{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,5,20]],"date-time":"2026-05-20T19:08:40Z","timestamp":1779304120839,"version":"3.51.4"},"reference-count":80,"publisher":"Springer Science and Business Media LLC","issue":"19","license":[{"start":{"date-parts":[[2000,10,2]],"date-time":"2000-10-02T00:00:00Z","timestamp":970444800000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.springer.com\/tdm"},{"start":{"date-parts":[[2000,10,2]],"date-time":"2000-10-02T00:00:00Z","timestamp":970444800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/www.springer.com\/tdm"}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["EMBO J"],"published-print":{"date-parts":[[2000,10,2]]},"DOI":"10.1093\/emboj\/19.19.5135","type":"journal-article","created":{"date-parts":[[2002,7,26]],"date-time":"2002-07-26T18:48:15Z","timestamp":1027709295000},"page":"5135-5147","update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":58,"title":["MDM2 induces hyperplasia and premalignant lesions when expressed in the basal layer of the epidermis"],"prefix":"10.1038","volume":"19","author":[{"given":"Gitali","family":"Ganguli","sequence":"first","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Joseph","family":"Abecassis","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Bohdan","family":"Wasylyk","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"297","published-online":{"date-parts":[[2000,10,2]]},"reference":[{"key":"BFEMBJ7593328_CR1","doi-asserted-by":"crossref","first-page":"1419","DOI":"10.1038\/sj.onc.1202448","volume":"18","author":"M Alkhalaf","year":"1999","unstructured":"Alkhalaf M, Ganguli G, Messaddeq N, Le Meur M and Wasylyk B (1999) MDM2 overexpression generates a skin phenotype in both wild type and p53 null mice. Oncogene, 18, 1419\u20131434.","journal-title":"Oncogene"},{"key":"BFEMBJ7593328_CR2","first-page":"F1","volume":"1333","author":"N Almog","year":"1997","unstructured":"Almog N and Rotter V (1997) Involvement of p53 in cell differentiation and development. Biochim Biophys Acta, 1333, F1\u2013F27.","journal-title":"Biochim Biophys Acta"},{"key":"BFEMBJ7593328_CR3","first-page":"52","volume":"3","author":"HN Ananthaswamy","year":"1998","unstructured":"Ananthaswamy HN, Loughlin SM, Ullrich SE and Kripke ML (1998) Inhibition of UV\u2010induced p53 mutations by sunscreens: implications for skin cancer prevention. J Investig Dermatol Symp Proc, 3, 52\u201356.","journal-title":"J Investig Dermatol Symp Proc"},{"key":"BFEMBJ7593328_CR4","doi-asserted-by":"crossref","first-page":"274","DOI":"10.1073\/pnas.93.1.274","volume":"93","author":"RJ Berg","year":"1996","unstructured":"Berg RJ, van Kranen HJ, Rebel HG, de Vries A, van Vloten WA, Van Kreijl CF, van der Leun JC and de Gruijl FR (1996) Early p53 alterations in mouse skin carcinogenesis by UVB radiation: immunohistochemical detection of mutant p53 protein in clusters of preneoplastic epidermal cells. Proc Natl Acad Sci USA, 93, 274\u2013278.","journal-title":"Proc Natl Acad Sci USA"},{"key":"BFEMBJ7593328_CR5","doi-asserted-by":"crossref","first-page":"235","DOI":"10.1007\/BF01535205","volume":"13","author":"L Cahilly\u2010Snyder","year":"1987","unstructured":"Cahilly\u2010Snyder L, Yang\u2010Feng T, Francke U and George DL (1987) Molecular analysis and chromosomal mapping of amplified genes isolated from a transformed mouse 3T3 cell line. Somat Cell Mol Genet, 13, 235\u2013244.","journal-title":"Somat Cell Mol Genet"},{"key":"BFEMBJ7593328_CR6","doi-asserted-by":"crossref","first-page":"410","DOI":"10.1038\/7747","volume":"21","author":"EF Chan","year":"1999","unstructured":"Chan EF, Gat U, McNiff JM and Fuchs E (1999) A common human skin tumour is caused by activating mutations in \u03b2\u2010catenin. Nature Genet, 21, 410\u2013413.","journal-title":"Nature Genet"},{"key":"BFEMBJ7593328_CR7","doi-asserted-by":"crossref","first-page":"2295","DOI":"10.1083\/jcb.109.5.2295","volume":"109","author":"PA Coulombe","year":"1989","unstructured":"Coulombe PA, Kopan R and Fuchs E (1989) Expression of keratin K14 in the epidermis and hair follicle: insights into complex programs of differentiation. J Cell Biol, 109, 2295\u20132312.","journal-title":"J Cell Biol"},{"key":"BFEMBJ7593328_CR8","first-page":"1899","volume":"149","author":"LM Coussens","year":"1996","unstructured":"Coussens LM, Hanahan D and Arbeit JM (1996) Genetic predisposition and parameters of malignant progression in K14\u2010HPV16 transgenic mice. Am J Pathol, 149, 1899\u20131917.","journal-title":"Am J Pathol"},{"key":"BFEMBJ7593328_CR9","doi-asserted-by":"crossref","first-page":"1123","DOI":"10.1038\/sj.onc.1200922","volume":"14","author":"JE Dazard","year":"1997","unstructured":"Dazard JE, Augias D, Neel H, Mils V, Marechal V, Basset\u2010Seguin N and Piette J (1997) MDM\u20102 protein is expressed in different layers of normal human skin. Oncogene, 14, 1123\u20131128.","journal-title":"Oncogene"},{"key":"BFEMBJ7593328_CR10","doi-asserted-by":"crossref","first-page":"397","DOI":"10.1152\/physrev.1997.77.2.397","volume":"77","author":"RL Eckert","year":"1997","unstructured":"Eckert RL, Crish JF and Robinson NA (1997) The epidermal keratinocyte as a model for the study of gene regulation and cell differentiation. Physiol Rev, 77, 397\u2013424.","journal-title":"Physiol Rev"},{"key":"BFEMBJ7593328_CR11","doi-asserted-by":"crossref","first-page":"817","DOI":"10.1016\/0092-8674(93)90500-P","volume":"75","author":"WS el\u2010Deiry","year":"1993","unstructured":"el\u2010Deiry WS et al. (1993) WAF1, a potential mediator of p53 tumor suppression. Cell, 75, 817\u2013825.","journal-title":"Cell"},{"key":"BFEMBJ7593328_CR12","doi-asserted-by":"crossref","first-page":"1565","DOI":"10.1002\/j.1460-2075.1991.tb07676.x","volume":"10","author":"SS Fakharzadeh","year":"1991","unstructured":"Fakharzadeh SS, Trusko SP and George DL (1991) Tumorigenic potential associated with enhanced expression of a gene that is amplified in a mouse tumor cell line. EMBO J, 10, 1565\u20131569.","journal-title":"EMBO J"},{"key":"BFEMBJ7593328_CR13","doi-asserted-by":"crossref","first-page":"8945","DOI":"10.1074\/jbc.275.12.8945","volume":"275","author":"S Fang","year":"2000","unstructured":"Fang S, Jensen JP, Ludwig RL, Vousden KH and Weissman AM (2000) Mdm2 is a RING finger\u2010dependent ubiquitin protein ligase for itself and p53. J Biol Chem, 275, 8945\u20138951.","journal-title":"J Biol Chem"},{"key":"BFEMBJ7593328_CR14","doi-asserted-by":"crossref","first-page":"5048","DOI":"10.1128\/MCB.16.9.5048","volume":"16","author":"TA Fiddler","year":"1996","unstructured":"Fiddler TA, Smith L, Tapscott SJ and Thayer MJ (1996) Amplification of MDM2 inhibits MyoD\u2010mediated myogenesis. Mol Cell Biol, 16, 5048\u20135057.","journal-title":"Mol Cell Biol"},{"key":"BFEMBJ7593328_CR15","first-page":"301","volume":"13","author":"CA Finlay","year":"1993","unstructured":"Finlay CA (1993) The mdm\u20102 oncogene can overcome wild\u2010type p53 suppression of transformed cell growth. Mol Cell Biol, 13, 301\u2013306.","journal-title":"Mol Cell Biol"},{"key":"BFEMBJ7593328_CR16","doi-asserted-by":"crossref","first-page":"7288","DOI":"10.1128\/MCB.18.12.7288","volume":"18","author":"DA Freedman","year":"1998","unstructured":"Freedman DA and Levine AJ (1998) Nuclear export is required for degradation of endogenous p53 by MDM2 and human papillomavirus E6. Mol Cell Biol, 18, 7288\u20137293.","journal-title":"Mol Cell Biol"},{"key":"BFEMBJ7593328_CR17","doi-asserted-by":"crossref","first-page":"96","DOI":"10.1007\/s000180050273","volume":"55","author":"DA Freedman","year":"1999","unstructured":"Freedman DA, Wu L and Levine AJ (1999) Functions of the MDM2 oncoprotein. Cell Mol Life Sci, 55, 96\u2013107.","journal-title":"Cell Mol Life Sci"},{"key":"BFEMBJ7593328_CR18","doi-asserted-by":"crossref","first-page":"123","DOI":"10.1146\/annurev.cb.11.110195.001011","volume":"11","author":"E Fuchs","year":"1995","unstructured":"Fuchs E (1995) Keratins and the skin. Annu Rev Cell Dev Biol, 11, 123\u2013153.","journal-title":"Annu Rev Cell Dev Biol"},{"key":"BFEMBJ7593328_CR19","doi-asserted-by":"crossref","first-page":"1033","DOI":"10.1016\/0092-8674(80)90094-X","volume":"19","author":"E Fuchs","year":"1980","unstructured":"Fuchs E and Green H (1980) Changes in keratin gene expression during terminal differentiation of the keratinocyte. Cell, 19, 1033\u20131042.","journal-title":"Cell"},{"key":"BFEMBJ7593328_CR20","doi-asserted-by":"crossref","first-page":"99","DOI":"10.1002\/mc.2940070208","volume":"7","author":"DA Greenhalgh","year":"1993","unstructured":"Greenhalgh DA, Rothnagel JA, Quintanilla MI, Orengo CC, Gagne TA, Bundman DS, Longley MA and Roop DR (1993) Induction of epidermal hyperplasia, hyperkeratosis and papillomas in transgenic mice by a targeted v\u2010Ha\u2010ras oncogene. Mol Carcinogen, 7, 99\u2013110.","journal-title":"Mol Carcinogen"},{"key":"BFEMBJ7593328_CR21","doi-asserted-by":"crossref","first-page":"227","DOI":"10.3109\/10428199709051772","volume":"26","author":"DS Haines","year":"1997","unstructured":"Haines DS (1997) The mdm2 proto\u2010oncogene. Leuk Lymph, 26, 227\u2013238.","journal-title":"Leuk Lymph"},{"key":"BFEMBJ7593328_CR22","first-page":"203","volume":"8","author":"PA Hall","year":"1993","unstructured":"Hall PA, McKee PH, Menage HD, Dover R and Lane DP (1993) High levels of p53 protein in UV\u2010irradiated normal human skin. Oncogene, 8, 203\u2013207.","journal-title":"Oncogene"},{"key":"BFEMBJ7593328_CR23","doi-asserted-by":"crossref","first-page":"805","DOI":"10.1016\/0092-8674(93)90499-G","volume":"75","author":"JW Harper","year":"1993","unstructured":"Harper JW, Adami GR, Wei N, Keyomarsi K and Elledge SJ (1993) The p21 Cdk\u2010interacting protein Cip1 is a potent inhibitor of G1 cyclin\u2010dependent kinases. Cell, 75, 805\u2013816.","journal-title":"Cell"},{"key":"BFEMBJ7593328_CR24","doi-asserted-by":"crossref","first-page":"296","DOI":"10.1038\/387296a0","volume":"387","author":"Y Haupt","year":"1997","unstructured":"Haupt Y, Maya R, Kazaz A and Oren M (1997) Mdm2 promotes the rapid degradation of p53. Nature, 387, 296\u2013299.","journal-title":"Nature"},{"key":"BFEMBJ7593328_CR25","doi-asserted-by":"crossref","first-page":"1473","DOI":"10.1038\/sj.onc.1203464","volume":"19","author":"R Honda","year":"2000","unstructured":"Honda R and Yasuda H (2000) Activity of MDM2, a ubiquitin ligase, toward p53 or itself is dependent on the RING finger domain of the ligase. Oncogene, 19, 1473\u20131476.","journal-title":"Oncogene"},{"key":"BFEMBJ7593328_CR26","doi-asserted-by":"crossref","first-page":"25","DOI":"10.1016\/S0014-5793(97)01480-4","volume":"420","author":"R Honda","year":"1997","unstructured":"Honda R, Tanaka H and Yasuda H (1997) Oncoprotein MDM2 is a ubiquitin ligase E3 for tumor suppressor p53. FEBS Lett, 420, 25\u201327.","journal-title":"FEBS Lett"},{"key":"BFEMBJ7593328_CR27","doi-asserted-by":"crossref","first-page":"1001","DOI":"10.1128\/MCB.20.3.1001-1007.2000","volume":"20","author":"MW Jackson","year":"2000","unstructured":"Jackson MW and Berberich SJ (2000) MdmX protects p53 from Mdm2\u2010mediated degradation. Mol Cell Biol, 20, 1001\u20131007.","journal-title":"Mol Cell Biol"},{"key":"BFEMBJ7593328_CR28","doi-asserted-by":"crossref","first-page":"4247","DOI":"10.1038\/sj.onc.1202789","volume":"18","author":"W Jiang","year":"1999","unstructured":"Jiang W, Ananthaswamy HN, Muller HK and Kripke ML (1999) p53 protects against skin cancer induction by UV\u2010B radiation. Oncogene, 18, 4247\u20134253.","journal-title":"Oncogene"},{"key":"BFEMBJ7593328_CR29","doi-asserted-by":"crossref","first-page":"14025","DOI":"10.1073\/pnas.93.24.14025","volume":"93","author":"AS Jonason","year":"1996","unstructured":"Jonason AS, Kunala S, Price GJ, Restifo RJ, Spinelli HM, Persing JA, Leffell DJ, Tarone RE and Brash DE (1996) Frequent clones of p53\u2010mutated keratinocytes in normal human skin. Proc Natl Acad Sci USA, 93, 14025\u201314029.","journal-title":"Proc Natl Acad Sci USA"},{"key":"BFEMBJ7593328_CR30","doi-asserted-by":"crossref","first-page":"206","DOI":"10.1038\/378206a0","volume":"378","author":"SN Jones","year":"1995","unstructured":"Jones SN, Roe AE, Donehower LA and Bradley A (1995) Rescue of embryonic lethality in Mdm2\u2010deficient mice by absence of p53. Nature, 378, 206\u2013208.","journal-title":"Nature"},{"key":"BFEMBJ7593328_CR31","doi-asserted-by":"crossref","first-page":"15608","DOI":"10.1073\/pnas.95.26.15608","volume":"95","author":"SN Jones","year":"1998","unstructured":"Jones SN, Hancock AR, Vogel H, Donehower LA and Bradley A (1998) Overexpression of Mdm2 in mice reveals a p53\u2010independent role for Mdm2 in tumorigenesis. Proc Natl Acad Sci USA, 95, 15608\u201315612.","journal-title":"Proc Natl Acad Sci USA"},{"key":"BFEMBJ7593328_CR32","doi-asserted-by":"crossref","first-page":"71","DOI":"10.1007\/BF03402141","volume":"5","author":"T Juven\u2010Gershon","year":"1999","unstructured":"Juven\u2010Gershon T and Oren M (1999) Mdm2: the ups and downs. Mol Med, 5, 71\u201383.","journal-title":"Mol Med"},{"key":"BFEMBJ7593328_CR33","doi-asserted-by":"crossref","first-page":"7701","DOI":"10.1038\/sj.onc.1202955","volume":"18","author":"Jr WG Kaelin","year":"1999","unstructured":"Kaelin WG,Jr (1999) The p53 gene family. Oncogene, 18, 7701\u20137705.","journal-title":"Oncogene"},{"key":"BFEMBJ7593328_CR34","doi-asserted-by":"crossref","first-page":"299","DOI":"10.1038\/387299a0","volume":"387","author":"MH Kubbutat","year":"1997","unstructured":"Kubbutat MH, Jones SN and Vousden KH (1997) Regulation of p53 stability by Mdm2. Nature, 387, 299\u2013303.","journal-title":"Nature"},{"key":"BFEMBJ7593328_CR35","doi-asserted-by":"crossref","first-page":"457","DOI":"10.1006\/excr.1999.4433","volume":"248","author":"S Lain","year":"1999","unstructured":"Lain S, Midgley C, Sparks A, Lane EB and Lane DP (1999) An inhibitor of nuclear export activates the p53 response and induces the localization of HDM2 and p53 to U1A\u2010positive nuclear bodies associated with the PODs. Exp Cell Res, 248, 457\u2013472.","journal-title":"Exp Cell Res"},{"key":"BFEMBJ7593328_CR36","doi-asserted-by":"crossref","first-page":"18","DOI":"10.1067\/mjd.2000.103340","volume":"42","author":"DJ Leffell","year":"2000","unstructured":"Leffell DJ (2000) The scientific basis of skin cancer. J Am Acad Dermatol, 42, 18\u201322.","journal-title":"J Am Acad Dermatol"},{"key":"BFEMBJ7593328_CR37","first-page":"486","volume":"8","author":"R Lersch","year":"1988","unstructured":"Lersch R and Fuchs E (1988) Sequence and expression of a type II keratin, K5, in human epidermal cells. Mol Cell Biol, 8, 486\u2013493.","journal-title":"Mol Cell Biol"},{"key":"BFEMBJ7593328_CR38","doi-asserted-by":"crossref","first-page":"189","DOI":"10.1016\/S0925-4773(98)00074-4","volume":"74","author":"T Leveillard","year":"1998","unstructured":"Leveillard T, Gorry P, Niederreither K and Wasylyk B (1998) MDM2 expression during mouse embryogenesis and the requirement of p53. Mech Dev, 74, 189\u2013193.","journal-title":"Mech Dev"},{"key":"BFEMBJ7593328_CR39","doi-asserted-by":"crossref","first-page":"1661","DOI":"10.1242\/jcs.113.10.1661","volume":"113","author":"M Levrero","year":"2000","unstructured":"Levrero M, De Laurenzi V, Costanzo A, Gong J, Wang JY and Melino G (2000) The p53\/p63\/p73 family of transcription factors: overlapping and distinct functions. J Cell Sci, 113, 1661\u20131670.","journal-title":"J Cell Sci"},{"key":"BFEMBJ7593328_CR40","doi-asserted-by":"crossref","first-page":"197","DOI":"10.1016\/S0962-8924(00)01736-0","volume":"10","author":"MA Lohrum","year":"2000","unstructured":"Lohrum MA and Vousden KH (2000) Regulation and function of the p53\u2010related proteins: same family, different rules. Trends Cell Biol, 10, 197\u2013202.","journal-title":"Trends Cell Biol"},{"key":"BFEMBJ7593328_CR41","doi-asserted-by":"crossref","first-page":"714","DOI":"10.1101\/gad.11.6.714","volume":"11","author":"K Lundgren","year":"1997","unstructured":"Lundgren K, Montes de Oca Luna R, McNeill YB, Emerick EP, Spencer B, Barfield CR, Lozano G, Rosenberg MP and Finlay CA (1997) Targeted expression of MDM2 uncouples S phase from mitosis and inhibits mammary gland development independent of p53. Genes Dev, 11, 714\u2013725.","journal-title":"Genes Dev"},{"key":"BFEMBJ7593328_CR42","doi-asserted-by":"crossref","first-page":"708","DOI":"10.1038\/19531","volume":"398","author":"AA Mills","year":"1999","unstructured":"Mills AA, Zheng B, Wang XJ, Vogel H, Roop DR and Bradley A (1999) p63 is a p53 homologue required for limb and epidermal morphogenesis. Nature, 398, 708\u2013713.","journal-title":"Nature"},{"key":"BFEMBJ7593328_CR43","doi-asserted-by":"crossref","first-page":"293","DOI":"10.1016\/0092-8674(95)90513-8","volume":"80","author":"T Miyashita","year":"1995","unstructured":"Miyashita T and Reed JC (1995) Tumor suppressor p53 is a direct transcriptional activator of the human bax gene. Cell, 80, 293\u2013299.","journal-title":"Cell"},{"key":"BFEMBJ7593328_CR44","doi-asserted-by":"crossref","first-page":"3453","DOI":"10.1093\/nar\/26.15.3453","volume":"26","author":"J Momand","year":"1998","unstructured":"Momand J, Jung D, Wilczynski S and Niland J (1998) The MDM2 gene amplification database. Nucleic Acids Res, 26, 3453\u20133459.","journal-title":"Nucleic Acids Res"},{"key":"BFEMBJ7593328_CR45","doi-asserted-by":"crossref","first-page":"15","DOI":"10.1016\/S0378-1119(99)00487-4","volume":"242","author":"J Momand","year":"2000","unstructured":"Momand J, Wu HH and Dasgupta G (2000) MDM2\u2014master regulator of the p53 tumor suppressor protein. Gene, 242, 15\u201329.","journal-title":"Gene"},{"key":"BFEMBJ7593328_CR46","doi-asserted-by":"crossref","first-page":"203","DOI":"10.1038\/378203a0","volume":"378","author":"R Montes de Oca Luna","year":"1995","unstructured":"Montes de Oca Luna R, Wagner DS and Lozano G (1995) Rescue of early embryonic lethality in mdm2\u2010deficient mice by deletion of p53. Nature, 378, 203\u2013206.","journal-title":"Nature"},{"key":"BFEMBJ7593328_CR47","doi-asserted-by":"crossref","first-page":"244","DOI":"10.1083\/jcb.97.1.244","volume":"97","author":"WG Nelson","year":"1983","unstructured":"Nelson WG and Sun TT (1983) The 50\u2010 and 58\u2010kDa keratin classes as molecular markers for stratified squamous epithelia: cell culture studies. J Cell Biol, 97, 244\u2013251.","journal-title":"J Cell Biol"},{"key":"BFEMBJ7593328_CR48","first-page":"247","volume":"89","author":"K Niederreither","year":"1998","unstructured":"Niederreither K and Dolle P (1998) In situ hybridization with 35S\u2010labeled probes for retinoid receptors. Methods Mol Biol, 89, 247\u2013267.","journal-title":"Methods Mol Biol"},{"key":"BFEMBJ7593328_CR49","doi-asserted-by":"crossref","first-page":"80","DOI":"10.1038\/358080a0","volume":"358","author":"JD Oliner","year":"1992","unstructured":"Oliner JD, Kinzler KW, Meltzer PS, George DL and Vogelstein B (1992) Amplification of a gene encoding a p53\u2010associated protein in human sarcomas. Nature, 358, 80\u201383.","journal-title":"Nature"},{"key":"BFEMBJ7593328_CR50","doi-asserted-by":"crossref","first-page":"857","DOI":"10.1038\/362857a0","volume":"362","author":"JD Oliner","year":"1993","unstructured":"Oliner JD, Pietenpol JA, Thiagalingam S, Gyuris J, Kinzler KW and Vogelstein B (1993) Oncoprotein MDM2 conceals the activation domain of tumour suppressor p53. Nature, 362, 857\u2013860.","journal-title":"Nature"},{"key":"BFEMBJ7593328_CR51","doi-asserted-by":"crossref","first-page":"817","DOI":"10.1126\/science.276.5313.817","volume":"276","author":"AE Oro","year":"1997","unstructured":"Oro AE, Higgins KM, Hu Z, Bonifas JM, Epstein EH,Jr and Scott MP (1997) Basal cell carcinomas in mice overexpressing sonic hedgehog. Science, 276, 817\u2013821.","journal-title":"Science"},{"key":"BFEMBJ7593328_CR52","doi-asserted-by":"crossref","first-page":"201","DOI":"10.1016\/S0002-9440(10)64720-7","volume":"156","author":"A Ouhtit","year":"2000","unstructured":"Ouhtit A, Muller HK, Davis DW, Ullrich SE, McConkey D and Ananthaswamy HN (2000) Temporal events in skin injury and the early adaptive responses in ultraviolet\u2010irradiated mouse skin. Am J Pathol, 156, 201\u2013207.","journal-title":"Am J Pathol"},{"key":"BFEMBJ7593328_CR53","doi-asserted-by":"crossref","first-page":"949","DOI":"10.1038\/sj.onc.1202031","volume":"17","author":"JM Paramio","year":"1998","unstructured":"Paramio JM, Lain S, Segrelles C, Lane EB and Jorcano JL (1998) Differential expression and functionally co\u2010operative roles for the retinoblastoma family of proteins in epidermal differentiation. Oncogene, 17, 949\u2013957.","journal-title":"Oncogene"},{"key":"BFEMBJ7593328_CR54","doi-asserted-by":"crossref","first-page":"1099","DOI":"10.1046\/j.1523-1747.1999.00780.x","volume":"113","author":"R Parsa","year":"1999","unstructured":"Parsa R, Yang A, McKeon F and Green H (1999) Association of p63 with proliferative potential in normal and neoplastic human keratinocytes. J Invest Dermatol, 113, 1099\u20131105.","journal-title":"J Invest Dermatol"},{"key":"BFEMBJ7593328_CR55","doi-asserted-by":"crossref","first-page":"429","DOI":"10.3109\/02841869609109917","volume":"35","author":"SM Picksley","year":"1996","unstructured":"Picksley SM, Spicer JF, Barnes DM and Lane DP (1996) The p53\u2013MDM2 interaction in a cancer\u2010prone family and the identification of a novel therapeutic target. Acta Oncol, 35, 429\u2013434.","journal-title":"Acta Oncol"},{"key":"BFEMBJ7593328_CR56","doi-asserted-by":"crossref","first-page":"1001","DOI":"10.1038\/sj.onc.1201432","volume":"15","author":"J Piette","year":"1997","unstructured":"Piette J, Neel H and Marechal V (1997) Mdm2: keeping p53 under control. Oncogene, 15, 1001\u20131010.","journal-title":"Oncogene"},{"key":"BFEMBJ7593328_CR57","doi-asserted-by":"crossref","first-page":"5","DOI":"10.1016\/S0092-8674(00)81774-2","volume":"95","author":"C Prives","year":"1998","unstructured":"Prives C (1998) Signaling to p53: breaking the MDM2\u2013p53 circuit. Cell, 95, 5\u20138.","journal-title":"Cell"},{"key":"BFEMBJ7593328_CR58","first-page":"147","volume":"10","author":"V Reinke","year":"1999","unstructured":"Reinke V, Bortner DM, Amelse LL, Lundgren K, Rosenberg MP, Finlay CA and Lozano G (1999) Overproduction of MDM2 in vivo disrupts S phase independent of E2F1. Cell Growth Differ, 10, 147\u2013154.","journal-title":"Cell Growth Differ"},{"key":"BFEMBJ7593328_CR59","doi-asserted-by":"crossref","first-page":"554","DOI":"10.1093\/emboj\/17.2.554","volume":"17","author":"J Roth","year":"1998","unstructured":"Roth J, Dobbelstein M, Freedman DA, Shenk T and Levine AJ (1998) Nucleo\u2010cytoplasmic shuttling of the hdm2 oncoprotein regulates the levels of the p53 protein via a pathway used by the human immunodeficiency virus rev protein. EMBO J, 17, 554\u2013564.","journal-title":"EMBO J"},{"key":"BFEMBJ7593328_CR60","doi-asserted-by":"crossref","first-page":"865","DOI":"10.1093\/carcin\/21.5.865","volume":"21","author":"M Serrano","year":"2000","unstructured":"Serrano M (2000) The INK4a\/ARF locus in murine tumorigenesis. Carcinogenesis, 21, 865\u2013869.","journal-title":"Carcinogenesis"},{"key":"BFEMBJ7593328_CR61","doi-asserted-by":"crossref","first-page":"94","DOI":"10.1016\/S0959-437X(99)00038-6","volume":"10","author":"CJ Sherr","year":"2000","unstructured":"Sherr CJ and Weber JD (2000) The ARF\/p53 pathway. Curr Opin Genet Dev, 10, 94\u201399.","journal-title":"Curr Opin Genet Dev"},{"key":"BFEMBJ7593328_CR62","doi-asserted-by":"crossref","first-page":"D538","DOI":"10.2741\/A211","volume":"2","author":"H Soehnge","year":"1997","unstructured":"Soehnge H, Ouhtit A and Ananthaswamy ON (1997) Mechanisms of induction of skin cancer by UV radiation. Frontiers Biosci, 2, D538\u2013D551.","journal-title":"Frontiers Biosci"},{"key":"BFEMBJ7593328_CR63","doi-asserted-by":"crossref","first-page":"1121","DOI":"10.1016\/S0002-9440(10)65215-7","volume":"155","author":"S Song","year":"1999","unstructured":"Song S and Lambert PF (1999) Different responses of epidermal and hair follicular cells to radiation correlate with distinct patterns of p53 and p21 induction. Am J Pathol, 155, 1121\u20131127.","journal-title":"Am J Pathol"},{"key":"BFEMBJ7593328_CR64","doi-asserted-by":"crossref","first-page":"427","DOI":"10.1083\/jcb.107.2.427","volume":"107","author":"A Stoler","year":"1988","unstructured":"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\u2013446.","journal-title":"J Cell Biol"},{"key":"BFEMBJ7593328_CR65","doi-asserted-by":"crossref","first-page":"2270","DOI":"10.1126\/science.282.5397.2270","volume":"282","author":"P Sun","year":"1998","unstructured":"Sun P, Dong P, Dai K, Hannon GJ and Beach D (1998) p53\u2010independent role of MDM2 in TGF\u2010\u03b21 resistance. Science, 282, 2270\u20132272.","journal-title":"Science"},{"key":"BFEMBJ7593328_CR66","doi-asserted-by":"crossref","first-page":"3077","DOI":"10.1073\/pnas.96.6.3077","volume":"96","author":"W Tao","year":"1999","unstructured":"Tao W and Levine AJ (1999) Nucleocytoplasmic shuttling of oncoprotein Hdm2 is required for Hdm2\u2010mediated degradation of p53. Proc Natl Acad Sci USA, 96, 3077\u20133080.","journal-title":"Proc Natl Acad Sci USA"},{"key":"BFEMBJ7593328_CR67","doi-asserted-by":"crossref","first-page":"16","DOI":"10.1177\/120347549800300104","volume":"3","author":"VA Tron","year":"1998","unstructured":"Tron VA, Trotter MJ, Ishikawa T, Ho VC and Li G (1998a) p53\u2010dependent regulation of nucleotide excision repair in murine epidermis in vivo. J Cutan Med Surg, 3, 16\u201320.","journal-title":"J Cutan Med Surg"},{"key":"BFEMBJ7593328_CR68","doi-asserted-by":"crossref","first-page":"579","DOI":"10.1016\/S0002-9440(10)65600-3","volume":"153","author":"VA Tron","year":"1998","unstructured":"Tron VA, Trotter MJ, Tang L, Krajewska M, Reed JC, Ho VC and Li G (1998b) p53\u2010regulated apoptosis is differentiation dependent in ultraviolet B\u2010irradiated mouse keratinocytes. Am J Pathol, 153, 579\u2013585.","journal-title":"Am J Pathol"},{"key":"BFEMBJ7593328_CR69","doi-asserted-by":"crossref","first-page":"280","DOI":"10.1177\/120347549900300511","volume":"3","author":"VA Tron","year":"1999","unstructured":"Tron VA, Li G, Ho V and Trotter MJ (1999) Ultraviolet radiation\u2010induced p53 responses in the epidermis are differentiation\u2010dependent. J Cutan Med Surg, 3, 280\u2013283.","journal-title":"J Cutan Med Surg"},{"key":"BFEMBJ7593328_CR70","doi-asserted-by":"crossref","first-page":"714","DOI":"10.1101\/gad.5.5.714","volume":"5","author":"R Vassar","year":"1991","unstructured":"Vassar R and Fuchs E (1991) Transgenic mice provide new insights into the role of TGF\u2010\u03b1 during epidermal development and differentiation. Genes Dev, 5, 714\u2013727.","journal-title":"Genes Dev"},{"key":"BFEMBJ7593328_CR71","doi-asserted-by":"crossref","first-page":"1563","DOI":"10.1073\/pnas.86.5.1563","volume":"86","author":"R Vassar","year":"1989","unstructured":"Vassar R, Rosenberg M, Ross S, Tyner A and Fuchs E (1989) Tissue\u2010specific and differentiation\u2010specific expression of a human K14 keratin gene in transgenic mice. Proc Natl Acad Sci USA, 86, 1563\u20131567.","journal-title":"Proc Natl Acad Sci USA"},{"key":"BFEMBJ7593328_CR72","first-page":"4643","volume":"12","author":"R Vassar","year":"1992","unstructured":"Vassar R, Hutton ME and Fuchs E (1992) Transgenic overexpression of transforming growth factor \u03b1 bypasses the need for c\u2010Ha\u2010ras mutations in mouse skin tumorigenesis. Mol Cell Biol, 12, 4643\u20134653.","journal-title":"Mol Cell Biol"},{"key":"BFEMBJ7593328_CR73","doi-asserted-by":"crossref","first-page":"3417","DOI":"10.1128\/MCB.20.10.3417-3424.2000","volume":"20","author":"D Wang","year":"2000","unstructured":"Wang D, Russell JL and Johnson DG (2000) E2F4 and E2F1 have similar proliferative properties but different apoptotic and oncogenic properties in vivo. Mol Cell Biol, 20, 3417\u20133424.","journal-title":"Mol Cell Biol"},{"key":"BFEMBJ7593328_CR74","doi-asserted-by":"crossref","first-page":"219","DOI":"10.1073\/pnas.94.1.219","volume":"94","author":"X Wang","year":"1997","unstructured":"Wang X, Zinkel S, Polonsky K and Fuchs E (1997) Transgenic studies with a keratin promoter\u2010driven growth hormone transgene: prospects for gene therapy. Proc Natl Acad Sci USA, 94, 219\u2013226.","journal-title":"Proc Natl Acad Sci USA"},{"key":"BFEMBJ7593328_CR75","first-page":"2271","volume":"10","author":"WC Weinberg","year":"1995","unstructured":"Weinberg WC, Azzoli CG, Chapman K, Levine AJ and Yuspa SH (1995) p53\u2010mediated transcriptional activity increases in differentiating epidermal keratinocytes in association with decreased p53 protein. Oncogene, 10, 2271\u20132279.","journal-title":"Oncogene"},{"key":"BFEMBJ7593328_CR76","doi-asserted-by":"crossref","first-page":"1397","DOI":"10.1083\/jcb.98.4.1397","volume":"98","author":"RA Weiss","year":"1984","unstructured":"Weiss RA, Eichner R and Sun TT (1984) Monoclonal antibody analysis of keratin expression in epidermal diseases: a 48\u2010 and 56\u2010kdalton keratin as molecular markers for hyperproliferative keratinocytes. J Cell Biol, 98, 1397\u20131406.","journal-title":"J Cell Biol"},{"key":"BFEMBJ7593328_CR77","first-page":"313","volume":"9","author":"W Xie","year":"1998","unstructured":"Xie W, Wu X, Chow LT, Chin E, Paterson AJ and Kudlow JE (1998) Targeted expression of activated erbB\u20102 to the epidermis of transgenic mice elicits striking developmental abnormalities in the epidermis and hair follicles. Cell Growth Differ, 9, 313\u2013325.","journal-title":"Cell Growth Differ"},{"key":"BFEMBJ7593328_CR78","first-page":"5075","volume":"59","author":"CH Yam","year":"1999","unstructured":"Yam CH, Siu WY, Arooz T, Chiu CH, Lau A, Wang XQ and Poon RY (1999) MDM2 and MDMX inhibit the transcriptional activity of ectopically expressed SMAD proteins. Cancer Res, 59, 5075\u20135078.","journal-title":"Cancer Res"},{"key":"BFEMBJ7593328_CR79","doi-asserted-by":"crossref","first-page":"714","DOI":"10.1038\/19539","volume":"398","author":"A Yang","year":"1999","unstructured":"Yang A et al. (1999) p63 is essential for regenerative proliferation in limb, craniofacial and epithelial development. Nature, 398, 714\u2013718.","journal-title":"Nature"},{"key":"BFEMBJ7593328_CR80","doi-asserted-by":"crossref","first-page":"773","DOI":"10.1038\/372773a0","volume":"372","author":"A Ziegler","year":"1994","unstructured":"Ziegler A, Jonason AS, Leffell DJ, Simon JA, Sharma HW, Kimmelman J, Remington L, Jacks T and Brash DE (1994) Sunburn and p53 in the onset of skin cancer. Nature, 372, 773\u2013776.","journal-title":"Nature"}],"container-title":["The EMBO Journal"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/api.wiley.com\/onlinelibrary\/tdm\/v1\/articles\/10.1093%2Femboj%2F19.19.5135","content-type":"unspecified","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1093\/emboj\/19.19.5135.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/article\/10.1093\/emboj\/19.19.5135\/fulltext.html","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/full\/10.1093\/emboj\/19.19.5135","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1093\/emboj\/19.19.5135.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,5,20]],"date-time":"2026-05-20T18:35:12Z","timestamp":1779302112000},"score":1,"resource":{"primary":{"URL":"https:\/\/link.springer.com\/10.1093\/emboj\/19.19.5135"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2000,10,2]]},"references-count":80,"journal-issue":{"issue":"19","published-print":{"date-parts":[[2000,10,2]]}},"alternative-id":["BFEMBJ7593328"],"URL":"https:\/\/doi.org\/10.1093\/emboj\/19.19.5135","relation":{},"ISSN":["0261-4189","1460-2075"],"issn-type":[{"value":"0261-4189","type":"print"},{"value":"1460-2075","type":"electronic"}],"subject":[],"published":{"date-parts":[[2000,10,2]]},"assertion":[{"value":"26 June 2000","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"3 August 2000","order":2,"name":"revised","label":"Revised","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"3 August 2000","order":3,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"2 October 2000","order":4,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"This content has been made available to all.","name":"free","label":"Free to read"}]}}