VRTN及NR6A1基因多态性与苏尼特羊胸椎数变异的相关性及组织表达分析
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摘要
胸椎数性状是生产中一类重要的经济性状,在绵羊(Ovis aries)上已确定的候选基因很少。本研究旨在研究脊椎发育相关基因(vertnin,VRTN)及核受体辅抑制子1基因(nuclear receptor subfamily 6 group A member 1, NR6A1)多态性与苏尼特羊胸椎数变异之间的联系及两个基因的组织表达水平。研究利用PCR研究了苏尼特羊VRTN和NR6A1基因的多态性及两个基因的SNP位点与多胸椎数性状的相关性,利用qRTPCR技术检测了VRTN和NR6A1基因在不同胸椎数(胸椎数14根为T14组;胸椎数13根为T13组)苏尼特羊部分组织中的表达。结果表明,VRTN基因g.82533904C>T位点、NR6A1基因g.11206763T>C位点和g.11210075T>C位点与苏尼特羊胸椎数之间无显著关联(P>0.05),VRTN基因g.82533661C>A位点与苏尼特羊胸椎数之间存在极显著关联(P<0.01)。NR6A1基因在不同胸椎数(T14组;T13组)苏尼特羊的心、肝、脾、肺、肾、肌肉和脂肪组织中均有表达,NR6A1基因在2种不同胸椎数苏尼特羊的心脏、肺脏、肾脏、肌肉和脂肪组织中的表达差异不显著(P>0.05),在脾脏组织中T13组与T14组之间表达差异处于显著水平(P<0.05),在肝脏组织中T13组与T14组之间表达差异处于极显著水平(P<0.01);VRTN基因在苏尼特羊的心、肝、脾、肺、肾、肌肉和脂肪组织中均有表达,在T14组中表达量高于T13组,T14组肌肉中表达量显著高于T13组(P<0.05),其他组织在不同胸椎数苏尼特羊之间的表达差异不显著(P>0.05)。NR6A1基因在两种不同胸椎数苏尼特羊的组织中表达没有明显的规律,以上实验结果表明,绵羊VRTN基因g.82533661C>A位点对于选育多胸椎的苏尼特羊群体有一定指导意义,同时,NR6A1基因与苏尼特羊胸椎数性状之间没有明显关系,而VRTN基因可能是影响苏尼特羊胸椎数的重要基因之一。确定绵羊胸椎数候选基因有利于选育多胸椎数绵羊群体,能够有效提高肉羊产业的经济效益。
The number of thoracic vertebrae is an important economic trait in production, but few candidate genes have been identified in sheep(Ovis aries). The aim of this study is to explore the association between polymorphism of VRTN(vertnin) and NR6 A1(nuclear receptor subfamily 6 group A member 1) genes and thoracic vertebral number, and tissue expression levels of 2 genes were also detected. The association analysis of VRTN and NR6 A1 polymorphisms with thoracic vertebral number were performed by PCR, and expression levels of VRTN and NR6 A1 genes in Sunite sheep with different thoracic vertebral numbers(14 thoracic vertebral number, T14; 13 thoracic vertebral number, T13) were tested by qRT-PCR. The results showed that there were no significant association between g.82533904 C>T, g.11206763 T>C and g.11210075 T>C and the thoracic vertebral number in Sunite sheep(P>0.05), while there was a highly significant association between g.82533661 C>A and the thoracic vertebral number in Sunite sheep(P<0.01). NR6 A1 expressed in heart, liver,spleen, lung, kidney, muscle and fat of Sunite sheep. There were no significant difference of expression levels between sheep with T14 and T13 in heart, lung, kidney, muscle and fat(P>0.05), while there were significant difference in spleen(P<0.05) and extremely significant difference in liver(P<0.01) of Sunite sheep with different thoracic vertebral number. The expression levels of VRTN were all higher in Sunite sheep with T14 than those of T13 in heart, liver, spleen, lung and kidney, the difference of muscle was significant(P<0.05),the difference of the other tissues were not significant(P>0.05). There was no uniform expression pattern of Sunite sheep with different thoracic vertebral number. In summary, g.82533661 C>A had potential application value for the selection of the Sunite sheep population with more thoracic vertebral number. At the same time,there was no significant association between NR6 A1 and the thoracic vertebral trait of the Sunite sheep, and VRTN may be one of the candidate genes affecting the thoracic vertebral numbers in Sunite sheep.Determining candidate genes for thoracic vertebrae number trait in sheep is conducive to the selection of multiple thoracic vertebrae sheep population, which can effectively improve the economic benefits of the sheep industry.
The number of thoracic vertebrae is an important economic trait in production, but few candidate genes have been identified in sheep(Ovis aries). The aim of this study is to explore the association between polymorphism of VRTN(vertnin) and NR6 A1(nuclear receptor subfamily 6 group A member 1) genes and thoracic vertebral number, and tissue expression levels of 2 genes were also detected. The association analysis of VRTN and NR6 A1 polymorphisms with thoracic vertebral number were performed by PCR, and expression levels of VRTN and NR6 A1 genes in Sunite sheep with different thoracic vertebral numbers(14 thoracic vertebral number, T14; 13 thoracic vertebral number, T13) were tested by qRT-PCR. The results showed that there were no significant association between g.82533904 C>T, g.11206763 T>C and g.11210075 T>C and the thoracic vertebral number in Sunite sheep(P>0.05), while there was a highly significant association between g.82533661 C>A and the thoracic vertebral number in Sunite sheep(P<0.01). NR6 A1 expressed in heart, liver,spleen, lung, kidney, muscle and fat of Sunite sheep. There were no significant difference of expression levels between sheep with T14 and T13 in heart, lung, kidney, muscle and fat(P>0.05), while there were significant difference in spleen(P<0.05) and extremely significant difference in liver(P<0.01) of Sunite sheep with different thoracic vertebral number. The expression levels of VRTN were all higher in Sunite sheep with T14 than those of T13 in heart, liver, spleen, lung and kidney, the difference of muscle was significant(P<0.05),the difference of the other tissues were not significant(P>0.05). There was no uniform expression pattern of Sunite sheep with different thoracic vertebral number. In summary, g.82533661 C>A had potential application value for the selection of the Sunite sheep population with more thoracic vertebral number. At the same time,there was no significant association between NR6 A1 and the thoracic vertebral trait of the Sunite sheep, and VRTN may be one of the candidate genes affecting the thoracic vertebral numbers in Sunite sheep.Determining candidate genes for thoracic vertebrae number trait in sheep is conducive to the selection of multiple thoracic vertebrae sheep population, which can effectively improve the economic benefits of the sheep industry.
引文
范家萌.2015.NR6A1、PLAG1、LCORL基因在民猪群体内的单倍型及连锁不平衡分析[D].博士学位论文,东北农业大学,导师:刘娣.pp.8-10.(Fan J M.2015.The Studies on the haplotypes of NR6A1,PLAG1 and LCORLand the association on min pig population[D].Thesis for Ph.D.,Northeast Agricultural University,Supervisor:Liu D,pp.8-10.)
范家萌,于浩,刘娣.2016.民猪NR6A1基因的单核苷酸多态性检测及连锁不平衡研究[J].黑龙江畜牧兽医,(1):87-89.(Fan J M,Yu H,Liu D.2016.Single nucleotide polymorphism detection and linkage disequilibrium study of NR6A1 gene in min pigs[J].Heilongjiang Animal Science and Veterinary Medicine,(1):87-89.)
韩浩园,王绍强,黄洁萍,等.2013.秦川牛多脊椎性状调查[J].中国牛业科学,39(6):78-79.(Han H Y,Wang S Q,Huang J P,et al.2013.Investigation of multi-vertebral trait in Qinchuan cattle[J].China Cattle Science,39(6):78-79.)
韩立霞.2009.河北小尾寒羊多脊柱调控基因Btg2和NR6A1的多态性分析[D].硕士学位论文,河北农业大学,导师:孙少华.pp.37-39.(Han L X.2009.Polymorphism analysis of multi-vertebrae regulatory genes Btg2and NR6A1 in Hebei Small Tail Han Sheep[D].Thesis for M.S.,Hebei Agricultural University,Supervisor:Sun S H,pp.37-39.)
黄洁萍,张明明,马云,等.2017.家畜多脊椎性状研究进展[J].中国畜牧兽医,44(8):2386-2391.(Huang J P,Zhang M M,Ma Y,et al.2017.Research progress on multi-vertebral trait in livestock[J].China Animal Husbandry&Veterinary Medicine,44(8):2386-2391.)
刘建明,孙少华,韩立霞,等.2010.ActRⅡB基因单核苷酸多态性与小尾寒羊多脊椎变异的关联研究[J].畜牧兽医学报,41(8):951-954.(Liu J M,Sun S H,Han L X,et al.2010.Association between single nucleotide polymorphism of ActRⅡB gene and vertebra number variation in Small Tail Han Sheep[J].Chinese Journal of Animal&Veterinary Sciences,41(8):951-954.)
刘晓雯,刘华伟,王伟龙,等.2018.孤儿核受体Nr6a1的生物学功能与表达调控机制[J].激光生物学报,27(3):199-204.(Liu X W,Liu H W,Wang W L,et al.2018.Biological function and expression regulation mechanism of orphan nuclear receptor nr6a1[J].Acta Laser Biology Sinica,27(3):199-204.)
孙庆,贺小云,周梅,等.2018.家畜脊椎数性状相关基因研究进展[J].农业生物技术学报,26(12):2142-2149.(Sun Q,He X Y,Zhou M,et al.2018.Advances in genes associated with vertebral number trait in livestock[J].Journal of Agricultural Biotechnology,26(12):2142-2149.)
吴一尘,杜星,李平华,等.2016.苏淮猪VRTN基因克隆、组织表达特征与多态性分析[J].中国农业科学,49(18):3639-3648.(Wu Y C,Du X,Li P H,et al.2016.Sequence cloning,tissue expression profile and polymorphism of VRTN gene in Suhuai pig[J].Scientia Agricultura Sinica,49(18):3639-3648.)
张立岭.2000.蒙古羊的发育遗传和群体遗传研究[D].博士学位论文,中国农业大学,导师:张沅,pp.40-46.(Zhang L L.2000.Study on developmental and population genetics in mongolia sheep[D].Thesis for D.S.,China Agriculture University,Supervisor:Zhang Y,pp.40-46.)
张立岭,张世铨.1998.多脊椎蒙古羊的胸腰椎长度及其产肉性能分析[J].内蒙古农业大学学报(自然科学版),(3):1-5.(Zhang L L,Zhang S Q.1998.The lengths of thoracic and lumbar vertebrae and the performance of Mongolia sheep[J].Journal of Inner Mongola Institute of Agriculture&Animal Husbandry,(3):1-5.)
张亚辉,汪雄,吴胜英,等.2017.核受体NR6A1通过上调RIPK3基因表达诱导血管平滑肌细胞凋亡[J].中国病理生理杂志,33(9):1581-1586.(Zhang Y H,Wang X,Wu S Y,et al.2017.NR6A1 promotes vascular smooth muscle cell apoptosis by up-regulating RIPK3 gene expression[J].Chinese Journal of Pathophysiology,33(9):1581-1586.)
张震,徐盼,张慧,等.2015.VRTN Promoter-hsp68-LacZ表达载体的构建与鉴定[J].江西农业大学学报,(6):1080-1085.(Zhang Z,Xu P,Zhang H,et al.2015.Construction and identificationof VRTN promoterhsp68-LacZ expression vector[J].Acta Agriculturae Universitatis Jiangxiensis,(6):1080-1085.)
赵静,张立岭,陈琦,等.2011.蒙古羊Hoxc8基因甲基化与胸椎数量的关系[J].黑龙江畜牧兽医,(5):5-8.(Zhao J,Zhang L L,Chen Q,et al.2011.Relationship between methylation of Hoxc8 gene and the numbers of thoracic vertebrae in Mongolia sheep[J].Heilongjiang Animal Science&Veterinary Medicine,(5):5-8.)
周世华,李丹,刘晓雯,等.2017.miR-181b-5p靶向调控Nr6a1抑制GC-1spg细胞的增殖与迁移[J].生命科学研究,(04):59-64.(Zhou S H,Li D,Liu X W,et al.2017.miR-181b-5p inhibits proliferation and migration of GC-1spg cells through targeting nr6a1[J].Life Science Research,(04):59-64.)
周梅,曹晓涵,贺小云,等.2018a.绵羊FGF7基因组织表达及其多态性与产羔数之间的关系[J].畜牧兽医学报,49(03):525-533.(Zhou M,Cao X H,He X Y,et al.2018.Tissue expression and polymorphism of sheep FGF7 gene and their association with litter size[J].Chinese Journal of Animal&Veterinary Sciences,49(03):525-533.)
周梅,曹晓涵,贺小云,等.2018b.不同产羔数小尾寒羊FOXL2和FIGLA基因的表达研究[J].家畜生态学报,39(01):14-18.(Zhou M,Cao X H,He X Y,et al.2018.Study on the expression of FOXL2 and FIGLA genes in small tail han sheep with different litter size[J].Journal of Domestic Animal Ecology,39(01):14-18.)
Borchers N,Reinsch N,Kalm E.2015.The number of ribs and vertebrae in a Piétrain cross:Variation,heritability and effects on performance traits[J].Journal of Animal Breeding&Genetics,121(6):392-403.
Burgos C,Latorre P,Altarriba J,et al.2015.Allelic frequencies of NR6A1 and VRTN,two genes that affect vertebrae number in diverse pig breeds:A study of the effects of the VRTN insertion on phenotypic traits of a Duroc Landrace-Large White cross[J].Meat Science,100:150-155.
Chung A C,Katz D,Pereira F A,et al.2001.Loss of orphan receptor germ cell nuclear factor function results in ectopic development of the tail bud and a novel posterior truncation[J].Molecular&Cellular Biology,21(2):663-677.
Dall'olio S,Ribani A,Moscatelli G,et al.2018.Teat number parameters in Italian Large White pigs:Phenotypic analysis and association with vertnin(VRTN)gene allele variants[J].Livestock Science,210:68-72.
Donaldson C L,Lambe N R,Maltin C A,et al.2013.Between-and within-breed variations of spine characteristics in sheep[J].Journal of Animal Science,91(2):995-1004.
Duan Y,Zhang H,Zhang Z,et al.2018.VRTN is required for the development of thoracic vertebrae in mammals[J].International Journal of Biological Sciences,14(6):667-681.
Hirose K,Mikawa S,Okumura N,et al.2013.Association of swine vertnin(VRTN)gene with production traits in Duroc pigs improved using a closed nucleus breeding system[J].Animal Science Journal,84(3):213-221.
Ibrahim D A,Myung K S,Skaggs D L.2013.Ten percent of patients with adolescent idiopathic scoliosis have variations in the number of thoracic or lumbar vertebrae[J].The Journal of Bone&Joint Surgery,95(9):828-833.
Katz D,Niederberger C,Slaughter G R,et al.1997.Characterization of germ cell-specific expression of the orphan nuclear receptor,germ cell nuclear factor[J].Endocrinology,138(10):4364-4372.
King J W B,Roberts R C.1960.Carcass length in the bacon pig;its association with vertebrae numbers and prediction from radiographs of the young pig[J].Animal Production,2(1):59-65.
Mikawa S,Morozumi T,Shimanuki S,et al.2007.Fine mapping of a swine quantitative trait locus for number of vertebrae and analysis of an orphan nuclear receptor,germ cell nuclear factor(NR6A1)[J].Genome Research,17(5):586-593.
Mikawa S,Sato S,Nii M,et al.2011.Identification of a second gene associated with variation in vertebral number in domestic pigs[J].BMC Genetics,12(1):5-17.
Nakano H,Sato S,Uemoto Y,et al.2015.Effect of VRTNgene polymorphisms on Duroc pig production and carcass traits,and their genetic relationships[J].Animal Science Journal,86(2):125-131.
Narita Y,Kuratani S.2005.Evolution of the vertebral formulae in mammals:A perspective on developmental constraints[J].Journal of Experimental Zoology Part B Molecular&Developmental Evolution,304B(2):91-106.
Tang J,Zhang Z,Yang B,et al.2017.Identification of loci affecting teat number by genome-wide association studies on three pig populations[J].Asian-Australasian Journal of Animal Sciences,30(1):1-7.
Wang Y,Zhang Y,Dai X,et al.2015.NR6A1 couples with cAMP response element binding protein and regulates vascular smooth muscle cell migration[J].International Journal of Biochemistry&Cell Biology,69:225-232.
Yang J,Huang L,Yang M,et al.2016.Possible introgression of the VRTN mutation increasing vertebral number,carcass length and teat number from Chinese pigs into European pigs[J].Scientific Rep,6:1-8.
Zhang Z,Sun Y,Wei D,et al.2017.Effects of vertebral number variations on carcass traits and genotyping of vertnin candidate gene in Kazakh sheep[J].Asian-Australasian Journal of Animal Sciences,30(9):1234-1238.
范家萌,于浩,刘娣.2016.民猪NR6A1基因的单核苷酸多态性检测及连锁不平衡研究[J].黑龙江畜牧兽医,(1):87-89.(Fan J M,Yu H,Liu D.2016.Single nucleotide polymorphism detection and linkage disequilibrium study of NR6A1 gene in min pigs[J].Heilongjiang Animal Science and Veterinary Medicine,(1):87-89.)
韩浩园,王绍强,黄洁萍,等.2013.秦川牛多脊椎性状调查[J].中国牛业科学,39(6):78-79.(Han H Y,Wang S Q,Huang J P,et al.2013.Investigation of multi-vertebral trait in Qinchuan cattle[J].China Cattle Science,39(6):78-79.)
韩立霞.2009.河北小尾寒羊多脊柱调控基因Btg2和NR6A1的多态性分析[D].硕士学位论文,河北农业大学,导师:孙少华.pp.37-39.(Han L X.2009.Polymorphism analysis of multi-vertebrae regulatory genes Btg2and NR6A1 in Hebei Small Tail Han Sheep[D].Thesis for M.S.,Hebei Agricultural University,Supervisor:Sun S H,pp.37-39.)
黄洁萍,张明明,马云,等.2017.家畜多脊椎性状研究进展[J].中国畜牧兽医,44(8):2386-2391.(Huang J P,Zhang M M,Ma Y,et al.2017.Research progress on multi-vertebral trait in livestock[J].China Animal Husbandry&Veterinary Medicine,44(8):2386-2391.)
刘建明,孙少华,韩立霞,等.2010.ActRⅡB基因单核苷酸多态性与小尾寒羊多脊椎变异的关联研究[J].畜牧兽医学报,41(8):951-954.(Liu J M,Sun S H,Han L X,et al.2010.Association between single nucleotide polymorphism of ActRⅡB gene and vertebra number variation in Small Tail Han Sheep[J].Chinese Journal of Animal&Veterinary Sciences,41(8):951-954.)
刘晓雯,刘华伟,王伟龙,等.2018.孤儿核受体Nr6a1的生物学功能与表达调控机制[J].激光生物学报,27(3):199-204.(Liu X W,Liu H W,Wang W L,et al.2018.Biological function and expression regulation mechanism of orphan nuclear receptor nr6a1[J].Acta Laser Biology Sinica,27(3):199-204.)
孙庆,贺小云,周梅,等.2018.家畜脊椎数性状相关基因研究进展[J].农业生物技术学报,26(12):2142-2149.(Sun Q,He X Y,Zhou M,et al.2018.Advances in genes associated with vertebral number trait in livestock[J].Journal of Agricultural Biotechnology,26(12):2142-2149.)
吴一尘,杜星,李平华,等.2016.苏淮猪VRTN基因克隆、组织表达特征与多态性分析[J].中国农业科学,49(18):3639-3648.(Wu Y C,Du X,Li P H,et al.2016.Sequence cloning,tissue expression profile and polymorphism of VRTN gene in Suhuai pig[J].Scientia Agricultura Sinica,49(18):3639-3648.)
张立岭.2000.蒙古羊的发育遗传和群体遗传研究[D].博士学位论文,中国农业大学,导师:张沅,pp.40-46.(Zhang L L.2000.Study on developmental and population genetics in mongolia sheep[D].Thesis for D.S.,China Agriculture University,Supervisor:Zhang Y,pp.40-46.)
张立岭,张世铨.1998.多脊椎蒙古羊的胸腰椎长度及其产肉性能分析[J].内蒙古农业大学学报(自然科学版),(3):1-5.(Zhang L L,Zhang S Q.1998.The lengths of thoracic and lumbar vertebrae and the performance of Mongolia sheep[J].Journal of Inner Mongola Institute of Agriculture&Animal Husbandry,(3):1-5.)
张亚辉,汪雄,吴胜英,等.2017.核受体NR6A1通过上调RIPK3基因表达诱导血管平滑肌细胞凋亡[J].中国病理生理杂志,33(9):1581-1586.(Zhang Y H,Wang X,Wu S Y,et al.2017.NR6A1 promotes vascular smooth muscle cell apoptosis by up-regulating RIPK3 gene expression[J].Chinese Journal of Pathophysiology,33(9):1581-1586.)
张震,徐盼,张慧,等.2015.VRTN Promoter-hsp68-LacZ表达载体的构建与鉴定[J].江西农业大学学报,(6):1080-1085.(Zhang Z,Xu P,Zhang H,et al.2015.Construction and identificationof VRTN promoterhsp68-LacZ expression vector[J].Acta Agriculturae Universitatis Jiangxiensis,(6):1080-1085.)
赵静,张立岭,陈琦,等.2011.蒙古羊Hoxc8基因甲基化与胸椎数量的关系[J].黑龙江畜牧兽医,(5):5-8.(Zhao J,Zhang L L,Chen Q,et al.2011.Relationship between methylation of Hoxc8 gene and the numbers of thoracic vertebrae in Mongolia sheep[J].Heilongjiang Animal Science&Veterinary Medicine,(5):5-8.)
周世华,李丹,刘晓雯,等.2017.miR-181b-5p靶向调控Nr6a1抑制GC-1spg细胞的增殖与迁移[J].生命科学研究,(04):59-64.(Zhou S H,Li D,Liu X W,et al.2017.miR-181b-5p inhibits proliferation and migration of GC-1spg cells through targeting nr6a1[J].Life Science Research,(04):59-64.)
周梅,曹晓涵,贺小云,等.2018a.绵羊FGF7基因组织表达及其多态性与产羔数之间的关系[J].畜牧兽医学报,49(03):525-533.(Zhou M,Cao X H,He X Y,et al.2018.Tissue expression and polymorphism of sheep FGF7 gene and their association with litter size[J].Chinese Journal of Animal&Veterinary Sciences,49(03):525-533.)
周梅,曹晓涵,贺小云,等.2018b.不同产羔数小尾寒羊FOXL2和FIGLA基因的表达研究[J].家畜生态学报,39(01):14-18.(Zhou M,Cao X H,He X Y,et al.2018.Study on the expression of FOXL2 and FIGLA genes in small tail han sheep with different litter size[J].Journal of Domestic Animal Ecology,39(01):14-18.)
Borchers N,Reinsch N,Kalm E.2015.The number of ribs and vertebrae in a Piétrain cross:Variation,heritability and effects on performance traits[J].Journal of Animal Breeding&Genetics,121(6):392-403.
Burgos C,Latorre P,Altarriba J,et al.2015.Allelic frequencies of NR6A1 and VRTN,two genes that affect vertebrae number in diverse pig breeds:A study of the effects of the VRTN insertion on phenotypic traits of a Duroc Landrace-Large White cross[J].Meat Science,100:150-155.
Chung A C,Katz D,Pereira F A,et al.2001.Loss of orphan receptor germ cell nuclear factor function results in ectopic development of the tail bud and a novel posterior truncation[J].Molecular&Cellular Biology,21(2):663-677.
Dall'olio S,Ribani A,Moscatelli G,et al.2018.Teat number parameters in Italian Large White pigs:Phenotypic analysis and association with vertnin(VRTN)gene allele variants[J].Livestock Science,210:68-72.
Donaldson C L,Lambe N R,Maltin C A,et al.2013.Between-and within-breed variations of spine characteristics in sheep[J].Journal of Animal Science,91(2):995-1004.
Duan Y,Zhang H,Zhang Z,et al.2018.VRTN is required for the development of thoracic vertebrae in mammals[J].International Journal of Biological Sciences,14(6):667-681.
Hirose K,Mikawa S,Okumura N,et al.2013.Association of swine vertnin(VRTN)gene with production traits in Duroc pigs improved using a closed nucleus breeding system[J].Animal Science Journal,84(3):213-221.
Ibrahim D A,Myung K S,Skaggs D L.2013.Ten percent of patients with adolescent idiopathic scoliosis have variations in the number of thoracic or lumbar vertebrae[J].The Journal of Bone&Joint Surgery,95(9):828-833.
Katz D,Niederberger C,Slaughter G R,et al.1997.Characterization of germ cell-specific expression of the orphan nuclear receptor,germ cell nuclear factor[J].Endocrinology,138(10):4364-4372.
King J W B,Roberts R C.1960.Carcass length in the bacon pig;its association with vertebrae numbers and prediction from radiographs of the young pig[J].Animal Production,2(1):59-65.
Mikawa S,Morozumi T,Shimanuki S,et al.2007.Fine mapping of a swine quantitative trait locus for number of vertebrae and analysis of an orphan nuclear receptor,germ cell nuclear factor(NR6A1)[J].Genome Research,17(5):586-593.
Mikawa S,Sato S,Nii M,et al.2011.Identification of a second gene associated with variation in vertebral number in domestic pigs[J].BMC Genetics,12(1):5-17.
Nakano H,Sato S,Uemoto Y,et al.2015.Effect of VRTNgene polymorphisms on Duroc pig production and carcass traits,and their genetic relationships[J].Animal Science Journal,86(2):125-131.
Narita Y,Kuratani S.2005.Evolution of the vertebral formulae in mammals:A perspective on developmental constraints[J].Journal of Experimental Zoology Part B Molecular&Developmental Evolution,304B(2):91-106.
Tang J,Zhang Z,Yang B,et al.2017.Identification of loci affecting teat number by genome-wide association studies on three pig populations[J].Asian-Australasian Journal of Animal Sciences,30(1):1-7.
Wang Y,Zhang Y,Dai X,et al.2015.NR6A1 couples with cAMP response element binding protein and regulates vascular smooth muscle cell migration[J].International Journal of Biochemistry&Cell Biology,69:225-232.
Yang J,Huang L,Yang M,et al.2016.Possible introgression of the VRTN mutation increasing vertebral number,carcass length and teat number from Chinese pigs into European pigs[J].Scientific Rep,6:1-8.
Zhang Z,Sun Y,Wei D,et al.2017.Effects of vertebral number variations on carcass traits and genotyping of vertnin candidate gene in Kazakh sheep[J].Asian-Australasian Journal of Animal Sciences,30(9):1234-1238.