基于ITS和matK序列的獐牙菜亚族 (龙胆科龙胆族)分子系统学

为探讨獐牙菜亚族（subtribe Swertiinae）各属之间和一些属内的系统关系,本研究选取了该亚族14属68种1变种,采用最大简约法（maximum parsimony）和贝叶斯法（Bayesian inference）对样品核基因ITS和叶绿体基因matK的两个片段进行独立和联合分析。结果显示：Bartonia位于亚族的最基部;喉毛花属（Comastoma）、肋柱花属（Lomatogonium）和假龙胆属（Gentianella）都非单系,处于同一个较为进化的分支中;獐牙菜属折皱组（Swertia sect. Rugosa）和獐牙菜组（S. sect. Swertia）亲缘关系最近,宽丝组（S. sect. Platynema）和藏獐牙菜组（S. sect. Kingdon Wardia）亲缘关系最近;口药花属（Jaeschkea）与獐牙菜属多枝组（S. sect. Ophelia）的大籽獐牙菜（Smacrosperma）亲缘关系最近。同时讨论了獐牙菜亚族形态分类与分子数据不一致的原因。     

肋柱花属的系统研究

本文对肋柱花属的属下分类、系统发育和地理分布等方面进行了深入研究。文用分支系统学的方法和原理,用计算机PAUP程序处理,获得了几个最简约的支序图。 肋柱花属属于龙胆亚族辐状花冠群,在这群中,论亲缘关系它与辐花属最近,与獐牙菜属次之,而与黄秦艽属关系较远。 獐牙菜属在进化程度上较肋柱花属低,因此它被选为肋柱花属的外类群。 经过支序分析,肋柱花属的18个种根据Hennig的“共近裔性原则”可组合为三个组,其中肉质根茎组为较原始的组,肋柱花组为中级进化水平的组,合萼组是进化程度最高的组。 肋柱花属是北温带分布型的属,分布于亚洲、欧洲及北美洲,直达北极。从种的地理分布型分析,表明秦岭一横断山区是本属的起源与分化中心。 随文报道了一个新组、一个新种和一个新变种。 研究了全部种类的命名模式。     

基于不同DNA序列对四数獐牙菜系统位置分析

以扁蕾属植物为外类群,基于trnL-trnF序列、rpl16序列以及 ITS序列,采用最大简约法对四数獐牙菜的系统位置进行了分析.结果显示:尽管在ITS系统发育树、trnL-trnF系统发育树和rpl16系统发育树中所用的种和数量有所不同,所得到的3种系统发育树的拓扑结构也不一致,但四数獐牙菜的系统位置在3种系统发育树中的结果基本一致.在trnL-trnF系统发育树中,四数獐牙菜与花锚属的H.elliptica、H.brevicornis和H.weddelliana形成了一个单系群;而在ITS和rpl16系统发育树中,四数獐牙菜与歧散獐牙菜先聚在一起,再与花锚属的植物聚为一支支持率较高的单系支.研究表明,四数獐牙菜与Anagalidium属的歧散獐牙菜的亲缘关系最近,其次是与花锚属的植物,而与獐牙菜属植物的亲缘关系较远.本实验系统发育的结果支持将四数獐牙菜从獐牙菜属中分出而归入Anagalidium属的观点.     

龙胆族(龙胆科)分类与进化研究进展

龙胆是著名的高山植物,广布世界温带地区,不仅可作药用,也可用于园艺观赏。龙胆科分为6个族,其中龙胆族占龙胆科物种数的一半以上;龙胆族包括龙胆亚族和獐牙菜亚族,分类与进化方面的相关研究众多。随着分子生物学在龙胆族研究中的应用,近些年对多个类群进行了研究并发表了多个新属和新种。该文从属间和属内两个层次梳理了龙胆族近年来在分类和进化研究中的最新进展,总结了新发表的类群。并分析了现存的问题,重点强调在今后的研究中用特别注意以下几点:(1)形态性状的准确掌握对龙胆族的研究至关重要;(2)发表新类群应当谨慎;(3)单拷贝核基因有助于解析龙胆族复杂的系统发育关系;(4)关注杂交和多倍化等在龙胆族进化中的作用。     

四数獐牙菜的胚胎学及其系统学意义

四数獐牙菜花药四室,药壁发育为双子叶型;绒毡层腺质型。小孢子母细胞减数分裂为同时型, 四分体的排列方式为四面体形;成熟花粉为3-细胞。子房2心皮,2心皮连接处强烈内凸,4列胚珠。薄 珠心,单珠被,直生胚珠。胚囊发育为蓼型。胚乳发育为核型。胚胎发育为茄型酸浆I变型。反足细胞 在胚囊成熟时期宿存。果实成熟时,种子只发育至球型胚阶段。承珠盘存在。与已研究的獐牙菜种类 的胚胎特征相比较,该种存在三个显著的特点：直生胚珠,心皮连接处强烈内凸,承珠盘存在。这三个特 征目前仅在龙胆族中的花锚属中发现。经过性状分析,它们均是演征,此三个共有演征表明四数獐牙菜 与花锚属是一单系群,它们的亲缘关系最近。这与Yun &; Kupfer(1995)的分子证据部分吻合,因为分 子系统树上的花锚单系支包括花锚、四数獐牙菜和北美的密花属(广义獐牙菜属中的密花组),而后者的 胚胎学特征未有任何报道。胚胎学研究暗示四数獐牙菜的分类学等级有待重新评价,广义的獐牙菜属 有可能不是一单系发生群。     

珍珠菜属系统发育关系的初步研究

本文运用形态学性状对珍珠菜属（Lysimachia)进行系统发育分析。内类群包括珍珠菜属29个代表种以及珍珠菜族其它单种、寡种属;仙客来属（Cylclamen)被选作外类群。最简约性分析表明,珍珠菜属并不构成－自然分类群;在其严格一致化树的二岐分支中,异花珍珠菜（L.crispidens)单独构成一支,其它所有内类群构成一支。香草亚属（subgen.Idiophyton)、木黄连花亚属（subgne.Lysimachiopsis)以及珍珠菜亚属（subgen.palladia)均表现为单系群,而黄连花亚属（subgen.Lysimachia)则为一异形的并系群。球尾花亚属（subgen.Naumburgia)仍“内藏”于珍珠菜属的主体之中。而喉鳞花亚属（subgen.Seleucia)则偏离出来而与七瓣莲属（Trien talis)构成姐妹群。如何准确地界定珍珠菜属和进行属下分类群的划分,还需进一步研究。     

藏药提宗龙胆和线叶龙胆花中四种苦苷类成分的HPLC测定

应用HPLC分析方法同时测定藏药提宗龙胆和线叶龙胆两种植物花中落干酸、獐牙菜苦苷、龙胆苦苷、獐牙菜苷4种苦苷类成分的含量.采用Econosphere C18色谱柱(250×4.6 mm,5 μm),以甲醇-0.5%乙酸为流动相进行梯度洗脱,流速为1.0 mL/min;检测波长为245 nm.结果表明,除提宗龙胆中未检出獐牙菜苦苷外,其它成分均在两种植物中存在,但含量存在一定的差异.     

西藏色季拉山区龙胆科植物种子微形态特征及其系统分类学意义

利用扫描电镜和解剖镜对西藏色季拉山区分布的龙胆科6属20种植物种子微形态进行观察研究。结果显示：这20种植物的种子颜色有黑色、黄色、黑褐色、黄褐色、灰褐色和红棕色6种;种子形状分为长卵球形、卵球形、椭圆球形、矩圆形和近球形;表皮纹饰可分为4种类型,即网纹型、条纹型、疣状型和嚼烂型,其中网纹型又细分为6个亚型。根据种子微形态特征,笔者对龙胆科相关属的进化趋势、分类学意义和分类鉴定等进行了讨论,认为獐牙菜属（Swertia L.）和龙胆属（Gentiana（Tourn.）L.）进化程度较高;从种子微形态特征上看肋柱花属（Lomatogonium A.Br.）、花锚属（Halenia Borkh.）和假龙胆属（Gentianella Moench.）间有较近的亲缘关系;龙胆科各属种子表皮纹饰特征与分类系统基本吻合。本研究表明种子微形态在植物分类研究中具有十分重要的意义,可作为植物系统划分、鉴定和进化研究的参考性状。     

苦龙胆酯苷的研究进展

苦龙胆酯苷是一种裂环烯醚萜苷类化合物,又称为苦龙苷或龙胆苦酯苷。其分子式为:C27H28O14,分子量为576.52,是已知最苦的裂环烯醚萜苷类化合物。目前已证明可从川东獐牙菜、印度獐牙菜、龙胆草以及辐花肋柱花中提取。辐花肋柱花是最新证明的可提取苦龙胆酯苷的植物。苦龙胆酯苷具有助消化,保肝,抗皮肤肿瘤,抗黑热病等药理活性。在古代印度传统医药以及中药藏药中,苦龙胆酯苷的来源植物是治疗消化系统相关疾病的一味常见的草药,如保肝、抗糖尿病等。在体内药代动力学研究中,兔静脉注射苦龙胆酯苷,在血中有较快的清除率(2.62±0.41 L/h/kg)和广泛的体内组织分布(1.08±0.44 L/kg);采用游离、脂质体和囊泡体形式给仓鼠用药具有明显保护肝肾功能且未发现明显不良反应。     

獐牙菜属植物花蜜腺形态及解剖学

在扫描电镜下观察了獐牙菜属Swertia L.10组30种植物花蜜腺的数目,位置,形态和附属物等特征;同时还利用光镜对各组代表种的蜜腺结构进行了解剖学观察。结果表明獐牙菜属花蜜腺外部形态多种多,但在组与组之间无明显间断,演化序列呈梯度变化;内部结构基本相同,为不具维管束的结构蜜腺,且均为淀粉型蜜腺。因此,从花蜜腺的角度不支持将獐牙菜属划分为小属的观点,同时,还结合其它证据讨论了花蜜腺特征的演化趋势。     

Pollen morphology of the Swertiinae (Gentianaceae): phylogenetic implications

Recent molecular surveys of the Swertiinae (Gentianaceae–Gentianeae) revealed unexpected phylogenetic relationships, including polyphyly of the genera Gentianella , Jaeschkea , Lomatogonium and Swertia . To find new non-molecular characters supporting the phylogeny, we examined the exine variation of 73 species of all major lineages of subtribe Swertiinae using environmental scanning electron miscroscopy supplementing older, mainly light microscopical, studies. In contrast to previous studies, we were able to pick out taxa from phylogenetic key positions with particular focus on Swertia . Many distantly related taxa such as parts of Frasera , Gentianopsis , Halenia , Gentianella , Megacodon and several lineages of Swertia share a striate–reticulate or reticulate exine pattern. This is interpreted as the plesiomorphic character state of Swertiinae. There is also considerable variation of derived patterns; for example, different types of microechinate or almost smooth pollen was repeatedly observed in distantly related groups. Another extreme was the ring-shaped reticulation found in a North American species of Gentianopsis . Unfortunately, major relationships as revealed by molecular analyses were rarely supported because of the abundance of the plesiomorphic type and homoplasy even on low taxonomic levels; for example, within Lomatogonium . Exine variation was particularly useful in characterizing independent lineages of Swertia . For example, according to pollen characters and in agreement with other data, the Asian Swertia cuneata is a sister group of a strongly diversified African lineage and Swertia yunnanensis , which is rather aberrant in flower morphology, seems close to parts of Lomatogonium .  © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society , 2008, 157 , 323–341.     

The Origin,Dispersal and Formation of the Distribution Pattern of Swertia L. (Gentianaceae)

The genus Swertia is one of the large genera in Gentianaceae, including 154 species, 16 series and 11 sections. It is disjunctly distributed in Europe, Asia, Africa and N. America, but entirely absent from Oceania and S. America. According to Takhtajan’s (1978) regionalization of the world flora, Swertia is found in 14 regions. Eastern Asiatic region with 86 species, of which 58 are local endemics, 13 series and 9 sections, ranks the first among all the regions. The highest concentration of the taxa and endemics in Eastern Asiatic region occurs in SW China-Himalayan area (Sikang-Yunnan P. , W. Sichuan, W. Yunnan-Guichou Plateau of China and NE. Burma, N. Burmense P. , E. Himalayan P. and Khasi-Manipur P. ). In this area there are 74 species (48 endemics), 12 series, and 9 sections; thus about half species of the world total, three quarters of series and 82% of sections occur in this small area. Besides, the taxa at different evolutionary stages in Swertia also survive here. It is an indication that SW. China-Himalayan area is a major distribution centre of the genus Swertia. In addition, Sudan-Zambezian Region in Africa, with 22 species, 4 series and 2 sections, is a second distribution centre. The primitive type of the genus Swertia is Sect. Rugosa which consists of 2 series and 23 species. It is highly centred in the mountains of SW. China (Yunnan, Sichuan, Guizhou and SE. Xizang) where 2 series and 16 species occur. Among them 15 species of Ser. Rugosae were considered as the most primitive groups in this genus. From our study, the outgroup of Swertia is the genus Latouchea Frahch. , which is distributed in Yunnan, Sichuan, Guizhou, Hunan, Guangdong, Guangxi and Fujian. The two groups overlap in distribution in SW. China. According to the principle of common origin, the ancestor of two genera ap peared most probably in this overlapping area. It was inferred that SW. China Was the birth-place of the genus Swertia. Four sections of Swertia have different disjunct distribution patterns: Sect. Ophelia is of Tropic Asia, Africa and Madagascar disjunct distribution; sect. Swertia is of north temperate distribution; sect. Spinosisemina is in Tropical Asia (Trop. India to S. China and Philipines); sect. Platynema also is in Tropical Asia (Java, Sumatra, Himalayas to SW. China). These disjunct patterns indicate that the Swertia floras between the continents or between continent and islands have a connection with each other. From paleogeographical analysis, Swertia plants dispersed to Madagascar before the Late Cretaceous, to SE. Asian Islands in the Pleistocene, to North America in the Miocene. The distribution of Swertia in Madagascar might be later than that in Asia. Therefore the origin time of the genus Swertia was at least not later than the Late Cretaceous, and might be back to the Mid-Cretaceous. The genus Swertia first fully developed and differentiated, forming some taxa at different evolutionary stages (Rugosa, Swertia, Poephila, Ophelia and Platynema etc. ) in the original area, and these taxa quickly dispersed in certain directions during the Late Cretaceous-Middle Tertiary when the global climate was warm and no much change. There seem to be three main dispersal routes from the origin area to different continents; (1) The westward route i. e. from SW. China, along the Himalayas area to Kashmir, Pakistan, Afghanistan and Iran, and then southwestwards into Africa throuth Arabia. Four sections (Poephila, Macranthos, Kingdon-Wardia and Ophelia) took this dispersal route. Most species of sect. Ophelia dispersed along this route, but a few along southern route and north ern route. Sect. Ophelia greatly differentiated in Africa and the African endemic sectionSect. Montana was derived from it. The two sections form there a second distribution center of Swertia. (2) The southward route, i. e. towards S. India through the Himalayas, and towards SE. Asian islands through C. and S. China, Indo-China. Along this dispersal route sect. Platynema, Sect. Spinosisemina and a few species of Sect. Ophelia dispersed; (3) The northward rout, i. e. northwards across N. China, C. Asia to a high latitude of Euasia, and also through E. Asia into N. America. The following groups took this route: sect. Rugosa, sect. Swertia, sect. Frasera, sect. Heteranthos and sect. Ophelia ser. Dichotomae. Therefore, it seems that the genus Swertia originated in SW. China and then dispersed from there to N. and S. Asia, Africa, Europe and North America and formed the moderndistribution pattern of this genus.     

Secretory hairs in Gentiana and allied genera (Gentianaceae, subtribe Gentianinae) from the Iberian Peninsula

Eleven species from six different sections of the genus Gentiana , as well as one species each of genera Gentianella (G. campestris), Gentianopsis (G. ciliata), Comastoma (C. tenellum) and Swertia (S. perennis) have been studied by light microscopy for the presence of hairs in floral as well as in vegetative parts. Hairs are produced in the calyx and vegetative leaves of all of them, and also in the corollae of the last three species. They fall into two different types: those found in the corolla of Gentianopsis ciliata are non-secretory, while in the rest of the species studied, and also in the calyx and leaves of G. ciliata , they produce a mucilaginous secretion. Calycine and foliar hairs are always produced in the adaxial epidermis at the base of the foliar organ, and are considered as mucilage secreting colleters. The presence of colleters in vegetative organs has not been adequately considered in previous taxonomic accounts, in spite of their presumed significance.     

堇菜属组间的系统发育与亲缘关系：基于trnL-trnF、psbA trnH、rpL16、ITS序列，细胞学以及形态学证据

以鼠鞭草(Hybanthus enneaspermus)、鳞隔堇(Scyphellandra pierrei)、雷诺木(Rinorea bengalensis)作为外类群,对堇菜属(Viola)20个类群的trnL trnF序列,17个类群的psbA trnH、rpL16序列以及1个类群的nrDNA ITS序列进行了测定,并从GenBank下载相应的序列,运用最大简约法以及贝叶斯推论法进行系统分析,构建系统发育树。结果表明：堇菜亚属(subgen.Viola)不是一个单系类群,并明确了堇菜属部分组间类群的亲缘关系。本文还结合形态与细胞学证据对堇菜属进行性状演化的推测。结果表明：1）直立茎较匍匐茎、莲座状茎（叶基生）原始;2）托叶边缘长流苏状与托叶1/2~3/4合生分别是鸟嘴柱头堇菜组（sect.Trigonocarpae）和合生托叶组（sect.Adnatae）演化路线的重要性状标志;3）花柱样式从柱头无喙演化至柱头有喙,并由柱头简单演化至柱头复杂,再趋向于柱头简化。     

Pollen morphology of selected species of Allium (Alliaceae) distributed in Iran

Pollen morphology of 19 species of Allium representing six subgenera and 12 sections, were studied. The following characters were recognized as important for separating taxa at different taxonomic ranks: the sulcus, presence or absence of perforations on the pollen surface, density of perforations, size of perforations and size of the pollen grains. Rugulate, rugulate–striate, and simple–perforate exine ornamentation were distinguished. The type of sulcus is very characteristic in A. subgen. Allium sect. Allium . The extended sulcus was not observed in any species the other sections of subgen. Allium studied here, such as sect. Caerulea ( A. capitellatum ), sect. Codonprasum ( A. lenkoranicum and A. stamineum ) and sect. Avulsea ( A. rubellum and A. umblicatum ). Imperforated pollen grains were observed in representatives of A. subgen. Reticulatobulbosa and subgen. Polyprason .     

堇菜属组间的系统发育与亲缘关系：基于trnL-trnF、psbA-trnH、rpL16、ITS序列,细胞学以及形态学证据

以鼠鞭草（Hybanthus enneaspermus）、鳞隔堇（Scyphellandra pierrei）、雷诺木（Rinorea benga-lensis）作为外类群,对堇菜属（Viola）20个类群的trnL-trnF序列,17个类群的psbA-trnH、rpL16序列以及1个类群的nrDNAITS序列进行了测定,并从GenBank下载相应的序列,运用最大简约法以及贝叶斯推论法进行系统分析,构建系统发育树。结果表明：堇菜亚属（subgen. Viola）不是一个单系类群,并明确了堇菜属部分组间类群的亲缘关系。本文还结合形态与细胞学证据对堇菜属进行性状演化的推测。结果表明：1）直立茎较匍匐茎、莲座状茎（叶基生）原始;2）托叶边缘长流苏状与托叶1/2～3/4合生分别是鸟嘴柱头堇菜组（sect. Trigonocarpae）和合生托叶组（sect. Adnatae）演化路线的重要性状标志;3）花柱样式从柱头无喙演化至柱头有喙,并由柱头简单演化至柱头复杂,再趋向于柱头简化。     

A new species of Swertia L. (Gentianaceae) from Nepal

Swertia barunensis P. Chassot sp. nov. from Nepal is described and illustrated. It was collected in 1997 in an alpine meadow in the Makalu Barun National Park at an elevation of 4200 m. It belongs to Swertia section Macranthos T.-N. Ho & S.-W. Liu and resembles S. pseudohookeri H. Smith, from which it differs mainly by the shape of the nectary and the exomorphic seed structure. A key to all the species of sect. Macranthos is provided. The affinities of S. barunensis with some other taxa in the subtribe Swertiinae (Griseb.) Rchb. are also briefly discussed.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 141, 389–394.     

Taxonomic notes on Astragalus sect. Malacothrix (Fabaceae) from Iran

Astragalus porphyrogrammus , occurring in the Zarin-Abad of Zanjan (Iran), is described and illustrated. This species belongs to A. sect. Malacothrix and seems to be distinct and interesting among the Iranian species. In this research, A. sect. Grammocalyx which was very artificial placed in A. subgen. Calycophysa transferred to A. subgen. Hypoglottis. Moreover, relationships between A. subgen. Hypoglottis and the closely related subgenera in genus Astragalus are discussed. Also A. lineatus is recorded for Iran.