龙胆科獐牙菜属新分类纲要

獐牙菜属是龙胆科中的一个大属,广泛分布于亚洲、北美洲、欧洲和非洲.本文报道獐牙菜属下的两个新组(sect.Montana和sect.Echinulata)和七个新系(ser.Repentes,ser.Kilimandscharicae,ser.Coombosae,ser.Japonicae,ser.Swertopsis,ser.Pumilae和ser.Abyssinicae).同时对獐牙菜属中的一些类群进行了分类修订,本分类纲要将被獐牙菜属世界专著采用.     

Embryology of Swertia (Gentianaceae) relative to taxonomy

The embryological features of three species of Swertia ( s.l.) – S. erythrosticta , S. franchetiana , and S. tetraptera – were characterized, and the observations were used, together with previously gathered data on other species, to evaluate a recently proposed polyphyly, based on molecular data, of Swertia s.l. Comparisons of species within the genus showed that they have diversified embryologically, and there are significant between-species differences. Notable features that vary between species include the number of cell layers that form the anther locule wall, the construction of the wall of the mature anther, tapetum origin, the cell number in mature pollen grains, the structure of the fused margins of the two carpels, the ovule numbers in placental cross-sections, the shape of the mature embryo sac, the degree of ovule curvature, antipodal variation and the presence of a hypostase, and seed appendages. They share characters that are widely distributed in the tribe Gentianeae, such as a dicotyledonous type of anther wall formation, a glandular tapetum with uninucleate cells, simultaneous cytokinesis following the meiosis of the microsporocytes, tetrahedral microspore tetrads, superior, bicarpellary and unilocular ovaries, unitegmic and tenuinucellar ovules, Polygonum -type megagametophytes, progamous fertilization, nuclear endosperm, and Solanad-type embryogeny. The presence of variation in embryological characters amongst the species of Swertia s.l. strongly supports the view that Swertia s.l. is not a monophyletic group. Frasera is better separated from Swertia s.l. as an independent genus, and is only distantly related to Swertia s.s. judging from the numerous differences in embryology. Swertia tetraptera is very closely related to Halenia , as they show identical embryology.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 155 , 383–400.     

西南獐牙菜的胚胎学及其系统学意义

报道了西南獐牙菜的大、小孢子发生及雌,雄配子体发育过程,并以此讨论了獐牙菜属宽丝组和多枝组的分类等级和系统演化关系,西南獐牙菜花药四室;药壁发育为双子叶型;绒毡层二型起源,腺质型;中层细胞2层;药室内壁纤维状加厚,药壁表皮宿存,小孢子母细胞减数分裂为同时型;小孢子四分体的排列为四面体形,成熟花粉为3-细胞型,孔子房为3心皮,1室,胚珠12列,故为超侧膜胎座,薄珠心,单珠被,倒弯生胚珠,大孢子母细胞减数分裂形成4个大孢子呈直线形排列,合点端的大孢子具功能,胚囊发育为蓼型,两极核在受精前融合为次生核,3个反足细胞次生增殖为5-8个,宿存,比较宽丝组和多枝组的胚胎党性一状表明宽丝组从多枝组中分出是合理的,在系统位置上宽丝组较多枝组进化。     

獐牙菜属5种植物的核型研究

首次报道了中国５种獐芽菜属植物的染色体数目和核型。它们染色体中期核和相对长度组分分别是：四数獐牙菜为２ｎ＝１４＝４ｍ＋８ｓｍ＋２ｓｔ＝２Ｌ＋６Ｍ２＋４Ｍ１＋２Ｓ;华北獐牙菜２ｎ＝２８＝１２ｍ＋１４ｓｍ＋２ｓｔ＝６Ｌ＋８Ｍ１＋＋６Ｓ;二叶獐牙菜为２ｎ＝２８＝１４ｍ＋４ｓｍ＋１０ｓｔ＝２Ｌ＋１４Ｍ２＋１０１＋ｓＳ;抱茎獐 菜为２ｎ＝６ｍ＋１２ｓｍ＋２ｓｔ＝８Ｍ２＋１２Ｍ;浙江獐牙菜为２ｎ＝２０＝８ｍ＋     

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。     

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.     

Embryology of Swertia tetraptera Maxim. (Gentianaceae) and its systematic implication

The embryology of Swertia tetraptera is reported for the first time. The anther is tetrasporangiate and its wall, of which the development conforms to the dicotyledonous type, is composed of epidermis, endothecium, middle layer and tapetum at the mature stage. The tapetum has dual origin and is similar to the glandular type. Cytokinesis is of the simultaneous type and microspore tetrads are mostly tetrahedral. Pollen grains are mostly 3-celled when shed. The ovary is bicarpellate and unilocular with four series of ovules. The fused margins of two carpels intensively protrude into the locule. The ovule is unitegmic, tenuinucellate and orthotropous. The development of embryo sac is of the Polygonum type. The development of endosperm conforms to the nuclear type and the embryogeny corresponds with the Physalis I variation of Solanad type. Three antipodal cells persist at the mature embryo sac. The embryo is at the globular stage when seeds are released from the capsule. Compared with the other studied species in Swertia, S. tetraptera shows differences from them in three distinctive embryological traits: enlargement and protrusion of the fused margins of two carpels into the ovary locule; orthotropous ovules and a developed hypostase. These three characters were previously only found in the genus Halenia of the tribe Gentianinae in the Gentianaceae,and thus suggest that S. tetraptera should be related to Halenia. The results partly correspond with the molecular evidence (Yuan & Kupfer, 1995). The monophyletic Halenia clade in Yuan & Kupfer's molecular cladogram includes Halenia, S. tetraptera and Frasera (section Frasera in Swertia s. l.); nevertheless, up to now, no embryological study on Frasera has been reported. The embryological results imply that the taxonomic status of S. tetraptera needs to be revaluated and the genus Swertia s. l. may not be a mono-phyletic group. Decontaminated thianthrene disproportion. 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Nectary tracks as pollinator manipulators: The pollination ecology of Swertia bimaculata (Gentianaceae)

Floral nectaries are closely associated with biotic pollination, and the nectar produced by corolla nectaries is generally enclosed in floral structures. Although some Swertia spp. (Gentianaceae), including S. bimaculata, evolved a peculiar form of corolla nectaries (known as “gland patches”) arranged in a conspicuous ring on the rotate corolla and that completely expose their nectar, little is known about the pollination of these plants. Two hypotheses were made concerning the possible effects of gland patches: visual attraction and visitor manipulation. The floral traits, mating system, and insect pollination of S. bimaculata were examined, and the pollination effects of gland patches were evaluated. A comparative study was made using Swertia kouitchensis, a species with fimbriate nectaries. Swertia bimaculata flowers were protandrous, with obvious stamen movement leading to herkogamy in the female phase and to a significant reduction in nectary–anther distance. The species is strongly entomophilous and facultatively xenogamous. The daily reward provided per flower decreased significantly after the male phase. The most effective pollinators were large dipterans, and the visiting proportion of Diptera was significantly higher in S. bimaculata than in S. kouitchensis. Most visitors performed “circling behavior” in S. bimaculata flowers. Removing or blocking the nectaries caused no reduction in visiting frequency but a significant reduction in visit duration, interrupting the circling behavior. The circling behavior was encouraged by nectar abundance and promoted pollen dispersal. Visitor species with small body size had little chance to contact the anthers or stigma, revealing a filtration effect exerted by the floral design. These results rejected the “visual attraction” hypothesis and supported the “visitor manipulation” hypothesis. The nectary whorl within a flower acted like a ring‐shaped track that urged nectar foragers to circle on the corolla, making pollination in S. bimaculata flowers more orderly and selective than that in classically generalist flowers.     

Lack of altitudinal trends in phytochemical constituents of Swertia franchetiana (Gentianaceae)

Diverse biological characters commonly vary with altitude in species that have a wide altitudinal distribution, partly at least as a result of adaptation to differences in aridity, but whether such variation exists for phytochemical constituents remains unknown. Therefore, levels of seven important phytochemical constituents of Swertia franchetiana (swertiamarin, oleanolic acid, swertisin, mangiferin, 1,5,8-trihydroxy-3-methoxyxanthone, 1,8-dihydroxy-3,7-dimethoxyxanthone and 1,8-dihydroxy-3,5-dimethoxyxanthone) were studied and statistically compared, using materials collected from sites ranging from 2200 to 3960 m in altitude. Swertiamarin was the most abundant in all samples, then mangiferin, oleanolic acid and the other three xanthones. Throughout the distributional range of this species, no altitudinal trend was detected for other constituents except 1,8-dihydroxy-3,7-dimethoxyxanthone, which showed a negative correlation with altitude. However, the concentration of 1,8-dihydroxy-3,7-dimethoxyxanthone and mangiferin showed a significantly latitudinal and longitudinal correlation.     

青藏高原特有植物祁连獐牙菜（龙胆科）的花综合征与虫媒传粉

雌雄异熟和异型花在一些被子植物中的共同存在被认为是一种有效避免两性花植物自交并促进异交的机制。本文报道青藏高原特有植物祁连獐牙菜Swertia przewalskii Pissjauk.的花同时具有雌雄异熟和异型花这两种特征。在此基础上,我们详细观察了该植物访花昆虫的访花行为,主要探讨这种促进异花传粉的机制是否能完全避免株内异花传粉的自交。繁育系统试验证明祁连獐牙菜单花套袋不结实,说明雌雄异熟和异型花的花特征完全避免了单花的自花传粉,自然结实必须依赖昆虫传粉才能完成。然而,该植物却是高度自交亲和的,因为人     

Floral character states of the Northeast and Tropical East African Swertia species (Gentianaceae)

The floral gross morphological and anatomical character states of 23 African Swertia species are described. Colleters are frequent in Swertia, especially at the lower rim of calyx lobes and are usually placed in a single row except in S. crassiuscula (they are not "typical" colleters). The aestivation of corolla lobes is found to be uniform except in S. engleri var. engieri. Each corolla lobe is beset with either one or two foveae. Foveae are naked or either partly or wholly fimbriated along their margins. No vascular tissue was observed entering the foveae. The taxonomic importance of the the floral character states is discussed.     

Cannabidiol: an overview of some chemical and pharmacological aspects. Part I: chemical aspects

Over the last few years considerable attention has focused on cannabidiol (CBD), a major non-psychotropic constituent of Cannabis. In Part I of this review we present a condensed survey of the chemistry of CBD; in Part II, to be published later, we shall discuss the anti-convulsive, anti-anxiety, anti-psychotic, anti-nausea and anti-rheumatoid arthritic properties of CBD. CBD does not bind to the known cannabinoid receptors and its mechanism of action is yet unknown. In Part II we shall also present evidence that it is conceivable that, in part at least, its effects are due to its recently discovered inhibition of anandamide uptake and hydrolysis and to its anti-oxidative effect.     

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.     

３种獐牙菜属植物花蜜腺的发育解剖学研究

獐牙菜属的红直獐牙菜、抱茎獐牙菜和四数獐牙菜３种植物花蜜腺都属花被蜜腺,其结构相似,均由分泌表皮和产蜜组织组成,为结构蜜腺,是花冠其部薄壁组织恢复分和能力形成的,分泌表皮无气孔器,原蜜汁由蜜腺周围的维管束提供,经产蜜组织加工后,由分泌表皮外薄的角质层泌出。四数獐牙菜花蜜腺裸露,凸起,而另２化蜜腺凹限为囊状、;红直獐牙菜为脱落蜜腺、而抱茎獐牙菜和四数獐牙菜为宿存蜜腺,其花蜜腺的性状基本印证了３种獐牙菜属植物的系统位置。     

Comparative and phylogenetic analyses of Swertia L. (Gentianaceae) medicinal plants (from Qinghai,China) based on complete chloroplast genomes

Swertia L. is a large genus in Swertiinae (Gentianaceae). In China, many Swertia species are used as traditional Tibetan medicines, known as “Zangyinchen” or “Dida”. However, the phylogenetic relationships among Swertia medicinal plants and their wild relatives have remained unclear. In this study, we sequenced and assembled 16 complete chloroplast (cp) genomes of 10 Swertia species, mainly distributed in Qinghai Province, China. The results showed that these species have typical structures and characteristics of plant cp genomes. The sizes of Swertia cp genomes are ranging from 149,488 bp to 154,097 bp. Most Swertia cp genomes presented 134 genes, including 85 protein coding genes, eight rRNA genes, 37 tRNA genes, and four pseudogenes. Furthermore, the GC contents and boundaries of cp genomes are similar among Swertia species. The phylogenetic analyses indicated that Swertia is a complex polyphyletic group. In addition, positive selection was found in psaI and petL genes, indicating the possible adaptation of Qinghai Swertia species to the light environment of the Qinghai-Tibet plateau. These new cp genome data could be further investigated to develop DNA barcodes for Swertia medicinal plants and for additional systematic studies of Swertia and Swertiinae species.     

Mesua sp.: chemical aspects and pharmacological relevance of prenylated polyphenols

Phytochemistry Reviews - The genus Mesua L. (Calophyllaceae) comprises approximately 50 species that grow in the restrictive area of South East Asia. Investigations of Mesua species reported the...     

Genetic diversity and population structure of Swertia tetraptera (Gentianaceae), an endemic species of Qinghai-Tibetan Plateau

Swertia tetraptera Maxim is an annual alpine herb endemic to the Qinghai-Tibetan Plateau (QTP). Its populations are locally scattered as isolated patches throughout this region. Genetic variation within and among thirty-four populations of this species was assessed using ISSR fingerprinting with 10 primers. High levels of genetic diversity exist within species (P = 98.9%, I = 0.3475; He = 0.2227), while the within-population diversity is low (P = 32.7%, I = 0.177; He = 0.12). High levels of genetic differentiation were detected among populations based on various statistics, including Nei’s genetic diversity analysis (G_ST = 0.4608), Bayesian analysis (θ^B = 0.476) and AMOVA (F_ST = 0.57). That is, populations shared low levels of genetic identity (I = 0.2622–0.0966). This genetic structure was probably due to severe genetic drift, breeding system and limited gene flow. The observed genetic structure of the populations implies that different populations across the distribution range of the species should be sampled to maintain high genetic diversity when a conservation strategy is implemented.     

Isozyme variability of the wetland specialist Swertia perennis (Gentianaceae) in relation to habitat size, isolation, and plant fitness

We examined the effects of size and spatial isolation of fens on the isozyme variability of 17 populations of Swertia perennis. This long-lived perennial is a locally abundant fen specialist in Switzerland, where wetlands have been strongly fragmented. Isozyme variability was comparable to other outcrossing plants (A = 1.53, AP(p) = 2.01, P(p) = 42.5, H(o) = 0.113, H(e) = 0.139). F statistics indicated both inbreeding within and differentiation between populations (F(IS) = 0.076, F(IT) = 0.194, F(ST) = 0.128), with moderate gene flow between populations (N(e)m = 1.703). Populations in small, isolated fens had reduced genetic variability and the highest within-population inbreeding coefficients (F(IS)). Isozyme variability was significantly associated with vegetative fitness traits (MANOVA), and the magnitude of leaf herbivory decreased as the percentage of polymorphic loci increased. These data suggest that the reduced genetic variability of S. perennis in small, isolated populations may reduce plant fitness, thereby increasing susceptibility to herbivore damage. Our study also shows that habitat fragmentation can reduce the genetic variability of populations of fairly common habitat specialists, which so far have attracted less conservation attention than rare species.     

条纹龙胆的胚胎学研究

首次报道了条纹龙胆（Ｇｅｎｔｉａｎａ ｔｓｒｉａｔａ Ｍａｘｉｍ．）的胚胎学特征,研究结果用以讨论龙胆属狭蕊组（Ｇｅｎｔｉａｎａ Ｓｅｃｔ．Ｓｔｅｎｏｇｙｎｅ）的系统演化关系。主要研究结果如下：花药四室;药壁发育为双子叶型;绒毡层细胞仅来源于初生壁细胞,故绒毡层起源属单型起源,细胞单核,原位退化,属腺质型绒毡层,药隔处的绒毡层细胞经多次平周分裂形成２层至多层的绒毡层细胞,其余部位的绒毡层细胞仍为１