-N to nitrate N. Our findings suggested that the negligible response of R s to grazing could be directly attributed to that of respiration substrate and that soil N may indirectly affect R s by its effect on BGB.

Responses of ecosystem CO 2 fluxes to short-term experimental warming and nitrogen enrichment in an Alpine meadow, northern Tibet Plateau.

Over the past decades, the Tibetan Plateau has experienced pronounced warming, yet the extent to which warming will affect alpine ecosystems depends on how warming interacts with other influential global change factors, such as nitrogen (N) deposition. A long-term warming and N manipulation experiment was established to investigate the interactive effects of warming and N deposition on alpine meadow. Open-top chambers were used to simulate warming. N addition, warming, N addition × warming, and a control were set up. In OTCs, daytime air and soil temperature were warmed by 2.0°C and 1.6°C above ambient conditions, but soil moisture was decreased by 4.95 m3 m−3. N addition enhanced ecosystem respiration (Reco); nevertheless, warming significantly decreased Reco. The decline of Reco resulting from warming was cancelled out by N addition in late growing season. Our results suggested that N addition enhanced Reco by increasing soil N availability and plant production, whereas warming decreased Reco through lowering soil moisture, soil N supply potential, and suppression of plant activity. Furthermore, season-specific responses of Reco indicated that warming and N deposition caused by future global change may have complicated influence on carbon cycles in alpine ecosystems.

Biomass Allocation Patterns of Alpine Grassland Species and Functional Groups along a Precipitation Gradient on the Northern Tibetan Plateau

Variations in the fractions of biomass allocated to functional components are widely considered as plant responses to resource availability for grassland plants. Observations indicated shoots isometrically relates to roots at the community level but allometrically at the species level in Tibetan alpine grasslands. These differences may result from the specific complementarity of functional groups between functional components, such as leaf, root, stem and reproductive organ. To test the component complementary responses to regional moisture variation, we conducted a multi-site transect survey to measure plant individual size and component biomass fractions of common species belonging to the functional groups: forbs, grasses, legumes and sedges on the Northern Tibetan Plateau in peak growing season in 2010. Along the mean annual precipitation (MAP) gradient, we sampled 70 species, in which 20 are in alpine meadows, 20 in alpine steppes, 15 in alpine desert-steppes and 15 in alpine deserts, respectively. Our results showed that the size of alpine plants is small with individual biomass mostly lower than 1.0 g. Plants keep relative conservative component fractions across alpine grasslands at the individual level. However, the complementary responses between functional components to moisture variations specifically differ among functional groups. These results indicate that functional group diversity may be an effective tool for scaling biomass allocation patterns from individual up to community level. Therefore, it is necessary and valuable to perform intensive and systematic studies on identification and differentiation the influences of compositional changes in functional groups on ecosystem primary services and processes.

Litter species traits, but not richness, contribute to carbon and nitrogen dynamics in an alpine meadow on the Tibetan Plateau

AimsLitter, as afterlife of plants, plays an important role in driving belowground decomposition processes. Here we tested effects of litter species identity and diversity on carbon (C) and nitrogen (N) dynamics during litter decomposition in N-limited alpine meadow soil from the Qinghai–Tibet Plateau.MethodsWe incubated litters of four meadow species, a sedge (“S”, Kobresia humilis), a grass (“G”, Elymus nutans), a herb (“H”, Saussurea superba), and a legume (“L”, Oxytropis falcata), in monoculture and in mixture with meadow soil. CO2 release was measured 21 times during the incubation, and soil available N and microbial biomass C and N were measured before and after the experiment.ResultsThe organic C decay rate did not differ much among soils amended with monocultures or mixtures of litter, except in the H, S, L, and S+H treatments, which had much higher decay rates. Potential decomposable C pools were lowest in the control, highest in the L treatment, and intermediate in the S treatment. Mineralized N was completely immobilized by soil microbes in all treatments except the control, S+L, and S+G+L treatments. Litter mixtures had both additive and non-additive effects on CO2-C emission (mainly antagonistic effects), net N mineralization (mainly synergistic), and microbial biomass C and N (both). Overall, these parameters were not significantly correlated with litter species richness. Similarly, microbial C or N was not significantly correlated with litter N content or C/N. However, cumulative CO2-C emission and net N mineralization were positively correlated with litter N content and negatively correlated with litter C/N.ConclusionsLitter N content and C/N rather than litter species richness drove the release of CO2-C and net available N in this ecosystem. The antagonistic effects of litter mixtures contributed to a modest release of CO2-C, but their synergistic effects enhanced net available N. We suggest that in alpine meadow communities, balancing species with high and low N contents will benefit soil carbon sequestration and plant competition for available N with soil microbes.

Correlation Between CO2 Efflux and Net Nitrogen Mineralization and Its Response to External C or N Supply in an Alpine Meadow Soil

In nutrient-limited alpine meadows, nitrogen (N) mineralization is prior to soil microbial immobilization; therefore, increased mineral N supply would be most likely immobilized by soil microbes due to nutrient shortage in alpine soils. In addition, low temperature in alpine meadows might be one of the primary factors limiting soil organic matter decomposition and thus N mineralization. A laboratory incubation experiment was performed using an alpine meadow soil from the Tibetan Plateau. Two levels of NH4NO3 (N) or glucose (C) were added, with a blank without addition of C or N as the control, before incubation at 5, 15, or 25 ◦ C for 28 d. CO2 efflux was measured during the 28-d incubation, and the mineral N was measured at the beginning and end of the incubation, in order to test two hypotheses: 1) net N mineralization is negatively correlated with CO2 efflux for the control and 2) the external labile N or C supply will shift the negative correlation to positive. The results showed a negative correlation between CO2 efflux and net N immobilization in the control. External inorganic N supply did not change the negative correlation. The external labile C supply shifted the linear correlation from negative to positive under the low C addition level. However, under the high C level, no correlation was found. These suggested that the correlation of CO2 efflux to net N mineralization strongly depend on soil labile C and C:N ratio regardless of temperatures. Further research should focus on the effects of the types and the amount of litter components on interactions of C and N during soil organic matter decomposition.