中国玉兰种质资源调查及亲缘关系的研究
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摘要
玉兰亚属植物(Magnolia Subgenus of Yulania)是中国重要的传统花卉。本研究重点对我国玉兰亚属及白玉兰(Magnolia denudata)的栽培品种资源开展了详细的整理与调查工作,获得了玉兰品种和玉兰亚属部分种的现状情况；建立了玉兰亚属部分种和玉兰品种的AFLP指纹图谱；通过比较各品种间形态学上的差异,结合孢粉学和AFLP分子生物学手段,梳理了玉兰亚属种和玉兰品种的分类情况,探讨了其之间可能的亲缘关系和演化情况。结论如下：
1、经调查和整理,目前有记载的玉兰亚属植物共52种,明确认定18种,近来文献提及的新种24个,尚存有争议的10种；我国作为玉兰亚属植物的起源中心与现代分布中心,共有48个本地种(包括文献提及及有争议种,另4个为国外种)。
经调查整理,初步调查到我国白玉兰和二乔玉兰(M×soulangeana)品种40个,其中白玉兰类26个,二乔玉兰类14个。根据形态学比较分析,本研究提出将‘黄脉’和‘玉蝶’等12个具典型特征的玉兰类型暂作为新品种处理。
运用形态学数量聚类分析将其中37个玉兰类型很清晰地划分为呈间隔分布的5个白玉兰和二乔玉兰品种群,在印证了二乔玉兰与白玉兰在许多外部性状上均存在着广泛交叉和逐渐过渡情况的同时,也表明形态学聚类分析是一种开展玉兰品种分类研究较好的方法。R型聚类显示,30个性状的量化处理较为合理,保证了最终研究结果的准确性和科学性。通过主成分分析,根据贡献率大小确定了小枝粗细、叶芽颜色、花被颜色等18个影响较大的性状因子,为以后开展更为系统、深入的中国玉兰品种分类研究提供了参考依据。
2、对37个样本35个玉兰品种的孢粉学研究显示,玉兰品种的花粉形态相似,均为左右对称,异极,具单沟,超长球形(P/E>2),大小(52.9-61.3μm)×(21.3-27.6μm),属于NPC花粉分类系统中的N1P3C3类型,与玉兰业属植物花粉共同的典型特征一致,但表面纹饰的细微形态结构(穿孔和孔穴的分布密度、纹饰类型等)存在差异,本次研究结果表明花粉的外壁纹饰可以作为区分玉兰品种的参考依据。
孢粉学的数量聚类图中,供试玉兰品种可以较清晰的划分出白玉兰类群与二乔玉兰类群,说明孢粉学性状可以作为种一级的分类方法,但用于对玉兰品种一级的分类并不十分敏锐,可以作为品种分类辅助的参考标准。
3、建立了玉兰业属16个种和玉兰39个样本37个品种的AFLP指纹图谱和聚类关系图,确定了DICE (Jaccard)和SM算法是分别适用于种和品种的高信度聚类途径。
16个种的聚类结果支持将天目玉兰(M. amoena)和望春玉兰归为同一组；厚叶玉兰(M. crassifolius)和红花玉兰(M. wufengensis)亲缘关系相近,其可能的亲缘关系还需进一步验证；建议将景宁玉兰(M. sinostellata)单独作为种的分类处理,并建议将景宁玉兰、天目玉兰与望春玉兰同归于望春玉兰组；不支持将伏牛山玉兰(Mfuniushanensis)并入望春玉兰及作为宝华玉兰的变种处理；AFLP分子标记的研究结果还显示出以萼片状花被片、花被片颜色作为玉兰亚属分组依据存在欠缺和不足。
39个样品大部分能被较为准确地聚类为白玉兰和二乔玉兰两大类,该结果支持了将种源作为玉兰品种的一级分类标准；聚类结果显示,品种之间的亲缘关系与花型的相关性较小。AFLP聚类结果还显示出同一地区的品种聚于一起,可能的原因是玉兰品种大多由当地习用的种或品种育成。
研究结果提示我们,在充分了解种或品种种质形成渊源的基础上,AFLP技术是开展玉兰亚属种或品种分类鉴定有效的辅助和参考手段。
4、形态及孢粉数量分类法和AFLP技术均能将供试的玉兰品种较为清晰地区分为白玉兰和二乔玉兰两大品种群,因此,三种方法均是开展玉兰亚属种或品种分类、鉴定有效的辅助和参考手段。
5、同花色玉兰品种在孢粉学及AFLP聚类图中大部分能聚于一处,而多瓣类玉兰品种在AFLP聚类图和孢粉学聚类图中则呈间隔分布,表明对于玉兰品种的分类实践来说,花色性状是居于花型(重瓣)性状更高一层次分类等级的位置,但花色、花型性状是否能作为玉兰品种二级分类标准,尚需要更多的品种参与及更深入的研究。
6、玉兰最早在我国2500年前就已开始栽植,并在漫长的栽培历史中形成了独特的玉兰花文化,发展出了包括诗词、绘画和民俗等在内的多种艺术表现形式,成为中国传统花文化的重要组成内容之一。
以上研究为掌握我国玉兰植物资源现状,深入了解其生物学特性以及之间的亲缘关系,完善玉兰业属种和玉兰品种分类系统等提供了可靠的科学数据和理论依据,并为玉兰植物资源保护、收集和新种质创制等工作打下了坚实基础。
Magnolia Subgenus of Yulania is an important traditional flower of China. In this study, We gained the current status of species about subgenus of Yulania and cultivated varieties of Magnolia denudata by conducting detailed on-the-spot investigations of the germplasm resources. AFLP fingerprints were made successfully for some species of subgenus Yulania and some varieties of M. denudata. We worked on the classification of species about subgenus Yulania and varieties of M. denudata,and made discussions about the genetic relationship and the evolutional relationship between them, according to the morphology comparisons among the varieties, in conjunction with palynology researches and AFLP molecular-biology means. The principal conclusions as follows:
1. According the collection and collation work in our study, there were 52 species of subgenus of Yulania on record at present, in which 18 species were authorization approval,24 new species were be published in scientific literature and 10 species were under suspicion. China as the origin center of subgenus of Yulania in the world, as well as one of the present distribution centers, has 48 native species and 4 alien species.
We preparatory acknowledged that there are 40 varieties of M. denudata in China, in which 26 varieties belong to M. denudata, and 14 varieties belong to M.×soulangeana. According to the morphology comparisons among the varieties, we published 11 types of M. denudata which have representative characteristic such as M. denudata'Huangmai'and M. denudata "Yudie" to be as new varieties in this study.
We made a clearly classification that 37 varieties of M. denudata were divided into 5 M. denudata and M.×soulangeana groups alternately using a numerical taxonomic method, the results gave us the ideas that extensive characteristics intersections and gradual characteristic transitions among M. denudata and M.×soulangeana were universal situation, on the other way. they also showed that numerical taxonomic analysis might be a great means in the classification researches about varieties of M. denudata.
The dendrogram of R clustering analysis showed that characteristics of M. denudata varieties were dispersed independently and had poor relationships among them, which suggested that,30 characteristics of M. denudata varieties we choosed for R clustering analysis were correct, quantization treatments of those characteristics were suitable which given a guarantee for the accuracy and scientificity of the results. Accorrding the contributor ratio from principal component analysis，18 characteristics such as thickness of young branches, leaf bud color and perianth color etc. were cognizanced as the most impacted factors in numerical taxonomic analysis, all of those might made a fine base for the classification researches about varieties of M. denudata.
2. Palynology researches of 35 varieties of M. denudata showed that they have similar configuration which their pollen gains were bilateral symmetry, hemimorphic,1-colpate, perprolate(P/E >2), with the size 52.9-61.3μm×21.3-27.6μm, which all belong to pollen gains type N1P3C3, what more, those are the typical characteristic of Subgenus Yulania, but there are still some tiny different were examined about their pollen gains'exine ornamentation (such as the density of perforations and caves, type of exine ornamentation etc.). M. denudata has a cave-shaped exine ornamentation, otherwise, M.×soulangeana. has striate-shaped, helminth-shaped or spine-shaped exine ornamentation. Those results showed that the morphological characteristic of pollen gains exine ornamentation could surely be used as the criteria of classification among varieties of M. denudata.
The clustering result of palynology numerical taxonomic showed that varieties in this research were divided into 2 groups clearly:M. denudata and M×soulangeana, which suggested that palynology might be a fine method for species classification, whereas, it was not so sensitive in variety classification which might be as an assistant method only. 3. AFLP fingerprints and dendrogram with high diversity and rich polymorphism were made successfully for 16 species of subgenus Yulania and 39 varieties of M. denudata, what more, DICE or Jaccard coefficient and SM coefficient have been acknowledged as the more authentic methods for dendrogram computing of species and varieties of subgenus Yulania and M. denudata independently.
The phylogenetics clustering result of 16 species approved of the classification treatment that M.amoena and M.biondii were in the same group.We acknowledged that M.crassifolius was a mutation of M. wufengensis firstly because of that they have a more closer relationship according the clustering result. Our study didn't approve of the classification treatment David made that M. sinostellata was merged into M. tomentosa, we tended to deal with it as a species of subgenus Yulania individually and be as the same section with M.amoena and M.biondii. We didn't approve of the treatment that M funiushanensis to be as a variety of M.zenii and be merged into M.biondii. The results of AFLP suggested that the sepal-like petal and petal color were defective and not enough for devided sections of subgenus Yulania.
The clustering result that 39 varieties were divided into 2 groups:M. denudata and M.×soulangeana, which approve of that provenance might be the first-class standard for variety classification. The clustering also showed that the petal shape had a poor correlation between the similarities among the varities. Varieties from the same areas showed to be clustered together in this study which a conceivable reason might be that they were bred from the same species or varieties of the same areas.
Results above showed that AFLP technique could surely be used as an efficacious assistant and referenced method for the classification and identification researches among species and varieties of Yulania on the base of complete understanding of their germplasm development course.
4. Varieties in our research were divided into 2 groups (M. denudata and M.×soulangeana) clearly, according to morphology numerical taxonomic analysis, palynology numerical taxonomic analysis and AFLP technique, all of these results were approved of the treatment for M. denudata classification nowadays, which suggested that these 3 methods were fine assistant means for species classification of Magnolia Subgenus of Yulania and varieties classification of M. denudata.
5. Varieties with the same petal color were clustering closely in the dendrograms of palynology numerical taxonomic analysis and AFLP analysis, varieties with the multiple petals were clustering scatteredly however, which suggested that petal color were in a higher lever against to petal shape (multiple petals) as the standard of varieties classification, what more, plenty of research works were indispensable to make petal color and as the secondary classification standard in the future.
6. The planting history of Chinese Subgenus Yulania was discovered as early as 2500 years ago, and the culture of Chinese Subgenus Yulania which be as an important content of Chinese traditional flower-culture then has a great development with all kind forms of artistic expression (such as poetry, painting and folk-custom etc).
Our research provided a large amount of; useful information not only for thorough comprehending the present resource situation, the biological characteristics and the genetic relationships of subgenus Yulania, but also for supplying credible science data and theory evidences in improving and perfecting the classification system of subgenus and varieties of M. denudata. What more, it also made a substantial foundation for the following work such as resource protection, collection and new germplasm resource development etc. of subgenus Yulania in China.
1、经调查和整理,目前有记载的玉兰亚属植物共52种,明确认定18种,近来文献提及的新种24个,尚存有争议的10种；我国作为玉兰亚属植物的起源中心与现代分布中心,共有48个本地种(包括文献提及及有争议种,另4个为国外种)。
经调查整理,初步调查到我国白玉兰和二乔玉兰(M×soulangeana)品种40个,其中白玉兰类26个,二乔玉兰类14个。根据形态学比较分析,本研究提出将‘黄脉’和‘玉蝶’等12个具典型特征的玉兰类型暂作为新品种处理。
运用形态学数量聚类分析将其中37个玉兰类型很清晰地划分为呈间隔分布的5个白玉兰和二乔玉兰品种群,在印证了二乔玉兰与白玉兰在许多外部性状上均存在着广泛交叉和逐渐过渡情况的同时,也表明形态学聚类分析是一种开展玉兰品种分类研究较好的方法。R型聚类显示,30个性状的量化处理较为合理,保证了最终研究结果的准确性和科学性。通过主成分分析,根据贡献率大小确定了小枝粗细、叶芽颜色、花被颜色等18个影响较大的性状因子,为以后开展更为系统、深入的中国玉兰品种分类研究提供了参考依据。
2、对37个样本35个玉兰品种的孢粉学研究显示,玉兰品种的花粉形态相似,均为左右对称,异极,具单沟,超长球形(P/E>2),大小(52.9-61.3μm)×(21.3-27.6μm),属于NPC花粉分类系统中的N1P3C3类型,与玉兰业属植物花粉共同的典型特征一致,但表面纹饰的细微形态结构(穿孔和孔穴的分布密度、纹饰类型等)存在差异,本次研究结果表明花粉的外壁纹饰可以作为区分玉兰品种的参考依据。
孢粉学的数量聚类图中,供试玉兰品种可以较清晰的划分出白玉兰类群与二乔玉兰类群,说明孢粉学性状可以作为种一级的分类方法,但用于对玉兰品种一级的分类并不十分敏锐,可以作为品种分类辅助的参考标准。
3、建立了玉兰业属16个种和玉兰39个样本37个品种的AFLP指纹图谱和聚类关系图,确定了DICE (Jaccard)和SM算法是分别适用于种和品种的高信度聚类途径。
16个种的聚类结果支持将天目玉兰(M. amoena)和望春玉兰归为同一组；厚叶玉兰(M. crassifolius)和红花玉兰(M. wufengensis)亲缘关系相近,其可能的亲缘关系还需进一步验证；建议将景宁玉兰(M. sinostellata)单独作为种的分类处理,并建议将景宁玉兰、天目玉兰与望春玉兰同归于望春玉兰组；不支持将伏牛山玉兰(Mfuniushanensis)并入望春玉兰及作为宝华玉兰的变种处理；AFLP分子标记的研究结果还显示出以萼片状花被片、花被片颜色作为玉兰亚属分组依据存在欠缺和不足。
39个样品大部分能被较为准确地聚类为白玉兰和二乔玉兰两大类,该结果支持了将种源作为玉兰品种的一级分类标准；聚类结果显示,品种之间的亲缘关系与花型的相关性较小。AFLP聚类结果还显示出同一地区的品种聚于一起,可能的原因是玉兰品种大多由当地习用的种或品种育成。
研究结果提示我们,在充分了解种或品种种质形成渊源的基础上,AFLP技术是开展玉兰亚属种或品种分类鉴定有效的辅助和参考手段。
4、形态及孢粉数量分类法和AFLP技术均能将供试的玉兰品种较为清晰地区分为白玉兰和二乔玉兰两大品种群,因此,三种方法均是开展玉兰亚属种或品种分类、鉴定有效的辅助和参考手段。
5、同花色玉兰品种在孢粉学及AFLP聚类图中大部分能聚于一处,而多瓣类玉兰品种在AFLP聚类图和孢粉学聚类图中则呈间隔分布,表明对于玉兰品种的分类实践来说,花色性状是居于花型(重瓣)性状更高一层次分类等级的位置,但花色、花型性状是否能作为玉兰品种二级分类标准,尚需要更多的品种参与及更深入的研究。
6、玉兰最早在我国2500年前就已开始栽植,并在漫长的栽培历史中形成了独特的玉兰花文化,发展出了包括诗词、绘画和民俗等在内的多种艺术表现形式,成为中国传统花文化的重要组成内容之一。
以上研究为掌握我国玉兰植物资源现状,深入了解其生物学特性以及之间的亲缘关系,完善玉兰业属种和玉兰品种分类系统等提供了可靠的科学数据和理论依据,并为玉兰植物资源保护、收集和新种质创制等工作打下了坚实基础。
Magnolia Subgenus of Yulania is an important traditional flower of China. In this study, We gained the current status of species about subgenus of Yulania and cultivated varieties of Magnolia denudata by conducting detailed on-the-spot investigations of the germplasm resources. AFLP fingerprints were made successfully for some species of subgenus Yulania and some varieties of M. denudata. We worked on the classification of species about subgenus Yulania and varieties of M. denudata,and made discussions about the genetic relationship and the evolutional relationship between them, according to the morphology comparisons among the varieties, in conjunction with palynology researches and AFLP molecular-biology means. The principal conclusions as follows:
1. According the collection and collation work in our study, there were 52 species of subgenus of Yulania on record at present, in which 18 species were authorization approval,24 new species were be published in scientific literature and 10 species were under suspicion. China as the origin center of subgenus of Yulania in the world, as well as one of the present distribution centers, has 48 native species and 4 alien species.
We preparatory acknowledged that there are 40 varieties of M. denudata in China, in which 26 varieties belong to M. denudata, and 14 varieties belong to M.×soulangeana. According to the morphology comparisons among the varieties, we published 11 types of M. denudata which have representative characteristic such as M. denudata'Huangmai'and M. denudata "Yudie" to be as new varieties in this study.
We made a clearly classification that 37 varieties of M. denudata were divided into 5 M. denudata and M.×soulangeana groups alternately using a numerical taxonomic method, the results gave us the ideas that extensive characteristics intersections and gradual characteristic transitions among M. denudata and M.×soulangeana were universal situation, on the other way. they also showed that numerical taxonomic analysis might be a great means in the classification researches about varieties of M. denudata.
The dendrogram of R clustering analysis showed that characteristics of M. denudata varieties were dispersed independently and had poor relationships among them, which suggested that,30 characteristics of M. denudata varieties we choosed for R clustering analysis were correct, quantization treatments of those characteristics were suitable which given a guarantee for the accuracy and scientificity of the results. Accorrding the contributor ratio from principal component analysis，18 characteristics such as thickness of young branches, leaf bud color and perianth color etc. were cognizanced as the most impacted factors in numerical taxonomic analysis, all of those might made a fine base for the classification researches about varieties of M. denudata.
2. Palynology researches of 35 varieties of M. denudata showed that they have similar configuration which their pollen gains were bilateral symmetry, hemimorphic,1-colpate, perprolate(P/E >2), with the size 52.9-61.3μm×21.3-27.6μm, which all belong to pollen gains type N1P3C3, what more, those are the typical characteristic of Subgenus Yulania, but there are still some tiny different were examined about their pollen gains'exine ornamentation (such as the density of perforations and caves, type of exine ornamentation etc.). M. denudata has a cave-shaped exine ornamentation, otherwise, M.×soulangeana. has striate-shaped, helminth-shaped or spine-shaped exine ornamentation. Those results showed that the morphological characteristic of pollen gains exine ornamentation could surely be used as the criteria of classification among varieties of M. denudata.
The clustering result of palynology numerical taxonomic showed that varieties in this research were divided into 2 groups clearly:M. denudata and M×soulangeana, which suggested that palynology might be a fine method for species classification, whereas, it was not so sensitive in variety classification which might be as an assistant method only. 3. AFLP fingerprints and dendrogram with high diversity and rich polymorphism were made successfully for 16 species of subgenus Yulania and 39 varieties of M. denudata, what more, DICE or Jaccard coefficient and SM coefficient have been acknowledged as the more authentic methods for dendrogram computing of species and varieties of subgenus Yulania and M. denudata independently.
The phylogenetics clustering result of 16 species approved of the classification treatment that M.amoena and M.biondii were in the same group.We acknowledged that M.crassifolius was a mutation of M. wufengensis firstly because of that they have a more closer relationship according the clustering result. Our study didn't approve of the classification treatment David made that M. sinostellata was merged into M. tomentosa, we tended to deal with it as a species of subgenus Yulania individually and be as the same section with M.amoena and M.biondii. We didn't approve of the treatment that M funiushanensis to be as a variety of M.zenii and be merged into M.biondii. The results of AFLP suggested that the sepal-like petal and petal color were defective and not enough for devided sections of subgenus Yulania.
The clustering result that 39 varieties were divided into 2 groups:M. denudata and M.×soulangeana, which approve of that provenance might be the first-class standard for variety classification. The clustering also showed that the petal shape had a poor correlation between the similarities among the varities. Varieties from the same areas showed to be clustered together in this study which a conceivable reason might be that they were bred from the same species or varieties of the same areas.
Results above showed that AFLP technique could surely be used as an efficacious assistant and referenced method for the classification and identification researches among species and varieties of Yulania on the base of complete understanding of their germplasm development course.
4. Varieties in our research were divided into 2 groups (M. denudata and M.×soulangeana) clearly, according to morphology numerical taxonomic analysis, palynology numerical taxonomic analysis and AFLP technique, all of these results were approved of the treatment for M. denudata classification nowadays, which suggested that these 3 methods were fine assistant means for species classification of Magnolia Subgenus of Yulania and varieties classification of M. denudata.
5. Varieties with the same petal color were clustering closely in the dendrograms of palynology numerical taxonomic analysis and AFLP analysis, varieties with the multiple petals were clustering scatteredly however, which suggested that petal color were in a higher lever against to petal shape (multiple petals) as the standard of varieties classification, what more, plenty of research works were indispensable to make petal color and as the secondary classification standard in the future.
6. The planting history of Chinese Subgenus Yulania was discovered as early as 2500 years ago, and the culture of Chinese Subgenus Yulania which be as an important content of Chinese traditional flower-culture then has a great development with all kind forms of artistic expression (such as poetry, painting and folk-custom etc).
Our research provided a large amount of; useful information not only for thorough comprehending the present resource situation, the biological characteristics and the genetic relationships of subgenus Yulania, but also for supplying credible science data and theory evidences in improving and perfecting the classification system of subgenus and varieties of M. denudata. What more, it also made a substantial foundation for the following work such as resource protection, collection and new germplasm resource development etc. of subgenus Yulania in China.
引文
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