毛茛科铁线莲属的分子系统学与繁育生物学研究
江南
学位类型博士
导师管开云
2010-01
学位授予单位中国科学院研究生院
学位专业植物学
摘要铁线莲属(Clematis L.)隶属于毛茛科(Ranunculaceae),共355种左右,为世界广布类群,主要分布于温带和亚热带。该属的属下分类等级划分一直倍受争议。本文通过分子手段,比较了分子系统与传统分类系统的异同,并探讨了属下分类等级等问题。此外还对该属植物的繁育生物学及萌发特性进行了研究。主要研究结果如下:1. 云南铁线莲属物种多样和地理分布横断山区可能是铁线莲属的起源、分化和特有中心,而滇西、滇西南以及滇东南地区也与该属的分化有一定联系。云南铁线莲属植物特有现象较为突出:含中国特有的56种(含变种),占云南总种数的67.2%;云南特有16种(含变种),占云南分布的中国特有种的35.0%。云南铁线莲属植物与国内邻近的四川关系最为密切,西藏的关系最弱;与国外邻近的缅甸具有一定的联系。2. 分子系统学研究:基于ITS、psbA-trnQ、atpB-rbcL和rpoB-trnC四个基因片段对铁线莲属及其近缘属79种138个样品进行了分子系统学研究。结果表明,铁线莲属是一个并系类群(paraphyly),其包含了锡兰莲属和互叶铁线莲属。在属下分类等级上,联合分析结果没有支持王文采和李良千系统以及任何一个传统分类系统,所研究的12个组多为多系(polyphyly)或并系类群,组间关系错综复杂。我们认为,这是由于该属与银莲花亚族分化时间较早,但属内各种间是在近期发生的快速分化的结果;同时,种间杂交可能也是主要因素之一。结合植株及幼苗性状,我们发现,单一以花部性状作为该属属下分类依据是不太合理的,应以幼苗叶序作为主要的鉴别特征,结合花瓣有无、心皮数量、花开放于新枝或老枝作为亚属或组等级划分标准。3. 命名学问题处理:对金毛铁线莲及罂粟银莲花进行了后选模式指定;订正了王文采和李良千系统中属下等级作者引证中的错误;同时,对非毛茛科的一些种类,尖叶酒饼簕、长茎金耳环、封开酒饼簕和霸王金橘进行了合格发表。4. 繁育生物学研究:本研究选取了三种不同花冠类型的铁线莲,进行传粉和繁育生物学研究。通过花部特征的测量、传粉者行为的观察、评估其近交衰退系数和花粉限制系数等研究,结果表明,蕊-雌蕊的比例可以像花粉-胚珠比一样在评价铁线莲繁育系统非常有价值;花冠形态分化与传粉昆虫种类和行为密切相关,平展型花冠是适应蝇类传粉,钟状和管状花冠是适应熊蜂传粉;花冠形状与传粉系统和繁育系统之间的关系非常密切。近交衰退系数与繁育系统之间密切相关。花萼、雄蕊和蜜腺(金毛铁线莲除外)在传粉吸引中扮演了不同的角色。5. 金毛铁线莲的花部性状:通过对金毛铁线莲花部性状的测量,包括萼片长、萼片宽、雄蕊数量、雌蕊数量、花药长、单个萼片大小、总萼片大小等,并采用不同年份单独分析和三年联合分析性状之间的相关性,来检验各性状年际间的稳定性,验证性配置理论。花部性状(除萼片长)在三年连续的研究中保持着较高的稳定性。部分花部性状之间的相关性在各年份单独分析和联合分析时一致,说明它们可能在演化过程关系密切;而相关性不一致或相反的花部性状,一方面可能是取样随机性误差所造成的,也有可能是它们之间存在一定的相互关系,但不密切相关。雄蕊数量与花药长不存在负相关关系(第一年呈显著正相关,后两年没有明显相关性);雄蕊数量和雌蕊数量之间呈显著正相关关系;雌蕊数量、总花药大小和总萼片大小三者之间呈显著的正相关关系。综合以上结果,我们发现金毛铁线莲的性资源配置不属于“补偿”理论模型,而是由基因和资源量决定。6. 种子萌发特性研究:本研究通过设置不同的预处理(砂纸打磨和不同时长的冲洗处理)、不同浓度的GA3浸种、不同光照和温度条件以及不同的土壤播种基质,对六种铁线莲进行了种子萌发特性的研究。冲洗处理可提高种子萌发率和缩短平均萌发时间。不同浓度的GA3对不同种的铁线莲影响不一,但总体来说,低浓度的GA3有一定的促萌作用。珍珠岩基质中的种子萌发率最高,可达80%以上,是较理想的铁线莲播种基质
资助项目The genus Clematis, with about 355 species, is one of the largest genera of Ranunculaceae. The genus is distributed through out the world except Antarctica, primarily in temperate and subtropical zone. Numerous infra-generic classification systems have been proposed, whereas the infra-generic ranks were confusing. In the present study, we used four molecular regions, ITS, psbA-trnH, atpB-rbcL and rpoB-trnC to reconstruct the phylogeny of Clematis. On the basis of comparison with traditional classification system, criteria of the infra-generic ranks are discussed. In addition, investigations of reproductive biology and seed germination were also conducted. The main important results and conclusions are as follows: 1. Species diversity and geographical distribution of the Genus Clematis in Yunnan province,According to the latest system of Wang and Li in 2005, there are 59 species and 24 varieties in 6 sections of 3 subgenera recognized within the genus of Clematis in Yunnan, which is the richest province in terms of species numbers in China. The Hengduan Mountains is regarded as an origination, differentiation and endemic center of the genus. West, Northwest and Northeast of Yunnan have some relationship with its differentiation. There are 56 species (including varieties) endemic to China, accounting for 67.2% of Yunnan’s total number of species. There are 16 species (including varieties) endemic to Yunnan, accounting for 35.0% of China’s endemic number of species distributed in Yunnan. Among the adjacent provinces, Sichuan has the largest common number of Clematis to Yunnan, while Tibet has the smallest. The floristic links with Myanmar is also relatively weak.2. Molecular phylogeny of Clematis,Phylogenetic relationships within and out of Clematis were analysed using nucleotide sequences of chloroplast DNA psbA-trnQ, atpB-rbcL and rpoB-trnC, and the nuclear ITS regions. The phylogenetic trees suggested that the genus Clematis was paraphyletic. The genera Archiclematis and Naravelia were found to be nested within the genus Clematis, and should be included in it. The molecular results did not agree with Wang and Li’s taxonomic system and other classifications of Clematis at the phylogeneric level. And the sections we studied were found to be paraphyletic or polyphyletic. The complicated relationship between sections and species indicated the recent radiation of the genus and the possibility of hybrid origin. Our results suggested that flower characters which have been used for previous infrageneric classification do not correlate with the phylogenic tree. Phyllotaxy of seedlings as the primarily diagnostic characters may reflect the phylogeny within Clematis, the following characters should be considered at the same time: presence/absence of petals, carpel number, and flowering at the new-born\former-born branches. 3. Nomenclature notes of Clematis and other plants,Two gatherings of Clematis chrysocoma Franchet and Anemoclama glaucifolia (Franchet) W. T. Wang were cited in the protologues, whereas none of them was indicated as type or holotype. After examining the cited specimens, we designate one specimen of them as lectotype in according with ICBN. We list the incorrect author(s) citation of infra-generic names by Wang and Li in their new Clematis classification systems. Additionally, four names, Atalantia acuminata C. C. Huang, Asarum longerhizomatosum C. F. Liang & C. S. Yang, Atalantia fongkaica C. C. Huang and Fortunella wawangica C. C. Huang, are validated.4. Reproductive biology of three Clematis species,An integrative study was undertaken, focusing on pollination and reproductive biology of three species representing all three floral types of Clematis. Floral traits were measured, and pollinator assemblages were observed in the field. Bagging, hand-pollination and removal treatments were used to examine breeding system. The values of inbreeding depression and pollen limitation were calculated based on experimental data obtained from breeding system. Their floral traits were distinctly different. Floral traits were highly associated with pollination syndrome and breeding system. Among them, Clematis akebioides and C. rehderiana were facultative autogamy (the former was delayed selfing, and the later competing selfing), and C. chrysocoma was nearly obligate outcrossing. This conclusion was reflected by their stamen-pistil ratios. The levels of inbreeding depression negatively associated with autonomous self-pollination. Selfing evolution of C. akebioides and C. rehderiana, and reproduction of the three Clematis species no pollen limitation were discussed. Present study integrating with previous results will help us to comprehensively recognize and understand pollination system and reproductive characteristics of Clematis.5. Correlation among floral traits of Clematis chrysocoma, Floral traits of C. chrysocoma including sepal length and width and anther length were measured, stamen and pistil number were counted, single and total sepal size, and total anther size were calculated in three continued years from 2007 to 2009. These traits of three years were analyzed as united and independent data, respectively, which was used to compare the stability of floral traits among three years, and to test sexual allocation theory. Our results show that floral traits except sepal length are stable in three continued years, correlation among floral traits of three years as whole and independence, respectively, some of which are consistent, and some of which are inconsistent or negative. The consistent traits demonstrate that they highly co-evolve, whereas the inconsistent or negative traits may be results from the stochastically errors or that they may have some correlation but not close. Association of stamen number with anther length is not negative, but with pistil number is significantly positive. Correlation among pistil number, total anther size and total sepal size, all of that are significantly positive. Therefore, sexual allocation strategies of C. chrysocoma should be determined by genetics and resource pool in contrary to the model of “compensation” as traditionally view.6. Seed germination of six Clematis species,Seed germination of six species of Clematis were studied by setting different conditions including pretreatment, GA3 treatments, light and temperature regimes, and substrates. The results of the test were as follows: flushing treatment had a significant promoting on seed germination percentage and mean germination time (MGT); and germination of the seeds soaked in low concentrations GA3 was a little higher than those of others. The seeds in perlite had the highest germination percentage than those in other substrates.
语种中文
文献类型学位论文
条目标识符http://ir.kib.ac.cn/handle/151853/16176
专题昆明植物所硕博研究生毕业学位论文
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江南. 毛茛科铁线莲属的分子系统学与繁育生物学研究[D]. 中国科学院研究生院,2010.
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