Evolution of the tectogene concept, 1930-1965

The tectogene, or crustal downbuckle, was proposed in the early 1930s by F.A. Vening Meinesz to explain the unexpected belts of negative gravity anomalies in island arcs. He attributed the isostatic imbalance to a deep sialic root resulting from the action of subcrustal convection currents. Vening Meinesz's model was initially corroborated experimentally by P.H. Kuenen, but additional experiments by D.T. Griggs and geological analysis by H.H. Hess in the late 1930s led to substantial revision in detail. As modified, the tectogene provided a plausible model for the evolution of island arcs into alpine mountain belts for another two decades. Additional revisions became necessary in the early 1950s to accommodate the unexpected absence of sialic crust in the Caribbean and the marginal seas of the western Pacific. By 1960 the cherished analogy between island arcs and alpine mountain belts had collapsed under the weight of the detailed field investigations by Hess and his students in the Caribbean region. Hess then incorporated a highly modified form of the tectogene into his sea-floor spreading hypothesis. Ironically, this final incarnation of the concept preserved some of the weaker aspects of the 1930s original, such as the ad hoc explanation for the regular geometry of island arcs.

The terrestrial paleorecord of climate change over the past thousand years in the Canadian Rockies [abstract]

EXTRACT (SEE PDF FOR FULL ABSTRACT): High alpine environments provide a variety of paleorecords based on physical (glaciers, glacio-lacustrine sedimentation) and biological systems (tree rings, tree-line fluctuations). These records have varying temporal resolution and contain different climate-related signals but, in concert, provide a more comprehensive reconstruction of past climates than is possible from any single archive.

青藏苔草和紫花针茅形态结构及其生态适应性

为了研究高原植物对青藏高原特殊环境的适应机制，本文选取青藏高原高寒草原优势物种紫花针茅和青藏苔草作为研究物种，运用数量分析的方法对它们的形态结构及生态适应性进行了研究。试图通过对环境因子和这些形态结构变化之间的关系进行分析以找出这两种植物是如何适应严酷的环境以及这两种具有不同克隆生长方式的植物是否具有不同的适应机制。 2004年8月，我们沿青藏公路设置一条从西大滩到羊八井的样带，并进行取样。样带的生态特征从南到北变化较大：海拔从4586m上升到4901m，生长季降水量从384.0cm下降到202.2cm，生长季月均温从5.1℃下降到1.4℃，生长季月均相对湿度从65%，下降到54%，生长季蒸发量从1242cm下降到798cm，生长季月均风速从2.4m/s增大到4.0m/s。将采集的植物叶片材料制成石蜡切片，采用番红固绿对染，在显微镜下观察测量。同时研究了青藏苔草的根茎特征。数据分析中运用了变异系数比较，多重比较，相关分析和回归分析被用来分析结构变化及其与生态因子的关系。 结果表明，两物种的叶片结构特征在不同环境中的种群间存在显著差异。总体上，青藏苔草的保护组织、光合组织以及综合指标变异系数明显大于紫花针茅的，仅输导组织相关指标的变异系数小于紫花针茅的。这些结果表明，在叶片结构水平上青藏苔草比紫花针茅具有较大的可塑性，也意味着青藏苔草对生态因子变化的潜在适应能力可能较紫花针茅强。 紫花针茅和青藏苔草的大部分叶片结构特征与其生存环境的生态因子间存在着线性回归关系：紫花针茅叶肉细胞大小随土壤有效K含量增高而减小;下表皮厚度和韧皮部面积随生长季云盖度增高而增大;单一导管半径和导管平均面积随生长季月均湿度的增加而增大;青藏苔草上表皮细胞厚度随生长季月均最低温的降低而增厚;泡状细胞厚度随大陆度的增强而增加;上表皮细胞大小随土壤pH的增大而增大;导管总数和韧皮部面积随土壤速效P含量的增高而增加;气腔总面积随土壤有效K含量增高而增大。 青藏苔草分枝方式为典型的合轴分枝。分株间由埋藏在地下的匍匐根状茎连接，其扩展方式不呈直线形。青藏苔草喜沙质土，其根茎一般分布在沙土中10cm以下，在部分通气性能良好的风成沙丘或沙地，深度可超过50cm，间隔子长度依生境的不同而变化很大，短的仅约3cm，而长的可达2m多。 青藏苔草的间隔子长度和节间长度随有效温度的增加而减小，根茎每间隔子节间数随降水的增加而减少，在较低有效温度下和较低水分供给的情况下单位面积分布的克隆分株数量较少，在较高有效温度下和较高水分供给的情况下单位面积的克隆分株数量分布较多。这种变化趋势有助于青藏苔草有效迅速地占领条件适合的生境，充分利用生境内的各种资源，扩大种群;在不利条件下，青藏苔草则减少单位面积内克隆分株的分布，以适应不良生境。

增温与土地利用变化对青藏高原高寒草甸生态系统温室气体通量影响的研究

随着青藏高原气候变暖进程及放牧压力与开垦面积的逐年加大，非常有必要评价青藏高原高寒草甸生态系统温室气体通量对气候变暖、放牧（包括粪尿斑）和开垦的响应和反馈。与此同时，我们进行了粪尿斑及开垦对土壤理化性质影响研究。本研究对于青藏高原高寒草甸C、N循环关键过程对全球变化响应的认识具有重要意义。 本文通过2年（2006-2007）增温（白天+1.2℃，晚上+1.7℃）与放牧耦合试验，在生长季观测了增温与放牧对高寒草甸生态系统温室气体通量的影响。研究初步表明，增温试验显著增加10 cm土壤温度1℃左右；在放牧频度相对较高的2007年，放牧也显著增加了土壤温度1℃左右。总体上，增温对土壤湿度的影响不显著。 增温可以促进土壤对CH4的吸收，而放牧对CH4通量影响不大或起到一定程度抑制作用。在较低放牧频度的2006年，放牧显著降低了植物-土壤系统CO2总释放量，增温促进了CO2的释放。而在放牧频度相对较高的2007年生长季，没有发现增温及放牧对土壤-植被系统CO2释放显著的影响。2006年放牧后，增温促进了高寒草甸土壤N2O的释放；2007年，增温不放牧小区N2O总通量较对照增加了24.6%，同时放牧处理也促进了N2O释放。基于土壤温度和土壤湿度的线性回归模型可以解释55%-89%的CH4通量变异，而土壤湿度较土壤温度对CH4通量影响更大。土壤温度是影响CO2和N2O通量的主要因子，通过拟合的指数型曲线，土壤温度可以分别解释43%-63%CO2通量变异与65%-81%N2O通量变异。 在2005年与2006年夏季放牧期间，对牦牛粪尿斑处理对高寒草甸CH4、CO2和N2O通量进行了观测。牛粪小区2年观测期内CH4平均通量为687 μg m-2 h-1，而尿斑和对照土壤吸收CH4（平均通量分别为-34 μg m-2 h-1和-39 μg m-2 h-1）。牛粪小区CO2在2005年和2006年观测期内累积释放量较对照分别增加了35.8%和49.7%，而牛尿小区与对照累积释放量差异不显著。牛尿与牛粪小区N2O累积释放量显著高于对照，在2005年牛尿与牛粪小区N2O累积释放量较对照分别增加了3.7和3.5倍，而在2006年分别增加了2.1和1.8倍。因此，在估算放牧高寒草甸生态系统N2O释放时，来自牛粪斑释放量是不能被忽略的。但在中等放牧强度下（1.45头ha-1 y-1），有粪尿斑覆盖的高寒草甸在观测期内全球变暖潜势较相同面积没有粪尿斑覆盖的草甸仅增加了1%。土壤水分孔隙度（WFPS）可以解释牛尿小区35%和对照小区36%CH4通量变异。土壤温度是控制CO2释放的主要因子之一，它可以解释所有处理40-75% CO2的变异。牛尿处理（34%）、牛粪处理（48%）及对照（56%）N2O时间变异则同时受土壤温度和WFPS的驱动。在观测期内，牛尿可以显著提高土壤的pH值。粪尿斑对土壤微生物量碳氮没有产生显著性影响，但在一定时段内能显著增加土壤无机氮含量。 通过在青藏高原高寒草甸开展的人工草地试验，初步探讨了不同土地利用方式（种植燕麦、开垦后闲置及自然恢复）对CO2、CH4和N2O通量，以及土壤无机氮和微生物量氮的影响。燕麦地、自然恢复草地及开垦闲置地与天然草地相比，吸收CH4的能力均表现为增强（CH4的吸收总量分别增加了31.9%、57.2%和71.0%）。由于燕麦地生物量低于天然草地与恢复草地，造成了燕麦地土壤-植被CO2释放量低于天然草地和恢复草地。而闲置地几乎没有植被覆盖，其CO2释放量显著低于天然草地。草地恢复8年后，CO2释放基本恢复到天然草地的水平。闲置地N2O总通量显著高于天然草地，较天然草地增加了60.5%。观测期内燕麦地与天然草地相比，N2O总通量增加了24.3%，但没有达到显著性水平。开垦及种植燕麦，增加了土壤硝态氮的含量，而自然恢复地、燕麦地、天然草地和闲置地在观测期内土壤铵态氮平均含量没有观察到显著性差异。燕麦地土壤微生物量氮平均含量最低（119.7 mg N kg-1），而自然恢复草地、天然草地和闲置地土壤微生物量氮差异不大。

高山植物唐古特红景天（Rhodiola algida Var.tangutica）抗冻蛋白的研究

抗冻蛋白（AFP）或热滞蛋白（THP）最早是从极区鱼和昆虫中发现的一类特殊蛋白质，其共同特性是能阻止体液内冰核的形成和生长，降低体液的冰点，使其冰点低于熔点（热滞效应）。近几年的研究表明，AFP在某些植物、细菌、真菌中也存在。生长在极端低温环境中的高山植物体内是否也存在AFP，据我们掌握的资料，还未见这方面的报道。为此，我们选择了青海高原海拔4000米高山生长的唐古特红景天为实验材料，并在这种植物中发现了抗冻蛋白。主要实验结果概括如下： 1. 唐古特红景天叶片有很强的抗冻性，可耐受-26.5 ℃的冰冻温胁迫。在北京地区夏季（7月）驯化20天后，半致死温度（LT_(50)）升高为-15.5 ℃。叶片质外体蛋白抽提液有明显的热滞效应（0.2 ℃），在AFP中存在糖基。SDS-PAGE分析表明，质外体AFP为分子量在43-85KD范围内的五条多肽。 2. 光镜组织化学切片显示，在红景天叶片中存在蛋白质量丰富的细胞;电镜细胞化学研究揭示，在细胞壁外层及细胞间隙中分布着明显的经钌红特异染色的糖蛋白，这种糖蛋白因环境温度升高而减少。结合前述的实验结果，确认这种细胞壁外层及细胞间隙中的糖蛋白，即抗冻蛋白。 3. 以唐古特红景天叶片为外植体，在MS+BA_2+NAA_(0.2)固体培养基上可诱导出黄绿色，松脆愈伤组织。愈伤组织细胞在同样成分的液体培养基中培养获得成功。在悬浮培养液中可检测到分泌蛋白的存在。经SDS-PAGE分析表明，经低温锻炼或ABA诱导后，细胞分泌蛋白的多肽谱带数增加。与此相对应的是，细胞的抗冻能力也明显提高。PAS染色揭示，多肽中均含有糖基。 4. 通过测定热滞值，确信细胞分泌蛋白是具有抗冻活性的糖蛋白。

A survey of the gomphonemaceae and cymbellaceae (Bacillariophyta) from the Jolmolungma Mountain (Everest) region of China

Gomphonemaceae and Cymbellaceae from Jolmolungma Mountain region of China, comprised 79 taxa belonging to five genera. The dominant species were Gomphonema angustatum var. productum, G. gracile var. gracile, G. parvulum var. parvulum, G. hedini, Cymbella aequalis var. pisciculus, C. delicatula, C. cistula, C. cistula var. hebetata, C. cesatii, C. cymbiformis, C microcephala, C. minuta var. minuta., C. minuta var. silesiaca, C. pusilla, C. affinis. Some arctic and alpine forms also occurred, and the following taxa were unique to this region: Cymbella delicatula, C. naviculiformis, C. gracilis, C. minuta var. minuta, C. affinis, C. cistula, C. cymbiformis, C. cistula var. hebetata, C. arctica, Didymosphenia geminata, Gomphonema hedinii, G. olivaceum. Numbers of taxa in each samples varied greatly, It appears that species diversity of diatoms is related not only to macro-environments (e.g., geographic zonation) but also to microenvironments (e.g., microhabitats and microclimates). Cymbella spp. are not only comparatively widespread in the region but also outnumber Gomphonema spp. A large number of species found in the Jolmolungma Mountain region are also found in Tibet and the headwaters of Yangtze River.

Floral surveys on Gomphonemaceae and Cymbellaceae (Bacillariophyta) from the headwaters of the Yangtze River, Qinghai, China

Gomphonemaceae and Cymbellaceae from the headwaters of the Yangtze River, Qinghai Province, China, comprised 84 taxa belonging to four genera. The dominant species were Gomphonema kaznakowi Mer., G. hedini Hust., G. olivaceum (Lyngbye) Kutz., Cymbella cistula (Ehr.) Kirchn. var. cistula and C. minuta Hilse ex Rabh. var. minuta. Some arctic and alpine forms also occurred, and the following taxa were unique to this region: C. cistula var. asiatica Mer., C. cistula var. capitata Grun., C. yabe Skvortzow var. punctata Li and Shi, G. olivaceum (Lyngbye) Kutzing var. brevistriatum Li and Shi and G. staurophorum (Pant.) Cleve-Euler var. oblongum Li and Shi. Different morphological forms of G. kaznakowi Mer. may be related to the upheaval of the plateau. Species diversity of the diatoms appears to be related not only to macro-environment (e.g., geographic zonation) but also to microhabitat and microclimate.

增温对川西北高寒草甸植物群落结构及几种主要植物生长和生理的影响

由于人类活动所引起的地球大气层中温室气体的富集已导致全球地表平均温度在20世纪升高了0.6 ℃，并预测在本世纪将上升1.4-5.8 ℃。气候变暖对陆地植物和生态系统影响深远，并已成为全球变化研究的重要议题。高海拔、高纬度地带的生态系统对气候变化最敏感。而在高原和高山极端环境影响下所形成的高寒草甸生态系统极其脆弱，对由于温室效应引起的全球气候变化极其敏感，对这些变化的响应更具有超前性。 本研究以川西北高寒草甸植物群落及几种主要物种为研究对象，采用国际山地综合研究中心（ITEX）普遍所采用的增温方法-----开顶式生长室（OTC）模拟气候变暖来研究增温对高寒草甸植物群落结构、物质分配及其主要物种生长和生理的影响，以探讨高寒草甸植物响应与适应气候变暖的生物学和生态学机制。主要研究结论如下： 1、OTC的增温效果 由于地温、地表温度和气温的平均值在OTC内分别高出对照样地0.28℃、0.46℃和1.4℃，这说明本研究所采用的开顶式生长室（OTC）起到了增温的作用；同时，由于温室内与温室外接受的降水量相同，温室内由于热量条件的改善，土壤蒸发和植被的蒸腾作用增强，直接导致了OTC内土壤表层相对湿度的减少。 2、群落结构对增温的响应 由于增温时间较短，增温内外样地的物种组成并未发生改变；但增温后一定程度上改变了植物群落的小气候环境，从而导致物种间的竞争关系被破坏，种间竞争关系的破坏引起群落优势种组成发生相应的改变，在对照样地，鹅绒委陵菜、甘青老鹳草、遏蓝菜和蚤缀是占绝对优势的物种，而在OTC内，小米草、尼泊尔酸模、垂穗披碱草、发草和羊茅的重要性显著增加。 禾草和杂草由于对增温的生物学特性及其资源利用响应的不同，加之增温造成土壤含水量下降等环境因子的改变。与对照样地相比较，OTC内禾草的盖度及生物量都显著增加，而杂草的盖度和生物量则显著下降。 3、植物生长期对增温的响应 OTC内立枯和调落物的生物量在生长季末（10月份）都要小于对照样地的立枯和调落物生物量，而OTC内的地上鲜体生物量在10月份却略高于对照样地。这说明OTC内植物的衰老或死亡得以延缓，而植物的生长期得以延长。 4、群落生物量及分配对增温的响应 OTC内的地上鲜体生物量(10月份除外)和地下0-30cm的根系生物量与对照样地相比较，都出现了不同程度的减少；土壤根系的分配格局也发生了明显的改变，其中，OTC内0-10cm土层的生物量分配比例增加，而20-30cm土层生物量分配比例的减少。 5、群落碳、氮对增温的响应 增温后，OTC内植物群落地上活体和地下活根的碳浓度不同程度的高于对照样地，植物群落的碳库在OTC内也略高于对照样地；而OTC内植物群落地上活体和地下活根的氮浓度不同程度的低于对照样地，其植物群落的氮库与对照样地相比也略有下降。 6、几种主要植物的生长及物质分配对增温的响应 垂穗披碱草在增温后株高、比叶面积和地上生物量均显著地增加；尼泊尔酸模在增温后比叶面积和单株平均生物量积累显著地增加，而各组分中，增温处理使叶的生物量显著增加，而根的生物量却显著下降；鹅绒委陵菜在增温后株高、比叶面积和单株平均生物量积累显著地减少，而各组分中，增温处理使叶和茎的生物量显著减少，根的生物量却显著地增加。 尼泊尔酸模的LMR、RMR、R/S、根部碳含量、碳和氮在叶片与根部的分配比例在增温后显著地增加，而SMR、根部氮含量、碳和氮在茎部的分配比例在增温后却显著地降低；鹅绒委陵菜的RMR、R/S、碳和氮在根部的分配比例在增温后显著地增加，而SMR、LMR、碳在叶片的分配比例在增温后却显著地降低 7、几种主要植物的光合生理过程对增温的响应 增温使垂穗披碱草和尼泊尔酸模叶片中的叶绿素a、叶绿素b、总叶绿素含量显著增加；而鹅绒委陵菜叶片的叶绿素a、叶绿素b、总叶绿素含量在增温后显著减少，类胡萝卜素含量在增温后却显著增加。 增温对3种植物的气体交换产生了显著影响。其中，垂穗披碱草和尼泊尔酸模叶片的光响应曲线在增温后明显高于对照处理，A、E、gs、Pmax、、Rday、AQY和LSP显著增加，而LCP则显著降低；鹅绒委陵菜的光响应曲线在增温后则明显的低于对照处理，A、E、gs、Pmax、、Rday、AQY和LSP显著减少，而LCP则显著增加。 增温后垂穗披碱草和尼泊尔酸模叶片的Fv/Fm、Yield和qP显著增加；而鹅绒委陵菜叶片的Fv/Fm、Yield和qP则显著减少，qN却显著地增加。 8、几种主要植物的抗氧化酶系统对增温的响应 增温使垂穗披碱草和尼泊尔酸模体内抗氧化酶活性和非酶促作用有所提高，植物膜脂过氧化作用降低；鹅绒委陵菜叶片中酶促反应和非酶促反应在增温后也显著提高，但可能由于增温后的土壤干旱超过了鹅绒委陵菜叶的抗氧化保护能力，抗氧化酶活性及非酶促反应（脯氨酸、类胡萝卜素）的提高不足以完全清除干旱诱导形成的过量活性氧，因此叶片的膜脂过氧化程度仍然显著提高。 Enrichment of atmospheric greenhouse gases resulted from human activities such as fossil fuel burning and deforestation has increased global mean temperature by 0.6 ℃ in the 20th century and is predicted to increase in this century by 1.4-5.8 ℃. The global warming will have profound, long-term impacts on terrestrial plants and ecosystems. The ecoologcial consequences arising from global warming have also become the very important issuses of global change research. The terrestrial habitats of high-elevation and high-latitude ecosystems are regarded as the most sensitive to changing climate. The alpine meadow ecosystme, which resulted from the composite effects of mountain extreme climatic factors in Tibetan Plateau, is thus thought to be especially vulnerable and sensitive to global warming. In this paper, the response of plant community and several main species in the alpine meadow of Northewst Sichuan to experimemtal warming was studied by using open-top chambers (OTC). The aim of the this study was to research the warming effects on plant community structure, substance allocation, growth and physiological processes of several mian species, and to explore the biological and ecological mechanism of how the alpine meadow plants acclimate and adapt to future global warming. The results were as follows: 1. Warming effects of OTC The mean soil temperature, soil surface temperature and air temperature in OTC manipulation increased by 0.28℃、0.46℃ and 1.4℃ compared to the control during the growing season. This suggested that the OTC used in our study had increased temperature there. Meanwhile, the OTC manipulation slightly altered thermal conditions, but the same amount of precipitation was supplied to both the OTC manipulation and the control, so higher soil evaporation and plant transpiration in OTC manipulation directly lead to the decrease of soil surface water content. 2. The reponse of community structure to experimental warming The species richness was not changed by the short-term effect of OTC manipulation. However, experimental warming changed the microenvironment of plant community, therefore competitive balances among species were shift, leading to changes in species dominance. In the present study, the dominant plant species in the control plots were some forbs including Potentilla anserine, Geranium pylzowianum, Thlaspi arvense and Arenaria serpyllifolia, however, the importance value of some gramineous grasses including Elymus nutans, Deschampsia caespitosa, Festuca ovina, and some forbs including Euphrasia tatarica and Rumex acetosa significantly increased in OTC. The different biology characteristics and resource utilizations between gramineous grasses and forbs, and enhanced temperature caused change in some environment factors such as soil water content. As a result, the coverage and biomass of gramineous grasses significantly increased in OTC compared to the control, however, the coverage and biomass of forbs singnifciantly decreased in OTC compared to the control. 3. The reponse of plant growing season to experimental warming Both the standing dead and fallen litter biomass in OTC were lower than those in the control in October, and the biomass of aboveground live-vegetation in OTC was higher than that of the control. The results indicated that the senescence of plants was postponed, and the growing season was prolonged in our research. 4. The reponse of community biomass accumulation and its allocation to experimental warming Experimental warming caused the decrease of aboveground live biomass and belowground root biomass except for the aboveground live biomass in October. Experimental warming also had pronounced effects on the pattern of root biomass allocation. In the present study, the root biomass in 0-10cm soil layer increased in OTC manipulation compared to the control, however, the root biomass in the 20-30cm soil layer decreased in OTC manipulation compared to the control. 5. The reponse of community C and N content to experimental warming The C concentration and stock in aboveground live and belowground root both increased in OTC manipulation compared to the control. However, the N concentration and stock in aboveground live and belowground root both decreased in OTC manipulation compared to the control. 6. The reponse of gowth and biomass, C and N alloction of several species to experimental warming Experimental warming significantly increased the height, SLA (specific leaf area) and aboveground biomass of Elymus nutans in OTC manipulation compared to the control. The SLA and total biomass of Rumex acetosa also significantly increased in OTC manipulation compared to control, among the different components of Rumex acetosa, leaf biomass significantly increased, but root biomass significantly decreased in OTC manipulation compared to the control. However, the height, SLA and total biomass of Potentilla anserina significantly decreased in OTC manipulation compared to the control, among the different component of Potentilla anserina, leaf and stem biomass significantly decreased, but root biomass significantly increased in OTC manipulation compared to the control. The LMR (leaf mass ratio), RMR (root mass ratio), R/S (shoot/root biomass ration) and root C concentration of Rumex acetosa significantly increased in OTC manipulation compared to outside control, also, Rumex acetosa allocated relatively more C and N content to leaf and root in response to experimental warming, however, the SMR (stem mass ration) and root N concentration of Rumex acetosa significantly decreased in OTC manipulation compared to outside control, also, Rumex acetosa allocated relatively less C and N content to stem in response to experimental warming. The RMR and R/S of Potentilla anserina significantly increased in OTC manipulation compared to outside control, also, Potentilla anserina allocated relatively more C and N content to root in response to experimental warming, however, the SMR and LMR of Potentilla anserina significantly decreased in OTC manipulation compared to outside control, also, Potentilla anserina allocated relatively less C and N content to leaf in response to experimental warming. 7. The reponse of physiological processes of several species to experimental warming Experimental warming significantly increased chlorophyll a, chlorophyll b and total chlorophyll of Elymus nutans and Rumex acetosa in OTC manipulation compared to outside control. However, chlorophyll a, chlorophyll b, total chlorophyll and carotenoid of Potentilla anserina in OTC manipulation significantly decreased compared to outside control. Experimental warming had pronounced effects on gas exchange of Elymus nutans, Rumex acetosa and Potentilla anserine. In the present study, warming markedly increased the light response curves of Elymus nutans and Rumex acetosa in OTC manipulation compared to outside control, and also singnificantly increased A (net photosynthesis rate), E (transpiration rate), gs (stomatal conductance), Pmax (maximum net photosynthetic rate), Rday (dark respiration rate), AQY (apparent quantum yield) and LSP (light saturation point), but LCP (photosynthetic light compensation) of Elymus nutans and Rumex acetosa in OTC manipulation singnificantly decreased compared to outside control. However, warming markedly decreased the light response curves of Potentilla anserina in OTC manipulation compared to outside control, and also singnificantly decreased A, E, gs, Pmax, Rday, AQY and LSP, but LCP of Potentilla anserina in OTC manipulation singnificantly increased compared to outside control. Experimental warming singnificantly increased the chlorophyll fluorescence kinetics parameters such as Fv/Fm, Yield and qP of Elymus nutans and Rumex acetosa and qN of Potentilla anserina in OTC manipulation, but Fv/Fm, Yield and qP of Potentilla anserina in OTC manipulation singnificantly decreased. 8. The reponse of antioxidative systems of several species to experimental warming Experimental warming tended to increase the activities of antioxidative enzymes and stimulate the role of non-enzymes of Elymus nutans and Rumex acetosa. As a result, MDA content of Elymus nutans and Rumex acetosa decreased. The activities of antioxidative enzymes and non-enzymes of Potentilla anserina also significantly increased in OTC manipulation, but more O2- was produced because of lower soil water content, and the O2- accumulation exceeded the defense ability of antioxidative systems and non-enzymes fuctions. As a result, MDA content of Potentilla anserine still increased in OTC manipulation compared to outside control.

青藏高原东部混农季节性放牧及其对高山林线格局的影响

混农季节性放牧（agropastoral transhumance）通过作物种植和畜牧生产相结合的方式对不同海拔高度带上的资源进行相互补充利用，在亚洲兴都库什地区、青藏高原、横断山、东部及南部非洲、南美安第斯地区等具有悠久的历史。这种传统的生计系统几千年以来一直是居住在该地区的人类社会和自然生态系统相互作用的主要形式之一。这种传统的资源利用方式与山地自然植被以及特殊的山地人类文化和社会特征具有密切的协同演变关系。认识和理解这一关系，是山地生态学和人类学的核心科学问题之一。近年来，山地生态系统的多重功能性及动态演变对山区社会经济可持续发展的重要意义受到人们的不断关注。本文通过对云南省德钦县的12个自然村的混农季节性放牧以及对云南德钦、四川壤塘等山地植被格局特别是高海拔地带植被格局的的详细调查，探讨青藏高原东缘地区混农季节性放牧的主要特征、系统构成及相互关系，及其在全球变化、经济全球化和市场化及现代化过程中的变化趋势，分析混农季节性放牧与高山林线格局及生态系统的互动关系，旨在探讨山地地区人类活动与自然生态系统之间的互动关系，从而为山区社会经济可持续发展、环境建设和生物多样性保护等国家战略提供理论依据。 调查结果表明，混农季节性放牧是一种适应青藏高原东部高山峡谷地区环境因子及自然资源呈明显的垂直分布、资源数量稀少而时空分布异质性极高的生存环境的一种传统经济形式。这种传统的畜牧业的主要生产目的仍然是提供当地基本生存所需的产品，饲养牲口的种类和数量取决于农户的当地需求并且受资源的限制，因而维持在比较低的水平的。分布在不同海拔高度的放牧资源在一年中被牲口利用的时间也不同，互为补充，共同构成混农季节性放牧的资源基础。根据各社区永久居住点的位置和该村的土地资源特别是牧草地资源的分布范围，牲口迁移的距离和格局有较大的差异。。天然牧场仍然是最主要的畜牧业生产资源。混农季节性放牧中的农业系统和牧业系统互为补充，共同构成调查地区完整的的生计系统，农耕活动为放牧活动提供精饲料如粮食等和冬季饲料如秸秆， 其数量往往成为家庭畜牧业生产规模的主要决定因子之一。 通过对牲口数量和结构、牲口的时空迁移格局、牧业活动在整个经济活动中的相对重要性以及牧业活动和作物种植的关系方面的研究分析，混农季节性放牧在近几十年发生了深刻的变化。主要表现在牲口数量总体下降，牲口组成发生变化，牲口移动性降低、牧业活动的经济重要性下降以及牧业活动和种植活动之间的相互依存度降低等。上述变化的根本驱动力是发生在当地、地区及全球尺度上的环境、政治、社会经济、技术和文化等的变化，从而造成当地群众畜牧生产目标、土地利用和劳动力的分布等发生了变化。当地生计系统发生的改变可能会带来对方面而深刻的政治、社会经济、文化和生态影响。 混农季节性放牧这种古老的传统生计策略面临着许多挑战，如冬季饲料短缺、草场退化、缺乏市场竞争力、经济重要性降低、对年轻人缺乏吸引力、国家缺乏专门的政策指导等。与此同时，经济全球化、市场经济、新技术的应用、替代生计机会的增加、国家对于山地生态系统的作用的重新定位等也为传统生计系统转型、实现社会与生态共赢创造了机遇。 混农季节性放牧活动对亚高山及树线交错带生态系统系统的互动方式主要体现在以下几个方面：（1）牲口啃食、践踏等影响森林群落更新，改变森林群落的组成和结构，从而影响森林群落的演替进程和植被格局。林线边缘是搭建夏棚的首选地点，因此林线及树线交错地带就成了牲口活动的主要场所之一；（2）利用火烧开辟、维持和改良高山牧场； 3）在亚高山火灾迹地的放牧活动能够阻止火烧迹地的顺向演替； 4）牧民在林线附近获取建材和薪材等活动影响高山林线附近森林的结构和功能。 在调查区域，梅里雪山、白马雪山、甲午雪山的林线海拔高度在4200-4300m之间； 四川雅江、理塘一线，林线位置多在4300-4400m；四川壤塘二林场一带的林线主体在4100-4200m，在个别地区达到4300m； 在贡嘎山的南坡和东坡一带，林线位置在3600-3700m；而在四川松潘一带，林线位置主体在3700-3800米左右。树线高度的分布趋势和林线一致。混农季节性放牧及其有关人类利用活动使研究地区很多地方高山林线降低、树线交错带宽变窄或消失。在研究地区，总体情况是，阳坡和半阳坡（南坡、西南坡等）的林线和树线比阴坡和半阴坡（北坡、东北坡等）低，变化幅度达20-200m。这种差异主要是为了开辟牧场而人为清除了南向坡自然林线及其以上的植被从而使林线位置下降所致。在南坡自然林线保留得比较好的地方，林线和树线依然可以达到甚至超过北坡林线和树线的高度。放牧活动抑制了高山林线带火烧迹地的天然更新，从而使林线位置保持在目前的位置。 放牧活动对高山林线带森林群落更新的影响是显著的。自然林线内的乔木个体密度特别是新生苗和幼苗的密度大大高于非自然林线。没有放牧的自然林线及树线交错带内的I级个体（新生苗）密度达到725-2917株/公顷，而与之相对的处理样地内I级个体的密度只有0-228株/公顷；II级个体（高度10-50cm）也表现出类似的趋势，在没有放牧的自然林线及树线交错带样方内，其密度达到550-5208株/，而在放牧处理样方内只有14-321株/公顷。在非自然林线带样地内，在有正常放牧的样地内，完全缺乏I级个体。 从相对比例来看，没有放牧的样方内的I、II级个体在全部个体中所占的比例显著高于有放牧活动的样方。放牧使林线交错带的乔木幼苗数量显著减少，从而影响林线及树线交错带森林群落的天然更新过程。林线和树线交错带的灌木对乔木幼苗具有重要的保护作用，能够为树线树种如冷杉等幼苗的定居体提供有利的微气候环境，同时保护苗免受牲口的啃食和践踏。火烧以后接着进行放牧能够100%地抑制高山林线带的幼苗更新。 高山牧场放牧强度降低、使用时间缩短而低海拔地带放牧强度增加是研究地区混农季节性放牧系统的一个显著变化。这种变化也必然会引起各海拔带上的生态系统的变化。放牧强度的降低、生产性用火的停止将导致原来通过人工火烧而降低并通过进一步的火烧和放牧活动来维持的林线及其以上地带的灌木盖度和高度的增加，从而为林线森林群落的扩张创造条件。 青藏高原东部高山峡谷地区是我国重要的山地生态系统，在我国的生物多样性保护、生态环境建设、社会经济可持续发展战略中具有举足轻重的作用。正确认识人类特别是当地传统的生计系统与生态环境系统的互动关系是实现上述战略目标的前提。决策者必须以综合、系统的的视角协调促进社会经济可持续发展、保护生物及文化多样性和维持人、牲口和生态系统之间的平衡的多重目标。 Agropastoral transhumance, which makes a complementary exploitation of the natural resources at different altitudinal belts through a combination of migratory animal husbandry and crop cultivation, has a long history in Hindu-Kush Himalaya, Tibet Plateau, Hengduan Ranges, eastern and southern Africa and the Andes region of south America.For millennia, this traditional livelihood strategy has been one of the main forms of interaction between human societies inhabiting in these regions and their natural ecocystems. A close co-evolutionary relationship has been developed between this indigenous resources management systems and the mountain vegetation systems on the one hand and a unique set of cultural values and social features on the other. Understanding this relationship has been one of the core scientific issues in mountain ecology and anthropology. In recent years, the importance of the multiple functions of the mountain ecosystems and their dynamic changes in the sustainable socio-economic development of the mountain regions has gained increasing attention. This paper, which is based on a detailed study on the agropastoral practices of the 12 natural villages in Deqin County of Yunnan, and the mountainnn vegetation patterns in Deqin of Yunnan and Rangtang County of Sichuan, intends to reveal the major characteristics, system composition and the inter-relations of the subsystems of the agropastoral transhumance in Eastern Tibetan Plateau as well as the trends of changes of the system within the context of global changes, economic globalization and modernity process of China and analyze the relations between agropastoral transhumance and alpine ecosystem, ao as to understand the interactions between human activities and natural ecosystems of the mountains and provide theoretical basis for the national strategies in eocioeconomic development, environmental reconstruction and biodiversity conservation in the mountain regions. Results of the survey indicate that agropastoral transhumance in the investigated area is a traditional economic form that is highly adapted to the eastern Tibet Plateau where the topography features high peaks and deep gorges and where the highly variable environmental parameters and scanty natural resources exhibit a distinct vertical spectrum of distribution and great temporal and spatial heterogeneity. The main objective of pastoral management is still aimed at the production of basic goods and services of local people and thus the type and size of animals raised for each household mainly depend on local needs and are limited by the availability of natural resources. The scale of production is relatively low. Pastoral resources at different altidudinal belts are complementarily used at different seasons of a year and thus form the resources basis for agropastoral production of the study area. Migration distances and patterns vary with the location of the permanent settlements, the elevational distribution range of the resources of the villages concerned. Natural pastures (rangelands) are the main fodder resources and sumplement feedings only account for less than 5% of the total fodder consumption. Crop cultivation and pastoral activities support each other to form a complete livelihood system. The ability of the farmig lands (crop cultivation) to provide the pastoral activities with concentrates and sumplements often becomes a main factor limiting the scale of livestock production at household level. Agropastoral transhumance is experiencing drastic changes in recent decades as is reflected in the size and composition of animals, the seasonal migration pattern, the relative importance of pastoralism in the household economy and the interplays of agricultural and pastoral elements of the system. In general, there is a decline in animal population and mobility, a shift in animal composition to meet new needs arising from changed macro-economic situation, a decrease in the relative importance in the household economy and an increasing decoupling of agro&pastoral relations. The fundamental divers of these changes can be traced to environmental, social, economic, technological and cultural changes from local to global levels and such changes have further caused local changes in livestock management objectives, land use and distribution of labor forces. Changes in local livelihood systems could have profound political, socioeconomic, cultural and ecological conseuquences. Agropastoral transhumance, as an age-old traditional livelihood strategy, is facing multifacet challenges, such as winter fodder shortage, rangeland degradation, lack of market competitiveness, decrease in economic importance, lack of appreciation among the young generation and adequate policies from the government. At the same time, economic globalization, market economy, intrdoctution of new technologies, increase of alternative income generating opportunities and the national re-oreitation of policies on mountain ecosystems have all brought about new opportunities for the transformation of the traditional livelihood system and the synchronized development of local society and the environment. Agropastoral transhumance interacts with the ecosystems at the timberline and treeline ecotone mainly through the following aspects: 1)Animal browsing and stamping affect the regeneration process of the forest communities and alters the composition and structure of the forest which in turn affect the succession process and vegetation pattern of the forest communities. Forest edges are the priority locations for summer houses and therefore the timeline and treeline area becomes the major venues of aninal activities; （2）herders create, maintain and improve pastures through burning that remove the forest communities at the timeline and treeline ecotone; 3）immediate grazing on the fire sites can significantly prevent the fire sites from perogressive succession; and 4）herders harvesting of construction timber and firewoods affects the structure and functions of the forest communities at the timberline and treeline zone. Timberline position in the survey region shows geographical variations. It is around 4200-4300m in Meilixueshan, Baimaxueshan and Jiawuxueshan in Northwest of Yunnan and rises to 4300-4400m in Yajiang County and Litang County of Sichuan. In Rangtang of Sichuan, it is between 4100-4200m, though reaching 4300m in localized sites. In the southern and eastern slopes of Gongga Mountain, the timberline is only between 3600m and 3700m and in Songpan County at the upper reach of the Minjiang River the timberline is around 3700-3800m.Treeline pattern follows similar trend. In many places, agropastoral transhumance and related human activities have lowered the timberline and treeline and narrowed or removed the treeline ecotone. In the area of survey, generally speaking, timberlines and treelines are lower on the southern slopes than on the northern slopes, with a difference between 20 and 200m. This is mainly because that the use of fires to crerate pastures has removed the forest vegetation at the previous timberline and above. In fact, in many places, well-preserved forests on the south slopes have even high timberline position that the corresponding northern slopes. At subalpine zone, grazing activities could have prohibited the natural regeneration of many forest fire sites and maintained the forest position at the present level. Grazing has a significant impact on the regernation process of forest communities at the timberline zone. Natural timberline and treeline ecotone has much higher density of treeline species individuals especially the emergents and seedlings than the timberlines that are maintained by human activities. In natural timberline and treelien ecotone without grazing interference, the density of the I Class seedlings (less than 10cm in height) ranges 725-2917 /hm2; while that in the treatment plots (with grazing disturbance) is only 0-228//hm2；II Class seedlings (10-50cm）exhibit similar density trends, reaching 550-5208//hm2 in natural timberline without grazing but only 14-321//hm2 in the plots with grazing treatment. In the man-created timberlines, there is no I Class seedling at all in plots with normal grazing activities. In relative terms, in plots without grazing activities, the propotion of I Class and II Class seedlings is much higher than that in plots with grazing. Grazing activities have significantly reduced the number of seedlings in the timberline ane treeline ecotone, and thus affect the natural regeneration process of the forests. Shrubs at the timberline and treeline ecotone can effectively protect the seedlings from severe climate and animal tramping, thus increasing the survival rate of the seedlings. Grazing following fires can completely inhibit forest regeneration process at timberline. Changes in agropastoral transhumance will have great impact on the timberline and treeline pattern of the studied area. The decrease in grazing intensity on alpine pastrues and the cessation of the use of fires will result an increase in the cover and height of shrubs above the present human-maintained treeline, which will create further condition for the expansion of timberline forest communities. Eastern Tibet Plateau harbors some most important mountain ecosystems of China that are of vital importance to the country’s strategy in biodiversity conservation, environmental construction and sustainable sociaoeconomic development. A proper knowledge of the interactions between traditional livelihood systems and the ecosystems in the region is a precondition to the realization of the above strategic goals. Therefore, the decision-makers must have a holistic and systemic perspective so as to integrate the multiple objectives of promoting sustainable socioeconomic development, conserving biological and cultural diversity and maintaining the balances among people, animal population and the ecosystems.

环境CO2浓度和群落密度对红桦(Betula albosinensis Burk.)幼苗碳氮吸收和分配的影响

当前大气CO2浓度升高是全球变化的主要趋势之一，CO2浓度升高还会引起全球变暖等其它环境问题，因而CO2浓度浓度升高对植物影响的研究已经成为全球变化领域的焦点。红桦是川西亚高山地区暗针叶林演替初期的先锋树种和演替后期的建群种，在群落演替过程中它对环境因子的响应决定红桦群落的演替进程。本文通过控制CO2浓度的气候室试验，研究了CO2浓度倍增环境下，不同密度水平红桦碳氮固定、分配可能发生的改变，并探讨了升高大气CO2浓度对群体内部竞争的影响。以期通过本研究明确川西亚高山地区代表性物种红桦对未来气候变化的响应，为今后采取措施应对气候变化、妥善进行森林管理提供理论依据和科学指导。主要研究结果如下： 1.升高CO2浓度对红桦幼苗生长的影响以及树皮、树干响应的不同 (1) CO2浓度升高显著促进红桦幼苗的生物量、株高、基茎的生长，同时也改变生物量在体内的分配格局，主要是增加根和主茎、减少叶在总生物量中的比重。(2)树皮和树干对升高CO2浓度的影响有差异，它们对CO2浓度升高的反应程度不同，但反应方向一致。 2.密度的副效应 (1) 增加种植密度对单株生物量、株高和基径的生长具有副效应，也降低升高CO2浓度对红桦生长的正效应。(2) 增加种植密度，显著增加红桦幼苗的群体生物量，从而使红桦群体固定更多的大气CO2气体。可见密度在决定红桦生物量及固碳能力方面具有重要意义。探索适合未来大气CO2浓度升高条件下植物生长的密度，对未来的森林经济生产、生态恢复具有重要意义。 3. 升高CO2浓度对红桦幼苗苗冠结构及冠层内部竞争的影响 (1) 冠幅、冠高、苗冠表面积和苗冠体积等树冠特征均受CO2浓度升高的影响而增加，但是受密度增加的影响而降低。(2) 单位苗冠投影面积叶片数(LDcpa)和单位苗冠体积叶片数(LDcv)均低于相应的现行CO2浓度处理，这主要是由于冠幅和冠高的快速生长所造成的。(3) LDcpa和LDcv的降低表明，红桦在升高CO2浓度的条件下，会作出积极的响应，从而缓解由于群体和个体生长的增加所引起的竞争压力的增加。 4. 升高CO2浓度对红桦幼苗养分元素吸收与分配的影响 (1) CO2浓度升高，植株各器官N、P含量降低，但单株N、P总吸收量均增加。红桦幼苗体内N、P浓度的下降是由于生物量迅速增加引起的稀释效应造成的。(2) CO2浓度升高，N、P向主茎和根的分配增加，向叶片的分配减少，主要是由于前者在总生物量中的比重增加，而后者减少了。(3) CO2浓度升高，氮磷利用效率(NUE和PUE)提高，氮磷累积速率(NAcR和PAcR)显著增加。而NUE和PUE的提高可以有效缓解CO2浓度升高后，亚高山和高山地区森林土壤中养分元素不足对森林生产力的限制。 5. 升高CO2浓度对红桦幼苗群体碳平衡的影响 (1) 升高CO2浓度对植物的光合作用、呼吸速率和生长均具有促进作用。(2) 土壤有机碳含量在实验前期迅速增加，后期积累速率下降。(3) 升高CO2浓度以后，土壤呼吸显著增强；土壤呼吸还具有明显的季节变化。(4) 红桦群体日固碳量受到升高CO2浓度的促进作用。结果(1)-(4)说明所研究群落的碳动态对现行的气候波动是敏感的；所研究群落在作为大气CO2气体的源-汇关系方面至少存在季节间的源汇飘移。(5)种植密度的升高显著增加了群体固碳量。 6. 升高CO2浓度对红桦幼苗生长后期叶片衰老的影响 升高CO2浓度有利于减缓红桦幼苗叶片生长季节末期的衰老。生长季节末期，随着CO2浓度的升高光合速率和可溶性蛋白含量均呈上升趋势，同时MDA（丙二醛）含量下降，保护酶SOD（超氧化物岐化酶）、CAT（过氧化氢酶）活性升高。由此说明，升高CO2浓度有利于减缓生长季节后期叶片的衰老，使叶片维持较高的光合速率，也从生理学的角度支持了本文及前人有关CO2浓度升高促进植物光合和生长的假说及结果。 The increased CO2 concentration is one of the most important problems among global changes. The increase of CO2 will also cause other environmental problems, such as global warming, etc. So the effects of elevated CO2 on plant have drawn sights of many scientists in the research field of global change. Red birch (Betula albosinensis) usually emerges as the pioneer species in initial stage and as constructive species in later stages of forest community succession of the dark coniferous forests in Western Sichuan, China. It’s response to elevated CO2 may determine the succession process of the community where it lives in. By controlling CO2 at the ambient and twice as the ambient level (ambient + 350 umol mol-1) using enclosed-top chambers (ETC), possible effects of elevated CO2 on carbon fixation and allocation under two plantation densities are investigated. The effects of elevated CO2 on competition within canopy of red birch seedlings are also observed in the present paper. We hope to make sure of the effects of elevated CO2 on the representative species, red birch. And so that, our results could provide a strong theoretical evidence and scientific direction for forest management and afforestation under a future, CO2 elevated world. The results are as fowllows: 1. The effects of elevated CO2 on growth and the different responses of wood and bark of red birch seedlings (1) Elevated CO2 increases the growth of seedling biomass, seedling height and basal diameter of red birch. It also changed the biomass allocation in red birch seedlings. The ratio of root and main stem to all biomass is increased and the ratio of leaf is decreased. (2) Tree bark and wood show different response degree but similar response direction to elevated CO2. 2. Negative effects of planting density (1) The increase of planting density showes negative effects on the individual growth of seedling biomass, seedling height and basal diameter of red birch. It also eliminates the positive effects of elevated CO2 on growth of red birch seedlings. (2) Community biomass is increased by the elevated planting density, which means that the high density red birch community could fix more CO2 than the low density one. These results show that planting density plays an important role in determining biomass and carbon fixation ability of red birch community. Thus, exploring proper planting density becomes economically important for the future, CO2 elevated word. 3. The effects of elevated CO2 on crown architecture and competition within canopy of red birch seedlings (1) Crown width, crown depth, crown surface area and crown volume are all increased under the influence of elevated CO2. (2) Leaf number per unit area of projected crown area (LDcpa) and per unit volume of crown volume (LDcv) are lower under elevated CO2. This is resulted from the stimulated growth of tree crown features. (3) The decrease of LDcpa and LDcv indicate that plants will respond forwardly to reduce the possible increase of competition resulted from stimulated growth of individual plant and collectives in conditions of elevated CO2. 4. The effects of elevated CO2 on nutrition accumulation and allocation of red birch seedlings (1) Contents of N and P decrease due to the prompt increase of biomass of plant organs caused by elevated CO2. However, their accumulations increase under elevated CO2. (2) Elevated CO2 increases the allocation of N, P to main stem but reduced its allocation to leaf for that dry weight of the former increased but the dry weight of the later decreased. (3) Using efficiencies of N, P (NUE and PUE) and their accumulation rates (NAcR and PAcR) are found to increase under elevated CO2. Soil nutrition contents are always the limiting factors for plant growth at subalpine and alpine region. The increased NUE and PUE are helpful to eliminate the nutrition limitation in this area in the future world, when CO2 concentration doubles the ambient. 5. The effects of elevated CO2 on carbon balance of red birch communities (1) Net photosynthetic rates (Pn), dark respiration rates (Rd) and growth are all stimulated by elevated CO2. (2) Content soil organic carbon increases sharply at the primary stage of experiments and then the increasing rates decrease to a low level at later stages. (3) Soil respiration rates increase significantly with the elevation of CO2 concentration. (4) The daily carbon fixations of whole community are heightened by elevated CO2. The results (1)-(4) suggest that, the community being studied are sensitive to current climate change; the studied community, as a sink of atmospheric CO2, is pool-sink alternative between seasons. (5) The carbon fixations are increased along the increase of planting densities. 6. The effects of elevated CO2 on physiological features of leaf senescences of red birch seedlings at the later stage of growing season Elevated CO2 helps to postpone the leaf senescences of red birch at the end of the growth season. CO2 enrichment increases the photosynthetic rates, contents of soluble proteins and photosynthetic pigments. And meanwhile contents of malondialdehyde (MDA) decreases and activities of superoxide dismutase (SOD) and catalase (CAT) are both increased. These results suggest that the senescences of red birch leaves are delayed by elevated CO2, which keep the photosynthetic rates at relatively high levels. Our results lend supports to hypothesis and results on stimulated photosynthetic rates and growth from both other researchers and the present paper.

岷江上游高山林线附近优势树种土壤种子库和幼苗更新特征的研究

岷江上游地区高山/亚高山植被分布的坡向性分异显著，阴阳坡高山林线不仅物种组成差异明显，并且分布海拔呈现出阴坡高阳坡低的格局．阳坡林线树种主要是圆柏属乔木，林线类型多为渐变型，海拔高度大约在3 400m~3 800m；阴坡林线树种主要是冷杉，林线类型多为骤变型，海拔高度约在3 800m~4 400m．本研究采用土壤种子库物理筛选、室内萌发实验及野外群落调查等方法，对岷江上游地区阴坡岷江冷杉和阳坡祁连圆柏两类林线树种不同海拔梯度上土壤种子库以及幼苗库特征进行了调查，从土壤种子库和幼苗更新特征的角度对林线乔木树种种群更新特征进行了分析，进而对该地区高山林线在阴阳坡分布差异的原因进行了探讨，结果显示： 1．土壤种子库 阴坡：阴坡高山林线附近岷江冷杉土壤种子的平均密度大约为50.96粒/m2，其中树线以上10m处土壤种子密度为1.00粒/m2，树线处大约19.33粒/m2，林线交错带内土壤种子密度最高为136.83粒/m2，郁闭林内种子密度小于林线交错带，只有30.50粒/m2，种子平均空壳率为52％，霉变率达34%，完好种子只有6%．土壤种子库垂直分布特征为地被物层含种子比重最大，大约在67.50%左右；其次为0~2cm层，约18.84%左右；2~5cm层所占种子比例最小，约13.66%左右．霉变种子数量与土壤深度呈负相关． 阳坡：阳坡祁连圆柏土壤种子的平均密度为60.16粒/m2．树线以上10m处密度为1.92粒/m2，树线位置大约108.16粒/m2，林线交错带内平均为75.80粒/m2，郁闭林内种子密度小于林线交错带，只有20.00粒/m2．种子平均空壳率为36％，完好种子占49%，霉变率较低，大约为10%．阴阳坡林线树种土壤种子库垂直分布特征为：地被物层含种子最多，其次为0~4cm层，4~10cm层所占种子比例最小，霉变种子数量与土壤深度也呈负相关． 2． 幼苗库调查 阳坡：在树线以上区域没有发现幼苗，林线交错带内幼苗密度平均达3 250株/hm2，郁闭林内仅2 750株/ hm2．整个样地内1～2a幼苗很少甚至没有出现，3～10a的幼苗相对较多．空间分布上，祁连圆柏幼苗在林线交错带内接近随机分布，郁闭林内则介于随机分布和均匀分布之间． 阴坡：在树线以上幼苗密度为1 250株/ hm2，全部为1~2a幼苗，林线交错带内幼苗密度平均达7 000株/ hm2，郁闭林内达6 250株/ hm2．林线附近岷江冷杉幼苗丰富度以及幼苗的出现频率明显高于祁连圆柏，年龄结构也较祁连圆柏完整．岷江冷杉幼苗空间分布除了树线处幼苗的分布为随机分布，其他海拔则为集群分布． 3．从不同土壤深度的种子总量和幼苗数量的相关性检验发现，当年生幼苗数量跟表层种子总量相关性极显著, 但是两年生幼苗的数量与底层种子数量相关性显著．土壤种子在土壤中的垂直分布格局从一定程度上可以反映种子库的年际特征．岷江冷杉土壤种子库较丰富，种子散布后的存活力随着时间的变化逐渐下降，属于季节性瞬时种子库；祁连圆柏土壤种子散布格局为集群型分布，成熟种子大部分散布在母株冠幅内，属于永久性土壤种子库． 4．在阴坡林线交错带及以上区域还存在较为丰富的乔木土壤种子，并且在树线以上区域还发现了少量的岷江冷杉幼苗．从样地乔木的年龄结构发现，在林线交错带内上部到树线位置主要以幼龄林为主，且年龄结构完整，基本符合入侵性林线特征；阳坡林线交错带内幼苗出现频率很低，树线以上区域虽然存在种子库，但是没有幼苗出现，在林线交错带内乔木径级差距很大，年龄结构异常不完整，这种特征的林线将会面临两个可能结果：一种是维持现有状态，保持平衡；另外一种就是退化，但阳坡林线的实际动态趋势还有待长期定点研究． Treelines on the upper region of Minjiang River differ between the north aspect and the south aspect in their appearances, altitudes and tree species. On the north aspect, trees of Abies form a sharp and abrupt treeline ranging from 3800m to 4400m, while on the south the treeline is generally lower(3 400~3 800m), more open and gradual and mostly composed of Sabina. In this study, we examined the altitudinal gradients of soil seed banks and seedling recruitments at the treeline ecotones of a N-aspect and a S-aspect by using soil sieving, germination experiment and field investigations, analyzed the characteristics of population regeneration of tree species at the transitional zone and presented a analysis of the causes to the aspect-related difference in treeline patterns in the study area. Major results of our study include: 1． Soil seed bank N-aspect: Of the 50 plots investigated, the average density of soil seeds is 50.96/m2, in which well-formed seeds account for 6%, empty seeds 52%, parasitized seeds34%, and seeds damaged by animals 8%. The size of soil seed bank varies along altitude, being 1.00 seeds /m2 at the 10m above the treeline and ca.19.33 seeds/m2 at the upper limit of treeline. The highest density (136.83 seeds/m2) occurs at the treeline ecotone. By contrast, the density of soil seed for the closed forest is only 30.50 seeds/m2. In terms of vertical strata, 67.50% of the total seeds are at the surface layer, 18.84% at the middle layer (0~2cm) and 13.66% at deeper layer (2~5cm). The number of parasitized seeds is negatively correlated to soil depth. S-aspect: Of the 50 plots investigated, the average density of soil seeds is 60.16 seeds/m2, and the well-formed seeds account for 49%, empty seeds 36%, parasitized seeds10%, and seeds damaged by animals 1%. The size of soil seed bank varies along altitude, with 1.92 seeds/m2 recorded at the10m above the treeline，108.16 seeds/m2 at the upper limit of treeline, and 75.80 seeds/m2 at the treeline ecotone, while that for the closed forest is 20.00 seeds/m2. The number of seeds decreases with the depth of soil. As is on the N-aspect, the size of soil bank, from large to small, follows the order of the surface layer, the middle layer (0~4cm) and the bottom layer (4~10cm). The number of parasitized seeds is also negatively correlated to the depth of the soil. 2． Seedling bank N-aspect: A mean maximum seedling abundance of 31 000 seedlings/hm2 was recorded near alpine treeline at growing season. The density of seedlings is 1 250 seedlings/ha (all being 1 or 2 years old) at the alpine meadow 10m away above treeline, 7 000 seedlings/ha at treeline ecotone and 6 250 seedlings/ha for closed forest．The spatial distribution of Abies faxoniana seedlings is random at the upper limit of the treeline but clumped at other altitudes． S-aspect: No seedlings were found at the alpine meadow 10m away from the treeline. The density of seedlings was 3 250 seedlings/ha at treeline ecotone and 2 750 seedlings/ha for the closed forest．Hardly any 1 year current and 2 year-old seedlings appeared at the plots. The spatial distribution of Sabina przewalskii seedlings is random at treeline ecotone and between “random” and “even” forest closed forest． 3．Correlation tests of seedling population and seed bank at different soil layers indicated that the emergents were strongly correlated to seed bank at surface layer while the number of two-year seedlings was significantly correlated to the seed bank at the bottom of soil layer, indicating that germination mainly occurs at the soil surface while the middle or bottom layer was the reserve for non-germination or dead seeds. It can thus be postulated that Abies faxoniana soil seed bank is of seasonal transient type. By contrast, the soil seed bank of Sabina przewalskii is of persistent type and the soil seeds and seedlings of this species occurred more frequently near the islands of adult trees. 4．A good many soil seeds of both tree species were found near the treeline ecotone and above at N- and S-aspects. A few young seedlings were found above the Abies treeline. Investigation of five altitudinal transects respectively on N- and S-aspects indicated that Abies faxoniana has a more complete age structure than the stands of Sabina przewalskii. The age of firs decreased from closed forest to the upper limit of treeline, which suggests that the Abies treeline is advancing to higher altitude. While on the south aspect, only few Sabina przewalskii soil seeds and nearly no seedlings were found above the treeline ecotone. The stands exhibit extremely great difference in diameter classes with significantly incomplete age structure. This would lead to two possible results for the treelines: maintaining an equilibrium state at the current position or degenerating. But more studies should be carried out at longer time scales or larger spatial scales to understand whether the Sabina treeline is degenerating.

中国卧龙自然保护区不同海拔川滇高山栎(Quercus aquifolioides)群体的遗传变异

海拔梯度造成的环境异质性，如崎岖的地形、复杂的植被结构以及花期延迟等可能会极大地影响到物种的形态和遗传变异格局。理解物种形态和遗传变异的海拔格局对于物种多样性的管理和保护是非常重要的。尽管植物群体遗传学是一个飞速发展的研究领域，然而与海拔相关的形态变异、遗传变异及群体间遗传差异的研究却很少。到目前为止，还不清楚遗传变异与海拔之间是否必然的相关性。 川滇高山栎是一种重要的生态和经济型树种，广泛分布于中国西南的四川、西藏、贵州和云南省的高海拔地区，在保持水土、调节气候方面起着十分重要的作用。尽管主要受阳光限制而仅分布于阳坡，但其海拔梯度范围较大，表明川滇高山栎对不同的环境具有很强的适应性。本文通过叶型及生理响应、微卫星分子标记和扩增性片段长度多态性方法，试图探索川滇高山栎叶沿海拔梯度的形态和生理响应及其沿海拔梯度的遗传变异格局，为川滇高山栎的保护和利用提供进一步的遗传学理论依据和技术指导。 对叶形、含氮量及碳同位素的试验结果表明，平均比叶面积、气孔密度、气孔长度和气孔指数等气孔参数随海拔的升高呈非线性变化。在海拔大于2800 m时，川滇高山栎的比叶面积、气孔长度和气孔指数都随海拔升高而降低，但是在海拔小于2800 m时，这些指标都随海拔的升高而增大。相对而言，单位叶面积的含氮量和碳同位素则表现出相反的变化模式。另外，比叶面积是决定碳同位素沿海拔梯度变化的最重要参数。本研究结果表明，海拔2800 m附近是川滇高山栎生长和发育的最适地带，在这里生长的植物叶片厚度更薄、气孔更大、叶碳同位素值更小。 利用六对微卫星引物对五个不同海拔川滇高山栎群体遗传多样性进行研究，结果表明，群体内表现出较高的遗传多样性，平均每位点等位基因数11.33个，平均期望杂合度达0.820。群体间差异较小，分化仅为6.6％。聚类分析也并没有显示出明显的海拔格局。然而低频率等位基因却与海拔呈显著性正相关(R2=0.97, P < 0.01)，表明在高海拔处，川滇高山栎以更多的稀有基因来适应恶劣的环境条件。本试验结果表明由海拔梯度形成的选择性压力对川滇高山栎群体的遗传变异影响并不明显。 为了进一步探讨川滇高山栎群体遗传变异与海拔之间的相互关系，我们还对其进行了扩增性片段长度多态性分析。结果表明：(1)随海拔的升高(从群体WL2到群体WL5)，群体内遗传变异降低，而群体间遗传差异增加；(2)低海拔群体WL1表现出最低的遗传变异性(HE = 0.181)，同时与其余四个群体间呈现出最大的遗传差异性(平均FST = 0.0596)；(3)在除去低海拔群体WL1后，Mantel检测表明群体间遗传距离与海拔距离之间表现出正相关性。另外，研究结果还表明，遗传变异受生境条件(过度的湿热环境)及人为干扰(火烧、砍伐和放牧)的影响，这一点至少在低海拔群体WL1上发生了作用。 通过叶形态、生理及DNA分子水平的研究，结果表明叶形态特征和碳同位素与海拔紧密相关，与海拔之间呈非线性变化，海拔2,800 m附近是川滇高山栎生长和发育的最适地带。海拔梯度在一定程度上会影响到川滇高山栎群体的遗传变异结构，但在这样一个狭窄的地理分布区域里，这种影响并不足以导致群体间较大的遗传分化。同时生境条件及人为干扰也是影响遗传变异的限制性因子，不容忽视。 Altitudinal gradients impose heterogeneous environmental conditions, such as rugged topography, a complex pattern of vegetation and flowering delay, and they likely furthermore markedly affect the morphological and genetic variation pattern of a species. Understanding altitudinal pattern of morphological and genetic variation at a species is important for the management and conservation of species diversity. Although plant population genetics is a fast growing field of research, there are only few recent investigations, which analyzed the genetic differentiation and changes of intra-population variation along altitudinal gradients. At present, it is still unclear whether there are some common patterns of morphological and genetic variation with altitude. Quercus aquifolioides Rehder & E.H. Wilson, which is an important ecological and economical endemic woody plant species, is widely distributed in the Yunnan and Sichuan provinces, Southwest China. Its large range of habitat across different altitudes implies strong adaptation to different environments, although it is mainly restricted to sunny, south facing slopes. It plays a very important role in preventing soil erosion, soil water loss and regulating climate, as well as in retaining ecological stability. In this paper, we tried to understand the altitudinal pattern of morphological and genetic variation along altitudinal gradients through the experiments of leaf morphological and physiological responses, microsatellite analysis and AFLP markers. In leaf morphological and physiological responses experiment, we measured leaf morphology, nitrogen content and carbon isotope composition (as an indicator of water use efficiency) of Q. aquifolioides along an altitudinal gradient. We found that these leaf morphological and physiological responses to altitudinal gradients were non-linear with increasing altitude. Specific leaf area, stomatal length and index increased with increasing altitude below 2,800 m, but decreased with increasing altitude above 2,800 m. In contrast, leaf nitrogen content per unit area and carbon isotope composition showed opposite change patterns. Specific leaf area seemed to be the most important parameter that determined the carbon isotope composition along the altitudinal gradient. Our results suggest that near 2,800 m in altitude could be the optimum zone for growth and development of Q. aquifolioides, and highlight the importance of the influence of altitude in research on plant physiological ecology. Genetic variation and differentiation were investigated among five natural populations of Q. aquifolioides occurring along an altitudinal gradient that varied from 2,000 to 3,600 m above sea level in the Wolong Natural Reserve of China, by analyzing variation at six microsatellite loci. The results showed that the populations were characterized by relatively high intra-population variation with the average number of alleles equaling 11.33 per locus and the average expected heterozygosity (HE) being 0.779. The amount of genetic variation varied only little among populations, which suggests that the influence of altitude factors on microsatellite variation is limited. However, there is a significantly positive correlation between altitude and the number of low-frequency alleles (R2=0.97, P < 0.01), which indicates that Q. aquifolioides from high altitudes has more unique variation, possibly enabling adaptation to severe conditions. F statistics showed the presence of a slight deficiency of heterozygosity (FIS=0.136) and a low level of differentiation among populations (FST=0.066). The result of the cluster analysis demonstrates that the grouping of populations does not correspond to the altitude of the populations. Based on the available data, it is likely that the selective forces related to altitude are not strong enough to significantly differentiate the populations of Q. aquifolioides in terms of microsatellite variation. To further elucidate genetic variation pattern of Q. aquifolioides populations under sub-alpine environments, genetic variation and differentiation were investigated along altitudinal gradients using AFLP markers. The altitudinal populations with an average altitude interval of 400 m, i.e. WL1, WL2, WL3, WL4 and WL5, correspond to the altitudes 2,000, 2,400, 2,800, 3,200 and 3,600 m, respectively. Our results were as follows: (i) decreasing genetic variation (ranging from 0.253 to 0.210) and increasing genetic differentiation with altitude were obtained from the WL2 to the WL5 population; (ii) the WL1 population showed the lowest genetic variation (HE = 0.181) and the highest genetic differentiation (average FST = 0.0596) with the other four populations; (iii) the positive correlation was obtained using Mantel tests between genetic and altitude distances except for the WL1 population. Our results suggest that altitudinal gradients may have influenced the genetic variation pattern of Q. aquifolioides populations to some extent. In addition, habitat environments (unfavorable wet and hot conditions) and human disturbances (burning, grazing and felling) were possible influencing factors, especially to the low-altitude WL1 population. The present study shows that there were close correlations between morphological features and carbon isotope composition in our data. This indicates that a coordinated plant response modified these parameters simultaneously across different altitudes. Around 2,800 m altitude there seems to be an optimum zone for growth and development of Q. aquifolioides, as indicated by thinner leaves, larger stomata and more negative d13C values. All available evidence indicates altitudinal gradients may have influenced the genetic variation pattern of Q. aquifolioides to some extent. Decreasing genetic variation and increasing genetic differentiation with altitude was obtained except for the WL1 population. And the environment of habitats and human disturbances were also contributing factors, which impact genetic variation pattern, especially to the low-altitude WL1 population.

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.

不同放牧强度下高山草甸生态系统碳氮分布格局和循环过程研究

本研究针对川西北高山草甸缺乏科学管理，过度放牧导致草场退化，并由此引发的一系列生态环境问题，选取红原县瓦切乡1996 年草地承包后形成的四个放牧强度草场，即不放牧、轻度（1.2 头牦牛hm-1）、中度（2.0 头牦牛hm-1）和重度放牧（2.9 头牦牛hm-1），作为研究对象，研究了不同放牧强度对草地植物-土壤系统中碳、氮这两个最基本物质的分布格局和循环过程的影响，并探讨了放牧干扰下高山草甸生态系统的管理。 1．放牧对草地植物群落物种组成，尤其是优势种，产生了明显的影响。不放牧、轻度、中度和重度放牧草地群落物种数分别为22，23，26，20 种，群落盖度分别是不放牧96.2%＞中度93.6%＞轻度89.7%＞重度73.6%。随放牧强度的增加， 原植物群落中的优势种垂穗鹅冠草（ Roegneria nutans ）、发草（Deschampsia caespitosa）和垂穗披碱草（Elymus nutans）等禾草逐渐被莎草科的川嵩草（Kobresia setchwanensis）和高山嵩草（Kobresia pygmaea）所取代成为优势种。同时，随放牧强度的增加，高原毛茛（Ranunculus brotherusii）、狼毒（Stellera chamaejasme）、鹅绒委陵菜（Potentilla anserina）和车前（Plantagodepressa）等杂类草的数量也随之增加。 2．生长季6～9 月份，草地植物地上和地下生物量（0～30cm）都是从6 月份开始增长，8 月份达到最高值，9 月份开始下降。每个月份，通常地上生物量以不放牧为最高，重度放牧总是显著小于不放牧；地下生物量随放牧强度的增加表现为增加的趋势，通常重度和中度放牧显著高于不放牧和轻度放牧草地。不放牧、轻度、中度和重度放牧草地6～9 月份4 个月的植物总生物量平均值分别是1543、1622、2295 和2449 g m-2，但随放牧强度的增加越来越来多的生物量被分配到了地下部分，地下生物量占总生物量比例的大小顺序分别是重度88%＞中度82%＞轻度76%＞不放牧69%。生物量这种变化主要是由于放牧使得群落优势种发生改变而引起的，其分配比例的变化体现了草地植物对放牧干扰的适应策略。 3．植物碳氮贮量的季节变化类似与生物量的变化。每个月份，不同放牧强度间植物地上碳氮的贮量有所不同，一般重度放牧会显著减少植物地上碳氮贮量。植物根系(0～30cm)碳氮贮量随放牧强度的增加表现为增加的趋势，通常重度和中度放牧显著高于不放牧和轻度放牧草地。不放牧、轻度、中度和重度放牧草地6～9 月份4 个月的植物总碳平均值分别是547、586、847 和909 g m-2，根系碳贮量占植物总碳的比例大小顺序分别是重度88%＞中度82%＞轻度76%＞不放牧69%；放牧、轻度、中度和重度放牧草地6～9 月份4 个月的植物总氮平均值分别是17、17、23 和26 g m-2，根系氮贮量占植物总氮的比例大小顺序分别是重度79%＞轻度71%＞中度70%＞不放牧65%。 4. 土壤有机碳贮量（0～30cm）的季节变化表现为7 月份略有下降，8 月开始增加，9 月份达到的最大值。土壤氮贮量的季节变化表现为随季节的推移逐渐增加的趋势。增加的放牧强度不同程度的增加土壤有机碳氮的贮量。不放牧、轻度、中度和重度放牧6～9 月份4 个月的土壤有机碳贮量的平均值分别是9.72、10.36、10.62 和11.74 kg m-2，土壤氮贮量分别为1.45、1.56、1.66 和1.83 kg m-2。土壤中有机碳（氮）的贮量都占到了植物-土壤系统有机碳（氮）的90%以上，但不同放牧强度之间的差异不明显。 5. 土壤氮的总硝化和反硝化，温室气体N2O 和CO2 的释放率的季节变化表现为从6 月份开始增加，7 月份达到最大值，8 月份开始下降，9 月份降为最小值。增加的放牧强度趋向于增加土壤氮的总硝化和反硝化作用，温室气体N2O和CO2 的释放率，通常情况下，中度放牧和重度放牧显著地加强了这些过程。 6．垂穗鹅冠草（Roegneria nutans）和川嵩草（Kobresia setchwanensis）凋落物在不同放牧强度下经过1 年的分解，两种凋落物的失重率及其碳氮的损失率3都随放牧增加表现为增加的趋势。在同一放牧强度下，川嵩草凋落物的失重率和碳氮的损失率都高于垂穗鹅冠草凋落物。 7. 尽管重度放牧显著增加了土壤碳氮的贮量，但同时也显著降低了植被群落盖度，降低了植物地上生物量，因此，久而久之会减少植物向土壤中的碳氮归还率；与不放牧和轻度放牧相比，重度放牧又显著增加了土壤CO2 和NO2 的排放量，这是草地生态系统碳氮损失的重要途径。由此可见，对于这些地处青藏高原的非常脆弱的高山草甸生态系统，长期重度放牧不仅导致植物生产力降低，而且将导致草地生态系统退化，甚至造成土壤中碳氮含量减少。 Long-term overgrazing has resulted in considerable deterioration in alpine meadowof the northwest Sichan Province. In order to explore management strategies for thesustainability of these alpine meadows, we selected four grasslands with differentgrazing intensity (no grazing-NG: 0, light grazing-LG: 1.2, moderate grazing-MG: 2.0,and heavy grazing-HG: 2.9 yaks ha-1) to evaluate carbon, nitrogen pools and cyclingprocesses within the plant-soil system in Waqie Village, Hongyuan County, Sichuan Province. 1. Grazing obviously changed the plant species composition, especially ondominant plant species. Total number of species is 22, 23, 26, and 20 for NG, LG, MGand HG, respectively. Vegetation coverage under different grazing intensity ranked inthe order of 96.2% for HG＞93.6% for MG＞89.7% for LG＞73.6% for NG. Thedominator of HG community shifted from grasses-Roegneria nutans andDeschampsia caespitosa dominated in the NG and LG sites into sedges-Kobresiapygmaea and K. setchwanensis. At the same time, with the increase of grazingintensity, the numbers of forbs, such as Ranunculus brotherusii, Stellera chamaejasme,Potentilla anserine and Plantago depressa, increased with grazing intensity. 2. Over the growing season, aboveground and belowground biomass showed a 5single peak pattern with the highest biomass in August. For each month, abovegroundbiomass usually was the highest in the NG site and lowest in the HG site.Belowground biomass showed a trend of increase as grazing intensity increased and itwas significantly higher in the HG and MG site than in the NG and LG sites. Totalplant biomass averaged over the growing season is 1543, 1622, 2295 and 2449 g m-2for NG, LG, MG and HG, respectively. The proportion of biomass to total plantbiomass for NG, LG, MG and HG is 88%, 82%, 76% and 69%, respectively. Higherallocation ratio for is an adaptive response of plant to grazing. 3. Carbon and nitrogen storage in plant components followed the similar seasonalpatterns as their biomass under different grazing intensities. Generally, heavy grazingsignificantly decreases aboveground biomass carbon and nitrogen compared to nograzing. Carbon and nitrogen storage in root tended to increase as grazing increasedand they are significantly higher in the HG and MG sites compared to the LG and NGsite. Total Carbon storage in plant system averaged over the growing season is 547,586, 847 and 909 g m-2 for NG, LG, MG and HG, respectively, while 17, 17, 23 and 26g m-2 for nitrogen. The proportion of carbon storage in root to total plant carbon forNG, LG, MG and HG is 88%, 82%, 76%, 69%, respectively, while 65%, 71%, 70%and 79% for nitrogen. 4. Carbon storage in soil (0-30cm) decreased slightly in July, then increased inAugust and peaked in September. Nitrogen storage in soil tended to increase withseason and grazing intensity. Total Carbon storage in soil averaged over the growingseason is 9.72, 10.36, 10.62 and11.74 kg m-2 for NG, LG, MG and HG, respectively,while 1.45, 1.56, 1.66 and 1.83 for nitrogen. The proportion of carbon (nitrogen)storage in soil to plant-soil system carbon (nitrogen) storage for NG, LG, MG and HGis more than 90%, which is not markedly different among different grazing intensities. 5. Gross nitrification, denitrification, CO2 and N2O flux rates in soil increasedfrom June to July and then declined until September, all of which tended to increasewith the increase of grazing intensity. Generally, heavy and moderate grazing intensitysignificantly enhanced these process compared to no and light grazing intensity. 6. After decomposing in situ for a year, relative weight, carbon and nitrogen loss in the litter of Roegneria nutans and Kobresia setchwanensis tended to increase asgrazing intensity increased. Under the same grazing intensity, relative weight, carbonand nitrogen loss in the litter of Kobresia setchwanensis were higher than these in thelitter of Roegneria nutans. 7. Although heavy grazing intensity resulted in higher levels of carbon andnitrogen in plant and soil, it decreased vegetation coverage and aboveground biomass,which are undesirable for livestock production and sustainable grassland development.What is more, heavy grazing could also introduce potential carbon and nitrogen lossvia increasing CO2 and N2O emission into the atmosphere. Grazing at moderateintensity resulted in a plant community dominated by forage grasses with highaboveground biomass productivity and N content. The alpine meadow ecosystems inTibetan Plateau are very fragile and evolve under increasing grazing intensity by largeherbivores; therefore, deterioration of the plant-soil system, and possible declines insoil C and N, are potential without proper management in the future.

若尔盖高原花湖湖滨湿地甲烷排放研究

若尔盖高原湿地位于青藏高原东北部地区，平均海拔3,400-3,600m，是长江和黄河的自然分水区，区内发育了大面积的草本沼泽以及高寒沼泽化草甸、高寒湖泊。由于它所处的位置海拔高、气候波动较大，并处于我国三大自然区的交错过渡带，因而被认为是我国最为典型的脆弱湿地生态系统之一。由于地处偏远、自然环境条件恶劣等多方面的原因，针对若尔盖湿地的科学研究资料一直以来还非常缺乏。本文对国内外近年来在湿地生态系统甲烷排放过程、研究方法，以及关于湿地生态系统甲烷排放的影响因素进行了综述，并采用静态箱－气相色谱法，从湿地环境格局、湿地甲烷排放等方面，对若尔盖高原典型高寒湖泊湖滨不同类型湿地甲烷排放特征进行了研究，并进一步探讨了影响若尔盖高原高寒湖泊湖滨带甲烷排放的因素。得到如下结果：1．若尔盖高原花湖湖滨湿地在植物生长季（6 至8 月），甲烷排放平均速率为0.315 mg·m-2·h-1；不同月份间甲烷排放速率存在差异，分别为：-0.054、0.471、0.493 mg·m-2·h-1。不同类型湿地甲烷排放速率亦表现出差异，两栖蓼（Polygonum amphibium）湿地、滩涂和藏嵩草（Kobresia tibetica）草甸甲烷排放速率分别为：0.464、0.477、0.005mg·m-2·h-1。2．若尔盖高原花湖湖滨湿地甲烷排放速率与土壤10cm 温度显著相关。土壤温度是影响若尔盖高原花湖湖滨不同类型湿地甲烷排放的重要因素之一。随着土壤温度的升高，土壤微生物活性增强，使土壤中的氧消耗加快，氧化还原电位下降，有利于产甲烷菌的生长，从而增加土壤的甲烷产生量。3．地表水位与若尔盖高原花湖湖滨湿地甲烷排放速率相关性不显著。地表水覆盖，使得湿地土壤缺氧状况得到加强，增强了土壤中产甲烷菌的活性，促进甲烷形成，再通过植物、气泡或扩散的形式释放出土壤。但水层的加深，也使土壤中已产生的甲烷在通过气泡或扩散形式穿越水层时，被氧化的量增加，从而减少了甲烷向大气中的排放。4．植被高度以及植被地上生物量与若尔盖高原花湖湖滨带甲烷排放速率的相关性不显著。植物主要通过凋落物以及根系分泌物的输入为产甲烷菌提供基质，并作为土壤与大气之间的甲烷气体交换的传输途径；与其他环境因素共同影响湿地生态系统甲烷排放。The Zoige wetland on the eastern fringe of Qinghai-Tibetan Plateau, with averagealtitude between 3,400 and 3,600m, is the watershed of Yangtze River and YellowRiver. There are large area of peatland, subalpine meadow and lakes in this region.Due to its high elevation, transitional topology and high fluctuation of climate, theZoige wetlands represent one of the most fragile wetland ecosystems in China. And asa result of remote location and harsh environment conditions, the researches on theZoige wetland are relatively rare, especially the researches on the methane emissionfrom littoral zone of alpine lakes. Variations of methane emission rates as measuredby the method of static chamber – gas chromatography (GC) were detected fromlittoral zone of alpine lake on the Zoige Plateau. Relationships between methaneemission rates and environmental factors were analyzed. It is concluded that:1．The average methane emission rate in the littoral zone of Huahu Lake, ZoigePlateau is 0.315 mg·m-2·h-1, with evident spatial and temporal variations. The littoralzone has different methane effluxes with -0.054, 0.471, and 0.493 mg·CH4·m-2·h-1in June, July and August respectively. Different types of wetland have differentmethane emission rates, with value of 0.464, 0.477, and 0.005 mg·CH4·m-2·h-1 forPolygonum amphibium wetland ( PA ), Shoal ( S ) and Kobresi tibetica meadow ( KT ), respectively.2． The soil temperature at 10cm is significantly correlated with the methane effluxesin littoral zone of Huahu Lake, Zoige Plateau, and which is one of the most important factors influencing the methane emission from this region. The activities of soilmicroorganisms rise under higher soil temperature and increases oxygen consumptionand decreases Eh, which is in favor of the methanogensis, and enhances theproduction of methane in soil.3． The correlation between the standing water and methane effluxes from littoralzone of Huahu Lake is not significant. Because of the standing water, the anaerobicconditions of wetland soil have been enhanced, and are favor to the decomposition oforganic matter. And the anaerobic conditions strengthen the methanogensis’ activities,thus the methane production, which release to the atmosphere by diffusion, ebullitionand aerenchymal plants. With the water level’s increase, more methane produced insoil which is transferred by ebullitions or diffusion are oxidated, thus reduce themethane release to the atmosphere.4． The height and aboveground biomass of vegetation are not significant related tothe methane effluxes from littoral zone of Huahu Lake, Zoige Plateau. The vegetationprovides substrates for methanogensis by litter and root exudates; act as thetransportation way of methane between soil and atmosphere; influence the methaneemission of wetland ecosystems with other environment factors.