Regulation of cytokines by small RNAs during skin inflammation

Journal of Biomedical Science

Volumn 17, Issue 1, 2010, Pages

  Bak, Rasmus O ^a   Mikkelsen, Jacob G ^a  

^a AARHUS UNIVERSITY   (Denmark)

Author keywords

[No Author keywords available]

Indexed keywords

ADALIMUMAB; ALEFACEPT; CORTICOSTEROID; CYCLOSPORIN A; CYTOKINE; ETANERCEPT; GRANULOCYTE MACROPHAGE COLONY STIMULATING FACTOR; INFLIXIMAB; INTERLEUKIN 17; INTERLEUKIN 22; INTERLEUKIN 23; INTERLEUKIN 4; INTERLEUKIN 6; KERATOLYTIC AGENT; MESSENGER RNA; METHOTREXATE; MICRORNA; MITOGEN ACTIVATED PROTEIN KINASE P38; PSORALEN; RETINOID DERIVATIVE; SHORT HAIRPIN RNA; SMALL INTERFERING RNA; SUPPRESSOR OF CYTOKINE SIGNALING 1; TRISTETRAPROLIN; TUMOR NECROSIS FACTOR ALPHA; TUMOR NECROSIS FACTOR ALPHA INHIBITOR; UNINDEXED DRUG; UNTRANSLATED RNA; USTEKINUMAB; VITAMIN D DERIVATIVE;

CLINICAL TRIAL; CONGENITAL PACHYONYCHIA; CROHN DISEASE; CYTOKINE PRODUCTION; GENE CONTROL; GENE EXPRESSION; HUMAN; KERATINOCYTE; NONHUMAN; PHOTOTHERAPY; PRIORITY JOURNAL; PSORIASIS; REVIEW; RHEUMATOID ARTHRITIS; RNA DEGRADATION; RNA INTERFERENCE; SIGNAL TRANSDUCTION; SKIN; SKIN INFLAMMATION; TRANSCRIPTION REGULATION; ULCERATIVE COLITIS; GENE EXPRESSION REGULATION; IMMUNOLOGY; IMMUNOTHERAPY; INNATE IMMUNITY; METABOLISM; METHODOLOGY;

CYTOKINES; GENE EXPRESSION REGULATION; HUMANS; IMMUNITY, INNATE; IMMUNOTHERAPY; MICRORNAS; PSORIASIS; RNA INTERFERENCE; RNA, MESSENGER; SIGNAL TRANSDUCTION;

EID: 77954054512     PISSN: 10217770     EISSN: 14230127     Source Type: Journal    
DOI: 10.1186/1423-0127-17-53     Document Type: Review

References (162)

1. Pathogenesis and clinical features of psoriasis. CE Griffiths JN Barker, Lancet 2007 370 263 271 10.1016/S0140-6736(07)61128-3 17658397
2. The prevalence of previously diagnosed and undiagnosed psoriasis in US adults: results from NHANES 2003-2004. SK Kurd JM Gelfand, Journal of the American Academy of Dermatology 2009 60 218 224 10.1016/j.jaad.2008.09.022 19022533
3. Psoriasis. MP Schon WH Boehncke, N Engl J Med 2005 352 1899 1912 10.1056/NEJMra041320 15872205
4. Psoriasis. C Sand NK Veien, Book Psoriasis (Editor ed. eds.), vol. LEO TEMABOG City: Lep Pharma 2005
5. The psychosocial and occupational impact of chronic skin disease. J Hong B Koo J Koo, Dermatol Ther 2008 21 54 59 10.1111/j.1529-8019.2008.00170.x 18318886
6. Quality of life in 6497 Nordic patients with psoriasis. R Zachariae H Zachariae K Blomqvist S Davidsson L Molin C Mork B Sigurgeirsson, The British journal of dermatology 2002 146 1006 1016 10.1046/j.1365-2133.2002.04742.x 12072069
7. Psoriasis. FO Nestle DH Kaplan J Barker, N Engl J Med 2009 361 496 509 10.1056/NEJMra0804595 19641206
8. Clearance of severe psoriasis after allogenic bone marrow transplantation. DJ Eedy D Burrows JM Bridges FG Jones, BMJ 1990 300 908 10.1136/bmj.300.6729.908 2337714
9. Psoriasis after allogeneic bone marrow transplantation. M Gardembas-Pain N Ifrah C Foussard M Boasson JP Saint Andre JL Verret, Arch Dermatol 1990 126 1523 10.1001/archderm.126.11.1523 2241215
10. Novel mechanisms of T-cell and dendritic cell activation revealed by profiling of psoriasis on the 63,100-element oligonucleotide array. X Zhou JG Krueger MC Kao E Lee F Du A Menter WH Wong AM Bowcock, Physiol Genomics 2003 13 69 78 12644634
11. Th22 cells represent a distinct human T cell subset involved in epidermal immunity and remodeling. S Eyerich K Eyerich D Pennino T Carbone F Nasorri S Pallotta F Cianfarani T Odorisio C Traidl-Hoffmann H Behrendt SR Durham CB Schmidt-Weber A Cavani, J Clin Invest 2009 119 3573 3585 19920355
12. Regulatory T cells and autoimmune disease. S Paust H Cantor, Immunol Rev 2005 204 195 207 10.1111/j.0105-2896.2005.00247.x 15790360
13. Psoriasis vulgaris lesions contain discrete populations of Th1 and Th17 T cells. MA Lowes T Kikuchi J Fuentes-Duculan I Cardinale LC Zaba AS Haider EP Bowman JG Krueger, J Invest Dermatol 2008 128 1207 1211 10.1038/sj.jid.5701213 18200064
14. Interleukin-4 therapy of psoriasis induces Th2 responses and improves human autoimmune disease. K Ghoreschi P Thomas S Breit M Dugas R Mailhammer W van Eden R van der Zee T Biedermann J Prinz M Mack U Mrowietz E Christophers D Schlöndorff G Plewig CA Sander M Röcken, Nat Med 2003 9 40 46 10.1038/nm804 12461524
15. Expanding the effector CD4 T-cell repertoire: the Th17 lineage. LE Harrington PR Mangan CT Weaver, Curr Opin Immunol 2006 18 349 356 10.1016/j.coi.2006.03.017 16616472
16. Immunology News: A New Type of T Cell Is Discovered. C Helliwell, ACNR 2007 6
17. Circulating Th17, Th22, and Th1 cells are increased in psoriasis. S Kagami HL Rizzo JJ Lee Y Koguchi A Blauvelt, The Journal of investigative dermatology 2010 130 1373 1383 10.1038/jid.2009.399 20032993
18. Human Langerhans cells induce distinct IL-22-producing CD4+ T cells lacking IL-17 production. H Fujita KE Nograles T Kikuchi J Gonzalez JA Carucci JG Krueger, Proceedings of the National Academy of Sciences of the United States of America 2009 106 21795 21800 10.1073/pnas.0911472106 19996179
19. Mechanisms underlying lineage commitment and plasticity of helper CD4+ T cells. JJ O'Shea WE Paul, Science 327 1098 1102 10.1126/science.1178334 20185720
20. Immunopathogenesis of psoriasis. R Sabat S Philipp C Hoflich S Kreutzer E Wallace K Asadullah HD Volk W Sterry K Wolk, Exp Dermatol 2007 16 779 798 10.1111/j.1600-0625.2007.00629.x 17845210
21. The immunologic basis for the treatment of psoriasis with new biologic agents. JG Krueger, J Am Acad Dermatol 2002 46 1 23 quiz 23-26 10.1067/mjd.2002.120568 11756941
22. Innate immunity and antimicrobial defense systems in psoriasis. AS Buchau RL Gallo, Clin Dermatol 2007 25 616 624 10.1016/j.clindermatol.2007.08.016 18021900
23. Topical preparations for the treatment of psoriasis: results of a retrospective study over 15 years. A Zampetti A Barone D Antuzzi P Amerio A Tulli C Feliciani, J Dermatolog Treat 2008 19 134 140 10.1080/09546630701846087 18569269
24. Biological therapies in the systemic management of psoriasis: International Consensus Conference. W Sterry J Barker WH Boehncke JD Bos S Chimenti E Christophers M De La Brassinne C Ferrandiz C Griffiths A Katsambas K Kragballe C Lynde A Menter JP Ortonne K Papp J Prinz B Rzany J Ronnevig JH Saurat M Stahle FM Stengel P Van De Kerkhof J Voorhees, Br J Dermatol 2004 151 Suppl 69 3 17 10.1111/j.1365-2133.2004.06070.x 15265063
25. Biologic therapy for psoriasis: an update on the tumor necrosis factor inhibitors infliximab, etanercept, and adalimumab, and the T-cell-targeted therapies efalizumab and alefacept. JM Weinberg CJ Bottino J Lindholm R Buchholz, J Drugs Dermatol 2005 4 544 555 16167412
26. Review and expert opinion on prevention and treatment of infliximab-related infusion reactions. LL Lecluse G Piskin JR Mekkes JD Bos MA de Rie, Br J Dermatol 2008 159 527 536 10.1111/j.1365-2133.2008.08728.x 18627374
27. Management of severe psoriasis with TNF antagonists. Adalimumab, etanercept and infliximab. R Mossner K Reich, Curr Probl Dermatol 2009 38 107 136 full-text 19710553
28. Management of moderate to severe plaque psoriasis (part I): clinical update on antitumor necrosis factor agents. JM Sobell RE Kalb JM Weinberg, J Drugs Dermatol 2009 8 147 154 19213230
29. Increased expression of IL-12p70 and IL-23 by multiple dendritic cell and macrophage subsets in plaque psoriasis. N Yawalkar GG Tscharner RE Hunger AS Hassan, J Dermatol Sci 2009 54 99 105 10.1016/j.jdermsci.2009.01.003 19264456
30. Differential regulation of interleukin 12 and interleukin 23 production in human dendritic cells. F Gerosa B Baldani-Guerra LA Lyakh G Batoni S Esin RT Winkler-Pickett MR Consolaro M De Marchi D Giachino A Robbiano M Astegiano A Sambataro RA Kastelein G Carra G Trinchieri, J Exp Med 2008 205 1447 1461 10.1084/jem.20071450 18490488
31. In vitro and in situ expression of IL-23 by keratinocytes in healthy skin and psoriasis lesions: enhanced expression in psoriatic skin. G Piskin RM Sylva-Steenland JD Bos MB Teunissen, J Immunol 2006 176 1908 1915 16424222
32. Interleukin-12 and the regulation of innate resistance and adaptive immunity. G Trinchieri, Nat Rev Immunol 2003 3 133 146 10.1038/nri1001 12563297
33. IL-12 and IL-23: master regulators of innate and adaptive immunity. CL Langrish BS McKenzie NJ Wilson R de Waal Malefyt RA Kastelein DJ Cua, Immunol Rev 2004 202 96 105 10.1111/j.0105-2896.2004.00214.x 15546388
34. Interleukin-17 and interferon-gamma synergize in the enhancement of proinflammatory cytokine production by human keratinocytes. MB Teunissen CW Koomen R de Waal Malefyt EA Wierenga JD Bos, J Invest Dermatol 1998 111 645 649 10.1046/j.1523-1747.1998.00347.x 9764847
35. A large-scale genetic association study confirms IL12B and leads to the identification of IL23R as psoriasis-risk genes. M Cargill SJ Schrodi M Chang VE Garcia R Brandon KP Callis N Matsunami KG Ardlie D Civello JJ Catanese DU Leong JM Panko LB McAllister CB Hansen J Papenfuss SM Prescott TJ White MF Leppert GG Krueger AB Begovich, Am J Hum Genet 2007 80 273 290 10.1086/511051 17236132
36. Clinical improvement in chronic plaque-type psoriasis lesions after narrow-band UVB therapy is accompanied by a decrease in the expression of IFN-gamma inducers - IL-12, IL-18 and IL-23. G Piskin U Tursen RM Sylva-Steenland JD Bos MB Teunissen, Exp Dermatol 2004 13 764 772 10.1111/j.0906-6705.2004.00246.x 15560760
37. TNF inhibition rapidly down-regulates multiple proinflammatory pathways in psoriasis plaques. AB Gottlieb F Chamian S Masud I Cardinale MV Abello MA Lowes F Chen M Magliocco JG Krueger, J Immunol 2005 175 2721 2729 16081850
38. Alefacept reduces infiltrating T cells, activated dendritic cells, and inflammatory genes in psoriasis vulgaris. F Chamian MA Lowes SL Lin E Lee T Kikuchi P Gilleaudeau M Sullivan-Whalen I Cardinale A Khatcherian I Novitskaya KM Wittkowski JG Krueger, Proc Natl Acad Sci USA 2005 102 2075 2080 10.1073/pnas.0409569102 15671179
39. Trial watch: novel biologic for psoriasis shows superiority over current best-seller. Nat Rev Drug Discov 2008 7 880 881
40. Interleukin-20 plays a critical role in maintenance and development of psoriasis in the human xenograft transplantation model. K Stenderup C Rosada A Worsaae F Dagnaes-Hansen T Steiniche E Hasselager LF Iversen S Zahn H Wöldike HL Holmberg J Rømer K Kragballe JT Clausen TN Dam, Br J Dermatol 2009 160 284 296 10.1111/j.1365-2133.2008.08890.x 18945296
41. Transcriptional control of the inflammatory response. R Medzhitov T Horng, Nat Rev Immunol 2009 9 692 703 10.1038/nri2634 19859064
42. Cytokine signaling modules in inflammatory responses. JJ O'Shea PJ Murray, Immunity 2008 28 477 487 10.1016/j.immuni.2008.03.002 18400190
43. ERK and p38 MAPK-activated protein kinases: a family of protein kinases with diverse biological functions. PP Roux J Blenis, Microbiol Mol Biol Rev 2004 68 320 344 10.1128/MMBR.68.2.320-344.2004 15187187
44. ARED 2.0: an update of AU-rich element mRNA database. T Bakheet BR Williams KS Khabar, Nucleic Acids Res 2003 31 421 423 10.1093/nar/gkg023 12520039
45. MicroRNA targets in immune genes and the Dicer/Argonaute and ARE machinery components. AJ Asirvatham CJ Gregorie Z Hu WJ Magner TB Tomasi, Mol Immunol 2008 45 1995 2006 10.1016/j.molimm.2007.10.035 18061676
46. AU-rich elements: characterization and importance in mRNA degradation. CY Chen AB Shyu, Trends Biochem Sci 1995 20 465 470 10.1016/S0968-0004(00)89102-1 8578590
47. The role of the AU-rich elements of mRNAs in controlling translation. E Espel, Semin Cell Dev Biol 2005 16 59 67 10.1016/j.semcdb.2004.11.008 15659340
48. The exosome: a multipurpose RNA-decay machine. M Schmid TH Jensen, Trends Biochem Sci 2008 33 501 510 10.1016/j.tibs.2008.07.003 18786828
49. Human 5' - >3' exonuclease Xrn2 promotes transcription termination at co-transcriptional cleavage sites. S West N Gromak NJ Proudfoot, Nature 2004 432 522 525 10.1038/nature03035 15565158 (Pubitemid 39576535)
50. Mitogen-activated protein kinase-activated protein kinase 2 regulates tumor necrosis factor mRNA stability and translation mainly by altering tristetraprolin expression, stability, and binding to adenine/uridine-rich element. E Hitti T Iakovleva M Brook S Deppenmeier AD Gruber D Radzioch AR Clark PJ Blackshear A Kotlyarov M Gaestel, Mol Cell Biol 2006 26 2399 2407 10.1128/MCB.26.6.2399-2407.2006 16508014
51. Feedback inhibition of macrophage tumor necrosis factor-alpha production by tristetraprolin. E Carballo WS Lai PJ Blackshear, Science 1998 281 1001 1005 10.1126/science.281.5379.1001 9703499
52. Tristetraprolin and its family members can promote the cell-free deadenylation of AU-rich element-containing mRNAs by poly(A) ribonuclease. WS Lai EA Kennington PJ Blackshear, Mol Cell Biol 2003 23 3798 3812 10.1128/MCB.23.11.3798-3812.2003 12748283
53. TTP and BRF proteins nucleate processing body formation to silence mRNAs with AU-rich elements. TM Franks J Lykke-Andersen, Genes Dev 2007 21 719 735 10.1101/gad.1494707 17369404
54. Recruitment and activation of mRNA decay enzymes by two ARE-mediated decay activation domains in the proteins TTP and BRF-1. J Lykke-Andersen E Wagner, Genes Dev 2005 19 351 361 10.1101/gad.1282305 15687258
55. A pathogenetic role for TNF alpha in the syndrome of cachexia, arthritis, and autoimmunity resulting from tristetraprolin (TTP) deficiency. GA Taylor E Carballo DM Lee WS Lai MJ Thompson DD Patel DI Schenkman GS Gilkeson HE Broxmeyer BF Haynes PJ Blackshear, Immunity 1996 4 445 454 10.1016/S1074- 7613(00)80411-2 8630730
56. RNA stabilization by the AU-rich element binding protein, HuR, an ELAV protein. SS Peng CY Chen N Xu AB Shyu, EMBO J 1998 17 3461 3470 10.1093/emboj/17.12.3461 9628881
57. Nuclear export of NF90 is required for interleukin-2 mRNA stabilization. J Shim H Lim RY J M Karin, Mol Cell 2002 10 1331 1344 10.1016/S1097-2765(02) 00730-X 12504009
58. ELAV proteins stabilize deadenylated intermediates in a novel in vitro mRNA deadenylation/degradation system. LP Ford J Watson JD Keene J Wilusz, Genes Dev 1999 13 188 201 10.1101/gad.13.2.188 9925643
59. A KH domain RNA binding protein, KSRP, promotes ARE-directed mRNA turnover by recruiting the degradation machinery. R Gherzi KY Lee P Briata D Wegmuller C Moroni M Karin CY Chen, Mol Cell 2004 14 571 583 10.1016/j.molcel.2004.05.002 15175153
60. RNA trafficking and stabilization elements associate with multiple brain proteins. M Snee GJ Kidd TP Munro R Smith, J Cell Sci 2002 115 4661 4669 10.1242/jcs.00137 12415010
61. Bringing the role of mRNA decay in the control of gene expression into focus. CJ Wilusz J Wilusz, Trends Genet 2004 20 491 497 10.1016/j.tig.2004.07. 011 15363903
62. Mitogen-activated protein (MAP) kinase pathways: regulation and physiological functions. G Pearson F Robinson T Beers Gibson BE Xu M Karandikar K Berman MH Cobb, Endocr Rev 2001 22 153 183 10.1210/er.22.2.153 11294822
63. MAP kinase and pain. RR Ji RWt Gereau M Malcangio GR Strichartz, Brain Res Rev 2009 60 135 148 10.1016/j.brainresrev.2008.12.011 19150373
64. A protein kinase involved in the regulation of inflammatory cytokine biosynthesis. JC Lee JT Laydon PC McDonnell TF Gallagher S Kumar D Green D McNulty MJ Blumenthal JR Heys SW Landvatter,, et al. Nature 1994 372 739 746 10.1038/372739a0 7997261
65. Differential expression and activation of p38 mitogen-activated protein kinase alpha, beta, gamma, and delta in inflammatory cell lineages. KK Hale D Trollinger M Rihanek CL Manthey, J Immunol 1999 162 4246 4252 10201954
66. The p38 MAP kinase pathway and its biological function. L New J Han, Trends Cardiovasc Med 1998 8 220 228 10.1016/S1050-1738(98)00012-7 14987568
67. p38 MAP kinases: key signalling molecules as therapeutic targets for inflammatory diseases. S Kumar J Boehm JC Lee, Nat Rev Drug Discov 2003 2 717 726 10.1038/nrd1177 12951578
68. MAPKAP kinase-2; a novel protein kinase activated by mitogen-activated protein kinase. D Stokoe DG Campbell S Nakielny H Hidaka SJ Leevers C Marshall P Cohen, EMBO J 1992 11 3985 3994 1327754
69. MAPKAP kinase 2 is essential for LPS-induced TNF-alpha biosynthesis. A Kotlyarov A Neininger C Schubert R Eckert C Birchmeier HD Volk M Gaestel, Nat Cell Biol 1999 1 94 97 10.1038/10061 10559880
70. MK2 targets AU-rich elements and regulates biosynthesis of tumor necrosis factor and interleukin-6 independently at different post-transcriptional levels. A Neininger D Kontoyiannis A Kotlyarov R Winzen R Eckert HD Volk H Holtmann G Kollias M Gaestel, J Biol Chem 2002 277 3065 3068 10.1074/jbc.C100685200 11741878
71. The p38 MAPK pathway inhibits tristetraprolin-directed decay of interleukin-10 and pro-inflammatory mediator mRNAs in murine macrophages. C Tudor FP Marchese E Hitti A Aubareda L Rawlinson M Gaestel PJ Blackshear AR Clark J Saklatvala JL Dean, FEBS Lett 2009 583 1933 1938 10.1016/j.febslet.2009. 04.039 19416727
72. The p38 mitogen-activated kinase pathway regulates the human interleukin-10 promoter via the activation of Sp1 transcription factor in lipopolysaccharide-stimulated human macrophages. W Ma W Lim K Gee S Aucoin D Nandan M Kozlowski F Diaz-Mitoma A Kumar, J Biol Chem 2001 276 13664 13674 11278848
73. The kinases MSK1 and MSK2 act as negative regulators of Toll-like receptor signaling. O Ananieva J Darragh C Johansen JM Carr J McIlrath JM Park A Wingate CE Monk R Toth SG Santos L Iversen JS Arthur, Nat Immunol 2008 9 1028 1036 10.1038/ni.1644 18690222
74. Phosphorylation of HSF1 by MAPK-activated protein kinase 2 on serine 121, inhibits transcriptional activity and promotes HSP90 binding. X Wang MA Khaleque MJ Zhao R Zhong M Gaestel SK Calderwood, J Biol Chem 2006 281 782 791 10.1074/jbc.M505822200 16278218
75. Heat shock proteins and regulation of cytokine expression. Y Xie CM Cahill A Asea PE Auron SK Calderwood, Infect Dis Obstet Gynecol 1999 7 26 30 10.1002/(SICI)1098-0997(1999)7:1/2<26::AID-IDOG6>3.0.CO;2-Q 10231005
76. Phosphorylation of tristetraprolin, a potential zinc finger transcription factor, by mitogen stimulation in intact cells and by mitogen-activated protein kinase in vitro. GA Taylor MJ Thompson WS Lai PJ Blackshear, J Biol Chem 1995 270 13341 13347 10.1074/jbc.270.22.13341 7768935
77. Mitogen-activated protein kinase p38 controls the expression and posttranslational modification of tristetraprolin, a regulator of tumor necrosis factor alpha mRNA stability. KR Mahtani M Brook JL Dean G Sully J Saklatvala AR Clark, Mol Cell Biol 2001 21 6461 6469 10.1128/MCB.21.9.6461-6469.2001 11533235
78. MAPKAP kinase 2 phosphorylates tristetraprolin on in vivo sites including Ser178, a site required for 14-3-3 binding. CA Chrestensen MJ Schroeder J Shabanowitz DF Hunt JW Pelo MT Worthington TW Sturgill, J Biol Chem 2004 279 10176 10184 10.1074/jbc.M310486200 14688255
79. MK2-induced tristetraprolin: 14-3-3 complexes prevent stress granule association and ARE-mRNA decay. G Stoecklin T Stubbs N Kedersha S Wax WF Rigby TK Blackwell P Anderson, EMBO J 2004 23 1313 1324 10.1038/sj.emboj.7600163 15014438
80. A role for mitogen-activated protein kinase activation by integrins in the pathogenesis of psoriasis. I Haase RM Hobbs MR Romero S Broad FM Watt, J Clin Invest 2001 108 527 536 11518726
81. Suprabasal integrin expression in the epidermis of transgenic mice results in developmental defects and a phenotype resembling psoriasis. JM Carroll MR Romero FM Watt, Cell 1995 83 957 968 10.1016/0092-8674(95)90211-2 8521519
82. The mitogen-activated protein kinases p38 and ERK1/2 are increased in lesional psoriatic skin. C Johansen K Kragballe M Westergaard J Henningsen K Kristiansen L Iversen, Br J Dermatol 2005 152 37 42 10.1111/j.1365-2133.2004. 06304.x 15656798
83. Protein expression of TNF-alpha in psoriatic skin is regulated at a posttranscriptional level by MAPK-activated protein kinase 2. C Johansen AT Funding K Otkjaer K Kragballe UB Jensen M Madsen L Binderup T Skak-Nielsen MS Fjording L Iversen, J Immunol 2006 176 1431 1438 16424170
84. p38(MAPK): stress responses from molecular mechanisms to therapeutics. LR Coulthard DE White DL Jones MF McDermott SA Burchill, Trends Mol Med 2009 15 369 379 10.1016/j.molmed.2009.06.005 19665431
85. The selectivity of protein kinase inhibitors: a further update. J Bain L Plater M Elliott N Shpiro CJ Hastie H McLauchlan I Klevernic JS Arthur DR Alessi P Cohen, Biochem J 2007 408 297 315 10.1042/BJ20070797 17850214
86. Inhibition of HSP27 phosphorylation by a cell-permeant MAPKAP Kinase 2 inhibitor. LB Lopes C Flynn P Komalavilas A Panitch CM Brophy BL Seal, Biochem Biophys Res Commun 2009 382 535 539 10.1016/j.bbrc.2009.03.056 19289101
87. Protein kinases as small molecule inhibitor targets in inflammation. M Gaestel A Mengel U Bothe K Asadullah, Curr Med Chem 2007 14 2214 2234 10.2174/092986707781696636 17896971
88. Pyrrolo-pyrimidones: a novel class of MK2 inhibitors with potent cellular activity. A Schlapbach R Feifel S Hawtin R Heng G Koch H Moebitz L Revesz C Scheufler J Velcicky R Waelchli C Huppertz, Bioorg Med Chem Lett 2008 18 6142 6146 10.1016/j.bmcl.2008.10.039 18945615
89. Pyrrolopyridine inhibitors of mitogen-activated protein kinase-activated protein kinase 2 (MK-2). DR Anderson MJ Meyers WF Vernier MW Mahoney RG Kurumbail N Caspers GI Poda JF Schindler DB Reitz RJ Mourey, J Med Chem 2007 50 2647 2654 10.1021/jm0611004 17480064
90. Potent and specific genetic interference by double-stranded RNA in Caenorhabditis elegans. A Fire S Xu MK Montgomery SA Kostas SE Driver CC Mello, Nature 1998 391 806 811 10.1038/35888 9486653 (Pubitemid 28099681)
91. Characterization of DGCR8/Pasha, the essential cofactor for Drosha in primary miRNA processing. KH Yeom Y Lee J Han MR Suh VN Kim, Nucleic Acids Res 2006 34 4622 4629 10.1093/nar/gkl458 16963499
92. The nuclear RNase III Drosha initiates microRNA processing. Y Lee C Ahn J Han H Choi J Kim J Yim J Lee P Provost O Radmark S Kim VN Kim, Nature 2003 425 415 419 10.1038/nature01957 14508493
93. Nuclear export of microRNA precursors. E Lund S Guttinger A Calado JE Dahlberg U Kutay, Science 2004 303 95 98 10.1126/science.1090599 14631048
94. MicroRNA maturation: stepwise processing and subcellular localization. Y Lee K Jeon JT Lee S Kim VN Kim, EMBO J 2002 21 4663 4670 10.1093/emboj/cdf476 12198168
95. Running interference: prospects and obstacles to using small interfering RNAs as small molecule drugs. DM Dykxhoorn J Lieberman, Annu Rev Biomed Eng 2006 8 377 402 10.1146/annurev.bioeng.8.061505.095848 16834561
96. Distinct passenger strand and mRNA cleavage activities of human Argonaute proteins. B Wang S Li HH Qi D Chowdhury Y Shi CD Novina, Nat Struct Mol Biol 2009 16 1259 1266 10.1038/nsmb.1712 19946268
97. siRNAs can function as miRNAs. JG Doench CP Petersen PA Sharp, Genes Dev 2003 17 438 442 10.1101/gad.1064703 12600936
98. Revisiting the principles of microRNA target recognition and mode of action. P Brodersen O Voinnet, Nat Rev Mol Cell Biol 2009 10 141 148 10.1038/nrm2619 19145236
99. RNAi and the P-body connection. JJ Rossi, Nat Cell Biol 2005 7 643 644 10.1038/ncb0705-643 15990892
100. Biological basis for restriction of microRNA targets to the 3' untranslated region in mammalian mRNAs. S Gu L Jin F Zhang P Sarnow MA Kay, Nat Struct Mol Biol 2009 16 144 150 10.1038/nsmb.1552 19182800
101. Conserved seed pairing, often flanked by adenosines, indicates that thousands of human genes are microRNA targets. BP Lewis CB Burge DP Bartel, Cell 2005 120 15 20 10.1016/j.cell.2004.12.035 15652477
102. MicroRNAs: target recognition and regulatory functions. DP Bartel, Cell 2009 136 215 233 10.1016/j.cell.2009.01.002 19167326
103. The widespread impact of mammalian MicroRNAs on mRNA repression and evolution. KK Farh A Grimson C Jan BP Lewis WK Johnston LP Lim CB Burge DP Bartel, Science 2005 310 1817 1821 10.1126/science.1121158 16308420
104. Principles of microRNA-target recognition. J Brennecke A Stark RB Russell SM Cohen, PLoS Biol 2005 3 85 10.1371/journal.pbio.0030085 15723116
105. Computational challenges in miRNA target predictions: to be or not to be a true target? C Barbato I Arisi ME Frizzo R Brandi L Da Sacco A Masotti, J Biomed Biotechnol 2009 2009 803069 19551154
106. MicroRNA in the immune system, microRNA as an immune system. LF Lu A Liston, Immunology 2009 127 291 298 10.1111/j.1365-2567.2009.03092.x 19538248
107. Aberrant T cell differentiation in the absence of Dicer. SA Muljo KM Ansel C Kanellopoulou DM Livingston A Rao K Rajewsky, J Exp Med 2005 202 261 269 10.1084/jem.20050678 16009718
108. Requirement of bic/microRNA-155 for normal immune function. A Rodriguez E Vigorito S Clare MV Warren P Couttet DR Soond S van Dongen RJ Grocock PP Das EA Miska D Vetrie K Okkenhaug AJ Enright G Dougan M Turner A Bradley, Science 2007 316 608 611 10.1126/science.1139253 17463290
109. Foxp3-dependent microRNA155 confers competitive fitness to regulatory T cells by targeting SOCS1 protein. LF Lu TH Thai DP Calado A Chaudhry M Kubo K Tanaka GB Loeb H Lee A Yoshimura K Rajewsky AY Rudensky, Immunity 2009 30 80 91 10.1016/j.immuni.2008.11.010 19144316
110. SOCS1 restricts dendritic cells' ability to break self tolerance and induce antitumor immunity by regulating IL-12 production and signaling. K Evel-Kabler XT Song M Aldrich XF Huang SY Chen, J Clin Invest 2006 116 90 100 10.1172/JCI26169 16357940
111. Altered expression of MicroRNA in synovial fibroblasts and synovial tissue in rheumatoid arthritis. J Stanczyk DM Pedrioli F Brentano O Sanchez-Pernaute C Kolling RE Gay M Detmar S Gay D Kyburz, Arthritis Rheum 2008 58 1001 1009 10.1002/art.23386 18383392 (Pubitemid 351564001)
112. NF-kappaB-dependent induction of microRNA miR-146, an inhibitor targeted to signaling proteins of innate immune responses. KD Taganov MP Boldin KJ Chang D Baltimore, Proc Natl Acad Sci USA 2006 103 12481 12486 10.1073/pnas.0605298103 16885212
113. Modulation of miR-155 and miR-125b levels following lipopolysaccharide/ TNF-alpha stimulation and their possible roles in regulating the response to endotoxin shock. E Tili JJ Michaille A Cimino S Costinean CD Dumitru B Adair M Fabbri H Alder CG Liu GA Calin CM Croce, J Immunol 2007 179 5082 5089 17911593
114. Upregulated miR-146a expression in peripheral blood mononuclear cells from rheumatoid arthritis patients. KM Pauley M Satoh AL Chan MR Bubb WH Reeves EK Chan, Arthritis Res Ther 2008 10 101 10.1186/ar2344 18759964
115. Expression of microRNA-146 in rheumatoid arthritis synovial tissue. T Nakasa S Miyaki A Okubo M Hashimoto K Nishida M Ochi H Asahara, Arthritis Rheum 2008 58 1284 1292 10.1002/art.23429 18438844
116. Silencing of microRNA-155 in mice during acute inflammatory response leads to derepression of c/ebp Beta and down-regulation of G-CSF. J Worm J Stenvang A Petri KS Frederiksen S Obad J Elmen M Hedtjarn EM Straarup JB Hansen S Kauppinen, Nucleic Acids Res 2009 37 5784 5792 10.1093/nar/gkp577 19596814
117. High serum and synovial fluid granulocyte colony stimulating factor (G-CSF) concentrations in patients with rheumatoid arthritis. H Nakamura Y Ueki S Sakito K Matsumoto M Yano S Miyake T Tominaga M Tominaga K Eguchi, Clin Exp Rheumatol 2000 18 713 718 11138333
118. MicroRNAs: novel regulators in skin inflammation. E Sonkoly M Stahle A Pivarcsi, Clin Exp Dermatol 2008 33 312 315 10.1111/j.1365-2230.2008.02804.x 18419608
119. MicroRNAs: novel regulators involved in the pathogenesis of Psoriasis? E Sonkoly T Wei PC Janson A Sääf L Lundeberg M Tengvall-Linder G Norstedt H Alenius B Homey A Scheynius M Ståhle A Pivarcsi, PLoS One 2007 2 610 10.1371/journal.pone.0000610 17622355
120. SOCS3 negatively regulates IL-6 signaling in vivo. BA Croker DL Krebs JG Zhang S Wormald TA Willson EG Stanley L Robb CJ Greenhalgh I Förster BE Clausen NA Nicola D Metcalf DJ Hilton AW Roberts WS Alexander, Nat Immunol 2003 4 540 545 10.1038/ni931 12754505
121. Stat3 links activated keratinocytes and immunocytes required for development of psoriasis in a novel transgenic mouse model. S Sano KS Chan S Carbajal J Clifford M Peavey K Kiguchi S Itami BJ Nickoloff J DiGiovanni, Nat Med 2005 11 43 49 10.1038/nm1162 15592573
122. miR-203 represses 'stemness' by repressing DeltaNp63. AM Lena R Shalom-Feuerstein P Rivetti di Val Cervo D Aberdam RA Knight G Melino E Candi, Cell Death Differ 2008 15 1187 1195 10.1038/cdd.2008.69 18483491
123. microRNAs in inflammation. E Sonkoly A Pivarcsi, Int Rev Immunol 2009 28 535 561 10.3109/08830180903208303 19954362
124. Regulation of suppressor of cytokine signaling 3 (SOCS3) mRNA stability by TNF-alpha involves activation of the MKK6/p38MAPK/MK2 cascade. C Ehlting WS Lai F Schaper ED Brenndorfer RJ Matthes PC Heinrich S Ludwig PJ Blackshear M Gaestel D Haussinger JG Bode, J Immunol 2007 178 2813 2826 17312125
125. miRNPs: versatile regulators of gene expression in vertebrate cells. JA Steitz S Vasudevan, Biochem Soc Trans 2009 37 931 935 10.1042/BST0370931 19754429
126. Switching from repression to activation: microRNAs can up-regulate translation. S Vasudevan Y Tong JA Steitz, Science 2007 318 1931 1934 10.1126/science.1149460 18048652
127. Natural mRNA is required for directing Met-tRNA(f) binding to 40S ribosomal subunits in animal cells: involvement of Co-eIF-2A in natural mRNA-directed initiation complex formation. AL Roy D Chakrabarti B Datta RE Hileman NK Gupta, Biochemistry 1988 27 8203 8209 10.1021/bi00421a033 3233204
128. Specific sequences in the fragile X syndrome protein FMR1 and the FXR proteins mediate their binding to 60S ribosomal subunits and the interactions among them. MC Siomi Y Zhang H Siomi G Dreyfuss, Mol Cell Biol 1996 16 3825 3832 8668200
129. Fragile X-related protein FXR1P regulates proinflammatory cytokine tumor necrosis factor expression at the post-transcriptional level. J Garnon C Lachance S Di Marco Z Hel D Marion MC Ruiz MM Newkirk EW Khandjian D Radzioch, J Biol Chem 2005 280 5750 5763 10.1074/jbc.M401988200 15548538
130. Involvement of microRNA in AU-rich element-mediated mRNA instability. Q Jing S Huang S Guth T Zarubin A Motoyama J Chen F Di Padova SC Lin H Gram J Han, Cell 2005 120 623 634 10.1016/j.cell.2004.12.038 15766526
131. RNAi therapeutics: principles, prospects and challenges. L Aagaard JJ Rossi, Adv Drug Deliv Rev 2007 59 75 86 10.1016/j.addr.2007.03.005 17449137
132. RNAi-based therapeutics-current status, challenges and prospects. K Tiemann JJ Rossi, EMBO Mol Med 2009 1 142 151 10.1002/emmm.200900023 20049714
133. Duplexes of 21-nucleotide RNAs mediate RNA interference in cultured mammalian cells. SM Elbashir J Harborth W Lendeckel A Yalcin K Weber T Tuschl, Nature 2001 411 494 498 10.1038/35078107 11373684
134. Viruses and interferons. GC Sen, Annu Rev Microbiol 2001 55 255 281 10.1146/annurev.micro.55.1.255 11544356
135. Lentiviral-mediated RNA interference. T Abbas-Terki W Blanco-Bose N Deglon W Pralong P Aebischer, Hum Gene Ther 2002 13 2197 2201 10.1089/104303402320987888 12542850
136. A system for stable expression of short interfering RNAs in mammalian cells. TR Brummelkamp R Bernards R Agami, Science 2002 296 550 553 10.1126/science.1068999 11910072
137. Widespread siRNA "off-target" transcript silencing mediated by seed region sequence complementarity. AL Jackson J Burchard J Schelter BN Chau M Cleary L Lim PS Linsley, RNA 2006 12 1179 1187 10.1261/rna.25706 16682560
138. Cutaneous short-interfering RNA therapy. B Geusens N Sanders T Prow M Van Gele J Lambert, Expert Opin Drug Deliv 2009 6 1333 1349 10.1517/ 17425240903304032 19941411
139. Amelioration of psoriasis by anti-TNF-alpha RNAi in the xenograft transplantation model. M Jakobsen K Stenderup C Rosada B Moldt S Kamp TN Dam TG Jensen JG Mikkelsen, Mol Ther 2009 17 1743 1753 10.1038/mt.2009.141 19568223
140. Sustainable systemic delivery via a single injection of lentivirus into human skin tissue. SC Baek Q Lin PB Robbins H Fan PA Khavari, Hum Gene Ther 2001 12 1551 1558 10.1089/10430340152480276 11506697
141. Lentivectors for regulated and reversible cutaneous gene delivery. Z Siprashvili PA Khavari, Mol Ther 2004 9 93 100 10.1016/j.ymthe.2003.09.016 14741782
142. Structure of the skin barrier and its modulation by vesicular formulations. JA Bouwstra PL Honeywell-Nguyen GS Gooris M Ponec, Prog Lipid Res 2003 42 1 36 10.1016/S0163-7827(02)00028-0 12467638
143. The skin barrier in healthy and diseased state. JA Bouwstra M Ponec, Biochim Biophys Acta 2006 1758 2080 2095 10.1016/j.bbamem.2006.06.021 16945325
144. Efficient expression of naked plasmid DNA in mucosal epithelium: prospective for the treatment of skin lesions. UR Hengge W Pfutzner M Williams M Goos JC Vogel, The Journal of investigative dermatology 1998 111 605 608 10.1046/j.1523-1747.1998.00353.x 9764840
145. Expression of naked DNA in human, pig, and mouse skin. UR Hengge PS Walker JC Vogel, J Clin Invest 1996 97 2911 2916 10.1172/JCI118750 8675706
146. Single-nucleotide-specific siRNA targeting in a dominant-negative skin model. RP Hickerson FJ Smith RE Reeves CH Contag D Leake SA Leachman LM Milstone WH McLean RL Kaspar, The Journal of investigative dermatology 2008 128 594 605 17914454
147. Delivery and inhibition of reporter genes by small interfering RNAs in a mouse skin model. Q Wang H Ilves P Chu CH Contag D Leake BH Johnston RL Kaspar,
148. Development of therapeutic siRNAs for pachyonychia congenita. FJ Smith RP Hickerson JM Sayers RE Reeves CH Contag D Leake RL Kaspar WH McLean, The Journal of investigative dermatology 2008 128 50 58 10.1038/sj.jid.5701040 17762855
149. Therapeutic siRNAs for dominant genetic skin disorders including pachyonychia congenita. SA Leachman RP Hickerson PR Hull FJ Smith LM Milstone EB Lane SJ Bale DR Roop WH McLean RL Kaspar, Journal of dermatological science 2008 18495438
150. First-in-human mutation-targeted siRNA phase Ib trial of an inherited skin disorder. SA Leachman RP Hickerson ME Schwartz EE Bullough SL Hutcherson KM Boucher CD Hansen MJ Eliason GS Srivatsa DJ Kornbrust FJ Smith WI McLean LM Milstone RL Kaspar, Mol Ther 2010 18 442 446 10.1038/mt.2009.273 19935778
151. Microporation techniques for enhanced delivery of therapeutic agents. TR Singh MJ Garland CM Cassidy K Migalska YK Demir S Abdelghany E Ryan AD Woolfson RF Donnelly, Recent Pat Drug Deliv Formul 4 1 17 10.2174/187221110789957174 19807682
152. Microneedles and other physical methods for overcoming the stratum corneum barrier for cutaneous gene therapy. S Coulman C Allender J Birchall, Crit Rev Ther Drug Carrier Syst 2006 23 205 258 17206925
153. Minimally invasive cutaneous delivery of macromolecules and plasmid DNA via microneedles. SA Coulman D Barrow A Anstey C Gateley A Morrissey N Wilke C Allender K Brain JC Birchall, Curr Drug Deliv 2006 3 65 75 10.2174/ 156720106775197510 16472095
154. Sonophoresis: recent advancements and future trends. R Rao S Nanda, J Pharm Pharmacol 2009 61 689 705 10.1211/jpp.61.06.0001 19505359
155. Modulation of scratching behavior by silencing an endogenous cyclooxygenase-1 gene in the skin through the administration of siRNA. T Inoue M Sugimoto T Sakurai R Saito N Futaki Y Hashimoto Y Honma I Arai S Nakaike, The journal of gene medicine 2007 9 994 1001 10.1002/jgm.1091 17703497
156. Therapeutic RNA interference of malignant melanoma by electrotransfer of small interfering RNA targeting Mitf. N Nakai T Kishida M Shin-Ya J Imanishi Y Ueda S Kishimoto O Mazda, Gene therapy 2007 14 357 365 10.1038/sj.gt.3302868 17024102
157. Vesicular systems for delivering conventional small organic molecules and larger macromolecules to and through human skin. GM El Maghraby AC Williams, Expert Opin Drug Deliv 2009 6 149 163 10.1517/17425240802691059 19239387
158. Therapeutic silencing of an endogenous gene by siRNA cream in an arthritis model mouse. M Takanashi K Oikawa K Sudo M Tanaka K Fujita A Ishikawa S Nakae RL Kaspar M Matsuzaki M Kudo M Kuroda, Gene Ther 2009 16 982 989 10.1038/gt.2009.66 19474812
159. Topical application of cream-emulsified CD86 siRNA ameliorates allergic skin disease by targeting cutaneous dendritic cells. P Ritprajak M Hashiguchi M Azuma, Mol Ther 2008 16 1323 1330 10.1038/mt.2008.91 18461054
160. Silencing of microRNAs in vivo with 'antagomirs'. J Krutzfeldt N Rajewsky R Braich KG Rajeev T Tuschl M Manoharan M Stoffel, Nature 2005 438 685 689 10.1038/nature04303 16258535
161. Lentivirus-mediated antagomir expression for specific inhibition of miRNA function. M Scherr L Venturini K Battmer M Schaller-Schoenitz D Schaefer I Dallmann A Ganser M Eder, Nucleic Acids Res 2007 35 149 10.1093/nar/gkm971 18025036
162. Therapeutic silencing of miR-10b inhibits metastasis in a mouse mammary tumor model. L Ma F Reinhardt E Pan J Soutschek B Bhat EG Marcusson J Teruya-Feldstein GW Bell RA Weinberg, Nat Biotechnol 2010