106 resultados para CLIMATIC ZONING


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In Asia, especially in China, our knowledge of the distribution of testate amoebae is still limited. In this paper, the geographical distribution of testate amoebae in Tibetan Plateau and northwestern Yunnan Plateau, southwest China and their relationships with the climatic factors have been studied. We found testate amoebae shifted in the most dominant species and increased in species (or genus) richness from northwest to southeast. Further, the linear regression analyses revealed that both species richness and genus richness have higher positive correlations with the mean temperature of the warmest month and annual mean precipitation as contrasted with the mean altitude, which showed weak negative correlation. This indicates that the temperature and precipitation are more significant influences on the richness than the altitude. The cluster analysis based on the community structure, defined by Sorenson's coefficient matrix, suggested four groups from the 10 physiographical regions. This geographical distribution pattern was also closely related with the climatic regionalization. The present climatic regionalization pattern of the study area originated from the uplift of Tibetan Plateau and mainly occurred in or after the late Pleistocene. Therefore, the geographical distribution of testate amoebae in our study area may have experienced complicated and drastic changes corresponding to the variation of the climate caused by the geological events.

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During the twentieth century evidence was presented which suggested the presence of various strains and races of the parasite Ichthyophthirius multifiliis Fouquet. However, ecological profiles of various parasite isolates from different climatic zones are sparse. Such stringent characterizations of parasite development at defined abiotic conditions could provide valuable criteria for the different races: profile comparison from various localities is one way to differentiate these strains. Baseline investigations were therefore performed on the associations between abiotic factors (temperature/salinity) and the development of theronts in tomocysts of I. multifiliis isolated from rainbow trout in a Danish trout farm. It was shown that tomocyst formation and theront development took place between 5 and 30degreesC. Development rates and sizes of theronts were clearly affected by temperature: theronts escaped tomocysts already after 16-27 h at 25degreesC and 30degreesC, whereas this process took 8-9 days at 5degreesC. Likewise, theront size decreased steadily from a maximum of 57.4 x 28.6 mum at 5degreesC to 28.6 x 20.0 mum at 30degreesC. This size variation was only partly associated with the number of theronts that appeared at different temperatures. The lowest number of theronts escaping from one tomocyst was indeed found at 5-7degreesC (mean 329-413). At 11.6, 17.0 and 21degreesC. the highest number of theronts appeared (mean 546-642). However, at 25 and 30degreesC, the number decreased (458 and 424, respectively). Additional studies on the salinity dependent development of the parasite (at 11.6degreesC) showed that salinities above 5 p.p.t. totally inhibited development. Even at 5 p.p.t. the developmental time significantly increased and the number of theronts produced from one tomocyst decreased.

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Unequivocal Eocene suckers from China are for the first time reported here. This discovery demonstrates that catostomids of the Eocene Epoch (some 55-35 Ma ago) are scattered widely on mainland Asia as well as western North America. The present day disjunct distribution pattern of catostomids, with 68 extant species widespread in North America and the northern part of Middle America and only two in the restricted areas of Asia, is the result of their post-Eocene decline in Asia due to the competitive pressure from cyprinids, their Late Cenozoic radiation in North America, and the vicariant and dispersal events triggered by the changed biogeographic landscape. All of these prove to be a historical product of the geological, biological, and climatic changes throughout the Cenozoic.

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A non-linear perturbation model for river flow forecasting is developed, based on consideration of catchment wetness using an antecedent precipitation index (API). Catchment seasonality, of the form accounted for in the linear perturbation model (the LPM), and non-linear behaviour both in the runoff generation mechanism and in the flow routing processes are represented by a constrained nan-linear model, the NLPM-API. A total of ten catchments, across a range of climatic conditions and catchment area magnitudes, located in China and in other countries, were selected for testing daily rainfall-runoff forecasting with this model. It was found that the NLPM-API model was significantly more efficient than the original linear perturbation model (the LPM). However, restric tion of explicit nan-linearity to the runoff generation process, in the simpler LPM-API form of the model, did not produce a significantly lower value of the efficiency in flood forecasting, in terms of the model efficiency index R-2. (C) 1997 Elsevier Science B.V.

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Principal Component and Canonical Correlation Analysis of the Environmental Factors Influencing the Growth of Caragana korshinskii Kom. in Grassland

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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.

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大气CO2浓度的增加已经成为不可争议的事实。预计本世纪末大气CO2浓度将增加到约700µmol mol-1。森林年光合产量约占陆地生态系统年光合产量的70%。森林树木是一个巨大的生物碳库,约占全球陆地生物碳库的85%。森林树木对CO2的固定潜力是缓解由大气CO2浓度升高引起的未来全球气候变化问题的决定性因子之一。红桦(Betula albosinensis Burk.)是川西亚高山采伐迹地自然或人工恢复的重要树种。本研究以1a红桦幼苗为模式植物,采用人工模拟的方法,研究CO2浓度升高对不同种内竞争强度(种群水平)下红桦幼苗的生理特征、生长、干物质积累及其分配的影响,探讨在种内竞争生长条件下红桦幼苗的“光合适应机理”与生长特征,为西南亚高山森林生产力对未来全球变化的预测提供重要参考。 本研究的主要结果如下: 1)在种内竞争生长条件下红桦幼苗经过CO2浓度升高熏蒸4个月后,叶片出现“光合适应”现象。与对照相比,低种植密度(28株m-2)和高种植密度(84株m-2)条件下的红桦幼苗净光合速率(A)、气孔导度(gs)、蒸腾速率(E)、表观量子产量(AQY)和羧化速率(CE)显著降低,而水分利用效率(WUE)则显著提高。CO2浓度升高处理的红桦幼苗叶片Rubisco活性、单位叶面积N浓度、叶绿素a、叶绿素b和类胡萝卜素浓度都显著降低。但CO2浓度对红桦幼苗的叶绿素a与叶绿素b的比值没有显著影响。CO2浓度升高显著增加红桦幼苗单位叶面积的非结构性碳水化合物(TNC)浓度,结果是红桦幼苗的比叶面积(SLA,cm2 g-1)显著降低。 2)与对照相比,CO2浓度升高处理的红桦幼苗高、基径、单叶面积和侧枝的相对生长速率(R GR)显著提高,尤其在试验处理的早期。CO2浓度升高既增加单株红桦幼苗总叶片数量又增加单叶面积,结果是单株红桦幼苗的总叶面积比对照显著增加。 3)CO2浓度升高处理显著增加红桦幼苗干物质积累(尤其是细根生物量),改变了红桦幼苗生物量的分配格局。与对照相比,CO2浓度升高处理的红桦幼苗叶重比(LWR)、叶面积比(LAR)、叶根重比(Wl/Wr)和源汇重比(leaf weight to non-leaf weight ratio, Wsource/Wsink)显著下降(高种植密度的LWR除外),而根冠比(R/S)则显著增加。在两种种植密度条件下,CO2浓度升高显著增加红桦幼苗根生物量的分配比率,显著降低叶片的生物量分配比率,对主茎、侧枝以及地上生物量的分配比率不变或约有下降。 总之,长期生长在CO2浓度升高条件下的红桦幼苗光合能力下降,并伴随Rubisco活性、叶N浓度、光合色素浓度的显著降低以及TNC浓度的显著增加。支持树木光合速率下降与Rubisco活性、叶N浓度下降以及TNC浓度增加紧密相关的假设。CO2浓度升高处理红桦幼苗的早期相对生长速率大大高于对照,而后期迅速下降,说明红桦幼苗生物量的显著增加主要归功于CO2浓度升高的早期促进作用和叶面积的显著增加。CO2浓度升高显著增加红桦幼苗根系生物量和根冠比,表明红桦幼苗“额外”固定的C向根系转移。 The steady increae of atmospheric CO2 concentration([CO2])has been inevitable fact. Models predict that the atmospheric [CO2] will increase to about 700µmol mol-1 at the end of the twenty-first century. As trees constitute a majoor carbon reservoir–85% of total plant carbon is found in forest, and their ability to sequester carbon is a key determinant of future global change problems caused by increases in atmospheric CO2. In addition to the role of forests in the global carbon cycle, inceased growth could be of economic benefit, for example, offsetting deleterious effects of climatic changes. Betula albosinensis (Burk.) usually emerges as the pioneer species in initial stage and as constructive species in later stages of forest community succession of mountain forest area, and also is one of important tree species for afforestation in logged area, in southwesten China. In this experinment, Betula albosinensis seedling (one-year-old) was used as the model plant. B. albosinensis seedlings were grown under two all-day [CO2], ambient (about 350 µmol·mol-1) and elevated [CO2] (about 700 µmol·mol-1), and two planting densities of 28 plants per m2 and 84 plants per m2. The objectives were to characterize birch mature leaf photosynthesis, growth, mass accumulation and allocation responses to long-tern elevated growth [CO2] under the influences of neighbouring plants, and to assess whether elevated [CO2] regulated birch mature leaf photosynthetic capacity, in terms of leaf nitrogen concentration (leaf [N]), activity of ribulose bisphosphate carboxygenase (Rubisco), Rubisco photosynthetic efficiency, and total nonstructural carbohydrates (TNC) concentration, and also to provide a strong reference to predict the productivity of subalpine forests under the future global changes. The results are as follows: 1) B.albosinensis seedlings exposed to elevated [CO2] for 120 days, photosynthetic acclimation phenomena occurred. At two planting densities, leaves of birch seedlings grown under elevated [CO2] had lower net photosynthetic rate (A), stomatal conductance (gs), transpiration (E), apparent quantum yield (AQY) and carboxylated efficiency (CE) and higher water use efficiency (WUE), compared to those of B.albosinensis seedlings grown under ambient [CO2]. Based on the leaf area, leaf [N], Rubisco activity and photosynthetic pigments concentrations of B. albosinensis seedlings grown under elevated [CO2] were significantly lower than those grown under ambient [CO2]. The ratio of chlorophyll a to chlorophyll b concentration was not affected by elevated [CO2]. Under elevated [CO2], the TNC concentration per unit leaf area significantly increased, resulting in significant decrease in specific leaf area. Thus leaf photosynthetic capacity of B. albosinensis seedlings would perform worse under rising atmospheric [CO2] and the influences of neighbouring plants. 2) Under elevated [CO2], the relative growth rate (RGR) of B. albosinensis seedlings height, basal diameter, a leaf area and branch length significantly increased, especially at the initial stage of exposure to elevated [CO2], and a leaf area and leaf numbers per B. albosinensis seedling also significantly increased. Thus the total leaf area per B. albosinensis seedling was significantly increased under elevated [CO2]. 3) As the increase of RGR and total leaf area, biomass of B. albosinensis seedling grown elevated [CO2] was higher, compared to that of B.albosinensis seedlings grown at ambient [CO2]. Elevated [CO2] changed the biomass allocation pattern of B. albosinensis seedling. At two planting densities, B. albosinensis seedlings grown elevated [CO2] had lower leaf weight to total weight ratio (LWR), leaf area to total weight ratio (LAR) and leaf weight to non-leaf weight ratio (Wsource/Wsink), but higher root weight to shoot weight ratio (R/S), compared to those of B.albosinensis seedlings grown at ambient [CO2]. Under elevated [CO2], roots biomass to total biomass ratio was signigicantly increased, leaves biomass to total biomass ratio was significantly decreased. The main stem and branch biomass to total biomass ratio were not affected by elevated [CO2]. In conclusion, our results supported the hypothesis that the decline in photosynthetic capacity of C3 plants will appear after long-term exposure to elevated [CO2], accompanying with the significant decrease in Rubisco activity, leaf N concentration, photosynthetic pigments concentration, and significant increase in total non-structural carbohydrates concentration. Our results also have shown that the increase of biomass of B. albosinensis seedlings should be attributed to initial stimulation on RGR and total leaf area resulted from elevated [CO2]. Under elevated [CO2], the extra carbon sequestered by B.albosinensis seedlings transferred into under-ground part because of increase in root biomass and R/S.

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青杨作为一个本土树种,能较好的适应潮湿和寒冷的环境,对中国西部的人工造林有着重要的参考价值。在本实验中,选取7个中国西南地区分布的自然群体,用ISSR(inter-simple sequence repeats)作为分子标记研究其遗传多样性水平和遗传结构。通过筛选的8个ISSR引物,获得了158条清晰可重复的DNA条带,其中有156条具有多样性(占98.7%)。平均的Nei’s遗传多样性(h)为0.331;遗传分化系数(GST)为0.477,这表明有47.7%的遗传多样性发生在群体间。这种高水平的分化可能是由于当地复杂多变的地形和气候特点阻碍了基因流而引起的。此外在这7个青杨群体中,遗传距离和地理距离并未体现出有显著相关性(r=0.3122, P>0.05)。联合遗传距离和地理距离分析,鉴定出两处低水平基因交流的地区, 探讨其遗传障碍形成原因。 As a native species to China, Populus cathayana Rehd is well-adapted to the wet and cold environments where it occurs. It is considered to be an important reforestation species in western China. In the present study, we surveyed the level of genetic variation and the pattern of genetic structure in seven natural populations of P. cathayana, originating from the southeastern Qinghai-Tibetan Plateau of China, by using ISSR (inter-simple sequence repeats) markers. Based on eight primers, 158 clear and reproducible DNA fragments were generated, of which 156 (98.7%) were polymorphic. The average value of Nei's gene diversity (h) equaled 0.331. The coefficient of genetic differentiation (GST) equaled 0.477, which means that 47.7% of the total molecular variance existed among populations. Such a high level of divergence present among populations may be caused by the complex topography and variable climatic conditions present in the southeastern Qinghai-Tibetan Plateau which effectively restrict gene flow. Moreover, there is a lack of significant association between genetic and geographical distances (r=0.3122, P>0.05) in the populations of P. cathayana. The application of a novel method, which combines geographical coordinates and genetic differentiation to detect barriers for gene flow, allowed us to identify two zones of lowered gene flow.

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青杨(Populus cathayana Rehd.)是青杨派杨树的主要树种之一,为我国特有乡土树种,其主要分布区之一是我国的青藏高原,集中分布地带在甘肃省中部及青海省东部,四川省西北部岷江上游和松潘等地区。本研究以青藏高原东缘青杨天然分布区的6个群体143个个体为材料,用AFLP、SSR和叶绿体SSR分子标记分析青杨天然群体的遗传多样性,分析其遗传结构和分化,比较6个群体间遗传多样性的高低和群体间的遗传关系。旨在为青杨基因资源评价、保护与保存、遗传改良策略制定等提供科学理论依据。通过以上研究,得出如下主要研究结果: 1 AFLP分子标记研究结果 采用4对选择性引物对6个青杨天然群体143个个体进行分析,扩增谱带分析共检测到175个位点,其中173个位点表现为多态,多态位点百分率高达98.9%。从整体上表现出较高的遗传多样性,Nei’s基因多样度(h)水平为0.306。从青杨天然群体位点分布来看,有高达20%的位点(32位点)为群体所特有,仅有9.14%的位点(16位点)在所有群体中存在。群体间的遗传分化极大,所有遗传变异中,有48.9%的遗传变异存在于群体间。在个体群丛(Individuals cluster)和主坐标(PCO analysis)分析中,青杨各群体未呈现任何地理模式,Mantel检测也显示各群体间遗传距离与地理距离无明显相关。研究认为,由于地理和空间上大尺度的隔离和地形地貌复杂使得群体间无法进行基因交流,导致群体间遗传分化极大,另外各群体在不同的选择压力下,经历各自独立的进化历程,这些都可能导致群体间遗传距离与地理距离的不相关。 2 SSR分子标记研究结果 在SSR分析中,7个位点在6个青杨天然群体143个个体中共检测到79个等位基因,每位点检测到的等位基因数在5-16之间,平均11.3个,总体上多态位点百分率达100%。平均观察杂合度和期望杂合度分别为0.792和0.802。Hardy-Weinberg平衡检验表明青杨大部分群体都处于非平衡状态,群体大部分位点都是偏离哈迪-温伯格平衡(76.3%),只有23.7%的测验满足哈迪-温伯格平衡。分析青杨天然群体内和群体间的遗传变异,基因分化系数(GST)为0.373,即有62.7%的遗传变异存在群体内,37.3%的遗传变异存在群体间。群体内的遗传变异高于群体间水平。根据各群体遗传距离UPGMA聚类分析,有来自相临分布区、近似气候类型的群体聚在一起的趋势,但Mantel检测反映遗传距离与地理距离间并无明显相关性。 3 cpSSR分子标记研究结果 分析来自青藏高原东缘6个青杨天然群体,所用cpSSR引物中有5对cpSSR引物(CCMP2、CCMP5、SCUO01、SCU03、SCU07)都表现较高的多态性,单个引物检测的片段数都在4以上。5对cpSSR引物共检测片段数26个,组成了12种叶绿体DNA单倍型。各群体的单倍型分布和频率有较大差异,群体单倍型多样性范围为0-0.4926,TS、JZ、PW和SHY群体单倍型多样性高于QHY和LED群体水平。本研究发现,分布在青藏高原东缘的青杨天然群体,群体间不存在共享的单倍型,各群体间存在极大的遗传分化(GST=0.9223)。从青藏高原东缘地区经历的地质历史事件来看,第四纪的冰期气候变迁可能是造成青杨现今遗传结构模式的主要因素之一。根据单倍型在各群体的分布情况,进行青杨群体聚类分析结果,各群体无明显的分组现象,青杨各群体也未呈现任何清晰地理模式。 由于不同分子标记在对群体遗传多样性检测能力与效率上存在差异,所以三种标记检测的青藏高原东缘青杨天然群体遗传多性水平也不尽一致,但在与用同种方法检测其它物种或同一物种不同种源群体比较,三种分子标记方法都揭示了青藏高原东缘青杨天然群体具有中等偏上的遗传多样性水平。结果分析表明,群体间遗传分化极大,这是由于青杨天然群体分布于青藏高原东缘,既有高原又有高山峡谷,由于地理和空间上大尺度的隔离和地形地貌复杂导致了基因流物理上的阻隔。三种分子标记研究结果经Mantel分析检测,遗传距离与地理距离之间都无明显相关性。较为一致的解释是,青杨分布区域地理和空间上大尺度的隔离和和地形地貌复杂导致群体之间不存在均匀扩散现象,另外各群体在不同的选择压力下,经历各自独立的进化历程,这些都可能导致群体间遗传距离与地理距离的不相关。 The wide geographical and climatic distribution of P. cathayana Rehd. indicates that there is a large amount of genetic diversity available, which can be exploited for conservation, breeding programs and afforestation schemes. The results are as follows: 1 Research results of AFLP genetic diversity In present study, genetic diversity was evaluated in the natural populations of P. cathayana originating from southern and eastern edge of the Qinghai-Tibetan Plateau of China by means of AFLP markers. For four primer combinations, a total of 175 bands were obtained, of which 173 (98.9%) were polymorphic. Six natural populations of P. cathayana possessed different levels of genetic diversity, high level of genetic differentiation existed among populations (GST=0.489) of P. cathayana. Individuals cluster and PCO analysis based on Jaccard’s similarity coefficient also showed evident population genetic structure with high level population genetic differentiation. The long evolutionary process coupled with genetic drift within populations, rather than contemporary gene flow, are the major forces shaping genetic structure of P. cathayana populations. Moreover, there is no correspondence between geographical and genetic distances in the populations of P. cathayana, seldom gene exchange among populations and different selection pressures may be the causes. Our finding of different levels of genetic diversity within population and high level of genetic differentiation among populations provided promising condition for further breeding or conservation programs. 2 Research results of SSR genetic diversity In this study, the genetic diversity of P. cathayana was investigated using microsatellite markers. In a total of 150 individuals collected from six natural populations in the southeastern part of the Qinghai-Tibetan Plateau in China, a high level of microsatellite polymorphism was detected. At the seven investigated microsatellite loci, the number of alleles per locus ranged from 5 to 16, with a mean of 11.3, the observed heterozygosities across populations ranged from 0.408 to 0.986, with a mean of 0.792, and the expected heterozygosities across populations ranged from 0.511 to 0.891, with a mean of 0.802. The proportion of genetic differentiation among populations accounted for 37.3% of the whole genetic diversity. The presence of such a high level of genetic diversity could be attributed to the features of the species and the habitats where the sampled populations occur: The southeastern part of the Qinghai-Tibetan Plateau is regarded as the natural distribution and variation center of the genus Populus in China. Variation in environmental conditions and selection pressures in different populations, and topographic dispersal barriers could be factors associated with the high level of genetic differentiation found among populations. The populations possessed significant heterozygosity excesses, which may be due to extensive population mixing at the local scale. The cluster analysis showed that the populations are not strictly grouped according to their geographic distances but the habitat characteristics also influence the divergence pattern. In addition, we suggest that population SHY should be regarded as an ecologically divergent species of P. cathayana. 3 Research results of cpSSR genetic diversity Genetic diversity of six natural populations of P. cathayana originating from the southeastern part of the Qinghai-Tibetan Plateau in China was studied by use of cpSSR markers. Based on 5 pairs of polymorphic primers screened from 12 pairs of primers, twenty-six different length fragments and twelve different kinds of haplotypes were reduced in 143 samples. There were significant variant haplotypes among the populations.There were no shared haplotypes found among populations, analysis of molecular variance indicated that a high proportion of the total genetic variance was attributable to variations among populations (92.23%). The pattern of genetic structure which is associated with spatial separation, variation in environmental conditions and selection pressures in different populations, is also the result of geological historical factor. A molecular phylogenetic tree based on the 12 haplotypes showed that the populations are not strictly grouped according to their geographic distances.

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本研究通过粗枝云杉不同种群进行的温室半控制试验,采用植物生态学、生理学和生物化学的研究方法,系统地研究了粗枝云杉不同种群抗旱性的生长、形态、生理和生化机理,并结合有关研究进行综合分析,得出主要研究结论如下: 1.粗枝云杉对干旱胁迫的综合反应 粗枝云杉在干旱胁迫下的适应机制为:(1)相对生长速率及植株结构的调整:干旱胁迫下虽然植株相对生长速率显著降低,且有相对较多的生物量向根部分配,但并未发现细根/总根比增加。(2)粗枝云杉对干旱胁迫的光合作用表现为:干旱胁迫显著地降低了控制的理想条件下的气体交换,但干旱胁迫对PSII最大光化学效率(Fv/Fm)没有影响,表明干旱并未影响到光合机构。(3)干旱还影响了很多生理生化过程,包括渗透调解物质(游离脯氨酸)、膜脂过氧化产物、脱落酸(ABA)含量的增加,以及保护酶活性的升高。这些结果证明植物遭受干旱胁迫后发生了一系列的形态、生理和生化响应,这些变化能提高干旱时期植物的存活和生长能力。 2.粗枝云杉不同种群对干旱胁迫反应的种群差异 粗枝云杉三个种群-干旱种群(四川丹巴和甘肃迭部)和湿润种群(四川黑水)对干旱适应不同,这种不同应归因于它们采用的用水策略不同:在水分良好和干旱胁迫条件下,受试种群在相对生长速率和水分利用效率(WUE)方面都表现出显著的种群间差异。与湿润种群相比,干旱种群在两种水分条件下有更高的WUE。粗枝云杉不同种群的碳同位素组分(δ13C)只在干旱胁迫下有显著差异,并且这种差异在水分良好时比干旱胁迫条件下小,说明生理响应和干旱适应性之间的关系受植物内部抗旱机制和外部环境条件(如水分可利用性)或两者互作效应的影响。这些结果说明干旱种群和湿润种群所采用的用水策略不同。干旱种群有更强的抗旱能力,采用的是节水型的用水策略,而湿润种群抗旱能力较弱,采用的是耗水型的用水策略。 3. 遮荫对粗枝云杉不同种群抗旱性影响 干旱胁迫显著降低了全光条件下叶相对含水量(RWC)、相对生长速率、气体交换参数、PSII的有效量子产量(Y),提高了非光化学猝灭效率(qN)、水分利用效率、脯氨酸(PRO)积累、脱落酸(ABA)含量及保护酶活性。然而这种变化在遮荫条件下不明显。我们得出结论适度遮荫降低了干旱对植物的胁迫作用。另一方面,在干旱条件下,与湿润种群相比,干旱种群抗旱性更强,表现在干旱种群净光合速率与单位重量上叶氮含量(Nmass)降低较少。另外,干旱种群表现出更为敏感的气孔导度,更高的热耗散能力(qN)能力、用水效率、ABA积累、保护酶活性,以及更低的总用水量、相对生长速率。这一结果表明这两种群采用不同的生理策略对干旱和遮荫做出反应。许多生长和生理反应差异与这两个种群原产地气候条件相适应。 4. 外源脱落酸(ABA)喷施对粗枝云杉不同种群抗旱性影响 外源ABA喷施在干旱和水分良好条件下均不同程度地提高了根/茎比,表明根和茎对ABA敏感程度不同。实验结果还表明,外源ABA喷施对这两个种群在干旱胁迫期间影响不同。干旱胁迫期间,伴随着ABA喷施,湿润种群净光合速率(A)显著降低,而干旱种群净光合速率变化不明显。另一方面,外源ABA喷施显著提高了干旱条件下干旱种群的单位叶面积重(LMA)、根/茎比、细根/总根(Ft)比、水分利用效率(WUE)、ABA含量, 以及保护酶活性。然而,外源ABA喷施对湿润种群的上述测定指标没有显著影响。这一结果表明干旱种群对外源ABA喷施更为敏感, 反应在更大的气孔导度降低,更高的生物量可塑性,及更高的水分利用效率、ABA含量和保护酶活性。综上所述,我们得出结论,粗枝云杉对外源ABA敏感性因种群的不同而不同。该研究结果可为两个明显不同种群在适应分化方面提供强有力的证据。 Arid or semi-arid land covers more than half of China's land territory. In arid systems, severe shortages of soil water often coincide with periods of high temperatures and high solar radiation, producing multiple stresses on plant performance. Protection from high radiation loads in shaded microenvironments during drought may compensate for a loss of productivity due to reduced irradiance when water is available. Additionally, ABA, a well-known stress-inducible plant hormone, has long been studied as a potential mediator for induction of drought tolerance in plants. Picea asperata Mast., which is one of the most important tree species used for the production of pulp wood and timber, is a prime reforestation species in western China. In this experiment, different population of P. asperata were used as experiment material to study the adaptability to drought stress and population differences in adaptabiliy, and the effects of shade and exogenous abscisic acid (ABA) application on the drought tolerance. Our results cold provide a strong theoretical evidence and scientific direction for the afforestation, and rehabilitation of ecosystem in the arid and semi-arid area, and provide a strong evidence for adaptive differentiation of different populations, and so may be used as criteria for species selection and tree improvement. The results are as follows: 1. A large set of parallel response to drought stress Drought stress caused pronounced inhibition of the growth and increased relatively dry matter allocation into the root; drought stress also caused pronounced inhibition of photosynthesis, while drought showed no effects on the maximal quantum yield of PSII photochemistry (Fv/Fm) in dark-adapted leaves, indicating that drought had no effects on the primary photochemistry of PSII. However, in light-adapted leaves, drought reduced the quantum yield of PSII electron transport (Y) and increased the non-photochemical quenching (qN). Drought also affected many physiological and biochemical processes, including increases in superoxide dismutase (SOD), ascorbate peroxidase (APX) activities, malondialdehyde and ABA content. These results demonstrate that there are a large set of parallel changes in the morphological, physiological and biochemical responses when plants are exposed to drought stress; these changes may enhance the capability of plants to survive and grow during drought periods. 2. Difference in adaptation to drought stress between contrasting populations of Picea asperata There were significant population differences in growth, dry matter allocation and water use efficiency. Compared with the wet climate population (Heishui), the dry climate population (Dan ba and Jiebu) showed higher LMA, fine root/total root ratio and water use efficiency under drought-stressed treatments. The results suggested that there were different water-use strategies between the dry population and the wet population. The dry climate population with higher drought tolerance may employ a conservative water-use strategy, whereas the wet climate population with lower drought tolerance may employ a prodigal water-use strategy. These variations in drought responses may be used as criteria for species selection and tree improvement. 3. The effects of shade on the drought tolerance For both populations tested, drought resulted in lower needle relative water content (RWC), relative growth rate (RGR), gas exchange parameters and effective PSII quantum yield (Y), and higher non-photochemical quenching (qN), water use efficiency (WUE), proline (PRO) and abscisic acid (ABA) accumulation, superoxide dismutase (SOD), ascorbate peroxidase (APX) activities as well as malondialdehyde (MDA) levels and electrolyte leakage in sun plants, whereas these changes were not significant in shade plants. Our study results implied that shade, applied together with drought, ameliorated the detrimental effects of drought. On the other hand, compared with the wet climate population, the dry climate population was more tolerant to drought in the sun treatment, as indicated by less decreases in A and mass-based leaf nitrogen content (Nmass), more responsive stomata, greater capacity for non-radiative dissipation of excitation energy as heat (analysed by qN), and higher WUE,higher level of antioxidant enzyme activities,higher ABA accumulation as well as lower MDA content and electrolyte leakage. Many of the differences in growth and physiological responses reported here are consistent with the climatic differences between the locations of the populations of P. asperata. 4. The effects of exogenous abscisic acid (ABA) application on the drought tolerance For both populations tested, exogenous ABA application increased root/shoot ratio (Rs) under well-watered and drought-stressed conditions, indicating that there was differential sensitivity to ABA in the roots and shoots. However, it appeared that ABA application affected the two P. asperata populations very differently during drought. CO2 assimilation rate (A) was significantly decreased in the wet climate population, but only to a minor extent in the dry climate population following ABA application during soil drying. On the other hand, ABA application significantly decreased stomatal conductance (gs), transpiration rate (E) and malondialdehyde (MDA) content, and significantly increased leaf mass per area (LMA), Rs, fine root/total root ratio (Ft), water use efficiency (WUE), ABA contents, superoxide dismutase (SOD), ascorbate peroxidase (APX) and catalase (CAT) activities under drought condition in the dry climate population, whereas ABA application did not significantly affect these parameters in the wet population plants. The results clearly demonstrated that the dry climate population was more responsive to ABA application than the wet climate population, as indicated by the strong stomata closure and by greater plasticity of LMA and biomass allocation, as well as by higher WUE, ABA content and anti-oxidative capacity to defense against oxidative stress, possibly predominantly by APX. We concluded that sensitivity to exogenous ABA application is population dependent in P. asperata. Our results provide strong evidence for adaptive differentiation between populations of P. asperata.

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中国沙棘是一种雌雄异株、风媒传粉的灌木或乔木,在中国西南的卧龙自然保护区有广泛的分布。本研究以采集于四川卧龙自然保护区5 个海拔(1800 m、2200 m、2600 m、3000 m、3400 m)梯度的中国沙棘天然群体为材料,以ISSR 和AFLP 标记技术研究其遗传多样性水平及其遗传结构,旨在了解卧龙地区中国沙棘天然群体的遗传多样性水平以及遗传多样性在群体间、群体内以及雌雄亚群体间的分布和特征,为中国沙棘树种的遗传改良及种质资源保存提供遗传研究背景与实验依据。同时探讨ISSR、AFLP 和RAPD三种标记对中国沙棘天然群体的遗传变异水平和群体间遗传结构的评估能力和各自的优缺点。研究得出以下主要结论: 1. ISSR和AFLP分析都表明卧龙自然保护区的中国沙棘群体拥有较高的遗传变异水平(h = 0.249,HT = 0.305)。出现这种结果的主要原因可能与卧龙自然保护区多变的气候条件和生境的异质度大有关。 2. ISSR 和AFLP 都揭示出卧龙自然保护区中国沙棘群体的遗传多样性随着海拔的增加发生显著的变化,表现为中海拔群体(2200 m 和2600 m)比高海拔群体(3000 m 和3400 m)和低海拔群体(1800 m)有更高的遗传多样性的趋势。出现这种趋势的可能解释是低海拔群体处在相对高温和相对干旱的环境,高海拔群体受到低温和紫外线胁迫,而中海拔群体存在中国沙棘生长的适宜环境。 3. ISSR 和AFLP 分析都表明:卧龙自然保护区中国沙棘的遗传结构遵循分布范围广、交配系统以异交为主的木本植物的通常模式,即大多数的遗传变异存在于群体内,只有少部分的遗传变异存在于群体间。 4. 经Mantel 检测表明,卧龙自然保护区中国沙棘群体间的海拔距离和对应遗传距离之间存在显著的正相关关系,即随着垂直海拔距离的增加,群体间的遗传距离也随之增加。Mantel 检测结果以及聚类分析将卧龙自然保护区5 个不同海拔的中国沙棘群体分为低、中、高海拔群体三组的研究结果都表明,海拔很可能是限制群体间基因交流的主要因素。 5. ISSR 分析发现同一海拔的雌雄亚群体首先聚类的研究结果表明,同一海拔的雌雄亚群体在遗传上最相似。方差分析结果表明只有3.8%的总遗传变异存在于雌雄亚群体间,这可能与雌雄植株间的交配和遗传物质的混合有关。 6. ISSR、AFLP 和RAPD 分析都表明卧龙自然保护区不同海拔的中国沙棘天然群体的遗传多样性水平较高。它们的分析结果估算得到的Nei's 平均基因多样度(h)分别为0.249、0.214 和0.170。从该结果可以看出ISSR 和AFLP 比RAPD 检测到更多的遗传多态性,这很可能是不同标记检测的基因组的位点不同所致。 7. 依据对不同标记系统的比较分析,认为ISSR、AFLP 和RAPD 三种分子标记系统都能成功地用于调查卧龙自然保护区不同海拔的中国沙棘群体的遗传变异水平及遗传变异结构,提供关于中国沙棘天然群体多态性水平和遗传变异分布的有用信息。在三者中,AFLP 具有最高效能指数和标记指数,在确定种间分类关系或鉴别个体方面是一种比较理想的标记。 Hippophae rhamnoides subsp. sinensis, a dioecious and deciduous shrub species,occupies a wide range of habitats in the Wolong Nature Reserve, Southwest China. Ourpresent study investigated the pattern of genetic variation and differentiation among fivenatural populations of H. rhamnoides subsp. sinensis, occurring along an altitudinal gradientthat varied from 1,800 to 3,400 m above sea level in the Wolong Natural Reserve, by usingISSR and AFLP markers to guide its genetic improvement and germplasm conservation. And,comparative study of ISSR, AFLP and RAPD was performed to detect their capacity toestimating the level and pattern of genetic variation occurring among the five elevationpopulations of H. rhamnoides subsp. sinensis, and to discuss their application to the study onplant genetics. The results were list following: 1. The ISSR and AFLP analysis conducted for the H. rhamnoides subsp. sinensispopulations located in the Wolong Natural Reserve of China revealed the presence of highlevels of genetic variation (h = 0.249, HT = 0.305). Besides such features as relatively widedistribution, dominantly outcrossing mating system, and effective seed dispersal by small animals and birds, it is sometimes argued that hard climatic conditions and heterogeneous habitats may also contribute to high levels of diversity. 2. Genetic diversity of H. rhamnoides subsp. sinensis populations was found to varysignificantly with changing elevation, showing a trend that mid-elevation populations (2,200m and 2,600 m) were genetically more diverse than both low-elevation (1,800 m) andhigh-elevation populations (3,000 m and 3,400 m). H. rhamnoides subsp. sinensis is thoughtto be stressed by drought and high temperature at low elevations, and by low temperature athigh elevations. The high genetic variability present in the mid-elevation populations of H.rhamnoides subsp. sinensis is assumed to be related to a greater plant density in the middlealtitudinal zone, where favorable ecological conditions permit its continuous distributioncovering the zone from 2,200 m to 2,600 m above sea level. 3. The genetic structure of H. rhamnoides subsp. sinensis revealed by ISSRs andAFLPs followed the general pattern detected in woody species with widespread distributionsand outcrossing mating systems. Such plants possess more genetic diversity withinpopulations and less variation among populations than species with other combinations oftraits. 4. In the present study, Mantel tests showed positive correlations between altitudinaldistances and genetic distances among populations or subpopulations. The observedrelationship between altitude and genetic distances, and the result of the cluster analysisincluding populations or male subpopulations and classifying the groups into three altitudeclusters suggest that altitude is a major factor that restricts gene flow between populationsand subpopulations. 5. The analysis of molecular variance showed that only 3.8% of the variability residedbetween female and male subpopulations. Such a very restricted proportion of the totalmolecular variance between female and male subpopulations is due to common sexuality andmixing of genetic material between females and males. 6. The analysis based on ISSRs, AFLPs and RAPDs all revealed relatively high levelsof genetic variation among different altitudinal populations of H. rhamnoides subsp. sinensisin Wolong Natural Reserve of China. Their estimates of mean Nei’s gene diversity is equal to0.249, 0.214 and 0.170 respectively, suggesting the higher capacity of detecting geneticvariation of ISSR and AFLP than RAPD. It might be ascribed to their distinct sensitivity todifferent type of genetic variation. 7. Based on the coparative study on ISSR, AFLP and RAPD, we drew a conclusion thatthey all successfully reveal some useful information concerning the level and pattern ofgenetic vatiation occurring among different elevation populations of H. rhamnoides subsp.sinensis. AFLP is a ideal tool to taxonomic study and individual identification for theirhighest efficiency index and marker index among the three marker systems.

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常绿阔叶林以其富饶的生物资源、丰富的生物多样性和巨大的生态与环境效益引起了人们越来越大的重视,它的研究已成为国际植被科学界关注的主题之一。我国分布着世界上面积最大的亚热带常绿阔叶林,在世界植被中具有重要地位,它的分布表现出明显的地带性差异,存在着多样的植物群系及其对应的气候特征。但是在植物功能性状领域,与全球范围其它生物群系相比,常绿阔叶林物种的研究较少,其功能性状间、功能性状与环境间的关系尚不清晰。 本研究以常绿阔叶林木本植物的当年生小枝为对象,试图从小枝水平上的生物量分配格局、叶片大小与数量的权衡关系、小枝茎的构型效应、叶片元素化学计量学,以及小枝大小的成本与效益分析等方面,较为系统地揭示小枝水平上的植物功能性状间及其与气候间的关系。因此,在华西雨屏带内部的不同纬度设置峨眉-青城-雷波-平武的温度梯度进行比较,并对有降水差异的川西南偏湿性(雷波)与偏干性常绿阔叶林(西昌)进行对比研究,同时在不同山体进行不同海拔梯度的比较研究。 本文主要研究结果如下: (1)小枝生物量分配格局叶水平上,叶片重-叶柄重(Y轴vs.X轴,下同)呈斜率小于1的异速生长关系,表明叶柄对叶内部的生物量分配影响显著。小枝水平上,叶和茎的生物量以及它们与小枝总生物量间基本呈等速生长关系,表明大的小枝或大叶物种不一定在叶生物量的分配上占优势。不同生活型间,在小枝或者茎的生物量一定时,常绿物种叶片的生物量比例较落叶物种稍高。与温度和水分较优越(峨眉及其低海拔)的生境相比,在相对低湿(螺髻)与低温(平武)的生境中的植物会减少对叶的投入而增加对支撑部分的投资比例。 (2)小枝叶片大小与数量的权衡无论是不同气候带还是不同生活型以及不同海拔梯度,叶片大小与出叶强度基本都是呈负的等速生长关系,表明了叶片大小-数量在小枝水平上的权衡。在不同气候梯度的对比中,叶片数量(出叶强度)一定时,高温和高水分生境(峨眉)比低温(平武)和低湿(螺髻山)生境中的物种的叶片大小(质量和面积)更大,表明不同生境的比较中,小的叶片可能具有较高的出叶强度和更高的适合度收益。“出叶强度优势”(Leafingintensitypremium)假说可能不适宜解释不同生境物种叶片大小差异。 (3)小枝茎的构型效应虽然茎长和茎径与叶片大小都呈正相关关系,与出叶强度都呈负相关关系,但茎长/茎径比与叶/茎生物量之比呈负相关关系;与叶片的大小呈负相关关系,与出叶强度呈正相关关系。这说明小枝构型能影响小枝叶/茎生物量分配和叶大小-数量的权衡关系。其影响机制可能是小枝内部的顶端优势。另外,茎长/茎径比在低湿和低温等不利生境中的植物中较高,而在降水和温度较适宜环境中较低。 (4)叶片C、N、P化学计量学N含量和P含量,C/N比和比叶重(LMA,leafmassperarea)呈正的等速生长关系,而N和LMA,P和LMA呈负的等速生长关系。在LMA一定时,C/N比随着生境胁迫压力的增加而降低,N、P含量随着生境压力的增加而增加。在P含量一定时,N含量随着生境压力的增加而降低,即N/P比在生境条件较优(峨眉及其低海拔)时较高。常绿和落叶植物叶片的N/P比没有差异,在LMA一定时,常绿植物的N、P含量较高、C/N比较低。总之,植物的C、N、P化学计量学特征受叶片属性如LMA与气候,及其相互作用的影响。 (5)小枝大小的代价与效益分析、TLA与小枝总重总叶面积(TLA,totalleafarea,Y轴,下同)与总叶重(X轴)均呈斜率小于1的异速生长关系,TLA与小枝横切面积呈斜率为1的等速生长关系。表明叶片面积的增加总是小于叶重和小枝总重的增加,随着小枝的增大,它的叶面积支撑效率下降。在热量和降水优越的生境(峨眉及其低海拔)中,相同小枝重或者相同茎横切面积的小枝,其叶面积支撑效率较低湿与低温环境下(螺髻山、平武及高海拔)的高。 总体上,本文初步研究了小枝水平上可能存在的以下三种权衡关系:叶-茎生物量分配权衡;叶片大小-数量的权衡;小枝茎长-茎径的权衡关系,以及气候要素等对这三种权衡关系的影响。在此基础上,我们还讨论了这些权衡关系的可能形成机制,及其与物种生态适应的联系。本研究丰富了生活史对策中关于权衡关系的研究内容,为我国常绿阔叶林功能生态学研究积累了材料。 Evergreen broad-leaved forests are attracting much more attention from vegetation ecologists than ever before because of their abundant nature resource and biological diversity, and also great ecological benefits. China has the largest distribution of subtropical evergreen broad-leaved forests (temperate rainforests) that are typical and representative in the world. The forests span over more than ten degrees in latitude and more than 30 degrees in longitude, providing an ideal place to study plant functional ecology, i.e., the climatic effect on plant functional traits and the relationship between the traits. However, relative to the other biomes, there are few studies addressing functional ecology of the plant species from subtropical evergreen broad-leaved forests. In this study, I focused on the leaf size-twig size spectrum of the woody species of subtropical evergreen broad-leaved forests in southwestern china. I collected data on leaf size and number, twig size in terms of both mass and volume, and stem architecture from five temperate mountains, and then I analyzed the relationships between leaf and stem biomass and between leaf size and number, the effect of stem length/diameter ratio on biomass allocation and on the relationship between leaf size and number, leaf C:N:P stoichiometry, and the twig efficiency of supporting leaf area in relation to twig size. I also addressed the climate effect on the spectrum. The temperature gradient from warm to cool sites was represented by Emei Mountain, Qingchengshan, Leibo, and Pingwu, and the rainfall gradient was assumed to emerge from the comparison between Leibo (High) and Luojishan (Low). In addition, altitudinal effects were analyzed with comparisons between low and high altitudes for each mountains. My main results are as follows. Isometric relationships were found between leaf mass and twig mass and between lamina mass and twig mass, suggesting that the biomass allocation to leaves or laminas was independent of twig mass. Petiole mass disproportionably increase with respect to lamina mass and twig mass, indicating the importance of leaf petioles to the within-twig biomass allocation. In addition, the investigated species tended to have a larger leaf and lamina mass, but a smaller stem mass at a given twig mass at favorable environments including warm and humid sites or at low altitude than unfavorable habitats, which might be due to the large requirements in physical support and transporting safety for the species living at unfavorable conditions. Moreover, the evergreen species invested more in leaves and laminas than the deciduous at given stem or twig biomass within any specified habitats. Negative, isometric scaling relationships between leaf number and size broadly existed in the species regardless of climate, altitude, and life forms, suggesting a leaf size/number trade-off within twigs. Along the climatic gradients, at given leaf number or leafing intensity, the leaves were larger in the favorable environments than the poor habitats. This suggested that the fitness benefit gained by small leaves could be larger than that with high leafing intensity in the stressful sites. I concluded that the “leafing intensity premium” hypothesis was not appropriate to interpreting between-habitat variation in leaf size. Both stem length and diameter were positively correlated to leaf size but negatively correlated to leafing intensity. The ratio of stem length to diameter was negatively correlated to leaf mass fraction, and it was negatively correlated to leaf size but positively correlated to leafing intensity. This suggested that the stem architecture influenced twig biomass allocation and the relationship between leaf size and number. The mechanism underlying the architectural effect might lie in the apical dominance within twig. Moreover, the ratio was greater in unfavorable habitats but smaller in favorable environments. Positive, isometric relationships were found between N and P contents per leaf mass, and between C/N ratio and leaf mass per area (LMA), but N and P contents scaled negatively to LMA. C/N ratio decreased but N and P increased with increasing habitat stress at a given LMA. N content declined with increasing habitat stress at given P content. These indicated that N/P and C/N were higher but LMA was lower in favorable habitats than in the other circumstances. The evergreen and deciduous species were non-heterogeneous in N/P, but the evergreen species have higher N and P contents and lower C/N than the deciduous ones. In general, C:N:P stoichiometry were related to both climatic conditions and other important functional traits like LMA. Total leaf area (TLA) allometricly scaled to leaf mass with a slope shallower than 1, similar to the relationship between TLA and total twig mass (leaf mass plus stem mass), suggesting that TLA failed to keep pace with the increase of leaf mass and twig size. However, TLA scaled isometricly to twig cross-sectional area. Thus, it could be inferred that the twig efficiency of displaying leaf area decreased with increasing twig size. In addition, the efficiency at a given twig size was large in favorable than unfavorable habitats. In general, in this preliminary study, I studied three tradeoff relationships within twigs, i.e., between leaf and stem biomass, between leaf number and size, and between stem length and diameter, as well as the climatic effect on the relationships. I discussed the mechanisms underlying the tradeoff relationships in view of biophysics and eco-physiology of plants. I believe that this study can serve as important materials advancing plant functional ecology of subtropical forest and that it will improve the understanding of life history strategies of plants from this particular biome.

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青藏高原东南缘由于特殊的生态地理条件,有着丰富的森林资源,这些资源是长江上游涵养水源、保持水土的生态屏障,是生物多样性的资源宝库。但随着过量的森林采伐,使该区曾经丰富的生物多样性资源遭受了前所未有的破坏,天然林的质量严重下降,生态系统退化,功能减弱。与此同时,许多物种的种群规模正在锐减,物种的遗传多样性也严重丧失。川西云杉是西部地区分布最广的云杉树种之一,在较高海拔的地区有着重要的生态学功能,是一种适应性很强的乡土树种。本项目采用简单序列重复标记(SSR)和特定序列位点(STS)研究不同生境条件下川西云杉群体的遗传变异及其时空分布格局,考察遗传变异与复杂的山地生态环境间的潜在联系,系统地揭示川西云杉天然群体与环境系统相互作用的生态适应与分子进化机制。研究成果能有效地为该树种遗传资源的科学保护与合理利用提供理论依据和科学指导,可为中国西南部亚高山天然林的可持续经营及退化生态系统的恢复与重建提供依据。主要研究结果如下: 1 STS和SSR两种分子标记的研究结果表明:川西云杉群体拥有中等水平的遗传多样性(基于SSR标记,平均He = 0.640;基于STS标记,平均He = 0.553)。造成这种中等水平的遗传多样性,可能是由于历史原因,川西云杉天然林被大量采伐,导致了遗传多样性的丧失。两种方法都检测出群体BT有着最高水平的遗传多样性。 2 两种标记的结果都一致说明:检测的10个川西云杉群体间遗传分化比较高,其存在群体间的遗传变异比例要明显地高于广泛分布的挪威云杉、横贯大陆的黑云杉以及兼有连续分布和不连续分布的西加云杉等,但要低于分布范围狭窄且呈不连续分布的粗枝云杉。青藏高原东南部的片段化生境可能是导致高水平遗传分化的主要原因。 3 基于两种标记的UPGMA 聚类和PCA分析结果,以及基于SSR标记的FCA分析结果都显示:群体BY遗传上明显区别于其它群体,其可能原因是青藏高原东南缘山脉阻隔而导致的生殖隔离的结果。 4 根据软件Bottleneck 1.2.02的检测以及不依赖哈迪一温伯格平衡的M比率检测结果:川西云杉种群很有可能经历了近期的遗传瓶颈效应。在本研究中遗传瓶颈效应并没有显著影响到物种的遗传多样性。然而,在这些片断化群体未来的子代群体中很可能出现遗传瓶颈导致的遗传多样性下降的效应。 Due to the extremely complex topography and climatic conditions, the southeast of the Qinghai-Tibet Plateau is region abundant in forest resource, which is benefit to the upper reaches of the Yangtze River water conservation, and protecting natural environment and biodiversity resources. However, by the reason of a plenty of trees were cut in these yeas, the biodiversity resource was destroy with degraded ecosystem and imperfect funcation. Picea balfouriana is one of the regionally distributed conifer species in the southeast of the Qinghai-Tibet Plateau and considered as a constructive species within its distribution area. It is an optimal species for the production of biomass. Moreover, it is well adapted to stressful environments at high altitude, especially to cold and drought conditions which are generally harsh for other trees. In our study, two types of molecular markers (SSR and STS) were used to estimate the genetic diversity and genetic structure of P. balfouriana populations originating from the southeast of the Qinghai-Tibet Plateau with varying climatic and geographical conditions. The results will not only provide a deep insight into its genetic diversity and population genetic structure but also valuable information for further management and breeding programs in P. balfouriana. In summary, the results obtained in this study revealed that: 1 A moderate degree of genetic variation is present in P. balfouriana in the southeast of the Qinghai-Tibet Plateau and it may caused by many trees were cut in the past. Population BT owns the highest level of genetic diversity by both two types of markers. 2 Considerable population differentiation exists among the ten P. balfouriana populations based on both SSR and STS markers, possibly caused by habitat fragmentation and heterogeneous environments. 3 The UPGMA clustering and PCA analyses based on SSR and STS marker, and FCA analyses based on SSR marker congruously separated population BY from other populations, which is likely due to the presence of mountain barriers. 4 The results of bottleneck analysis indicated that P. balfouriana populations that have undergone recent declines. In our research, the bottleneck effect don’t have a significant impact on the genetic diversity of species, however, the level of genetic diversity of P. balfouriana offspring populations growing in the fragmented habitats will decline in the future.

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除植被冠层的光合作用之外,土壤的呼吸作用是陆地生态系统碳收支中最大的通量。土壤呼吸即使发生较小的变化也能显著地减缓或加剧大气中CO2浓度的增加,从而明显影响到全球气候变化。土壤呼吸速率变化与否以及变化的方向可以反映生态系统对环境变化的敏感程度和响应模式。尽管如此,土壤呼吸仍是一个为人们了解不多的生态系统过程。 草地生态系统是陆地生态系统的一个重要组成部分。针对草地土壤呼吸进行野外实验研究和相应方法论的探讨将对区域乃至全球碳源汇性质的准确估算具有重要的科学意义。然而,近几年来关于草地土壤呼吸的主要研究工作都集中在温带草原和部分热带草原,而针对高寒草甸生态系统土壤呼吸的研究报道还很少。 2008年4月至2009年4月期间,我分别在2008年6、8、10、12月和2009年2月和4月分6次对川西北的典型高寒草甸群落的土壤呼吸进行观测,分析了不同类型高寒草甸群落土壤呼吸的季节变化特征以及环境因子和放牧模式对其影响。主要研究结果如下: 1)该地区高寒草甸生态系统在生长季(6月~8月)土壤呼吸速率较大(6.07~9.30μmolCO2¡m-2¡s-1 ) , 在非生长季( 12 月~ 2 月) 较小( 0.16 ~0.49μmolCO2¡m-2¡s-1 ) 。土壤CO2 年累积最大释放量为3963 ~ 5730gCO2¡m-2¡yr-1,其中,生长季土壤CO2的释放量占年总释放量的85%~90%。非生长季占10%~15%。非生长季所占比例略小于冬季积雪覆盖地区的冬季土壤呼吸占年土壤呼吸量的比例(14%~30%)。温度,尤其地温,是影响该地区高寒草甸生态系统土壤呼吸速率的最主要环境因子。土壤呼吸速率与地上生物量和土壤水分之间没有显著相关性,但是土壤含水量过大会导致土壤呼吸速率下降。 2)在观测期内,草丘区的土壤呼吸显著高于对照区的土壤呼吸,其最大土壤呼吸速率为16.77μmolCO2¡m-2¡s-1,土壤CO2 年累积最大释放量为8145gCO2¡m-2¡yr-1,是对照区的近2 倍。由于草丘在高寒草甸中占有较大的面积比例(近30%),因此,它将对高寒草甸生态系统的碳循环起着重要的作用。 3)放牧模式不仅可以影响高寒草甸群落的土壤CO2 排放,而且还可以改变土壤呼吸的温度敏感性(Q10)。本研究表明,在生长季有长期放牧活动干扰时将会增加土壤向大气中释放二氧化碳的速度,促使土壤碳库中碳的流失。禁牧样地的土壤呼吸速率在刚禁牧时先迅速增大,随着禁牧时间的延长土壤呼吸速率将会下降。此外,与其它放牧模式相比,冬季放牧将高寒草甸群落土壤呼吸速率在生长季达到最大值的时间明显向后推迟。不同放牧模式下高寒草甸群落土壤呼吸的Q10 值大小顺序为:禁牧一年群落>冬季放牧群落>禁牧三年群落>夏季放牧群落>自由放牧群落。 4)基于呼吸室技术的观测方法中,测量前的剪草处理可以明显改变该地区高寒草甸群落的土壤温度和土壤呼吸速率。在生长季,剪草处理将使土壤呼吸速率的瞬时响应增加90%左右。由于剪草处理明显增加了剪草样方白天的土壤温度,而土壤温度与土壤呼吸之间存在着极显著的指数相关关系,因而剪草处理导致土壤呼吸速率迅速增加。因此,在高寒地区基于呼吸室技术观测的土壤呼吸应当进行校正。 综上所述,川西北高寒草甸生态系统土壤呼吸速率在生长季较高,而在非生长季较低。土壤温度是影响该地区土壤呼吸的最主要环境因子。在实验观测期,草丘区土壤呼吸速率显著高于对照区的,是对照区土壤呼吸速率的近2倍。由于测量前的剪草处理可以明显改变待测点的土壤呼吸速率,因此,应对在高寒地区基于呼吸室技术观测的土壤呼吸进行校正。 Soil respiration is the second largest component (less than plant phtotosynthesis) of carbon dioxide flux between terrestrial ecosystems and the atmosphere. A minor change in soil respiration rate can significantly slow down or accelerate the increase of atmospheric CO2 concentration that is closely related to global climatic change. In turn, the change in the flux direction and rate of soil respiration may indicate the elasticity and stability of ecosystems to global changes and human disturbance. However, soil respiration is still an ecosystem process that has been poorly understood. Grassland ecosystem is an important component of the terrestrial ecosystem. Accurately estimating the CO2 flux from soil to atmosphere in situ is the key to evaluating the carbon resource and sink regionally or globally. Despite of extensive studies on the temperate and tropic grasslands, the soil respiration of alpine meadows has not substantially been measured. In the current study, soil respiration was measured for an annual cycle from April, 2008 to April, 2009 for the alpine meadow in northwestern Sichuan Province of China to determine the seasonal variation of soil respiration for the typical plant communities. The results are shown as follows: 1) Large seasonal variation of soil respiration was observed in the alpine meadow. The rate of soil respiration was the greatest (6.07~9.30μmolCO2¡m-2¡s-1) in June and the smallest (0.16 ~ 0.49μmolCO2¡m-2¡s-1) occurred from December to February in the non-growing season. The total emission of soil CO2 was estimated as 3963~5730 gCO2¡m-2¡yr-1, 85%~90% of which was released during the growing season, and 10%~15% during the non-growing season which was slightly less than the ratio of winter and annual CO2 flux from soil. Temperature, particularly the soil temperature, was the major environmental factor regulating the soil respiration. Significant and positive relationships were not found between soil respiration and soil moisture and between soil respiration and plant above-ground biomass, but excessive soil water content would decrease in the rate of soil respiration. 2) The rate of soil respiration in grass hummock communities was up to 16.77μmolCO2¡m-2¡s-1, which was about twice as great as in the controls (communities located in low and even sites). Considering the large proportion (about 30% on average) of hummock area in the meadow, it can be concluded that the hummocks played an important role in the carbon cycling of the study ecosystem. 3) Grazing patterns affected the flux of CO2 emission and the temperature sensitivity of soil respiration (Q10) in the alpine meadow. Grazing during growing season increased the rate of soil respiration. The rate of soil respiration increased significantly immediately after the alpine meadow being fenced, but thereafter decreased. In addition, grazing in winter delayed the peak respiration rate relative to the non-grazing mode. The Q10 value was the largest in the non-grazed area for one year, and next came the area with grazing in winter, followed by the non-grazed area for three years, the area with grazing in summer, and the non-limited grazed area. 4) In the chamber-based techniques, clipping manipulation before each measurement increased the transient rate of soil respiration by about 90% in the summer of the alpine meadow. As increase in soil temperature at daytime in the clipped plots by clipping and the exponential relationship between soil respiration and temperature, clipping manipulation led to increase in the rate of soil respiration. This suggested that a correction should be done for the techniques if employed in alpine and cold regions. In summary, the rate of soil respiration in the alpine meadow was the greatest in June and the smallest occurred from ecember to February in the non-growing season. Soil temperature was the major environmental factor regulating the soil respiration. The rate of soil respiration in grass hummock communities was up to 16.77μmolCO2¡m-2¡s-1, which was about twice as great as in the controls. A correction should be done for the techniques if employed in alpine and cold regions, because of the effect of clipping manipulation on soil temperature and respiration.

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中国拥有92466 Km2的各类高原湿地,具有湿地退化、过度放牧等相似特征,保护与利用矛盾突出。高寒湿地保护区尽管在制度上以核心区、缓冲区来约束当地的放牧等外来干扰行为,但在实际管理中却不能起到应有的作用。 本研究以四川若尔盖湿地国家级自然保护区为例,应用3S技术,建立保护区多功能动态分区工作流模型,通过不同植被类型的识别和空间特征分析、不同动物类群在上述植被生境中的时空分布特征分析、保护区主要干扰因素的时空分布特征分析,突出对保护区主要保护对象(湿地生态系统)的保护,对保护区进行管理分区,依据野生动物利用特征和植被生长特征对核心区进行年周期动态利用,缓解保护与发展的矛盾,促进保护区的优化管理。 应用归一化植被指数(NDVI)与植被盖度的相关性,将归一化植被指数(NDVI)转化为植被盖度指数(MDVI),结合保护区牧场划分和时空利用特征专家经验,结果表明,MDVI值在1-139之间主要代表着水体、裸地、沙地等;MDVI140-256为草地和高山灌丛;MDVI210是当地夏牧场和秋冬牧场的划分区间值。 合理的区划需要资金、技术和政策的支持,为保证保护区多核心动态分区的实施,本研究提出了生态工程、牧业发展方式转变、湿地特色产业发展、湿地政策、社区参与和科技支撑等六大保障措施。 In China, 92466 Km2 highland or frigid wetlands are (were) facing major management problems, such as wetland degradation and overgazing. Conflict between conservation and utilization on those wetlands can be found anywhere today. Although many nature reserves have been setup for protection of frigid wetland, and core and buffer zone has been declared to forbid any kinds of disturbance, local farmers still use these areas for grazing. As an example by Sichuan Roige Wetlands National Nature Reserve(SRWNRR), we set up a 3S flow model to analyze the character of year-round distribution patters of vegetation, wildlife, and grazing. Combined and overlapped these characters together, we select multi-core zone and buffer zone, then define a dynamic management period in different zone to optimize protection wetland in the reserve. Normalized Difference Vegetation Index(NDVI)is highly related with coverage of vegetation. When convert NDVI to MDVI (coverage index, 1-256), index 139 and 210 can be as inflexion to distinguish among water/sand/bared land, summer pasture, and autumn / winter pasture. We use these to select different layers and analyze grazing pattern. To be more realistic, we put forward some strategies to support our multi-core and dynamic management of wetland in Roige, including ecological restoration engineering, changing of stock raising industry, changing of wetland policy, community based management and technology renovation supports.