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melanin natural pigment melanogenesis oxidation tyrosine polymerization melanocytes radiation eumelanin pheomelanin neuromelanin allomelanin pyomelanin parkinson' malignant melanoma cancer melanin cells skin tone color iris eye stria vascularis ear medulla pigment-bearing neurons brainstem locus coeruleus zona reticularis adrenal gland basal layer epidermis albinism trichochromes metabolic pathway catecholaminergic apoptosis parkinson's disease cephalopods biosynthetic pathways keratinocyte melanosomes cellular vesicles ultraviolet rays 3d