青藏高原东缘高山林线乔木种群生态特征研究

青藏高原东部分布着世界最高的林线，该区域也是由欧亚北温带物种形成的林线的南界。在大面积野外踏勘的基础上，选择青藏高原东部具有典型高山林线分布的三个地点（滇西北白马雪山、川西北鹧鸪山及岷江源地区）作为研究区，从种群的结构、生存特征、分布格局及分形特征等方面对青藏高原东缘高山林线乔木种群生态学特征进行了研究，并在此基础上探讨了人类活动对林线种群生态特征及林线格局的影响。结果表明，林线区乔木树种多以单种群形式存在，林线区群落结构简单，乔木层多为单一树种组成，其生长型较之郁闭林发生了急剧的变化：树木高度急剧下降，而发展多茎多分枝的生长型。生长型的转变是高山林线乔木对恶劣自然条件的形态适应。 研究发现，在青藏高原东缘，阴坡林线乔木主要是冷杉（Abies spp.），阳坡主要由圆柏（Sabina spp.）组成，少数地方还有云杉（Picea spp.）。阴坡乔木种群结构多表现为增长型，幼苗和幼树在种群中占较大比重，种群潜在自然更新能力较强，但幼龄个体死亡率非常高，存活曲线多接近Deevey-Ⅲ型；阳坡乔木种群幼苗个体数极少，幼树相对增加。野外调查表明，人为活动较频繁的阳坡林线区幼苗数量极少甚至缺失，而受人为活动干扰较小的样地中幼苗和幼树数量明显增多，从一个侧面说明放牧等人类活动可能对林线种群的更新带来较大影响，而对卡卡沟围栏内外的样地分析也进一步证明了这一结果。 所研究林线乔木种群各龄级的空间格局在不同尺度上表现为聚集、随机和均匀分布，以聚集分布为主；各龄级在不同尺度上表现出显著的相关性，幼苗通常与另外两个龄级的关联性较密切。各龄级间显著的相关性表明不同龄级个体在空间交错分布，有利于对各种资源的充分利用，对种群的生存和发展非常有利，反映了高山生态系统恶劣生境中种群的一种适应对策。 林线乔木种群各龄级分布格局的计盒维数有差别，林线种群的计盒维数总是小于郁闭林种群的计盒维数。另外，郁闭林各龄级计盒维数通常也高于林线各龄级，表明不同海拔或者不同群落类型中的乔木树种具有不同的水平空间占据能力。林线区种群分布格局的计盒维数都很低，占据现实水平空间的程度较低，具有相对较高的生态间隙维，其潜在占据空间的能力较高，群落还可提供给种群的最大空间限度较大，但实际上由于受群落中种内、种间的竞争及林线区恶劣的生态环境条件的限制，其潜在空间占据能力可能难以表现出来。 青藏高原东缘高海拔地带以季节性游牧为主要的资源利用和生产方式，阳坡森林郁闭度低于阴坡，灌丛数量和种类较阴坡少，融雪早且积雪时间短，所以阳坡包括高山林线区成为当地牧民游牧路线的必经之地。牲畜的践踏、啃食使得幼龄乔木树种个体数量大大减少，严重阻碍了林线乔木种群的自然更新，同时种群占据空间的能力也明显降低。因此可以认为，在青藏高原东部地区，山地游牧等人为干扰叠加于恶劣的自然条件，阳坡林线的自然更新潜力受到抑制，其生存状态较之阴坡林线显著恶化，并可使阳坡林线高度逐渐降低。高山林线区森林一旦破坏在短时间内很难有效更新和恢复，因此，对于处于恶劣高山生境中的乔木种群应加强保护，同时适度控制人为干扰强度和幅度以减少其直接和间接破坏，防止阳坡林线退化并促进高山生态系统的自然恢复。 Eastern Qinghai-Tibetan Plateau has the highest timberline of the world. On the basis of field surveys and literature reviews, three typical alpine timberlines were chosen for in-depth studies, i.e., Baima Snow Mountain in northwest Yunnan, Zhegu Mountain and the waterhead area of Minjiang River in west Sichuan. Using the methodologis of population ecology, we analyzed the population structure, survival characteristics, spatial point patterns and fractal dimensions of the timberline tree populations and discussed the impacts of grazing on the structure and spatial pattern of alpine timberline. Compared with closed forests, the community structure of timberline is simpler, usually with one or two species constituting the tree layer. Differences also exist in the growth forms: the trees were significantly shorter with more stems and branches, reflecting morphological adaptation of trees to the severe conditions at timberline. In the eastern Qinghai-Tibetan Plateau, Abies spp. often formed alpine timberline in the north-facing slope while Sabina spp. and sometimes Picea spp. in the south- facing slope. The population structures of north-facing slope showed an increasing trend, with numerous seedlings and saplings. However, the survival curves tend to follow Deevy-III because of high dead ratio of young individuals. There are only few seedlings in the south-facing slope with heavy grazing, demonstrating that human disturbance may prevent regeneration at alpine timberline, which was confirmed by comparisons between fenced enclosures and control plots in the Kaka Valley. Depending on the spatial scales on consideration, the individuals of different age-classes showed clumping, random or even distribution, but mostly with clumping distribution. At all scales, individuals in different age-classes were all significantly correlated with each other while the seedlings were usually more correlated to two other age classes. This high degree of correlation among different age classes indicates that individuals of different age classes are spatially interlocked with each other, which helps sufficient utilization of various resources and is conducive to the survival and development of population. It is another adaptation strategy for trees at the severe environment. The spatial patterns of different age classes had different box dimension. In general, the box dimensions of total individuals and each age class at timberline are always smaller than that of closed forests, suggesting that space occupation capacity is not the same for populations at different altitude or in different communities. Populations on both the south- and the north-facing slopes had a very low box dimensions (far away from the max., 2), however, the lower the box dimension, the bigger the potential space provided by community. In fact, because of inner- and inter- competition as well as the severe conditions at timberline, this kind of potential ability can hardly be realized. Mountain pastoralism is the major type of as well as the only most effective way of resource uses in the high elevation regions of the eastern Qinghai-Tibetan Plateau. Due to lower canopy cover, less bushes and short snow-cover time, south-facing slopes became the favorite pastures. Damages from livestock through tramping, browsing and others have greatly reduced the number of young individuals. As a result, the potential of timberline trees to regenerate and their ability to occupy more space are greatly inhitibted. We conclude that human disturbances (mountain pastoralism) as well as harsh environmental conditions co-worked to inhibit the regeneration of tree populations in the south-facing slope and made south slopes more difficult than the north-facing slopes for trees to survive and develop, resulting a gradual retreat of timberline in the north-facing slopes. Forests at alpine timberline are susceptible to disturbance and difficult to regenerate and restore once damaged and controlling human disturbances is important for protecting the forest ecosystems at the timberline area.

克隆植物东方草莓对异质生境的适应对策

克隆植物被认为比非克隆植物更宜于利用异质性环境。在复杂的空间异质环境中，克隆植物可能形成了各种有效利用环境异质性的适应对策。对于克隆植物适应机制的研究，前人已做了大量的工作，特别是从形态和生物量分配等方面对简单异质生境下克隆植物的克隆整合和克隆分工进行了详细的研究。本研究以分布广泛的克隆植物东方草莓（Fragaria orientalis）作为研究对象，应用野外调查和实验生态学方法，采用多对比度单资源模型和不同向双资源模型，从形态和生理生态的角度，研究复杂异质生境下克隆植物的整合和分工及其耗益问题，分析不同类型的生境对克隆植物整合和分工的修饰作用，进而探讨克隆植物对异质生境的适应策略。克隆构型和分株种群特征是植物克隆生长及其生态适应对策研究的基本内容。本文通过野外调查，研究在不同光照条件下东方草莓克隆构型、分株种群特征以及点分布格局。结果表明：东方草莓的克隆构型随光照发生相应的变化，低光照下其匍匐茎节间长和分枝角度均增大而分枝强度减小；随光照减弱，东方草莓分株种群的生物量、根冠比和分株种群密度显著降低；不同光照下东方草莓分株均以随机分布为主但不同尺度下有所差异，其分布格局强度依次为旷地<林缘<林下。结合克隆植物对资源的利用对策，探讨了克隆构型和分株种群特征以及分布格局随环境条件变化的生态适应意义。不同生境斑块条件下克隆植物可能采取不同的适应对策。采用盆栽实验，研究不同水分对比度下克隆整合及其生理生态特征，并对单向和交互资源中东方草莓的克隆整合做了对比研究。结果显示：高的水分对比度能够促进东方草莓的克隆整合，并能刺激相连分株增加光合作用，东方草莓体内的氧化—抗氧化系统也II随对比度做出相应的反应。耗-益分析表明胁迫分株的受益是以供给分株的损耗为代价的，但从克隆片段总体来说是受益的。单向资源中东方草莓生长的绝对值高于交互资源，但耗-益分析表明生长于交互资源下东方草莓的克隆整合获益大于生长于单向资源下东方草莓的克隆整合获益。长期生长于特定生境的克隆植物，在进化过程中其克隆整合和克隆分工在对资源异质性的适应策略方面可能有所侧重。采用盆栽实验对来自不同海拔梯度的东方草莓的克隆整合和克隆分工对异质资源的适应对策进行了研究。实验结果表明，来自高海拔的东方草莓可塑性较差。来自两个海拔的东方草莓对切断匍匐茎的表现有所差异，总体上切断匍匐茎对来自高海拔的东方草莓影响更大些。另外，来自高海拔的东方草莓表现出更高的克隆分工。IIIClonal plants are known to be more suitable for the habitats of heterogeneousresources than nonclonal plants, perhaps due to their well developed adaptivestrategies to environmental heterogeneity. Many studies have been done on theadaptive mechanisms of clonal plants, especially on the clonal integration anddivision of labor with morphology and biomass allocation under simpleheterogeneous habitats. Based on field surveys, laboratory experiments, multi-contrastunidirectional resource model and reciprocal resource model, Fragaria orientalis, aRosaceae stoloniferous herb that widely distributes in China, was used to study thisplant’s morphological and physiological responses to complicated heterogeneoushabitats in terms of its clonal integration, division of labor and cost-benefit, as well astheir modifications by different habitats, so as to better understand the adaptivestrategies of clonal plants under heterogeneous environments.Clonal architecture and ramet population characteristics are of the major concernin the studies on growth and adaptive strategies of clonal plants. Clonal architecture,ramet population characteristics and spatial point pattern of F. orientalis underdifferent light intensity were studied with field observations. The results showed that,clonal architecture changed with light availability: Internode-lengths and branchangels of stolons were larger while branch intensities were smaller under lower lightintensity than those under higher light intensity; Biomass of ramet population,root-shoot ratio and density of ramet population decreased significantly with reduce oflight intensity; Under all light intensities, spatial pattern of ramets was mainlyrandomly distributed but it changed with different scales, with pattern intensity as:open space < forest edge < understory. Adaptation significance of the clonal architecture, the ramet population characteristics and the spatial pattern changing withdifferent environments was discussed according to these results.Clonal plants may take different adaptive strategies under different patches. Withpot culture, clonal integration and physiological parameters of F. orientalis underdifferent water contrasts were studied, and clonal integration under unilateralresources and reciprocal resources were also compared. The results suggested that,high water contrast improve the clonal integration of F. orientalis and increase thephotosynthesis of connected ramets. Oxidative and antioxidative system of F.orientalis also responded with changing water contrasts. According to cost-benefitanalysis, the drought-stressed ramets obtained benefits from the connectedwell-watered ramets, and as a whole, the clonal fragment could also get benefits.Growth of F. orientalis in homogeneous resources was better than that inheterogeneous resources, but the whole plant got more benefit through clonalintegration in heterogeneous resources than in homogeneous resources.Pot culture experiments were also used to study the adaptive strategies inutilizing heterogeneous resources by the plant populations from different altitudes.The results showed that, F. orientalis from alpine zones were shorter and lessexpanded with poorer clonal plasticity than those from middle mountains. F.orientalis from two different altitudes showed different responses to stolon severing,and as a whole, stolon severing had more influence on F. orientalis from alpine zones.In addition, F. orientalis from alpine zones exhibited higher division of labor, whichsuggested that clonal plants from different habitats develop their own adaptivemechanisms in their clonal integration and division of labor in response toenvironmental heterogeneity.

Did an extensive forest ever develop on the Chinese Loess Plateau during the past 130 ka?: a test using soil carbon isotopic signatures

IEECAS SKLLQG

Nitrogen-15 signals of leaf-litter-soil continuum as a possible indicator of ecosystem nitrogen saturation by forest succession and N loads

National Key Research and Development Program [2010CB833502]; National Natural Science Foundation of China [30600071, 40601097, 30590381]; Chinese Academy of Sciences [KZCX2-YW-432, O7V70080SZ, LENOM07LS-01]; GUCAS [O85101PM03]

Impact of cloudiness on net ecosystem exchange of carbon dioxide in different types of forest ecosystems in China

Asia 3 Foresight Program [30721140307]; National Key Research and Development Program [2010CB833500]; National Natural Science Foundation of China [30590381, 30900198];

Effects of multiple environmental factors on CO2 emission and CH4 uptake from old-growth forest soils

National Key Research and Development Program [2010CB833502]; National Natural Science Foundation of China [30600071, 40601097, 30590381, 30721140307]; Knowledge Innovation Project of the Chinese Academy of Sciences [KZCX2-YW-432, O7V70080SZ, LENOM07LS-01

Use of Ecosystem Flux Data and a Simulation Model to Examine Seasonal Drought Effects on a Subtropical Coniferous Forest

National Natural Science Foundation of China (NSFC) [30670384, 30590381]

Environmental controls over carbon exchange of three forest ecosystems in eastern China

National Natural Science Foundation of China [30590381, 30670384]; Knowledge Innovation Project of the Chinese Academy of Sciences [KZCX2-YW-432]; National Key Research and Development Program [2002CB412501]; 'Hundred Talents' Program of the Chinese Acade