基于REMAP和IRAP分子标记的梅（Prunus mume Sieb. et Zucc.）遗传多样性分析
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摘要
梅(Prunus mume Sieb. et Zucc.)原产中国,是我国著名观赏和加工保健食用的特产树种。其栽培历史悠久,种质资源丰富。逆转座子通过RNA为中介进行转座,在植物基因组中分布广,拷贝数多,异质高,在种内和种间表现出较高的序列差异性和丰富的插入多态性,引起稳定突变,基于这些特点开发出的几种分子标记与常规分子标记比较,显示出基因组覆盖面广、多态性丰富的优点。本研究通过表型数量性状分类与DNA分子标记的梅遗传多样性特征分析,探讨梅种质资源的遗传多样性和亲缘关系,为今后有效开展种质资源的开发及创新利用提供理论依据。主要研究结果如下：
1.基于重要表型性状数量分类的梅遗传多样性分析
以南京农业大学国家梅种质资源圃保存的84个梅品种为试材,运用SPSS 11.5对叶花果等31个重要表型性状进行Q聚类和主成分及因子分析,84个梅品种分为5组,其中‘长农17’等13个果梅品种为第一组；‘红梅’等30个果梅品种为第二组；‘重叶梅’等15个花梅品种为第三组；‘银红朱砂’等9个花梅品种为第四组；‘小绿萼’等17个花梅品种为第五组。果梅品种与花梅品种分别聚类。果梅中的‘透骨红’介于花梅和果梅之间。31个表型性状中果实光亮度、果实黄蓝偏差、果实横径、单果重、果实侧径、单核重、花瓣数、花瓣层数、a果、果实纵径共10个性状在梅品种数量分类中起重要作用。2.基于REMAP标记技术的梅遗传多样性分析
通过DNA纯化方法、5因素4水平L16(45)的完全随机正交试验、不同浓度PAGE胶、引物3’的碱基等的研究,建立了适合梅REMAP标记的技术体系。体系总体积25μl：基因组DNA40 ng、2.5 mmol/L MgCl2 1.5μL、2.5 mmol/L dNTPs 2.0μL、10×Buffer 2.5μL、TaqDNA聚合酶1.5 U、10μmol/LLTR引物1.0μL、10μmol/L SSR引物1.0 gL,加ddH2O至25μL。以一次裂解,抽提后加RNase获取的DNA质量好；REMAP体系的SSR引物的3’端加碱基明显提高REMAP的扩增效果,但碱基数量无明显影响。浓度为5%的PAGE胶电泳条带数量多、清晰地反映品种间的多态性,且制胶方便,不易碎胶。
筛选出5对LTR-SSR引物组合,用于84个梅品种的REMAP遗传多样性分析,共产生122个多态性位点。5对引物组合的香浓信息指数I在0.3897-0.6364之间,Nei's遗传多样性指数He在0.2395-0.4455之间；对REMAP扩增结果进行UPGMA聚类,在相似系数0.766处,84个梅品种可分为18组,果梅与花梅分别聚类；‘小叶猪肝’与‘大叶猪肝’呈现出明显的多态性,‘小叶猪肝’在198 bp和600 bp出现多态性位点。用PopGen32分析梅遗传多样性,推测日本果梅可能引自中国浙江,日本花梅引自中国江苏；花瓣3层是梅花瓣数量引起遗传变异的起点,3-5层间遗传基本稳定,果梅花瓣1层、花梅花瓣5层具有较明显的遗传多样性。品种间的遗传距离,84个梅品种中,最近的是‘双粉垂枝’与‘粉皮宫粉’为0.059,最远的是‘云南红梅’与‘大粒’为0.630；果梅品种中,最近的是‘黄小大’与‘福建白粉’为0.104,最远的是‘四月梅’与‘信侬小梅’为0.487；花梅品种中,最近的是‘双粉垂枝’与‘粉皮宫粉’为0.059,最远的是‘中玉宫粉’与‘云南丰后’为0.556。基因流强度的群体每代迁移数梅品种群最大。3.基于IRAP标记技术的梅遗传多样性分析
通过5因素4水平L16(45)的完全随机正交试验、不同浓度PAGE胶等的研究,建立了适合梅IRAP标记的技术体系。体系总体积25μ1：基因组DNA30ng,2.5 mmol/L MgCl2 2.0μL,2.5 mmol/L dNTPs 2.5μL,10×Buffer 2.5μL, Taq DNA聚合酶1.0 U,10μmol/L LTR引物1.0μL,加ddH20至25μL。6%和8%浓度的非变性PAGE胶适于IRAP多态性检测。
筛选出6条LTR引物,用于84个梅品种的IRAP遗传多样性分析,共产生99个多态性位点。6条引物的香浓信息指数I在0.3359-0.5807,Nei's遗传多样性指数He在0.1967-0.3976之间；对IRAP扩增结果进行UPGMA聚类,在相似系数0.766处,84个梅品种聚为41组,果梅的‘长农17’与花梅宫粉型的‘粉皮宫粉’、‘香雪宫粉’、‘云南红梅’、‘小玉宫粉’聚一起,花梅的‘美人梅’与果梅的‘信侬小梅’聚一起,其余都是果梅与花梅分别聚类。日本果梅品种的‘玉英’、‘白加贺’、‘月世界’、‘古城’、‘节田梅’、‘花香实’聚在一起。‘大叶猪肝’与‘小叶猪肝’不聚在一组。应用PopGen32分析了梅遗传多样性,日本、江苏、浙江品种群亲缘关系最近,推测日本果梅引自中国浙江,日本花梅引自中国江苏；花瓣3层是梅花瓣数量变异引起遗传变异的起点,3-5层间遗传基本稳定。84个梅品种的遗传距离,最近的是‘白加贺’和‘月世界’为0.107,也是果梅中最近的,最远的是‘甲州小梅’和‘骨红垂枝’为0.683；果梅品种中最远的是‘东山李梅’和‘横核’为0.588；花梅品种中,最近的是‘香雪宫粉’和‘云南红梅’为0.118,最远的是‘云南丰后’与‘骨红垂枝’为0.663。
4.基于REMAP-IRAP分子标记的梅遗传多样性分析
综合REMAP和IRAP进行梅遗传多样性分析,梅群体的观测等位基因数1.9955,有效等位基因数为1.4887,Nei's遗传多样性为0.2910,Shannon信息指数为0.4465。UPGMA聚类,84个梅品种在相似系数0.766处被分为30组；在相似系数0.752处,被分成20组。果梅与花梅分别聚类,说明果梅与花梅间基因渗透较少。日本果梅亲缘关系较近的有‘白加贺’、‘月世界’与‘古城’；‘养老3号’与‘莺宿’；‘甲州最小’、‘玉英’与‘花香实’；青梅分组聚一起,但有部分红梅聚在青梅一起,白梅与青梅接近；‘大叶猪肝’与‘小叶猪肝’较远。花梅品种聚类中,朱砂型紧密在一组,垂枝型梅松散聚一组,玉碟型(除‘龙泉玉蝶’)在一组；春后、绿萼类型的品种群分散聚在一起,宫粉型分为3组,表明遗传背景复杂,进化程度不一。PopGen32分析,推测日本果梅品种群引自中国浙江,日本花梅品种群引自中国江苏。梅花瓣数3层为多层明显变化的层数,1、2层的遗传基因与3-5层有明显的不同,3-5层基本稳定。各品种间的遗传距离,梅品种中最近的是‘白加贺’与‘月世界’为0.136,最远的为‘丰后’与‘横核’为0.602；果梅品种中,最近的是‘白加贺’与‘月世界’为0.136,最远的是‘横核’与‘东山李梅’为0.464；花梅品种中,最近的是‘粉皮宫粉’与‘香雪宫粉’及‘银红朱砂’与‘南京红须’为0.141,最远的是‘骨红垂枝’与‘云南丰后’为0.508。
Mei(Prunus mume Sieb. et Zucc.), native to China, is famous for viewing and processing of special healthy food species. The cultivation has a long history and rich germplasm resource. Retrotransposons transpose through an RNA intermediate by a'copy and paste'mechanism and widely distribute and copy in the genome with high heterogeneous populations. It shows high sequence difference and rich insert polymorphism in one or multiple different species which exhibits stable mutation. Compared with normal molecular marker technique, retrotransposons polymorphism molecular marker developed according to above-mentioned characteristics shows the benefits of genome-wide coverage and rich polymorphism. This study aims to explore the genetic diversity, kinship and to provide theoretical basis for effective development and innovation of germplasm resources from the DNA molecule level of germplasm. The results are as follows:
1. The genetic diversity of mei analysis based on the mathematic classification of important morphologic characters
84 mei varieties (including 43 fruiting mei and 41 flowering mei) located in National mei germplasm pool of Nanjing Agricultural University were taken as test materials. Q clustering, principal component and factor analysis to 31 important morphologic characters such as leaves, flower and fruit with SPSS 11.5,84 mei varieties were divided into 5 groups, including 13 fruiting mei for the first group such as'Changnong17' ,30 fruiting mei for the second group such as'Hongmei' ,15 flowering mei for the third group such as 'Chongyemei',9 flowering mei for the fourth group such as'Yinhongzhusha',17 flowering mei for the fifth group such as'Xiaolve'. Fruiting mei and flowering mei clustered respectively. The'Touguhong'of fruiting mei gaultheria preference flowering mei.10 morphologic characters among 31 morphologic characters, like L-fruit, b-fruit, fruit across diameter, weight of per fruit, fruit side diameter, stone weight, petal numbers, petal layers, a-fruit and fruit high play an important role to mei mathematic classification.
2. The genetic diversity analysis of mei based on REMAP marker
Through studying on DNA purification methods, five factors L16 (45) 4 levels completely random orthogonal test, the various PAGE concentrations, base of primer 3', and so on, the final optimal system of REMAP marker technique was established. REMAP system was totally in 25μl, composed of template DNA 40 ng,2.5 mmol/L MgCl2 1.5μL, 2.5 mmol/L dNTPs 2.0μL,10×Buffer 2.5μL, Taq DNA polymerase 1.5 U,10μmol/L LTR primer 1.0μL,10μmol/L SSR primer 1.0μL. Once cleavage was the best of the four treatments in quality of template DNA. The selective base at 3'-end of SSR primer influenced on REMAP notably. The best was one base on 3'-end of SSR primer.5% PAGE was the best concentration of non-denaturing polyacrylamide gel, showing the most and clearest electrophoresis line and could present genetic diversity and had the characteristic of convenience and no crash.5 couples of LTR-SSR primers are screened out for REMAP genetic diversity analysis to 84 mei varieties and produced a total of 122 polymorphisms. For 5 couples of primers, the Shannon's information index was from 0.3897 to 0.6364, Nei's gene diversity from 0.2395 to 0.4455. UPGMA clustering to REMAP amplification results,84 mei varieties can be divided into 18 groups at the similarity coefficient of 0.766, fruiting mei and flowering mei were clustered in different group. 'Xiaoyezhugan' was not in the same cluster with 'Dayezhugan' because of distinct genetic differentiation. 'Xiaoyezhugan' had spesical bands of 198bp and 600bp bands under REMAP, which may related to the fact that retrotransposon insertion causing the differentiation. It was proved by PopGen32 that Japanese fruiting mei ware introduced from Zhejiang province and Japanese flowering mei from Jiangsu province of China. Three layers of petal was the jumping-off point for genetic variation,3-5 layers genetic stability. One layer of petal in fruiting mei and 5 layers in flowering mei had obvious genetic diversity. Among 84 mei varieties, the least of genetic distance was 0.059 between'Shuangfencuizhi'and 'Fenpigongfen'and the farmost one was 0.63 between'Yunnanhongmei'and'Dali'. Among fruiting mei, the least one was 0.104 between'Huangxiaoda'and'Fujianbaifen'and the farmost one was 0.487 between'Siyuemei'and'Koumeshinano'. Among flowering mei, the least one was 0.059 between'Shuangfencuizhi'and'Fenpigongfen'and the farmost one was 0.556 between'Zhongyugongfen'and'Yunnanfenghou'. The genetic flower strength group transfer number for varieties group of mei was the largest, which illustrated varieties group of mei had genetic diversity in abundance.
3. The genetic diversity of mei analysis based on IRAP marker
Through five factors four levels L16 (45) completely random orthogonal test, IRAP molecular marker system was established for mei. IRAP system was total in 25μl, composed of template DNA 30 ng,2.5 mmol/L MgCl2 2.0μL,2.5 mmol/L dNTPs 2.5μL, 10×Buffer 2.5μL, Taq DNA polymerase 1.0 U,10μmol/L LTR primer 1.0μL.6% and 8% PAGE were the better concentration of non-denaturing polyacrylamide gel and could present genetic diversity. Six LTR primers were screened out for IRAP genetic diversity analysis to 84 mei and produced a total of 99 polymorphisms, and the average polymorphism ratio was 83.20%. For 6 primers, the Shannon's information index was in the range from 0.3359 to 0.5807, Nei's gene diversity in the range from 0.1967 to 0.3976. UPGMA clustering to IRAP amplification results,84 mei could be divided into 41 groups at the similarity coefficient of 0.766.'Changnong17'of fruiting mei was clustered together with'Fenpigongfen','Xiangxuegongfen','Yunnanhongmei'and'Xiaoyugongfen', Pink double form of flowering mei and'Meirenmei'of flowering mei was clustered together with'Koumeshinano'. The rest fruiting mei and flowering mei were clustered in different group, respectively. Among Japanese fruiting mei,'Gyokouei','Shirokaga','Gessekai', 'Gojirou','Setsuda'and'Hanakami'were clustered together.'Dayezhugan'was not clustered in the same group with'Xiaoyezhugan'. By PopGen32, Japanese mei was the most similar to Zhejiang mei and Jiangsu mei. It could be confered Japanese fruiting mei was introduced from Zhejiang province in China and Japanese flowering mei was introduced from Jiangsu province in China. Three layers of petal was the jumping-off point for genetic variation,3-5 layers genetic stability. Within 84 mei varieties, the least of genetic distance was 0.107 between'Shirokaga'and'Gessekai'and the farmost one was 0.683 between'Koume'and'Benishidale'. Within fruiting mei, the least one was 0.107 between'Shirokaga'and'Gessekai'and the farmost one was 0.588 between 'Dongshanlimei'and'Henghe'. Within flowering mei, the least one was 0.118 between 'Xiangxuegongfen'and'Yunnanhongmei'and the farmost one was 0.663 between 'Yunnanfenghou'and'Benishidale'.
4. The genetic diversity of analysis based on the REMAP and IRAP molecular markers
The genetic diversity analysis combined REMAP with IRAP technique showed that the observed number of alleles was 1.9955 in mei populations, effective number of alleles was 1.4887, Nei's gene diversity was 0.2910, Shannon's information index was 0.4465. UPGMA cluster, at the similarity coefficient of 0.766,84 mei varieties could be divided into 30 groups, and 20 groups at the similarity coefficient of 0.752. Fruiting mei and flowering mei were clustered respectively showed less genes penetration occur between fruiting mei and flowering mei. Among Japanese fruiting mei,'Shirokaga','Gessekai' and'Gojirou','Yourou3' and 'Gyokouei','Koume','Gyokouei' and 'Hanakami' were similar. Green mei gather together as groups, but part of red mei were assembled into green mei, white mei was similar to green mei, but 'Dayezhugan' is not similar to group with 'Xiaoyezhugan' . Within flowering groups, cinnabar purple form was tightly as the same group, pendant form was loosely as one group, albo-Plena Form was (except for 'Longquanyudie' ) one group, after-spring form and green calyx form as one group separately, pink double form assembles as three groups which showed the complexity of inheritance and evolution extent was different. PopGen32 analysis showed Japanese fruiting mei were introduced from Zhejiang province of China and Japanese flowering mei from Jiangsu province of China. Three layers mei club was obvious change for multilayer ones, gene of 1 layer and 2 layer had big different from 3-5 layer ones. Within 84 mei varieties, the least of genetic distance was 0.136 between 'Shirokaga' and 'Gessekai' and the farmost one was 0.602 between 'Fenghou' and 'Henghe' . Within fruiting mei, the least one was 0.136 between 'Shirokaga' and 'Gessekai' and the farmost one was 0.464 between 'Dongshanlimei' and 'Henghe' . Within flowering mei, the least one was 0.141 between 'Xiangxuegongfen' and 'Fenpigongfen' & 'Yinhongzhusha' and 'Nanjinghongxu' and the farmost one was 0.508 between 'Benishidale' and 'Yunnanfenghou' .
1.基于重要表型性状数量分类的梅遗传多样性分析
以南京农业大学国家梅种质资源圃保存的84个梅品种为试材,运用SPSS 11.5对叶花果等31个重要表型性状进行Q聚类和主成分及因子分析,84个梅品种分为5组,其中‘长农17’等13个果梅品种为第一组；‘红梅’等30个果梅品种为第二组；‘重叶梅’等15个花梅品种为第三组；‘银红朱砂’等9个花梅品种为第四组；‘小绿萼’等17个花梅品种为第五组。果梅品种与花梅品种分别聚类。果梅中的‘透骨红’介于花梅和果梅之间。31个表型性状中果实光亮度、果实黄蓝偏差、果实横径、单果重、果实侧径、单核重、花瓣数、花瓣层数、a果、果实纵径共10个性状在梅品种数量分类中起重要作用。2.基于REMAP标记技术的梅遗传多样性分析
通过DNA纯化方法、5因素4水平L16(45)的完全随机正交试验、不同浓度PAGE胶、引物3’的碱基等的研究,建立了适合梅REMAP标记的技术体系。体系总体积25μl：基因组DNA40 ng、2.5 mmol/L MgCl2 1.5μL、2.5 mmol/L dNTPs 2.0μL、10×Buffer 2.5μL、TaqDNA聚合酶1.5 U、10μmol/LLTR引物1.0μL、10μmol/L SSR引物1.0 gL,加ddH2O至25μL。以一次裂解,抽提后加RNase获取的DNA质量好；REMAP体系的SSR引物的3’端加碱基明显提高REMAP的扩增效果,但碱基数量无明显影响。浓度为5%的PAGE胶电泳条带数量多、清晰地反映品种间的多态性,且制胶方便,不易碎胶。
筛选出5对LTR-SSR引物组合,用于84个梅品种的REMAP遗传多样性分析,共产生122个多态性位点。5对引物组合的香浓信息指数I在0.3897-0.6364之间,Nei's遗传多样性指数He在0.2395-0.4455之间；对REMAP扩增结果进行UPGMA聚类,在相似系数0.766处,84个梅品种可分为18组,果梅与花梅分别聚类；‘小叶猪肝’与‘大叶猪肝’呈现出明显的多态性,‘小叶猪肝’在198 bp和600 bp出现多态性位点。用PopGen32分析梅遗传多样性,推测日本果梅可能引自中国浙江,日本花梅引自中国江苏；花瓣3层是梅花瓣数量引起遗传变异的起点,3-5层间遗传基本稳定,果梅花瓣1层、花梅花瓣5层具有较明显的遗传多样性。品种间的遗传距离,84个梅品种中,最近的是‘双粉垂枝’与‘粉皮宫粉’为0.059,最远的是‘云南红梅’与‘大粒’为0.630；果梅品种中,最近的是‘黄小大’与‘福建白粉’为0.104,最远的是‘四月梅’与‘信侬小梅’为0.487；花梅品种中,最近的是‘双粉垂枝’与‘粉皮宫粉’为0.059,最远的是‘中玉宫粉’与‘云南丰后’为0.556。基因流强度的群体每代迁移数梅品种群最大。3.基于IRAP标记技术的梅遗传多样性分析
通过5因素4水平L16(45)的完全随机正交试验、不同浓度PAGE胶等的研究,建立了适合梅IRAP标记的技术体系。体系总体积25μ1：基因组DNA30ng,2.5 mmol/L MgCl2 2.0μL,2.5 mmol/L dNTPs 2.5μL,10×Buffer 2.5μL, Taq DNA聚合酶1.0 U,10μmol/L LTR引物1.0μL,加ddH20至25μL。6%和8%浓度的非变性PAGE胶适于IRAP多态性检测。
筛选出6条LTR引物,用于84个梅品种的IRAP遗传多样性分析,共产生99个多态性位点。6条引物的香浓信息指数I在0.3359-0.5807,Nei's遗传多样性指数He在0.1967-0.3976之间；对IRAP扩增结果进行UPGMA聚类,在相似系数0.766处,84个梅品种聚为41组,果梅的‘长农17’与花梅宫粉型的‘粉皮宫粉’、‘香雪宫粉’、‘云南红梅’、‘小玉宫粉’聚一起,花梅的‘美人梅’与果梅的‘信侬小梅’聚一起,其余都是果梅与花梅分别聚类。日本果梅品种的‘玉英’、‘白加贺’、‘月世界’、‘古城’、‘节田梅’、‘花香实’聚在一起。‘大叶猪肝’与‘小叶猪肝’不聚在一组。应用PopGen32分析了梅遗传多样性,日本、江苏、浙江品种群亲缘关系最近,推测日本果梅引自中国浙江,日本花梅引自中国江苏；花瓣3层是梅花瓣数量变异引起遗传变异的起点,3-5层间遗传基本稳定。84个梅品种的遗传距离,最近的是‘白加贺’和‘月世界’为0.107,也是果梅中最近的,最远的是‘甲州小梅’和‘骨红垂枝’为0.683；果梅品种中最远的是‘东山李梅’和‘横核’为0.588；花梅品种中,最近的是‘香雪宫粉’和‘云南红梅’为0.118,最远的是‘云南丰后’与‘骨红垂枝’为0.663。
4.基于REMAP-IRAP分子标记的梅遗传多样性分析
综合REMAP和IRAP进行梅遗传多样性分析,梅群体的观测等位基因数1.9955,有效等位基因数为1.4887,Nei's遗传多样性为0.2910,Shannon信息指数为0.4465。UPGMA聚类,84个梅品种在相似系数0.766处被分为30组；在相似系数0.752处,被分成20组。果梅与花梅分别聚类,说明果梅与花梅间基因渗透较少。日本果梅亲缘关系较近的有‘白加贺’、‘月世界’与‘古城’；‘养老3号’与‘莺宿’；‘甲州最小’、‘玉英’与‘花香实’；青梅分组聚一起,但有部分红梅聚在青梅一起,白梅与青梅接近；‘大叶猪肝’与‘小叶猪肝’较远。花梅品种聚类中,朱砂型紧密在一组,垂枝型梅松散聚一组,玉碟型(除‘龙泉玉蝶’)在一组；春后、绿萼类型的品种群分散聚在一起,宫粉型分为3组,表明遗传背景复杂,进化程度不一。PopGen32分析,推测日本果梅品种群引自中国浙江,日本花梅品种群引自中国江苏。梅花瓣数3层为多层明显变化的层数,1、2层的遗传基因与3-5层有明显的不同,3-5层基本稳定。各品种间的遗传距离,梅品种中最近的是‘白加贺’与‘月世界’为0.136,最远的为‘丰后’与‘横核’为0.602；果梅品种中,最近的是‘白加贺’与‘月世界’为0.136,最远的是‘横核’与‘东山李梅’为0.464；花梅品种中,最近的是‘粉皮宫粉’与‘香雪宫粉’及‘银红朱砂’与‘南京红须’为0.141,最远的是‘骨红垂枝’与‘云南丰后’为0.508。
Mei(Prunus mume Sieb. et Zucc.), native to China, is famous for viewing and processing of special healthy food species. The cultivation has a long history and rich germplasm resource. Retrotransposons transpose through an RNA intermediate by a'copy and paste'mechanism and widely distribute and copy in the genome with high heterogeneous populations. It shows high sequence difference and rich insert polymorphism in one or multiple different species which exhibits stable mutation. Compared with normal molecular marker technique, retrotransposons polymorphism molecular marker developed according to above-mentioned characteristics shows the benefits of genome-wide coverage and rich polymorphism. This study aims to explore the genetic diversity, kinship and to provide theoretical basis for effective development and innovation of germplasm resources from the DNA molecule level of germplasm. The results are as follows:
1. The genetic diversity of mei analysis based on the mathematic classification of important morphologic characters
84 mei varieties (including 43 fruiting mei and 41 flowering mei) located in National mei germplasm pool of Nanjing Agricultural University were taken as test materials. Q clustering, principal component and factor analysis to 31 important morphologic characters such as leaves, flower and fruit with SPSS 11.5,84 mei varieties were divided into 5 groups, including 13 fruiting mei for the first group such as'Changnong17' ,30 fruiting mei for the second group such as'Hongmei' ,15 flowering mei for the third group such as 'Chongyemei',9 flowering mei for the fourth group such as'Yinhongzhusha',17 flowering mei for the fifth group such as'Xiaolve'. Fruiting mei and flowering mei clustered respectively. The'Touguhong'of fruiting mei gaultheria preference flowering mei.10 morphologic characters among 31 morphologic characters, like L-fruit, b-fruit, fruit across diameter, weight of per fruit, fruit side diameter, stone weight, petal numbers, petal layers, a-fruit and fruit high play an important role to mei mathematic classification.
2. The genetic diversity analysis of mei based on REMAP marker
Through studying on DNA purification methods, five factors L16 (45) 4 levels completely random orthogonal test, the various PAGE concentrations, base of primer 3', and so on, the final optimal system of REMAP marker technique was established. REMAP system was totally in 25μl, composed of template DNA 40 ng,2.5 mmol/L MgCl2 1.5μL, 2.5 mmol/L dNTPs 2.0μL,10×Buffer 2.5μL, Taq DNA polymerase 1.5 U,10μmol/L LTR primer 1.0μL,10μmol/L SSR primer 1.0μL. Once cleavage was the best of the four treatments in quality of template DNA. The selective base at 3'-end of SSR primer influenced on REMAP notably. The best was one base on 3'-end of SSR primer.5% PAGE was the best concentration of non-denaturing polyacrylamide gel, showing the most and clearest electrophoresis line and could present genetic diversity and had the characteristic of convenience and no crash.5 couples of LTR-SSR primers are screened out for REMAP genetic diversity analysis to 84 mei varieties and produced a total of 122 polymorphisms. For 5 couples of primers, the Shannon's information index was from 0.3897 to 0.6364, Nei's gene diversity from 0.2395 to 0.4455. UPGMA clustering to REMAP amplification results,84 mei varieties can be divided into 18 groups at the similarity coefficient of 0.766, fruiting mei and flowering mei were clustered in different group. 'Xiaoyezhugan' was not in the same cluster with 'Dayezhugan' because of distinct genetic differentiation. 'Xiaoyezhugan' had spesical bands of 198bp and 600bp bands under REMAP, which may related to the fact that retrotransposon insertion causing the differentiation. It was proved by PopGen32 that Japanese fruiting mei ware introduced from Zhejiang province and Japanese flowering mei from Jiangsu province of China. Three layers of petal was the jumping-off point for genetic variation,3-5 layers genetic stability. One layer of petal in fruiting mei and 5 layers in flowering mei had obvious genetic diversity. Among 84 mei varieties, the least of genetic distance was 0.059 between'Shuangfencuizhi'and 'Fenpigongfen'and the farmost one was 0.63 between'Yunnanhongmei'and'Dali'. Among fruiting mei, the least one was 0.104 between'Huangxiaoda'and'Fujianbaifen'and the farmost one was 0.487 between'Siyuemei'and'Koumeshinano'. Among flowering mei, the least one was 0.059 between'Shuangfencuizhi'and'Fenpigongfen'and the farmost one was 0.556 between'Zhongyugongfen'and'Yunnanfenghou'. The genetic flower strength group transfer number for varieties group of mei was the largest, which illustrated varieties group of mei had genetic diversity in abundance.
3. The genetic diversity of mei analysis based on IRAP marker
Through five factors four levels L16 (45) completely random orthogonal test, IRAP molecular marker system was established for mei. IRAP system was total in 25μl, composed of template DNA 30 ng,2.5 mmol/L MgCl2 2.0μL,2.5 mmol/L dNTPs 2.5μL, 10×Buffer 2.5μL, Taq DNA polymerase 1.0 U,10μmol/L LTR primer 1.0μL.6% and 8% PAGE were the better concentration of non-denaturing polyacrylamide gel and could present genetic diversity. Six LTR primers were screened out for IRAP genetic diversity analysis to 84 mei and produced a total of 99 polymorphisms, and the average polymorphism ratio was 83.20%. For 6 primers, the Shannon's information index was in the range from 0.3359 to 0.5807, Nei's gene diversity in the range from 0.1967 to 0.3976. UPGMA clustering to IRAP amplification results,84 mei could be divided into 41 groups at the similarity coefficient of 0.766.'Changnong17'of fruiting mei was clustered together with'Fenpigongfen','Xiangxuegongfen','Yunnanhongmei'and'Xiaoyugongfen', Pink double form of flowering mei and'Meirenmei'of flowering mei was clustered together with'Koumeshinano'. The rest fruiting mei and flowering mei were clustered in different group, respectively. Among Japanese fruiting mei,'Gyokouei','Shirokaga','Gessekai', 'Gojirou','Setsuda'and'Hanakami'were clustered together.'Dayezhugan'was not clustered in the same group with'Xiaoyezhugan'. By PopGen32, Japanese mei was the most similar to Zhejiang mei and Jiangsu mei. It could be confered Japanese fruiting mei was introduced from Zhejiang province in China and Japanese flowering mei was introduced from Jiangsu province in China. Three layers of petal was the jumping-off point for genetic variation,3-5 layers genetic stability. Within 84 mei varieties, the least of genetic distance was 0.107 between'Shirokaga'and'Gessekai'and the farmost one was 0.683 between'Koume'and'Benishidale'. Within fruiting mei, the least one was 0.107 between'Shirokaga'and'Gessekai'and the farmost one was 0.588 between 'Dongshanlimei'and'Henghe'. Within flowering mei, the least one was 0.118 between 'Xiangxuegongfen'and'Yunnanhongmei'and the farmost one was 0.663 between 'Yunnanfenghou'and'Benishidale'.
4. The genetic diversity of analysis based on the REMAP and IRAP molecular markers
The genetic diversity analysis combined REMAP with IRAP technique showed that the observed number of alleles was 1.9955 in mei populations, effective number of alleles was 1.4887, Nei's gene diversity was 0.2910, Shannon's information index was 0.4465. UPGMA cluster, at the similarity coefficient of 0.766,84 mei varieties could be divided into 30 groups, and 20 groups at the similarity coefficient of 0.752. Fruiting mei and flowering mei were clustered respectively showed less genes penetration occur between fruiting mei and flowering mei. Among Japanese fruiting mei,'Shirokaga','Gessekai' and'Gojirou','Yourou3' and 'Gyokouei','Koume','Gyokouei' and 'Hanakami' were similar. Green mei gather together as groups, but part of red mei were assembled into green mei, white mei was similar to green mei, but 'Dayezhugan' is not similar to group with 'Xiaoyezhugan' . Within flowering groups, cinnabar purple form was tightly as the same group, pendant form was loosely as one group, albo-Plena Form was (except for 'Longquanyudie' ) one group, after-spring form and green calyx form as one group separately, pink double form assembles as three groups which showed the complexity of inheritance and evolution extent was different. PopGen32 analysis showed Japanese fruiting mei were introduced from Zhejiang province of China and Japanese flowering mei from Jiangsu province of China. Three layers mei club was obvious change for multilayer ones, gene of 1 layer and 2 layer had big different from 3-5 layer ones. Within 84 mei varieties, the least of genetic distance was 0.136 between 'Shirokaga' and 'Gessekai' and the farmost one was 0.602 between 'Fenghou' and 'Henghe' . Within fruiting mei, the least one was 0.136 between 'Shirokaga' and 'Gessekai' and the farmost one was 0.464 between 'Dongshanlimei' and 'Henghe' . Within flowering mei, the least one was 0.141 between 'Xiangxuegongfen' and 'Fenpigongfen' & 'Yinhongzhusha' and 'Nanjinghongxu' and the farmost one was 0.508 between 'Benishidale' and 'Yunnanfenghou' .
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