Epidermal keratinocytes - genes and their regulation

Cell Death and Differentiation

Volumn 3, Issue 4, 1996, Pages 373-383

  Eckert, Richard L ^a   Welter, Jean F ^a  

^a CASE WESTERN RESERVE UNIVERSITY SCHOOL OF MEDICINE   (United States)

Author keywords

Activator; AP1; c fos; c jun; Epidermis; Gene regulation; Interferon; Involucrin; Keratin; Keratinocyte differentiation; Keratinocyte proliferation; Loricin; OCT factors; POU domain proteins; Protein 1; Retinoids; Vitamin D

Indexed keywords

BINDING PROTEIN; CYTOKERATIN; EPIDERMAL GROWTH FACTOR; IMMUNOGLOBULIN ENHANCER BINDING PROTEIN; INVOLUCRIN; PHORBOL ESTER; PROTEIN GLUTAMINE GAMMA GLUTAMYLTRANSFERASE; RETINOID; SULTAMICILLIN; TRANSCRIPTION FACTOR; TRANSFORMING GROWTH FACTOR BETA; VITAMIN D;

ANIMAL CELL; APOPTOSIS; CELL DIFFERENTIATION; EPIDERMIS CELL; GENE ACTIVATION; GENE CONTROL; HUMAN; HUMAN CELL; KERATINOCYTE; NONHUMAN; ONCOGENE C FOS; ONCOGENE C JUN; PHYSIOLOGY; PRIORITY JOURNAL; REVIEW;

ANIMALIA;

EID: 0029826817     PISSN: 13509047     EISSN: None     Source Type: Journal    
DOI: None     Document Type: Review

References (154)

1. Abernethy JL, Hill RL and Goldsmith LA (1977) epsilon-(gamma-Glutamyl)lysine cross-links in human stratum corneum. J. Biol. Chem. 252: 1837-1839
2. Agarwal C and Eckert RL (1990) Immortalization of human keratinocytes by simian virus 40 large T-antigen alters keratin gene response to retinoids. Cancer Res. 50: 5947-5953
3. Agarwal C, Hembree JR, Rorke EA and Eckert RL (1994) Interferon and retinoic acid suppress the growth of human papillomavirus type 16 immortalized cervical epithelial cells, but only interferon suppresses the level of the human papillomavirus transforming oncogenes Cancer Res. 54: 2108-2112
4. Agarwal C, Rorke EA, Boyce M, Howard J, Crish J, Hufeisen S and Eckert RL (1993) Retinoid-dependent transcriptional suppression of cytokeratin gene expression in human epidermal squamous cell carcinoma cells Differentiation. 52: 185-191
5. Agarwal C, Rorke EA, Irwin JC and Eckert RL (1991) Immortalization by human papillomavirus type 16 alters retinoid regulation of human ectocervical epithelial cell differentiation. Cancer Res 51: 3982-3989
6. Albers K and Fuchs E (1992) The molecular biology of intermediate filament proteins, Int Rev Cytol 134: 243-279
7. Andersen B, Schonemann MD, Flynn SE, Pearse RV 2d, Singh H and Rosenfeld MG (1993) Skn-1a and Skn-1i: two functionally distinct Oct-2-related factors expressed in epidermis. Science 260: 78-82
8. Angel P, Imagawa M, Chiu R, Stein B, Imbra RJ, Rahmsdorf HJ, Jonat C, Herrlich P and Karin M (1987) Phorbol ester-inducible genes contain a common cis element recognized by a TPA-modulated trans-acting factor. Cell 49: 729-739
9. Barber JR, Sassone-Corsi P and Verma IM (1987) Proto-oncogene fos: factors affecting expression and covalent modification of the gene product. Ann. N Y Acad. Sci. 511: 117-130
10. Barrandon Y and Green H (1987) Cell migration is essential for sustained growth of keratinocyte colonies: the roles of transforming growth factor-alpha and epidermal growth factor. Cell 50: 1131-1137
11. Bernerd F, Magnaldo T, Freedberg IM and Blumenberg M (1993) Expression of the carcinoma-associated keratin K6 and the role of AP-1 proto-oncoproteins. Gene Expr. 3: 187-199
12. Blessing M, Jorcano JL and Franke WW (1989) Enhancer elements directing cell-type-specific expression of cytokeratin genes and changes of the epithelial cytoskeleton by transfections of hybrid cytokeratin genes. Embo J. 8: 117-126
13. Bollag WB, Ducote J and Harmon CS (1993) Effects of the selective protein kinase C inhibitor, Ro 31-7549, on the proliferation of cultured mouse epidermal keratinocytes. J. Invest. Dermatol. 100: 240-246
14. Boyle WJ, Smeal T, Defize LH, Angel P, Woodgett JR, Karin M and Hunter T (1991) Activation of protein kinase C decreases phosphorylation of c-Jun at sites that negatively regulate its DNA-binding activity. Cell 64: 573-584
15. Brown LJ, Geesin JC, Rothnagel JA, Roop DR and Gordon JS (1994) Retinoic acid suppression of loricrin expression in reconstituted human skin cultured at the liquid-air interface. J. Invest. Dermatol. 102: 886-890
16. Byrne C and Fuchs E (1993) Probing keratinocyte and differentiation specificity of the human K5 promoter in vitro and in transgenic mice. Mol. Cell. Biol. 13: 3176-3190
17. Carroll JM, Albers KM, Garlick JA, Harrington R and Taichman LB (1993) Tissue-and stratum-specific expression of the human involucrin promoter in transgenic mice. Proc. Natl. Acad. Sci. USA 90: 10270-10274
18. Casatorres J, Navarro JM, Blessing M and Jorcano JL (1994) Analysis of the control of expression and tissue specificity of the keratin 5 gene, characteristic of basal keratinocytes. Fundamental role of an AP-1 element. J. Biol. Chem. 269: 20489-20496
19. Chakravarty R and Rice RH (1989) Acylation of keratinocyte transglutaminase by palmitic and myristic acids in the membrane Anchorage region. J. Biol. Chem. 264: 625-629
20. Chakravarty R, Rong XH and Rice RH (1990) Phorbol ester-stimulated phosphorylation of keratinocyte transglutaminase in the membrane anchorage region. Biochem. J. 271: 25-30
21. Chan WK, Chong T, Bernard HU and Klock G (1990) Transcription of the transforming genes of the oncogenic human papillomavirus-16 is stimulated by tumor promotors through AP1 binding sites. Nucleic Acids Res. 18: 763-769
22. Chen JD, Kim JP, Zhang K, Sarret Y, Wynn KC, Kramer RH and Woodley DT (1993) Epidermal growth factor (EGF) promotes human keratinocyte locomotion on collagen by increasing the alpha 2 integrin subunit. Exp. Cell Res. 209: 216-223
23. Chen TC, Persons K, Liu WW, Chen ML and Holick MF (1995) The antiproliferative and differentiative activities of 1,25-dihydroxyvitamin D3 are potentiated by epidermal growth factor and attenuated by insulin in cultured human keratinocytes. J. Invest. Dermatol. 104: 113-117
24. Cline PR and Rice RH (1983) Modulation of involucrin and envelope competence in human keratinocytes by hydrocortisone, retinyl acetate, and growth arrest. Cancer Res. 43: 3203-3207
25. Coffey RJ Jr, Sipes NJ, Bascom CC, Graves-Deal R, Pennington CY, Weissman BE and Moses HL (1988) Growth modulation of mouse keratinocytes by transforming growth factors. Cancer Res. 48: 1596-1602
26. Cripe TP, Alderborn A, Anderson RD, Parkkinen S, Bergman P, Haugen TH, Pettersson U and Turek LP (1990) Transcriptional activation of the human papillomavirus-16 P97 promoter by an 88-nucleotide enhancer containing distinct cell-dependent and AP-1 -responsive modules. New Biol. 2: 450-463
27. Crish JF, Howard JM, Zaim TM, Murthy S and Eckert RL (1993) Tissue-specific and differentiation-appropriate expression of the human involucrin gene in transgenic mice: an abnormal epidermal phenotype. Differentiation 53: 191-200
28. Darnell JE Jr, Kerr IM and Stark GR (1994) Jak-STAT pathways and transcriptional activation in response to IFNs and other extracellular signaling proteins. Science 264: 1415-1421
29. DiSepio D, Jones A, Longley MA, Bundman D, Rothnagel JA and Roop DR (1995) The proximal promoter of the mouse loricrin gene contains a functional AP-1 element and directs keratinocyte-specific but not differentiation-specific expression. J. Biol. Chem. 270: 10792-10799
30. Dlugosz AA (1995) Biochemical regulation of keratinocyte differentiation and neoplastic transformation. In Skin cancer: Mechanisms and human relevance, Mukhtar H, (Ann Arbor: CRC Press) 199-206
31. Dlugosz AA, Mischak H, Mushinski JF and Yuspa SH (1992) Transcripts encoding protein kinase C-alpha, -delta, -epsilon, -zeta, and -eta are expressed in basal and differentiating mouse keratinocytes in vitro and exhibit quantitative changes in neoplastic cells. Mol. Carcinog. 5: 286-292
32. Dlugosz AA and Yuspa SH (1993) Coordinate changes in gene expression which mark the spinous to granular cell transition in epidermis are regulated by protein kinase. C. J. Cell. Biol. 120: 217-225
33. Dlugosz AA and Yuspa SH (1994) Protein kinase C regulates keratinocyte transglutaminase (TGK) gene expression in cultured primary mouse epidermal keratinocytes induced to terminally differentiate by calcium. J. Invest. Dermatol. 102: 409-414
34. Eckert RL (1989) Structure, function, and differentiation of the keratinocyte. Physiol. Rev. 69: 1316-1346
35. Eckert RL and Green H (1984) Cloning of cDNAs specifying vitamin A-responsive human keratins. Proc. Natl. Acad. Sci. USA 81: 4321-4325
36. Eckert RL and Green H (1986) Structure and evolution of the human involucrin gene. Cell 46: 583-589
37. Eckert RL, Yaffe MB, Crish JF, Murthy S, Rorke EA and Welter JF (1993) Involucrin-structure and role in envelope assembly. J. Invest. Dermatol. 100: 613-617
38. Elder JT, Astrom A, Pettersson U, Voorhees JJ and Trent JM (1992) Assignment of the human CRABP-II gene to chromosome 1q21 by nonisotopic in situ hybridization. Hum. Genet. 89: 487-490
39. Elias PM, Menon GK, Grayson S and Brown BE (1988) Membrane structural alterations in murine stratum corneum: relationship to the localization of polar lipids and phospholipases. J. Invest. Dermatol. 91: 3-10
40. Faus I, Hsu HJ and Fuchs E (1994) Oct-6: a regulator of keratinocyte gene expression in stratified squamous epithelia. Mol. Cell. Biol. 14: 3263-3275
41. Fietz MJ, Rogers GE, Eyre HJ, Baker E, Callen DF and Sutherland GR (1992) Mapping of the trichohyalin gene: co-localization with the profilaggrin, involucrin, and loricrin genes. J. Invest. Dermatol. 99: 542-544
42. Fisher GJ, Talwar HS, Xiao JH, Datta SC, Reddy AP, Gaub MP, Rochette-Egly C, Chambon P and Voorhees JJ (1994) Immunological identification and functional quantitation of retinoic acid and retinoid X receptor proteins in human skin. J. Biol. Chem. 269: 20629-20635
43. Fuchs E (1990) Epidermal differentiation: the bare essentials. J. Cell. Biol. 111: 2807-2814
44. Giguere V, Ong ES, Segui P and Evans RM (1987) Identification of a receptor for the morphogen retinoic acid. Nature 330: 624-629
45. Gilfix BM and Eckert RL (1985) Coordinate control by vitamin A of keratin gene expression in human keratinocytes. J. Biol. Chem. 260: 14026-14029
46. Green H (1977) Terminal differentiation of cultured human epidermal cells. Cell 11: 405-416
47. Green H, Fuchs E and Watt F (1982) Differentiated structural components of the keratinocyte. Cold Spring Harb. Symp. Quant. Biol. 46 Pt 1: 293-301
48. Greenberg CS, Birckbichler PJ and Rice RH (1991)Transglutaminases:multifunctional cross-linking enzymes that stabilize tissues. FASEB J. 5: 3071-3077
49. Griffiths CE, Rosenthal DS, Reddy AP, Elder JT, Astrom A, Leach K, Wang TS, Finkel LJ, Yuspa SH, Voorhees JJ, et al (1992) Short-term retinoic acid treatment increases in vivo, but decreases in vitro, epidermal transglutaminase-K enzyme activity and immunoreactivity. J. Invest. Dermatol. 99: 283-288
50. Haussler MR, Mangelsdorf DJ, Komm BS, Terpening CM, Yamaoka K, Allegretto EA, Baker AR, Shine J, McDonnell DP, Hughes M, Weigel NL, O'Malley BW and Pike JW (1988) Molecular biology of the vitamin D hormone. Recent Prog. Horm. Res. 44: 263-305
51. Hennings H, Michael D, Cheng C, Steinert P, Holbrook K and Yuspa SH (1980) Calcium regulation of growth and differentiation of mouse epidermal cells in culture. Cell 19: 245-254
52. Hohl D (1993) Expression patterns of loricrin in dermatological disorders. Am. J. Dermatopathol. 15: 20-27
53. Hohl D, Lichti U, Breitkreutz D, Steinert PM and Roop DR (1991a) Transcription of the human loricrin gene in vitro is induced by calcium and cell density and suppressed by retinoic acid. J. Invest. Dermatol. 96: 414-418
54. Hohl D, Mehrel T, Lichti U, Turner ML, Roop DR and Steinert PM (1991b) Characterization of human loricrin. Structure and function of a new class of epidermal cell envelope proteins. J. Biol. Chem. 266: 6626-6636
55. Hohl D, Ruf-Olano B, de-Viragh PA, Huber M, Detrisac CJ, Schnyder UW and Roop DR (1993) Expression patterns of loricrin in various species and tissues. Differentiation 54: 25-34
56. Holbrook KA and Wolff K (1987) The structure and development of skin. In Dermatology in general medicine, Fitzpatrick TB, Lesen AZ, Wolff K, Freedberg IM and Austen KF, (New York: McGraw Hill) 93-131
57. Holick MF (1994) McCollum Award Lecture, 1994: vitamin D-new horizons for the 21st century. Am. J. Clin. Nutr. 60: 619-630
58. Hosomi J, Hosoi J, Abe E, Suda T and Kuroki T (1983) Regulation of terminal differentiation of cultured mouse epidermal cells by 1 alpha,25-dihydroxyvitamin D3. Endocrinology 113: 1950-1957
59. Huff CA, Yuspa SH and Rosenthal D (1993) Identification of control elements 3′ to the human keratin 1 gene that regulate cell type and differentiation-specific expression. J. Biol. Chem. 268: 377-384
60. Ishida-Yamamoto A, Hohl D, Roop DR, lizuka H and Eady RA (1993) Loricrin immunoreactivity in human skin: localization to specific granules (L-granules) in acrosyringia. Arch Dermatol. Res. 285: 491-498
61. Jiang CK, Flanagan S, Ohtsuki M, Shuai K, Freedberg IM and Blumenberg M (1994) Disease-activated transcription factor: allergic reactions in human skin cause nuclear translocation of STAT-91 and induce synthesis of keratin K17. Mol. Cell. Biol. 14: 4759-4769
62. Jiang CK, Magnaldo T, Ohtsuki M, Freedberg IM, Bernerd F and Blumenberg M (1993) Epidermal growth factor and transforming growth factor alpha specifically induce the activation- and hyperproliferation-associated keratins 6 and 16. Proc. Natl. Acad. Sci. USA 90: 6786-6790
63. Johnson RS, Spiegelman BM and Papaioannou V (1992) Pleiotropic effects of a null mutation in the c-fos proto-oncogene. Cell 71: 577-586
64. Jones KT and Sharpe GR (1994) Intracellular free calcium and growth changes in single human keratinocytes in response to vitamin D and five 20-epi-analogues. Arch Dermatol. Res. 286: 123-129
65. Juhlin L, Magnoldo T and Darmon M (1992) Expression of loricrin in skin disorders. Acta Derm. Venereol. 72: 407-409
66. Kachinskas DJ, Phillips MA, Qin Q, Stokes JD and Rice RH (1994) Arsenate perturbation of human keratinocyte differentiation. Cell Growth Differ. 5: 1235-1241
67. Kim HC, Lewis MS, Gorman JJ, Park SC, Girard JE, Folk JE and Chung SI (1990) Protransglutaminase E from guinea pig skin. Isolation and partial characterization. J. Biol. Chem. 265: 21971-21978
68. Kopan R, Traska G and Fuchs E (1987) Retinoids as important regulators of terminal differentiation: examining keratin expression in individual epidermal cells at various stages of keratinization. J. Cell Biol. 105: 427-440
69. Lavker RM, Miller SJ and Sun TT (1993) Epithelial stem cells, hair follicles, and tumor formation Recent. Results. Cancer Res. 128: 31-43
70. Leask A, Byrne C and Fuchs E (1991) Transcription factor AP2 and its role in epidermal-specific gene expression. Proc. Natl. Acad. Sci. USA 88: 7948-7952
71. Leask A, Rosenberg M, Vassar R and Fuchs E (1990) Regulation of a human epidermal keratin gene: sequences and nuclear factors involved in keratinocyte-specific transcription. Genes Dev. 4: 1985-1998
72. Li L, Tucker RW, Hennings H and Yuspa SH (1995) Chelation of intracellular Ca2+ inhibits murine keratinocyte differentiation in vitro. J. Cell. Physiol. 163: 105-114
73. Liew FM and Yamanishi K (1992) Regulation of transglutaminase 1 gene expression by 12-O-tetradecanoylphorbol-13-acetate, dexamethasone, and retinoic acid in cultured human keratinocytes. Exp. Cell. Res. 202: 310-315
74. Lu B, Rothnagel JA, Longley MA, Tsai SY and Roop DR (1994) Differentiation-specific expression of human keratin 1 is mediated by a composite AP-1/steroid hormone element. J. Biol. Chem. 269: 7443-7449
75. Mack DH and Laimins LA (1991 ) A keratinocyte-specific transcription factor, KRF-1, interacts with AP-1 to activate expression of human papillomavirus type 18 in squamous epithelial cells. Proc. Natl. Acad. Sci. USA 88: 9102-9106
76. MacLaughlin JA, Cantley LC and Holick MF (1990) 1,25-dihydroxyvitamin D3 increases calcium and phosphatidylinositol metabolism in differentiating cultured human keratinocytes. J Nutr Biochem 1: 81-87
77. Magnaldo T, Bernerd F, Asselineau D and Darmon M (1992) Expression of loricrin is negatively controlled by retinoic acid in human epidermis reconstructed in vitro. Differentiation 49: 39-46
78. Magnaldo T, Bernerd F, Freedberg IM, Ohtsuki M and Blumenberg M (1993) Transcriptional regulators of expression of K#16, the disease-associated keratin DNA-Cell Biol. 12: 911-923
79. Magnaldo T, Pommes L, Asselineau D and Darmon M (1990) Isolation of a GC-rich cDNA identifying mRNA present in human epidermis and modulated by calcium and retinoic acid in cultured keratinocytes. Homology with murine loricrin mRNA. Mol. Biol. Rep. 14: 237-246
80. MagnaldoT, Vidal RG, Ohtsuki M, Freedberg IM and Blumenberg M (1993) On the role of AP2 in epithelial-specific gene expression. Gene Expr. 3: 307-315
81. Malmqvist KG, Carlsson LE, Forslind B, Roomans GM and Akselsson KR (1984) Proton and electron microprobe analysis of human skin. Nucl. Instrum. Meth. Phys. Res. 63: 611-617
82. Mangelsdorf DJ (1994) Vitamin A receptors. Nutr. Rev. 52: S32-S44
83. Mariniello L, Qin Q, Jessen BA and Rice RH (1996) Keratinocyte transglutaminase promoter analysis: identification of a functional response element. J. Biol. Chem. (in press)
84. Marvin KW, George MD, Fujimoto W, Saunders NA, Bernacki SH and Jetten AM (1992) Cornifin, a cross-linked envelope precursor in keratinocytes that is down-regulated by retinoids. Proc. Natl. Acad. Sci. USA 89: 11026-11030
85. Matoltsy AG and Matoltsy MN (1966) The membrane protein of horny cells. J. Invest. Dermatol. 46: 127-129
86. MehrelT, Hohl D, Rothnagel JA, Longley MA, Bundman D, Cheng C, Lichti U, Bisher ME, Steven AC, Steinert PM, Yuspa SH and Roop DR (1990) Identification of a major keratinocyte cell envelope protein, loricrin. Cell 61: 1103-1112
87. Menon GK, Grayson S and Elias PM (1985) Ionic calcium reservoirs in mammalian epidermis:ultrastructural localization by ion-capture cytochemistry. J. Invest. Dermatol. 84: 508-512
88. Milde P, Hauser U, Simon T, Mall G, Ernst V, Haussler MR, Frosch P and Rauterberg EW (1991) Expression of 1,25-dihydroxyvitamin D3 receptors in normal and psoriatic skin. J. Invest. Dermatol. 97: 230-239
89. Mitchell PJ, Timmons PM, Hebert JM, Rigby PW and Tjian R (1991) Transcription factor AP-2 is expressed in neural crest cell lineages during mouse embryogenesis. Genes Dev. 5: 105-119
90. Monuki ES, Kuhn R and Lemke G. (1993) Repression of the myelin P0 gene by the POU transcription factor SCIP. Mech. Dev. 42: 15-32
91. Murthy S, Crish JF, ZaimTM and Eckert RL (1993) A dual role for involucrin in the epidermis-ultrastructural localization in epidermis and hair follicle in humans and transgenic mice. J. Struct. Biol. 111: 68-76
92. Navarro JM, Casatorres J and Jorcano JL (1995) Elements controlling the expression and induction of skin hyperproliferation-associated keratin K6. J. Biol. Chem. 270: 21362-21367
93. Nicholson RC, Mader S, Nagpal S, Leid M, Rochette-Egly C and Chambon P (1990) Negative regulation of the rat stromelysin gene promoter by retinoic acid is mediated by an AP1 binding site. EMBO J. 9: 4443-4454
94. Nishizuka Y (1992) Intracellular signaling by hydrolysis of phospholipids and activation of protein kinase. C. Science 258: 607-614
95. Offord EA and Beard P (1990) A member of the activator protein 1 family found in keratinocytes but not in fibroblasts required for transcription from a human papillomavirus type 18 promoter. J. Virol. 64: 4792-4798
96. Ohtsuki M, Flanagan S, Freedberg IM and Blumenberg M (1993) A cluster of five nuclear proteins regulates keratin gene transcription. Gene Expr. 3: 201-213
97. Ohtsuki M, Tomic-Canic M, Freedberg IM and Blumenberg M (1992) Nuclear proteins involved in transcription of the human K5 keratin gene. J. Invest. Dermatol. 99: 206-215
98. Oshima RG, Abrams L and Kulesh D (1990) Activation of an intron enhancer within the keratin 18 gene by expression of c-fos and c-jun in undifferentiated F9 embryonal carcinoma cells. Genes Dev. 4: 835-848
99. Piepkorn M, Lo C and Plowman G (1994) Amphiregulin-dependent proliferation of cultured human keratinocytes: autocrine growth, the effects of exogenous recombinant cytokine, and apparent requirement for heparin-like glycosaminoglycans. J. Cell. Physiol. 159: 114-120
100. Pietenpol JA, Holt JT, Stein RW and Moses HL (1990a) Transforming growth factor beta 1 suppression of c-myc gene transcription: role in inhibition of keratinocyte proliferation. Proc. Natl. Acad. Sci. USA 87: 3758-3762
101. Pietenpol JA, Stein RW, Moran E, Yaciuk P, Schlegel R, Lyons RM, Pittelkow MR, Munger K, Howley PM and Moses HL (1990b) TGF-beta 1 inhibition of c-myc transcription and growth in keratinocytes is abrogated by viral transforming proteins with pRB binding domains. Cell 61: 777-785
102. Polakowska R, Herting E and Goldsmith LA (1991) Isolation of cDNA for human epidermal type I transglutaminase. J. Invest. Dermatol. 96: 285-288
103. Pulverer BJ, Kyriakis JM, Avruch J, Nikolakaki E and Woodgett JR (1991) Phosphorylation of c-jun mediated by MAP kinases. Nature 353: 670-674
104. Ransone LJ and Verma IM (1990) Nuclear proto-oncogenes fos and jun. Annu. Rev. Cell. Biol. 6: 539-557
105. Reiss M and Sartorelli AC (1987) Regulation of growth and differentiation of human keratinocytes by type beta transforming growth factor and epidermal growth factor. Cancer Res. 47: 6705-6709
106. Rheinwald JG and Green H (1975) Serial cultivation of strains of human epidermal keratinocytes: the formation of keratinizing colonies from single cells. Cell 6: 331-343
107. Rheinwald JG and Green H (1977) Epidermal growth factor and the multiplication of cultured human epidermal keratinocytes. Nature 265: 421-424
108. Rice RH and Green H (1977) The cornified envelope of terminally differentiated human epidermal keratinocytes consists of cross-linked protein. Cell 11: 417-422
109. Rice RH and Green H (1979) Presence in human epidermal cells of a soluble protein precursor of the cross-linked envelope: activation of the cross-linking by calcium ions. Cell 18: 681-694
110. Rosenthal DS, Griffiths CE, Yuspa SH, Roop DR and Voorhees JJ (1992) Acute or chronic topical retinoic acid treatment of human skin in vivo alters the expression of epidermal transglutaminase, loricrin, involucrin, filaggrin, and keratins 6 and 13 but not keratins 1, 10, and 14. J. Invest. Dermatol. 98: 343-350
111. Rothnagel JA, Greenhalgh DA, Gagne TA, Longley MA and Roop DR (1993) Identification of a calcium-inducible, epidermal-specific regulatory element in the 3′-flanking region of the human keratin 1 gene. J. Invest. Dermatol. 101: 506-513
112. Salbert G, Fanjul A, Piedrafita FJ, Lu XP, Kirn SJ, Tran P and Pfahl M (1993) Retinoic acid receptors and retinoid X receptor-alpha down-regulate the transforming growth factor-beta 1 promoter by antagonizing AP-1 activity. Mol. Endocrinol. 7: 1347-1356
113. Saunders NA, Bernacki SH, Vollberg TM and Jetten AM (1993) Regulation of transglutaminase type I expression in squamous differentiating rabbit tracheal epithelial cells and human epidermal keratinocytes: effects of retinoic acid and phorbol esters. Mol. Endocrinol. 7: 387-398
114. Saunders NA and Jetten AM (1994) Control of growth regulatory and differentiation-specific genes in human epidermal keratinocytes by interferon gamma. Antagonism by retinoic acid and transforming growth factor beta 1. J. Biol. Chem. 269: 2016-2022
115. Shipley GD, Pittelkow MR, Wille JJ Jr, Scott RE and Moses HL (1986) Reversible inhibition of normal human prokeratinocyte proliferation by type beta transforming growth factor-growth inhibitor in serum-free medium. Cancer Res. 46: 2068-2071
116. Simon M and Green H (1984) Participation of membrane-associated proteins in the formation of the cross-linked envelope of the keratinocyte. Cell 36: 827-834
117. Simon M and Green H (1985) Enzymatic cross-linking of involucrin and other proteins by keratinocyte particulates in vitro. Cell 40: 677-683
118. Simon M and Green H (1988) The glutamine residues reactive in transglutaminase-catalyzed cross-linking of involucrin. J. Biol. Chem. 263: 18093-18098
119. Smith EL, Walworth NC and Holick MF (1986) Effect of 1 alpha,25-dihydroxyvitamin D3 on the morphologic and biochemical differentiation of cultured human epidermal keratinocytes grown in serum-free conditions. J. Invest. Dermatol. 86: 709-714
120. Steinert PM (1995) A model for the hierachical structure of the human epidermal cornified cell envelope. Cell Death Differ. 2: 33-40
121. Steinert PM and Marekow LN (1995) The proteins elafin, filaggrin, keratin intermediate filaments, loricrin and small proline-rich proteins 1 and 2 are isodipeptide cross linked components of the human epidermal keratinocyte cornified envelope. J. Biol. Chem. 270: 17702-17711
122. Steinert PM, Steven AC and Roop DR (1985)The molecular biology of intermediate filaments. Cell 42: 411-420
123. Steven AC, Bisher ME, Roop DR and Steinert PM (1990) Biosynthetic pathways of filaggrin and loricrin-two major proteins expressed by terminally differentiated epidermal keratinocytes. J. Struct. Biol. 104: 150-162
124. Steven AC and Steinert PM (1994) Protein composition of cornified cell envelopes of epidermal keratinocytes. J. Cell. Sci. 107: 693-700
125. Stoler A, Kopan R, Duvic M and Fuchs E (1988) Use of monospecific antisera and cRNA probes to localize the major changes in keratin expression during normal and abnormal epidermal differentiation. J. Cell. Biol. 107: 427-446
126. Studzinski GP, McLane JA and Uskokovic MR (1993) Signaling pathways for vitamin D-induced differentiation: implications for therapy of proliferative and neoplastic diseases. Crit. Rev. Eukaryot. Gene. Expr. 3: 279-312
127. Su MJ, Bikle DD, Mancianti ML and Pillai S (1994) 1,25-Dihydroxyvitamin D3 potentiates the keratinocyte response to calcium. J. Biol. Chem. 269: 14723-14729
128. Sun TT, Eichner R, Nelson WG, Tseng SC, Weiss RA, Jarvinen M and Woodcock-Mitchell J (1983) Keratin Classes: molecular markers for different types of epithelial differentiation. J. Invest. Dermatol. 81: 109s-115s
129. Suzuki T, Okuno H, Yoshida T, Endo T, Nishina H and Iba H (1991) Difference in transcriptional regulatory function between c-Fos and Fra-2. Nucleic Acids Res. 19: 5537-5542
130. Takahashi H and Iizuka H (1993) Analysis of the 5′-upstream promoter region of human involucrin gene: activation by 12-O-tetradecanoylphorbol-13-acetate. J. Invest. Dermatol. 100: 10-15
131. Takemoto Y, Fujimura Y, Matsumoto M, Tamai Y, Morita T, Matsushiro A and Nozaki M (1991) The promoter of the endo A cytokeratin gene is activated by a 3′ downstream enhancer. Nucleic Acids Res. 19: 2761-2765
132. Tamai Y, Takemoto Y, Matsumoto M, Morita T, Matsushiro A and Nozaki M (1991) Sequence of EndoA gene encoding mouse cytokeratin and its methylation state in the CpG-rich region. Gene 104: 169-176
133. Tang W, Ziboh VA, Isseroff RR and Martinez D (1987) Novel regulatory actions of 1 alpha,25-dihydroxyvitamin D3 on the metabolism of polyphosphoinositides in murine epidermal keratinocytes. J. Cell. Physiol. 132: 131-136
134. Thacher SM and Rice RH (1985) Keratinocyte-specific transglutaminase of cultured human epidermal cells: relation to cross-linked envelope formation and terminal differentiation. Cell 40: 685-695
135. Thierry F, Spyrou G, Yaniv M and Howley P (1992) Two AP1 sites binding junB are essential for human papillomavirus type 18 transcription in keratinocytes. J. Virol. 66: 3740-3748
136. Volz A, Korge BP, Compton JG, Ziegler A, Steinert PM and Mischke D (1993) Physical mapping of a functional cluster of epidermal differentiation genes on chromosome 1q21. Genomics 18: 92-99
137. Wegner M, Drolet DW and Rosenfeld MG (1993) POU-domain proteins: structure and function of developmental regulators. Curr. Opin. Cell. Biol. 5: 488-498
138. Weiss RA, Eichner R and Sun TT (1984) Monoclonal antibody analysis of keratin expression in epidermal diseases: a 48- and 56-kdalton keratin as molecular markers for hyperproliferative keratinocytes. J. Cell. Biol. 98: 1397-1406
139. Welter JF, Crish JF, Agarwal C and Eckert RL (1995) Fos-related antigen (Fra-1), junB, and junD activate human involucrin promoter transcription by binding to proximal and distal AP1 sites to mediate phorbol ester effects on promoter activity. J. Biol. Chem. 270: 12614-12622
140. Welter JF and Eckert RL (1995) Differential expression of the fos and jun family members c-fos, fosB, fra-1 fra-2, c-jun, junB and junD during human epidermal keratinocyte differentiation. Oncogene 11: 2681-2687
141. Welter JF, Gali H, Crish JF and Eckert RL (1996) Regulation of human involucrin promoter activity by POU domain proteins. J. Biol. Chem. 271: 14727-14733
142. Williams T and Tjian R (1991) Analysis of the DNA-binding and activation properties of the human transcription factor AP-2. Genes Dev. 5: 670-682
143. Wu RL, Chen TT and Sun TT (1994) Functional importance of an Sp1- and an NFkB-related nuclear protein in a keratinocyte-specific promoter of rabbit K3 keratin gene. J. Biol. Chem. 269: 28450-28459
144. Yaffe MB, Beegen H and Eckert RL (1992) Biophysical characterization of involucrin reveals a molecule ideally suited to function as an intermolecular cross-bridge of the keratinocyte cornified envelope. J. Biol. Chem. 267: 12233-12238
145. Yaffe MB, Murthy S and Eckert RL (1993) Evidence that involucrin is a covalently linked constituent of highly purified cultured keratinocyte cornified envelopes. J. Invest. Dermatol. 100: 3-9
146. Yamada K, Yamanishi K, Kakizuka A, Kibe Y, Doi H and Yasuno H (1994) Transcriptional regulation of human transglutaminase1 gene by signaling systems of protein kinase C, RAR/RXR and Jun/Fos in keratinocytes. Biochem. Mol. Biol. Int. 34: 827-836
147. Yang JS, Lavker RM and Sun TT (1993) Upper human hair follicle contains a subpopulation of keratinocytes with superior in vitro proliferative potential. J. Invest. Dermatol. 101: 652-659
148. Yoneda K, McBride OW, Korge BP, Kim IG and Steinert PM (1992) The cornified cell envelope: loricrin and transglutaminases. J. Dermatol. 19: 761-764
149. Yoneda K and Steinert PM (1993) Overexpression of human loricrin in transgenic mice produces a normal phenotype. Proc. Natl. Acad. Sci. USA 90: 10754-10758
150. Younus J and Gilchrest BA (1992) Modulation of mRNA levels during human keratinocyte differentiation. J. Cell. Physiol. 152: 232-239
151. Yuspa SH (1994)The pathogenesis of squamous cell cancer: lessons learned from studies of skin carcinogenesis-thirty-third GHA Clowes Memorial Award Lecture. Cancer Res. 54: 1178-1189
152. Yuspa SH, Kilkenny AE, Steinert PM and Roop DR (1989) Expression of murine epidermal differentiation markers is tightly regulated by restricted extracellular calcium concentrations in vitro. J Cell Biol 109: 1207-1217
153. Zhang XK, Hoffmann B, Tran PB, Graupner G and Pfahl M (1992) Retinoid X receptor is an auxiliary protein for thyroid hormone and retinoic acid receptors. Nature 355: 441-446
154. Ziboh VA, Isseroff RR and Pandey R (1984) Phospholipid metabolism in calcium-regulated differentiation in cultured murine keratinocytes. Biochem. Biophys. Res. Commun. 122: 1234-1240