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。     

Strong impacts of daily minimum temperature on the green‐up date and summer greenness of the Tibetan Plateau

Understanding vegetation responses to climate change on the Tibetan Plateau (TP) helps in elucidating the land–atmosphere energy exchange, which affects air mass movement over and around the TP. Although the TP is one of the world's most sensitive regions in terms of climatic warming, little is known about how the vegetation responds. Here, we focus on how spring phenology and summertime greenness respond to the asymmetric warming, that is, stronger warming during nighttime than during daytime. Using both in situ and satellite observations, we found that vegetation green‐up date showed a stronger negative partial correlation with daily minimum temperature (T_min) than with maximum temperature (T_max) before the growing season (‘preseason’ henceforth). Summer vegetation greenness was strongly positively correlated with summer T_min, but negatively with T_max. A 1‐K increase in preseason T_min advanced green‐up date by 4 days (P < 0.05) and in summer enhanced greenness by 3.6% relative to the mean greenness during 2000–2004 (P < 0.01). In contrast, increases in preseason T_max did not advance green‐up date (P > 0.10) and higher summer T_max even reduced greenness by 2.6% K^?1 (P < 0.05). The stimulating effects of increasing T_min were likely caused by reduced low temperature constraints, and the apparent negative effects of higher T_max on greenness were probably due to the accompanying decline in water availability. The dominant enhancing effect of nighttime warming indicates that climatic warming will probably have stronger impact on TP ecosystems than on apparently similar Arctic ecosystems where vegetation is controlled mainly by T_max. Our results are crucial for future improvements of dynamic vegetation models embedded in the Earth System Models which are being used to describe the behavior of the Asian monsoon. The results are significant because the state of the vegetation on the TP plays an important role in steering the monsoon.     

内蒙古主要草原类型植物物候对气候波动的响应

物候是气候变化的指示者,由于不同地区植被类型不同,导致其对气候波动的响应方式不同。利用2004—2013年内蒙古草原区生态监测站群落优势种物候观测资料和同时段的气象资料,分析了不同草原类型区优势种物候期变化及其与气候因子间的相互关系,结果表明:(1)2004—2013年内蒙古草原区各时段气候波动趋势均不显著,返青前以气温降低、降水增加趋势为主;黄枯前草甸草原、典型草原以气温降低、降水增加趋势为主,荒漠草原变化趋势相反。(2)2004—2013年典型草原植物返青期平均提前4.01 d,黄枯推后10.35 d,生长季延长14.36 d;草甸草原返青期提前2.04 d,黄枯期推后12.68 d,生长季延长14.72 d;荒漠草原物候变化趋势最小,返青期平均提前了1.32 d,黄枯期平均推后了9.58 d,生长季延长了10.90 d。(3)内蒙古草原区植物返青期主要受气温波动的影响,草甸草原返青期与前3个月平均气温的负相关最为显著,气温每升高1℃,返青期约提前1.123 d;典型草原、荒漠草原返青期与前2个月平均气温的负相关最为显著气,气温每升高1℃,返青期约提前1.137 d和1.743 d。(4)典型草原区植物黄枯期受前1—2月平均气温和累积降水的共同影响,与夏季平均气温和当月降水量的相关最为显著,夏季气温每升高1℃,黄枯期约提前2.250 d,当月降水每增加1 mm,黄枯期约推后0.119 d。草甸草原、荒漠草原植物黄枯期与各时段降水、气温的相关均不显著,影响黄枯机制比较复杂。     

气候因子通过土壤微生物生物量氮促进青藏高原高寒草地地上生态系统功能

近年来, 在人类活动和气候变化的影响下, 物种多样性丧失趋势不断加剧, 对生态系统功能带来严重后果。目前, 关于生态系统功能的研究, 忽略了土壤和微生物碳氮养分循环过程对地上生态系统功能(AEF)的重要驱动作用, 而土壤碳氮要素和微生物的任何变化都有可能改变地下群落对生态系统功能的维持作用。该研究旨在探究高寒草地AEF的主要控制因子, 以及其关键要素对AEF的作用机理。2015年7-8月, 对青藏高原地区115个样点进行了草地群落和土壤属性等要素样带调查; 综合植物地上生物量, 叶片碳、氮和磷含量等参数计算AEF值, 分析地下土壤有机碳含量、全氮含量、生物量等关键要素对AEF值的影响。结合取样点年降水量和年平均气温, 深入探讨影响AEF的主要控制因子和作用机理。结果表明降水对AEF有较大影响, 而气温影响相对较低。年降水量、土壤微生物生物量碳含量和干旱指数对AEF值的相对重要性贡献较高(重要值分别为21.1%、10.9%和10.1%), 控制青藏高原高寒草地AEF值的关键是土壤因子。在气候因子对土壤养分和微生物的作用下, 土壤微生物生物量氮含量在调控高寒草地AEF值方面发挥重要作用。     

Responses of phenology and biomass production of boreal fens to climate warming under different water‐table level regimes

Climate change affects peatlands directly through increased air temperatures and indirectly through changes in water‐table level (WL). The interactions of these two still remain poorly known. We determined experimentally the separate and interactive effects of temperature and WL regime on factors of relevance for the inputs to the carbon cycle: plant community composition, phenology, biomass production, and shoot:root allocation in two wet boreal sedge‐dominated fens, “southern” at 62°N and “northern” at 68°Ν. Warming (1.5°C higher average daily air temperature) was induced with open‐top chambers and WL drawdown (WLD; 3–7 cm on average) by shallow ditches. Total biomass production varied from 250 to 520 g/m², with belowground production comprising 25%–63%. Warming was associated with minor effects on phenology and negligible effects on community composition, biomass production, and allocation. WLD clearly affected the contribution of different plant functional types (PFTs) in the community and the biomass they produced: shrubs benefited while forbs and mosses suffered. These responses did not depend on the warming treatment. Following WLD, aboveground biomass production decreased mainly due to reduced growth of mosses in the southern fen. Aboveground vascular plant biomass production remained unchanged but the contribution of different PFTs changed. The observed changes were also reflected in plant phenology, with different PFTs showing different responses. Belowground production increased following WLD in the northern fen only, but an increase in the contributions of shrubs and forbs was observed in both sites, while sedge contribution decreased. Moderate warming alone seems not able to drive significant changes in plant productivity or community composition in these wet ecosystems. However, if warming is accompanied by even modest WL drawdown, changes should be expected in the relative contribution of PFTs, which could lead to profound changes in the function of fens. Consequently, hydrological scenarios are of utmost importance when estimating their future function.     

The responses of grassland plants to experimentally simulated climate change depend on land use and region

Macroclimatic niche properties derived from species distribution ranges are fundamental for projections of climate change impacts on biodiversity. However, it has been recognized that changes in regional or local distribution patterns also depend on interactions with land use. The reliability and transferability of large scale geographic predictions to small scale plant performance need to be tested experimentally. Thus, we asked how grassland plant species pairs with different macroclimatic niche properties respond to increased spring temperature and decrease summer precipitation in three different land‐use types. An experiment was carried out in the framework of the German Biodiversity Exploratories simulating climate change in 45 experimental plots in three geographical regions (Schorfheide‐Chorin, Hainich‐Dün, Schwäbische Alb) and three grassland management types (meadow, pasture, mown pasture). We planted six plant species as phytometers, each two of them representing congeneric species with contrasting macroclimatic niches and recorded plant survival and growth over 1 year. To quantify the species macroclimatic niches with respect to drought tolerance, the species’ distribution ranges were mapped and combined with global climate data. The simulated climate change had a general negative effect on plant survival and plant growth, irrespective of the macroclimatic niche characteristics of the species. Against expectation, species with ranges extending into drier regions did not generally perform better under drier conditions. Growth performance and survival was best in mown pastures, representing a quite intensive type of land use in all study regions. Species with higher macroclimatic drought tolerance were generally characterized by lower growth rates and higher survival rates in land‐use types with regular mowing regimes, probably because of reduced competition in the growing season. In conclusion, plant species with similar climatic niche characteristics cannot be expected to respond consistently over different regions owing to complex interactions of climate change with land use practices.     

天山北坡不同海拔梯度山地草原生态系统地上净初级生产力对气候变化及放牧的响应

以天山北坡三工河流域为例,利用改进后的Biome-BGC模型分别模拟了仅气候变化（Clm）、气候变化与放牧联合作用（ClmGra）下研究区不同海拔梯度3种山地草原生态系统（低山干旱草原,LAG;森林草甸草原,FMG;高寒草甸草原,AMG）1959—2009年地上净初级生产力（Aboveground Net Primary Production,ANPP）的动态,并通过假设27种放牧强度情景（0—8 羊/ha）模拟了其ANPP随放牧强度增加的变化趋势。近50年气候变化致使研究区各海拔梯度草原生态系统ANPP整体均呈上升趋势,但在放牧联合作用下,不同草原类型ANPP变化趋势差异显著;放牧导致FMG和AMG的ANPP呈下降态势,分别减少30.0%和33.2%,对比之下,由于1980前较低放牧强度促进了LAG的ANPP,放牧导致其ANPP整体增加1.3%。随着放牧强度增加,LAG的ANPP呈先增后减趋势,且在干旱年份最为显著;而FMG和AMG的ANPP呈显著非线性递减趋势。这些结果表明,近50年气候波动可能有利于中亚干旱区山地草原生态系统生产力的提高,但日益增强的放牧活动导致其净初级生产力显著降低;放牧对FMG与AMG生产力的负面效应随放牧强度增加而增强,但适度放牧可能促进LAG净初级生产力,尤其在干旱年份。     

植物功能群去除对高寒草甸群落结构、多样性及生产力的影响

群落中物种的丧失在干扰下普遍存在,但对生态系统过程和功能的影响仍存在较大不确定性。选取青藏高原东缘典型高寒草甸为对象,开展优势植物功能群的梯度去除试验,以模拟长期过牧干扰下物种的损失。经过连续两个生长季的功能群去除,我们对群落的物种组成、结构、多样性和生物量等特征进行了分析,探讨了上述指标的响应过程和机制。研究结果表明：（1）功能群的去除降低了群落高度,增加了物种均匀度,并显著影响了禾草、杂草优势比以及功能群多样性和优势度;（2）同时,去除操作显著减小了凋落物量与禾草生物量,并显著影响了群落地上生物量;（3）进一步分析还发现,禾草、莎草和杂草功能群之间存在显著的竞争关系,群落生产力主要取决于禾草功能群并随物种均匀度的增大而显著减小。上述结果表明,禾草在高寒草甸群落中占据竞争优势地位,植物功能群的损失主要通过改变种间竞争关系、引起有机物质丢失影响群落过程和功能。     

模拟增温改变青藏高原植物繁殖物候及植株高度

气候变化显著影响了高寒植物物候期及生长模式, 从而改变了高寒生态系统功能。而高寒植物物候期和生长状况对气候变化的响应程度, 与其自身资源分配策略有关。为了更好地探究气候变化下高寒植物繁殖物候及生长的规律, 该研究以青藏高原高寒草甸为研究对象, 按生物量从高到低选取15种常见植物, 其生物量之和占样地总生物量80%以上, 采用红外辐射器模拟增温的方法, 利用同质园实验, 观测无种间竞争条件下, 增温2年间植物返青、现蕾、开花以及结实物候, 并监测了植株高度。研究结果表明: (1)在功能群水平上, 增温使豆科类植物的返青、现蕾和开花时间分别显著提前了(8.21 ± 1.81)、(9.14 ± 2.41)和(10.14 ± 2.05) d, 使其开花持续时间显著延长了(6.14 ± 1.52) d, 而增温对其他功能群物候事件无显著影响。增温对高寒植物物候的影响存在种间及年际间差异, 但总体上增温使大多数高寒植物繁殖物候提前并且开花持续时间延长, 将更多的资源更多地分配到繁殖生长上。(2)增温显著降低了杂类草植物的植株高度(平均降低(3.58 ± 0.96) cm), 但对豆科类、禾草类及莎草类功能群植株高度没有显著影响。 增温对高寒植物植株高度的影响存在显著的种间差异以及年际差异。综上所述, 未来气候变暖背景下, 青藏高原高寒植物群落可能更早进入繁殖阶段, 从而降低在营养生长上的资源分配。另外, 由于各物种繁殖能力和营养生长对温度变化响应的差异, 气候变暖将导致高寒植物群落中各物种盖度的变化, 进而改变群落物种组成, 从而影响高寒生态系统的功能。     

近30年来青藏高原高寒草地NDVI动态变化对自然及人为因子的响应

草地生态系统是陆地生态系统的重要组成部分,在调节气候、水土保持、防风固沙、保护生物多样性等方面发挥着重要作用。青藏高原是全球海拔最高的独特地域单元,平均海拔超过4000 m,素有“世界第三极”之称,亦是我国重要的生态安全屏障,其对气候变化敏感且易受人类活动的影响,属于气候变化敏感区和生态脆弱带。近年来,由于气候变化和人类活动的不断加剧,青藏高原区域气候和环境发生了重大变化,气候变暖、水污染、草地退化和沙化等问题已严重阻碍了当地社会经济的可持续发展。高寒草地是青藏高原主要的植被类型,在气候变化和人类活动加剧的背景下,青藏高原高寒草地植被的动态变化受到人们的广泛关注。归一化植被指数（Normalized difference vegetation index, NDVI）因能有效地反映植被覆盖程度和生长状况而被广泛应用于植被动态的研究中。气温与降水被认为是影响青藏高原植被动态的主要气候因子,放牧强度与人口数量则是主要人为因子。因此,研究高寒草地植被对气候变化和人类活动的响应机制对预测未来草地变化有着重要的意义。基于青藏高原生长季草地的NDVI、气温、降水、放牧强度及人口数量等数据,在县区尺度上,采用趋势分析法探究了1982—2013年青藏高原143个县区生长季草地NDVI动态变化、气候变化及人类活动的变化,同时采用面板数据模型分析了32年来青藏高原143个县区气候、人为因子变化对草地NDVI变化的相对贡献。研究结果显示：（1）青藏高原高寒草地生长季NDVI总体呈增长趋势,草地植被生长状态呈现“整体改善、局部退化”趋势;（2）青藏高原生长季平均气温与降水量整体增加,气候呈现“暖湿化”趋势;（3）在长时间尺度上,气候因子主导了青藏高原高寒草地NDVI的变化,降雨和气温的增加促进草地NDVI的增加,放牧强度的持续增加则导致草地NDVI的减少。     

Plant functional groups mediate effects of climate and soil factors on species richness and community biomass in grasslands of Mongolian Plateau

植物功能群在调控气候和土壤因子对蒙古高原草原群落物种丰富度和生物量影响中的作用 植物功能群组成主要受环境因素驱动,同时植物功能群组成也是影响草地生物多样性和生产力的主要因素之一。因此,理解植物功能群在调控环境因素对生态系统功能和生物多样性影响中可能发挥的作用至关重要。通过对蒙古高原草原65个样点的植物生物量和物种丰富度的调查,将157种多年生草本植物分为两种植物功能群(即禾草和杂类草)。通过随机森林模型和普通最小二乘回归,确定与植物功能群物种丰富度和地上生物量显著相关的环境因素(即干燥度、土壤总氮和pH),并利用结构方程模型探讨筛选出的环境因素与群落物种丰富度和生物量间的关系,以及植物功能群在驱动这种关系中发挥的作用。干燥度与禾草、杂类草以及整个群落的地上生物量和物种丰富度均呈显著的单峰关系。所有的物种丰富度和生物量指标均与土壤总氮和pH值显著相关。禾草在维持蒙古高原草原生态系统群落生物量中起着关键作用,并受气候因素的直接影响。而杂类草物种丰富度决定了群落总丰富度,并受到土壤因素直接的调控。因此,群落组成在调控环境因素对群落生物量和植物多样性的影响中起着关键作用。     

Stream ecosystem responses to the 2007 spring freeze in the southeastern United States: unexpected effects of climate change

Some expected changes in climate resulting from human greenhouse gas emissions are clear and well documented, but others may be harder to predict because they involve extreme weather events or heretofore unusual combinations of weather patterns. One recent example of unusual weather that may become more frequent with climate change occurred in early spring 2007 when a large Arctic air mass moved into the eastern United States following a very warm late winter. In this paper, we document effects of this freeze event on Walker Branch, a well‐studied stream ecosystem in eastern Tennessee. The 2007 spring freeze killed newly grown leaf tissues in the forest canopy, dramatically increasing the amount of light reaching the stream. Light levels at the stream surface were sustained at levels considerably above those normal for the late spring and summer months due to the incomplete recovery of canopy leaf area. Increased light levels caused a cascade of ecological effects in the stream beginning with considerably higher (two–three times) rates of gross primary production (GPP) during the late spring and summer months when normally low light levels severely limit stream GPP. Higher rates of stream GPP in turn resulted in higher rates of nitrate (NO₃^?) uptake by the autotrophic community and lower NO₃^? concentrations in stream water. Higher rates of stream GPP in summer also resulted in higher growth rates of a dominant herbivore, the snail Elimia clavaeformis. Typically, during summer months net NO₃^? uptake and snail growth rates are zero to negative; however, in 2007 uptake and growth were maintained at moderate levels. These results show how changes in forest vegetation phenology can have dramatic effects on stream productivity at multiple trophic levels and on nutrient cycling as a result of tight coupling of forest and stream ecosystems. Thus, climate change‐induced changes in canopy structure and phenology may lead to large effects on stream ecosystems in the future.     

Biodiversity as a solution to mitigate climate change impacts on the functioning of forest ecosystems

Forest ecosystems are critical to mitigating greenhouse gas emissions through carbon sequestration. However, climate change has affected forest ecosystem functioning in both negative and positive ways, and has led to shifts in species/functional diversity and losses in plant species diversity which may impair the positive effects of diversity on ecosystem functioning. Biodiversity may mitigate climate change impacts on (I) biodiversity itself, as more‐diverse systems could be more resilient to climate change impacts, and (II) ecosystem functioning through the positive relationship between diversity and ecosystem functioning. By surveying the literature, we examined how climate change has affected forest ecosystem functioning and plant diversity. Based on the biodiversity effects on ecosystem functioning (B→EF), we specifically address the potential for biodiversity to mitigate climate change impacts on forest ecosystem functioning. For this purpose, we formulate a concept whereby biodiversity may reduce the negative impacts or enhance the positive impacts of climate change on ecosystem functioning. Further B→EF studies on climate change in natural forests are encouraged to elucidate how biodiversity might influence ecosystem functioning. This may be achieved through the detailed scrutiny of large spatial/long temporal scale data sets, such as long‐term forest inventories. Forest management strategies based on B→EF have strong potential for augmenting the effectiveness of the roles of forests in the mitigation of climate change impacts on ecosystem functioning.     

Alteration of the phenology of leaf senescence and fall in winter deciduous species by climate change: effects on nutrient proficiency

Leaf senescence in winter deciduous species signals the transition from the active to the dormant stage. The purpose of leaf senescence is the recovery of nutrients before the leaves fall. Photoperiod and temperature are the main cues controlling leaf senescence in winter deciduous species, with water stress imposing an additional influence. Photoperiod exerts a strict control on leaf senescence at latitudes where winters are severe and temperature gains importance in the regulation as winters become less severe. On average, climatic warming will delay and drought will advance leaf senescence, but at varying degrees depending on the species. Warming and drought thus have opposite effects on the phenology of leaf senescence, and the impact of climate change will therefore depend on the relative importance of each factor in specific regions. Warming is not expected to have a strong impact on nutrient proficiency although a slower speed of leaf senescence induced by warming could facilitate a more efficient nutrient resorption. Nutrient resorption is less efficient when the leaves senesce prematurely as a consequence of water stress. The overall effects of climate change on nutrient resorption will depend on the contrasting effects of warming and drought. Changes in nutrient resorption and proficiency will impact production in the following year, at least in early spring, because the construction of new foliage relies almost exclusively on nutrients resorbed from foliage during the preceding leaf fall. Changes in the phenology of leaf senescence will thus impact carbon uptake, but also ecosystem nutrient cycling, especially if the changes are consequence of water stress.