用形态标记和分子标记研究蜡梅栽培种质的遗传多样性
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摘要
蜡梅(Chimonanthus praecox)是蜡梅科蜡梅属落叶灌木，我国的传统观赏花木，具有重要的观赏价值和经济价值。在长达千年的栽培过程中，蜡梅产生了丰富的变异。但是，由于现有的蜡梅品种记载与描述缺乏统一的标准与规范，现有的品种分类体系中选用的性状及标准不一样，造成了品种的描述与记载相当混乱，同名异物及同物异名的现象时有发生，品种分类系统也有多个，这对蜡梅种质资源的利用、保存与交流带来极大的不方便。
武汉和南京特别适宜蜡梅的栽培与生产，而且从目前来看，所收集保存的种质资源也最为丰富，本研究对武汉和南京两地栽培的蜡梅种质资源进行了多年的调查与观测，用形态标记、RAPD标记和ISSR标记对这些种质进行了遗传多样性的研究，探寻蜡梅种质形态性状的变异规律、宏观的形态性状与分子标记之间的关系，为蜡梅的种质(或品种)鉴定、品种分类提供依据。
1、形态学研究
(1)蜡梅形态性状的变异幅度分析、年份间及蜡梅种质间的双因素方差检验表明：花期、花蕾色、花色、花被片是否内含、是否波皱、是否反卷、中被片长、中被片长宽比、中被片面积、内被片长、内被片长宽比、内被片面积、中内被片数目和、雄蕊数目和内被片紫纹颜色深浅这15个性状在同一蜡梅种质内表现稳定，在不同蜡梅种质之间差异显著，在不同年份之间变化小，适合用于蜡梅的鉴定和性状评价。
(2)供试的蜡梅种质在形态上具有丰富的多样性，8个数值性状的变异系数为9.54％-28.30％，最大值与最小值之间的差距一般都在2倍以上。15个性状的平均形态多样性指数为0.59。武汉蜡梅种质的形态多样性指数比南京的高。
(3) Q型聚类把供试蜡梅种质分为绿蕾类、卷瓣类、大花波皱类、狭瓣磬口类、大花类、大花磬口类、小花类和小花狗牙类，各类别之间在形态上差异明显。内被片紫纹颜色深浅对蜡梅聚类起的作用不是特别重要。
(4) R型聚类中反映出花蕾是否为绿色与花被片是否波皱有一定的相关性。
(5)主成分分析中得到前8个主成分累积贡献率达86.37％，分别反映的是蜡梅花的大小、花被片形状、花被片是否波皱、花被片数目及是否内含、花被片是否反卷、花色及花蕾色、内被片颜色和花期。这些主成分基本上体现出了Q型聚类中的各类别的主要特点。主成分聚类的结果与Q型聚类的结果相关性极高(r=0.98)，表明15个形态性状选择合理，它们之间的相关性没有对形态聚类产生太多影响。
(6)Bayes判别证明了所选的15个性状都对蜡梅的归类有极强的辨别能力。Bayes判别公式可用于今后蜡梅种质的电子数据库管理。
2、RAPD和ISSR研究
(1)利用19个RAPD引物得到165条扩增带，其中105(63.63％)为多态性条带。利用11个ISSR引物扩增出115条带，其中90条(78.26％)为多态性条带。RAPD或ISSR标记都可以区分出所有供试蜡梅种质。
(2) RAPD和ISSR的Nei指数分别为h_(RAPD)=0.21，h_(ISSR)=0.25，显示出蜡梅栽培种质有较高的遗传多样性，RAPD和ISSR均表明武汉蜡梅种质的遗传多样性比南京的高。
(3) RAPD和ISSR的聚类、主坐标分析均表明，蜡梅的聚类与其地理来源有紧密联系。
(4) RAPD表明有86.75％的遗传变异存在于蜡梅栽培群体内部，ISSR表明有85.68％的遗传变异存在于蜡梅栽培群体内部；仅有少数遗传变异存在于武汉蜡梅群体与南京蜡梅群体之间。进一步的AMOVA和HOMOVA分析表明，两个蜡梅群体之间的遗传差异显著(RAPD的Φ_(ST)=0.132，p＜0.001：ISSR的Φ_(ST)=0.143，p＜0.001)。
(5) RAPD和ISSR都显示出花蕾是否为绿色这一形态性状与花被片是否波皱这一性状存在一定相关性。内被片紫纹深浅这一性状在分子聚类中不重要。
(6)分子标记与形态标记的相关性较低，RAPD标记与ISSR标记有较密切的相关性。
形态标记与分子标记的综合结果表明，蜡梅在长期栽培的过程中，产生了丰富的遗传多样性。由于各地选种的标准不同，可能在不同地域形成了独特的基因库，使蜡梅栽培种质产生了一定的遗传分化。在今后的蜡梅育种中，应有意识地选择来自于应不同地域的亲本，以充分利用现有种质的遗传多样性。
Wintersweet (Calycanthaceae, Chimonanthus praecox), a kind of perennial deciduous shrub, is an traditional ornamental flowering plant endemic to China and is of great ornamental and economic value in practice. During its long history of cultivation with more than one thousand years, abundant variations arose, producing a variety of germplasms. However, at present, due to lack of uniformed standards and rules for identification and documented description of wintersweet cultivars, various characters as well as different criteria for their assessment were employed in the current cultivar classification system, which caused considerable confusions in cultivar identification and description. Furthermore, because there existed several different cultivar classification systems, homonyms and synonyms had occurred from time to time, which brought a great deal of inconvenience in the exchange, utilization and conservation of germplasms resources of wintersweet.
Wuhan and Nanjing are two regions especially suitable for the cultivation and production of wintersweet. In addition, for the'moment, these two regions appear to hold the most abundant germplasms resources of wintersweet germplasms. In this research, the germplasms resources of wintersweet cultivated in Wuhan and Nanjing were observed and surveyed for years, and as well the genetic diversity of these germplasms was investigated by means of morphological, RAPD and ISSR markers. The purposes of this research were to explore for the rules of variations in morphological traits of wintersweet germplasms, identify the potential relationships between morphological traits and molecular markers with respect to their utilizations in cultivar classification, and eventually provide a basis for germplasm or cultivar identification and classification for wintersweet.
Firstly, morphological research was carded out.
(1) Based on the analysis of extent of variation and two-way ANOVA among years and genotypes of wintersweet for morphological traits, it was demonstrated that the following 15 characters were suitable to be used for identification, description and evaluation in classification system in that they exhibited significant differences among different germplasm categories but gave stable performance within a category, with minor variance between years. The 15 characters include: flower period, flower color, flower bud color, flower urn-shaped or not, tepal margin crisped or not, the tip of the tepal recurved or not, medium tepal length, medium tepal length/width, area size of medium tepal, inner tepal length, inner tepal length/width, area size of inner tepal, number of medium and inner tepals, stamen number and color of inner tepal
(2) There were rich diversities on morphology for the wintersweet germplasms under the study. The variation of 8 numerical characters ranged from 9.54% to 28.3%, with over 2 times difference between maximum and minimum values. The average morphological diversity index for the 15 characters reached 0.59, while the morphological diversity index for the genotypes from Wuhan was higher than that for the genotypes from Nanjing.
(3) By means of Q cluster analysis, the wintersweet germplasms collected for this study were categorized into 8 groups, i. e. green flower bud group, big-sized and crisped tepal group, urn-shaped and narrow tepal group, big-sized group, big-sized and urn-shaped group, small-sized group and patens group. They were distinctive on morphology between each other.
(4) It was shown by R cluster that there was some degree of correlation between the characters of flower bud (green or not) and tepal margin (crisped or not).
(5) In principal component analysis, the first 8 principal components, from which the cumulative contribution reached 86.37%, corresponded to the following 8 characters, i. e. flower size, tepal shape, tepal margin crisped or not, number of tepals and flower urn-shaped or not, tepal recurved or not, flower and flower bud color, color of inner tepal and flower period. Moreover, these principal components basically reflected the major attributes of each of the groups generated in Q cluster analysis. The extremely high correlation (r=0.98) between the results produced by principal component analysis and Q cluster analysis showed that it was reasonable to choose the 15 morphological characters and the correlation among them had little effect on morphological clustering.
(6) It was proven by Bayes discriminant analysis that the selected 15 characters had strong discriminative power for classification of wintersweet. The discriminant formulae can be used for the management of wintersweet germplasms in the future.
Secondly, RAPD and ISSR molecular marker analyses were carried out.
(1) 19 RAPD primers amplified 165 bands, of which 105 (63.63%) were polymorphic, while 11 ISSR primers generated 115 bands, with 90 (78.26%) polymorphic. Either ISSR or RAPD markers were sufficient to distinguish all the genotypes surveyed.
(2) The Nei index calculated for RAPD or ISSR was h_RAPD=0.21 or h_ISSR=0.25 respectively, showing the relatively high level of genetic diversity existing in cultivated wintersweet germplasms. Meanwhile, both analyses indicated that wintersweet from Wuhan had higher genetic diversity compared to that from Nanjing.
(3) Clustering and principal coordinate analysis using both RAPD and ISSR data revealed that the clustering pattern of wintersweet was tightly linked with their geographical origins.
(4) Most of the genetic variation (86.75% with RAPD data and 85.68% with ISSR data) occurred among genotypes within each region, with the remaining existing between two regions. However, it was indicated by further AMOVA and HOMOVA analyses that the genetic difference between Wuhan and Nanjing groups was statistically significant (Φ_ST =0.132, p＜0.001, with RAPD data;Φ_ST=0.143, p＜0.001, with ISSR data).
(5) In consistent with the result by the morphological analysis, both RAPD and ISSR analyses revealed that the two morphological characters, flower bud color (green or not) and tepal margin shape (crisped or not), had very close relationship.
(6) By comparison, with respect to clustering, while the correlation between morphological and each molecular marker was low, two kinds of molecular markers were closely related.
The comprehensive results by the above analyses demonstrated that abundant genetic diversity was produced during the long history of wintersweet cultivation. Due to the distinctive criteria adopted for breeding, the unique gene pools were gradually formed in different geographical regions, resulting in the genetic differentiation of cultivated wintersweet germplasms. Consequently, in the future breeding program, parental lines from diverse regions should be utilized on purpose so as to make the most of the genetic diversity reserved in the current germplasms.
武汉和南京特别适宜蜡梅的栽培与生产，而且从目前来看，所收集保存的种质资源也最为丰富，本研究对武汉和南京两地栽培的蜡梅种质资源进行了多年的调查与观测，用形态标记、RAPD标记和ISSR标记对这些种质进行了遗传多样性的研究，探寻蜡梅种质形态性状的变异规律、宏观的形态性状与分子标记之间的关系，为蜡梅的种质(或品种)鉴定、品种分类提供依据。
1、形态学研究
(1)蜡梅形态性状的变异幅度分析、年份间及蜡梅种质间的双因素方差检验表明：花期、花蕾色、花色、花被片是否内含、是否波皱、是否反卷、中被片长、中被片长宽比、中被片面积、内被片长、内被片长宽比、内被片面积、中内被片数目和、雄蕊数目和内被片紫纹颜色深浅这15个性状在同一蜡梅种质内表现稳定，在不同蜡梅种质之间差异显著，在不同年份之间变化小，适合用于蜡梅的鉴定和性状评价。
(2)供试的蜡梅种质在形态上具有丰富的多样性，8个数值性状的变异系数为9.54％-28.30％，最大值与最小值之间的差距一般都在2倍以上。15个性状的平均形态多样性指数为0.59。武汉蜡梅种质的形态多样性指数比南京的高。
(3) Q型聚类把供试蜡梅种质分为绿蕾类、卷瓣类、大花波皱类、狭瓣磬口类、大花类、大花磬口类、小花类和小花狗牙类，各类别之间在形态上差异明显。内被片紫纹颜色深浅对蜡梅聚类起的作用不是特别重要。
(4) R型聚类中反映出花蕾是否为绿色与花被片是否波皱有一定的相关性。
(5)主成分分析中得到前8个主成分累积贡献率达86.37％，分别反映的是蜡梅花的大小、花被片形状、花被片是否波皱、花被片数目及是否内含、花被片是否反卷、花色及花蕾色、内被片颜色和花期。这些主成分基本上体现出了Q型聚类中的各类别的主要特点。主成分聚类的结果与Q型聚类的结果相关性极高(r=0.98)，表明15个形态性状选择合理，它们之间的相关性没有对形态聚类产生太多影响。
(6)Bayes判别证明了所选的15个性状都对蜡梅的归类有极强的辨别能力。Bayes判别公式可用于今后蜡梅种质的电子数据库管理。
2、RAPD和ISSR研究
(1)利用19个RAPD引物得到165条扩增带，其中105(63.63％)为多态性条带。利用11个ISSR引物扩增出115条带，其中90条(78.26％)为多态性条带。RAPD或ISSR标记都可以区分出所有供试蜡梅种质。
(2) RAPD和ISSR的Nei指数分别为h_(RAPD)=0.21，h_(ISSR)=0.25，显示出蜡梅栽培种质有较高的遗传多样性，RAPD和ISSR均表明武汉蜡梅种质的遗传多样性比南京的高。
(3) RAPD和ISSR的聚类、主坐标分析均表明，蜡梅的聚类与其地理来源有紧密联系。
(4) RAPD表明有86.75％的遗传变异存在于蜡梅栽培群体内部，ISSR表明有85.68％的遗传变异存在于蜡梅栽培群体内部；仅有少数遗传变异存在于武汉蜡梅群体与南京蜡梅群体之间。进一步的AMOVA和HOMOVA分析表明，两个蜡梅群体之间的遗传差异显著(RAPD的Φ_(ST)=0.132，p＜0.001：ISSR的Φ_(ST)=0.143，p＜0.001)。
(5) RAPD和ISSR都显示出花蕾是否为绿色这一形态性状与花被片是否波皱这一性状存在一定相关性。内被片紫纹深浅这一性状在分子聚类中不重要。
(6)分子标记与形态标记的相关性较低，RAPD标记与ISSR标记有较密切的相关性。
形态标记与分子标记的综合结果表明，蜡梅在长期栽培的过程中，产生了丰富的遗传多样性。由于各地选种的标准不同，可能在不同地域形成了独特的基因库，使蜡梅栽培种质产生了一定的遗传分化。在今后的蜡梅育种中，应有意识地选择来自于应不同地域的亲本，以充分利用现有种质的遗传多样性。
Wintersweet (Calycanthaceae, Chimonanthus praecox), a kind of perennial deciduous shrub, is an traditional ornamental flowering plant endemic to China and is of great ornamental and economic value in practice. During its long history of cultivation with more than one thousand years, abundant variations arose, producing a variety of germplasms. However, at present, due to lack of uniformed standards and rules for identification and documented description of wintersweet cultivars, various characters as well as different criteria for their assessment were employed in the current cultivar classification system, which caused considerable confusions in cultivar identification and description. Furthermore, because there existed several different cultivar classification systems, homonyms and synonyms had occurred from time to time, which brought a great deal of inconvenience in the exchange, utilization and conservation of germplasms resources of wintersweet.
Wuhan and Nanjing are two regions especially suitable for the cultivation and production of wintersweet. In addition, for the'moment, these two regions appear to hold the most abundant germplasms resources of wintersweet germplasms. In this research, the germplasms resources of wintersweet cultivated in Wuhan and Nanjing were observed and surveyed for years, and as well the genetic diversity of these germplasms was investigated by means of morphological, RAPD and ISSR markers. The purposes of this research were to explore for the rules of variations in morphological traits of wintersweet germplasms, identify the potential relationships between morphological traits and molecular markers with respect to their utilizations in cultivar classification, and eventually provide a basis for germplasm or cultivar identification and classification for wintersweet.
Firstly, morphological research was carded out.
(1) Based on the analysis of extent of variation and two-way ANOVA among years and genotypes of wintersweet for morphological traits, it was demonstrated that the following 15 characters were suitable to be used for identification, description and evaluation in classification system in that they exhibited significant differences among different germplasm categories but gave stable performance within a category, with minor variance between years. The 15 characters include: flower period, flower color, flower bud color, flower urn-shaped or not, tepal margin crisped or not, the tip of the tepal recurved or not, medium tepal length, medium tepal length/width, area size of medium tepal, inner tepal length, inner tepal length/width, area size of inner tepal, number of medium and inner tepals, stamen number and color of inner tepal
(2) There were rich diversities on morphology for the wintersweet germplasms under the study. The variation of 8 numerical characters ranged from 9.54% to 28.3%, with over 2 times difference between maximum and minimum values. The average morphological diversity index for the 15 characters reached 0.59, while the morphological diversity index for the genotypes from Wuhan was higher than that for the genotypes from Nanjing.
(3) By means of Q cluster analysis, the wintersweet germplasms collected for this study were categorized into 8 groups, i. e. green flower bud group, big-sized and crisped tepal group, urn-shaped and narrow tepal group, big-sized group, big-sized and urn-shaped group, small-sized group and patens group. They were distinctive on morphology between each other.
(4) It was shown by R cluster that there was some degree of correlation between the characters of flower bud (green or not) and tepal margin (crisped or not).
(5) In principal component analysis, the first 8 principal components, from which the cumulative contribution reached 86.37%, corresponded to the following 8 characters, i. e. flower size, tepal shape, tepal margin crisped or not, number of tepals and flower urn-shaped or not, tepal recurved or not, flower and flower bud color, color of inner tepal and flower period. Moreover, these principal components basically reflected the major attributes of each of the groups generated in Q cluster analysis. The extremely high correlation (r=0.98) between the results produced by principal component analysis and Q cluster analysis showed that it was reasonable to choose the 15 morphological characters and the correlation among them had little effect on morphological clustering.
(6) It was proven by Bayes discriminant analysis that the selected 15 characters had strong discriminative power for classification of wintersweet. The discriminant formulae can be used for the management of wintersweet germplasms in the future.
Secondly, RAPD and ISSR molecular marker analyses were carried out.
(1) 19 RAPD primers amplified 165 bands, of which 105 (63.63%) were polymorphic, while 11 ISSR primers generated 115 bands, with 90 (78.26%) polymorphic. Either ISSR or RAPD markers were sufficient to distinguish all the genotypes surveyed.
(2) The Nei index calculated for RAPD or ISSR was h_RAPD=0.21 or h_ISSR=0.25 respectively, showing the relatively high level of genetic diversity existing in cultivated wintersweet germplasms. Meanwhile, both analyses indicated that wintersweet from Wuhan had higher genetic diversity compared to that from Nanjing.
(3) Clustering and principal coordinate analysis using both RAPD and ISSR data revealed that the clustering pattern of wintersweet was tightly linked with their geographical origins.
(4) Most of the genetic variation (86.75% with RAPD data and 85.68% with ISSR data) occurred among genotypes within each region, with the remaining existing between two regions. However, it was indicated by further AMOVA and HOMOVA analyses that the genetic difference between Wuhan and Nanjing groups was statistically significant (Φ_ST =0.132, p＜0.001, with RAPD data;Φ_ST=0.143, p＜0.001, with ISSR data).
(5) In consistent with the result by the morphological analysis, both RAPD and ISSR analyses revealed that the two morphological characters, flower bud color (green or not) and tepal margin shape (crisped or not), had very close relationship.
(6) By comparison, with respect to clustering, while the correlation between morphological and each molecular marker was low, two kinds of molecular markers were closely related.
The comprehensive results by the above analyses demonstrated that abundant genetic diversity was produced during the long history of wintersweet cultivation. Due to the distinctive criteria adopted for breeding, the unique gene pools were gradually formed in different geographical regions, resulting in the genetic differentiation of cultivated wintersweet germplasms. Consequently, in the future breeding program, parental lines from diverse regions should be utilized on purpose so as to make the most of the genetic diversity reserved in the current germplasms.
引文
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