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题名: 粉花绣线菊复合群及其相关类群的生物地理学研究
作者: 张朝阳
学位类别: 博士
答辩日期: 2003
授予单位: 中国科学院昆明植物研究所
授予地点: 中国科学院昆明植物研究所
导师: 周渐昆
学位专业: 植物学
中文摘要: 本文综合多学科的研究手段从种,居群,个体,细胞直到分子的各个生物学层次,描述了粉花绣线菊及其相关类群的生物学特性,揭示了该类群的遗传变异式样(遗传结构),并与植物化学和植物地理学相结合深入探讨了该类群遗传变异式样的形成机制和过程。以粉花绣线菊及其相关类群为例,从种群遗传学角度论述了东亚物理环境异质区的客观实在性及其生物学效应。细胞遗传学分析表明粉花绣线菊复合体绝大多数变异类型为二倍体,并且不同变异类型具有多样化的核型,细胞遗传学趋异主要不是染色体数目或倍性的变化,而在于染色体组织结构上的重组。西南山地高原的抬升与粉花绣线菊多倍化水平没有明显的相关性。rDNA ITS区比较序列分析和AFLP分析都表明粉花绣线菊复合体各变异类型间已发生了一定程度的分化,确认了基于形态证据对粉花绣线菊复合体内各变异类型(变种)的认知,同时也表明工TS序列和AFLP是探讨绣线菊属或亚科种下或近缘种间系统发育关系的有效分子标记。基于ITS序列的系统发育树具有高度的地理结构性:所有东部居群聚合成的分支与所有西部居群形成的分支构成姐妹群关系,这表明粉花绣线菊复合体的祖先类群曾经发生过一次一分为二谱系分化。这种东西一分为二的谱系分化也得到植化证据的支持。所有东部居群的二帖生物碱均为Hetisine型,而绝大多数西部居群的二帖生物碱为Atisine型,仅有一个可能为低海拔残遗类型的居群为Hetisine型。由于粉花绣线菊复合体东西分支分布区的分野在西南山地高原的边缘地带,与东亚大的物理环境的分异(第一级台阶和第二级台阶)相一致,提示在东亚,东西物理环境的差异是影响植物系统发育模式的一个重要和基本的因子。化石资料表明绣线菊属的起源最迟不晚于晚白至世,并且由于其化石均见于高纬地区,因此,它是东亚植物区系中典型的北极第三纪成分。系统学分析表明粉花绣线菊是绣线菊属中较原始和保守的变异类型,因此,粉花绣线菊应在该属起源后的早期就已经出现,以后随第三纪气候变凉,不断向南迁移,并且随古地中海退却,喜马拉雅一横断山的隆升,东亚中部干旱带的退却,发生过从东北向西南的迁移,其分布区达到现代的东喜马拉雅和横断山地区。据分子钟推侧,粉花绣线菊东西地理分支的分歧时间约为中新世阿启坦期(23Ma.),与喜马拉雅-横断山地区隆升成山地高原的时期大体一致,说明该类群一分为二的谱系分化与山地高原的形成密切相关。分布在西南山地高原的西部分支的众多变异类型可能是随着山地高原的隆升适应环境的产物。物种分化与其所处的物理,气候环境密切相关,同时也是其所属植物区系演化发展的产物和体现者,因此,粉花绣线菊复合体的东西谱系分异时间可能也反映了中国一日本和中国一喜马拉雅森林植物亚区分异和成型的大致时间,即中新世附近。AFLP分析揭示了粉花绣线菊复合体及其相关类群东西地理集群在遗传变异模式和遗传结构上的差异性和各自的特点。西部集群居群间具有更大的遗传分化值和更大的遗传分化值变动范围,提示西部物理环境的复杂性导致更多的小区域隔离,小的基因流和更大的绝对成种速率,这与西部的物理环境的复杂性是相协调的,并且可能是造成西部山地高原地区生物多样性异常丰富的微观过程和机制。北极第三纪成分南移是现代东亚植物区系的一个极其重要的区系来源,绣线菊属是典型的北极第三纪成分,粉花绣线菊及其相关类群的演化历史轨迹,现代生物多样性格局形成的机制和过程很可能反映了众多东亚植物区系中北极第三纪成分的共同模式。从种群的角度为理解东亚植物区系演化发展的历史过程提供了一个个例。同时,粉花绣线菊及其相关类群生物地理学研究也是从种群遗传学角度对西南生物多样性中心在生物多样性保育中的重要意义和地位的进一步说明。为加强该地区生物多样性管理的科技投入提供了进一步的理论支持。
英文摘要: Spiraea japonica L. f. and its alliance are a species complex widely spread across the East Asia from Japan to east Himalayas. With multiple variations and typical distribution pattern, the complex can be a representative element of Eastern Asian flora. Studies of morphology, cytogenetics, comparative sequencing of ITS, AFLP, phytochimistry biogeography and paleobotany were performed on the complex at present dissertation, trying to reveal the divergence pattern and relationship among different geographical populations within the complex, and more importantly, to disclose the correlation between the diversification pattern of the complex and the historical physical environmental changes to further our understanding of the biological effects of historical physical environmental changes. Cytogenetical analysis revealed no polyploidy series within the complex, but a series of diversified karyotypes. With most populations being of diploidy, cytogenetical divergence within the complex lied mainly in chromosome size and structure. The fact that polyploidization did not occur very often for variations in Southwest China was against viewpoint that polyploidization level in this area is higher than that of other distribution areas due to the elevation of mountains and plateau in southwest China. Cytologically, chromosome structural re-arrangement seemed to play a key role in the adaptation of Spiraea japonica complex to the elevation. 16 populations representing eight of the nine varieties in S. japonica complex and S. fritchina were analyzed for comparative sequencing of ITS. Monophyly of Spiraea japonica complex including S. fritchina was strongly supported, and the strongly supported phylogenetic tree of the complex was highly geographically structured. All populations from east China were grouped into a single Clade, sister to the one comprising all populations from Southwest China. S. fritchina, a taxon distributed in east China, was nested into the East Clade indicating it would be a member of the complex. Moreover, divergence timing of the West Clade from the East Clade of S. japonica complex was estimated to be Aquitanian of Miocene(23Ma) after a test of consistency of molecular mutation ratio. Because distribution boundary between the East Clade and the West Clade was almost the same for the division line of physical environmental heterogenicity between the East and the West within the East Asia (division line between the second and third stage in accordance of topographical disjunction), it was speculated that physical environmental heterogenicity between the East and the West had played a key role in the phylogeny of the complex. Phylogenetic tree reconstructed by AFLP displayed similar tree topography, with the only exception of the location of Tengchong population which was nested into the East Clade. Genetic structure of populations from the east and the west was significantly different. Higher average genetic divergence ratio, wider range of genetic divergence ratio and smaller gene flow had been observed for populations from the West. Furthermore, AMOVA analysis based on AFLP indicated that most genetic divergence was allocated between varieties while the genetic divergence between the east group and the west group was considerable. Based on the observed genetic divergence pattern, it was speculated that elevation of mountains and plateau in southwest China had complicated the physical environment and resulted into more geographical isolation among small territories within the West, leading to fewer gene flow and higher absolute speciation speed. And this could be the micro-process and machinism controlling formation of the biology diversity center of southwest China. Phytochemical data was reviewed. Eight of the nine varieties of S. japonica complex as well as S. fritchina and other Spiraea had been investigated for Phytochemistry. Alkaloids were chemical constituents peculiar for S. japonica complex and its closely related alliances implying raonophyly nature of S. japonica complex and its alliances as a whole. Moreover, the fact that all populations from the East produced alkaloids of Hetisine type while alkaloids produced by populations from the West (except for the Tengchong population) were of Atisine type was in consonance with the phylogeny reconstructed from ITS sequences and AFLP. Fossil records of Spiraea were reviewed as well and origin of Spiraea was dated back to late Cretaceous. Based on multiple evidences from morphology, cytogenetics, ITS, AFLP, phytochemistry and paleobotany, a historical evolutionary hypothesis was developed explaining the spatio-temporal pattern of biodiversity of the S. japonica complex. Since all fossil records of the genus Spiraeawere found in area of high latitude, it was presumed that Spiraea was element of Arctic-Tertiary Flora. As a group with multiple primitive characters of Spiraea, S. japonica probably occurred very early soon after the origin of the genus. With the climate becoming cool in Tertiary, it probably had, with other elements of Arctic-Tertiary Flora, migrated southwards in East Asia. And with the Tethys retreating, a migration from the Northeast to the Southwest might have happened during Miocene so that S. japonica had reached the mountain and plateau area of Southwest China. Because divergence timing of the West Clade from the East Clade of S. japonica complex was estimated to be Aquitanian of Miocene(23 Ma), during which the mountains and plateau of Southwest China were formatted, the divergence of the West Clade from the East Clade of S. japonica complex was presumably correlated closely with the uplifting of mountains and plateau of Southwest China. The multiple variations of the West Clade of S. japonica complex might be production of adaptation to uplifting mountains and plateau and complicated environment. Elements of Arctic-Tertiary Flora are extremely important resources of Eastern Asian flora, of which S. Japonica is a representative. Evolutionary history of 5". japonica complex might reflect the common historical evolutionary process of multiple elements of Arctic-Tertiary Flora. The divergence timing of the west clad from the east clad might imply the early stage of the divergence of Sino-Himalaya forest Subkindom from the Sino-Japanese forest Subkindom. Also, the raachinisra controlling diversification of 5". japonica complex might be the same for the formation of the biological diversification center in Southwest China. Whatever, the presumed evolutionary history of 5". japonica complex was an important clue for the secret of natural history of the Eastern Asian flora.
语种: 中文
内容类型: 学位论文
URI标识: http://ir.kib.ac.cn/handle/151853/688
Appears in Collections:昆明植物所硕博研究生毕业学位论文_学位论文

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粉花绣线菊复合群及其相关类群的生物地理学研究.张朝阳[d].中国科学院昆明植物研究所,2003.20-25
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