955 resultados para Yellow birch.
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Secondary metabolites produced by water-blooming cyanobacteria in eutrophic waters include some potent hepatotoxins, These compounds also have tumour-promoting properties, attributable to their inhibition and activation of protein phosphatases and kinases respectively. The inhibitory effect of these toxins on protein phosphatases have been employed in a commonly used radiometric assay, involving the use of a P-32-labeled substrate, for the detection and quantitation of these compounds. This paper investigates and describes a colorimetric method in which the activity of protein phosphatase 2A is determined by measuring the rate of colour production from the release of yellow p-nitrophenol using p-nitrophenyl phosphate as the substrate. Results of this study suggest that the colorimetric protein phosphatase inhibition assay is a simple, inexpensive tool for screening substances that may have tumour-promoting characteristics in aquatic systems. The detection limit of the colorimetric method is comparable to the radiometric assay. (C) 1998 Elsevier Science Ltd. All rights reserved.
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Thick GaN films were grown on GaN/sapphire template in a vertical HVPE reactor. Various material characterization techniques,including AFM, SEM, XRD, RBS/Channeling, CL, PL, and XPS, were used to characterize these GaN epitaxial films. It was found that stepped/terraced structures appeared on the film surface,which were indicative of a nearly step-flow mode of growth for the HVPE GaN despite the high growth rate. A few hexagonal pits appeared on the surface, which have strong light emission. After being etched in molten KOH, the wavy steps disappeared and hexagonal pits with {1010} facets appeared on the surface. An EPD of only 8 ×10~6cm~(-2) shows that the GaN film has few dislocations. Both XRD and RBS channeling indicate the high quality of the GaN thick films. Sharp band-edge emission with a full width at half maximum(FWHM)of 67meV was observed, while the yellow and infrared emissions were also found. These emissions are likely caused by native defects and C and O impurities.
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The persistent photoconductivity(PPC) phenomena in n-type GaN Films grown by metalorganic chemical vapor deposition(MOCVD) have been studied. After using some testing and analysis methods, such as the double crystal X-ray diffraction(DCXRD), the photolumineseence(PL) spectra, etc, it is found that the issue which influences PPC in n-type GaN is not relative to the dislocations and yellow band (YB), and is caused by the doping level of Si most likely.
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We observed yellow colored light emission bands from multiwalled carbon nanotubes in photo-luminescence (PL) experiments. The light emission band features indicate that the PL bands are associated with the electronic properties inherent to the carbon nanotube (CNT) structures.
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Photoluminescence (PL) was investigated in undoped GaN from 4.8 K to room temperature. The 4.8 K spectra exhibited recombinations of free exciton, donor-acceptor pair (DAP), blue and yellow bands (Ybs). The blue band (BB) was also identified to be a DAP recombination. The YB was assigned to a recombination from deep levels. The energy-dispersive X-ray spectroscopy show that C and O are the main residual impurities in undoped GaN and that C concentration is lower in the epilayers with the stronger BB. The electronic structures of native defects, C and O impurities, and their complexes were calculated using ab initio local-density-functional (LDF) methods with linear muffin-tin-orbital and 72-atomic supercell. The theoretical analyses suggest that the electron transitions from O-N states to C-N and to V-Ga states are responsible for DAP and the BB, respectively, and the electron transitions between the inner levels of the C-N-O-N complex may be responsible for the YB in our samples. (C) 2002 Elsevier Science B.V. All rights reserved.
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GaN epilayers grown on pre-nitridated (0001) sapphire substrates by metallorganic vapor phase epitaxy were investigated by wavelength dispersive X-ray spectroscopy and energy dispersive S-ray spectroscopy. Precipitates were observed to mainly consist of O impurity whose strengths were weaker than surrounding matrix. The precipitates were larger in size and distributed more sparsely and inhomogeneously in < 11-20 > directions of the epilayers grown on substrates pre-nitridated for longer periods. The larger precipitates often joined to cracks in the TEM specimens. The crack formation seems to be attributed to the compressive stress concentration at edge angles of the larger precipitates. Yellow luminescence of the epilayers was imaged by cathodoluminescence. The distribution similarity between the cathodoluminescence and the precipitates suggested that the precipitates were responsible for the yellow luminescence band. (C) 2000 Elsevier Science S.A, All rights reserved.
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随着工业化的发展,大气中二氧化碳的浓度(CO2)预测从现在的平均350μmol·mol-1升高到2030年的570μmol·mol-1,其增温作用将持续多个世纪。植被在大气二氧化碳减排以及调控区域水热状况过程中起重要作用,而其机理过程目前十分不清楚。本实验应用自控、封闭、独立生长室,研究了CO2浓度和温度升高对红桦根、茎、叶和枝可溶性蛋白含量和分配的影响,从蛋白水平上来解释川西亚地区的建群种-红桦对CO2升高和温度升高及其交互作用的响应规律,为全球气候变暖川西亚高山的植被保护和恢复提供理论依据。研究结果表明: 1. CO2浓度升高增加了可溶性蛋白的总量,改变了可溶性蛋白分配模式,即,可溶性蛋白分配到根的比例增加,分配到茎、枝、叶的比例减少。可能意味:在CO2浓度升高条件下,红桦根系的生长和营养物质吸收功能将会增强。 2. CO2浓度升高增加了根和茎的清蛋白含量,降低了叶片的清蛋白含量,叶片的球蛋白含量、醇溶蛋白含量和谷蛋白含量均增加。表明CO2浓度升高增加了清蛋白在根中积累,球蛋白、醇溶蛋白和谷蛋白大量在叶片中积累;前人研究所指出的CO2浓度升高使植物叶片可溶性蛋白的含量降低可能仅仅是由于清蛋白含量的降低造成的。 3. 温度升高使红桦幼苗整株所含可溶性蛋白总量增加,但可溶性蛋白总量的分配因红桦幼苗器官的不同而异。温度升高下根、茎、叶和枝的分配量分别占总可溶性蛋白的27.74%、35.57%、23.00%、13.68%,即茎>根>叶>枝。对照的根茎叶枝的分配量分别占总可溶性蛋白的21.01%、41.41%、23.08%、14.50%,即茎>叶>根>枝。表明温度升高使可溶性蛋白分配到根的比例增加,有利于根的可溶性蛋白的积累,增强了根吸收水分和矿质营养的能力,从而有利于根系的生长。 4. 温度升高处理下清蛋白和球蛋白在根中含量升高,在茎、叶和枝中含量下降,但没有达到显著水平;醇溶蛋白在根和叶中含量显著增加;谷蛋白在茎中的含量显著降低。表明温度升高增加清蛋白和球蛋白在红桦幼苗根部的积累,也有利于根和叶醇溶蛋白的积累,但不利于谷蛋白在茎的积累;温度升高条件下叶片可溶性蛋白升高是醇溶蛋白在叶片中积累的结果。 5. CO2浓度和温度同时升高条件下红桦幼苗的可溶性蛋白总量增加很少,只有分配到茎的可溶性蛋白比例增加,并且对可溶性蛋白分配规律没有影响。CO2和温度同时升高下红桦幼苗枝的可溶性蛋白含量的降低是可溶性蛋白总量的降低而不是碳水化合物稀释的结果,并且CO2和温度同时升高对红桦幼苗的生长没有明显的促进作用。 6. CO2和温度同时升高处理对可溶性蛋白含量有显著影响。清蛋白含量在根、茎、叶和枝中均降低,球蛋白含量在根中显著降低,醇溶蛋白含量在根、茎、叶和枝中均降低,谷蛋白含量在根中显著降低。表明CO2浓度和温度同时升高对根的影响显著,即降低了根的可溶性蛋白含量,可能导致根的吸收能力下降。 7. 因此,CO2和温度同时升高对可溶性蛋白影响不能简单地通过CO2和温度单因子影响机理来解释。 It is well known that atmospheric CO2 concentration and temperature are increasing as a consequence of human activities. Atmospheric CO2 concentration are predicted to increase from 350μmol·mol-1 now to 570μmol·mol-1 2030. And temperature will continue to increase for several centuries as a result of CO2 enrichment. Vegetation play a key role in reducing atmospheric CO2 and adapting and controlling warter and energy process in a certain region, while the underlying mechanism are not clear, yet. Betula albo-sinensis, as the dominating tree species of subalpine dark coniferous forest in west Sichuan province, play an important role in determing structure and function of forest ecosystem. In our study, effects of elevated atmospheric CO2 concentration (ambient±350±25μmol·mol-1), increased temperature (ambient±2.0±0.5℃) and their combination on contents and allocation of soluble protein were studied in independent and enclosed-top chamber system under high-frigid conditions. Chambers with ambient CO2 concentration and temperature are taken as control. The results are as the following, 1) Elevated atmospheric CO2 increased the accumulation of total weight of soluble protein in whole plant and changed allocation of soluble protein in red birch by increasing its allocation to roots and reducing its allocation to stem. This caused much more accumulation of soluble protein in roots which might help to prompt growth, development and nutrient absorption ability of roots. 2) Treatment EC increased content of albumin in roots and stems, reduced the content of albumin in leaves, and increased the content of globulin, promalin and glutenin in leaves. That is to say EC increased the accumulation of albumin in roots and accumulation of globulin, promalin and glutenin in leaves. The reduced soluble protein contents in plant leaves by EC, as reported by former researchers, are mainly resulted from the reduced content of albumin in leaves. 3) Elevated temperature increased the total of soluble proteins, but its allocation was dependent on organs. In treatment ET, roots, stems, leaves and branches take 27.74%, 35.57%, 23.00% and 13.68% of total weight of soluble protein. In treatment CK, roots, stems, leaves and branches take 21.01%, 41.41%, 23.08% and 14.50%. Elevated temperature changed allocation of soluble proteins in that it stimulated soluble proteins accumulation in roots and improved the uptake of water in roots. 4) Treatment ET increased the content of albumin and globulin in roots, and reduced the content of albumin and globulin in stems, leaves and branches. The content of promalin in roots and leaves was increased significantly, and the content of glutenin in stems was reduced significant. This suggested that ET stimulated the accumulation of albumin and globulin in roots and accumulation of promalin in leaves and roots; that treatment ET increased content of soluble protein in leaves was mainly resulted from the increased promalin content in leaves. 5) Regarding treatment ETC, the total of weight of soluble proteins increased, but not significantly; but increased in stems. So the combination of elevated atmospheric CO2 and temperature had not changed the allocation of soluble proteins in red birch seedling and reduced soluble proteins in branches were not the result of increased carbohydrate. 6) Treatment ETC reduced the content of albumin and promalin in roots, stems, leaves and branches, reduced the content of globulin and glutenin in roots significantly. That is to say elevated atmospheric CO2 and temperature reduced the content of soluble proteins in roots significantly which might help to prompt growth, development and nutrient absorption ability of roots. 7) The effects of elevated atmospheric CO2 and temperature on soluble protein cannot be simply interpreted through their mechanism that obtained when they were imposed on plant separately.
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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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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.
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辐射传输研究是贯穿森林生态系统的纽带,太阳辐射为植物的生长发育提供光合能量、适宜的环境温度以及发育信息。一方面,气候变化使到达地面辐射能的质和量发生变化,影响到植被的生长发育,改变森林的结构,而森林结构的变化又会影响林冠内辐射能的分配和质量,这些变化会进一步影响到林下土壤温度,改变森林根系活性以及土壤营养转化的效率;连锁反应的结果有可能会使森林生态系统的生产力发生变化,改变碳素和氮素源库的调节方向,从而反馈影响地球气候系统。另一方面,人类作为生态系统的成员,必然需要森林生态系统为其提供更多的原材料和更好的生态服务功能,如何实现这些目标,就需要人类适度调整干预方式和频度,达到预期的目的。本文在建立适合于川西亚高山森林的叶面积测量技术、光照辐射模型和土壤温度变化模型的基础上,对川西亚高山地带森林生态系统的辐射传输特征进行了分析,并从森林结构的角度探讨了林分内的辐射分布以及对土壤温度的影响。主要成果如下: 1. 提出了一种照相法测量叶面积的方法。通过对摆放在平面上的叶片照相,利用投影变化,把非正射图像转化为正射图像,然后经过计算机图像处理得到每一片叶片的面积、周长、长度、宽度等信息。这种方法可使用户以任意方向和距离拍摄处于平面上的叶片,能同时处理大量的叶片,适于野外离体或活体叶片测量。叶片面积分辨率可调,分辨率可以与常用的激光叶面积仪相近甚至更高,而且叶片图像可以存档查询。 2. 提出一种模拟林内光照变化的模型。利用林冠半球照片,记录视点以上半球内的林冠构件空间分布,作为林冠子模型;天空辐射子模型采用国际照明委员会(CIE)的标准晴天和阴天以及插值模型。该模型能够模拟林下某一位点处的实时光斑变化。 3. 提出一种土壤温度变化模型。把土壤视为具有容量和阻力性质的结构,利用电阻和电容器件构建土壤能量分布模型。外界太阳辐射能经过植被以及其它一些能量分配器后进入土壤,其中有一部分转化为土壤势能,即土壤温度。土壤温度的变化类似于电池的充放电过程。在已知模型参数的情况下,可以从太阳辐射计算土壤温度的变化。在模型参数未知的情况下,通过输入和输出值推算模型的参数,而模型参数中的时间常数与土壤组成和含水量有关,这样就可以知道土壤水分的变化情况。 4. 从王朗亚高山森林典型样地林分结构的测量获得林地三维结构图、树冠形态、叶面积密度等参数,这些参数输入到Brunner (1998)开发的tRAYci 模型中计算出一段时间内林分任意位置处的光照值。与林下辐射计测量值以及半球照片计算结果的比较,该模型基本上能够满足对林分光环境了解的要求。 5. 从川西亚高山森林生产力的角度,探讨了森林生产力研究的方法以及川西地区的研究历史和成果,发现了其中的一些规律和问题,特别是在叶面积测量上,还没有使用标准的叶面积指数定义。综合来看,川西地区针叶林叶面积指数(单位土地面积上植物冠层总叶面积的一半) 应在4-5 之间。降雨丰富的华西雨屏带是川西地区森林生产力最高的地区,而向西北森林生产力逐渐降低。川西地区云冷杉林森林生产力平均约为600 gDM m-2 a-1,但是根据辐射能计算的潜在生产力则达到1800 gDM m-2 a-1。实际与潜在森林生产力的巨大差异说明其它因子对生产力的影响。 6. 王朗亚高山3 个典型森林林分中,白桦林样地(BF) 林下草本以糙野青茅、牛至、紫菀等喜阳性物种为主,林下透光度较高;冷杉林样地(FF) 林下透光度最低,以喜阴性物种水金凤、蟹甲草、囊瓣芹等为主;而云杉林样地(SF)林分林龄最大,林下透光度介于冷杉林和白桦林之间,草本层仍然以喜阴性物种东方草莓、紫花碎米芥、酢浆草等为主。冷杉林和云杉林的灌木层也很丰富,卫矛属、五加属、茶藨子属、忍冬属植物很丰富,而在白桦林则以栒摘要子属、榛子属、鹅耳枥属等植物为主。藓类植物在云杉林中最丰富,并且形成毯状层,其它两个林分则很稀少。3 个样地林分结构与林下光环境有很强的相关性,从光环境特征可以在一定程度上推测林分的结构。各样地单纯从乔木层材积推算的NPP 排列顺序为BF>FF>SF,与林下辐射透射率和林分年龄的顺序相同,暗示辐射对群落演替过程的驱动作用。 7. 用半球照相法测得BF、FF 和SF 3 个样地的有效叶面积指数以SF 样地最高,BF 最低。如果考虑针叶树叶片在小枝上的丛聚分布,利用北方针叶林的数值进行校正,则SF 样地LAI 显著增加(达到89%),其它样地的LAI 基本不变甚至有所下降。校正后的数值与文献中地面测量的结果较相近,说明在使用半球照相法测量川西亚高山针叶林LAI 时必须加以校正。 8. 在3 个样地中,白桦、岷江冷杉和方枝柏种群为丛聚分布,紫果云杉在FF和SF 样地中基本上为随机分布。3 个物种出现丛聚分布的最短距离约为2m,在最短距离以内则为随机分布。最短距离可能与树冠大小有关,种子传播特征以及对光照的需求状况可能是造成这种分布格局类型的原因。 Radiative transfer plays a key role in forest ecosystems. Solar radiation providesenergy for photosynthesis, appropriate ambient temperature and development informationfor plants. However, quality and quantity of radiation reaching land surface are affected byweather and subsequently influence the growth and development of plants, which in turnchanges the budget of radiation in forest. Soil temperature changes with the variation ofradiation under forest canopy and influences the activity of roots and rate of nutrientturnover. Thus, any changes of radiation will induce chain reactions in the entireecosystem and display in the value of net primary productivity which will possibly shiftthe relationship between carbon source and sink at local or regional scale and feed back tothe global climate system. On the other hand, as a component of ecosystems, humanbeings of course need to demand more materials and better service from ecosystems. Forthese purpose, man must adapt their pattern and frequency of interference to ecosystems.This paper aims to research on the canopy structure, the radiation distribution and theirinfluence on soil temperature from the process of radiative transfer in subalpine forestecosystem of western Sichuan. The main results are: 1 Present a new photogrammetric method for leaf area. The main idea is to convertnon-vertically taken images of planar leaves to orthoimages through projectivetransformation. The resultant images are used to get leaf morphological parametersthrough image processing. This method enables users to take photos at almost anyorientation and distance if only the leaves are placed on same plane, and to processlarge quantity of leaves in a short time, which is suitable for field measurement. Theresolution of leaf area is adjustable to fit for special requirement. 2 A model using hemispherical photos combining with solar tracks and radiation courseis provided to simulate light variation in forest. The hemispherical photos of canopyrecord the real spatial distribution of each element of plants viewed from a point. Skyradiance is simulated with CIE standard clear sky or cloudy sky model. This modelcan be used to simulate real time light variation under canopy. 3 Present a soil temperature model. Soil could be regarded as a body of resistor andcapacitor. Some of the budget of solar radiation in soil body is transformed into soilpotential energy, the soil temperature. Variation of soil temperature is driven by solarradiation, vegetation, soil properties, etc. This model has two parameters, one of whichis time constant and is related to soil water content. The inversed model can be used tosimulate the variation of soil water. 4 By using model tRAYci developed by Brunner (1998), the 3-D distribution of light inthree subalpine forest stands of Wanglang Nature Reserve has been simulated andvalidated with value of radiometers in these stands. This model can basically satisfythe need for understanding light regimes of these stands. 5 Present some principles and questions of NPP (net primary of productivity) researchesin western Sichuan. The standard leaf area index (LAI) defined by Chen and Black(1997) has not been used in this region. Total leaf area and projected leaf area indexare still used in NPP researches which may differ around 1-fold in magnitude. Thestandard LAI which is a half of total leaf area above unit land area should be between4 and 5 for typical subalpine coniferous forest of western Sichuan concluded fromliteratures. The maximum forest NPP occurs in West China rain belt and decreasesnorthwestwards. Average NPP of spruce-fir forest in western Sichuan is about600gDM m-2 a-1, which is below the potential NPP of 1800gDM m-2 a-1 based onmeasured radiation in this region. The significant difference between potential and realNPP suggests that other factors influence the growth of stands. 6 In the three subalpine forest stands of Wanglang Nature Reserve, herbage layer ofAbstractbirch stand (BF) with age of 40 is dominated by heliophytes of Deyeuxia scabrescens,Origanum vulgare, Aster tongoloa etc.. However, both of the other two stands aredominated by shade tolerent species, such as Impatiens noli-tangere, Impatiensdicentra, Cacalia deltophylla and Pternopetalum tanakae etc. in fir stand (FF) withage of 180 and Fragaria orientalis, Cardamine tangutorum and Oxalis corniculata etc.in spruce stand (SF) with age of 330. Shrub species in the latter two stands arerelatively rich, typical dominant genera being Euonymus, Acanthopanax, Ribes andLonicera. Birch stand has relatively sparse shrubs dominated by genera of Cotoneaster,Corylus and Carpinus. Mosses are significant only in spruce stand. The canopystructure controls the light regime of stand, which influence the composition of herblayers beneath the canopy. This light regime-community structure relationship can beused to infer the herb community from canopy structure. The NPP derived from timbervolume of arbor layer of the three stands decreases from BF to SF, which is in thesame order of transmitted total radiation under canopy and age of these stands,suggesting the driving effect of radiation in the succession of community. 7 The highest effective LAI of the three stands obtained by hemispherical photos is inplot SF and lowest in plot BF. After rectification of the clumping effect of leaves onshoot, the real LAI in plot SF increases significantly (89%) and approximate to theaverage LAI of coniferous forest in western Sichuan. Therefore, the LAI obtainedfrom hemispherical photos needs rectification for clumping effect. 8 Spatial distribution pattern for Betula platyphylla, Abies faxoniana and Sabinasaltuaria is clumpy, but Picea purpurea almost random in plot FF and SF. The shortestdistance for clumpy distribution for Betula platyphylla and Sabina saltuaria is 1.5m,and 2m for Abies faxoniana. And random pattern for these trees is exhibited within thisrange which almost coincides with the diameter of crown. Seed dispersalcharacteristics and light requirement may be the reason for different spatial pattern.
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碳水化合物按其存在的形式可分为结构性碳水化合物和非结构性碳水化合物两种。前者主要用于植物体的形态建成;后者是参与植物生命代谢的重要物质。迄今为止,有关CO2浓度升高对植物叶片中的碳水化合物含量的研究较多,而对其它器官中碳水化合物含量以及碳水化合物在植物体内的分配响应研究较少。碳水化合物含量在植物各器官中的变化能够反映光合同化产物在叶和茎、枝和根中的转运情况;碳水化合物的分配与植物的生长模式相关,它的变化会对植物的生长情况产生影响。因此,为全面认识植物生理生化与生长过程对大气CO2浓度升高响应情况,需要对CO2浓度升高条件下植物体内碳水化合物的含量及分配变化进行深入的研究与探讨。本文应用自控、独立、封闭的生长室系统,研究了红桦幼苗根、茎、叶和枝的碳水化合物含量以及分配格局对大气CO2浓度升高(环境CO2浓度+350 µmol·mol-1) 的响应。研究结果表明:1) CO2浓度升高使红桦幼苗叶片中的非结构性碳水化合物含量显著增加。这可能会对光合作用造成反馈抑制,降低光合速率。2) CO2浓度升高使红桦幼苗根、茎和枝中的还原糖、蔗糖、总可溶性糖、淀粉和总的非结构性碳水化合物(TNC) 含量显著增加。说明CO2浓度升高促进了碳水化合物由叶片向枝、茎和根中的运输转移,支持了Finn和Brun的假设。3) 在总的非结构性碳水化合物(TNC) 中,淀粉所占比例最大。同样地,CO2浓度升高使TNC含量增加的部分中,淀粉所占的比例也最大。在叶片、枝、茎和根中淀粉含量增加部分占TNC含量增加部分的91.45%、88.23%、83.23%和82.01%。4) CO2浓度升高使红桦幼苗根、茎、叶和枝内的纤维素含量有增加的趋势,但未达到显著水平。需要进一步研究长期CO2浓度升高下,纤维素含量的响应程度。5) CO2浓度升高使碳水化合物在红桦幼苗体内的分配发生了改变。红桦幼苗体内碳水化合物分配变化的一致趋势是由地上部分向地下部分分配转移。其中,测定的所有碳水化合物均向根中分配增多。同时,CO2浓度升高使红桦幼苗的根冠比显著增大;根系干重显著增加。这些结果支持了Gorissen 和Cotrufo的假设,即碳水化合物向根中分配增多是根冠比增大的主要原因。6) CO2浓度升高使红桦幼苗体内的氮含量显著下降。氮含量的下降可能主要是由生长的加快和TNC (主要是淀粉) 含量的增加对氮的稀释造成的。Carbohydrates found in plants are frequently grouped into two different classes:structural carbohydrates and non-structural carbohydrates. The former mainlyconstruct the plant basic framework, while the latter are essential for plant growth andmetabolism. As yet there is lack of information on the effects of elevated CO2concentration on carbohydrate contents in stem, branch and root of plant, and oncarbohydrate allocation in organs of plant although there have been many reports onthe responses of carbohydrate contents to elevated CO2 concentration in plant foliages.A shift of carbohydrate contents in plant reflects a change in transporting ofphotosynthetic production from leaf to stem, branch and root of plant. The allocationof carbohydrates that is correlated to plant growth patterns affects plant growth. Thus,in order to understand the influences of elevated CO2 on biochemical process,physiological change and plant growth well, the response of carbohydrate contentsand allocation in plant to elevated CO2 should be further investigated. In our study, theeffects of elevated CO2 on carbohydrate contents and their allocation between leaf,stem, branch and root tissue of Betula albosinensis seedlings were determined. Theseedlings were grown in independent and enclosed-top chambers. Chambers werecontrolled to reproduce ambient (CK) and ambient + 350 µmol·mol-1 CO2 (EC)concentration for 1 year. The results here showed that,1) Elevated CO2 significantly increased non-structural carbohydrate contents in leafof red birch seedlings. This will reduce photosynthetic rate.2) Elevated CO2 also significantly increased non-structural carbohydrate contentsin root, stem and branch of red birch seedlings. These findings supported thehypothesis that elevated CO2 accelerated carbohydrates from leaf to branch, stem androot.3) Starch makes up the largest parts of total non-structural carbohydrate. In thesame way, the increase of starch plays a main role in the increase of totalnon-structural carbohydrate under elevated CO2. In leaf, branch, stem and root, theincrements of starch contents comprised 91.45%, 88.23%, 83.23% and 82.01% of theincrements of total non-structural carbohydrate contents.4) Under elevated CO2 the cellulose contents have an increasing tendency in redbirch seedlings. It is needed to investigate the effects of long-term elevated CO2 oncellulose contents in plant.5) There are significant CO2 effects on the allocation of carbohydrate in organs ofred birch seedlings. Under elevated CO2 more carbohydrates were allocated to root.Moreover, CO2 enrichment significantly increased the root to shoot ratio of red birchseedlings and the dry weight of roots. These results supported Gorissen and Cotrufo ‘shypothesis that increase of carbohydrate allocation to root mostly contributed to theincrease of root to shoot ratio.6) Elevated CO2 brought about a reduction in the nitrogen contents of leaf, stem,branch and root. The decline of nitrogen contents under elevated CO2 is mainlycaused by the dilution effects of increasing starch level and growth of red birchseedlings.
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对隆肛蛙属的物种构成进行了订正,建立新属肛刺蛙属Yerana gen. nov.;订正后的隆肛蛙属现仅隶2种, 即隆肛蛙F. quadrana和太行隆肛蛙F. taihangnicus。运用形态学分析探讨了隆肛蛙属物种及种群的形态差异和分类关系,通过分子系统学研究探讨了隆肛蛙属物种及种群的分类和系统发育关系,运用动物地理学方法结合系统发育关系探讨了隆肛蛙属种群的地理分布格局成因与历史过程。主要结果和推论如下: 1.隆肛蛙属物种构成的订正及一新属建立 建立新属肛刺蛙属,将隆肛蛙属中的原叶氏隆肛蛙F. yei归隶新属肛刺蛙属并更名为叶氏肛刺蛙Y. yei,,新属建立的主要依据为:(1)雄性肛部隆起,肛孔下方有两个布满黑刺的大的白色球形隆起,具单咽下内声囊, 第一指具婚刺;(2)形态量度分析表明叶氏肛刺蛙与隆肛蛙和太行隆肛蛙的形态差异远大于后两者之间的差异;(3)叶氏肛刺蛙的分布区与隆肛蛙和太行隆肛蛙的分布区距离较远且呈隔离状态;(4)分子系统学研究资料(Jiang et al.,2005)证明叶氏肛刺蛙与隆肛蛙和太行隆肛蛙非单系发生;叶氏肛刺蛙在第二支中位于基部。因此,隆肛蛙属现仅隶2种,即隆肛蛙和太行隆肛蛙。 2.隆肛蛙属种群形态学研究 对隆肛蛙属中隆肛蛙和太行隆肛蛙的15个地理种群565只标本的28项形态性状进行了测量,运用典型判别分析法对其分析的结果表明:(1)太行隆肛蛙与隆肛蛙形态差异明显,支持其为不同的物种;(2)原隆肛蛙河南伏牛山种群和山西中条山种群应为太行隆肛蛙的地理种群;(3)隆肛蛙不同地理种群之间形态差异明显,其中四川安县种群、陕西周至种群和湖北利川种群与模式产地重庆巫山种群的差异可能达到了亚种或亚种以上分化水平。对隆肛蛙属量度分析的15个种群进行定性形态分析表明其分为三种形态型,对应隆肛蛙、过渡型和太行隆肛蛙,其变异特征主要为内跗褶、雄性肛部隆起及疣粒分布、第五趾外侧缘膜等,这与量度分析结果相似。 3.隆肛蛙属种群分子系统学研究 测定隆肛蛙属Feirana的2种19种群的线粒体12S rRNA和16S rRNA基因片段、ND2基因的DNA序列,比对后共计1953bps。(1)遗传多样性与距离分析:结果表明,隆肛蛙属种群具很高的遗传多样性,19个种群样品表现出19种单倍型(遗传多样性指数Hd=1.0); ND2基因的进化信息含量远高于12SrRNA和16SrRNA。隆肛蛙属2种群组内的种群间的遗传距离远小于两种群组间的距离,种群在不同基因上的遗传距离表现的关系与对应的系统树一致。(2)系统发育关系分析:结果表明,不同基因片断基于不同方法构建的隆肛蛙属种群系统发育树结构基本一致,基本表明隆肛蛙属种群为单系发生;它们在系统树中分为两大支,分别对应于隆肛蛙和太行隆肛蛙;支持中条山种群(沁水、历山和济源种群)和伏牛山种群(栾川和内乡种群)为太行隆肛蛙的地理种群,而原隆肛蛙秦岭中东段的部分种群(柞水、宁陕、长安大坝沟种群)也应为太行隆肛蛙的地理种群。(3)亚种分化分析:根据遗传距离分析和系统发育关系分析结果,并考虑形态上的差异情况以及地理分布信息,隆肛蛙所隶种群组可分为2亚种,即隆肛蛙指名亚种F. quadrana quadrana包括四川盆地东缘大巴山东段-巫山-武陵山北麓种群和秦岭中段(周至板房子和长安广货街)种群,他们在系统关系树上聚为一支;安县亚种F. quadrana anxianensis包括四川盆地西缘岷山东麓-龙门山-大巴山和秦岭西段的种群(安县、青川、文县、南江和凤县种群),他们在系统关系树上聚为一支。太行隆肛蛙所隶种群组也可分为2亚种,即太行隆肛蛙指名亚种F. taihangnicus taihangnicus包括中条山的种群(沁水、历山和济源种群)和中东秦岭的部分种群(柞水、长安大坝沟和宁陕种群),他们在系统关系树上聚为一支;太行隆肛蛙伏牛亚种F. taihangnicus funiuensis,为伏牛山地区的种群(栾川和内乡种群),他们在系统关系树上聚为一支。 4.隆肛蛙属种群动物地理学研究 隆肛蛙属19种群的分歧年代分析: 以长江巫山段和黄河三门峡段的形成历史时期为参考点,根据已测隆肛蛙属19种群及其外群包括N. pleski、P. yunnanesis、P. robertingeri、F. limnocharis的1953bps DNA序列构建分子钟,获得各支系的分歧年代。结果表明:①棘蛙族在70Ma左右开始其独立演化历程,这与Roelants et al.(2004)的分析结果~60±15Ma左右开始分化基本一致,后者印证了本文的分子钟。②隆肛蛙属的起始分化年代较早,隆肛蛙和太行隆肛蛙两种群组的最近祖先种群大概在46Ma~50Ma左右;隆肛蛙和太行隆肛蛙种群组内的种群分化年代相对两种群组间晚得多, 隆肛蛙种群组内两亚种分化起始年代约为10Ma左右,而太行隆肛蛙种群组内两亚种分化起始年代约为6Ma。 隆肛蛙属种群分布格局形成过程分析: ①隆肛蛙属的系统关系与地理分布格局密切相关,大部分系统分支分级与地理距离成正比;②隆肛蛙属最近祖先种群的分化中心可能位于秦岭中部地区, 隆肛蛙属的种群分布格局的形成表现为隔离分化与扩散相结合的机制,由隔离分化产生的隆肛蛙祖先种群主要从秦岭中部向西南方向扩散,后隔离分化为两亚种;太行隆肛蛙祖先种群向东北方向扩散也分化为两亚种。 隆肛蛙属种群分布区域地质历史的探讨:本文所建分子钟和种群分化方式印证了该区域的几次主要地质事件,包括岷山-龙门山-西秦岭等地区的快速差异隆起、第四纪冰期等。 The specific composition of the genus Feirana should be revised. A new genus Yerana gen. nov.(Ranidae:Dicroglossinae)was established based on morphological data-set and molecular phylogeny, as a result, only two species F. quadrana and F. taihangnicus are classified into Feirana now. Morphological differences and taxonomy of populations of Feirana were investigated based on morphological and morphometric data; phylogenetic relationships and taxonomy of populations of Feirana were elucidated using molecular data, and then the proceeding of the distribution pattern of populations of Feirana were discussed. The main results and conclusions and proposals were presented as following: 1. Revising of the specific composition of the genus Feirana and establishment of a new genus The new genus Yerana, only containing the type species Y. yei, was established based on the following evidences: (1) In adult male, distinct up-heaved circular vesicle presents around the anal, and under anal there are two white balls on which black spines exist, black horny spines scatter on the upper side of first finger, and internal single subgular vocal sac presents; (2) there is obvious morphometric differences between Yerana and Feirana; (3) Yerana is distributed far from Feirana; (4) evidences of molecular phylogeny(Jiang et al.,2005)suggested that Yerana take a special phylogenetic clade which is different from other genus included in the tribe Paini. As a result, there are only two species in Feirana, i.e., F. quadrana and F. taihangnicus. 2. Morphological research of populations of Feirana Twenty-eight characters of 565 individuals of 15 populations of the genus Feirana were measured, the results of Canonical Discriminant analysis of the morphometric data-set indicated that: (1) there are very prominent differences between the two species F. quadrana and F. taihangnicus. The validity of species F. taihangnicus was approved here; (2) Mt. Funiu population and Mt. Zhongtiao population should belong to the species F. taihangnicus; (3) Obvious differences exist among 12 populations of F. quadrana, the differentiation among Zhouzhi population, Anxian population, Lichuan population, and Wushan population together with the others probably reach sub-specific or specific level. Result of morphological comparison between 15 different populations show that 3 morphological types are recogenized in according with F. quadrana, F. taihangnicus and intergradation, this result conform to the result of morphometric analysis. 3. Molecular phylogenetic study on populaions of Feirana Fragment of 12SrRNA and 16SrRNA genes, and ND2 gene of 19 populations of two species of Feirana were sequenced and aligned, from which 1953 bps were received. (1) analyses of genetic distance and hereditary diversity indicated that: genetic distance between populations in each group were less than distance between two groups of Feirana, 19 haplotypes were recognized from 19 samples of 19 populations, so the hereditary diversity of populations of Feirana was very high (Hd=1.0), phylogenetic information in ND2 gene is more than fragment sequence of 12SrRNA and 16SrRNA genes. (2) Result of molecular phylogeny indicate that the phylogenetic trees constructed using different methods based on different sequence data sets showed the revised genus Feirana is monophyletic since the 19 populations of Feirana were firstly clustered together as one large clade, which was further clustered into two major clades, corresponding to F. quadrana(GroupⅠ) and F. taihangnicus(GroupⅡ), respectively. So populations of Qinshui and Lishan in Mt. Zhongtiao, populations of Luanchuan and Neixiang in Mt. Funiu, and populations of Zhashui, Dabagou of Chang’an and Ningshan in eastern Mt. Qinling should belong to the species F. taihangnicus; (3) Subspecific differentiation. on the basis of genetic distance, phylogenetic trees and geographical distribution, F. quadrana should have two subspecies, i.e., F. quadrana qudadrana, consisting of the populations Guanghuojie of Chang’an and Zhouzhi in Mid-Mt. Qinling, populations in Wushan area and northern Mt. Wuling (Lichuan), and F. qudadrana anxianensis, consisting of the populations in eastern Mt. Ming shan-Mt. Longmen-western Mt. Daba-western Mt. Qinling (Anxian, Qingchuan, Wenxian, Nanjiang and Fengxian); F. taihangnicus should also has two subspecies, i.e., F. taihangnicus taihangnicus, consisting of the populations in Mt. Zhongtiao and eastern Mt. Qinling, and F. taihangnicus funiuensis, consisting of the populations in Mt. Funiu. 4. Zoogeography of populaions of Feirana Analysis for divergent time of 19 populations of Feirana: Using the dates of run-through of Wushan segment of Changjiang River as the time when the population of Lichuan started differentiated from the populations of Wushan and Shennongjia, and the dates of Sanmenxia segment of Yellow River as the time when the populations in Mt. Zhongtiao started differentiated from the population of Dabagou in Chang’an, molecular clock was established using sequences with 1953 bps of 19 populations of Feirana and outgroup including N. pleski, P. yunnanesis, P. robertingeri, F. limnocharis in order to estimate divergent time of all clades. Result of that indicated that: ① the tribe Paini started to evolve independently at about 70Ma when is in consistent with that estimated by Roelants et al.(2004)with result of about ~60±15Ma, they were corroborated by each other, this confirms the validity of this molecular clock; ② divergent time for speciation of Feriana is early, ancestral populations of F. quadrana and F. taihangnicus were found about 46Ma~50Ma; differentiation of populations within species is greatly late to the divergence of the two species, divergent time for F. quadrana is 10Ma and divergent time for F. taihangnicus is 6Ma. Proceeding of distribution pattern of Feirana. Phylogenetic relationships of populations of Feirana matched quite with distribution pattern of them, the relationships among clades showed in phylogenetic trees is direct ratio to geographical distance of them; the estimated date of speciation between two species of Feirana was as early as speciation of Paa yunnanesis and Nanara pleski; middle part of Mt. Qinling is the center of speciation of Feirana, combination of mult-events of dispersal and vicariance are probably the mechanism of speciation of Feirana, F. quadrana colonized the mid-Mt. Qinling and then differentiated into two subspecies in southwest direction, ancestral population of F. taihangnicus colonized the mid-Mt. Qinling and then differentiated into two subspecies in northeast direction. On geological history of the distribution of Feirana. According to molecular clock and speciation model of populations of Feirana, some geological events are confirmed, including special rise of Mt. Minshan- Mt. Longmen-western Mt. Qinling, glacial age.
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沙蜥属(Phrynocephalus)的卵胎生类群主要分布在我国青藏高原,包括南疆沙蜥(P. forsythii)、西藏沙蜥(P. theobaldi)、红尾沙蜥(P. erythrurus)、贵德沙蜥(P. putjatia)和青海沙蜥(P. vlangalii)。其卵胎生生殖方式适应了高寒生境,与青藏高原隆升有关。纵观前人的研究,上述几种卵胎生沙蜥的分类、系统发育关系以及生物地理都还存在疑问。本文研究了分布在若尔盖湿地的青海沙蜥红原亚种(P. v hongyuanensis)以及分布在黄河上游其它地区青海沙蜥种组的地理分布格局,并探讨了其形成机制。 青海沙蜥在黄河上游主要分布于若尔盖湿地以及青海湖周边地区。若尔盖湿地青海沙蜥红原亚种的生境由于沼泽的形成被切割成不连续的斑块,通过遗传分析可以推测这种特殊生境对它们遗传结构的影响。其次,贵德沙蜥、青海沙蜥的青海湖周边各居群以及若尔盖湿地居群之间的系统地理格局还未见报道。因此本文以居群为单位,将它们作为一个复合体,通过系统地理研究,可以了解其种群遗传结构,据此分析相关的地质历史事件对其分布的影响。主要结果如下: 1. 若尔盖湿地青海沙蜥红原亚种的种群遗传结构: 共研究了三个地理单元(红原(HY)、辖曼(XM)、玛曲(MQ))的7个采集点的72个个体。所有ND4-tRNALeu序列比对得到785 bp的片断,定义了9种单倍型。结果显示总的核苷酸多样性较低,单倍型多样性较高。分子变异分析(AMOVA)显示3个单元间差异显著(P<0.01),遗传变异主要存在于地理单元间,占62.61%。除MQ单元,XM各居群与HY居群混杂在一起,单倍型网络图没有显示出单倍型和地理位置的对应关系。XM单元单倍型的不配对分布(Mismatch distribution)为明显左移的单峰,且Fu’s Fs test为负值,表明XM单元可能经历了近期种群扩张,有足够的时间积累单倍型的多态性,还不足以大幅提高核苷酸多样性,这是其单倍型多样性较高和核苷酸多样性较低的原因。MQ单元遗传多样性低而与其他单元显著分化,推测这与3万年前黄河在若尔盖玛曲之间贯通有关。近期沼泽的形成对XMb居群的隔离时间短,使得其遗传多样性低但还不足以形成大的遗传差异。无论黄河的贯通还是沼泽的形成其隔离形成的时间都不长,其作用改变了单倍型出现的频率,也出现了一些特有单倍型,但共享单倍型还广泛存在,还不足以使得不同居群之间形成较大的遗传距离。 2. 黄河上游青海沙蜥种组的分布格局与地史过程的关系: 黄河上游青海沙蜥种组包括贵德沙蜥、青海沙蜥指名亚种的青海湖周边各居群、青海沙蜥红原亚种若尔盖湿地居群、以及青海湖以西的部分居群(序列由Genbank下载获得),总计22个居群189个样品。所有ND4-tRNALeu序列比对得到703个位点,定义了39种单倍型。以南疆沙蜥为外群构建的贝叶斯树以及MP法构建的无根树,都分为A、B两大组。其中A包括若尔盖湿地居群以及玛多居群(A1)、青海湖以西的居群和兴海居群(A2)、西藏沙蜥;B包括青海湖以南的居群和天祝居群(B1)、青海湖以东北的居群(B2)。单倍型网络图分别对应了系统发育树上的各支。按照系统发育结果分组进行分子变异分析,得到组间变异占88.63%,各组间差异显著(P=0.000)。种群遗传结构分析得到,A1和B2可能经历了近期的种群扩张,前者扩张时间约为0.105-0.189 Ma B.P.(million years before present),后者为0.057-0.102 Ma B.P.,可能与末次间冰期的气候变暖有关。A2和B1对应的两个地理单元都具有较强的种群遗传结构,较为稳定。 青海沙蜥种组A、B两大支之间遗传距离大,分化明显,分化大约发生在4.29-2.38 Ma B.P.,推测青藏运动的A幕运动后复杂的地形变化可能是它们产生分化的原因。B1和B2分化大约发生在1.73-0.96 Ma B.P.,这与湟水流域构造运动发生的时间相符。在早、中更新世时期,B1支内部各居群可能有交流,中更新世末共和盆地出现的抬升以及河流溯源改道等事件可能是引起这支内部多个单倍型丢失的原因。A1、A2支的分化可能与倒数第三次冰期降临之后气候变冷、阿尼玛卿山的大冰帽有关。 The viviparous group of genus Phrynocephalus is mainly distributed in the Qinghai –Tibetan Plateau, including P. forsythii、P. theobaldi、P. erythrurus、P. putjatia and P. vlangalii. These species are adapted well to the cold clime there, and the origin of this group was the result of a vicariance event associated with the uplifting of the Qinghai -Tibetan Plateau. Although many works have been done, there are still several questions about classification、phylogenetic relationships and the biogeography of this group. The phylogeographic pattern of the P. vlangalii complex on the upper reaches of the Yellow River and the P. v. hongyuanensis in Zoige Wetland were studied in this thesis. On the upper reaches of the Yellow River, P. vlangalii complex are distributed in Zoige Wetland and the southeast and northeast region of Kuku-noor Lake. Because of the forming of the wetland in Zoige, the habitats for sand lizards are divided into many discontinuous ones, and it is necessary to analyze genetic structure in these unique habitats. The phylogeographic patter among P. putjatia、populations of P. vlangalii in the southeast region of Kuku-noor Lake and populations of P. vlangalii in Zoige Wetland hasn’t been studied yet, and the complicated geological events of the Plateau may play an important role in the populations’ diversity and species forming there. So these populations were gathered as a complex, and phylogeographic analysis were used to clarify these doubts. According to the two topics above, this thesis has two parts of results as follows: 1. Three geographic units of P. vlangalii hongyuanensis in Zoige Wetland were defined, and they were Xiaman (XM)、Hongyuan (HY) and Maqu (MQ). 785bp fragments of the mtDNA ND4-tRNAleu were determined from 72 samples and nine haplotypes were identified. As a whole, the nucleotide diversity was low,but the haplotype diversity was high. Analysis of molecular variance (AMOVA) showed that the three units were distinctly different(P<0.01),and 62.61% of the total genetic diversity was attributable to variation among units. There were 3 haplotypes shared among XM and HY,and no geographic clustering was observed except MQ from the TCS network. The results from the mismatch distribution analysis and Fu’s Fs test implied that there might be a recent population expansion in the XM unit, and this may be the reason why XM had a high haplotype diversity but a low nucleotide diversity. We estimate that the MQ and XMb have lower diversities because of some very recent geographic events, such as the formation of the Yellow river’s upriver and the Zoige Wetland. Although they are distinctly different, not enough time has passed for them to have diverged a great genetic distance. 2. 189 samples in 22 populations of P. vlangalii complex were collected, including P. putjatia、populations of P. vlangalii in the southeast and northeast region of Kuku-noor Lake、 populations of P. vlangalii in Zoige Wetland and the data from Genbank. 703bp ND4-tRNALeu sequences identified 39 haplotypes. P. forsythii was selected as outgroup, and both the Bayesian tree and the MP unrooted tree were divided into two groups(A、B). A included populations in Zoige Wetland and Xinghai(A1)、populations in the west of Kuku-noor Lake(A2)、P. theobaldi, and B included populations in the southeast of Kuku-noor Lake and Tianzhu(B1)、populations in the northeast of Kuku-noor Lake(B2). The haplotype network agreed with these groups. AMOVA showed that these five groups were distinctly different(P<0.01), and 88.63% of the total genetic diversity was attributable to variation among groups. There might be recent population expansion in A1 and A2, which corresponded to the dry climate of the last interglacial period. The expansion times were 0.189-0.105 Ma B.P. and 0.102-0.057 Ma B.P., respectively. A2 and B1 had strong genetic structure. The large genetic distance between A and B showed that they had been separated from each other for a long time(about 4.29-2.38 Ma B.P.), and it corresponded to the A phase of Qingzang Movement. The diversity between B1 and B2 at 1.73-0.96 Ma B.P. may be caused by the geological event in Huangshui valley. In early Pleistocene, populations in B1 may have gene flow because of geographic linkage, and later the uplift of the Plateau and the change of river route there made a few haplotypes lost. A1 and A2 were divided into two parts by A’nyemaqen Mountains at 0.66-0.37 Ma B.P., which maybe corresponded to glaciations at about 0.7 Ma B.P.
Resumo:
本研究对自育小麦白粉病抗源“07鉴126”和条锈病抗源CD1437、CD0534-5进行抗性遗传分析和微卫星引物的筛选,建立了与PmCD1和YrCD抗病基因连锁的SSR分子标记,主要研究结果如下: 1.小麦白粉抗源“07鉴126”抗白粉病基因的鉴定和分子标记的建立 品系“07鉴126”对我国目前白粉菌强优势生理小种E09、E11和其它多种小种表现免疫或高度抵抗。Pm-sus是07鉴126的自然突变感病株。利用“07鉴126”和Pm-sus的F2抗病性分离群体进行抗条锈病性遗传分析和分子标记定位,结果表明,“07鉴126”的白粉抗性为显性单基因控制的全生育期抗性,暂命名为PmCD1;并筛选到了与PmCD1共分离的显性SSR分子标记Xbarc183。系谱分析和分子标记分析表明PmCD1来源于荆州黑麦。抗谱分析表明PmCD1不同于已知的黑麦抗白粉基因,是一个新的抗白粉病基因。Xbarc183这一分子标记的建立为PmCD1的分子标记辅助选择和抗病基因累加提供了方便。 2.小麦条锈抗源CD1437抗条锈病基因的鉴定和分子标记的建立 利用对优势条锈菌小种条中32免疫的小麦品系CD1437及其自然突变感病株Yr-sus杂交构建F2、F3抗病性分离群体。抗条锈病性遗传分析结果显示,1437的抗条锈性为显性单基因控制的全生育期抗性,该基因暂命名为YrCD。SSR分析发现,位于1B染色体上的7个SSR标记Xcfd65、Xgwm11、Xgwm18、Xbarc187、Xwmc406、 Xwmc419、Xwmc216依次分布在YrCD的一侧,与YrCD的遗传距离在1.7 cM至9.2 cM。YrCD和YrCH42的等位性分析显示二者可能为等位基因。YrCD和Yr24、Yr26的抗谱相似。系谱分析和分子标记分析表明贵农20是YrCD的供体。本研究推测YrCD、Yr24、Yr26和YrCH42可能是等位基因,并推测Yr-sus是缺失突变体。 3. 小麦条锈抗源CD0534-5抗条锈病基因的鉴定 利用对条中32免疫的小麦抗条锈病品系CD0534-5及其感病重组自交系CD0534-4建立F2抗病性分离群体。抗条锈病性遗传分析表明,CD0534-5的条锈抗性由两对独立的显性主效基因控制。用BSK法分析,发现其中一对基因与SSR分子标记Xgwm11、Xgwm18、Xwmc128、Xwmc419连锁,该基因是来源于贵农20的YrCD。另一抗性基因来源贵农19,是极有利用价值的未知抗性基因。 This study focused on the investigation and identification of a novel powdery mildew resistant gene PmCD1 in wheat lines 07jian126 and stripe rust resistant gene YrCD in wheat lines CD1482 and CD0534-5, and screened SSR markers tightly linked to them. The main results were as follows: 1.Identification and SSR markers screening of a novel powdery mildew PmCD1 in wheat line 07jian126. Using a Pm resistant wheat line 07jian126 and its Pm susceptible mutant, a F2 population was constructed. Pedigree and genetic analyses indicated that the Pm resistance in 07jian126 was tranderred from rye (Secale cereale L.) cv. Jinzhou and was controlled by a single dominant gene. Differential test using 21 Bgt isolates revealed that the Pm resistant gene in 07jian126 is novel and was temporarily designated as PmCD1. A dominant SSR marker Xbarc183/130 bp was found co-segregated with PmCD1 in the F2 population. The diagnostic band of Xbarc183/130 bp co-segregating with PmCD1 could be used as an ideal marker in marker-assisted-selection during wheat breeding program. 2. Identification and SSR markers mapping of yellow rust resistant gene YrCD in wheat line CD1437. Wheat line CD1437 was highly resistant to predominant Chinese stripe rust race CYR32 at both seedling and adult stages. A F2 population was developed from the cross of CD1437 and its Yr susceptible mutant Yr-sus. Genetic analysis indicated line CD1437 contains a single dominant gene, temporarily designated YrCD. Seven SSR markers on the chromosome 1BS including Xcfd65, Xgwm11, Xgwm18, Xbarc187, wmc406, Xwmc419and Xwmc216 were close linked to YrCD with a genetic dsitance 1.7 cM to 9.2 cM. YrCD came from wheat cultivar Guinong 20. Allelic test of CD1437 and Chinese cultivar Chuanmai 42 indicated that YrCD and YrCH42 were allelic. Reaction patterns of YrCD and Yr24, Yr26 to 21 PST isolates were the same. These results suggested that YrCD and Yr24, Yr26, YrCH42 might be allelic. 3.Detection and identification of yellow rust resistance genes in wheat line CD0534-5 Wheat line CD0534-5 was highly resistant to predominant Chinese stripe rust race CYR32, while its recombinant inbred line CD0534-4 was susceptible. Genetic analysis with a F2 population developed from the cross of CD0534-5 and CD0534-4 indicated line CD0534-5 contains two independent dominant genes. Four SSR markers on the chromosome 1BS including Xgwm11, Xgwm18, Xwmc128, Xwmc419 were found to linked with one gene in CD0534-5. According the locations of makers and pedigree, this gene in CD0534-5 was YrCD, from cultivar Guinong 20. Another resistant gene was from cultivar Guinong 19, different with those genes on 1B such as Yr10, Yr15, Yr5 etc, was a valuable resistant gene in wheat breeding.