荷花的民族植物学及河南地区品种资源研究
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摘要
荷花是最为古老的栽培植物之一,据考证,我国已有3000多年的栽培历史,有着丰富的栽培品种及乡土资源,许多优良野生及乡土荷花资源还未被有效地开发利用。同时荷花在被子植物的演化中占据着相当独特的地位。荷花是被子植物中起源最早的植物之一,素有“活化石”之称,是冰期以后幸存的孑遗植物,荷花花粉具3条萌发沟,与睡莲的1条萌发沟完全不同,胚胎学研究表明,胚芽构造特殊,叶绿体的发育途径也与睡莲差异较大。本文对荷花民族植物学进行了系统研究,并从形态学和DNA分子水平上对河南引进和乡土荷花品种资源遗传多样性作出评价。民族植物学研究的重点区域选择我国河南为基础,兼顾陕西、甘肃、山西、山东和其它黄河文明发祥地。
1、以汉文化主要古籍及现代研究文献为研究材料,经过区域调查,详细的论证了荷花的历史渊源、传承关系及品种起源、栽培技术发展演变的脉络,在5000年以前的河南仰韶文化遗址考古中,发现室内台面上碳化的莲子,表明人们早已将莲子作为食物;而在3000多年前的《周书》中则记载着:“薮泽已竭,即莲掘藕”,表明西周时期我国人民已经开始种植食用莲藕。据苏州灵岩山“玩花池”遗址考证,荷花作为观赏植物栽植,最早是在2500年前,吴王夫差为西施赏荷而引种。盆栽荷花历史记载最早见于距今1600余年的东晋,书法家王羲之作《柬书堂贴》,“敝于今岁植得千叶者数盆。亦便发花,相继不绝,今已开二十余枝,颇有可观”。1400年前北魏贾思勰的《齐民要术》中记载着“种藕法”,这种用荷花的根茎繁殖,缩短荷花生长年限,提高产量的方法至今仍有借鉴意义。
2、研究荷花文化的形成、历史演变发现:(1)莲与民众日常生活密切关联,由于应用范围的不断扩大,种植技术不断进步,品种不断增多,生产性活动逐步具有娱乐性,莲逐步成为观赏植物。(2)在漫长的历史过程中,莲的思想内涵和文化意蕴不断深化发展,莲的美好、吉祥和圣洁的象征意义不断得到升华和扩大,在中国特有的民族文化环境和宗教艺术体系中代代相传。(3)从远古到近代,在我国悠久文化传承中,叙述、赞美莲的美好,莲也逐渐成为一种文化象征,它代表着一种高尚的文人节操。(4)随着文化发展,莲也成为重要的宗教圣物,尤其与佛教、道教、儒教等宗教的关系特别密切。莲的文化特殊性足以说明它在中国传统文化的特殊地位,渗透在生活中的方方面面,给人们带来感官的审美与愉悦,精神的寄托与满足,并成为艺术创作汲取不尽的物质源泉。
3、研究古代利用荷花观赏、食用和药用的方法及历史传承结果表明:夏商周的先人很早就观赏荷花,秦汉魏晋隋唐宋荷花在中国古园林中得到传承与发展,到了明清,在君子比德的思想影响下,荷花在江南私家园林中得到发扬光大。荷花的莲藕、莲子、花瓣均可食用。在古朴的食荷花传统文化中,蕴含着关于荷花植物资源利用和保护丰富的传统知识,并且存在着对荷花生物多样性管理和利用的朴素的科学内涵;本草学考证荷花、莲子、莲衣、莲房、莲须、莲子心、荷叶、荷梗、藕节等均可药用,荷花能活血止血,莲子能养心,莲须益肾,荷叶能减肥瘦身,藕节能散瘀、解热毒,荷梗通气行水、泻火清心。
4、调查河南荷花地方品种(系)以及引种栽培的主要品种,研究各品种的地理分布,研究荷花种质资源在河南的保护方法,促进荷花遗传多样性的保护与合理利用。确定河南荷花已有及引种成功、性状稳定的品种有96个,还有一些品种正处于引种驯化阶段。在种植规模上,以生产需要的藕莲栽培为主,藕莲品种有8个,主要分布在黄淮流域。花莲品种主要以引进为主,性状稳定的有88个,主要分布在我省各城市公园和风景名胜区。作为生产需要的籽莲几乎没有,现有的籽莲品种均与花莲一起作观赏用。地方品种(系)31个,其中野生、半野生品系15个,主要分布在黄河以南各个地市乡村的水域与湿地。河南荷花资源欠缺,类型较少,并且各类型所分布的品种数很不平衡。在资源调查的基础上进一步探讨了利用传统文化进行现代生物多样性保护的可能性、必要性和可行性。
5、在河南荷花品种资源调查的基础上,本文选择了在河南生长性状稳定的引进品种96个,地方品种(系)31个选取立叶长、立叶宽、叶柄高、群体花期、着花密度、花蕾形、花蕾色、花态、花径、花色、花型、花瓣数、花瓣长、花瓣宽、花柄高15个主要观赏性状进行评比筛选,建立了河南主栽品种资源的评价体系。根据灰色系统理论中的灰色局势决策原理,利用性状加权局势决策方法,对127个河南主要栽培荷花品种(系)的主要性状指标进行分析,并列出了各品种的优劣次序,综合观赏性状排在前十位的依次分别为‘大洒锦’,‘雪里红’,‘傲霜’,‘鄂莲4号’,‘冬红’,灵宝鼎湖湾荷花品系,‘眼儿媚’,息县陈棚汪湖荷花品系,‘曙光’,罗山高湾村荷花品系,有3个地方品种(系)进入前十,说明在荷花育种时乡土品种是荷花重要的亲本资源。为荷花品种的综合评价提供了一种较为客观的分析方法。
6、通过对‘娃娃莲’、‘淡霞’、‘小台红’等96个河南主栽荷花品种及‘九孔阌莲’、‘期思红莲’等31个地方品种(系)的26个性状主成分分析认为影响荷花品种分类的几个主要指标分别为:植株大小和花径、花型、花色和花态。植株大小为荷花品种分类的第一级标准,花型为第二级标准,花色为第三级标准,花态为第四级标准。剔除2个近似的形态性状,数量分类学用24个性状对127个河南主要栽培荷花品种(系)进行了Q型聚类分析。通过分析,将其划分在结合线L0=3.7453处将荷花品种分为两大类,即大株型群(Ⅰ类)和中小株型群(Ⅱ类);结合线L2=2.8234又将大株型群分为三大类,即A组、B组、C组。结合线L4=2.5671又将中小株型群分为三类,即重台类(D组)、重瓣类(E组和F组)、少瓣和半重瓣类(G组)。这些大类下面又可分为若干小类。在荷花育种和生产需求中,根据各类的特点可以对品种进行相关选择,以减少育种和生产的盲目性和工作量。
7、研究荷花遗传多样性分子评价。构建ISSR种群遗传距离树状图;补充和完善荷花品种的分类系统。结果如下:
(1)ISSR反应程序的确立,ISSR-PCR最佳扩增反应程序为:94℃预变性5 min,94℃变性30 s,49～54℃(退火温度随引物不同而定)退火30s,72℃延伸90s,进行30个循环,72℃延伸7 min,10℃保存。
(2)引物筛选,本实验共筛选了5条引物,扩增出44条谱带,其中25条多态性条带,平均每个引物扩增出5.0条多态性条带,引物多态性信息含量变幅为26.7%～66.7%,平均为56.8%。其中3个引物UBC811的6个多态性条带(850bp、710bp、590bp、460bp、329bp、283bp)、UBC840的4个多态性条带(1235bp、875bp、691bp、210bp)以及UBC880的1个多态性条带(406bp)可以将33个荷花品种(系)完全区分开,建立了33个荷花品种(系)的DNA指纹图谱。
(3)遗传相似性分析,根据DNA扩增结果计算的品种间遗传相似系数绝大部分集中在0.5～0.8之间,通过ISSR分子标记分析结果亦可以看出,荷花丰富的变异和较大遗传多样性应该是中国荷花长时间进化和演化的一个结果。
(4)UPGMA法聚类分析,用UPGMA法对33个荷花品种(系)进行聚类,大致可分为4大类群,即第Ⅰ类群由采自灵宝鼎湖湾、灵宝后地、三门峡天鹅湖、民权林七水库、虞城小乔集黑龙潭、虞城小乔集潭坑的6个品系以及‘落霞映雪’、‘娇容醉杯’、‘笑靥’、‘案头青’、‘丽霞’、‘长白藕’、‘小3735’、‘楚天祥云’8个品种组成;第Ⅱ类群由‘滴翠莲’、‘紫玉莲’、‘露丰唇’、‘小栀子’、‘蝶恋花’、‘丹顶玉阁’、‘露华浓’、‘白衣战士’、‘醉梨花’、‘粉青莲’、‘丽质芳姿’、‘赛佛座’、‘霜晨月’、‘黄鹂’、‘夜明珠’、‘小碧莲’16品种组成;‘白鹅’为第Ⅲ类群;由‘天高云淡’和‘翠微夕照’组成第Ⅳ类群。
ABSTRACT According to textual research, Lotus as one of the most ancient cultivated plants, which have been cultivated for more than 3,000 years in China. There were abundant locus cultivars and native resources in China, but many of these resources have not been developed and utilized effectively. Meanwhile lotus plays an important role in the process of angiosperm evolution. Lotus was one of the ancient angiosperms, known as the“living fossil”, which was the surviving relic plant after ice age. Lotus pollens with three germination ditches were completely different from that of water lily with one germination ditch, and embryology studies also suggested there were large differences between locus and water lily on germ structure and chloroplast growth pathway. This paper made a systematic research on lotus national botany, and an evaluation to genetic diversity of lotus resources in Henan local areas and introduced to Henan on morphology and DNA molecular level. Key area of national botanical research was Henan province, including Shanxi province,Gansu province, Shanxi province, Shandong province and other Yellow River civilization birthplace.
1、Taking the ancient books of the Han dynasty culture and modern research literature as research materials, the regional investigation was conducted and it was proved in detail that historical origin, inheriting relationship, cultivation origin, development and evolution of cultivation technology. It was found that there was carbonized lotus seed indoor in Henan Yangshao culture archaeological site 5,000 years ago, which suggested that people had took lotus seed as food. And over 3,000 years ago in《Zhou Shu》it was recorded,“when river was dry, lotus root appeared and was digged”, which suggested that people in our country had started to plant edible lotus root. According to Suzhou“play flower-spinning pool”of Lingyan mountain textual ruins, lotus was introduced and planted as an ornamental plant 2,500 years ago because the king of Wu dynasty entertaining Xishi. The earliest historical record of potted lotus was the saying“I plant several basins of flowers with many leaves this year, and over 20 of them flower, which is considerable”in the book of calligrapher Wang xizhi in Eastern Jin dynasty 1,600 years ago.《QiMinYaoShu》written by Jia Sixie of Northern Wei dynasty 1400 years ago recorded the method of planting lotus root, which can short its age of growth and increase production. The method is also useful for us today.
2、By researching formation and historical evolution of lotus culture some conclusions are drawn. (1) Lotus was close to people’s daily life. As the extensive applications, progress of cultivation technology, increase of cultivars and entertainment of production activities lotus became an ornamental plant. (2)During the long historical period, cultural content and meaning of lotus developed further. Symbolic significance of lotus’s beauty, lucky, holy and purity constantly sublimate and expland, which are handed from generation to generation in China under the unique national cultural environment and religious art system. (3)Literature and folk tales praise the merits of lotus in the long cultural inheritance from ancient to modern times. Lotus become a cultural symbol gradually and represent a kind of noble literati moral integrity. (4) Along with the lotus cultural development lotus also became an important religious holy thing and has close relations especially with Buddhism, Taoism and Confucianism. The cultural specialness of locus suggest its important status in Chinese traditional culture. It has permeated in the people’life as the sensual happiness and spiritual sustenance, also provides never-failing material sources for artistic creation.
3、According to the research of how lotus can be appreciable, edible and officinal in ancient times, the results suggested ancestor in the Xia, Shang and Zhou dynasties began to appreciate lotus; lotus was used widely in Chinese ancient garden in the Qin, Han, Wei, Jin, Sui, Tang and Song dynasties; to the Ming and Qing dynasties lotus developed greatly in Jiangnan private gardens. Lotus root, seed and petal are all edible. The simple traditional culture of edible lotus contain a lot of knowledge about resource utilization and protection of lotus and the scientific connotation of lotus biodiversity management and use. According to herbalism textual research, many parts of lotus are used medicinally. Lotus petals can invigorate circulation of blood and stop bleeding, lotus seed can nourish heart, lotus stamen can be beneficial for kidney, lotus leaves can lose weight, lotus root can relieve bruises and fire toxin, lotus peduncle can discharge fire.
4、In order to protect lotus genetic diversity and use it reasonably the paper investigated and introduction lotus cultivars(strains) in Henan province, researched their geographical distribution and conservation method of lotus germ plasm resources. There were 96 cultivars whose characters were steady including Henan native cultivars and successfully introduction cultivars. some other were during the process of introduction and domestication. On the production scale, it was the lotus root cultivation mainly and there were 8 lotus root cultivars distributing in the Huanghuai River. 88 ornamental lotus cultivars were mainly introduced distributing parks and scenic spots in Henan province. There was almost no seed lotus for production and the existing seed lotus cultivars were used to appreciate as ornamental lotus. There were 31 native cultivars including 15 wild and half-wild cultivars distributing in the water areas and wetlands of many villages in the south of Yellow River. Henan lotus resources and types were fewer, and it’s unbalanced among the cultivar number of certain kinds of types. Based on the resource investigation, it was further discussed that possibility, necessity and feasibility to protect modern biological diversity using traditional culture.
5、Based on the investigation to lotus cultivar resources in Henan province, this paper chose 96 introduction cultivars and 31 native cultivars(strains) which characters were stable in Henan province and established the evaluation system of lotus cultivar sources in Henan. The system selected 15 main ornamental characters, which were length of ripe leaves, width of ripe leaves, height of leave petiole, flowering phase of group, density of flowering, shape of flower-bud, color of flower-bud, form of flower, flower diameter, color of flower, type of flower, number of petals, length of petal,width of petals and height of flower stalk. According to the grey situation decision-making principle of grey system theory the main characters of above 127 lotus cultivars(strains) were analyzed using method of character weighting and these cultivars were sequenced. The best ten cultivars of 127 lotus cultivars(strains) on general ornamental characters were‘Dasajin’,‘Xuelihong’,‘Aoshuang’,‘Eliansihao’,‘Donghong’, Lingbao Dinghu bay lotus strains,‘Yanermei’, Xixian Chenpeng Wang lake lotus strains,‘Shuguang’, Luoshan Gaowan village lotus strains. Among the best ten there were three native cultivars(strains) which suggested the native lotus cultivars were important parental resources for breeding. The result was in accordance with actual production. The evaluation system provided an objective analysis method to overall evaluation of lotus.
6、By a principal component analysis to 26 characters of lotus including 96 main cultivars in Henan such as‘Wawalian’,‘Danxia’,‘Xiaotaihong’, and 31 native cultivars(strains) such as‘Jiukongwenlian’,‘Qisihonglian’, the several main indexes affecting classification of lotus cultivars were size of plants, flower diameter, type of flowers, color of flowers and form of flowers. As classification standard of lotus cultivars size of plants was class I, flower diameter was class II, types of flowers was classⅢ, and color of flowers was class IV. Taking out of two similar morphological characters, Q type clustering analysis was made to 127 main lotus cultivars(strains) in Henan on 24 characters by quantity classification method. By comprehensive comparison, sample clustering is very suitable for lotus cultivar classification. Through analysis, these cultivars were divided into two groups at the combination line L_0 =3.7453, which were large size type (groupⅠ), small and medium-sized type (groupⅡ) . Large size type was divided into three groups at the combination line L_2=2.8234, which were group A, group B, group C. Small and medium-sized type was divided into three groups at the combination line L_4=2.5671, which were duplicate-petalled type (group D), double-petalled type (group E and group F), few-petalled and half double-petalled type (group G). These types can be divided into several small groups. In lotus breeding and production, proper cultivars were chosen according to characters of the different kinds of lotus in order to reduce the blindness and workload.
7、The molecular evaluation of lotus genetic diversity was studied. Genetic distance tree diagram of ISSR population was constructed and the classification systems of lotus cultivars were supplemented and completed.
(1) The best ISSR-PCR amplification reaction procedures was 94℃for 5min as pre-degeneration; 94℃for 30s as degeneration; 49~54℃for 30s as annealing(specific temperature was due to the different primers.); 72℃for 90s as extension; 30 circulation; 72℃for 7min as extension; 10℃for preservation.
(2)This experiment chose five primers and got 44 amplification spectral bands, of which there were 25 polymorphism bands. There were 5 polymorphism bands per primer on average with 26.7% ~ 66.7% of primer polymorphism information and the average was 56.8%. 33 Lotus cultivars(strains) were completely separated by 3 primers and the DNA fingerprint of these cultivars(strains) was established, which were primer UBC811 with six polymorphism bands (850bp, 710bp, 590bp, 460bp,329bp,283bp), primer UBC840 with four polymorphism bands(1235bp,875bp, 691bp,210bp) and primer UBC880 with 1 polymorphism (406bp).
(3)According to results of DNA amplification genetic similarity coefficient of cultivars was from 0.5 to 0.8. The results of ISSR molecular markers also showed the abundant variation and greater genetic diversity of Chinese lotus should be an outcome of evolution for a long time.
(4) Using UPGMA cluster analysis to 33 lotus cultivars(strains), they were roughly divided into four groups. GroupⅠincluded 14 cultivars which were‘LuoXiaYingXue’,‘JiaoRongZuiBei’,‘XiaoYan’,‘AnTouQing’,‘LiXia’,‘ChangBaiOu’,‘Small 3735’,‘ChuTianXiangYun’and other 6 strains from Lingbao Dinghu bay, Lingbao Houdi, Sanmenxia swan lake, Minquan Linqi reservoir, Yucheng Xiaoqiaoji black dragon spring; Yucheng Xiaoqiaoji pool; groupⅡincluded 16 cultivars, which were‘Dicuilian’,‘Ziyulian’,‘Lufengchun’,‘Smallzhizi’,‘Dielianhua’,‘Dandingyuge’,‘Luhuanong’,‘Baiyizhanshi’,‘Zuilihua’,‘Fenqinglian’,‘Lizhifangzi’,‘Saifozuo’,‘Shuangchenyue’,‘Huangli’,‘Huangli’,‘Yemingzhu’,‘Xiaobilian’;‘Baie’belonged to groupⅢ; groupⅣincluded‘Tiangaoyundan’and‘Cuiweixizhao’.
1、以汉文化主要古籍及现代研究文献为研究材料,经过区域调查,详细的论证了荷花的历史渊源、传承关系及品种起源、栽培技术发展演变的脉络,在5000年以前的河南仰韶文化遗址考古中,发现室内台面上碳化的莲子,表明人们早已将莲子作为食物;而在3000多年前的《周书》中则记载着:“薮泽已竭,即莲掘藕”,表明西周时期我国人民已经开始种植食用莲藕。据苏州灵岩山“玩花池”遗址考证,荷花作为观赏植物栽植,最早是在2500年前,吴王夫差为西施赏荷而引种。盆栽荷花历史记载最早见于距今1600余年的东晋,书法家王羲之作《柬书堂贴》,“敝于今岁植得千叶者数盆。亦便发花,相继不绝,今已开二十余枝,颇有可观”。1400年前北魏贾思勰的《齐民要术》中记载着“种藕法”,这种用荷花的根茎繁殖,缩短荷花生长年限,提高产量的方法至今仍有借鉴意义。
2、研究荷花文化的形成、历史演变发现:(1)莲与民众日常生活密切关联,由于应用范围的不断扩大,种植技术不断进步,品种不断增多,生产性活动逐步具有娱乐性,莲逐步成为观赏植物。(2)在漫长的历史过程中,莲的思想内涵和文化意蕴不断深化发展,莲的美好、吉祥和圣洁的象征意义不断得到升华和扩大,在中国特有的民族文化环境和宗教艺术体系中代代相传。(3)从远古到近代,在我国悠久文化传承中,叙述、赞美莲的美好,莲也逐渐成为一种文化象征,它代表着一种高尚的文人节操。(4)随着文化发展,莲也成为重要的宗教圣物,尤其与佛教、道教、儒教等宗教的关系特别密切。莲的文化特殊性足以说明它在中国传统文化的特殊地位,渗透在生活中的方方面面,给人们带来感官的审美与愉悦,精神的寄托与满足,并成为艺术创作汲取不尽的物质源泉。
3、研究古代利用荷花观赏、食用和药用的方法及历史传承结果表明:夏商周的先人很早就观赏荷花,秦汉魏晋隋唐宋荷花在中国古园林中得到传承与发展,到了明清,在君子比德的思想影响下,荷花在江南私家园林中得到发扬光大。荷花的莲藕、莲子、花瓣均可食用。在古朴的食荷花传统文化中,蕴含着关于荷花植物资源利用和保护丰富的传统知识,并且存在着对荷花生物多样性管理和利用的朴素的科学内涵;本草学考证荷花、莲子、莲衣、莲房、莲须、莲子心、荷叶、荷梗、藕节等均可药用,荷花能活血止血,莲子能养心,莲须益肾,荷叶能减肥瘦身,藕节能散瘀、解热毒,荷梗通气行水、泻火清心。
4、调查河南荷花地方品种(系)以及引种栽培的主要品种,研究各品种的地理分布,研究荷花种质资源在河南的保护方法,促进荷花遗传多样性的保护与合理利用。确定河南荷花已有及引种成功、性状稳定的品种有96个,还有一些品种正处于引种驯化阶段。在种植规模上,以生产需要的藕莲栽培为主,藕莲品种有8个,主要分布在黄淮流域。花莲品种主要以引进为主,性状稳定的有88个,主要分布在我省各城市公园和风景名胜区。作为生产需要的籽莲几乎没有,现有的籽莲品种均与花莲一起作观赏用。地方品种(系)31个,其中野生、半野生品系15个,主要分布在黄河以南各个地市乡村的水域与湿地。河南荷花资源欠缺,类型较少,并且各类型所分布的品种数很不平衡。在资源调查的基础上进一步探讨了利用传统文化进行现代生物多样性保护的可能性、必要性和可行性。
5、在河南荷花品种资源调查的基础上,本文选择了在河南生长性状稳定的引进品种96个,地方品种(系)31个选取立叶长、立叶宽、叶柄高、群体花期、着花密度、花蕾形、花蕾色、花态、花径、花色、花型、花瓣数、花瓣长、花瓣宽、花柄高15个主要观赏性状进行评比筛选,建立了河南主栽品种资源的评价体系。根据灰色系统理论中的灰色局势决策原理,利用性状加权局势决策方法,对127个河南主要栽培荷花品种(系)的主要性状指标进行分析,并列出了各品种的优劣次序,综合观赏性状排在前十位的依次分别为‘大洒锦’,‘雪里红’,‘傲霜’,‘鄂莲4号’,‘冬红’,灵宝鼎湖湾荷花品系,‘眼儿媚’,息县陈棚汪湖荷花品系,‘曙光’,罗山高湾村荷花品系,有3个地方品种(系)进入前十,说明在荷花育种时乡土品种是荷花重要的亲本资源。为荷花品种的综合评价提供了一种较为客观的分析方法。
6、通过对‘娃娃莲’、‘淡霞’、‘小台红’等96个河南主栽荷花品种及‘九孔阌莲’、‘期思红莲’等31个地方品种(系)的26个性状主成分分析认为影响荷花品种分类的几个主要指标分别为:植株大小和花径、花型、花色和花态。植株大小为荷花品种分类的第一级标准,花型为第二级标准,花色为第三级标准,花态为第四级标准。剔除2个近似的形态性状,数量分类学用24个性状对127个河南主要栽培荷花品种(系)进行了Q型聚类分析。通过分析,将其划分在结合线L0=3.7453处将荷花品种分为两大类,即大株型群(Ⅰ类)和中小株型群(Ⅱ类);结合线L2=2.8234又将大株型群分为三大类,即A组、B组、C组。结合线L4=2.5671又将中小株型群分为三类,即重台类(D组)、重瓣类(E组和F组)、少瓣和半重瓣类(G组)。这些大类下面又可分为若干小类。在荷花育种和生产需求中,根据各类的特点可以对品种进行相关选择,以减少育种和生产的盲目性和工作量。
7、研究荷花遗传多样性分子评价。构建ISSR种群遗传距离树状图;补充和完善荷花品种的分类系统。结果如下:
(1)ISSR反应程序的确立,ISSR-PCR最佳扩增反应程序为:94℃预变性5 min,94℃变性30 s,49～54℃(退火温度随引物不同而定)退火30s,72℃延伸90s,进行30个循环,72℃延伸7 min,10℃保存。
(2)引物筛选,本实验共筛选了5条引物,扩增出44条谱带,其中25条多态性条带,平均每个引物扩增出5.0条多态性条带,引物多态性信息含量变幅为26.7%～66.7%,平均为56.8%。其中3个引物UBC811的6个多态性条带(850bp、710bp、590bp、460bp、329bp、283bp)、UBC840的4个多态性条带(1235bp、875bp、691bp、210bp)以及UBC880的1个多态性条带(406bp)可以将33个荷花品种(系)完全区分开,建立了33个荷花品种(系)的DNA指纹图谱。
(3)遗传相似性分析,根据DNA扩增结果计算的品种间遗传相似系数绝大部分集中在0.5～0.8之间,通过ISSR分子标记分析结果亦可以看出,荷花丰富的变异和较大遗传多样性应该是中国荷花长时间进化和演化的一个结果。
(4)UPGMA法聚类分析,用UPGMA法对33个荷花品种(系)进行聚类,大致可分为4大类群,即第Ⅰ类群由采自灵宝鼎湖湾、灵宝后地、三门峡天鹅湖、民权林七水库、虞城小乔集黑龙潭、虞城小乔集潭坑的6个品系以及‘落霞映雪’、‘娇容醉杯’、‘笑靥’、‘案头青’、‘丽霞’、‘长白藕’、‘小3735’、‘楚天祥云’8个品种组成;第Ⅱ类群由‘滴翠莲’、‘紫玉莲’、‘露丰唇’、‘小栀子’、‘蝶恋花’、‘丹顶玉阁’、‘露华浓’、‘白衣战士’、‘醉梨花’、‘粉青莲’、‘丽质芳姿’、‘赛佛座’、‘霜晨月’、‘黄鹂’、‘夜明珠’、‘小碧莲’16品种组成;‘白鹅’为第Ⅲ类群;由‘天高云淡’和‘翠微夕照’组成第Ⅳ类群。
ABSTRACT According to textual research, Lotus as one of the most ancient cultivated plants, which have been cultivated for more than 3,000 years in China. There were abundant locus cultivars and native resources in China, but many of these resources have not been developed and utilized effectively. Meanwhile lotus plays an important role in the process of angiosperm evolution. Lotus was one of the ancient angiosperms, known as the“living fossil”, which was the surviving relic plant after ice age. Lotus pollens with three germination ditches were completely different from that of water lily with one germination ditch, and embryology studies also suggested there were large differences between locus and water lily on germ structure and chloroplast growth pathway. This paper made a systematic research on lotus national botany, and an evaluation to genetic diversity of lotus resources in Henan local areas and introduced to Henan on morphology and DNA molecular level. Key area of national botanical research was Henan province, including Shanxi province,Gansu province, Shanxi province, Shandong province and other Yellow River civilization birthplace.
1、Taking the ancient books of the Han dynasty culture and modern research literature as research materials, the regional investigation was conducted and it was proved in detail that historical origin, inheriting relationship, cultivation origin, development and evolution of cultivation technology. It was found that there was carbonized lotus seed indoor in Henan Yangshao culture archaeological site 5,000 years ago, which suggested that people had took lotus seed as food. And over 3,000 years ago in《Zhou Shu》it was recorded,“when river was dry, lotus root appeared and was digged”, which suggested that people in our country had started to plant edible lotus root. According to Suzhou“play flower-spinning pool”of Lingyan mountain textual ruins, lotus was introduced and planted as an ornamental plant 2,500 years ago because the king of Wu dynasty entertaining Xishi. The earliest historical record of potted lotus was the saying“I plant several basins of flowers with many leaves this year, and over 20 of them flower, which is considerable”in the book of calligrapher Wang xizhi in Eastern Jin dynasty 1,600 years ago.《QiMinYaoShu》written by Jia Sixie of Northern Wei dynasty 1400 years ago recorded the method of planting lotus root, which can short its age of growth and increase production. The method is also useful for us today.
2、By researching formation and historical evolution of lotus culture some conclusions are drawn. (1) Lotus was close to people’s daily life. As the extensive applications, progress of cultivation technology, increase of cultivars and entertainment of production activities lotus became an ornamental plant. (2)During the long historical period, cultural content and meaning of lotus developed further. Symbolic significance of lotus’s beauty, lucky, holy and purity constantly sublimate and expland, which are handed from generation to generation in China under the unique national cultural environment and religious art system. (3)Literature and folk tales praise the merits of lotus in the long cultural inheritance from ancient to modern times. Lotus become a cultural symbol gradually and represent a kind of noble literati moral integrity. (4) Along with the lotus cultural development lotus also became an important religious holy thing and has close relations especially with Buddhism, Taoism and Confucianism. The cultural specialness of locus suggest its important status in Chinese traditional culture. It has permeated in the people’life as the sensual happiness and spiritual sustenance, also provides never-failing material sources for artistic creation.
3、According to the research of how lotus can be appreciable, edible and officinal in ancient times, the results suggested ancestor in the Xia, Shang and Zhou dynasties began to appreciate lotus; lotus was used widely in Chinese ancient garden in the Qin, Han, Wei, Jin, Sui, Tang and Song dynasties; to the Ming and Qing dynasties lotus developed greatly in Jiangnan private gardens. Lotus root, seed and petal are all edible. The simple traditional culture of edible lotus contain a lot of knowledge about resource utilization and protection of lotus and the scientific connotation of lotus biodiversity management and use. According to herbalism textual research, many parts of lotus are used medicinally. Lotus petals can invigorate circulation of blood and stop bleeding, lotus seed can nourish heart, lotus stamen can be beneficial for kidney, lotus leaves can lose weight, lotus root can relieve bruises and fire toxin, lotus peduncle can discharge fire.
4、In order to protect lotus genetic diversity and use it reasonably the paper investigated and introduction lotus cultivars(strains) in Henan province, researched their geographical distribution and conservation method of lotus germ plasm resources. There were 96 cultivars whose characters were steady including Henan native cultivars and successfully introduction cultivars. some other were during the process of introduction and domestication. On the production scale, it was the lotus root cultivation mainly and there were 8 lotus root cultivars distributing in the Huanghuai River. 88 ornamental lotus cultivars were mainly introduced distributing parks and scenic spots in Henan province. There was almost no seed lotus for production and the existing seed lotus cultivars were used to appreciate as ornamental lotus. There were 31 native cultivars including 15 wild and half-wild cultivars distributing in the water areas and wetlands of many villages in the south of Yellow River. Henan lotus resources and types were fewer, and it’s unbalanced among the cultivar number of certain kinds of types. Based on the resource investigation, it was further discussed that possibility, necessity and feasibility to protect modern biological diversity using traditional culture.
5、Based on the investigation to lotus cultivar resources in Henan province, this paper chose 96 introduction cultivars and 31 native cultivars(strains) which characters were stable in Henan province and established the evaluation system of lotus cultivar sources in Henan. The system selected 15 main ornamental characters, which were length of ripe leaves, width of ripe leaves, height of leave petiole, flowering phase of group, density of flowering, shape of flower-bud, color of flower-bud, form of flower, flower diameter, color of flower, type of flower, number of petals, length of petal,width of petals and height of flower stalk. According to the grey situation decision-making principle of grey system theory the main characters of above 127 lotus cultivars(strains) were analyzed using method of character weighting and these cultivars were sequenced. The best ten cultivars of 127 lotus cultivars(strains) on general ornamental characters were‘Dasajin’,‘Xuelihong’,‘Aoshuang’,‘Eliansihao’,‘Donghong’, Lingbao Dinghu bay lotus strains,‘Yanermei’, Xixian Chenpeng Wang lake lotus strains,‘Shuguang’, Luoshan Gaowan village lotus strains. Among the best ten there were three native cultivars(strains) which suggested the native lotus cultivars were important parental resources for breeding. The result was in accordance with actual production. The evaluation system provided an objective analysis method to overall evaluation of lotus.
6、By a principal component analysis to 26 characters of lotus including 96 main cultivars in Henan such as‘Wawalian’,‘Danxia’,‘Xiaotaihong’, and 31 native cultivars(strains) such as‘Jiukongwenlian’,‘Qisihonglian’, the several main indexes affecting classification of lotus cultivars were size of plants, flower diameter, type of flowers, color of flowers and form of flowers. As classification standard of lotus cultivars size of plants was class I, flower diameter was class II, types of flowers was classⅢ, and color of flowers was class IV. Taking out of two similar morphological characters, Q type clustering analysis was made to 127 main lotus cultivars(strains) in Henan on 24 characters by quantity classification method. By comprehensive comparison, sample clustering is very suitable for lotus cultivar classification. Through analysis, these cultivars were divided into two groups at the combination line L_0 =3.7453, which were large size type (groupⅠ), small and medium-sized type (groupⅡ) . Large size type was divided into three groups at the combination line L_2=2.8234, which were group A, group B, group C. Small and medium-sized type was divided into three groups at the combination line L_4=2.5671, which were duplicate-petalled type (group D), double-petalled type (group E and group F), few-petalled and half double-petalled type (group G). These types can be divided into several small groups. In lotus breeding and production, proper cultivars were chosen according to characters of the different kinds of lotus in order to reduce the blindness and workload.
7、The molecular evaluation of lotus genetic diversity was studied. Genetic distance tree diagram of ISSR population was constructed and the classification systems of lotus cultivars were supplemented and completed.
(1) The best ISSR-PCR amplification reaction procedures was 94℃for 5min as pre-degeneration; 94℃for 30s as degeneration; 49~54℃for 30s as annealing(specific temperature was due to the different primers.); 72℃for 90s as extension; 30 circulation; 72℃for 7min as extension; 10℃for preservation.
(2)This experiment chose five primers and got 44 amplification spectral bands, of which there were 25 polymorphism bands. There were 5 polymorphism bands per primer on average with 26.7% ~ 66.7% of primer polymorphism information and the average was 56.8%. 33 Lotus cultivars(strains) were completely separated by 3 primers and the DNA fingerprint of these cultivars(strains) was established, which were primer UBC811 with six polymorphism bands (850bp, 710bp, 590bp, 460bp,329bp,283bp), primer UBC840 with four polymorphism bands(1235bp,875bp, 691bp,210bp) and primer UBC880 with 1 polymorphism (406bp).
(3)According to results of DNA amplification genetic similarity coefficient of cultivars was from 0.5 to 0.8. The results of ISSR molecular markers also showed the abundant variation and greater genetic diversity of Chinese lotus should be an outcome of evolution for a long time.
(4) Using UPGMA cluster analysis to 33 lotus cultivars(strains), they were roughly divided into four groups. GroupⅠincluded 14 cultivars which were‘LuoXiaYingXue’,‘JiaoRongZuiBei’,‘XiaoYan’,‘AnTouQing’,‘LiXia’,‘ChangBaiOu’,‘Small 3735’,‘ChuTianXiangYun’and other 6 strains from Lingbao Dinghu bay, Lingbao Houdi, Sanmenxia swan lake, Minquan Linqi reservoir, Yucheng Xiaoqiaoji black dragon spring; Yucheng Xiaoqiaoji pool; groupⅡincluded 16 cultivars, which were‘Dicuilian’,‘Ziyulian’,‘Lufengchun’,‘Smallzhizi’,‘Dielianhua’,‘Dandingyuge’,‘Luhuanong’,‘Baiyizhanshi’,‘Zuilihua’,‘Fenqinglian’,‘Lizhifangzi’,‘Saifozuo’,‘Shuangchenyue’,‘Huangli’,‘Huangli’,‘Yemingzhu’,‘Xiaobilian’;‘Baie’belonged to groupⅢ; groupⅣincluded‘Tiangaoyundan’and‘Cuiweixizhao’.
引文
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