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Melanosome-enriched fractions isolated from human melanocytes expressed the Cdc42 effector proteins PAK1 and N-WASP by western blotting. Expression of constitutively active Cdc42(Cdc42V12) in melanocytes co-cultured with keratinocytes induced a highly dendritic phenotype with extensive contacts between melanocytes and keratinocytes through filopodia, many of which contained melanosomes. These results suggest a unique role for filopodia in organelle transport and, in combination with our previous work showing the presence of SNARE proteins and rab3a on melanosomes, suggest a novel model system for melanosome transfer to keratinocytes.<\/jats:p>","DOI":"10.1242\/jcs.115.7.1441","type":"journal-article","created":{"date-parts":[[2021,4,24]],"date-time":"2021-04-24T06:48:03Z","timestamp":1619246883000},"page":"1441-1451","source":"Crossref","is-referenced-by-count":130,"title":["Filopodia are conduits for melanosome transfer to keratinocytes"],"prefix":"10.1242","volume":"115","author":[{"given":"Glynis","family":"Scott","sequence":"first","affiliation":[{"name":"Department of Dermatology, University of Rochester School of Medicine and Dentistry, Rochester, NY, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sonya","family":"Leopardi","sequence":"additional","affiliation":[{"name":"Department of Dermatology, University of Rochester School of Medicine and Dentistry, Rochester, NY, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Stacey","family":"Printup","sequence":"additional","affiliation":[{"name":"Department of Dermatology, University of Rochester School of Medicine and Dentistry, Rochester, NY, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Brian C.","family":"Madden","sequence":"additional","affiliation":[{"name":"Department of Dermatology, University of Rochester School of Medicine and Dentistry, Rochester, NY, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"237","reference":[{"key":"2021042517340582900_REF1","doi-asserted-by":"crossref","unstructured":"Altun-Gultekin, Z. F. and Wagner, J. A. (1996). Src, Ras, and Rac mediate the migratory response elicited by NGF and PMA in PC12 cells. J. Neuroscience Res.44, 308-327.","DOI":"10.1002\/(SICI)1097-4547(19960515)44:4<308::AID-JNR2>3.0.CO;2-G"},{"key":"2021042517340582900_","doi-asserted-by":"crossref","unstructured":"Bagrodia, S., Taylor, S. J., Creasy, C. L., Chernoff, J. and Cerione, R. A. (1995). Identification of a mouse p21Cdc42\/Rac activated kinase. J. Biol. Chem.270, 22731-22737.","DOI":"10.1074\/jbc.270.39.22731"},{"key":"2021042517340582900_REF3","doi-asserted-by":"crossref","unstructured":"Bahadoran, P., Aberdam, E., Mantoux, F., Busca, R., Bille, K.,Yalman, N., de Saint-Basile, G., Casaroli-Marano, R., Ortonne, J. P. and Ballotti, R. (2001). Rab27a: A key to melanosome transport in human melanocytes. J. Cell Biol.152, 843-850.","DOI":"10.1083\/jcb.152.4.843"},{"key":"2021042517340582900_REF4","doi-asserted-by":"crossref","unstructured":"Banzai, Y., Miki, H., Yamaguchi, H. and Takenawa, T.(2000). Essential role of neural Wiskott-Aldrich syndrome protein in neurite extension in PC12 cells and rat hippocampal primary culture cells.J. Biol. Chem.275, 11987-11992.","DOI":"10.1074\/jbc.275.16.11987"},{"key":"2021042517340582900_REF5","doi-asserted-by":"crossref","unstructured":"Brown, A., O'Sullivan, A. J. and Gomperts, B. D.(1998). Induction of exocytosis from permeabilized mast cells by the guanosine triphosphatases Rac and Cdc42. Mol. Biol. Cell9, 1053-1063.","DOI":"10.1091\/mbc.9.5.1053"},{"key":"2021042517340582900_REF6","doi-asserted-by":"crossref","unstructured":"Brown, M. D., Cornejo, B. J., Kuhn, T. B. and Bamburg, J. R.(2000). Cdc42 stimulates neurite outgrowth and formation of growth cone filopodia and lamellipodia. J. Neurobiol.43, 352-364.","DOI":"10.1002\/1097-4695(20000615)43:4<352::AID-NEU4>3.0.CO;2-T"},{"key":"2021042517340582900_REF7","doi-asserted-by":"crossref","unstructured":"Burbelo, P. D., Dreschsel, D. and Hall, A.(1995). A conserved binding motif defines numerous candidate target proteins for both Cdc42 and Rac GTPases. J. Biol. Chem.270, 29071-29074.","DOI":"10.1074\/jbc.270.49.29071"},{"key":"2021042517340582900_REF8","doi-asserted-by":"crossref","unstructured":"Busca, R., Bertolotto, C., Abbe, P., Englaro, W., Ishizaki, T.,Narumiya, S., Boquet, P., Ortonne, J. P. and Ballotti, R.(1998). Inhibition of Rho is required for cAMP-induced melanoma cell differentiation. Mol. Biol. Cell9, 1367-1378.","DOI":"10.1091\/mbc.9.6.1367"},{"key":"2021042517340582900_REF9","doi-asserted-by":"crossref","unstructured":"Chung, C. Y., Potikyan, G. and Firtel, R. A.(2001). Control of cell polarity and chemotaxis by Akt\/PKB and PI3 kinase through the regulation of PAK\u03b1. Mol. Cell7, 937-947.","DOI":"10.1016\/S1097-2765(01)00247-7"},{"key":"2021042517340582900_REF10","doi-asserted-by":"crossref","unstructured":"Cohen, J. and Szabo, G. (1968). Study of pigment donation in vitro. Exp. Cell. Res.50, 418-434.","DOI":"10.1016\/0014-4827(68)90460-6"},{"key":"2021042517340582900_REF11","unstructured":"Corcuff, P., Chaussepied, C., Madry, G. and Hadjur, C.(2001). Skin optics revisited by in vivo confocal microscopy:Melanin and sun exposure. J. Cosmet Sci.52, 91-102."},{"key":"2021042517340582900_REF12","doi-asserted-by":"crossref","unstructured":"Dan, C., Kelly, A., Bernard, O. and Minden, A.(2001). Cytoskeletal changes regulated by the PAK4 serine\/threonine kinase are mediated by LIMK and cofilin. J. Biol. Chem.276, 32115-32121.","DOI":"10.1074\/jbc.M100871200"},{"key":"2021042517340582900_REF13","doi-asserted-by":"crossref","unstructured":"Davenport, R. W., Dou, P., Rehder, V. and Kater, S. B.(1993). 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Cell Biol.119, 1541-1557.","DOI":"10.1083\/jcb.119.6.1541"},{"key":"2021042517340582900_REF17","doi-asserted-by":"crossref","unstructured":"Frost, J. A., Khokhlatchev, A., Stippec, S., White, M. A. and Cobb, M. H. (1998). Differential effects of PAK1-activating mutations reveal activity-dependent and -independent effects on cytoskeletal regulation. J. Biol. Chem.273, 28191-28198.","DOI":"10.1074\/jbc.273.43.28191"},{"key":"2021042517340582900_REF18","doi-asserted-by":"crossref","unstructured":"Frost, J. A., Swantek, J. L., Stippec, S., Yin, M. J., Gaynor,R. and Cobb, M. H. (2000). Stimulation of NFkappa B activity by multiple signaling pathways requires PAK1. J. Biol. Chem.275, 19693-19699.","DOI":"10.1074\/jbc.M909860199"},{"key":"2021042517340582900_REF19","doi-asserted-by":"crossref","unstructured":"Fukuoka, M., Miki, H. and Takenawa, T. (1997). 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EMBO J.15, 5326-5335."},{"key":"2021042517340582900_REF41","doi-asserted-by":"crossref","unstructured":"Miki, H., Sasaki, T., Takai, Y. and Takenawa, T.(1998). Induction of filopodium formation by a WASP-related actin-depolymerizing protein N-WASP. Nature391, 93-96.","DOI":"10.1038\/34208"},{"key":"2021042517340582900_REF42","doi-asserted-by":"crossref","unstructured":"Minwalla, L., Zhao, Y., Cornelius, J., Babcock, G. F., Wickett,R. R., LePoole, I. C. and Boissy, R. E. (2001). Inhibition of melanosome transfer from melanocytes to keratinocytes by lectins and neoglycoproteins in an in vitro model system. Pigment Cell Res.14, 185-194.","DOI":"10.1034\/j.1600-0749.2001.140308.x"},{"key":"2021042517340582900_REF43","doi-asserted-by":"crossref","unstructured":"Montecucco, C., Pozzan, T. and Rink, T. (1979)Dicarbocyanine fluorescent probes of membrane potential block lymphocyte capping, deplete cellular ATP and inhibit respiration of isolated mitochondria. Biochim. Biophys. 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Cell8, 1971-1988.","DOI":"10.1091\/mbc.8.10.1971"},{"key":"2021042517340582900_REF47","unstructured":"Pathak, M. A., Jimbow, K. and Szabo, G. (1978). Sunlight and melanin pigmentation. In Smith KC, ed. Photochem Photobiol Reviews, Vol. 1. New York:Plenum Press, 221-239."},{"key":"2021042517340582900_REF48","doi-asserted-by":"crossref","unstructured":"Provance, D. W., Wei, M., Ipe, V. and Mercer, J. A.(1996). Cultured melanocytes from dilute mutant mice exhibit dendritic morphology and altered melanosome distribution. Proc. Natl. Acad. Sci USA93, 14554-14558.","DOI":"10.1073\/pnas.93.25.14554"},{"key":"2021042517340582900_REF49","doi-asserted-by":"crossref","unstructured":"Puls, A., Eliopoulos, G. G., Nobes, C. D., Bridges, T., Young,L. S. and Hall, A. (1999). Activation of the small GTPase Cdc42 by the inflammatory cytokines TNF(\u03b1) and IL-1, and by the Epstein-Barr virus transforming protein LMP1. J. 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(2000b). Inhibition of melanosome transfer results in skin lightening. J. Invest. Dermatol.115, 162-167.","DOI":"10.1046\/j.1523-1747.2000.00035.x"},{"key":"2021042517340582900_","doi-asserted-by":"crossref","unstructured":"Sells, M. A., Knaus, U. G., Bagrodia, S., Ambrose, D. M.,Bokoch, G. M. and Chernoff, J. (1997). Human p21-activated kinase (Pak1) regulates actin organization in mammalian cells.Curr. Biol.7, 202-210.","DOI":"10.1016\/S0960-9822(97)70091-5"},{"key":"2021042517340582900_REF63","doi-asserted-by":"crossref","unstructured":"Sells, M. A., Boyd, J. T. and Chernoff, J.(1999). P21-activated kinase 1 (PAK1) regulates cell motility in mammalian fibroblasts. J. Cell Biol.145, 837-849.","DOI":"10.1083\/jcb.145.4.837"},{"key":"2021042517340582900_REF64","doi-asserted-by":"crossref","unstructured":"Sharlow, E. R., Paine, C., Babiarz, L., Eisinger, M., Shapiro,S. S. and Seiberg, M. (2000). The protease activated receptor-2 upregulates keratinocyte phagocytosis. J. 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E., Espreafico, E. M. and Wolenski, J. S. (1999). Subcellular localization of GFP-myosin-V in live mouse melanocytes.J. Cell Sci.112, 2853-2865.","DOI":"10.1242\/jcs.112.17.2853"},{"key":"2021042517340582900_REF74","unstructured":"Wang F.-S., Wolenski, J. S., Cheney, R. E., Mooseker, M. S. and Jay, D. G. (1996). Function of myosin V in filopodia extension of neuronal growth cones. Science273, 660-663."},{"key":"2021042517340582900_REF75","doi-asserted-by":"crossref","unstructured":"Wang, J., Frost, J. A., Cobb, M. H. and Ross, E. M.(1999). Reciprocal signaling between heterotrimeric G proteins and the p21-stimulated protein kinase. J. Biol. Chem.274, 31641-34147.","DOI":"10.1074\/jbc.274.44.31641"},{"key":"2021042517340582900_REF76","doi-asserted-by":"crossref","unstructured":"Wei, Q., Wu, X. and Hammer, J. A., III (1997). The predominant defect in dilute melanocytes is in melanosome distribution and not cell shape, supporting a role for myosin V in melanosome transport.J. Muscle Res. Cell Mot.18, 517-527.","DOI":"10.1023\/A:1018659117569"},{"key":"2021042517340582900_REF77","doi-asserted-by":"crossref","unstructured":"Wilson, S., Yip, R., Swing, D. A., O'Sullivan, N., Zhang, Y.,Novak, E. D., Swank, R. T., Russell, L. B., Copeland, N. G. and Jenkins, N. A. (2000). A mutation in the rab27a causes the vesicle transport defects observed in ashen mice. Proc. Natl. Acad. Sci.97, 7933-7938.","DOI":"10.1073\/pnas.140212797"},{"key":"2021042517340582900_REF78","doi-asserted-by":"crossref","unstructured":"Wojciak-Stothard, B., Entwistle, A., Garg, R. and Ridley, A. J. (1998). Regulation of TNF-alpha-induced reorganiztion of the actin cytoskelton and cell-celljunctions by Rho, Rac, and Cdc42 in human endothelial cells. J. Cell. Physiol.179, 150-165.","DOI":"10.1002\/(SICI)1097-4652(199807)176:1<150::AID-JCP17>3.0.CO;2-B"},{"key":"2021042517340582900_REF79","unstructured":"Wolff, K. (1973). Melanocyte-keratinocyte interactions in vivo: the fate of melanosomes. Yale J. Biol. Med.46, 384-396."},{"key":"2021042517340582900_REF80","doi-asserted-by":"crossref","unstructured":"Wu, W. J., Erickson, J. W., Lin, R. and Cerione, R. A.(2000). The \u03b3-subunit of the coatomer complex binds Cdc42 to mediate transformation. Nature405, 800-803.","DOI":"10.1038\/35015585"},{"key":"2021042517340582900_REF81","doi-asserted-by":"crossref","unstructured":"Wu, X., Bowers, B., Wei, Q., Kocher, B. and Hammer, J. A.(1997). Myosin V associates with melanosomes in mouse melanocytes: evidence that myosin-V is an organelle motor. J. Cell Sci.110, 847-859.","DOI":"10.1242\/jcs.110.7.847"},{"key":"2021042517340582900_REF82","doi-asserted-by":"crossref","unstructured":"Wu, X. F., Bowers, B., Rao, K., Wei, Q. and Hammer, J. A.,III (1998). Visualization of melanosome dynamics within wild-type and dilute melanocytes suggests a paradigm for myosin V function in vivo. J. Cell Biol.143, 1899-1918.","DOI":"10.1083\/jcb.143.7.1899"},{"key":"2021042517340582900_REF83","doi-asserted-by":"crossref","unstructured":"Wu, X., Rao, K., Bowers, M. B., Copeland, N. G., Jenkins, N. A. and Hammer, J. A. (2001). Rab27a enables myosin Va-dependent melanosome capture by recruiting the myosin to the organelle. J. Cell Sci.114, 1091-1100.","DOI":"10.1242\/jcs.114.6.1091"},{"key":"2021042517340582900_REF84","doi-asserted-by":"crossref","unstructured":"Yamamoto, O. and Bhawan, J. (1994). Three modes of melanosome transfer in Caucasian facial skin: hypothesis based on an ultrastructural study. Pig. 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