{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,10]],"date-time":"2026-10-10T15:00:42Z","timestamp":1791644442822,"version":"4.3.4"},"reference-count":85,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2026,1,12]],"date-time":"2026-01-12T00:00:00Z","timestamp":1768176000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":["frontiersin.org"],"crossmark-restriction":true},"short-container-title":["Front. Cell Dev. Biol."],"abstract":"<jats:p>Lysosome-related organelles (LROs) encompass specialized intracellular compartments that share features with lysosomes while fulfilling distinct physiological roles, with melanosomes representing the best-studied example. Melanosome biogenesis relies on coordinated trafficking, sorting, and membrane remodeling mechanisms that diverge from the canonical endolysosomal pathways. These organelles ultimately serve as the primary sites of melanin synthesis and deposition. In the skin, melanin is produced by melanocytes and transferred to keratinocytes, where it achieves its essential photoprotective role. Melanin is a remarkably diverse and ancient polymer, with eumelanin, pheomelanin, and neuromelanin constituting the major mammalian forms. Understanding melanin biology also requires tracing the origins of melanocytes, which were once thought to derive exclusively from the neural crest but are now known to arise from multiple embryonic lineages. This expanded view of melanocyte ontogeny has revealed unexpected pigment cell populations in several internal organs. Beyond these developmental aspects, melanin performs multifaceted physiological functions that extend far beyond photoprotection of the skin. Here, we discuss the current knowledge on the origin of melanosomes from endosomal precursors, the transfer of melanin from melanocytes to keratinocytes, and its fate in these recipient cells within the epidermis. Additionally, the intriguing mysteries surrounding melanosomes in the retinal pigment epithelium are addressed, as well as the broader diversity, origins, and physiological roles of melanin in other cell types. Taken together, these perspectives highlight the melanosome as both a model LRO and an organellar hub for deciphering melanin diversity, cellular origins, and the wide-ranging physiological roles of this pigment in vertebrate biology.<\/jats:p>","DOI":"10.3389\/fcell.2025.1758081","type":"journal-article","created":{"date-parts":[[2026,1,12]],"date-time":"2026-01-12T08:12:56Z","timestamp":1768205576000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":4,"title":["Insights into lysosome-related organelle biogenesis: melanosome as a model organelle"],"prefix":"10.3389","volume":"13","author":[{"given":"Duarte C.","family":"Barral","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"C\u00e9dric","family":"Delevoye","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Lionel","family":"Larue","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Miguel C.","family":"Seabra","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Gra\u00e7a","family":"Raposo","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Subba Rao Gangi","family":"Setty","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1965","published-online":{"date-parts":[[2026,1,12]]},"reference":[{"key":"B1","doi-asserted-by":"publisher","first-page":"366","DOI":"10.1016\/j.cell.2009.07.049","article-title":"Schwann cell precursors from nerve innervation are a cellular origin of melanocytes in skin","volume":"139","author":"Adameyko","year":"2009","journal-title":"Cell"},{"key":"B2","doi-asserted-by":"publisher","first-page":"5622","DOI":"10.1073\/pnas.1600108113","article-title":"TPC2 controls pigmentation by regulating melanosome pH and size","volume":"113","author":"Ambrosio","year":"2016","journal-title":"Proc. Natl. Acad. Sci. U. S. A."},{"key":"B3","doi-asserted-by":"publisher","first-page":"332","DOI":"10.1034\/j.1600-0749.2000.130505.x","article-title":"Small gtpase rab3A is associated with melanosomes in melanoma cells","volume":"13","author":"Araki","year":"2000","journal-title":"Pigment. Cell Res."},{"key":"B4","doi-asserted-by":"publisher","first-page":"1097","DOI":"10.1098\/rsif.2009.0174","article-title":"Melanocyte pigmentation stiffens murine cardiac tricuspid valve leaflet","volume":"6","author":"Balani","year":"2009","journal-title":"J. R. Soc. Interface"},{"key":"B5","doi-asserted-by":"publisher","first-page":"8630","DOI":"10.3390\/ijms26178630","article-title":"Melanosome transport and processing in skin pigmentation: mechanisms and targets for pigmentation modulation","volume":"26","author":"Bao","year":"2025","journal-title":"Int. J. Mol. Sci."},{"key":"B6","doi-asserted-by":"publisher","DOI":"10.1038\/s41556-025-01817-4","article-title":"Keratin intermediate filaments mechanically position melanin pigments for genome photoprotection","author":"Benito-Mart\u00ednez","year":"2025","journal-title":"Nat. Cell Biol"},{"key":"B7","doi-asserted-by":"publisher","first-page":"11289","DOI":"10.3390\/ijms241411289","article-title":"Melanin's journey from melanocytes to keratinocytes: uncovering the molecular mechanisms of melanin transfer and processing","volume":"24","author":"Bento-Lopes","year":"2023","journal-title":"Int. J. Mol. Sci."},{"key":"B8","doi-asserted-by":"publisher","first-page":"510","DOI":"10.1086\/500850","article-title":"Loss-of-function mutations in the keratin 5 gene lead to dowling-degos disease","volume":"78","author":"Betz","year":"2006","journal-title":"Am. J. Hum. Genet."},{"key":"B9","doi-asserted-by":"publisher","first-page":"102669","DOI":"10.1016\/j.jbc.2022.102669","article-title":"OCA7 is a melanosome membrane protein that defines pigmentation by regulating early stages of melanosome biogenesis","volume":"298","author":"Beyers","year":"2022","journal-title":"J. Biol. Chem."},{"key":"B10","doi-asserted-by":"publisher","first-page":"1438","DOI":"10.3390\/ijms17091438","article-title":"PMEL amyloid fibril formation: the bright steps of pigmentation","volume":"17","author":"Bissig","year":"2016","journal-title":"Int. J. Mol. Sci."},{"key":"B11","doi-asserted-by":"publisher","first-page":"61","DOI":"10.1016\/j.exer.2014.01.016","article-title":"Studying melanin and lipofuscin in RPE cell culture models","volume":"126","author":"Boulton","year":"2014","journal-title":"Exp. Eye Res."},{"key":"B12","doi-asserted-by":"publisher","first-page":"404","DOI":"10.1111\/tra.12646","article-title":"The road to lysosome-related organelles: insights from Hermansky-Pudlak syndrome and other rare diseases","volume":"20","author":"Bowman","year":"2019","journal-title":"Traffic"},{"key":"B13","doi-asserted-by":"publisher","first-page":"539","DOI":"10.1111\/j.1751-1097.2007.00226.x","article-title":"The protective role of melanin against UV damage in human skin","volume":"84","author":"Brenner","year":"2008","journal-title":"Photochem Photobiol."},{"key":"B14","doi-asserted-by":"publisher","first-page":"292","DOI":"10.1016\/j.amjmed.2015.10.015","article-title":"Mining retrospective data for virtual prospective drug repurposing: L-DOPA and age-related macular degeneration","volume":"129","author":"Brilliant","year":"2016","journal-title":"Am. J. Med."},{"key":"B15","doi-asserted-by":"publisher","first-page":"464","DOI":"10.1111\/j.1755-148X.2008.00459.x","article-title":"Timeline and distribution of melanocyte precursors in the mouse heart","volume":"21","author":"Brito","year":"2008","journal-title":"Pigment. Cell Melanoma Res."},{"key":"B16","doi-asserted-by":"publisher","first-page":"16","DOI":"10.4161\/sgtp.22349","article-title":"Cell type-specific Rab32 and Rab38 cooperate with the ubiquitous lysosome biogenesis machinery to synthesize specialized lysosome-related organelles","volume":"4","author":"Bultema","year":"2013","journal-title":"Small GTPases"},{"key":"B17","first-page":"2864","article-title":"Dynamic analyses reveal cytoprotection by RPE melanosomes against non-photic stress","volume":"17","author":"Burke","year":"2011","journal-title":"Mol. Vis."},{"key":"B18","doi-asserted-by":"publisher","first-page":"813","DOI":"10.1046\/j.1523-1747.2003.12481.x","article-title":"Role of cytoplasmic dynein in perinuclear aggregation of phagocytosed melanosomes and supranuclear melanin cap formation in human keratinocytes","volume":"121","author":"Byers","year":"2003","journal-title":"J. Invest Dermatol"},{"key":"B19","doi-asserted-by":"publisher","first-page":"1736","DOI":"10.1038\/sj.jid.5700760","article-title":"Requirement of dynactin p150(Glued) subunit for the functional integrity of the keratinocyte microparasol","volume":"127","author":"Byers","year":"2007","journal-title":"J. Invest Dermatol"},{"key":"B20","doi-asserted-by":"publisher","first-page":"100139","DOI":"10.1016\/j.xjidi.2022.100139","article-title":"RAB3A regulates melanin exocytosis and transfer induced by keratinocyte-conditioned medium","volume":"2","author":"Cabaco","year":"2022","journal-title":"JID Innov."},{"key":"B21","doi-asserted-by":"publisher","first-page":"3143","DOI":"10.3390\/ijms22063143","article-title":"Melanin distribution in human skin: influence of Cytoskeletal, polarity, and centrosome-related machinery of stratum basale keratinocytes","volume":"22","author":"Castellano-Pellicena","year":"2021","journal-title":"Int. J. Mol. Sci."},{"key":"B22","doi-asserted-by":"publisher","first-page":"406","DOI":"10.1203\/PDR.0b013e31822a59f5","article-title":"Long-chain polyunsaturated fatty acid supplementation in infancy reduces heart rate and positively affects distribution of attention","volume":"70","author":"Colombo","year":"2011","journal-title":"Pediatr. Res."},{"key":"B23","doi-asserted-by":"publisher","first-page":"dev194407","DOI":"10.1242\/dev.194407","article-title":"Stabilization of beta-catenin promotes melanocyte specification at the expense of the Schwann cell lineage","volume":"149","author":"Colombo","year":"2022","journal-title":"Development"},{"key":"B24","doi-asserted-by":"publisher","first-page":"637","DOI":"10.1016\/j.jid.2017.09.042","article-title":"Melanin transferred to keratinocytes resides in nondegradative endocytic compartments","volume":"138","author":"Correia","year":"2018","journal-title":"J. Invest Dermatol"},{"key":"B25","doi-asserted-by":"publisher","first-page":"336","DOI":"10.1186\/s12967-024-05113-x","article-title":"Melanocytes in regenerative medicine applications and disease modeling","volume":"22","author":"Coutant","year":"2024","journal-title":"J. Transl. Med."},{"key":"B26","doi-asserted-by":"publisher","first-page":"606","DOI":"10.1111\/j.1600-0749.2006.00338.x","article-title":"Relationship between skin response to ultraviolet exposure and skin color type","volume":"19","author":"Del Bino","year":"2006","journal-title":"Pigment. Cell Res."},{"key":"B27","doi-asserted-by":"publisher","first-page":"247","DOI":"10.1083\/jcb.200907122","article-title":"AP-1 and KIF13A coordinate endosomal sorting and positioning during melanosome biogenesis","volume":"187","author":"Delevoye","year":"2009","journal-title":"J. Cell Biol."},{"key":"B28","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1016\/j.cub.2015.11.020","article-title":"BLOC-1 brings together the actin and microtubule cytoskeletons to generate recycling endosomes","volume":"26","author":"Delevoye","year":"2016","journal-title":"Curr. Biol."},{"key":"B29","doi-asserted-by":"publisher","first-page":"147","DOI":"10.1016\/j.ceb.2019.05.003","article-title":"Lysosome-related organelles as functional adaptations of the endolysosomal system","volume":"59","author":"Delevoye","year":"2019","journal-title":"Curr. Opin. Cell Biol."},{"key":"B30","doi-asserted-by":"publisher","first-page":"563","DOI":"10.1083\/jcb.201410026","article-title":"BLOC-2 targets recycling endosomal tubules to melanosomes for cargo delivery","volume":"209","author":"Dennis","year":"2015","journal-title":"J. Cell Biol."},{"key":"B31","doi-asserted-by":"publisher","first-page":"293","DOI":"10.1083\/jcb.201605090","article-title":"BLOC-1 and BLOC-3 regulate VAMP7 cycling to and from melanosomes via distinct tubular transport carriers","volume":"214","author":"Dennis","year":"2016","journal-title":"J. Cell Biol."},{"key":"B32","doi-asserted-by":"publisher","first-page":"112","DOI":"10.1111\/j.1755-148X.2009.00662.x","article-title":"Inducible expression of (V600E) braf using tyrosinase-driven Cre recombinase results in embryonic lethality","volume":"23","author":"Dhomen","year":"2010","journal-title":"Pigment. Cell Melanoma Res."},{"key":"B33","doi-asserted-by":"publisher","first-page":"4027","DOI":"10.1091\/mbc.e06-05-0379","article-title":"BLOC-1 interacts with BLOC-2 and the AP-3 complex to facilitate protein trafficking on endosomes","volume":"17","author":"Di Pietro","year":"2006","journal-title":"Mol. Biol. Cell"},{"key":"B34","first-page":"195","article-title":"Aging human RPE: morphometric analysis of macular, equatorial, and peripheral cells","volume":"25","author":"Feeney-Burns","year":"1984","journal-title":"Invest Ophthalmol. Vis. Sci."},{"key":"B35","doi-asserted-by":"publisher","first-page":"786","DOI":"10.1111\/pcmr.12982","article-title":"Genetics of non-syndromic and syndromic oculocutaneous albinism in human and mouse","volume":"34","author":"Fernandez","year":"2021","journal-title":"Pigment. Cell Melanoma Res."},{"key":"B36","doi-asserted-by":"publisher","first-page":"12432","DOI":"10.1074\/jbc.C200005200","article-title":"Slac2-a\/melanophilin, the missing link between Rab27 and myosin Va: implications of a tripartite protein complex for melanosome transport","volume":"277","author":"Fukuda","year":"2002","journal-title":"J. Biol. Chem."},{"key":"B37","doi-asserted-by":"publisher","first-page":"458","DOI":"10.1034\/j.1600-0749.2000.130608.x","article-title":"Melanosome capping of keratinocytes in pigmented reconstructed epidermis--effect of ultraviolet radiation and 3-isobutyl-1-methyl-xanthine on melanogenesis","volume":"13","author":"Gibbs","year":"2000","journal-title":"Pigment. Cell Res."},{"key":"B38","doi-asserted-by":"publisher","first-page":"149","DOI":"10.3389\/fnana.2015.00149","article-title":"Meningeal melanocytes in the mouse: distribution and dependence on mitf","volume":"9","author":"Gudjohnsen","year":"2015","journal-title":"Front. Neuroanat."},{"key":"B39","doi-asserted-by":"publisher","first-page":"3111","DOI":"10.1242\/jcs.010207","article-title":"Rab27a and MyoVa are the primary Mlph interactors regulating melanosome transport in melanocytes","volume":"120","author":"Hume","year":"2007","journal-title":"J. Cell Sci."},{"key":"B40","doi-asserted-by":"publisher","first-page":"19726","DOI":"10.1073\/pnas.0803488105","article-title":"Electron tomography of early melanosomes: implications for melanogenesis and the generation of fibrillar amyloid sheets","volume":"105","author":"Hurbain","year":"2008","journal-title":"Proc. Natl. Acad. Sci. U. S. A."},{"key":"B41","doi-asserted-by":"publisher","first-page":"647","DOI":"10.1016\/j.jid.2017.09.039","article-title":"Melanosome distribution in keratinocytes in different skin types: melanosome clusters are not degradative organelles","volume":"138","author":"Hurbain","year":"2018","journal-title":"J. Invest Dermatol"},{"key":"B42","doi-asserted-by":"publisher","first-page":"5177","DOI":"10.1242\/jcs.109314","article-title":"Functional involvement of Rab1A in microtubule-dependent anterograde melanosome transport in melanocytes","volume":"125","author":"Ishida","year":"2012","journal-title":"J. Cell Sci."},{"key":"B43","doi-asserted-by":"publisher","first-page":"8238","DOI":"10.1038\/srep08238","article-title":"Rab1A regulates anterograde melanosome transport by recruiting kinesin-1 to melanosomes through interaction with SKIP","volume":"5","author":"Ishida","year":"2015","journal-title":"Sci. Rep."},{"key":"B44","doi-asserted-by":"publisher","first-page":"e202110132","DOI":"10.1083\/jcb.202110132","article-title":"PI4P and BLOC-1 remodel endosomal membranes into tubules","volume":"221","author":"Jani","year":"2022","journal-title":"J. Cell Biol."},{"key":"B45","doi-asserted-by":"publisher","first-page":"e399","DOI":"10.1111\/j.1755-3768.2012.02423.x","article-title":"Comparison of macular pigment in patients with age-related macular degeneration and healthy control subjects - a study using spectral fundus reflectance","volume":"90","author":"Kaya","year":"2012","journal-title":"Acta Ophthalmol."},{"key":"B46","doi-asserted-by":"publisher","first-page":"403","DOI":"10.1111\/pcmr.12838","article-title":"Autophagy induction can regulate skin pigmentation by causing melanosome degradation in keratinocytes and melanocytes","volume":"33","author":"Kim","year":"2020","journal-title":"Pigment. Cell Melanoma Res."},{"key":"B47","doi-asserted-by":"publisher","first-page":"95","DOI":"10.1111\/pcmr.12645","article-title":"The patterns of birthmarks suggest a novel population of melanocyte precursors arising around the time of gastrulation","volume":"31","author":"Kinsler","year":"2018","journal-title":"Pigment. Cell Melanoma Res."},{"key":"B48","doi-asserted-by":"publisher","first-page":"373","DOI":"10.1016\/j.ophtha.2005.12.013","article-title":"Prevalence of age-related macular degeneration in 4 racial\/ethnic groups in the multi-ethnic study of atherosclerosis","volume":"113","author":"Klein","year":"2006","journal-title":"Ophthalmology"},{"key":"B49","doi-asserted-by":"publisher","first-page":"806","DOI":"10.1046\/j.1523-1747.1998.00178.x","article-title":"Supranuclear melanin caps reduce ultraviolet induced DNA photoproducts in human epidermis","volume":"110","author":"Kobayashi","year":"1998","journal-title":"J. Invest Dermatol"},{"key":"B50","doi-asserted-by":"publisher","first-page":"168","DOI":"10.1016\/j.jdermsci.2019.11.005","article-title":"TLR3 stimulation induces melanosome endo\/phagocytosis through RHOA and CDC42 in human epidermal keratinocyte","volume":"96","author":"Koike","year":"2019","journal-title":"J. Dermatol Sci."},{"key":"B51","doi-asserted-by":"publisher","first-page":"1195","DOI":"10.1038\/ncb1197","article-title":"Rab27A-binding protein Slp2-a is required for peripheral melanosome distribution and elongated cell shape in melanocytes","volume":"6","author":"Kuroda","year":"2004","journal-title":"Nat. Cell Biol."},{"key":"B52","doi-asserted-by":"publisher","first-page":"1517","DOI":"10.1093\/icb\/icab078","article-title":"Melanosome biogenesis in the pigmentation of Mammalian skin","volume":"61","author":"Le","year":"2021","journal-title":"Integr. Comp. Biol."},{"key":"B53","doi-asserted-by":"publisher","first-page":"3420","DOI":"10.1172\/JCI39109","article-title":"Melanocyte-like cells in the heart and pulmonary veins contribute to atrial arrhythmia triggers","volume":"119","author":"Levin","year":"2009","journal-title":"J. Clin. Invest"},{"key":"B54","doi-asserted-by":"publisher","first-page":"10300","DOI":"10.1038\/s41467-025-65221-0","article-title":"Atomic structure and in situ visualization of native PMEL lamellae in melanosomes","volume":"16","author":"Ma","year":"2025","journal-title":"Nat. Commun."},{"key":"B55","doi-asserted-by":"publisher","first-page":"495","DOI":"10.1016\/j.ceb.2013.04.008","article-title":"Lysosome-related organelles: unusual compartments become mainstream","volume":"25","author":"Marks","year":"2013","journal-title":"Curr. Opin. Cell Biol."},{"key":"B56","doi-asserted-by":"publisher","first-page":"45","DOI":"10.1247\/csf.19039","article-title":"Rab7B\/42 is functionally involved in protein degradation on Melanosomes in keratinocytes","volume":"45","author":"Marubashi","year":"2020","journal-title":"Cell Struct. Funct."},{"key":"B57","doi-asserted-by":"publisher","first-page":"341","DOI":"10.1046\/j.0022-202x.2001.01411.x","article-title":"Keratinocytes play a role in regulating distribution patterns of recipient melanosomes in vitro","volume":"117","author":"Minwalla","year":"2001","journal-title":"J. Invest Dermatol"},{"key":"B58","doi-asserted-by":"publisher","first-page":"366","DOI":"10.1111\/pcmr.12840","article-title":"The exocyst is required for melanin exocytosis from melanocytes and transfer to keratinocytes","volume":"33","author":"Moreiras","year":"2020","journal-title":"Pigment. Cell Melanoma Res."},{"key":"B59","doi-asserted-by":"publisher","first-page":"331","DOI":"10.1111\/tra.12843","article-title":"Melanocore uptake by keratinocytes occurs through phagocytosis and involves protease-activated receptor-2 internalization","volume":"23","author":"Moreiras","year":"2022","journal-title":"Traffic"},{"key":"B60","doi-asserted-by":"publisher","first-page":"2416","DOI":"10.1038\/jid.2013.165","article-title":"Autophagy has a significant role in determining skin color by regulating melanosome degradation in keratinocytes","volume":"133","author":"Murase","year":"2013","journal-title":"J. Invest Dermatol"},{"key":"B61","doi-asserted-by":"publisher","first-page":"25481","DOI":"10.1038\/s41598-024-73673-5","article-title":"Multiple cell types including melanocytes contribute to elastogenesis in the developing murine aortic valve","volume":"14","author":"Nasim","year":"2024","journal-title":"Sci. Rep."},{"key":"B62","doi-asserted-by":"publisher","first-page":"1155","DOI":"10.1016\/j.jid.2024.08.023","article-title":"Photoprotective melanin is maintained within keratinocytes in storage lysosomes","volume":"145","author":"Neto","year":"2025","journal-title":"J. Invest Dermatol"},{"key":"B63","doi-asserted-by":"publisher","first-page":"15835","DOI":"10.1038\/ncomms15835","article-title":"Routing of the RAB6 secretory pathway towards the lysosome related organelle of melanocytes","volume":"8","author":"Patwardhan","year":"2017","journal-title":"Nat. Commun."},{"key":"B64","doi-asserted-by":"publisher","first-page":"131","DOI":"10.1016\/j.exer.2017.10.018","article-title":"Visualizing melanosomes, lipofuscin, and melanolipofuscin in human retinal pigment epithelium using serial block face scanning electron microscopy","volume":"166","author":"Pollreisz","year":"2018","journal-title":"Exp. Eye Res."},{"key":"B65","doi-asserted-by":"publisher","first-page":"809","DOI":"10.1083\/jcb.152.4.809","article-title":"Distinct protein sorting and localization to premelanosomes, melanosomes, and lysosomes in pigmented melanocytic cells","volume":"152","author":"Raposo","year":"2001","journal-title":"J. Cell Biol."},{"key":"B66","doi-asserted-by":"publisher","first-page":"2709","DOI":"10.1083\/jcb.201709055","article-title":"Myosin VI and branched actin filaments mediate membrane constriction and fission of melanosomal tubule carriers","volume":"217","author":"Ripoll","year":"2018","journal-title":"J. Cell Biol."},{"key":"B67","doi-asserted-by":"publisher","first-page":"e70036","DOI":"10.1111\/boc.70036","article-title":"Functional and morphological plasticity of the endolysosomal System: pigment organelles at the crossroads of physiology and pathology","volume":"117","author":"Salavessa","year":"2025","journal-title":"Biol. Cell"},{"key":"B68","doi-asserted-by":"publisher","first-page":"193","DOI":"10.1111\/j.1600-0749.1993.tb00602.x","article-title":"Does melanin turnover occur in the eyes of adult vertebrates?","volume":"6","author":"Schraermeyer","year":"1993","journal-title":"Pigment. Cell Res."},{"key":"B69","doi-asserted-by":"publisher","first-page":"237","DOI":"10.1006\/exer.1998.0596","article-title":"Melanin granules of retinal pigment epithelium are connected with the lysosomal degradation pathway","volume":"68","author":"Schraermeyer","year":"1999","journal-title":"Exp. Eye Res."},{"key":"B70","doi-asserted-by":"publisher","first-page":"296","DOI":"10.1046\/j.1523-1747.2001.01221.x","article-title":"Rab3a and SNARE proteins: potential regulators of melanosome movement","volume":"116","author":"Scott","year":"2001","journal-title":"J. Invest Dermatol"},{"key":"B71","doi-asserted-by":"publisher","first-page":"768","DOI":"10.1091\/mbc.e06-12-1066","article-title":"BLOC-1 is required for cargo-specific sorting from vacuolar early endosomes toward lysosome-related organelles","volume":"18","author":"Setty","year":"2007","journal-title":"Mol. Biol. Cell"},{"key":"B72","doi-asserted-by":"publisher","first-page":"e45918","DOI":"10.15252\/embr.201845918","article-title":"Rab22A recruits BLOC-1 and BLOC-2 to promote the biogenesis of recycling endosomes","volume":"19","author":"Shakya","year":"2018","journal-title":"EMBO Rep."},{"key":"B73","doi-asserted-by":"publisher","first-page":"3093","DOI":"10.1242\/jcs.113.17.3093","article-title":"The protease-activated receptor-2 upregulates keratinocyte phagocytosis","volume":"113","author":"Sharlow","year":"2000","journal-title":"J. Cell Sci."},{"key":"B74","doi-asserted-by":"publisher","first-page":"25423","DOI":"10.1074\/jbc.M202574200","article-title":"A family of Rab27-binding proteins. Melanophilin links Rab27a and myosin Va function in melanosome transport","volume":"277","author":"Strom","year":"2002","journal-title":"J. Biol. Chem."},{"key":"B75","doi-asserted-by":"publisher","first-page":"1081","DOI":"10.1038\/2221081a0","article-title":"Racial differences in the fate of melanosomes in human epidermis","volume":"222","author":"Szabo","year":"1969","journal-title":"Nature"},{"key":"B76","doi-asserted-by":"publisher","first-page":"1056","DOI":"10.1038\/jid.2013.432","article-title":"Rab11b mediates melanin transfer between donor melanocytes and acceptor keratinocytes via coupled exo\/endocytosis","volume":"134","author":"Tarafder","year":"2014","journal-title":"J. Invest Dermatol"},{"key":"B77","doi-asserted-by":"publisher","first-page":"5356","DOI":"10.1091\/mbc.e05-07-0626","article-title":"Functions of adaptor protein (AP)-3 and AP-1 in tyrosinase sorting from endosomes to melanosomes","volume":"16","author":"Theos","year":"2005","journal-title":"Mol. Biol. Cell"},{"key":"B78","doi-asserted-by":"publisher","first-page":"498","DOI":"10.1046\/j.1365-2133.2003.05473.x","article-title":"The patterns of melanosome distribution in keratinocytes of human skin as one determining factor of skin colour","volume":"149","author":"Thong","year":"2003","journal-title":"Br. J. Dermatol"},{"key":"B79","doi-asserted-by":"publisher","first-page":"708","DOI":"10.1016\/j.devcel.2011.08.019","article-title":"The tetraspanin CD63 regulates ESCRT-independent and -dependent endosomal sorting during melanogenesis","volume":"21","author":"Van Niel","year":"2011","journal-title":"Dev. Cell"},{"key":"B80","doi-asserted-by":"publisher","first-page":"43","DOI":"10.1016\/j.celrep.2015.08.057","article-title":"Apolipoprotein E regulates amyloid Formation within endosomes of pigment cells","volume":"13","author":"Van Niel","year":"2015","journal-title":"Cell Rep."},{"key":"B81","doi-asserted-by":"publisher","first-page":"300","DOI":"10.1111\/pcmr.12067","article-title":"PMEL: a pigment cell-specific model for functional amyloid formation","volume":"26","author":"Watt","year":"2013","journal-title":"Pigment. Cell Melanoma Res."},{"key":"B82","doi-asserted-by":"publisher","first-page":"471","DOI":"10.1111\/j.1755-148X.2008.00483.x","article-title":"The location of heart melanocytes is specified and the level of pigmentation in the heart may correlate with coat color","volume":"21","author":"Yajima","year":"2008","journal-title":"Pigment. Cell Melanoma Res."},{"key":"B83","doi-asserted-by":"publisher","first-page":"e53183","DOI":"10.1371\/journal.pone.0053183","article-title":"A subpopulation of smooth muscle cells, derived from melanocyte-competent precursors, prevents patent ductus arteriosus","volume":"8","author":"Yajima","year":"2013","journal-title":"PLoS One"},{"key":"B84","doi-asserted-by":"publisher","first-page":"2829","DOI":"10.1096\/fj.06-6845com","article-title":"Functional analysis of keratinocytes in skin color using a human skin substitute model composed of cells derived from different skin pigmentation types","volume":"21","author":"Yoshida","year":"2007","journal-title":"FASEB J."},{"key":"B85","doi-asserted-by":"publisher","first-page":"e202110114","DOI":"10.1083\/jcb.202110114","article-title":"Type II phosphatidylinositol 4-kinases function sequentially in cargo delivery from early endosomes to melanosomes","volume":"221","author":"Zhu","year":"2022","journal-title":"J. Cell Biol."}],"container-title":["Frontiers in Cell and Developmental Biology"],"original-title":[],"link":[{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fcell.2025.1758081\/full","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,1,12]],"date-time":"2026-01-12T08:12:57Z","timestamp":1768205577000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fcell.2025.1758081\/full"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,1,12]]},"references-count":85,"alternative-id":["10.3389\/fcell.2025.1758081"],"URL":"https:\/\/doi.org\/10.3389\/fcell.2025.1758081","relation":{},"ISSN":["2296-634X"],"issn-type":[{"value":"2296-634X","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,1,12]]},"article-number":"1758081"}}