Changes in plant biomass and species composition of alpine Kobresia meadows along altitudinal gradient on the Qinghai-Tibetan Plateau

Alpine Kobresia meadows are major vegetation types on the Qinghai-Tibetan Plateau. There is growing concern over their relationships among biodiversity, productivity and environments. Despite the importance of species composition, species richness, the type of different growth forms, and plant biomass structure for Kobresia meadow ecosystems, few studies have been focused on the relationship between biomass and environmental gradient in the Kobresia meadow plant communities, particularly in relation to soil moisture and edaphic gradients. We measured the plant species composition, herbaceous litter, aboveground and belowground biomass in three Kobresia meadow plant communities in Haibei Alpine Meadow Ecosystem Research Station from 2001 to 2004. Community differences in plant species composition were reflected in biomass distribution. The total biomass showed a decrease from 13196.96 +/- 719.69 g/m(2) in the sedge-dominated K. tibetica swamp to 2869.58 +/- 147.52 g/m(2) in the forb and sedge dominated K. pygmaea meadow, and to 2153.08 +/- 141.95 g/m(2) in the forbs and grasses dominated K. humilis along with the increase of altitude. The vertical distribution of belowground biomass is distinct in the three meadow communities, and the belowground biomass at the depth of 0-10 cm in K. tibetica swamp meadow was significantly higher than that in K. humilis and K. pygmaea meadows (P < 0.01). The herbaceous litter in K. tibetica swamp was significantly higher than those in K. pygnaeca and K. humilis meadows. The effects of plant litter are enhanced when ground water and soil moisture levels are raised. The relative importance of litter and vegetation may vary with soil water availability. In the K. tibetica swamp, total biomass was negatively correlated to species richness (P < 0.05); aboveground biomass was positively correlated to soil organic matter, soil moisture, and plant cover (P < 0.05); belowground biomass was positively correlated with soil moisture (P < 0.05). However, in the K. pygnaeca and K. humilis meadow communities, aboveground biomass was positively correlated to soil organic matter and soil total nitrogen (P < 0.05). This suggests that the distribution of biomass coincided with soil moisture and edaphic gradient in alpine meadows.

Effect of imitated global warming on Delta C-13 values in seven plant species growing in Tibet alpine meadows

Open-top chambers were used to estimate the possible effects of global warming on delta C-13 of seven plant species grown in alpine meadow ecosystem. The delta C-13 values of plant species were lower after long-term growth in open-top chambers. In the course of experiment, temperature significantly increased inside the chambers by 4 degrees C. Plant species grown at a lower elevation above sea level had higher delta C-13 values as compared to those grown at a higher elevation. This was in accordance with the effect of open-top chamber on delta C-13 values in plants. Greater availability of CO2 and lower water vapor as indicated by an increase in discrimination against (CO2)-C-13, probably result in more negative delta C-13 values of plants because higher stomatal conductance increases availability of CO2 and causes greater discrimination against (CO2)-C-13. The plant species studied could be the indicator species for testing global warming by the change in carbon isotope ratios at the two growth temperatures.

Population dynamics and responses to management of plateau pikas Ochotona curzoniae

1. Plateau pikas Ochotona curzoniae are considered a pest species on the Tibetan Plateau because they compete with livestock for forage and their burrowing could contribute to soil erosion. The effectiveness of pest control programmes in Tibet has not been measured, and it is not known whether changes in livestock management have exacerbated problems with plateau pikas or compromised their control. This study measured the impact of control programmes and livestock management for forage conservation on populations of plateau pikas in alpine meadow in Naqu District, central Tibet, during 2004 and 2005.2. Current techniques for controlling plateau pikas in spring cause large reductions in abundance, but high density-dependent rates of increase result in no differences between treated and untreated populations by the following autumn. Rates of increase from spring to autumn are not influenced by standing plant biomass or concurrent grazing by yaks Bos grunniens and Tibetan sheep Ovis aries.3. In autumn there was significantly lower biomass outside fenced areas with year-round livestock grazing compared with inside fenced areas with equivalent or higher numbers of plateau pikas but predominantly winter grazing by livestock. Inside fenced areas, control of plateau pikas in spring produced no detectable effect on standing plant biomass at the end of the following summer compared with uncontrolled populations of plateau pikas.4. Regardless of their initial density, populations of plateau pikas declined rapidly over winter outside fenced areas where there was very low standing plant biomass in autumn. However, inside fenced areas with higher plant biomass in autumn, low-density populations of plateau pikas declined more slowly than high-density populations.5. Synthesis and applications. Current control programmes have limited effect because populations of plateau pikas can recover in one breeding season. There was no apparent increase in forage production in areas where plateau pikas were controlled. However, plateau pikas appear to benefit from changes in grazing management, with low-density populations declining less over winter inside fenced areas than elsewhere. It was not evident that control programmes are warranted or that they will improve the livelihoods of Tibetan herders.

Modeling gross primary production of alpine ecosystems in the Tibetan Plateau using MODIS images and climate data

The eddy covariance technique provides measurements of net ecosystem exchange (NEE) Of CO2 between the atmosphere and terrestrial ecosystems, which is widely used to estimate ecosystem respiration and gross primary production (GPP) at a number Of CO2 eddy flux tower sites. In this paper, canopy-level maximum light use efficiency, a key parameter in the satellite-based Vegetation Photosynthesis Model (VPM), was estimated by using the observed CO2 flux data and photosynthetically active radiation (PAR) data from eddy flux tower sites in an alpine swamp ecosystem, an alpine shrub ecosystem and an alpine meadow ecosystem in Qinghai-Tibetan Plateau, China. The VPM model uses two improved vegetation indices (Enhanced Vegetation Index (EVI), Land Surface Water Index (LSWI)) derived from the Moderate Resolution Imaging Spectral radiometer (MODIS) data and climate data at the flux tower sites, and estimated the seasonal dynamics of GPP of the three alpine grassland ecosystems in Qinghai-Tibetan Plateau. The seasonal dynamics of GPP predicted by the VPM model agreed well with estimated GPP from eddy flux towers. These results demonstrated the potential of the satellite-driven VPM model for scaling-up GPP of alpine grassland ecosystems, a key component for the study of the carbon cycle at regional and global scales. (c) 2006 Elsevier Inc. All rights reserved.

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.

Enrichment of stable carbon and nitrogen isotopes of plant populations and plateau pikas along altitudes

Stable carbon and nitrogen isotope ratios were measured in plant populations and plateau pikas to determine enrichment in stable isotopes of three alpine meadow ecosystems at different elevations in the Qinghai-Tibet Plateau. The results indicated that stable carbon isotope signatures of plant populations at the three locations showed significant variations, delta C-13 of plant populations showed an enrichment of 0.86 parts per thousand per 1000 in over the linear proportion of the altitudinal response, while stable nitrogen isotopes showed no apparent difference. The stable nitrogen isotopes of plateau pikas became significantly isotopically heavier along altitudinal gradients and showed an enrichment of 3.17 parts per thousand/km. Stable carbon isotopes showed no significance, however, and the enrichment of 0.448 parts per thousand/km. delta C-13 and delta N-15 in plateau pikas were not significantly correlated. There appeared to be segregation between the metabolism of stable carbon and nitrogen isotopes of plateau pikas. Variances in metabolic rate at lower water availability and temperatures are presumed to be the main cause of enrichment of stable nitrogen isotopes along altitudinal gradients. Attention should be paid to construct trophic positions and to trace food chain information based oil an isotopic enrichment model.

Distribution and species diversity of plant communities along transect on the Northeastern Tibetan plateau

The distribution and species diversity of plant communities along a 600 km transect through the northeastern Tibetan Plateau (32 degrees 42'-35 degrees 07' N, 101 degrees 02'-97 degrees 38' E) with altitudes from 3255 to 4460 m are described. The transect started from the Youyi Bridge of Banma through Dari, Maqin and Maduo to Zaling Lake. The data from 47 plots along the transect are summarized and analyzed. The mean annual temperature, the mean annual rainfall and the length of growing season decreases from 2.6 to -4.5 degrees C, from 767.2 to 240.1 mm, from 210 to 140 days, respectively, along the transect from the southeastern Banma to northwestern Zaling Lake. The number of vascular plant species recorded in 47 plots is 242 including 2 tree, 34 shrub, 206 herb species. Main vegetation types on the transect from southeast to northwest are: Sabina convallium forest, Picea likiangensis forest, Pyracantha fortuneana + Spiraea alpina shrub, Hippophae neurocarpu shrub, Sibiraea angustata + Polygonum viviparum shrub, Stellera chamaejasme herb meadow, Potentilla fruticosa + Salix obscura + Carex sp. Shrub, Kobresia capillifolia meadow, P. froticosa + Kobresia humilis shrub, Caragana jubata + S. obscura shrub, Kobresia tibetica meadow, Kobresia pygmaea meadow, K. pygmaea + Stipa purpurea steppe meadow, Stipa purpurea steppe. Plant richness and diversity index all showed a decreasing trend with increasing of elevation along transect from southeast to northwest. Detailed information on altitudinal ranges and distribution of the alpine vegetation, vascular flora and environments over the alpine zone at northeastern Tibetan Plateau provides baseline records relevant to future assessment of probable effects of global climate changes.

Characterization of photosynthetic pathway of plant species growing in the eastern Tibetan plateau using stable carbon isotope composition

The photosynthetic pathway of plant species collected at Menyuan, Henan, and Maduo sites, east of Tibetan Plateau, China, during the growing season were studied using stable carbon isotopes in leaves. The 232 samples leaves analyzed belonged to 161 species, 30 families, and 94 genera. The delta(13)C values (from -24.6 to -29.2 %o) indicated that all the considered species had a photosynthetic C-3 pathway. The absence of plant species with C-4 photosynthetic pathway might be due to the extremely low air temperature characterizing the Tibetan Plateau. The average delta(13)C value was significantly (p < 0.05) different between annuals and perennials at the three considered study sites. Hence the longer-lived species had greater water-use efficiency (WUE) than shorter-lived species, that is, longer-lived species are better adapted to the extreme environmental conditions of the Tibetan Plateau.

The effect of land management on carbon and nitrogen status in plants and soils of alpine meadows on the Tibetan plateau

Large-scale grassland rehabilitation has been carried out on the severely degraded lands of the Tibetan plateau. The grasslands created provide a useful model for evaluating the recovery of ecosystem properties. The purposes of this research were: (1) to examine the relative influence of various rehabilitation practices on carbon and nitrogen in plants and soils in early secondary succession; and (2) to evaluate the degree to which severely degraded grassland altered plant and soil properties relative to the non-disturbed native community. The results showed: (1) The aboveground tissue C and N content in the control were 105-97 g m(-2) and 3.356gm(-2), respectively. The aboveground tissue C content in the mixed seed treatment, the single seed treatment, the natural recovery treatment and the severely degraded treatment was 137 per cent, 98 per cent, 49 per cent and 38 per cent, respectively, of that in the control. The corresponding aboveground tissue N content was 109 per cent, 84 per cent, 60 per cent and 47 per cent, respectively, of that in the control. (2) Root C and N content in 0-20 cm depths of the control had an 2 2 average 1606 gm(-2) and 30-36 gm(-2) respectively. Root C and N content in the rehabilitation treatments were in the range of 26-36 per cent and 35-53 per cent, while those in the severely degraded treatment were only 17 per cent and 26 per cent of that in the control. (3) In the control the average soil C and N content at 0-20 cm was 11307 gm(-2) and 846 gm(-2), respectively. Soil C content in the uppermost 20 cm in the seeded treatments, the natural recovery treatment and the severely degraded treatment was 67 per cent, 73 per cent and 57 per cent, respectively, while soil N content in the uppermost 20cm was 72 per cent, 82 per cent and 79 per cent, respectively, of that in the control. The severely degraded land was a major C source. Restoring the severely degraded lands to perennial vegetation was an alternative approach to sequestering C in former degraded systems. N was a limiting factor in seeding grassland. It is necessary for sustainable utilization of seeding grassland to supply extra N fertilizer to the soil or to add legume species into the seed mix. Copyright (c) 2005 John Wiley & Sons, Ltd.

Phylogeography of the Qinghai-Tibetan Plateau endemic Juniperus przewalskii (Cupressaceae) inferred from chloroplast DNA sequence variation

The vegetation of the northeast Qinghai-Tibetan Plateau is dominated by alpine meadow and desert-steppe with sparse forests scattered within it. To obtain a better understanding of the phylogeography of one constituent species of the forests in this region, we examined chloroplast trnT-trnF and trnS-trnG sequence variation within Juniperus przewalskii, a key endemic tree species. Sequence data were obtained from 392 trees in 20 populations covering the entire distribution range of the species. Six cpDNA haplotypes were identified. Significant population subdivision was detected (G(ST) = 0.772, N-ST = 0.834), suggesting low levels of recurrent gene flow among populations and significant phylogeographic structure (N-ST > G(ST), P < 0.05). Eight of the nine disjunct populations surveyed on the high-elevation northeast plateau were fixed for a single haplotype (A), while the remaining, more westerly population, contained the same haplotype at high frequency together with two low frequency haplotypes (C and F). In contrast, most populations that occurred at lower altitudes at the plateau edge were fixed or nearly fixed for one of two haplotypes, A or E. However, two plateau edge populations had haplotype compositions different from the rest. In one, four haplotypes (A, B, D and E) were present at approximately equivalent frequencies, which might reflect a larger refugium in the area of this population during the last glacial period. Phylogenetic analysis indicated that the most widely distributed haplotype A is not ancestral to other haplotypes. The contrasting phylogeographic structures of the haplotype-rich plateau edge area and the almost haplotype-uniform plateau platform region indicate that the plateau platform was recolonized by J. przewalskii during the most recent postglacial period. This is supported by the findings of a nested clade analysis, which inferred that postglacial range expansion from the plateau edge followed by recent fragmentation is largely responsible for the present-day spatial distribution of cpDNA haplotypes within the species.

Nitrogen deposition and carbon sequestration in alpine meadows

Nitrogen deposition experiments were carried out in alpine meadow ecosystems in Qinghai-Xizang Plateau in China, in order to explore the contribution of nitrogen deposition to carbon sequestration in alpine meadows. Two methods were used in this respect. First, we used the allocation of N-15 tracer to soil and plant pools. Second, we used increased root biomass observed in the nitrogen-amended plots. Calculating enhanced carbon storage, we considered the net soil CO2 emissions exposed to nitrogen deposition in alpine meadows. Our results show that nitrogen deposition can enhance the net soil CO2 emissions, and thus offset part of carbon uptake by vegetation and soils. It means that we have to be cautious to draw a conclusion when we estimate the contribution of nitrogen deposition to carbon sequestration based on the partitioning of N-15 tracer in terrestrial ecosystems, in particular in N-limited ecosystems. Even if we assess the contribution of nitrogen deposition to carbon sequestration based on increased biomass exposed to nitrogen deposition in terrestrial ecosystems, likewise, we have to consider the effects of nitrogen deposition on the soil CO2 emissions.

Analysis of diets of upland buzzards using stable carbon and nitrogen isotopes

We measured the stable carbon and nitrogen isotope ratios for muscles of the upland buzzards (Buteo hemilasius) and their potential food sources, plateau pikas (Ochotona curzoniae), Qinghai voles (Lasiopodomys fuscus), plateau zokors (Myospalax fontanierii), and several passerine bird species at the alpine meadow in Maduo county, Guoluo prefecture of Qinghai province, People's Republic of China, to provide diet information of upland buzzards, highlighting different diet composition of upland buzzards exposed to different locations. The results demonstrated that stable carbon isotope ratios of upland buzzards, passerine birds, plateau pikas, plateau zokors, and Qinghai voles were -24.42 +/- 0.25parts per thousand, -22.89 +/- 1.48parts per thousand, -25.30 +/- 1.47parts per thousand, -25.78 +/- 0.22parts per thousand, and -25.41 +/- 0.01parts per thousand, respectively, and stable nitrogen isotope ratios were 7.89 +/- 0.38parts per thousand, 8.37 +/- 2.05parts per thousand, 5.83 +/- 1.10parts per thousand, 5.23 +/- 0.34parts per thousand, and 8.86 +/- 0.06parts per thousand, respectively. Fractionation of stable carbon and nitrogen isotope ratios between upland buzzards and their food were 1.03parts per thousand and 2.11parts per thousand, respectively. Based on mass balance principle of stable isotopes and the Euclidean distance mixing model, upland buzzards depended mainly on plateau pikas as food (74.56%). Plateau zokors, Qinghai voles, and passerine birds only contributed a small proportion (25.44%) to diets of upland buzzards. The results were closely accordant with analyses of stomach contents and food pellets, which firmly supported the feasibility of using stable carbon and nitrogen isotope ratios to investigate diet information of upland buzzards. Another study based on stable carbon isotopes showed that upland buzzards living in the Haibei prefecture (another prefecture located in the southeast Qinghai province) mainly preyed on passerine birds (64.96% or more) as food supply. We were alarmed by the preliminary results that widespread poisoning activities of small mammals could reshape the food composition of upland buzzards, influencing the stability and sustainability of the alpine meadow. Bio-control on rodent pests should be carried out rather than the chemical measures.

Photosynthesis of Saussurea superba and Gentiana straminea is not reduced after long-term enhancement of UV-B radiation

Experiments were conducted in an alpine Kobresia humilis meadow near Haibei Alpine Meadow Ecosystem Research Station (37degrees29'-37degrees45'N, 101degrees12'-101degrees33'E; altitude 3200 m). Effects of enhanced ultraviolet-B (UV-B) radiation on photosynthesis of the alpine plants of Saussurea superba and Gentiana straminea were investigated. Both species were exposed to a UV-B-BE density at 15.80 kJ m(-2) per day, simulating nearly 14% ozone (O-3) reduction during the plant growing season. Neither photosynthetic CO2 uptake rate nor photosynthetic O-2 evolution rate were decreased after a long period of enhanced UV-B radiation treatment. On the contrary, there was a tendency to increase of both parameters in both species. The photosynthetic pigments were also increased, when expressed on a leaf area basis. UV-B absorbing compounds, detected by the absorbance values at 300 mm, had a tendency to increase in both species after enhanced UV-B radiation. After long-term exposure of plants to enhanced UV-B radiation, leaf morphology was also affected. Leaf thickness in both S. superba and G. straminea were increased significantly (P < 0.001). This supports our hypothesis that the increase of leaf thickness in both species after long-term exposure of enhanced UV-B radiation could compensate for the photodestruction of photosynthetic pigments when light passes through the leaf. Therefore, photosynthesis is not reduced in either species when expressed on leaf area basis. (C) 2003 Elsevier B.V. All rights reserved.

Social and ecological influences on dispersal and philopatry in the plateau pika (Ochotona curzoniae)

Benefits and costs of dispersal and philopatry of the social plateau pika (Ochotona curzoniae) were studied on the Tibetan plateau for 3 years. Although short-lived, plateau pikas live in cohesive family groups that occupy burrow systems in sedge meadow habitat Most (57.8%) plateau pikas were philopatric, and dispersal movements were extremely restricted. No juvenile females or adult pikas moved more than two family ranges between years; the greatest observed dispersal distances were by two juvenile males that moved five family ranges from the family of their birth. Traversing unfamiliar habitat was not a cost of pika dispersal because most dispersers settled in families that they could easily visit before dispersal. Dispersal movements appeared to result in equalization of density among pika families, an expected result if competition for environmental resources influenced dispersal. Males did not disperse to gain advantages in competition for mates, as evidenced by their moving to families with significantly fewer females. Females, however, moved to families with significantly more males. Males provide abundant paternal care, and significantly more offspring per female survived to become adults from families with more adult males per adult female. Evidence concerning the influence of inbreeding avoidance on natal dispersal was indirect. Some males exhibited natal philopatry; thus some families had opportunity for close inbreeding. Males and females that dispersed had no opposite-sex relatives in their new families. Philopatric pikas may have benefited by remaining in families that exhibited low local densities, and philopatric females might have benefited from social cooperation with relatives.

碳、氧稳定同位素分析在古环境研究中的应用

A number of proxy records of paleoenvironment using stable isotopes could show the history of past environmental changes. These archives include peat and lake sediments, loess-paleosot sequence, fossil mammals and stalagmite, and so on. The stable isotopic composition of carbonate and organic matter and frequency magnetic susceptibility from Tianshuigou and Yuanlei loess-palesol sequence can be used to give estimates of the paleoenvironmental history of Dali, and even of the whole Chinese Loess Plateau during the last 250ka. Features of the High Temperature and Large Precipitation Event in the Tibet Plateau and its adjacent area during 40~30kaBP had been studied by Professor Shi Y. In this dissertation, its impact on Chinese Loess Plateau has been discussed again. Carbon and oxygen isotopic ratios, magnetic susceptibility and frequency magnetic susceptibility in Tianshuigou and Yuanlei profiles show that the Event in this area is not so stronger as the Tibet Plateau. The carbon isotopic composition of organic matter in Tianshuigou, Yuanlei, dingcun and Jingcun loess-palesol sequences are indicative of major changes in the paleovagetation between terrace and plain of the Chinese Loess Plateau. Water is one of the most important factors adjusting the relative biomass of C4 plant in terrestrial ecosystems. Stable carbon isotope ratio of vertebrate tooth enamel is used increasingly to reconstruct environmental and ecological information modern and ancient ecosystems. The SI3C value of tooth enamel bioapatites can distinguish between browsers and grazers. Data from typical grassland of Inter Mongolia, the Alpine meadow of Qinghai-Tibet Plateau and the Yaluzangbu Great Canyon indicate that diets of mammals could record the relative biomass of C4 plant only in the C4 dominated ecosystem. In a C3 dominated ecosystem, diet of mammals would include more C3 plants than vegetation. According to Professor Cerling, proxy records from North and South America, Africa and Pakistan show that at the end of the Miocene (between 8Ma to 6 Ma) there was a global expansion of CA biomass, probably when atmospheric CO2 levels declined. Thus, "C4 world" and "CO2 starvation" are put forward. In this dissertation, carbon isotopes of fossil tooth such as Equus sanmeniensis and Hipparion chiai from Linxia, China reveal that there is a C3 dominated ecosystem in the late Miocene. Diets of ancient mammals in Linxia are not evidence of global expansion of C4 biomass.