摘要
为了进一步明确我国地方绵羊品种的起源、分类及遗传关系,本研究从中国甘肃、宁夏、陕西、山东、河南、新疆、青海等省、市自治区采集了兰州大尾羊(Lanzhou large tail sheep,LLT)、滩羊(Tan sheep,TS)、同羊(Tong sheep,TTS)、小尾寒羊(Small-tail han sheep,STH)、大尾寒羊(Large-tail han sheep,LTH)、阿勒泰羊(Aletail sheep,ALT)、西藏羊(Tibetan sheep,TBS)、多浪羊(Duolang sheep,DL)8个地方品种,并在陕西榆林采集了2个外来纯种萨福克绵羊(Suffolk sheep,SFK)和道塞特绵羊(Dorset sheep,DST)作为对照。本试验采用上游引物L15387 :5′AGCCCCACTATCAACACC3′和下游引物H16149:5′AAATAGTTACCCCCACAGTTAG3′。具体扩增序列位置是靠近tRNA pro一侧的D-loop 763 bp ,其中包含了D-loop上游50 bp。并通过测序分析D-loop高变区序列对我国的绵羊品种进行起源进化和多态性研究,结果如下:
1.本试验所研究序列已提交GenBank,登录号:EF494774~EF494792,EF494802~EF494826,EF494875~EF600931,EF494842~EF600924,EF494866~EF600929,EF600912~EF600917。供试绵羊mtDNA D-loop HVI序列A、T含量占优势,其中国绵羊品种与外来品种mtDNAD-loop序列碱基组成没有明显的差异,说明了绵羊mtDNAD-loop为A、T富集区。
2.供试绵羊mtDNA D-loop序列的长度范围为522 bp~683 bp,共有19种分子类型。这种长度变异除了少数碱基的缺失和插入之外,主要是由于D-loop区存在2~5个75 bp的串连重复序列所致。但这种序列长度的变异在单倍型中的出现是随机的,并不能作为单倍型的序列特征。
3.本试验所获得的93条绵羊mtDNA D-loop序列,共发现了63种单倍型,共有73个变异位点,其中单一多态位点有30个,简约信息位点有43个。除了缺失和插入位点外,还包括22处碱基转换和1处碱基颠换,转换和颠换之比为14.8。说明绵羊线粒体碱基替换存在转换偏倚现象。
4.本试验所获得的93条绵羊mtDNA D-loop序列单倍型多样度和核苷酸多样性分别为0.981、0.02900,平均核苷酸差异数K为9.309,反映了中国绵羊品种遗传多样性较贫乏。
5.对D-loop区所测个体的所有单倍型利用MEGA 4.0进行NJ, UPGMA和ME聚类时,中国绵羊mtDNA D-loop单倍型序列聚类成A、B、和C三大支系。为了增加结果的可靠性,借用GenBank己发表的摩佛伦羊、赤羊、羱羊、盘羊和家绵羊的D-loop区序列与本研究中的所有序列一同聚类,结果摩佛伦羊聚在B单倍型组中,说明摩佛伦羊与家绵羊的关系较近。而赤羊自成一类,并没有出现在3个单倍型组中。说明现代家绵羊存在3个母系起源,且第三个母系起源地可能就在中国。
In order to ascertain origin, sort and genetic relation of locality sheep breed in China, samples from 79 individuals of 8 different sheep breeds were collected from Gansu province
(Lanzhou large tail sheep , LLT ), Ningxia province(Tan sheep , TS), Shanxxi province(Tong sheep , TTS),Shandong province(Small-tail han sheep , STH),Henan province(Large-tail han sheep,LTH), Xinjiang province(Aletail sheep,ALT; Small-tail han sheep,STH; Duolang sheep,DL),Qinghai province(Tibetan sheep,TBS) . And the Suffolk sheep and Dorset sheep from Yulin city in Shanxxi province as a control. Studying on the phylogeny evolution and population diversities among Chinese Domestic sheep breeds through analysising sequence of mtDNA D-loop HVI . the results showed that:
1. All sequence data to GenBank,GenBank accession numbers:EF494774~EF494792,EF494802~EF494826,EF494875~EF600931,EF494842~EF600924,EF494866~EF600929,EF600912~EF600917。From the mean nucleotide compositions it could be concluded that there were enriched A and T in the mtDNA D-loop HVI .
2. The length of the mtDNA D-loop sequences was considerable variable between 522bp and 683 bp, and nineteen molecule model. Except for minor insertions and deletions, the observed length variations were caused by different copy numbers of a 75 bp tandem repeat sequence. But it could not be considered as the characteristic sequence of haplotypes.
3. The 93 individual sheep mtDNA D-loop HVI sequences found 63 haplotypes.and contained 73 variable sites, among them, singleton variable sites was 30, parsimony informative sites of two variants was 43. Except for insertions and deletions sites,the transition sites were 22 and transversion sites were 1, the ratio of transitions to transversions was 14.8,suggesting Chinese sheep has the high transition bias.
4. The haplotype diversity and nucleotide diversity 93 individual sheep mtDNA D-loop HVI sequences were 0.981、0.02900.The average number of nucleotide differences variation was 9.309.The results indicate that the genetic diversity of Chinese sheep is very abundant.
5. The NJ, UPGMA and ME phylogenetic trees was constructed by using MEGA 4.0 with goat as outgroup, three major lineages appeared in the 8domestic sheep, which termed A, B and C lineage. In order to improve the reliability of theresults, all the D-loop complete sequences were used to clustering which was O.musimon, O. vignei, O.aries from GenBank together with the sequences obtained in this study. The results showed that: the relationship between O.musimon and O.aries was nearer, because they were all in B lineage. All the analysis results indicated that the modern domestic sheep have the third maternal origin, and the genesis might be in China.
引文
[1] 马月辉,陈幼春,冯维棋.中国家养动物多样性概况[J].畜牧兽医学报,2000, 31(5) : 394~399.
[2] 马月辉,陈幼春,冯维棋.畜禽种质资源评价[J].家畜生态,2002, 23 (2) : 1~5.
[3] McNeely J A. Conserving the World's Biological Diversity [C].Washington, D. C. and Gland.Switzerland. 1990.
[4] 胡玉华.谈生物多样性的内涵[J].生物学通报,1999, 34 (3) : 19
[5] 马克平,米湘成,魏伟等.生物多样性研究中的几个热点问题.中国生物多样性保护与研究进展—第五届全国生物多样性保护与持续利用研讨会论文集[A].北京:气象出版社,2004,15~36.
[6] 陈幼春.中国家畜多样性保护的意义.生物多样性[J].1995,3(3):143~146.
[7] Hammond,P.M."The Current Magnitude of Biodiversity." In Global Biodiversity Assessment.Ed.V.H.Heywood.Cambridge:Cambridge Univ.Press[J].1995,113~128.
[8] Anderson S,De Bruin MHL,Coulson A R et al..Complete sequence of bovine mitochondrial DNA. Conserved features of the mammalian mitochondrial genome.J Mol Evol [J].1982,156:683~717.
[9] Hecht W.Studies on mitochondrial DNA in farm animals.In:Genome analysis in domestic animals[A].1990,259~268
[10] Lin Chich-Sheng,Sun Yu-Lin,Liu Chang-Yeu et al..Complete nucleotide sequence of pig(Sus scrofa) mitochondrial genome and dating evolutionary divergence within Artiodactyla.Genetics[J].1999, 236,107~114.
[11] Bjǒ M.Uring,Ulfur Arnason.The complete mitochondrial DNA sequence of the pig(Sus scrofa).J Mol.Evol.[J].1998,47:302~306
[12] Hiendleder S,Lewalski H,Wassmuth R et al..The complete mitochondrial DNA sequence of the domestic sheep(Ovis aries)and comparison with the other major ovine haplotype.J Mol Evol[J]. 1998a,47:441~448.
[13] Xu X, Amason U.The complete mitochondrial DNA sequence of the horse,Equus caballus; Extensive heteroplasmy of the control region.Gene[J].1994,148:357~362.
[14] Pietro P,Maria F,Gianfranco G et al..The complete nucleotide sequence of goat(Capra hircus)mitochondrial genome.DNA sequence[J].2003,14(3):199~203.
[15] Bibb M J,Van Etten R A,Wright C T et al.Sequence and gene organization of mouse mitochondrial DNA.Cell[J].1981,26:167~180.
[16] Roe B A,Ma DP,Wilson RKet al..The complete nucleotide sequence of the Xenopus laevis mitochondrial genome.J Biol.Chem[J].1985,260(17):9759~9774.
[17] Janke A,Xu Xiufeng,Arnason Ulfur.The complete mitochondrial genome of the wallaroo(Macropus robustus)and the phylogenetic relationship among Monotremata,Marsupialia,and Eutheria.Proc. Natl.Acad.Sci.USA[J].1997,94:1276~1281.
[18] Crease T J.The complete sequence of the mitochondrial genome of Daphnia pulex(Cladocera: Crustacea).Gene[J].1999,233(1-2):89~99.
[19] Broughton R E,Milam J E,Roe B A.The complete sequence of the Zebrafish(Danio rerio) mitochondrial genome and evolutionary patterns in vertebrate mitochondrial DNA.Genome Research[J].2001,11:1958~1967.
[20] Vivian W,Jaime V,Alvaro M et al..The complete sequence of the mitochondrial genome of the Chinook salmon,Oncorhynchus tshawytscha.Biol.Res.[J].2003,36(2):223~231.
[21] Kutsyi M P,Gouliaeva N A,Kuznetsova E A et al mitochondria.Mitochondrion[J].2005,5(1):35~44.
[22] Taanman J W.The mitochondrial genome:Structure, Biochim Biophys Acta[J].1999,1410:103~123.
[23] Dawid I. B., Blacker A. W. Matenal and cytoplasmic inheritance of mtDNA in xenopus. Developm.Bio1.1972.29:152~161.
[24] Hutchinson C.A. etal. 1980. Maternal inheritance of mammalian mitochondria DNA [J]. Nature,251:536~538.
[25]Hill E.W., Bradley D.G,, Al-Barody M., Ertugrul O., Splan R.K., Zakharov L., Cunningham E.P 2002. History and integrity of thoroughbred dam lines revealed in equine mtDNA variation.Anim Genet. 33(4):287~94.
[26]Shitara H, Hayashi J I, Takahama S, et al. Maternal inheritance of mouse mtDNA in interspecific hybrids: segregation of the leaked paternal mtDNA followed by the prevention of subsequent paternal leakage. Genetics, 1998, 148(2): 851~857.
[27]Gyllensten U, Wharton D, Josefsson A, et al. Paternal inheritance of mitochondrial DNA in mice. Nature, 1991, 352(6 332):255~257.
[28]Kennedy P, Nachman M W. Deleterious moutation at the mitochondrial ND3 gene in south American marsh rates (Holochilus) [J]. Genetics 1998 , 150(1):359~368.
[29] 王静波,胡长龙,徐宏发.线粒体 DNA(mtDNA)多态性在动物保护生物学中的应用.生物多样性[J].2001,9(2):181~187.
[30] Pietro P, Maria F, Gianfranco G etal.. The complete nucleotide sequence of goat (Capra hircus) mitochondrial genome. DNA sequence [J]. 2003, 14(3):199 一 203.
[31]. Rand, D M. Endotherms, ectotherms, and mitochondrial genome-size variation. J. Mol. Evol.[J].1993, 37: 281~295.
[32]兰蓉,洪琼花,高源汉等.云南绵羊线粒体 DNA 遗传多样性研究.遗传[J].1998,20(1):20-23.
[33] Chen H and Leibenguth F.Restriction endonuclease analysis of mitochondrial DNA of three farm animal species:Cattle,sheep and goat.Comp.BIochem.Physiol.Part B:Biochemical and molecular Biology[J].1995,111(4):643~649.
[34]涂正超,张亚平,邱怀.中国牦牛线粒体 DNA 多态性及遗传分化.遗传学报[J].1998,25(3): 205~212.
[35].赵兴波,冯继东,李宁等.绵羊(Ovis aries)线粒体 DNA 的遗传变异类型研究.自然科学进展 [J].2001,11(12):1326~1328.
[36] 陈宏,F.Leibenguth,邱怀.家畜线粒体 DNA 的研究.黄牛杂志[J].1995,21(1):7~17.
[37] Wallace D C,Brown M D,Lott M.Mitochondrial DNA variation in human evolution and disease.Gene[J].1999,238:211~230.
[38].Tengan C H,Claudia Ferreiro-Barros,Marina Cardeal et al..Frequency of duplications in the D-loop in patients with mitochondrial DNA deletions.Biochimica et Biophysica Acta[J].2002,1588:65~70.
[39].Mckenzie D,Bua E,Mckiernan S et al..Mitochondrial DNA deletion mutations:A causal role in sarcopenia.Eur.J Biochem[J].2002,269:2010~2015
[40] Brown,W. M. Po lymo rph ism in m itochondrial DNA of humans as revealed by restriction endonuclease analysis. PN A S USA , 1980, 77: 3605~3609. [41 ] Horai, S. A nd M atsunage E. m tDNA po lymo rph ism in Japanese analysis w ith restriction enzymesof four o r five base pair recognition [J ]. Hum. Genet. , 1986, 72: 105~117.
[42]王勇,张俊武.线粒体DNA 突变与线粒体脑肌病[J ].生理科学进展,199728 (1): 70~72.
[43] Botto Nicoletta, Berti Sergio, Manfredi Samantha et al..Detection of mtDNA with 4977 bp deletion in blood cells and atherosclerotic lesions of patients with coronary artery disease.Mutaion research/fundamental and molecular mechanisms of mutagenesis [J].2004,570(1):81~88.
[44].Obermaier-Kusser B, Müller-Hǒcker J,Nelson I et al..Different copy numbers of apparently identically deleted mitochondrial DNA in tissues from a patient with Kearns-Sayre syndrome detected by PCR.Biochemical and biophysical research communications [J].1990, 169(3):1007~1015.
[45].Zeviani M and DiDonato S. Neurological disorders due to mutations of the mitochondrial genome.Neuromuscular Disorders [J].1991, 1(3):165~172.
[46].Lightowlers R N, Chinnery P F, Turnbul D M et al..Mammalian mitochondrial genetics: heredity, heteroplasmy and disease.Trends in Genetics [J].1997, 3(11):450~455.
[47].Taanman J W, Bodnar A G, Cooper J M et al. .Molecular mechanisms in mitochondrial DNA depletion syndrome.Human Molecular Genetics [J].1997, 6:935~942.
[48].Taanman J W,Muddle J R,Muntau A C.Mitochondrial DNA depletion can be prevented by dGMP and dAMP supplementation in a resting culture of deoxyguanosine kinase-deficient fibroblasts.Human Molecular Genetics[J].2003,12(15):1839~1845.
[49] Spinazzola A, Carrara F, Marina M et al..Mitochondrial myopathy and ophthalmoplegia in asporadic patient with the mitochondrial DNA mutation.Neuromuscular Disorders[J].2004,14(12):815~817.
[50] Masuyama M, Iida R, Takatsuka H et al.. Quantitative change in mitochondrial DNA content in various mouse tissues during aging. Biochimica et Biophysica Acta (BBA)[J]. 2005, Article in press.
[51] 宫澜, 曾溢滔. 线粒体DNA 突变相关疾病的基因治疗[J ]. 生命科学, 1997 (4) : 154~ 157.
[52] 黄学文,赵琪,陈道祯等.肝癌组织中线粒体DNA D-loop区碱基变异与ROS水平.遗传[J] 2005,27(1):014-020.
[53] Watanabe T,Masangkay J S ,Wakana S et al .Mitochondrial DNA polymorphism in native Philippine cattle based on restriction endonuclease cleavage patterns [J]. Biochem Genetic.1989,27:431~438.
[54] Tu Z C,Qiu H,Zhang Y P.Polymorphism in mitochondrial DNA(mtDNA)of Yak(Bos grunniens).Biochemical Genetics[J].2002,40(5-6):187-193.
[55] 雷初朝,陈宏,杨公社等.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.
[56] Loftus RT,MacHugh D E,Bradley D G et al. Evidence for two independent domestications of cattle. Pro.Natl. Acad.Sci. [J].1994a, 91:2757-2761.
[57] Loftus R T, MacHugh D E,Ngere L O et al..Mitochondrial genetic variation in European, African and Indian cattle populations.Animal Genetics [J].1994b,25,265-271.
[58] 蒙世杰,王静,刘佩等.羚牛细胞色素b基因序列分析和系统进化研究.西北大学学报(自然科学版)[J].2001, 31(4):347-354.
[59] 涂正超,张亚平,邱怀.中国牦牛线粒体DNA多态性及遗传分化.遗传学报[J].1998, 25(3): 205-212.
[60] 俞 英,文际坤,朱开贤等,1996.云南文山黄牛和迪庆黄牛的遗传多样性比较研究.黄牛杂志,22:50~55.(Yu Y,Weng J K,Zhu K X et al,1996.Study on the genetic polymorphism in Wenshan yellow cattle and Diqing yellow cattle in Yunnan.Journal of Yellow Cattle Science,22:50-55.)
[61] Brown W.M. Evolution of animal mitochondrial DNA, in: Nei M., Koehn R.K. (Eds.), Evolution ofgenes and proteins, Sinauer Associates, Sunderland, MA [A]. 1983, 63一88.
[62] 黄勇富.猪mtDNA多态性与中国地方猪种起源分化的关系[[J].遗传学报.1998, 25(4): 322-329.
[63] watanabe T et a1.Pig mitochondrial DNA polymorphism,restriction map orientation andsequence data.Biochen [J].Gene. 1986. 24:85-96.
[64] 兰宏,王文,施立明.西南地区家猪和野猪mtDNA遗传多样性研究.遗传学报[J].1995, 22(1):28-32.
[65].黄勇富,张亚平,邱祥聘等.猪线粒体DNA多态性与中国地方猪种起源分化的关系.遗传学报[J].1998,25 (4):322-329.
[66] Kijas J M H,Anderson L.A phylogenetic study of the origin of the domestic pig estimated from the near-complete mtDNA genome.J Mol Evol[J].2001,52:302-308.
[67] 蒋思文,Giuffra Elisabetta,Andersson Leif等.6个中国猪地方品种和3个瑞典猪DNA分子系统发育相关关系.遗传学报[J].2001,28(12):1120-1128.
[68] David E.MacHugh and Daniel G.Bradley.Livestock gengetic origins: Goats buck the trend.ProcNatl Acid Sci USA[J].2001,98(10):5382-5384.
[69] 赵兴波,李宁.猪线粒体DNAW D—loop PCR—RFLP分析[J].
[70] Yang J,Wang J,Kijas J.Genetic diversity present within the near-complete mtDNA genome of 17 breeds of indigenous Chinese pigs.Journal of Heredity[J],2003,94(5):381-385.
[71] Larson Greger, Dobney Keith,Albarella Umberto,Fang Meiying,et al.2005.Worldwide phylogeography of wild boar reveals multiple centers of pig domenstication [J]. Science.307:1618~1621.
[72]段子渊,从 mtDNA 序列分析中国家猪的起源与遗传多样性[D].西北农林科技大学 .杨凌,2000.
[73]叶绍辉,彭和禄,王文,等.云南保种山羊mtDNA限制性酶切研究[J].中国畜牧杂志,1998, 34 (3):11一12.
[74]贾永红,史宪伟,简承松,等.贵州四个山羊品种mtDNA多态性及其起源分化,动物学研究,1999, 20 (2):88一92.
[75]中国家畜家禽品种志编委会.中国羊品种志[M]. 上海科学技术出版社,1989.
[76]李样龙,田庆义,刘金福,等.河北省山羊品种mtDNA遗传多样性研究[J] 中国畜牧杂志,2000,36 (3):9一11.
[77]谢成侠著.中国养牛羊史(附养鹿简史)[M ].北京:中国农业出版社.1985.
[78]冯维棋.我国占代绵羊品种形成初考.农业考古[J],1991, (3):338345.
[79]Ryder M L. Domestication history and breed evolution in sheep. In: Genetic resources of pig, sheep and goat[A]. Elsevier science publishers B.V..1991, 365一375.
[80]涂正超,张亚平.藏绵羊线粒体DNA遗传多样性研究[J].畜牧兽医学报,1998, 29( 2) :132-135.
[81] Hiendleder S, Hecht W, Dzapo V et al.. Ovine mitochondrial DNA: restriction enzyme analysis, mapping and sequencing data. Animal Genetics [J]. 1991, 23,151一160.
[82] Hiendleder S, Kaupe B, Wassmuth R et al.. Molecular analysis of wild and domestic sheep questions current nomenclature and p-ovides evidence for domestication from two different subspecies. Froc.R. Soc. Lond[J]. 2002, B269,893一904.
[83] Hiendleder S, Mainz K, Plante Y et al.. 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. J Hered[J]. 1998c, 89(2):113一120.
[84] Hiendleder S, Phua S H, Hecht W et al.. A diagnostic assay discriminating between two major Oviscries mitochondrial DNA haplogroups. Animal Genetics [J]. 1999, 30(3):211-213.
[85]贾永红,简承松,朱文适等.贵州绵羊线粒体DNA (mtDNA)多态性研究.西南农业学报[[J].1998, 12(1):100-105.
[86]李祥龙,川庆义,刘铮铸等.九个绵羊品种线粒体DNA限制性片段长度多态性比较研究.畜牧兽医学报[J]. 2001, 32(4):295-298.
[87]贾斌,陈杰,赵茹茜等.新疆8个绵羊品种遗传多样性和系统发生关系的微卫星分析.遗传学报[J]. 2003, 30(9):847一854.
[88]赵兴波,李宁,吴常信.动物线粒体核质基因4_作的研究进展.遗传[[J]. 2001, 23(1):81-85.
[89]陈玉林.中国绵羊的分了进化与遗传多样性研究.西北农林科技大学博士学位论文[[D].中国杨凌,2000.
[90]郭军.中国部分绵羊品种mtNDA多态性及其遗传分化的研究[[D ].北京.中国农业科学院硕士学位论文.
[91] Ivankovic A, Dovc P, Kavar T et al.. Genetic cha2racterisation of the Pag island sheep breed based on microsatellite and mtDNA data. Small Ruminant Research [J]. 2005, 57(2-3):167-174.
[92] Boyce W M, Ramey R R, Rodwell T C et al.. Population subdivision among desert bighorn sheep (Ovis Canadensis) ewes revealed by mitochondrial DNA analysis. Molecular Evoution[J]. 1999,8:99106.
[93] 邹平, 张廷钦, 王文等. 云南几个地方鸡种的mtDNA限制性片段多态性研究.中国畜牧杂志, 1997, 33(6): 26~27
[94] 史宪伟 张亚平.中国主要家鹅品种的遗传分化研究〔J〕.遗传学报,2005,32(10):1053~1054.
[95] Nei M, Li W H.endonucleases [J].Mathem atical model Proc.Nad.Acad.Sci.USA for study genetic variation in teens of restriction.1979, 76(10): 5269~5273.
[96] Nei M, Li W H.endonucleases [J].Mathem atical model Proc.Nad.Acad.Sci.USA for study genetic variation in teens of restriction.1979, 76(10): 5269~5273.
[97] Fitch , W. M. and E. Margoliash. Construction of phylogenetic trees. Science 1967 155: 279~2841.
[98] Saitou N. and M. Nei. The neighbor joining method: A new method for reconstructing phylogenetictrees[J]. Mol. Biol. Evol.,1987, 4: 406-425.
[99] Studier J. A. and K. J. Keppler. A note on the neighbor joining algorithm of Saitou and Nei [J]. Mol.Biol. Evol. 1988 5: 729-731.
[100] Eck R. V and M. O. Dayhoff. Atlas of protein sequence and structure [J]. National Biomedical Research Roundation, Silver Springs, MD. 1966.
[101] Fitch W. M. Toward defining the course of evolution: Minimum change for a specific tree topology[J]. Syst. Zool. ,1971, 20: 406-416.
[102] Hartigan J.A. Minimum evolution fits to a given tree [J]. Biometrics, 1973, 29: 53-65.
[103] Felsenstein J. Evolutionary trees from DNA sequences: A maximum likelihood approach [J]. J. Mol. Evol. 1990, 31:151一160.
[104] Kishino H.,T. Miyata and M. Hasegawa. Maximum likelihood inference of protein phylogeny and the origin of chloroplasts [J]. J. Mol. Evol. 1990, 31:151一160.
[105] Cavalli-Sforza L. L. and A. W. F. Edwards. Phylogenetic analysis: Models and estimation procedures[J]. Am.J. Hum. Genet., 1967, 19: 233-257.
[106] Rozas J, Sanchez-Delbarrio J C, Messeguer X and Rozas R. DNA Sequence Polymorphism Version4.10.1. 2005.
[107] Kumar S., Tamura K.,&Nei M. MEGA3: integrated software for molecular evolutionary genetics analysis and sequence alignment [M]. Briefings in Bioinformatics. 2004, 5, 150-160.
[108] Bandelt H J, Forster P, Sykes B C et al.. Mitochondrial portraits of human populations. Genetics[J].1995, 141:743-753.
[109] Bandelt H J, Foister P and Rohl A. Median-joining networks for inferring interspecific phylogenies.Mol. Bio. Evo1[J]. 1999, 16:37-48
[110] Tobias Polzin, Siavash Vahdati, Network4.1.0.8. Fluxus Technology Ltd. 2004.
[111] Li Wen-Hsiung, D.an Graur. Foundamentals of molecular evolution [M].[0] Sinauer Associates, Inc.Sunderland, Massachusetts. 1991.
[112] Hiendleder S.Kaupe B,Wassmuth R,Janke A.Molecuhr ana卜sisofwild and domestic sheep questons currentnom enchlure and provides evidence for domesticaton from domestication from two different subspecies ProcR SocLand (B).2002,269: 893 ~904.
[113] 成述儒,韩建林,Olivier Hanotte等.中国绵羊群体mtDNA D-Loop的遗传多样性分析[J]. 甘肃农业大学学报,2005,4:440~447.
[114] H iendlederS, Mainz K,Plante Y, Lew a lski H. Ana lys is o f m itochond rial DNA indicates tha t domestic sheep are derive d from two different ancestral maternal sources: no e vidence for contributions from urial and argali sheep[J].Hered,1998,89:113~120.
[115] Wu C H,Zhang Y P,Bunch T D,Wang S,Wang W.Mitocho nd ria l control region sequence va riation with in the argali wild sheep(Ovis ammon):evolution and conservation relevance.Mammalia,2003,67:109~118.
[116] Hiendleder S, Mainz K, Plante Y et al.. 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. J Hered[J]. 1998c, 89(2):113~120.
[117] Wood NJ,P hua S H.Va riation in the control refion sequence of the sheep mitochondrial genome[J]. Anim Genet,1996,27:25~33.
[118] 李祥龙,张增利,巩元芳等.我国主要地方绵羊品种遗传亲缘关系[J].中国兽医学报,2004,24(5):508~510.
[119] 《中国羊品种志》组编.中国羊品种志[M].上海:上海科学技术出版社,1989.4.
[120] 常洪主编.家畜遗传资源纲要.北京:中国农业出版社,1995.
[121] 雷雪芹,陈宏,陈玉林等.六个中国固有绵羊品种的群体遗传学分析[J].遗传改良,2001~2002年全国养羊生产与学术研讨会论文集.2003,81~83.
[122] 马海明,门正明,黄生强等.兰州大尾羊血液蛋白多态性研究[J].湖南农业大学学报(自然科学版),2004,30(4):351~354.
[123] Nei M, Miller J C. A simple method for estimating average number of nucleotide substitutions within and between populations from restriction data. Genetics[J]. 1990, 125:873-879.