青藏高原黑土型退化草地成因与恢复

对青藏高原“黑土型”退化草地的成因和生态建设进行评述,认为“黑土型”退化草地是草原退化生态学行为在江河源区高寒草地的特有体现形式,由气候变暖、冰川退缩、过度放牧、鼠害等综合因子共同导致。生态恢复不仅要从草地建设入手,还要对草地资源进行合理管理与规划。加强牧区文化教育投资、实行草地长期承包、加强《草原法》建设等措施是我国高寒草地畜牧业健康发展的重要保证。青藏高原生态系统特有的“惰性”特征可能是导致其草地生态系统自我更新能力差、系统结构脆弱、生态恢复困难的重要原因,应深入开展该方面研究,为青藏高原生态环境建设提供依据。     

青藏高原多年冻土区高寒植被物种多样性和地上生物量

基于样方调查统计了青藏高原多年冻土区高寒草地生态系统植物的科属组成,计算了多样性指数和均匀度指数,探讨了多年冻土退化对高寒草地物种多样性的影响。结果表明:多年冻土退化过程中物种组成在属和物种丰富度上呈现降低趋势,湿、中生植物逐渐被旱中生和旱生植物替代;青藏高原多年冻土退化会导致高寒草地生态系统的物种多样性和初级生产力的降低,影响高寒草地生态系统的稳定性;物种多样性与初级生产力具有密切的抛物线型关系。     

History and evolution of alpine plants endemic to the Qinghai-Tibetan Plateau: Aconitum gymnandrum (Ranunculaceae)

How Quaternary climatic oscillations affected range distributions and intraspecific divergence of alpine plants on the Qinghai‐Tibetan Plateau (QTP) remains largely unknown. Here, we report a survey of chloroplast DNA (cpDNA) and nuclear ribosomal internal transcribed spacer (ITS) DNA variation aimed at exploring the phylogeographical history of the QTP alpine endemic Aconitum gymnandrum. We sequenced three cpDNA fragments (rpl20–rps12 intergenic spacer, the trnV intron and psbA‐trnH spacer) and also the nuclear (ITS) region in 245 individuals from 23 populations sampled throughout the species’ range. Two distinct lineages, with eastern and western geographical distributions respectively, were identified from a phylogenetic analysis of ITS sequence variation. Based on a fast substitution rate, these were estimated to have diverged from each other in the early Pleistocene approximately 1.45 Ma. The analysis of cpDNA variation identified nine chlorotypes that clustered into two major clades that were broadly congruent in geographical distribution with the two ITS lineages. The east–west split of cpDNA divergence was supported by an amova which partitioned approximately half of the total variance between these two groups of populations. Analysis of the spatial distribution of chlorotypes showed that each clade was subdivided into two groups of populations such that a total of four population groups existed in the species. It is suggested that these different groups derive from four independent glacial refugia that existed during the Last Glacial Maximum (LGM), and that three of these refugia were located at high altitude on the QTP platform itself at that time. Coalescent simulation of chlorotype genealogies supported both an early Pleistocene origin of the two main cpDNA clades and also the ‘four‐refugia’ hypothesis during the LGM. Two previous phylogeographical studies of QTP alpine plants indicated that such plants retreated to refugia at the eastern/south‐eastern plateau edge during the LGM and/or previous glacial maxima. However, the results for A. gymnandrum suggest that at least some of these cold‐tolerant species may have also survived centrally on the QTP platform throughout the Quaternary.     

青藏高原高寒草原生态系统植被碳密度分布规律及其与气候因子的关系

根据青藏高原高寒草原生态系统中以降水量为主要驱动力的东西样带和以气温为主要驱动力的南北样带内植被土壤的实测数据,分析了这一区域植被碳密度的分布特征及其与气候因子之间的关系.结果表明,在南北样带内(北纬28°46′～31°40′),植被碳密度首先随纬度的增加而增加,当纬度达到约北纬30°16′处,植被碳密度达到最大值0.873 1 kg·m-2,之后,则随纬度的增加而减少,植被碳密度总体上呈现出南北低、中间高的分布特征;在东西样带内(东经80°02′～91°50′),植被碳密度随经度的增加而增加,呈现出东高西低的分布特征.在南北样带内植被碳密度与年均降水量和年均气温之间的偏相关系数均达到极显著水平,而在东西样带内植被碳密度与年均降水量和年均气温之间的偏相关系数也均达到显著水平;在南北样带内植被碳密度先随年均气温和年均降水量的增加而增加,当年均气温达到约-1.5 ℃、年均降水量达到约497.0 mm时,植被碳密度达到最大值1.329 6 kg·m-2,之后,随年均气温和年均降水量的增加而减少;在东西样带内植被碳密度也先随年均气温和年均降水量的增加而增加,当年均气温达到约0.7 ℃、年均降水量达到约409.0 mm时,植被碳密度达到最大值1.208 3 kg·m-2,之后,随年均气温和年均降水量的增加而减少.研究结果显示,青藏高原高寒草原生态系统南北样带和东西样带内的植被碳密度分布均是年均气温和年均降水量综合作用的结果,且年均降水量的作用大于年均气温.     

No temperature acclimation of soil extracellular enzymes to experimental warming in an alpine grassland ecosystem on the Tibetan Plateau

Alpine grassland soils store large amounts of soil organic carbon (SOC) and are susceptible to rising air temperature. Soil extracellular enzymes catalyze the rate-limiting step in SOC decomposition and their catalysis, production and degradation rates are regulated by temperature. Therefore, the responses of these enzymes to warming could have a profound impact on carbon cycling in the alpine grassland ecosystems. This study was conducted to measure the responses of soil extracellular enzyme activity and temperature sensitivity (Q₁₀) to experimental warming in samples from an alpine grassland ecosystem on the Tibetan Plateau. A free air-temperature enhancement system was set up in May 2006. We measured soil microbial biomass, nutrient availability and the activity of five extracellular enzymes in 2009 and 2010. The Q₁₀ of each enzyme was calculated using a simple first-order exponential equation. We found that warming had no significant effects on soil microbial biomass C, the labile C or N content, or nutrient availability. Significant differences in the activity of most extracellular enzymes among sampling dates were found, with typically higher enzyme activity during the warm period of the year. The effects of warming on the activity of the five extracellular enzymes at 20 °C were not significant. Enzyme activity in vitro strongly increased with temperature up to 27 °C or over 30 °C (optimum temperature; T_opt). Seasonal variations in the Q₁₀ were found, but the effects of warming on Q₁₀ were not significant. We conclude that soil extracellular enzymes adapted to seasonal temperature variations, but did not acclimate to the field experimental warming.     

青藏高原东缘高寒草甸植物群落的开花物候

利用2008年整个生长季的地面花期物候观测数据,初步研究了青藏高原东缘高寒草甸植物群落开花物候的动态、格局、参数间的联系及其与物种共存的关系.结果表明:植物群落的花期从5月初开始,8月上旬进入开花数目的高峰期,到10月初结束,遍及整个生长季节;不同物种进入花期的时间、峰值期及持续时间不同,各物种花期物候生态位相分离,但群落中大部分物种开花峰值期主要集中在7-8月;不同物种的开花峰值期时间与花期持续时间有负相关关系:开花越早的植物其花期持续时间越长;物种间的开花物候差异主要是由植物本身的特性和环境条件共同决定的,而群落水平上的开花物候格局主要受非生物因素的控制.

Abstract：

Based on the observation on the flowering phenology in the whole growth season of 2008, the dynamics, patterns, and relationships among various parameters of flowering phenology of alpine meadow plant community in eastern Qinghai-Tibetan Plateau were studied, and the potential effects of above-mentioned factors on the species coexistence of the plant community were approached. The flowering period of the plant community started from early May, came into peak period in the first ten days of August, and ended in early October, extending all over whole growth period. Different species had their different first flowering date, peak flowering period, and flowering duration, and their flowering phenologieal niches differentiated obviously. Most species had their peak flowering period centralized in July-August. There was a negative rela-tionship between peak flowering period and flowering duration: the earlier the flowering, the lon-ger the flowering duration. The differences in flowering phonology among the species were mainly determined by species per se characteristics and environmental conditions, but the flowering phe-nological pattern on community level was primarily controlled by abiotic factors.     

羌塘高原寒草地生态系统生产力动态

基于实测气象数据和遥感数据,分析了藏北地区气候变化趋势,并采用植被-气候综合模型和CASA模型模拟分析了藏北草地潜在和现实净第一性生产力(NPP)的动态变化和空间格局.结果表明:1955-2004年间,羌塘高原年平均气温上升了1.37 ℃,降水量增加了63 mm,中、东部区域的气候趋于暖湿化,西部区域趋于暖干化,目前气候变化尚未引起草地退化.草地潜在NPP平均值为东部＞中部＞西部.1982-2004年,由于水热条件的变化,中部区域的潜在NPP增加值最高,达0.55 t·hm-2·a-1,东部和西部分别为0.51和0.21 t·hm-2·a-1;东、中、西部现实NPP增量分别为-0.19、-0.03和0.20 t·hm-2·a-1.超载过牧是东、中部草地退化的主要原因,中部是草地保护恢复工程的最佳实施区域.     

Reciprocal interactions between plants and soil in an upland grassland

Through the production of litter, plants with different life history strategies are predicted to both affect and be affected by the properties of soil. Competitive species are expected to increase the fertility of, and have a positive growth feedback with, soil, whereas stress-tolerant species should decrease fertility but show no growth feedback. We maintained monocultures of competitive (Lolium perenne and Agrostis capillaris) and stress-tolerant (Festuca ovina and Nardus stricta) grasses on an unproductive grassland for six years. The Nardus soil developed significantly greater inorganic nitrogen than the Agrostis and Festuca soil, and significantly greater soil moisture content than the Festuca soil. However, there were no differences in organic matter content, phosphate or bulk density between the soil types. In a greenhouse assay, each species was grown in soil cores from the different monocultures as well as natural turf. There were significant differences in growth between plant species and soil types. As expected, L. perenne produced the greatest amount of biomass. However, plants grown on Nardus soil were twice as large and had a 21% lower root allocation than plants grown on any of the other soil types. Lolium perenne, A. capillaris and F. ovina had significant negative growth feedbacks with their own soil (−0.460, −0.821 and −0.792, respectively) and N. stricta had a significant positive feedback (0.560). This study highlights the difficulties of predicting how plant traits will affect soil properties.     

Floral closure in response to temperature and pollination in Gentiana straminea Maxim. (Gentianaceae), an alpine perennial in the Qinghai-Tibetan Plateau

Floral closure may be induced by pollination and various other factors, but is rarely studied comprehensively. Different kinds of floral closure should have various effects on reproductive fitness of plants. Two contrasting types of floral closure were observed in the flowers of Gentiana straminea Maxim. in the eastern Qinghai-Tibetan Plateau. The first type occurred prior to pollination during both gender phases, in response mainly to decreasing air temperatures. Flowers closed when decreasing temperatures approached 20°C and subsequently began to reopen the following day during mid-morning when air temperatures warmed to approximately 13–15 °C. This kind of floral closure can protect pollen grains on either stamens or stigmas, increasing fitness of both male and female. Following pollination, permanent floral closure occurred, although there was a delay between the dates of pollination and permanent closure, during which flowers continued to show temporary closure in response to low temperature episodes. The time required for permanent, pollination-induced closure varied according to the age of the gender phase, including a prolonged time before closure if pollination occurred early in the female phase. The retaining of permanent closed flowers increased both approaching (to inflorescences) and visiting (to unpollinated flowers) frequencies of individual plants when with fewer open flowers and the persisting corolla is further beneficial for seed sets of these pollinated flowers. Thus, two separate types of floral closure, one in response to environmental cues and the other in response to the age of each gender stage, appeared to have a strong influence on reproductive fitness in this species. These results revealed a different adaptive strategy of alpine plants in the sexual reproduction assurance in addition to the well-known elevated floral longevity, dominant role of more effective pollinators and increased reproduction allocation in the arid habitats.     

Contribution of arbuscular mycorrhizal fungi of sedges to soil aggregation along an altitudinal alpine grassland gradient on the Tibetan Plateau

The diversity of arbuscular mycorrhizal fungi (AMF) in sedges on the Tibetan Plateau remains largely unexplored, and their contribution to soil aggregation can be important in understanding the ecological function of AMF in alpine ecosystems. Roots of Kobresia pygmaea C.B. Clarke and Carex pseudofoetida Kük. in alpine Kobresia pastures along an elevational transect (4149–5033 m) on Mount Mila were analysed for AMF diversity. A structural equation model was built to explore the contribution of biotic factors to soil aggregation. Sedges harboured abundant AMF communities covering seven families and some operational taxonomic units are habitat specific. The two plant species hosted similar AMF communities at most altitudes. The relative abundance of the two sedges contributed largely to soil macroaggregates, followed by extraradical mycorrhizal hyphae (EMH) and total glomalin‐related soil protein (T‐GRSP). The influence of plant richness was mainly due to its indirect influence on T‐GRSP and EMH. There was a strong positive correlation between GRSP and soil total carbon and nitrogen. Our results indicate that mycorrhization might not be a major trait leading to niche differentiation of the two co‐occurring sedge species. However, AMF contribute to soil aggregation and thus may have the potential to greatly influence C and N cycling in alpine grasslands.     

Fine root response to soil resource heterogeneity differs between grassland and forest

Soil resource heterogeneity has clear effects on plant root development and overall plant performance. Here we test whether contrasting vegetation types have similar or different responses to soil patches of differing resource availability. We examined the fine root responses of grassland and forest vegetation at the northern edge of the Great Plains to transplanted patches of resource-poor and resource-rich soils, using rhizotron imaging. Every aspect of measured root behavior, including root length, production, mortality, turnover, variability and size distribution, varied significantly between patch types, and most aspects also varied between vegetation types. Most importantly, differential responses to patches between grassland and forest were shown by significant interactions between patch type and vegetation for two response variables. First, root length variability was significantly lower in resource-rich compared to resource-poor patches in forest but not grassland. Second, the proportion of very fine roots was significantly greater in resource-rich than resource-poor patches in forests but not grassland. Thus, compared to grassland, forest more fully occupied resource-rich patches relative to resource-poor patches by allocating more growth to very fine roots. We report the first example of significant differences between vegetation types (grassland and forest) in root responses to soil resource heterogeneity measured in a field experiment. The relatively high ability of forest roots to more fully occupy resource-rich patches is consistent with the global expansion of woody vegetation and associated increases in soil heterogeneity.     

Responses in leaf functional traits and resource allocation of a dominant alpine sedge (Kobresia pygmaea) to climate warming in the Qinghai-Tibetan Plateau permafrost region

Assessing the influence of warming on leaf traits, carbon, and nutrient concentrations above and below ground to understand how the dominant sedge Kobresia pygmaea (C. B. Clarke) C. B. Clarke may respond and adapt to extant and future climate in the alpine meadow of the Qinghai-Tibetan Plateau. A warming experiment was conducted in the permafrost region of the Qinghai-Tibetan Plateau from 2008 to 2009. Two 2-year warming treatments (T1, annual warming of 2.1°C; T2, annual warming of 4.4°C) were used, and responses of leaf traits and above- and belowground carbon, nitrogen, and phosphorus concentrations of K. pygmaea were examined. The results show that both moderate (T1) and more extensive (T2) warming decreased leaf mass, leaf thickness, and vascular bundle size, and increased the mass-based photosynthetic rate (A_mass) and photosynthetic nitrogen use efficiency (PNUE). A moderate warming significantly decreased leaf carbon (C), nitrogen (N), and phosphorus (P), and root C and N concentrations of K. pygmaea. These decreases were even more pronounced under the more extensive warming. The decreases in leaf N and P were significantly larger than the decrease in leaf C concentration. Root P concentration increased more under the extensive than the moderate warming. The observed increase in leaf C:N ratio in the warming treatment indicates that enhanced temperature may increase the long-term nitrogen use efficiency of K. pygmaea leaves. This again suggests that K. pygmaea might adapt well to future climate warming, and that nitrogen might be a more important factor for K. pygmaea dominated alpine meadows under future climate warming.     

高寒山区混播草地燕麦和毛苕子种间的竞争关系

植物通过调整生长发育阶段的生长行为和形态特征来适应种间竞争关系的变化.在祁连山区建立了禾本科牧草燕麦与豆科毛苕子混播草地,以相对产量(RY)、相对密度(RD)和相对产量总值(RYT)为指标,研究了这2种1年生植物在不同物候期的种间竞争关系.结果表明:随着物候期的变化,种间竞争关系发生着动态变化,在出苗期,二者存在激烈的种间竞争,2个物种限制彼此相对数量的发展,株高、叶片数等增长缓慢;分蘖期,种间竞争转换为种间协作,分蘖数、叶片数等迅速增大,相对产量总值不断积累;进入拔节期,毛苕子开始扩大相对数量,限制了燕麦的生长,导致了激烈的种内竞争;到结实期,种内竞争再次转化为种间竞争.不同物候期的燕麦和毛苕子通过对植物形态可塑性的调整,更好地响应了种间竞争关系的转换.     

Net primary productivity and spatial distribution of vegetation in an alpine wetland,Qinghai-Tibetan Plateau

To initially describe vegetation structure and spatial variation in plant biomass in a typical alpine wetland of the Qinghai-Tibetan Plateau, net primary productivity and vegetation in relationship to environmental factors were investigated. In 2002, the wetland remained flooded to an average water depth of 25 cm during the growing season, from July to mid-September. We mapped the floodline and vegetation distribution using GPS (global positioning system). Coverage of vegetation in the wetland was 100%, and the vegetation was zonally distributed along a water depth gradient, with three emergent plant zones (Hippuris vulgaris-dominated zone, Scirpus distigmaticus-dominated zone, and Carex allivescers-dominated zone) and one submerged plant zone (Potamogeton pectinatus-dominated zone). Both aboveground and belowground biomass varied temporally within and among the vegetation zones. Further, net primary productivity (NPP) as estimated by peak biomass also differed among the vegetation zones; aboveground NPP was highest in the Carex-dominated zone with shallowest water and lowest in the Potamogeton zone with deepest water. The area occupied by each zone was 73.5% for P. pectinatus, 2.6% for H. vulgaris, 20.5% for S. distigmaticus, and 3.4% for C. allivescers. Morphological features in relationship to gas-transport efficiency of the aerial part differed among the emergent plants. Of the three emergent plants, H. vulgaris, which dominated in the deeper water, showed greater morphological adaptability to deep water than the other two emergent plants.     

土壤-牧草氮素供需状况变化对高寒草甸植被演替与草地退化的影响

在中国科学院海北高寒草甸生态系统定位站 ,选取处于不同退化阶段的具有典型代表意义的草甸草场为研究对象 ,通过对其土壤氮素矿化补给能力、牧草对氮素的需求量的研究 ,探讨土壤 牧草氮素供需状况变化对高寒草甸植被演替与草地退化的影响。结果表明 ,在牧草生长季 5～ 8月 ,高寒草甸土壤的氮素矿化补给量为 15 86 g·m-2 ,而随着高寒草甸退化程度的加重 ,植物群落中优势种群由禾草演替为禾草 苔草 嵩草、嵩草至杂类草 ,其牧草生长需要的总氮量分别为 2 2 86、2 4 87、37 3、14 96g·m-2 ,只有在杂类草草甸阶段 ,其牧草生长对氮素的需求才与其土壤氮素供求相适配 ,可见养分是高寒草甸植被演替与草场退化的重要驱动因子之一。     

The negative interaction between the degradation of phenanthrene and tricyclazole in medium,soil and soil/compost mixture

To assess the co-catabolism of phenanthrene and tricyclazole in different samples, the interaction during the degradation of phenanthrene and tricyclazole were investigated in medium, soil and soil/spent mushroom compost (SMC) mixture. Generally, tricyclazole showed a negative influence on the activity of phenanthrene dioxygenase and it inhibited the degradation of phenanthrene prominently, both in cultures of phenanthrene catabolic isolates (Sphingomonas paucimobilis ZX4 and the mixed flora M1) and soils, while a similar inhibition caused by phenanthrene was also found for the degradation of tricyclazole in soil/SMC. However, the inhibition effect on phenanthrene degradation was eliminated in soil/SMC mixture, due to the abundant microorganisms and enormous catabolic potential in SMC. Furthermore, it was proved that the negative influence between phenanthrene and tricyclazole was most likely derived from the molecular similarity and it tended to decrease with improved microbial diversity in environment.     

The impact of land degradation on the C pools in alpine grasslands of the Qinghai-Tibet Plateau

Aims

To determine the effect of grassland degradation on the soil carbon pool in alpine grassland.

Methods

In this study, we calculated the carbon pool in the above-and below-ground biomass, the soil microbial biomass carbon pool, the total organic carbon pool and the soil total carbon.

Results

Grassland degradation has resulted in decreases in biomass and carbon content and has changed the ratio of roots to shoots. However, there was less influence of degradation on dead root biomass. There was most likely a lag effect of changes in dead root biomass following grassland degradation. In the alpine grassland ecosystem, the carbon pool in soil accounts for more than 92 % of the total carbon both in vegetation and soil. The carbon in alpine grassland is stored primarily in the form of total organic carbon below-ground. As organic carbon decreases, the ratio of the microbial biomass carbon pool to the total organic carbon pool increases and then declines with increasing degradation level. Along the grassland degradation gradient, the total vegetation biomass (above-and below-ground) and the soil carbon pool (microbial biomass C, total organic C and total C) all decreased.     

Methane emission by plant communities in an alpine meadow on the Qinghai-Tibetan Plateau: a new experimental study of alpine meadows and oat pasture

Recently, plant-derived methane (CH₄) emission has been questioned because limited evidence of the chemical mechanism has been identified to account for the process. We conducted an experiment with four treatments (i.e. winter-grazed, natural alpine meadow; naturally restored alpine meadow eight years after cultivation; oat pasture and bare soil without roots) during the growing seasons of 2007 and 2008 to examine the question of CH₄ emission by plant communities in the alpine meadow. Each treatment consumed CH₄ in closed, opaque chambers in the field, but two types of alpine meadow vegetation reduced CH₄ consumption compared with bare soil, whereas oat pasture increased consumption. This result could imply that meadow vegetation produces CH₄. However, measurements of soil temperature and water content showed significant differences between vegetated and bare soil and appeared to explain differences in CH₄ production between treatments. Our study strongly suggests that the apparent CH₄ production by vegetation, when compared with bare soil in some previous studies, might represent differences in soil temperature and water-filled pore space and not the true vegetation sources of CH₄.     

青藏高原草地生态系统对气候变化的响应

青藏高原高寒草地生态系统对气候变化高度敏感,其如何响应和反馈气候变化一直以来受到极大关注.本文系统综述了近5年来有关青藏高原草地生态系统在物候、生产力、碳循环等方面对气候变化的响应过程以及应对气候变化的适应性管理的最新研究成果,发现气候变化对高寒草地生态系统的诸多影响还存在很大的不确定性.多数研究结果表明,增温使高寒草甸的植被物候提前和初级生产力水平提高,而高寒草原有相反的影响趋势,说明不同地域、不同群落类型对不同季节温度变化的响应模式不同.而气候变化对物种多样性和碳循环有关过程的影响结果尚没有一致的结论,时空尺度和方法上的差异可能是导致不同结果的主要原因.因此,建议在增强时空异质性的响应与反馈研究的同时,更需要加强生态过程和机理的研究.     

The contribution of grazing to plant diversity in alpine grassland and heath

Abstract Grazing on transplants of a grass, a forb and a tree was examined in low-diversity grassland and more diverse heath in Australia's Snowy Mountains. Transplants were surrounded by 2 mm mesh netting. In one grassland plot, grazers (probably soil invertebrates) attacked 40–90% of tree and forb seedlings but no grass seedlings. In heath, which had about half the grass cover of grassland, grazers consumed grasses but not trees or forbs. The results suggest that grazers can depress diversity in grassland by attacking species other than grass. In heath, they may promote diversity by attacking only grass and releasing other species from competition.