The cutaneous serotoninergic/melatoninergic system: Securing a place under the sun

FASEB Journal

Volumn 19, Issue 2, 2005, Pages 176-194

  Slominski, Andrzej ^a   Wortsman, Jacobo ^b   Tobin, Desmond J ^c  

^a UNIVERSITY OF TENNESSEE HEALTH SCIENCE CENTER   (United States)

^b SOUTHERN ILLINOIS UNIVERSITY SCHOOL OF MEDICINE   (United States)

^c UNIVERSITY OF BRADFORD   (United Kingdom)

Author keywords

Endocrinology of the skin; Melatonin; N acetylserotonin; Serotonin; Skin

Indexed keywords

MELATONIN; MELATONIN RECEPTOR; SEROTONIN; SEROTONIN RECEPTOR;

ANIMAL CELL; ANIMAL EXPERIMENT; ANIMAL MODEL; ANIMAL TISSUE; AUTOCRINE EFFECT; BIODEGRADATION; CELL DIFFERENTIATION; CELL METABOLISM; CELL POPULATION; CELL PROLIFERATION; CIRCADIAN RHYTHM; CONTROLLED STUDY; DRUG ACTIVITY; ENVIRONMENTAL FACTOR; FIBROBLAST; HAIR FOLLICLE; HOMEOSTASIS; HUMAN; HUMAN CELL; HUMAN TISSUE; KERATINOCYTE; MAMMAL; MELANOCYTE; MELATONINERGIC SYSTEM; NONHUMAN; NUCLEOTIDE SEQUENCE; OXIDATIVE STRESS; PARACRINE SIGNALING; PHENOTYPE; PINEAL BODY; PRIORITY JOURNAL; PROTEIN EXPRESSION; RECEPTOR INTRINSIC ACTIVITY; REVIEW; RODENT; SEROTONIN METABOLISM; SEROTONINERGIC SYSTEM; SKIN; SKIN FUNCTION; SKIN STIMULATION;

ANIMALS; HOMEOSTASIS; HUMANS; MELATONIN; SEROTONIN; SKIN; SOLAR SYSTEM; SUNLIGHT;

ANIMALIA; MAMMALIA; RODENTIA;

EID: 13244267262     PISSN: 08926638     EISSN: None     Source Type: Journal    
DOI: 10.1096/fj.04-2079rev     Document Type: Review

References (193)

1. Slominski, A., and Wortsman, J. (2000) Neuroendocrinology of the skin. Endocr. Rev. 21, 457-487
2. Slominski, A., Wortsman, J., Luger, T., Paus, R., and Salomon, S. (2000) Corticotropin releasing hormone and proopiomelanocortin involvement in the cutaneous response to stress. Physiol. Rev. 80, 979-1020
3. Slominski, A., Tobin, D. J., Shibahara, S., and Wortsman, J. (2004) Melanin pigmentation in mammalian skin and its hormonal regulation. Physiol. Rev. In press
4. Bos, J. D. (1997) Skin Immune System (SIS), CRH Press, Boca Raton, Florida
5. Stenn, K. S., and Paus, R. (2001) Controls of hair follicle cycling. Physiol. Rev. 81, 449-494
6. Slominski, A., Wortsman, J., Pisarchik, A., Zbytek, B., Linton, E. A., Mazurkiewicz, J. E., and Wei, E. T. (2001) Cutaneous expression of corticotropin-releasing hormone (CRH), urocortin, and CRH receptors. FASEB J. 15, 1678-1693
7. Slominski, A., Pisarchik, A., Tobin, D. J., Mazurkiewicz, J. E., and Wortsman, J. (2004) Differential expression of a cutaneous corticotropin- releasing hormone system. Endocrinology 145, 941-950
8. Chen, W., Thiboutot, D., and Zouboulis, C. C. (2002) Cutaneous androgen metabolism: basic research and clinical perspectives. J. Invest. Dermatol. 119, 992-1007
9. Holick, M. F. (2003) Vitamin D: A millennium perspective. J. Cell. Biochem. 88, 296-307
10. Schallreuter, K. U., Lemke, K. R., Pittelkow, M. R., Wood, J. M., Korner, C., and Malik, R. (1995) Catecholamines in human keratinocyte differentiation. J. Invest. Dermatol. 104, 953-957
11. Grando, S. A., and Horton, R. M. (1997) The keratinocyte cholinergic system with acetylcholine as an epidermal cytotransmitter. Curr. Opin. Dermatol. 1997, 462-468
12. Schallreuter, K. U., Schulz-Douglas, V., Bunz., A., Beazley, W. D., and Korner, C. (1997) Pteridines in the control of pigmentation. J. Invest. Dermatol. 109, 31-35
13. Gillbro, J. M., Marles, L. K., Hibberts, N. A., and Schallreuter, K. U. (2004) Autocrine catecholamine biosynthesis and the beta-adrenoceptor signal promote pigmentation in human epidermal melanocytes. J. Invest. Dermatol. 123, 346-353
14. Schallreuter, K. U. (1999) A review of recent advances on the regulation of pigmentation in the human epidermis. Cell. Mol. Biol. (Noisy-le-grand) 45, 943-949
15. Mockus, S. M., and Vrana, K. E. (1998) Advances in the molecular characterization of tryptophan hydroxylase. J. Mol. Neurosci. 10, 163-179
16. Yu, H. S., and Reiter, R. J. (1993) Melatonin Biosynthesis, Physiological Effects, and Clinical Implications, CRC Press, Boca Raton
17. Kema, I. P., de Vries, E. G. E., and Muskiet, F. A. J. (2000) Clinical chemistry of serotonin and metabolites. J. Chromatogr. B 747, 33-48
18. Boularand, S., Darmon, M. C., and Mallet, J. (1995) The human tryptophan hydroxylase gene. J Biol. Chem. 270, 3748-3756
19. Wang, G. A., Coon, S. L., and Kaufman, S. (1998) Alternative splicing at the 3′-cDNA of human tryptophan hydroxylase. J. Neurochem. 71, 1769-1772
20. Yang, X. J., and Kaufman, S. (1994) High-level expression and deletion mutagenesis of human tryptophan hydroxylase. Proc. Natl. Acad. Sci. USA 91, 6659-6663
21. Walther, D. J., Peter, J. U., Bashammakh, S., Hortnagl, H., Voits, M., Fink, H., and Bader, M. (2003) Synthesis of serotonin by a second tryptophan hydroxylase isoform. Science 299, 76
22. Zhang, X., Beaulieum, J. M., Sotnikova, T. D., Gainetdinov, R. R., and Caron, M. G. (2004) Tryptophan hydroxylase-2 controls brain serotonin synthesis. Science 305, 217
23. Slominski, A., Pisarchik, A., Semak, I., Sweatman, T., Worstman, J., Szczesniewski, A., Slugocki, G., McNulty, J., Kauser, S., and Tobin, D. J. (2002) Serotoninergic and melatoninergic systems are fully expressed in human skin. FASEB J. 16, 896-898
24. Matsuda, M., Imaoka, T., Vomachka, A. J., Gudelsky, G. A., Hou, Z., Mistry, M., Bailey, J. P., Nieport, K. M., Walther, D. J., Bader, M., et al. (2004) Serotonin regulates mammary gland development via an autocrine-paracrine loop. Dev. Cell 6, 193-203
25. Gordon, J., and Barnes, N. M. (2003) Lymphocytes transport serotonin and dopamine: agony or ecstasy? Trends Immunol. 24, 438-443
26. Leon, J., Acuna-Castrovicjo, D., Sainz, R. M., Mayo, J. C., Tan, D. X., and Reiter, R. J. (2004) Melatonin and mitochondrial function. Life Sci. 75, 765-790
27. Reiter, R. J., and Tan, D. X. (2002) Melatonin: an antioxidant in edible plants. Ann. N.Y. Acad. Sci. 957, 341-344
28. Tan, D. X., Manchester, L. C., Hardeland, R., Lopez-Burillo, S., Mayo, J. C., Sainz, R. M., and Reiter, R. J. (2003) Melatonin: a hormone, a tissue factor, an autocoid, a paracoid, and an antioxidant vitamin. J. Pineal Res. 34, 75-78
29. Tilden, A. R., Becker, M. A., Amma, L. L., Arciniega, J., and McGaw, A. K. (1997) Melatonin production in an aerobic photosynthetic bacterium: an evolutionarily early association with darkness. J. Pineal Res. 22, 102-106
30. Reiter, R. J. (1991) Pineal melatonin: cell biology of its synthesis and of its physiological interactions. Endocr. Rev. 12, 151-180
31. Abe, M., Itoh, M. T., Miyata, M., Ishikawa, S., and Sumi, Y. (1999) Detection of melatonin, its precursors and related enzyme activities in rabbit lens. Exp. Eye Res. 68, 255-262
32. Conti, A., Conconi, S., Hertens, E., Skwarlo-Sonta, K., Markowska, M., and Maestroni, J. M. (2000) Evidence for melatonin synthesis in mouse and human bone marrow cells. J. Pineal Res. 28, 193-202
33. Finocchiaro, L. M., Nahmod, V. E., and Launay J. M. (1991) Melatonin biosynthesis and metabolism in peripheral blood mononuclcar leucocytes. Biochem. J. 280, 727-731
34. Finocchiaro, L. M., and Glikin, G. C. (1998) Intracellular melatonin distribution in cultured cell lines. J. Pineal Res. 24, 22-34
35. Itoh, M. T., Ishizuka, B., Kuribayashi, Y., Amemiya, A., and Sumi, Y. (1999) Melatonin, its precursors, and synthesizing enzyme activities in the human ovary. Mol. Hum. Reprod. 5, 402-408
36. Carrillo-Vico, A., Calvo, J. R., Abreu, P., Lardone, P. J., Garcia-Maurino, S., Reiter, R. J., and Guerrero, J. M. (2004) Evidence of melatonin synthesis by human lymphocytes and its physiological significance: possible role as intracrine, autocrine, and/or paracrine substance. FASEB J. 18, 537-539
37. Yaga, K., Reiter, R. J., and Richardson, B. A. (1993) Tryptophan loading increases daytime serum melatonin levels in intact and pinealectomized rats. Life Sci. 52, 1231-1238
38. Bubenik, G. A. (2002) Gastrointestinal melatonin: localization, function, and clinical relevance. Dig. Dis. Sci. 47, 2336-2348
39. Kvetnoy, I. M., Ingel, I. E., Kvetnaia, T. V., Malinovskaya, N. K., Rapoport, S. I., Raikhlin, N. T., Trofimov, A. V., and Yuzhakov, V. V. (2002) Gastrointestinal melatonin: cellular identification and biological role. Neuroendocrinol. Lett. 23, 121-132
40. Stehle, J. H., von Gall, C., and Korf, H. W. (2002) Organisation of the circadian system in melatonin-proficient C3H and melatonin-deficient C57BL mice: a comparative investigation. Cell Tissue Res. 309, 173-182
41. Coon, S. L., Roseboom, P. H., Baler, R., Weller, J. L., Namboodiri, M. A., Koonin, E. V., and Klein, D. C. (1995) Pineal serotonin N-acetyltransferase: expression cloning and molecular analysis. Science 270, 1681-1683
42. Bernard, M., Donohue, S. J., and Klein, D. C. (1995) Human hydroxyindole-O-methyltransferase in pineal gland, retina and Y79 retinoblastoma cells. Brain Res. 696, 37-48
43. Coon, S. L., Mazuruk, K., Bernard, M., Roseboom, P. H., Klein, D. C., and Rodriguez., I. R. (1996) The human serotonin N-acetyltransferase (EC 2.3.1.87) gene (AANAT): structure, chromosomal localization, and tissue expression. Genomics 34, 76-84
44. Donohue, S. J., Roseboom, P. H., Illnerova, H., Weller, J. L., and Klein, D. C. (1993) Human hydroxyindole-O-methyltransferase: presence of LINE-1 fragment in a cDNA clone and pineal mRNA. DNA Cell Biol. 12, 715-727
45. Rodriguez, I. R., Mazuruk, K., Schoen, T. J., and Chader, G. J. (1994) Structural analysis of the human hydroxyindole-O-methyltransferase gene. Presence of two distinct promoters. J. Biol. Chem. 269, 31969-31977
46. Kopin, I. J., Pare, C. M., Axelrod, J., and Weissbach, H. (1960) 6-Hydroxylation, the major metabolic pathway for melatonin. Biochim. Biophys. Acta 40, 377-378
47. Kopin, I. J., Pare, C. M., Axelrod, J., and Weissbach, H. (1961) The fate of melatonin in animals. J Biol. Chem. 236, 3072-3075
48. Vakkuri, O., Tervo, J., Luttinen, R., Ruotsalainen, H., Rahkamaa, E., and Leppaluoto, J. (1987) A cyclic isomer of 2-hydroxymelatonin: a novel metabolite of melatonin. Endocrinology 120, 2453-2459
49. Tan, D. X., Manchester, L. C., Reiter, R. J., Plummer, B. F., Hardies, L. J., and Weintraub, S. T., Vijayalaxmi, and Shepherd, A. M. (1998) A novel melatonin metabolite, cyclic 3-hydroxymelatonin: a biomarker of in vivo hydroxyl radical generation. Biochem. Biophys. Res. Commun. 253, 614-620
50. Arendt, J. (1988) Melatonin. Clin. Endocrinol. (Oxford) 29, 205-229
51. Facciola, G., Hidestrand, M., von Bahr, C., and Tybring, G. (2001) Cytochrome P450 isoforms involved in melatonin metabolism in human liver microsomes. Eur. J. Clin. Pharmacol. 56, 881-888
52. Rogawski, M. A., Roth, R. H., and Aghajanian, G. K. (1979) Melatonin: deacetylation to 5-methoxytryptamine by liver but not brain aryl acylamidase. J. Neurochem. 32, 1219-1226
53. Grace, M. S., Cahill, G. M., and Besharse, J. C. (1991) Melatonin deacetylation: retinal vertebrate class distribution and Xenopus laevis tissue distribution. Brain Res. 559, 56-63
54. Hardeland, R., Reiter, R. J., Poeggeler, B., and Tan, D. X. (1993) The significance of the metabolism of the neurohormone melatonin: antioxidative protection and formation of bioactive substances. Neurosci. Biobehav. Rev. 17, 347-357
55. Tesoriere, L., Avellone, G., Ceraulo, L., D'Arpa, D., Allegra, M., and Livrea, M. A. (2001) Oxidation of melatonin by oxoferryl hemoglobin: a mechanistic study. Free Radic. Res. 35, 633-642
56. Allegra, M., Furtmuller, P. G., Regelsberger, G., Turco-Liveri, M. L., Tesoriere, L., Perretti, M., Livrea, M. A., and Obinger, C. (2001) Mechanism of reaction of melatonin with human myeloperoxidase. Biochem. Biophys. Res. Commun. 282, 380-386
57. Tan, D. X., Manchester, L. C., Reiter, R. J., Plummer, B. F., Limson, J., Weintraub, S. T., and Qi, W. (2000) Melatonin directly scavenges hydrogen peroxide: a potentially new metabolic pathway of melatonin biotransformation. Free Radic. Biol. Med. 29, 1177-1185
58. Hoyer, D., Hannon, J. P., and Martin, G. R. (2002) Molecular, pharmacological and functional diversity of 5-HT receptors. Pharmacol. Biochem. Behav. 71, 533-554
59. Kroeze, W. K., Kristiansen, K., and Roth, B. L. (2002) Molecular biology of serotonin receptors structure and function at the molecular level. Curr. Top. Med. Chem. 2, 507-528
60. Richerson, G. B. (2004) Serotonergic neurons as carbon dioxide sensors that maintain pH homeostasis. Nat. Rev. Neurosci. 5, 449-461
61. Seuwen, K., and Pouyssegur, J. (1990) Serotonin as a growth factor. Biochem. Pharmacol. 39, 985-990
62. Buznikov, G. A., Lambert, H. W., and Lander, J. M. (2001) Serotonin and serotonin-like substances as regulators of early embryogenesis and morphogenesis. Cell Tissue Res. 305, 177-186
63. Talley, N. J. (2001) Serotoninergic neuroenteric modulators. Lancet 358, 2061-2068
64. Frohlich, P. F., and Meston, C. M. (2000) Evidence that serotonin affects female sexual functioning via peripheral mechanisms. Physiol. Behav. 71, 383-393
65. Nebigil, C. G., Launay, J. M., Hickel, P., Tournois, C., and Maroteaux, L. (2000) 5-Hydroxytryptamine 2B receptor regulates cell-cycle progression: cross-talk with tyrosine kinase pathways. Proc. Natl. Acad. Sci. USA 97, 2591-2596
66. Nebigil, C. G., Choi, D. S., Dierich, A., Hickel, P., Le Meur, M., Messaddeq, N., Launay, J. M., and Maroteaux, L. (2000) Serotonin 2B receptor is required for heart development. Proc. Natl. Acad. Sci. USA 97, 9508-9513
67. Lefebvre, H., Compagnon, P., Contesse, V., Delarue, C., Thuillez, C., Vaudry, H., and Kuhn, J. M. (2001) Production and metabolism of serotonin (5-HT) by the human adrenal cortex: paracrine stimulation of aldosterone secretion by 5-HT. J. Clin. Endocrinol. Metab. 86, 5001-5007
68. Kawano, H., Tsuji, H., Nishimura, H., Kimura, S., Yano, S., Ukimura, N., Kunieda, Y., Yoshizumi, M., Sugano, T., Nakagawa, K., et al. (2001) Serotonin induces the expression of tissue factor and plasminogen activator inhibitor-1 in cultured rat aortic endothelial cells. Blood 97, 1697-1702
69. Vesela, J., Rehak, P., Mihalik, J., Czikkova, S., Pokorny, J., and Koppel, J. (2003) Expression of serotonin receptors in mouse oocytes and preimplantation embryos. Physiol. Res. 52, 223-228
70. Goldberg, J. I., and Kater, S. B. (1989) Expression and function of the neurotransmitter serotonin during development of the Helisoma nervous system. Dev. Biol. 131, 483-495
71. Julius, D. (1991) Molecular biology of serotonin receptors. Annu. Rev. Neurosci. 14, 335-360
72. Nebigil, C. G., Etienne, N., Messaddeq, N., and Maroteaux, L. (2003) Serotonin is a novel survival factor of cardiomyocytes: mitochondria as a target of 5-HT2B receptor signaling. FASEB J. 17, 1373-1375
73. Zilkha-Falb, R., Ziv, I., Nardi, N., Offen, D., Melamed, E., and Barzilai, A. (1997) Monoamine-induced apoptotic neuronal cell death. Cell. Mol. Neurobiol. 17, 101-118
74. Serafeim, A., Grafton, G., Chamba, A., Gregory, C. D., Blakely, R. D., Bowery, N. G., Barnes, N. M, and Gordon, J. (2002) 5-Hydroxytryptamine drives apoptosis in biopsylike Burkitt lymphoma cells: reversal by selective serotonin reuptake inhibitors. Blood 99, 2545-2553
75. Reiter, R. J. (2003) Melatonin: clinical relevance. Best Pract. Res. Clin. Endocrinol. Metab. 17, 273-285
76. Reiter, R. J. (2002) Potential biological consequences of excessive light exposure: melatonin suppression, DNA damage, cancer and neurodegenerative diseases. Neuroendocrinol. Lett. 23, Suppl. 2, 9-13
77. Dubocovich, M. L., Masana, M. I., and Benloucif, S. (1999) Molecular pharmacology and function of melatonin receptor subtypes. Adv. Exp. Med. Biol. 460, 181-190
78. Sugden, D., Pickering, H., Teh, M. T., and Garratt, P. J. (1997) Melatonin receptor pharmacology: toward subtype specificity. Biol. Cell. 89, 531-537
79. Carlberg, C. (2000) Gene regulation by melatonin. Ann. N. Y. Acad. Sci. 917, 387-396
80. Nosjean, O., Nicolas, J. P., Klupsch, F., Delagrange, P., Canet, E., and Boulin, J. A. (2001) Comparative pharmacological studies of melatonin receptors: MT1, MT2 and MT3/QR2. Tissue distribution of MT3/QR2. Biochem. Pharmacol. 61, 1369-1379
81. Allegra, M., Reiter, R. J., Tan, D. X., Gentile, C., Tesoriere, L., and Livrea, M. A. (2003) The chemistry of melatonin's interaction with reactive species. J. Pineal Res. 34, 1-10
82. Reiter, R. J. (1996) Functional diversity of the pineal hormone melatonin: its role as an antioxidant. Exp. Clin. Endocrinol. Diabetes 104, 10-16
83. Mayo, J. C., Sainz, R. M., Antoli, I., Herrera, F., Martin, V., and Rodriguez, C. (2002) Melatonin regulation of antioxidant enzyme gene expression. Cell. Mol. Life Sci. 59, 1706-1713
84. Rodriguez, C., Mayo, J. C., Sainz, R. M., Antolin, I., Herrera, F., Martin, V., and Reiter, R. J. (2004) Regulation of antioxidant enzymes: a significant role for melatonin. J. Pineal Res. 36, 1-9
85. Martin, M., Macias, M., Escames, G., Leon, J., and Acuna-Castroviejo, D. (2000) Melatonin but not vitamins C and E maintains glutathione homeostasis in t-butyl hydroperoxide-induced mitochondrial oxidative stress. FASEB J. 14, 1677-1679
86. Karbownik, M., Lewinski, A., and Reiter, R. J. (2001) Anticarcinogenic actions of melatonin which involve antioxidative processes: comparison with other antioxidants. Int. J. Biochem. Cell Biol. 33, 735-753
87. Skwarlo-Sonta, K. (2002) Melatonin in immunity: comparative aspects. Neuroendocrinol. Lett. 23, Suppl. 1, 61-66
88. Cos, S., and Sanchez-Barcelo, E. J. (2000) Melatonin and mammary pathological growth. Front. Neuroendocrinol. 21, 133-170
89. Gupta, D., Atanasio, A., and Reiter, R. J. (1988) The Pineal Gland and Cancer, Brain Research Promotion, Oxford
90. Reppert, S. M., Weaver, D. R., and Ebisawa, T. (1994) Cloning and characterization of a mammalian melatonin receptor that mediates reproductive and circadian responses. Neuron 13, 1177-1185
91. Reppert, S. M., Godson, C., Mahle, C. D., Weaver, D. R., Slaugenhaupt, S. A., and Gusella, J. F. (1995) Molecular characterization of a second melatonin receptor expressed in human retina and brain: the Mellb melatonin receptor. Proc. Natl. Acad. Sci. USA 92, 8734-8738
92. Witt-Enderby, P. A., Bennett, J., Jarzynka, M. J., Firestine, S., and Melan, M. A. (2003) Melatonin receptors and their regulation: biochemical and structural mechanisms. Life Sci. 72, 2183-2198
93. Dubocovich, M. L., Masana, M. I., Iacob, S., and Sauri, D. M. (1997) Melatonin receptor antagonists that differentiate between the human Mella and Mellb recombinant subtypes are used to assess the pharmacological profile of the rabbit retina MLl presynaptic heteroreceptor. Naunyn Schmiedebergs Arch. Pharmacol. 355, 365-375
94. Brydon, L., Petit, L., Delagrange, P., Strosberg, A. D., and Jockers, R. (2001) Functional expression of MT2 (Mellb) melatonin receptors in human PAZ6 adipocytes. Endocrinology 142, 4264-4271
95. Brydon, L., Roka, F., Petit, L., de Coppet, P., Tissot, M., Barrett, P., Morgan, P. J., Nanoff, C., Strosberg, A. D., and Jockers, R. (1999) Dual signaling of human Mella melatonin receptors via G(i2), G(i3), and G(q/11) proteins. Mol. Endocrinol. 13, 2025-2038
96. Nelson, C. S., Ikeda, M., Gompf, H. S., Robinson, M. L., Fuchs, N. K., Yoshioka, T., Neve, K. A., and Allen, C. N. (2001) Regulation of melatonin 1a receptor signaling and trafficking by asparagine-124. Mol. Endocrinol. 15, 1306-1317
97. Wiesenberg, I., Missbach, M., Kahlen, J. P., Schrader, M., and Carlberg, C. (1995) Transcriptional activation of the nuclear receptor RZR alpha by the pineal gland hormone melatonin and identification of CGP 52608 as a synthetic ligand. Nucleic Acids Res. 23, 327-333
98. Carlberg, C., Hooft van Huijsduijnen, R., Staple, J. K., De Lamarter, J. F., and Becker-Andre, M. (1994) RZRs, a new family of retinoid-related orphan receptors that function as both monomers and homodimers. Mol. Endocrinol. 8, 757-770
99. Nosjean, O., Ferro, M., Coge, F., Beauverger, P., Henlin, J. M., Lefoulon, F., Fauchere, J. L., Delagrange, P., Canet, E., and Boulin, J. A. (2000) Identification of the melatonin-binding site MT3 as the quinone reductase 2. J. Biol. Chem. 275, 31311-31317
100. Slominski, A., Pisarchik, A., Johansson, O., Jing, C., Semak, I., Slugocki, G., and Wortsman. J. (2003) Tryptophan hydroxylase (TPH) expression in human skin cells. Biochim. Biophys. Acta 1639, 80-86
101. Finocchiaro, L. M., Arzt, E. S., Fernandez-Castelo, S., Criscuolo, M., Finkielman, S., and Nahmod, V. E. (1988) Serotonin and melatonin synthesis in peripheral blood mononuclear cells: stimulation by interferon-gamma as part of an immunomodulatory pathway. J. Interferon Res. 8, 705-716
102. Iida, Y., Sawabe, K., Kojima, M., Oguro, K., Nakanishi, N., and Hasegawa, H. (2002) Proteasome-driven turnover of tryptophan hydroxylase is triggered by phosphorylation in RBL2H3 cells, a serotonin producing mast cell line. Eur. J. Biochem. 269, 4780-4788
103. Johansson, O., Liu, P. Y., Bondesson, L., Nordlind, K., Olsson, M. J., Lontz, W., Verhofstad, A., Liang, Y., and Gangi, S. (1998) A serotonin-like immunoreactivity is present in human cutaneous melanocytes. J. Invest. Dermatol. 111, 1010-1014
104. Semak, I., Korik, E., Naumova, M., Wortsman, J., and Slominski, A. (2004) Serotonin metabolism in rat skin: characterization by liquid chromatography-mass spectrometry. Arch. Biochem. Biophys. 421, 61-66
105. Slominski, A., Pisarchik, A., Semak, I., Sweatman, T., and Wortsman, J. (2003) Characterization of the serotoninergic system in the C57BL/6 mouse skin. Eur. J. Biochem. 270, 3335-3344
106. Coon, S. L., Weller, J. L., Korf, H. W., Namboodiri, M. A., Rollag, M., and Klein, D. C. (2001) cAmp regulation of arylalkylamine N-acetyltransferase (AANAT, EC 2.3.1.87): a new cell line (1E7) provides evidence of intracellular AANAT activation. J. Biol. Chem. 276, 24097-24107
107. Ferry, G., Loynel, A., Kucharczyk, N., Bertin, S., Rodriguez, M., Delagrange, P., Galizzi, J. P., Jacoby, E., Volland, J. P., Lesieur, D., et al. (2000) Substrate specificity and inhibition studies of human serotonin N-acetyltransferase. J. Biol. Chem. 275, 8794-8805
108. Slominski, A., Semak, I., Pisarchik, A., Sweatman, T., Szczesniewski, A., and Worstman. J. (2002) Conversion of L-tryptophan to serotonin and melatonin in melanoma cells. FEBS Lett. 511, 102-106
109. Cahill, G. M., and Besharse, J. C. (1989) Retinal melatonin is metabolized within the eye of Xenopus laevis. Proc. Natl. Acad. Sci. USA 86, 1098-1102
110. Slominski, A., Pisarchik, A., Scmak, I., Sweatman, T., Szczesniewski, A., and Wortsman, J. (2002) Serotoninergic system in hamster skin. J. Invest. Dermatol. 119, 934-942
111. Kojima, M., Oguro, K., Sawabe, K., Iida, Y., Ikeda, R., Yamashita, A., Nakanishi, N., and Hasegawa, H. (2000) Rapid turnover of tryptophan hydroxylase is driven by proteasomes in RBL2H3 cells, a serotonin producing mast cell line. J. Biochem. (Tokyo) 127, 121-127
112. Theoharides, T. C. (1996) The mast cell: a neuroendocrine master player. Int. J. Tissue React. 18, 1-21
113. Tachibana, T., Taniguchi, S., Furukawa, F., Miwa, S., and Imamura, S. (1990) Serotonin metabolism in the arthus reaction. J. Invest. Dermatol. 94, 120-125
114. Khalil, E. M., De Angelis, J., and Cole, P. A. (1998) Indoleamine analogs as probes of the substrate selectivity and catalytic mechanism of serotonin N-acetyltransferase. J. Biol. Chem. 273, 30321-30327
115. Hickman, A. B., Namboodiri, M. A., Klein, D. C., and Dyda, F. (1999) The structural basis of ordered substrate binding by serotonin N-acetyltransferase: enzyme complex at 1.8 A resolution with a bisubstrate analog. Cell 97, 361-369
116. Gaudet, S. J., Slominski, A., Etminan, M., Pruski, D., Paus, R., and Namboordiri, M. A. A. (1993) Identification and characterization of two isozymic forms of arylamine N-acetyltransferase in Syrian hamster skin. J. Invest. Dermatol. 101, 660-665
117. Slominski, A., Baker, J., Rosano, T., Guisti, L. W., Ermak, G., Grande, M., and Gaudet, S. J. (1996) Metabolism of serotonin to N-acetylserotonin, melatonin, and 5-methoxytryptamine in hamster skin culture. J. Biol. Chem. 271, 12281-12286
118. Roseboom, P. H., Namboodiri, M. A., Zimonjic, D. B., Popescu, N. C., Rodriguez, I. R., Gastel, J. A., and Klein, D. C. (1998) Natural melatonin 'knockdown' in C57BL/6J mice: rare mechanism truncates serotonin N-acetyltransferase. Mol. Brain Res. 63, 189-197
119. Sadrieh, N., Davis, C. D., and Snyderwine, E. G. (1996) N-acetyltransferase expression and metabolic activation of the food-derived heterocyclic amines in the human mammary gland. Cancer Res. 56, 2683-2687
120. Kawakubo, Y., Merk, H. F., Masaoudi, T. A., Sieben, S., and Blomeke, B. (2000) N-Acetylation of paraphenylenediamine in human skin and keratinocytes. J. Pharmacol. Exp. Ther. 292, 150-155
121. Williams, J. A., and Phillips, D. H. (2000) Mammary expression of xenobiotic metabolizing enzymes and their potential role in breast cancer. Cancer Res. 60, 4667-4677
122. Siraki, A. G., and O'Brien, P. J. (2002) Prooxidant activity of free radicals derived from phenol-containing neurotransmitters. Toxicology 177, 81-90
123. Silva, S. O., Ximenes, V. F., Catalani, L, H., and Campa, A. (2000) Myeloperoxidase-catalyzed oxidation of melatonin by activated neutrophils. Biochem. Biophys. Res. Commun. 279, 657-662
124. Lundeberg, L., El-Nour, H., Mohabbati, S., Morales, M., Azmitia, E., and Nordlind, K. (2002) Expression of serotonin receptors in allergic contact eczematous human skin. Arch. Dermatol. Res. 294, 393-398
125. Slominski, A., Pisarchik, A., Zbytek, B., Tobin, D. J., Kauser, S., and Wortsman, J. (2003) Functional activity of serotoninergic and melatoninergic systems expressed in the skin. J. Cell. Physiol. 196, 144-153
126. Weisshaar, E., Ziethen, B., and Gollnick, H. (1997) Can a serotonin type 3 (5-HT3) receptor antagonist reduce experimentally-induced itch? Inflamm. Res. 46, 412-416
127. McEwan, M., and Parsons, P. G. (1987) Inhibition of melanization in human melanoma cells by a serotonin uptake inhibitor. J. Invest. Dermatol. 89, 82-86
128. Belmonte, C., and Cervero, F. (1996) Neurobiology of Nociceptors, Oxford University Press, New York
129. Fitzpatrick, T. B., Eisen, A. Z., Wolff, K., Freedberg, I. M., and Austen, K. F. (1997) Dermatology in General Medicine, McGraw-Hill, New York
130. Symoens, J. (1990) Ketanserin: a novel cardiovascular drug. Blood Coagul. Fibrinolysis 1, 219-224
131. Balaskas, E. V., Bamihas, G. I., Karamouzis, M., Voyiatzis, G., and Tourkantonis, A. (1998) Histamine and serotonin in uremic pruritus: effect of ondansetron in CAPD-pruritic patients. Nephron 78, 395-402
132. Hagermark, O. (1992) Peripheral and central mediators of itch. Skin Pharmacol. 5, 1-8
133. Kam, P. C., and Tan, K. H. (1996) Pruritus-itching for a cause and relief? Anaesthesia 51, 1133-1138
134. Ashkenazy-Shahar, M., and Beitner, R. (1997) Serotonin decreases cytoskeletal and cytosolic glycolytic enzymes and the levels of ATP and glucose 1,6-bisphosphate in skin, which is prevented by the calmodulin antagonists thioridazine and clotrimazole. Biochem. Mol. Med. 60, 187-193
135. Colditz, I. G. (1991) The induction of plasma leakage in skin by histamine, bradykinin, activated complement, platelet-activating factor and serotonin. Immunol. Cell Biol. 69, 215-219
136. Fujii, E., Irie, K., Uchida, Y., Tsukahara, F., and Muraki, T. (1994) Possible role of nitric oxide in 5-hydroxytryptamine-induced increase in vascular permeability in mouse skin. Naunyn Schmiedebergs Arch. Pharmacol 350, 361-364
137. Khalil, Z., and Helme, R. D. (1990) Serotonin modulates substance P-induced plasma extravasation and vasodilatation in rat skin by an action through capsaicin-sensitive primary afferent nerves. Brain Res. 527, 292-298
138. Paus, R., Maurer, M., Slominski, A., and Czarnetzki, B. M. (1994) Mast cell involvement in murine hair growth. Dev. Biol. 163, 230-240
139. English, K. B., Wang, Z. Z., Stayner, N., Stensaas, L. J., Martin, H., and Tuckett, R. P. (1992) Serotonin-like immunoreactivity in Merkel cells and their afferent neurons in touch domes from the hairy skin of rats. Anat. Rec. 232, 112-120
140. Garcia-Caballero, T., Gallego, R., Roson, E., Basanta, D., Morel, G., and Beiras, A. (1989) Localization of serotonin-like immunoreactivity in the Merkel cells of pig snout skin. Anal. Rec. 225, 267-271
141. Maurer, M., Opitz, M., Henz, B. M., and Paus, R. (1997) The mast cell products histamine and serotonin stimulate and TNF-alpha inhibits the proliferation of murine epidermal keratinocytes in situ. J. Dermatol. Sci. 16, 79-84
142. Slominski, A., and Pruski, D. (1993) Melatonin inhibits proliferation and melanogenesis in rodent melanoma cells. Exp. Cell Res. 206, 189-194
143. Carlton, S. M., and Coggeshall, R. E. (1997) Immunohistochemical localization of 5-HT2A receptors in peripheral sensory axons in rat glabrous skin. Brain Res. 763, 271-275
144. Branchek, T. A., Mawe, G. M., and Gershon, M. D. (1988) Characterization and localization of a peripheral neural 5-hydroxytryptamine receptor subtype (5-HT1P) with a selective agonist, 3H-5-hydroxyindalpine. J. Neurosci. 8, 2582-2595
145. Hadley, M. (1996) Endocrinology, Prentice Hall, Englewood Cliffs, New Jersey
146. Filadelfi, A. M., and Castrucci, A. M. (1994) Melatonin desensitizing effects on the in vitro responses to MCH, alpha-MSH, isoproterenol and melatonin in pigment cells of a fish (S. marmoratus), a toad (B. ictericus), a frog (R. pipiens), and a lizard (A. carolinensis), exposed to varying photoperiodic regimens. Comp. Biochem. Physiol. A Physiol. 109, 1027-1037
147. Weatherhead, B. (1982) The pineal gland and pigmentation. In The Physiology and Pathophysiology of the Skin (Jarrett, A., ed) Vol. 7, pp. 2165-2179, Academic Press, NY
148. Lerchl, A., and Schlatt, S. (1993) Influence of photoperiod on pineal melatonin synthesis, fur color, body weight, and reproductive function in the female Djungarian hamster, Phodopus sungorus. Neuroendocrinology 57, 359-364
149. Allain, D., Thebault, R. G., Rougeot, J., and Martinet, L. (1994) Biology of fibre growth in mammals producing fine fibre and fur in relation to control by day length: relationship with other seasonal functions. Eur. Fine Fibre Network 2, 23-39
150. Xiao, Y., Forsberg, M., Laitinen, J. T., and Valtonen, M. (1995) Effects of melatonin implants in spring on testicular regression and moulting in adult male raccoon dogs (Nyctereutes procynoides). J. Reprod. Fertil. 105, 9-15
151. Nixon, A. J., Choy, V. J., Parry, A. L., and Pearson, A. J. (1993) Fiber growth initiation in hair follicles of goats treated with melatonin. J. Exp. Zool. 267, 47-56
152. Allain, D., and Rougeot, J. (1980) Induction of autumn moult in mink (Mustela vison Peale and Beauvois) with melatonin. Reprod. Nutr. Dev. 20, 197-201
153. Rose, J., Oldfield, J., and Stormshak, F. (1987) Apparent role of melatonin and prolactin in initiating winter fur growth in mink. Gen. Comp. Endocrinol. 65, 212-215
154. McCloghry, E., Foldes, A., Hollis, D., Rintoul, A., Maxwell, C., Downing, J., Baker, P., Kennedy, J., and Wynn, P. (1992) Effects of pinealectomy on wool growth and wool follicle density in merino sheep. J. Pineal Res. 13, 139-144
155. Gurlek, A., Aydogan, H., Parlakpinar, H., Bay-Karabulut, A., Celik, M., Sezgin, N., and Acet, A. (2004) Protective effect of melatonin on random pattern skin flap necrosis in pinealectomized rat. J. Pineal Res. 36, 58-63
156. Kumar, C. A., and Das, U. N. (2000) Effect of melatonin on two stage skin carcinogenesis in Swiss mice. Med. Sci. Monit. 6, 471-475
157. Logan, A., and Weatherhead, B. (1980) Post-tyrosinase inhibition of melanogenesis by melatonin in hair follicles in vitro. J. Invest. Dermatol. 74, 47-50
158. Weatherhead, B., and Logan, A. (1981) Interaction of alpha-melanocyte- stimulating hormone, melatonin, cyclic AMP and cyclic GMP in the control of melanogenesis in hair follicle melanocytes in vitro. J. Endocrinol. 90, 89-96
159. Slominski, A., Chassalerris, N., Mazurkiewicz, J., and Paus, R. (1994) Murine skin as a target for melatonin bioregulation. Exp. Dermatol. 3, 45-50
160. Valverde, P., Benedito, E., Solano, F., Oaknin, S., Lozano, J. A., and Garcia-Borron, J. C. (1995) Melatonin antagonizes alpha-melanocyte-stimulating hormone enhancement of melanogenesis in mouse melanoma cells by blocking the hormone-induced accumulation of the c locus tyrosinase. Eur. J. Biochem. 232, 257-263
161. Kadekaro, A. L., Andrade, L. N., Floeter-Winter, L. M., Rollag, M. D., Virador, V., Vieira, W., and Castrucci, A. M. (2004) MT-1 melatonin receptor expression increases the antiproliferative effect of melatonin on S-91 murine melanoma cells. J. Pineal Res. 36, 204-211
162. Ibraheem, M., Galbraith, H., Scaife, J., and Ewen, S. (1994) Growth of secondary hair follicles of the Cashmere goat in vitro and their response to prolactin and melatonin. J. Anat. 185, 135-142
163. Drobnik, J., and Dabrowski, R. (1999) Pinealectomy-induced elevation of collagen content in the intact skin is suppressed by melatonin application. Cytobios 100, 49-55
164. Drobnik, J., and Dabrowski, R. (1996) Melatonin suppresses the pinealectomy-induced elevation of collagen content in a wound. Cytobios 85, 51-58
165. Dicks, P., Morgan, C. J., Morgan, P. J., Kelly, D., and Williams, L. M. (1996) The localisation and characterisation of insulin-like growth factor-I receptors and the investigation of melatonin receptors on the hair follicles of seasonal and non-seasonal fibre-producing goats. J. Endocrinol. 151, 55-63
166. Shui-Wang, Y., Niles, L. P., and Crocker, C. (1993) Human malignant melanoma cells express high-affinity receptors for melatonin: antiproliferative effects of melatonin and 6-chloromelatonin. Eur. J. Pharmacol. 246, 89-96
167. Eison, A. S., and Mullins, U. L. (1993) Melatonin binding sites are functionally coupled to phosphoinositide hydrolysis in Syrian hamster RPMI 1846 melanoma cells. Life Sci. 53, PL393-PL398
168. Ying, S. W., Niles, L. P., and Crocker, C. (1993) Human malignant melanoma cells express high-affinity receptors for melatonin: antiproliferative effects of melatonin and 6-chloromelatonin. Eur. J. Pharmacol. 246, 89-96
169. Nash, M. S., and Osborne, N. N. (1995) Pertussis toxin-sensitive melatonin receptors negatively coupled to adenylate cyclase associated with cultured human and rat retinal pigment epithelial cells. Invest. Ophthalmol. Vis. Sci. 36, 95-102
170. Pawelek, J., Fleischman, R. A., McLane, J., Guillette, B., Emanuel, J. R., Korner, A., Bergstrom, A., and Murray, M. (1985) Studies on growth and pigmentation of cloudman S91 melanoma cells. In Biological, molecular and clinical aspects of pigmentation (Bagnara, J. T., Klaus, S. N., Paul, E., and Schartl, M., eds) pp. 521-533, University of Tokyo Press, Tokyo
171. Roberts, J. E., Wiechmann, A. F., and Hu, D.-N. (2000) Melatonin receptors in human uveal melanocytes and melanoma cells. J. Pineal Res. 28, 165-171
172. Mengeaud, V., Skene, D., Pevet, P., and Ortonne, J. P. (1994) No high affinity melatonin binding sites are detected in murine melanoma cells and in normal human melanocytes cultured in vitro. Melanoma Res. 4, 87-91
173. Bartsch, M., Bartsch, C., and Flehmig, B. (1986) Differential effect of melatonin on slow and fast growing passages of a human melanoma cell line. Neuroendocrinol. Lett. 8, 289-293
174. Meyskens, F. L., Jr., and Salmon, S. E. (1981) Modulation of clonogenic human melanoma cells by follicle-stimulating hormone, melatonin, and nerve growth factor. Br. J. Cancer 43, 111-115
175. Hu, D.-N., McCormick, S. A., and Roberts, J. E. (1998) Effects of melatonin, its precursors and derivatives on the growth of cultured human uveal melanoma cells. Melanoma Res. 8, 205-210
176. Cos, S., and Blask, D. E. (1990) Effects of the pineal hormone melatonin on the anchorage-independent growth of human breast cancer cells (MCF-7) in a clonogenic culture system. Cancer Lett. 50, 115-119
177. Iyengar, B. (2000) Melatonin and melanocyte functions. Biol. Signals Recept. 9, 260-266
178. Kim, B. C., Shon, B. S., Ryoo, Y. W., Kim, S. P., and Lee, K. S. (2001) Melatonin reduces X-ray irradiation-induced oxidative damages in cultured human skin fibroblasts. J. Dermatol. Sci. 26, 194-200
179. Ryoo, Y. W., Suh, S. I., Mun, K. C., Kim, B. C., and Lee, K. S. (2001) The effects of the melatonin on ultraviolet-B irradiated cultured dermal fibroblasts. J. Dermatol. Sci. 27, 162-169
180. Nickel, A., and Wohlrab, W. (2000) Melatonin protects human keratinocytes from UVB irradiation by light absorption. Arch. Dermatol. Res. 292, 366-368
181. Lee, K. S., Lee, W. S., Suh, S. I., Kim, S. P., Lee, S. R., Ryoo, Y. W., and Kim, B. C. (2003) Melatonin reduces ultraviolet-B induced cell damages and polyamine levels in human skin fibroblasts in culture. Exp. Mol. Med. 35, 263-268
182. Kilanczyk, E., and Bryszewska, M. (2003) The effect of melatonin on antioxidant enzymes in human diabetic skin fibroblasts. Cell. Mol. Biol. Lett. 8, 333-336
183. Mozzanica, N., Tadini, G., Radaelli, A., Negri, M., Pigatto, P., Morelli, M., Frigerio, U., Finzi, A., Esposti, G., Rossi, D., et al. (1988) Plasma melatonin levels in psoriasis. Acta Derm. Venereol. 68, 312-316
184. Schwarz, W., Birau, N., Hornstein, O. P., Heubeck, B., Schonberger, A., Meyer, C., and Gottschalk, J. (1988) Alterations of melatonin secretion in atopic eczema. Acta Derm. Venereol. 68, 224-229
185. Maietta, G., Rongioletti, F., and Rebora, A. (1991) Seborrheic dermatitis and daylight. Acta Derm. Venereol. 71, 538-539
186. Bangha, E., Elsner, P., and Kistler, G. S. (1997) Suppression of UV-induced erythema by topical treatment with melatonin (N-acetyl-5- methoxytryptamine). Influence of the application time point. Dermatology 195, 248-252
187. Dreher, F., Gabard, B., Schwindt, D. A., and Maibach, H. I. (1998) Topical melatonin in combination with vitamins E and C protects skin from ultraviolet-induced erythema: a human study in vivo. Br. J. Dermatol. 139, 332-339
188. Nordlund, J. J., and Lerner, A. B. (1977) The effects of oral melatonin on skin color and on the release of pituitary hormones. J. Clin. Endocrinol. Metab. 45, 768-774
189. McElhinney, D. B., Hoffman, S. J., Robinson, W. A., and Ferguson, J. (1994) Effect of melatonin on human skin color. J. Invest. Dermatol. 102, 258-259
190. Bangha, E., Lauth, D., Kistler, G. S., and Elsner, P. (1997) Daytime serum levels of melatonin after topical application onto the human skin. Skin Pharmacol. 10, 298-302
191. Fischer, T. W., Burmeister, G., Schmidt, H. W., and Elsner, P. (2004) Melatonin increases anagen hair rate in women with androgenetic alopecia or diffuse alopecia: results of a pilot randomized controlled trial. Br. J. Dermatol. 150, 341-345
192. Bardak, Y., Ozerturk, Y., Ozguner, F., Durmus, M., and Delibas, N. (2000) Effect of melatonin against oxidative stress in ultraviolet-B exposed rat lens. Curr. Eye Res. 20, 225-230
193. Gambichler, T., Bader, A., Vojvodic, M., Bechara, F. G., Sauermann, K., Altmeyer, P., and Hoffmann, K. (2002) Impact of UVA exposure on psychological parameters and circulating serotonin and melatonin. BMC Dermatol. 2, 6