长江源区高山嵩草物候与生物量动态及其对气候变化的响应

利用青藏高原腹地江河源头的曲麻莱县气象站1994～2004年观测的高寒嵩草草甸优势种高山嵩草的生育期、高度、产量等指标与同期气象资料,通过定量分析研究较长时段的物候及生物量变化特征,以揭示其对气候变化的响应.结果表明:(1)高山嵩草返青期和开花期的变化总体均呈"W"形,在区域气候变暖背景下,植物物候表现为返青期提前,开花期和枯黄期推迟,整个生长季延长.(2)高山嵩草的生物量变化在10年间呈明显的波动趋势,各月最高生物量均出现在1999年,最低生物量出现在1994年.(3)高山嵩草物候期与生长季各月气温呈显著的正相关关系(P<0.001),与月降水量呈弱正相关关系(P>0.05);月均气温成为本区草本植物发芽生长的先决条件,且6～8月气温与植物萌动的相关性最大.(4)生长季6～8月高山嵩草生物量鲜重和干重均与月均气温呈显著正相关,鲜重仅与月降水量呈显著正相关关系,在青藏高原的高寒区,温度比降水对植物产量的影响更大.     

高山嵩草气孔导度对环境因子的响应模拟

以青藏高原地区高寒草甸常见建群种高山嵩草(Kobresia pygmaea)为研究对象,利用光合测定仪于2018年7—9月测定高山嵩草气体交换参数与环境因子的日变化值,利用土壤水分测量仪及环刀法、透膜法测得0—10 cm土壤水分数据作为模型变量。首先,采用3种气孔导度模型对高山嵩草气孔导度进行拟合和检验,其次,用Jarvis模型、Leuning模型和Gao模型对高山嵩草气孔导度在不同月份(7,8,9月)日变化的模拟结果进行分析,最后,总结并讨论了气孔导度对3个主要环境因子(光合有效辐射(Photosynthetically active radiation, PAR)、气温(T_air)、饱和水汽压差(Vapor pressure deficit, VPD))的响应特征。主要研究结果:(1)3个气孔导度模型都可以较好地模拟高山嵩草的气孔导度变化,Leuning模型表现最好(R²=0.726),Jarvis模型次之(R²=0.659),Gao模型准确度最低(R²=0.624)。(2)高山嵩草叶片气孔导度...     

高原鼠兔有效洞穴密度对高山嵩草群落及其主要种群空间分布特征的影响

高原鼠兔(Ochotona curzoniae)有效洞穴密度变化会引发高山嵩草(Kobresia pygmaea)草甸植物群落及其种群分布格局发生变化。采用野外调查法研究了高原鼠兔有效洞穴密度对高山嵩草群落特征及其主要种群分布格局的影响。结果表明:随高原鼠兔有效洞穴密度增加,高山嵩草草甸植物群落的优势种没发生明显变化,部分伴生种发生更替;高度、盖度、多样性指数和均匀度指数呈现降低态势;地上生物量和丰富度指数变化不明显;高山嵩草和矮火绒草(Leontopodium nanum)种群的盖度、密度以及生物量均呈现降低态势,而达乌里秦艽(Gentiana dahurica)和小花草玉梅(Anemone rivularis var.flore-minors)种群的盖度、密度和生物量呈增加趋势。高山嵩草和矮火绒草的种群分布格局从8个/625m2和14个/625m2的聚集分布分别变为34个/625m2时的均匀分布和随机分布,达乌里秦艽和小花草玉梅种群从8个/625m2和14个/625m2的随机分布变为23个/625m2和34个/625m2时的聚集分布,这说明高原鼠兔有效洞穴密度变化改变了高山嵩草群落的特征和主要植物种群的空间分布格局,而对应群落特征和种群分布格局改变的有效洞穴密度为14个/625m2和23个/625m2。     

2种嵩草属植物形态特征对生境干旱化的响应

该研究于西藏自治区东南部的东达山高山草甸沿生境干旱化梯度设置10个样地,采用线性回归方法分析优势种高山嵩草(Kobresia pygmaea)和矮生嵩草(K.humilis)叶片数量与其它构件数值(分株数量、茎基直径、根系数量和根系长度)之间的关系,并采用线性回归斜率测度分株功效、叶片萌生能力、根系分生功效和根系伸长功效,以探讨嵩草的分株能力、茎基生长和根系生长对生境干旱化过程的响应机制。结果显示:(1)2种嵩草植物的叶片数量与4种构件数值均为显著线性正相关关系。(2)随着生境干旱化程度增加,高山嵩草分株数量增加,矮生嵩草分株受干旱抑制程度高,分株数量呈下降趋势;2种嵩草分株功效下降,即单株叶片数量因干旱化程度增加而减少。(3)2种嵩草的茎基直径、叶片萌生能力随着生境干旱化程度增加而下降;高山嵩草叶片萌生能力的变化与生境干旱化梯度一致,具有连续性;矮生嵩草叶片萌生能力对生境干旱化的适应性弱,干旱到一定程度发生骤降。(4)随着生境干旱化,高山嵩草的根系数量和长度均增加,矮生嵩草根系长度增加,但数量却无规律变化;2种嵩草的根系分生功效和根系伸长功效均下降。研究表明:嵩草属植物分布的最适宜生境为表面稍有积水的沼生生境。为适应生境干旱化,高山嵩草降低叶片萌生率以减少蒸腾作用,增加分株数量以增强对空间的占有能力,并增加根系数量和长度来提高对土壤水分的吸收能力,因此对干旱有较强适应性且分布范围广;矮生嵩草只通过降低叶片萌生率,增加根系长度响应干旱化生境,其分布范围较窄。     

若尔盖高原沙化沼泽区植物群落物种组成及其驱动因素

若尔盖高原沼泽区土壤沙化过程中植物群落的变化及其成因机制是亟待解决的区域关键科学问题之一。在若尔盖高原红原县沙化沼泽区,研究了土壤沙化梯度上植物群落种类组成、物种多样性、地上生物量的变化及其与土壤湿度、容重和孔隙度的关系。随着土壤沙化程度增加,群落特征种组替代顺序为华扁穗草(Blysmus sinocompressus)种组→高山嵩草(Kobresia pygmaea)种组→四川嵩草(K.setchwanensi)种组→黑褐穗苔草(Carex atrofusca subsp.minor)种组→粗壮嵩草(Kobresia robusta)种组;植物种数(单位样方物种丰富度和总种数)和地上生物量均表现为单峰变化格局,但植物种数峰值出现在极轻度沙化的高山嵩草群落,地上生物量峰值出现在中度沙化的四川嵩草-黑褐穗苔草群落;β多样性表现为"U"型变化格局,最高值出现在未沙化的华扁穗草群落,最低值出现在极轻度沙化的高山嵩草群落;土壤湿度对植物种多度影响最大,可解释总方差的24.8%,其次为土壤容重,可解释总方差的1.4%;沙化过程中土壤湿度是影响植物群落生态分布和种类构成的关键因素之一,尤其影响群落...     

Small-scale switch in cover–perimeter relationships of patches indicates shift of dominant species during grassland degradation

草地生态系统普遍经受着全球气候变化和不可持续的土地利用方式的胁迫,这些环境胁迫往往引起草地生态系统状态在多稳态间 跃变,甚至诱发灾变性跃变,直至荒漠化的发生。研究表明,植被斑块的空间构型可以指示较大空间尺度的生态系统状态跃变。本文展示了小尺度 斑块构型如何指示草地生态系统的状态跃变。在高寒草甸生态系统选取7种草甸群落类型作为研究对象,这7种类型的草甸群落 具有高寒草甸退化过程不同阶段的典型群落特征。在每一种群落类型内,采用样带方法进行植被斑块调查,沿样带采用邻接格子样方法 记录样方内的斑块数量、斑块周长和斑块面积,并测定了每个斑块内的物种多度。研究结果表明,沿着草甸退化阶段七种 类型的草甸群落(即未退化的禾草-嵩草群落、轻度退化的高山嵩草-禾草群落、中度退化的高山嵩草群落、重度退化草皮开裂的高 山嵩草群落、侵蚀期的高山嵩草-杂草群落、严重侵蚀期的杂草-高山嵩草群落、零星杂草的黑土滩裸地),斑块数量和斑块周长随斑块面积的下降呈现先增加的变化趋势,而后,随斑块面积的急剧降低呈下降的趋势。斑块数量、周长与面积关系变化的拐点出现在植被总面积约占68%的状态处。在这一拐点附近,斑块上的植物组成呈现出高山嵩草或杂草占优势两种状态往复波动的不稳定状态,表明草地状态处于向荒漠化裸地跃变的边缘。本研究展示了如何基于小尺度植被斑块空间构型为草地退化、状态跃变提供数量化的早期预警信号,有望为嵩草草甸退化至黑土滩裸地的早期预警提供判定指标。     

3-吲哚乙酸对铜钱草去除氮磷的影响

迅猛发展的水产养殖业在带动经济发展的同时，也引发了一定的环境问题，如养殖废水任意排放，引起河流、湖泊、水库等水体富营养化。水生植物可通过植物根系的吸附、转化、富集等作用降低水中氮磷的浓度。目前已有一些利用铜钱草吸收氮磷营养盐进而改良水质的研究，但主要集中在对不同水质的氮磷吸收率方面。基于此，文章将研究铜钱草在不同浓度吲哚乙酸的作用下对氮、磷的去除效率和蛋白质生理指标的变化，以期为铜钱草作为废水处理植物的应用提供参考。     

不同退化程度高寒草甸高山嵩草的构件变化

对不同退化程度下高山嵩草的构件变化进行了初步研究.结果表明:随着退化程度的加剧,高山嵩草无性系复合主分蘖数、单分蘖数以及构件的叶片数均明显下降,退化程度加剧不利于高山嵩草的营养生长;除极度退化草地外,其余样地各构件的生物量大小依次是:根茎>根>叶>秆,并且随退化程度的加剧,各构件生物量显著下降;重度退化草地中,莎草科植物生殖枝数占植物群落总生殖枝的比例最大,每个有效生殖枝平均产生3.4粒种子.     

蛋白质组研究中的化学"探针"

随着蛋白质组学概念的提出以及诸如血浆蛋白质组等有影响力的计划开展, 蛋白质组研究迅速发展起来, 这门基于分析化学和物理化学的领域也逐渐为广大生物学家所关注, 同时也相应地在细胞生物学、生物化学等领域的研究中崭露头角。蛋白质表达量的变化以及各种各样的修饰无不反映出机体对环境变化的应激和自身功能的需要。因此, 定量蛋白质组和修饰化的蛋白质组成为了目前蛋白质组研究的重要领域之一。文章着重从采用化学标记实现定量和修饰化研究这个角度来介绍近些年来在这方面取得的进展, 希望对生物学领域的研究有所借鉴。     

高原鼠兔有效洞穴密度对青藏高原高寒草甸群落植物生态位的影响

高原鼠兔(Ochotona curzoniae)有效洞穴密度扩增会引起高寒草甸植物群落组分的变化。采用野外调查方法研究了高原鼠兔有效洞穴密度对青藏高原高寒草甸群落植物生态位的影响。结果表明:随高原鼠兔有效洞穴密度增加,样方内植物种数逐渐增多,但主要是毒杂草,有效洞穴密度超过15个/625m2时豆科植物出现。优势种高山嵩草(Kobresia pygmaea)的重要值随有效洞穴密度增加呈单峰曲线,峰值出现在15个/625m2。有效洞穴密度低于或等于21个/625m2时,优势种高山嵩草生态位宽度位居第一,但有效洞穴密度达31个/625m2时,其生态位宽度降至并列第三,低于小米草(Euphrasia pectinata)和钝裂银莲花(Anemone obtusiloba),等同于小花草玉梅(Anemone rivularis)和乳浆大戟(Euphorbia esula)。有效洞穴密度分别为10个/625m2,15个/625m2,21个/625m2,31个/625m2时,与高山嵩草生态位重叠度最大的物种分别为莓叶委陵菜(Potentilla fragarioides)、圆叶筋骨草(Ajuga ovalifolia)、乳浆大戟、条叶垂头菊(Cremanthodium lineare)。有效洞穴密度为10个/625m2时,高山嵩草是群落内种群间竞争最激烈的一方,但有效洞穴密度超过或等于15个/625m2时,高山嵩草则退出了种群间竞争最激烈的双方。说明高原鼠兔有效密度增加严重影响了高寒草甸群落组分、重要值、生态位宽度和生态位重叠值。     

青藏高原高寒矮嵩草草甸碳增汇潜力估测方法

以广布于青藏高原的高寒矮嵩草草甸为研究对象,研究了草甸碳储存的场所、碳库容量随草甸演替的变化过程及其碳增汇潜力的空间分布格局,同时探讨了高寒草甸碳增汇潜力估测的困惑与解决方法。结果表明,高寒草甸生态系统碳增汇潜力空间分布格局差异极大,主要受到土层厚度和草地演化进程的影响。高寒草甸碳主要贮存于草毡表层,其增汇潜力在于退化草地草毡表层的恢复与重建。保持适宜厚度的草毡表层是协调高寒草甸生产与碳生态服务功能的关键。随着退化高寒草甸的恢复,土壤容重呈现下降趋势,计算其系统碳增汇潜力,需要用根土体积比进行土层深度的校正。高寒草甸具有较大的固碳潜力,但其潜力的发挥受到气候和草地恢复与管理措施的影响,比较漫长。     

人类活动对青藏高原高寒矮嵩草草甸碳过程的影响

随着人类活动干扰(放牧)的增加,青藏高原高寒嵩草甸的退化演替过程依次为禾草-矮嵩草群落、矮嵩草群落、小嵩草群落和杂类草-黑土滩4个阶。其中小嵩草群落又可划分为草毡表层加厚、开裂与塌陷3个子阶段。采用时空转换的方法,研究了人类活动对青藏高原高寒矮嵩草草甸碳过程的影响。结果表明,随着人类干扰强度的增加,植物群落地上部分有机碳储量逐渐降低,由禾草-矮嵩草群落的(134.7±17.3)gC/m2逐渐降低到杂类草-黑土滩次生裸地(18.96±6.18)gC·m-2。土壤、植物地下部分有机碳贮量呈单峰曲线变化,草毡表层开裂子阶段最高,分别为(49.7±0.83)gC·kg-1和(3596.7±179.8)gC·m-2。;杂类草-黑土滩阶段最低,分别为(19.2±1.13)gC·kg-1和(121.6±6.1)gC·m-2。受植物地下部贮碳的影响,土壤-植被系统呈现逐渐降低的变化特征。随人类活动干扰的加强,高寒嵩草草地植物有机碳地下/地上分配比发生巨大改变,草地草毡表层厚度不高于4.3cm是保证草地生产与生态服功能双赢的重要指标。     

Leaf δ13C reflects ecosystem patterns and responses of alpine plants to the environments on the Tibetan Plateau

Leaf δ¹³C is an indicator of water-use efficiency and provides useful information on the carbon and water balance of plants over longer periods. Variation in leaf δ¹³C between or within species is determined by plant physiological characteristics and environmental factors. We hypothesized that variation in leaf δ¹³C values among dominant species reflected ecosystem patterns controlled by large-scale environmental gradients, and that within-species variation indicates plant adaptability to environmental conditions. To test these hypotheses, we collected leaves of dominant species from six ecosystems across a horizontal vegetation transect on the Tibetan Plateau, as well as leaves of Kobresia pygmaea (herbaceous) throughout its distribution and leaves of two coniferous tree species ( Picea crassifolia, Abies fabri ) along an elevation gradient throughout their distribution in the Qilian Mountains and Gongga Mountains, respectively. Leaf δ¹³C of dominant species in the six ecosystems differed significantly, with values for evergreen coniferous13C values of the dominant species and of K. pygmaea were negatively correlated with annual precipitation along a water gradient, but leaf δ¹³C of A. fabri was not significantly correlated with precipitation in habitats without water-stress. This confirms that variation of δ¹³C between or within species reflects plant responses to environmental conditions. Leaf δ¹³C of the dominant species also reflected water patterns on the Tibetan Plateau, providing evidence that precipitation plays a primary role in controlling ecosystem changes from southeast to northwest on the Tibetan Plateau.     

Mixed reproduction strategy and polyploidy facilitate dominance of Kobresia pygmaea on the Tibetan Plateau

Aims The sedge Kobresia pygmaea is the dominant species of high-altitude pastures in Tibet, and it is the most important source of forage in animal husbandry. We present the first comprehensive reproduction study for this perennial key species that adopts a molecular approach and tests how sexual and vegetative reproduction, as well as ploidy, relate to survival and dominance under harsh conditions.Methods We assessed inflorescence numbers of K. pygmaea across two Tibetan alpine pastures with differing grazing regimes. Germination was tested in untreated diaspores and then following mechanical and chemical scarification. In a 4-year experiment, we assessed diaspore viability and seed bank formation. Using eight microsatellite markers, we recorded multilocus genotypes in hierarchical grids and measured their ploidy using flow cytometry. Adjusted analysis of variance models were used to analyse data on sexual reproduction, while the complement of the Simpson index and the Shannon diversity index were used to characterize the spatial distribution of multilocus genotypes and clonal richness.Important findings Inflorescence production was high and differed significantly between years (2010: 617±460 SD; 2012: 2015±1213 SD) but not between grazing regimes. Diaspore viability was high (94%) and gradually decreased after 3 and 4 years of storage in the soil. Diaspores not exposed to further scarification failed to germinate, while mechanical and chemical (H 2 SO 4) scarification increased germination to 9 and 44%, respectively. Clonal diversity was high, although in situ germination was rarely observed. Multilocus genotypes intersected and covered a mean area of 0.74 m 2. Most individuals were found to be tetraploid, with only 0.8% of all ramets being triploid. We conclude that K. pygmaea survives on the Tibetan Plateau by employing a mixed reproduction strategy involving both sexual and clonal propagation. The species' adaptability and dominance is further facilitated by its polyploidy. As pasture restoration using diaspores would be difficult, existing Kobresia pasture should be managed more sustainably.     

青藏高原小型哺乳动物的生理生态学研究：从个体到生态系统

本文回顾了青藏高原小型哺乳动物生理生态学研究取得的主要成就,包括能量代谢特征与环境适应性、适应性产热与体温调节、能量平衡与体重调节、生理极限值和种群能流估算等,总结了近年一些新兴领域的最新进展,包括双标记水方法测定能量消耗、肠道菌群的功能、地理生理学、种群生理学、植物次生代谢产物及其生理功能等。有些工作引领了中国动物生理生态学的发展,如生态能量学、适应性产热和生理适应等。本文对未来需要发展的领域和深入方面提出了建议,希望能建立青藏高原小型哺乳动物生理生态学学科体系。     

Comparative Physiological and Molecular Analyses of Intraspecific Differences of Stipa purpurea (Poaceae) Response to Drought

Stipa purpurea Griseb, the dominant species of alpine steppe, is widely distributed across the large precipitation gradient on the Tibetan Plateau. It is possible that because of local adaptation, Spurpurea populations from different habitats along this precipitation gradient respond differently to drought, which may affect their responses to climate change. To explore the problem, in the present study, we investigated the physiological and molecular response of Spurpurea seedlings from two different populations (Pulan & Cuoqin) to 14 d drought stress and subsequent recovery. The results showed that the relative water content, chlorophyll fluorescence, content of osmoticant proline and malondialdehyde (indicator of oxidative stress), accumulation of reactive oxygen species, antioxidant enzyme activities and the expression of drought related genes all changed under drought treatment and went back to the control levels in the subsequent recovery in plants from Pulan. However, these patterns were quite different in plants from Cuoqin, in which these traits changed by inconsistent degrees and did not return to pretreatment levels after rewatering. The results demonstrated that the plants from Pulan had greater resistance to drought stress compared with those from Cuoqin, which had a larger mortality rate ultimately. Combating the differences of offspring in response to drought and the habitat distribution of parents, we considered that genetic basis has been obtained in response to precipitation difference among Spurpurea populations. The results help to understand the adaptation and evolution of Spurpurea to the special environment and the effect of climate change to this botanical system.     

Current challenges in distinguishing climatic and anthropogenic contributions to alpine grassland variation on the Tibetan Plateau

Quantifying the impact of climate change and human activities on grassland dynamics is an essential step for developing sustainable grassland ecosystem management strategies. However, the direction and magnitude of climate change and human activities in driving alpine grassland dynamic over the Tibetan Plateau remain under debates. Here, we systematically reviewed the relevant studies on the methods, main conclusions, and causes for the inconsistency in distinguishing the respective contribution of climatic and anthropogenic forces to alpine grassland dynamic. Both manipulative experiments and traditional statistical analysis show that climate warming increase biomass in alpine meadows and decrease in alpine steppes, while both alpine steppes and meadows benefit from an increase in precipitation or soil moisture. Overgrazing is a major factor for the degradation of alpine grassland in local areas with high level of human activity intensity. However, across the entire Tibetan Plateau and its subregions, four views characterize the remaining controversies: alpine grassland changes are primarily due to (1) climatic force, (2) nonclimatic force, (3) combination of anthropogenic and climatic force, or (4) alternation of anthropogenic and climatic force. Furthermore, these views also show spatial inconsistencies. Differences on the source and quality of remote sensing products, the structure and parameter of models, and overlooking the spatiotemporal heterogeneity of human activity intensity contribute to current disagreements. In this review, we highlight the necessity for taking the spatiotemporal heterogeneity of human activity intensity into account in the models of attribution assessment, and the importance for accurate validation of climatic and anthropogenic contribution to alpine grassland variation at multiple scales for future studies.     

Comparative analyses of leaf anatomy of dicotyledonous species in Tibetan and Inner Mongolian grasslands

Knowledge of the leaf anatomy of grassland plants is crucial for understanding how these plants adapt to the environment. Tibetan alpine grasslands and Inner Mongolian temperate grasslands are two major grassland types in northern China. Tibetan alpine grasslands occur in high-altitude regions where the low temperatures limit plant growth. Inner Mongolian temperate grasslands are found in arid regions where moisture is the limiting factor. Few comparative studies concerning the leaf anatomy of grassland plants of the Tibetan Plateau and Inner Mongolian Plateau have been conducted. We examined leaf characteristics at 71 sites and among 65 species, across the alpine grasslands of the Tibetan Plateau and the temperate grasslands of the Inner Mongolian Plateau. We compared the leaf structures of plants with different life forms and taxonomies, and their adaptation to arid or cold environments. We explored relationships among leaf features and the effects of climatic factors (i.e., growing season temperature and precipitation) on leaf characteristics. Our results showed that (i) there were significant differences in leaf anatomy between Tibetan alpine and Inner Mongolian temperate grasslands. Except for mesophyll cell density, the values obtained for thickness of leaf tissue, surface area and volume of mesophyll cells were larger on the Tibetan Plateau than on the Inner Mongolian Plateau. (ii) Within the same family or genus, leaf anatomy showed significant differences between two regions, and trends were consistent with those of whole species. (iii) Leaf anatomy of woody and herbaceous plants also showed significant differences between the regions. Except for mesophyll cell density, the values obtained for the thickness of leaf tissue, and the surface area and volume of mesophyll cells were larger in herbaceous than in woody plants. (iv) Leaf anatomical traits changed accordingly. Total leaf thickness, thicknesses of lower and upper epidermal cells, and surface area and volume of mesophyll cells were positively correlated, while mesophyll cell density was negatively associated with those traits. (v) Growing season temperature had stronger effects on leaf anatomy than growing season precipitation. Although the communities in Tibetan and Inner Mongolian grasslands were similar in appearance, leaf anatomy differed; this was probably due to the combined effects of evolutionary adaptation of plants to environment and environmental stress induced by climatic factors.     

青藏高原高寒草甸生态系统净二氧化碳交换量特征

高寒草甸是青藏高原广泛分布的植被类型之一,面积约120万km2,地处青藏高原腹地的当雄草原站即位于该类植被的典型分布区。以2003年8~10月中旬在该站用涡度相关法连续观测的CO2通量数据资料为基础,分析了高寒草甸生态系统8~10月份净二氧化碳交换量(NEE)的日变化规律,及其与光合有效辐射、降水、温度等环境因子之间的关系。结果表明,8~10月份的日均NEE有明显的日变化,表现为单峰型,通常在地方时11:00~12:00左右达到碳吸收的最大值,平均为-0.2680mgCO2/(m2·s)(-6.0800μmolCO2/(m2·s))。白天的NEE与光合有效辐射之间符合很好的直角双曲线关系,表观量子产额平均为0.0203μmolCO2/μmolPAR,表观最大光合速率平均为9.7411μmolCO2/(m2·s)。夜晚的NEE与5cm地温有很好的指数函数关系。