乌骨鸡黑色素沉积机理及相关候选基因
|
摘要
乌骨鸡的乌色度与其营养特性和药用价值密切相关,是乌骨鸡重要的经济性状之一。乌骨鸡乌色度受机体黑色素沉积影响。本文对黑色素发生、沉积机理进行较为全面的综述,并讨论了影响乌骨鸡黑色素沉积的候选基因,旨在增进对乌骨鸡黑色素性状及相关分子选育的理解和研究。
The blackness of silky fowls,as one of the most important economic traits,is often considered to be very relevant to the nutritional quality and medicinal value. The degree of blackness in silky fowls is mainly affected by the deposition level of melanin. This article reviewed the mechanisms of the synthesis and deposition of melanin,as well as the candidate genes involving melanin deposition,for the purpose of improving outstanding for melanin and promoting molecular breeding plan of silky fowls.
The blackness of silky fowls,as one of the most important economic traits,is often considered to be very relevant to the nutritional quality and medicinal value. The degree of blackness in silky fowls is mainly affected by the deposition level of melanin. This article reviewed the mechanisms of the synthesis and deposition of melanin,as well as the candidate genes involving melanin deposition,for the purpose of improving outstanding for melanin and promoting molecular breeding plan of silky fowls.
引文
[1]李瑞成,尚永彪,管俊峰.乌骨鸡黑色素及其研究进展[J].肉类研究,2010(10):54-59.
[2]REEDY M V,FARACO C D,ERICKSON C A.Specification and migration of melanoblasts at the vagal level and in hyperpigmented silkie chickens[J].Developmental Dynamics,1998,213(4):476-485.
[3]ADAMEYKO I,LALLEMEND F,AQUINO J B,et al.Schwann cell precursors from nerve innervation are a cellular origin of melanocytes in skin[J].Cell,2009,139(2):366-379.
[4]MOTOHASHI T,KUNISADA T.Chapter three-extended multipotency of neural crest cells and neural crest-derived cells[J].Current Topics in Developmental Biology,2015,111:69-95.
[5]HENION P D,WESTON J A.Timing and pattern of cell fate restrictions in the neural crest lineage[J].Development,1997,124(21):4351-4359.
[6]LORING J F,ERICKSON C A.Neural crest cell migratory pathways in the trunk of the chick embryo[J].Developmental Biology,1987,121(1):220-236.
[7]DOTTORI M,GROSS M K,LABOSKY P,et al.The winged-helix transcription factor Foxd3 suppresses interneuron differentiation and promotes neural crest cell fate[J].Development,2001,128(21):4127-4138.
[8]KOS R,REEDY M V,JOHNSON R L,et al.The winged-helix transcription factor FoxD3 is important for establishing the neural crest lineage and repressing melanogenesis in avian embryos[J].Development,2001,128(8):1467-1479.
[9]MARTINA J A,DIAB H I,LI H Q,et al.Novel roles for the Mitf/TFE family of transcription factors in organelle biogenesis,nutrient sensing,and energy homeostasis[J].Cellular and Molecular Life Sciences,2014,71(13):2483-2497.
[10]MAYANIL C S K,GEORGE D,MANIA-FARNELL B,et al.Overexpression of murine Pax3 increases NCAM polysialylation in a human medulloblastoma cell line[J].Journal of Biological Chemistry,2000,275(30):23259-23266.
[11]周洋,崔英霞.PAX3对黑色素细胞形成的调控[J].中国优生与遗传杂志,2012,20(3):10-11,128.
[12]PANI L,HORAL M,LOEKEN M R,et al.Rescue of neural tube defects in Pax-3-deficient embryos by p53loss of function:implications for Pax-3-dependent development and tumorigenesis[J].Genes&Development,2002,16(6):676-680.
[13]田颖刚,徐德利,廖春艳.泰和乌骨鸡黑色素细胞形态与分布特征[J].中国家禽,2015,37(1):5-8.
[14]蒋明.乌鸡黑色素沉积机理及分子标记筛选的研究[D].硕士学位论文.长沙:湖南农业大学,2016:11-16.
[15]PARK H Y,KOSMADAKI M G,YAAR M,et al.Cellular mechanisms regulating human melanogenesis[J].Cellular and Molecular Life Sciences,2009,66(9):1493-1506.
[16]熊渺.α-MSH和cAMP对泰和乌骨鸡黑色素细胞增殖及黑色素合成的影响研究[D].硕士学位论文.南昌:江西农业大学,2014:2-5.
[17]TAPIA J C D,BAGUTTI C,COTTI R,et al.Induction of constitutive melanogenesis in amelanotic mouse melanoma cells by transfection of the human melanocortin-1 receptor gene[J].Journal of Cell Science,1996,109(8):2023-2030.
[18]THODY A J.α-MSH and the regulation of melanocyte function[J].Annals of the New York Academy of Sciences,1999,885(1):217-229.
[19]FURUMURA M,SAKAI C,ABDELMALEK Z A,et al.The interaction of agouti signal protein and melanocyte stimulating hormone to regulate melanin formation in mammals[J].Pigment Cell Research,1996,9(4):191-203.
[20]SAKAI C,OLLMANN M M,KOBAYASHI T,et al.Modulation of murine melanocyte function in vitro by agouti signal protein[J].The EMBO Journal,1997,16(12):3544-3552.
[21]杨朋坤.鸡真皮黑色素抑制基因精细定位及相关质量性状分子标记鉴定研究[D].博士学位论文.郑州:河南农业大学,2014:7-11.
[22]HARA M,YAAR M,BYERS H R,etal.Kinesin participates in melanosomal movement along melanocyte dendrites[J].Journal of Investigative Dermatology,2000,114(3):438-443.
[23]WESTBROEK W,LAMBERT J,NAEYAERT J M.The Dilute locus and griscelli syndrome:gateways towards a better understanding of melanosome transport[J].Pigment Cell Research,2001,14(5):320-327.
[24]WESTBROEK W,LAMBERT J,DE SCHEPPER S,et al.Rab27b is up-regulated in human griscelli syndrome typeⅡmelanocytes and linked to the actin cytoskeleton via exon F-myosin Va transcripts[J].Pigment Cell Research,2004,17(5):498-505.
[25]WEI Q,WU X F,HAMMER J A.The predominant defect in dilute melanocytes is in melanosome distribution and not cell shape,supporting a role for myosin V in melanosome transport[J].Journal of Muscle Research and Cell Motility,1997,18(5):517-527.
[26]SEIBERG M.Keratinocyte-melanocyte interactions during melanosome transfer[J].Pigment Cell Research,2001,14(4):236-242.
[27]GREATENS A,HAKOZAKI T,KOSHOFFER A,et al.Effective inhibition of melanosome transfer to keratinocytes by lectins and niacinamide is reversible[J].Experimental Dermatology,2005,14(7):498-508.
[28]VAN DEN BOSSCHE K V,NAEYAERT J M,LAMBERT J,et al.The quest for the mechanism of melanin transfer[J].Traffic,2006,7(7):769-778.
[29]VAEZ M,FOLLETT S A,BED’HOM B,et al.A single point-mutation within the melanophilin gene causes the lavender plumage colour dilution phenotype in the chicken[J].BMC Genetics,2008,9(1):7.
[30]BED’HOM B,VAEZ M,COVILLE J,et al.The lavender plumage colour in Japanese quail is associated with a complex mutation in the region of MLPH that is related to differences in growth,f eed consumption and body temperature[J].BMC Genomics,2012,13:442.
[31]CIRERA S,MARKAKIS M N,CHRISTENSEN K,et al.New insights into the melanophilin(MLPH)gene controlling coat color phenotypes in American mink[J].Gene,2013,527(1):48-54.
[32]XU J G,XIE M G,ZOU S Y,et al.Interactions of allele E of the MC1R gene with FM and mutations in the MLPH gene cause the five-gray phenotype in the Anyi tile-like gray chicken[J].Genetics and Molecular Research,2016,15(2),doi:10.4238/gmr.15027633.
[33]郑嫩珠,辛清武,缪中纬,等黑色素基因在半番鸭及番鸭不同羽色中的差异表达研究[J].福建农业学报,2013,28(5):427-431.
[34]XU Y,ZHANG X H,PANG Y Z,et al.Association of tyrosinase(TYR)and tyrosinase-related protein 1(TYRP1)with melanic plumage color in Korean quails(Coturnix coturnix)[J].Asian-Australasian Journal of Animal Sciences,2013,26(11):1518-1522.
[35]DENOVAN-WRIGHT E M,RAMSEY N B,MCCORMICK C J,et al.Nucleotide sequence of transferrin cDNAs and tissue-specific expression of the transferrin gene in Atlantic cod(Gadus morhua)[J].Comparative Biochemistry and Physiology Part B:Biochemistry and Molecular Biology,1996,113(2):269-273.
[36]郑嫩珠,李丽,辛清武,等.酪氨酸酶(TYR)、小眼畸形相关转录因子(Mitf)和刺鼠信号蛋白(ASIP)基因对白绒乌骨鸡黑色素沉积的遗传效应[J].农业生物技术学报,2015,23(8):1076-1083.
[37]陈志强,邓学梅,周军,等.鸡的酪氨酸酶基因5'调控区的单链构象多态性分析[J].农业生物技术学报,2005,13(2):191-194.
[38]ALALUF S,BARRETT K,BLOUNT M,et al.Ethnic variation in tyrosinase and TYRP1 expression in photoexposed and photoprotected human skin[J].Pigment Cell Research,2003,16(1):35-42.
[39]LI B,HE X L,ZHAO Y P,et al.Tyrosinase-related protein 1(TYRP1)gene polymorphism and skin differential expression related to coat color in Mongolian horse[J].Livestock Science,2014,167:58-64.
[40]刘薇.丝羽乌骨鸡BAC文库的构建和黑色素相关基因TYRP1和ID的研究[D].博士学位论文.北京:中国农业大学,2004:78-82.
[41]刘甲斐,仇雪梅.黑色素及其相关基因的研究进展[J].生物技术通报,2007(4):55-58.
[42]RAPOSO G,MARKS M S.Melanosomes-dark organelles enlighten endosomal membrane transport[J].Nature Reviews Molecular Cell Biology,2007,8(10):786-797.
[43]HELLSTROM A R,WATT B,FARD S S,et al.Inactivation of Pmel alters melanosome shape but has only a subtle effect on visible pigmentation[J].PLoS Genetics,2011,7(9):e1002285.
[44]KWON B S,HALABAN R,PONNAZHAGAN S,et al.Mouse silver mutation is caused by a single base insertion in the putative cytoplasmic domain of Pmel17[J].Nucleic Acids Research,1995,23(1):154-158.
[45]MARTINEZ-ESPARZA M,JIMENEZ-CERVANTES C,BENNETT D C,et al.The mouse silver locus encodes a single transcript truncated by the silver mutation[J].Mammalian Genome,1999,10(12):1168-1171.
[46]BRUNBERG E,ANDERSSON L,COTHRAN G,et al.A missense mutation in Pmel17 is associated with the Silver coat color in the horse[J].BMC Genetics,2006,7(1):46-55.
[47]KERJE S,SHARMA P M,GUNNARSSON U,et al.The Dominant white,Dun and Smoky color variants in chicken are associated with insertion/deletion polymorphisms in the Pmel17 gene[J].Genetics,2004,168(3):1507-1518.
[48]TAKEUCHI S,SUZUKI H,YABUUCHI M,et al.Molecular cloning and sequence analysis of the chick melanocortin-1 receptor gene[J].Biochimica et Biophysica Acta(BBA):Gene Structure and Expression,1996,1306(2/3):122-126.
[49]LU D,WILLARD D,PATEL I R,et al.Agouti protein is an antagonist of the melanocyte-stimulating hormone receptor[J].Nature,1994,371(6500):799-802.
[50]杜晓明.边鸡MC1R和TYR基因序列及mRNA表达研究[D].硕士学位论文.太谷:山西农业大学,2013:29-41.
[51]杨永升,邓学梅,李宁,等.MC1R是控制鸡黑色素形成的候选主效基因[J].生物化学与生物物理进展,2004,31(6):500-505.
[52]LING M K,LAGERSTRM M C,FREDRIKSSON R,et al.Association of feather colour with constitutively active melanocortin 1 receptors in chicken[J].The FEBS Journal,2003,270(7):1441-1449.
[53]迟良,李兰,潘庆杰,等.乌骨鸡黑色素皮质激素受体-1基因的克隆、序列分析与原核表达[J].中国农业科学,2012,45(5):966-972.
[54]VACHTENHEIM J,BOROVANSK Y'J.“Transcription physiology”of pigment formation in melanocytes:central role of Mitf[J].Experimental Dermatology,2010,19(7):617-627.
[55]ZHU Z,HE J,JIA X,et al.MicroRNA-25 functions in regulation of pigmentation by targeting the transcription factor Mitf in Alpaca(Lama pacos)skin melanocytes[J].Domestic Animal Endocrinology,2010,38(3):200-209.
[56]LI S J,WANG C,YU W H,et al.Identification of genes related to white and black plumage formation by RNA-Seq from white and black feather bulbs in ducks[J].PLoS One,2012,7(5):e36592.
[57]MINVIELLE F,BED’HOM B,COVILLE J,et al.The“silver”Japanese quail and the Mitf gene:causal mutation,a ssociated traits and homology with the“blue” chicken plumage[J].BMC Genetics,2010,11:15.
[58]WANG Y,LI S M,HUANG J,et al.Mutations of TYR and Mitf genes are associated with plumage colour phenotypes in geese[J].Asian-Australasian Journal of Animal Sciences,2014,27(6):778-783.
[59]郑嫩珠,辛清武,朱志明,等.白绒乌鸡Mitf基因的cDNA克隆、表达及其对黑色素沉积的影响[J].中国农业科学,2015,48(18):3711-3718.
[60]ISHIDA Y,DAVID V A,EIZIRIK E,et al.A homozygous single-base deletion in MLPH causes the dilute coat color phenotype in the domestic cat[J].Genomics,2006,88(6):698-705.
[61]PHILIPP U,HAMANN H,MECKLENBURG L,et al.Polymorphisms within the canine MLPH gene are associated with dilute coat color in dogs[J].BMC Genetics,2005,6:34.
[62]FONTANESI L,SCOTTI E,ALLAIN D,et al.A frameshift mutation in the melanophilin gene causes the dilute coat colour in rabbit(Oryctolagus cuniculus)breeds[J].Animal Genetics,2014,45(2):248-255.
[63]刘杰,刘毅,王翠,等.鹅MLPH基因分子克隆、组织表达及SNP检测[J].中国家禽,2016,38(12):4-9.
[2]REEDY M V,FARACO C D,ERICKSON C A.Specification and migration of melanoblasts at the vagal level and in hyperpigmented silkie chickens[J].Developmental Dynamics,1998,213(4):476-485.
[3]ADAMEYKO I,LALLEMEND F,AQUINO J B,et al.Schwann cell precursors from nerve innervation are a cellular origin of melanocytes in skin[J].Cell,2009,139(2):366-379.
[4]MOTOHASHI T,KUNISADA T.Chapter three-extended multipotency of neural crest cells and neural crest-derived cells[J].Current Topics in Developmental Biology,2015,111:69-95.
[5]HENION P D,WESTON J A.Timing and pattern of cell fate restrictions in the neural crest lineage[J].Development,1997,124(21):4351-4359.
[6]LORING J F,ERICKSON C A.Neural crest cell migratory pathways in the trunk of the chick embryo[J].Developmental Biology,1987,121(1):220-236.
[7]DOTTORI M,GROSS M K,LABOSKY P,et al.The winged-helix transcription factor Foxd3 suppresses interneuron differentiation and promotes neural crest cell fate[J].Development,2001,128(21):4127-4138.
[8]KOS R,REEDY M V,JOHNSON R L,et al.The winged-helix transcription factor FoxD3 is important for establishing the neural crest lineage and repressing melanogenesis in avian embryos[J].Development,2001,128(8):1467-1479.
[9]MARTINA J A,DIAB H I,LI H Q,et al.Novel roles for the Mitf/TFE family of transcription factors in organelle biogenesis,nutrient sensing,and energy homeostasis[J].Cellular and Molecular Life Sciences,2014,71(13):2483-2497.
[10]MAYANIL C S K,GEORGE D,MANIA-FARNELL B,et al.Overexpression of murine Pax3 increases NCAM polysialylation in a human medulloblastoma cell line[J].Journal of Biological Chemistry,2000,275(30):23259-23266.
[11]周洋,崔英霞.PAX3对黑色素细胞形成的调控[J].中国优生与遗传杂志,2012,20(3):10-11,128.
[12]PANI L,HORAL M,LOEKEN M R,et al.Rescue of neural tube defects in Pax-3-deficient embryos by p53loss of function:implications for Pax-3-dependent development and tumorigenesis[J].Genes&Development,2002,16(6):676-680.
[13]田颖刚,徐德利,廖春艳.泰和乌骨鸡黑色素细胞形态与分布特征[J].中国家禽,2015,37(1):5-8.
[14]蒋明.乌鸡黑色素沉积机理及分子标记筛选的研究[D].硕士学位论文.长沙:湖南农业大学,2016:11-16.
[15]PARK H Y,KOSMADAKI M G,YAAR M,et al.Cellular mechanisms regulating human melanogenesis[J].Cellular and Molecular Life Sciences,2009,66(9):1493-1506.
[16]熊渺.α-MSH和cAMP对泰和乌骨鸡黑色素细胞增殖及黑色素合成的影响研究[D].硕士学位论文.南昌:江西农业大学,2014:2-5.
[17]TAPIA J C D,BAGUTTI C,COTTI R,et al.Induction of constitutive melanogenesis in amelanotic mouse melanoma cells by transfection of the human melanocortin-1 receptor gene[J].Journal of Cell Science,1996,109(8):2023-2030.
[18]THODY A J.α-MSH and the regulation of melanocyte function[J].Annals of the New York Academy of Sciences,1999,885(1):217-229.
[19]FURUMURA M,SAKAI C,ABDELMALEK Z A,et al.The interaction of agouti signal protein and melanocyte stimulating hormone to regulate melanin formation in mammals[J].Pigment Cell Research,1996,9(4):191-203.
[20]SAKAI C,OLLMANN M M,KOBAYASHI T,et al.Modulation of murine melanocyte function in vitro by agouti signal protein[J].The EMBO Journal,1997,16(12):3544-3552.
[21]杨朋坤.鸡真皮黑色素抑制基因精细定位及相关质量性状分子标记鉴定研究[D].博士学位论文.郑州:河南农业大学,2014:7-11.
[22]HARA M,YAAR M,BYERS H R,etal.Kinesin participates in melanosomal movement along melanocyte dendrites[J].Journal of Investigative Dermatology,2000,114(3):438-443.
[23]WESTBROEK W,LAMBERT J,NAEYAERT J M.The Dilute locus and griscelli syndrome:gateways towards a better understanding of melanosome transport[J].Pigment Cell Research,2001,14(5):320-327.
[24]WESTBROEK W,LAMBERT J,DE SCHEPPER S,et al.Rab27b is up-regulated in human griscelli syndrome typeⅡmelanocytes and linked to the actin cytoskeleton via exon F-myosin Va transcripts[J].Pigment Cell Research,2004,17(5):498-505.
[25]WEI Q,WU X F,HAMMER J A.The predominant defect in dilute melanocytes is in melanosome distribution and not cell shape,supporting a role for myosin V in melanosome transport[J].Journal of Muscle Research and Cell Motility,1997,18(5):517-527.
[26]SEIBERG M.Keratinocyte-melanocyte interactions during melanosome transfer[J].Pigment Cell Research,2001,14(4):236-242.
[27]GREATENS A,HAKOZAKI T,KOSHOFFER A,et al.Effective inhibition of melanosome transfer to keratinocytes by lectins and niacinamide is reversible[J].Experimental Dermatology,2005,14(7):498-508.
[28]VAN DEN BOSSCHE K V,NAEYAERT J M,LAMBERT J,et al.The quest for the mechanism of melanin transfer[J].Traffic,2006,7(7):769-778.
[29]VAEZ M,FOLLETT S A,BED’HOM B,et al.A single point-mutation within the melanophilin gene causes the lavender plumage colour dilution phenotype in the chicken[J].BMC Genetics,2008,9(1):7.
[30]BED’HOM B,VAEZ M,COVILLE J,et al.The lavender plumage colour in Japanese quail is associated with a complex mutation in the region of MLPH that is related to differences in growth,f eed consumption and body temperature[J].BMC Genomics,2012,13:442.
[31]CIRERA S,MARKAKIS M N,CHRISTENSEN K,et al.New insights into the melanophilin(MLPH)gene controlling coat color phenotypes in American mink[J].Gene,2013,527(1):48-54.
[32]XU J G,XIE M G,ZOU S Y,et al.Interactions of allele E of the MC1R gene with FM and mutations in the MLPH gene cause the five-gray phenotype in the Anyi tile-like gray chicken[J].Genetics and Molecular Research,2016,15(2),doi:10.4238/gmr.15027633.
[33]郑嫩珠,辛清武,缪中纬,等黑色素基因在半番鸭及番鸭不同羽色中的差异表达研究[J].福建农业学报,2013,28(5):427-431.
[34]XU Y,ZHANG X H,PANG Y Z,et al.Association of tyrosinase(TYR)and tyrosinase-related protein 1(TYRP1)with melanic plumage color in Korean quails(Coturnix coturnix)[J].Asian-Australasian Journal of Animal Sciences,2013,26(11):1518-1522.
[35]DENOVAN-WRIGHT E M,RAMSEY N B,MCCORMICK C J,et al.Nucleotide sequence of transferrin cDNAs and tissue-specific expression of the transferrin gene in Atlantic cod(Gadus morhua)[J].Comparative Biochemistry and Physiology Part B:Biochemistry and Molecular Biology,1996,113(2):269-273.
[36]郑嫩珠,李丽,辛清武,等.酪氨酸酶(TYR)、小眼畸形相关转录因子(Mitf)和刺鼠信号蛋白(ASIP)基因对白绒乌骨鸡黑色素沉积的遗传效应[J].农业生物技术学报,2015,23(8):1076-1083.
[37]陈志强,邓学梅,周军,等.鸡的酪氨酸酶基因5'调控区的单链构象多态性分析[J].农业生物技术学报,2005,13(2):191-194.
[38]ALALUF S,BARRETT K,BLOUNT M,et al.Ethnic variation in tyrosinase and TYRP1 expression in photoexposed and photoprotected human skin[J].Pigment Cell Research,2003,16(1):35-42.
[39]LI B,HE X L,ZHAO Y P,et al.Tyrosinase-related protein 1(TYRP1)gene polymorphism and skin differential expression related to coat color in Mongolian horse[J].Livestock Science,2014,167:58-64.
[40]刘薇.丝羽乌骨鸡BAC文库的构建和黑色素相关基因TYRP1和ID的研究[D].博士学位论文.北京:中国农业大学,2004:78-82.
[41]刘甲斐,仇雪梅.黑色素及其相关基因的研究进展[J].生物技术通报,2007(4):55-58.
[42]RAPOSO G,MARKS M S.Melanosomes-dark organelles enlighten endosomal membrane transport[J].Nature Reviews Molecular Cell Biology,2007,8(10):786-797.
[43]HELLSTROM A R,WATT B,FARD S S,et al.Inactivation of Pmel alters melanosome shape but has only a subtle effect on visible pigmentation[J].PLoS Genetics,2011,7(9):e1002285.
[44]KWON B S,HALABAN R,PONNAZHAGAN S,et al.Mouse silver mutation is caused by a single base insertion in the putative cytoplasmic domain of Pmel17[J].Nucleic Acids Research,1995,23(1):154-158.
[45]MARTINEZ-ESPARZA M,JIMENEZ-CERVANTES C,BENNETT D C,et al.The mouse silver locus encodes a single transcript truncated by the silver mutation[J].Mammalian Genome,1999,10(12):1168-1171.
[46]BRUNBERG E,ANDERSSON L,COTHRAN G,et al.A missense mutation in Pmel17 is associated with the Silver coat color in the horse[J].BMC Genetics,2006,7(1):46-55.
[47]KERJE S,SHARMA P M,GUNNARSSON U,et al.The Dominant white,Dun and Smoky color variants in chicken are associated with insertion/deletion polymorphisms in the Pmel17 gene[J].Genetics,2004,168(3):1507-1518.
[48]TAKEUCHI S,SUZUKI H,YABUUCHI M,et al.Molecular cloning and sequence analysis of the chick melanocortin-1 receptor gene[J].Biochimica et Biophysica Acta(BBA):Gene Structure and Expression,1996,1306(2/3):122-126.
[49]LU D,WILLARD D,PATEL I R,et al.Agouti protein is an antagonist of the melanocyte-stimulating hormone receptor[J].Nature,1994,371(6500):799-802.
[50]杜晓明.边鸡MC1R和TYR基因序列及mRNA表达研究[D].硕士学位论文.太谷:山西农业大学,2013:29-41.
[51]杨永升,邓学梅,李宁,等.MC1R是控制鸡黑色素形成的候选主效基因[J].生物化学与生物物理进展,2004,31(6):500-505.
[52]LING M K,LAGERSTRM M C,FREDRIKSSON R,et al.Association of feather colour with constitutively active melanocortin 1 receptors in chicken[J].The FEBS Journal,2003,270(7):1441-1449.
[53]迟良,李兰,潘庆杰,等.乌骨鸡黑色素皮质激素受体-1基因的克隆、序列分析与原核表达[J].中国农业科学,2012,45(5):966-972.
[54]VACHTENHEIM J,BOROVANSK Y'J.“Transcription physiology”of pigment formation in melanocytes:central role of Mitf[J].Experimental Dermatology,2010,19(7):617-627.
[55]ZHU Z,HE J,JIA X,et al.MicroRNA-25 functions in regulation of pigmentation by targeting the transcription factor Mitf in Alpaca(Lama pacos)skin melanocytes[J].Domestic Animal Endocrinology,2010,38(3):200-209.
[56]LI S J,WANG C,YU W H,et al.Identification of genes related to white and black plumage formation by RNA-Seq from white and black feather bulbs in ducks[J].PLoS One,2012,7(5):e36592.
[57]MINVIELLE F,BED’HOM B,COVILLE J,et al.The“silver”Japanese quail and the Mitf gene:causal mutation,a ssociated traits and homology with the“blue” chicken plumage[J].BMC Genetics,2010,11:15.
[58]WANG Y,LI S M,HUANG J,et al.Mutations of TYR and Mitf genes are associated with plumage colour phenotypes in geese[J].Asian-Australasian Journal of Animal Sciences,2014,27(6):778-783.
[59]郑嫩珠,辛清武,朱志明,等.白绒乌鸡Mitf基因的cDNA克隆、表达及其对黑色素沉积的影响[J].中国农业科学,2015,48(18):3711-3718.
[60]ISHIDA Y,DAVID V A,EIZIRIK E,et al.A homozygous single-base deletion in MLPH causes the dilute coat color phenotype in the domestic cat[J].Genomics,2006,88(6):698-705.
[61]PHILIPP U,HAMANN H,MECKLENBURG L,et al.Polymorphisms within the canine MLPH gene are associated with dilute coat color in dogs[J].BMC Genetics,2005,6:34.
[62]FONTANESI L,SCOTTI E,ALLAIN D,et al.A frameshift mutation in the melanophilin gene causes the dilute coat colour in rabbit(Oryctolagus cuniculus)breeds[J].Animal Genetics,2014,45(2):248-255.
[63]刘杰,刘毅,王翠,等.鹅MLPH基因分子克隆、组织表达及SNP检测[J].中国家禽,2016,38(12):4-9.