Abstract
Solar ultraviolet (UV) radiation is a major environmental factor that causes DNA damage, inflammation, erythema, sunburn, immunosuppression, photoaging, gene mutations, and skin cancer. p38 mitogen activated protein kinase (MAPK) are strongly activated by UV radiation, and play important roles in regulating cellular responses to UV. In this review, we examine the role played by p38 MAPK in mediating UV-induced cell cycle, apoptosis, inflammation, and skin tanning response. We review the role played by p38 MAPK in transcriptional regulation of key downstream genes that have been implicated in the regulation of cellular responses to UV radiation. Understanding this will undoubtedly help in the prevention and control of UV-induced damage and the development of novel therapeutic strategies.
Similar content being viewed by others
References
de Gruijl F.R. (2000) Photocarcinogenesis: UVA vs. UVB Methods Enzymol. 319:359–366
Matsui M.S., DeLeo V.A. (1991) Longwave ultraviolet radiation and promotion of skin cancer. Cancer Cells 3:8–12
de Gruijl F.R., Sterenborg H.J., Forbes P.D., Davies R.E., Cole C., Kelfkens G., van Weelden H., Slaper H., van der Leun J.C. (1993) Wavelength dependence of skin cancer induction by ultraviolet irradiation of albino hairless mice. Cancer Res. 53:53–60
Beissert S., Schwarz T. (1999) Mechanisms involved in ultraviolet light-induced immunosuppression. J. Invest. Dermatol. Symp. Proc. 4:61–64
Fisher G.J., Datta S.C., Talwar H.S., Wang Z.Q., Varani J., Kang S., Voorhees J.J. (1996) Molecular basis of sun-induced premature skin ageing and retinoid antagonism. Nature 379:335–339
Whitmarsh A.J., Davis R.J. (1996) Transcription factor AP-1 regulation by mitogen-activated protein kinase signal transduction pathways. J. Mol. Med. 74:589–607
Bachelor M.A., Cooper S.J., Sikorski E.T., Bowden G.T. (2005) Inhibition of p38 mitogen-activated protein kinase and phosphatidylinositol 3-kinase decreases UVB-induced activator protein-1 and cyclooxygenase-2 in a SKH-1 hairless mouse model. Mol. Cancer Res. 3:90–99
Black H.S., de Gruijl F.R., Forbes P.D., Cleaver J.E., Ananthaswamy H.N., de Fabo E.C., Ullrich S.E., Tyrrell R.M. (1997) Photocarcinogenesis: an overview. J. Photochem. Photobiol. B 40:29–47
Ananthaswamy H.N., Pierceall W.E. (1992) Molecular alterations in human skin tumors. Prog. Clin. Biol. Res. 376:61–84
Katiyar S.K., Perez A., Mukhtar H. (2000) Green tea polyphenol treatment to human skin prevents formation of ultraviolet light B-induced pyrimidine dimers in DNA. Clin. Cancer Res. 10:3864–3869
Katiyar S.K., Afaq F., Azizuddin K., Mukhtar H. (2001) Inhibition of UVB-induced oxidative stress-mediated phosphorylation of mitogen-activated protein kinase signaling pathways in cultured human epidermal keratinocytes by green tea polyphenol (-)-epigallocatechin-3-gallate. Toxicol. Appl. Pharmacol. 176:110–117
Katiyar S.K., Afaq F., Perez A., Mukhtar H. (2001) Green tea polyphenol (-)-epigallocatechin-3-gallate treatment of human skin inhibits ultraviolet radiation-induced oxidative stress. Carcinogenesis 22:287–294
Sander C.S., Chang H., Hamm F., Elsner P., Thiele J.J. (2004) Role of oxidative stress and the antioxidant network in cutaneous carcinogenesis. Int. J. Dermatol. 43:326–335
Ichihashi M., Ueda M., Budiyanto A., Bito T., Oka M., Fukunaga M., Tsuru K., Hotikawa T. (2003) UV-induced skin damage. Toxicology 189:21–39
Cerutti P.A. (1985) Prooxidant states and tumor promotion. Science 227:375–381
Peus D., Vasa R.A., Beyerle A., Meves A., Krautmacher C., Pittelkow M.R. (1999) UVB activates ERK1/2 and p38 signaling pathways via reactive oxygen species in cultured keratinocytes. J. Invest. Dermatol. 112:751–756
Kabuyama Y., Homma M.K., Sekimata M., Homma Y. (2001) Wavelength-specific activation of MAP kinase family proteins by monochromatic UV irradiation. Photochem. Photobiol. 73:147–152
Ding M., Li J., Leonard S.S., Shi X., Costa M., Castranova V., Vallyathan V., Huang C. (2002) Differential role of hydrogen peroxide in UV-induced signal transduction. Mol. Cell Biochem. 234–235:81–90
Gotoh Y., Cooper J.A. (1998) Reactive oxygen species- and dimerization-induced activation of apoptosis signal-regulating kinase 1 in tumor necrosis factor-alpha signal transduction. J. Biol. Chem. 273:17477–17482
Seo M., Lee Y.I., Cho C.H., Bae C.D., Kim I.H., Juhnn Y.S. (2002) Bi-directional regulation of UV-induced activation of p38 kinase and c-Jun N-terminal kinase by G protein βγ-subunits. J. Biol. Chem. 277:24197–24203
Seo M., Cho C.H., Lee Y.I., Shin E.Y., Park D., Bae C.D., Lee J.W., Lee E.S., Juhnn Y.S. (2004) Cdc42-dependent mediation of UV-induced p38 activation by G protein βγ-subunits. J. Biol. Chem. 279:17366–17375
Derijard B., Raingeaud J., Barrett T., Wu I.H., Han J., Ulevitch R.J., Davis R.J. (1995) Independent human MAP-kinase signal transduction pathways defined by MEK and MKK isoforms. Science. 267:682–685
Stein B., Brady H., Yang M.X., Young D.B., Barbosa M.S. (1996) Cloning and characterization of MEK6, a novel member of the mitogen-activated protein kinase kinase cascade. J. Biol. Chem. 271:11427–11433
Rouse J., Cohen P., Trigon S., Morange M., Alonso-Llamazares A., Zamanillo D., Hunt T., Nebreda A.R. (1994) A novel kinase cascade triggered by stress and heat shock that stimulates MAPKAP kinase-2 and phosphorylation of the small heat shock proteins. Cell 78:1027–1037
McLaughlin M.M., Kumar S., McDonnell P.C., Van Hom S., Lee J.C., Livi G.P., Young P.R. (1996) Identification of mitogen-activated protein (MAP) kinase-activated protein kinase-3, a novel substrate of CSBP p38 MAP kinase. J. Biol. Chem. 271:8488–8492
Waskiewicz A.J., Flynn A., Proud C.G., Cooper J.A. (1997) Mitogen-activated protein kinases activate the serine/threonine kinases Mnk1 and Mnk2. EMBO J. 16:1909–1920
Fukunaga R., Hunter T. (1997) MNK1, a new MAP kinase-activated protein kinase, isolated by a novel expression screening method for identifying protein kinase substrates. EMBO J. 16:1921–1933
New L., Jiang Y., Zhao M., Liu K., Zhu W., Flood L.J., Kato Y., Parry G.C., Han J. (1998) PRAK, a novel protein kinase regulated by the p38 MAP kinase. EMBO J. 17:3372–3384
Deak M., Clifton A.D., Lucocq L.M., Alessi D.R. (1998) Mitogen- and stress-activated protein kinase-1 (MSK1) is directly activated by MAPK and SAPK2/p38, and may mediate activation of CREB. EMBO J. 17:4426–4441
Iordanov M., Bender K., Ade T., Schmid W., Sachsenmaier C., Engel K., Gaestel M., Rahmsdorf H.J., Herrlich P. (1997) CREB is activated by UVC through a p38/HOG-1-dependent protein kinase. EMBO J. 16:1009–1022
Robinson M.J., Cobb M.H. (1997) Mitogen-activated protein kinase pathways. Curr. Opin. Cell Biol. 9:180–186
Martin-Blanco E. (2000) p38 MAPK signaling cascades: ancient roles and new functions. Bioessays 22:637–645
Maltzman W., Czyzyk L. (1984) UV irradiation stimulates levels of p53 cellular tumor antigen in nontransformed mouse cells. Mol. Cell. Biol. 4:1689–1694
Ouhtit A., Muller H.K., Davis D.W., Ullrich S.E., McConkey D., Ananthaswamy H.N. (2000) Temporal events in skin injury and the early adaptive responses in ultraviolet-irradiated mouse skin. Am. J. Pathol. 156:201–207
Smith M.L., Fornace A.J. Jr. (1997) p53-mediated protective responses to UV irradiation. Proc. Natl. Acad. Sci. USA 94:12255–12257
Hall P.A., McKee P.H., Menage H.D., Dover R., Lane D.P. (1993) High levels of p53 protein in UV-irradiated normal human skin. Oncogene 8:203–207
McKay B.C., Francis M.A., Rainbow A.J. (1997) Wildtype p53 is required for heat shock and ultraviolet light enhanced repair of a UV-damaged reporter gene. Carcinogenesis 18:245–249
Meek D.W. (1998) Multisite phosphorylation and the integration of stress signals at p53. Cell Signal. 10:159–166
Giaccia A.J., Kastan M.B. (1998) The complexity of p53 modulation: emerging patterns from divergent signals. Genes Dev. 12:2973–2983
Sakaguchi K., Herrera J.E., Saito S., Miki T., Bustin M., Vassilev A., Anderson C.W., Appella E. (1998) DNA damage activates p53 through a phosphorylation-acetylation cascade. Genes Dev. 12:2831–2841
Shieh S.Y., Taya Y., Prives C. (1999) DNA damage-inducible phosphorylation of p53 at N-terminal sites including a novel site, ser20, requires tetramerization. EMBO J. 18:1815–1823
Unger T., Sionov R.V., Moallem E., Yee C.L., Howley P.M., Oren M., Haupt Y. (1999) Mutations in serines 15 and 20 of human p53 impair its apoptotic activity. Oncogene 18:3205–3212
She Q.B., Chen N., Dong Z. (2000) ERKs and p38 kinase phosphorylate p53 protein at serine 15 in response to UV radiation. J. Biol. Chem. 275:20444–20449
Bulavin D.V., Saito S., Hollander M.C., Sakaguchi K., Anderson C.W., Appella E., Fornace A.J. Jr. (1999) Phosphorylation of human p53 by p38 kinase coordinates N-terminal phosphorylation and apoptosis in response to UV radiation. EMBO J. 18:6845–6854
Huang C., Ma W.Y., Maxiner A., Sun Y., Dong Z. (1999) p38 kinase mediates UV-induced phosphorylation of p53 protein at serine 389. J. Biol. Chem. 274:12229–12235
Keller D., Zeng X., Li X., Kapoor M., Iordanov M.S., Taya Y., Lozano G., Magun B., Lu H. (1999) The p38MAPK inhibitor SB203580 alleviates ultraviolet-induced phosphorylation at serine 389 but not serine 15 and activation of p53. Biochem. Biophys. Res. Commun. 261:464–471
Devary Y., Gottlieb R.A., Lau L.F., Karin M. (1991) Rapid and preferential activation of the c-jun gene during the mammalian UV response. Mol. Cell Biol. 11:2804–2811
Angel P., Karin M. (1991) The role of Jun, Fos and the AP-1 complex in cell proliferation and transformation. Biochim. Biophys. Acta 1072:129–157
Cooper S.J., MacGowan J., Ranger-Moore J., Young M.R., Colburn N.H., Bowden G.T. (2003) Expression of dominant negative c-jun inhibits ultraviolet B-induced squamous cell carcinoma number and size in an SKH-1 hairless mouse model. Mol. Cancer Res. 1:848–854
Karin M. (1995) The regulation of AP-1 activity by mitogen-activated protein kinases. J. Biol. Chem. 270:16483–16486
Whitmarsh A.J., Davis R.J. (1996) Transcription factor AP-1 regulation by mitogen-activated protein kinase signal transduction pathways. J. Mol. Med. 74:589–607
Price M.A., Cruzalegui F.H., Treisman R. (1996) The p38 and ERK MAP kinase pathways cooperate to activate Ternary Complex Factors and c-fos transcription in response to UV light. EMBO J. 15:6552–6563
Chen W., Bowden G.T. (1999) Activation of p38 MAP kinase and ERK are required for ultraviolet-B induced c-fos gene expression in human keratinocytes. Oncogene 18:7469–7476
Silvers A.L., Bachelor M.A., Bowden G.T. (2003) The role of JNK and p38 MAPK activities in UVA-induced signaling pathways leading to AP-1 activation and c-Fos expression. Neoplasia 5:319–329
Tanos T., Marinissen M.J., Leskow F.C., Hochbaum D., Martinetto H., Gutkind J.S., Coso O.A. (2005) Phosphorylation of c-Fos by members of the p38 MAPK family. Role in the AP-1 response to UV light. J. Biol. Chem. 280:18842–18852
Chen W., Bowden G.T. (2000) Role of p38 mitogen-activated protein kinases in ultraviolet-B irradiation-induced activator protein 1 activation in human keratinocytes. Mol. Carcinog. 28:196–202
Bodero A.J., Ye R., Lees-Miller S.P. (2003) UV-light induces p38 MAPK-dependent phosphorylation of Bcl10. Biochem. Biophys. Res. Commun. 301:923–926
Kovarik P., Stoiber D., Eyers P.A., Menghini R., Neininger A., Gaestel M., Cohen P., Decker T. (1999) Stress-induced phosphorylation of STAT1 at Ser727 requires p38 mitogen-activated protein kinase whereas IFN-gamma uses a different signaling pathway. Proc. Natl. Acad. Sci. USA 96:13956–13961
Kovarik P., Mangold M., Ramsauer K., Heidari H., Steinborn R., Zotter A., Levy D.E., Muller M., Decker T. (2001) Specificity of signaling by STAT1 depends on SH2 and C-terminal domains that regulate Ser727 phosphorylation, differentially affecting specific target gene expression. EMBO J. 20:91–100
Nurse P. (1997) Checkpoint pathways come of age. Cell 91:865–867
Bartek J., Lukas J. (2001) Pathways governing G1/S transition and their response to DNA damage. FEBS Lett. 490:117–122
Gottlieb T.M., Oren M. (1996) p53 in growth control and neoplasia. Biochim. Biophys. Acta 1287:77–102
Zhan Q., Antinore M.J., Wang X.W. (1999) Association with Cdc2 and inhibition of Cdc2/Cyclin B1 kinase activity by the p53-regulated protein Gadd45. Oncogene 18:2892–2900
Bulavin D.V., Higashimoto Y., Popoff I.J., Gaarde W.A., Basrur V., Potapova O., Appella E., Fornace A.J. Jr. (2001) Initiation of a G2/M checkpoint after UV radiation requires p38 kinase. Nature (Lond.) 411:102–107
Norbury C., Blow J., Nurse P. (1991) Regulatory phosphorylation of the p34cdc2 protein kinase in vertebrates. EMBO J. 10:3321–3329
Parker L.L., Piwnica-Worms H. (1992) Inactivation of the p34cdc2-cyclin B complex by the human WEE1 tyrosine kinase. Science 257:1955–1957
Mueller P., Coleman T., Kumagai A., Dunphy W. (1995) Myt1: a membrane-associated inhibitory kinase that phosphorylates Cdc2 on both threonine-14 and tyrosine-15. Science 270:86–90
Liu F., Stanton J.J., Wu Z., Piwnica-Worms H. (1997) The human Myt1 kinase preferentially phosphorylates Cdc2 on threonine 14 and localizes to the endoplasmic reticulum and Golgi complex. Mol. Cell. Biol. 17:571–583
Gautier J., Solomon M.J., Booher R.N., Bazan J.F., Kirschner M.W. (1991) cdc25 is a specific tyrosine phosphatase that directly activates p34cdc2. Cell 67:197–211
Kumagai A., Dunphy W. (1991) The cdc25 protein controls tyrosine dephosphorylation in a cell-free system. Cell 64:903–914
Strausfeld U., Labbe J.C., Fesquet D., Cavadore J.C., Picard A., Sadhu K., Russell P., Doree M. (1991) Dephosphorylation and activation of a p34cdc2/cyclin B complex in␣vitro by human CDC25 protein. Nature 351:242–245
Galaktionov K., Beach D. (1991) Specific activation of cdc25 tyrosine phosphatases by B-type cyclins: evidence for multiple roles of mitotic cyclins. Cell 67:1181–1194
Nagata A., Igarashi M., Jinno S., Suto K., Okayama H. (1991) An additional homolog of the fission yeast cdc25+ gene occurs in humans and is highly expressed in some cancer cells. New Biol. 3:959–968
Sadhu K., Reed S.I., Richardson H., Russell P. (1990) Human homolog of fission yeast cdc25 mitotic inducer is predominantly expressed in G2. Proc. Natl. Acad. Sci. USA 87:5139–5143
Donzelli M., Draetta G.F. (2003) Regulating mammalian checkpoints through Cdc25 inactivation. EMBO Rep. 4:671–677
Davezac N., Baldin V., Gabrielli B., Forrest A., Theis-Febvre N., Yashida M., Ducommun B. (2000) Regulation of CDC25B phosphatases subcellular localization. Oncogene 19:2179–2185
Forrest A., Gabrielli B. (2001) Cdc25B activity is regulated by 14-3-3. Oncogene 20:4393–4401
Lindqvist A., Kallstrom H., Karlsson Rosenthal C. (2004) Characterisation of Cdc25B localisation and nuclear export during the cell cycle and in response to stress. J. Cell Sci. 117:4979–4990
Manke I.A., Nguyen A., Lim D., Stewart M.Q., Elia A.E., Yaffe M.B. (2005) MAPKAP kinase-2 is a cell cycle checkpoint kinase that regulates the G2/M transition and S phase progression in response to UV irradiation. Mol. Cell 17:37–48
Kulms D., Schwarz T. (2002) Molecular mechanisms involved in UV-induced apoptotic cell death. Skin Pharmacol. Appl. Skin Physiol. 15:342–347
Kim D.S., Hwang E.S., Lee J.E., Kim S.Y., Park K.C. (2003) Sphingosine-1-phosphate promotes mouse melanocyte survival via ERK and Akt activation. Cell. Signal. 15:919–926
Shimizu H., Banno Y., Sumi N., Naganawa T., Kitajima Y., Nozawa Y. (1999) Activation of p38 mitogen-activated protein kinase and caspases in UVB-induced apoptosis of human keratinocyte HaCaT cells. J Invest. Dermatol. 112:769–774
Assefa Z., Vantieghem A., Garmyn M., Declercq W., Vandenabeele P., Vandenheede J.R., Bouillon R., Merlevede W., Agostinis P. (2000) p38 mitogen-activated protein kinase regulates a novel, caspase-independent pathway for the mitochondrial cytochrome c release in ultraviolet B radiation-induced apoptosis. J. Biol. Chem. 275:21416–21421
Van Laethem A., Van Kelst S., Lippens S., Declercq W., Vandenabeele P., Janssens S., Vandenheede J.R., Garmyn M., Agostinis P. (2004) Activation of p38 MAPK is required for Bax translocation to mitochondria, cytochrome c release and apoptosis induced by UVB irradiation in human keratinocytes. FASEB J. 18:1946–1948
Chen H.Y., Zhu L., Zhan S.M., Han Z.W., Du W., Wang Y.J., Cui R.Y., Wang C.B. (2005) Polypeptide from Chlamys farreri inhibits murine thymocytes apoptosis and modulates UVB induced signaling pathway activation. Life Sci. 77:768–779
Hildesheim J., Awwad R.T., Fornace A.J. Jr. (2004) p38 mitogen-activated protein kinase inhibitor protects the epidermis against the acute damaging effects of ultraviolet irradiation by blocking apoptosis and inflammatory responses. J. Invest. Dermatol. 122:497–502
Papoutsaki M., Moretti F., Lanza M., Marinari B., Sartorelli V., Guerrini L., Chimenti S., Levrero M., Costanzo A. (2005) A p38-dependent pathway regulates DeltaNp63 DNA binding to p53-dependent promoters in UV-induced apoptosis of keratinocytes. Oncogene 24:6970–6975
Chouinard N., Valerie K., Rouabhia M., Huot J. (2002) UVB-mediated activation of p38 mitogen-activated protein kinase enhances resistance of normal human keratinocytes to apoptosis by stabilizing cytoplasmic p53. Biochem. J. 365:133–145
Ivanov V.N., Ronai Z. (2000) p38 protects human melanoma cells from UV-induced apoptosis through down-regulation of NF-kappaB activity and Fas expression. Oncogene 19:3003–3012
Nemoto S., Xiang J., Huang S., Lin A. (1998) Induction of apoptosis by SB202190 through inhibition of p38β mitogen-activated protein kinase. J. Biol. Chem. 273:16415–16420
Bachelor M.A., Bowden G.T. (2004) Ultraviolet A-induced modulation of bcl-xl by p38 MAPK in human keratinocytes: post-translational regulation through the 3’untranslated region. J. Biol. Chem. 279:42658–42668
Konnikov N., Pincus S.H., Dinarello C.A. (1989) Elevated plasma interleukin-1 levels in humans following ultraviolet light therapy for psoriasis. J. Invest. Dermatol. 92:235–239
Kock A., Schwarz T., Kirnbauer R., Urbanski A., Perry Ansel J.C., Luger T.A. (1990) Human keratinocytes are a source for tumor necrosis factor alpha: evidence for synthesis and release upon stimulation with endotoxin or ultraviolet light. J. Exp. Med. 172:1609–1614
Kirnbauer R., Kock A., Neuner P., Forster E., Krutmann J., Urbanski A., Schauer E., Ansel J.C., Schwarz T., Luger T.A. (1991) Regulation of epidermal cell interleukin-6 production by UV light and corticosteroids. J. Invest. Dermatol. 96:484–489
Kondo S., Kono T., Sauder D.N., McKenzie R. (1993) IL-8 gene expression and production in human keratinocytes and their modulation by UVB. J. Invest. Dermatol. 101:690–694
Kondo S. (1999) The roles of keratinocyte-derived cytokines in the epidermis and their possible responses to UVA-irradiation. J. Invest. Dermatol. Symp. Proc. 4:177–183
Allen M., Svensson L., Roach M., Hambor J., McNeish J., Gabel C.A. (2000) Deficiency of the stress kinase p38alpha results in embryonic lethality: characterization of the kinase dependence of stress responses of enzyme-deficient embryonic stem cells. J. Exp. Med. 191:859–870
Kim A.L., Labasi J.M., Zhu Y., Tang X., McClure K., Gabel C.A., Athar M., Bickers D.R. (2005) Role of p38 MAPK in UVB-induced inflammatory responses in the skin of SKH-1 hairless mice. J. Invest. Dermatol. 124:1318–1325
Vane J.R., Bakhle Y.S., Botting R.M. (1998) Cyclooxygenases 1 and 2. Annu. Rev. Pharmacol. Toxicol. 38:97–120
Chen W., Tang Q., Gonzales M.S., Bowden G.T. (2001) Role of p38 MAP kinases and ERK in mediating ultraviolet-B induced cyclooxygenase-2 gene expression in human keratinocytes. Oncogene 20:3921–3926
Buckman S.Y., Gresham A., Hale P., Hruza G., Anast J., Masferrer J., Pentland A.P. (1998) COX-2 expression is induced by UVB exposure in human skin: implications for the development of skin cancer. Carcinogenesis 19:723–729
Wilgus T.A., Koki A.T., Zweifel B.S., Kusewitt D.F., Rubal P.A., Oberyszyn T.M. (2003) Inhibition of cutaneous ultraviolet light B-mediated inflammation and tumor formation with topical celecoxib treatment. Mol. Carcinog. 38:49–58
Wilgus T.A., Parrett M.L., Ross M.S., Tober K.L., Robertson F.M., Oberyszyn T.M. (2002) Inhibition of ultraviolet light B-induced cutaneous inflammation by a specific cyclooxygenase-2 inhibitor. Adv. Exp. Med. Biol. 507:85–92
Bachelor M.A., Silvers A.L., Bowden G.T. (2002) The role of p38 in UVA induced cyclooxygenase-2 expression in the human keratinocyte cell line, HaCaT. Oncogene 21:7092–7099
Galibert M.D., Carreira S., Goding C.R. (2001) The Usf-1 transcription factor is a novel target for the stress-responsive p38 kinase and mediates UV-induced Tyrosinase expression. EMBO J. 20:5022–5031
Corre S., Primot A., Sviderskaya E., Bennett D.C., Vaulont S., Goding C.R., Galibert M.D. (2004) UV-induced expression of key component of the tanning process, the POMC and MC1R genes, is dependent on the p38-activated upstream stimulating factor-1 (USF-1) J. Biol. Chem. 279:51226–51233
Abdel-Malek Z., Swope V.B., Suzuki I., Akcali C., Harriger M.D., Boyce S.T., Urabe K., Hearing V.J. (1995) Mitogenic and melanogenic stimulation of normal human melanocytes by melanotropic peptides. Proc. Natl. Acad. Sci. 92:1789–1793
Acknowledgements
The work presented here was funded by the National Science Natural foundation of China (No. 30471458) and Science Natural Foundation of Shandong province (No. Y2003c02).
Author information
Authors and Affiliations
Corresponding author
Rights and permissions
About this article
Cite this article
Jinlian, L., Yingbin, Z. & Chunbo, W. p38 MAPK in regulating cellular responses to ultraviolet radiation. J Biomed Sci 14, 303–312 (2007). https://doi.org/10.1007/s11373-007-9148-4
Received:
Accepted:
Published:
Issue Date:
DOI: https://doi.org/10.1007/s11373-007-9148-4