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题名: 广义鹿药属的系统学与民族植物学研究
作者: 孟盈
学位类别: 博士
答辩日期: 2006-12-26
授予单位: 中国科学院昆明植物研究所
授予地点: 昆明植物研究所
导师: 孙航
关键词: 鹿药属 ; 舞鹤草属 ; 叶表皮 ; 细胞地理学 ; 分子系统发育 ; 地理分布格局 ; 东亚 ; 种群生态 ; 民族植物学
学位专业: 植物学
中文摘要: 本论文主要通过形态学、细胞学和分子生物学手段对广义鹿药属(Maianthemum s.l.)进行系统发育与生物地理进化方面的研究以及对滇西北地区该属植物利用与保护方面进行民族植物学与种群生态学的综合研究,主要结论如下: 1. 系统学研究方面 鹿药属及其邻近属30个种的叶表皮进行了光镜和电镜下的形态学研究。结果表明叶表皮微形态特征在鹿药属内没有显著差别,不适合作为种间划分的主要依据,但黄精族内不同属间的叶表皮有一定的差异。传统鹿药属和舞鹤草属的叶表皮细胞形状、垂周壁式样、气孔的类型与分布、气孔外拱盖内缘等特征均无显著差别,从形态学方面再一次证明了二者具有很近的亲缘关系。但扫描电镜下垂周壁式样和角质膜的形态有一定的分类学意义,特别在鉴别一些形态变异比较大的种如M. purpureum时有一定的参考价值。另外,扫描电镜下黄精族内各属间的角质膜及气孔的形状也有明显差别。黄精属的角质膜多为鳞片状,气孔保卫细胞狭长形;而竹根七属的角质膜呈不明显的波状隆起,气孔保卫细胞卵形,气孔器明显的凸出叶面。 染色体数目和核型的研究对揭示本属植物的系统关系和进化有一定的参考价值。本文对10个种的本属植物进行了细胞学研究。其中,M. lichiangense、M. forrestii、M. nanchuanense染色体数目和核型为首次报道,M. atropurpureum首次发现多倍体现象(2n=72)。目前该属有染色体数目报道的约19种,大多数的种染色体数目为2n=36,支持本属的染色体基数为x=18。而通常认为染色体基数大于11为高基数,鹿药属的染色体高基数可能是在它们的进化历史中通过多倍化后而形成的古多倍体,即古多倍化。鹿药属内目前仅发现四个种(M. bifolium, M. atropurpureum, M. fuscum, M. racemosum)有不同的倍性变化,多倍体在鹿药属中并不普遍。鹿药属植物的染色体核型变异程度非常大,一般可分为三种类型:一型,两型和三型。如分布于我国东北的M. dahuricum,在长度上逐渐减小,没有明显的梯度变化,称之为一型染色体,二型染色体见于M. tatsienense。而分布于我国西南的绝大多数种的染色体长度都表现为二个梯度三组不同类型的长度,我们称之为三型染色体。分子系统发育的研究结果也表明具有一型染色体的种是相对原始的,三型的种相对比较进化。这种核型变化也表现在染色体核型不对称性方面。最为对称(2B)的种M. dahuricum分布于东北,较为原始(2C)的种M. japonicum和M. nanchuanense分布于东北至华中地区,而相对进化的(3B)种都分布在我国西南部横断山区,因此我们推测鹿药属的祖先类群应该分布于东北,然后由我国北方地区迁移到华中,再到西南横断山区。青藏高原抬升以后形成的特殊的地形地貌致使该属进一步分化,成为鹿药属在东亚的现代多样化分布中心。鹿药属的核型发展方向可视为2B→2C→3B→3C,由对称到不对称,对称性核型代表了相对较原始的类型。总的说来,鹿药属染色体核型(不对称性核型及长度上的变异特征)揭示了一定的地理及系统演化关系,分子数据的研究结果也支持同样的观点。 通过对广义鹿药属各分布中心代表种的核基因序列ITS和六个叶绿体片段(psbA-trnH, rps16, trnL-F, rpl16, ndhF, and rbcL)的综合分析,结果表明广义鹿药属是一个单系类群,得到了叶绿体基因和核基因数据的很高的支持率(BP=100,PP=100)。在以前的形态学与分子生物学研究中,本类群的单系地位都得到了一致认可,我们的结果在更全面的取样条件下也无容置疑地支持把这两个传统的属合并为一个属。分子证据也揭示本属过去认为重要的分类学性状并不能真正反映本类群的系统进化关系。根状茎、花冠裂片的离合程度、花颜色、花序等方面都没有发现明显的规律符合我们的分子结论。另一方面,我们也发现本属内种间分子系统发育关系与其地理分布格局比较一致,可以看出一定的规律,特别是分布于东亚的类群,体现在同一区域的物种很好的聚在一起,如我国西南地区本属的种绝大多数聚在一起,且各种分析方法的支持率均很高(分支A)。分布于日本的种与我国东北至华中的种较好的聚在一起(分支C)。M. tatsienense 和M. stenolobum分布也处于横断山区一直到华中地区,形态也比较特殊,分子证据也支持单独成一支(分支B),但系统位置不定,可能是联系前面这两分支的中间类群的后裔。而传统舞鹤草属广布于北温带北部的广大地区,也很好的独立成一支(分支D),成为本属较为进化或特化的一个分支。总的说来,本属分布于北方的尽管种类不多,但系统发育关系比较复杂,没有单独成支,不像西南一支种类更多却完全聚成很好的一支。这可能正好说明了本属可能起源于在北温带靠北的区域,以及它们进化过程的古老性和复杂性。另外也发现一些形态学特征和与我们的分析结果大体相吻合。如舞鹤草属(Maianthemum s. str.)的单系地位得到非常强烈的分子证据支持。另外我们还发现花丝与花药在长度上的比例以及柱头裂开的程度与我们的分子数据和地理分布格局有一定的联系:分支A中的所有种,即分布于我国西南部的种柱头均深裂,花丝与花药近等长;而分支C中的种,所有种柱头均浅裂或不裂,花丝长,花药小,花丝长与花药长之比大于2。 鹿药属在东亚的这种地理分布格局与系统发育关系的一致性得到了细胞学、分子系统学以及部分形态学证据的支持。这种地理分布格局与系统发育关系也体现在东亚的地质演化历史方面。一般认为凡分布于旧大陆温带、北温带的属,大多起源于劳亚古陆(古北大陆)。我们推断旧世界的鹿药植物可能起源于北方森林地区,属于北极—第三纪类型,后来从北向南迁移扩散,到达横断山区后,由于此地区复杂多变的地形和气候条件为本属植物的快速分化与发展提供了得天独厚的条件,从而成为鹿药属在东亚的现代多样化分布中心。 2. 鹿药属植物的利用与保护方面 鹿药属植物的研究价值与重要性也体现在它的开发与利用方面,本属植物具有很好的食用和药用价值。在滇西北地区俗称“竹叶菜”的鹿药属植物有M. atropurpureum, M. purpureum, 在维西还发现市场上有M. tatsienense、M. forrestii和M. henryi出售。由于近年来迪庆州旅游业的兴起,加剧了对竹叶菜的需求,竹叶菜价格一直居高不下,整个市场处于供不应求的状态;而交通相对不发达的怒江州人口稀少,竹叶菜的市场并不是很大,加上气候等得天独厚的条件,竹叶菜资源相当丰富,价格也相对较低,市场长期处于供过于求的形势。一方面在中甸等地极大的经济利益吸引更多的人去采集竹叶菜,进一步加剧了竹叶菜资源的枯竭;另一方面在竹叶菜资源丰富的地区却由于市场需求量小而造成了资源的浪费。近年来由于生境的破坏和过度采集而造成其野生资源量急剧减少。 为了评估滇西北地区生境破坏及采集利用对鹿药属主要利用种—高大鹿药野生资源的影响,我们调查了贡山和中甸两个点各5个样方的种群特征,结果表明在中甸由于过度采集和严重的生境破坏,无论是在种群密度、植株平均高度、开花植株数目、生物量等方面都远远低于贡山。另外在中甸,高大鹿药的分布非常不均匀,有的地方只有幼苗,而有的地方仍有少数长势良好的植株。总的来说中甸高大鹿药野生资源正处于日益减少的趋势,而贡山的野生资源则非常丰富,有进一步开发利用的良好前景。
英文摘要: To understand the systematics and evolution of the genus Maianthemum s. l. (include Maianthemum s. str. and Smilacina), an integrative study was undertaken by means of morphological, cytological and molecular phylogenetic approaches. Moreover, both ethnobotanical survey of Maianthemum and population ecology of Maianthemum atropurpureum were conducted in northwestern Yunnan Province, SW China. The primary results are summarized as follows: 1. Systematics Leaf epidermis of 30 species from Maianthemum and related genera were investigated with both light microcopy (LM) and scanning electron microscopy (SEM) methods. No distinct epidermal variation was observed within Maianthemum species, whereas differences were found among genera of the tribe Polygonatae. The shapes of adaxial epidermal cells of Maianthemum are usually polygonal or irregular, with the anticlinal walls straight, arched, while those of daxial epidermal cells are usually irregular with the anticlinal walls simulate, sinuous to sinuate. The stomatal apparatus in all species were observed only on abaxial epidermis except for M. trifolium, without accessory cells. Under SEM, the inner margin of the outer stomatal rim is nearly smooth, sinulate or sinuous, and the cuticular membrane of the leaf epidermis is striated, sometimes striated to wrinkled, occasionally granular or scalelike. The stomatal apparatus are sunken, protuberant or equal to epidermis cells. Chromosomal numbers and karyotypes of 10 species of Maianthemum from China were reported in this study. Three species were investigated cytologically for the first time. All species have a chromosome set of 2n = 36, except four species having polyploids in some populations: M. bifolium, M. atropurpureum, M. fuscum, and M. racemosum. The basic chromosome number of the genus was confirmed to be x = 18 and the genus is suggested to have been paleo-polyploidized. The results also showed striking variation of karyotype in different species, as in the numbers of metacentric, submetacentric, and acrocentric chromosomes as well as in the number of satellites. Three modes of unimodal, bimodal, and trimodal were recognized based on the number of clear gaps in chromosome length variation. The unimodal variation (without gap) was found in M. dahuricum from NE China, the bimodal variation (with one gap) in M. tatsienense, and almost the species from SW China have the trimodal variation (with two gaps). Our molecular study also supported similar results. In addition, the chromosome variation is also reflected from the karyotypic asymmetry. Maianthemum dahuricum from NE China has a 2B-type, M. japonicum and M. nanchuanense from NE to C China have a 2C-type, while species from SW China almost show 3B and 3C-types. Results of this cytological study provide a better understanding of phylogenetic relationships as well as chromosome evolution in the genus. Forty-eight samples representing twenty-eight species of Maianthemum together with eleven from close taxa as outgroup were sequenced including both chloroplast and nuclear markers (trnL-F, rps16, rpl16, psbA_trnH, rbcL, ndhF, and ITS). Phylogenetic analysis of the molecular data robustly supported the monophyly of Maianthemum including both Maianthemum sensu stricto and Smilacina, and provided a moderate phylogeny for several clades within the genus. Species from the high mountains of southwestern China to the Himalayas formed a well supported clade, which characterized by large and colorful flowers, hairy stems and leaves. Maianthemum tatsienense and M. stenolobum form a clade which has a distribution from southwestern to central China. Species with linear petals form a well supported clade, which distributed from Japan, northeastern to central China. The North American M. racemosum is sister to a clade including the tropic American species with a relatively low support. Many important morphological characters were not consistent with the molecular data, e.g. rhizome, representing a sympodial aggregation consisting of several individual rhizome units. But we also found some characters, such as the ratio of filament and anther in length, the degree of stigma dehiscent, were related to DNA evidences. Maianthemum showed recent or fast radiation in the high mountains of southwestern China, and complex intercontinental relationships in the Northern Hemisphere. 2. Ethnobotany In Gongshan and Zhongdian county of Yunnan, M. atropurpureum was the most common species collected traditionally as so-called “Zhu-ye-cai” by local residents. In order to evaluate on the impacts of harvesting, population ecology of M. atropurpureum, including population density, height and basic diameter of ramets, biomass, ramets flowering/m2, were investigated in ten patches of Gongshan and Zhongdian. The results reveal that populations of M. atropurpureum in Gongshan are less affected by local collection and gathering than those in Zhongdian. In Northwest Yunnan, including Lujiang Prefecture (Liuku, Fugong, Gongshan) and Deqing Prefecture (Weixi, Zhongdian and Deqin), three kinds of key informants, that is, collectors, vendors and restaurateurs, were interviewed. Vernacular name(s), price, harvest, frequency, and trade mechanisms and amount, were recorded. The availability, seasonal and yearly changes of Maianthemum resources and marketing were documented through semi-structured interviews. In Fugong, a participant observation was undertaken for understanding the collection behavior and process. M. atropurpureum and M. purpureum are generally harvested and sold in Northwest Yunnan. In Weixi, however, M. tatsienense, M. forrestii and M. henryi were also found in the marketplace. In local communities, M. atropurpureum was more popular and better sold than M. purpureum. Seasonality and resource availability were considered as the main reasons resulting in price instability, supply-and-demand balance, and tourism as the other factors. The price of Zhu-ye-cai fluctuated from 1-16 yuan RMB per kilogram. In Zhongdian, the resource of Zhu-ye-cai was declining at an unexpected rate, being speeded up by degradation and destruction of fir old-growth. To some extent, local tourism stimulates and motivates the existing over-harvesting and consumption. In comparison, there was a great exploitive potential, with abundant resource in Fugong and Gongshan.
语种: 中文
内容类型: 学位论文
URI标识: http://ir.kib.ac.cn/handle/151853/260
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Recommended Citation:
广义鹿药属的系统学与民族植物学研究.孟盈[d].中国科学院昆明植物研究所,2006.20-25
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