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Вавиловский журнал генетики и селекции

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Молекулярно-генетические основы формирования окраски оперения у кур

https://doi.org/10.18699/VJ19.499

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Аннотация

Окраска оперения – важный признак у птиц, нередко определяющий принадлежность к тому или иному виду или породе. Окраска является результатом действия веществ, которые поглощают определенную длину волны и формируют так называемые пигментные цвета, и оптическим эффектом, обусловленным интерференцией света, отраженного биологическими микроструктурами пера. Основой для формирования окраски служит синтез меланина. Эумеланин ответственен за черные и коричневые оттенки, а феомеланин отвечает за красновато-коричневые оттенки. Молекулярно-генетический механизм появления того или иного типа окраски еще до конца не изучен, поскольку на один и тот же признак могут влиять несколько генов. Первичная пигментация оперения определяется взаимодействием полиморфных вариантов гена MC1R и генов, участвующих в регуляции меланогенеза. Гены-модификаторы вызывают изменение окраски любого генотипа по локусу E и могут как уменьшать или увеличивать экспрес сию эумеланина, так и разрушать меланоциты. Вторичная пигментация оперения определяется белыми пятнами или специфическим распределением эумеланина на отдельных перьях. Современные методы анализа ДНК, такие как секвенирование, полногеномный анализ с использованием чипов различной плотности, анализ экспрессии генов, позволяют получать новые данные о генах, определяющих окраску оперения.

Об авторах

А. В. Макарова
Всероссийский научно-исследовательский институт генетики и разведения сельскохозяйственных животных – филиал Федерального научного центра животноводства – ВИЖ им. академика Л.К. Эрнста
Россия


О. В. Митрофанова
Всероссийский научно-исследовательский институт генетики и разведения сельскохозяйственных животных – филиал Федерального научного центра животноводства – ВИЖ им. академика Л.К. Эрнста
Россия


А. Б. Вахрамеев
Всероссийский научно-исследовательский институт генетики и разведения сельскохозяйственных животных – филиал Федерального научного центра животноводства – ВИЖ им. академика Л.К. Эрнста
Россия


Н. В. Дементьева
Всероссийский научно-исследовательский институт генетики и разведения сельскохозяйственных животных – филиал Федерального научного центра животноводства – ВИЖ им. академика Л.К. Эрнста
Россия


Список литературы

1. Alekseevich L.A., Barabanova L.V., Suller I.L. Genetics of Domesticated Animals. St. Petersburg–Lomonosov, 2000. (in Russian)

2. Ancans J., Tobin D.J., Hoogduijn M.J., Smit N.P., Wakamatsu K., Thody A.J. Melanosomal pH controls rate of melanogenesis, eumelanin/phaeomelanin ratio and melanosome maturation in melanocytes and melanoma cells. Exp. Cell Res. 2001;268:26-35. DOI 10.1006/excr.2001.5251.

3. Baiao P.C., Schreiber E., Parker P.G. The genetic basis of the plumage polymorphism in red-footed boobies (Sula sula): a melanocortin-1 receptor (MC1R) analysis. J. Hered. 2007;98(4):287-292. DOI 10.1093/jhered/esm030.

4. Bed’hom B., Vaez M., Coville J.L., Gourichon D., Chastel O., Follett S., Burke T., Minvielle F. 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, feed consumption and body temperature. BMC Genomics. 2012;13:442. DOI 10.1186/ 1471-2164-13-442.

5. Blum A., Hartmann K., Rutten A. Braunliche Verfarbung der linken Brustwarze bei einer 60-jahrigen Patientin. Der Hautarzt. 2010;61: 64-68. DOI 10.1007/s00105-009-1885-z.

6. Brelsford A., Toews D.P.L., Irwin D.E. Admixture mapping in a hybrid zone reveals loci associated with avian feather coloration. Proc. Biol. Sci. 2017;284:1866. DOI 10.1098/rspb.2017.1106.

7. Campo J.L. Use of the sex-linked barring (B) gene for chick sexing on an eumelanotic Columbian background. Poult. Sci. 1991;70(7):1469- 1473. DOI 10.3382/ps.0701469.

8. Chang C.M., Coville J.L., Coquerelle G., Gourichon D., OulmoudenA., Tixier-Boichard M. Complete association between a retroviral insertion in the tyrosinase gene and the recessive white mutation in chickens. BMC Genomics. 2006;5:7-19. DOI 10.1186/1471-2164-7-19.

9. Cooke T.F., Fischer C.R., Wu P., Jiang T.X., Xie K.T., Kuo J., Doctorov E., Zehnder A., Khosla C., Chuong C.M., Bustamante C.D. Genetic mapping and biochemical basis of yellow feather pigmentation in budgerigars. Cell. 2017;171(2):427-439.e21. DOI 10.1016/j. cell.2017.08.016.

10. Cott H.B. Adaptive Coloration in Animals. London, UK: Methuen & Co., Ltd, Oxford Univ. Press, 1940. Crawford R.D. Poultry Breeding and Genetics. Ser.: Developments in Animal and Veterinary Sciences; 22, Elsevier, 1991. D’Alba L., Kieffer L., Shawkey M.D. Relative contributions of pigments and biophotonic nanostructures to natural color production: a case study in budgerigar (Melopsittacus. undulatus) feathers. J. Exp. Biol. 2012;215:1272-1277. DOI 10.1242/jeb.064907.

11. Davila S.G., Gil M.G., Resino-Talavan P., Campo J.L. Association between polymorphism in the melanocortin 1 receptor gene and E locus plumage color phenotype. Poult. Sci. 2014;93(5):1089-1096. DOI 10.3382/ps.2013-03611. Dorshorst B.J., Ashwell C.M. Genetic mapping of the sex-linked barring gene in the chicken. Poult. Sci. 2009;88(9):1811-1817. DOI 10.3382/ps.2009-00134.

12. Dorshorst B., Molin A.M., Rubin C.J., Johansson A.M., Strömstedt L., Pham M.H., Chen C.F., Hallböök F., Ashwell C., Andersson L. A complex genomic rearrangement involving the endothelin 3 locus causes dermal hyperpigmentation in the chicken. PLoS Genet. 2011;7(12):e1002412. DOI 10.1371/journal.pgen.1002412.

13. Doucet S.M., Shawkey M.D., Rathburn M.K., Mays H.L., Montgomerie R. Concordant evolution of plumage colour, feather microstructure and a melanocortin receptor gene between mainland and island populations of a fairy–wren. Proc. R. Soc. Lond. B, Biol. Sci. 2004; 271(1549):1663-1670. DOI 10.1098/rspb.2004.2779.

14. Dupin E., Le Douarin N.M. Development of melanocyte precursors from the vertebrate neural crest. Oncogene. 2003;22(20):3016-3023. Eising C.M., Muller W., Groothuis T.G.G. Avian mothers create different phenotypes by hormone deposition in their eggs. Biol. Lett. 2006;2:20-22. DOI 10.1098/rsbl.2005.0391.

15. Galeotti P., Rubolini D., Dunn P.O., Fasola M. Colour polymorphism in birds: causes and functions. J. Evol. Biol. 2003;16(4):635-646. DOI 10.1046/j.1420-9101.2003.00569.x.

16. Galvan I., Garcia-Campa J., Negro J.J. Complex plumage patterns can be produced only with the contribution of melanins. Physiol. Biochem. Zool. 2017;90(5):600-604. DOI 10.1086/693962. Galvan I., Solano F. Bird integumentary melanins: biosynthesis, forms, function and evolution. Int. J. Mol. Sci. 2016;17(4):520. DOI 10.3390/ijms17040520.

17. Garcia-Borron J.C., Sanchez-Laorden B.L., Jimenez-Cervantes C. Melanocortin-1 receptor structure and functional regulation. Pigment Cell Res. 2005;18(6):393-410. Gaudet P., Livstone M.S., Lewis S.E., Thomas P.D. Phylogenetic-based propagation of functional annotations within the Gene Ontology consortium. Rief. Bioinform. 2011;12(5):449-462. DOI 10.1093/ bib/bbr042.

18. Gudbjartsson D.F., Sulem P., Stacey S.N., Goldstein A.M., Rafnar T., Sigurgeirsson B., Benediktsdottir K.R., Thorisdottir K., Ragnarsson R., Sveinsdottir S.G. Magnusson V., Lindblom A., Kostulas K., Botella-Estrada R., Soriano V., Juberias P., Grasa M., Saez B., Andres R., Scherer D., Rudnai P., Gurzau E., Koppova K., Kiemeney L.A., Jakobsdottir M., Steinberg S., Helgason A., Gretarsdottir S., Tucker M.A., Mayordomo J.I., Nagore E., Kumar R., Hansson J., Olafsson J.H., Gulcher J., Kong A., Thorsteinsdottir U., Stefansson K. ASIP and TYR pigmentation variants associate with cutaneous melanoma and basal cell carcinoma. Nat. Genet. 2008;40: 886-891. DOI 10.1038/ng.161.

19. Guernsey M.W., Ritscher L., Miller M.A., Smith D.A., Schoneberg T., Shapiro M.D. AVal85Met mutation in melanocortin-1 receptor is associated with reductions in eumelanic pigmentation and cell surface expression in domestic rock pigeons (Columba livia). PLoS One. 2013;8(8):e74475. DOI 10.1371/journal.pone.0074475.

20. Gunnarsson U., Hellstrom A.R., Tixier-Boichard M., Minvielle F., Bed’hom B., Ito S., Jensen P., Rattink A., Vereijken A., Andersson L. Mutations in SLC45A2 cause plumage color variation in chicken and Japanese quail. Genetics. 2007;175(2):867-877. DOI 10.1534/ genetics.106.063107.

21. Gunnarsson U., Kerje S., Bed’hom B., Sahlqvist A.S., Ekwall O., Tixier-Boichard M., Kampe O., Andersson L. The Dark brown plumage color in chickens is caused by an 8.3-kb deletion upstream of SOX10. Pigment Cell Melanoma Res. 2011;24(2):268-274. DOI 10.1111/j.1755-148X.2011.00825.x.

22. Hellström A.R., Sundström E., Gunnarsson U., Bed’Hom B., TixierBoichard M., Honaker C.F., Sahlqvist A.S., Jensen P., Kämpe O., Siegel P.B., Kerje S., Andersson L. Sex-linked barring in chickens is controlled by the CDKN2A/B tumour suppressor locus. Pigment Cell Melanoma Res. 2010;23(4):521-530. DOI 10.1111/j.1755-148X. 2010.00700.x.

23. Hellström A.R., Watt B., Fard S.S., Tenza D., Mannström P., Narfström K., Ekesten B., Ito S., Wakamatsu K., Larsson J., Ulfendahl M., Kullander K., Raposo G., Kerje S., Hallböök F., Marks M.S., Andersson L. Inactivation of Pmel alters melanosome shape but has only a subtle effect on visible pigmentation. PLoS Genet. 2011;7(9):e1002285. DOI 10.1371/journal. pgen.1002285.

24. Hoque M.R., Jin S., Heo K.N., Kang B.S., Jo C., Lee J.H. Investigation of MC1R SNPs and their relationships with plumage colors in Korean native chicken. Asian Australas. J. Anim. Sci. 2013;26(5):625- 629. DOI 10.5713/ajas.2012.12581.

25. Ibarrola-Villava M., Hu H.-H., Guedj M., Fernandez L.P., Descamps V., Basset-Seguin N., Bagot M., Benssussan A., Saiag P., Fargnoli M.C., Peris K., Aviles J.A., Lluch A., Ribas G., Soufir N. MC1R, SLC45A2 and TYR genetic variants involved in melanoma susceptibility in Southern European populations: results from a meta-analysis. Eur. J. Cancer. 2012;48:2183-2191. DOI 10.1016/j.ejca.2012.03.006.

26. Ito S., Wakamatsu K., Ozeki H. Chemical analysis of melanins and its application to the study of the regulation of melanogenesis. Pigment Cell Res. 2000;13(8):103-109.

27. Jerome F.N. Autosex linkage in Barred Plymouth Rock. Poult Sci. 1939;(18):437-440. Johnson J.A., Ambers A.D., Burnham K.K. Genetics of plumage color in the Gyrfalcon (Falco rusticolus): analysis of the melanocortin 1 receptorgene. J. Hered. 2012;103:315-321. DOI 10.1093/jhered/ess023.

28. Karlsson A.C., Kerje S., Andersson L., Jensen P. Genotype at the PMEL17 locus affects social and explorative behaviour in chickens. Br. Poult. Sci. 2010;51(2):170-177. DOI 10.1080/00071661 003745802.

29. Keeling L., Andersson L., Schutz K.E., Kerje S., Fredriksson R., Carlborg O., Cornwallis C.K., Pizzari T., Jensen P. Chicken genomics: feather-pecking and victim pigmentation. Nature. 2004;431(7009): 645-646. DOI 10.1038/431645a.

30. Kerje S., Lind J., Schutz K., Jensen P., Andersson L. Melanocortin 1-receptor (MC1R) mutations are associated with plumage colour in chicken. Anim. Genet. 2003;34(4):241-248. DOI 10.1046/j.1365- 2052.2003.00991.x.

31. Kerje S., Sharma P., Gunnarsson U., Kim H., Bagchi S., Fredriksson R., Schutz K., Jensen P., von Heijne G., Okimoto R., Andersson L. The Dominant white, Dun and Smoky color variants in chicken are associated with insertion/deletion polymorphisms in the PMEL17 gene. Genetics. 2004;168(3):1507-1518. DOI 10.1534/genetics.104. 027995.

32. Kinoshita K., Akiyama T., Mizutani M., Shinomiya A., Ishikawa A., Younis H.H., Tsudzuki M., Namikawa T., Matsuda Y. Endothelin receptor B2 (EDNRB2) is responsible for the tyrosinase-independent recessive white (mow) and mottled (mo) plumage phenotypes in the chicken. PLoS One. 2014;9(1):e86361. DOI 10.1371/journal. pone.0086361.

33. Kogan S.M. Features of the Exterior and Interior of Chickens. Novosibirsk, 1979. (in Russian) Kuliawat R., Santambrogio L. A mutation within the transmembrane domain of melanosomal protein Silver (Pmel17) changes lumenal fragment interactions. Eur. J. Cell Biol. 2009;88(11):653-667. DOI 10.1016/j.ejcb.2009.07.001.

34. Land E.J., Riley P.A. Spontaneous redox reactions of dopaquinone and the balance between the eumelanic and phaeomelanic pathways. Pigment Cell Res. 2000;13:273-277.

35. Lerner A.B., Fitzpatrick T.B. Biochemistry of melanin formation. Physiol. Rev. 1950;30(1):91-126.

36. Ling M.K., Lagerstrom M.C., Fredriksson R., Okimoto R., Mundy N.I., Takeuchi S., Schioth H.B. Association of feather colour with constitutively active melanocortin 1 receptors in chicken. Eur. J. Biochem. 2003;270(7):1441-1449. DOI 10.1046/j.1432-1033.2003.03506.x.

37. Liu W.B., Chen S.R., Zheng J.X., Qu L.J., Xu G.Y., Yang N. Developmental phenotypic-genotypic associations of tyrosinase and melanocortin 1 receptor genes with changing profiles in chicken plumage pigmentation. Poult. Sci. 2010;89(6):1110-1114. DOI 10.3382/ps. 2010-00628.

38. Lopes R.J., Johnson J.D., Toomey M.B., Ferreira M.S., Araujo P.M., Melo-Ferreira J., Andersson L., Hill G.E., Corbo J.C., Carneiro M. Genetic basis for red coloration in birds. Curr. Biol. 2016;26(11): 1427-1434. DOI 10.1016/j.cub.2016.03.076.

39. Maia R., Rubenstein D.R., Shawkey M.D. Key ornamental innovations and diversification in an avian radiation. Proc. Natl. Acad. Sci. USA. 2013;110(26):10687-10692. DOI 10.1073/pnas.1220784110.

40. McGraw K.J., Safra R.J., Evans M.R., Wakamatsu K. European barn swallows use melanin pigments to color their feathers brown. Behav. Ecol. 2004;15:889-891.

41. Mitrofanova O.V., Dementeva N.V., Krutikova A.A., Yurchenko O.P., Vakhrameev A.B., Terletskiy V.P. Association of polymorphic variants in MSTN, PRL, and DRD2 genes with intensity of young animal growth in pushkin breed chickens. Cytol. Genet. 2017;51(3):179- 184. DOI 10.3103/S0095452717030082.

42. Moiseyeva I.G., Romanov M.N., NikiforovA.A., Avrutskaya T.B. Studies in chicken genetics. Commemorating the 120th anniversary of the outstanding soviet geneticist A.S. Serebrovsky (1892–1948). Russ. J. Genet. 2012;48(9):869-885. DOI 10.1134/S1022795412090074.

43. Mundy N.I. A window on the genetics of evolution: MC1R and plumage colouration in birds. Proc. R. Soc. Lond. B, Biol. Sci. 2005;272: 1633-1640. DOI 10.1098/rspb.2005.3107.

44. Natt D., Kerje S., Andersson L., Jensen P. Plumage color and feather pecking – behavioral differences associated with PMEL17 genotypes in chicken (Gallus gallus). Behav. Genet. 2007;37(2):399-407. DOI 10.1007/s10519-006-9125-0.

45. Negro J.J., Blasco R., Rosell J., Finlayson C. Potential exploitation of avian resources by fossil hominins: an overview from ethnographic and historical data. Quat. Int. 2016;421:6-11. DOI 10.1016/j.quaint. 2015.09.034.

46. Niwa T., Mochii M., Nakamura A., Shiojiri N. Plumage pigmentation and expression of its regulatory genes during quail development – histochemical analysis using Bh (black at hatch) mutants. Mech. Dev. 2002;118(1-2):139-146. DOI 10.1016/S0925-4773(02)00256-3.

47. Nordén K.K., Faber J., Babarović F., Stubbs T.L., Selly T., Schiffbauer J.D., Štefanić P.P., Mayr G., Smithwick F., Vinther J. Melanosome diversity and convergence in the evolution of iridescent avian feathers – implications for paleocolor reconstruction. Evolution. 2018;73-1:15-27. DOI 10.1111/evo.13641.

48. Oribe E., Fukao A., Yoshihara C., Mendori M., Rosal K.G., Takahashi S., Takeuchi S. Conserved distal promoter of the agouti signaling protein (ASIP) gene controls sexual dichromatism in chickens. Gen. Comp. Endocrinol. 2012;177(2):231-237. DOI 10.1016/j. ygcen.2012.04.016.

49. Ozeki H., Ito S., Wakamatsu K., Ishiguro I. Chemical characterization of pheomelanogenesis starting from dihydroxyphenylalanine or tyrosine and cysteine. Effects of tyrosinase and cysteine concentrations and reaction time. Biochim. Biophys. Acta. 1997;1336(3):539-548.

50. Park M.N., Choi J.A., Lee K.T., Lee H.J., Choi B.H., Kim H., Kim T.H., Cho S., Lee T. Genome-wide association study of chicken plumage pigmentation. Asian Australas. J. Anim. Sci. 2013;26(11):1523- 1528. DOI 10.5713/ajas.2013.13413.

51. Rees J.L., Birch-Machin M., Flanagan N., Healy E., Phillips S., Todd C. Genetic studies of the human melanocortin-1 receptor. Ann. N.Y. Acad. Sci. 1999;885:134-142. Roulin A. The evolution, maintenance and adaptive function of genetic colour polymorphism in birds. Biol. Rev. 2004;79:815-848. DOI 10.1017/S1464793104006487.

52. Roulin A., Ducrest A.L. Genetics of colouration in birds. Semin. Cell Dev. Biol. 2013;24(6-7):594-608. DOI 10.1016/j.semcdb.2013.05.005.

53. Rzepka Z., Buszman E., Beberok A., Wrześniok D. From tyrosine to melanin: signaling pathways and factors regulating melanogenesis. Postepy Hig. Med. Dosw. 2016;70:695-708. DOI 10.5604/ 17322693.1208033.

54. Sazanov A., Masabanda J., Ewald D., Takeuchi S., Tixier-Boichard M., Buitkamp J., Fries R. Evolutionarily conserved telomeric location of BBC1 and MC1R on a microchromosome questions the identity of MC1R and a pigmentation locus on chromosome 1 in chicken. Chromosome Res. 1998;6(8):651-654. DOI 10.1023/A:1009269830117.

55. Schiaffino M.V. Signaling pathways in melanosome biogenesis and pathology. Int. J. Biochem. Cell Biol. 2010;42(7):1094-1104. DOI 10.1016/j.biocel.2010.03.023.

56. Schiöth H.B. The physiological role of melanocortin receptors. Vitam. Horm. 2001;63:195-232. DOI 10.1016/S0083-6729(01)63007-3.

57. Schiöth H.B., Raudsepp T., Ringholm A., Fredriksson R., Takeuchi S., Larhammar D., Chowdhary B.P. Remarkable synteny conservation of melanocortin receptors in chicken, human, and other vertebrates. Genomics. 2003;81(5):504-509.

58. Schwochow-Thalmann D. Molecular Identification of Colour Pattern Genes in Birds. Uppsala: Swed. Univ. Agric. Sci., 2018;9:1652-6880.

59. Schwochow Thalmann D., Ring H., Sundstrom E., Cao X., Larsson M., Kerje S., Höglund A., Fogelholm J., Wright D., Jemth P., Hallböök F., Bed’Hom B., Dorshorst B., Tixier-Boichard M., Andersson L. The evolution of Sex-linked barring alleles in chickens involves both regulatory and coding changes in CDKN2A. PLoS Genet. 2017;13(4):1006665. DOI 10.1371/journal.pgen.1006665.

60. Serebrovsky A.S. Genetics of Domestic Chicken. Moscow, 1926. (in Russian) Shan X., Zhang Y., Peng W., Wang Z., Xie D. Molecular mechanism for jasmonate-induction of anthocyanin accumulation in Arabidopsis. J. Exp. Bot. 2009;60:3849-3860.

61. Smit N.P., Van der Meulen H., Koerten H.K., Kolb R.M., Mommaas A.M., Lentjes E.G., Pavel S. Melanogenesis in cultured melanocytes can be substantially influenced by L-tyrosine and L-cysteine. J. Invest Dermatol. 1997;109:796-800.

62. Smyth J.R. Genetics of Plumage, Skin and Eye Pigmentation in Chickens. In: Crawford R.D. (Ed.). Poultry Breeding and Genetics. Ser.: Developments in Animal and Veterinary Sciences; 22. Elsevier, 1990;109-168.

63. Somes R.G. The mottling gene, the basis of six plumage color patterns in the domestic fowl. Poult. Sci. 1980;59(7):1370-1374. DOI 10.3382/ps.0591370.

64. Somes R.G., Jr. International Registry of Poultry Genetic Stocks. Storrs Agric. Exp. Station. 29. 1988. https://opencommons.uconn.edu/saes/29

65. Strasser R., Schwabl H. Yolk testosterone organizes behavior and male plumage coloration in house sparrows (Passer domesticus). Behav. Ecol. Sociobiol. 2004;56:491-497. DOI 10.1007/s00265- 004-0810-9.

66. Takeuchi S., Suzuki H., Yabuuchi M. A possible involvement of melanocortin 1-receptor in regulating feather color pigmentation in the chicken. Biochim. Biophys. Acta.1996;14(1308(2)):164-168. DOI 10.1016/0167-4781(96)00100-5.

67. Takeuchi S., Takahashi S. Melanocortin receptor genes in the chickentissue distributions. Gen. Comp. Endocrinol. 1998;112(2):220-231.

68. Takeuchi S., Teshigawara K., Takahashi S. Widespread expression of Agouti-related protein (AGRP) in the chicken: a possible involvement of AGRP in regulating peripheral melanocortin systems in the chicken. Biochim. Biophys. Acta. 2000;1496:261-269.

69. Tang X.-F., Zhang Z., Hu D.-Y., Xu A.-E., Zhou H.-S., Sun L.-D., Gao M., Gao T.-W., Gao X.-H., Chen H.-D. Association analyses identify three susceptibility loci for vitiligo in the Chinese Han population. J. Invest. Dermatol. 2012;133:403-410.

70. Tobita-Teramoto T., Jang G.Y., Kino K., Salter D.W., Brumbaugh J., Akiyama T. Autosomal albino chicken mutation (c a/c a) deletes hexanucleotide (–ΔGACTGG817) at a copper-binding site of the tyrosinase gene. Poult. Sci. 2000;79(1):46-50.

71. Tsao H., Chin L., Garraway L.A., Fisher D.E. Melanoma: from mutations to medicine. Genes Dev. 2012;26:1131-1155.

72. Uy J.A., Moyle R.G., Filardi C.E., Cheviron Z.A. Difference in plumage color used in species recognition between incipient species is linked to a single amino acid substitution in the melanocortin 1 receptor. Am. Nat. 2009;174:244-254. DOI 10.1086/600084.

73. Vaez M., Follett S.A., Bed’hom B., Gourichon D., Tixier-Boichard M., Burke T. A single point-mutation within the melanophilin gene causes the lavender plumage colour dilution phenotype in the chicken. BMC Genet. 2008;9:7. DOI 10.1186/1471-2156-9-7.

74. Wolff G.L. Regulation of yellow pigment formation in mice: a historical perspective. Pigment Cell Res. 2003;16:2-15.

75. Xu J.G., Xie M.G., Zou S.Y., Liu X.F., Li X.H., Xie J.F., Zhang X.Q. Interactions of allele E of the MC1R gene with FM and mutations in the MLPH gene cause the five-gray phenotype in the Anyi tilelike gray chicken. Genet. Mol. Res. 2016;15(2). DOI 10.4238/gmr. 15027633.

76. Yang L., Du X., Wei S., Gu L., Li N., Gong Y., Li S. Genome-wide association analysis identifies potential regulatory genes for eumelanin pigmentation in chicken plumage. Anim. Genet. 2017;48(5):611- 614.

77. Yu S., Liao J., Tang M., Wang Y., Wei X., Mao L., Zeng C. A functional single nucleotide polymorphism in the tyrosinase gene promoter affects skin color and transcription activity in the black-boned chicken. Poult. Sci. 2017;96(11):4061-4067. DOI 10.3382/ps/pex217.

78. Yurchenko O.P., Vakhrameev A.B., Makarova A.V. Additive interactions of genes in the formation of plumage colors in chickens. Genetika i Razvedenie Zhivotnykh = Genetics and Animal Breeding. 2015;4:41-45. (in Russian)

79. Zhang Y., Gou W., Ma J., Zhang H., Zhang Y., Zhang H. Genome methylation and regulatory functions for hypoxic adaptation in Tibetan chicken embryos. Peer J. 2017;5:e3891. DOI 10.7717/peerj. 3891.

80. Zucca F.A., Basso E., Cupaioli F.A., Ferrari E., Sulzer D., Casella L., Zecca L. Neuromelanin of the human substantia nigra: an update. Neurotox Res. 2014;25(1):13-23. DOI 10.1007/s12640-013- 9435-y.


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