地黄属分类学与系统学研究
摘要
地黄属(Rehmannia)目前记载共约6种,除作为常用中药的地黄(R.glutinosa)为东亚—日本分布外,均为我国特有植物。鉴于国内外研究偏重于地黄,疏于地黄属其它物种的现状,因此该属内各物种存在很多尚未解决的问题,如中药地黄的种质鉴定、高地黄(R.elata)、裂叶地黄(R.piasezkii)的分类学地位、属的系统发育等,生药种质资源和系统分类学基础研究均存在信息盲区,为此本文针对这些问题从分类学和系统学视角出发进行了探索。
研究首先从考证腊叶标本记载入手,结合野外考察数据,对地黄属各物种的地理分布和生境特征进行整理,获得了较为详细的地黄属植物生态习性资料,为分类学研究提供了生态学依据。接着对地黄属的果实、种子、幼苗、块根、叶、茎、花、花粉、胎座进行了形态学及解剖学观察,首次获得了各物种的详细鉴别特征及种内变异范围数据,为分类学修订提供了基础依据。果实和种子的形态学性状、花粉粒特征的特征性分析显示地黄属可能与Gratiola、Digitalis、Paederota、Wulfenia、Picrorhiza、Mazus、Buddleja等属有一定联系。
为了确定地黄属有关物种间的亲缘关系,研究进行了种间杂交育性实验,结果表明地黄与其余种的亲缘关系较远,裂叶地黄和高地黄在与其它种的杂交中表现完全一致;细胞学观察发现中药地黄是多倍体植物,并首次发现了4个种的体细胞染色体数目均为28条,与56条染色体的地黄相差1倍:植物化学分析表明,地黄属各物种都含有梓醇和桃叶珊瑚苷等环烯醚萜类成分,其中以地黄块根中的梓醇含量最为丰富。
研究为确定地黄属的系统位置,继续对属内各物种进行了核糖体内转录基因间区ITS的测序,首次获得5个种ITS区的碱基序列,Gene Bank登陆认证,登录号分别为:DQ069312(地黄)、DQ069313(天目地黄R.chingii)、DQ069314(茄叶地黄R.solanifolia)、DQ069315(高地黄)、DQ069316(裂叶地黄),序列长度均为613bp(不包括18S和26S部分的序列)。
在获得地黄属ITS序列信息的基础上,研究选择唇形目中关系较近的40个属进行分子系统发育分析,采用最大简约法、邻接法、贝叶斯法分别获得的3种系统树大致相同,结果显示由醉鱼草属(Buddleja)、藏玄参属(Oreosolen)等构成的分支与由地黄属的全部物种构成的分支之间的姐妹群关系固定不变。
将地黄属按6种植物经系统发育分析显示,地黄属是一个稳定的单系,共有3支:天目地黄与湖北地黄构成第1支,高地黄与裂叶地黄构成第2支,地黄和
There are about 6 species in Rehmannia, of which, Chinese traditional medicine R. glutinosa distributed from East Asia to Japan, all others are endemic to China. As worldwide studies emphasized on R. glutinosa but neglected other species, it leaves us many unsettled problems related to Rehmannia, such as, how to differentiate R. glutinosa from similar species, where is R. elata, and how about the phylogenetic development of the genus. This thesis is an exploration aimed to all those problems from the view of taxonomy and phylogeny.Start with specimen researches on Rehmannia geographic distribution and environmental adaptive traits both from herbaria and from field environments, detailed species data have been acquired, which provide ecological evidences for taxonomy. Next, morphological and anatomical studies performed on fruits, seeds, seedlings, root tubers, leaves, stems, flowers, pollens and placentas of Rehmannia, and many particular identifying characters and variations of each species acquired for the first time. By comparing fruit and seed morphology, pollen characters and placenta types with other genera, it shows that Rehmannia has some possible relationships with Gratiola、 Digitalis、 Paederota、 Wulfenia、 Picrorhiza、 Mazus and Buddleja.To explore the relationship among the species of Rehmannia, cross-fertilization have been performed. The results indicate that R. glutinosa exhibits wider distances . from other species, whereas R. piasezkii and R. elata are among the closest species. Cytological observation brings out the first records on the same chromosome number 2n=28 of R. piasezkii, R. elata, R. henry and R. chingii, and suggested polyploid nature in R. glutinosa. Phytochemical analysis reveals existence of iridoid glycoside (catalpol and aucubin), with the most abundant catalpol content in root tubers of R. glutinosa.Internal transcribed spacer (ITS) sequences of all the Rehmannia species have been sequenced, among which, 5 sequences (R.glutinosa, DQ069312; R. chingii, DQ069313; R. solanifolia, DQ069314; R. etata, DQ069315; R. piasezkii, DQ069316 are accepted by Gene Bank of NCBI.Molecular phylogenetic analysis using ITS sequences from Rehmannia and other 39 related genera have been performed under PAUP* 4.0bl0 and Mrbayes 3.0b4.
Three kind of consensus trees (Maximum Parsimony tree, Neighbor Joining tree and Bayesian Inference tree) show approximately similar branches with Rehmannia always a sister group of Buddleja and Oreosolen.For species within Rehmannia, phylogenetic analysis shows that it's a monophyletic group with 3 clades: clade 1, R. chingii and R. henryi; clade 2, R. elata and R. piasezkii; clade 3, R. glutinosa and R. solanifolia. Herein, the two species of clade 2 deserve to be considered one species because of the highest bootstrap percentages and morphological resemblance and their overlapping distribution, and clade 3 an advanced group considering its polyploid nature.For systematic position of Rehmannia in Lamiales, it shows that Rehmannia may be considered a member of Scrophulariaceae (sensu stricto), be close to tribe Gratiolieae and Rhinantheae of Veronicaceae, be far from Gesneriaceae and tribe Digitalieae of Veronicaceae, with supports from phylogenetic analysis, phytochemical studies, morphological and anatomical observes.Lastly, a revision of Rehmannia taxonomy achieved: (1) a new synonym of Rehmannia piasezkii Maxim.—Rehmannia elata N. E. Brown (syn. nov.), (2) an amended description of the genus and species, (3) a new key for species, and (4) a detailed feature collection for identifying Chinese traditional medicine R. glutinosa.
研究首先从考证腊叶标本记载入手,结合野外考察数据,对地黄属各物种的地理分布和生境特征进行整理,获得了较为详细的地黄属植物生态习性资料,为分类学研究提供了生态学依据。接着对地黄属的果实、种子、幼苗、块根、叶、茎、花、花粉、胎座进行了形态学及解剖学观察,首次获得了各物种的详细鉴别特征及种内变异范围数据,为分类学修订提供了基础依据。果实和种子的形态学性状、花粉粒特征的特征性分析显示地黄属可能与Gratiola、Digitalis、Paederota、Wulfenia、Picrorhiza、Mazus、Buddleja等属有一定联系。
为了确定地黄属有关物种间的亲缘关系,研究进行了种间杂交育性实验,结果表明地黄与其余种的亲缘关系较远,裂叶地黄和高地黄在与其它种的杂交中表现完全一致;细胞学观察发现中药地黄是多倍体植物,并首次发现了4个种的体细胞染色体数目均为28条,与56条染色体的地黄相差1倍:植物化学分析表明,地黄属各物种都含有梓醇和桃叶珊瑚苷等环烯醚萜类成分,其中以地黄块根中的梓醇含量最为丰富。
研究为确定地黄属的系统位置,继续对属内各物种进行了核糖体内转录基因间区ITS的测序,首次获得5个种ITS区的碱基序列,Gene Bank登陆认证,登录号分别为:DQ069312(地黄)、DQ069313(天目地黄R.chingii)、DQ069314(茄叶地黄R.solanifolia)、DQ069315(高地黄)、DQ069316(裂叶地黄),序列长度均为613bp(不包括18S和26S部分的序列)。
在获得地黄属ITS序列信息的基础上,研究选择唇形目中关系较近的40个属进行分子系统发育分析,采用最大简约法、邻接法、贝叶斯法分别获得的3种系统树大致相同,结果显示由醉鱼草属(Buddleja)、藏玄参属(Oreosolen)等构成的分支与由地黄属的全部物种构成的分支之间的姐妹群关系固定不变。
将地黄属按6种植物经系统发育分析显示,地黄属是一个稳定的单系,共有3支:天目地黄与湖北地黄构成第1支,高地黄与裂叶地黄构成第2支,地黄和
There are about 6 species in Rehmannia, of which, Chinese traditional medicine R. glutinosa distributed from East Asia to Japan, all others are endemic to China. As worldwide studies emphasized on R. glutinosa but neglected other species, it leaves us many unsettled problems related to Rehmannia, such as, how to differentiate R. glutinosa from similar species, where is R. elata, and how about the phylogenetic development of the genus. This thesis is an exploration aimed to all those problems from the view of taxonomy and phylogeny.Start with specimen researches on Rehmannia geographic distribution and environmental adaptive traits both from herbaria and from field environments, detailed species data have been acquired, which provide ecological evidences for taxonomy. Next, morphological and anatomical studies performed on fruits, seeds, seedlings, root tubers, leaves, stems, flowers, pollens and placentas of Rehmannia, and many particular identifying characters and variations of each species acquired for the first time. By comparing fruit and seed morphology, pollen characters and placenta types with other genera, it shows that Rehmannia has some possible relationships with Gratiola、 Digitalis、 Paederota、 Wulfenia、 Picrorhiza、 Mazus and Buddleja.To explore the relationship among the species of Rehmannia, cross-fertilization have been performed. The results indicate that R. glutinosa exhibits wider distances . from other species, whereas R. piasezkii and R. elata are among the closest species. Cytological observation brings out the first records on the same chromosome number 2n=28 of R. piasezkii, R. elata, R. henry and R. chingii, and suggested polyploid nature in R. glutinosa. Phytochemical analysis reveals existence of iridoid glycoside (catalpol and aucubin), with the most abundant catalpol content in root tubers of R. glutinosa.Internal transcribed spacer (ITS) sequences of all the Rehmannia species have been sequenced, among which, 5 sequences (R.glutinosa, DQ069312; R. chingii, DQ069313; R. solanifolia, DQ069314; R. etata, DQ069315; R. piasezkii, DQ069316 are accepted by Gene Bank of NCBI.Molecular phylogenetic analysis using ITS sequences from Rehmannia and other 39 related genera have been performed under PAUP* 4.0bl0 and Mrbayes 3.0b4.
Three kind of consensus trees (Maximum Parsimony tree, Neighbor Joining tree and Bayesian Inference tree) show approximately similar branches with Rehmannia always a sister group of Buddleja and Oreosolen.For species within Rehmannia, phylogenetic analysis shows that it's a monophyletic group with 3 clades: clade 1, R. chingii and R. henryi; clade 2, R. elata and R. piasezkii; clade 3, R. glutinosa and R. solanifolia. Herein, the two species of clade 2 deserve to be considered one species because of the highest bootstrap percentages and morphological resemblance and their overlapping distribution, and clade 3 an advanced group considering its polyploid nature.For systematic position of Rehmannia in Lamiales, it shows that Rehmannia may be considered a member of Scrophulariaceae (sensu stricto), be close to tribe Gratiolieae and Rhinantheae of Veronicaceae, be far from Gesneriaceae and tribe Digitalieae of Veronicaceae, with supports from phylogenetic analysis, phytochemical studies, morphological and anatomical observes.Lastly, a revision of Rehmannia taxonomy achieved: (1) a new synonym of Rehmannia piasezkii Maxim.—Rehmannia elata N. E. Brown (syn. nov.), (2) an amended description of the genus and species, (3) a new key for species, and (4) a detailed feature collection for identifying Chinese traditional medicine R. glutinosa.
引文
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