Genetics of Hair and Skin Color

Annual Review of Genetics

Volumn 37, Issue , 2003, Pages 67-90

  Rees, Jonathan L ^a  

^a UNIVERSITY OF EDINBURGH   (United Kingdom)

Author keywords

Eumelanin; Melanin; Melanocortin 1 receptor (MC1R); Pheomelanin; Red hair

Indexed keywords

AGOUTI RELATED PROTEIN; ALPHA INTERMEDIN; COPPER CONTAINING ENZYME TYROSINASE; DOPA; MELANIN; MELANOCORTIN 1 RECEPTOR; MONOPHENOL MONOOXYGENASE; UNCLASSIFIED DRUG; VITAMIN D;

ALBINISM; ALLELE; CARCINOGENESIS; CORRELATION ANALYSIS; ERYTHEMA; GENE MUTATION; GENE SEQUENCE; GENETICS; GENOTYPE; HAIR COLOR; HAPLOTYPE; HIGH PERFORMANCE LIQUID CHROMATOGRAPHY; HUMAN; KERATINOCYTE; LIGHT SCATTERING; MELANOCYTE; MELANOGENESIS; NONHUMAN; OCULOCUTANEOUS ALBINISM; PHENOTYPE; PIGMENT DISORDER; PRIORITY JOURNAL; PROMOTER REGION; REVIEW; RICKETS; SKIN CANCER; SKIN COLOR; ULTRAVIOLET RADIATION;

ALPHA-MSH; GENOTYPE; HAIR COLOR; HUMANS; INTERCELLULAR SIGNALING PEPTIDES AND PROTEINS; MELANINS; MELANOCYTES; PHENOTYPE; RECEPTOR, MELANOCORTIN, TYPE 1; SKIN PIGMENTATION; STRUCTURE-ACTIVITY RELATIONSHIP; VARIATION (GENETICS);

AGOUTI;

EID: 0346786217     PISSN: 00664197     EISSN: None     Source Type: Book Series    
DOI: 10.1146/annurev.genet.37.110801.143233     Document Type: Review

References (126)

1. Abdel-Malek ZA, Scott MC, Furumura M, Lamoreux ML, Ollmann M, et al. 2001. The melanocortin 1 receptor is the principal mediator of the effects of agouti signaling protein on mammalian melanocytes. J. Cell Sci. 114:1019-24
2. Alaluf S, Atkins D, Barrett K, Blount M, Carter N, Heath A. 2002. Ethnic variation in melanin content and composition in photoexposed and photoprotected human skin. Pigment Cell Res. 15:112-18
3. Bagnara JT. 1998. Comparative anatomy and physiology of pigment cells in non-mammalian tissues. See Ref. 84, 2:9-40
4. Barsh G. 1999. From Agouti to Pomc - 100 years of fat blonde mice. Nat. Med. 5:984-85
5. Barsh GS. 1996. The genetics of pigmentation: from fancy genes to complex traits. Trends Genet. 12:299-305
6. Bastiaens M, ter Huurne J, Gruis N, Bergman W, Westendorp R, et al. 2001. The melanocortin-1-receptor gene is the major freckle gene. Hum. Mol. Genet. 10: 1701-8
7. Bastiaens MT, Huurne JA, Kielich C, Gruis NA, Westendorp RG, et al. 2001. Melanocortin-1 receptor gene variants determine the risk of nonmelanoma skin cancer independently of fair skin and red hair. Am. J. Hum. Genet. 68:884-94
8. Bhardwaj RS, Luger TA. 1994. Proopiomelanocortin production by epidermal cells: evidence for an immune neuroendocrine network in the epidermis. Arch. Dermatol. Res. 287:85-90
9. Bodmer WF, Cavalli-Sforza LL. 1976. Racial differentiation. In Genetics, Evolution, and Man. 19:559-604. San Francisco: Freeman
10. Box NF, Duffy DL, Irving R, Russell A, Chen W, et al. 2001. Melanocortin 1 receptor genotype is a risk factor for basal and squamous cell carcinoma. J. Invest. Dermatol. 116:224-29
11. Box NF, Wyeth JR, O'Gorman LE, Martin NG, Sturm RA. 1997. Characterization of melanocyte stimulating hormone receptor variant alleles in twins with red hair. Hum. Mol. Genet. 6:1891-97
12. Bruls WA, Slaper H, van der Leun JC, Berrens L. 1984. Transmission of human epidermis and stratum corneum as a function of thickness in the ultraviolet and visible wavelengths. Photochem. Photobiol. 40:485-94
13. Bruls WA, van Weelden H, van der Leun JC. 1984. Transmission of UV-radiation through human epidermal layers as a factor influencing the minimal erythema dose. Photochem. Photobiol. 39:63-67
14. Byard PJ, Lees FC. 1981. Estimating the number of loci determining skin colour in a hybrid population. Ann. Hum. Biol. 8: 49-58
15. Chardon A, Cretois I, Hourseau C. 1991. Skin colour typology and suntanning pathways. Int. J. Cosmet. Sci. 13:191-208
16. Chen WB, Kelly MA, Opitz-Araya X, Thomas RE, Low MJ, Cone RD. 1997. Exocrine gland dysfunction in MC5-R-deficient mice: evidence for coordinated regulation of exocrine gland function by melanocortin peptides. Cell 91:789-98
17. Chhajlani V, Muceniece R, Wikberg JE. 1993. Molecular cloning of a novel human melanocortin receptor. Biochem. Biophys. Res. Commun. 195:866-73. Erratum. 1996. Biochem. Biophys. Res. Commun. 218 (2):638
18. Chuang TY, Reizner GT, Elpern DJ, Stone JL, Farmer ER. 1995. Nonmelanoma skin cancer in Japanese ethnic Hawaiians in Kauai, Hawaii: an incidence report. J. Am. Acad. Dermatol. 33:422-26
19. Cohn B. 2002. Sunlight, skin color, and folic acid. J. Am. Acad. Dermatol. 46: 317-18
20. Cone RD, Lu D, Koppula S, Vage DI, Klungland H, et al. 1996. The melanocortin receptors: agonists, antagonists, and the hormonal control of pigmentation. Rec. Prog. Horm. Res. 51:287-317
21. Cone RD, Mountjoy KG, Robbins LS, Nadeau JH, Johnson KR, et al. 1993. Cloning and functional characterization of a family of receptors for the melanotropic peptides. Ann. NY Acad. Sci. 680: 342-63
22. Davenport CC, Davenport CB. 1909. Heredity of hair color in man. Am. Nat. 43: 193-211
23. De Luca M, Siegrist W, Bondanza S, Mathor M, Cancedda R, Eberle AN. 1993. Alpha melanocyte stimulating hormone (alpha MSH) stimulates normal human melanocyte growth by binding to high-affinity receptors. J. Cell Sci. 105:1079-84
24. Diamond JM. 1994. Race without color. Discovery. Nov. 83-89
25. Diffey BL. 1999. Human exposure to ultraviolet radiation. See Ref. 39a, 2:5-24
26. Diffey BL, Elwood JM. 1994. Tables of ambient solar ultraviolet radiation for use in epidemiological studies of malignant melanoma and other diseases. In Epidemiological Aspects of Cutaneous Malignant Melanoma, ed. RP Gallagher, JM Elwood. 19:81-105. Boston: Kluwer
27. Diffey BL, Jansen CT, Urbach F, Wulf HC. 1997. The standard erythema dose: a new photobiological concept. Photodermatol. Photoimmunol. Photomed. 13:64-66
28. Diffey BL, Oliver RJ, Farr PM. 1984. A portable instrument for quantifying erythema induced by ultraviolet radiation. Br. J. Dermatol. 111:663-72
29. Farr PM, Diffey BL. 1986. The vascular response of human skin to ultraviolet radiation. Photochem. Photobiol. 44:501-7
30. Fitzpatrick TB. 1988. The validity and practicality of sun-reactive skin types I through VI. Arch. Dermatol. 124:869-71
31. Flanagan N, Healy E, Ray A, Philips S, Todd C, et al. 2000. Pleiotropic effects of the melanocortin 1 receptor (MC1R) gene on human pigmentation. Hum. Mol. Genet. 9:2531-37
32. Flanagan N, Ray AJ, Todd C, Birch-Machin MA, Rees JL. 2001. The relation between melanocortin 1 receptor genotype and experimentally assessed ultraviolet radiation sensitivity. J. Invest. Dermatol. 117:1314-17
33. Frändberg PA, Doufexis M, Kapas S, Chhajlani V. 1998. Human pigmentation phenotype: A point mutation generates nonfunctional MSH receptor. Biochem. Biophys. Res. Commun. 245:490-92
34. Friedmann PS, Wren F, Buffey J, MacNeil S. 1990. Alpha-MSH causes a small rise in cAMP but has no effect on basal or ultraviolet-stimulated melanogenesis in human melanocytes. Br. J. Dermatol. 123:145-51
35. Gantz I, Yamada T, Tashiro T, Konda Y, Shimoto Y, et al. 1994. Mapping of the gene encoding the melanocortin-1 (alpha-melanocyte stimulating hormone) receptor (MC1R) to human chromosome 16q24.3 by fluorescence in situ hybridization. Genomics 19:394-95
36. Gilchrest BA, Blog FB, Szabo G. 1979. Effects of ageing and chronic sun exposure on melanocytes in human skin. J. Invest. Dermatol. 73:141-43
37. Harding RM, Healy E, Ray AJ, Ellis NS, Flanagan N, et al. 2000. Evidence for variable selective pressures at the human pigmentation locus, MC1R. Am. J. Hum. Genet. 66:1351-61
38. Harrison GA. 1973. Differences in human pigmentation: measurement, geographic variation, and causes. J. Invest. Dermatol. 60:418-26
39. Hatta N, Dixon C, Ray AJ, Phillips SR, Cunliffe WJ, et al. 2001. Expression, candidate gene, and population studies of the melanocortin 5 receptor. J. Invest. Dermatol. 116:564-70
40. Hawk JLM, ed. 1999.Photodermatology. London: Arnold
41. Healy E, Birch-Machin MA, Rees JL. 2000. The human melanocortin-1 receptor. In The Human Melanocortin-1 Receptor, ed. RD Cone. 11:341-60. Totawa, NJ: Humana
42. Healy E, Flannagan N, Ray A, Todd C, Jackson U, et al. 2000. Melanocortin-1-receptor gene and sun sensitivity in individuals without red hair. Lancet 355:1072-73
43. Healy E, Jordan SA, Budd PS, Suffolk R, Rees JL, Jackson LT. 2001. Functional variation of MClRalleles from red-haired individuals. Hum. Mol. Genet. 10:2397-402
44. Healy E, Todd C, Jackson IJ, Birch-Machin M, Rees JL. 1999. Skin type, melanoma, and melanocortin 1 receptor variants. J. Invest. Dermatol. 112:512-13
45. Hedley SJ, Gawkrodger DJ, Weetman AP, Morandini R, Boeynaems JM, et al. 1998. α-melanocyte stimulating hormone inhibits tumour necrosis factor - a stimulated intercellular adhesion molecule-1 expression in normal cutaneous human melanocytes and in melanoma cell lines. Br. J. Dermatol. 138:536-43
46. Hill HZ. 1992. The function of melanin or six blind people examine an elephant. BioEssays 14(1):49-56
47. Hoffmann K, Kaspar K, Altmeyer P, Gambichler T. 2000. UV transmission measurements of small skin specimens with special quartz cuvettes. Dermatology 201:307-11
48. Holick MF. 2001. Sunlight "D" ilemma: risk of skin cancer or bone disease and muscle weakness. Lancet 357:4-6
49. Hunt G, Kyne S, Ito S, Wakamatsu K, Todd C, Thody A. 1995. Eumelanin and pheomelanin contents of human epidermis and cultured melanocytes. Pigment Cell Res. 8:202-8
50. Hunt G, Todd C, Cresswell JE, Thody AJ. 1994. Alpha-melanocyte stimulating hormone and its analogue Nle4DPhe7 alpha-MSH affect morphology, tyrosinase activity and melanogenesis in cultured human melanocytes. J. Cell Sci. 107(Pt. 1):205-11
51. Ichii-Jones F, Lear JT, Heagerty AHM, Smith AG, Hutchinson PE, et al. 1998. Susceptibility to melanoma: influence of skin type and polymorphism in the melanocyte stimulating hormone receptor gene. J. Invest. Dermatol. 111:218-21
52. Ito S. 1998. Advances in chemical analysis of melanins. See Ref. 84, pp. 439-50
53. Ito S, Ozeki H, Wakamatsu K. 2000. Spectrophotometric and HPLC characterization of hair melanins. In Melanogenesis and Malignant Melanoma: Biochemistry, Cell Biology, Molecular Biology, Pathophysiology, Diagnosis and Treatment, ed. K Hori, VJ Hearing, J Nakayama, pp. 63-72. Oxford: Oxford Univ. Press
54. Jablonski NG, Chaplin G. 2000. The evolution of human skin coloration. J. Hum. Evol. 39:57-106
55. Jackson IJ. 1991. Mouse coat color mutations: a molecular genetic resource which spans the centuries. BioEssays 13: 439-46
56. Jackson IJ. 1994. Molecular and developmental genetics of mouse coat color. Annu. Rev. Genet. 28:189-217
57. Jackson IJ. 1997. Homologous pigmentation mutations in human, mouse and other model organisms. Hum. Mol. Genet. 6:1613-24
58. Jimenez-Cervantes C, Olivares C, Gonzalez P, Morandini R, Ghanem G, Carlos Garcia-Borron J. 2001. The pro162 variant is a loss-of-function mutation of the human melanocortin 1 receptor gene. J. Invest. Dermatol. 117:156-58
59. Joerg H, Fries HR, Meijerink E, Stranzinger GF. 1996. Red coat color in Holstein cattle is associated with a deletion in the MSHR gene. Mamm. Genome 7:317-18
60. Jones FT, Ramachandran S, Lear J, Smith A, Bowers B, et al. 1999. The melanocyte stimulating hormone receptor polymorphism: association of the V92M and A294H alleles with basal cell carcinoma. Clin. Chim. Acta 282:125-34
61. Kanetsky PA, Swoyer J, Panossian S, Holmes R, Guerry D, Rebbeck TR. 2002. A polymorphism in the agouti signaling protein gene is associated with human pigmentation. Am. J. Hum. Genet. 70:770-75
62. Kennedy C, ter Huurne J, Berkhout M, Gruis N, Bastiaens M, et al. 2001. Melanocortin 1 receptor (MC1R) gene variants are associated with an increased risk for cutaneous melanoma which is largely independent of skin type and hair color. J. Invest. Dermatol. 117:294-300
63. Kijas JMH, Wales R, Törnsten A, Chardon P, Moller M, Andersson L. 1998. Melanocortin receptor 1 (MC1R) mutations and coat color in pigs. Genetics 150:1177-85
64. King RA, Hearing VJ, Creel DJ, Oetting WS. 2001. Albinism. In The Metabolic and Molecular Bases of Inherited Disease, ed. CR Scriver, AL Beaudet, WS Sly, D Valle, B Childs, KW Kinzler, B Vogelstein. 200:5587-28. New York: McGraw Hill. 8th ed.
65. Klungland H, Vage DI, Gomez-Raya L, Adalsteinsson S, Lien S. 1995. The role of melanocyte-stimulating hormone (MSH) receptor in bovine coat color determination. Mamm. Genome 6:636-39
66. Kollias N, Sayre RM, Zeise L, Chedekel MR. 1991. Photoprotection by melanin. J. Photochem. Photobiol. B 9:135-60
67. Kollias N, Stamatas GN. 2002. Optical non-invasive approaches to diagnosis of skin diseases. J. Investig. Dermatol. Symp. Proc. 7:64-75
68. Koppula SV, Robbins LS, Lu D, Baack E, White CRJ, et al. 1997. Identification of common polymorphisms in the coding sequence of the human MSH receptor (MCIR) with possible biological effects. Hum. Mutat. 9:30-36
69. Krude H, Biebermann H, Luck W, Horn R, Brabant G, Grüters A. 1998. Severe early-onset obesity, adrenal insufficiency and red hair pigmentation caused by POMC mutations in humans. Nat. Genet. 19:155-57
70. Lerner AB. 1993. The discovery of the melanotropins. Ann. NY Acad. Sci. 680:1-12
71. Little MA, Wolff ME. 1981. Skin and hair reflectance in women with red hair. Ann. Hum. Biol. 8:231-41
72. Lock-Andersen J, Therkildsen P, de Fine O, Gniadecka M, Dahlstrom K, et al. 1997. Epidermal thickness, skin pigmentation and constitutive photosensitivity. Photodermatol. Photoimmunol. Photomed. 13:153-58
73. Lock-Andersen J, Wulf HC. 1997. Seasonal variation of skin pigmentation. Acta Derm. Venereol. 77:219-21
74. Lookingbill DP, Lookingbill GL, Leppard B. 1995. Actinic damage and skin cancer in albinos in northern Tanzania: findings in 164 patients enrolled in an outreach skin care program. J. Am. Acad. Dermatol. 32:653-58
75. Luger TA, Bhardwaj RS, Grabbe S, Schwarz T. 1996. Regulation of the immune response by epidermal cytokines and neurohormones. J. Dermatol. Sci. 13:5-10
76. Makova KD, Ramsay M, Jenkins T, Li WH. 2001. Human DNA sequence variation in a 6.6-kb region containing the melanocortin 1 receptor promoter. Genetics 158:1253-68
77. Manga P, Kromberg JGR, Box NF, Sturm RA, Jenkins T, Ramsay M. 1997. Rufous oculocutaneous albinism in southern African Blacks is caused by mutations in the TYRP1 gene. Am. J. Hum. Genet. 61: 1095-101
78. Marklund L, Moller MJ, Sandberg K, Andersson L. 1996. A missense mutation in the gene for melanocyte-stimulating hormone receptor (MC1R) is associated with the chestnut coat color in horses. Mamm. Genome 7:895-99
79. McKenzie CA, Harding RM, Tomlinson JB, Ray AJ, Wakamatsu K, Rees JL. 2003. Phenotypic expression of melanocortin-1 receptor (MC1R) mutations in Black Jamaicans. J. Invest. Dermatol. In press
80. Miller MW, Duhl DM, Vrieling H, Cordes SP, Ollmann MM, et al. 1993. Cloning of the mouse agouti gene predicts a secreted protein ubiquitously expressed in mice carrying the lethal yellow mutation. Genes Dev. 7:454-67
81. Montagna W, Prota G, Kenney JA. 1993. Black Skin: Structure and Function. London: Academic
82. Morandini R, Boeynaems JM, Hedley SJ, MacNeil S, Ghanem G. 1998. Modulation of ICAM-1 expression by a-MSH in human melanoma cells and melanocytes. J. Cell. Physiol. 175:276-82
83. Mountjoy KG, Robbins LS, Mortrud MT, Cone RD. 1992. The cloning of a family of genes that encode the melanocortin receptors. Science 257:1248-51
84. Newton JM, Wilkie AL, He L, Jordan SA, Metallinos DL, et al. 2000. Melanocortin 1 receptor variation in the domestic dog. Mamm. Genome 11:24-30
85. Nordlund JJ, Boissy RE, Hearing VJ, King RA, Ortonne JP, eds. 1998. The Pigmentary System: Physiology and Pathophysiology. New York: Oxford Univ. Press
86. Ozeki H, Ito S, Wakamatsu K, Thody AJ. 1996. Spectrophotometric characterization of eumelanin and pheomelanin in hair. Pigment Cell Res. 9:265-70
87. Palmer JS, Duffy DL, Box NF, Aitken JF, O'Gorman LE, et al. 2000. Melanocortin-1 receptor polymorphisms and risk of melanoma: Is the association explained solely by pigmentation phenotype? Am. J. Hum. Genet. 66:176-86
88. Price JS, Burton JL, Shuster S, Wolff K. 1976. Control of scrotal colour in the vervet monkey. J. Med. Primatol. 5:296-304
89. Prota G. 1992. Melanins and Melanogenesis. San Diego: Academic
90. Prota G, d'Ischia M, Napolitano A. 1998. The chemistry of melanins and related metabolites. See Ref. 84, pp. 307-32
91. Quevedo WC, Holstein TJ. 1998. General biology of mammalian pigmentation. See Ref. 84, pp. 43-58
92. Rampen FH, Fleuren BA, de Boo TM, Lemmens WA. 1988. Unreliability of self-reported burning tendency and tanning ability. Arch. Dermatol. 124:885-88
93. Rana BK, Hewett-Emmett D, Jin L, Chang BHJ, Sambuughin N, et al. 1999. High polymorphism at the human melanocortin 1 receptor locus. Genetics 151:1547-57
94. Rees JL. 2000. The melanocortin 1 receptor (MC1R): more than just red hair. Pigment Cell Res. 13:135-40
95. Rees JL. 2002. Molecular phototypes. In Mechanisms of Suntanning, ed. Jean-Paul Ortonne, Robert Ballotti. 31:333-339. London: Martin Dunitz
96. Robbins LS, Nadeau JH, Johnson KR, Kelly MA, Roselli-Rehfuss L, et al. 1993. Pigmentation phenotypes of variant extension locus alleles result from point mutations that alter MSH receptor function. Cell 72:827-34
97. Roberts DF, Kromberg JG, Jenkins T. 1986. Differentiation of heterozygotes in recessive albinism. J. Med. Genet. 23: 323-27
98. Robins AH. 1991. The evolution of skin colour. In Biological Perspectives on Human Pigmentation, pp. 187-212. Cambridge: Cambridge Univ. Press
99. Sarna T, Swartz HM. 1998. The physical properties of melanins. See Ref. 84, 25:333-57
100. Schioth HB, Muceniece R, Larsson M, Mutulis F, Szardenings M, et al. 1997. Binding of cyclic and linear MSH core peptides to the melanocortin receptor subtypes. Eur. J. Pharmacol. 319:369-73
101. Schioth HB, Muceniece R, Wikberg JE. 1996. Characterisation of the melanocortin 4 receptor by radioligand binding. Pharmacol. Toxicol. 79:161-65
102. Schioth HB, Muceniece R, Wikberg JE, Chhajlani V. 1995. Characterisation of melanocortin receptor subtypes by radioligand binding analysis. Eur. J. Pharmacol. 288:311-17
103. Schioth HB, Phillips S, Rudzish R, Birch-Machin M, Wikberg J, Rees JL. 1999. Loss of function mutations of the human melanocortin 1 receptor are common and associated with red hair. Biochem. Biophys. Res. Commun. 260:488-91
104. Shriver MD, Parra EJ. 2000. Comparison of narrow-band reflectance spectroscopy and tristimulus colorimetry for measurements of skin and hair color in persons of different biological ancestry. Am. J. Phys. Anthropol. 112:17-27
105. Silvers WK. 1979. Coat Colors of Mice. New York: Springer-Verlag
106. Smith R, Healy E, Siddiqui S, Flanagan N, Steijlen PM, et al. 1998. Melanocortin 1 receptor variants in an Irish population. J. Invest. Dermatol. 111:119-22
107. Sturm RA, Box NF, Ramsay M. 1998. Human pigmentation genetics: the difference is only skin deep. BioEssays 20:712-21
108. Sturm RA, Teasdale RD, Box NF. 2001. Human pigmentation genes: identification, structure and consequences of polymorphic variation. Gene 277:49-62
109. Suzuki I, Cone RD, Im S, Nordlund J, Abdel-Malek ZA. 1996. Binding of melanotropic hormones to the melanocortin receptor MC1R on human melanocytes stimulates proliferation and melanogenesis. Endocrinology 137:1627-33
110. Takeuchi S, Suzuki H, Yabuuchi M, Takahashi S. 1996. A possible involvement of melanocortin 1-receptor in regulating feather color pigmentation in the chicken. Biochim. Biophys. Acta 1308:164-68
111. Tan CP, McKee KK, Weinberg DH, MacNeil T, Palyha OC, et al. 1999. Molecular analysis of a new splice variant of the human melanocortin-1 receptor. FEBS Lett. 451:137-41
112. Theron E, Hawkins K, Bermingham E, Ricklefs RE, Mundy NI. 2001. The molecular basis of an avian plumage polymorphism in the wild: a melanocortin-1-receptor point mutation is perfectly associated with the melanic plumage morph of the bananaquit, Coereba flaveola. Curr. Biol. 11:550-57
113. Thody AJ, Graham A. 1998. Does α-MSH have a role in regulating skin pigmentation in humans. Pigment Cell Res. 11:265-74
114. Thody AJ, Higgins EM, Wakamatsu K, Ito S, Burchill SA, Marks JM. 1991. Pheomelanin as well as eumelanin is present in human epidermis. J. Invest. Dermatol. 97:340-44
115. Thody AJ, Ridley K, Penny RJ, Chalmers R, Fisher C, Shuster S. 1983. MSH peptides are present in mammalian skin. Peptides 4:813-16
116. Urbach F. 1999. The cumulative effects of ultraviolet radiation on the skin: photocarcinogenesis. See Ref. 39a, 7:89-111
117. Våge DI, Lu DS, Klungland H, Lien S, Adalsteinsson S, Cone RD. 1997. A non-epistatic interaction of agouti and extension in the fox, Vulpes vulpes. Nat. Genet. 15:311-15
118. Valverde P, Healy E, Jackson I, Rees JL, Thody AJ. 1995. Variants of the melanocyte-stimulating hormone receptor gene are associated with red hair and fair skin in humans. Nat. Genet. 11:328-30
119. Valverde P, Healy E, Sikkink S, Haldane F, Thody AJ, et al. 1996. The Asp84Glu variant of the melanocortin 1 receptor (MC1R) is associated with melanoma. Hum. Mol. Genet. 5:1663-66
120. Van den Berghe PL Frost P. 1986. Skin color preference, sexual dimorphism, and sexual selection: a case of gene-culture co-evolution. Ethn. Racial Stud. 9:87-113
121. Voisey J, Box NF, van Daal A. 2001. A polymorphism study of the human Agouti gene and its association with MC1R. Pigment Cell Res. 14:264-57
122. Vrieling H, Duhl DM, Millar SE, Miller KA, Barsh GS. 1994. Differences in dorsal and ventral pigmentation result from regional expression of the mouse agouti gene. Proc. Natl. Acad. Sci. USA 91:5667-71
123. Wagner JK, Jovel C, Norton HL, Parra EJ, Shriver MD. 2002. Comparing quantitative measures of erythema, pigmentation and skin response using reflectometry. Pigment Cell Res. 15:379-84
124. Wagner JK, Parra EJ, Norton L, Jovel C, Shriver MD. 2002. Skin responses to ultraviolet radiation: effects of constitutive pigmentation, sex, and ancestry. Pigment Cell Res. 15:385-90
125. Wakamatsu K, Ito S. 2002. Advanced chemical methods in melanin determination. Pigment Cell Res. 15:174-83
126. Wakamatsu K, Ito S, Rees JL. 2002. Usefulness of 4-amino-3- hydroxyphenylalanine as a specific marker of pheomelanin. Pigment Cell Res. 15:225-32