北方地区地方品种牛线粒体DNA多态性和亲缘关系研究
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摘要
线粒体DNA D-Loop环区是动物线粒体的调控区和唯一的非编码区,它是线粒体基因组中序列和长度变异最大的区域。D-loop的碱基替换率比mtDNA分子的其他区域高5～10倍,是核单拷贝基因的25～100倍。细胞色素B(Cyt B)是线粒体呼吸链上电子传递的细胞色素之一,它包含8个跨膜区的螺旋,通过膜内或膜间区域相连结,哺乳动物线粒体细胞色素B(Cyt B)是构成线粒体氧化磷酸化系统复合体Ⅲ的蛋白质之一,也是其中唯一由线粒体基因组编码的蛋白质。
本试验以延边黄牛、鲁西黄牛、草原红牛和日本黑牛为研究对象,根据牛线粒体DNA非编码区D-loop环全长序列和细胞色素B (Cyt B)基因部分编码区序列,结合生物信息学方法,检测了北方地区地方品种牛的线粒体D-Loop和细胞色素B基因的部分序列的多态性。通过DNAsp、MEGA3.1等分子进化软件分析发现,在11头日本黑牛群体中,存在9种D-loop单倍型,3种Cyt B单倍型,8头鲁西黄牛发现7种D-loop单倍型,2种Cyt B单倍型,11头草原红牛发现11种D-loop单倍型,5种Cyt B单倍型,10头延边黄牛9种D-loop单倍型,3种Cyt B单倍型。Kimura双参数遗传距离分析和构建分子进化树结果表明日本黑牛与延边黄牛、草原红牛存在较近的亲缘关系,而与鲁西黄牛亲缘关系较远,这与鲁西牛是由北方蒙古利亚牛系和南方瘤牛的混血种,鲁西牛与东南亚牛种亲缘关系较近的研究结论基本一致。
日本黑牛489号个体与鲁西黄牛中D-loop单倍型H18、H19、H21有较近的亲缘关系。
Mitochondron is unique extranuclear heredity factor in mammal and also the center of energy metabolism, which can biosynthesis fatty acid and some protein.
Mitochondrial D-loop region is the unique noncoding region, which have maximal variation in mitochondrial genome, including sequence composition and length. Cytochrome B (CytB) is in charge of transferring electron on mitochondrial respiratory chain. It is also the unique coding protein in mitochondrial genome. In field of phylogenic evolution research, it is always an available molecular labeling. More and more mammalian phylogenesis was analyzed with CytB gene polymorphism recently.
Mitochondrial DNA polymorphisms contributed to nucleotide and number variation that existed in mitochondrial DNA hypervariable region. There are five types of mutation, including transition, transversion, insertion, deletion and the coexistance of transition and transvertion.
The kimura 2-parameter distances and base substitution, composition, variation were analyzed with MEGA software, the percentage differences among them were calculated with DNAstar software, and the polymorphic sites were analyzed with DNAsp software. The phylogenic trees constructed by three methods (UPGMA-tree,NJ-tree,ME-tree) which supported almost the same topology. The topology structures of different phylogenic trees were similar and discrepancies are not significant. It indicated that phylogenic trees wereaccurate.
The polymorphism of D-loop complete sequence and CytB gene partial sequence with northern cattle strains including Luxi yellow, YanBian yellow, CaoYuan red and alien Japan black cattle were studied in this research. The relationship of four populations was analyzed with bioinformation methods.
The results of mtDNA D-loop(CytB) polymorphism analysis showed that there were 7(2) haplotypes in 8 Luxi Yellow, the average haplotype diversity was 0.964±0.006 (0.536±0.123); 9(3) haplotypes in 11 Japan Black, the average haplotype diversity was 0.964±0.003(0.473±0.162); 9(3) haplotypes in 10 YanBian yellow, the average haplotype diversity was 0.978±0.003 (0.182±0.144); 11(5) haplotypes in 11 CaoYuan red, the average haplotype diversity was 1.000±0.005(0.644±0.152).
D-loop Kimura 2-parameter distance between breeds were 0.0162, 0.0190, 0.0196, 0.0308, 0.0333, and 0.0354 respectively. By pairwise comparison, the closest distance was 0.0162 between Japan Black and YanBian yellow; the farest distance was 0.0354 between Luxi yellow and YanBian yellow. The phylogenic analysis revealed that the Luxi yellow cattle is the farest relation to the other breeds, the Japan black is more closely related to the YanBian yellow than to that of the other breeds.
Cyt B Kimura 2-parameter distance between breeds is 0.0025, 0.0035, 0.0040, 0.0063, 0.0077, and 0.0078 respectively. By pairwise comparison,the Luxi yellow is also the most farthest relation to the other breeds. Japan black cattle 489,463 is special compared with the other cattle intraspecies. It suggests that maybe they are hybrid individuals.
本试验以延边黄牛、鲁西黄牛、草原红牛和日本黑牛为研究对象,根据牛线粒体DNA非编码区D-loop环全长序列和细胞色素B (Cyt B)基因部分编码区序列,结合生物信息学方法,检测了北方地区地方品种牛的线粒体D-Loop和细胞色素B基因的部分序列的多态性。通过DNAsp、MEGA3.1等分子进化软件分析发现,在11头日本黑牛群体中,存在9种D-loop单倍型,3种Cyt B单倍型,8头鲁西黄牛发现7种D-loop单倍型,2种Cyt B单倍型,11头草原红牛发现11种D-loop单倍型,5种Cyt B单倍型,10头延边黄牛9种D-loop单倍型,3种Cyt B单倍型。Kimura双参数遗传距离分析和构建分子进化树结果表明日本黑牛与延边黄牛、草原红牛存在较近的亲缘关系,而与鲁西黄牛亲缘关系较远,这与鲁西牛是由北方蒙古利亚牛系和南方瘤牛的混血种,鲁西牛与东南亚牛种亲缘关系较近的研究结论基本一致。
日本黑牛489号个体与鲁西黄牛中D-loop单倍型H18、H19、H21有较近的亲缘关系。
Mitochondron is unique extranuclear heredity factor in mammal and also the center of energy metabolism, which can biosynthesis fatty acid and some protein.
Mitochondrial D-loop region is the unique noncoding region, which have maximal variation in mitochondrial genome, including sequence composition and length. Cytochrome B (CytB) is in charge of transferring electron on mitochondrial respiratory chain. It is also the unique coding protein in mitochondrial genome. In field of phylogenic evolution research, it is always an available molecular labeling. More and more mammalian phylogenesis was analyzed with CytB gene polymorphism recently.
Mitochondrial DNA polymorphisms contributed to nucleotide and number variation that existed in mitochondrial DNA hypervariable region. There are five types of mutation, including transition, transversion, insertion, deletion and the coexistance of transition and transvertion.
The kimura 2-parameter distances and base substitution, composition, variation were analyzed with MEGA software, the percentage differences among them were calculated with DNAstar software, and the polymorphic sites were analyzed with DNAsp software. The phylogenic trees constructed by three methods (UPGMA-tree,NJ-tree,ME-tree) which supported almost the same topology. The topology structures of different phylogenic trees were similar and discrepancies are not significant. It indicated that phylogenic trees wereaccurate.
The polymorphism of D-loop complete sequence and CytB gene partial sequence with northern cattle strains including Luxi yellow, YanBian yellow, CaoYuan red and alien Japan black cattle were studied in this research. The relationship of four populations was analyzed with bioinformation methods.
The results of mtDNA D-loop(CytB) polymorphism analysis showed that there were 7(2) haplotypes in 8 Luxi Yellow, the average haplotype diversity was 0.964±0.006 (0.536±0.123); 9(3) haplotypes in 11 Japan Black, the average haplotype diversity was 0.964±0.003(0.473±0.162); 9(3) haplotypes in 10 YanBian yellow, the average haplotype diversity was 0.978±0.003 (0.182±0.144); 11(5) haplotypes in 11 CaoYuan red, the average haplotype diversity was 1.000±0.005(0.644±0.152).
D-loop Kimura 2-parameter distance between breeds were 0.0162, 0.0190, 0.0196, 0.0308, 0.0333, and 0.0354 respectively. By pairwise comparison, the closest distance was 0.0162 between Japan Black and YanBian yellow; the farest distance was 0.0354 between Luxi yellow and YanBian yellow. The phylogenic analysis revealed that the Luxi yellow cattle is the farest relation to the other breeds, the Japan black is more closely related to the YanBian yellow than to that of the other breeds.
Cyt B Kimura 2-parameter distance between breeds is 0.0025, 0.0035, 0.0040, 0.0063, 0.0077, and 0.0078 respectively. By pairwise comparison,the Luxi yellow is also the most farthest relation to the other breeds. Japan black cattle 489,463 is special compared with the other cattle intraspecies. It suggests that maybe they are hybrid individuals.
引文
[1] 赵兴波,李宁,吴常信. 线粒体 DNA 指纹技术的研究及其应用[J].高技术通讯,2004,5:94
[2] 刘振湘,唐晓玲.我国家养动物多样性与畜牧业持续发展.畜牧与兽医[J]. 1997,177～180
[3] 陈振文等. 用 DNA 指纹技术分析 WISTAR 大鼠的遗传距离.浙江畜牧兽医[J].1996,21(3):11～12
[4] Anderson S , de Bruijn M H , Coulson A R , Eperon I C , Sanger F , Young I G. Complete sequence of bovine mitochondrial DNA. Conserved features of the mammalian mitochondrial genome. J Mol Biol[J].1982,156 (4) :683～717
[5] 廖顺尧,鲁成. 动物线粒体基因组研究进展.生物化学与生物物理进展[J].2000,27(5):508～511
[6] Wolstenholme D R, Int Rev Cytol[M]. 1992, 141:173
[7] Kikkawa Y, Yonekawa H , Suzuki H , Amano T. Analysis of genetic diversity of domestic water buffaloes and anoas based on variations in the mitochondrial gene for cytochrome b. Anim Genet[J]. 1997.28 :195～201
[8] Weir B.S.(徐云碧等译). 遗传学数据分析—群体遗传学离散型数据分析方法[M]. 北京:中国农业出版社,1996,184～196
[9] Masatoshi Nei, Sudhir Kumar. 分子进化与系统发育[M]. 北京:高等教育出版社. 2004,5:20～180
[10] 丁波.DNA指纹技术及其在畜禽遗传育种中的应用.青海畜牧兽医杂志[J]. 1994,(4):42～44
[11] Cao,Y.,J.Adachi and M.Hasegawa.(1998)Comment on the quarter puzzlingmethod for finding maximum-likelihood tree topologiesMol.Bio.Evol[J].15:87～89
[12] 徐其放,陈嘉昌,朱世杰,申煜,刘运忠,常弘. 动物线粒体DNA 的特异结构及应用分子系统学分析的方法. 中国比较医学杂志[J]. 2005,10,15(5):515～518
[13] 张 丽,侯水生,刘小林,徐铁山. DNA 指纹技术分析畜禽亲缘关系的原理及应用.家畜生态[J]. 2003,24(4):48～50
[14] Jorge S B, Jerry W, Tinnin S, Yates L.Phylogeny and evolution of the neotropical rodent genus calomys:inferences from mitochondrial DNA sequence data. Mol Phylogenet Evol[J]. 2001,20(2):173～184.
[15] XIAO Bing,MA Fei,SUN Yi,LI Qing-Wei. Comparative Analysis of Complete Mitochondrial DNA Control Region of Four Species of Strigiformes.遗传学报[J]. 2006, 33 (11):965～974
[16] 陈宏等.家畜线粒体(mtDNA)的研究.黄牛杂志,1995,21(2):7～13
[17] 王文,施立明.一种改进的动物线粒体DNA提取方法.动物学研究[J].1993,14(2):197～198
[18] 曹丽荣,王小明,方盛国. 从细胞色素 b 基因全序列差异分析岩羊和矮岩羊的系统进化关系.动物学报[J].2003.49(2):198～204
[19] Hiendleder S, Mainz K, Plante Y, Lewalski H.Analysis of mitochondrial DNA indicates that domestic sheep are derived from two different ancestral maternal sources: no evidence for contributions from Urial and Argali Sheep.Journal of Heredity [J]. 1998,89:113
[20] 庄 轩,丁少雄,郭 丰,王 军,苏永全,张其永,李祺福. 基于细胞色素b基因片段序列研究 中国近海石斑鱼类系统进化关系. 中国科学 C 辑 生命科学[J] 2006, 36(1): 27～34
[21] 孙宗炎等. 应用DNA 指纹图谱法对湘白猪群体遗传结构分析.畜牧兽医学报[J]. 1995,26(1): 29～30
[22] 雷初潮,陈宏,杨公社等.蒙古赫斯坦牛线粒体DNA-Dloop高变区序列分析.黄牛杂志[J]. 2002,28(3):6～7
[23] 雷初潮,陈宏,杨公社等.秦川牛线粒体DNA高变区序列分析.黄牛杂志[J]. 2002,28(2):5～6
[24] 李丽娜,陆师义.mtDNA概述.生物学通报[J]. 1987,(11):5～7
[25] 涂正超,张亚平,邱怀.中国牦牛线粒体DNA多态性及其分化.遗传学报[J]. 1998,25(3):205～212
[26] 张文平,张志和,沈富军,岳碧松. 华南虎、东北虎、孟加拉虎的D-loop 和ND5 序列及其在系统进化分析中的应用.四川动物[J]. 2005.24(1).315～318
[27] 王存芳 曾勇庆 等. 线粒体DNA(mtDNA)的研究进展. 动物科学与动物医学[J]. 2001,18(1):16～18
[28] 郭松长,刘建全,祁得林,杨洁,赵新全. 牦牛的分类学地位及起源研究: mtDNA D-loop 序列的分析. 兽类学报[J]. 2006,26 (4):325～330
[29] 常 洪,苗泽荣,耿社民,武 彬,Hash iguch i T, M eaeda Y. 中亚以东南家牛亲缘系统研究.江苏农业研究[J]. 1999,20(1):42～47
[30] Eleda, J.W.(1996). Concerted evolution of multigenetic families. In Evolution of genes and proteins(M.Nei and R.K Koehn,eds.),pp[J].38～58
[31] 雷初潮.中国四个畜种mtDNA遗传多样性研究,西北农林科技大学博士学位论文(2002)[M]
[32] Mundy N I, Helbig A J. Origin and evolution of tandem repeats in the mitochondrial DNA control region of Shrikes (Lsnius spp.). J Mol Evol[J]. 2004, 59: 250～257
[33]Masatoshi Nei. Sudhir Kumar. Molecular Evolution and Phylogenetics[M]. 北京:高等教育出版社,2002:76～163
[34] SHI Yan-Feng,SHAN Xiang-Nian,LI Jian,SHI Ting-Yan1,ZHENG Ai-Ling. Sequence Analysis and Phylogeny of Deer (Cervidae) mtDNAControl Regions. 遗传学报[J].2004.31(4):395～402
[35] 涂正超.家牛mtDNA多态性研究进展.黄牛杂志[J]. 1997,23(1):11～12
[36] Ravago R G, Monje V D, Juinio-Menez M A. Length and sequence variability in mitochondrial control region of the Milkfish, Chanos chanos. Mar Biotechnol[J]. 2002, 4: 40～50
[37] Carroll,S.B.(1995)Homeotic genes and the evolution of arthropods and chordates. Nature[J] 376:479～485
[38] G.H. 比尔. J.K.C 诺尔斯. 核外遗传学[M].北京:科学出版社,1984:9～55
[39] 雷初朝,陈宏,杨公社,宋林生,雷雪芹,孙维斌,李瑞彪,刘小林 中国部分黄牛品种mtDNA遗传多态性研究.遗传学报[J]. 2004,31(1):57～62
[40] 鲍淑琴.河北畜牧兽医.国外黑色和牛发展动态[J].2001,17(8).12.
[41] 王作洲,戈新.日本和牛.山东畜牧兽医[J].2000,7:39～40.
[42] 刘文生,旭日干,扎鲁达,张锁链. 黑毛和牛在内蒙古呼和浩特地区的繁育.黄牛杂志[J].2004,30(7):13～14
[43] 潘 军 . 日 本 和 牛 品 种 形 成 、 特 征 与 评 价 . 黑 龙 江 动 物 繁殖.1994.2(1):4～5
[44] 郭瑞芬.美国对“日本和牛”的探索. 河北畜牧兽医[J]. 2001,17(5):39～40
[45] 董传河.鲁西黄牛在日本.山东畜牧兽医[J].1998,5:9～11
[46] 孟彦,许尚忠,昝林森,高雪,陈金宝,任红艳,王兴平. 10 个牛品种线粒体12S rRNA 基因多态性分析. 遗传[J].2006,28(4):422～426
[47] 马月辉,曹红鹤,陈幼春,王栋. 部分黄牛品种(群体) 遗传多样性分析. 中国农业科学[J]. 2003,36(6):696～699
[48] 王朝锋,陈宏,雷初朝,孙维斌,党瑞华,张志清.延边牛线粒体DNA D- loop 序列多态性研究. 西北农林科技大学学报[J].2005,33(7):9～11
[49] 吴长庆,于洪春,张国良. 中国草原红牛品种资源现状及展望. 黄牛杂志[J].2000,12,26(6):44～46
[50] 杨国忠,张嘉保,赵玉民,胡成华,任文陟,张国梁. 8 种微卫星DNA 标 记 在 草 原 红 牛 杂 交 群 体 中 多 态 性 的 研 究 . 草 食 家 畜[J].2004,9,3:23～25
[51] 李向阳,张嘉保,赵玉民,胡成华. 草原红牛微卫星DNA 多态性的研究. 中国草食动物[J].2004,24(3):3～5
[52] 杨国忠,任文陟,张嘉保. 草原红牛微卫星DNA群体遗传变异分析研究. 奶牛肉牛[J].2000,4:20～21
[53] 赵玉民,杨国忠,任文陟,张嘉保,胡成华,张国梁. 草原红牛及其杂 交 群 体 中 微 卫 星 DNA 多 态 性 研 究 . 中 国 牛 业 科 学[J].2006,32(6):34～37
[54] Carroll,S.B.(1995)Homeotic genes and the evolution of arthropods and chordates. Nature[J]. 376:479～485
[55] K L Kim 等著,郭军摘译,马月辉校. 通过线粒体DNA D-loop环序列多态性决定亚洲和欧洲猪品种间的系统发育关系. Animal Genetics[J]. 2002, 33:19～25
[56] г.г. 高 泽.线粒体DNA[M].北京:科学出版社,1982:100～113.
[57] 郭松长,刘建全,祁得林,杨洁,赵新全. 牦牛的分类学地位及起源研究: mtDNA D-loop 序列的分析. 兽类学报[J]. 2006,26 (4):325～330
[58] Carroll,S.B.(1995)Homeotic genes and the evolution of arthropods and chordates. Nature[J]. 376:479～485
[59] Bhattacharyya,A.(1946) On a measure of divergence between twomultinomial populations. Sankhya[J].7:401～406
[60] Cann,R.L.,M.Stoneking and A.C. Wilson. (1987)Mitochondrial DNA and human evolution. Nature[J]. 325:31～36
[61] Cao,Y.,J.Adachi and M.Hasegawa.(1998)Comment on the quarter puzzling method for finding maximum-likelihood tree topologies. Mol.Biol. Evol [J] 5:87～89
[62] 涂正超. 动物线粒体 DNA 多态性及其在畜牧科学中的应用. 黑龙江畜牧兽医[J].1994,12:38～40
[63] Wolstenholme D R, Int Rev Cytol[M]. 1992, 141:173
[64] Boettcher P J, Steverink D W B, Beitz D C. et al, J Dairy Sci[J]. 1996,79(4):655
[65] 赵兴波,李宁,吴常信.奶牛产乳性状的核外基因效应. 高科技通讯[J]. 2000,1(10):97～99
[66] Ron M, Genis I, Ezra E. et al, Anim. Biotech. [J]. 1992, 3(2):201
[67] Ron M, Yoffe O and Weller J I, Anim Genet[J]. 1993, 24(3):183
[68] 毛永江,常 洪,杨章平,许 明,宋卫涛,张 柳,常国斌,王冬蕾. 鲁西黄牛遗传多样性及其系统地位的研究. 中国农业科学[J]. 2005, 38(8):1658～1663
[2] 刘振湘,唐晓玲.我国家养动物多样性与畜牧业持续发展.畜牧与兽医[J]. 1997,177～180
[3] 陈振文等. 用 DNA 指纹技术分析 WISTAR 大鼠的遗传距离.浙江畜牧兽医[J].1996,21(3):11～12
[4] Anderson S , de Bruijn M H , Coulson A R , Eperon I C , Sanger F , Young I G. Complete sequence of bovine mitochondrial DNA. Conserved features of the mammalian mitochondrial genome. J Mol Biol[J].1982,156 (4) :683～717
[5] 廖顺尧,鲁成. 动物线粒体基因组研究进展.生物化学与生物物理进展[J].2000,27(5):508～511
[6] Wolstenholme D R, Int Rev Cytol[M]. 1992, 141:173
[7] Kikkawa Y, Yonekawa H , Suzuki H , Amano T. Analysis of genetic diversity of domestic water buffaloes and anoas based on variations in the mitochondrial gene for cytochrome b. Anim Genet[J]. 1997.28 :195～201
[8] Weir B.S.(徐云碧等译). 遗传学数据分析—群体遗传学离散型数据分析方法[M]. 北京:中国农业出版社,1996,184～196
[9] Masatoshi Nei, Sudhir Kumar. 分子进化与系统发育[M]. 北京:高等教育出版社. 2004,5:20～180
[10] 丁波.DNA指纹技术及其在畜禽遗传育种中的应用.青海畜牧兽医杂志[J]. 1994,(4):42～44
[11] Cao,Y.,J.Adachi and M.Hasegawa.(1998)Comment on the quarter puzzlingmethod for finding maximum-likelihood tree topologiesMol.Bio.Evol[J].15:87～89
[12] 徐其放,陈嘉昌,朱世杰,申煜,刘运忠,常弘. 动物线粒体DNA 的特异结构及应用分子系统学分析的方法. 中国比较医学杂志[J]. 2005,10,15(5):515～518
[13] 张 丽,侯水生,刘小林,徐铁山. DNA 指纹技术分析畜禽亲缘关系的原理及应用.家畜生态[J]. 2003,24(4):48～50
[14] Jorge S B, Jerry W, Tinnin S, Yates L.Phylogeny and evolution of the neotropical rodent genus calomys:inferences from mitochondrial DNA sequence data. Mol Phylogenet Evol[J]. 2001,20(2):173～184.
[15] XIAO Bing,MA Fei,SUN Yi,LI Qing-Wei. Comparative Analysis of Complete Mitochondrial DNA Control Region of Four Species of Strigiformes.遗传学报[J]. 2006, 33 (11):965～974
[16] 陈宏等.家畜线粒体(mtDNA)的研究.黄牛杂志,1995,21(2):7～13
[17] 王文,施立明.一种改进的动物线粒体DNA提取方法.动物学研究[J].1993,14(2):197～198
[18] 曹丽荣,王小明,方盛国. 从细胞色素 b 基因全序列差异分析岩羊和矮岩羊的系统进化关系.动物学报[J].2003.49(2):198～204
[19] Hiendleder S, Mainz K, Plante Y, Lewalski H.Analysis of mitochondrial DNA indicates that domestic sheep are derived from two different ancestral maternal sources: no evidence for contributions from Urial and Argali Sheep.Journal of Heredity [J]. 1998,89:113
[20] 庄 轩,丁少雄,郭 丰,王 军,苏永全,张其永,李祺福. 基于细胞色素b基因片段序列研究 中国近海石斑鱼类系统进化关系. 中国科学 C 辑 生命科学[J] 2006, 36(1): 27～34
[21] 孙宗炎等. 应用DNA 指纹图谱法对湘白猪群体遗传结构分析.畜牧兽医学报[J]. 1995,26(1): 29～30
[22] 雷初潮,陈宏,杨公社等.蒙古赫斯坦牛线粒体DNA-Dloop高变区序列分析.黄牛杂志[J]. 2002,28(3):6～7
[23] 雷初潮,陈宏,杨公社等.秦川牛线粒体DNA高变区序列分析.黄牛杂志[J]. 2002,28(2):5～6
[24] 李丽娜,陆师义.mtDNA概述.生物学通报[J]. 1987,(11):5～7
[25] 涂正超,张亚平,邱怀.中国牦牛线粒体DNA多态性及其分化.遗传学报[J]. 1998,25(3):205～212
[26] 张文平,张志和,沈富军,岳碧松. 华南虎、东北虎、孟加拉虎的D-loop 和ND5 序列及其在系统进化分析中的应用.四川动物[J]. 2005.24(1).315～318
[27] 王存芳 曾勇庆 等. 线粒体DNA(mtDNA)的研究进展. 动物科学与动物医学[J]. 2001,18(1):16～18
[28] 郭松长,刘建全,祁得林,杨洁,赵新全. 牦牛的分类学地位及起源研究: mtDNA D-loop 序列的分析. 兽类学报[J]. 2006,26 (4):325～330
[29] 常 洪,苗泽荣,耿社民,武 彬,Hash iguch i T, M eaeda Y. 中亚以东南家牛亲缘系统研究.江苏农业研究[J]. 1999,20(1):42～47
[30] Eleda, J.W.(1996). Concerted evolution of multigenetic families. In Evolution of genes and proteins(M.Nei and R.K Koehn,eds.),pp[J].38～58
[31] 雷初潮.中国四个畜种mtDNA遗传多样性研究,西北农林科技大学博士学位论文(2002)[M]
[32] Mundy N I, Helbig A J. Origin and evolution of tandem repeats in the mitochondrial DNA control region of Shrikes (Lsnius spp.). J Mol Evol[J]. 2004, 59: 250～257
[33]Masatoshi Nei. Sudhir Kumar. Molecular Evolution and Phylogenetics[M]. 北京:高等教育出版社,2002:76～163
[34] SHI Yan-Feng,SHAN Xiang-Nian,LI Jian,SHI Ting-Yan1,ZHENG Ai-Ling. Sequence Analysis and Phylogeny of Deer (Cervidae) mtDNAControl Regions. 遗传学报[J].2004.31(4):395～402
[35] 涂正超.家牛mtDNA多态性研究进展.黄牛杂志[J]. 1997,23(1):11～12
[36] Ravago R G, Monje V D, Juinio-Menez M A. Length and sequence variability in mitochondrial control region of the Milkfish, Chanos chanos. Mar Biotechnol[J]. 2002, 4: 40～50
[37] Carroll,S.B.(1995)Homeotic genes and the evolution of arthropods and chordates. Nature[J] 376:479～485
[38] G.H. 比尔. J.K.C 诺尔斯. 核外遗传学[M].北京:科学出版社,1984:9～55
[39] 雷初朝,陈宏,杨公社,宋林生,雷雪芹,孙维斌,李瑞彪,刘小林 中国部分黄牛品种mtDNA遗传多态性研究.遗传学报[J]. 2004,31(1):57～62
[40] 鲍淑琴.河北畜牧兽医.国外黑色和牛发展动态[J].2001,17(8).12.
[41] 王作洲,戈新.日本和牛.山东畜牧兽医[J].2000,7:39～40.
[42] 刘文生,旭日干,扎鲁达,张锁链. 黑毛和牛在内蒙古呼和浩特地区的繁育.黄牛杂志[J].2004,30(7):13～14
[43] 潘 军 . 日 本 和 牛 品 种 形 成 、 特 征 与 评 价 . 黑 龙 江 动 物 繁殖.1994.2(1):4～5
[44] 郭瑞芬.美国对“日本和牛”的探索. 河北畜牧兽医[J]. 2001,17(5):39～40
[45] 董传河.鲁西黄牛在日本.山东畜牧兽医[J].1998,5:9～11
[46] 孟彦,许尚忠,昝林森,高雪,陈金宝,任红艳,王兴平. 10 个牛品种线粒体12S rRNA 基因多态性分析. 遗传[J].2006,28(4):422～426
[47] 马月辉,曹红鹤,陈幼春,王栋. 部分黄牛品种(群体) 遗传多样性分析. 中国农业科学[J]. 2003,36(6):696～699
[48] 王朝锋,陈宏,雷初朝,孙维斌,党瑞华,张志清.延边牛线粒体DNA D- loop 序列多态性研究. 西北农林科技大学学报[J].2005,33(7):9～11
[49] 吴长庆,于洪春,张国良. 中国草原红牛品种资源现状及展望. 黄牛杂志[J].2000,12,26(6):44～46
[50] 杨国忠,张嘉保,赵玉民,胡成华,任文陟,张国梁. 8 种微卫星DNA 标 记 在 草 原 红 牛 杂 交 群 体 中 多 态 性 的 研 究 . 草 食 家 畜[J].2004,9,3:23～25
[51] 李向阳,张嘉保,赵玉民,胡成华. 草原红牛微卫星DNA 多态性的研究. 中国草食动物[J].2004,24(3):3～5
[52] 杨国忠,任文陟,张嘉保. 草原红牛微卫星DNA群体遗传变异分析研究. 奶牛肉牛[J].2000,4:20～21
[53] 赵玉民,杨国忠,任文陟,张嘉保,胡成华,张国梁. 草原红牛及其杂 交 群 体 中 微 卫 星 DNA 多 态 性 研 究 . 中 国 牛 业 科 学[J].2006,32(6):34～37
[54] Carroll,S.B.(1995)Homeotic genes and the evolution of arthropods and chordates. Nature[J]. 376:479～485
[55] K L Kim 等著,郭军摘译,马月辉校. 通过线粒体DNA D-loop环序列多态性决定亚洲和欧洲猪品种间的系统发育关系. Animal Genetics[J]. 2002, 33:19～25
[56] г.г. 高 泽.线粒体DNA[M].北京:科学出版社,1982:100～113.
[57] 郭松长,刘建全,祁得林,杨洁,赵新全. 牦牛的分类学地位及起源研究: mtDNA D-loop 序列的分析. 兽类学报[J]. 2006,26 (4):325～330
[58] Carroll,S.B.(1995)Homeotic genes and the evolution of arthropods and chordates. Nature[J]. 376:479～485
[59] Bhattacharyya,A.(1946) On a measure of divergence between twomultinomial populations. Sankhya[J].7:401～406
[60] Cann,R.L.,M.Stoneking and A.C. Wilson. (1987)Mitochondrial DNA and human evolution. Nature[J]. 325:31～36
[61] Cao,Y.,J.Adachi and M.Hasegawa.(1998)Comment on the quarter puzzling method for finding maximum-likelihood tree topologies. Mol.Biol. Evol [J] 5:87～89
[62] 涂正超. 动物线粒体 DNA 多态性及其在畜牧科学中的应用. 黑龙江畜牧兽医[J].1994,12:38～40
[63] Wolstenholme D R, Int Rev Cytol[M]. 1992, 141:173
[64] Boettcher P J, Steverink D W B, Beitz D C. et al, J Dairy Sci[J]. 1996,79(4):655
[65] 赵兴波,李宁,吴常信.奶牛产乳性状的核外基因效应. 高科技通讯[J]. 2000,1(10):97～99
[66] Ron M, Genis I, Ezra E. et al, Anim. Biotech. [J]. 1992, 3(2):201
[67] Ron M, Yoffe O and Weller J I, Anim Genet[J]. 1993, 24(3):183
[68] 毛永江,常 洪,杨章平,许 明,宋卫涛,张 柳,常国斌,王冬蕾. 鲁西黄牛遗传多样性及其系统地位的研究. 中国农业科学[J]. 2005, 38(8):1658～1663