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题名: 红豆杉科的系统学研究
作者: 周其兴
学位类别: 博士
答辩日期: 2001
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: 吴征镒
关键词: 红豆杉科 ; 三尖杉科 ; 罗汉松科 ; 系统学
学位专业: 植物学
中文摘要: 本文对红豆杉科系统学的研究历史进行了综述,对红豆杉科、三尖杉科、罗汉松科和南洋杉科的代表类群开展了叶表皮和叶片及叶柄横切、细胞学、等位酶和分子系统学实验,并尝试用分支分析的方法将多学科资料进行综合分析,以探讨红豆杉科的系统位置、分类学范畴和属间演化关系问题。主要研究结果和结论如下:1.形态解剖学对上述类群的11属23种进行了叶片和叶柄的比较形态解剖学研究。结果表明叶结构和表皮形态特征在属一级比较一致,但在属间具有明显的差异。红豆杉科中红豆杉属、白豆杉属和南紫杉属的叶片结构最为相似,表现在它们的叶片中均具有中脉,而不具有树脂道,但叶表皮上气孔类型不同,气孔之间的细胞在是否具有颗粒状突起点上也不同。穗花杉属和榧树属的叶片结构比较相似,叶片气孔类型都为单环型,中脉下均有1条树脂道。但榧树属气孔带外具有独特的角质钉,气孔带上的气孔排列紧密。三尖杉属的叶片结构与穗花杉属在中脉和气孔类型上十分相似。尽管罗汉松科4个属的叶片结构在叶型、树脂道排列方式上各不相同,但气孔类型、叶表皮细胞均相似或相同。竹柏属同贝壳杉属的叶片在外形上很相似,并且均为平行脉,但叶片内部树脂道排列方式和叶肉组织分化情况迥然不同。2.细胞学报道了红豆杉科、三尖杉科和罗汉松科8属的核形态结构。红豆杉科4属的间期核构型和前期染色体分别都为复杂染色中心型和中间型。红豆杉属和白豆杉的体细胞中期染色体数目都为24。其中,都有2条T染色体和20条左右的m染色体,核型不对称性都属于2B型,N.F.值为46。穗花杉属的体细胞分裂中期染色体数目为2n=36,核型都为2n=36=8m+28T,核型不对称性为1B型,N.F.值为44。榧树属的中期染色体核型多为2n=22=22m(2sat),仅个别种为2n=22=20m(2sat)+2sm,但N.F.值也为44,核型不对称性为lA型。三尖杉科的间期核和前期染色体特征同红豆杉科一致,分别为复杂染色中心型和中间型,体细胞分裂中期染色体数目为24,核型不对称性属于2A或在篦齿三尖杉中为2B型。三尖杉的染色体结构都是1至22号染色体为m染色体,23和24号染色体为sm染色体并具随体,其随体较大而稳定,N.F.值为48。在罗汉松科中,鸡毛松属、竹柏属和罗汉松属的有丝分裂前期染色体都为中间型,但间期核类型各不相同。它们的也不同,鸡毛松属为2n=20,基数为X=10;竹柏属为2n=26,基数为X=13;罗汉松属为2n=38,基数为X=19。3.等位酶分析采用水平式淀粉凝胶电泳技术,对中国产红豆杉属3种1变种以及白豆杉共12个居群的遗传多样性进行了检测。对7个酶系统9个酶位点的分析结果表明,红豆杉属和白豆杉的种内遗传多样性水平较高,各类群的多态位点比率(P)为44.4-77.8%,等位基因平均数(A)为1.6-2.1,平均期望杂合度(He)和实际观察杂合度(Ho)分别为0.065-0.152和0.068-0.111。除须弥红豆杉与红豆杉属其它几个种的遗传一致度稍低外(I=0.727-0.804),红豆杉属的其它几个种的遗传一致度较高(O.906-0.995),体现其亲缘关系很近。我们建议红豆杉属的这几个种合并为一个种并根据地理分布及不同生境发生的变异在种下设地理宗。同时根据等位酶分析结果,对白豆杉的系统位置进行了讨论。4.分子系统学测定了红豆杉科所有5属及相关类群三尖杉科、罗汉松科、南洋杉科部分代表类群的叶绿体trnL/F的两个非编码基因序列。运用PAUP软件,分别用最大简约法、最大似然法和邻接距离法进行分析。3种方法分析所得的分支图构型基本一致。主要有2个明显的大分支,其中之一包括红豆杉科和三尖杉科,另一支包括罗汉松科和南洋杉科的代表类群。松科的2个代表类群作为外类群位于内类群的基部。在红豆杉科和三尖杉科一支中,红豆杉属、白豆杉属和南紫杉属形成了一支,尤其是白豆杉和红豆杉属靠得最近。穗花杉属和榧属以及三尖杉属形成了另外一支。罗汉松科4属形成了较稳定的单系类群。5.分支系统学共选用来自形态学、解剖学、胚胎学和细胞学等多学科的127个性状进行分析。分析结果表明:1)红豆杉科中,红豆杉属和白豆杉属关系比较近,穗花杉属和榧属关系也比较近,南紫杉属可能与红豆杉属和白豆杉属的关系更近,三尖杉属为红豆杉科5属的姐妹群。2)罗汉松科的4属也组成了一个单系类群。3)南洋杉科的2属聚在一起,与罗汉松科有相对较近的亲缘联系。与基于叶绿体基因trnL/F所构建的系统发育关系不同的是,三尖杉属的位置在分子证据支持下置于红豆杉科中,与穗花杉属和榧属有很近的关系,而基于形态学等多学科证据的分析结果表明三尖杉属与红豆杉科5属构成了姐妹群关系。6.结论综合各学科证据表明,红豆杉科置于松柏类是合适的,在红豆杉科内,红豆杉属(包括白豆杉)与南紫杉属组成了红豆杉族,穗花杉属和榧属组成了榧族。三尖杉属与红豆杉科具有比较近的亲缘关系。中国罗汉松科4属为很好的单系类群,不支持竹柏属独立为科的观点。
英文摘要: The yew family, Taxaceae Gray generally comprises 5 genera, i.e. Amentotaxus Pilger, Austrotaxus Compton, Pseudotaxus Cheng, Taxus Linn, and Torreya Am. The systematic works of the family were carried out by means of leaf morphology and anatomy, cytology, allozyme analysis and molecular systematics in the present study. Also, a cladistic analysis of 127 characters from multi-discipline was conducted. The main results are given as follows: Anatomy The results of leaf morphology and anatomy from 11 genera in the Taxaceae, Cephalotaxaceae, Podocarpaceae and Araucariaceae indicate that Taxus, Pseudotaxus and Austrotaxus are very alike within the Taxaceae, and Amentotaxus and Torreya are very close to each other. Also, Amentotaxus is similar to Cephalotaxus in that their resin canals, but the epidermal cells are somewhat different though their stomata were of the same type. 4 Chinese genera of the Podocarpaceae are quite different from each other, although they have the same type of stoma and similar epidermal cells. Obviously, Nageia and Agathis are very similar in the external leaf morphology, but distinctly different in the arrangement of resin canals in the leaves and petioles. Agathis is similar to Araucaria within the Araucariaceae. Cytology The representative species of 8 genera of the Taxaceae, Cephalotaxaceae and Podocarpaceae were karyomorphologicaly investigated. All the genera of Taxaceae and Cephalotaxaceae have the similar resting nuclei and prophase chromosome. Taxus and Pseudotaxus have the same metaphase chromosome number (2n=24) and very similar karyotypes. Although Amentotaxus and Torreya have a different metaphase chromosome number, which 2n=36 for the former and 2n=22 for the latter, but they share the same N.F. (number of fundamental) value. And this may be coursed by Robertson variation. Cephalotaxus has the same metaphase chromosome number of Taxus, but have different characteristic chromosomes (the last pair chromosomes). The prophase chromosomes of 7 species in 3 genera of Podocarpaceae are of the same type, but their interphase nuclei are of different. And the metaphase chromosome numbers are also found different from each other, 2n=20 is counted for Dacrycarpus, 2n=26 for Nageia, and 2n=38 for Podocarpus, respectively. Allozyme Analysis The genetic diversity of 12 populations of four Taxus taxa and and Pseudotaxus chienii in China were studied vising the method of horizontal starch gel electrophoresis. Genetic diversity within 5 taxa is relatively high. The results from allozyme analysis suggested that there were very close relationships between three species and one variety in Taxus, with high genetic identities (I) between them (from 0.727 to 0.995) except those between T. wallichiana var. wallichiana and other taxa in Taxus (0.727-0.804). Consequently, we proposed that these taxa in Taxus should be treated as a single species. Also the taxonomic position of Pseudotaxus chienii was discussed from the allozymic data. Molecular systematics Phylogenetic analysis of Taxaceae and its allies, together with 2 genera in the Pinaceae as outgroups, was performed based on two non-coding chloroplast sequences, the trnL (UAA) intron and intergenic spacer between trnL (UAA) and trriF (GAA). The analysis using PAUP 4.0 strongly identified two main ingroup clades: one clade contains 5 genera in Taxaceae and the only genus in Cephalotaxaceae, the other clade contains the genera of Podocarpaceae and Araucariaceae. In the first clade, Taxus, Pseudotaxus and Austrotaxus form a subclade, Taxus and Pseudotaxus were the closest groups, and Austrotaxus was the basal group to them. The second subclade comprised Amentotaxus, Torreya and Cephalotaxus. In the second clade, Nageia and Podocarpus had the closest relationships within the Podocarpaceae. In terms of Agathis in the Araucariaceae, it's the basal group to the Podocarpaceae. Based on these results, the systematic positions of the family Taxaceae and the genus Nageia were discussed. Ciadistics A cladistic analysis based on 127 chracteres from external morphology, wood and leaf anatomy, cytology, embryology, palynology, and chemistry was carried on using Maximum parsimony and Neighber-joining methods. The results show that: 1) Taxus and Pseudotaxus are very close in Taxaceae, while Amentotaxus and Torreya are very close to each other. Austrotaxus may be closer to Taxus and Pseudotaxus within Taxaceae. Cephalotaxaceae is the sister group to Taxaceae. 2) 4 Chinese genera in Podocarpaceae are a monophyletic group. And 3) the 2 genera of Araucariaceae also make up of a monophyletic group with relatively close relationship to Podocarpaceae. 6. Conclusion As inferred from the evidences from morphology, anatomy, cytology, embryology, molecular systematics and other disciplines, Taxaceae is better the member of conifers. And 2 tribes, namely Trib. Taxeae (Taxus, Austrotaxus) and Trib. Torreyeae (Amentotaxus and Torreya) may be included in the Taxaceae. Pseudotaxus is merged into Taxus as a subgenus. Cephalotaxus is temperaly be considered a sister group to Taxaceae. The generic status of 4 genera of Podocarpaceae was also supported.
语种: 中文
内容类型: 学位论文
URI标识: http://ir.kib.ac.cn/handle/151853/636
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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