22 resultados para Polygonum multiflorum

em Chinese Academy of Sciences Institutional Repositories Grid Portal


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The 70% EtOH extract of Polygonum cuspidatum showed inhibitory action against HIV-1-induced syncytium formation at non-cytotoxic concentrations in vitro with a 50% effective concentration (EC50) of 13.94 +/- 3.41 mu g/mL. Through bioactivity-guided fractionation, 20 phenolic compounds, including eight stilbenoids, were isolated from the roots of Polygonum cuspidatum, and their anti-HIV-1 activities were evaluated. Results showed that compounds 1, 13, 14, and 16 demonstrated fairly strong antiviral activity against HIV-1-induced cytopathic effects in C8166 lymphocytes at non-cytotoxic concentrations, with EC50 values of 4.37 +/- 1.96 mu g/mL, 19.97 +/- 5.09, 14.4 +/- 1.34 mu g/mL, and 11.29 +/- 6.26 mu g/mL and therapeutic index (TI) values of 8.12, > 10.02, > 13.89, and > 17.71, respectively. Other compounds showed either weak or no effects. Compound 6 also showed weak inhibition (153.42 +/- 19.25 mu g/mL); however, it possesses very good water solubility and showed almost no cytotoxicity (> 2000 mu g/mL), therefore achieving a fairly good TI (13.04). The activities of the two compounds (3 and 18) from Polygonum multiflorum were also assayed. The relationship between molecular structures and their bioactivities was also discussed.

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Study on the antioxidant systems of Polygonum viviparumgrown at two different altitudes indicated plants grown at Haibei Research Station at 3200 m altitude as compared with plants grown in Xining at 2300 m altitude had apparently higher contents of ultroviolet-absorbing compounds and ascorbic acid, and significantly higher activities of superoxide dismutase, peroxidase and ascorbic peroxidase. Higher contents of superoxide radical anions and malonadehyde were also found in plants at Haibei Research Station as compared with the plants grown in Xining which have been transplanted from Haibei Research Station for at least four years. The differences in antioxidant system reflect a long term of time of adaptation to different environments.

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Grazing by domestic herbivores is generally recognized as a major ecological factor and an important evolutionary force in grasslands. Grazing has both extensive and profound effects on individual plants and communities. We investigated the response patterns of Polygonum viviparum species and the species diversity of an alpine shrub meadow in response to long-term livestock grazing by a field manipulative experiment controlling livestock numbers on the Qinghai-Tibet Plateau in China. Here, we hypothesize that within a range of grazing pressure, grazing can alter relative allocation to different plant parts without changing total biomass for some plant species if there is life history trade-offs between plant traits. The same type of communities exposed to different grazing pressures may only alter relative species' abundances or species composition and not vary species diversity because plant species differ in resistant capability to herbivory. The results show that plant height and biomass of different organs differed among grazing treatments but total biomass remained constant. Biomass allocation and absolute investments to both reproduction and growth decreased and to belowground storage increased with increased grazing pressure, indicating the increasing in storage function was attained at a cost of reducing reproduction of bulbils and represented an optimal allocation and an adaptive response of the species to long-term aboveground damage. Moreover, our results showed multiform response types for either species groups or single species along the gradient of grazing intensity. Heavy grazing caused a 13.2% increase in species richness. There was difference in species composition of about 18%-20% among grazing treatment. Shannon-Wiener (H') diversity index and species evenness (E) index did not differ among grazing treatments. These results support our hypothesis.

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Ⅰ 虎杖聚酮类化合物生物合成相关基因的克隆及功能分析 虎杖 (Polygonum cuspidatum Sieb. et Zucc) 属于蓼科蓼属多年生草本植物,在中国和日本民间曾被广泛用于动脉粥样硬化、高血压、咳嗽、化脓性皮肤炎以及淋病的治疗,具有祛风利湿、散瘀定痛、止咳化痰等功效。而在现代医学上最令人瞩目和具有发展前景的是其在抗肿瘤、心血管保护、抗氧化方面的作用,相关疗效主要来自于虎杖中结构迥异、种类丰富的聚酮化合物及其衍生物资源。这些聚酮类化合物主要包括蒽醌、大黄素、大黄素-甲醚、大黄酚、芪类以及类黄酮化合物等。其中,大部分聚酮类化合物生物合成途径机制尚不明确,但可以肯定的是植物类型III聚酮合酶type III polyketide synthases (PKSs) 在这些聚酮化合物的生物合成起始反应中行使着关键的作用。因此,除了我们所熟悉的类黄酮化合物、芪类化合物之外,进一步分离和分析虎杖中其它重要聚酮类化合物生物合成所涉及的类型III聚酮合酶基因的是非常值得期待的。 目前,已经有14个植物类型III PKS基因被克隆和功能分析。植物类型III PKS的共同特征包括基因结构、序列相似性、保守的活性中心、酶学性质以及共同的催化机制等。显花植物(裸子植物和被子植物)中,植物类型III PKS的基因结构绝对保守,除了一个早期报道的金鱼草(Antirrhinum majus)查尔酮合酶chalcone synthase (CHS) 含有第二个内含子外,迄今为止所有已知的植物类型III PKS基因均含有一个内含子且该内含子位置保守。有趣的是,在本研究中,两个含有3个内含子的类型III PKS基因从虎杖中被分离,且两个基因3个内含子的位置完全保守,这是三内含子类型III PKS基因首次得到分离。除了新奇的基因结构外,体外功能分析显示上述两个基因还具有特殊的酶学性质和功能。 本论文围绕上述2个三内含子基因开展了以下工作: 虎杖中一个由三内含子基因编码的新型类型III聚酮合酶 一个类型III PKS的cDNA及其相应的基因(PcPKS2)从药用植物虎杖中被克隆。序列分析结果表明,PcPKS2的开放阅读框被3个内含子分隔,这是一个出人意料的发现,因为截至到目前为止,除了金鱼草一个CHS基因外,所有已知的类型III PKS基因均在固定位置上含有一个内含子。除了特殊的基因结构外,PcPKS2显示了一些有趣的特性:(i) CHS“守卫”苯丙氨酸——Phe215和Phe265在PcPKS2中双双缺失,它们分别被亮氨酸和半胱氨酸取代;(ii) 体外功能分析结果表明,当酶促反应体系的pH值为6.5-8.5时,大肠杆菌中过表达的重组PcPKS2高效地合成丁烯酮非环化产物——4-香豆酰甘油酸内酯(4-coumaroyltriacetic acid lactone (CTAL))为主产物,而丙烯酮非环化产物bis-noryangonin (BNY) 以及苯亚甲基丙酮为副产物;而当酶促反应体系的pH值为9.0时,PcPKS2高效地合成苯亚甲基丙酮为主产物,而CTAL、BNY为副产物。另外,除了上述3种产物外,在不同的pH条件下,还有痕量的柚皮素查尔酮能被检测到。此外,在4-香豆酰辅酶A(4-coumaroyl-CoA)的类似化合物中,除了4-香豆酰辅酶A外,只有feruloyl-CoA能够被PcPKS2接受作为起始底物。PcPKS2不接受脂肪酰辅酶A——异丁酰基辅酶A(isobutyryl-CoA)、异戊酰基辅酶A(isovaleryl-CoA)以及乙酰辅酶A(acetyl-CoA)作为起始底物。Southern blot杂交结果表明,在虎杖基因组中存在2-4个PcPKS2基因的拷贝。Northern blot杂交结果表明,在根茎和幼叶中,PcPKS2表达量很高,而在根中无表达。叶中的PcPKS2的表达受病原菌诱导,但不受伤诱导。 虎杖中一个编码双功能类型III聚酮合酶的三内含子基因的鉴定 显花植物中,所有已知的类型III PKS 基因均含有一个内含子且位置绝对保守。本研究中,综合运用PCR技术,从富含聚酮类化合物的植物虎杖中克隆得到一个类型III PKS 基因(PcPKS1)及其cDNA。序列分析结果表明,PcPKS1含有3个内含子。系统发育分析结果表明,PcPKS1与其它植物的CHSs归为一类。然而,体外功能分析结果表明,当酶促反应体系pH值为7.0时,大肠杆菌中过表达的重组PcPKS1高效地合成柚皮素查尔酮(naringenin)为单一产物;而当pH值为9.0时,苯亚甲基丙酮(p-hydroxybenzalacetone)几乎为重组PcPKS1的唯一产物。后续的研究表明,与典型的CHSs相比,PcPKS1具有另外一些不同的特点:在pH值为9.0时(PcPKS1的苯亚甲基丙酮合成活性最适pH值),在4-香豆酰辅酶A的类似化合物中,只有feruloyl-CoA能够被PcPKS1接受作为起始底物。与CHSs展现出的对脂肪酰辅酶A宽泛的底物特异性不同,在不同的pH条件下,PcPKS1不接受异丁酰基辅酶A(isobutyryl-CoA)、异戊酰基辅酶A(isovaleryl-CoA)以及乙酰辅酶A(acetyl-CoA)作为起始底物。以上数据指出重组PcPKS1是一个具有查尔酮合酶(CHS)和苯亚甲基丙酮合酶(BAS)活性的双功能酶。Southern blot杂交结果表明,在虎杖基因组中存在2-4个PcPKS1基因的拷贝。Northern blot杂交结果表明,PcPKS1可能在防御病原菌和草食动物方面起着重要作用。PcPKS1和PcPKS2共同从虎杖中被分离的事实极有可能暗示了苯丁烷类化合物(phenylbutanoid)及其衍生物存在于虎杖中。 Ⅱ 高山红景天酪醇生物合成代谢途径机制研究 高山红景天(Rhodiola sachalinensis A. Bor)是景天科(Crassulaceae)红景天属多年生草本植物,作为一种适应原性中草药在中国的应用史已经超过800年。最近红景天提取物作为一种重要的商业药用制剂资源,其应用遍及欧洲、亚洲和美国,其主要治疗范围包括抗变应性和消炎,提高心理机敏性等。目前已经非常明确,红景天甙(salidroside)和甙元酪醇(tyrosol)是红景天属植物的主要功效成分,主要分布于这类植物的根中并且具有抗缺氧、抗疲劳、延缓衰老、预防紫外线辐射伤害等功效。红景天甙为酪醇8-O-β-D葡萄糖甙,是酪醇在葡萄糖基转移酶UDP-glucosyltransferase (UGT) 的催化下糖基化后形成的,可以认为是酪醇在植物体内的贮存形式。酪醇作为一种重要的活性分子,同样存在于橄榄树和葡萄酒中。 虽然已经非常明确酪醇来自于莽草酸代谢途径,然而其具体的生物合成途径及其调控仍不明确。总结以往的报道,在酪醇的生物合成上主要存在两种观点:一是酪醇可能来自于苯丙烷代谢途径产生的4-香豆酸(4-coumaric acid)前体;二是来自于酪氨酸的酪胺(tyramine)可能是酪醇生物合成的直接前体。我们的工作兴趣主要围绕着鉴别高山红景天中的酪醇生物合成途径展开: 高山红景天内源苯丙氨酸解氨酶PALrs1的过表达对红景天甙积累的影响 红景天甙是来自于药用植物高山红景天的一种适应原性新型药物,其生物合成途径可能起始于苯丙氨酸或酪氨酸。由于高山红景天野生植物资源的匮乏和相对含量很低,阐明红景天甙的生物合成途径对于增加红景天甙的供给至关重要。在我们以前的工作中,运用cDNA末端快速扩增技术(RACE),一个编码苯丙氨酸解氨酶phenylalanine ammonia-lyase (PAL)的cDNA从高山红景天中被克隆,命名为PALrs1。在本研究中,PALrs1置于35S启动子+Ω增强子序列的控制下通过农杆菌(Agrobacterium tumefaciens)介导法转化回高山红景天。PCR 和 PCR–Southern blot分析结果表明,PALrs1已经整合到了转基因植物的基因组上。Northern blot杂交结果表明,PALrs1已经获得在转录水平上的高水平表达。与预期的结果相同,高效液相色谱High-performance liquid chromatography (HPLC)测定结果显示PALrs1的过表达引起4-香豆酸含量增长3.3倍。然而,与之相反的是,酪醇和红景天甙含量与对照相比反而分别下降4.7和7.7倍。此外,我们发现PALrs1的过表达造成酪氨酸含量下降2.6倍。这些数据暗示着PALrs1的过表达和4-香豆酸的积累并不能促进酪醇的生物合成。酪醇,作为一种苯乙烷类衍生物并非来自苯丙氨酸,而酪氨酸含量的下降则极有可能是酪醇生物合成和红景天甙积累大规模下降的直接原因。

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植物源杀菌剂的开发应用以及从植物中寻找杀菌活性物质作为先导化合物,是目前杀菌研究领域的热点之一。本文以蓼科植物虎杖(Polygonum cuspidatum Sieb.et Zucc.) 为材料,研究了虎杖提取物的杀菌活性和作用机理,确定了虎杖中的有效杀菌活性成分,并以此为先导化合物进行了衍生物合成与结构活性关系的研究。 不同溶剂提取物制备与杀菌活性测定结果表明,乙醇适合作为虎杖植物杀菌剂的提取溶剂,提取率高,对多种植物病原真菌具有广谱的杀菌和抑菌活性,除对黄瓜白粉病(Sphaerotheca fuliginea)表现出很好的防治效果外,对苹果腐烂病菌(Valsa mali)、玉米小斑病菌(Helminthosporium maydis)、葡萄炭疽病菌(Colletotrichum gloeosporioides)、小麦赤霉病菌(Fusarium graminearum)、油菜菌核病菌(Sclerotinia sclerotiorum)、水稻纹枯病菌(Rhizoconia solani)等也具有很好的抑制作用。虎杖回流提取物对黄瓜白粉病的杀菌作用以保护作用为主,兼具一定的治疗作用,并且具有一定的内吸活性,持效期约为4-7 d。温室试验结果表明,虎杖乙醇回流提取物10%可溶性液剂对黄瓜白粉病的EC90值为172.83 mg/L,田间小区试验表明该制剂在800-1600 mg/L的浓度下,对黄瓜白粉病的防效达到76.3-93.4%,具有较好的应用前景。 对苹果腐烂病菌的抑菌作用机理表明,虎杖乙醇提取物对该病原菌有明显的抑制作用,能够抑制蛋白质、葡萄糖等菌体细胞内物质的合成,从而使病菌代谢速度减慢,抑制其生长。虎杖提取物还能够使几丁质酶和β-1,3葡聚糖酶这两种细胞壁相关水解酶的活性升高,降解细胞壁而破坏菌体结构,使菌体自溶。 过测定虎杖乙醇回流提取物对黄瓜体内等一些防御酶和病程相关蛋白活性的影响,表明在40 mg/L和400 mg/L浓度下,虎杖乙醇提取物能够使黄瓜叶片内的过氧化物酶(POD)、多酚氧化酶(PPO)、苯丙氨酸解氨酶(PAL)、几丁质酶等不同程度的升高,从而在一定程度上提高植物对病原真菌的抗病能力。 通过生物活性跟踪测定以及pH梯度提取法确定了虎杖中的主要杀菌活性成分为蒽醌类化合物大黄素(emodin)和大黄素甲醚(physcion),结构通过了HPLC-MS和1H NMR确认,并且通过HPLC确定了虎杖乙醇回流提取物中二者的含量分别为3.28%和1.11%。 以虎杖中的有效成份之一的大黄素为原料,通过羟基的甲基化反应合成了包括已知物大黄素甲醚在内的11个大黄素衍生物,其中5个化合物为首次报道,并进行了初步结构活性关系研究。结果表明通过对大黄素3-OH位置以短直链烷基取代,其衍生物对黄瓜白粉病的活性大大提高,其中以甲基取代的衍生物大黄素甲醚的活性为母体大黄素的16.7倍,而以取代苄基修饰的衍生物的活性没有明显提高。一些目标化合物的活性明显优于三唑酮。研究中还意外发现大黄素的甲基化衍生物三甲氧基大黄素在4000 mg/L时能够明显抑制甜菜夜蛾幼虫的取食与生长发育,而大黄素和大黄素甲醚则无此作用。

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

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瑞香狼毒(Stellera chamaejasme L.)是瑞香科(Thymelaeaceae)狼毒属的一种多年生野草,有毒。据调查,从20 世纪60 年代开始至今,狼毒在青藏高原东缘的高寒草甸上不断蔓延、密度不断变大,在一些地段甚至成为优势物种。有关狼毒在高寒草甸蔓延的生态系统效应的研究尚未见报道。本文从系统碳、氮循环的角度,分别研究狼毒在生长和非生长季节对高寒草甸生态系统的影响。同时,从花粉化感的角度,深入研究狼毒对当地同花期物种有性繁殖的影响。系统地研究高寒草甸生态系统物质循环过程,特别是非生长季节微生物和土壤碳氮库的动态变化,有助于揭示狼毒在系统物质循环方面的“物种效应”以及这种效应的季节变化,为丰富有关高海拔生态系统,特别是其非生长季的物质循环的科学理论做出贡献。同时,碳氮循环和花粉化感的研究还有助于深刻地理解狼毒作为一种入侵性很强的杂草的特殊的蔓延机制,从而为狼毒的有效防治、高寒草甸的科学管理提供依据。 针对狼毒在青藏高原高寒草甸上蔓延的生态系统碳氮循环方面的影响,开展以下2 方面的研究:(1)在生长季,研究松潘县尕米寺附近(北纬32°53',东经103°40',海拔3190 m)的两种地形(平地和阳坡)条件下狼毒对土壤碳氮循环影响及可能的原因。狼毒和其它几个主要物种(圆穗蓼(Polygonummacrophyllum D. Don var. Macrophyllum),草地早熟禾(Poa pretensis L.),四川嵩草(Kobresia setchwanensis Hand.-Maizz.),鹅绒委陵菜(Potentilla anserina L.var. anserine)和鳞叶龙胆(Gentiana squarrosa Ledeb.)的地上凋落物产量以及地上凋落物和根的化学组成被测量。在有-无狼毒斑块下,各种土壤的库(比如,铵态氮、硝态氮、无机磷和微生物生物量)和周转率(包括净矿化、净硝化、总硝化、反硝化和微生物呼吸速率)被测量和比较。(2)在非生长季节,尤其是春季冻融交替期,选取了两个研究地点——尕米寺和卡卡沟(北纬32°59',东经103°41',海拔3400 m),分别测定有狼毒和无狼毒斑块下土壤微生物生物量碳和氮、可溶性有机碳和氮以及铵态氮和硝态氮的动态变化。同时,分别在上述两个地点有-无狼毒的样地上,研究6 个主要物种(狼毒、圆穗蓼、草地早熟禾、四川嵩草、鹅绒委陵菜和鳞叶龙胆)从秋季开始、为期1 年的凋落物分解过程。 针对狼毒花粉化感对同花期其它物种可能的花粉化感作用开展以下工作:在实验室中,用一系列浓度的狼毒花粉水浸提液对与它同花期的其它物种以及自身花粉进行测试,测定花粉萌发率;在野外自然条件下的其它物种的柱头上施用上述浓度的狼毒花粉水浸提液,观测种子结实率,同时,观察狼毒花粉的种间花粉散布数量。 生长季节的研究结果表明,狼毒地上凋落物氮含量比其它几个主要物种更高,而木质素-总氮比更低。狼毒显著地增加其斑块下表层土壤中有机质的含量,而有-无狼毒的亚表层土壤在有机碳和总磷方面没有显著差异。狼毒表土中硝态氮含量在平地和阳坡比无狼毒土壤分别高113%和90%。狼毒表土中微生物生物量碳和氮量显著高于无狼毒表土。无论是平地还是阳坡,狼毒土壤的总硝化和微生物呼吸速率显著高于无狼毒土壤;而它们的反硝化速率只在平地有显著的差异。狼毒与其它物种间地上凋落物的产量和质量的差异可能是导致有-无狼毒土壤碳氮循环差异的原因。我们假设,狼毒可能通过增加贫氮生态系统土壤中的有效氮含量提高其入侵能力。 非生长季的研究结果表明,在青藏高原东缘的高寒草甸上,土壤微生物生3物量在11 月的秋-冬过渡期达到第一个峰值;在春季的冻融交替期,微生物生物量达到第二个峰值后又迅速降低。无机氮以及可溶性有机碳氮与微生物生物量有相似的变化过程。微生物碳氮比呈现显著的季节性变化。隆冬季节的微生物生物量碳氮比显著高于生长旺季的微生物碳氮比。这种变化可能暗示冬、夏季微生物的群落组成和对资源的利用有所不同。有-无狼毒斑块下土壤微生物和土壤碳、氮库一般只在秋-冬过渡期有显著差异,有狼毒土壤微生物生物量和土壤碳、氮库显著高于无狼毒土壤;而在之后的冬季和春季没有显著差异。所有6 个物种凋落物在非生长季分解率为24%-50%,均高于生长季的10%到30%。其中在秋-冬过渡期,凋落物开始埋藏的两周时间内,分解最快,达10%-20%。不同物种凋落物全年的分解率和分解过程有显著差异。圆穗蓼在全年的分解都较缓慢(非生长季26%,生长季15%),草地早熟禾和四川嵩草等全年的分解速率比较均匀(非生长季和生长季均为30%,非生长季略高),而狼毒在非生长季分解较快(约50%),而在接下来的生长季分解变得缓慢(约12%)。所有物种的凋落物氮含量在非生长季下降,而在随后的生长季上升。 实验室的花粉萌发试验证明,狼毒花粉对自身花粉萌发没有自毒作用,而其它受试的所有物种(圆穗蓼,秦艽(Gentiana macrophylla Pall. var. fetissowii),湿生扁蕾(Gentianopsis paludosa (Hook. f.) Ma var. paludosa),鳞叶龙胆,椭圆叶花锚(Halenia elliptica D. Don var. elliptica),蓝钟花(Cyananthus hookeri C. B.Cl. var. grandiflorus Marq.),小米草(Euphrasia pectinata Ten.),川西翠雀花(Delphinium tongolense Franch.),高原毛茛(Ranunculus tanguticus (Maxim.)Ovcz. var. tanguticus)和鹅绒委陵菜)的花粉萌发率随着狼毒花粉浸提液浓度的增加呈显著的非线性降低。大约3 个狼毒花粉的浸提液就可以抑制受试的多数物种的50%的花粉萌发。在鳞叶龙胆和小米草柱头上狼毒花粉的数量分别为5.76 个和3.35 个。狼毒花粉散布数量的差异最可能的原因在于是否有共同的传粉昆虫。花的形状(辐射对称VS 左右对称)、植株或花的密度以及花期重叠性可以部分解释这种差异。在野外试验中,我们发现6 个物种(秦艽、湿生扁蕾、鳞叶龙胆、椭圆叶花锚、蓝钟花和小米草)的种子结实率随狼毒花粉浸提液浓度的增加呈显著的非线性降低。鳞叶龙胆和小米草柱头上狼毒花粉的数量(分别是5.76 个和3.35 个)分别达到了抑制它们63%和55%种子结实率的水平。因此,狼毒对鳞叶龙胆和小米草可能存在明显的花粉化感抑制作用。狼毒周围的物种可能通过花期在季节或昼夜上的分异避免受到狼毒花粉化感的影响或者通过无性繁殖来维持种群繁衍,因此狼毒通过花粉化感作用对其周围物种繁殖的影响程度还需要进一步地研究。如果狼毒的花粉化感抑制作用确实存在,那么它可能成为一种自然选择压力,进而影响物种的进化。 Stellera chamaejasme L., a perennial toxic weed, has emerged and quicklydominated and spread in the high-frigid meadow on the eastern Tibetan Plateau ofChina since the 1960s. In the present study, effects of S. chamaejasme on carbon andnitrogen cycles on the high-frigid meadow on the eastern Qinghai-Tibetan Plateau ingrowing and non-growing season, and its pollen allelopathic effects on the sympatricspecies were determined. The present study that focused on carbon and nitrogencycles, especially on microbial biomass and pools of carbon and nitrogen innon-growing season, could profoundly illuminate plant-species effects on carbon andnutrient cycles and its seasonal pattern and help to understand spread mechanism ofS. chamaejasme as an aggressive weed. The present study also contributed to furtherunderstand carbon and nutrient cycles on alpine regions in non-growing season andprovide a basis on weed control of S. chamaejasme and scientific management in thehigh-frigid ecosystem. Effects of S. chamaejasme on carbon and nitrogen cycles on the high-frigidmeadow on the eastern Qinghai-Tibetan Plateau were determined. The study couldbe divided into two parts. (1) In the growing season, we quantified the effects of S.chamaejasme on carbon and nitrogen cycles in two types of topographic habitats, theflat valley and the south-facing slope, where S. chamaejasme was favored to spreadlitter and root were measured to explain the likely effects of S. chamaejasme on soilcarbon and nutrient cycles. The sizes of various soil pools, e.g. nitrite, ammonium,inorganic phosphorus and microbial biomass, and turnover rates including netmineralization, gross nitrification, denitrification and microbial respiration weredetermined. (2) In the non-growing season study, microbial biomass carbon andnitrogen, soluble organic carbon and nitrogen, ammonium and nitrate weredetermined through the non-growing season, especially in the processes offreeze-thaw of spring in two high-frigid sites, i.e. Kaka valley and Gami temple, onthe eastern Qinghai-Tibetan Plateau. Meanwhile, litter decomposition of six commonspecies, including Stellera chamaejasme L., Polygonum macrophyllum D. Don var.Macrophyllum, Poa pretensis L., Kobresia setchwanensis L., Potentilla anserina L.var. anserine and Gentiana squarrosa Ledeb., were also examined under theabove-mentioned experimental design through one whole-year, which began in theautumn in 2006. In the study of pollen allelopathy, several work, including in vitro study oneffects of extract of pollen from S. chamaejasme on sympatric species and pollenfrom itself, field experiments on effects of pollen extract with the same regime ofconcentrations on seed set and field observation on heterospecific pollen transfer ofS. chamaejasme to six of those sympatric species has been done. The results in the growing season showed that aboveground litter of S.chamaejasme had higher tissue nitrogen and lower lignin: nitrogen ratio than thoseco-occurring species. S. chamaejasme significantly increased topsoil organic matter,whereas no significant differences were found for organic C and total P in subsoilbetween under-Stellera and away-Stellera locations. The nitrate in Stellera topsoilwas 113% and 90% higher on the flat valley and on the south-facing slope,respectively. Both microbial biomass C and N were significantly higher in Stelleratopsoil. Gross nitrification and microbial respiration were significantly higher inStellera topsoil both on the flat valley and on the south-facing slope, whereassignificant differences of denitrification were found only on the flat valley. Thedifferences in the quantity and quality of aboveground litter are a likely mechanismresponsible for the changes of soil variables. We assumed that S. chamaejasme couldenhance their spread by increasing nutrient availability in N-deficient ecosystems. The results in the non-growing season showed that microbial biomass achievedthe first summit in late autumn and early winter on the eastern Qinghai-TibetanPlateau. In the stages of freeze-thaw of spring, microbial biomass firstly achieved thesecond summit and subsequently sharply decreased. Inorganic nitrogen, solubleorganic carbon and nitrogen had a similar dynamics with that of microbial biomass.Ratio of microbial biomass carbon and nitrogen had an obviously seasonal pattern.The highest microbial C: N were in the non-growing season, which weresignificantly higher than those in the growing season. The seasonal pattern inmicrobial biomass C: N suggested that large changes in composition of microbialpopulation and in resources those used by microbes between summer and winter.Generally, microbial biomass and pools size of carbon and nitrogen in Stellera soilwere significantly higher than those under adjacent locations in late autumn andearly winter, but there were not significant differences in winter and in spring. Litterof all the focal species (Stellera chamaejasme L., Polygonum macrophyllum D. Donvar. Macrophyllum, Poa pretensis L., Kobresia setchwanensis Hand.-Maizz.,Potentilla anserina L. var. anserine and G. squarrosa Ledeb.) decomposed about24%-50% in the non-growing season, which were higher than those in the growingseason (ranged from 10% to 30%). Litter decomposed 10%-20% within the first twoweeks in late autumn and early winter. Significant differences in the whole-yeardecomposition rate and in the processes of decomposition were found among species.Polygonum macrophyllum decomposed slowly through the whole year (26% and15% in the non-growing season and in the growing season, respectively). Certainspecies, such as P. pretensis L. and K. setchwanensis, decomposed at a similar rate(30% both in the non-growing and in the growing season, slightly higher in the8growing season than those in the growing season), whereas S. chamaejasmedecomposed more rapidly (about 50%) in the non-growing season and subsequentlydecomposition became slow (about 12%) in the growing season. Litter nitrogencontents of all the focal species firstly decreased in the non-growing season and thenincreased in the growing season. In vitro experiments of pollen allelopathy, the results showed that pollen from S.chamaejasme was not autotoxic, whereas pollen germination in all the sympatricspecies (Polygonum macrophyllum D. Don var. Macrophyllum, Gentianamacrophylla Pall. var. fetissowii, Gentianopsis paludosa (Hook. f.) Ma var. paludosa,Gentiana squarrosa Ledeb., Halenia elliptica D. Don var. elliptica, Cyananthushookeri C. B. Cl. var. grandiflorus Marq., Euphrasia pectinata Ten., Delphiniumtongolense Franch., Ranunculus tanguticus (Maxim.) Ovcz. var. tanguticus andPotentilla anserina L. var. anserina) decreased nonlinearly as the increasingconcentrations of extract of pollen from S. chamaejasme. Pollen Extract of threepollens from S. chamaejasme generally inhibited 50% pollen germination of most ofthe focal species. 5.76 and 3.35 pollens from S. chamaejasme were observed in fieldon stigmas of G. squarrosa and E. pectinata, respectively. Differences inheterospecific pollen transfer of S. chamaejasme could be attributed to the primaryreason whether they shared common pollinators. Flower morphology (e.g.zygomorphic or actinomorphic), plant or floral density and concurrence in floweringphonologies could explain, in part, the differences in heterospecific pollen transfer.In field experiments, the results showed that seed set in six sympatric species(Gentiana macrophylla Pall. var. fetissowii, Gentianopsis paludosa (Hook. f.) Mavar. paludosa, Gentiana squarrosa Ledeb., Halenia elliptica D. Don var. elliptica,Cyananthus hookeri C. B. Cl. var. grandiflorus Marq. and Euphrasia pectinata Ten.)decreased nonlinearly as the increasing concentrations of extract of pollen from S.chamaejasme. According to the nonlinear curves, the amounts of pollens from S.chamaejasme on stigmas of G. squarrosa and of E. pectinata (i.e. 5.76 grains and3.35 grains, respectively) could reduce 63% and 55% seed set of G. squarrosa and ofE. pectinata, respectively. Thus, allelopathic effects of S. chamaejasme on G.squarrosa and E. pectinata could be realistic. The sympatric species of S.chamaejasme could avoid pollen allelopathy of S. chamaejasme to sustainthemselves. This highlights the need to study how much pollen allelopathy in S.chamaejasme influences the sympatric species through divergence in seasonal ordiurnal flowering phonologies or through asexual reproduction. If pollen allelopathyin S. chamaejasme was confirmed, it could be as a pressure of natural selection andthus play an important role in species evolution.

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本文以青藏高原东部的高山草甸为研究对象,设置早融、中间及晚融三个融雪部位,采用实验室测量、野外测量、野外样方调查相结合的 方法,从个体、种群和群落的水平上比较研究了高山雪场植物在同一雪场样地中不同融雪梯度上的特征变异及适应,结果表明: 从早融到晚融的梯度上,随着融雪时间的逐渐推迟,表土日温差降低,冻融交替的强度减弱,土壤水份逐渐增加,总N、总P、总K 以及 可溶性的N、P 和pH 变化不明显,土壤有机质及可溶性的K 和Ca 逐渐降低。冻融交替强度上的差异以及土壤水分差异被认为是融雪梯度上 影响植物生长的主要原因。 从早融到晚融的梯度上,伴随着生态因子的改变,几种常见植物的个体特征也发生相应的变化。首先,物候期推迟。植物开始生长的时间 一般要推迟将近二十天,但同一种植物在不同的融雪部位上的衰老期趋于一致,这预示着在晚融部位同一植物的生长期要缩短。其次,个体生 长特性发生改变。黑褐穗苔草(Carex atrofusca subsp. minor (Boott) T.Koyama)和西北黄芪(Astragalus fenzelianus Pet.-Stib.)的个体生长(株高、单株叶数、单叶面积和地上生物量)表现为逐渐增加的趋势;斑唇马先蒿(Pedicularis longiflora Rudolph var. tubiformis (Klotz.) Tsoong)和川西小黄菊(Pyrethrum tatsienense (Bur. et Franch.) Ling ex Shih.)则表现为逐渐降低的趋势;长叶火绒草(Leontopodium longifolium Ling)在融雪梯度上的变化趋势不明显。再次,从繁殖特性来看,大卫马先蒿(Pedicularis davidii var. pentodon Tsoong)的单株花数、单花种子数、种子千粒重及种子萌发率随融雪的推迟呈现为逐渐增加的趋势;圆穗蓼(Polygonum macrophyllum D.Don)的种子(小坚果)千粒重和萌发率也表现为逐渐增加,其余繁殖特征变化不明显。 在种群层次上,几个常见物种的分布格局随着融雪的推迟都发生一定的变化,基本上表现为从早融的集群分布到中间或晚融部位的随机分布。物种间的联结性也发生较大的变化,由早融部位的总体上的正关联逐步过度到晚融部位上的总体上的负关联。特定种对间的联结性也发生较大的变化。恶劣环境条件(如剧烈的冻融交替)的影响以及对恶劣条件适应被认为是分布格局及种间联结性发生变化的主要原因。 在群落层次上,物种多样性的变化表现为单峰曲线的格局,即在中间部位多样性最高。早融部位强烈的冻融交替和晚融部位缩短的生长季是早融及晚融部位物种多样性不高的重要原因。几乎所有的只出现在一个融雪部位(雪深级别)上的物种都发生在中间融雪部位。这说明,中等的雪深更有利于许多高山植物的存活,而过浅过深的积雪都不利于植物的生存。另外,相距较近的融雪梯度之间的物种相似性较大,而相距较远的梯度之间物种的替代率较高,物种的相似性较小。在群落的生物量方面,地上生物量随融雪的推迟而升高,地下生物量随融雪的推迟而下降,地上与地下生物量之总和随着融雪的推迟而下降,地下生物量与地上生物量之比随着融雪的推迟而下降。早融部位的地上生物量主要集中于地上0-10cm 的范围内,表明在早融部位植物地上部分有变矮的趋势;早融部位的地下生物量在土壤各深度分布相对较均一,而晚融部位地下生物量则主要集中于地下0-10cm 的范围内。生物量的变化趋势主要与雪场中各部位的土壤水分含量及地表日温度差异有关,是植物适应特定环境的结果。 To detect the plants’ responses to snow-cover gradients in an alpine meadow of eastern Tibetan plateau, laboratory method and field sample plot method were employed, and three gradeients (early-, medium and late-melting)were established in a natural snowbed. The measurements were carried out for two years and was done on three levels——individual, population and community. The results are shown as follows : From early- to late-melting gradients, daily ground temperature difference between day and night decreased, amplitude of freeze-thaw alternation weakened, soil organic matter contents and soluble K and Ca decreased, while soil water content increased. Total N, total P, total K,pH soluble N and soluble P kept constant from early- to late-melting portions. Among these factors, the changes of intense freeze-thaw alternation and soil water contents were considered as main factors affecting plants’ growth. From early- to late-melting portions, all phenological phases postponed, e.g. phase of plant emergence postponed almost twenty days. However, the same species’ individuals at different portions withered in step, which implied that the individuals at late-melting portion possessed shorter growing season length. Along the same gradient, both Carex atrofusca subsp. minor (Boott) T. Koyama and Astragalus fenzelianus Pet.-Stib. increased their individual growth, whereas Pedicularis longiflora Rudolph var. tubiformis (Klotz.) Tsoong and Pyrethrum tatsienense (Bur. et Franch.) Ling ex Shih. decreased their individual growth. Unlike the four plants mentioned above, Leontopodium longifolium L. did not show any evident change. As to reproductive charateristics, the flowers per individual, the number of seeds per flower, the thousand seed weight and the seed germination rate of Pedicularis davidii var. pentodon showed an increasing trend; and Polygonum macrophyllum D.Don also increased its thousand seed weight and seed germination rate along the same gradient. However, the other reproductive charateristics of Polygonum macrophyllum D.Don did not change significantly. At population level, the distribution pattern of several selected species changed from cluster pattern to random pattern as the snowmelt postponed. Overall association among the species changed from positive to negative along the same gradient. Further, interspecific association also changed evidently. Adverse circumstances such as intense freeze-thaw alternation were considered as primary factors resulting in changes of population distribution pattern and interspecific association. At the level of community, species diversity showed a pattern of a unimodal trend, i.e. the highest diversity occurred at medium snow depth,perhaps because of intense freeze-thaw alternation at early-melting portions and the shortest growing season at late-melting portions. Almost all species that only appeared at one snowmelt portion occurred at medium portion, indicating that medium snow depth was more suitable for many species’ survival. Species replacement from one snowmelt portion to its neighboring portion seldom took place. However, while distance between two portions became farther, species replacement between the two portions occurred more frequently. As for biomass, aboveground biomass increased from early- to late-melting portions, whereas belowground biomass, total biomass and the ratio of belowground to aboveground all decreased along the same snow gradient. A majority of aboveground biomass distributed in a height range of 0-10 cm, suggesting that height of plants inhabiting early-melting portion be shorter compared with other portions. In addition, belowground biomass at early-melting portion was evenly distributed at different soil depth in comparison with aboveground biomass, whereas belowground biomass at late-melting portion concentrated 0-10cm soil layer below ground. The changing trend of biomass was also related to two factors. One was soil water content, and the other topsoil temperature difference between day and night.

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通过秋水仙素诱导获得同源四倍体水稻10个株系,包括6个恢复系、3个保持系和1个不育系,这些株系具有加倍的染色体组。田间观察表明10个株系具有特殊的农艺性状:茎杆变粗壮、植株颜色加深、叶片变厚、叶宽适度增加、分蘖数减少、有效分蘖的比率下降等。根尖有丝分裂鉴定表明,同源四倍体水稻10个株系具有正常的有丝分裂,观察细胞的染色体数目皆为2n=48。花粉母细胞减数分裂鉴定表明10个株系具有比较理想的减数分裂行为,后期I染色体滞后、末期I微核生成和末期II异常小孢子比率较低,能较好的完成减数分裂过程,其中后期I染色体滞后比率约为10%-20%,末期I微核生成比率约为1%-6%,末期II异常小孢子比率约为1%-8%。这提示,染色体联合和分离不规则导致三价体、单价体 和落后染色体等产生,并进一步导致在后期和末期不均横分离产生异常小孢子,这可能是同源四倍体株系结实率不高的原因之一。 同源四倍体水稻正常胚囊为蓼型,变异胚囊具有多种类型,其比率显著高于二倍体对照,变化范围为39.62%-69.85%。按变异胚囊的结构特点和形成方式,分为四种类型:退化型,结构变异型,无融合生殖型和反足细胞增殖型。退化型胚囊的平均比率为29.17%,包括小胚囊(15.04%)和完全退化胚囊(14.13%),前者仍有较小胚囊腔而后者胚囊腔缺失。结构变异胚囊包括结构缺失、结构重复和位置异常,反映了蓼型胚囊八核七细胞结构的变异,其在各株系的平均比率为18.96%。无融合生殖胚囊发生比率极低,平均比率为1.77%,类型包括反足胚和卵细胞胚。反足细胞增殖胚囊是反足细胞团频繁增殖形成,伴随上述三种变异发生使异常胚囊的多样性进一步增加,其在各株系的平均比率为10.62%。相关分析表明,同源四倍体水稻结实可能主要来自三部分:正常胚囊、正常型小胚囊和反足细胞增殖型胚囊。这三种胚囊具有相对完整的蓼型结构,可能具有较好的育性,其对结实率的贡献程度估计值分别为72.44%、15.12%、12.44%。此外,完全退化型胚囊和位置异常型胚囊对结实率分别表现出显著(-0.66)和极显著(-0.92)的负相关,这表明二者可能是结实性的抑制因素。 Ten autotetraploid strains, which include six restoring lines, three maintaining lines and a sterile line, are derived from artificial induction by colchicine treatments. Variations of agronomical traits are observed which include large organs, sturdy plants, long panicle length and deep leaf color and so on. It has been confirmed that autotetraploid strains exhibit normal chromosome behaviors in mitosis and the chromosome numbers are all 48. Moreover, abnormal chromosome behaviors are investigated in meiosis including univalent, trivalent, quatrivalent, chromosome lagging and microkernel and so on. It evaluates that the percentage of chromosome lagging in anaphase I is about 10%-20%, the percentage of microkernel in telophase I is about 1%-6% and the percentage of abnormal microspore in telophase II is about 1%-8%. In all, abnormal behaviors of chromosomes could induce univalent, trivalent and et al. and subsequently induce infertile microspore. That may be one of the causes of low seed sets in autotetraploid strains. Embryo sacs of autotetraploid strains are formed according to the Polygonum type. However, these strains exhibit variations of abnormal embryo sacs at high frequencies (39.62% - 69.85%). The variations are frequently involved in the spikelets of the main panicles and the first tillers, leading to obvious decreases of the percentages of normal embryo sacs among the strains. Four types of abnormal embryo sacs are classified basing on their different structures and origins: degenerated embryo sac (DES), structure variation (SV), apomixis (Apo) and antipodal cell proliferation (ACP). Embryo sacs of DES (29.17%) exhibit small embryo sacs (15.04%) or no embryo sac (14.13%), the former showing embryo sacs with decreased size and the latter showing no sac. Embryo sacs of AS (18.96%) include three subtypes: structure deletion, structure duplication and location variation, which suggests abnormalities of the eight nuclei, seven celled pattern of the Polygonum type. Embryo sacs of Apo (only 1.77%) include two origins of apomictic embryos: antipodal embryo and egg embryo. Embryo sacs of ACP are observed frequently (10.62%) in autotetraploid strains which subsequently increase the variations of abnormal embryo sacs. It evaluates by the Pearson’s correlation analysis that seed set is probably contributed by three origins of embryo sacs: normal embryo sacs, small embryo sacs (normal pattern) and embryo sacs of ACP. These three origins exhibit comparatively good structure of the Polygonum type and could account for seed set at a percentage of 72.44%, 15.12%, 12.44%, respectively. Moreover, the subtype of no embryo sac (NES) negatively related to seed set at the P>0.01 level (-0.92) and the subtype of location variation (LV) negatively related to seed set at the P>0.05 level (-0.66). Which suggest the two subtypes may have strong stress on seed set.

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在实验室内进行了虎杖(Polygonum cuspidatum Sieb.etZucc.)根提取物及其活性组分分离研究,并且对黄瓜白粉病进行了活性测定及温室小区试验。结果表明,无论提取率还是作用效果,乙醇较适合于做为虎杖根提取物的提取溶剂。虎杖根乙醇提取物和乙醚提取物对黄瓜白粉病的EC90值分别为537.85、35.38mg/L,而乙醚提取后残留物的活性很弱,说明乙醇提取物中具有杀菌作用的成分主要存在于乙醚提取物中。用pH梯度分离法对乙醚提取物进行分离,得到3个不同组分,其中以氢氧化钠分离得到的组分对黄瓜白粉病显示了极高的杀菌活性。在温室条件下虎杖乙醇提取物在800~1600mg/L下对黄瓜白粉病的防效相当或优于对照药剂三唑酮。

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通过研究围栏封育1年后中度退化的矮嵩草草甸(夏季牧场)群落结构的变化,探讨主要植物种群在小尺度(50 cm×50 cm)上的空间分布格局,并从生活史特征和生态适应对策等角度探讨产生和维持这些格局的机理。结果表明,围封1年显著降低了群落中主要种群矮嵩草(Kobresia humilis)、高山唐松草(Thalictrum alpinum)和雪白委陵菜(Potentilla nivea)的重要值,而增加了线叶龙胆(Gentiana farreri)的重要值;显著增加了群落的地上生物量和总生物量,但对地下生物量和群落多样性的影响不显著。围封使退化矮嵩草草甸主要种群矮嵩草、高山唐松草、珠芽蓼(Polygonum viviparum)、线叶嵩草(Kobresia capillifolia)和金露梅(Potentilla fruticosa)等空间分布格局从放牧后的随机分布向聚集分布发展,而雪白委陵菜、重齿风毛菊(S.katochaeteMaxim)、矮火绒草(Leonto-podium nanum)和美丽风毛菊(Saussurea pulchra)等种群的空间分布格局没有发生改变;但黑褐苔草(Carex atro-fusca)的空间格局从放牧后的聚集分布转向随机分布。因此,退化的矮嵩草草甸在围封的初始阶段,由于避免了家畜的选择性采食及其践踏作用,首先可能是使主要种群的空间分布格局有从随机分布向聚集分布变化的趋势,从而使小尺度的种间隔离来降低种间的竞争强度,从而改变了不同物种对资源和空间的竞争能力,进而推动群落物种组成和结构的恢复演替。