杨山牡丹遗传多样性与江南牡丹品种资源研究
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摘要
杨山牡丹(Paeonia ostii)属于芍药科牡丹组革质花盘亚组,我国特产的传统名花和药用植物,是江南牡丹品种群的主要原种之一。因其抗性强,结实力强,是牡丹育种和抗性研究的优良材料。但对于杨山牡丹的资源分布、遗传多样性等尚没有系统的研究。目前,牡丹根据地域分为4个的牡丹品种群,中原、西北、江南和西南牡丹品种群。江南牡丹品种群具有耐湿热的优良特点,古时江南牡丹栽培繁荣,现在,江南品种资源稀缺,有必要进行系统的调查和研究。
为此,本文以杨山牡丹和江南牡丹品种群的品种为研究对象,采用表型与分子手段相结合的方法,对杨山牡丹和江南地区牡丹品种的资源历史和现状、遗传多样性分析研究,揭示其种质资源遗传背景。主要结果如下:
1.历时3年在河南、安徽、湖北、湖南、甘肃、浙江6个省16个地点进行了杨山牡丹的野生分布和栽培分布实地调查。调查包括了记载中除陕西和重庆的所有省份,具有典型性和代表性。调查结果,杨山牡丹分布范围,水平分布于北纬28°12′53.1″—35°33′37.0″,东经106°33′090″—118°01′49.0″。野生杨山牡丹的垂直分布于海拔901—1742m范围,主要分布于河南、安徽和湖北省。栽培凤丹分布范围很广,遍布全国。
2.杨山牡丹居群遗传多样性研究,形态学数量分类在遗传相似系数0.03—0.14范围内的聚类结果显示,杨山牡丹表型变异范围很小,不同居群在表型上很难区分。不同居群间的花粉形态差异较大,说明在孢粉学水平,不同居群间具有较高的遗传多样性,且同一居群内花粉形态也有差别。AFLP分子标记研究结果:在物种水平上,总的多态带百分率(P)为93.23%、基因多样性指数(H)为0.3376、香农指数(1)为0.5025,而在居群水平上平均遗传多样性水平则降低(P=78.76%、H=0.2615、I=0.3942)。居群间的基因分化系数Gst=0.1991,说明其遗传多样性19.91%存在于居群间,80.09%存在于居群内,即其遗传结构变异主要存在于居群内:基因流Nm=2.0116＞1,表明居群间基因交流较少。其中安徽亳州居群与河南宝天曼居群亲缘关系最高,地理距离与遗传距离之间不存在明显相关性。
3.历史上,江南牡丹的分布范围从南朝永嘉到唐代的杭州,到宋代浙江东部及吴中一带(今苏州),再到明清时期的江苏、浙江、上海,分布范围呈现先扩大后趋于平稳。品种数量呈现由少到多,再由多到少的动态消长趋势。现在,江南牡丹品种群主要分布地为上海、安徽东南部、浙江大部、江苏中南部,次要分布地为湖南西北部等。
4.江南牡丹品种群的牡丹品种主要指‘凤丹'和江南传统品种,现存的品种都是特别适应江南气候的优秀品种。因江南地区气候特点以及牡丹本身不太耐湿热的生物学特性等原因,导致江南地区牡丹品种稀缺,且花型花色单一,实际品种数量约20个,分别为:‘凤丹'、‘西施'、‘粉莲'、‘玉楼春'、‘昌红'、‘呼红'、‘羽红'、‘雀好'、‘玫红'、‘四旋'、‘轻罗'、‘云芳'、‘黑楼紫'、‘湘西紫'、‘湘西粉'、‘香丹'、‘玉楼'、‘凤尾'、‘盐城红'和‘盐城粉'。这些品种存在同物异名现象,根据形态和分子证据,对其进行归并处理:‘西施'、‘粉莲'、‘玉楼春'合并,统称‘玉楼春';‘昌红'、‘呼红'、‘羽红'、‘雀好'、‘玫红'、‘四旋'、‘轻罗'合并,统称‘徽紫';‘玉楼'、‘凤尾'合并,统称‘玉楼'。归并整理后,江南现有品种11种。
5.江南地区古牡丹资源丰富。实地调查结果,古牡丹主要分布于上海、浙江、江苏、安徽3省1市,27处,保留于民间私人院落及寺庙中。主要品种有4个,粉花的‘玉楼春'、紫红色花(无名)、‘黑楼紫'和‘盐城红',说明江南地区栽培牡丹历史悠久。有的古牡丹出现衰弱或消失现象,亟需复壮和养护,进行有效地保护和综合利用。
6.江南牡丹的可能起源,主要直接来自中原牡丹,兼有西南和西北牡丹品种。杨山牡丹对江南牡丹品种的起源起间接作用。提出了江南牡丹资源育种与利用的策略及建议,确定江南培育耐湿热、根系浅、观赏性好的育种目标,利用杨山牡丹和江南现有传统品种及古牡丹与其他优良的耐湿热资源,如日本牡丹、西南牡丹品种,通过杂交等方法,选育适合江南地区的新品种。
以上研究为种质资源保护利用、江南牡丹品种分类、江南牡丹新品种选育等研究奠定了理论基础。
Paeonia ostii,which belongs to Paeoniaceae,is one of the famous ornamental and medicinal plants in China.It is also the main species to Jiangnan Mudan Group.Postii is a good material for breeding and researching because of its strong resistance and fruitfulness.However,there were few reports to its distribution,and the studies on the genetic diversity of P.ostii also need to be systematic.According to its area distribution,tree peony can be divided into four main variety groups,they are Zhongyuan (Central Plains),Xibei(Northwest),Jiangnan(Southern Yangtse) and Xinan(Southwest).The Jiangnan Mudan(southern Yangtse tree peony) is adaptive for rainy,moist and warm weather.As early as the old dynasties,there were many cultivars in the Group grew in Southern China.Now,the number of cultivars of this group has decreased,which seriously need systematic investigation and research.
Therefore,this paper used Yangshan Nudan and Jiangnan Mudan as material,based on the method of configuration analysis with molecular data gained by ISSR and AFLP,to study.The main results were as follows:
1.The field distributions of wild and cultivated P.ostii had been surveyed in Henan,Anhui,Hubei, Hunan,Gansu and Zhejiang provinces including 16 spots during 3 years.The investigation results were typical and representative.It covers all provinces in recorded except Shanxi and Chongqing.The investigation shows on-the-spot distribution range investigation of P.ostii,the level distributes in the north latitude 28°12′53.1″- 35°33′37.0″,east longitude 106°33′090″-118°01′49.0″. The wild P.ostii vertical distribution in the elevation 901-1742m scope,mainly distributes in Henan, Anhui and Hubei Province.The distribution range of cultivation P.ostii is very broad,proliferates the nation.Because the medicinal value of P.ostii,the artificial destruction was serious,which lead to serious the wild resources loss.
2.According to the analysis on Genetic Diversity of Different Populations in P.ostii,the results showed that there was an low genetic diversity in morphological characteristics of populations based on the genetic distance from0.03-0.14,different population of P.ostii could not be differentiate by the morphological characteristics.The pollen shape was different Amony and within P.ostii population indicated that there was an abundant genetic diversity in pollen characteristics of populations. Conservation genetics research by AFLP indicated that P.ostii had high genetic diversity at species level (P=93.23%,H=0.3376,I=0.5025),but relatively low levels of polymorphism at the population level (P=78.76%,H=0.2615,I=0.3942);the coefficient of genetic differentiation among populations(Gst) was 0.1991 and if showed that 19.91%of the total genetic variation among populations while the rest 80.09%within populations;the level of gene flow(Nm) was estimated to be 2.0116＞1.Henan Baotianman and Anhui Bozhou populations had the closest relationship,and the geographic distance was not positively correlated with genetic distance.
3.In history,the distribution of Jiangnan tree peony cultivars distributed covered from South Dynasty in Yongjia to the Tang Dynasty in Hangzhou,the Song Dynasety in eastern of Zhejiang and Wuzhong area (now Suzhou),and then the Ming and Qing Dynasties in JiangSu,Zhejiang,Shanghai.The number of cultivars showed the trend of dynamic growth with fluctuation trend.And now,Jiangnan Mudan Group presently mainly distributed in Shanghai,northwest of Zhejiang,south of Jiangsu,southeast of Anhui,the secondary distribution was north-western of Hunan.
4.The tree peony cultivars of Jiangnan Mudan Group mainly meaned to the P.ostii and its traditional cultivars.The existing cultivars had good adaptability to the climatic conditions in Jiangnan area.But due to the moisture climatic characteristics and the sticky soil of Jiangnan regions,as well as the biological characteristics of tree peonies,the number of tree peony cultivars was deficient in Jiangnan area.It was unitary in flower color and shape.The number of actual cultivars was less than 20, including 'Fengdan','Xishi','Fenlian','Yulouchun','Changhong','Huhong','Yuhong','Quehao', 'Meihong','Sixuan','Qingluo','Yunfang','Heilouzi','Xiangxizi','Xiangxifen','Xiangdan','Yulou', 'Fengwei','Yanchenghong','Yancbengfen'.According to morphological and molecular evidence,to resolve the problem of synonym:'Xishi','Fenlian','Yulouchun' were merged and called 'Yulouchun'; 'Changhong','Huhong','Yuhong','Quehao','Meihong','Sixuan','Qingluo',were merged and named 'Huizi';'Yulou','Fengwei',were merged and called 'Yulou'.Then,there were only 11 tree peony o cultivars in Jiangnan area after reorganization.
5.The resources of ancient tree peony cultivars in Jiangnan area was richful.On-the-spot investigation, the ancient tree peony cultivars mainly distributed in 27 places of Shanghai,Zhejiang,Jiangsu,and Anhui Provinces.They were distributed in the private courtyard and old temples.There were mainly four cultivars,'Yulouchun',the purple-red flower(nameless),'Heilouzi' and 'Yanchenghong'.It explained that the history of Jiangnan Mudan was glorious.Some of the ancient tree peonies appeard weak or died.Rejuvenation and conservation,protection and effective utilization were necessary for ancient tree peony.
6.According to the studies,the possible origin of Jiangnan Mudan Cultivar Group,mainly directly came from the tree peony cultivars form Zhongyuan,Xinan and Xibei Group.P.ostii had the indirect participation the formation of tradition cultivars.The strategies of breeding Jiangnan Mudan were proposed.Firstly,the breeding aim of Strong resistance and high value was ascertained.Secondly,using P.ostii and existing wild species and the ancient traditional tree peony from Jiangnan and other excellent heat resistance resources,such as P.×suffruticosa Japan Group,P.×suffruticosa Tianpeng Group,P.rockii Gansu Grou,through hybridization and other means,to breed the excellent cultivars for Jiangnan Area.
This study could be a base theory for protecting and use the germplasm resourses,identifying relationship of different cultivars from Jiangnan Area,selecting and breed excellent cultivars.
为此,本文以杨山牡丹和江南牡丹品种群的品种为研究对象,采用表型与分子手段相结合的方法,对杨山牡丹和江南地区牡丹品种的资源历史和现状、遗传多样性分析研究,揭示其种质资源遗传背景。主要结果如下:
1.历时3年在河南、安徽、湖北、湖南、甘肃、浙江6个省16个地点进行了杨山牡丹的野生分布和栽培分布实地调查。调查包括了记载中除陕西和重庆的所有省份,具有典型性和代表性。调查结果,杨山牡丹分布范围,水平分布于北纬28°12′53.1″—35°33′37.0″,东经106°33′090″—118°01′49.0″。野生杨山牡丹的垂直分布于海拔901—1742m范围,主要分布于河南、安徽和湖北省。栽培凤丹分布范围很广,遍布全国。
2.杨山牡丹居群遗传多样性研究,形态学数量分类在遗传相似系数0.03—0.14范围内的聚类结果显示,杨山牡丹表型变异范围很小,不同居群在表型上很难区分。不同居群间的花粉形态差异较大,说明在孢粉学水平,不同居群间具有较高的遗传多样性,且同一居群内花粉形态也有差别。AFLP分子标记研究结果:在物种水平上,总的多态带百分率(P)为93.23%、基因多样性指数(H)为0.3376、香农指数(1)为0.5025,而在居群水平上平均遗传多样性水平则降低(P=78.76%、H=0.2615、I=0.3942)。居群间的基因分化系数Gst=0.1991,说明其遗传多样性19.91%存在于居群间,80.09%存在于居群内,即其遗传结构变异主要存在于居群内:基因流Nm=2.0116＞1,表明居群间基因交流较少。其中安徽亳州居群与河南宝天曼居群亲缘关系最高,地理距离与遗传距离之间不存在明显相关性。
3.历史上,江南牡丹的分布范围从南朝永嘉到唐代的杭州,到宋代浙江东部及吴中一带(今苏州),再到明清时期的江苏、浙江、上海,分布范围呈现先扩大后趋于平稳。品种数量呈现由少到多,再由多到少的动态消长趋势。现在,江南牡丹品种群主要分布地为上海、安徽东南部、浙江大部、江苏中南部,次要分布地为湖南西北部等。
4.江南牡丹品种群的牡丹品种主要指‘凤丹'和江南传统品种,现存的品种都是特别适应江南气候的优秀品种。因江南地区气候特点以及牡丹本身不太耐湿热的生物学特性等原因,导致江南地区牡丹品种稀缺,且花型花色单一,实际品种数量约20个,分别为:‘凤丹'、‘西施'、‘粉莲'、‘玉楼春'、‘昌红'、‘呼红'、‘羽红'、‘雀好'、‘玫红'、‘四旋'、‘轻罗'、‘云芳'、‘黑楼紫'、‘湘西紫'、‘湘西粉'、‘香丹'、‘玉楼'、‘凤尾'、‘盐城红'和‘盐城粉'。这些品种存在同物异名现象,根据形态和分子证据,对其进行归并处理:‘西施'、‘粉莲'、‘玉楼春'合并,统称‘玉楼春';‘昌红'、‘呼红'、‘羽红'、‘雀好'、‘玫红'、‘四旋'、‘轻罗'合并,统称‘徽紫';‘玉楼'、‘凤尾'合并,统称‘玉楼'。归并整理后,江南现有品种11种。
5.江南地区古牡丹资源丰富。实地调查结果,古牡丹主要分布于上海、浙江、江苏、安徽3省1市,27处,保留于民间私人院落及寺庙中。主要品种有4个,粉花的‘玉楼春'、紫红色花(无名)、‘黑楼紫'和‘盐城红',说明江南地区栽培牡丹历史悠久。有的古牡丹出现衰弱或消失现象,亟需复壮和养护,进行有效地保护和综合利用。
6.江南牡丹的可能起源,主要直接来自中原牡丹,兼有西南和西北牡丹品种。杨山牡丹对江南牡丹品种的起源起间接作用。提出了江南牡丹资源育种与利用的策略及建议,确定江南培育耐湿热、根系浅、观赏性好的育种目标,利用杨山牡丹和江南现有传统品种及古牡丹与其他优良的耐湿热资源,如日本牡丹、西南牡丹品种,通过杂交等方法,选育适合江南地区的新品种。
以上研究为种质资源保护利用、江南牡丹品种分类、江南牡丹新品种选育等研究奠定了理论基础。
Paeonia ostii,which belongs to Paeoniaceae,is one of the famous ornamental and medicinal plants in China.It is also the main species to Jiangnan Mudan Group.Postii is a good material for breeding and researching because of its strong resistance and fruitfulness.However,there were few reports to its distribution,and the studies on the genetic diversity of P.ostii also need to be systematic.According to its area distribution,tree peony can be divided into four main variety groups,they are Zhongyuan (Central Plains),Xibei(Northwest),Jiangnan(Southern Yangtse) and Xinan(Southwest).The Jiangnan Mudan(southern Yangtse tree peony) is adaptive for rainy,moist and warm weather.As early as the old dynasties,there were many cultivars in the Group grew in Southern China.Now,the number of cultivars of this group has decreased,which seriously need systematic investigation and research.
Therefore,this paper used Yangshan Nudan and Jiangnan Mudan as material,based on the method of configuration analysis with molecular data gained by ISSR and AFLP,to study.The main results were as follows:
1.The field distributions of wild and cultivated P.ostii had been surveyed in Henan,Anhui,Hubei, Hunan,Gansu and Zhejiang provinces including 16 spots during 3 years.The investigation results were typical and representative.It covers all provinces in recorded except Shanxi and Chongqing.The investigation shows on-the-spot distribution range investigation of P.ostii,the level distributes in the north latitude 28°12′53.1″- 35°33′37.0″,east longitude 106°33′090″-118°01′49.0″. The wild P.ostii vertical distribution in the elevation 901-1742m scope,mainly distributes in Henan, Anhui and Hubei Province.The distribution range of cultivation P.ostii is very broad,proliferates the nation.Because the medicinal value of P.ostii,the artificial destruction was serious,which lead to serious the wild resources loss.
2.According to the analysis on Genetic Diversity of Different Populations in P.ostii,the results showed that there was an low genetic diversity in morphological characteristics of populations based on the genetic distance from0.03-0.14,different population of P.ostii could not be differentiate by the morphological characteristics.The pollen shape was different Amony and within P.ostii population indicated that there was an abundant genetic diversity in pollen characteristics of populations. Conservation genetics research by AFLP indicated that P.ostii had high genetic diversity at species level (P=93.23%,H=0.3376,I=0.5025),but relatively low levels of polymorphism at the population level (P=78.76%,H=0.2615,I=0.3942);the coefficient of genetic differentiation among populations(Gst) was 0.1991 and if showed that 19.91%of the total genetic variation among populations while the rest 80.09%within populations;the level of gene flow(Nm) was estimated to be 2.0116＞1.Henan Baotianman and Anhui Bozhou populations had the closest relationship,and the geographic distance was not positively correlated with genetic distance.
3.In history,the distribution of Jiangnan tree peony cultivars distributed covered from South Dynasty in Yongjia to the Tang Dynasty in Hangzhou,the Song Dynasety in eastern of Zhejiang and Wuzhong area (now Suzhou),and then the Ming and Qing Dynasties in JiangSu,Zhejiang,Shanghai.The number of cultivars showed the trend of dynamic growth with fluctuation trend.And now,Jiangnan Mudan Group presently mainly distributed in Shanghai,northwest of Zhejiang,south of Jiangsu,southeast of Anhui,the secondary distribution was north-western of Hunan.
4.The tree peony cultivars of Jiangnan Mudan Group mainly meaned to the P.ostii and its traditional cultivars.The existing cultivars had good adaptability to the climatic conditions in Jiangnan area.But due to the moisture climatic characteristics and the sticky soil of Jiangnan regions,as well as the biological characteristics of tree peonies,the number of tree peony cultivars was deficient in Jiangnan area.It was unitary in flower color and shape.The number of actual cultivars was less than 20, including 'Fengdan','Xishi','Fenlian','Yulouchun','Changhong','Huhong','Yuhong','Quehao', 'Meihong','Sixuan','Qingluo','Yunfang','Heilouzi','Xiangxizi','Xiangxifen','Xiangdan','Yulou', 'Fengwei','Yanchenghong','Yancbengfen'.According to morphological and molecular evidence,to resolve the problem of synonym:'Xishi','Fenlian','Yulouchun' were merged and called 'Yulouchun'; 'Changhong','Huhong','Yuhong','Quehao','Meihong','Sixuan','Qingluo',were merged and named 'Huizi';'Yulou','Fengwei',were merged and called 'Yulou'.Then,there were only 11 tree peony o cultivars in Jiangnan area after reorganization.
5.The resources of ancient tree peony cultivars in Jiangnan area was richful.On-the-spot investigation, the ancient tree peony cultivars mainly distributed in 27 places of Shanghai,Zhejiang,Jiangsu,and Anhui Provinces.They were distributed in the private courtyard and old temples.There were mainly four cultivars,'Yulouchun',the purple-red flower(nameless),'Heilouzi' and 'Yanchenghong'.It explained that the history of Jiangnan Mudan was glorious.Some of the ancient tree peonies appeard weak or died.Rejuvenation and conservation,protection and effective utilization were necessary for ancient tree peony.
6.According to the studies,the possible origin of Jiangnan Mudan Cultivar Group,mainly directly came from the tree peony cultivars form Zhongyuan,Xinan and Xibei Group.P.ostii had the indirect participation the formation of tradition cultivars.The strategies of breeding Jiangnan Mudan were proposed.Firstly,the breeding aim of Strong resistance and high value was ascertained.Secondly,using P.ostii and existing wild species and the ancient traditional tree peony from Jiangnan and other excellent heat resistance resources,such as P.×suffruticosa Japan Group,P.×suffruticosa Tianpeng Group,P.rockii Gansu Grou,through hybridization and other means,to breed the excellent cultivars for Jiangnan Area.
This study could be a base theory for protecting and use the germplasm resourses,identifying relationship of different cultivars from Jiangnan Area,selecting and breed excellent cultivars.
引文
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