应用微卫星标记分析部分中国地方山羊群体的遗传多样性
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摘要
家山羊是中国重要的畜种。近年来由于商业品种的引入导致地方品种山羊数量的锐减,甚至有些品种处于濒危状态。我国地方山羊品种的遗传多样性研究势在必行。本研究利用微卫星标记对中国10个地方山羊品种(河西绒山羊、中卫山羊、陕南白山羊、黄槐山羊、伏牛白山羊、沂蒙黑山羊、鲁北白山羊、太行山羊、吕梁黑山羊、济宁青山羊)及1个引进品种波尔山羊共计530个个体的遗传多样性进行了分析研究。结果如下:
1.采用SRCRSP5、MAF209、TGLA53、ETH10、OarAE54、CSRD247、SRCRSP15、SRCRSP3、MAF065、SRCRSP8、SPS113、INRABERN172、OarFCB20、McM527、ILSTS087和DRBP1共计16个微卫星座位对11个山羊品种进行了遗传多样性分析,共检测到181个等位基因,平均每个位点检测到11.31个等位基因;品种特有等位基因总计28个。表明这16个FAO—ISAG推荐的微卫星座位能用于我国地方山羊遗传多样性的检测,也表明了我国山羊品种遗传多样性丰富。
2.计算了16个微卫星座位在11个山羊品种的等位基因频率,并以其为基础获得了各群体的平均杂合度、平均多态信息含量和平均有效等位基因数和群体内近交系数,分别为0.605、0.630、3.348和0.126,表明群体内的遗传变异丰富,也表明我国地方山羊品种的遗传多样性比引进品种波尔山羊高。
3群体间基因流分析结果表明太行山羊和河西绒山羊之间有较大的基因流(17.866),太行山羊和黄淮山羊间的基因流次之(16.76),而其它的群体之间的基因流基本在一个较高水平(均值和标准偏差为8.46±4.81):分子方差分析也显示95.79%的遗传变异来源于群体内个体间,只有4.11%的遗传变异来源于不同群体间的分化。上述结果表明这10个地方山羊群体间变异小。
4.本研究的16个微卫星座位在11个山羊群体中总体上极显著偏离哈代-温伯格平衡,总体上处于杂合子缺失状态(P＜0.01);从群体看,16个座位总体上在每个山羊群体都偏离哈代-温伯格平衡。从座位看,11个山羊总体上在CSRD247、ETH10、OARAE54、SRCRSP5、INRABERN172和MCM527座位处于HW平衡(P＞0.05),MAF209、SRCRSP15、ILSTS087和MAF65座位显著的偏离HW平衡(0.05＜P＜0.01),SRCRSP3 TGLA53 DRBP1、OARFCB20、SPS113和SRCRSP8座位极显著的偏离哈代-温伯格平衡(P＜0.01)。连锁平衡检验结果表明在1320对座位配对/群体中,有638对显著或者极显著(P＜0.05或P＜0.01)偏离连锁平衡,但没有任何座位配对出现持续的连锁平衡偏离。
5.采用了基于遗传距离的聚类分析、主成分分析和基于贝叶斯模型的聚类分析三种方法研究了10个地方山羊群体间的遗传关系。计算了每两群体间的Nei氏标准遗传距离(D_S)和,用非加权组对算术平均法(UPGMA)和邻近结合法(NJ)构建了4个11个中外山羊群体的系统进化树图(D_A/NJ、D_s/NJ、D_A/UPGMA和D_s/UPGM图)并得到较高的Bootstrap检验聚类值,4种聚类图大同小异。主成分分析结果得到10个成分,第一主成分和第二主成分分别占45.35%和17.35%,散点图表明引进品种与10个中国地方品种明显分开,10个地方山羊品种遗传分化较低,按主成分分两类,并可细分为4类。基于贝叶斯模型的群体结构分别分析11个山羊群体中的群体和个体的的遗传结构,K=2时,聚类图中波尔山羊很明显的与10个中国地方山羊群体区分开。k=3时,可根据聚类图中的不同颜色填充区域范围将10个中国地方山羊群体划分为两类。K=4,k=5时,群体或者个体Q值普遍变小,难以聚类。基于遗传距离的聚类分析中的Ds/NJ系统进化树图、主成分分析的前两个主成份分析和基于贝叶斯模型聚类分析中的按K=3时个体Q值所得的遗传结构分析这三个结果一致表明,陕南白山羊、济宁青山羊、鲁北白山羊、黄淮山羊聚为一类,沂蒙黑山羊、太行山羊、伏牛白山羊、河西绒山羊、吕梁黑山羊、中卫山羊聚为一类,波尔山羊单独聚为一类。
The domestic goat is one of the important livestock species in China.The size of each breed dramatilly and many of them were endangred in recent years because of introduced commercial breeds.Studying on the genetie diversity of Chinese indigenous goat breeds have been carryed out in nowadays.The microsatellite loci were used in this experiment to analysis the genetic diversity of 530 goats from ten indigenous breeds (Shannan White,Hexi cashmere,Zhongwei,Funiu White,Huanghuai,Jining Grey, Yimeng Black,Lubei White,Lvliang Black,Taihang goat) and one introduced breed (Boer Goat).The research results of this study are as the followings;
1.Based on the 16 microsatellites loci,we analyze the genetic variation of 11 goat populations.Among loci,the mean number of alleles was 11.31,which proved that the microsatellites recommended by FAO-ISAG are useful for the biodiversity studies in Chinese indigenous goat breeds.
2.The observed heterozygosities(Ho),polymorphism information content(PIC), Effective number of alleles(Ne) and inbreeding coefficients(Fis) calculated besed on allele frequency in this 11 breeds is 0.605、0.630、3.348 and 0.126,respectively.The results showed that the genetic variations are abundan in Chinese indigenous goat populations.
3.The level of gene flow from Taihang goat into Hexi cashmere goat is very high, followed that between Taihang goat and Huanghuai goat.the gene flow among others is almost in a same higher level(Mean±SD is 8.46±4.81).The mean Fst(0.64) for only Chinese native goat populations suggests that 93.6%of the total genetic variation was from genetic differentiation within each breed and 6.4%of the genetic variation existed among populations,which was also supported by Analysis of Molecular Variance.
4.The HWE test showed that all loci gave a deviation when analyzed across population.Deviations from HWE across the loci were present in all of the 11 populations. Deviations from HWE across the population were present in MAF209,SRCRSP15, ILSTS087,MAF65,SRCRSP3,TGLA53,DRBP1,OARFCB20,SPS113 and SRCRSP8 (P＜0.05).Out of 1320 locus pair/population combinations,Exact tests for LE between all marker pairs were carried out within each of the populations and resulted in 638 significant(P＜0.05) outcomes,which were sporadically distributed among the populations,but no specific locus pairshowed a consistent deviation from LE that would have been in each,or even in most,of the populations.
5.Three approaches were employed to investigate the genetic relationships among the populations.Firstly,DA and Ds distance matrices were used to build phylogenetic trees with UPGMA and NJ methods.The four Phylogenetic trees are nearly the same. Secondly,in a Principal Component Analysis,the first and second principal component accounted for 45.35%and 17.35%of the variance.The scatterplot revealed that the introduced population is distant from the indigenous population,genetic differentiation among the indigenous breeds is in low level,and the 10 indigenous breeds were clustered into two distinct groups.Thirdly,Population structure analysis based on a Bayesian clustering method showed that a splitting solution was found for K=2 runs in which Boer Goat were identified as distinctive.At K=3,additional structure was detected and the new splitting appeared in Chinese indigeous goat populations.At K=4 or K=5, Inferred membership coefficients of population or individual is very small and could not be applied to clustering well.In a word,the results from Ds/NJ genetic distance cluster, Principal Component Analysis and population genetic structure analysis indicated that the 10 indigenous populations can be clustered into two distinct groups:one group includes Shannan White,Jining Grey,Lubei White,Huanghuai Goat;the other group includes Yimeng Black,Taihang Goat,Funiu White,Hexi cashmere,Lvliang Black,Zhongwei Goat.
1.采用SRCRSP5、MAF209、TGLA53、ETH10、OarAE54、CSRD247、SRCRSP15、SRCRSP3、MAF065、SRCRSP8、SPS113、INRABERN172、OarFCB20、McM527、ILSTS087和DRBP1共计16个微卫星座位对11个山羊品种进行了遗传多样性分析,共检测到181个等位基因,平均每个位点检测到11.31个等位基因;品种特有等位基因总计28个。表明这16个FAO—ISAG推荐的微卫星座位能用于我国地方山羊遗传多样性的检测,也表明了我国山羊品种遗传多样性丰富。
2.计算了16个微卫星座位在11个山羊品种的等位基因频率,并以其为基础获得了各群体的平均杂合度、平均多态信息含量和平均有效等位基因数和群体内近交系数,分别为0.605、0.630、3.348和0.126,表明群体内的遗传变异丰富,也表明我国地方山羊品种的遗传多样性比引进品种波尔山羊高。
3群体间基因流分析结果表明太行山羊和河西绒山羊之间有较大的基因流(17.866),太行山羊和黄淮山羊间的基因流次之(16.76),而其它的群体之间的基因流基本在一个较高水平(均值和标准偏差为8.46±4.81):分子方差分析也显示95.79%的遗传变异来源于群体内个体间,只有4.11%的遗传变异来源于不同群体间的分化。上述结果表明这10个地方山羊群体间变异小。
4.本研究的16个微卫星座位在11个山羊群体中总体上极显著偏离哈代-温伯格平衡,总体上处于杂合子缺失状态(P＜0.01);从群体看,16个座位总体上在每个山羊群体都偏离哈代-温伯格平衡。从座位看,11个山羊总体上在CSRD247、ETH10、OARAE54、SRCRSP5、INRABERN172和MCM527座位处于HW平衡(P＞0.05),MAF209、SRCRSP15、ILSTS087和MAF65座位显著的偏离HW平衡(0.05＜P＜0.01),SRCRSP3 TGLA53 DRBP1、OARFCB20、SPS113和SRCRSP8座位极显著的偏离哈代-温伯格平衡(P＜0.01)。连锁平衡检验结果表明在1320对座位配对/群体中,有638对显著或者极显著(P＜0.05或P＜0.01)偏离连锁平衡,但没有任何座位配对出现持续的连锁平衡偏离。
5.采用了基于遗传距离的聚类分析、主成分分析和基于贝叶斯模型的聚类分析三种方法研究了10个地方山羊群体间的遗传关系。计算了每两群体间的Nei氏标准遗传距离(D_S)和,用非加权组对算术平均法(UPGMA)和邻近结合法(NJ)构建了4个11个中外山羊群体的系统进化树图(D_A/NJ、D_s/NJ、D_A/UPGMA和D_s/UPGM图)并得到较高的Bootstrap检验聚类值,4种聚类图大同小异。主成分分析结果得到10个成分,第一主成分和第二主成分分别占45.35%和17.35%,散点图表明引进品种与10个中国地方品种明显分开,10个地方山羊品种遗传分化较低,按主成分分两类,并可细分为4类。基于贝叶斯模型的群体结构分别分析11个山羊群体中的群体和个体的的遗传结构,K=2时,聚类图中波尔山羊很明显的与10个中国地方山羊群体区分开。k=3时,可根据聚类图中的不同颜色填充区域范围将10个中国地方山羊群体划分为两类。K=4,k=5时,群体或者个体Q值普遍变小,难以聚类。基于遗传距离的聚类分析中的Ds/NJ系统进化树图、主成分分析的前两个主成份分析和基于贝叶斯模型聚类分析中的按K=3时个体Q值所得的遗传结构分析这三个结果一致表明,陕南白山羊、济宁青山羊、鲁北白山羊、黄淮山羊聚为一类,沂蒙黑山羊、太行山羊、伏牛白山羊、河西绒山羊、吕梁黑山羊、中卫山羊聚为一类,波尔山羊单独聚为一类。
The domestic goat is one of the important livestock species in China.The size of each breed dramatilly and many of them were endangred in recent years because of introduced commercial breeds.Studying on the genetie diversity of Chinese indigenous goat breeds have been carryed out in nowadays.The microsatellite loci were used in this experiment to analysis the genetic diversity of 530 goats from ten indigenous breeds (Shannan White,Hexi cashmere,Zhongwei,Funiu White,Huanghuai,Jining Grey, Yimeng Black,Lubei White,Lvliang Black,Taihang goat) and one introduced breed (Boer Goat).The research results of this study are as the followings;
1.Based on the 16 microsatellites loci,we analyze the genetic variation of 11 goat populations.Among loci,the mean number of alleles was 11.31,which proved that the microsatellites recommended by FAO-ISAG are useful for the biodiversity studies in Chinese indigenous goat breeds.
2.The observed heterozygosities(Ho),polymorphism information content(PIC), Effective number of alleles(Ne) and inbreeding coefficients(Fis) calculated besed on allele frequency in this 11 breeds is 0.605、0.630、3.348 and 0.126,respectively.The results showed that the genetic variations are abundan in Chinese indigenous goat populations.
3.The level of gene flow from Taihang goat into Hexi cashmere goat is very high, followed that between Taihang goat and Huanghuai goat.the gene flow among others is almost in a same higher level(Mean±SD is 8.46±4.81).The mean Fst(0.64) for only Chinese native goat populations suggests that 93.6%of the total genetic variation was from genetic differentiation within each breed and 6.4%of the genetic variation existed among populations,which was also supported by Analysis of Molecular Variance.
4.The HWE test showed that all loci gave a deviation when analyzed across population.Deviations from HWE across the loci were present in all of the 11 populations. Deviations from HWE across the population were present in MAF209,SRCRSP15, ILSTS087,MAF65,SRCRSP3,TGLA53,DRBP1,OARFCB20,SPS113 and SRCRSP8 (P＜0.05).Out of 1320 locus pair/population combinations,Exact tests for LE between all marker pairs were carried out within each of the populations and resulted in 638 significant(P＜0.05) outcomes,which were sporadically distributed among the populations,but no specific locus pairshowed a consistent deviation from LE that would have been in each,or even in most,of the populations.
5.Three approaches were employed to investigate the genetic relationships among the populations.Firstly,DA and Ds distance matrices were used to build phylogenetic trees with UPGMA and NJ methods.The four Phylogenetic trees are nearly the same. Secondly,in a Principal Component Analysis,the first and second principal component accounted for 45.35%and 17.35%of the variance.The scatterplot revealed that the introduced population is distant from the indigenous population,genetic differentiation among the indigenous breeds is in low level,and the 10 indigenous breeds were clustered into two distinct groups.Thirdly,Population structure analysis based on a Bayesian clustering method showed that a splitting solution was found for K=2 runs in which Boer Goat were identified as distinctive.At K=3,additional structure was detected and the new splitting appeared in Chinese indigeous goat populations.At K=4 or K=5, Inferred membership coefficients of population or individual is very small and could not be applied to clustering well.In a word,the results from Ds/NJ genetic distance cluster, Principal Component Analysis and population genetic structure analysis indicated that the 10 indigenous populations can be clustered into two distinct groups:one group includes Shannan White,Jining Grey,Lubei White,Huanghuai Goat;the other group includes Yimeng Black,Taihang Goat,Funiu White,Hexi cashmere,Lvliang Black,Zhongwei Goat.
引文
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35.钟金城,王杰,江明峰,晏昭丽,石敏.山谷型藏山羊染色体核型及带型的研究.西南民族学院学报,1995,21(1):40-42
36.Arranz J J,Boyon Y,Primitivo J S Genetic relationships among Spanish sheep using microsate llite, Animal Genetics, 1998, 29: 435-440
37. Barendse W, Armitage S M, Kossarek L M A genetic linkage map of the bovine genome. Nature Genetics, 1994, 6: 227-238
38. Barker J S, Bradley D G,Fries R. An integrated global programme to establish the genetic relationships among the breeds of each domestica animal species Rome:FAO Report, 1993
39. Barker, J S F, Tan S G, Moore, S S ; Mukherjee, T K ; Matheson, J -L ; Selvaraj, O S. Genetic variation within and relationships among populations of Asian goats (Capra hircus). Anim Breed Genet. 2001, 118: 213-233
40. Barker, Tan, Moore, Mukherjee, Matheson, Selvaraj Genetic variation within and relationships among populations of Asian goats (Capra hircus). Animal Breeding and Genetics, 2001, 118 (4):213-233
41. Beckmann, J S , Soller M Toward a Unified approach to genetic mapping of evkaryotes based on sequence tagged microsatellite sites. Genome, 1990, 8: 930-932
42. Bin Fan, Jian-Lin Han, Shi-Lin Chen, David N. Mburu, Olivier Hanotte, Qi-Kang Chen, Shu-Hong Zhao, Kui Li Individual-breed assignments in caprine populations using microsatellite DNA analysis. Small Ruminant Research, 2008, 75: 154-161
43. Buchanan F C, Adams L J, Littlejohn R P, Maddox J F, Crawford A M Detection of evolutionary relationships among sheep breeds using microsatellites. Genomics, 1994, 22: 397-403
44. Canon J D, Garcia M A, Garcia-Atance G, Obexer-Ruff J A, Lenstra P, Ajmone-Marsan, Dunner S, The ECONOGENE Consortium. Geographical partitioning of goat diversity in Europe and the Middle East. Anim Genet, 2006, 37(4): 327-334
45. Cavalli-Sporza L L. The history and geography of Human genes Princeton University Press, Princeton, N J, 1994
46. Chenyambuga S W. Genetic Characterization of indigenous goats of sub-saharan Africa using Microsatellite DNA Markers.Asian-Australasian. Animal Sciences, 2004, 17(4): 1445 .1452
47. Crawford, A M , Dodds, et al An autosomal genetic linkage map of the sheep genome Genetics, 1995, 140: 703-724
48. Davies N, Rraneis X, Villablanea, Roderick G K. Determining the source of individuals: multilocus genotyping in nonequilibrium population genetics. Trends in Ecology and Evolution, 1999, 14(1): 17-21
49. Ellegren H. DNA fingerprinting in horses using a simple (TG)n probe and its application to population comparisons. Animal Genetics, 1992, 23: 1-9
50. Excoffier, L G, Schneider S. Arlequin version3.0: An integrated software package for population genetics data analysis. Evolutionary Bioinformatics Online, 2005, 1: 47-50
51. Falush D, Stephens Mand, pritehard J K. Inference of population strueture using multilocus genotypes data: linked loci and correlated allele frequeneies. Genetics, 2003,164: 1727-1738
52. Felsenstein J, Kishino H. Is there something wrong with the bootstrap. Evolution, 1993(39):783-793
53. Goudet J. FSTAT, a program to estimate and test gene diversities and fixation indices (version 2 9 3) ,2001 [online]:http://www.unil.ch/izea/softwares/fstat.html
54. Guo S W, ThomPson E A. Performing the exact test of Hardy-weinberg proportions for multiple alleles.Biometri, 1992,48: 361-372
55. Hedrick P W. Genetics of Populations. Science Books International, Boston,1983
56. Henderson LM, Bruere A N. Conservation of nucleolus organizer regions during evolution in sheep, goat, cattle and aoudad. Genetic Cytology, 1979, 21: 1-8
57. Huelsenbeck, J P, Ronquist F, Nielsen R, Bollback J P. Bayesian inference of phylogeny and its impact on evolutionary. biology Science, 2001, 94:2310-2214
58. Iamartino D, Bruzzone A, Lanza A, Blasi M, Pilla F. Genetic diversity of Southern Italian goat populations assessed by microsatellite markers. Small Ruminant Research, 2005, 57: 2499-2555
59. Jonathan K, Pritchard, Matthew Stephens, Peter Donnelly. Inference of Population Structure Using Multilocus Genotype Data. Genetics, 2000, 155: 945-959.
60. Ken Nozawa,Yoshiaki Kikkawa, Kenji Tsunoda. Gene constitution of the Yunnan native goat in China. Report of the society for researches on native livestock, 1995, 15
61. Kim, K S, J S Yeo, J W Lee, J W Kim, C B Choi. Genetic giversity of goats from Korea and China using Microsatellite Analysis. Asian-Australasian. Animal Sciences, 2002,(5): 1461-1465
62. Kimura M, Weiss G H. The stepping stone model of population structure and the decrease of genetic correlation with distance. Genetics, 1964,49: 561-576
63. Li M H, Zhao S H, Bian C, Wang H S, Wei H, Liu B, Yu M, Fan B, Chen S L, Zhu M J, Li S J, Xiong T A, Li K Genetic relationship s among twelve Chinese indigenous goat populations based on microsatellite analysis. Genet Sel Evol, 2002, 34: 729-744.
64. Li X L, Valentini A. Genetic diversity of Chinese indigenous goat breeds based on microsatellite markers. Animal Breeding and Genetics,2004,21(5): 350-355
65. Luikart G, Biju-Duval MP, Ertugrul O, Zagdsuren Y, Maudet C, Taberlet P Power of 22 microsatellite markers in fluorescent multiplexes for parentage testing in goats (Capra hircus). Anim Genet, 1999, 30(6): 431-438
66. Maudet C, Miller C, Bassano B, Breitenmoser-Wursten C, Gauthier D, Obexer-Ruff G, Michallet J, Taberlet P, Luikart G. Microsatellite DNA and recent statistical methods in wildlife conservation management: applications in Alpine ibex [Capra ibex (ibex)]. Molecular Ecology, 2002,11(3): 421
67. Moore S S, Sargeant L L, King T G. The conservation of dinucleotide microsatellites among mammaliam genomes allows the use of heterologous PCR primer pars in closely related species. Genomics, 1991,10: 654-670
68. Mullis K, Faloona F, Scharf S, Saiki R, Horn G, Erlich H. Specific enzymatic amplification of DNA in vitro: the polymerase chain reaction. Cold Spring Harb Symp Quant Biol. 1986,51(1):263-273
69. Murra V, Monchawin C, England P R. The determination of the sequences present in the shadow bands of a dinucleotide repeatPCR. Nucleic Acids Research, 1993, 21(10): 2395-2398
70. Nei M, Grauer D. Extent of protein polymorphism and the neutral mutation theory. Evol Biol, 1984, 17:3-118.
71. Nei M, Li W H. Linkage disequilibrium in subdivided populations. Genetics, 1973, 75:213-219.
72. Nei M. Molecular Evolutionary Genetics. Columbia University Press, New York, USA 1987: 176-187
73. Olufowote J O, Xu Y, Chen X, Park WD, Beachell HM, Dilday RH, Goto M, McCouch SR. Comparative evaluation of within-cultivar variation of rice (Oryza sativa L) using microsatellite and RFLP markers. Genome. 1997, 40(3): 370-378.
74. PeL NguyenT C.Blood groups and Protein Polymorphisms in five goat breeds (Capra heraeus). Animal Geneties. 1994, 23: 333-336
75. Pritchard,J K ,M Stephens,and P Donnelly,Inference of population structure using multilocus genotype data. Genetics,2000 15 5(2): 94 -95
76. R Z Ma, Russ I, Park C, Heyen D W, Beever J E, Green C A, Lewin H A. Isolation and characterization of 45 polymorphic microsatellites from the bovine' genome. Animal Genetics, 1996, 27(1): 43-47
77. Rannala B and Z Yang 1996 Probability distribution of molecular evolutionary trees : a new method of phylogenetic inference. Mol Evol, 43 : 304-311
78. Raymond M., Rousset F, GENEPOP (version 1.2): population genetics software for exact tests and ecumenicism. Heredity. 1995,86:248-249
79. Rout, P K, M B Joshi, et al (2008) Microsatellite-based phylogeny of Indian domestic goats. BMC Genet, 9(1): 11
80. Saitbekova N, Gaillard C, Obexer-Ruff G, Dolf G. Genetic diversity in Swiss goat breeds based on microsatellite analysis . Anim Genet, 1999, 30(1): 36-41
81. Schaller, G B. Mountain monarchs: wild sheep and goats of the Himalaya Univ Chicago Press, 1977:425
82. Shriver M D VNIT allele frequency distributions under the stepwise mutation model: a computer simulation approach. Genetics, 1993, 24: 345-348
83. Slatkin, M.. Gene flow and the geographic structure of natural populations. Science 1987, 236(6): 787-791.
84. Slatkin, M. and N.H. Barton. A comparison of three indirect methods for estimating average levels of gene flow. Evolution, 1989, 43(7):1349-1368.
85. Slatkin M Isolation by distance in equilibrium and non-equilibrium population Evolution,1993,47: 262-279
86. Sun W, Chang H, Ren Z J, Yang Z P, Geng R Q, Lu S X, Du L, Tsunoda K. Genetic diffentiation between sheep and goat based on Microsatellite DNA. Aust J Anim Sci, 2004, 17: 583-587
87. Sunden S L , Fstone R T, Bishop M D ,et al A highly polymorphic bovine microsatellite locus BM2113. Anim Genet, 1993, 24: 69-73
88. Tautz D Hypervariability of simple sequences as a general source for polymorphic DNA markers. Nuleic Acids Res, 1989,17(6): 6463-6467
89. Tucker E M, Young J C Genetic variation in the purine nucleioside phosphorlase activity of good red cells Biochemical Genetics. Animal Blood Groups, 1976, 7: 109-117
90. Tunon M J, Gonzale Z P, Vallejo M Genetic relationships between fourteen native Spanish breeds of goat. Animal Genetics, 1989, 20: 205-212
91. Upholt W B, David I B Mapping of mitochondrial DNA of individual sheep and goats: Rapid evolution in the D loop region .Cell, 1977, 11:571-583
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98. Yang, Z , Rannala B. Bayesian phylogenetic inference using DNA sequences : A Markov chain Monte Carlo method. Mol Evol, 1997,14: 717-724
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33.中国畜禽遗传资源状况编委会.中国畜禽遗传资源状况.中国农业出版社,2004:164-179
34.中国羊品种志编写组.中国羊品种志.上海上海科学技术出版社,1989:15-19,27-58
35.钟金城,王杰,江明峰,晏昭丽,石敏.山谷型藏山羊染色体核型及带型的研究.西南民族学院学报,1995,21(1):40-42
36.Arranz J J,Boyon Y,Primitivo J S Genetic relationships among Spanish sheep using microsate llite, Animal Genetics, 1998, 29: 435-440
37. Barendse W, Armitage S M, Kossarek L M A genetic linkage map of the bovine genome. Nature Genetics, 1994, 6: 227-238
38. Barker J S, Bradley D G,Fries R. An integrated global programme to establish the genetic relationships among the breeds of each domestica animal species Rome:FAO Report, 1993
39. Barker, J S F, Tan S G, Moore, S S ; Mukherjee, T K ; Matheson, J -L ; Selvaraj, O S. Genetic variation within and relationships among populations of Asian goats (Capra hircus). Anim Breed Genet. 2001, 118: 213-233
40. Barker, Tan, Moore, Mukherjee, Matheson, Selvaraj Genetic variation within and relationships among populations of Asian goats (Capra hircus). Animal Breeding and Genetics, 2001, 118 (4):213-233
41. Beckmann, J S , Soller M Toward a Unified approach to genetic mapping of evkaryotes based on sequence tagged microsatellite sites. Genome, 1990, 8: 930-932
42. Bin Fan, Jian-Lin Han, Shi-Lin Chen, David N. Mburu, Olivier Hanotte, Qi-Kang Chen, Shu-Hong Zhao, Kui Li Individual-breed assignments in caprine populations using microsatellite DNA analysis. Small Ruminant Research, 2008, 75: 154-161
43. Buchanan F C, Adams L J, Littlejohn R P, Maddox J F, Crawford A M Detection of evolutionary relationships among sheep breeds using microsatellites. Genomics, 1994, 22: 397-403
44. Canon J D, Garcia M A, Garcia-Atance G, Obexer-Ruff J A, Lenstra P, Ajmone-Marsan, Dunner S, The ECONOGENE Consortium. Geographical partitioning of goat diversity in Europe and the Middle East. Anim Genet, 2006, 37(4): 327-334
45. Cavalli-Sporza L L. The history and geography of Human genes Princeton University Press, Princeton, N J, 1994
46. Chenyambuga S W. Genetic Characterization of indigenous goats of sub-saharan Africa using Microsatellite DNA Markers.Asian-Australasian. Animal Sciences, 2004, 17(4): 1445 .1452
47. Crawford, A M , Dodds, et al An autosomal genetic linkage map of the sheep genome Genetics, 1995, 140: 703-724
48. Davies N, Rraneis X, Villablanea, Roderick G K. Determining the source of individuals: multilocus genotyping in nonequilibrium population genetics. Trends in Ecology and Evolution, 1999, 14(1): 17-21
49. Ellegren H. DNA fingerprinting in horses using a simple (TG)n probe and its application to population comparisons. Animal Genetics, 1992, 23: 1-9
50. Excoffier, L G, Schneider S. Arlequin version3.0: An integrated software package for population genetics data analysis. Evolutionary Bioinformatics Online, 2005, 1: 47-50
51. Falush D, Stephens Mand, pritehard J K. Inference of population strueture using multilocus genotypes data: linked loci and correlated allele frequeneies. Genetics, 2003,164: 1727-1738
52. Felsenstein J, Kishino H. Is there something wrong with the bootstrap. Evolution, 1993(39):783-793
53. Goudet J. FSTAT, a program to estimate and test gene diversities and fixation indices (version 2 9 3) ,2001 [online]:http://www.unil.ch/izea/softwares/fstat.html
54. Guo S W, ThomPson E A. Performing the exact test of Hardy-weinberg proportions for multiple alleles.Biometri, 1992,48: 361-372
55. Hedrick P W. Genetics of Populations. Science Books International, Boston,1983
56. Henderson LM, Bruere A N. Conservation of nucleolus organizer regions during evolution in sheep, goat, cattle and aoudad. Genetic Cytology, 1979, 21: 1-8
57. Huelsenbeck, J P, Ronquist F, Nielsen R, Bollback J P. Bayesian inference of phylogeny and its impact on evolutionary. biology Science, 2001, 94:2310-2214
58. Iamartino D, Bruzzone A, Lanza A, Blasi M, Pilla F. Genetic diversity of Southern Italian goat populations assessed by microsatellite markers. Small Ruminant Research, 2005, 57: 2499-2555
59. Jonathan K, Pritchard, Matthew Stephens, Peter Donnelly. Inference of Population Structure Using Multilocus Genotype Data. Genetics, 2000, 155: 945-959.
60. Ken Nozawa,Yoshiaki Kikkawa, Kenji Tsunoda. Gene constitution of the Yunnan native goat in China. Report of the society for researches on native livestock, 1995, 15
61. Kim, K S, J S Yeo, J W Lee, J W Kim, C B Choi. Genetic giversity of goats from Korea and China using Microsatellite Analysis. Asian-Australasian. Animal Sciences, 2002,(5): 1461-1465
62. Kimura M, Weiss G H. The stepping stone model of population structure and the decrease of genetic correlation with distance. Genetics, 1964,49: 561-576
63. Li M H, Zhao S H, Bian C, Wang H S, Wei H, Liu B, Yu M, Fan B, Chen S L, Zhu M J, Li S J, Xiong T A, Li K Genetic relationship s among twelve Chinese indigenous goat populations based on microsatellite analysis. Genet Sel Evol, 2002, 34: 729-744.
64. Li X L, Valentini A. Genetic diversity of Chinese indigenous goat breeds based on microsatellite markers. Animal Breeding and Genetics,2004,21(5): 350-355
65. Luikart G, Biju-Duval MP, Ertugrul O, Zagdsuren Y, Maudet C, Taberlet P Power of 22 microsatellite markers in fluorescent multiplexes for parentage testing in goats (Capra hircus). Anim Genet, 1999, 30(6): 431-438
66. Maudet C, Miller C, Bassano B, Breitenmoser-Wursten C, Gauthier D, Obexer-Ruff G, Michallet J, Taberlet P, Luikart G. Microsatellite DNA and recent statistical methods in wildlife conservation management: applications in Alpine ibex [Capra ibex (ibex)]. Molecular Ecology, 2002,11(3): 421
67. Moore S S, Sargeant L L, King T G. The conservation of dinucleotide microsatellites among mammaliam genomes allows the use of heterologous PCR primer pars in closely related species. Genomics, 1991,10: 654-670
68. Mullis K, Faloona F, Scharf S, Saiki R, Horn G, Erlich H. Specific enzymatic amplification of DNA in vitro: the polymerase chain reaction. Cold Spring Harb Symp Quant Biol. 1986,51(1):263-273
69. Murra V, Monchawin C, England P R. The determination of the sequences present in the shadow bands of a dinucleotide repeatPCR. Nucleic Acids Research, 1993, 21(10): 2395-2398
70. Nei M, Grauer D. Extent of protein polymorphism and the neutral mutation theory. Evol Biol, 1984, 17:3-118.
71. Nei M, Li W H. Linkage disequilibrium in subdivided populations. Genetics, 1973, 75:213-219.
72. Nei M. Molecular Evolutionary Genetics. Columbia University Press, New York, USA 1987: 176-187
73. Olufowote J O, Xu Y, Chen X, Park WD, Beachell HM, Dilday RH, Goto M, McCouch SR. Comparative evaluation of within-cultivar variation of rice (Oryza sativa L) using microsatellite and RFLP markers. Genome. 1997, 40(3): 370-378.
74. PeL NguyenT C.Blood groups and Protein Polymorphisms in five goat breeds (Capra heraeus). Animal Geneties. 1994, 23: 333-336
75. Pritchard,J K ,M Stephens,and P Donnelly,Inference of population structure using multilocus genotype data. Genetics,2000 15 5(2): 94 -95
76. R Z Ma, Russ I, Park C, Heyen D W, Beever J E, Green C A, Lewin H A. Isolation and characterization of 45 polymorphic microsatellites from the bovine' genome. Animal Genetics, 1996, 27(1): 43-47
77. Rannala B and Z Yang 1996 Probability distribution of molecular evolutionary trees : a new method of phylogenetic inference. Mol Evol, 43 : 304-311
78. Raymond M., Rousset F, GENEPOP (version 1.2): population genetics software for exact tests and ecumenicism. Heredity. 1995,86:248-249
79. Rout, P K, M B Joshi, et al (2008) Microsatellite-based phylogeny of Indian domestic goats. BMC Genet, 9(1): 11
80. Saitbekova N, Gaillard C, Obexer-Ruff G, Dolf G. Genetic diversity in Swiss goat breeds based on microsatellite analysis . Anim Genet, 1999, 30(1): 36-41
81. Schaller, G B. Mountain monarchs: wild sheep and goats of the Himalaya Univ Chicago Press, 1977:425
82. Shriver M D VNIT allele frequency distributions under the stepwise mutation model: a computer simulation approach. Genetics, 1993, 24: 345-348
83. Slatkin, M.. Gene flow and the geographic structure of natural populations. Science 1987, 236(6): 787-791.
84. Slatkin, M. and N.H. Barton. A comparison of three indirect methods for estimating average levels of gene flow. Evolution, 1989, 43(7):1349-1368.
85. Slatkin M Isolation by distance in equilibrium and non-equilibrium population Evolution,1993,47: 262-279
86. Sun W, Chang H, Ren Z J, Yang Z P, Geng R Q, Lu S X, Du L, Tsunoda K. Genetic diffentiation between sheep and goat based on Microsatellite DNA. Aust J Anim Sci, 2004, 17: 583-587
87. Sunden S L , Fstone R T, Bishop M D ,et al A highly polymorphic bovine microsatellite locus BM2113. Anim Genet, 1993, 24: 69-73
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