中国木瓜属观赏品种调查和分类研究
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摘要
木瓜属(Chaenomeles)植物既是著名的观赏花木,又是我国的传统中药,其栽培历史悠久,种质资源丰富,深受人们喜爱,但其品种分类一直比较混乱。本文对我国主要栽培区的木瓜属观赏品种进行了系统研究,野外调查根据编制的木瓜属植物品种调查表进行性状记载,并采集标本,拍摄照片;根据比较形态学方法,在综合分析木瓜属植物品种性状变异和演化规律的基础上,提出分类原则和依据,按照国际栽培植物命名法规ICNCP进行品种分类和整理,并在泰安市建立了种质资源圃,收集木瓜属观赏品种37个。主要研究结果如下:
(1)在分析形态演化和变异规律的基础上,制定了描述术语规范;按照自然演化的规律,植物一般是从木本进化为草本,乔木进化为灌木,在木瓜属内,木瓜为小乔木,木瓜海棠为大灌木至小乔木,贴梗海棠为灌木,而倭海棠为低矮灌木,傲大贴梗海棠是贴梗海棠和倭海棠的杂交种,介于二者之间,因此,我们认为种间的演化规律为,木瓜最原始,其次为木瓜海棠、贴梗海棠、傲大贴梗海棠、倭海棠。
(2)对品种分类原则和分类等级进行了探讨,提出了木瓜属观赏品种的分类方案,并对我国木瓜属观赏品种进行了系统分类和整理,详细记载了各品种的形态特征,共记载木瓜属观赏品种37个,其中贴梗海棠品种9个,包括‘红艳’、‘秀美’、‘多彩’、‘风扬’、‘凤凰木’、‘夕照’、‘红星’、‘沂红’、‘沂锦’;倭海棠品种5个,包括‘单粉’、‘单白’、‘矮红’、‘日落’、‘四季红’;木瓜品种5个,包括‘粗皮剩花’、‘豆青’、‘小狮子头’、‘细皮剩花’、‘玉兰’;木瓜海棠品种7个,包括‘长俊’、‘红霞’、‘金陵粉’、‘罗扶’、‘一品香’、‘醉杨妃’、‘蜀红’;傲大贴梗海棠品种11个,包括‘早春’、‘碧雪’、‘长寿乐’、‘猩红与金黄’、‘大富贵’、‘复长寿’、‘红宝石’、‘绿宝石’、‘沂橙’、‘红舞’、‘紫衣’;命名了7个新品种,‘风扬’、‘夕照’、‘沂红’、‘沂橙’、‘红舞’、‘紫衣’、‘单粉’;根据ICNCP对品种名称进行了考证,对品种分类和命名违反国际栽培植物命名法规的3个品种‘红贴梗海棠’、‘报春’、‘四季贴梗海棠’进行了订正,现分别命名为‘红艳’、‘早春’、‘四季红’;对同名异物和同物异名的品种进行了考证,将‘复色海棠’、‘复色贴梗海棠’归入‘多彩’,将‘东洋锦’归入‘猩红与金黄’,将‘世界一’归入‘大富贵’,将‘长寿冠’归入‘红宝石’,将‘银长寿’归入‘绿宝石’;根据调查,对于一些品种的植物学归属做了相应的调整,将王嘉祥等(2004)放入贴梗海棠的‘长俊’、‘红霞’、‘罗扶’等改隶于木瓜海棠,将王永慧(2004)放入木瓜海棠的‘绿宝石’、‘长寿乐’改隶于傲大贴梗海棠,将胡忠惠(2004)放入木瓜海棠的‘红宝石’改隶于傲大贴梗海棠。
(3)对木瓜属植物19个品种的花粉亚显微形态进行了扫描电镜观察。结果表明,其花粉属于N3P4C3型,为辐射对称的长球形至超长球形,大小为极轴长48.64 um-39.95 um、赤道轴长25.28 um-20.97 um,外壁纹饰为条纹状或绉波状纹饰,具覆盖层穿孔。花粉亚显微形态对于木瓜属品种分类具有很大价值,根据花粉形态编制了品种分类检索表。聚类分析结果表明,贴梗海棠、倭海棠和傲大贴梗海棠的亲缘关系最近,而木瓜与其他种类的亲缘关系疏远,这与本属的经典分类结果一致。
(4)选取木瓜属18个栽培品种和4个近缘种的24个形态学性状,利用SPSS10.0软件进行聚类分析,探讨了品种间的亲缘关系。Q型聚类分析显示供试样本可分为2类,‘玉兰’和‘豆青’聚合为一类,与其他种类和品种关系疏远;其余种类和品种聚合成一类。第2类分为4个亚类,贴梗海棠、木瓜海棠、倭海棠以及5个杂交品种分别聚合。根据聚类结果,‘猩红与金黄’和‘碧雪’等品种的分类地位值得进一步探讨。数量分类结果能够反映品种之间的亲缘关系。R型聚类结果表明,各分类性状之间没有明显的逻辑关系,24个形态学性状对品种的演化具有较独立的意义。主成分分析表明,前6个主成分的累计贡献率达85.9%,根据其与性状间的相关性可知,影响力较大的性状有花径、花梗颜色、花瓣形状、花瓣宽、瓣爪长、幼叶颜色、花托和果实形状、重瓣性和花瓣数目等。
Chaenomeles is not only a well-known ornamental flower, but also a traditional Chinese medicine plant; they have a long cultural history and rich germplasm resources, which is loved deeply by people. But the cultivar classification system of Chaenomeles is not very clear. A cultivar investigation was conducted at the China's major plants cultivated areas. Record the traits according to the investigation form of Chaenomeles cultivars, collecting the samples, and taking photographs. In accordance with comparative morphology, in the comprehensive analysis of variation and evolution of traits on the basis of principles and to classify basis, in accordance with international cultivated plants named regulations ICNCP and compiling a species classification, and in Tai'an City establishmented a garden germplasm resource, including 37 cultivars are recognized, the main results are summarized as follows:
1. A criteria and terminology for describing cultivars are revised based on a morphologic analysis of the variation and evolutionary trend. In accordance with the laws of natural evolution, the plants generally evolutde from woody to herb, and from tree to shrub, C. sinensis are trees, C. cathayensis are large shrubs or small trees, C. speciosa are shrubs, and C. japonica are low shrubs, C.×superba= C. japonica×C. speciosa, between the two species, therefore, we believe that the evolution of species is, C. sinensis is the most primitive, and next is C. cathayensis, C. speciosa, C.×superba, C. japonica.
2. Discussed the Chaenomeles cultivars base on approaches of principies and ranks for cultivar classification. Detailed records the morphological characteristics of all species, 37 cultivars are recognized, including 9 cultivars in C. speciosa, such as‘Hongyan’,‘Moerloosei’, 'Toyo Nishiki' etc; 5 cultivars in C. japonica, such as‘Danbai’,‘Siji Hong’,‘Danfen’etc; 7 cultivars in C. cathayensis, such as‘Changjun’,‘Hongxia’,‘Jinling Fen’etc; 5 cultivars in C. sinensis, such as‘Cupi Shenghua’,‘Douqing’,‘Xiao Shizitou’,‘Xipi Shenghua’etc; 11 cultivars in C.×superba, such as‘Zaochun’,‘Bixue’,‘Changshou Le’etc; 7 cultivars names are recorded and described as new, such as‘Xizhao’,‘Yihong’,‘Yijin’etc; According to ICNCP, revised 3 Chaenomeles cultivars, and also revised some Chaenomeles cultivars which of the same name bur not the same cultivars or a cultivar but has more than one name; According to the survey, adjusting some cultivars the corresponding species.
3 The pollen grains of 19 cultivars in Chaenomeles were observed under SEM. The conclusion showed that they all belonged to N3P4C3 type, and were of prolate or perprolate and of radio symmetric. The size varied within (48.64-39.95) um×(25.28-20.97) um. The ornamentation is regulated or striate, having pores in the tectum. Pollen morphology is valuable in cultivar classification in this genus. A key was compiled for cultivar classification based on pollen morphology difference. The result with clustering analysis suggested the close relationship among C. speciosa, C. japonica and C.×superba, and the alienation between C. sinensis and the other 4 species, which was consistent with traditional classification result.
4 Based on twenty-four morphological characteristics, eighteen cultivars and four related wild species of flowering quince (Chaenomeles) were investigated systematically and analyzed by cluster analysis with SPSS10.0 program in order to determine their relationships. The Q-cluster analysis showed that the twenty-two specimen were classified into two groups. The first group contains C. sinensis‘Yulan’and C. sinensis‘Douqing’, and the others were categorized into another group, reflecting the alienation between C. sinensis and the other species; the second group could be divided into four subgroups, and every cultivar could be categorized into the right species. Cultivars‘Crimson and Gold’and‘Bixue’, however, should be transferred into C. speciosa and C. japonica respectively. The differences among cultivars in different species of Chaenomeles were significant with a high level of diversity. The numerical taxonomic result could reflect cultivar relationships. According to R-cluster analysis, the selection of testing characteristics was correct, and no characteristics had obvious logical relationship with one another. The twenty-four morphological characteristics had independent effect on the cultivar evolution. The principal component analysis showed that the 24 characteristics were integrated into 6 principal components and their accumulative contributor ratio amounted to 85.9%, and some major discriminating characteristics were selected as flower diameter, pedicel color, petal shape, petal width, petal claw length, young leaf color, shapes of receptacle and fruit, double flower and petal number, etc.
(1)在分析形态演化和变异规律的基础上,制定了描述术语规范;按照自然演化的规律,植物一般是从木本进化为草本,乔木进化为灌木,在木瓜属内,木瓜为小乔木,木瓜海棠为大灌木至小乔木,贴梗海棠为灌木,而倭海棠为低矮灌木,傲大贴梗海棠是贴梗海棠和倭海棠的杂交种,介于二者之间,因此,我们认为种间的演化规律为,木瓜最原始,其次为木瓜海棠、贴梗海棠、傲大贴梗海棠、倭海棠。
(2)对品种分类原则和分类等级进行了探讨,提出了木瓜属观赏品种的分类方案,并对我国木瓜属观赏品种进行了系统分类和整理,详细记载了各品种的形态特征,共记载木瓜属观赏品种37个,其中贴梗海棠品种9个,包括‘红艳’、‘秀美’、‘多彩’、‘风扬’、‘凤凰木’、‘夕照’、‘红星’、‘沂红’、‘沂锦’;倭海棠品种5个,包括‘单粉’、‘单白’、‘矮红’、‘日落’、‘四季红’;木瓜品种5个,包括‘粗皮剩花’、‘豆青’、‘小狮子头’、‘细皮剩花’、‘玉兰’;木瓜海棠品种7个,包括‘长俊’、‘红霞’、‘金陵粉’、‘罗扶’、‘一品香’、‘醉杨妃’、‘蜀红’;傲大贴梗海棠品种11个,包括‘早春’、‘碧雪’、‘长寿乐’、‘猩红与金黄’、‘大富贵’、‘复长寿’、‘红宝石’、‘绿宝石’、‘沂橙’、‘红舞’、‘紫衣’;命名了7个新品种,‘风扬’、‘夕照’、‘沂红’、‘沂橙’、‘红舞’、‘紫衣’、‘单粉’;根据ICNCP对品种名称进行了考证,对品种分类和命名违反国际栽培植物命名法规的3个品种‘红贴梗海棠’、‘报春’、‘四季贴梗海棠’进行了订正,现分别命名为‘红艳’、‘早春’、‘四季红’;对同名异物和同物异名的品种进行了考证,将‘复色海棠’、‘复色贴梗海棠’归入‘多彩’,将‘东洋锦’归入‘猩红与金黄’,将‘世界一’归入‘大富贵’,将‘长寿冠’归入‘红宝石’,将‘银长寿’归入‘绿宝石’;根据调查,对于一些品种的植物学归属做了相应的调整,将王嘉祥等(2004)放入贴梗海棠的‘长俊’、‘红霞’、‘罗扶’等改隶于木瓜海棠,将王永慧(2004)放入木瓜海棠的‘绿宝石’、‘长寿乐’改隶于傲大贴梗海棠,将胡忠惠(2004)放入木瓜海棠的‘红宝石’改隶于傲大贴梗海棠。
(3)对木瓜属植物19个品种的花粉亚显微形态进行了扫描电镜观察。结果表明,其花粉属于N3P4C3型,为辐射对称的长球形至超长球形,大小为极轴长48.64 um-39.95 um、赤道轴长25.28 um-20.97 um,外壁纹饰为条纹状或绉波状纹饰,具覆盖层穿孔。花粉亚显微形态对于木瓜属品种分类具有很大价值,根据花粉形态编制了品种分类检索表。聚类分析结果表明,贴梗海棠、倭海棠和傲大贴梗海棠的亲缘关系最近,而木瓜与其他种类的亲缘关系疏远,这与本属的经典分类结果一致。
(4)选取木瓜属18个栽培品种和4个近缘种的24个形态学性状,利用SPSS10.0软件进行聚类分析,探讨了品种间的亲缘关系。Q型聚类分析显示供试样本可分为2类,‘玉兰’和‘豆青’聚合为一类,与其他种类和品种关系疏远;其余种类和品种聚合成一类。第2类分为4个亚类,贴梗海棠、木瓜海棠、倭海棠以及5个杂交品种分别聚合。根据聚类结果,‘猩红与金黄’和‘碧雪’等品种的分类地位值得进一步探讨。数量分类结果能够反映品种之间的亲缘关系。R型聚类结果表明,各分类性状之间没有明显的逻辑关系,24个形态学性状对品种的演化具有较独立的意义。主成分分析表明,前6个主成分的累计贡献率达85.9%,根据其与性状间的相关性可知,影响力较大的性状有花径、花梗颜色、花瓣形状、花瓣宽、瓣爪长、幼叶颜色、花托和果实形状、重瓣性和花瓣数目等。
Chaenomeles is not only a well-known ornamental flower, but also a traditional Chinese medicine plant; they have a long cultural history and rich germplasm resources, which is loved deeply by people. But the cultivar classification system of Chaenomeles is not very clear. A cultivar investigation was conducted at the China's major plants cultivated areas. Record the traits according to the investigation form of Chaenomeles cultivars, collecting the samples, and taking photographs. In accordance with comparative morphology, in the comprehensive analysis of variation and evolution of traits on the basis of principles and to classify basis, in accordance with international cultivated plants named regulations ICNCP and compiling a species classification, and in Tai'an City establishmented a garden germplasm resource, including 37 cultivars are recognized, the main results are summarized as follows:
1. A criteria and terminology for describing cultivars are revised based on a morphologic analysis of the variation and evolutionary trend. In accordance with the laws of natural evolution, the plants generally evolutde from woody to herb, and from tree to shrub, C. sinensis are trees, C. cathayensis are large shrubs or small trees, C. speciosa are shrubs, and C. japonica are low shrubs, C.×superba= C. japonica×C. speciosa, between the two species, therefore, we believe that the evolution of species is, C. sinensis is the most primitive, and next is C. cathayensis, C. speciosa, C.×superba, C. japonica.
2. Discussed the Chaenomeles cultivars base on approaches of principies and ranks for cultivar classification. Detailed records the morphological characteristics of all species, 37 cultivars are recognized, including 9 cultivars in C. speciosa, such as‘Hongyan’,‘Moerloosei’, 'Toyo Nishiki' etc; 5 cultivars in C. japonica, such as‘Danbai’,‘Siji Hong’,‘Danfen’etc; 7 cultivars in C. cathayensis, such as‘Changjun’,‘Hongxia’,‘Jinling Fen’etc; 5 cultivars in C. sinensis, such as‘Cupi Shenghua’,‘Douqing’,‘Xiao Shizitou’,‘Xipi Shenghua’etc; 11 cultivars in C.×superba, such as‘Zaochun’,‘Bixue’,‘Changshou Le’etc; 7 cultivars names are recorded and described as new, such as‘Xizhao’,‘Yihong’,‘Yijin’etc; According to ICNCP, revised 3 Chaenomeles cultivars, and also revised some Chaenomeles cultivars which of the same name bur not the same cultivars or a cultivar but has more than one name; According to the survey, adjusting some cultivars the corresponding species.
3 The pollen grains of 19 cultivars in Chaenomeles were observed under SEM. The conclusion showed that they all belonged to N3P4C3 type, and were of prolate or perprolate and of radio symmetric. The size varied within (48.64-39.95) um×(25.28-20.97) um. The ornamentation is regulated or striate, having pores in the tectum. Pollen morphology is valuable in cultivar classification in this genus. A key was compiled for cultivar classification based on pollen morphology difference. The result with clustering analysis suggested the close relationship among C. speciosa, C. japonica and C.×superba, and the alienation between C. sinensis and the other 4 species, which was consistent with traditional classification result.
4 Based on twenty-four morphological characteristics, eighteen cultivars and four related wild species of flowering quince (Chaenomeles) were investigated systematically and analyzed by cluster analysis with SPSS10.0 program in order to determine their relationships. The Q-cluster analysis showed that the twenty-two specimen were classified into two groups. The first group contains C. sinensis‘Yulan’and C. sinensis‘Douqing’, and the others were categorized into another group, reflecting the alienation between C. sinensis and the other species; the second group could be divided into four subgroups, and every cultivar could be categorized into the right species. Cultivars‘Crimson and Gold’and‘Bixue’, however, should be transferred into C. speciosa and C. japonica respectively. The differences among cultivars in different species of Chaenomeles were significant with a high level of diversity. The numerical taxonomic result could reflect cultivar relationships. According to R-cluster analysis, the selection of testing characteristics was correct, and no characteristics had obvious logical relationship with one another. The twenty-four morphological characteristics had independent effect on the cultivar evolution. The principal component analysis showed that the 24 characteristics were integrated into 6 principal components and their accumulative contributor ratio amounted to 85.9%, and some major discriminating characteristics were selected as flower diameter, pedicel color, petal shape, petal width, petal claw length, young leaf color, shapes of receptacle and fruit, double flower and petal number, etc.
引文
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