Melanin - Knowledge

74: 1147: 1218:, which is mostly an autosomal recessive disorder. Certain ethnicities have higher incidences of different forms. For example, the most common type, called oculocutaneous albinism type 2 (OCA2), is especially frequent among people of black African descent and white Europeans. People with OCA2 usually have fair skin, but are often not as pale as OCA1. They (OCA2 or OCA1? see comments in History) have pale blonde to golden, strawberry blonde, or even brown hair, and most commonly blue eyes. 98.7–100% of modern Europeans are carriers of the derived allele 848: 453:. This selenium analogue of pheomelanin has been successfully synthesized through chemical and biosynthetic routes using selenocystine as a feedstock. Due to selenium's higher atomic number, the obtained selenomelanin can be expected to provide better protection against ionising radiation as compared to the other known forms of melanin. This protection has been demonstrated with radiation experiments on human cells and bacteria, opening up the possibility of applications in space travel. 62: 474: 1330:. Effective in treating white skin, in general, lasers are less successful in removing port-wine stains in people of Asian or African descent. Higher concentrations of melanin in darker-skinned individuals simply diffuse and absorb the laser radiation, inhibiting light absorption by the targeted tissue. In a similar manner, melanin can complicate laser treatment of other dermatological conditions in people with darker skin. 691:, convert resources to melanin to enhance their thermoregulation. As the wood tiger moth has populations over a large range of latitudes, it has been observed that more northern populations showed higher rates of melanization. In both yellow and white male phenotypes of the wood tiger moth, individuals with more melanin had a heightened ability to trap heat but an increased predation rate due to a weaker and less effective 1171: 785: 1451:, and possibly others). The first modern humans had darker skin as the indigenous people of Africa today. Following migration and settlement in Asia and Europe, the selective pressure dark UV-radiation protecting skin decreased where radiation from the sun was less intense. This resulted in the current range of human skin color. Of the two common gene variants known to be associated with pale human skin, 839: 412: 42: 1135: 539:. Although, in general, human beings possess a similar concentration of melanocytes in their skin, the melanocytes in some individuals and ethnic groups produce variable amounts of melanin. The ratio of eumelanin (74%) and pheomelanin (26%) in the epidermis is constant regardless of the degree of pigmentation. Some humans have very little or no melanin synthesis in their bodies, a condition known as 1159: 7094: 1183: 421: 320: 336:. Pheomelanin is also present in the skin, and redheads consequently often have a more pinkish hue to their skin as well. Exposure of the skin to ultraviolet light increases pheomelanin content, as it does for eumelanin; but rather than absorbing light, pheomelanin within the hair and skin reflect yellow to red light, which may increase damage from UV radiation exposure. 242:, causing the skin to darken. Eumelanin is an effective absorbent of light; the pigment is able to dissipate over 99.9% of absorbed UV radiation. Because of this property, eumelanin is thought to protect skin cells from UVA and UVB radiation damage, reducing the risk of folate depletion and dermal degradation. Exposure to UV radiation is associated with increased risk of 264: 601:. In invertebrates, a major aspect of the innate immune defense system against invading pathogens involves melanin. Within minutes after infection, the microbe is encapsulated within melanin (melanization), and the generation of free radical byproducts during the formation of this capsule is thought to aid in killing them. Some types of fungi, called 7106: 828:. This can be thought of as a single monomer that accounts for the measured elemental composition and some properties of melanin, but is unlikely to be found in nature. Solano claims that this misleading trend stems from a report of an empirical formula in 1948, but provides no other historical detail. 436:

that form a 'corona', that are responsible for the substance's solubility. The peptide chains are linked to the L-DOPA core polymer via peptide bonds. This lead to a proposed biosynthetic process involving the hydroxylation of tyrosinylated peptides formed via proteases during sporogenesis, which are

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As with peoples having migrated northward, those with light skin migrating toward the equator acclimatize to the much stronger solar radiation. Nature selects for less melanin when ultraviolet radiation is weak. Most people's skin darkens when exposed to UV light, giving them more protection when it

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metal ions through its carboxylate and phenolic hydroxyl groups, often much more efficiently than the powerful chelating ligand ethylenediaminetetraacetate (EDTA). Thus, it may serve to sequester potentially toxic metal ions, protecting the rest of the cell. This hypothesis is supported by the fact

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Pheomelanins (or phaeomelanins) impart a range of yellowish to reddish colors. Pheomelanins are particularly concentrated in the lips, nipples, glans of the penis, and vagina. When a small amount of eumelanin in hair (which would otherwise cause blond hair) is mixed with pheomelanin, the result is

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Not all the effects of pigmentation are advantageous. Pigmentation increases the heat load in hot climates, and dark-skinned people absorb 30% more heat from sunlight than do very light-skinned people, although this factor may be offset by more profuse sweating. In cold climates dark skin entails

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reduction or absence of melanin pigment in the skin, hair, and eyes. The estimated frequency of OCA2 among African-Americans is 1 in 10,000, which contrasts with a frequency of 1 in 36,000 in white Americans. In some African nations, the frequency of the disorder is even higher, ranging from 1 in

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strands that form feathers. Pheomelanin synthesis in birds implies the consumption of cysteine, a semi‐essential amino acid that is necessary for the synthesis of the antioxidant glutathione (GSH) but that may be toxic if in excess in the diet. Indeed, many carnivorous birds, which have a high

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Because melanin is an aggregate of smaller component molecules, there are many different types of melanin with different proportions and bonding patterns of these component molecules. Both pheomelanin and eumelanin are found in human skin and hair, but eumelanin is the most abundant melanin in

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In natural conditions, DHI and DHICA often co-polymerize, resulting in a range of eumelanin polymers. These polymers contribute to the variety of melanin components in human skin and hair, ranging from light yellow/red pheomelanin to light brown DHICA-enriched eumelanin and dark brown or black

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synthesis. While no correlation between race and the level of neuromelanin in the substantia nigra has been reported, the significantly lower incidence of Parkinson's in blacks than in whites has "prompt some to suggest that cutaneous melanin might somehow serve to protect the neuromelanin in

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Ocular albinism affects not only eye pigmentation but visual acuity, as well. People with albinism typically test poorly, within the 20/60 to 20/400 range. In addition, two forms of albinism, with approximately 1 in 2,700 most prevalent among people of Puerto Rican origin, are associated with

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Up until the 1960s, melanin was classified into eumelanin and pheomelanin. However in 1955 a melanin associated with nerve cells was discovered, neuromelanin. In 1972 a water-soluble form, pyomelanin was discovered. In 1976, allomelanin, the fifth form of the melanins was found in nature.

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availability, which is transported into the melanosome, reacting with dopaquinone to form cys-dopa. Cys-dopa then undergoes several transformations before forming pheomelanin. In chemical terms, pheomelanins differ from eumelanins in that the oligomer structure incorporates

1415:. Since pigmentation appears to be not entirely advantageous to life in the tropics, other hypotheses about its biological significance have been advanced; for example a secondary phenomenon induced by adaptation to parasites and tropical diseases. 1260:

in North America. The incidence of albinism in Hopi Indians has been estimated as approximately 1 in 200 individuals. Similar patterns of albinism and deafness have been found in other mammals, including dogs and rodents. However, a lack of melanin

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in the brain. Humans have the largest amount of NM, which is present in lesser amounts in other primates, and totally absent in many other species. The biological function remains unknown, although human NM has been shown to efficiently bind

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evolved dark skin color, as an adaptation to a loss of body hair that increased the effects of UV radiation. Before the development of hairlessness, early humans might have had light skin underneath their fur, similar to that found in other

283:(DHICA). DHI-derived eumelanin is dark brown or black and insoluble, and DHICA -derived eumelanin which is lighter and soluble in alkali. Both eumelanins arise from the oxidation of tyrosine in specialized organelles called 306:

V and VI) skin finds that DHI-eumelanin comprises the largest portion, approximately 60–70%, followed by DHICA-eumelanin at 25–35%, and pheomelanin only 2–8%. Notably, while an enrichment of DHI-eumelanin occurs in during

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Alaluf S, Heath A, Carter N, Atkins D, Mahalingam H, Barrett K, et al. (2001). "Variation in Melanin Content and Composition in Type V and VI Photoexposed and Photoprotected Human Skin: The Dominant Role of DHI".

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does not appear to be directly responsible for deafness associated with hypopigmentation, as most individuals lacking the enzymes required to synthesize melanin have normal auditory function. Instead, the absence of

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functioning, there is decreased neuromelanin in the substantia nigra and locus coeruleus as a consequence of specific dropping out of dopaminergic and noradrenergic pigmented neurons. This results in diminished

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Millar SE, Miller MW, Stevens ME, Barsh GS (October 1995). "Expression and transgenic studies of the mouse agouti gene provide insight into the mechanisms by which mammalian coat color patterns are generated".

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In addition to melanin deficiency, the molecular weight of the melanin polymer may be decreased by various factors such as oxidative stress, exposure to light, perturbation in its association with melanosomal

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SEM micrograph of Aspergillus niger (strain: melanoliber) conodiophore possessing a large number of small conidospores (colorized). These spores release peptidomelanin into the surrounding medium during

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of melanoma cells is crucial to metastasis and growth; non-pigmented tumors were larger than pigmented tumors, and spread far more easily. Pigmented and non-pigmented cells are both present in melanoma

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Greco G, Panzella L, Verotta L, d'Ischia M, Napolitano A (April 2011). "Uncovering the structure of human red hair pheomelanin: benzothiazolylthiazinodihydroisoquinolines as key building blocks".

1231:, who are of primarily Swiss and German ancestry. People with this IB variant of the disorder commonly have white hair and skin at birth, but rapidly develop normal skin pigmentation in infancy. 2649: 1531:

are more at risk of sun-related eye problems. Furthermore, the ocular lens yellows with age, providing added protection. However, the lens also becomes more rigid with age, losing most of its

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has an affinity for melanin-containing tissues because of its precursor function in melanin synthesis or its irreversible binding of melanin. This has been suggested to underlie the increased

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Fedorow H, Tribl F, Halliday G, Gerlach M, Riederer P, Double KL (2005). "Neuromelanin in human dopamine neurons: comparison with peripheral melanins and relevance to Parkinson's disease".

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It is now understood that melanins do not have a single structure or stoichiometry. Nonetheless, chemical databases such as PubChem include structural and empirical formulae; typically

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Pheomelanins, unlike euemanins, are rare in lower organisms with claims they are an "evolutionary innovation in the tetrapod lineage" but recent research finds them also in some fish.

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have larger quantities of eumelanin in their skins. This makes their skins brown or black and protects them against high levels of exposure to the sun, which more frequently result in

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Meier-Davis SR, Dines K, Arjmand FM, et al. (December 2012). "Comparison of oral and transdermal administration of rasagiline mesylate on human melanoma tumor growth in vivo".

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tint given to some skin or hair colors. Neuromelanin is found in the brain. Research has been undertaken to investigate its efficacy in treating neurodegenerative disorders such as

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which regulates the distribution of melanin. The mechanisms of the gene have been extensively studied in mice to provide an insight into the diversity of mammalian coat patterns.

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can transform into either 5,6-dihydroxyindole (DHI) or 5,6-dihydroxyindole-2-carboxylic acid (DHICA). DHI and DHICA are oxidized and then polymerize to form the two eumelanins.

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Higher eumelanin levels also can be a disadvantage, however, beyond a higher disposition toward vitamin D deficiency. Dark skin is a complicating factor in the laser removal of

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Mosse IB, Dubovic BV, Plotnikova SI, Kostrova LN, Molophei V, Subbot ST, et al. (20–25 May 2001). Obelic B, Ranogajev-Komor M, Miljanic S, Krajcar Bronic I (eds.).

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evolved in Africa between 200,000 and 100,000 years ago, and then populated the rest of the world through migration between 80,000 and 50,000 years ago, in some areas

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Melanin is brown, non-refractile, and finely granular with individual granules having a diameter of less than 800 nanometers. This differentiates melanin from common

3444: 3348: 5530: 215:. Eumelanin is the most common type. Pheomelanin, which is produced when melanocytes are malfunctioning due to derivation of the gene to its recessive format, is a 3108:

Gunderson AR, Frame AM, Swaddle JP, Forsyth MH (1 September 2008). "Resistance of melanized feathers to bacterial degradation: is it really so black and white?".

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pigments. Feathers that contain melanin are also 39% more resistant to abrasion than those that do not because melanin granules help fill the space between the

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Berth-Jones, J. (2010). "Constitutive pigmentation, human pigmentation and the response to sun exposure". In Burns T, Breathnach S, Cox N, Griffiths C (eds.).

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King G, Yerger VB, Whembolua GL, Bendel RB, Kittles R, Moolchan ET (June 2009). "Link between facultative melanin and tobacco use among African Americans".

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Cable J, Huszar D, Jaenisch R, Steel KP (February 1994). "Effects of mutations at the W locus (c-kit) on inner ear pigmentation and function in the mouse".

6634: 5974: 5336: 246:, a cancer of melanocytes (melanin cells). Studies have shown a lower incidence for skin cancer in individuals with more concentrated melanin, i.e. darker 73: 5964: 5920: 5770: 5435: 3965: 1436: 1252:

observed that "cats which are entirely white and have blue eyes are generally deaf". In humans, hypopigmentation and deafness occur together in the rare

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are one of the few groups in which melanin has not been easily detected, though researchers found data suggesting spiders do in fact produce melanin.

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Lamason RL, Mohideen MA, Mest JR, et al. (December 2005). "SLC24A5, a putative cation exchanger, affects pigmentation in zebrafish and humans".

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Rodríguez-Martínez S, Galván I (2020). "Juvenile pheomelanin-based plumage coloration has evolved more frequently in carnivorous species".

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of fruits such as bananas. Chestnut shell melanin can be used as an antioxidant and coloring agent. Biosynthesis involves the oxidation of

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and the development of melanoma, a potentially deadly form of skin cancer, as well as other health problems related to exposure to strong

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Ito S, Wakamatsu K (December 2011). "Diversity of human hair pigmentation as studied by chemical analysis of eumelanin and pheomelanin".

6140: 5737: 5460: 2848: 6846: 6766: 5940: 5430: 4495: 3547:"To quiver or to shiver: increased melanization benefits thermoregulation, but reduces warning signal efficacy in the wood tiger moth" 1691: 579:), and biochemical threats (such as host defenses against invading microbes). Therefore, in many pathogenic microbes (for example, in 5066:

Sarangarajan R, Apte SP (2005). "Melanin aggregation and polymerization: possible implications in age-related macular degeneration".

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Slominski A, Tobin DJ, Shibahara S, Wortsman J (October 2004). "Melanin pigmentation in mammalian skin and its hormonal regulation".

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leading to the formation of catechol melanin. Despite this many plants contain compounds which inhibit the production of melanins.

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Sköld HN, Aspengren S, Cheney KL, Wallin M (2016). "Chapter Four - Fish Chromatophores—From Molecular Motors to Animal Behavior".

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as the eumelanins and pheomelanins, but unlike those molecules they have low molecular weight. They occur in some red human hair.

6485: 5465: 2712: 6432: 1323:, an important monotherapy drug in Parkinson's disease, has melanin binding properties, and melanoma tumor reducing properties. 428:

Peptidomelanin is another water-soluble form of melanin. It was found to be secreted into the surrounding medium by germinating

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Liu Y, Hong L, Kempf VR, Wakamatsu K, Ito S, Simon JD (June 2004). "Ion-exchange and adsorption of Fe(III) by Sepia melanin".

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Recent research suggests that melanin may serve a protective role other than photoprotection. Melanin is able to effectively

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2,000 to 1 in 5,000. Another form of Albinism, the "yellow oculocutaneous albinism", appears to be more prevalent among the

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that the loss of neuromelanin, observed in Parkinson's disease, is accompanied by an increase in iron levels in the brain.

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may lead to formation of pro-oxidant melanin with lower-molecular-weight, implicated in the causation and progression of

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of birds owe their color to melanin and are less readily degraded by bacteria than unpigmented ones or those containing

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are formed where there is a localized concentration of melanin in the skin. They are highly associated with pale skin.

6885: 6088: 6039: 5506: 2217: 6905: 6571: 620:, melanin occurs not only in the skin but also in internal organs such as eyes. Most fish species use eumelanin, but 555:

Melanins have very diverse roles and functions in various organisms. A form of melanin makes up the ink used by many

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Sarangarajan R, Apte SP (2005). "Melanization and phagocytosis: implications for age related macular degeneration".

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Peracha MO, Cosgrove FM, Garcia-Valenzuela E, Eliott D (5 November 2015). Roy H Sr, Talavera F, Rowsey JJ (eds.).

3759:"Melanin-Like Nanomaterials for Advanced Biomedical Applications: A Versatile Platform with Extraordinary Promise" 1222:, a known cause of nonsyndromic oculocutaneous albinism. It is an autosomal recessive disorder characterized by a 7126: 6747: 6695: 6452: 6447: 6135: 6130: 6083: 6044: 5689: 4342: 1524: 1253: 6700: 6093: 5670: 1244: 6654: 3164: 4263:"Microbial Tyrosinases: Promising Enzymes for Pharmaceutical, Food Bioprocessing, and Environmental Industry" 3467: 136: 6761: 6397: 5824: 5799: 5727: 5709: 5470: 4805: 4189:"Melanins: Skin Pigments and Much More—Types, Structural Models, Biological Functions, and Formation Routes" 2983:"Ionizing radiation changes the electronic properties of melanin and enhances the growth of melanized fungi" 2279:"Melanins: Skin Pigments and Much More—Types, Structural Models, Biological Functions, and Formation Routes" 1215: 581: 568: 6788: 6644: 6490: 6211: 6004: 5857: 4658: 1311:. In its pro-oxidant state, melanin has been suggested to be involved in the causation and progression of 1203: 606: 391: 129: 3642: 1170: 670:

of alternating refractive index. When the scale of this pattern matches the wavelength of visible light,

432:(strain: melanoliber) spores. Peptidomelanin is formed as a copolymer between L-DOPA eumelanin and short 7131: 6327: 6306: 6261: 6206: 5570: 2624: 2158: 688: 671: 6900: 2736:"5,6-Dihydroxyindole eumelanin content in human skin with varying degrees of constitutive pigmentation" 5110: 4514: 6924: 6816: 6752: 6355: 5877: 5651: 5384: 5374: 5360: 5219: 4950: 4895: 4761: 3844: 3601:. IRPA Regional Congress on Radiation Protection in Central Europe: Radiation Protection and Health. 2994: 1730: 1632: 1616: 1580: 1476:. Dark-skinned people, who produce more skin-protecting eumelanin, have a greater protection against 1399:

rays. In general, people whose ancestors lived for long periods in the regions of the globe near the

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in the Soviet era, was increasingly supported by government funding in the wake of the discovery of

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such as iron, as well as other potentially toxic molecules. Therefore, it may play crucial roles in

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Mouchet SR, Cortesi F, Bokic B, Lazovic V, Vukusic P, Marshall NJ, et al. (1 November 2023).

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Micrograph of Melanin pigment (light refracting granular material—center of image) in a pigmented

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Cerenius L, Söderhäll K (April 2004). "The prophenoloxidase-activating system in invertebrates".

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Part of the structural formula of pheomelanin. "(COOH)" can be COOH or H, or (more rarely) other

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Double KL (2006). "Functional effects of neuromelanin and synthetic melanin in model systems".

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Part of the structural formula of eumelanin. "(COOH)" can be COOH or H, or (more rarely) other

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Melanin produced by plants are sometimes referred to as 'catechol melanins' as they can yield

462: 429: 383: 378: 303: 5111:"Etiologic pathogenesis of melanoma: a unifying hypothesis for the missing attributable risk" 1182: 908: 6710: 6273: 6180: 5245: 5227: 5125: 5075: 5040: 5005: 4968: 4958: 4903: 4858: 4850: 4777: 4769: 4624: 4581: 4529: 4469: 4284: 4274: 4233: 4200: 4148: 4140: 4099: 4089: 3978: 3922: 3852: 3792: 3776: 3763: 3726: 3710: 3646: 3566: 3558: 3517: 3401: 3303: 3254: 3246: 3209: 3199: 3148: 3117: 3088: 3049: 3012: 3002: 2963: 2920: 2883: 2840: 2812: 2804: 2763: 2747: 2694: 2684: 2576: 2543: 2527: 2484: 2441: 2396: 2326: 2290: 2248: 2201: 2124: 2073: 2030: 1985: 1977: 1930: 1883: 1873: 1804: 1796: 1738: 1670: 1572: 1485: 1380: 1053: 950: 705: 576: 536: 517: 489: 481: 87: 4499: 3829:

Mosse IB (2012). "Genetic effects of ionizing radiation – some questions with no answers".

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Kolipakala RS, Basu S, Sarkar S, Biju BM, Salazar D, Reddy L, et al. (7 August 2024).

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Meredith P, Riesz J (2004). "Radiative relaxation quantum yields for synthetic eumelanin".

1862:"Neuromelanin, one of the most overlooked molecules in modern medicine, is not a spectator" 7110: 7052: 6772: 6730: 6715: 6685: 6659: 6556: 6402: 5756: 5632: 5515: 4365: 3706: 3077:"Morphological and Optical Modification of Melanosomes in Fish Integuments upon Oxidation" 1641: 1612: 1481: 1327: 888: 721: 667: 628: 571:. Melanin also protects against damage from high temperatures, chemical stresses (such as 513: 505: 5031:

Donatien PD, Orlow SJ (August 1995). "Interaction of melanosomal proteins with melanin".

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Castelvecchi D (26 May 2007). "Dark Power: Pigment seems to put radiation to good use".

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Melanin is Effective Radioprotector against Chronic Irradiation and Low Radiation Doses

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Neuromelanin (NM) is an insoluble polymer pigment produced in specific populations of

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Harding RM, Healy E, Ray AJ, Ellis NS, Flanagan N, Todd C, et al. (April 2000).

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dopaquinone + cysteine → 2-S-cysteinyldopa → benzothiazine intermediate → pheomelanin

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dopaquinone + cysteine → 5-S-cysteinyldopa → benzothiazine intermediate → pheomelanin

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The first step of the biosynthetic pathway for both eumelanins and pheomelanins is

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Trichochromes (formerly called trichosiderins) are pigments produced from the same

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Cao W, Zhou X, McCallum NC, Hu Z, Ni QZ, Kapoor U, et al. (9 February 2021).

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of about 4.93 kPa, compared to non-pigmented cells, with a value of 0.98 kPa. The

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Sarna M, Krzykawska-Serda M, Jakubowska M, Zadlo A, Urbanska K (26 June 2019).

4078:"Comparison of Antioxidant Activities of Melanin Fractions from Chestnut Shell" 3371: 2888: 2871: 2188:

Thody AJ, Higgins EM, Wakamatsu K, Ito S, Burchill SA, Marks JM (August 1991).

2128: 1742: 784: 7037: 7008: 6954: 6690: 6509: 6417: 6350: 6289: 6265: 6189: 6165: 6034: 6012: 5994: 5984: 5955: 5804: 5497: 5388: 5352: 4628: 4366:"Ocular Manifestations of Albinism: Background, Pathophysiology, Epidemiology" 4144: 4094: 3651: 3628: 2488: 2015: 2013: 2011: 2009: 1501: 1497: 1493: 1448: 1372: 1320: 1223: 1104: 1057: 984: 899: 700: 675: 663: 640: 610: 556: 528: 357: 299:

DHI-enriched eumelanin. These final polymers differ in solubility and color.

288: 284: 205: 5241: 4722: 4046: 3990: 3936: 3864: 3788: 3722: 3714: 3660: 3268: 2759: 2689: 2588: 2563:

Cao W, McCallum NC, Ni QZ, Li W, Boyce H, Mao H, et al. (22 July 2020).

2539: 2531: 2520:"Fungal Peptidomelanin: A Novel Biopolymer for the Chelation of Heavy Metals" 2410: 2338: 2085: 2042: 1878: 1818: 1750: 724:

come from the laboratories and research groups of Irma Mosse. Mosse began in

6599: 6380: 6193: 6054: 5873: 5869: 5844: 5732: 5718: 5704: 5680: 4963: 4907: 4773: 4392: 3772: 3606: 3405: 2808: 1701: 1543: 1528: 1412: 733: 586: 509: 501: 420: 387: 247: 201: 24: 5259: 5189: 5139: 5087: 5017: 4982: 4915: 4872: 4791: 4636: 4593: 4543: 4298: 4247: 4162: 4113: 3998: 3872: 3806: 3780: 3757:

Liu H, Yang Y, Liu Y, Pan J, Wang J, Man F, et al. (7 February 2020).

3740: 3668: 3629:"Melanin pigments of fungi under extreme environmental conditions (Review)" 3580: 3562: 3531: 3223: 3061: 3026: 2932: 2897: 2826: 2777: 2708: 2596: 2496: 2453: 2418: 2385:"Genome sequence and genetic diversity of the common carp, Cyprinus carpio" 2346: 2260: 2189: 2136: 2093: 2050: 1999: 1942: 1897: 1826: 1809: 1758: 5052: 4481: 4279: 4205: 4188: 4063: 3413: 3188:"Bird Integumentary Melanins: Biosynthesis, Forms, Function and Evolution" 2383:

Xu P, Zhang X, Wang X, Li J, Liu G, Kuang Y, et al. (November 2014).

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occurs when melanocytes produce little melanin. This albino girl is from

446: 340: 333: 216: 208: 169:. Melanin pigments are produced in a specialized group of cells known as 166: 154: 66: 20: 3259: 6949: 6821: 6737: 6677: 6278: 5747: 5642: 5619: 5594: 5585: 5421: 3522: 3505: 3160: 3093: 3076: 2751: 2330: 1784: 1576: 1536: 1516: 1489: 1477: 1459: 1428: 1400: 1271: 1219: 1113: 997: 920: 644: 636: 450: 433: 263: 196:. Eumelanin is produced through a multistage chemical process known as 158: 41: 5079: 4346: 4182: 4180: 3250: 2791:

Cichorek M, Wachulska M, Stasiewicz A, Tymińska A (20 February 2013).

2252: 7059: 5911: 5598: 5397: 5364: 5303: 5284: 4750:"The genetic structure and history of Africans and African Americans" 4445: 4431: 1654: 1505: 1068: 652: 648:

protein content in their diet, exhibit pheomelanin‐based coloration.

547:

humans, as well as the form most likely to be deficient in albinism.

437:

then incorporated autoxidatively into a growing L-DOPA core polymer.

235:. Allomelanin and pyomelanin are two types of nitrogen free melanin. 228: 142: 3370:

Griffiths AJ, Miller JH, Suzuki DT, Lewontin RC, Gelbart WM (2000).

3152: 2401: 2384: 1785:"Unraveling the Structure and Function of Melanin through Synthesis" 1575:. Suboptimal conditions for the effective polymerization of melanin 1411:

more heat loss by radiation. Pigmentation also hinders synthesis of

1028:

Except where otherwise noted, data are given for materials in their

4854: 3963:

Mosse I (18 January 2022). "Radiobiology in my life – Irma Mosse".

3136: 1307:

has been proposed to turn the normally anti-oxidant polymer into a

1112:

dopaquinone → leucodopachrome → dopachrome → 5,6-dihydroxyindole →

838: 5775: 4806:"A Single Migration From Africa Populated the World, Studies Find" 4376: 1274:

impairment, though the reasons for this are not fully understood.

1228: 783: 472: 419: 410: 353: 318: 280: 262: 72: 60: 1367:

Melanocytes insert granules of melanin into specialized cellular

3903:

Mosse I, Kilchevsky A, Nikolova N, Zhelev N (14 December 2016).

3376: 2190:"Pheomelanin as well as eumelanin is present in human epidermis" 1966:"The protective role of melanin against UV damage in human skin" 1567:. It has been proposed that the ability of melanin to act as an 1453: 1257: 1121: 617: 492:. It is also found in hair, the pigmented tissue underlying the 6234: 5325: 4748:

Tishkoff SA, Reed FA, Friedlaender FR, et al. (May 2009).

2673:"Biochemical sites of gene action for melanogenesis in mammals" 708:

against DNA damage from non-UV radiation. Important studies in

655:

pigmentation. The coat pattern of mammals is determined by the

2066:

Journal of the European Academy of Dermatology and Venereology

1388: 1242:

is well known, though poorly understood. In his 1859 treatise

224: 4688:. Vol. 3 (8th ed.). Wiley-Blackwell. p. 58.9. 3290:

Jimbow K, Quevedo WC J, Fitzpatrick TB, Szabo G (July 1976).

1721:

Casadevall A (2018). "Melanin triggers antifungal defences".

1383:. The melanosomes in each recipient cell accumulate atop the 1107:→ 5,6-dihydroxyindole-2-carboxylic acid → quinone → eumelanin 1022:

450 to 550 °C (842 to 1,022 °F; 723 to 823 K)

666:

has been observed to be deposited in layers thus producing a

238:

In the human skin, melanogenesis is initiated by exposure to

3334:

Genetic Analysis: Genes, Genomes, and Networks in Eukaryotes

2565:"Selenomelanin: An Abiotic Selenium Analogue of Pheomelanin" 1571:

is directly proportional to its degree of polymerization or

1457:

does not appear to have undergone positive selection, while

1611:

by the RPE. This phenomenon has been attributed in part to

1300: 102: 96: 4939:"Human skin pigmentation as an adaptation to UV radiation" 77:

Micrograph of the epidermis, with melanin labeled at left.

108: 2646:"New biomaterial could shield against harmful radiation" 832:

3,8-Dimethyl-2,7-dihydrobenzodiindole-4,5,9,10-tetrone

585:, a fungus) melanins appear to play important roles in 4375:. Additional contributions from Kilbourn Gordon, III. 3905:"Some problems and errors in cytogenetic biodosimetry" 865:

3,8-Dimethyl-2,7-dihydrobenzodiindole-4,5,9,10-tetrone

826:

3,8-Dimethyl-2,7-dihydrobenzodiindole-4,5,9,10-tetrone

7082: 5109:

Meyskens FL, Farmer PJ, Anton-Culver H (April 2004).

4023:

Dadachova E, Casadevall A (2011). Horikoshi K (ed.).

3622: 3620: 111: 105: 5970:

Branched-chain alpha-keto acid dehydrogenase complex

5926:

Branched-chain alpha-keto acid dehydrogenase complex

5441:

Branched-chain alpha-keto acid dehydrogenase complex

4515:"Redox regulation in human melanocytes and melanoma" 2981:

Dadachova E, Bryan RA, Huang X, et al. (2007).

1270:

in the stria vascularis of the inner ear results in

99: 7030: 7007: 6978: 6940: 6873: 6864: 6807: 6781: 6668: 6612: 6593: 6584: 6539: 6508: 6463: 6416: 6388: 6373: 6341: 6320: 6297: 6286: 6272: 6188: 6178: 6116: 6106: 6072: 6053: 5983: 5954: 5910: 5895: 5833: 5813: 5784: 5746: 5717: 5702: 5679: 5660: 5641: 5593: 5578: 5569: 5544: 5496: 5420: 5396: 5383: 4839:"Evidence for Variable Selective Pressures at MC1R" 4663:

The Smithsonian Institution's Human Origins Program

3898: 3896: 3894: 3892: 3890: 3888: 3886: 3884: 3882: 3688: 3686: 3592: 3590: 3545:Hegna RH, Nokelainen O, Hegna JR, Mappes J (2013). 1082:by two pathways to benzothiazines and pheomelanins 93: 90: 51: 34: 4364: 3042:International Review of Cell and Molecular Biology 1551:Physical properties and technological applications 6706:Inherited patterned lentiginosis in black persons 4018: 4016: 3752: 3750: 3137:"Melanin and the Abrasion Resistance of Feathers" 504:. In the brain, tissues with melanin include the 488:In humans, melanin is the primary determinant of 16:Group of natural pigments found in most organisms 6915:Eczematid-like purpura of Doucas and Kapetanakis 5279:. Department of Computer Science and Technology. 4513:Meyskens FL, Farmer P, Fruehauf JP (June 2001). 3627:Gessler NN, Egorova AS, Belozerskaya TA (2014). 1472:is needed. This is the physiological purpose of 1277:In Parkinson's disease, a disorder that affects 1120:Detailed metabolic pathways can be found in the 932: 508:and pigment-bearing neurons within areas of the 327:. The arrows denote where the polymer continues. 271:. The arrow denotes where the polymer continues. 5531:Aminocarboxymuconate-semialdehyde decarboxylase 4943:Proceedings of the National Academy of Sciences 1099:and follow two more pathways to the eumelanins 1012:< −20 °C (−4 °F; 253 K) 887: 3910:Biotechnology & Biotechnological Equipment 2734:Del Bino S, Ito S, Sok J, Wakamatsu K (2022). 2313:Kottler VA, Künstner A, Schartl M (May 2015). 1667:, a syndrome characterised by hypopigmentation 339:Pheomelanin production is highly dependent on 223:portions that are largely responsible for the 6567:Yemenite deaf-blind hypopigmentation syndrome 6246: 5337: 5201: 5199: 3958: 3956: 3954: 3824: 3822: 3820: 3818: 3816: 3504:Hsiung BK, Blackledge TA, Shawkey MD (2015). 796:on alkali fusion. It is commonly seen in the 740:. Her most significant contribution is Mosse 738:Belarusian Institute of Genetics and Cytology 424:The biochemical composition of peptidomelanin 8: 4358: 4356: 3292:"Some aspects of melanin biology: 1950–1975" 2677:Annales de Génétique et de Sélection Animale 2308: 2306: 352:units that are produced, instead of DHI and 6635:Reticular pigmented anomaly of the flexures 5975:3-hydroxy-2-methylbutyryl-CoA dehydrogenase 3372:"Gene interaction in coat color of mammals" 3192:International Journal of Molecular Sciences 2793:"Skin melanocytes: biology and development" 287:. This reaction is catalyzed by the enzyme 6870: 6609: 6590: 6385: 6294: 6283: 6253: 6239: 6231: 6185: 6113: 5965:Branched-chain amino acid aminotransferase 5921:Branched-chain amino acid aminotransferase 5907: 5714: 5590: 5575: 5436:Branched-chain amino acid aminotransferase 5393: 5344: 5330: 5322: 3966:International Journal of Radiation Biology 1778: 1776: 1555:Evidence exists for a highly cross-linked 1235:mortality beyond melanoma-related deaths. 907: 830: 161:, which among other functions provide the 40: 5946:Methylmalonate semialdehyde dehydrogenase 5249: 5231: 5129: 4972: 4962: 4862: 4781: 4533: 4288: 4278: 4261:Zaidi KU, Ali AS, Ali SA, Naaz I (2014). 4237: 4204: 4152: 4103: 4093: 3926: 3796: 3730: 3693:Nenoi M, Wang B, Vares G (12 June 2014). 3650: 3611:Croatian Radiation Protection Association 3570: 3521: 3433:. Springer Science & Business Media. 3307: 3258: 3213: 3203: 3092: 3016: 3006: 2887: 2816: 2767: 2698: 2688: 2547: 2400: 2378: 2376: 2294: 1989: 1924: 1887: 1877: 1860:Haining RL, Achat-Mendes C (March 2017). 1808: 716:, 1985. Much of our understanding of the 6911:Doucas and Kapetanakis pigmented purpura 6640:Naegeli–Franceschetti–Jadassohn syndrome 5536:Aminomuconate-semialdehyde dehydrogenase 3296:The Journal of Investigative Dermatology 2569:Journal of the American Chemical Society 2194:The Journal of Investigative Dermatology 1789:Journal of the American Chemical Society 1627:Heavily pigmented melanoma cells have a 1291:substantia nigra from external toxins." 788:Chemical structure of indole-5,6-quinone 605:, appear to be able to use melanin as a 7089: 6650:X-linked reticulate pigmentary disorder 6630:Reticulate acropigmentation of Kitamura 6625:Pigmentatio reticularis faciei et colli 5767:1-Pyrroline-5-carboxylate dehydrogenase 5308:Kyoto Encyclopedia of Genes and Genomes 5289:Kyoto Encyclopedia of Genes and Genomes 5285:"Tyrosine metabolism—Reference pathway" 4937:Jablonski NG, Chaplin G (11 May 2010). 2797:Advances in Dermatology and Allergology 1713: 1354:rates in darker pigmented individuals. 1130: 744:, 2000 on mice but also includes Mosse 527:The melanin in the skin is produced by 332:orange hair, which is typically called 176:There are five basic types of melanin: 7022:Dyschromatosis universalis hereditaria 6827:Familial progressive hyperpigmentation 6027:Betaine-homocysteine methyltransferase 4843:The American Journal of Human Genetics 4711:"Why Humans and Their Fur Parted Ways" 4617:Pharmacology Biochemistry and Behavior 4323:from the original on 25 September 2017 3832:Journal of Environmental Radioactivity 2622: 2220:from the original on 25 September 2022 1375:. These are then transferred into the 1214:There are approximately nine types of 1095:Also, dopaquinone can be converted to 674:arises: giving a number of species an 445:It is possible to enrich melanin with 31: 7017:Dyschromatosis symmetrica hereditaria 6852:Transient neonatal pustular melanosis 5488:Template:Leucine metabolism in humans 3634:Applied Biochemistry and Microbiology 3486:from the original on 12 November 2023 2611:from the original on 6 September 2023 2361:from the original on 22 February 2024 2272: 2270: 2169:from the original on 11 February 2020 736:, and as of 2022 continues under the 281:5,6-dihydroxyindole-2-carboxylic acid 7: 6842:Photoleukomelanodermatitis of Kobori 6620:Dermatopathia pigmentosa reticularis 5628:Guanidinoacetate N-methyltransferase 4133:Cellular and Molecular Life Sciences 3469:Natural Photonics and Bioinspiration 3167:from the original on 23 January 2020 2870:Hamilton AJ, Gomez BL (March 2002). 2740:Pigment Cell & Melanoma Research 2319:Pigment Cell & Melanoma Research 1599:(RPE) also may be implicated in the 1539:crosslinking caused by UV exposure. 1238:The connection between albinism and 1210:Genetic disorders and disease states 613:and harness this energy for growth. 149: 'black, dark') is a family of 6141:4-Hydroxyphenylpyruvate dioxygenase 5738:Formiminotransferase cyclodeaminase 5461:Isovaleryl coenzyme A dehydrogenase 4553:from the original on 30 August 2024 4267:Biochemistry Research International 3506:"Spiders do have melanin after all" 3447:from the original on 30 August 2024 3351:from the original on 30 August 2024 3186:Galván I, Solano F (8 April 2016). 2715:from the original on 30 August 2024 2652:from the original on 30 August 2024 1841:from the original on 30 August 2024 1256:, predominantly observed among the 1125: 923: 46:One possible structure of Eumelanin 6847:Postinflammatory hyperpigmentation 6767:Poikiloderma vasculare atrophicans 5941:3-hydroxyisobutyrate dehydrogenase 5431:3-Hydroxybutyryl-CoA dehydrogenase 5157:from the original on 3 August 2020 5045:10.1111/j.1432-1033.1995.tb20794.x 4474:10.1111/j.1600-0749.1994.tb00015.x 4383:from the original on 28 March 2017 2163:MetaCyc Metabolic Pathway Database 1935:10.1111/j.1751-1097.2004.tb00012.x 1692:Risks and benefits of sun exposure 1463:has undergone positive selection. 820:Interpretation as a single monomer 593:by protecting the microbe against 275:Eumelanin has two forms linked to 14: 6552:Postinflammatory hypopigmentation 6496:Progressive macular hypomelanosis 5936:3-hydroxyisobutyryl-CoA hydrolase 5853:Gamma-glutamylcysteine synthetase 5762:Pyrroline-5-carboxylate reductase 4729:from the original on 18 June 2009 4535:10.1034/j.1600-0749.2001.140303.x 2851:from the original on 17 July 2020 2671:Prota, G., Searle, A. G. (1978). 2035:10.1034/j.1600-0749.2001.140505.x 1387:, where they protect the nuclear 687:Some moth species, including the 7104: 7092: 6486:Idiopathic guttate hypomelanosis 5277:"Absorption spectrum of melanin" 5033:European Journal of Biochemistry 5010:10.1111/j.1600-0749.2004.00140.x 3695:"In vivo radioadaptive response" 3430:Biology of the Arthropod Cuticle 3122:10.1111/j.0908-8857.2008.04413.x 2925:10.1111/j.0105-2896.2004.00116.x 2078:10.1111/j.1468-3083.2011.04278.x 1982:10.1111/j.1751-1097.2007.00226.x 1206:is an effective melanin bleach. 1181: 1169: 1157: 1145: 1133: 1103:dopaquinone → leucodopachrome → 968: 962: 846: 837: 181: 86: 6970:Titanium metallic discoloration 6726:Partial unilateral lentiginosis 5608:Serine hydroxymethyltransferase 4818:from the original on 2 May 2019 4574:Cutaneous and Ocular Toxicology 4226:Journal of Biological Chemistry 4127:Kim YJ, Uyama H (15 May 2005). 4076:Yao ZY, Qi JH (22 April 2016). 3510:Journal of Experimental Biology 2876:Journal of Medical Microbiology 2446:10.1016/j.pneurobio.2005.02.001 1970:Photochemistry and Photobiology 1913:Photochemistry and Photobiology 1032:(at 25 °C , 100 kPa). 6743:Erythema dyschromicum perstans 6022:Homocysteine methyltransferase 6000:Methionine adenosyltransferase 5481:Methylglutaconyl-CoA hydratase 5476:Methylcrotonyl-CoA carboxylase 4686:Rook's Textbook of Dermatology 3472:(1st ed.), Artech House, 3466:Mouchet SR, Deparis O (2021), 2872:"Melanins in fungal pathogens" 1964:Brenner M, Hearing VJ (2008). 974: 956: 302:Analysis of highly pigmented ( 219:-derivative that contains poly 177: 1: 6932:Hemosiderin hyperpigmentation 6891:Pigmented purpuric dermatosis 6837:Periorbital hyperpigmentation 6333:Vogt–Koyanagi–Harada syndrome 6151:Fumarylacetoacetate hydrolase 6146:Homogentisate 1,2-dioxygenase 5863:Gamma-glutamyl transpeptidase 5526:3-hydroxyanthranilate oxidase 5466:α-Ketoisocaproate dioxygenase 5131:10.1158/1078-0432.ccr-03-0638 4239:10.1016/S0021-9258(18)35614-X 3983:10.1080/09553002.2022.2026517 3928:10.1080/13102818.2016.1259018 3857:10.1016/j.jenvrad.2012.05.009 2629:: CS1 maint: date and year ( 1391:from mutations caused by the 1078:Dopaquinone can combine with 651:Melanin is also important in 609:that enables them to capture 6799:Shiitake mushroom dermatitis 4659:"Human Skin Color Variation" 4586:10.3109/15569527.2012.676119 3973:(3: Women in Radiobiology). 3309:10.1111/1523-1747.ep12512500 3054:10.1016/bs.ircmb.2015.09.005 3008:10.1371/journal.pone.0000457 2968:10.1002/scin.2007.5591712106 2206:10.1111/1523-1747.ep12480680 1866:Neural Regeneration Research 1525:high-frequency visible light 1511:Melanin in the eyes, in the 6886:Iron metallic discoloration 6562:Vagabond's leukomelanoderma 6089:Methylmalonyl CoA epimerase 6040:Cystathionine beta synthase 5507:Indoleamine 2,3-dioxygenase 4949:(Supplement 2): 8962–8968. 3337:. Oxford University Press. 2648:. Northwestern University. 2644:Mark Heiden (8 July 2020). 2241:Journal of Natural Products 1640:, so that they can both be 1623:Role in melanoma metastasis 1407:in lighter-skinned people. 720:effects of melanin against 7153: 6832:Pallister–Killian syndrome 6481:Albinism–deafness syndrome 6013:regeneration of methionine 5995:generation of homocysteine 5795:Ornithine aminotransferase 5511:Tryptophan 2,3-dioxygenase 5412:Glutaryl-CoA dehydrogenase 5407:Saccharopine dehydrogenase 5233:10.1038/s41598-019-45643-9 2889:10.1099/0022-1317-51-3-189 2129:10.1152/physrev.00044.2003 1743:10.1038/d41586-018-02370-x 1597:retinal pigment epithelium 1529:blue, green, and grey eyes 1433:Anatomically modern humans 370: 135: 18: 6748:Lichen planus pigmentosus 6696:Centrofacial lentiginosis 6453:Griscelli syndrome type 3 6448:Griscelli syndrome type 2 6428:Hermansky–Pudlak syndrome 6136:Tyrosine aminotransferase 6131:Phenylalanine hydroxylase 6084:Propionyl-CoA carboxylase 6045:Cystathionine gamma-lyase 5690:L-threonine dehydrogenase 5371: 4709:Wade N (19 August 2003). 4629:10.1016/j.pbb.2009.02.011 4343:"Oculocutaneous Albinism" 4145:10.1007/s00018-005-5054-y 4095:10.3390/molecules21040487 3652:10.1134/s0003683814020094 2841:"oculocutaneous albinism" 2489:10.1007/s00702-006-0450-5 1026: 943: 871: 859: 854: 845: 836: 632:use pheomelanin instead. 531:, which are found in the 379:catecholaminergic neurons 39: 6701:Generalized lentiginosis 6433:Chédiak–Higashi syndrome 6094:Methylmalonyl-CoA mutase 5671:D-cysteine desulfhydrase 5118:Clinical Cancer Research 4317:pubchem.ncbi.nlm.nih.gov 3715:10.1177/0960327114537537 3110:Journal of Avian Biology 2690:10.1186/1297-9686-10-1-1 2532:10.1021/acsomega.4c03704 1879:10.4103/1673-5374.202928 1245:On the Origin of Species 1200:blood breakdown pigments 516:. It also occurs in the 55:Heterogeneous biopolymer 19:Not to be confused with 6762:Poikiloderma of Civatte 6398:Oculocutaneous albinism 5825:Glutamate dehydrogenase 5800:Ornithine decarboxylase 5728:Histidine ammonia-lyase 5471:Leucine 2,3-aminomutase 4964:10.1073/pnas.0914628107 4908:10.1126/science.1116238 4774:10.1126/science.1172257 4407:"Causes of Variability" 4027:. Tokyo New York City: 3406:10.1242/dev.121.10.3223 2845:Genetics Home Reference 2809:10.5114/pdia.2013.33376 1216:oculocutaneous albinism 804:by the tyrosinase type 712:models include Hopwood 582:Cryptococcus neoformans 569:reactive oxygen species 398:Other forms of melanins 291:. The initial product, 6906:Gougerot–Blum syndrome 6789:Incontinentia pigmenti 6645:Dyskeratosis congenita 6572:Wende–Bauckus syndrome 6491:Phylloid hypomelanosis 6262:Pigmentation disorders 6212:Aspartate transaminase 6035:conversion to cysteine 6005:Adenosylhomocysteinase 5858:Glutathione synthetase 4193:New Journal of Science 4031:. pp. xxix+1247. 4025:Extremophiles handbook 3781:10.1002/advs.201903129 3613:. p. 35 (of 268). 3563:10.1098/rspb.2012.2812 2315:"Pheomelanin in fish?" 2283:New Journal of Science 1563:to matrix scaffolding 1254:Waardenburg's syndrome 1204:potassium permanganate 1194:Microscopic appearance 789: 607:photosynthetic pigment 485: 425: 417: 356:, when the amino acid 334:"red" or "ginger" hair 328: 272: 78: 70: 6328:Alezzandrini syndrome 6307:Quadrichrome vitiligo 6207:Asparagine synthetase 5375:Essential amino acids 4998:Pigment Cell Research 4814:. 22 September 2016. 4522:Pigment Cell Research 4462:Pigment Cell Research 4273:: 1–16 (see Fig. 3). 2913:Immunological Reviews 2117:Physiological Reviews 2023:Pigment Cell Research 1519:, helps protect from 1048:Biosynthetic pathways 787: 672:structural coloration 476: 423: 414: 322: 266: 76: 64: 6925:Angioma serpiginosum 6817:Acanthosis nigricans 6356:Waardenburg syndrome 5878:Glutamine synthetase 5652:Alanine transaminase 4413:on 23 September 2006 4349:on 23 December 2008. 3975:Taylor & Francis 3919:Taylor & Francis 3205:10.3390/ijms17040520 2581:10.1021/jacs.0c05573 1801:10.1021/jacs.0c12322 1688:, melanin production 1617:macular degeneration 1595:melanization in the 1581:macular degeneration 1419:Evolutionary origins 1313:macular degeneration 861:Preferred IUPAC name 496:of the eye, and the 6655:Galli–Galli disease 6547:Nevus depigmentosus 6018:Methionine synthase 5931:Enoyl-CoA hydratase 5816:alpha-ketoglutarate 5733:Urocanate hydratase 5446:Enoyl-CoA hydratase 5224:2019NatSR...9.9280S 5068:Ophthalmic Research 4955:2010PNAS..107.8962J 4900:2005Sci...310.1782L 4766:2009Sci...324.1035T 4496:"Lewy Body Disease" 4280:10.1155/2014/854687 4206:10.1155/2014/498276 3849:2012JEnvR.112...70M 3643:Pleiades Publishing 3427:Neville AC (2012). 2999:2007PLoSO...2..457D 2575:(29): 12802–12810. 2296:10.1155/2014/498276 1735:2018Natur.555..319C 1660:Albinism in biology 1379:cells of the human 1348:nicotine dependence 992: g·mol 833: 392:Parkinson's disease 277:5,6-dihydroxyindole 6901:Majocchi's disease 6794:Scratch dermatitis 6721:Mucosal lentigines 6521:Vasospastic macule 6438:Griscelli syndrome 6064:Threonine aldolase 5613:Serine dehydratase 5357:Protein metabolism 5212:Scientific Reports 4811:The New York Times 4715:The New York Times 3557:(1755): 20122812. 3523:10.1242/jeb.128801 3331:Meneely P (2014). 3135:Bonser RH (1995). 3094:10.3390/opt4040041 2752:10.1111/pcmr.13062 2331:10.1111/pcmr.12359 1665:Griscelli syndrome 1589:Signaling pathways 1393:ionizing radiation 1036:Infobox references 831: 806:polyphenol oxidase 802:indole-5,6-quinone 798:enzymatic browning 790: 623:Stegastes apicalis 603:radiotrophic fungi 486: 426: 418: 329: 273: 244:malignant melanoma 79: 71: 7080: 7079: 7003: 7002: 6999: 6998: 6960:Arsenic poisoning 6896:Schamberg disease 6860: 6859: 6753:Café au lait spot 6580: 6579: 6504: 6503: 6443:Elejalde syndrome 6369: 6368: 6228: 6227: 6224: 6223: 6220: 6219: 6174: 6173: 6102: 6101: 5891: 5890: 5698: 5697: 5565: 5564: 5456:HMG-CoA reductase 5080:10.1159/000085533 4760:(5930): 1035–44. 4695:978-1-4051-6169-5 4220:Mason HS (1948). 4187:Solano F (2014). 4139:(15): 1707–1723. 4056:978-4-431-53897-4 4038:978-4-431-53898-1 3516:(22): 3632–3635. 3479:978-163-081-797-8 3400:(10): 3223–3232. 3251:10.1111/ibi.12770 2395:(11): 1212–1219. 2277:Solano F (2014). 2253:10.1021/np100740n 2072:(12): 1369–1380. 1729:(7696): 319–320. 1358:Human adaptations 1352:smoking cessation 1176:L-leucodopachrome 1044:Chemical compound 1042: 1041: 772:, 2007 and Mosse 698:Melanin protects 597:responses of its 567:from the sun and 463:metabolic pathway 430:Aspergillus niger 384:transition metals 59: 58: 7144: 7127:Human skin color 7109: 7108: 7107: 7097: 7096: 7088: 6871: 6711:Ink spot lentigo 6610: 6591: 6386: 6312:Vitiligo ponctué 6295: 6284: 6255: 6248: 6241: 6232: 6186: 6114: 5908: 5715: 5591: 5576: 5394: 5346: 5339: 5332: 5323: 5318: 5316: 5314: 5299: 5297: 5295: 5280: 5264: 5263: 5253: 5235: 5203: 5194: 5193: 5178:Molecular Vision 5173: 5167: 5166: 5164: 5162: 5156: 5133: 5115: 5106: 5100: 5099: 5063: 5057: 5056: 5028: 5022: 5021: 4993: 4987: 4986: 4976: 4966: 4934: 4928: 4927: 4894:(5755): 1782–6. 4883: 4877: 4876: 4866: 4849:(4): 1351–1361. 4834: 4828: 4827: 4825: 4823: 4802: 4796: 4795: 4785: 4745: 4739: 4738: 4736: 4734: 4706: 4700: 4699: 4681: 4675: 4674: 4672: 4670: 4655: 4649: 4648: 4612: 4606: 4605: 4569: 4563: 4562: 4560: 4558: 4552: 4537: 4519: 4510: 4504: 4503: 4502:on 21 July 2009. 4498:. Archived from 4492: 4486: 4485: 4457: 4451: 4443: 4437: 4429: 4423: 4422: 4420: 4418: 4409:. Archived from 4403: 4397: 4396: 4390: 4388: 4368: 4360: 4351: 4350: 4345:. Archived from 4339: 4333: 4332: 4330: 4328: 4309: 4303: 4302: 4292: 4282: 4258: 4252: 4251: 4241: 4217: 4211: 4210: 4208: 4199:(498276): 1–28. 4184: 4175: 4174: 4156: 4124: 4118: 4117: 4107: 4097: 4073: 4067: 4050: 4020: 4011: 4010: 3960: 3949: 3948: 3930: 3900: 3877: 3876: 3826: 3811: 3810: 3800: 3764:Advanced Science 3754: 3745: 3744: 3734: 3690: 3681: 3680: 3654: 3624: 3615: 3614: 3594: 3585: 3584: 3574: 3542: 3536: 3535: 3525: 3501: 3495: 3494: 3493: 3491: 3463: 3457: 3456: 3454: 3452: 3424: 3418: 3417: 3388: 3382: 3381: 3367: 3361: 3360: 3358: 3356: 3328: 3322: 3321: 3311: 3287: 3281: 3280: 3262: 3234: 3228: 3227: 3217: 3207: 3183: 3177: 3176: 3174: 3172: 3132: 3126: 3125: 3105: 3099: 3098: 3096: 3072: 3066: 3065: 3037: 3031: 3030: 3020: 3010: 2978: 2972: 2971: 2951: 2945: 2944: 2908: 2902: 2901: 2891: 2867: 2861: 2860: 2858: 2856: 2837: 2831: 2830: 2820: 2788: 2782: 2781: 2771: 2731: 2725: 2724: 2722: 2720: 2702: 2692: 2668: 2662: 2661: 2659: 2657: 2641: 2635: 2634: 2628: 2620: 2618: 2616: 2560: 2554: 2553: 2551: 2515: 2509: 2508: 2472: 2466: 2465: 2429: 2423: 2422: 2404: 2380: 2371: 2370: 2368: 2366: 2310: 2301: 2300: 2298: 2274: 2265: 2264: 2236: 2230: 2229: 2227: 2225: 2185: 2179: 2178: 2176: 2174: 2155: 2149: 2148: 2112: 2106: 2105: 2061: 2055: 2054: 2017: 2004: 2003: 1993: 1961: 1955: 1954: 1928: 1926:cond-mat/0312277 1908: 1902: 1901: 1891: 1881: 1857: 1851: 1850: 1848: 1846: 1812: 1795:(7): 2622–2637. 1780: 1771: 1770: 1718: 1671:Human skin color 1644:and metastatic. 1573:molecular weight 1486:photodegradation 1484:, including the 1328:port-wine stains 1185: 1173: 1161: 1149: 1137: 1002:1.6 to 1.8 g/cm 991: 976: 970: 964: 958: 951:Chemical formula 936: 925: 911: 891: 850: 841: 834: 577:oxidizing agents 518:zona reticularis 498:stria vascularis 482:Papua New Guinea 390:and the related 304:Fitzpatrick type 146: 139: 127: 126: 125: 124: 117: 114: 113: 110: 107: 104: 101: 98: 95: 92: 44: 32: 7152: 7151: 7147: 7146: 7145: 7143: 7142: 7141: 7117: 7116: 7115: 7105: 7103: 7091: 7083: 7081: 7076: 7026: 6995: 6974: 6942: 6936: 6881:Hemochromatosis 6866: 6856: 6809: 6803: 6777: 6773:Riehl melanosis 6731:PUVA lentigines 6716:Lentigo maligna 6686:Lentigo simplex 6670: 6664: 6660:Revesz syndrome 6603: 6598: 6576: 6557:Pityriasis alba 6535: 6526:Woronoff's ring 6513: 6500: 6459: 6420: 6412: 6403:Ocular albinism 6379: 6365: 6343: 6337: 6316: 6288: 6277: 6268: 6259: 6229: 6216: 6170: 6098: 6068: 6049: 5979: 5950: 5901: 5887: 5829: 5809: 5780: 5757:Proline oxidase 5742: 5710:α-ketoglutarate 5708: 5694: 5675: 5656: 5637: 5633:Creatine kinase 5583: 5561: 5540: 5516:Arylformamidase 5492: 5416: 5379: 5377:are in Capitals 5367: 5363:and catabolism 5350: 5312: 5310: 5304:"Melanogenesis" 5302: 5293: 5291: 5283: 5275: 5272: 5267: 5205: 5204: 5197: 5175: 5174: 5170: 5160: 5158: 5154: 5113: 5108: 5107: 5103: 5065: 5064: 5060: 5030: 5029: 5025: 4995: 4994: 4990: 4936: 4935: 4931: 4885: 4884: 4880: 4836: 4835: 4831: 4821: 4819: 4804: 4803: 4799: 4747: 4746: 4742: 4732: 4730: 4708: 4707: 4703: 4696: 4683: 4682: 4678: 4668: 4666: 4657: 4656: 4652: 4614: 4613: 4609: 4571: 4570: 4566: 4556: 4554: 4550: 4517: 4512: 4511: 4507: 4494: 4493: 4489: 4459: 4458: 4454: 4444: 4440: 4430: 4426: 4416: 4414: 4405: 4404: 4400: 4386: 4384: 4362: 4361: 4354: 4341: 4340: 4336: 4326: 4324: 4311: 4310: 4306: 4260: 4259: 4255: 4219: 4218: 4214: 4186: 4185: 4178: 4126: 4125: 4121: 4075: 4074: 4070: 4039: 4022: 4021: 4014: 3962: 3961: 3952: 3902: 3901: 3880: 3828: 3827: 3814: 3756: 3755: 3748: 3692: 3691: 3684: 3626: 3625: 3618: 3596: 3595: 3588: 3551:Proc. R. Soc. B 3544: 3543: 3539: 3503: 3502: 3498: 3489: 3487: 3480: 3465: 3464: 3460: 3450: 3448: 3441: 3426: 3425: 3421: 3390: 3389: 3385: 3369: 3368: 3364: 3354: 3352: 3345: 3330: 3329: 3325: 3289: 3288: 3284: 3236: 3235: 3231: 3185: 3184: 3180: 3170: 3168: 3153:10.2307/1369048 3134: 3133: 3129: 3107: 3106: 3102: 3074: 3073: 3069: 3039: 3038: 3034: 2980: 2979: 2975: 2953: 2952: 2948: 2910: 2909: 2905: 2869: 2868: 2864: 2854: 2852: 2839: 2838: 2834: 2790: 2789: 2785: 2733: 2732: 2728: 2718: 2716: 2670: 2669: 2665: 2655: 2653: 2643: 2642: 2638: 2621: 2614: 2612: 2562: 2561: 2557: 2517: 2516: 2512: 2477:J Neural Transm 2474: 2473: 2469: 2431: 2430: 2426: 2402:10.1038/ng.3098 2389:Nature Genetics 2382: 2381: 2374: 2364: 2362: 2312: 2311: 2304: 2276: 2275: 2268: 2238: 2237: 2233: 2223: 2221: 2187: 2186: 2182: 2172: 2170: 2157: 2156: 2152: 2123:(4): 1155–228. 2114: 2113: 2109: 2063: 2062: 2058: 2019: 2018: 2007: 1963: 1962: 1958: 1910: 1909: 1905: 1859: 1858: 1854: 1844: 1842: 1810:1854/LU-8699336 1782: 1781: 1774: 1720: 1719: 1715: 1711: 1706: 1650: 1629:Young's modulus 1625: 1553: 1482:solar radiation 1469: 1421: 1365: 1360: 1297:matrix proteins 1212: 1196: 1189: 1186: 1177: 1174: 1165: 1162: 1153: 1150: 1141: 1138: 1097:leucodopachrome 1050: 1045: 1038: 1033: 989: 979: 973: 967: 961: 953: 939: 926: 914: 894: 881: 867: 866: 822: 782: 722:gamma radiation 718:radioprotective 689:wood tiger moth 668:Bragg reflector 629:Cyprinus carpio 553: 551:Other organisms 514:locus coeruleus 471: 459: 443: 409: 400: 375: 369: 317: 261: 256: 211:is followed by 121: 120: 119: 89: 85: 47: 28: 17: 12: 11: 5: 7150: 7148: 7140: 7139: 7134: 7129: 7119: 7118: 7114: 7113: 7101: 7078: 7077: 7075: 7074: 7072:Depigmentation 7069: 7068: 7067: 7057: 7056: 7055: 7045: 7043:Skin whitening 7040: 7034: 7032: 7028: 7027: 7025: 7024: 7019: 7013: 7011: 7005: 7004: 7001: 7000: 6997: 6996: 6994: 6993: 6988: 6982: 6980: 6976: 6975: 6973: 6972: 6967: 6965:Lead poisoning 6962: 6957: 6952: 6946: 6944: 6938: 6937: 6935: 6934: 6929: 6928: 6927: 6922: 6917: 6908: 6903: 6898: 6888: 6883: 6877: 6875: 6868: 6862: 6861: 6858: 6857: 6855: 6854: 6849: 6844: 6839: 6834: 6829: 6824: 6819: 6813: 6811: 6805: 6804: 6802: 6801: 6796: 6791: 6785: 6783: 6779: 6778: 6776: 6775: 6770: 6764: 6755: 6750: 6745: 6740: 6734: 6733: 6728: 6723: 6718: 6713: 6708: 6703: 6698: 6693: 6688: 6674: 6672: 6666: 6665: 6663: 6662: 6657: 6652: 6647: 6642: 6637: 6632: 6627: 6622: 6616: 6614: 6607: 6588: 6582: 6581: 6578: 6577: 6575: 6574: 6569: 6564: 6559: 6554: 6549: 6543: 6541: 6537: 6536: 6534: 6533: 6531:Nevus anemicus 6528: 6523: 6517: 6515: 6506: 6505: 6502: 6501: 6499: 6498: 6493: 6488: 6483: 6478: 6473: 6471:Cross syndrome 6467: 6465: 6461: 6460: 6458: 6457: 6456: 6455: 6450: 6445: 6435: 6430: 6424: 6422: 6414: 6413: 6411: 6410: 6405: 6400: 6394: 6392: 6383: 6371: 6370: 6367: 6366: 6364: 6363: 6361:Tietz syndrome 6358: 6353: 6347: 6345: 6339: 6338: 6336: 6335: 6330: 6324: 6322: 6318: 6317: 6315: 6314: 6309: 6303: 6301: 6292: 6281: 6270: 6269: 6260: 6258: 6257: 6250: 6243: 6235: 6226: 6225: 6222: 6221: 6218: 6217: 6215: 6214: 6209: 6199: 6197: 6183: 6176: 6175: 6172: 6171: 6169: 6168: 6154: 6153: 6148: 6143: 6138: 6133: 6127: 6125: 6111: 6104: 6103: 6100: 6099: 6097: 6096: 6091: 6086: 6080: 6078: 6070: 6069: 6067: 6066: 6060: 6058: 6051: 6050: 6048: 6047: 6042: 6030: 6029: 6024: 6008: 6007: 6002: 5990: 5988: 5981: 5980: 5978: 5977: 5972: 5967: 5961: 5959: 5952: 5951: 5949: 5948: 5943: 5938: 5933: 5928: 5923: 5917: 5915: 5905: 5893: 5892: 5889: 5888: 5886: 5885: 5880: 5866: 5865: 5860: 5855: 5837: 5835: 5831: 5830: 5828: 5827: 5821: 5819: 5811: 5810: 5808: 5807: 5802: 5797: 5791: 5789: 5782: 5781: 5779: 5778: 5773: 5764: 5759: 5753: 5751: 5744: 5743: 5741: 5740: 5735: 5730: 5724: 5722: 5712: 5700: 5699: 5696: 5695: 5693: 5692: 5686: 5684: 5677: 5676: 5674: 5673: 5667: 5665: 5658: 5657: 5655: 5654: 5648: 5646: 5639: 5638: 5636: 5635: 5630: 5616: 5615: 5610: 5604: 5602: 5588: 5573: 5567: 5566: 5563: 5562: 5560: 5559: 5555: 5553: 5542: 5541: 5539: 5538: 5533: 5528: 5523: 5518: 5513: 5503: 5501: 5494: 5493: 5484: 5483: 5478: 5473: 5468: 5463: 5458: 5453: 5448: 5443: 5438: 5433: 5427: 5425: 5418: 5417: 5415: 5414: 5409: 5403: 5401: 5391: 5381: 5380: 5372: 5369: 5368: 5351: 5349: 5348: 5341: 5334: 5326: 5320: 5319: 5300: 5281: 5271: 5270:External links 5268: 5266: 5265: 5195: 5168: 5101: 5058: 5023: 4988: 4929: 4878: 4855:10.1086/302863 4829: 4797: 4740: 4701: 4694: 4676: 4665:. 20 June 2012 4650: 4607: 4564: 4505: 4487: 4452: 4438: 4424: 4398: 4352: 4334: 4304: 4253: 4212: 4176: 4119: 4068: 4037: 4012: 3950: 3878: 3812: 3746: 3682: 3616: 3586: 3537: 3496: 3478: 3458: 3439: 3419: 3383: 3362: 3343: 3323: 3282: 3245:(1): 238–244. 3229: 3178: 3147:(2): 590–591. 3127: 3116:(5): 539–545. 3100: 3087:(4): 563–562. 3067: 3032: 2973: 2946: 2903: 2862: 2832: 2783: 2746:(6): 622–626. 2726: 2663: 2636: 2555: 2510: 2483:(6): 751–756. 2467: 2440:(2): 109–124. 2434:Prog Neurobiol 2424: 2372: 2325:(3): 355–356. 2302: 2266: 2231: 2200:(2): 340–344. 2180: 2150: 2107: 2056: 2029:(5): 337–347. 2005: 1956: 1903: 1872:(3): 372–375. 1852: 1772: 1712: 1710: 1707: 1705: 1704: 1699: 1697:Skin whitening 1694: 1689: 1683: 1678: 1676:Melanin theory 1673: 1668: 1662: 1657: 1651: 1649: 1646: 1642:drug-resistant 1624: 1621: 1607:outer segment 1601:downregulation 1565:melanoproteins 1552: 1549: 1527:; people with 1468: 1465: 1420: 1417: 1364: 1361: 1359: 1356: 1305:ocular melanin 1288:norepinephrine 1250:Charles Darwin 1211: 1208: 1195: 1192: 1191: 1190: 1187: 1180: 1178: 1175: 1168: 1166: 1163: 1156: 1154: 1151: 1144: 1142: 1139: 1132: 1126:External links 1124:database (see 1118: 1117: 1109: 1108: 1093: 1092: 1088: 1087: 1076: 1075: 1049: 1046: 1043: 1040: 1039: 1034: 1030:standard state 1027: 1024: 1023: 1020: 1014: 1013: 1010: 1004: 1003: 1000: 994: 993: 987: 981: 980: 977: 971: 965: 959: 954: 949: 946: 945: 941: 940: 938: 937: 929: 927: 919: 916: 915: 913: 912: 904: 902: 896: 895: 893: 892: 884: 882: 877: 874: 873: 869: 868: 864: 863: 857: 856: 852: 851: 843: 842: 821: 818: 814:catecholamines 781: 778: 768:, 2006, Mosse 764:, 2002, Mosse 760:, 2001, Mosse 756:, 1998, Mosse 752:, 1997, Mosse 748:, 1994, Mosse 561:cephalopod ink 552: 549: 512:, such as the 470: 467: 458: 455: 442: 439: 408: 407:Peptidomelanin 405: 399: 396: 371:Main article: 368: 365: 316: 313: 260: 257: 255: 252: 213:polymerization 57: 56: 53: 49: 48: 45: 37: 36: 15: 13: 10: 9: 6: 4: 3: 2: 7149: 7138: 7135: 7133: 7130: 7128: 7125: 7124: 7122: 7112: 7102: 7100: 7095: 7090: 7086: 7073: 7070: 7066: 7063: 7062: 7061: 7058: 7054: 7051: 7050: 7049: 7046: 7044: 7041: 7039: 7036: 7035: 7033: 7029: 7023: 7020: 7018: 7015: 7014: 7012: 7010: 7006: 6992: 6991:Tar melanosis 6989: 6987: 6984: 6983: 6981: 6977: 6971: 6968: 6966: 6963: 6961: 6958: 6956: 6953: 6951: 6948: 6947: 6945: 6939: 6933: 6930: 6926: 6923: 6921: 6920:Lichen aureus 6918: 6916: 6912: 6909: 6907: 6904: 6902: 6899: 6897: 6894: 6893: 6892: 6889: 6887: 6884: 6882: 6879: 6878: 6876: 6872: 6869: 6863: 6853: 6850: 6848: 6845: 6843: 6840: 6838: 6835: 6833: 6830: 6828: 6825: 6823: 6820: 6818: 6815: 6814: 6812: 6806: 6800: 6797: 6795: 6792: 6790: 6787: 6786: 6784: 6780: 6774: 6771: 6768: 6765: 6763: 6759: 6756: 6754: 6751: 6749: 6746: 6744: 6741: 6739: 6736: 6735: 6732: 6729: 6727: 6724: 6722: 6719: 6717: 6714: 6712: 6709: 6707: 6704: 6702: 6699: 6697: 6694: 6692: 6689: 6687: 6683: 6679: 6676: 6675: 6673: 6671:circumscribed 6667: 6661: 6658: 6656: 6653: 6651: 6648: 6646: 6643: 6641: 6638: 6636: 6633: 6631: 6628: 6626: 6623: 6621: 6618: 6617: 6615: 6611: 6608: 6606: 6601: 6596: 6592: 6589: 6587: 6583: 6573: 6570: 6568: 6565: 6563: 6560: 6558: 6555: 6553: 6550: 6548: 6545: 6544: 6542: 6538: 6532: 6529: 6527: 6524: 6522: 6519: 6518: 6516: 6514:hypomelanosis 6511: 6507: 6497: 6494: 6492: 6489: 6487: 6484: 6482: 6479: 6477: 6476:ABCD syndrome 6474: 6472: 6469: 6468: 6466: 6462: 6454: 6451: 6449: 6446: 6444: 6441: 6440: 6439: 6436: 6434: 6431: 6429: 6426: 6425: 6423: 6419: 6415: 6409: 6406: 6404: 6401: 6399: 6396: 6395: 6393: 6391: 6387: 6384: 6382: 6377: 6372: 6362: 6359: 6357: 6354: 6352: 6349: 6348: 6346: 6340: 6334: 6331: 6329: 6326: 6325: 6323: 6319: 6313: 6310: 6308: 6305: 6304: 6302: 6300: 6296: 6293: 6291: 6285: 6282: 6280: 6275: 6271: 6267: 6263: 6256: 6251: 6249: 6244: 6242: 6237: 6236: 6233: 6213: 6210: 6208: 6204: 6201: 6200: 6198: 6195: 6191: 6187: 6184: 6182: 6177: 6167: 6163: 6159: 6156: 6155: 6152: 6149: 6147: 6144: 6142: 6139: 6137: 6134: 6132: 6129: 6128: 6126: 6123: 6119: 6118:PHENYLALANINE 6115: 6112: 6110: 6105: 6095: 6092: 6090: 6087: 6085: 6082: 6081: 6079: 6076: 6071: 6065: 6062: 6061: 6059: 6056: 6052: 6046: 6043: 6041: 6038: 6036: 6032: 6031: 6028: 6025: 6023: 6019: 6016: 6014: 6010: 6009: 6006: 6003: 6001: 5998: 5996: 5992: 5991: 5989: 5986: 5982: 5976: 5973: 5971: 5968: 5966: 5963: 5962: 5960: 5957: 5953: 5947: 5944: 5942: 5939: 5937: 5934: 5932: 5929: 5927: 5924: 5922: 5919: 5918: 5916: 5913: 5909: 5906: 5904: 5899: 5898:propionyl-CoA 5894: 5884: 5881: 5879: 5875: 5871: 5868: 5867: 5864: 5861: 5859: 5856: 5854: 5850: 5846: 5842: 5839: 5838: 5836: 5832: 5826: 5823: 5822: 5820: 5817: 5812: 5806: 5803: 5801: 5798: 5796: 5793: 5792: 5790: 5787: 5783: 5777: 5774: 5772: 5768: 5765: 5763: 5760: 5758: 5755: 5754: 5752: 5749: 5745: 5739: 5736: 5734: 5731: 5729: 5726: 5725: 5723: 5720: 5716: 5713: 5711: 5706: 5701: 5691: 5688: 5687: 5685: 5682: 5678: 5672: 5669: 5668: 5666: 5663: 5659: 5653: 5650: 5649: 5647: 5644: 5640: 5634: 5631: 5629: 5625: 5621: 5618: 5617: 5614: 5611: 5609: 5606: 5605: 5603: 5600: 5596: 5592: 5589: 5587: 5582: 5577: 5574: 5572: 5568: 5557: 5556: 5554: 5551: 5547: 5546:PHENYLALANINE 5543: 5537: 5534: 5532: 5529: 5527: 5524: 5522: 5519: 5517: 5514: 5512: 5508: 5505: 5504: 5502: 5499: 5495: 5491: 5489: 5482: 5479: 5477: 5474: 5472: 5469: 5467: 5464: 5462: 5459: 5457: 5454: 5452: 5451:HMG-CoA lyase 5449: 5447: 5444: 5442: 5439: 5437: 5434: 5432: 5429: 5428: 5426: 5423: 5419: 5413: 5410: 5408: 5405: 5404: 5402: 5399: 5395: 5392: 5390: 5386: 5382: 5378: 5376: 5370: 5366: 5362: 5358: 5354: 5347: 5342: 5340: 5335: 5333: 5328: 5327: 5324: 5309: 5305: 5301: 5290: 5286: 5282: 5278: 5274: 5273: 5269: 5261: 5257: 5252: 5247: 5243: 5239: 5234: 5229: 5225: 5221: 5217: 5213: 5209: 5202: 5200: 5196: 5191: 5187: 5183: 5179: 5172: 5169: 5153: 5149: 5145: 5141: 5137: 5132: 5127: 5124:(8): 2581–3. 5123: 5119: 5112: 5105: 5102: 5097: 5093: 5089: 5085: 5081: 5077: 5074:(3): 136–41. 5073: 5069: 5062: 5059: 5054: 5050: 5046: 5042: 5039:(1): 159–64. 5038: 5034: 5027: 5024: 5019: 5015: 5011: 5007: 5003: 4999: 4992: 4989: 4984: 4980: 4975: 4970: 4965: 4960: 4956: 4952: 4948: 4944: 4940: 4933: 4930: 4925: 4921: 4917: 4913: 4909: 4905: 4901: 4897: 4893: 4889: 4882: 4879: 4874: 4870: 4865: 4860: 4856: 4852: 4848: 4844: 4840: 4833: 4830: 4817: 4813: 4812: 4807: 4801: 4798: 4793: 4789: 4784: 4779: 4775: 4771: 4767: 4763: 4759: 4755: 4751: 4744: 4741: 4728: 4724: 4720: 4716: 4712: 4705: 4702: 4697: 4691: 4687: 4680: 4677: 4664: 4660: 4654: 4651: 4646: 4642: 4638: 4634: 4630: 4626: 4623:(4): 589–96. 4622: 4618: 4611: 4608: 4603: 4599: 4595: 4591: 4587: 4583: 4579: 4575: 4568: 4565: 4549: 4545: 4541: 4536: 4531: 4528:(3): 148–54. 4527: 4523: 4516: 4509: 4506: 4501: 4497: 4491: 4488: 4483: 4479: 4475: 4471: 4467: 4463: 4456: 4453: 4450: 4447: 4442: 4439: 4436: 4433: 4428: 4425: 4412: 4408: 4402: 4399: 4394: 4382: 4378: 4374: 4373: 4367: 4359: 4357: 4353: 4348: 4344: 4338: 4335: 4322: 4318: 4314: 4308: 4305: 4300: 4296: 4291: 4286: 4281: 4276: 4272: 4268: 4264: 4257: 4254: 4249: 4245: 4240: 4235: 4231: 4227: 4223: 4216: 4213: 4207: 4202: 4198: 4194: 4190: 4183: 4181: 4177: 4172: 4168: 4164: 4160: 4155: 4150: 4146: 4142: 4138: 4134: 4130: 4123: 4120: 4115: 4111: 4106: 4101: 4096: 4091: 4087: 4083: 4079: 4072: 4069: 4065: 4061: 4057: 4053: 4048: 4044: 4040: 4034: 4030: 4026: 4019: 4017: 4013: 4008: 4004: 4000: 3996: 3992: 3988: 3984: 3980: 3976: 3972: 3968: 3967: 3959: 3957: 3955: 3951: 3946: 3942: 3938: 3934: 3929: 3924: 3920: 3916: 3912: 3911: 3906: 3899: 3897: 3895: 3893: 3891: 3889: 3887: 3885: 3883: 3879: 3874: 3870: 3866: 3862: 3858: 3854: 3850: 3846: 3842: 3838: 3834: 3833: 3825: 3823: 3821: 3819: 3817: 3813: 3808: 3804: 3799: 3794: 3790: 3786: 3782: 3778: 3774: 3770: 3766: 3765: 3760: 3753: 3751: 3747: 3742: 3738: 3733: 3728: 3724: 3720: 3716: 3712: 3708: 3704: 3700: 3696: 3689: 3687: 3683: 3678: 3674: 3670: 3666: 3662: 3658: 3653: 3648: 3644: 3640: 3636: 3635: 3630: 3623: 3621: 3617: 3612: 3608: 3604: 3600: 3593: 3591: 3587: 3582: 3578: 3573: 3568: 3564: 3560: 3556: 3552: 3548: 3541: 3538: 3533: 3529: 3524: 3519: 3515: 3511: 3507: 3500: 3497: 3485: 3481: 3475: 3471: 3470: 3462: 3459: 3446: 3442: 3440:9783642809101 3436: 3432: 3431: 3423: 3420: 3415: 3411: 3407: 3403: 3399: 3395: 3387: 3384: 3379: 3378: 3373: 3366: 3363: 3350: 3346: 3344:9780199681266 3340: 3336: 3335: 3327: 3324: 3319: 3315: 3310: 3305: 3301: 3297: 3293: 3286: 3283: 3278: 3274: 3270: 3266: 3261: 3256: 3252: 3248: 3244: 3240: 3233: 3230: 3225: 3221: 3216: 3211: 3206: 3201: 3197: 3193: 3189: 3182: 3179: 3166: 3162: 3158: 3154: 3150: 3146: 3142: 3138: 3131: 3128: 3123: 3119: 3115: 3111: 3104: 3101: 3095: 3090: 3086: 3082: 3078: 3071: 3068: 3063: 3059: 3055: 3051: 3047: 3043: 3036: 3033: 3028: 3024: 3019: 3014: 3009: 3004: 3000: 2996: 2992: 2988: 2984: 2977: 2974: 2969: 2965: 2961: 2957: 2950: 2947: 2942: 2938: 2934: 2930: 2926: 2922: 2918: 2914: 2907: 2904: 2899: 2895: 2890: 2885: 2882:(3): 189–91. 2881: 2877: 2873: 2866: 2863: 2850: 2846: 2842: 2836: 2833: 2828: 2824: 2819: 2814: 2810: 2806: 2802: 2798: 2794: 2787: 2784: 2779: 2775: 2770: 2765: 2761: 2757: 2753: 2749: 2745: 2741: 2737: 2730: 2727: 2714: 2710: 2706: 2701: 2696: 2691: 2686: 2682: 2678: 2674: 2667: 2664: 2651: 2647: 2640: 2637: 2632: 2626: 2610: 2606: 2602: 2598: 2594: 2590: 2586: 2582: 2578: 2574: 2570: 2566: 2559: 2556: 2550: 2545: 2541: 2537: 2533: 2529: 2525: 2521: 2514: 2511: 2506: 2502: 2498: 2494: 2490: 2486: 2482: 2478: 2471: 2468: 2463: 2459: 2455: 2451: 2447: 2443: 2439: 2435: 2428: 2425: 2420: 2416: 2412: 2408: 2403: 2398: 2394: 2390: 2386: 2379: 2377: 2373: 2360: 2356: 2352: 2348: 2344: 2340: 2336: 2332: 2328: 2324: 2320: 2316: 2309: 2307: 2303: 2297: 2292: 2288: 2284: 2280: 2273: 2271: 2267: 2262: 2258: 2254: 2250: 2247:(4): 675–82. 2246: 2242: 2235: 2232: 2219: 2215: 2211: 2207: 2203: 2199: 2195: 2191: 2184: 2181: 2168: 2164: 2160: 2159:"pheomelanin" 2154: 2151: 2146: 2142: 2138: 2134: 2130: 2126: 2122: 2118: 2111: 2108: 2103: 2099: 2095: 2091: 2087: 2083: 2079: 2075: 2071: 2067: 2060: 2057: 2052: 2048: 2044: 2040: 2036: 2032: 2028: 2024: 2016: 2014: 2012: 2010: 2006: 2001: 1997: 1992: 1987: 1983: 1979: 1976:(3): 539–49. 1975: 1971: 1967: 1960: 1957: 1952: 1948: 1944: 1940: 1936: 1932: 1927: 1922: 1918: 1914: 1907: 1904: 1899: 1895: 1890: 1885: 1880: 1875: 1871: 1867: 1863: 1856: 1853: 1840: 1836: 1832: 1828: 1824: 1820: 1816: 1811: 1806: 1802: 1798: 1794: 1790: 1786: 1779: 1777: 1773: 1768: 1764: 1760: 1756: 1752: 1748: 1744: 1740: 1736: 1732: 1728: 1724: 1717: 1714: 1708: 1703: 1700: 1698: 1695: 1693: 1690: 1687: 1686:Melanogenesis 1684: 1682: 1679: 1677: 1674: 1672: 1669: 1666: 1663: 1661: 1658: 1656: 1653: 1652: 1647: 1645: 1643: 1639: 1634: 1630: 1622: 1620: 1618: 1614: 1610: 1606: 1602: 1598: 1594: 1590: 1586: 1582: 1578: 1574: 1570: 1566: 1562: 1558: 1557:heteropolymer 1550: 1548: 1545: 1540: 1538: 1534: 1533:accommodation 1530: 1526: 1522: 1518: 1514: 1509: 1507: 1503: 1499: 1495: 1491: 1487: 1483: 1479: 1475: 1466: 1464: 1462: 1461: 1456: 1455: 1450: 1446: 1442: 1441:archaic human 1439:with certain 1438: 1437:interbreeding 1434: 1430: 1425: 1418: 1416: 1414: 1408: 1406: 1402: 1398: 1395:of the sun's 1394: 1390: 1386: 1382: 1378: 1374: 1370: 1362: 1357: 1355: 1353: 1349: 1345: 1341: 1339: 1335: 1331: 1329: 1324: 1322: 1318: 1314: 1310: 1306: 1302: 1299:, changes in 1298: 1292: 1289: 1285: 1280: 1275: 1273: 1269: 1264: 1259: 1255: 1251: 1247: 1246: 1241: 1236: 1232: 1230: 1225: 1221: 1217: 1209: 1207: 1205: 1201: 1193: 1184: 1179: 1172: 1167: 1164:L-dopaquinone 1160: 1155: 1148: 1143: 1136: 1131: 1129: 1127: 1123: 1115: 1111: 1110: 1106: 1102: 1101: 1100: 1098: 1090: 1089: 1085: 1084: 1083: 1081: 1074: 1070: 1066: 1063: 1062: 1061: 1059: 1055: 1047: 1037: 1031: 1025: 1021: 1019: 1018:Boiling point 1016: 1015: 1011: 1009: 1008:Melting point 1006: 1005: 1001: 999: 996: 995: 988: 986: 983: 982: 955: 952: 948: 947: 942: 935: 931: 930: 928: 922: 918: 917: 910: 906: 905: 903: 901: 898: 897: 890: 886: 885: 883: 880: 876: 875: 870: 862: 858: 853: 849: 844: 840: 835: 829: 827: 819: 817: 815: 811: 807: 803: 799: 795: 786: 779: 777: 775: 771: 767: 763: 759: 755: 751: 747: 743: 739: 735: 731: 727: 723: 719: 715: 711: 707: 703: 702: 696: 694: 690: 685: 683: 679: 677: 673: 669: 665: 660: 658: 654: 649: 646: 642: 638: 633: 631: 630: 625: 624: 619: 614: 612: 608: 604: 600: 596: 592: 591:pathogenicity 588: 584: 583: 578: 574: 570: 566: 562: 558: 550: 548: 544: 542: 538: 534: 530: 525: 523: 522:adrenal gland 519: 515: 511: 507: 503: 499: 495: 491: 483: 479: 475: 468: 466: 464: 457:Trichochromes 456: 454: 452: 448: 441:Selenomelanin 440: 438: 435: 431: 422: 413: 406: 404: 397: 395: 393: 389: 385: 380: 374: 366: 364: 361: 359: 355: 351: 350:benzothiazole 347: 346:benzothiazine 342: 337: 335: 326: 321: 314: 312: 310: 305: 300: 296: 294: 290: 286: 282: 278: 270: 265: 258: 254:Melanin types 253: 251: 249: 245: 241: 236: 234: 230: 226: 222: 221:benzothiazine 218: 214: 210: 207: 203: 199: 198:melanogenesis 195: 191: 187: 183: 179: 174: 172: 168: 164: 160: 156: 153:organized as 152: 148: 145: 138: 134: 131: 130:Ancient Greek 123: 116: 83: 75: 68: 63: 54: 52:Material type 50: 43: 38: 33: 30: 26: 22: 7132:Skin anatomy 6758:Poikiloderma 6682:Lentiginosis 6594: 6375: 6203:Asparaginase 6181:oxaloacetate 6161: 6075:succinyl-CoA 6033: 6011: 5993: 5903:succinyl-CoA 5521:Kynureninase 5485: 5373: 5311:. Retrieved 5307: 5292:. Retrieved 5288: 5215: 5211: 5181: 5177: 5171: 5159:. Retrieved 5121: 5117: 5104: 5071: 5067: 5061: 5036: 5032: 5026: 5004:(3): 262–9. 5001: 4997: 4991: 4946: 4942: 4932: 4891: 4887: 4881: 4846: 4842: 4832: 4820:. Retrieved 4809: 4800: 4757: 4753: 4743: 4731:. Retrieved 4714: 4704: 4685: 4679: 4667:. Retrieved 4662: 4653: 4620: 4616: 4610: 4580:(4): 312–7. 4577: 4573: 4567: 4555:. Retrieved 4525: 4521: 4508: 4500:the original 4490: 4468:(1): 17–32. 4465: 4461: 4455: 4441: 4427: 4417:18 September 4415:. Retrieved 4411:the original 4401: 4391:– via 4385:. Retrieved 4370: 4347:the original 4337: 4327:25 September 4325:. Retrieved 4316: 4307: 4270: 4266: 4256: 4232:(1): 83–99. 4229: 4225: 4215: 4196: 4192: 4136: 4132: 4122: 4085: 4081: 4071: 4024: 3970: 3964: 3914: 3908: 3836: 3830: 3768: 3762: 3702: 3698: 3638: 3632: 3602: 3598: 3554: 3550: 3540: 3513: 3509: 3499: 3488:, retrieved 3468: 3461: 3449:. Retrieved 3429: 3422: 3397: 3393: 3386: 3375: 3365: 3353:. Retrieved 3333: 3326: 3302:(1): 72–89. 3299: 3295: 3285: 3260:10261/207451 3242: 3238: 3232: 3195: 3191: 3181: 3171:25 September 3169:. Retrieved 3144: 3140: 3130: 3113: 3109: 3103: 3084: 3080: 3070: 3045: 3041: 3035: 2990: 2986: 2976: 2959: 2956:Science News 2955: 2949: 2916: 2912: 2906: 2879: 2875: 2865: 2855:25 September 2853:. Retrieved 2844: 2835: 2803:(1): 30–41. 2800: 2796: 2786: 2743: 2739: 2729: 2717:. Retrieved 2680: 2676: 2666: 2654:. Retrieved 2639: 2625:cite journal 2613:. Retrieved 2572: 2568: 2558: 2523: 2513: 2480: 2476: 2470: 2437: 2433: 2427: 2392: 2388: 2363:. Retrieved 2322: 2318: 2286: 2282: 2244: 2240: 2234: 2224:25 September 2222:. Retrieved 2197: 2193: 2183: 2171:. Retrieved 2162: 2153: 2120: 2116: 2110: 2069: 2065: 2059: 2026: 2022: 1973: 1969: 1959: 1919:(2): 211–6. 1916: 1912: 1906: 1869: 1865: 1855: 1843:. Retrieved 1792: 1788: 1726: 1722: 1716: 1626: 1609:phagocytosis 1554: 1541: 1510: 1470: 1458: 1452: 1445:Neanderthals 1424:Early humans 1422: 1409: 1385:cell nucleus 1377:keratinocyte 1366: 1342: 1332: 1325: 1304: 1293: 1276: 1262: 1243: 1237: 1233: 1213: 1197: 1188:L-dopachrome 1119: 1094: 1077: 1051: 872:Identifiers 825: 823: 791: 773: 769: 765: 761: 757: 753: 749: 745: 741: 732:microbes in 730:radiotrophic 726:radiobiology 713: 709: 699: 697: 686: 680: 661: 650: 634: 627: 621: 615: 580: 573:heavy metals 565:UV radiation 554: 545: 526: 487: 460: 444: 427: 416:germination. 401: 376: 373:Neuromelanin 367:Neuromelanin 362: 360:is present. 338: 330: 325:substituents 301: 297: 274: 269:substituents 240:UV radiation 237: 200:, where the 186:neuromelanin 175: 151:biomolecules 143: 140: 133: 81: 80: 29: 6986:Carotenosis 6613:Reticulated 6344:development 6290:melanocytes 5883:Glutaminase 5849:glutathione 5558:(see below) 5218:(1): 9280. 4387:8 September 3977:: 474–478. 3921:: 460–468. 3775:: 1903129. 3709:: 272–283. 3645:: 105–113. 3609:(Croatia): 3394:Development 3048:: 171–219. 2993:(5): e457. 2962:(21): 325. 2365:22 February 1845:13 February 1615:sparing in 1569:antioxidant 1521:ultraviolet 1498:carotenoids 1494:riboflavins 1488:of certain 1474:sun tanning 1397:ultraviolet 1373:melanosomes 1309:pro-oxidant 1268:melanocytes 1116:→ eumelanin 1073:dopaquinone 944:Properties 662:Melanin in 657:agouti gene 635:The darker 557:cephalopods 533:basal layer 529:melanocytes 449:instead of 315:Pheomelanin 309:sun tanning 293:dopaquinone 285:melanosomes 233:Parkinson's 190:allomelanin 182:pheomelanin 171:melanocytes 128:; from 7137:Hair color 7121:Categories 7038:Skin color 7009:Dyschromia 6955:Chrysiasis 6691:Liver spot 6510:Leukoderma 6418:Melanosome 6351:Piebaldism 6342:Melanocyte 6266:Dyschromia 6190:asparagine 6166:Tyrosinase 5985:METHIONINE 5956:ISOLEUCINE 5805:Agmatinase 5498:TRYPTOPHAN 5389:acetyl-CoA 5353:Metabolism 5184:: 482–90. 4446:EntrezGene 4432:EntrezGene 4088:(4): 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