龙胆科獐牙菜亚族一些关键系统学问题的研究
其他题名Phylogeny of some group in subtribe Swertiinae of Gentianaceae-Gentianeae
薛春迎
学位类型博士
2003
学位授予单位中国科学院昆明植物研究所(昆明)
学位授予地点中国科学院昆明植物研究所(昆明)
关键词獐牙菜亚族 大钟花属 黄秦充属 獐牙菜属 胚胎学 花部解剖 花器官发生 发育 系统学
摘要獐牙菜亚族隶属于龙胆科龙胆族,约15个属,广泛分布于世界各地的高山,温带和高寒地带。獐牙莱亚族目前存在的主要系统学问题是:(1)獐牙菜亚族是否是一自然类群;(2)大钟花属是否为漳牙菜亚族的成员;(3)黄秦先属系统位置不确定;(4)獐牙菜亚族中一些属的检索用形态性状其系统意义不明确;(5)獐牙菜属的范畴。针对以上问题,本报告首先补充研究了大钟花属和黄秦充属的胚胎学和花部解剖学;同时首次报道了5属8种植物的花器官发生、发育;30余种植物的ITS和叩116序列;根据所获得的系统学资料,对津牙菜亚族上述系统学问题进行了讨论。主要实验结果和结论总结如下。1.胚胎学大钟花和黄秦式共同拥有药壁发育为双子叶型;绒毡层为异型起源,腺质型;胞质分裂为同时型,小抱子四分体排列主要为四面体型;薄珠心,单珠被;寥型胚囊;核型胚乳等胚胎学特征,但它们在药壁的层数;中层是否纤维状加厚,是否宿存;胎座类型,胚珠类型等方面存在差异。大钟花的胚胎学性状支持它为独立的属且为猪牙菜亚族的成员;黄秦式的胚胎学研究表明黄秦力属与猪牙菜属的四数獐牙菜关系较近。结合已有的胚胎学资料,发现与獐牙菜亚族相关的胚胎学性状为胚胎发育为茄型酸浆I变型,而与龙胆亚族相关的胚胎发育类型为茄型酸浆II变型。胚胎学性状表明猪牙菜亚族为一自然类群。2.花部解剖学大钟花属与黄秦式属的雌花部分花等维管束与花冠维管束来源于同一维管束这,而雄蕊维管束来源于雄蕊迹,每心皮具1条背维管束2条腹维管束,花被维管束为融合型;黄秦式属的雄花每个花窖、花瓣和雄蕊的维管束均来源于单个维管束迹,每心皮具1条背维管束2条腹维管束,属于基本型。从花部解剖结构看出,大钟花属与津牙菜亚族的关系较近;黄秦充属较漳牙菜属进化。3.花器官发生、发育利用扫描电镜观察了龙胆亚族和獐牙菜亚族5属8种植物的花器官发生、发育。它们共同具有(1)符合Hofmeister规律,即新器官的发生首先出现在花顶己经存在的器官之间;(2)花冠筒发育为"后合瓣"的发育模式,即花冠原基为分别起源,花冠筒的形成是后期分生组织活动的结果。(3)花警原基发育顺序为螺旋状相继发生,花冠原基,雄器原基为分别起源,发生、发育顺序为近同时型。但它们在花等原基的发生上存在差异:龙胆亚族龙胆属花等原基为一环状的分生组织,花警原基为共同起源,而其它獐牙菜亚族的类群花等原基为点状分生组织,花导原基为分别起源。獐牙莱亚族不同类群的花蜜腺为非同源性状。根据花器官早期发生、发育的相似性,推测湾牙菜亚族各种花冠类型的形成以及肋柱花属的柱头下延可能是由于生长比率的变化所至。花器官发生、发育研究结果支持獐牙菜亚族是一自然类群,扁蕾属为獐牙莱亚族的基部类群。4.分子系统学利用本研究得到的獐牙菜亚族8属34种植物的rp/16,5属18种的ITS1和ITS2序列,并结合GeneBank中得到的序列,对两套数据进行了单独分析和联合分析。结果表明,獐牙菜亚族作为一个单系群得到了100%的靴带支持率。大钟花属为獐牙菜亚族的成员。黄秦充属系统位置仍较为模糊。獐牙菜属为一高度的复系群,分别与喉毛花属,假龙胆属,肋柱花属和花锚属等网结在一起;从基因树中看出狭义漳牙菜属的范畴至少应包括何廷农系统中的招皱组和獐牙菜组的部分种。同时还讨论了一些类群的系统学。
其他摘要Subtribe Swertiinae of the Gentianaceae-Gentianeae is an almost cosmopolitan group of 15 genera growing mostly in alpine, temperate, or arctic habitats. Controversial systematic questions still remained of Swertiinae mainly include: (Y)Megacodon delimited in Swertiinae or not, subtribe Swertiinae is monophyletic or not, and the position of Vertrallia was ambiguous; (2)some key morphological characters systematic significance still didn't understand; (3)the concept and circumscription of Swertia. The disagreements of previous classification or systematic treatments of some taxa in the Swertiinae were mainly caused by incomplete data or by different understanding of the character evolutions. Therefore, the main aim of the present report was to find and collect new systematic data. Reported and recorded further in the present work were embryology and floral anatomy of 2 species representing 2 genera, floral development of 8 species representing 5 genera to 2 subtribe under SEM and ITS and rp/16 sequence of 34 species belonging to 2 subtribe. The characteristics obtained were compared among taxa and their systematic significance was valued. Based on our results, the systematic problems in Swertiinae were discussed. The main results and the essential points were summarized as follows. 1. Embryology Investigated in this chapter was the embryology of Megacodon stylophorus and Veratrilla baillonii representing 2 genera. The common embryological features of 2 species include: the dicotyledonous type of the anther wall; glandular tapetum which dual origin; simultaneous cytokinesis at meiosis of the microsporocytes; microspore tetrads nearly always tetrahedral; unitegmic and tenuinucellar; Polygonum type of megagametophytes; nuclear endosperm. However, they show variation in the following features: middle layers, ovary type and placentae type. The embryological data support Megacodon is independent genus and belonged to Swertiinae, Veratrilla is closed relationship with Swertia tetrapetra. The embryogeny corresponds to the Solanad Type physalis I variation of Swertiinae, to the Solanad Type physalis II variation of Gentianinae. Based on embryological data, Swertiinae is a monophyletic. 2. Floral anatomy The floral anatomy of Megacodon and female floral of Veratrilla are peculiar in the laterals of adjacent sepals arising conjointly with the petal traces. In male floral of Veralrilla, the petals and sepals receive one trace each directly from the stele. These group share similar characters: the stamens receive one trace each directly from the stele and each carpel receives 1 dorsal bundle and 2 ventral bundles. According to the floral anatomical characters, it is better treat Megacodon and Veralrilla as two distinct genera. Megacodon belongs to the "Gentianella" evolutionary line and Veratrilla is more advanced than Swertia 3. Floral development Floral organogenesis and development of 8 species in 5 genera of the tribe Gentian was studied by using scanning electron microscopy (SEM). They have common characters of floral organ initiation and development: (1) initiation of the floral organs in 8 species follows Hofmeister's rule; (2) the mode of corolla tube development belongs to the "last sympetaly" type; (3) the sepal primordia have spiral development, the petal and stamen primordia have simultaneous initiation. However, the sepal primordia initiating show different modes in these species. The sepals of species in Gentian are initiated on a common ring primodium, while the sepals of other species are initiated on five or four separated primodium. The characters of petal nectaries are homoplasy. The main flower types found in Swertiinae can be transformed into each other by simple development variation in proportion. The results suggest that Swertiinae is monophyletic and Gentianopsis is basal taxon of Swertiinae 4. Molecular systematics Phylogeny of some taxa of subtribe Swertiina was analysed based on rpl\6 and ITS sequence variation. The analyses of two separate data and combined data sets were performed using the parsimony method. Two species from Gentiana were used as outgroups. All three analyses strongly supported the monophyly of subtribe Swertiinae. Megacodon belonged to Swertiinae. The results suggested that Swertia is polyphyletic and should be limited to species of part of sect. Rugosa and sect. Swertia of Ho . The systematic position, relationships and the problems existing in some taxa were summarize and discussed.
页数113
语种中文
文献类型学位论文
条目标识符http://ir.kib.ac.cn/handle/151853/694
专题昆明植物所硕博研究生毕业学位论文
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薛春迎. 龙胆科獐牙菜亚族一些关键系统学问题的研究[D]. 中国科学院昆明植物研究所(昆明). 中国科学院昆明植物研究所(昆明),2003.
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