青藏高原微生物多样性研究

正青藏高原被誉为世界屋脊,其内部除平原外还有许多山峰、冰川、高山湖泊和高山沼泽,是生态环境最为奇特、生物资源最为丰富的自然资源宝库之一。同时,青藏高原的微生物群落结构及其多样性与其他区域存在巨大差异,因而具有极高的科学研究价值,并逐渐被人们所关注。研究发现气候变化对青藏高原高寒草地生态系统草丛-地境界面微生物会产生重要的影响[1]。冰川雪藻的研究主要在南部的Yala冰川开展,     

Precipitation impacts on vegetation spring phenology on the Tibetan Plateau

The ongoing changes in vegetation spring phenology in temperate/cold regions are widely attributed to temperature. However, in arid/semiarid ecosystems, the correlation between spring temperature and phenology is much less clear. We test the hypothesis that precipitation plays an important role in the temperature dependency of phenology in arid/semiarid regions. We therefore investigated the influence of preseason precipitation on satellite‐derived estimates of starting date of vegetation growing season (SOS) across the Tibetan Plateau (TP). We observed two clear patterns linking precipitation to SOS. First, SOS is more sensitive to interannual variations in preseason precipitation in more arid than in wetter areas. Spatially, an increase in long‐term averaged preseason precipitation of 10 mm corresponds to a decrease in the precipitation sensitivity of SOS by about 0.01 day mm^?1. Second, SOS is more sensitive to variations in preseason temperature in wetter than in dryer areas of the plateau. A spatial increase in precipitation of 10 mm corresponds to an increase in temperature sensitivity of SOS of 0.25 day °C^?1 (0.25 day SOS advance per 1 °C temperature increase). Those two patterns indicate both direct and indirect impacts of precipitation on SOS on TP. This study suggests a balance between maximizing benefit from the limiting climatic resource and minimizing the risk imposed by other factors. In wetter areas, the lower risk of drought allows greater temperature sensitivity of SOS to maximize the thermal benefit, which is further supported by the weaker interannual partial correlation between growing degree days and preseason precipitation. In more arid areas, maximizing the benefit of water requires greater sensitivity of SOS to precipitation, with reduced sensitivity to temperature. This study highlights the impacts of precipitation on SOS in a large cold and arid/semiarid region and suggests that influences of water should be included in SOS module of terrestrial ecosystem models for drylands.     

Phylogeography of Phrynocephalus erythrurus from the Qiangtang Plateau of the Tibetan Plateau

Phrynocephalus erythrurus of the Qiangtang Plateau occupies the highest regions of any reptile on earth. Here, we report mitochondrial DNA haplotypes sampled throughout the distribution of P. erythrurus and analyze patterns of genetic divergence among populations. The species diverged into two major lineages/subspecies at 3.67 mya corresponding to the Northern and Southern Qiangtang Plateau. The Northern Qiangtang lineage diverged into two subpopulations at 2.76 mya separated by the Beilu River Region and Wulanwula Mountains. Haplotypes from the southern Qiangtang lineage diverged 0.98 mya as a star-shaped pattern. Analyses of molecular variance indicated that most of the observed genetic variation occurred among populations/regions implying long-term interruptions to gene flow. There was no evidence of sudden recent range expansions within any of the clades/lineages. NCPA infers allopatric fragmentation and restricted gene flow as the most likely mechanisms of population differentiation. Our results also indicate the presence of at least three refugia since the Hongya glaciation. Mountain movement and glaciations since mid-Pliocene are considered to have shaped phylogenetic patterns of P. erythrurus. P. erythrurus parva is suggested as a valid subspecies of P. erythrurus. Using four calibration points, we estimate an evolutionary rate of 0.762% divergence per lineage per million years for a mitochondrial genomic segment comprising the genes encoding ND2, tRNA^Trp and tRNA^Ala.     

Modern pollen samples from alpine vegetation on the Tibetan Plateau

• 1 A set of 316 modern surface pollen samples, sampling all the alpine vegetation types that occur on the Tibetan Plateau, has been compiled and analysed. Between 82 and 92% of the pollen present in these samples is derived from only 28 major taxa. These 28 taxa include examples of both tree (AP) and herb (NAP) pollen types.
• 2 Most of the modern surface pollen samples accurately reflect the composition of the modern vegetation in the sampling region. However, airborne dust‐trap pollen samples do not provide a reliable assessment of the modern vegetation. Dust‐trap samples contain much higher percentages of tree pollen than non‐dust‐trap samples, and many of the taxa present are exotic. In the extremely windy environments of the Tibetan Plateau, contamination of dust‐trap samples by long‐distance transport of exotic pollen is a serious problem.
• 3 The most characteristic vegetation types present on the Tibetan Plateau are alpine meadows, steppe and desert. Non‐arboreal pollen (NAP) therefore dominates the pollen samples in most regions. Percentages of arboreal pollen (AP) are high in samples from the southern and eastern Tibetan Plateau, where alpine forests are an important component of the vegetation. The relative importance of forest and non‐forest vegetation across the Plateau clearly follows climatic gradients: forests occur on the southern and eastern margins of the Plateau, supported by the penetration of moisture‐bearing airmasses associated with the Indian and Pacific summer monsoons; open, treeless vegetation is dominant in the interior and northern margins of the Plateau, far from these moisture sources.
• 4 The different types of non‐forest vegetation are characterized by different modern pollen assemblages. Thus, alpine deserts are characterized by high percentages of Chenopodiaceae and Artemisia, with Ephedra and Nitraria. Alpine meadows are characterized by high percentages of Cyperaceae and Artemisia, with Ranunculaceae and Polygonaceae. Alpine steppe is characterized by high abundances of Artemisia, with Compositae, Cruciferae and Chenopodiaceae. Although Artemisia is a common component of all non‐forest vegetation types on the Tibetan Plateau, the presence of other taxa makes it possible to discriminate between the different vegetation types.
• 5 The good agreement between modern vegetation and modern surface pollen samples across the Tibetan Plateau provides a measure of the reliability of using pollen data to reconstruct past vegetation patterns in non‐forested areas.

     

Altitudinal patterns of maximum plant height on the Tibetan Plateau

Aims Several studies have shown that plant height changes along environmental gradients. However, altitudinal patterns of plant height across species are still unclear, especially in regions sensitive to climate change. As canopy height decreases dramatically near the tree line in alpine areas, we hypothesize that plant height across all species also decreases with increasing altitude, and distinct thresholds exist along this gradient.Methods Using a large dataset of maximum plant height and elevation range (400 to 6000 m a.s.l.) of 4295 angiosperms from the regional flora of the Tibetan Plateau, we regressed plant height for every 100 m belt against elevation to explore the relationships. To identify the approximate boundaries where dramatic changes in plant height occurs for herbaceous plants, shrubs, trees, woody plants and all angiosperms, we used piecewise linear regression. Phylogenetically independent contrast was used to test the potential evolutionary influences on altitudinal patterns at the family level.Important findings Results showed that for herbaceous plants, shrubs, trees, woody plants and all angiosperms, plant height decreases significantly as altitude increases. In addition, we found that altitude, a proxy for many environmental factors, had obvious thresholds (breakpoints) dictating patterns of plant height. The results of phylogenetically independent contrast also emphasized the importance of evolutionary history in determining the altitudinal patterns of plant height for some growth forms. Our results highlight the relative intense filtering effect of environmental factors in shaping patterns of functional traits and how this could vary for different ranges of environmental variables.     

Radiometric dating for recent lake sediments on the Tibetan Plateau

We present radiometric data from nine lakes across the Tibetan Plateau, and compare their reliability in relation to recent research. Unsupported ²¹⁰Pb profiles show, except for one particular lake, non-exponential decline of ²¹⁰Pb activity with sediment depth. Stratigraphic dates based on global atmospheric nuclear weapons maximum fallout of ¹³⁷Cs (1963) support the use of the constant rate of ²¹⁰Pb supply (CRS) model in four of the dated cores. The discrepancy in the others is likely due to recent increased input of catchment-derived ²¹⁰Pb. ²¹⁰Pb dates in this study suggest that post depositional diffusion of ¹³⁷Cs activity has been significant. The practice of assigning early 1950s dates (start of global atmospheric thermonuclear testing) to lake sediment sequences on the Tibetan Plateau should be used with caution. ¹³⁷Cs profiles from Tibetan lake sediment cores and their geographical distribution suggest that ¹³⁷Cs derived from the 1986 Chernobyl accident or atmospheric testing in China was not sufficient to form a significant peak effective for dating.     

Carbon Sequestration in Two Alpine Soils on the Tibetan Plateau

Soil carbon sequestration was estimated in a conifer forest and an alpine meadow on the Tibetan Plateau using a carbon- 14 radioactive label provided by thermonuclear weapon tests （known as bomb-^14C）. Soil organic matter was physically separated into light and heavy fractions. The concentration spike of bomb-^14C occurred at a soil depth of 4 cm in both the forest soil and the alpine meadow soil. Based on the depth of the bomb-^14C spike, the carbon sequestration rate was determined to be 38.5 g C/m^2 per year for the forest soil and 27.1 g C/m^2 per year for the alpine meadow soil. Considering that more than 60% of soil organic carbon （SOC） is stored in the heavy fraction and the large area of alpine forests and meadows on the Tibetan Plateau, these alpine ecosystems might partially contribute to ＂the missing carbon sink＂.     

Carbon Sequestration in Two Alpine Soils on the Tibetan Plateau

Soil carbon sequestration was estimated in a conifer forest and an alpine meadow on the Tibetan Plateau using a carbon-14 radioactive label provided by thermonuclear weapon tests (known as bomb-¹⁴C). Soil organic matter was physically separated into light and heavy fractions. The concentration spike of bomb-¹⁴C occurred at a soil depth of 4 cm in both the forest soil and the alpine meadow soil. Based on the depth of the bomb-¹⁴C spike, the carbon sequestration rate was determined to be 38.5 g C/m² per year for the forest soil and 27.1 g C/m² per year for the alpine meadow soil. Considering that more than 60% of soil organic carbon (SOC) is stored in the heavy fraction and the large area of alpine forests and meadows on the Tibetan Plateau, these alpine ecosystems might partially contribute to "the missing carbon sink".     

青藏高原气候变化与高寒草地

综述了近五十年来青藏高原气候和高寒草地的变化趋势,阐述了气候变化对高寒草地的可能影响。气候变化主要通过水、热过程及其诱导的环境变化对青藏高原高寒草地产生显著的影响。主要过程包括：气候变化对气候带、植被带、植物、植物群落、农业生产以及生态系统固碳潜力等的影响。从目前的观测和研究结果来看,有关青藏高原气候变化及其对高寒草地的可能影响都还很难得出一致的结论。因此,如何科学评价气候变化及其预测和评价对高寒草地结构和功能的潜在影响,以及如何将已经发生的变化纳入到全球变化模型或评价体系中,以便更加精确地评估气候变化的长期影响,将成为必须要回答的关键科学问题。     

Daphnia diversity on the Tibetan Plateau measured by DNA taxonomy

Daphnia on the Tibetan Plateau has been little studied, and information on species diversity and biogeography is lacking. Here, we conducted a 4‐year survey using the barcoding fragment of the mitochondrial COI gene to determine the distribution and diversity of Daphnia species found across the Plateau. Our results show that species richness is higher than previously thought, with total described and provisional species number doubling from 5 to 10. Six of the taxonomic units recovered by DNA taxonomy agreed well with morphology, but DNA barcoding distinguished three clades each for the D. longispina (D. galeata, D. dentifera, and D. longispina) and D. pulex (D. pulex, D. cf. tenebrosa, and D. pulicaria) complexes. The sequence divergence between congeneric species varied within a large range, from 9.25% to 30.71%. The endemic D. tibetana was the most common and widespread species, occurring in 12 hyposaline to mesosaline lakes. The lineage of D. longispina is the first confirmed occurrence in west Tibet.     

Permafrost nitrogen status and its determinants on the Tibetan Plateau

It had been suggested that permafrost thaw could promote frozen nitrogen (N) release and modify microbial N transformation rates, which might alter soil N availability and then regulate ecosystem functions. However, the current understanding of this issue is confined to limited observations in the Arctic permafrost region, without any systematic measurements in other permafrost regions. Based on a large‐scale field investigation along a 1,000 km transect and a laboratory incubation experiment with a ¹⁵N pool dilution approach, this study provides the comprehensive evaluation of the permafrost N status, including the available N content and related N transformation rates, across the Tibetan alpine permafrost region. In contrast to the prevailing view, our results showed that the Tibetan alpine permafrost had lower available N content and net N mineralization rate than the active layer. Moreover, the permafrost had lower gross rates of N mineralization, microbial immobilization and nitrification than the active layer. Our results also revealed that the dominant drivers of the gross N mineralization and microbial immobilization rates differed between the permafrost and the active layer, with these rates being determined by microbial properties in the permafrost while regulated by soil moisture in the active layer. In contrast, soil gross nitrification rate was consistently modulated by the soil content in both the permafrost and the active layer. Overall, patterns and drivers of permafrost N pools and transformation rates observed in this study offer new insights into the potential N release upon permafrost thaw and provide important clues for Earth system models to better predict permafrost biogeochemical cycles under a warming climate.     

藏鸡对高原环境的适应

藏鸡是我国青藏高原地区特有的地方品种,高原地区特有的高寒低压缺氧的地理环境,使其在高原适应性和机体抵抗力方面更胜于其他鸡种。本文综述了藏鸡对高原环境适应的研究现状,提出分子水平上探究藏鸡对高原环境的适应机制研究,为其标记辅助选育提供借鉴。     

Dietary Overlap of Przewalski's Gazelle,Tibetan Gazelle,and Tibetan Sheep on the Qinghai-Tibet Plateau

Abstract: We studied food composition and overlap among sympatric Przewalski's gazelle (Procapra przewalskii), Tibetan gazelle (P. picticaudata), and Tibetan domestic sheep (Ovis aries) in the Upper Buha River, Qinghai-Tibet Plateau, using microscopic fecal analysis. The 2 gazelles forage mainly on plants from Leguminosae and Compositae families and domestic sheep feed mainly on Gramineae and Cyperaceae. The dietary overlap index between the 2 gazelles and Tibetan domestic sheep increased from 0.43 to 0.58 during the plant-growing period to 0.76–0.77 during the plant-withering period, which indicated competition for foods intensified during the food-limited season. Although the 2 gazelle ate similar diets, they might avoid food competition by occupying different foraging areas. We suggest reducing numbers of overwintering Tibetan domestic sheep on pastures to lessen survival pressure on the endangered gazelles during winters.     

西藏高原拉萨河流域生态风险评估

随着我国西部崛起战略的实施,社会经济发展所引发的生态风险逐渐引起生态学者和政府管理部门的高度关注。拉萨河流域作为高原生物多样性的维持基地,也是我国及东亚的重要生态安全屏障区,为有效管控其社会经济发展中产生的生态环境风险,通过适合高寒流域特征的风险评估方法,识别了研究区风险源,选择风险受体,采用相对风险模型计算了各研究单元的风险等级。结果表明:城关区的生态风险最高,主要风险来源于城镇扩张和旅游娱乐;当雄县生态风险最低,风险来源于畜牧养殖;林周县、曲水县、达孜县、堆龙德庆区以及墨竹工卡县的首要风险源是农业污染。水体和湿地的风险值明显高于其他生态系统。风险值的空间分布表现为南高北低的特征,且风险等级以城关区为中心向外围逐渐降低。     

Seasonal Response of Grasslands to Climate Change on the Tibetan Plateau

Background

Monitoring vegetation dynamics and their responses to climate change has been the subject of considerable research. This paper aims to detect change trends in grassland activity on the Tibetan Plateau between 1982 and 2006 and relate these to changes in climate.

Methodology/Principal Findings

Grassland activity was analyzed by evaluating remotely sensed Normalized Difference Vegetation Index (NDVI) data collected at 15-day intervals between 1982 and 2006. The timings of vegetation stages (start of green-up, beginning of the growing season, plant maturity, start of senescence and end of the growing season) were assessed using the NDVI ratio method. Mean NDVI values were determined for major vegetation stages (green-up, fast growth, maturity and senescence). All vegetation variables were linked with datasets of monthly temperature and precipitation, and correlations between variables were established using Partial Least Squares regression. Most parts of the Tibetan Plateau showed significantly increasing temperatures, as well as clear advances in late season phenological stages by several weeks. Rainfall trends and significant long-term changes in early season phenology occurred on small parts of the plateau. Vegetation activity increased significantly for all vegetation stages. Most of these changes were related to increasing temperatures during the growing season and in some cases during the previous winter. Precipitation effects appeared less pronounced. Warming thus appears to have shortened the growing season, while increasing vegetation activity.

Conclusions/Significance

Shortening of the growing season despite a longer thermally favorable period implies that vegetation on the Tibetan Plateau is unable to exploit additional thermal resources availed by climate change. Ecosystem composition may no longer be well attuned to the local temperature regime, which has changed rapidly over the past three decades. This apparent lag of the vegetation assemblage behind changes in climate should be taken into account when projecting the impacts of climate change on ecosystem processes.     

Impact of historic grazing on steppe soils on the northern Tibetan Plateau

Aims

Few studies have focused on changes in the physical and chemical properties of soils that are induced by grazing at high altitudes. Our aim was to identify potential responses of soil to grazing pressure on the semiarid steppe of the northern Tibetan Plateau and their probable causes.

Methods

Fractal geometry to describe soil structure, soil dynamics, and physical processes within soil is becoming an increasingly useful tool that allows a better understanding of the performance of soil systems. In this study, we sampled four experimental areas in the northern part of the Tibetan Plateau under different grazing intensities: ungrazed, lightly grazed, moderately grazed and heavily grazed plots. Fractal methods were applied to characterise particle-size distributions and pore patterns of soils under different grazing intensities.

Results

Our results reveal a highly significant decrease in the fractal dimensions of particle size distributions (D ₁ ) and the fractal dimensions of all pores (D ₂ ) with increasing grazing intensity. Soil organic carbon (SOC), total N and total P concentrations increased significantly with decreasing grazing intensity. We did not find differences in soil pH in response to grazing.

Conclusions

Grazing induced a significant deterioration of the physical and chemical topsoil properties in the semiarid steppe of the northern Tibetan Plateau. Fractal dimensions can be a useful parameter for quantifying soil degradation due to human activities.     

Reliability of pollen ratios for environmental reconstructions on the Tibetan Plateau

Aim Pollen ratios are widely used to gain palaeovegetation and palaeoclimatic information from fossil pollen spectra, although their applicability has seldom been tested with modern pollen data. I used a data set of 113 lake‐surface sediments from the eastern Tibetan Plateau to test the reliability of several pollen ratios. Location The lake‐surface pollen spectra cover a wide range of vegetation types (temperate desert, temperate steppe, alpine desert, alpine steppe, high‐alpine meadow, sub‐alpine shrub, coniferous and mixed forest) and climatic conditions (mean July temperature, T_July: 4.0–17.4°C; mean annual precipitation, P_ann: 104–670 mm). Methods Lake‐surface sediments were analysed palynologically, and several pollen ratios were calculated. These ratios were interpreted with respect to vegetation and climatic conditions. Results The arboreal pollen sum (AP) was highest in samples from forested areas and was significantly correlated with P_ann (r² = 0.44). In non‐forested areas, samples from large lakes and from lakes surrounded by sparse vegetation had increased AP values, suggesting that AP is a useful vegetation density indicator. Artemisia/Chenopodiaceae (A/C) ratios were lowest in desert areas and were positively correlated with P_ann (r² = 0.25). The aridity pollen index was inappropriate for inferring (palaeo‐)climatic information from samples on the eastern Tibetan Plateau as it had no significant correlation with the environmental factors. Artemisia/Cyperaceae (A/Cy) ratios had a significant correlation with T_July (r² = 0.23), but only a weak correlation with P_ann, which indicates that the A/Cy ratio is applicable as a temperature indicator. Furthermore, it is a valuable tool for the differentiation of high‐alpine meadow from steppe vegetation. Main conclusions AP sum, A/C ratio and A/Cy ratio are useful tools for qualitative and semi‐quantitative palaeoenvironmental reconstruction on the Tibetan Plateau; however, the results obtained should not be interpreted quantitatively.     

青藏高原生态气候因子的空间格局

基于1966—2005年间青藏高原及其周边113个气象站的观测资料,对研究区最热月均温、最冷月均温、≥10 ℃日数、潜在蒸散、年降水量和干燥度指数的空间格局进行了分析.结果表明：青藏高原最热月均温在7 ℃～20 ℃,由东部周边地区向中部呈逐渐减小趋势;最冷月均温在-18.4 ℃～8 ℃,由南向北逐渐减小;青藏高原≥10 ℃日数由东部周边向中部地区逐渐减小,研究区东南部的≥10 ℃日数最大（150 d以上）,中部地区最少（<50 d）;研究区年均降水量由东南部向西北地区呈减少趋势;青藏高原的潜在蒸散（由Thornthwaite模型计算）由东南向西北逐渐减小,大部分地区在330～750 mm;青藏高原干旱地区所占比例较大,主要位于研究区西北部,湿润地区面积小,主要位于东部和东南部.青藏高原的植被分布与其生态气候限制因子有较好的一致性,生态气候因子可较好地刻画植被分布状况.     

青藏高原食草动物粪栖真菌的多样性

分别利用真菌通用引物和厌氧真菌特异引物构建了西藏地区3种反刍动物和1种单胃动物共8份新鲜粪样的ITS克隆文库,以通过系统发育分析解析其中好氧真菌与厌氧真菌的多样性。通用引物ITS文库测得324条真菌序列,分别属于子囊菌门Ascomycota 3个目、担子菌门Basidiomycota 2个目、接合菌门Zygomycota的1个目和严格厌氧的新丽鞭毛菌门Neocallimastigomycota,共24个OTUs。其中,子囊菌门相对丰度最高,占80.6%;新丽鞭毛菌门相对丰度最低,仅占0.6%。大部分OTUs与已知真菌属、种关系较远。厌氧真菌特异引物文库测得661条序列,全部属于新丽鞭毛菌门,包括所有已知的厌氧霉属Anaeromyces、盲肠菌属Caecomyces、肠霉属Cyllamyces、新丽鞭毛属Neocallimastix、奥式霉属Orpinomyces、胃梨囊霉属Piromyces 6个属和3个未培养的属级类群(NG9、NG10、NG11),共29个OTUs。其中3个已知的单中心属存在于所有反刍动物样品中,并以Piromyces相对丰度最高(37.4%)。单胃动物马粪样中全部为NG9类群。NG9是本研究新发现的属级类群,研究中同时揭示有多个未培养种和潜在的新种。研究结果证明青藏高原反刍动物粪栖真菌多样性较高,并存在丰富的未培养种和潜在的新属及新种。