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题名: 濒危药用植物云南黄连的保护生物学研究
作者: 黄骥
学位类别: 博士
答辩日期: 2004
授予单位: 中国科学院昆明植物研究所
授予地点: 中国科学院昆明植物研究所
导师: 裴盛基
关键词: 云南黄连 ; 黄连属 ; 保护生物学 ; 濒危原因 ; 生物学特性 ; 解剖生态学 ; 民族植物学 ; 繁殖生物学 ; 细胞学 ; 保护遗传学
学位专业: 植物学
中文摘要: 云南黄连是滇西北特有的一种药用植物,己被列为国家二级保护植物。本论文以高黎贡山东、西坡10个云南黄连种群和重庆黄水的1个中国黄连种群及1个日本黄连种群为材料,以保护生物学为核心,对云南黄连开展了生物学特性、形态学、民族植物学、解剖生态学、细胞学、繁她生物学和遗传多样性等方面的研究工作。以期对植物物种的生物学研究思路进行探讨,摸索濒危植物保护生物学的研究方法,同时为云南黄连的保护和可持续利用提供有价值的基础性资料。1.云南黄连的左目勿学特性云南黄连兼具有性生娘和无性繁殖。,在自然条件下,种群数量增长以发达的地下营养繁殖体(称为"觅养枝")行无性繁狱为主:高黎贡山西坡种群,特别是野生种群的一些个体,能以有性生娘方式形成实生苗,另外一些个体和东坡和}.群的几乎所有个体都以无{生系小株进行首夕仁繁娘。该物利1纵然开花、结实正常,但种子有一氏达半年的胚后熟过程,加上较低的种子传播效率,实生苗在种群中只占很少比例。成年植株的花芽在七月开始分化,次年1一2月开花,5月种子成熟,完成生活史,个体生命周期长达10余年。云南黄连种群较小,分布在狭小、呈片断化的特殊生境中。该物种生态幅极其狄窄,其本限于在温凉勃翻河的常绿阔「!一卜林一下陡坡生长。土壤和群落小气候是限制其分布的主要因:,公南黄连为中国一喜马拉雅特有种,基本分布于受印度洋季风影响、属于印度洋水系的儿条大河,如布拉马普特拉河、伊洛瓦底江、萨尔温江的分水岭地区海拔2200米以点温凉湿润的山地阔叶林下,野生种群已极为罕见,种质资源的保护刻不容缓。2.形态变异式样研究对高黎贡山东、西坡10个种群的20个性状的变异式样进行了研究。结果表明,西坡种群2。个性状中的大多数变异系数均大,东坡种群,在同一种群中,根茎数、根茎长、觅养枝数目、总长和节数等五个性状的变异系数均超过了39%。野生的西坡种群的平均变异系数最高。而东坡各种群的平均变异系数均小于西坡各种群,并且该区域(东坡)内各种群间的差异也不大。刘一功个种群的形态变异系数进行横向比较后发现:营养性状的变异系数集中分布在20%一8。%之间,平均CV值为42.15。就变异系数来看,总的变异l隔度为:觅养枝>根茎>叶。生殖性状的变异系数集中分布在10%一40%之间,平均Cv值为21.47。总的变异幅度为:花草>雌雄蕊>花曹花瓣。营养性状的平均Cv值(42.15)几乎是生殖性状平均cv值(21,47)的两倍,表明营养性状有较大的变异幅,、容易受到外界环境的影响,而生殖性状相对稳定。3.民族植物学研究运用民族植物学原理,采用野外面上调查、定点社区调查和文献研究相结合的办法,深入调查高黎贡山地区傈僳族和勒墨人对云南黄连的各种传统利用方式及历史,总结他们管理、保护和开发这一名贵药用植物的传统知识和经验,进而研究云南黄连产区的民族民间传统知识与利用、保护、栽培黄连的互动关系,通过民族植物学定点研究,确定人类活动对云南黄连种群、群落和生态系统的影响程度,结果表明:1)130年前,由于云南黄连的经济价值,过度的采挖使其野生种群几近枯竭,为了维持这一宝贵的经济来源,傈僳族发展了一套行之有效、并且具有深刻科学意义的种植、抚育和采收的资源可持续利用管理方法,不仅持续有效地利用了这一资源,并且也保护了其赖以生存的森林生境;2)傈僳族对云南黄连的引种栽培尽管更多是出于经济目的,但有效地保护了这一濒于灭绝的药用植物,他们建立和发展起来的种植、管理模式可以看作是我们现代保护学者所推崇的就地保护,它不仅是传统文化与植物资源直接相互作用的一个范例,也是当地人对生物多样性资源进行管理的一种方式,对野生种群很少的药用植物进行栽培和管理,对于当地生物多样性的保护和经济植物的可持续利用,无疑起到了积极的推动作用;3)在传统信仰文化的背景下,以经济利益为强大动力的对某一物种的利用,客观上起到了在多个层次上对生物多样性发生影响的作用,体现了各少数民族和植物之间的互动关系。传统文化通过对云南黄连的利用及引种栽培,在物种水平上发挥作用;通过对云南黄连适生生境的特定植物群落类型的利用和维护,使系统的结构和组成发生改变,从而在系统水平上发挥作用;通过对自然景观的塑造和文化一自然景观的建立而在景观水平上发挥作用;4)云南黄连混农林生态系统并不是孤立存在的,它是传统生产和保护系统的有机组成部分。通过乡土自然保护体系而发挥着综合的作用,并与现代的自然保护区存在着密切的联系,是以现代科学为基础的正式保护体系十分必要和积极的补充。它是传统文化与特定自然环境相互作用、相互统一和协同进化的结果。作为一种迄今仍然发挥着重要作用,并对群众日常生产和生活产生持续影响的自然保护体系,乡土自然保护体系体现了传统信仰文化对自然保护的积极贡献,是山地民族文化的一个组成部分,也是他们的祖先留给我们的宝贵遗产。4.解剖生态学研究黄连属植物的叶片由表皮、叶肉和叶脉构成,上下表皮细胞从纵切面上看,排列整齐,呈近长方形扁平状,具角质层;上下表皮内的叶肉薄壁细胞排列较整齐,类圆,没有明显分化为栅栏组织和海绵组织;叶脉为大型的薄壁细胞包围着一个维管束而成。这些特征都说明了黄连属植物不耐干旱的生物学特性和对温凉、湿润生境的生态学要求。从解剖特征来看,黄连属植物,特别是云南黄连存根、叶各器官的结构、功能上存在着明显的抗早和其它抗逆性弱点,致使种群适应力、扩展力较弱,从而可能导致种群逐步走向濒危。云南黄连和国产另外三种黄连及日本黄连的上、下表皮细胞的表面观以无规则形、垂周壁波状为主。气孔器类型为无规则型,即围绕保卫细胞的副卫细胞与表皮细胞无区别,这是毛莫科植物气孔器的典型类型。国产的四种黄连与日本黄连在叶表皮性状上存在着一个明显差异:国产的四种黄连上表皮均无气孔,而日本黄连的上表皮发现有稀疏的气孔存在。5.云南黄连的生殖生物学研究云南黄连都是两性花,但有雌雄蕊等长、雌蕊长于雄蕊和雄蕊长于雌蕊三种类型,可以肯定云南黄连也采用雌雄异位的方式来避免完全自交。而在日木黄连中不仅有正常的两性花,还发现有单雌性花或单雄性花的花序。云南黄连同一植株或同一花序上的花逐渐发育成熟,逐渐开放,逐渐结实,始花时间可相差近半个月,一个花序上最多只能成熟三个果序。j:l'.花期的差异与种群所处的海拔高度和坡向相关,海拔越高,开花期越早,海拔越低,开花期越晚;当海拔相近时,西坡种群开花期较东坡种群旱。同一种群内,不同个体的开花时间相差很大,最长可达1个月左右。云南黄连两性花的雌却至蕊异熟,混合传粉。在野外,云南黄连兼具有性生殖和无性生效2种繁欺方式。有性生娘通常在阳光相对·较充足的种群中发生频率较高,一个行有性生狱的个体产生数枚胚尚未发育的"成熟"种子,这些种子在离开毋株散落到其周围IIl的上壤中后,开始胚的发育,;直至半年后,月玉发育成熟,种子才开始萌发。无性生殖在阳光较难获得的荫蔽林下比较占优势。云南黄连的无性生娥是通过其顶芽和腋芽生出的觅养枝来进行,从一个腋芽生出的觅养枝可在离母株根茎30cm左右的范田内构成多分枝无性系。所研究的10个云南黄连种群中,由种子萌发形成的实生苗占全部植株数的5.12%,并且全部出现在西坡种群,东坡种群中没有发现实生苗。云南黄连种群的无性繁殖是其生殖中不可缺少的一环,也是云南黄连对半人工栽培环境和持续生存的一种适应。云南黄连更接近于S对策,即胁迫耐受者,它的主要胁迫因素可能为光照和湿度。6.云南黄连的细胞学研究细胞学研究中,首次报道了三角叶黄连(CoptisdeltoidaC.Ychenget.Hsiao)的核型,染色体数目为2下3x=27,为三倍体,核型公式为2下3x=27=3m+2lsm+3st,染色体长度比为1.61,As:K值为70.60%,核型类型属子st比bins的3B型。云南黄连1个种群的核型公式为2下x=18=10m十ssm,染色体长度比为1.30,As.K.值为62.05%。中国黄连核型公式为Zn=2舒18:IOm+ssm,染色体长度比为1.69,As.K,值为62,38%。日本黄连核型公式为2n=2护18=12m+6sm,染色体长度比为1.23,As,K.值为59.37%。它们的染色体数目均为18,与属的基数护9相吻合,核型比较相近,核型类型都是相当对称的ZA型。都具有5一6对中部着丝点染色体和3一4对近中部着丝点染色体,比较对称。在中国黄连黄水种群不足20个个体中,发现有一个三倍体植株,它较正常的二倍体植株高大,营养体发达,根状茎也更为粗壮。7.云南黄连的保护遗传学研究用直接扩增片断长度多态(DALP)方法对云南黄连10个种群的遗传多样性和种群遗传结构进行了研究。在六个随机引物组合中,10个种群以及两个外类群J(日本黄连)和C3(中国黄连)的样品在进行DALP扩增后的6个引物组合共检测到160个基因位点,平均每个引物产生26.7条带。10个种群的平均多态位6.45%,物种水平的多态位54.38%。其中西坡生态带缅甸漂草塘(WCT4)种群多态百分率最高为12,5%,其余的WCTI、WCT3、WCTS种群多态百分率最低均为3.75%,东坡生态带怒江沪水金满(ECT04)种群多态百分率最高为n.88%,ECT02种群多态百分率最低为3.75%。从DALP分析得到的数据来看,该物种的遗传多样性指数分别为PPB=54.38%,ht=01853,I=0.2784;种群水平上的遗传多样性指数分别为PPB=6.45%,he=0.0269,I=0,0301。种群间基因分化系数Gst为0.8547,有14.53%的遗传变异来自种群内,85.47%来自种群间,因此,云南黄连的多样性水平是较低的,且遗传分化主要来自于种群间。六种黄连在系统发育树上聚为两大分支:云南黄连和日本黄连的亲缘关系最近,在系统树的下部聚为一支,其次是五裂黄连;而中国黄连和三角叶黄连的亲缘关系最近,在系统树的上部聚为一支,然后是峨嵋黄连。8.云南黄连濒危机制和保护措施云南黄连是一个濒危物种(),主要表现在地理分布区域狭窄,并且伴随生态条件的恶化在不断地收缩,生境破坏较严重;云南黄连野生种群数量急剧下降,并呈衰减趋势;营养繁殖在部分野外种群繁殖中占较大优势。濒危的内在因素主要有果实成熟时种子并未发育完全,有长达半年的后熟期,在自然条件下由种子向幼苗的转化率极低;抗旱性,耐高温性差;遗传多样性较低。外在因素主要有云南黄连种群分布区地质构造复杂;人类活动的破坏;病虫及动物的危害。针对云南黄连严重的濒危状况,本文提出在继承前人就地保护(人工半栽培)成功模式的基础上,尽快开展以种子繁育为主的迁地保护·恢复残存种群·离体保存种质资源和归化自然等保护措施。
英文摘要: Coptis teeta is a rare medicinal plant and is endemic to Northwest of Yunnan Province, now it is listed as national II grand protect plant. In present study, we made our main aim on the conservation biology, many fields concerning with the conservation biology of C. teeta, such as biological characteristics, morphology, ethnobotany, anatomical ecology, cytology, reproductive biology and genetic diversity is also studied on the materials of 10 populations from eastern slope and western slope of Gaoligongshan Mountains, 1 population of C. chmensis and 1 population of C.japonica from Huang shui of Chongqing. Through this study, we want to probe the thought of biosystematics and the methods of conservation biology; meanwhile, this study will accumulate some valuable basic scientific information for the conservation and sustainable uses of C. teeta1. The biological characteristics of C. teetaC. teeta processes both sexual and asexual reproduction. In natural condition, some individuals of the populations on the west-slope, especially wild populations of C. teeta, produce seedlings from sexual reproduction, while some ones and almost entire individuals of populations on the east-slope of C. teeta produce ramets through vegetative reproduction. Either populations on the west-slope or on the east-slope, every individual flowers and seeds normally, but because seed germination needs an after-ripening process of one half year, associated with lower seed dispersal, so a little seedlings could be found. Differentiation of flower buds in the adult plant begins in July, but they flower in next January or February, and their seeds ripe in May, so one life cycle of an individual is always more than 10 years. It was found to be endemic to a small area, to occupy a very narrow habitat, to need a strict niche and to appear highly dispersed small populations. It was mostly confined to steep slopes of moist temperate, broad-leaved evergreen forests. Edaphic and community micro-climatic factors played a vital role in natural distribution, C. teeta is a species endemic to Sino-Himalaya, and mainly distributed in the watershed of several rivers (i.e. Brahmaputra R., Salween R., Irrawaddy R.s which feed into Indian Ocean). Owing to its wild populations are rare, so it is urgent to protect.2. Morphological variation patternThe morphological variation pattern of 20 characters of 10 populations on eastern andwestern slope of Gaoligongshan Mountains was studied. The results demonstrated that mostcoefficients of variation (c.v.) of populations on western slope are larger than those ofpopulations on eastern slope. In the same population, the values of c.v. of five charactersincluding rhizome numbers, rhizome length, ramet numbers, total length of ramet, andnumbers of ramet node are more than 39%. Wild populations on western slope have highestaverage c.v.. The values of average c.v. of populations on eastern slope are less than those ofwestern populations, moreover, the difference of variation among populations on easternslope is not obvious. By taking a comparison of c.v. of the 10 populations, we may find thatthe c.v. of vegetative characters is between 20% to 80%, average c.v. is 42.15%. On thewhole, ramet > rhizome> leaf. The c.v. of reproductive characters is between 10% to 40%,average c.v. is 21.47%, and the arrange of the variation is scape> pistil and stamen> petaland sepal. The average c.v of vegetative characters is almost double of the average c.v. ofreproductive character, This result demonstrates that vegetative characters have largervariation scale, they are more easy influenced by external conditions, but reproductivecharacter are more stable. 3. EthnobotanyBy using ethnobotanical principle, wild investigation and community investigation as well as information study, the traditional utilization model and history of C. teeta by native Lishu and Lemo people who lived in Gaoligongshan Mountain area has been studied. We summarized the traditional knowledge and experiences that they managed and protected this valuable medicinal plant resource, we also thoroughly investigated the interaction between the traditional knowledge and how to utilize, protect and cultivate C. teeta. Through the ethnobotanical study, we can revealed the level that human activities influence on the population, community and ecosystem of C. teeta. The results demonstrated that: 1) 130 years ago, the high economical value made the wild resource become dry up, in order to sustain this important cash income, Lishu people has developed a set of effective methods to cultivate, manage and harvest C. teeta. Traditional model for cultivating and managing C. teeta has profound scientific significance. By this model, this important resource is not only effectively used, the forest in which C. teeta grows, but also is protected. 2)Althrough introduction and cultivation of C. teeta by Lisu people is just for economic purpose firstly, in fact, their activities effectively conserved this endangered medicinal plant species. The cultivation and management that established and developed by local people are indeed in vitro conservation, moreover, they are canonized by modern conservators. This model not only reflects the interaction between traditional culture and plant resource, but reflects how local people to manage the biodiversity. To cultivate and manage medicinal plants whose population is very less will no doubt promote the conservation of local biodiversity and the sustainable uses of economic plants.
语种: 中文
内容类型: 学位论文
URI标识: http://ir.kib.ac.cn/handle/151853/758
Appears in Collections:昆明植物所硕博研究生毕业学位论文_学位论文

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濒危药用植物云南黄连的保护生物学研究.黄骥[d].中国科学院昆明植物研究所,2004.20-25
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