MC1R, eumelanin and pheomelanin: Their role in determining the susceptibility to skin cancer

Photochemistry and Photobiology

Volumn 91, Issue 1, 2015, Pages 188-200

  Nasti, Tahseen H ^a   Timares, Laura ^a,b  

^a UNIVERSITY OF ALABAMA AT BIRMINGHAM   (United States)

^b BIRMINGHAM VA MEDICAL CENTER   (United States)

Author keywords

[No Author keywords available]

Indexed keywords

EUMELANIN; MELANIN; MELANOCORTIN 1 RECEPTOR; PHEOMELANIN;

HUMAN; PATHOPHYSIOLOGY; PHYSIOLOGY; RADIATION INDUCED NEOPLASM; SKIN TUMOR;

HUMANS; MELANINS; NEOPLASMS, RADIATION-INDUCED; RECEPTOR, MELANOCORTIN, TYPE 1; SKIN NEOPLASMS;

EID: 84921504553     PISSN: 00318655     EISSN: 17511097     Source Type: Journal    
DOI: 10.1111/php.12335     Document Type: Article

References (133)

1. Briganti, S., and, M. Picardo, (2003) Antioxidant activity, lipid peroxidation and skin diseases. What's new J. Eur. Acad. Dermatol. Venereol. 17, 663-669.
2. Maresca, V., E. Flori, S. Briganti, E. Camera, M. Cario-Andre, A. Taieb, and, M. Picardo, (2006) UVA-induced modification of catalase charge properties in the epidermis is correlated with the skin phototype. J. Invest. Dermatol. 126, 182-190.
3. Brenner, M., and, V. J. Hearing, (2008) The protective role of melanin against UV damage in human skin. Photochem. Photobiol. 84, 539-549.
4. Costin, G. E., and, V. J. Hearing, (2007) Human skin pigmentation: melanocytes modulate skin color in response to stress. FASEB J. 21, 976-994.
5. Kadekaro, A. L., R. J. Kavanagh, K. Wakamatsu, S. Ito, M. A. Pipitone, and, Z. A. Abdel-Malek, (2003) Cutaneous photobiology. The melanocyte vs. the sun: who will win the final round? Pigment Cell Res. 16, 434-447.
6. Takeuchi, K., Y. Satou, H. Yamamoto, and, N. Satoh, (2005) A genome-wide survey of genes for enzymes involved in pigment synthesis in an ascidian, Ciona intestinalis. Zoolog. Sci. 22, 723-734.
7. Croucher, P. J., M. S. Brewer, C. J. Winchell, G. S. Oxford, and, R. G. Gillespie, (2013) De novo characterization of the gene-rich transcriptomes of two color-polymorphic spiders, Theridion grallator and T. californicum (Araneae: Theridiidae), with special reference to pigment genes. BMC Genomics 14, 862.
8. Scott, G., (2012) Demonstration of melanosome transfer by a shedding microvesicle mechanism. J. Invest. Dermatol. 132, 1073-1074.
9. Natarajan, V. T., P. Ganju, A. Ramkumar, R. Grover, and, R. S. Gokhale, (2014) Multifaceted pathways protect human skin from UV radiation. Nat. Chem. Biol. 10, 542-551.
10. Slominski, A., D. J. Tobin, S. Shibahara, and, J. Wortsman, (2004) Melanin pigmentation in mammalian skin and its hormonal regulation. Physiol. Rev. 84, 1155-1228.
11. Prota, G., (1992) The role of peroxidase in melanogenesis revisited. Pigment Cell Res. Suppl 2, 25-31.
12. Prota, G., (1980) Recent advances in the chemistry of melanogenesis in mammals. J. Invest. Dermatol. 75, 122-127.
13. Selinheimo, E., D. NiEidhin, C. Steffensen, J. Nielsen, A. Lomascolo, S. Halaouli, E. Record, D. O'Beirne, J. Buchert, and, K. Kruus, (2007) Comparison of the characteristics of fungal and plant tyrosinases. J. Biotechnol. 130, 471-480.
14. Bourquelot, E., and, A. Bertrand, (1895) Le bleuissement et le noircissement des champignons. Comp. Rend. Soc. Biol. 2, 582-584.
15. Lerner, A. B., T. B. Fitzpatrick, E. Calkins, and, W. H. Summerson, (1950) Mammalian tyrosinase; the relationship of copper to enzymatic activity. J. Biol. Chem. 187, 793-802.
16. Hearing, V. J., (2011) Determination of melanin synthetic pathways. J. Invest. Dermatol. 131, E8-E11.
17. Hearing, V. J. Jr, T. M. Ekel, P. M. Montague, and, J. M. Nicholson, (1980) Mammalin tyrosinase. Stoichiometry and measurement of reaction products. Biochim. Biophys. Acta 611, 251-268.
18. Lerner, A. B., T. B. Fitzpatrick, E. Calkins, and, W. H. Summerson, (1951) Mammalian tyrosinase; action on substances structurally related to tyrosine. J. Biol. Chem. 191, 799-806.
19. Lerch, K., (1983) Neurospora tyrosinase: structural, spectroscopic and catalytic properties. Mol. Cell. Biochem. 52, 125-138.
20. Raper, H. S., (1926) The tyrosinase-tyrosine reaction: production of l-3.4-dihydroxyphenylalanine from tyrosine. Biochem. J. 20, 735-742.
21. Mason, H. S., (1948) A classification of melanins. Ann. N. Y. Acad. Sci. 4, 399-404.
22. Riley, P. A., (1999) The great DOPA mystery: the source and significance of DOPA in phase I melanogenesis. Cell. Mol. Biol. 45, 951-960.
23. Orlow, S. J., B. K. Zhou, A. K. Chakraborty, M. Drucker, S. Pifko-Hirst, and, J. M. Pawelek, (1994) High-molecular-weight forms of tyrosinase and the tyrosinase-related proteins: evidence for a melanogenic complex. J. Invest. Dermatol. 103, 196-201.
24. Agrup, G., C. Hansson, H. Rorsman, A. M. Rosengren, and, E. Rosengren, (1979) Intracellular distribution of dopa and 5-S-cysteinyldopa in pigment cells with minimal pigment formation. Acta Derm. Venereol. 59, 355-356.
25. Ito, S., and, K. Fujita, (1985) Microanalysis of eumelanin and pheomelanin in hair and melanomas by chemical degradation and liquid chromatography. Anal. Biochem. 144, 527-536.
26. Thody, A. J., E. M. Higgins, K. Wakamatsu, S. Ito, S. A. Burchill, and, J. M. Marks, (1991) Pheomelanin as well as eumelanin is present in human epidermis. J. Invest. Dermatol. 97, 340-344.
27. Simon, J. D., D. Peles, K. Wakamatsu, and, S. Ito, (2009) Current challenges in understanding melanogenesis: bridging chemistry, biological control, morphology, and function. Pigment Cell Melanoma Res. 22, 563-579.
28. Ito, S., and, K. Wakamatsu, (2008) Chemistry of mixed melanogenesis-pivotal roles of dopaquinone. Photochem. Photobiol. 84, 582-592.
29. Inazu, M., and, Y. Mishima, (1993) Detection of eumelanogenic and pheomelanogenic melanosomes in the same normal human melanocyte. J. Invest. Dermatol. 100, 172S-175S.
30. Oyehaug, L., E. Plahte, D. I. Vage, and, S. W. Omholt, (2002) The regulatory basis of melanogenic switching. J. Theor. Biol. 215, 449-468.
31. Rees, J. L., and, R. M. Harding, (2012) Understanding the evolution of human pigmentation: recent contributions from population genetics. J. Invest. Dermatol. 132, 846-853.
32. Rees, J. L., (2011) The genetics of human pigmentary disorders. J. Invest. Dermatol. 131, E12-E13.
33. Ducrest, A. L., L. Keller, and, A. Roulin, (2008) Pleiotropy in the melanocortin system, coloration and behavioural syndromes. Trends Ecol. Evol. 23, 502-510.
34. Aberdam, E., C. Bertolotto, E. V. Sviderskaya, V. de Thillot, T. J. Hemesath, D. E. Fisher, D. C. Bennett, J. P. Ortonne, and, R. Ballotti, (1998) Involvement of microphthalmia in the inhibition of melanocyte lineage differentiation and of melanogenesis by agouti signal protein. J. Biol. Chem. 273, 19560-19565.
35. Vachtenheim, J., H. Novotna, and, G. Ghanem, (2001) Transcriptional repression of the microphthalmia gene in melanoma cells correlates with the unresponsiveness of target genes to ectopic microphthalmia-associated transcription factor. J. Invest. Dermatol. 117, 1505-1511.
36. Yasumoto, K., K. Yokoyama, K. Takahashi, Y. Tomita, and, S. Shibahara, (1997) Functional analysis of microphthalmia-associated transcription factor in pigment cell-specific transcription of the human tyrosinase family genes. J. Biol. Chem. 272, 503-509.
37. Slominski, A., A. Szczesniewski, and, J. Wortsman, (2000) Liquid chromatography-mass spectrometry detection of corticotropin-releasing hormone and proopiomelanocortin-derived peptides in human skin. J. Clin. Endocrinol. Metab. 85, 3582-3588.
38. Busca, R., and, R. Ballotti, (2000) Cyclic AMP a key messenger in the regulation of skin pigmentation. Pigment Cell Res. 13, 60-69.
39. Tachibana, M., (2000) MITF: a stream flowing for pigment cells. Pigment Cell Res. 13, 230-240.
40. Scott, G., S. Leopardi, S. Printup, N. Malhi, M. Seiberg, and, R. Lapoint, (2004) Proteinase-activated receptor-2 stimulates prostaglandin production in keratinocytes: analysis of prostaglandin receptors on human melanocytes and effects of PGE2 and PGF2alpha on melanocyte dendricity. J. Invest. Dermatol. 122, 1214-1224.
41. Starner, R. J., L. McClelland, Z. Abdel-Malek, A. Fricke, and, G. Scott, (2010) PGE(2) is a UVR-inducible autocrine factor for human melanocytes that stimulates tyrosinase activation. Exp. Dermatol. 19, 682-684.
42. Hara, M., M. Yaar, and, B. A. Gilchrest, (1995) Endothelin-1 of keratinocyte origin is a mediator of melanocyte dendricity. J. Invest. Dermatol. 105, 744-748.
43. Imokawa, G., Y. Yada, and, M. Kimura, (1996) Signalling mechanisms of endothelin-induced mitogenesis and melanogenesis in human melanocytes. Biochem. J. 314 (Pt 1), 305-312.
44. Swope, V. B., E. E. Medrano, D. Smalara, and, Z. A. Abdel-Malek, (1995) Long-term proliferation of human melanocytes is supported by the physiologic mitogens alpha-melanotropin, endothelin-1, and basic fibroblast growth factor. Exp. Cell Res. 217, 453-459.
45. Wu, C. S., C. C. Lan, M. H. Chiou, and, H. S. Yu, (2006) Basic fibroblast growth factor promotes melanocyte migration via increased expression of p125(FAK) on melanocytes. Acta Derm. Venereol. 86, 498-502.
46. Hachiya, A., A. Kobayashi, A. Ohuchi, Y. Takema, and, G. Imokawa, (2001) The paracrine role of stem cell factor/c-kit signaling in the activation of human melanocytes in ultraviolet-B-induced pigmentation. J. Invest. Dermatol. 116, 578-586.
47. Hirobe, T., R. Furuya, E. Hara, I. Horii, M. Tsunenaga, and, O. Ifuku, (2004) Granulocyte-macrophage colony-stimulating factor (GM-CSF) controls the proliferation and differentiation of mouse epidermal melanocytes from pigmented spots induced by ultraviolet radiation B. Pigment Cell Res. 17, 230-240.
48. Hirobe, T., M. Osawa, and, S. Nishikawa, (2004) Hepatocyte growth factor controls the proliferation of cultured epidermal melanoblasts and melanocytes from newborn mice. Pigment Cell Res. 17, 51-61.
49. Hirobe, T., (2002) Role of leukemia inhibitory factor in the regulation of the proliferation and differentiation of neonatal mouse epidermal melanocytes in culture. J. Cell. Physiol. 192, 315-326.
50. Hirobe, T., M. Osawa, and, S. Nishikawa, (2003) Steel factor controls the proliferation and differentiation of neonatal mouse epidermal melanocytes in culture. Pigment Cell Res. 16, 644-655.
51. Halder, R. M., and, K. M. Bang, (1988) Skin cancer in blacks in the United States. Dermatol. Clin. 6, 397-405.
52. Szabo, G., A. B. Gerald, M. A. Pathak, and, T. B. Fitzpatrick, (1969) Racial differences in the fate of melanosomes in human epidermis. Nature 222, 1081-1082.
53. van Nieuwpoort, F., N. P. Smit, R. Kolb, H. van der Meulen, H. Koerten, and, S. Pavel, (2004) Tyrosine-induced melanogenesis shows differences in morphologic and melanogenic preferences of melanosomes from light and dark skin types. J. Invest. Dermatol. 122, 1251-1255.
54. Pavel, S., F. van Nieuwpoort, H. van der Meulen, C. Out, K. Pizinger, P. Cetkovska, N. P. Smit, and, H. K. Koerten, (2004) Disturbed melanin synthesis and chronic oxidative stress in dysplastic naevi. Eur. J. Cancer 40, 1423-1430.
55. Mitra, D., X. Luo, A. Morgan, J. Wang, M. P. Hoang, J. Lo, C. R. Guerrero, J. K. Lennerz, M. C. Mihm, J. A. Wargo, K. C. Robinson, S. P. Devi, J. C. Vanover, J. A. D'Orazio, M. McMahon, M. W. Bosenberg, K. M. Haigis, D. A. Haber, Y. Wang, and, D. E. Fisher, (2012) An ultraviolet-radiation-independent pathway to melanoma carcinogenesis in the red hair/fair skin background. Nature 491, 449-453.
56. Greaves, M., (2014) Was skin cancer a selective force for black pigmentation in early hominin evolution? Proc. Biol. Sci. 281, 20132955.
57. Morgan, A. M., J. Lo, and, D. E. Fisher, (2013) How does pheomelanin synthesis contribute to melanomagenesis?: Two distinct mechanisms could explain the carcinogenicity of pheomelanin synthesis. BioEssays 35, 672-676.
58. Galvan, I., C. Alonso-Alvarez, and, J. J. Negro, (2012) Relationships between hair melanization, glutathione levels, and senescence in wild boars. Physiol. Biochem. Zool. 85, 332-347.
59. Galvan, I., and, A. P. Moller, (2011) Brain size and the expression of pheomelanin-based colour in birds. J. Evol. Biol. 24, 999-1006.
60. Ranadive, N. S., S. Shirwadkar, S. Persad, and, I. A. Menon, (1986) Effects of melanin-induced free radicals on the isolated rat peritoneal mast cells. J. Invest. Dermatol. 86, 303-307.
61. Ancans, J., D. J. Tobin, M. J. Hoogduijn, N. P. Smit, K. Wakamatsu, and, A. J. Thody, (2001) Melanosomal pH controls rate of melanogenesis, eumelanin/phaeomelanin ratio and melanosome maturation in melanocytes and melanoma cells. Exp. Cell Res. 268, 26-35.
62. Panzella, L., G. Szewczyk, M. d'Ischia, A. Napolitano, and, T. Sarna, (2010) Zinc-induced structural effects enhance oxygen consumption and superoxide generation in synthetic pheomelanins on UVA/visible light irradiation. Photochem. Photobiol. 86, 757-764.
63. Galvan, I., T. A. Mousseau, and, A. P. Moller, (2011) Bird population declines due to radiation exposure at Chernobyl are stronger in species with pheomelanin-based coloration. Oecologia 165, 827-835.
64. Barrenas, M. L., and, K. M. Holgers, (2000) Ototoxic interaction between noise and pheomelanin: distortion product otoacoustic emissions after acoustical trauma in chloroquine-treated red, black, and albino guinea pigs. Audiology 39, 238-246.
65. Srivastava, M. K., P. Sinha, V. K. Clements, P. Rodriguez, and, S. Ostrand-Rosenberg, (2010) Myeloid-derived suppressor cells inhibit T-cell activation by depleting cystine and cysteine. Cancer Res. 70, 68-77.
66. Kusmartsev, S., Y. Nefedova, D. Yoder, and, D. I. Gabrilovich, (2004) Antigen-specific inhibition of CD8+ T cell response by immature myeloid cells in cancer is mediated by reactive oxygen species. J. Immunol. 172, 989-999.
67. Gabrilovich, D. I., S. Ostrand-Rosenberg, and, V. Bronte, (2012) Coordinated regulation of myeloid cells by tumours. Nat. Rev. Immunol. 12, 253-268.
68. Zhang, Y., S. Choksi, K. Chen, Y. Pobezinskaya, I. Linnoila, and, Z. G. Liu, (2013) ROS play a critical role in the differentiation of alternatively activated macrophages and the occurrence of tumor-associated macrophages. Cell Res. 23, 898-914.
69. Saino, N., L. Canova, A. Costanzo, D. Rubolini, A. Roulin, and, A. P. Moller, (2013) Immune and stress responses covary with melanin-based coloration in the barn swallow. Evol. Biol. 40, 521-531.
70. Roulin, A., T. W. Jungi, H. Pfister, and, C. Dijkstra, (2000) Female barn owls (Tyto alba) advertise good genes. Proc. Biol. Sci. 267, 937-941.
71. Aranow, C., (2011) Vitamin D and the immune system. J. Investig. Med. 59, 881-886.
72. Liu, P. T., S. Stenger, H. Li, L. Wenzel, B. H. Tan, S. R. Krutzik, M. T. Ochoa, J. Schauber, K. Wu, C. Meinken, D. L. Kamen, M. Wagner, R. Bals, A. Steinmeyer, U. Zugel, R. L. Gallo, D. Eisenberg, M. Hewison, B. W. Hollis, J. S. Adams, B. R. Bloom, and, R. L. Modlin, (2006) Toll-like receptor triggering of a vitamin D-mediated human antimicrobial response. Science 311, 1770-1773.
73. Lemire, J. M., J. S. Adams, R. Sakai, and, S. C. Jordan, (1984) 1 alpha,25-dihydroxyvitamin D3 suppresses proliferation and immunoglobulin production by normal human peripheral blood mononuclear cells. J. Clin. Invest. 74, 657-661.
74. Chen, S., G. P. Sims, X. X. Chen, Y. Y. Gu, S. Chen, and, P. E. Lipsky, (2007) Modulatory effects of 1,25-dihydroxyvitamin D3 on human B cell differentiation. J. Immunol. 179, 1634-1647.
75. Bhalla, A. K., E. P. Amento, B. Serog, and, L. H. Glimcher, (1984) 1,25-Dihydroxyvitamin D3 inhibits antigen-induced T cell activation. J. Immunol. 133, 1748-1754.
76. Mattner, F., S. Smiroldo, F. Galbiati, M. Muller, P. Di Lucia, P. L. Poliani, G. Martino, P. Panina-Bordignon, and, L. Adorini, (2000) Inhibition of Th1 development and treatment of chronic-relapsing experimental allergic encephalomyelitis by a non-hypercalcemic analogue of 1,25-dihydroxyvitamin D(3). Eur. J. Immunol. 30, 498-508.
77. Boonstra, A., F. J. Barrat, C. Crain, V. L. Heath, H. F. Savelkoul, and, A. O'Garra, (2001) 1alpha,25-Dihydroxyvitamin d3 has a direct effect on naive CD4(+) T cells to enhance the development of Th2 cells. J. Immunol. 167, 4974-4980.
78. Gorman, S., L. A. Kuritzky, M. A. Judge, K. M. Dixon, J. P. McGlade, R. S. Mason, J. J. Finlay-Jones, and, P. H. Hart, (2007) Topically applied 1,25-dihydroxyvitamin D3 enhances the suppressive activity of CD4+CD25+ cells in the draining lymph nodes. J. Immunol. 179, 6273-6283.
79. Nasti, T. H., and, L. Timares, (2012) Inflammasome activation of IL-1 family mediators in response to cutaneous photodamage. Photochem. Photobiol. 88, 1111-1125.
80. Penna, G., and, L. Adorini, (2000) 1 Alpha, 25-dihydroxyvitamin D3 inhibits differentiation, maturation, activation, and survival of dendritic cells leading to impaired alloreactive T cell activation. J. Immunol. 164, 2405-2411.
81. Maaser, C., K. Kannengiesser, C. Specht, A. Lugering, T. Brzoska, T. A. Luger, W. Domschke, and, T. Kucharzik, (2006) Crucial role of the melanocortin receptor MC1R in experimental colitis. Gut 55, 1415-1422.
82. Gruis, N. A., and, R. van Doorn, (2012) Melanocortin 1 receptor function: shifting gears from determining skin and nevus phenotype to fetal growth. J. Invest. Dermatol. 132, 1953-1955.
83. Lerner, A. B., and, J. S. McGuire, (1961) Effect of alpha- and betamelanocyte stimulating hormones on the skin colour of man. Nature 189, 176-179.
84. Geschwind, I. I., (1966) Change in hair color in mice induced by injection of alpha-MSH. Endocrinology 79, 1165-1167.
85. Chhajlani, V., and, J. E. Wikberg, (1992) Molecular cloning and expression of the human melanocyte stimulating hormone receptor cDNA. FEBS Lett. 309, 417-420.
86. Mountjoy, K. G., L. S. Robbins, M. T. Mortrud, and, R. D. Cone, (1992) The cloning of a family of genes that encode the melanocortin receptors. Science 257, 1248-1251.
87. Gether, U., (2000) Uncovering molecular mechanisms involved in activation of G protein-coupled receptors. Endocr. Rev. 21, 90-113.
88. Subramaniam, S., and, R. Henderson, (1999) Electron crystallography of bacteriorhodopsin with millisecond time resolution. J. Struct. Biol. 128, 19-25.
89. Palczewski, K., T. Kumasaka, T. Hori, C. A. Behnke, H. Motoshima, B. A. Fox, I. Le Trong, D. C. Teller, T. Okada, R. E. Stenkamp, M. Yamamoto, and, M. Miyano, (2000) Crystal structure of rhodopsin: a G protein-coupled receptor. Science 289, 739-745.
90. Ringholm, A., J. Klovins, R. Rudzish, S. Phillips, J. L. Rees, and, H. B. Schioth, (2004) Pharmacological characterization of loss of function mutations of the human melanocortin 1 receptor that are associated with red hair. J. Invest. Dermatol. 123, 917-923.
91. Garcia-Borron, J. C., B. L. Sanchez-Laorden, and, C. Jimenez-Cervantes, (2005) Melanocortin-1 receptor structure and functional regulation. Pigment Cell Res. 18, 393-410.
92. Schaffer, J. V., and, J. L. Bolognia, (2001) The melanocortin-1 receptor: red hair and beyond. Arch. Dermatol. 137, 1477-1485.
93. Gerstenblith, M. R., A. M. Goldstein, M. C. Fargnoli, K. Peris, and, M. T. Landi, (2007) Comprehensive evaluation of allele frequency differences of MC1R variants across populations. Hum. Mutat. 28, 495-505.
94. Frandberg, P. A., M. Doufexis, S. Kapas, and, V. Chhajlani, (2001) Cysteine residues are involved in structure and function of melanocortin 1 receptor: substitution of a cysteine residue in transmembrane segment two converts an agonist to antagonist. Biochem. Biophys. Res. Commun. 281, 851-857.
95. Sanchez-Laorden, B. L., J. Sanchez-Mas, M. C. Turpin, J. C. Garcia-Borron, and, C. Jimenez-Cervantes, (2006) Variant amino acids in different domains of the human melanocortin 1 receptor impair cell surface expression. Cell. Mol. Biol. 52, 39-46.
96. Box, N. F., J. R. Wyeth, L. E. O'Gorman, N. G. Martin, and, R. A. Sturm, (1997) Characterization of melanocyte stimulating hormone receptor variant alleles in twins with red hair. Hum. Mol. Genet. 6, 1891-1897.
97. Smith, R., E. Healy, S. Siddiqui, N. Flanagan, P. M. Steijlen, I. Rosdahl, J. P. Jacques, S. Rogers, R. Turner, I. J. Jackson, M. A. Birch-Machin, and, J. L. Rees, (1998) Melanocortin 1 receptor variants in an Irish population. J. Invest. Dermatol. 111, 119-122.
98. Palmer, J. S., D. L. Duffy, N. F. Box, J. F. Aitken, L. E. O'Gorman, A. C. Green, N. K. Hayward, N. G. Martin, and, R. A. Sturm, (2000) Melanocortin-1 receptor polymorphisms and risk of melanoma: is the association explained solely by pigmentation phenotype? Am. J. Hum. Genet. 66, 176-186.
99. Bastiaens, M. T., J. A. ter Huurne, C. Kielich, N. A. Gruis, R. G. Westendorp, B. J. Vermeer, J. N. Bavinck, and T. Leiden Skin Cancer Study (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, 884-894.
100. Duffy, D. L., N. F. Box, W. Chen, J. S. Palmer, G. W. Montgomery, M. R. James, N. K. Hayward, N. G. Martin, and, R. A. Sturm, (2004) Interactive effects of MC1R and OCA2 on melanoma risk phenotypes. Hum. Mol. Genet. 13, 447-461.
101. Catania, A., N. Rajora, F. Capsoni, F. Minonzio, R. A. Star, and, J. M. Lipton, (1996) The neuropeptide alpha-MSH has specific receptors on neutrophils and reduces chemotaxis in vitro. Peptides 17, 675-679.
102. Loser, K., T. Brzoska, V. Oji, M. Auriemma, M. Voskort, V. Kupas, L. Klenner, C. Mensing, A. Hauschild, S. Beissert, and, T. A. Luger, (2010) The neuropeptide alpha-melanocyte-stimulating hormone is critically involved in the development of cytotoxic CD8+ T cells in mice and humans. PLoS One 5, e8958.
103. Andersen, G. N., M. Hägglund, O. Nagaeva, L. Frängsmyr, R. Petrovska, L. Mincheva-Nilsson, and, J. E. S. Wikberg, (2005) Quantitative measurement of the levels of melanocortin receptor subtype 1, 2, 3 and 5 and pro-opio-melanocortin peptide gene expression in subsets of human peripheral blood leucocytes. Scand. J. Immunol. 61, 279-284.
104. Catania, A., S. Gatti, G. Colombo, and, J. M. Lipton, (2004) Targeting melanocortin receptors as a novel strategy to control inflammation. Pharmacol. Rev. 56, 1-29.
105. Bhardwaj, R., E. Becher, K. Mahnke, M. Hartmeyer, T. Schwarz, T. Scholzen, and, T. A. Luger, (1997) Evidence for the differential expression of the functional alpha-melanocyte-stimulating hormone receptor MC-1 on human monocytes. J. Immunol. 158, 3378-3384.
106. Brzoska, T., D. H. Kalden, T. Scholzen, and, T. A. Luger, (1999) Molecular basis of the alpha-MSH/IL-1 antagonism. Ann. N. Y. Acad. Sci. 885, 230-238.
107. Ignar, D. M., J. L. Andrews, M. Jansen, M. M. Eilert, H. M. Pink, P. Lin, R. G. Sherrill, J. R. Szewczyk, and, J. G. Conway, (2003) Regulation of TNF-alpha secretion by a specific melanocortin-1 receptor peptide agonist. Peptides 24, 709-716.
108. Taherzadeh, S., S. Sharma, V. Chhajlani, I. Gantz, N. Rajora, M. T. Demitri, L. Kelly, H. Zhao, T. Ichiyama, A. Catania, and, J. M. Lipton, (1999) alpha-MSH and its receptors in regulation of tumor necrosis factor-alpha production by human monocyte/macrophages. Am. J. Physiol. 276, R1289-R1294.
109. Lipton, J. M., H. Zhao, T. Ichiyama, G. S. Barsh, and, A. Catania, (1999) Mechanisms of antiinflammatory action of alpha-MSH peptides. In vivo and in vitro evidence. Ann. N. Y. Acad. Sci. 885, 173-182.
110. Manna, S. K., and, B. B. Aggarwal, (1998) Alpha-melanocyte-stimulating hormone inhibits the nuclear transcription factor NF-kappa B activation induced by various inflammatory agents. J. Immunol. 161, 2873-2880.
111. Mandrika, I., R. Muceniece, and, J. E. Wikberg, (2001) Effects of melanocortin peptides on lipopolysaccharide/interferon-gamma-induced NF-kappaB DNA binding and nitric oxide production in macrophage-like RAW 264.7 cells: evidence for dual mechanisms of action. Biochem. Pharmacol. 61, 613-621.
112. Hassoun, H. T., L. Zou, F. A. Moore, R. A. Kozar, N. W. Weisbrodt, and, B. C. Kone, (2002) Alpha-melanocyte-stimulating hormone protects against mesenteric ischemia-reperfusion injury. Am. J. Physiol. Gastrointest. Liver Physiol. 282, G1059-G1068.
113. Ichiyama, T., T. Sakai, A. Catania, G. S. Barsh, S. Furukawa, and, J. M. Lipton, (1999) Inhibition of peripheral NF-kappaB activat-ion by central action of alpha-melanocyte-stimulating hormone. J. Neuroimmunol. 99, 211-217.
114. Robbins, L. S., J. H. Nadeau, K. R. Johnson, M. A. Kelly, L. Roselli-Rehfuss, E. Baack, K. G. Mountjoy, and, R. D. Cone, (1993) Pigmentation phenotypes of variant extension locus alleles result from point mutations that alter MSH receptor function. Cell 72, 827-834.
115. May, M. J., and, S. Ghosh, (1998) Signal transduction through NF-kappa B. Immunol. Today 19, 80-88.
116. Auriemma, M., T. Brzoska, L. Klenner, V. Kupas, T. Goerge, M. Voskort, Z. Zhao, T. Sparwasser, T. A. Luger, and, K. Loser, (2012) alpha-MSH-stimulated tolerogenic dendritic cells induce functional regulatory T cells and ameliorate ongoing skin inflammation. J. Invest. Dermatol. 132, 1814-1824.
117. Bhardwaj, R. S., A. Schwarz, E. Becher, K. Mahnke, Y. Aragane, T. Schwarz, and, T. A. Luger, (1996) Pro-opiomelanocortin-derived peptides induce IL-10 production in human monocytes. J. Immunol. 156, 2517-2521.
118. Murai, M., O. Turovskaya, G. Kim, R. Madan, C. L. Karp, H. Cheroutre, and, M. Kronenberg, (2009) Interleukin 10 acts on regulatory T cells to maintain expression of the transcription factor Foxp3 and suppressive function in mice with colitis. Nat. Immunol. 10, 1178-1184.
119. Fujii, S., K. Shimizu, T. Shimizu, and, M. T. Lotze, (2001) Interleukin-10 promotes the maintenance of antitumor CD8(+) T-cell effector function in situ. Blood 98, 2143-2151.
120. Yen, T. T., A. M. Gill, L. G. Frigeri, G. S. Barsh, and, G. L. Wolff, (1994) Obesity, diabetes, and neoplasia in yellow A(vy)/- mice: ectopic expression of the agouti gene. FASEB J. 8, 479-488.
121. Kanetsky, P. A., J. Swoyer, S. Panossian, R. Holmes, D. Guerry, and, T. R. Rebbeck, (2002) A polymorphism in the agouti signaling protein gene is associated with human pigmentation. Am. J. Hum. Genet. 70, 770-775.
122. Sulem, P., D. F. Gudbjartsson, S. N. Stacey, A. Helgason, T. Rafnar, M. Jakobsdottir, S. Steinberg, S. A. Gudjonsson, A. Palsson, G. Thorleifsson, S. Palsson, B. Sigurgeirsson, K. Thorisdottir, R. Ragnarsson, K. R. Benediktsdottir, K. K. Aben, S. H. Vermeulen, A. M. Goldstein, M. A. Tucker, L. A. Kiemeney, J. H. Olafsson, J. Gulcher, A. Kong, U. Thorsteinsdottir, and, K. Stefansson, (2008) Two newly identified genetic determinants of pigmentation in Europeans. Nat. Genet. 40, 835-837.
123. Gudbjartsson, D. F., P. Sulem, S. N. Stacey, A. M. Goldstein, T. Rafnar, B. Sigurgeirsson, K. R. Benediktsdottir, K. Thorisdottir, R. Ragnarsson, S. G. Sveinsdottir, V. Magnusson, A. Lindblom, K. Kostulas, R. Botella-Estrada, V. Soriano, P. Juberias, M. Grasa, B. Saez, R. Andres, D. Scherer, P. Rudnai, E. Gurzau, K. Koppova, L. A. Kiemeney, M. Jakobsdottir, S. Steinberg, A. Helgason, S. Gretarsdottir, M. A. Tucker, J. I. Mayordomo, E. Nagore, R. Kumar, J. Hansson, J. H. Olafsson, J. Gulcher, A. Kong, U. Thorsteinsdottir, and, K. Stefansson, (2008) ASIP and TYR pigmentation variants associate with cutaneous melanoma and basal cell carcinoma. Nat. Genet. 40, 886-891.
124. Lovett, M., Z. Y. Cheng, E. M. Lamela, T. Yokoi, and, C. J. Epstein, (1987) Molecular markers for the agouti coat color locus of the mouse. Genetics 115, 747-754.
125. Gunn, T. M., T. Inui, K. Kitada, S. Ito, K. Wakamatsu, L. He, D. M. Bouley, T. Serikawa, and, G. S. Barsh, (2001) Molecular and phenotypic analysis of Attractin mutant mice. Genetics 158, 1683-1695.
126. Ozeki, H., S. Ito, K. Wakamatsu, and, T. Hirobe, (1995) Chemical characterization of hair melanins in various coat-color mutants of mice. J. Invest. Dermatol. 105, 361-366.
127. Lamoreux, M. L., K. Wakamatsu, and, S. Ito, (2001) Interaction of major coat color gene functions in mice as studied by chemical analysis of eumelanin and pheomelanin. Pigment Cell Res. 14, 23-31.
128. Chintala, S., W. Li, M. L. Lamoreux, S. Ito, K. Wakamatsu, E. V. Sviderskaya, D. C. Bennett, Y. M. Park, W. A. Gahl, M. Huizing, R. A. Spritz, S. Ben, E. K. Novak, J. Tan, and, R. T. Swank, (2005) Slc7a11 gene controls production of pheomelanin pigment and proliferation of cultured cells. Proc. Natl Acad. Sci. USA 102, 10964-10969.
129. Jimbow, K., O. Ishida, S. Ito, Y. Hori, C. J. Witkop Jr, and, R. A. King, (1983) Combined chemical and electron microscopic studies of pheomelanosomes in human red hair. J. Invest. Dermatol. 81, 506-511.
130. Ito, S., and, K. Wakamatsu, (2003) Quantitative analysis of eumelanin and pheomelanin in humans, mice, and other animals: a comparative review. Pigment Cell Res. 16, 523-531.
131. Wakamatsu, K., and, S. Ito, (2002) Advanced chemical methods in melanin determination. Pigment Cell Res. 15, 174-183.
132. Strickland, F. M., H. K. Muller, L. C. Stephens, C. D. Bucana, C. K. Donawho, Y. Sun, and, R. P. Pelley, (2000) Induction of primary cutaneous melanomas in C3H mice by combined treatment with ultraviolet radiation, ethanol and aloe emodin. Photochem. Photobiol. 72, 407-414.
133. Elmets, C. A., N. Yusuf, S. Hamza, N. Iranikakh, J. Smith, A. L. Volk, H. Skelton, and, K. Smith, (2004) Topical application of dimethylbenz[a]anthracene results in the generation of multiple melanocytic nevi in C3H/HeN mice. Toxicol. Appl. Pharmacol. 195, 355-360.