杜鹃花品种资源多样性研究及品种分类体系构建
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摘要
杜鹃花,泛指杜鹃花科杜鹃花属植物,是我国十大传统名花之一,从唐代起就有栽培。该属下种质资源丰富,种类超过1000个,且花色花型丰富多彩,因此使其闻名于世。全世界约有800个种遍布北半球,从热带到极地都有分布,其中原产于我国的就超过500个。杜鹃花属植物可分为8个亚属,通常所指的杜鹃花(azalea)是由马银花亚属、映山红亚属和羊踯躅亚属下的种和品种等构成。全世界现有杜鹃花园艺品种28000个以上,我国目前保存的约有300个,其中很多来自日本和欧洲。目前我国使用的杜鹃花品种分类体系是由黄茂如和强鸿良于1984年提出的,他们根据表型和来源,将杜鹃花品种分成4类,1)东鹃(小叶,小花,双套)；2)西鹃(大花,常有半重瓣或重瓣品种)；3)毛鹃(花大、茎叶多毛)；4)夏鹃(初夏开花的品种)。
多年来,有关杜鹃花品种资源多样性和品种分类方面的研究一直比较缺乏。本研究搜集了130个国内保存的品种,利用表型性状的数量分类和AFLP、SSR分子标记,研究其亲缘关系和遗传多样性,并以“二元分类”为指导,构建了我国杜鹃花品种分类体系。主要研究结果如下：
1.杜鹃花国内主栽品种的数量分类研究
以66个国内常见杜鹃花品种为材料,仔细调查了30个表型性状,其中质量性状14个,数量性状16个。在NTSYS软件上计算各样品间的欧氏距离,利用非加权配对算术平均法(UPGMA)进行聚类分析。Mantel检验显示聚类结果和欧式距离矩阵的相关性较好。主坐标分析显示,前三个主向量共能代表49.94%的多样性。在所有表型性状中,有14个性状在前三个主向量上有较高载荷,其中与花相关的性状占了绝大多数,由此说明花部性状在数量分类中占有重要地位。聚类结果和主坐标分析都显示,东鹃和夏鹃能够与传统分类系统中的其他两类相区分,而西鹃和毛鹃从表型关系上看更为接近。
2.基于AFLP分子标记的杜鹃花品种亲缘关系分析
从各品种资源保护地和公园采集了130份杜鹃花种质,包括了东鹃、西鹃、毛鹃和夏鹃中的主要品种,部分未知品种以及近缘种,利用3对AFLP荧光标记引物组合进行分析,共获得408个有效的荧光标记片段,平均每对引物组合获得136个片段。在Treecon vl.3b软件上利用Dice系数计算样品间的遗传相似性,构建邻接树,并设置bootstrap为100计算置信度。基于Dice距离的AFLP标记主坐标分析在NTSYS软件上进行,前两个向量累计能解释15.6%的多样性。结果表明,AFLP标记在分析杜鹃花品种遗传多样性时十分有效。聚类结果显示,西鹃和夏鹃品种在传统分类体系里的区分度较好,而东鹃和毛鹃类界限不明,类别划分标准需要重新定义。
3.基于SSR分子标记的杜鹃花品种遗传多样性分析
利用Dendauw等开发的6对SSR荧光标记研究了130个杜鹃花种质的遗传多样性,共获得等位基因数98个,平均每对引物16.33个,片段大小在127-293bp之间。各引物在所有样品中的多态性都较好,多态信息含量(PIC)在0.73-0.92之间。引物AO2获得的等位基因数最多,达27个,其多态信息含量和香农指数也是6对引物中最高的。基于Dice系数的UPGMA法聚类结果基本支持传统分类中将杜鹃花品种分成东鹃、毛鹃、夏鹃、西鹃的观点,并且进一步认为西鹃和东鹃品种的来源较夏鹃和毛鹃复杂。
4.杜鹃花品种二元分类体系构建
综合品种资源搜集、数量分类和遗传多样性分析的研究结果,尝试以“二元分类”为原则,以品种来源和演化为基础、花型为辅,探讨以春鹃为例的杜鹃花品种分类体系。初步确立第一级分类标准为株型、枝条姿态,第二级分类标准为瓣类,第三级标准为花型,将所搜集的58个春鹃品种分为2类、6群、14型。
Rhododendrons belong to the family Ericaceae, and are among the ten famous traditional flowers in China with a long cultivation history since the Tang Dynasty. The genus Rhododendron (Ericaceae) is famous world-wide for the diversity of floral colors and forms of its more than1000species. Among them, over800species are distributed throughout the Northern Hemisphere, ranging from tropical to polar climates and more than500species are native to China. The genus is divided into eight subgenera (i.e. Azaleastrum, Candidastrum, Hymenanthes, Mumeazalea, Pentanthera, Rhododendron, Therorhodion and Tsutsusi). The two subgenera Azaleastrum and Tsutsusi together with the subgenus Pentanthera comprise the 'azaleas'. There are over28000cultivars in the world, but only about300cultivars in China, most of which were exotic cultivars arrived from Japan and Europe. The present classification system is put forward by Huang and Qiang (1984) according to the phenotype and the origin:(1) East azaleas, formed by cultivars with small leaves and hose-in-hose flowers;(2) West azaleas, including plants with large flowers and semi-double to double petals;(3) Hairy azaleas, characterized by large simple flowers and finely pubescent stems and leaves; and (4) Summer azaleas, cultivars blooming in the early summer.
Over the years, just a little work has been done on the cultivar diversity and classification. In this research, we collected130azaleas, analyzed the genetic relationships and diversity among them based on numerical taxonomy, AFLP and SSR markers, and managed to put up a new classification system based on the principles of "Dual Classification Method". The main results were as follows:
1. Numerical taxonomy on azalea cultivars in China
Sixty-six evergreen azalea cultivars were investigated in this study based on thirty morphological characters (14qualitative and16quantitative). Cluster analysis using unweighted pair group method with arithmetic average (UPGMA) and principal coordinate analysis (PCO) was performed then using NTSYS software. Mantel's test showed a good fitness between the cluster analysis and Euclidean distance matrix. The first three axes in PCO explained49.94%of the total variation, and revealed that14characters, most of which are related to flowers, played an important part in numerical taxonomy. Both cluster analysis and PCO suggested that cultivars in the East and Summer groups were distinct from other groups in the four-group classification system, but the West and Hairy groups were closer when concerned primarily with phenetic relationships.
2. Genetic relationships between azalea cultivars using AFLP markers
Amplified fragment length polymorphism (AFLP) markers were employed to obtain a clear view of azalea cultivars in China. One hundred and thirty genotypes collected from gardens and nurseries, including cultivars classified in the groups East, West, Hairy, and Summer, unknown cultivars and close species, were analyzed using three primer combinations. A total of408polymorphic fragments were used in data analysis with an average of136fragments per primer pair. Genetic similarities were generated based on Dice coefficients, used to construct a neighbour joining tree, and bootstrapped for100replicates in Treecon vl.3b. Principal coordinate analysis was performed based on Dice distances using NTSYS-pc software. The first two axes explained a total variation of15.6%. The AFLP technique was useful for analyzing genetic diversity in evergreen azaleas. Cluster analysis revealed that cultivars in the West and Summer groups were quite distinct from other groups in the four-group classification system and the East and Hairy groups should be redefined.
3. Genetic diversity analysis based on SSR markers
Six SSR loci developed by Dendauw et al. and labeled with fluorescent dyes were used to evaluate the genetic diversity of130azaleas. A total of98alleles were detected with an average of16.33alleles per locus. The size of alleles ranged from 127bp to293bp. All the loci were polymorphic with polymorphism information content (PIC) ranged from0.73to0.92. The largest number of alleles was27detected from locus A02, from which the highest PIC and Shannon's information index (I) were detected. UPGMA was used to analyze the genetic relationships based on Dice coefficients. Cluster analysis revealed that the four-group classification system was approximately reasonable, further more the cultivars in West or East group were more complicated than in Summer or Hairy group.
4. Classification system of azalea cultivars using "Dual Classification Method"
We did lots of work on cultivar collection, morphological character investigation, and genetic diversity analysis. Based on the principles of "Dual Classification Method" and these researches mentioned before, we managed to put up a new classification system of spring azalea cultivars, which took into consideration both cultivar origin and flower types. The three ranks were defined by branch habits, flower forms, and flower shapes separately. A total of fifty-eight spring azalea cultivars we investigated were classified into two groups, six subgroups, and fourteen forms according to the system.
多年来,有关杜鹃花品种资源多样性和品种分类方面的研究一直比较缺乏。本研究搜集了130个国内保存的品种,利用表型性状的数量分类和AFLP、SSR分子标记,研究其亲缘关系和遗传多样性,并以“二元分类”为指导,构建了我国杜鹃花品种分类体系。主要研究结果如下：
1.杜鹃花国内主栽品种的数量分类研究
以66个国内常见杜鹃花品种为材料,仔细调查了30个表型性状,其中质量性状14个,数量性状16个。在NTSYS软件上计算各样品间的欧氏距离,利用非加权配对算术平均法(UPGMA)进行聚类分析。Mantel检验显示聚类结果和欧式距离矩阵的相关性较好。主坐标分析显示,前三个主向量共能代表49.94%的多样性。在所有表型性状中,有14个性状在前三个主向量上有较高载荷,其中与花相关的性状占了绝大多数,由此说明花部性状在数量分类中占有重要地位。聚类结果和主坐标分析都显示,东鹃和夏鹃能够与传统分类系统中的其他两类相区分,而西鹃和毛鹃从表型关系上看更为接近。
2.基于AFLP分子标记的杜鹃花品种亲缘关系分析
从各品种资源保护地和公园采集了130份杜鹃花种质,包括了东鹃、西鹃、毛鹃和夏鹃中的主要品种,部分未知品种以及近缘种,利用3对AFLP荧光标记引物组合进行分析,共获得408个有效的荧光标记片段,平均每对引物组合获得136个片段。在Treecon vl.3b软件上利用Dice系数计算样品间的遗传相似性,构建邻接树,并设置bootstrap为100计算置信度。基于Dice距离的AFLP标记主坐标分析在NTSYS软件上进行,前两个向量累计能解释15.6%的多样性。结果表明,AFLP标记在分析杜鹃花品种遗传多样性时十分有效。聚类结果显示,西鹃和夏鹃品种在传统分类体系里的区分度较好,而东鹃和毛鹃类界限不明,类别划分标准需要重新定义。
3.基于SSR分子标记的杜鹃花品种遗传多样性分析
利用Dendauw等开发的6对SSR荧光标记研究了130个杜鹃花种质的遗传多样性,共获得等位基因数98个,平均每对引物16.33个,片段大小在127-293bp之间。各引物在所有样品中的多态性都较好,多态信息含量(PIC)在0.73-0.92之间。引物AO2获得的等位基因数最多,达27个,其多态信息含量和香农指数也是6对引物中最高的。基于Dice系数的UPGMA法聚类结果基本支持传统分类中将杜鹃花品种分成东鹃、毛鹃、夏鹃、西鹃的观点,并且进一步认为西鹃和东鹃品种的来源较夏鹃和毛鹃复杂。
4.杜鹃花品种二元分类体系构建
综合品种资源搜集、数量分类和遗传多样性分析的研究结果,尝试以“二元分类”为原则,以品种来源和演化为基础、花型为辅,探讨以春鹃为例的杜鹃花品种分类体系。初步确立第一级分类标准为株型、枝条姿态,第二级分类标准为瓣类,第三级标准为花型,将所搜集的58个春鹃品种分为2类、6群、14型。
Rhododendrons belong to the family Ericaceae, and are among the ten famous traditional flowers in China with a long cultivation history since the Tang Dynasty. The genus Rhododendron (Ericaceae) is famous world-wide for the diversity of floral colors and forms of its more than1000species. Among them, over800species are distributed throughout the Northern Hemisphere, ranging from tropical to polar climates and more than500species are native to China. The genus is divided into eight subgenera (i.e. Azaleastrum, Candidastrum, Hymenanthes, Mumeazalea, Pentanthera, Rhododendron, Therorhodion and Tsutsusi). The two subgenera Azaleastrum and Tsutsusi together with the subgenus Pentanthera comprise the 'azaleas'. There are over28000cultivars in the world, but only about300cultivars in China, most of which were exotic cultivars arrived from Japan and Europe. The present classification system is put forward by Huang and Qiang (1984) according to the phenotype and the origin:(1) East azaleas, formed by cultivars with small leaves and hose-in-hose flowers;(2) West azaleas, including plants with large flowers and semi-double to double petals;(3) Hairy azaleas, characterized by large simple flowers and finely pubescent stems and leaves; and (4) Summer azaleas, cultivars blooming in the early summer.
Over the years, just a little work has been done on the cultivar diversity and classification. In this research, we collected130azaleas, analyzed the genetic relationships and diversity among them based on numerical taxonomy, AFLP and SSR markers, and managed to put up a new classification system based on the principles of "Dual Classification Method". The main results were as follows:
1. Numerical taxonomy on azalea cultivars in China
Sixty-six evergreen azalea cultivars were investigated in this study based on thirty morphological characters (14qualitative and16quantitative). Cluster analysis using unweighted pair group method with arithmetic average (UPGMA) and principal coordinate analysis (PCO) was performed then using NTSYS software. Mantel's test showed a good fitness between the cluster analysis and Euclidean distance matrix. The first three axes in PCO explained49.94%of the total variation, and revealed that14characters, most of which are related to flowers, played an important part in numerical taxonomy. Both cluster analysis and PCO suggested that cultivars in the East and Summer groups were distinct from other groups in the four-group classification system, but the West and Hairy groups were closer when concerned primarily with phenetic relationships.
2. Genetic relationships between azalea cultivars using AFLP markers
Amplified fragment length polymorphism (AFLP) markers were employed to obtain a clear view of azalea cultivars in China. One hundred and thirty genotypes collected from gardens and nurseries, including cultivars classified in the groups East, West, Hairy, and Summer, unknown cultivars and close species, were analyzed using three primer combinations. A total of408polymorphic fragments were used in data analysis with an average of136fragments per primer pair. Genetic similarities were generated based on Dice coefficients, used to construct a neighbour joining tree, and bootstrapped for100replicates in Treecon vl.3b. Principal coordinate analysis was performed based on Dice distances using NTSYS-pc software. The first two axes explained a total variation of15.6%. The AFLP technique was useful for analyzing genetic diversity in evergreen azaleas. Cluster analysis revealed that cultivars in the West and Summer groups were quite distinct from other groups in the four-group classification system and the East and Hairy groups should be redefined.
3. Genetic diversity analysis based on SSR markers
Six SSR loci developed by Dendauw et al. and labeled with fluorescent dyes were used to evaluate the genetic diversity of130azaleas. A total of98alleles were detected with an average of16.33alleles per locus. The size of alleles ranged from 127bp to293bp. All the loci were polymorphic with polymorphism information content (PIC) ranged from0.73to0.92. The largest number of alleles was27detected from locus A02, from which the highest PIC and Shannon's information index (I) were detected. UPGMA was used to analyze the genetic relationships based on Dice coefficients. Cluster analysis revealed that the four-group classification system was approximately reasonable, further more the cultivars in West or East group were more complicated than in Summer or Hairy group.
4. Classification system of azalea cultivars using "Dual Classification Method"
We did lots of work on cultivar collection, morphological character investigation, and genetic diversity analysis. Based on the principles of "Dual Classification Method" and these researches mentioned before, we managed to put up a new classification system of spring azalea cultivars, which took into consideration both cultivar origin and flower types. The three ranks were defined by branch habits, flower forms, and flower shapes separately. A total of fifty-eight spring azalea cultivars we investigated were classified into two groups, six subgroups, and fourteen forms according to the system.
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