不同玉米自交系之间的遗传多样性分析
|
摘要
玉米是重要的粮食作物和饲用作物。我国是世界上第二大玉米生产国,自1996年后,玉米总产量和单位面积产量超过小麦,居禾谷类作物的第二位。从玉米总产发展的速度来看,玉米单产的提高起到不可忽视的作用。长期以来利用杂种优势已经成为增加玉米产量和改善玉米品质的一项重要措施。杂种优势利用是玉米育种研究的重要内容,其理论基础是杂种优势群和杂种优势模式。合理准确地划分杂种优势群,建立相应的杂种优势模式,才能有效地选配杂交组合。杂种优势理论建立在遗传差异之上,因而种质的遗传多样性研究是利用杂种优势的前提,有助于这些种质在育种实践中的有效利用。以往形态标记、系谱分析、配合力表现以及同工酶等方法相继被用于遗传多样性研究,但都表现出一定的局限性。近年来,建立在以DNA结构多态性基础上的分子标记技术克服了传统标记的许多缺点,已发展成为遗传多样性研究的重要工具。本实验采用了两种分子标记,即RAPD(randomamplified polymorphic DNA)分子标记和内转录间隔区(internal transcribed spacer,ITS)序列分析,对12个玉米自交系进行了初步分析。
本实验通过RAPD技术,在获得自交系高纯度总DNA的基础上,从40个随机引物中筛选出10个特异性引物,共扩增产生77条谱带,其中69条表现出多态性(89.6%),说明玉米自交系的基因组之间具有丰富的遗传多样性。根据得到的RAPD指纹图谱建立0、1型数据,利用phylip3.67程序包对所扩增的77个RAPD标记采用NJ法进行聚类分析,得到亲缘关系图。结果表明:除了自交系自330与Mo17亲缘关系较远,而与Hz32的亲缘关系较近,从而被划分到旅大红骨类群,与传统的系谱分析法将其划分到Lancaster类群不一致外,用RAPD标记建立的亲缘关系图与根据系谱来源分析得到的亲缘关系基本一致。
本实验利用直接测序克隆测序两种方法检测12个自交系中部分品种的ITS序列,所得序列利用Clustal W version1.7进行排序,结果表明:不同自交系间的ITS序列高度一致,具有很强的保守性。通过在GenBank中搜索,得到一系列玉米属中的ITS序列以及相关近缘种的18S和28rDNA序列,将其在Clustal W Version 1.7上进行排序对比,结果表明所测玉米自交系ITS序列长度约为505bp,这与报道过的一些玉米种类的ITS序列在592bp～602bp有较大的差异。
Corn is an important food crop and forage crop. China has been the world's second largest corn producer since 1996, corn production and yield per unit area than wheat ranking cereal crops second place. Looking from the corn total output development's speed, the enhancement of the corn per unit area yield can not be ignored。For a long time the use of hybrid vigor has become an important measure of increasing corn production and improving the quality of corn. Heterosis use is an important part of corn breeding research ,which theory is based on the heterotic group and heterosis model. Reasonably accurate division of heterotic group, the establishment of the corresponding heterosis pattern, can be effectively matching combinations. Heterosis theory is based on genetic differences, therefore the genetic diversity of germplasm is the premise of using hybrid vigor, contribute to these germplasm breeding practice in the effective utilization. in the past, form tag, pedigree analysis, Combining ability analysis and isozyme methods have been used to study genetic diversity, but have shown some limitations。In recent years, to establish the structure of DNA polymorphism on the basis of molecular marker technology has overcome many of the shortcomings of traditional markers, genetic diversity has become an important research tool. This study used two molecular markers, RAPD (random amplified polymorphic DNA) marker and ITS(internal transcribed spacer, ITS) sequence marker ,were utilized in analysis of maize inbred lines'pedigree.
This study by RAPD technology, on the basis of extraction of high purity DNA of the inbred lines , selected 10 specific from 40 random primers primers, and amplified a total of 77 bands, of which 69 showed polymorphism (89.6%), indicating that the genome of corn inbred line have a rich genetic diversity. in accordance with the RAPD fingerprint, the 0/1 database was generated ,and a dendrogram was constructed by adopting NJ method. The results showed that 6 clusters identified by RAPD markers confirmed the heterotic groups determined by their pedigree apart from the inbred line Zi330 close to Hz32 and far from Mo17,grouped into clade Lancaster.
This experiment used both the direct sequencing and the clone sequencing to detect some of ITS sequenes in 12 maize inbred lines .and aligned the sequences by Clustal W version1.7.The result showed that: the ITS sequence in the different lines is highly conservative and consistent. Searching through GenBank, a series of maize and related close species of 18 S and 28 rDNA sequences were found, and aligning with the ITS we had, the result showed that the measured corn inbreds ITS sequence length is of about To 505 bp, which has a greater difference compared with some of reported corn ITS sequences in the 592 bp～602bp.
本实验通过RAPD技术,在获得自交系高纯度总DNA的基础上,从40个随机引物中筛选出10个特异性引物,共扩增产生77条谱带,其中69条表现出多态性(89.6%),说明玉米自交系的基因组之间具有丰富的遗传多样性。根据得到的RAPD指纹图谱建立0、1型数据,利用phylip3.67程序包对所扩增的77个RAPD标记采用NJ法进行聚类分析,得到亲缘关系图。结果表明:除了自交系自330与Mo17亲缘关系较远,而与Hz32的亲缘关系较近,从而被划分到旅大红骨类群,与传统的系谱分析法将其划分到Lancaster类群不一致外,用RAPD标记建立的亲缘关系图与根据系谱来源分析得到的亲缘关系基本一致。
本实验利用直接测序克隆测序两种方法检测12个自交系中部分品种的ITS序列,所得序列利用Clustal W version1.7进行排序,结果表明:不同自交系间的ITS序列高度一致,具有很强的保守性。通过在GenBank中搜索,得到一系列玉米属中的ITS序列以及相关近缘种的18S和28rDNA序列,将其在Clustal W Version 1.7上进行排序对比,结果表明所测玉米自交系ITS序列长度约为505bp,这与报道过的一些玉米种类的ITS序列在592bp～602bp有较大的差异。
Corn is an important food crop and forage crop. China has been the world's second largest corn producer since 1996, corn production and yield per unit area than wheat ranking cereal crops second place. Looking from the corn total output development's speed, the enhancement of the corn per unit area yield can not be ignored。For a long time the use of hybrid vigor has become an important measure of increasing corn production and improving the quality of corn. Heterosis use is an important part of corn breeding research ,which theory is based on the heterotic group and heterosis model. Reasonably accurate division of heterotic group, the establishment of the corresponding heterosis pattern, can be effectively matching combinations. Heterosis theory is based on genetic differences, therefore the genetic diversity of germplasm is the premise of using hybrid vigor, contribute to these germplasm breeding practice in the effective utilization. in the past, form tag, pedigree analysis, Combining ability analysis and isozyme methods have been used to study genetic diversity, but have shown some limitations。In recent years, to establish the structure of DNA polymorphism on the basis of molecular marker technology has overcome many of the shortcomings of traditional markers, genetic diversity has become an important research tool. This study used two molecular markers, RAPD (random amplified polymorphic DNA) marker and ITS(internal transcribed spacer, ITS) sequence marker ,were utilized in analysis of maize inbred lines'pedigree.
This study by RAPD technology, on the basis of extraction of high purity DNA of the inbred lines , selected 10 specific from 40 random primers primers, and amplified a total of 77 bands, of which 69 showed polymorphism (89.6%), indicating that the genome of corn inbred line have a rich genetic diversity. in accordance with the RAPD fingerprint, the 0/1 database was generated ,and a dendrogram was constructed by adopting NJ method. The results showed that 6 clusters identified by RAPD markers confirmed the heterotic groups determined by their pedigree apart from the inbred line Zi330 close to Hz32 and far from Mo17,grouped into clade Lancaster.
This experiment used both the direct sequencing and the clone sequencing to detect some of ITS sequenes in 12 maize inbred lines .and aligned the sequences by Clustal W version1.7.The result showed that: the ITS sequence in the different lines is highly conservative and consistent. Searching through GenBank, a series of maize and related close species of 18 S and 28 rDNA sequences were found, and aligning with the ITS we had, the result showed that the measured corn inbreds ITS sequence length is of about To 505 bp, which has a greater difference compared with some of reported corn ITS sequences in the 592 bp～602bp.
引文
陈举林,马冲,邹仁峰等,2001,山东省主要玉米种质基础的现状分析[J].莱阳农学院学报,18(2):112-115
傅骏骅,李连城,刘新芝等,1997,玉米RAPD程序优化研究及其初步探讨[J].作物学报,23(1):56-61
高连明,杨俊波,2002,张长芹等,基于ITS序列分析探讨杜鹃属映山红亚属的组间关系.云南植物研究,24(30):313-320
黄益勤,李建生,2001,利用rflp标记划分45份玉米自交系杂种优势群的研究[J].中国农业科学[J],34(3):244-250
贾继增,1995,分子标记种质资源鉴定和分子标记育种[J].中国农业科学,29(4):1-10
李春香,杨群,2002,PCR产物直接测序还是克隆测序?—密叶杉属rDNAITS序列的测定方法[J].植物学通报,19(6):698-704.
刘宗华,汤继华等,2006,河南省主要玉米品种杂种优势利用模式分析[J].中国农业科学,39(8):1689-1696
刘新芝,彭泽斌,傅骏骅等,1996,RAPD在玉米类群划分研究中的应用.中国农业科学[J],30(3):44-51
刘新芝,彭泽斌,傅骏骅等,1998,采用RAPD分子标记、表型和杂种优势聚类分析法对、玉米自交系类群的划分[J],华北农学报,13(4):36-41
彭泽斌,刘新芝,傅骏骅等,1998,玉米自交系杂种优势类群与杂优模式建构的初步研究[J].作物学报,24(6):711-717
王莉花,殷富有,刘继梅等,2004,利用RAPD分析云南野生荞麦资源的多样性和亲缘关系[J],分子植物育种,2(6):807-815
王琳,1997,国外玉米品种类群杂种优势研究的回顾[J].玉米科学,5(3):78-80
王建波,张文驹,陈家宽,1999,核rDNA的ITS序列在被子植物系统与进化研究中的应用[J].植物分类学报,37(4):407-416.
吴景峰,1995,我国玉米杂交种发展的主要历程、差距和对策[J].玉米科学,3(1):1-5
徐芹,郭东全,刘金根,2005,玉米种质资源系谱研究[J].种子,24(10):44-47
袁力行,傅骏骅,张世煌等,2001,利用RFLP和SSR标记划分玉米自交系杂种优势群的研究[J].作物学报,27(2):149-156
曾三省,1990,中国玉米杂交种的种质基础[J].中国农业科学,23(4):1-9
赵剑峰,杨庆凯,2000,RAPD技术在国内玉米育种中的应用[J].玉米科学, 8(4) :18～19
A. H. Abu Assar, R. Uptmoor, A. A. Abdelmula et al.,2005,Genetic Variation in Sorghum Germplasm from Sudan, ICRISAT, and USA Assessed by Simple Sequence Repeats (SSRs)[J], Crop Sci. 45:1636-1644.
Badaeva E D ,Friebe B ,Gill B S.,1996, Genome differentiation in Aegilops. 2. Physical mapping of 5S and 18S-26S ribosomal RNA gene families in diploid species[J]. Genome, 39:1150-1158
Baldwin B G, Sanderson M J , Porter J M et al., 1995, The ITS region of nuclear ribosomal DNA: a valuable source of evidence on angiosperm phylogeny[J]. Ann Missouri Bot Gard, 82 : 247-277
Brandon S. Gaut et al.,2001,Patterns of Chromosomal Duplication in Maize and Their Implications for Comparative Maps of the Grasses[J], Genome Research,letter:55～66
Chi Shumin et al. 1995, Heterosis Group Analysis of Several Common Corn(Zea Mays L.) Inbred Lines[J], Journal of Hebei Agricultural Uniersity, 18(1), 22-26
Dai Xiaojun, Huang Guangwen, Liang Manzhong,2005,Comparison on ITS Sequences of Nuclear Ribosomal DNA of Hybrid Rices Parents[J]. Life Science Research, 9(2): 163-167
Edward S. Buckler IV and Timothy P. Holtsford,1996,Zea Systematics: Ribosomal ITS Evidence [J],Mol.Biol.Evol.,13(4):612～622
Edward S. Buckler IV and Timothy P. Holtsford,1996,Zea Ribosomal Repeat Evolution and Substitution Patterns[J].Mol.BioI.Evol.,13(4):623～632
Fu, H., Zheng, Z., and Dooner, H.K. 2002. Recombination rates between adjacent genic and retrotransposon regions in maize vary by 2 orders of magnitude[J]. Proc. Natl. Acad. Sci., USA 99:1082-1087.
Gao Jian, Liang Manzhong, Chen Liangbi. 2004,The Analysis Based on DNA Sequence from ITS Ribosomal DNA of Indica and Japonica Subspecies in Rice[J]. Biotechnology, 14(2): 1-3
Gaut, B.S., Le Thierry d'Ennequin, M., Peek, A.S., and Sawkins, M.C,2000,Maize as a model for the evolution of plant nuclear genomes[J]. Proc. Natl. Acad. Sci. 97: 7008-7015.
Haiming Li, Shihuang Zhang, Ruifa Hu,2005,Impacts of genetic uniformity on maize production in China[J]. Biodiversity Science, 13(2): 91-96
Huihua Fu, Wonkeun Park, Xianghe Yan et al.,2001,The highly recombinogenic bz locus lies in anunusually gene-rich region of the maize genome[J],PNAS,98(l 5):8903-8905
John GK. Williams, Anne R.Kubelik, Kenneth J.Livak, et al., 1990,DNA polymorphisms amplified by arbitrary primers are useful as genetic markers[J]. Nuclei Acids Research, 18(22): 6531-6535
Ju" rgen Jobst, Klaus King, Vera Hemleben,1998,Molecular Evolution of the Internal Transcribed Spacers(ITSl and ITS2) and Phylogenetic Relationships among Species of the Family Cucurbitaceae[J], Molecular Phylogenetics and Evolution,9(2):204～219
Kang, K.K., Sano, H., and Kameya, T. 1995,Characterization of cDNAs encoding small GTP-binding proteins from maize[J]. Plant Physiol, 107:275-276.
Kim Y D , Jansen R K, 1996,Phylogenetic implications of rbcL and ITS sequnece variation in the Berberidaceae[J]. Syst Bot, 21(3), 381 ～396
Li Xinhai, Fu Junhua, Zhang Shihuang, 2000, Genetic Variation of Inbred Lines of Maize Detected by SSR Markers[J]. Scientia Agricultural Sinica, 33(2) :1～9
Li Xinhai, Yuan Lixing, Li Xiaohui,2003, Heterotic Grouping of 70 Maize Inbred Line s by SSR Markers[J]. Scientia Agricultural Sinica, 36(6):622～627
Liu Zhonghua,Wang Qingdong et al., 2005,Cluster Analysis and Relation with Others of Elite Maize Inbred Line Yuzi87-1 [J]. Molecular Plant Breeding, 3(4): 525-530
LIU, J. S., and C. L. Schardl,1994, A conserved sequence in internal transcribed spacer-1 of plant nuclear ribosomal RNA genes[J]. Plant Mol. Biol. 26:775-778.
Luo Sulan, He Puchao, Zheng Xueqin,2001,Genetic Diversity in Wild Grapes Native to China Based on Randomly Amplified Polymorphic DNA(RAPD) Analysis[J]. Acta Botanica Sinica, 43(2): 158-163
Lynch,M. , Force,A.,2000, The probability of duplicate gene preservation by subfunctionalization [J]. Genetics ,154:459-473
Ma XiaoJun, Wang XiaoQuan, Xu ZhaoXi ,2000,RAPD Variation Within and Among Populations of Ginseng Cultivars[J]. Acta Botanica Sinica, Vol.42,No.6,586-590
Mao Jianing ,Duan Shihua, LI Shaoqing ,Zhu Yingguo, 2002,Genetic Analysis and Identification of Three Groups Three-line Hybrid Rice and Their Parents by RAPD Markers[J], 24(3):283～287
Masanori Yamasaki,Maud I. Tenaillon, Irie Vroh Bi,2005, A Large-Scale Screen for Artificial Selection in Maize Identifies Candidate Agronomic Loci for Domestication and Crop Improvement[J].The Plant Cell,17:2659-2872
Marianne Barrier, Robert H. Robichaux, and Michael D. Purugganan,2001,Accelerated regulatory gene evolution in an adaptive radiation[J],PNAS,98(18):10208-10213
Nei M, Li WH (1979) Mathematical model for studying genetic variation in terms of restriction endonucleases[J]. Proc Natl.Acad.Sci.,76:5269-5273
Rafalski JA, Tingey SV, Willams JGK ,1991, RAPD markers - a new technology for genetic mapping and plant breeding[J]. Agbiotech News and Info 3:645-648
R. J. Mailer, R. Scarth, B. Fristensky,1994,Discrimination among cultivars of rapeseed (Brassica napus L.) using DNA polymorphisms amplified from arbitrary primers[J], Theor.Appl.Genet. 87:697-704
Rogers S O ,Bendich A.J.,1987, Ribosomal RNA genes in plants :variability in copy number and in the intergenic spacer[J]. Plant Mol.Biol., 9 :509～520
Schultheis L M , Baldwin B G, 1994, Molecular systematics of Fouquieriaceae : evidence from nuclear rDNA ITS studies[J]. Amer J Bot, 81 (supplement, abstract): 199
Smith JSC ,1988,Identification of pedigrees of hybrid maize (Zea mays L.) cultivars by isozyme electrophoresis and reversedphase high-performance liquid chromatography[J], Euphytica, 39:199- 205
Song KM, Osborn TC, Williams PH (1988) Brassica taxonomy based on nuclear restriction fragment length polymorphisms (RFLPs)[J]. Theor Appl Genet,75:784-794
Stephen I. Wright, Irie Vroh Bi, Steve G. Schroeder,2005,The Effects of Artificial Selection on the Maize Genome[J].Science,308:1310～1314
Sun Yizhen et al .,2001, Analysis to Zea Mays Sinesis Inbred Lines Combining Abil ity and to Apply RAPD Technology to Predict the Herterosis of Zea Mays Sinesis[J]. Journal of Beijing Agriculture College, 16(3) 1-12
Susanna A ,Jacas N G,Soltis D E et al. ,1995,Phylogenetic relationships in tribe Cardueae (Asteraceae) based on ITS sequences[J]. Amer.J.Bot. ,82 :1056—1068
Tai TH, Tanksley SD (1990) A rapid and inexpensive method for isolation of total DNA from dehydrated plant tissue[J]. Plant.Mol.Biol., 8:297-303
Tenaillon, M.I., U'Ren, J., Tenaillon, O., and Gaut, B.S.,2004,Selection versus demography: A multilocus investigation of the domestication process in maize[J]. Mol. Biol. Evol. 21:1214-1225.
Wang Xiaoquan, Zou Yuping, Zhang Darning,1996,Problems in the use of RAPD to the study of genetic diversity and systematics[J].Acta Botanica Sinica, 38(12):954～962
Wang Xiaoquan, Hong Deyuan,1997, Progress in Molecular Systematics of Plants in Recent Five Years[J]. Acta Phytotaxonomica Smica,35(5): 465～480
Wang Xiaoquan, Li Zhenyu, 1998,The Application of Sequence Analysis of rDNA Fragment toThe Systematic Study of The Subfamily Cyrtandroideae (Gesneriaceae)[J]. Acta Phytotaxonomica Sinica,36 (2): 97～105
Wang Yibo , Wang Zhenhua, Lu Lixing,1998,Studies on Maize Germplasm Base ,Division of Hetero sis Groups and Utilizing Models of Heterosis in China[J].Maize Science,6(1): 9-15
Welsh J, McClelland M, 1990, Fingerprinting genomes using PCR with arbitrary primers. Nucl Acids Res, 18 :7213 -7218.
White T J, Bruns T, Lee S, Taylor J, 1990, Amplification and direct sequencing of rungal ribosomal RNA genes for phylogenetics. In : Innis M , Gelfand D , Sinisky J, White T eds. PCR Protocols: A Guide to Methods and Applications. San Diego, California : Academic Press. 315～322
Xu fen et al, 2006, Optimization of reaction condition for RAPD analysis in Mermithidae[J]. Journal Of Central China Normal University,Vol.40, No.3, 412-415
Xu Fen, Wang Guoxiu, Zha Yuping ,2005,Genetic Relationships Among Five Species of Mermithidae Based on RAPD[J]. Zoological Research,Vol.26,No.3,294-299
Yao Wenhua,TAN Jing,CHEN Hongmei,2006,The Advances of Studying on Maize Heterotic Group[J].Maize Science, 14(5):30～34
Zeng Jian, Zhang Li, Fan Xing, 2006, Comparison Analysis on Direct Sequencing of PCR Products and Sequencing of Cloning PCR Products of ITS Sequence of Nuclear rDNA in Kengyilia (Poaceae : Triticeae)[J]. Journal of Sichuan Agricultural University,24(4): 369-374
Zheng Shuyun et al., 2006, Heterotic Grouping of Maize Inbred Lines Classified by Using SSR Markers[J]. Journal of Maize Sciences,Vol.14, No.5,26-29
Zhu Jingjing, Jin Liang, Li Xinzheng. In Search of the Molecular Basis of Heterosis in Maize[J]. Biotechnology Bulletin, supplement
Zheng Degang, Jin Yi, Wang Lifeng, 2005,Methods of Classification of Maize Inbred Lines into Heterotic Groups[J]. Journal of Northeast Agricultural University, Vol.36,No.1, 1-4
Zhang Yingpei. Some Issues in Molecular Systematics[J]. Zoological Research, 1994, Vol.l5,No.1, 1-10
Zhou Yi, Zou Yuping, Hong Deyuan, 1996, ITS1 Sequences of Nuclear Ribosomal DNA in Wild Rices and Cultivated Rices of China and Their Phylogenetic Implications[J]. Acta Botanica Sinica ,38(10):785～ 791
傅骏骅,李连城,刘新芝等,1997,玉米RAPD程序优化研究及其初步探讨[J].作物学报,23(1):56-61
高连明,杨俊波,2002,张长芹等,基于ITS序列分析探讨杜鹃属映山红亚属的组间关系.云南植物研究,24(30):313-320
黄益勤,李建生,2001,利用rflp标记划分45份玉米自交系杂种优势群的研究[J].中国农业科学[J],34(3):244-250
贾继增,1995,分子标记种质资源鉴定和分子标记育种[J].中国农业科学,29(4):1-10
李春香,杨群,2002,PCR产物直接测序还是克隆测序?—密叶杉属rDNAITS序列的测定方法[J].植物学通报,19(6):698-704.
刘宗华,汤继华等,2006,河南省主要玉米品种杂种优势利用模式分析[J].中国农业科学,39(8):1689-1696
刘新芝,彭泽斌,傅骏骅等,1996,RAPD在玉米类群划分研究中的应用.中国农业科学[J],30(3):44-51
刘新芝,彭泽斌,傅骏骅等,1998,采用RAPD分子标记、表型和杂种优势聚类分析法对、玉米自交系类群的划分[J],华北农学报,13(4):36-41
彭泽斌,刘新芝,傅骏骅等,1998,玉米自交系杂种优势类群与杂优模式建构的初步研究[J].作物学报,24(6):711-717
王莉花,殷富有,刘继梅等,2004,利用RAPD分析云南野生荞麦资源的多样性和亲缘关系[J],分子植物育种,2(6):807-815
王琳,1997,国外玉米品种类群杂种优势研究的回顾[J].玉米科学,5(3):78-80
王建波,张文驹,陈家宽,1999,核rDNA的ITS序列在被子植物系统与进化研究中的应用[J].植物分类学报,37(4):407-416.
吴景峰,1995,我国玉米杂交种发展的主要历程、差距和对策[J].玉米科学,3(1):1-5
徐芹,郭东全,刘金根,2005,玉米种质资源系谱研究[J].种子,24(10):44-47
袁力行,傅骏骅,张世煌等,2001,利用RFLP和SSR标记划分玉米自交系杂种优势群的研究[J].作物学报,27(2):149-156
曾三省,1990,中国玉米杂交种的种质基础[J].中国农业科学,23(4):1-9
赵剑峰,杨庆凯,2000,RAPD技术在国内玉米育种中的应用[J].玉米科学, 8(4) :18～19
A. H. Abu Assar, R. Uptmoor, A. A. Abdelmula et al.,2005,Genetic Variation in Sorghum Germplasm from Sudan, ICRISAT, and USA Assessed by Simple Sequence Repeats (SSRs)[J], Crop Sci. 45:1636-1644.
Badaeva E D ,Friebe B ,Gill B S.,1996, Genome differentiation in Aegilops. 2. Physical mapping of 5S and 18S-26S ribosomal RNA gene families in diploid species[J]. Genome, 39:1150-1158
Baldwin B G, Sanderson M J , Porter J M et al., 1995, The ITS region of nuclear ribosomal DNA: a valuable source of evidence on angiosperm phylogeny[J]. Ann Missouri Bot Gard, 82 : 247-277
Brandon S. Gaut et al.,2001,Patterns of Chromosomal Duplication in Maize and Their Implications for Comparative Maps of the Grasses[J], Genome Research,letter:55～66
Chi Shumin et al. 1995, Heterosis Group Analysis of Several Common Corn(Zea Mays L.) Inbred Lines[J], Journal of Hebei Agricultural Uniersity, 18(1), 22-26
Dai Xiaojun, Huang Guangwen, Liang Manzhong,2005,Comparison on ITS Sequences of Nuclear Ribosomal DNA of Hybrid Rices Parents[J]. Life Science Research, 9(2): 163-167
Edward S. Buckler IV and Timothy P. Holtsford,1996,Zea Systematics: Ribosomal ITS Evidence [J],Mol.Biol.Evol.,13(4):612～622
Edward S. Buckler IV and Timothy P. Holtsford,1996,Zea Ribosomal Repeat Evolution and Substitution Patterns[J].Mol.BioI.Evol.,13(4):623～632
Fu, H., Zheng, Z., and Dooner, H.K. 2002. Recombination rates between adjacent genic and retrotransposon regions in maize vary by 2 orders of magnitude[J]. Proc. Natl. Acad. Sci., USA 99:1082-1087.
Gao Jian, Liang Manzhong, Chen Liangbi. 2004,The Analysis Based on DNA Sequence from ITS Ribosomal DNA of Indica and Japonica Subspecies in Rice[J]. Biotechnology, 14(2): 1-3
Gaut, B.S., Le Thierry d'Ennequin, M., Peek, A.S., and Sawkins, M.C,2000,Maize as a model for the evolution of plant nuclear genomes[J]. Proc. Natl. Acad. Sci. 97: 7008-7015.
Haiming Li, Shihuang Zhang, Ruifa Hu,2005,Impacts of genetic uniformity on maize production in China[J]. Biodiversity Science, 13(2): 91-96
Huihua Fu, Wonkeun Park, Xianghe Yan et al.,2001,The highly recombinogenic bz locus lies in anunusually gene-rich region of the maize genome[J],PNAS,98(l 5):8903-8905
John GK. Williams, Anne R.Kubelik, Kenneth J.Livak, et al., 1990,DNA polymorphisms amplified by arbitrary primers are useful as genetic markers[J]. Nuclei Acids Research, 18(22): 6531-6535
Ju" rgen Jobst, Klaus King, Vera Hemleben,1998,Molecular Evolution of the Internal Transcribed Spacers(ITSl and ITS2) and Phylogenetic Relationships among Species of the Family Cucurbitaceae[J], Molecular Phylogenetics and Evolution,9(2):204～219
Kang, K.K., Sano, H., and Kameya, T. 1995,Characterization of cDNAs encoding small GTP-binding proteins from maize[J]. Plant Physiol, 107:275-276.
Kim Y D , Jansen R K, 1996,Phylogenetic implications of rbcL and ITS sequnece variation in the Berberidaceae[J]. Syst Bot, 21(3), 381 ～396
Li Xinhai, Fu Junhua, Zhang Shihuang, 2000, Genetic Variation of Inbred Lines of Maize Detected by SSR Markers[J]. Scientia Agricultural Sinica, 33(2) :1～9
Li Xinhai, Yuan Lixing, Li Xiaohui,2003, Heterotic Grouping of 70 Maize Inbred Line s by SSR Markers[J]. Scientia Agricultural Sinica, 36(6):622～627
Liu Zhonghua,Wang Qingdong et al., 2005,Cluster Analysis and Relation with Others of Elite Maize Inbred Line Yuzi87-1 [J]. Molecular Plant Breeding, 3(4): 525-530
LIU, J. S., and C. L. Schardl,1994, A conserved sequence in internal transcribed spacer-1 of plant nuclear ribosomal RNA genes[J]. Plant Mol. Biol. 26:775-778.
Luo Sulan, He Puchao, Zheng Xueqin,2001,Genetic Diversity in Wild Grapes Native to China Based on Randomly Amplified Polymorphic DNA(RAPD) Analysis[J]. Acta Botanica Sinica, 43(2): 158-163
Lynch,M. , Force,A.,2000, The probability of duplicate gene preservation by subfunctionalization [J]. Genetics ,154:459-473
Ma XiaoJun, Wang XiaoQuan, Xu ZhaoXi ,2000,RAPD Variation Within and Among Populations of Ginseng Cultivars[J]. Acta Botanica Sinica, Vol.42,No.6,586-590
Mao Jianing ,Duan Shihua, LI Shaoqing ,Zhu Yingguo, 2002,Genetic Analysis and Identification of Three Groups Three-line Hybrid Rice and Their Parents by RAPD Markers[J], 24(3):283～287
Masanori Yamasaki,Maud I. Tenaillon, Irie Vroh Bi,2005, A Large-Scale Screen for Artificial Selection in Maize Identifies Candidate Agronomic Loci for Domestication and Crop Improvement[J].The Plant Cell,17:2659-2872
Marianne Barrier, Robert H. Robichaux, and Michael D. Purugganan,2001,Accelerated regulatory gene evolution in an adaptive radiation[J],PNAS,98(18):10208-10213
Nei M, Li WH (1979) Mathematical model for studying genetic variation in terms of restriction endonucleases[J]. Proc Natl.Acad.Sci.,76:5269-5273
Rafalski JA, Tingey SV, Willams JGK ,1991, RAPD markers - a new technology for genetic mapping and plant breeding[J]. Agbiotech News and Info 3:645-648
R. J. Mailer, R. Scarth, B. Fristensky,1994,Discrimination among cultivars of rapeseed (Brassica napus L.) using DNA polymorphisms amplified from arbitrary primers[J], Theor.Appl.Genet. 87:697-704
Rogers S O ,Bendich A.J.,1987, Ribosomal RNA genes in plants :variability in copy number and in the intergenic spacer[J]. Plant Mol.Biol., 9 :509～520
Schultheis L M , Baldwin B G, 1994, Molecular systematics of Fouquieriaceae : evidence from nuclear rDNA ITS studies[J]. Amer J Bot, 81 (supplement, abstract): 199
Smith JSC ,1988,Identification of pedigrees of hybrid maize (Zea mays L.) cultivars by isozyme electrophoresis and reversedphase high-performance liquid chromatography[J], Euphytica, 39:199- 205
Song KM, Osborn TC, Williams PH (1988) Brassica taxonomy based on nuclear restriction fragment length polymorphisms (RFLPs)[J]. Theor Appl Genet,75:784-794
Stephen I. Wright, Irie Vroh Bi, Steve G. Schroeder,2005,The Effects of Artificial Selection on the Maize Genome[J].Science,308:1310～1314
Sun Yizhen et al .,2001, Analysis to Zea Mays Sinesis Inbred Lines Combining Abil ity and to Apply RAPD Technology to Predict the Herterosis of Zea Mays Sinesis[J]. Journal of Beijing Agriculture College, 16(3) 1-12
Susanna A ,Jacas N G,Soltis D E et al. ,1995,Phylogenetic relationships in tribe Cardueae (Asteraceae) based on ITS sequences[J]. Amer.J.Bot. ,82 :1056—1068
Tai TH, Tanksley SD (1990) A rapid and inexpensive method for isolation of total DNA from dehydrated plant tissue[J]. Plant.Mol.Biol., 8:297-303
Tenaillon, M.I., U'Ren, J., Tenaillon, O., and Gaut, B.S.,2004,Selection versus demography: A multilocus investigation of the domestication process in maize[J]. Mol. Biol. Evol. 21:1214-1225.
Wang Xiaoquan, Zou Yuping, Zhang Darning,1996,Problems in the use of RAPD to the study of genetic diversity and systematics[J].Acta Botanica Sinica, 38(12):954～962
Wang Xiaoquan, Hong Deyuan,1997, Progress in Molecular Systematics of Plants in Recent Five Years[J]. Acta Phytotaxonomica Smica,35(5): 465～480
Wang Xiaoquan, Li Zhenyu, 1998,The Application of Sequence Analysis of rDNA Fragment toThe Systematic Study of The Subfamily Cyrtandroideae (Gesneriaceae)[J]. Acta Phytotaxonomica Sinica,36 (2): 97～105
Wang Yibo , Wang Zhenhua, Lu Lixing,1998,Studies on Maize Germplasm Base ,Division of Hetero sis Groups and Utilizing Models of Heterosis in China[J].Maize Science,6(1): 9-15
Welsh J, McClelland M, 1990, Fingerprinting genomes using PCR with arbitrary primers. Nucl Acids Res, 18 :7213 -7218.
White T J, Bruns T, Lee S, Taylor J, 1990, Amplification and direct sequencing of rungal ribosomal RNA genes for phylogenetics. In : Innis M , Gelfand D , Sinisky J, White T eds. PCR Protocols: A Guide to Methods and Applications. San Diego, California : Academic Press. 315～322
Xu fen et al, 2006, Optimization of reaction condition for RAPD analysis in Mermithidae[J]. Journal Of Central China Normal University,Vol.40, No.3, 412-415
Xu Fen, Wang Guoxiu, Zha Yuping ,2005,Genetic Relationships Among Five Species of Mermithidae Based on RAPD[J]. Zoological Research,Vol.26,No.3,294-299
Yao Wenhua,TAN Jing,CHEN Hongmei,2006,The Advances of Studying on Maize Heterotic Group[J].Maize Science, 14(5):30～34
Zeng Jian, Zhang Li, Fan Xing, 2006, Comparison Analysis on Direct Sequencing of PCR Products and Sequencing of Cloning PCR Products of ITS Sequence of Nuclear rDNA in Kengyilia (Poaceae : Triticeae)[J]. Journal of Sichuan Agricultural University,24(4): 369-374
Zheng Shuyun et al., 2006, Heterotic Grouping of Maize Inbred Lines Classified by Using SSR Markers[J]. Journal of Maize Sciences,Vol.14, No.5,26-29
Zhu Jingjing, Jin Liang, Li Xinzheng. In Search of the Molecular Basis of Heterosis in Maize[J]. Biotechnology Bulletin, supplement
Zheng Degang, Jin Yi, Wang Lifeng, 2005,Methods of Classification of Maize Inbred Lines into Heterotic Groups[J]. Journal of Northeast Agricultural University, Vol.36,No.1, 1-4
Zhang Yingpei. Some Issues in Molecular Systematics[J]. Zoological Research, 1994, Vol.l5,No.1, 1-10
Zhou Yi, Zou Yuping, Hong Deyuan, 1996, ITS1 Sequences of Nuclear Ribosomal DNA in Wild Rices and Cultivated Rices of China and Their Phylogenetic Implications[J]. Acta Botanica Sinica ,38(10):785～ 791