Experimental warming, not grazing, decreases rangeland quality on the Tibetan Plateau

We investigated experimental warming and simulated grazing ( clipping) effects on rangeland quality, as indicated by vegetation production and nutritive quality, in winter-grazed meadows and summer- grazed shrublands on the Tibetan Plateau, a rangeland system experiencing climatic and pastoral land use changes. Warming decreased total aboveground net primary productivity ( ANPP) by 40 g . m(-2) . yr(-1) at the meadow habitats and decreased palatable ANPP ( total ANPP minus non- palatable forb ANPP) by 10 g . m(-2) . yr(-1) at both habitats. The decreased production of the medicinal forb Gentiana straminea and the increased production of the non- palatable forb Stellera chamaejasme with warming also reduced rangeland quality. At the shrubland habitats, warming resulted in less digestible shrubs, whose foliage contains 25% digestible dry matter ( DDM), replacing more digestible graminoids, whose foliage contains 60% DDM. This shift from graminoids to shrubs not only results in lower- quality forage, but could also have important consequences for future domestic herd composition. Although warming extended the growing season in non- clipped plots, the reduced rangeland quality due to decreased vegetative production and nutritive quality will likely overwhelm the improved rangeland quality associated with an extended growing season.Grazing maintained or improved rangeland quality by increasing total ANPP by 20 - 40 g . m(-2) . yr(-1) with no effect on palatable ANPP. Grazing effects on forage nutritive quality, as measured by foliar nitrogen and carbon content and by shifts in plant group ANPP, resulted in improved forage quality. Grazing extended the growing season at both habitats, and it advanced the growing season at the meadows. Synergistic interactions between warming and grazing were present, such that grazing mediated the warming- induced declines in vegetation production and nutritive quality. Moreover, combined treatment effects were nonadditive, suggesting that we cannot predict the combined effect of global changes and human activities from single- factor studies.Our findings suggest that the rangelands on the Tibetan Plateau, and the pastoralists who depend on them, may be vulnerable to future climate changes. Grazing can mitigate the negative warming effects on rangeland quality. For example, grazing management may be an important tool to keep warming- induced shrub expansion in check. Moreover, flexible and opportunistic grazing management will be required in a warmer future.

CO2 fluxes of alpine shrubland ecosystem on the north-eastern Tibetan plateau

We measured ecosystem CO2 fluxes for an alpine shrubland on the north-eastern Tibetan Plateau, Qinghai, China. The study is to understand (1) the seasonal variation of CO2 flux and (2) how environmental factors affect the seasonality of CO2 exchange in the alpine ecosystem. Daytime ecosystem respiration was extrapolated from the relationship between temperature and nighttime CO2 fluxes under high turbulent conditions.Seasonal patterns of gross ecosystem production, ecosystem respiration and net ecosystem CO2 exchange followed highly the seasonal change of aboveground biomass in the alpine shrubland. The net ecosystem CO2 exchange was mainly controlled by the variation of photosynthetic photon flux density, while the ecosystem respiration was closely correlated to the soil temperature at 5-cm depth. Integrated values of gross ecosystem production, ecosystem respiration and net ecosystem CO2 exchange for the period from November 1, 2002 to October 31 2003 were estimated to be 1418, 1155 and 222 g CO2 m(-2) yr(-1), respectively.

Diurnal, seasonal and annual variation in net ecosystem CO2 exchange of an alpine shrubland on Qinghai-Tibetan plateau

Thus far, grassland ecosystem research has mainly been focused on low-lying grassland areas, whereas research on high-altitude grassland areas, especially on the carbon budget of remote areas like the Qinghai-Tibetan plateau is insufficient. To address this issue, flux of CO2 were measured over an alpine shrubland ecosystem (37 degrees 36'N, 101 degrees 18'E; 325 above sea level [a. s. l.]) on the Qinghai-Tibetan Plateau, China, for 2 years (2003 and 2004) with the eddy covariance method. The vegetation is dominated by formation Potentilla fruticosa L. The soil is Mol-Cryic Cambisols. To interpret the biotic and abiotic factors that modulate CO2 flux over the course of a year we decomposed net ecosystem CO2 exchange (NEE) into its constituent components, and ecosystem respiration (R-eco). Results showed that seasonal trends of annual total biomass and NEE followed closely the change in leaf area index. Integrated NEE were -58.5 and -75.5 g C m(-2), respectively, for the 2003 and 2004 years. Carbon uptake was mainly attributed from June, July, August, and September of the growing season. In July, NEE reached seasonal peaks of similar magnitude (4-5 g C m(-2) day(-1)) each of the 2 years. Also, the integrated night-time NEE reached comparable peak values (1.5-2 g C m(-2) day(-1)) in the 2 years of study. Despite the large difference in time between carbon uptake and release (carbon uptake time < release time), the alpine shrubland was carbon sink. This is probably because the ecosystem respiration at our site was confined significantly by low temperature and small biomass and large day/night temperature difference and usually soil moisture was not limiting factor for carbon uptake. In general, R-eco was an exponential function of soil temperature, but with season-dependent values of Q(10). The temperature-dependent respiration model failed immediately after rain events, when large pulses of R-eco were observed. Thus, for this alpine shrubland in Qinghai-Tibetan plateau, the timing of rain events had more impact than the total amount of precipitation on ecosystem R-eco and NEE.

Depression of net ecosystem CO2 exchange in semi-arid Leymus chinensis steppe and alpine shrub

Uptake and release of carbon in grassland ecosystems is very critical to the global carbon balance and carbon storage. In this study, the dynamics of net ecosystem CO2 exchange (FNEE) of two grassland ecosystems were observed continuously using the eddy covariance technique during the growing season of 2003. One is the alpine shrub on the Tibet Plateau, and the other is the sem-arid Leymus chinensis steppe in Inner Mongolia of China. It was found that the FNEE of both ecosystems was significantly depressed under high solar radiation. Comprehensive analysis indicates that the depression of FNEE in the L. chinensis steppe was the results of decreased plant photosynthesis and increased ecosystem respiration (R-eco) under high temperature. Soil water stress in addition to the high atmospheric demand under the strong radiation was the primary factor limiting the stomatal conductance. In contrast, the depression of FNEE in the alpine shrub was closely related to the effects of temperature on both photosynthesis and ecosystem respiration, coupled with the reduction of plant photosynthesis due to partial stomatal closure under high temperature at mid-day. The R,c of the alpine shrub was sensitive to soil temperature during high turbulence (u* > 0.2 m s(-1)) but its FNEE decreased markedly when the temperature was higher than the optimal value of about 12 degrees C. Such low optimal temperature contrasted the optimal value (about 20 degrees C) for the steppe, and was likely due to the acclimation of most alpine plants to the long-term low temperature on the Tibet Plateau. We inferred that water stress was the primary factor causing depression of the FNEE in the semi-arid steppe ecosystem, while relative high temperature under strong solar radiation was the main reason for the decrease of FNEE in the alpine shrub. This study implies that different grassland ecosystems may respond differently to climate change in the future. (c) 2006 Elsevier B.V All rights reserved.

Stability of alpine meadow ecosystem on the Qinghai-Tibetan Plateau

The meadow ecosystem on the Qinghai-Tibetan Plateau is considered to be sensitive to climate change. An understanding of the alpine meadow ecosystem is therefore important for predicting the response of ecosystems to climate change. In this study, we use the coefficients of variation (Cv) and stability (E) obtained from the Haibei Alpine Meadow Ecosystem Research Station to characterize the ecosystem stability. The results suggest that the net primary production of the alpine meadow ecosystem was more stable (Cv = 13.18%) than annual precipitation (Cv = 16.55%) and annual mean air temperature (Cv= 28.82%). The net primary production was insensitive to either the precipitation (E = 0.0782) or air temperature (E = 0.1113). In summary, the alpine meadow ecosystem on the Qinghai-Tibetan Plateau is much stable. Comparison of alpine meadow ecosystem stability with other five natural grassland ecosystems in Israel and southern African indicates that the alpine meadow ecosystem on the Qinghai-Tibetan Plateau is the most stable ecosystem. The alpine meadow ecosystem with relatively simple structure has high stability, which indicates that community stability is not only correlated with biodiversity and community complicity but also with environmental stability. An average oscillation cycles of 3-4 years existed in annual precipitation, annual mean air temperature, net primary production and the population size of consumers at the Haibei natural ecosystem. The high stability of the alpine meadow ecosystem may be resulting also from the adaptation of the ecosystem to the alpine environment.

Seasonal changes in weight and body composition of yak grazing on alpine-meadow grassland in the Qinghai-Tibetan plateau of China

Forty-five male yaks (born April 2001) were studied to determine how seasonal changes on the Qinghai-Tibetan plateau affected BW and body composition. Thirty yaks were weighed monthly from birth to 26 mo of age to determine seasonal changes in BW. The remaining 15 yaks were allocated randomly to five groups (three yaks per group), designated for slaughter at 13, 15, 18, 22, and 25 mo to measure seasonal effects on body chemical composition. All yaks were grazed on the alpine-meadow grassland of the plateau without any supplementation. All BW and body composition data were calculated on an individual basis. Body weight and body composition data were both compared across seven growth periods spanning 2 yr and defined by season. From April (birth) to December 2001 of the first growing season, yak BW increased (P < 0.01); however, during the subsequent cold season (December 2001 to May 2002), BW decreased (P < 0.01). The second growing season ran from May 2002 (13 mo of age) to October 2002 (18 mo of age), and the second live weight-loss season ran from October 2002 until May 2003. The weight loss experienced by yaks during the first weight loss season was 25.64% of the total weight gain in the first growing season. The weight loss experienced by yaks during the second weight-loss season was 29.73% of the total weight gain in the second growing season. Energy retention in the second growing season was 291.07 MJ, 50.8% of which was consumed during the subsequent cold season. Energy accumulation in the summer (from May to July) and fall (from July to October) of the second growing season did not differ (5.01 and 6.30 MJ/kg of EBW gain, respectively; P = 0.63). The energy mobilized during the second winter (from October 2002 to February 2003) was 16.49 MJ/kg of EBW, and in the second spring (from February to May 2003), it was 9.06 MJ/kg of EBW. These data suggest that the decrease in grazing yak BW during the first cold season is much less than during the second cold season, and that the energy content per unit of BW mobilized is greater (P = 0.02) in winter than in spring. Results from this study demonstrate highly efficient compensatory growth in grazing yaks following the first weight loss period during the first cold season. This benefit could be exploited by herders to improve yak production. Yaks may have developed a type of self-protection mechanism to overcome the long cold seasons in the Qinghai-Tibetan plateau.

A stable carbon isotopic approach for understanding the CO2 flux at the Haibei Alpine Meadow Ecosystem - A simple model

Although respiration of organisms and biomass as well as fossil fuel burning industrial production are identified as the major sources, the CO2 flux is still unclear due to the lack of proper measurements. A mass-balance approach that exploits differences in the carbon isotopic signature (delta(13)C) of CO2 Sources and sinks was introduced and may provide a means of reducing uncertainties in the atmospheric budget. delta(13)C measurements of atmospheric CO2 yielded an average of - 10.3 parts per thousand relative to the Peedee Belemnite standard; soil and plants had a narrow range from -25.09 parts per thousand to -26.51 parts per thousand and averaged at -25.80 parts per thousand. Based on the fact of steady fractionation and enrichment during respiration of mitochondria, we obtained the emission Of CO2 of 35.451 mol m(-2) a(-1) and CO2 flux of 0.2149 mu mol m(-2) s(-)1. The positive CO2 flux indicated the Haibei Alpine Meadow Ecosystem a source rather than a sink. The mass-balance model can be applied for other ecosystem even global carbon cycles because it neglects the complicated process of carbon metabolism, however just focuses on stable carbon isotopic compositions in any of compartments of carbon sources and sinks. (C) 2005 Elsevier B.V. All rights reserved.

Modeling trophic positions of the alpine meadow ecosystem combining stable carbon and nitrogen isotope ratios

Stable carbon and nitrogen isotope ratios of single tissues or whole bodies were analyzed to establish trophic positions of main consumers living at the alpine meadow ecosystem in the Tibetan Plateau. The results demonstrated that delta C-13 and delta N-15 values of vertebrates showed great variations and ranged from -26.83 to -22.51 parts per thousand and from 2.33 to 8.44 parts per thousand, respectively. Plateau pika, root vole, plateau hare, infants of rodents and hatchlings of passerine bird species had the lowest delta C-13 and delta N-15 values. delta C-13 and delta N-15 values of omnivorous and insectivorous birds and amphibians showed intermediate. Carnivorous species, steppe polecat and Upland buzzard, and omnivorous Robin accentor and White wagtail possessed extremely higher VC and delta N-15 values. Omnivorous birds captured in earlier year had significantly less negative delta C-13 and greater delta N-15 values than those captured later. Based on steady angular enrichment between trophic levels, an "alpha and vector model" combing delta C-13 and delta N-15 values was introduced to reveal trophic positions, the results indicated that Tibetan sheep, Tibetan yak, plateau pika, root vole, plateau hare, infants of small rodents showed the lowest trophic positions (TP 1.81-2.38). While omnivorous and insectivorous birds, their hatchlings and amphibians showed intermediate trophic positions (TP 2.06-2.89), carnivorous species steppe polecat and Upland buzzard, migrant birds possessed extremely higher trophic positions (TP 2.89-3.05). The isotopic investigation of organisms and the introduced "alpha and vector model" successfully demonstrated the same trophic positions and diet prediction of consumers as nitrogen enrichment model at the alpine meadow ecosystem. Besides of this information, the "alpha and vector model" can also be incorporated into multiple isotope signatures to infer trophic relationships. This angular enrichment model has the potential to address basic ecological questions, such as trophic structure, trophic dynamics, and energy flow in other terrestrial ecosystems of properly handled. (C) 2005 Elsevier B.V. All rights reserved.

Effect of long-term grazing on soil organic carbon content in semiarid steppes in Inner Mongolia

To clarify the response of soil organic carbon (SOC) content to season-long grazing in the semiarid typical steppes of Inner Mongolia, we examined the aboveground biomass and SOC in both grazing (G-site) and no grazing (NG-site) sites in two typical steppes dominated by Leymus chinensis and Stipa grandis, as well as one seriously degraded L. chinensis grassland dominated by Artemisia frigida. The NG-sites had been fenced for 20 years in L. chinensis and S. grandis grasslands and for 10 years in A. frigida grassland. Above-ground biomass at G-sites was 21-35% of that at NG-sites in L. chinensis and S. grandis grasslands. The SOC, however, showed no significant difference between G-site and NG-site in both grasslands. In the NG-sites, aboveground biomass was significantly lower in A. frigida grassland than in the other two grasslands. The SOC in A. frigida grassland was about 70% of that in L. chinensis grassland. In A. frigida grassland, aboveground biomass in the G-site was 68-82% of that in the NG-site, whereas SOC was significantly lower in the G-site than in the NG-site. Grazing elevated the surface soil pH in L. chinensis and A. frigida communities. A spatial heterogeneity in SOC and pH in the topsoil was not detected the G-site within the minimal sampling distance of 10 m. The results suggested that compensatory growth may account for the relative stability of SOC in G-sites in typical steppes. The SOC was sensitive to heavy grazing and difficult to recover after a significant decline caused by overgrazing in semiarid steppes.

Characterizing CO2 fluxes for growing and non-growing seasons in a shrub ecosystem on the Qinghai-Tibet Plateau

To assess carbon budget for shrub ecosystems on the Qinghai-Tibet Plateau, CO2 flux was measured with an open-path eddy covariance system for an alpine shrub ecosystem during growing and non-growing seasons. CO2 flux dynamics was distinct between the two seasons. During the growing season from May to September, the ecosystem exhibited net CO2 uptake from 08:00 to 19:00 (Beijing Standard Time), but net CO2 emission from 19:00 to 08:00. Maximum CO2 uptake appeared around 12:00 with values of 0.71, 1,19, 1.46 and 0.67 g CO2 m(-2) h(-1) for June, July, August and September, respectively. Diurnal fluctuation Of CO2 flux showed higher correlation with photosynthetic photon flux density than temperature. The maximum net CO2 influx occurred in August with a value of 247 g CO2 m(-2). The total CO2 uptake by the ecosystem was up to 583 g CO2 m(-2) for the growing season. During the non-growing season from January to April and from October to December, CO2 flux showed small fluctuation with the largest net CO2 efflux of 0.30 g CO2 m(-2) h(-1) in April. The diurnal CO2 flux was close to zero during most time of the day, but showed a small net CO2 eff lux from 11:00 to 18:00. Diurnal CO2 flux, is significantly correlated to diurnal temperature in the non-growing season. The maximum monthly net CO2 eff lux appeared in April, with a value of 105 g CO2 m(-2). The total net CO2 eff lux for the whole non-growing season was 356 g CO2 m(-2).

Energy exchange between the atmosphere and a meadow ecosystem on the Qinghai-Tibetan Plateau

To reveal the potential contribution of grassland ecosystems to climate change, we examined the energy exchange over an alpine Kobresia meadow on the northeastern Qinghai-Tibetan Plateau. The annual pattern of energy exchange showed a clear distinction between periods of frozen soil with the daily mean soil temperature at 5 cm (T-s5 &LE; 0 &DEG; C) and non-frozen soil (T-s5 > 0 &DEG; C). More than 80% of net radiation was converted to sensible heat (H) during the frozen soil period, but H varied considerably with the change in vegetation during the non-frozen soil period. Three different sub-periods were further distinguished for the later period: (1) the pre-growth period with Bowen ratio (β) > 1 was characterized by a high β of 3.0 in average and the rapid increase of net radiation associated with the increases of H, latent heat (LE) and soil heat; (2) during the Growth period when β &LE; 1, the LE was high but H fluxes was low with β changing between 0.3 and 0.4; (3) the post-growth period with average β of 3.6 when H increased again and reached a second maximum around early October. The seasonal pattern suggests that the phenology of the vegetation and the soil water content were the major factors affecting the energy partitioning in the alpine meadow ecosystem. © 2005 Elsevier B.V. All rights reserved.

Experimental warming causes large and rapid species loss, dampened by simulated grazing, on the Tibetan Plateau

We investigated the independent and combined effects of experimental warming and grazing on plant species diversity on the north-eastern Tibetan Plateau, a region highly vulnerable to ongoing climate and land use changes. Experimental warming caused a 26-36% decrease in species richness, a response that was generally dampened by experimental grazing. Higher species losses occurred at the drier sites where N was less available. Moreover, we observed an indirect effect of climate change on species richness as mediated by plant-plant interactions. Heat stress and warming-induced litter accumulation are potential explanations for the species' responses to experimental warming. This is the first reported experimental evidence that climate warming could cause dramatic declines in plant species diversity in high elevation ecosystems over short time frames and supports model predictions of species losses with anthropogenic climate change.

Seasonal patterns of gross primary production and ecosystem respiration in an alpine meadow ecosystem on the Qinghai-Tibetan Plateau

We measured the net ecosystem CO2 exchange (NEE) in an alpine meadow ecosystem (latitude 37degrees29'-45'N, longitude 101degrees12'-23'E, 3250 m above sea level) on the Qinghai-Tibetan Plateau throughout 2002 by the eddy covariance method to examine the carbon dynamics and budget on this unique plateau. Diurnal changes in gross primary production (GPP) and ecosystem respiration (R-e) showed that an afternoon increase of NEE was highly associated with an increase of R-e. Seasonal changes in GPP corresponded well to changes in the leaf area index and daily photosynthetic photon flux density. The ratio of GPP/R-e was high and reached about 2.0 during the peak growing season, which indicates that mainly autotrophic respiration controlled the carbon dynamics of the ecosystem. Seasonal changes in mean GPP and R-e showed compensatory behavior as reported for temperate and Mediterranean ecosystems, but those of GPP(max) and R-emax were poorly synchronized. The alpine ecosystem exhibited lower GPP (575 g C m(-2) y(-1)) than, but net ecosystem production (78.5 g C m(-2) y(-1)) similar to, that of subalpine forest ecosystems. The results suggest that the alpine meadow behaved as a CO2 sink during the 1-year measurement period but apparently sequestered a rather small amount of C in comparison with similar alpine ecosystems.

Influence of grazing intensity on performance of perennial grass mixtures in the alpine region of the Tibetan Plateau

Effects of grazing intensity on leaf photosynthetic rate (Pn), specific leaf area (SLA), individual tiller density, sward leaf area index (LAI), harvested herbage DM, and species composition in grass mixtures (Clinelymus nutans + Bromus inermis, Elymus nutans + Bromus inermis + Agropyron cristatum and Elymus nutans + Clinelymus nutans + Bromus inermis + Agropyron cristatum) were studied in the alpine region of the Tibetan Plateau. Four grazing intensities (GI), expressed as feed utilisation rates (UR) by Tibetan lambs were imposed as follows: (1) no grazing; (2) 30% UR as light grazing; (3) 50% UR as medium grazing; and (4) 70% UR as high grazing. Leaf Pn rate and tiller density of grasses increased (P < 0.05), while sward LAI and harvested herbage DM declined (P < 0.05) with the increments of GI, although no effect of GI on SLA was observed. With increasing GI, Elymus nutans and Clinelymus nutans increased but Bromus inermis and Agropyron cristatum decreased in swards, LAI and DM contribution. Whether being grazed or not, Elymus nutans + Clinelymus nutans + Bromus inermis + Agropyron cristatum was the most productive sward among the grass mixtures. Thus, two well-performed grass species (Elymus nutans and Clinelymus nutans) and the most productive mixture of four species should be investigated further as the new feed resources in the alpine grazing system of the Tibetan Plateau. Light grazing intensity of 30% UR was recommended for these grass mixtures when swards, LAI, herbage DM harvested, and species compatibility were taken into account.