436:
36:
428:
90:
692:. MAMPs, also known as PAMPs, are microbial associated molecular patterns, small molecular elements such as proteins, carbohydrates, and lipids present on or in a given pathogen. In addition, cytokine production by melanocytes can be triggered by cytokines secreted by other nearby immune cells.
727:
Tyrosine is the non-essential amino acid precursor of melanin. Tyrosine is converted to dihydroxyphenylalanine (DOPA) via the enzyme tyrosinase. Then DOPA is polymerized into melanin. The copper-ion based enzyme-catalyzed oxidative transformation of catechol derivative dopa to light absorbing
589:
In addition to their role as UV radical scavengers, melanocytes are also part of the immune system, and are considered to be immune cells. Although the full role of melanocytes in immune response is not fully understood, melanocytes share many characteristics with
514:: those in dark-toned skin are "packaged into peri-nuclear distributed, ellipsoid" melanosomes while those light-tone skin are "assembled into clustered small, circular melanosomes". There are also differences in the quantity and relative amounts of
763:. This is mainly a consequence of alpha-MSH being secreted along with the hormone associated with reproductive tendencies in primates. Alpha-MSH is a cleavage product of ACTH that has an equal affinity for the MC1 receptor on melanocytes as ACTH.
470:
There are both basal and activated levels of melanogenesis; in general, lighter-skinned people have low basal levels of melanogenesis. Exposure to UV-B radiation causes increased melanogenesis. The purpose of melanogenesis is to protect the
814:. When a hair is lost and the hair follicle regenerates, the stem cells are activated. These stem cells develop into both keratinocyte precursors and melanoblasts - and these melanoblasts supply both hair and skin (moving into the
494:
Typically, between 1000 and 2000 melanocytes are found per square millimeter of skin or approximately 5% to 10% of the cells in the basal layer of epidermis. Although their size can vary, melanocytes are typically 7 μm in length.
750:
Numerous stimuli are able to alter melanogenesis, or the production of melanin by cultured melanocytes, although the method by which it works is not fully understood. Increased melanin production is seen in conditions where
629:, a type of MHC expressed only by certain antigen presenting cells of the immune system, when stimulated by interactions with antigen or cytokines. All cells in any given vertebrate express MHC, but most cells only express
482:
and melanogenesis are virtually identical, they are assumed to be induced by the same mechanism. The agreement of the action spectrum with the absorption spectrum of DNA points towards the formation of
166:
862:(Chloasma) - Patchy hyperpigmentation of the skin Normal number of melanocytes with increased melanin production causing hyperpigmentation. Associated with pregnancy or oral contraceptive pill use.
475:, the layer under the skin, from damage by UV radiation. The color of the melanin is black, allowing it to absorb a majority of the UV light and block it from passing through the epidermis.
1622:
Plonka PM, Passeron T, Brenner M, Tobin DJ, Shibahara S, Thomas A, Slominski A, Kadekaro AL, Hershkovitz D, Peters E, Nordlund JJ, Abdel-Malek Z, Takeda K, Paus R, Ortonne JP, Hearing VJ,
822:). There is additionally evidence that melanocyte stem cells are present in cutaneous nerves, with nerve signals causing these cells to differentiate into melanocytes for the skin.
916:
656:
In addition to presenting antigen, one of the roles of melanocytes in the immune response is cytokine production. Melanocytes express many proinflammatory cytokines including
911:
842:
610:. Although melanocytes are dendritic in form and share many characteristics with dendritic cells, they derive from different cell lineages. Dendritic cells are derived from
1297:
Tadokoro, Taketsugu; Yamaguchi, Yuji; Batzer, Jan; Coelho, Sergio G.; Zmudzka, Barbara Z.; Miller, Sharon A.; Wolber, Rainer; Beer, Janusz Z.; Hearing, Vincent J. (2005).
570:
to name some. In all, already 17 types of oculocutaneous albinism have been recognized. Each gene is related to different protein having a role in pigment production.
1518:
Montoliu L, Grønskov K, Wei AH, Martínez-García M, Fernández A, Arveiler B, Morice-Picard F, Riazuddin S, Suzuki T, Ahmed ZM, Rosenberg T, Li W (January 2014).
707:, but they use their dendrites to interact with cells in other layers, and to capture pathogens that enter the epidermis. They likely work in concert with both
467:. This melanogenesis leads to a long-lasting pigmentation, which is in contrast to the pigmentation that originates from oxidation of already-existing melanin.
308:
411:
to induce pigmentation. Thus darker skin tones have more melanosomes present than lighter skin tones. Functionally, melanin serves as protection against
790:
will trigger melanogenesis and cause the melanocyte to produce melanosomes, which are then transferred by dendrites to the top layer of keratinocytes.
689:
1157:
Nichols SE, Reams WM (March 1960). "The occurrence and morphogenesis of melanocytes in the connective tissues of the PET/MCV mouse strain".
1578:
Gasque P, Jaffar-Bandjee MC (October 2015). "The immunology and inflammatory responses of human melanocytes in infectious diseases".
856:- Normal number of melanocytes, but decreased melanin production due to decreased tyrosinase activity or defective tyrosine transport
1114:
67:
1675:
Mackintosh JA (July 2001). "The antimicrobial properties of melanocytes, melanosomes and melanin and the evolution of black skin".
315:
574:
677:
1856:"Enhancement of DNA repair in human skin cells by thymidine dinucleotides: evidence for a p53-mediated mammalian SOS response"
523:
1774:
Tapia CV, Falconer M, Tempio F, Falcón F, López M, Fuentes M, Alburquenque C, Amaro J, Bucarey SA, Di Nardo A (July 2014).
984:
Cramer SF (February 1991). "The origin of epidermal melanocytes. Implications for the histogenesis of nevi and melanomas".
935:
303:
1995:
Millington GW (May 2006). "Proopiomelanocortin (POMC): the cutaneous roles of its melanocortin products and receptors".
681:
484:
1254:
Parrish JA, Jaenicke KF, Anderson RR (August 1982). "Erythema and melanogenesis action spectra of normal human skin".
752:
673:
435:
2076:
1184:
Theriault LL, Hurley LS (October 1970). "Ultrastructure of developing melanosomes in C57 black and pallid mice".
736:
is clearly seen following the polymerization to melanin, the color of the pigment ranges from red to dark brown.
622:. As such, although morphologically and functionally similar, melanocytes and dendritic cells are not the same.
291:
279:
946:
611:
2066:
2071:
1103:
Mayeaux EJ, Cox JT, et al. (American
Society for Colposcopy and Cervical Pathology) (2011-12-28).
539:
typically have a very low level of melanin production. Albinism is often but not always related to the
1815:
Abdel-Malek Z, Scott MC, Suzuki I, Tada A, Im S, Lamoreux L, Ito S, Barsh G, Hearing VJ (2000-01-01).
296:
2061:
1867:
1684:
1104:
865:
760:
756:
657:
645:, and melanocytes. Importantly, melanocytes stimulated by cytokines express surface proteins such as
1299:"Mechanisms of Skin Tanning in Different Racial/Ethnic Groups in Response to Ultraviolet Radiation"
901:
870:
498:
Both lightly and darkly pigmented skin contain similar numbers of melanocytes, with difference in
472:
2020:
1623:
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368:
1056:"The Effects of Genotype and Cell Environment on Melanoblast Differentiation in the House Mouse"
2012:
1983:
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1776:"Melanocytes and melanin represent a first line of innate immunity against Candida albicans"
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680:. Like other immune cells, melanocytes secrete these cytokines in response to activation of
499:
352:
175:
113:
1219:
Agar N, Young AR (April 2005). "Melanogenesis: a photoprotective response to DNA damage?".
969:
771:
712:
58:
850:- Decreased number of melanocytes due to autoimmune destruction causing decreased melanin
1966:
Ito S (June 2003). "The IFPCS presidential lecture: a chemist's view of melanogenesis".
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348:
344:
1351:"Was skin cancer a selective force for black pigmentation in early hominin evolution?"
1011:
Barden H, Levine S (June 1983). "Histochemical observations on rodent brain melanin".
2055:
2048: – Histology Learning System at Boston University - "Integument: pigmented skin"
2008:
1979:
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637:, is found only on "professional" antigen presenting cells such as dendritic cells,
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1817:"The melanocortin-1 receptor is a key regulator of human cutaneous pigmentation"
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Proceedings of the
National Academy of Sciences of the United States of America
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enzyme. Tyrosinase is required for melanocytes to produce melanin from the
526:
and ACTH peptides that are produced from the precursor proopiomelanocortin.
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2042: – Histology Learning System at Boston University - "Eye: fovea, RPE"
1899:
1275:
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532:
is a skin disease where people lack melanin in certain areas in the skin.
522:. Pigmentation including tanning is under hormonal control, including the
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Eller MS, Maeda T, Magnoni C, Atwal D, Gilchrest BA (November 1997).
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in addition to MHC class II, allowing for co-stimulation of T cells.
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384:
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1913:
Cichorek M, Wachulska M, Stasiewicz A, Tymińska A (February 2013).
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554:. Albinism may be caused by a number of other genes as well, like
434:
426:
380:
267:
1520:"Increasing the complexity: new genes and new types of albinism"
806:. In adults, stem cells are contained in the bulge area of the
685:
456:
388:
376:
360:
244:
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have a buildup of melanin granules due to abnormal function of
787:
699:
to be sentinels against harmful pathogens. They reside in the
356:
29:
1130:
Mintz B (1971). "Clonal basis of mammalian differentiation".
443:
Through a process called melanogenesis, melanocytes produce
1344:
1342:
782:. When ultraviolet rays penetrate the skin and damage DNA,
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196:
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155:
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119:
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1725:"Skin Immune Landscape: Inside and Outside the Organism"
1355:
Proceedings of the Royal
Society B: Biological Sciences
54:
917:
List of human cell types derived from the germ layers
778:
are organized as a cap protecting the nucleus of the
214:
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1628:"What are melanocytes really doing all day long...?"
399:. Once synthesized, melanin is contained in special
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912:
List of distinct cell types in the adult human body
843:
Melanocytic tumors of uncertain malignant potential
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1159:Journal of Embryology and Experimental Morphology
1132:Symposia of the Society for Experimental Biology
986:Archives of Pathology & Laboratory Medicine
618:. Melanocytes on the other hand originate from
8:
1915:"Skin melanocytes: biology and development"
786:dinucleotide (pTpT) fragments from damaged
695:Melanocytes are ideally positioned in the
439:Micrograph of melanocytes in the epidermis
88:
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68:Learn how and when to remove this message
1723:Abdallah F, Mijouin L, Pichon C (2017).
719:, to contribute to the immune response.
927:
625:Melanocytes are capable of expressing
415:. Melanocytes also have a role in the
319:
79:
1997:Clinical and Experimental Dermatology
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1109:. Lippincott Williams & Wilkins.
7:
1524:Pigment Cell & Melanoma Research
1303:Journal of Investigative Dermatology
1106:Modern Colposcopy Textbook and Atlas
1054:Markert CL, Silvers WK (May 1956).
770:that package the chemical inside a
407:which can be transported to nearby
27:Melanin-producing cells of the skin
1919:Postepy Dermatologii I Alergologii
1268:10.1111/j.1751-1097.1982.tb04362.x
715:, both of which are also actively
502:due to differences the packing of
447:, which is a pigment found in the
55:improve coverage for other species
25:
347:located in the bottom layer (the
2009:10.1111/j.1365-2230.2006.02128.x
1980:10.1034/j.1600-0749.2003.00037.x
1644:10.1111/j.1600-0625.2009.00912.x
1316:10.1111/j.0022-202x.2005.23760.x
606:; and production and release of
316:Anatomical terms of microanatomy
174:
112:
34:
1499:. National Institutes of Health
1477:. National Institutes of Health
1455:. National Institutes of Health
1433:. National Institutes of Health
1411:. National Institutes of Health
1256:Photochemistry and Photobiology
1833:10.1034/j.1600-0749.13.s8.28.x
1677:Journal of Theoretical Biology
970:Merriam-Webster.com Dictionary
834:- Cancer affecting melanocytes
598:capabilities; presentation of
57:and discuss this issue on the
1:
1309:(6). Elsevier BV: 1326–1332.
682:Pattern Recognition Receptors
485:cyclobutane pyrimidine dimers
478:Since the action spectrum of
1233:10.1016/j.mrfmmm.2004.11.016
1198:10.1016/0012-1606(70)90098-9
1025:10.1016/0361-9230(83)90218-6
755:(ACTH) is elevated, such as
431:Illustration of a melanocyte
1349:Greaves, Mel (2014-04-22).
753:adrenocorticotropic hormone
535:People with oculocutaneous
2093:
1592:10.1016/j.jinf.2015.06.006
743:
703:, the lowest layer of the
633:. The other class of MHC,
395:primarily responsible for
355:, the middle layer of the
2046:Histology image: 08103loa
2040:Histology image: 07903loa
1729:Mediators of Inflammation
1072:10.1093/genetics/41.3.429
802:of the melanocyte is the
585:Role in the immune system
314:
87:
49:excessively human-centric
1881:10.1073/pnas.94.23.12627
1632:Experimental Dermatology
1580:The Journal of Infection
612:hematopoietic stem cells
575:Chédiak–Higashi syndrome
1931:10.5114/pdia.2013.33376
1013:Brain Research Bulletin
947:Oxford University Press
688:(TLR4) which recognize
594:: branched morphology;
1697:10.1006/jtbi.2001.2331
1368:10.1098/rspb.2013.2955
440:
432:
1968:Pigment Cell Research
1821:Pigment Cell Research
1186:Developmental Biology
943:UK English Dictionary
826:Clinical significance
438:
430:
1742:10.1155/2017/5095293
686:Toll Like Receptor 4
391:. Melanin is a dark
1872:1997PNAS...9412627E
1827:(Suppl 8): 156–62.
1689:2001JThBi.211..101M
902:Mole (skin marking)
871:Nevus depigmentosus
43:This article about
1793:10.1093/mmy/myu026
1626:(September 2009).
1537:10.1111/pcmr.12167
1361:(1781): 20132955.
973:. Merriam-Webster.
838:Melanocytic tumors
734:indole-5,6-quinone
620:neural crest cells
441:
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369:vaginal epithelium
297:H2.00.03.0.01016
1221:Mutation Research
907:Tanning activator
808:outer root sheath
761:Cushing's disease
489:direct DNA damage
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16:(Redirected from
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2077:Epithelial cells
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949:. Archived from
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713:Langerhans cells
543:gene coding the
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684:(PRRs) such as
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1743:
1738:
1734:
1730:
1726:
1719:
1717:
1715:
1711:
1706:
1702:
1698:
1694:
1690:
1686:
1683:(2): 101–13.
1682:
1678:
1671:
1668:
1663:
1659:
1654:
1649:
1645:
1641:
1637:
1633:
1629:
1625:
1618:
1616:
1614:
1612:
1610:
1606:
1601:
1597:
1593:
1589:
1586:(4): 413–21.
1585:
1581:
1574:
1572:
1570:
1568:
1566:
1564:
1560:
1555:
1551:
1547:
1543:
1538:
1533:
1529:
1525:
1521:
1514:
1511:
1498:
1492:
1489:
1476:
1470:
1467:
1454:
1448:
1445:
1432:
1426:
1423:
1410:
1404:
1401:
1396:
1392:
1387:
1382:
1378:
1374:
1369:
1364:
1360:
1356:
1352:
1345:
1343:
1339:
1334:
1330:
1326:
1322:
1317:
1312:
1308:
1304:
1300:
1293:
1290:
1285:
1281:
1277:
1273:
1269:
1265:
1262:(2): 187–91.
1261:
1257:
1250:
1247:
1242:
1238:
1234:
1230:
1226:
1222:
1215:
1212:
1207:
1203:
1199:
1195:
1192:(2): 261–75.
1191:
1187:
1180:
1177:
1172:
1168:
1164:
1160:
1153:
1150:
1145:
1141:
1137:
1133:
1126:
1123:
1118:
1116:9781451153835
1112:
1108:
1107:
1099:
1096:
1091:
1087:
1082:
1077:
1073:
1069:
1066:(3): 429–50.
1065:
1061:
1057:
1050:
1047:
1042:
1038:
1034:
1030:
1026:
1022:
1019:(6): 847–51.
1018:
1014:
1007:
1004:
999:
995:
991:
987:
980:
977:
972:
971:
966:
960:
957:
952:
948:
944:
942:
937:
931:
928:
922:
918:
915:
913:
910:
908:
905:
903:
900:
898:
895:
892:
888:
885:(the pigment
884:
883:Chromatophore
881:
880:
876:
872:
869:
867:
864:
861:
858:
855:
852:
849:
846:
844:
841:
839:
836:
833:
830:
829:
825:
823:
821:
817:
813:
809:
805:
801:
793:
791:
789:
785:
781:
777:
773:
769:
764:
762:
758:
754:
747:
739:
737:
735:
731:
723:Melanogenesis
722:
720:
718:
714:
710:
709:keratinocytes
706:
702:
698:
693:
691:
687:
683:
679:
675:
671:
667:
663:
659:
654:
652:
648:
644:
640:
636:
632:
628:
623:
621:
617:
613:
609:
605:
601:
597:
593:
584:
582:
580:
576:
571:
569:
565:
561:
557:
553:
550:
546:
542:
538:
533:
531:
527:
525:
521:
517:
513:
512:keratinocytes
509:
505:
501:
496:
492:
490:
486:
481:
476:
474:
468:
466:
462:
458:
454:
450:
446:
437:
429:
422:
420:
418:
417:immune system
414:
410:
409:keratinocytes
406:
402:
398:
394:
390:
386:
382:
378:
374:
370:
366:
362:
358:
354:
350:
346:
342:
338:
334:
323:
317:
313:
310:
307:
305:
301:
298:
295:
293:
289:
286:
283:
281:
277:
274:
271:
269:
265:
260:
256:
253:
249:
246:
243:
239:
236:
233:
231:
227:
222:
167:
160:
110:
108:Pronunciation
106:
101:
97:
91:
86:
81:
72:
69:
60:
56:
50:
46:
41:
32:
31:
19:
18:Melanogenesis
2072:Skin anatomy
2000:
1996:
1974:(3): 230–6.
1971:
1967:
1925:(1): 30–41.
1922:
1918:
1908:
1863:
1859:
1849:
1824:
1820:
1810:
1783:
1779:
1769:
1732:
1728:
1680:
1676:
1670:
1635:
1631:
1583:
1579:
1527:
1523:
1513:
1501:. Retrieved
1491:
1479:. Retrieved
1469:
1457:. Retrieved
1447:
1435:. Retrieved
1425:
1413:. Retrieved
1403:
1358:
1354:
1306:
1302:
1292:
1259:
1255:
1249:
1224:
1220:
1214:
1189:
1185:
1179:
1162:
1158:
1152:
1135:
1131:
1125:
1105:
1098:
1063:
1059:
1049:
1016:
1012:
1006:
992:(2): 115–9.
989:
985:
979:
968:
965:"Melanocyte"
959:
951:the original
939:
936:"Melanocyte"
930:
891:poikilotherm
797:
780:keratinocyte
765:
749:
746:Melanocortin
726:
694:
655:
627:MHC Class II
624:
588:
579:microtubules
573:People with
572:
567:
563:
559:
555:
540:
534:
528:
497:
493:
477:
469:
461:nasal cavity
442:
413:UV radiation
341:neural crest
332:
331:
272:
235:Neural crest
64:
48:
2062:Human cells
1735:: 5095293.
1530:(1): 11–8.
816:basal layer
804:melanoblast
776:melanosomes
740:Stimulation
730:dopaquinone
639:macrophages
631:MHC class I
616:bone marrow
520:pheomelanin
508:melanosomes
405:melanosomes
339:-producing
333:Melanocytes
273:melanocytus
262:Identifiers
2056:Categories
1138:: 345–70.
923:References
794:Stem cells
717:phagocytic
596:phagocytic
549:amino acid
545:tyrosinase
500:skin color
473:hypodermis
401:organelles
397:skin color
257:production
83:Melanocyte
1453:"SLC45A2"
1377:0962-8452
1325:0022-202X
1165:: 24–32.
897:Eye color
889:found in
887:cell type
820:epidermis
800:precursor
784:thymidine
757:Addison's
705:epidermis
697:epidermis
608:cytokines
516:eumelanin
506:into the
504:eumelanin
487:(CPDs) -
465:inner ear
365:inner ear
353:epidermis
343:-derived
230:Precursor
59:talk page
2025:25213876
2017:16681590
1988:12753395
1949:24278043
1841:11041375
1802:24934806
1761:29180836
1705:11419954
1662:19659579
1600:26092350
1546:24066960
1503:25 March
1481:25 March
1459:25 March
1437:25 March
1395:24573849
1333:15955111
1284:38940583
1241:15748643
1171:14426921
1090:17247639
1060:Genetics
893:animals)
877:See also
854:Albinism
848:Vitiligo
832:Melanoma
768:vesicles
635:Class II
600:antigens
552:tyrosine
537:albinism
530:Vitiligo
423:Function
373:meninges
251:Function
241:Location
1940:3834696
1900:9356500
1868:Bibcode
1752:5664322
1685:Bibcode
1653:2792575
1554:7305884
1475:"TYRP1"
1415:23 June
1386:3953838
1276:7122713
1206:5476812
1144:4940552
1081:1209793
1041:4783099
1033:6616275
998:1992974
860:Melasma
818:of the
643:B cells
614:in the
604:T-cells
560:SLC45A2
480:sunburn
445:melanin
403:called
393:pigment
385:mammals
363:), the
337:melanin
285:D008544
255:Melanin
103:Details
96:melanin
53:Please
47:may be
45:biology
2023:
2015:
1986:
1947:
1937:
1898:
1888:
1839:
1800:
1759:
1749:
1703:
1660:
1650:
1598:
1552:
1544:
1497:"HPS1"
1431:"OCA2"
1393:
1383:
1375:
1331:
1323:
1282:
1274:
1239:
1204:
1169:
1142:
1113:
1088:
1078:
1039:
1031:
996:
941:Lexico
774:. The
676:, and
566:, and
463:, and
379:, and
2021:S2CID
1891:25061
1550:S2CID
1409:"TYR"
1280:S2CID
1037:S2CID
690:MAMPs
678:TGF-β
674:TNF-α
651:ICAM1
564:TYRP1
389:birds
381:heart
377:bones
359:(the
345:cells
320:[
309:70545
268:Latin
162:
2013:PMID
1984:PMID
1945:PMID
1896:PMID
1837:PMID
1798:PMID
1757:PMID
1733:2017
1701:PMID
1658:PMID
1596:PMID
1542:PMID
1505:2016
1483:2016
1461:2016
1439:2016
1417:2013
1391:PMID
1373:ISSN
1329:PMID
1321:ISSN
1272:PMID
1237:PMID
1202:PMID
1167:PMID
1140:PMID
1111:ISBN
1086:PMID
1029:PMID
994:PMID
798:The
759:and
711:and
670:IL-8
666:IL-6
662:IL-3
658:IL-1
649:and
647:CD40
568:HPS1
556:OCA2
518:and
457:hair
453:eyes
449:skin
387:and
361:uvea
335:are
280:MeSH
245:Skin
2005:doi
1976:doi
1935:PMC
1927:doi
1886:PMC
1876:doi
1829:doi
1788:doi
1747:PMC
1737:doi
1693:doi
1681:211
1648:PMC
1640:doi
1588:doi
1532:doi
1381:PMC
1363:doi
1359:281
1311:doi
1307:124
1264:doi
1229:doi
1225:571
1194:doi
1076:PMC
1068:doi
1021:doi
990:115
810:of
788:DNA
732:to
602:to
541:TYR
524:MSH
510:of
357:eye
304:FMA
172:or
2058::
2019:.
2011:.
2001:31
1999:.
1982:.
1972:16
1970:.
1943:.
1933:.
1923:30
1921:.
1917:.
1894:.
1884:.
1874:.
1864:94
1862:.
1858:.
1835:.
1825:13
1823:.
1819:.
1796:.
1784:52
1782:.
1778:.
1755:.
1745:.
1731:.
1727:.
1713:^
1699:.
1691:.
1679:.
1656:.
1646:.
1636:18
1634:.
1630:.
1608:^
1594:.
1584:71
1582:.
1562:^
1548:.
1540:.
1528:27
1526:.
1522:.
1389:.
1379:.
1371:.
1357:.
1353:.
1341:^
1327:.
1319:.
1305:.
1301:.
1278:.
1270:.
1260:36
1258:.
1235:.
1223:.
1200:.
1190:23
1188:.
1161:.
1136:25
1134:.
1084:.
1074:.
1064:41
1062:.
1058:.
1035:.
1027:.
1017:10
1015:.
988:.
967:.
945:.
938:.
672:,
668:,
664:,
660:,
641:,
581:.
562:,
558:,
491:.
459:,
455:,
451:,
419:.
375:,
371:,
367:,
292:TH
221:-/
218:oʊ
212:,-
206:aɪ
159:-/
156:oʊ
150:,-
144:aɪ
2027:.
2007::
1990:.
1978::
1951:.
1929::
1902:.
1878::
1870::
1843:.
1831::
1804:.
1790::
1763:.
1739::
1707:.
1695::
1687::
1664:.
1642::
1602:.
1590::
1556:.
1534::
1507:.
1485:.
1463:.
1441:.
1419:.
1397:.
1365::
1335:.
1313::
1286:.
1266::
1243:.
1231::
1208:.
1196::
1173:.
1163:8
1146:.
1119:.
1092:.
1070::
1043:.
1023::
1000:.
324:]
215:n
209:t
203:s
200:ˌ
197:ə
194:n
191:ə
188:l
185:ɛ
182:m
179:ˈ
176:/
153:n
147:t
141:s
138:ˌ
135:ə
132:n
129:æ
126:l
123:ˈ
120:ə
117:m
114:/
71:)
65:(
61:.
51:.
20:)
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