绵羊重要经济性状可用遗传标记研究进展

doi:10.16431/j.cnki.1671-7236.2023.08.012

摘要/Abstract

摘要： 绵羊作为重要的经济动物,是人类社会肉、奶和皮毛的来源之一。生长性状、胴体性状、繁殖性状、产毛性状、抗病及耐寒等都是绵羊生产和遗传育种研究领域的重要经济性状,了解这些性状的分子遗传基础,确定关键功能基因以及遗传标记位点有利于提高重要经济性状选择的准确性,进而加速绵羊经济性状的遗传改良。随着基因组重测序、转录组测序和甲基化测序等技术的发展以及全基因组关联分析技术的成熟,有关绵羊表型性状与基因型的关联分析报道较多,挖掘到一系列绵羊经济性状相关的候选基因和遗传标记。作者主要从生长发育、繁殖、尾脂沉积和羊毛生产4个角度对近几年筛选到的可用遗传标记基因进行总结,挑选出一些明星基因,如肌细胞增强因子(myocyte enhancer factor 2B,MEF2B)和生长抑素受体1(somatostatin receptor 1,SSTR1)影响绵羊体重;椎骨发育相关基因(vertebrae development associated,VRTN)是决定胸椎数量的主效基因;繁殖力基因(fecundity booroola,FecB)、骨形态发生蛋白15(bone morphogenetic protein 15,BMP15)和卵泡刺激素受体(follicle stimulating hormone receptor,FSHR)等与绵羊的繁殖性状相关;血小板衍生生长因子D(platelet derived growth factor D,PDGFD)和BMP2基因在肥尾羊中被强烈选择,在尾部脂肪沉积中有重要作用;Agouti信号蛋白(agouti signaling protein,ASIP)和黑皮质素1受体(melanocortin 1 receptor,MC1R)等与绵羊毛色显著相关。同时对影响绵羊经济性状的可用遗传标记基因突变位点、碱基变化及突变类型进行综述,为后续绵羊遗传育种研究提供了思路。

关键词: 绵羊; 候选基因; 全基因组; 遗传标记

Abstract: As an important economic animal,sheep is one of the sources of meat,milk and fur in human society.Growth traits,carcass traits,breeding traits,reproductive traits,disease resistance and cold resistance are important economic traits of production and genetic breeding in sheep,understanding the molecular genetic basis of these traits,determining the key functional genes and genetic markers are conducive to improve the accuracy of important economic trait selection,and accelerate the genetic improvement of economic traits in sheep.With the development of genome resequencing,transcriptome sequencing and methylation sequencing,as well as the maturity of genome-wide association analysis technologies,there were many reports on the association analysis of phenotypic traits and genotypes in sheep,and a series of candidate genes and genetic marker related to economic traits in sheep have been excavated.The author summarized the available genetic marker genes screened in recent years mainly from the perspectives of growth and development,reproduction,tail lipid deposition and wool production,and selected some star genes,such as myocyte enhancer factor 2B (MEF2B) and somatostatin receptor 1 (SSTR1) affected body weight in sheep.Vertebrae development associated gene (VRTN) was a major gene that determines the number of thoracic vertebrae.Fecundity booroola (FecB),bone morphogenetic protein 15 (BMP15) and follicle stimulating hormone receptor (FSHR) were related to the breeding characters of sheep.Platelet derived growth factor D (PDGFD) and BMP2 genes were strongly selected in Fat-tailed sheep,and played an important role in tail fat deposition.Signaling protein agouti (ASIP) and melanocortin 1 receptor (MC1R) were significantly associated with hair color in sheep.The mutation sites,base changes and mutation types of available genetic markers affecting economic traits in sheep were summarized in this paper,which provided ideas for the subsequent research on genetic breeding in sheep.

Key words: sheep; candidate gene; whole genome; genetic marker

中图分类号:

S813.3

李晓杰, 韩建刚, 梁奔梦, 叶绍辉, 蒋琳, 马月辉. 绵羊重要经济性状可用遗传标记研究进展[J]. 中国畜牧兽医, 2023, 50(8): 3142-3156.

LI Xiaojie, HAN Jiangang, LIANG Benmeng, YE Shaohui, JIANG Lin, MA Yuehui. Research Advance on Available Genetic Markers for Important Economic Traits in Sheep[J]. China Animal Husbandry and Veterinary Medicine, 2023, 50(8): 3142-3156.

参考文献

[1] MARTÍNEZ-MONTESÁ M,FERNÁNDEZ A,MUÑOZ M,et al.Using genome wide association studiesto identify common QTL regions in three different genetic backgrounds based on Iberian pig breed[J].PLoS One,2018,13(3):e0190184.
[2] VAN DEN BERG I,HAYES B J,CHAMBERLAIN A J,et al.Overlap between eQTL and QTL associated with production traits and fertility in dairy cattle[J].BMC Genomics,2019,20(1):291.
[3] LE MIGNON G,DÉSERT C,PITEL F,et al.Using transcriptome profiling to characterize QTL regions on chicken chromosome 5[J].BMC Genomics,2009,10:575.
[4] BOLORMAA S,SWAN A A,BROWN D J,et al.Multiple-trait QTL mapping and genomic prediction for wool traits in sheep[J].Genetics Selection Evolution,2017,49(1):62.
[5] HU X J,YANG J,XIE X L,et al.The genome landscape of Tibetan sheep reveals adaptive introgression from Argali and the history of early human settlements on the Qinghai-Tibetan Plateau[J].Molecular Biology and Evolution,2019,36(2):283-303.
[6] FAN B,ONTERU S K,DU Z Q,et al.Genome-wide association study identifies loci for body composition and structural soundness traits in pigs[J].PLoS One,2011,6(2):e14726.
[7] JIANG L,LIU J,SUN D,et al.Genome wide association studies for milk production traits in Chinese Holstein population[J].PLoS One,2010,5(10):e13661.
[8] DAI R,HUANG C,WU X,et al.Copy number variation (CNV) of the AHR gene in the Ashidan yak and its association with growth traits[J].Gene,2022,826:146454.
[9] CHENG J,CAO X,HANIF Q,et al.Integrating genome-wide CNVs into QTLs and high confidence GWAScore regions identified positional candidates for sheep economic traits[J].Frontiers in Genetics,2020,11:569.
[10] GEBRESELASSIE G,BERIHULAY H,JIANG L,et al.Review on genomic regions and candidate genes associated with economically important production and reproduction traits in sheep (Ovies aries)[J].Animals,2020,10(1):33.
[11] LU Z,YUE Y,YUAN C,et al.Genome-wide association study of body weight traits in Chinese Fine-wool sheep[J].Animals,2020,10(1):170.
[12] ZHANG L,LIU J,ZHAO F,et al.Genome-wide association studies for growth and meat production traits in sheep[J].PLoS One,2013,8(6):e66569.
[13] TAO L,HE X Y,PAN L X,et al.Genome-wide association study of body weight and conformation traits in neonatal sheep[J].Animal Genetics,2020,51(2):336-340.
[14] CHEN L,CHENG B,LI L,et al.The molecular characterization and temporal-spatial expression of myocyte enhancer factor 2 genes in the goat and their association with myofiber traits[J]. Gene,2015,555(2):223-230.
[15] ABOUSOLIMAN I,REYER H,OSTER M,et al.Genome-wide analysis for early growth-related traits of the locally adapted Egyptian Barki sheep[J].Genes,2021,12(8):1243.
[16] ZHANG L,MA X,XUAN J,et al.Identification of MEF2B and TRHDE gene polymorphisms related to growth traits in a new Ujumqin sheep population[J]. PLoS One,2016,11(7):e015950.
[17] CHENG J,ZHANG X,LI F,et al.Detecting single nucleotide polymorphisms in MEF2B and UCP3 and elucidating their association with sheep growth traits[J].DNA and Cell Biology,2021,40(12):1554-1562.
[18] PASANDIDEH M,GHOLIZADEH M,RAHIMI-MIANJI G.A genome-wide association study revealed five SNPs affecting 8-month weight in sheep[J].Animal Genetics,2020,51(6):973-976.
[19] WU H,CHEN Y,MIAO S,et al.Sperm associated antigen 8(SPAG8),a novel regulator of activator of CREM in testis during spermatogenesis[J].FEBS Letters,2010,584(13):2807-2815.
[20] CINAR M U,MOUSEL M R,HERNDON M K,et al.Association of TMEM8B and SPAG8 with mature weight in sheep[J].Animals,2020,10(12):2391.
[21] KOMINAKIS A,HAGER-THEODORIDES A L,ZOIDIS E,et al.Combined GWAS and'guilt by association'-based prioritization analysis identifies functional candidate genes for body size in sheep[J].Genetics Selection Evolution,2017,49(1):41.
[22] LOOS R J F,YEO G S H.The bigger picture of FTO:The first GWAS-identified obesity gene[J].Nature Reviews Endocrinology,2014,10(1):51-61.
[23] SZYDLOWSKI M,SALAMON S,GRZES M,et al.SNP in the 5'flanking region of the pig FTO gene is associated with fatness in Polish Landrace[J]. Livestock Science,2012,150(1-3):397-400.
[24] JEVSINEK SKOK D,KUNEJ T,KOVAC M,et al.FTO gene variants are associated with growth and carcass traits in cattle[J].Animal Genetics,2016,47(2):219-222.
[25] ZHANG G W,GAO L,CHEN S Y,et al.Single nucleotide polymorphisms in the FTO gene and their association with growth and meat quality traits in rabbits[J].Gene,2013,527(2):553-557.
[26] WANG S,LIU S,YUAN T,et al.Genetic effects of FTO gene insertion/deletion (InDel) on fat-tail measurements and growth traits in Tong sheep[J].Animal Biotechnology,2021,32(2):229-239.
[27] ZHAO Y,ZHANG D,ZHANG X,et al.Expression features of the ovine FTO gene and association between FTO polymorphism and tail fat deposition related-traits in Hu sheep[J].Gene,2022,826:146451.
[28] OSMAN N M,SHAFEY H I,ABDELHAFEZ M A,et al.Genetic variations in the myostatin gene affecting growth traits in sheep[J].Veterinary World,2021,14(2):475-482.
[29] KENDLER D B,ARAJO M L,ALENCAR R,et al.Somatostatin receptor subtype 1 might be a predictor of better response to therapy in medullary thyroid carcinoma[J].Endocrine,2017,58(3):474-480.
[30] JIN Q J,SUN J J,FANG X T,et al.Molecular characterization and polymorphisms of the caprine somatostatin (SST) and SST receptor 1(SSTR1) genes that are linked with growth traits[J].Molecular Biology Reports,2011,38(5):3129-3135.
[31] IIDA A,SAITO S,SEKINE A,et al.Catalog of 300 SNPs in 23 genes encoding G-protein coupled receptors[J].Journal of Human Genetics,2004,49(4):194-208.
[32] ZHAO F,ZHOU H,LI S,et al.Growth and carcass trait association with variation in the somatostatin receptor 1(SSTR1) gene in New Zealand Romney sheep[J].New Zealand Journal of Agricultural Research,2018,61(4):477-486.
[33] LI X,DING N,ZHANG Z,et al.Identification of somatostatin receptor subtype 1(SSTR1) gene polymorphism and their association with growth traits in Hulun Buir sheep[J].Genes,2022,13(1):77.
[34] WANG X,CAO X,WEN Y,et al.Associations of ORMDL1 gene copy number variations with growth traits in four Chinese sheep breeds[J].Archives Animal Breeding,2019,62(2):571-578.
[35] YANG Z,CAO X,MA Y,et al.Novel copy number variation of the BAG4 gene is associated with growth traits in three Chinese sheep populations[J].Animal Biotechnology,2021,32(4):461-469.
[36] TOREMURAT Z,IBRAHIM E E,HUANG Y Z,et al.Copy number variations of TOP2B gene are associated with growth traits in Chinese sheep breeds[J].Animal Biotechnology,2022,33(1):85-89.
[37] JIANG R,CHENG J,CAO X K,et al.Copy number variation of the SHE gene in sheep and its association with economic traits[J].Animals,2019,9(8):531.
[38] FENG Z,LI X,CHENG J,et al.Copy number variation of the PIGY gene in sheep and its association analysis with growth traits[J]. Animals,2020,10(4):688.
[39] ERDENEE S,LI J,KANG Z,et al.Sheep zinc finger proteins 395(ZNF395):Insertion/deletion variations,associations with growth traits,and mRNA expression[J].Animal Biotechnology,2020,31(3):237-244.
[40] ERDENEE S,AKHATAYEVA Z,PAN C,et al.An insertion/deletion within the CREB1 gene identified using the RNA-sequencing is associated with sheep body morphometric traits[J].Gene,2021,775:145444.
[41] LI J,ERDENEE S,ZHANG S,et al.Genetic effects of PRNP gene insertion/deletion (indel) on phenotypic traits in sheep[J].Prion,2018,12(1):42-53.
[42] ZHAO H,HE S,WANG S,et al.Two new insertion/deletion variants of the PITX2 gene and their effects on growth traits in sheep[J].Animal Biotechnology,2018,29(4):276-282.
[43] WANG S,YI X,WU M,et al.Detection of key gene InDels in TGF-β pathway and its relationship with growth traits in four sheep breeds[J].Animal Biotechnology,2021,32(2):194-204.
[44] LI W,WANG X,ZHANG X,et al.Polymorphism of sheep PRKAA2 gene and its association with growth traits[J].Animal Biotechnology,2021,Doi:10.1080/10495398.2021.2021215.Online ahead of print.
[45] SHAN H,SONG X,CAO Y,et al.Association of the melanocortin 4 receptor (MC4R) gene polymorphism with growth traits of Hu sheep[J].Small Ruminant Research,2020,192:106206.
[46] LIN C,LI F,ZHANG X,et al.Expression and polymorphisms of CD8B gene and its associations with body weight and size traits in sheep[J].AnimalBiotechnology,2021,Doi:10.1080/10495398.2021.2016432.Online ahead of print.
[47] ZHAO L,LI F,YUAN L,et al.Expression of ovine CTNNA3 and CAP2 genes and their association with growth traits[J].Gene,2022,807:145949.
[48] LI C,ZHANG X,CAO Y,et al.Multi-vertebrae variation potentially contribute to carcass length and weight of Kazakh sheep[J].Small Ruminant Research,2017,150:8-10.
[49] SHENGWEI H,CUNYUAN L,MING L,et al.Whole-genome resequencing reveals loci associated with thoracic vertebrae number in sheep[J].Frontiers in Genetics,2019,10:674.
[50] ZHANG X,LI C,LI X,et al.Association analysis of polymorphism in the NR6A1 gene with the lumbar vertebrae number traits in sheep[J].Genes and Genomics,2019,41(10):1165-1171.
[51] LI C,LIU K,DAI J,et al.Whole-genome resequencing to investigate the determinants of the multi-lumbar vertebrae trait in sheep[J]. Gene,2022,809:146020.
[52] CHEN S,GUO X,HE X,et al.Transcriptome analysis reveals differentially expressed genes and long non-coding RNAs associated with fecundity in sheep hypothalamus with different FecB genotypes[J].Frontiers in Cell and Developmental Biology,2021,9:633474.
[53] KUMAR S,MISHRA A K,KOLTE A P,et al.Effects of the Booroola (FecB) genotypes on growth performance,ewe's productivity efficiency and litter size in Garole×Malpura sheep[J].Animal Reproduction Science,2008,105(3-4):319-331.
[54] WANG X,GUO X,HE X,et al.Effects of FecB mutation on estrus,ovulation,and endocrine characteristics in Small-tail Han sheep[J].Frontiers in Veterinary Science,2021,8:709737.
[55] SEJIAN V,MAURYA V P,PRINCE L L L,et al.Effect of FecB status on the allometric measurements and reproductive performance of Garole×Malpura ewes under hot semi-arid environment[J].Tropical Animal Health and Production,2015,47(6):1089-1093.
[56] MAHDAVI M,NANEKARANI S,HOSSEINI S D.Mutation in BMPR-ⅠB gene is associated with litter size in Iranian Kalehkoohi sheep[J].Animal Reproduction Science,2014,147(3-4):93-98.
[57] WANG W,LIU S,LI F,et al.Polymorphisms of the ovine BMPR-ⅠB,BMP-15 and FSHR and their associations with litter size in two Chinese indigenous sheep breeds[J].International Journal of Molecular Sciences,2015,16(5):11385-11397.
[58] CHU M X,LIU Z H,JIAO C L,et al.Mutations in BMPR-ⅠB and BMP-15 genes are associated with litter size in Small-tailed Han sheep (Ovis aries)[J].Journal of Animal Science,2007,85(3):598-603.
[59] WANG F,CHU M,PAN L,et al.Polymorphism detection of GDF9 gene and its association with litter size in Luzhong mutton sheep (Ovis aries)[J].Animals,2021,11(2):571.
[60] IMRAN F S,AL-THUWAINI T M,AL-SHUHAIB M B S,et al.A novel missense single nucleotide polymorphism in the GREM1 gene is highly associated with higher reproductive traits in Awassi sheep[J].Biochemical Genetics,2021,59(2):422-436.
[61] MA H,FANG C,LIU L,et al.Identification of novel genes associated with litter size of indigenous sheep population in Xinjiang,China using specific-locus amplified fragment sequencing technology[J].PeerJ,2019,7:e8079.
[62] TAO J,ZHANG L,ZHANG X,et al.Effect of exogenous melatonin on the development of mice ovarian follicles and follicular angiogenesis[J].International Journal of Molecular Sciences,2021,22(20):11262.
[63] TIAN X,WANG F,ZHANG L,et al.Beneficial effects of melatonin on the in vitro maturation of sheep oocytes and its relation to melatonin receptors[J].International Journal of Molecular Sciences,2017,18(4):834.
[64] BASINI G,BUSSOLATI S,CICCIMARRA R,et al.Melatonin potentially acts directly on swine ovary by modulating granulosa cell function and angiogenesis[J].Reproduction,Fertility and Development,2017,29(12):2305-2312.
[65] LURIDIANA S,COSSO G,PULINAS L,et al.New polymorphisms at MTNR1A gene and their association with reproductive resumption in Sarda breed sheep[J].Theriogenology,2020,158:438-444.
[66] CALVO J H,SERRANO M,MARTINEZ-ROYO A,et al.SNP rs403212791 in exon 2 of the MTNR1A gene is associated with reproductive seasonality in the Rasa Aragonesa sheep breed[J].Theriogenology,2018,113:63-72.
[67] HE X,ZHANG Z,CHU M.The effect of SNP rs400827589 in exon 2 of the MTNR1B gene on reproductive seasonality and litter size in sheep[J].Veterinary Medicine and Science,2020,6(4):804-812.
[68] MOHAMMED M M,AL-THUWAINI T M,AL-SHUHAIB M B S.A novel p.K116Q SNP in the OLR1 gene and its relation to fecundity in Awassi ewes[J].Theriogenology,2022,184:185-190.
[69] CHEN S,TAO L,HE X,et al.Single-nucleotide polymorphisms in FLT3,NLRP5,and TGIF1 are associated with litter size in Small-tailed Han sheep[J].Archives Animal Breeding,2021,64(2):475-486.
[70] ESMAEILI-FARD S M,GHOLIZADEH M,HAFEZIAN S H,et al.Genome-wide association study and pathway analysis identify NTRK2 as a novel candidate gene for litter size in sheep[J].PLoS One,2021,16(1):e0244408.
[71] AKHATAYEVA Z,MAO C,JIANG F,et al.Indel variants within the PRL and GHR genes associated with sheep litter size[J].Reproduction in DomesticAnimals,2020,55(11):1470-1478.
[72] ZHANG Z,LIU Q,DI R,et al.Single nucleotide polymorphisms in BMP2 and BMP7 and the association with litter size in Small-tail Han sheep[J].Animal Reproduction Science,2019,204:183-192.
[73] ZHAO Y,PU Y,LIANG B,et al.A study using single-locus and multi-locus genome-wide association study to identify genes associated with teat number in Hu sheep[J].Animal Genetics,2022,53(2):203-211.
[74] COSSO G,NEHME M,LURIDIANA S,et al.Detection of polymorphisms in the mtnr1a gene and their association with reproductive performance in Awassi ewes[J].Animals,2021,11(2):583.
[75] LUO R,ZHANG X,WANG L,et al. GLIS1,a potential candidate gene affect fat deposition in sheep tail[J].Molecular Biology Reports,2021,48(5):4925-4931.
[76] PAN Z,LI S,LIU Q,et al.Rapid evolution of a retro-transposable hotspot of ovine genome underlies the alteration of BMP2 expression and development of fat tails[J].BMC Genomics,2019,20(1):261.
[77] YUAN Z,LIU E,LIU Z,et al.Selection signature analysis reveals genes associated with tail type in Chinese indigenous sheep[J].Animal Genetics,2017,48(1):55-66.
[78] LU Z,LIU J,HAN J,et al.Association between BMP2 functional polymorphisms and sheep tail type[J].Animals,2020,10(4):739.
[79] ZHU C,LI N,CHENG H,et al.Genome wide association study for the identification of genes associated with tail fat deposition in Chinese sheep breeds[J].Biology Open,2021,10(5):bio054932.
[80] LI Q,LU Z,JIN M,et al.Verification and analysis of sheep tail type-associated PDGF-D gene polymorphisms[J].Animals,2020,10(1):89.
[81] LI X,YANG J,SHEN M,et al.Whole-genome resequencing of wild and domestic sheep identifies genes associated with morphological and agronomic traits[J].Nature Communications,2020,11(1):2815.
[82] DONG K,YANG M,HAN J,et al.Genomic analysis of worldwide sheep breeds reveals PDGFD as a major target of fat-tail selection in sheep[J].BMC Genomics,2020,21(1):800.
[83] HAN J,MA S,LIANG B,et al.Transcriptome profiling of developing ovine fat tail tissue reveals an important role for mtfp1 in regulation of adipogenesis[J].Frontiers in Cell and Developmental Biology,2022,10:839731.
[84] WANG Z,ZHANG H,YANG H,et al.Genome-wide association study for wool production traits in a Chinese Merino sheep population[J].PLoS One,2014,9(9):e107101.
[85] WANG C,YUAN Z,HU R,et al.Association of SNPs within PTPN3 gene with wool production and growth traits in a dual-purpose sheep population[J].Animal Biotechnology,2022,Doi:10.1080/10495398.2022.2029465.Online ahead of print.
[86] LEE J,TUMBAR T.Hairy tale of signaling in hair follicle development and cycling[J]. Seminars in Cell and Developmental Biology,2012,23(8):906-916.
[87] ZHAO H,HU R,LI F,et al.Five SNPs within the FGF5 gene significantly affect both wool traits and growth performance in Fine-wool sheep (Ovis aries)[J].Frontiers in Genetics,2021,12:732097.
[88] ZHANG R,WU H,LIAN Z.Bioinformatics analysis of evolutionary characteristics and biochemical structure of FGF5 gene in sheep[J].Gene,2019,702:123-132.
[89] LIU Y X,SHI G Q,WANG H X,et al.Polymorphisms of KAP6,KAP7,and KAP8 genes in four Chinese sheep breeds[J].Genetics and Molecular Research,2014,13(2):3438-3445.
[90] SALLAM A M,GAD-ALLAH A A,AL-BITAR E M.Association analysis of the ovine KAP6-1 gene and wool traits in Barki sheep[J].Animal Biotechnology,2021,32(6):733-739.
[91] ZHOU H,GONG H,LI S,et al.A 57-bp deletion in the ovine KAP6-1 gene affects wool fibre diameter[J].Journal of Animal Breeding and Genetics,2015,132(4):301-307.
[92] GONG H,ZHOU H,HICKFORD J G H.Diversity of the glycine/tyrosine-rich keratin-associated protein 6 gene (KAP6) family in sheep[J].Molecular Biology Reports,2011,38(1):31-35.
[93] LI S,ZHOU H,GONG H,et al.Variation in the ovine KAP6-3 gene (KRTAP6-3) is associated with variation in mean fibre diameter-associated wool traits[J].Genes,2017,8(8):204.
[94] ULLAH F,JAMAL S M,EKEGBU U J,et al.Polymorphism in the ovine keratin-associated protein gene KRTAP7-1 and its association with wool characteristics[J]. Journal of Animal Science,2020,98(1):skz381.
[95] ZHAO H,GUO T,LU Z,et al.Genome-wide association studies detects candidate genes for wool traits by re-sequencing in Chinese Fine-wool sheep[J].BMC Genomics,2021,22(1):127.
[96] ZHANG Y,XUE X,LIU Y,et al.Genome-wide comparative analyses reveal selection signatures underlying adaptation and production in Tibetan and Poll Dorset sheep[J].Scientific Reports,2021,11(1):2466.
[97] ZHAO B,LUO H,HE J,et al.Comprehensive transcriptome and methylome analysis delineates the biological basis of hair follicle development and wool-related traits in Merino sheep[J].BMC Biology,2021,19(1):197.
[98] ZHAO B,LUO H,HUANG X,et al.Integration of a single-step genome-wide association study with a multi-tissue transcriptome analysis provides novel insights into the genetic basis of wool and weight traits in sheep[J].Genetics Selection Evolution,2021,53(1):56.
[99] NORRIS B J,WHAN V A.A gene duplication affecting expression of the ovine ASIP gene is responsible for white and black sheep[J].Genome Research,2008,18(8):1282-1293.
[100] LI M H,TIIRIKKA T,KANTANEN J.A genome-wide scan study identifies a single nucleotide substitution in ASIP associated with white versus non-white coat-colour variation in sheep (Ovis aries)[J]. Heredity,2014,112(2):122-131.
[101] KIJAS J W,LENSTRA J A,HAYES B,et al.Genome-wide analysis of the world's sheep breeds reveals high levels of historic mixture and strong recent selection[J].PLoS Biology,2012,10(2):e1001258.
[102] FARIELLO M I,SERVIN B,TOSSER-KLOPP G,et al.Selection signatures in worldwide sheep populations[J].PLoS One,2014,9(8):e103813.
[103] YAO L,BAO A,HONG W,et al.Transcriptome profiling analysis reveals key genes of different coat color in sheep skin[J].PeerJ,2019,7:e8077.
[104] FAN R,XIE J,BAI J,et al.Skin transcriptome profiles associated with coat color in sheep[J].BMC Genomics,2013,14:389.
[105] GAO J,LYU Y,ZHANG D,et al.Genomic characteristics and selection signatures in indigenous Chongming white goat (Capra hircus)[J].Frontiers in Genetics,2020,11:901.
[106] REN H,WANG G,JIANG J,et al.Comparative transcriptome and histological analyses provide insights into the prenatal skin pigmentation in goat (Capra hircus)[J].Physiological Genomics,2017,49(12):703-711.
[107] SHANG S,YU Y,ZHAO Y,et al.Synergy between MC1R and ASIP for coat color in horses (Equus caballus)1[J].Journal of Animal Science, 2019,97(4):1578-1585.
[108] BERTOLINI F,MOSCATELLI G,SCHIAVO G,et al.Signatures of selection are present in the genome of two close autochthonous cattle breeds raised in the North of Italy and mainly distinguished for their coat colours[J].Journal of Animal Breeding and Genetics,2022,139(3):307-319.
[109] YANG C W,RAN J S,YU C L,et al.Polymorphism in MC1R,TYR and ASIP genes in different colored feather chickens[J].3 Biotech,2019,9(5):203.
[110] ALSHANBARI F,CASTANEDA C,JURAS R,et al.Comparative FISH-mapping of MC1R,ASIP,and TYRP1 in new and old world camelids and association analysis with coat color phenotypes in the dromedary (Camelus dromedarius)[J].Frontiers in Genetics,2019,10:340.
[111] GEBRESELASSIE G,LIANG B,BERIHULAY H,et al.Genomic mapping identifies two genetic variants in the MC1R gene for coat colour variation in Chinese Tan sheep[J].PLoS One,2020,15,8:e0235426.
[112] YANG G L,FU D L,LANG X,et al.Mutations in MC1R gene determine black coat color phenotype in Chinese sheep[J].The Scientific World Journal,2013,2013:675382.
[113] DENG W D,SHU W,YANG S L,et al.Pigmentation in Black-boned sheep (Ovis aries):Association with polymorphism of the MC1R gene[J].Molecular Biology Reports,2009,36(3):431-436.
[114] MAHMOUD A H,MASHALY A M,RADY A M,et al.Allelic variation of melanocortin-1 receptor locus in Saudi indigenous sheep exhibiting different color coats[J].Asian-Australasian Journal of Animal Sciences,2017,30(2):154-159.
[115] HEPP D,GONÇALVES G L,MOREIRA G R,et al.Identification of the e allele at the extension locus (MC1R) in Brazilian Creole sheep and its role in wool color variation[J].Genetics and Molecular Research,2012,11(3):2997-3006.
[116] HAN J L,YANG M,YUE Y J,et al.Analysis of agouti signaling protein (ASIP) gene polymorphisms and association with coat color in Tibetan sheep (Ovis aries)[J].Genetics and Molecular Research,2015,14(1):1200-1209.
[117] ZHANG X,LI W,LIU C,et al.Alteration of sheep coat color pattern by disruption of ASIP gene via CRISPR Cas9[J].Scientific Reports,2017,7(1):8194.