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题名: 中国伞形科芹亚科的分子系统学研究
作者: 周静
学位类别: 博士
答辩日期: 2009-05-25
授予单位: 中国科学院昆明植物研究所
授予地点: 昆明植物研究所
导师: 彭华
关键词: 伞形科 ; 芹亚科 ; 细胞学 ; 分子系统学 ; 染色体演化模式 ; 生物地理 ; 系统演化
学位专业: 植物学
中文摘要: 芹亚科(Apioideae)是伞形科中最大也是分类学上最为复杂的一个亚科,包括404属,2827—2935种,中国分布有95属579种。本研究综合形态学、细胞学和分子系统学等多学科的手段和方法,以中国芹亚科为重点研究对象,集中探讨中国芹亚科类群的系统演化关系同时确立中国伞形科10个特有属的系统学位置。并在已有资料的基础上,对中国芹亚科的染色体演化模式以及生物地理学问题进行了初步的总结和讨论: 1.细胞学 报道了藁本属(Ligusticum)、环根芹属(Cyclorhiza)、东俄芹属(Tongoloa)、葛缕子属(Carum)、水芹属(Oenanthe)、囊瓣芹属(Pternopetalum)以及矮泽芹属(Chamaesium)共7属16个种的细胞学资料。藁本属10个种的染色体基数均为x= 11,但是基于核型公式以及核型的数量参数可以将它们区别开来。细苞藁本(L. capillaceum)和蕨叶藁本(L. pteridophyllum)为四倍体2n= 4x= 44,其余种类为二倍体2n= 2x= 22。我们通过对藁本属已进行过细胞学研究的种类进行分析,认为多倍化在藁本属类群的演化过程中所起的作用并不明显。在滇西北地区,藁本属不仅植物种类丰富,而且集中了该属叶柄解剖的全部类型,形态分化十分活跃,在核形态上也表现出一些变异,因此,该地区很可能是藁本属的最大变异中心。 2.分子系统学 基于伞形科87个属171个种的ITS,rpl16以及rps16序列的最大简约法 (MP)和贝叶斯法(BI)分析,得到如下结论:(1)确定中国伞形科10个特有属在伞形科中的系统学位置:环根芹属(Cyclorhiza)、明党参属(Changium)和川明参属(Chuanminshen)共同位于Komarovia Clade(Komarovieae);舟瓣芹属(Sinolimprichtia)、羌活属(Notopterygium)与其它一些具有中国-喜马拉雅分布的类群共同构成东亚分支(East-Asia Clade),羌活属不是一个单系,而是与单球芹属(Haplosphaera)和凹乳芹属(Vicatia)形成并系;紫伞芹属(Melanosciadium)与亮蛇床族(Selineae)的当归类群聚成一支;滇藏细叶芹属(Chaerophyllopsis)与针果芹族(Scandiceae)的针果芹亚族(Scandicinae)表现出很近的亲缘关系;细裂芹属(Harrysmithia)处于新确立的丝瓣芹分支(Acronema Clade);马蹄芹属(Dickinsia)被确认位于新建立的牵环花亚科;白苞芹属(Nothosmyrnium)的系统位置基于分析方法的不同而有所变化:在ITS系统树中,该属与茴芹族(Pimpinelleae)聚成一支,在ITS+cpDNA联合分析中,在临近茴芹族的位置形成一个比较孤立的分支,并且构成亮蛇床族+环翅芹族+滇芹分支(Selineae + Tordylieae + Sinodielsia Clade)的姐妹群,而在cpDNA系统树中,它形成亮蛇床族+环翅芹族polytomy中的一个分支或与环翅芹族的类群聚在一起;2)基于本研究新确立了三个分支,即丝瓣芹分支(Acronema Clade)、滇芹分支(Sinodielsia Clade)和矮泽芹分支(Chamaesium Clade)。丝瓣芹分支与针果芹族(Scandiceae)形成单系的姐妹群;滇芹分支与尖叶藁本(Ligusticum acuminatum)聚成一支或者与芹族(Apieae)形成并系;矮泽芹分支构成中国芹亚科中除外类群柴胡族以外其它所有类群的姐妹群。东亚分支(East-Asia Clade)的界限在本研究中得到极大扩充,它与Komarovia Clade(Komarovieae)构成得到单系支持的姐妹群;3)组成丝瓣芹分支的类群,如丝瓣芹属(Acronema)、囊瓣芹属 (Pternopetalum)、藏香叶芹属(Meeboldia)、丽江藁本(Ligusticum delavayi)、棱子芹属(Pleurospermum pro parte)以及细裂芹属等在形态上差异很大,但它们均为东亚特有。另外组成东亚分支(East-Asia clade)的类群在形态上也有很大的分化,但均为东亚分布。两个支系的分布区类型均属于东亚植物地理类型(East Asian Phytogeographycal type);4)一些通过形态特征不能很好界定的属,基于分子系统学的研究也不是单系,比如:藁本属(Ligusticum)、前胡属(Peucedanum)、滇芎属(Physospermopsis)、茴芹属(Pimpinella)、棱子芹属(Pleurospermum)、小芹属(Sinocarum)、东俄芹属(Tongoloa)以及糙果芹属(Trachyspermum)等;而一些在目前的研究当中确立为单系的属,如:丝瓣芹属、囊瓣芹属和水芹属(Oenanthe),仍需要通过更为广泛的取样,对其属下关系进行进一步的探讨;5)滇芹属(Sinodielsia) 与藏香叶芹属和凹乳芹属(Vicatia)是截然不同的;滇西前胡(Peucedanum delavayi)在系统发育上与滇芹属以及藏香叶芹属关系较远,而与亮蛇床族(Selineae)的藁本属类群聚在一起; 6)参棕亚科(Mackinlayoideae)、牵环花亚科(Azorelloideae)和变豆菜亚科(Saniculoideae)各自形成单系,并依次构成芹亚科的姐妹群。 本研究是第一次基于分子系统学的理论和方法对中国伞形科芹亚科的特有类群进行广泛的取样和研究。我们的结论不仅为后续的分类学修订构建了框架,同时也为进行形态性状的演化以及生物地理方面的研究奠定了基础。对于丝瓣芹分支(Acronema Clade)和东亚分支(East-Asia Clade)来说,尽管目前的分析确立了它们各自的单系性质及在芹亚科中的系统位置,但是分支内部类群之间的关系并没有得到很好的解决。下一步的工作有必要通过更为广泛的取样,并结合更多的分子片段对这两个大的分支开展进一步的研究。 3.分类学处理 在综合形态学、分子生物学及生物地理学证据的基础上,基于核和叶绿体DNA分析结果的一致,对Komarovia Clade和矮泽芹分支(Chamaesium Clade)进行分类学处理,分别被确立为新族:Komarovieae J. Zhou & S. R. Downie和Chamaesieae J. Zhou &F. D. Pu。根据矮泽芹族的系统位置,我们推测它可能是芹亚科在亚洲扩散形成的最早的谱系之一。 4. 生物地理学 伞形科广布于北半球温带、亚热带地区或热带高山上,是一个以温带为主的自然大科。根据已有的资料,伞形科有两个现代分化中心,一个位于地中海地区(Mediterranean area),另一个位于美国西部(western United States)以及墨西哥(Mexico);且南非构成芹亚科和变豆菜亚科的起源中心。 基于我们的分析,中国伞形科10个特有属分别位于7个主要的分支(丝瓣芹分支Acronema Clade、茴芹族Pimpinelleae、亮蛇床族Selineae、针果芹族Scandiceae、Komarovieae、东亚分支East-Asia Clade和牵环花亚科Azorelloideae),代表伞形科不同的演化阶段。马蹄芹属、明党参属、川明参属、环根芹属、舟瓣芹属、羌活属位于系统树近基部的位置,属于古特有属;滇藏细叶芹属、细裂芹属、紫伞芹属、白苞芹属位于系统树较为进化的位置,应属新特有属的范畴。根据各特有属的系统位置及地理分布,地处康滇古陆的云南和四川很可能是中国伞形科植物的次生演化中心。一方面由于青藏高原地形、气候复杂多变,形成天然避难所使科内的一些古老成分得以保留;另一方面由于第三纪末青藏高原的急剧隆起,促进该区域伞形科植物的分化并产生出新的类群。 5.芹亚科染色体演化模式初探 根据现有的细胞学资料,基于核和叶绿体联合分析的贝叶斯一致性树,我们对中国芹亚科的染色体演化模式进行了初步探讨:(1)在芹亚科内,从Komarovieae到亮蛇床族(Selineae)的大多数类群均保持了x= 11这一祖征;(2)基数x= 6的祖征在矮泽芹族(Chamaesieae)中得以保持,该性状在泽芹属(Sium)和山茉莉芹属(Oreomyrrhis)中均源自x=11的祖征;(3)染色体基数x= 8在城口东俄芹(Tongoloa silaifolia),峨参 (Anthriscus sylvestris)以及小窃衣(Torilis japonica)中独立起源,构成各自的自征(autopomorphy);(4)x= 9的基数构成棱子芹族(Pleurospermeae)、茴芹族(Pimpinelleae)和囊瓣芹属(Pternopetalum)各自的共有衍征(synapomorphy);(5)染色体基数x= 10在芹亚科中多次独立起源,且构成白苞芹(Nothosmyrnium japonicum)的自征;(6)茴芹属的类群基于分子系统学的分析,分散到三个分支当中: x= 11的类群与同样具有该基数的亮蛇床族(Selineae)聚成一支;具有x= 9基数的类群位于茴芹族;而x= 10的类群构成东亚分支的一个次级分支。因此,染色体分析的结果很好的支持了它们各自的系统位置;(7)棱子芹属(Pleurospermum)的类群在系统树上分别位于棱子芹族(x= 9)和丝瓣芹分支(x= 11),但是因为缺少细胞学资料,位于两个分支的该属类群是否能通过染色体基数区别开来还有待于进一步的研究。
英文摘要: Apiaceae (Umbelliferae) are a large and readily identifiable family of flowering plants, with subfamily Apioideae being the largest and most taxonomically complex (404 genera, 2827--2935 species). China is one of four major distribution centers of Apioideae, with 95 genera and 579 species recognized. In this study, we investigated the phylogenetic relationships of Apiaceae subfamily Apioideae mainly distributed in China including many endemics from the Hengduan Mountains, while ascertain or confirm the phylogenetic placements of all ten Chinese endemic genera by means of morphology, cytology and molecular systematics. Furthermore, chromosome evolution and biogeography of Chinese Apioideae were discussed. The main results are summarized as follows: 1.Cytology Karyological studies on seven genera, i.e. Ligusticum、Cyclorhiza、Tongoloa、Carum、Oenanthe、Pternopetalum and Chamaesium were carried out. All ten species of Ligusticum have the same basic chromosome number, x =11, but they could be differentiated by their karyotype formula and quantitative parameters of the karyotypes. L. capillaceum and L. pteridophyllum are tetraploid with the chromosome number of 2n = 4x = 44, other species are diploid with the chromosome number of 2n = 2x = 22. Our cytological results suggest that polyploidy did not play an important role in the chromosome evolution of Ligusticum. Ligusticum has great diversification in northwestern Yunnan of China, so this region is probably the most diversified center for it. 2.Molecular systematics Maximum parsimony and Bayesian analyses of partitioned nrDNA ITS and cpDNA data sets or the combined molecular data, revealed a robust and well resolved phylogeny: (1) The phylogenetic placements of all ten Chinese endemic genera are resolved: Chuanminshen allies with Changium and Cyclorhiza in tribe Komarovieae; Notopterygium and Sinolimprichtia along with many other taxa of Sino-Himalayan distribution comprise a well-supported East-Asia clade, Notopterygium is paraphyletic, with the Chinese endemic species Vicatia bipinnata and Haplosphaera phaea arising from within it; Chaerophyllopsis finds affinity within tribe Scandiceae subtribe Scandicinae; Harrysmithia falls within the Acronema Clade; Melanosciadium embeds in Angelica in tribe Selineae, and Dickinsia is confirmed as a member of Apiaceae subfamily Azorelloideae; On the basis of phylogenetic analyses of ITS data, Nothosmyrnium allies strongly with tribe Pimpinelleae, while the results of MP and BI analysis of cpDNA data are equivocal in its placement; (2) Three major clades heretofore unrecognized in the subfamily were revealed (Acronema Clade, Chamaesium Clade, Sinodielsia Clade) and one (East Asia Clade, or the Physospermopsis Clade) was expanded considerably from its previous circumscription. The Acronema Clade was a well-supported sister group to tribe Scandiceae; the Chamaesium Clade occupied a basal and isolated position within the subfamily; the Sinodielsia Clade was either sister group to Ligusticum acuminatum Franch., or paraphyletic to tribe Apieae depending upon the analysis; and the East Asia Clade was a sister group to the Komarovia Clade; (3) The Acronema Clade includes species with an almost exclusively east Asian distribution, although its members are morphologically very heterogeneous and taxonomically complex, the same for the East-Asia Clade. These two Clades all belong to the East Asian Phytogeographycal type; (4) Several genera were confirmed as monophyletic, e.g. Pternopetalum, Chamaesium, Cyclorhiza. A large number of genera, which are notoriously difficult to define, having diffuse generic boundaries and heterogeneous patterns of variation (e.g., Ligusticum、Peucedanum、Physospermopsis、Pimpinella、Pleurospermum、Sinocarum、Tongoloa and Trachyspermum etc.) were not monophyletic based on our present study; (5) Sinodielsia、Meeboldia and Vicatia are distinct and distantly related, Peucedanum delavayi, phylogenetically distant from Sinodielsia and Meeboldia and united with three Ligusticum species in tribe Selineae; (6) Apiaceae subfamilies Saniculoideae, Azorelloideae, and Mackinlayoideae are each monophyletic, and show successive sister group relationships to subfamily Apioideae. This is the first study to incorporate a broad sampling of Chinese endemics from Apiaceae subfamily Apioideae. Aside from providing a framework for taxonomic revisions, the phylogenetic structure recovered in this study for subfamily Apioideae will lay the foundation for future investigations of evolutionary patterns of morphological characters and biogeography. For Acronema clade and East-Asia clade, further sampling is necessary, as well as the acquisition of additional sequence data from the chloroplast genome to resolve distal relationships within the subfamily, before a comprehensive classification can be obtained. 3.Taxonomic treatment Based on evidence from morphology, molecular phylogeny and biogeography, while considerring the congruency of relationships and high branch support, Komarovia Clade and Chamsesium Clade are recognized at the tribal rank, i.e. Komarovieae J. Zhou & S. R. Downie and Chamaesieae J. Zhou & F. D. Pu. The monotypic tribe Chamaesieae represents one of the earliest diverging lineages of subfamily Apioideae in Asia. 4.Biogeography The family Apiaceae is widely distributed across temperate regions and especially in Central Asia, with two distribution centers (one is Mediterranean area, the other is western United States and Mexico). Based on previous studies, the ancetstor of the two largest subfamiles of Apiaceae (Apioideae and Saniculoideae) may have evolvd in southern Africa. In our analyses, ten Chinese endemic genera were dispersed into seven major clades (Pimpinelleae, Acronema Clade, Selineae, Komarovieae, Scandiceae, East- Asia clade and Azorelloideae), represented different evolutionary stage of Apiaceae. The phylogenetic placements of Dickinsia, Changium, Chuanminshen, Cyclorhiza, Sinolimprichita and Notopterygium within Apiaceae are relatively basal, so belived to be paleo-endemic genera; Chaerophyllopsis, Harrysmithia, Mlanosciadium and Nothosmyrnium are in a position relatively distal, and may belong to neo-endemic genera. According to the phylogenetic placement and distribution of endemic genera, Yunnan and Sichuan Provinces of Kangdian Ancient Land are probable the secondary evolutionary centers for Chinese Apiaceae. One the one hand, the Tibetan plateau, with its dramatic variations in topography and climate, reserved the ancient elements of the family; on the other hand, the uplift of Tibetan plateau in Tertiary, contribute to the diversity of Apiaceae and some new taxa occurrence. 5. Chromosome evolution in subfamily Apioideae Optimization of currently available chromosome information for subfamily Apioideae onto the phylogenies indicates: (1) x= 11(plesiomorphy)occurred in most genera of Komarovieae through Selineae; (2) The plesiomorphy x= 6 is maintained in tribe Chamaesieae, and the state reverses to x= 6 from the ancestor with x= 11 in Sium and Oreomyrrhis; (3) In Tongoloa silaifolia, Anthriscus sylvestris and Torilis japonica, x= 8 is autopomorphic; (4) x= 9 is synapomorphic in Pleurospermeae, Pimpinelleae and Pternopetalum; (5) x= 10 originated multiple times independently in Apioideae, and is autopomorphic for Nothosmyrnium; (6) In phylogenetic trees, Pimpinella species were dispersed into three major clades: taxa with x= 11 united with Selineae; taxa with x= 9 fell into the tribe Pimpinelleae; and taxa with x= 10 comprise a subclade (Chinese Pimpinella subclade) of East-Asia Clade. That is, the basic chromosome number of each coincide with their phylogenetic placement; (7) In our analysis, Pleurospermum species were fell into Pleurospermeae with x= 9 and Acronema Clade with x= 11. However, the chromosome information for Pleurospermum is insufficient to discern its infrageneric taxonomic treatment.
语种: 中文
内容类型: 学位论文
URI标识: http://ir.kib.ac.cn/handle/151853/480
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中国伞形科芹亚科的分子系统学研究.周静[d].中国科学院昆明植物研究所,2009.20-25
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