The Human Melanocortin-1 Receptor

  • Eugene Healy
  • Mark Birch-Machin
  • Jonathan L. Rees
Part of the The Receptors book series (REC)


One of the more obvious features that distinguishes one human from another are the pigmentatory characteristics (including skin type, hair and eye color) of the individual. Although it had been suspected (as a result of investigations into murine coat color) that several genes were likely to be involved in human pigmentation, and, although it had been known for some time that proopiomelanocortin (POMC) peptides such as alpha-melanocyte stimulating hormone (α-MSH) and adrenocorticotropic hormone (ACTH) can alter cutaneous pigmentation, it has only been during the past 10 years that molecular biologic/ genetic approaches have offered some insight into the complexities of human pigmentation (1–3). The detection of mutations within the genes responsible for type I and type II oculocutaneous albinism and piebaldism provided evidence for genotypic/phenotypic relationships in a subset of individuals with pigmentatory disorders, but did little to explain the wide variability in the pigmentatory characteristics of the vast majority of individuals (4–6).However, a basis for understanding “normal” human pigmentation became possible with the initial cloning of the human melanocortin 1 receptor (MC1R) gene by three separate groups who isolated this gene on the basis of its similarity to other G protein-coupled receptors (7,8) , and the subsequent identification of variant alleles within the murine homolog of this gene (mc1r) which could differentially activate adenylyl cyclase and which were associated with various coat colors in mice (9).


Melanoma Cell Skin Type Hair Color Human Melanocyte MC1R Gene 
These keywords were added by machine and not by the authors. This process is experimental and the keywords may be updated as the learning algorithm improves.


Unable to display preview. Download preview PDF.

Unable to display preview. Download preview PDF.


  1. 1.
    Nelson, D. H., Meakin, J. W., and Thorn, G. W. (1960) ACTH—producing pituitary tumors following adrenalectomy for Cushing’s syndrome. Ann. Intern. Med. 52, 561–569.Google Scholar
  2. 2.
    Lerner, A. B. and McGuire, J. S. (1961) Effect of alpha— and beta—melanocyte stimulating hormones on the skin colour of man. Nature 189, 176–179.PubMedCrossRefGoogle Scholar
  3. 3.
    Jackson, I. J. (1991) Mouse coat colour mutations: a molecular genetic resource which spans the centuries. BioEssays 13, 439–446.Google Scholar
  4. 4.
    Tomita, Y., Takeda, A., Okinaga, S., Tagami, H., and Shibahara, S. (1989) Human oculocutaneous albinism caused by single base insertion in the tyrosinase gene. Biochem. Biophys. Res. Commun. 164, 990–996.PubMedCrossRefGoogle Scholar
  5. 5.
    Giebel, L. B. and Spritz, R. A. (1991) Mutation of the KIT (mast/stem cell growth factor receptor) protooncogene in human piebaldism. Proc. Natl. Acad. Sci. U. S. A. 88, 8696–8699.PubMedCrossRefGoogle Scholar
  6. 6.
    Rinchik, E. M., Bultman, S. J., Horsthemke, B., Lee, S. —T., Strunk, K. M., Spritz, R. A., Avidano, K. M., Jong, M. T. C., and Nicholls, R. D. (1993) A gene for the mouse pink—eyed dilution locus and for human type II oculocutaneous albinism. Nature 361, 72–76.PubMedCrossRefGoogle Scholar
  7. 7.
    Chhajlani, V. and Wikberg, J. E. S. (1992) Molecular cloning and expression of the human melanocyte stimulating hormone receptor cDNA. FEBS Lett. 309, 417–420.PubMedCrossRefGoogle Scholar
  8. 8.
    Mountjoy, K. G., Robbins, L. S., Mortrud, M. T., and Cone, R.D. (1992) The cloning of a family of genes that encode the melanocortin receptors. Science 257, 1248–1251.PubMedCrossRefGoogle Scholar
  9. 9.
    Robbins, L. S., Nadeau, J. H., Johnson, K. R., Kelly, M. A., Roselli—Rehfuss, L., Baack, E., Mountjoy, K. G., and Cone, R. D. (1993) Pigmentation phenotypes of variant extension locus alleles result from point mutations that alter MSH receptor function. Cell 72, 827–834.Google Scholar
  10. 10.
    Gantz, I., Yamada, T., Tashiro, T., Konda, Y., Shimoto, Y., Miwa, H., and Trent, J. M. (1994) Mapping of the gene encoding the melanocortin-1 (a—melanocyte stimulating hormone) receptor (MC1R) to human chromosome 16q24.3 by fluorescence in situ hybridization. Genomics 19, 394–395.PubMedCrossRefGoogle Scholar
  11. 11.
    Marklund, L., Johansson, M., Sandberg, K., and Andersson, L. (1996) A missense mutation in the gene for melanocyte—stimulating hormone receptor (MCIR) is associated with the chestnut coat color in horses. Mamm. Genome 7, 895–899.PubMedCrossRefGoogle Scholar
  12. 12.
    Klungland, H., Vage, D. I., Gomezraya, L., Adalsteinsson, S., and Lien, S. (1995) The role of melanocyte—stimulating hormone (MSH) receptor in bovine coat colour determination. Mamm. Genome 6, 636–639.PubMedCrossRefGoogle Scholar
  13. 13.
    Joerg, H., Fries, H. R., Meijerink, E., and Stranzinger, G.F. (1996) Red coat color in holstein cattle is associated with a deletion in the MSHR gene. Mamm. Genome 7, 317–319.PubMedCrossRefGoogle Scholar
  14. 14.
    Takeuchi, S., Suzuki, H., Yabuuchi, M., and Takahashi, S. (1996) A possible involvement of melanocortin 1—receptor in regulating feather color pigmentation in the chicken. Biochim. Biophys. Acta 1308, 164–168.PubMedCrossRefGoogle Scholar
  15. 15.
    Vage, D. I., Lu, D., Klungland, H., Lien, S., Adalsteinsson, S., and Cone, R. D. (1997) A non—epistatic interaction of agouti and extension in the fox, Vulpes vulpes. Nat. Genet. 15, 311–315.CrossRefGoogle Scholar
  16. 16.
    Wong, G., Pawelek, J., Sansone, M., and Morowitz, J. (1974) Response of mouse melanoma cells to melanocyte stimulating hormone. Nature 248, 351–354.PubMedCrossRefGoogle Scholar
  17. 17.
    Fuller, B. B. and Meyskens, F. L. Jr. (1981) Endocrine responsiveness in human melanocytes and melanoma cells in culture. J. Natl. Cancer Inst. 66, 799–802.PubMedGoogle Scholar
  18. 18.
    Tatro, J. B., Atkins, M., Mier, J. W., Hardarson, S., Wolfe, H., Smith, T., Entwistle, M. L., and Reichlin, S. (1990) Melanotropin receptors demonstrated in situ in human melanoma. J. Clin. Invest. 85, 1825–1832.PubMedCrossRefGoogle Scholar
  19. 19.
    Cone, R. D., Mountjoy, K. M., Robbins, L. S., Nadeau, J. H., Johnson, K. R., Roselli—Rehfuss, L., and Mortrud, M. T. (1993) Cloning and functional characterization of a family of receptors for the melanotropic peptides. Ann. N.Y. Acad. Sci. 680, 342–363.Google Scholar
  20. 20.
    Eberle, A. N., Siegrist, W., Bagutti, C., Chluba—de Tapia, J., Solca, F., Wikberg, J. E. S., and Chhajlani, V. (1993) Receptors for melanocyte—stimulating hormone on melanoma cells. Ann. N.Y. Acad. Sci. 680, 320–341.CrossRefGoogle Scholar
  21. 21.
    Siegrist, W., Stutz, S., and Eberle, A. N. (1994) Homologous and heterologous regulation of a—melanocyte stimulating hormone receptors in human and mouse melanoma cell lines. Cancer Res. 54, 2604–2610.PubMedGoogle Scholar
  22. 22.
    Suzuki, I., Cone, R. C., Im, S., Nordlund, J., and Abdel—Malek, Z. A. (1996) Binding of melanotropic hormones to the melanocortin receptor MC1R on human melanocytes stimulates proliferation and melanogenesis. Endocrinology 137, 1627–1633.PubMedCrossRefGoogle Scholar
  23. 23.
    Chakraborty, A. and Pawelek, J. (1993) MSH receptors in immortalized human epidermal keratinocytes: a potential mechanism for coordinate regulation of the epidermal—melanin unit. J. Cell. Physiol. 157, 344–350.PubMedCrossRefGoogle Scholar
  24. 24.
    Sharma, S. D., Jiang, J., Hadley, M. E., Bentley, D. L., and Hruby, V. J. (1996) Melanotropic peptide—conjugated beads for microscopic visualization and characterization of melanoma melanotropin receptors. Proc. Natl. Acad. Sci. U. S. A. 93, 13715–13720.PubMedCrossRefGoogle Scholar
  25. 25.
    Bhardwaj, R.S., Becher, E., Mahnke, K., Hartmeyer, M., Scholzen, T., and Luger, T.A. (1996) Evidence of the expression of a functional melanocortin receptor 1 by human keratinocytes. J. Invest. Dermatol. 106, 817.Google Scholar
  26. 26.
    Xia, Y., Skoog, V., Muceniece, R., Chhajlani, V., and Wikberg, J. E. S. (1995a) Polyclonal antibodies against human melanocortin MC, receptor: preliminary immunohistochemical localisation of melanocortin MC1 receptor to malignant melanoma cells. Eur. J. Pharmacol. 288, 277–283.PubMedCrossRefGoogle Scholar
  27. 27.
    Friedmann, P. S., and Gilchrest, B. A. (1987) Ultraviolet radiation directly induces pigment production by cultured human melanocytes. J. Cell. Physiol. 133, 88–94.PubMedCrossRefGoogle Scholar
  28. 28.
    Friedmann, P. S., Wren, F., Buffey, J., and MacNeil, S. (1990a) a—MSH causes a small rise in cAMP but has no effect on basal or ultraviolet—stimulated melanogenesis in human melanocytes. Br. J. Dermatol. 123, 145–151.Google Scholar
  29. 29.
    Friedmann, P.S., Wren, F.E., Matthews, J.N. (1990b) Ultraviolet stimulated melanogenesis by human melanocytes is augmented by di—acyl glycerol but not TPA. J. Cell. Physiol. 142, 334–341.PubMedCrossRefGoogle Scholar
  30. 30.
    Eller, M. S., Yaar, M., Gilchrest, B. A. (1994) DNA damage and melanogenesis. Science 372, 413–414.Google Scholar
  31. 31.
    Chakraborty, A., Slominski, A., Ermak, G., Hwang, J., and Pawelek, J. (1995) Ultraviolet B and melanocyte—stimulating hormone (MSH) stimulate mRNA production for aMSH receptors and proopiomelanocortin—derived peptides in mouse melanoma cells and transformed keratinocytes: J. Invest. Dermatol. 105, 655–659.PubMedCrossRefGoogle Scholar
  32. 32.
    Rajora, N., Ceriani, G., Catania, A., Star, R. A., Murphy, M. T., and Lipton, J. M. Alpha—MSH production, receptors, and influence on neopterin in a human monocyte/macrophage cell line. J. Leukocyte Biol. 59, 248–253.Google Scholar
  33. 33.
    Hartmeyer, M., Scholzen, T., Becher, E., Bhardwaj, R. S., Fastrich, M., Schwarz, T., and Luger, T. A. Human microvascular endothelial cells (HMEC-1) express the melanocortin receptor type 1 and produce increased levels of IL-8 upon stimulation with a—MSH. J. Invest.Dermatol. 106, 809.Google Scholar
  34. 34.
    Xia, Y., Wikberg, J. E. S., and Chhajlani, V. (1995b) Expression of the melanocortin 1 receptor in periaqueductal grey matter. Neuroreport 6, 2193–2196.PubMedCrossRefGoogle Scholar
  35. 35.
    Vanetti, M., Schonrock, C., Meyerhof, W., and Hollt, V. (1994) Molecular cloning of a bovine MSH receptor which is highly expressed in the testis. FEBS Lett. 348, 268–272.PubMedCrossRefGoogle Scholar
  36. 36.
    Chhajlani, V. (1996) Distribution of cDNA for melanocortin receptor subtypes in human tissues. Biochem. Mol. Biol. Int. 38, 73–80.PubMedGoogle Scholar
  37. 37.
    Schioth, H. B., Muceniece, R., Wikberg, J.E.S., and Chhajlani, V. (1995) Characterisation of melanocortin receptor subtypes by radioligand binding analysis. Eur. J. Pharmacol. 288, 311–317.Google Scholar
  38. 38.
    Thody, A. J., Ridley, K., Penny, R. J., Chalmers, R., Fisher, C., and Shuster, S. (1983) MSH peptides are present in mammalian skin. Peptides 4, 813–816.PubMedCrossRefGoogle Scholar
  39. 39.
    Farooqui, J. Z., Medrano, E. E., Abdel—Malek, Z., and Nordlund, J. (1993) The expression of proopiomelanocortin and various POMC—derived peptides in mouse and human skin. Ann. N. Y. Acad. Sci. 680, 508–510.CrossRefGoogle Scholar
  40. 40.
    Slominski, A., Ermak, G., Hwang, J., Chakraborty, A., Mazurkiewicz, J. E., and Mihm, M. (1995) Proopiomelanocortin, corticotropin releasing hormone and corticotropin releasing hormone receptor genes are expressed in human skin. FEBS Lett. 374, 113–116.PubMedCrossRefGoogle Scholar
  41. 41.
    Wintzen, M., Yaar, M., Burbach, J. P. H., and Gilchrest, B. A. (1996) Proopiomelanocortin gene product regulation in keratinocytes. J. Invest. Dermatol. 106, 673–678.PubMedCrossRefGoogle Scholar
  42. 42.
    Kazumasa, W., Graham, A., Cook, D., and Thody, A. J. (1997) Characterisation of ACTH peptides in human skin and their activation of the melanocortin-1 receptor. Pigment Cell Res. 10, 288–297.CrossRefGoogle Scholar
  43. 43.
    Buffey, J., Thody, A. J., Bleehen, S. S., and MacNeil, S. (1992) a—Melanocytestimulating hormone stimulates protein kinase C activity in murine B 16 melanoma. J. Endocrinol. 133, 333–340.Google Scholar
  44. 44.
    Kuzumaki, T., Matsuda, A., Wakamatsu, K., Ito, S., and Ishikawa, K. (1993) Eumelanin biosynthesis is regulated by coordinate expression of tyrosinase and tyrosinase—related protein-1 genes. Exp. Cell Res. 207, 33–40.PubMedCrossRefGoogle Scholar
  45. 45.
    Park, H. —Y., Russakovsky, V., Ao, Y., Fernandez, E., and Gilchrrest, B. A. (1996) a—Melanocyte stimulating hormone—induced pigmentation is blocked by depletion of protein kinase C. Exp. Cell Res. 227, 70–79.Google Scholar
  46. 46.
    Prota, G. and Thomson, R. H. (1976) Melanin pigmentation in mammals. Endeavour 35, 32–38.PubMedCrossRefGoogle Scholar
  47. 48.
    Halaban, R., Tyrrell, L., Longley, J., Yarden, Y., and Rubin, J. (1993) Pigmentation and proliferation of human melanocytes and the effects of melanocyte—stimulating hormone and ultraviolet B light. Ann. N. Y. Acad. Sci. 680, 290–301.PubMedCrossRefGoogle Scholar
  48. 47.
    Thody, A. J., Higgins, E. M., Wakamatsu, K., Ito, S., Burchill, S. A., and Marks, J. M. (1991) Pheomelanin as well as eumelanin is present in human epidermis. J. Invest. Dermatol. 97, 340–344.PubMedCrossRefGoogle Scholar
  49. 49.
    Hunt, G., Donatien, P. D., Cresswell, J. E., and Thody, A. J. (1993) The effect of alpha—MSH on the attachment of human melanocytes to laminin and fibronectin. Ann. N. Y. Acad. Sci. 680, 549–551.PubMedCrossRefGoogle Scholar
  50. 50.
    Spiro, J., Parker, S., Oliver, I., Fraser, C., Marks, J. M., and Thody, A. J. (1987) Effect of PUVA on plasma and skin immunoreactive alpha—melanocyte stimulating hormone concentrations. Br. J. Dermatol. 117, 703–707.PubMedCrossRefGoogle Scholar
  51. 51.
    Kwon, B. S., Haq, A. K., Pomerantz, S. H., and Halaban, R. (1987) Isolation and sequence of a cDNA clone for human tyrosinase that maps at the mouse c—albino locus. Proc. Natl. Acad. Sci. U. S. A. 84, 7473–7477.PubMedCrossRefGoogle Scholar
  52. 52.
    Tsukamoto, K., Jackson, I. J., Urabe, K., Montague, P. M., and Hearing, V. J. (1992) A second tyrosinase—related protein, TRP-2, is a melanogenic enzyme termed DOPAchrome tautomerase. EMBO J. 11, 519–526.Google Scholar
  53. 53.
    Jimenez—Cervantes, C., Solano, F., Kobayashi, T., Urabe, K., Hearing, V. J., Lozano, J. A., and Garcia—Borron, J. C. (1994) A new enzymatic function in the melanogenic pathway. The 5,6—dihydroxyindole-2—carboxylic acid oxidase activity of tyrosinase—related protein-1 (TRP-1). J. Biol. Chem. 269, 17,993–18, 000.Google Scholar
  54. 54.
    Bertolotto, C., Bille, K., Ortonne, J.—P., and Ballotti, R. (1996) Regulation of tyrosinase gene expression by cAMP in B16 melanoma cells involves two CATGTG motifs surrounding the TATA box: implication of the microphthalmia gene product. J. Cell Biol. 134, 747–755.PubMedCrossRefGoogle Scholar
  55. 55.
    Yasumoto, K., Mahalingam, H., Suzuki, H., Yoshizawa, M., Yokoyama, K. (1995) Transcriptional activation of the melanocyte-specific genes by the human homolog of the mouse Microphthalmia protein. J. Biochem. 118, 874–881.PubMedCrossRefGoogle Scholar
  56. 56.
    Aroca, P., Urabe, K., Kobayashi, T., Tsukamoto, K., and Hearing, V.J. (1993) Melanin biosynthesis patterns following hormonal stimulation. J. Biol. Chem. 268, 25,650–25, 655.Google Scholar
  57. 57.
    Hunt, G., Donatien, P. D., Lunec, J., Todd, C., Kyne, S., and Thody, A. J. (1994) Cultured human melanocytes respond to MSH peptides and ACTH. Pigment Cell Res. 7, 217–221.PubMedCrossRefGoogle Scholar
  58. 58.
    Konda, Y., Gantz, I., DelValle, J., Shimoto, Y., Miwa, H., and Yamada, T. (1994) Interaction of dual intracellular signaling pathways activated by the melanocortin3 receptor. J. Biol. Chem. 269, 13,162–13, 166.Google Scholar
  59. 59.
    Chuang, T. Y., Reizner, G. T., Elpern, D. J., Stone, J. L., and Farmer, E. R. (1995) Nonmelanoma skin cancer in Japanese ethnic Hawaiians in Kauai, Hawaii: an incidence report. J. Am. Acad. Dermatol. 33, 422–426.PubMedCrossRefGoogle Scholar
  60. 60.
    Elder, D.E. (1995) Skin cancer: melanoma and other specific nonmelanoma skin cancers. Cancer 75 (Suppl. 1), 245–256.PubMedCrossRefGoogle Scholar
  61. 61.
    Halder, R. M. and Bridgeman—Shah, S. (1995) Skin cancer in African Americans. Cancer 75 (Suppl. 2), 667–673.PubMedCrossRefGoogle Scholar
  62. 62.
    Luande, J., Henschke, C. I., and Mohammed, N. (1985) The Tanzanian human albino skin. Cancer 55, 1823–1828.PubMedCrossRefGoogle Scholar
  63. 63.
    Diffey, B.L., Healy, E., Thody, A.J., and Rees, J.L. (1995) Melanin, melanocytes, and melanoma. Lancet 346, 1713.PubMedCrossRefGoogle Scholar
  64. 64.
    Lookingbill, D. P., Lookingbill, G. L., and Leppard, B. Actinic damage and skin cnacer in albinos in northern Tanzania: findings in 164 patients enrolled in an outreach skin care program. J. Am. Acad. Dermatol. 32, 653–658.Google Scholar
  65. 65.
    Kaidbey, K. H., Agin, P. P., Sayre, R. M., and Kligman, A. M. (1979) Photoprotection by melanin: a comparison of black and Caucasin skin. J. Am. Acad. Dermatol. 1, 249–260.PubMedCrossRefGoogle Scholar
  66. 66.
    Bodmer, W. F. and Cavalli-Sforza, L. L. (1976) Racial differentiation, in Genetics, Evolution and Man. ( Bodmer, W. F. and Cavalli-Sforza, L. L., eds.) Freeman, New York, pp. 559–604.Google Scholar
  67. 67.
    Kingdon, J. (1993) Self-Made Man and His Undoing. Simon and Schuster, London.Google Scholar
  68. 68.
    Valverde, P., Healy, E., Jackson, I., Rees, J. L., and Thody, A. J. (1995) Variants of the melanocyte-stimulating hormone receptor gene are associated with red hair and fair skin in humans: Nat. Genet. 11, 328–330.PubMedCrossRefGoogle Scholar
  69. 69.
    Box, N. F., Wyeth, J. R., O’Gorman, L. E., Martin, N. G., and Sturm, R. A. (1997) Characterisation of melanocyte-stimulating hormone receptor variant alleles in twins with red hair. Hum. Mol. Genet. 11, 1891–1897.CrossRefGoogle Scholar
  70. 70.
    Prota, G., Lamoreux, M. L., Muller, J., Kobayashi, T., Napolitano, A., Vincensi, M. R., Sakai, C., and Hearing, V. J. (1995) Comparative analysis of melanins and melanosomes produced by various coat color mutants. Pigment Cell Res. 8, 153–163.PubMedCrossRefGoogle Scholar
  71. 71.
    Barsh, G.S. (1996) The genetics of pigmentation: from fancy genes to complex traits. Trends Genet. 12, 299–305.PubMedCrossRefGoogle Scholar
  72. 72.
    Beim, S. F., Judge, P., Urbach, F., MacCon, C. F., and Martin, F. (1970) Skin cancer in County Galway, Ireland. Proc. Natl. Cancer Conf. 6, 489–500.Google Scholar
  73. 73.
    Fitzpatrick, T.B. (1988) The validity and practicality of sun-reactive skin types I through VI. Arch. Dermatol. 124, 869–871.PubMedCrossRefGoogle Scholar
  74. 74.
    Rampen, F. H., Fleuren, B. A., de Boo, T. M., and Lemmens, W. A. (1988) Unreliability of self-reported burning tendency and tanning ability. Arch. Dermatol. 124, 885–888.PubMedCrossRefGoogle Scholar
  75. 75.
    Jimbow, K., Ishida, O., Ito, S., Hori, Y., Witkop, C.J., and King, R.A. (1983) Combined chemical and electron microscopic studies of pheomelanosomes in human red hair. J. Invest. Dermatol. 81, 506–511.PubMedCrossRefGoogle Scholar
  76. 76.
    Schioth, H. B., Muceniece, R., Szardenings, M., Prusis, P., Lindeberg, G., Sharma, S. D., Hruby, V. J., and Wikberg, J. E. S. (1997) Characterisation of D117A and H260A mutations in the melanocortin 1 receptor. Mol. Cell. Endocrinol. 126, 213–219.PubMedCrossRefGoogle Scholar
  77. 77.
    Xu, X., Thornwall, M., Luhdin, L. G., Chhajlani, V. (1996) Va192Met variant of the melanocyte stimulating hormone receptor gene. Nat. Genet. 14, 384.Google Scholar
  78. 78.
    Koppula, S. V., Robbins, L. S., Lu, D., Baack, E., White, C. R.Jr., Swanson, N. A., and Cone, R. D. (1997) Identification of common polymorhpisms in the coding sequence of the human MSH receptor (MC1R) with possible biological effects. Hum. Mutat. 9, 30–36.PubMedCrossRefGoogle Scholar
  79. 79.
    Valverde, P., Healy, E., Sikkink, S., Haldane, F., Thody, A. J., Carothers, A., Jackson, I. J., and Rees, J. L. (1996) The Asp84Glu variant of the melanocortin 1 receptor (MC1R) is associated with melanoma. Hum. Mol. Genet. 5, 1663–1666.Google Scholar
  80. 80.
    Chluba-de Tapia, J., Bagutti, C., Cotti, R., and Eberle, A. N. (1996) Induction of constitutive melanogenesis in amelanotic mouse melanoma cells by transfection of the human melanocortin-1 receptor gene. J. Cell. Sci. 109, 2023–2030.Google Scholar
  81. 81.
    Loir, B., Bouchard, B., Morandini, R., Del Marmol, V., Deraemaecker, R., GarciaBorron, J. C., and Ghanem G. (1997) Immunoreactive a—melanotropin as an autocrine effector in human melanoma cells. Eur. J. Biochem. 244, 923–930.PubMedCrossRefGoogle Scholar
  82. 82.
    Hunt, G., Todd, C., and Thody, A. J. (1996) Unresponsiveness of human epidermal melanocytes to melanocyte—stimulating hormone and its association with red hair. Mol. Cell. Endocrinol. 116, 131–136.PubMedCrossRefGoogle Scholar
  83. 83.
    Singleton, W. R. and Ellis, B. (1964) Inheritance of red hair for six generations. J. Hered. 55, 261 + 266.Google Scholar
  84. 84.
    Eiberg, H. and Mohr, J. (1987) Major locus for red hair color linked to MNS blood groups on chromosome 4. Clin. Genet. 32, 125–128.PubMedCrossRefGoogle Scholar
  85. 85.
    Armstrong, B. K. and Kricker, A. (1996) Epidemiology of sun exposure and skin cancer, in Cancer Surveys Skin Cancer, Vol. 26 ( Leigh, I. M., Newton Bishop, J. A., and Kripke, M. L., eds.). Cold Spring Harbor Laboratory Press, New York, pp. 133–153.Google Scholar
  86. 86.
    Menon, I. A., Persad, S., Haberman, H. F., and Kurian, C. J. (1983) A comparative study of the physical and chemical properties of melanins isolated from human black and red hair. J. Invest. Dermatol. 80, 202–206.PubMedCrossRefGoogle Scholar
  87. 87.
    Persad, S., Menon, I. A., and Haberman, H. F. (1983) Comparison of the effects of UV—visible irradiation of melanins and melanin—hematoporphyrin complexes from human black and red hair. Photochem. Photobiol. 37, 63–68.PubMedCrossRefGoogle Scholar
  88. 88.
    Hunt, G., Kyne, S., Ito, S., Wakamatsu, K., Todd, C., and Thody, A. J. (1995) Eumelanin and phaeomelanin contents of human epidermis and cultured melanocytes. Pigment Cell Res. 8, 202–208.PubMedCrossRefGoogle Scholar
  89. 89.
    Lunec, J., Pieron, C., Sherbet, G.V., and Thody, A.J. (1990) Alpha—melanocytestimulating hormone immunoreactivity in melanoma cells. Pathobiology 58, 193–197.PubMedCrossRefGoogle Scholar
  90. 90.
    Lunec, J., Pieron, C., and Thody, A.J. (1992) MSH receptor expression and the relationship to melanogenesis and metastatic activity in B16 melanoma. Melanoma Res. 2, 5–12.PubMedCrossRefGoogle Scholar
  91. 91.
    Bhate, S. M., Sharpe, G. R., Marks, J. M., Shuster, S., Ross, W.M. (1993) Prevalence of skin and other cancers in patients with psoriasis. Clin. Exp. Dermatol. 18, 401–404.PubMedCrossRefGoogle Scholar
  92. 92.
    Stern, R. S., Nichols, K. T., and Vakeva, L. H. (1997) Malignant melanoma in patients treated for psoriasis with methoxsalen (psoralen) and ultraviolet A radiation (PUVA). N. Engl. J. Med. 336, 1041–1045.PubMedCrossRefGoogle Scholar
  93. 93.
    Saunder, D. N., and Nordlund, J. J. (1989) Alpha—melanocyte stimulating hormone modulates contact hypersensitivity responsiveness in C57/B16 mice. J. Invest. Dermatol. 93, 511–517.PubMedCrossRefGoogle Scholar
  94. 94.
    Smith, R., Healy, E., Siddiqui, S., Flanagan, N., Steijlen, P., Rosdahl, I., Rogers, S., etal. (1998) Melanocortin 1 receptor variants in an Irish population. J. Invest. Dermatol. 111, 101–104.Google Scholar

Copyright information

© Springer Science+Business Media New York 2000

Authors and Affiliations

  • Eugene Healy
  • Mark Birch-Machin
  • Jonathan L. Rees

There are no affiliations available

Personalised recommendations