CLPG和BMP15基因多态性与北疆7个绵羊群体生产性能关系的初步研究
摘要
本试验分析了中国美利奴母羔羊在4月龄断奶前6项体尺性状之间的关系，结果显示在不同的日龄时羔羊的胸围、胸宽、管围和体重之间有较高水平的正相关，相关系数在0.5-1.0之间。对不同日龄各项体尺与体重建立的了回归方程，结果表明，60日龄回归模型的决定系数R~2=0.452，其它6个回归模型的相关指数均在0.7以上，其中方程1、5、6、7的相关指数都在0.9以上，表明这些回归方程中各项体尺指标与体重之间相关密切，根据胸围、管围、体高和胸宽对体重的选择都是有效的，具有一定的参考作用。
本试验对6个绵羊群体(中国美利奴、萨福克羊、萨福克羊×新疆细毛羊母羊F1代杂种羊、多赛特羊、多赛特公羊×新疆细毛羊母羊F1代杂种羊和夏洛来羊)中3个微卫星位点(OY3、BMS1516和IDVGA30)的遗传变异(基因频率、杂合度、PIC和有效等位基因数)进行了初步的分析研究。结果显示3个微卫星在6个群体中的PIC、杂合度和有效等位基因数分别为0.5253-0.7243、0.6425-0.7682和2.1624-4.4705。根据这3个微卫星多态计算的各群体间的遗传距离绘制的系统聚类图，陶赛特羊和萨福克羊先分别和各自与新疆细毛羊的杂一代聚在一起(0.0117、0.0145)，然后它们再与夏洛来羊聚在一起，中国美利奴与夏洛来羊的遗传距离最大(0.6621)。所建系统聚类图符合其来源分类和育成史，能比较准确地反映这6个绵羊群体的地理分布和它们之间的亲缘关系。
通过对3个微卫星不同基因型与中国美利奴羔羊各项体尺指标进行方差分析和多重比较，结果表明3个微卫星的不同基因型对中国美利奴不同日龄的体尺和体重在0.1水平上有显著影响。其中微卫星座位IDVGA30 AB基因型90日龄平均体重最大(19.73kg)，DD基因型90日龄平均体长最大(56.00cm)；微卫星座位BMS1516各基因型对90日龄中国美利奴羔羊体尺的影响较大，其中基因型EF的胸深最大(22.44cm)，基因型CD胸围最大(62.280m)，管围最大的为CD(8.07cm)：微卫星座位OY3不同基因型初生重差异显著(P＜0.1)，基因型EF的初生重最大(4.10kg)，CD基因型初生重最小(3.42kg)。
本研究采用PCR-RFLP、SSCP分析了CLPG基因的两个SNP位点，CLPG基因的两个位点均表现出SSCP多态性。对影响绵羊排卵率的BMP15用Forced-RFLP分析后未发现突变型个体。
The body sizes data (body weight, body high, body length, heart girth, chest width, chest depth and circumference of cannon bone) of Chinese Merino lamb before being weaned were analyzed with the methods of correlations and regressions. The result showed that the single correlation between the characters of weight and heart girth, chest width and circumference of cannon bone was significantly different. The correlation between weight and heart girth, chest width, circumference of cannon bone was highly positive correlation. Among seven regression models, the correlation index of six regression models was over 0.7, but that of 60 days' was 0.452. The result suggested that selecting body weight was efficient through heart girth, body high, chest width and circumference of cannon bone.
The genetic variation (gene frequency, heterozygosity, polymorphic information content(PIC) and effective number of alleles) of 3 microsatellite loci(IDVGA30, BMS1516, OY3) was analyzed in 6 sheep populations (Chinese Merino, Suffolk, Dorset, FI of Suffolk crossed with Xinjiang Fine Wool, F] of Doset crossed with Xinjiang Fine Wool and Charolais sheep). The result showed that PIC, heterozygosity and effective number of alleles of 3 microsatellite loci ranged from 0.5253 to 0.7243, from 0.6425 ti 0.7682, from 2.1624 to 4.4705, respectively. The clustering dendrogram from genetic distance of the three microsatellites could correctly reflect the geographical distribution of and the relationship within the six sheep populations. The genetic distances of Chinese Merino between foreign sheep breeds and cultivated breeds bearing foreign bloodline were relatively large. Consequently, they clustered in two groups. The phylogenetic relationship between different sheep breeds was in accordance with their resource, bree
ding history, differentiation and locations.
The relationship between different genetype of 3 microsatellites and body sizes in Chinese Merino lamb was analyzed. The mean weight of genetype AB(19.73kg) was the highest and the mean body length of genetype DD(56.00cm ) was the longest in microsatellite IDVGA30 in 90 days old. In microsatellite BMS1516, the chest high of genetype EF(22.44cm), the heart girth (62.28cm)and the circumference of cannon bone (8.07cm) of genetype CD were the highest in Chinese Merino lamb of 90 days old. In microsatellite OY3, the born weight of genetype EF was the highest(4.10kg).
Tow SNP locus of CLPG gene were analyzed through the method of PCR-RFLP and PCR-SSCP. They all showed polymorphism of SSCP. The mutatant genetype was
not detected through the Forced-RFLP when analyzing the gene BMPl5.
本试验对6个绵羊群体(中国美利奴、萨福克羊、萨福克羊×新疆细毛羊母羊F1代杂种羊、多赛特羊、多赛特公羊×新疆细毛羊母羊F1代杂种羊和夏洛来羊)中3个微卫星位点(OY3、BMS1516和IDVGA30)的遗传变异(基因频率、杂合度、PIC和有效等位基因数)进行了初步的分析研究。结果显示3个微卫星在6个群体中的PIC、杂合度和有效等位基因数分别为0.5253-0.7243、0.6425-0.7682和2.1624-4.4705。根据这3个微卫星多态计算的各群体间的遗传距离绘制的系统聚类图，陶赛特羊和萨福克羊先分别和各自与新疆细毛羊的杂一代聚在一起(0.0117、0.0145)，然后它们再与夏洛来羊聚在一起，中国美利奴与夏洛来羊的遗传距离最大(0.6621)。所建系统聚类图符合其来源分类和育成史，能比较准确地反映这6个绵羊群体的地理分布和它们之间的亲缘关系。
通过对3个微卫星不同基因型与中国美利奴羔羊各项体尺指标进行方差分析和多重比较，结果表明3个微卫星的不同基因型对中国美利奴不同日龄的体尺和体重在0.1水平上有显著影响。其中微卫星座位IDVGA30 AB基因型90日龄平均体重最大(19.73kg)，DD基因型90日龄平均体长最大(56.00cm)；微卫星座位BMS1516各基因型对90日龄中国美利奴羔羊体尺的影响较大，其中基因型EF的胸深最大(22.44cm)，基因型CD胸围最大(62.280m)，管围最大的为CD(8.07cm)：微卫星座位OY3不同基因型初生重差异显著(P＜0.1)，基因型EF的初生重最大(4.10kg)，CD基因型初生重最小(3.42kg)。
本研究采用PCR-RFLP、SSCP分析了CLPG基因的两个SNP位点，CLPG基因的两个位点均表现出SSCP多态性。对影响绵羊排卵率的BMP15用Forced-RFLP分析后未发现突变型个体。
The body sizes data (body weight, body high, body length, heart girth, chest width, chest depth and circumference of cannon bone) of Chinese Merino lamb before being weaned were analyzed with the methods of correlations and regressions. The result showed that the single correlation between the characters of weight and heart girth, chest width and circumference of cannon bone was significantly different. The correlation between weight and heart girth, chest width, circumference of cannon bone was highly positive correlation. Among seven regression models, the correlation index of six regression models was over 0.7, but that of 60 days' was 0.452. The result suggested that selecting body weight was efficient through heart girth, body high, chest width and circumference of cannon bone.
The genetic variation (gene frequency, heterozygosity, polymorphic information content(PIC) and effective number of alleles) of 3 microsatellite loci(IDVGA30, BMS1516, OY3) was analyzed in 6 sheep populations (Chinese Merino, Suffolk, Dorset, FI of Suffolk crossed with Xinjiang Fine Wool, F] of Doset crossed with Xinjiang Fine Wool and Charolais sheep). The result showed that PIC, heterozygosity and effective number of alleles of 3 microsatellite loci ranged from 0.5253 to 0.7243, from 0.6425 ti 0.7682, from 2.1624 to 4.4705, respectively. The clustering dendrogram from genetic distance of the three microsatellites could correctly reflect the geographical distribution of and the relationship within the six sheep populations. The genetic distances of Chinese Merino between foreign sheep breeds and cultivated breeds bearing foreign bloodline were relatively large. Consequently, they clustered in two groups. The phylogenetic relationship between different sheep breeds was in accordance with their resource, bree
ding history, differentiation and locations.
The relationship between different genetype of 3 microsatellites and body sizes in Chinese Merino lamb was analyzed. The mean weight of genetype AB(19.73kg) was the highest and the mean body length of genetype DD(56.00cm ) was the longest in microsatellite IDVGA30 in 90 days old. In microsatellite BMS1516, the chest high of genetype EF(22.44cm), the heart girth (62.28cm)and the circumference of cannon bone (8.07cm) of genetype CD were the highest in Chinese Merino lamb of 90 days old. In microsatellite OY3, the born weight of genetype EF was the highest(4.10kg).
Tow SNP locus of CLPG gene were analyzed through the method of PCR-RFLP and PCR-SSCP. They all showed polymorphism of SSCP. The mutatant genetype was
not detected through the Forced-RFLP when analyzing the gene BMPl5.
引文
[1] 刘守仁，吕维斌．《绵羊学》[M]．新疆科技卫生出版社：1996年6月．2-15．
[2] 傅衍等．中国家鸡的起源探讨[J]．遗传学报，2001，28(5)：411-417．
[3] 张果平等．绵羊FecX~Ⅰ基因对排卵率调控的研究进展[J]．四川畜牧兽医，2002，29(8)：141-143．
[4] 郭万库，吴常信等．肉用动物双肌性状的研究现状[J]．国外畜牧科技．1999，26(3)：34-38．
[5] Freking B. A. Identification of the single base change causing the Callipyge muscle hypertrophy phenotype the only known example of polar overdominance in mammals [J]. Genome Research, 2002, 12: 1496-1506.
[6] Cockett, N, Jackson, S P, et al, Chromosomal localization of the callipyge gene in sheep (Ovisaries) using bovine DNA markers [J]. Proc. Natl. Acad. Sci. 1994, 91: 3019-3023.
[7] Cockett N, Shay T L, Localization of the locus causing spider lamb syndrome to the distal end of ovine chromosome 6 [J]. Mammal Genome. 1999, 10: 35-38.
[8] Fahrenkrug S C, Freking B A, Comparative mapping of the ovine CLPG Iocus[J]. Mammal Genome. 2000, 11: 871-876.
[9] Freking B A, Keele J W, Evaluation of the ovine callipyge locus: Ⅱ. Genotypic effects on growth, slaughter, and carcass traits [J]. Journal of Animal Science. 1998, 76: 2549-2559.
[10] Jackson, S P, Green R D. Muscle trait inheritance growth performance and feed efficiency of sheep exhibiting a muscle hypertrophy phenotype[J]. Journal of Animal Science. 1993, 71: 241(Abstract)
[11] Carpenter C E, Histology and composition of muscles from normal and callipyge lambs[J]. Journal of Animal Science. 1997, 75: 14-18
[12] Shey T, Berghmans S, Segers K. Fine-mapping and construction of a bovine contig spannig the ovine callipyge locus. Mammal Genome. 2001, 12: 141-149.
[13] Lien S, Cockett N E, Klungland H. High-resolution gametic map of the sheep callipyge region: linkage heterogeneity among rams detected by sperm type [J]. Animal Genetics, 1999, 30: 42-46
[14] Davis GH, Evidence for the presence of a major gene influencing ovulation rate on the X chromosome of sheep [J]. Biology of Reproduction. 1991, 44: 620-624
[15] Davis GH, Infertility due to bilateral ovarian hypoplasia in sheep homozygous (FecX~Ⅰ FecX~Ⅰ) for the Inverdale prolificacy gene located on the X chromosome [J]. Biology of Reproduction. 1992, 46: 636-640
[16] Galloway S M, Bmp 15 mutations and ovarian function [J]. Molecular and Cellular Endocrinology, 2002, 191, 15-18
[17] Galloway SM, A linkage map of the ovine X chromosome [J]. Genome Research. 1996, 6: 667-677
[18] 管锋，杨利国，石国庆等．绵羊多胎性状基因调控研究进展[J]．黑龙江畜牧兽医，2003，11(3)：12—17
[19] Jennifer L, Dube, Pei Wang, The bone Morphogenetic Protein 15 geneis X-Linked and Expressed in Oocytes [J]. Molecular Endocrinology. 2002, 12(12): 1809-1817.
[20] 张果平，黄永宏，李峰，绵羊多胎性FecB基因研究新进展[J]．动物科学与动物医学，2002，19(9)：5—8
[21] 石国庆，缪礼维等．新疆细毛羊发展回顾与展望．中国草食动物[J]．2001-2002年全国养羊生产与学术研讨会议论文集：18-23
[22] 徐宏发等．分子系统学研究进展[J]，生态学杂志．2001．20(3)：41-46。
[23] 李学斌．分子标记回归模型与显性分子标记资料[J]．动物科学与动物医学．2002，19(6)：5-9
[24] 赵宗胜，李大全等．杂交羔羊体尺与断奶体重间的多元回归分析[J]．黑龙江畜牧兽医．1998，12：5-6
[25] 赵宗胜，李大全等．杂交肉羊不同日龄体重间的回归分析[J]．黑龙江畜牧兽医．1998，10：11-14
[26] 王大广，苏秀侠等．萨福克肉用种羊线性相关与回归分析[J]。吉林农业科学．2002，27(1)：47-49
[27] 王同军．建昌黑山羊体尺体重相关性研究[J]．四川畜牧兽医．2003，30(9)：22-23．26
[28] 买买提明巴拉提等．麦盖提羊羔羊初生体重、体尺及其相关性的研究[J]．黑龙江畜牧兽医．2001，5：9-10
[29] 赵素君．微卫星标记及其与畜禽遗传育种中的应用[J]．西南民族大学学报(自然科学版)．2003，29(4)：499-502
[30] 樊斌，李奎，彭中镇，1999，微卫星标记检测遗传多样性的原理和方法[J]．第十次全国动物遗传育种学术讨论会论文．北京：中国农业科技出版社P32-36．
[31] 杨章平，常洪等．绵羊和山羊基于两种标记的遗传分化初步研究[J]．扬州大学学报．2003，24(4)：27-31
[32] Takezake N, Genetic Distance and Reconstruction of Phylogenetic Tree From Microsatellite DNA [J]. Genetics. 1996, 144: 389-399.
[33] Zeveren, A V, Peelman L, Weghe A, A genetic study of four Belgian pig populations
by means of seven microsatellite loci [J]. Anita. Breed. Genet. 1995, 112: 191-204
[34] 候冠玉，马月辉．微卫星DNA在草食动物遗传育种中的研究与应用[J]．中国草食动物．2003，23(3)：39-40
[35] 储明星，王吉振等，小尾寒羊五个微卫星基因座遗传多态性研究[J]．遗传学报，2002，2(6)：502—506
[36] 杨章平，常洪等．结构基因座和微卫星标记应用于绵(山)羊群体遗传分化的比较研究[J]．畜牧兽医学报．2003，34(5)：427-433
[37] 代江生，刘云芳等．五个绵羊品种的RAPD初步分析[J]．草食家畜．1998，1：11-13
[38] Jia Bin, Chen Jie et ai. , Microsatellite analysis of genetic diversity and phylogenetic relationship of eight sheep breeds in Xinjiang [J]. 遗传学报. 2003, 30 (9): 847-854.
[39] Berghmans, S., Segers, K., et al., Breakpioint mapping positions the callipyge gene within a 450-kilobase chromosome segment containing the DLK1 and GTL2 gene [J]. Mammal Genome. 2001, 12: 183-185.
[40] Bidwell, C. A., Shay, T. L. , et al. , Differential expression of the GTL2gene within the callipyge region ofovine chromosome 18 [J]. Animal Genetics. 2001, 32: 248-256.
[41] Charlier, C., Segers, K., et al., The callipyge mutation enhances the expression of coregulated imprinted genes in cis without affecting their imprinting status [J]. Mammal Genetic. 2001, 27: 367-369.
[42] Maria Smit, Karin Segers, et al., Mosaicism of Solid Gold Supports the causality of a moneoding A-to-G transition in the eterminism of the Callipyge phenotype [J]. Genetics. 2003, 163: 453-456.
[43] Segers, K., Vaiman, D. et al., Construction and characterizaton of an ovine BAC eontig spanning the eallipyge locus [J]. Animal Genetic. 2001, 31: 352-359.
[44] Hayashi K．PCR—SSCP．一种检测突变的方法[J]．国外医学卫生学分册．1994，21(4)：204-205
[45] Fahrenkrug, S C, Freking B A, Single nucleotide polymorphism (SNP) discovery in porcine expressed genes [J]. Animal Genetic. 2002, 33: 186-195.
[46] Freking B A, Keele J W, Evaluation of the ovine eallipyge locus: I. Relative chromosomal position and gene action [J] . Journal of Animal Science. 1998, 76: 2062-2071.
[47] Mulsant P, Lecerf F, Mutation in borne morphogenetic protein receptor-IB is associated with increased ovulation rate in Booroola Merino ewes[J]. Proc. Natl. Acad. Sci. 2001, 98: 5104-5109.
[48] Susan M. Galloway, Mutation in an oocyte-derived growth factor gene (BMP 15) can increased ovulation rate and infertileity in a dosage-sensitive manner [J]. Nature Genetic, 2000, 25.
[49] 柳淑芳，闫艳春，杜立新，FecB基因及BMPR—IB基因的突变特性与生物学意义[J]．遗传，
2003，25(1)：93—96．
[50] 王根林，毛鑫智等，DNA分析了现我国湖羊和小尾寒羊存在Booroola(FecB)多胎基因[J]．南京农业大学学报，2003，26(1)：104—108
[51] 钟发刚等．绵羊BMPR-IB基因多态性与其产羔数的相关研究[J]．草食家畜[J]．2003，2：20-23
[52] 陈晓军，钟发刚，罗淑萍，王新华，多浪羊BMPR-IB基因多态性的初步研究[J]．新疆农业科学．2004，41(1)：6—9．
[53] 储明星，程国华等，微卫星标记OarAE101和BM1329在五个绵羊品种中的初步研究[J]．遗传学报，2001，28(6)：510—517
[54] 储明星，王吉振等，小尾寒羊4个微卫星座位的克隆及序列分析[J]．遗传学报，2002，29(5)：402—405
[2] 傅衍等．中国家鸡的起源探讨[J]．遗传学报，2001，28(5)：411-417．
[3] 张果平等．绵羊FecX~Ⅰ基因对排卵率调控的研究进展[J]．四川畜牧兽医，2002，29(8)：141-143．
[4] 郭万库，吴常信等．肉用动物双肌性状的研究现状[J]．国外畜牧科技．1999，26(3)：34-38．
[5] Freking B. A. Identification of the single base change causing the Callipyge muscle hypertrophy phenotype the only known example of polar overdominance in mammals [J]. Genome Research, 2002, 12: 1496-1506.
[6] Cockett, N, Jackson, S P, et al, Chromosomal localization of the callipyge gene in sheep (Ovisaries) using bovine DNA markers [J]. Proc. Natl. Acad. Sci. 1994, 91: 3019-3023.
[7] Cockett N, Shay T L, Localization of the locus causing spider lamb syndrome to the distal end of ovine chromosome 6 [J]. Mammal Genome. 1999, 10: 35-38.
[8] Fahrenkrug S C, Freking B A, Comparative mapping of the ovine CLPG Iocus[J]. Mammal Genome. 2000, 11: 871-876.
[9] Freking B A, Keele J W, Evaluation of the ovine callipyge locus: Ⅱ. Genotypic effects on growth, slaughter, and carcass traits [J]. Journal of Animal Science. 1998, 76: 2549-2559.
[10] Jackson, S P, Green R D. Muscle trait inheritance growth performance and feed efficiency of sheep exhibiting a muscle hypertrophy phenotype[J]. Journal of Animal Science. 1993, 71: 241(Abstract)
[11] Carpenter C E, Histology and composition of muscles from normal and callipyge lambs[J]. Journal of Animal Science. 1997, 75: 14-18
[12] Shey T, Berghmans S, Segers K. Fine-mapping and construction of a bovine contig spannig the ovine callipyge locus. Mammal Genome. 2001, 12: 141-149.
[13] Lien S, Cockett N E, Klungland H. High-resolution gametic map of the sheep callipyge region: linkage heterogeneity among rams detected by sperm type [J]. Animal Genetics, 1999, 30: 42-46
[14] Davis GH, Evidence for the presence of a major gene influencing ovulation rate on the X chromosome of sheep [J]. Biology of Reproduction. 1991, 44: 620-624
[15] Davis GH, Infertility due to bilateral ovarian hypoplasia in sheep homozygous (FecX~Ⅰ FecX~Ⅰ) for the Inverdale prolificacy gene located on the X chromosome [J]. Biology of Reproduction. 1992, 46: 636-640
[16] Galloway S M, Bmp 15 mutations and ovarian function [J]. Molecular and Cellular Endocrinology, 2002, 191, 15-18
[17] Galloway SM, A linkage map of the ovine X chromosome [J]. Genome Research. 1996, 6: 667-677
[18] 管锋，杨利国，石国庆等．绵羊多胎性状基因调控研究进展[J]．黑龙江畜牧兽医，2003，11(3)：12—17
[19] Jennifer L, Dube, Pei Wang, The bone Morphogenetic Protein 15 geneis X-Linked and Expressed in Oocytes [J]. Molecular Endocrinology. 2002, 12(12): 1809-1817.
[20] 张果平，黄永宏，李峰，绵羊多胎性FecB基因研究新进展[J]．动物科学与动物医学，2002，19(9)：5—8
[21] 石国庆，缪礼维等．新疆细毛羊发展回顾与展望．中国草食动物[J]．2001-2002年全国养羊生产与学术研讨会议论文集：18-23
[22] 徐宏发等．分子系统学研究进展[J]，生态学杂志．2001．20(3)：41-46。
[23] 李学斌．分子标记回归模型与显性分子标记资料[J]．动物科学与动物医学．2002，19(6)：5-9
[24] 赵宗胜，李大全等．杂交羔羊体尺与断奶体重间的多元回归分析[J]．黑龙江畜牧兽医．1998，12：5-6
[25] 赵宗胜，李大全等．杂交肉羊不同日龄体重间的回归分析[J]．黑龙江畜牧兽医．1998，10：11-14
[26] 王大广，苏秀侠等．萨福克肉用种羊线性相关与回归分析[J]。吉林农业科学．2002，27(1)：47-49
[27] 王同军．建昌黑山羊体尺体重相关性研究[J]．四川畜牧兽医．2003，30(9)：22-23．26
[28] 买买提明巴拉提等．麦盖提羊羔羊初生体重、体尺及其相关性的研究[J]．黑龙江畜牧兽医．2001，5：9-10
[29] 赵素君．微卫星标记及其与畜禽遗传育种中的应用[J]．西南民族大学学报(自然科学版)．2003，29(4)：499-502
[30] 樊斌，李奎，彭中镇，1999，微卫星标记检测遗传多样性的原理和方法[J]．第十次全国动物遗传育种学术讨论会论文．北京：中国农业科技出版社P32-36．
[31] 杨章平，常洪等．绵羊和山羊基于两种标记的遗传分化初步研究[J]．扬州大学学报．2003，24(4)：27-31
[32] Takezake N, Genetic Distance and Reconstruction of Phylogenetic Tree From Microsatellite DNA [J]. Genetics. 1996, 144: 389-399.
[33] Zeveren, A V, Peelman L, Weghe A, A genetic study of four Belgian pig populations
by means of seven microsatellite loci [J]. Anita. Breed. Genet. 1995, 112: 191-204
[34] 候冠玉，马月辉．微卫星DNA在草食动物遗传育种中的研究与应用[J]．中国草食动物．2003，23(3)：39-40
[35] 储明星，王吉振等，小尾寒羊五个微卫星基因座遗传多态性研究[J]．遗传学报，2002，2(6)：502—506
[36] 杨章平，常洪等．结构基因座和微卫星标记应用于绵(山)羊群体遗传分化的比较研究[J]．畜牧兽医学报．2003，34(5)：427-433
[37] 代江生，刘云芳等．五个绵羊品种的RAPD初步分析[J]．草食家畜．1998，1：11-13
[38] Jia Bin, Chen Jie et ai. , Microsatellite analysis of genetic diversity and phylogenetic relationship of eight sheep breeds in Xinjiang [J]. 遗传学报. 2003, 30 (9): 847-854.
[39] Berghmans, S., Segers, K., et al., Breakpioint mapping positions the callipyge gene within a 450-kilobase chromosome segment containing the DLK1 and GTL2 gene [J]. Mammal Genome. 2001, 12: 183-185.
[40] Bidwell, C. A., Shay, T. L. , et al. , Differential expression of the GTL2gene within the callipyge region ofovine chromosome 18 [J]. Animal Genetics. 2001, 32: 248-256.
[41] Charlier, C., Segers, K., et al., The callipyge mutation enhances the expression of coregulated imprinted genes in cis without affecting their imprinting status [J]. Mammal Genetic. 2001, 27: 367-369.
[42] Maria Smit, Karin Segers, et al., Mosaicism of Solid Gold Supports the causality of a moneoding A-to-G transition in the eterminism of the Callipyge phenotype [J]. Genetics. 2003, 163: 453-456.
[43] Segers, K., Vaiman, D. et al., Construction and characterizaton of an ovine BAC eontig spanning the eallipyge locus [J]. Animal Genetic. 2001, 31: 352-359.
[44] Hayashi K．PCR—SSCP．一种检测突变的方法[J]．国外医学卫生学分册．1994，21(4)：204-205
[45] Fahrenkrug, S C, Freking B A, Single nucleotide polymorphism (SNP) discovery in porcine expressed genes [J]. Animal Genetic. 2002, 33: 186-195.
[46] Freking B A, Keele J W, Evaluation of the ovine eallipyge locus: I. Relative chromosomal position and gene action [J] . Journal of Animal Science. 1998, 76: 2062-2071.
[47] Mulsant P, Lecerf F, Mutation in borne morphogenetic protein receptor-IB is associated with increased ovulation rate in Booroola Merino ewes[J]. Proc. Natl. Acad. Sci. 2001, 98: 5104-5109.
[48] Susan M. Galloway, Mutation in an oocyte-derived growth factor gene (BMP 15) can increased ovulation rate and infertileity in a dosage-sensitive manner [J]. Nature Genetic, 2000, 25.
[49] 柳淑芳，闫艳春，杜立新，FecB基因及BMPR—IB基因的突变特性与生物学意义[J]．遗传，
2003，25(1)：93—96．
[50] 王根林，毛鑫智等，DNA分析了现我国湖羊和小尾寒羊存在Booroola(FecB)多胎基因[J]．南京农业大学学报，2003，26(1)：104—108
[51] 钟发刚等．绵羊BMPR-IB基因多态性与其产羔数的相关研究[J]．草食家畜[J]．2003，2：20-23
[52] 陈晓军，钟发刚，罗淑萍，王新华，多浪羊BMPR-IB基因多态性的初步研究[J]．新疆农业科学．2004，41(1)：6—9．
[53] 储明星，程国华等，微卫星标记OarAE101和BM1329在五个绵羊品种中的初步研究[J]．遗传学报，2001，28(6)：510—517
[54] 储明星，王吉振等，小尾寒羊4个微卫星座位的克隆及序列分析[J]．遗传学报，2002，29(5)：402—405