The biology and functions of Th22 cells

Advances in Experimental Medicine and Biology

Volumn 841, Issue , 2014, Pages 209-230

  Jia, Lei ^a   Wu, Changyou ^a  

^a SUN YAT SEN UNIVERSITY   (China)

Author keywords

Disease; IL 22 STAT; Th22 cell

Indexed keywords

BINDING PROTEIN; INTERLEUKIN 22; INTERLEUKIN 22 BINDING PROTEIN; INTERLEUKIN 22 RECEPTOR; INTERLEUKIN 22 RECEPTOR 1; RECEPTOR PROTEIN; UNCLASSIFIED DRUG; INTERLEUKIN DERIVATIVE; INTERLEUKIN RECEPTOR; INTERLEUKIN-22; INTERLEUKIN-22 RECEPTOR;

ALLERGIC ASTHMA; ARTICLE; ATOPIC DERMATITIS; CONFORMATIONAL TRANSITION; CYTOKINE PRODUCTION; EPITHELIUM CELL; FIBROBLAST; HEPATITIS; HUMAN; INFECTION; INFLAMMATORY BOWEL DISEASE; LIVER CELL; LYMPHOCYTE DIFFERENTIATION; LYMPHOCYTE FUNCTION; MALIGNANT NEOPLASTIC DISEASE; MOLECULAR PATHOLOGY; NONHUMAN; PANCREAS CELL; PANCREATITIS; PRIORITY JOURNAL; PROTEIN FUNCTION; PSORIASIS; REGULATORY MECHANISM; RHEUMATOID ARTHRITIS; SIGNAL TRANSDUCTION; SKIN ALLERGY; SYSTEMIC LUPUS ERYTHEMATOSUS; SYSTEMIC SCLEROSIS; TH22 CELL; ANIMAL; AUTOIMMUNE DISEASE; CELL DIFFERENTIATION; HELPER CELL; IMMUNOLOGY; NEOPLASM; PHYSIOLOGY;

ANIMALS; AUTOIMMUNE DISEASES; CELL DIFFERENTIATION; HUMANS; INTERLEUKINS; NEOPLASMS; PANCREATITIS; PSORIASIS; RECEPTORS, INTERLEUKIN; T-LYMPHOCYTES, HELPER-INDUCER;

EID: 84922518641     PISSN: 00652598     EISSN: 22148019     Source Type: Book Series    
DOI: 10.1007/978-94-017-9487-9_8     Document Type: Article

References (133)

1. Aggarwal S, Xie MH, Maruoka M, Foster J, Gurney AL (2001) Acinar cells of the pancreas are a target of interleukin-22. J Interferon Cytokine Res 21:1047–1053
2. Andoh A, Zhang Z, Inatomi O, Fujino S, Deguchi Y, Araki Y, Tsujikawa T, Kitoh K, Kim- Mitsuyama S, Takayanagi A et al (2005) Interleukin-22, a member of the IL-10 subfamily, induces inflammatory responses in colonic subepithelialmyofibroblasts. Gastroenterology 129:969–984
3. Aujla SJ, Chan YR, Zheng M, Fei M, Askew DJ, Pociask DA, Reinhart TA, McAllister F, Edeal J, Gaus K et al (2008) IL-22 mediates mucosal host defense against gram-negative bacterial pneumonia. Nat Med 14:275–281
4. Besnard AG, Sabat R, Dumoutier L, Renauld JC, Willart M, Lambrecht B, Teixeira MM, Charron S, Fick L, Erard F et al (2011) Dual Role of IL-22 in allergic airway inflammation and its cross-talk with IL-17A. Am J Respir Crit Care Med 183:1153–1163
5. Bleicher L, de Moura PR, Watanabe L, Colau D, Dumoutier L, Renauld JC, Polikarpov I (2008) Crystal structure of the IL-22/IL-22R1 complex and its implications for the IL-22 signaling mechanism. Febs Lett 582:2985–2992
6. Boniface K, Guignouard E, Pedretti N, Garcia M, Delwail A, Bernard FX, Nau F, Guillet G, Dagregorio G, Yssel H et al (2007) A role for T cell-derived interleukin 22 in psoriatic skin inflammation. Clin Exp Immunol 150:407–415
7. Brand S, Beigel F, Olszak T, Zitzmann K, Eichhorst ST, Otte JM, Diepolder H, Marquardt A, Jagla W, Popp A et al (2006) IL-22 is increased in active Crohn’s disease and promotes proinflammatory gene expression and intestinal epithelial cell migration. Am J Physiol Gastrointest Liver Physiol 290:827–838
8. Braun RK, Martin A, Shah S, Iwashima M, Medina M, Byrne K, Sethupathi P, Wigfield CH, Brand DD, Love RB (2010) Inhibition of bleomycin-induced pulmonary fibrosis through pre-treatment with collagen type V. J Heart Lung Transplant 29:873–880
9. Buonocore S, Ahern PP, Uhlig HH, Ivanov II, Littman DR, Maloy KJ, Powrie F (2010) Innate lymphoid cells drive interleukin-23-dependent innate intestinal pathology. Nature 464:1371–1375
10. Caproni M, Antiga E, Melani L, Volpi W, Del BE, Fabbri P (2009) Serum levels of IL-17 and IL-22 are reduced by etanercept, but not by acitretin, in patients with psoriasis: a randomized-controlled trial. J Clin Immunol 29:210–214
11. Cella M, Fuchs A, Vermi W, Facchetti F, Otero K, Lennerz JK, Doherty JM, Mills JC, Colonna M (2009) A human natural killer cell subset provides an innate source of IL-22 for mucosal immunity. Nature 457:722–725
12. Chang H, Hanawa H, Liu H, Yoshida T, Hayashi M, Watanabe R, Abe S, Toba K, Yoshida K, Elnaggar R et al (2006) Hydrodynamic-based delivery of an interleukin-22-Ig fusion gene ameliorates experimental autoimmune myocarditis in rats. J Immunol 177:3635–3643
13. Cheng F, Guo Z, Xu H, Yan D, Li Q (2009) Decreased plasma IL22 levels, but not increased IL17 and IL23 levels, correlate with disease activity in patients with systemic lupus erythematosus. Ann Rheum Dis 68:604–606
14. Chestovich PJ, Uchida Y, Chang W, Ajalat M, Lassman C, Sabat R, Busuttil RW, Kupiec- Weglinski JW (2012) Interleukin-22: implications for liver ischemia-reperfusion injury. Transplantation 93:485–492
15. Da RLJ, Duarte AL, Dantas AT, Mariz HA, Pitta IR, Galdino SL, Pitta MG (2012) Increased serum interleukin 22 in patients with rheumatoid arthritis and correlation with disease activity. J Rheumatol 39:1320–1325
16. Dambacher J, Beigel F, Zitzmann K, Heeg MH, Goke B, Diepolder HM, Auernhammer CJ, Brand S (2008) The role of interleukin-22 in hepatitis C virus infection. Cytokine 41:209–216
17. De Luca A, Zelante T, D’Angelo C, Zagarella S, Fallarino F, Spreca A, Iannitti RG, Bonifazi P, Renauld JC, Bistoni F et al (2010) IL-22 defines a novel immune pathway of antifungal resistance. Mucosal Immunol 3:361–373
18. deMoura PR, Watanabe L, Bleicher L, Colau D, Dumoutier L, Lemaire MM, Renauld JC, Polikarpov I (2009) Crystal structure of a soluble decoy receptor IL-22BP bound to interleukin-22. FebsLett 583:1072–1077
19. Dige A, Stoy S, Rasmussen TK, Kelsen J, Hvas CL, Sandahl TD, Dahlerup JF, Deleuran B, Agnholt J (2013) Increased levels of circulating Th17 cells in quiescent versus active Crohn’s disease. J Crohns Colitis 7:248–255
20. Duhen T, Geiger R, Jarrossay D, Lanzavecchia A, Sallusto F (2009) Production of interleukin 22 but not interleukin 17 by a subset of human skin-homing memory T cells. Nat Immunol 10:857–863
21. Dumoutier L, Leemans C, Lejeune D, Kotenko SV, Renauld JC (2001) Cutting edge: STAT activation by IL-19, IL-20 and mda-7 through IL-20 receptor complexes of two types. J Immunol 167:3545–3549
22. Dumoutier L, Lejeune D, Colau D, Renauld JC (2001) Cloning and characterization of IL-22 binding protein, a natural antagonist of IL-10-related T cell-derived inducible factor/IL-22. J Immunol 166:7090–7095
23. Dumoutier L, Louahed J, Renauld JC (2000) Cloning and characterization of IL-10-related T cell-derived inducible factor (IL-TIF), a novel cytokine structurally related to IL-10 and inducible by IL-9. J Immunol 164:1814–1819
24. Dumoutier L, Van Roost E, Ameye G, Michaux L, Renauld JC (2000) IL-TIF/IL-22: genomic organization and mapping of the human and mouse genes. Genes Immun 1:488–494
25. Dumoutier L, Van Roost E, Colau D, Renauld JC (2000) Human interleukin-10-related T cell-derived inducible factor: molecular cloning and functional characterization as an hepatocyte-stimulating factor. Proc Natl Acad Sci USA 97:10144–10149
26. Eyerich S, Eyerich K, Pennino D, Carbone T, Nasorri F, Pallotta S, Cianfarani F, Odorisio T, Traidl-Hoffmann C, Behrendt H et al (2009) Th22 cells represent a distinct human T cell subset involved in epidermal immunity and remodeling. J Clin Invest 119:3573–3585
27. Farfariello V, Amantini C, Nabissi M, Morelli MB, Aperio C, Caprodossi S, Carlucci A, Bianchi AM, Santoni G (2011) IL-22 mRNA in peripheral blood mononuclear cells from allergic rhinitic and asthmatic pediatric patients. Pediatr Allergy Immunol 22:419–423
28. Feng D (2014) Interleukin-22, liver progenitor cells, and liver cancer. Hepatology 60:427–428
29. Feng D, Kong X, Weng H, Park O, Wang H, Dooley S, Gershwin ME, Gao B (2012) Interleukin-22 promotes proliferation of liver stem/progenitor cells in mice and patients with chronic hepatitis B virus infection. Gastroenterology 143:188–198
30. Ferran M, Galvan AB, Rincon C, Romeu ER, Sacrista M, Barboza E, Gimenez-Arnau A, Celada A, Pujol RM, Santamaria-Babi LF (2013) Streptococcus induces circulating CLA(+) memory T-cell-dependent epidermal cell activation in psoriasis. J Invest Dermatol 133:999–1007
31. Forster R, Braun A, Worbs T (2012) Lymph node homing of T cells and dendritic cells via afferent lymphatics. Trends Immunol 33:271–280
32. Fujita H, Nograles KE, Kikuchi T, Gonzalez J, Carucci JA, Krueger JG (2009) Human Langerhans cells induce distinct IL-22-producing CD4+ T cells lacking IL-17 production. Proc Natl Acad Sci U S A 106:21795–21800
33. Geboes L, Dumoutier L, Kelchtermans H, Schurgers E, Mitera T, Renauld JC, Matthys P (2009) Proinflammatory role of the Th17 cytokine interleukin-22 in collagen-induced arthritis in C57BL/6 mice. Arthritis Rheum 60:390–395
34. Gessner MA, Werner JL, Lilly LM, Nelson MP, Metz AE, Dunaway CW, Chan YR, Ouyang W, Brown GD, Weaver CT et al (2012) Dectin-1-dependent interleukin-22 contributes to early innate lung defense against Aspergillus fumigatus. Infect Immun 80:410–417
35. Ghoreschi K, Laurence A, Yang XP, Tato CM, McGeachy MJ, Konkel JE, Ramos HL, Wei L, Davidson TS, Bouladoux N et al (2010) Generation of pathogenic T(H)17 cells in the absence of TGF-beta signalling. Nature 467:967–971
36. Graham AC, Carr KD, Sieve AN, Indramohan M, Break TJ, Berg RE (2011) IL-22 production is regulated by IL-23 during Listeria monocytogenes infection but is not required for bacterial clearance or tissue protection. PLoS ONE 6:e17171
37. Guilloteau K, Paris I, Pedretti N, Boniface K, Juchaux F., Huguier V, Guillet G, Bernard FX, Lecron JC, Morel F (2010) Skin Inflammation Induced by the synergistic action of IL-17A, IL-22, Oncostatin M, IL-1{alpha}, and TNF-{alpha} recapitulates some features of psoriasis. J Immunol 184:5263–5270
38. Guttman-Yassky E, Nograles KE, Krueger JG (2011) Contrasting pathogenesis of atopic dermatitis and psoriasis–part II: immune cell subsets and therapeutic concepts. J Allergy Clin Immunol 127:1420–1432
39. Hamada H, Garcia-Hernandez ML, Reome JB, Misra SK, Strutt TM, McKinstry KK, Cooper AM, Swain SL, Dutton RW (2009) Tc17, a unique subset of CD8 T cells that can protect against lethal influenza challenge. J Immunol 182:3469–3481
40. Harper EG, Guo C, Rizzo H, Lillis JV, Kurtz SE, Skorcheva I, Purdy D, Fitch E, Iordanov M, Blauvelt A (2009) Th17 cytokines stimulate CCL20 expression in keratinocytes in vitro and in vivo: implications for psoriasis pathogenesis. J Invest Dermatol 129:2175–2183
41. Hayashida S, Uchi H, Takeuchi S, Esaki H, Moroi Y, Furue M (2011) Significant correlation of serum IL-22 levels with CCL17 levels in atopic dermatitis. J Dermatol Sci 61:78–79
42. Hill NR, Levy JC, Matthews DR (2013) Expansion of the homeostasis model assessment of beta-cell function and insulin resistance to enable clinical trial outcome modeling through the interactive adjustment of physiology and treatment effects: iHOMA2. Diabetes Care 36:2324–2330
43. Huber S, Gagliani N, Zenewicz LA, Huber FJ, Bosurgi L, Hu B, Hedl M, Zhang W, O’Connor WJ, Murphy AJ et al (2012) IL-22BP is regulated by the inflamma some and modulates tumorigenesis in the intestine. Nature 491:259–263
44. Ikeuchi H, Kuroiwa T, Hiramatsu N, Kaneko Y, Hiromura K, Ueki K, Nojima Y (2005) Expression of interleukin-22 in rheumatoid arthritis: potential role as a proinflammatory cytokine. Arthritis Rheum 52:1037–1046
45. Jiang R, Tan Z, Deng L, Chen Y, Xia Y, Gao Y, Wang X, Sun B (2011) Interleukin-22 promotes human hepatocellular carcinoma by activation of STAT3. Hepatology 54:900–909
46. Jiang R, Wang H, Deng L, Hou J, Shi R, Yao M, Gao Y, Yao A, Wang X, Yu L et al (2013) IL-22 is related to development of human colon cancer by activation of STAT3. BMC Cancer 13:59
47. Jones BC, Logsdon NJ, Walter MR (2008) Crystallization and preliminary X-ray diffraction analysis of human IL-22 bound to the extracellular IL-22R1 chain. Acta Crystallogr Sect F Struct Biol Cryst Commun 64:266–269
48. Kagami S, Rizzo HL, Kurtz SE, Miller LS, Blauvelt A (2010) IL-23 and IL-17A, but not IL-12 and IL-22, are required for optimal skin host defense against Candida albicans. J Immunol 185:5453–5462
49. Kagami S, Rizzo HL, Lee JJ, Koguchi Y, Blauvelt A (2010) Circulating Th17, Th22, and Th1 cells are increased in psoriasis. J Invest Dermatol 130:1373–1383
50. Kanda N, Watanabe S (2012) Increased serum human beta-defensin-2 levels in atopic dermatitis: relationship to IL-22 and oncostatin M. Immunobiology 217:436–445
51. Ki SH, Park O, Zheng M, Morales-Ibanez O, Kolls JK, Bataller R, Gao B (2010) Interleukin- 22 treatment ameliorates alcoholic liver injury in a murine model of chronic-binge ethanol feeding: role of signal transducer and activator of transcription 3. Hepatology 52:1291–1300
52. Kim CJ, Nazli A, Rojas OL, Chege D, Alidina Z, Huibner S, Mujib S, Benko E, Kovacs C, Shin LY et al (2012) A role for mucosal IL-22 production and Th22 cells in HIV-associated mucosal immunopathogenesis. Mucosal Immunol 5:670–680
53. Kim KW, Kim HR, Park JY, Park JS, Oh HJ, Woo YJ, Park MK, Cho ML, Lee SH (2012) Interleukin-22 promotes osteoclastogenesis in rheumatoid arthritis through induction of RANKL in human synovial fibroblasts. Arthritis Rheum 64:1015–1023
54. Kobold S, Volk S, Clauditz T, Kupper NJ, Minner S, Tufman A, Duwell P, Lindner M, Koch I, Heidegger S et al (2013) Interleukin-22 is frequently expressed in small- and large-cell lung cancer and promotes growth in chemotherapy-resistant cancer cells. J Thorac Oncol 8:1032–1042
55. Kotenko SV, Izotova LS, Mirochnitchenko OV, Esterova E, Dickensheets H, Donnelly RP, Pestka S (2001) Identification, cloning, and characterization of a novel soluble receptor that binds IL-22 and neutralizes its activity. J Immunol 166:7096–7103
56. Larsen JM, Bonefeld CM, Poulsen SS, Geisler C, Skov L (2009) IL-23 and T(H)17-mediated inflammation in human allergic contact dermatitis. J Allergy Clin Immunol 123:486–492
57. Lee JS, Cella M, McDonald KG, Garlanda C, Kennedy GD, Nukaya M, Mantovani A, Kopan R, Bradfield CA, Newberry RD et al (2012) AHR drives the development of gut ILC22 cells and postnatal lymphoid tissues via pathways dependent on and independent of Notch. Nat Immunol 13:144–151
58. Leipe J, Schramm MA, Grunke M, Baeuerle M, Dechant C, Nigg AP, Witt MN, Vielhauer V, Reindl CS, Schulze-Koops H et al (2011) Interleukin 22 serum levels are associated with radiographic progression in rheumatoid arthritis. Ann Rheum Dis 70:1453–1457
59. Lejeune D, Dumoutier L, Constantinescu S, Kruijer W, Schuringa JJ, Renauld JC (2002) Interleukin-22 (IL-22) activates the JAK/STAT, ERK, JNK, and p38 MAP kinase pathways in a rat hepatoma cell line. Pathways that are shared with and distinct from IL-10. J BiolChem 277:33676–33682
60. Liang SC, Nickerson-Nutter C, Pittman DD, Carrier Y, Goodwin DG, Shields KM, Lambert AJ, Schelling SH, Medley QG, Ma HL et al (2010) IL-22 induces an acute-phase response. J Immunol 185:5531–5538
61. Liang SC, Tan XY, Luxenberg DP, Karim R, Dunussi-Joannopoulos K, Collins M, Fouser LA (2006) Interleukin (IL)-22 and IL-17 are coexpressed by Th17 cells and cooperatively enhance expression of antimicrobial peptides. J Exp Med 203:2271–2279
62. Lo RS, Dumoutier L, Couillin I, Van Vyve C, Yakoub Y, Uwambayinema F, Marien B, van den Brule S, Van Snick J, Uyttenhove C et al (2010) IL-17A-producing gammadelta T and Th17 lymphocytes mediate lung inflammation but not fibrosis in experimental silicosis. J Immunol 184:6367–6377
63. Logsdon NJ, Jones BC, Allman JC, Izotova L, Schwartz B, Pestka S, Walter MR (2004) The IL-10R2 binding hot spot on IL-22 is located on the N-terminal helix and is dependent on N-linked glycosylation. J Mol Biol 342:503–514
64. Logsdon NJ, Jones BC, Josephson K, Cook J, Walter MR (2002) Comparison of interleukin- 22 and interleukin-10 soluble receptor complexes. J Interferon Cytokine Res 22:1099–1112
65. Luci C, Reynders A, Ivanov II, Cognet C, Chiche L, Chasson L, Hardwigsen J, Anguiano E, Banchereau J, Chaussabel D et al (2009) Influence of the transcription factor ROR gammat on the development of NKp46+ cell populations in gut and skin. Nat Immunol 10:75–82
66. Marijnissen RJ, Koenders MI, Smeets RL, Stappers MH, Nickerson-Nutter C, Joosten LA, Boots AM, van den Berg WB (2011) Increased expression of interleukin-22 by synovial Th17 cells during late stages of murine experimental arthritis is controlled by interleukin-1 and enhances bone degradation. Arthritis Rheum 63:2939–2948
67. Martin B, Hirota K, Cua DJ, Stockinger B, Veldhoen M (2009) Interleukin-17-producing gammadelta T cells selectively expand in response to pathogen products and environmental signals. Immunity 31:321–330
68. Martin JC, Beriou G, Heslan M, Chauvin C, Utriainen L, Aumeunier A, Scott CL, Mowat A, Cerovic V, Houston SA et al (2014) Interleukin-22 binding protein (IL-22BP) is constitutively expressed by a subset of conventional dendritic cells and is strongly induced by retinoic acid. Mucosal Immunol 7:101–113
69. Mastelic B, Do RA, Veldhoen M, Renauld JC, Jarra W, Sponaas AM, Roetynck S, Stockinger B, Langhorne J (2012) IL-22 protects against liver pathology and lethality of an experimental blood-stage malaria infection. Front Immunol 3:85
70. Mathian A, Parizot C, Dorgham K, Trad S, Arnaud L, Larsen M, Miyara M, Hie M, Piette JC, Frances C et al (2012) Activated and resting regulatory T cell exhaustion concurs with high levels of interleukin-22 expression in systemic sclerosis lesions. Ann Rheum Dis 71:1227–1234
71. McGee HM, Schmidt BA, Booth CJ, Yancopoulos GD, Valenzuela DM, Murphy AJ, Stevens S, Flavell RA, Horsley V (2013) IL-22 promotes fibroblast-mediated wound repair in the skin. J Invest Dermatol 133:1321–1329
72. Mitra A, Raychaudhuri SK, Raychaudhuri SP (2012) IL-22 induced cell proliferation is regulated by PI3 K/Akt/mTOR signaling cascade. Cytokine 60:38–42
73. Nagakawa H, Shimozato O, Yu L, Takiguchi Y, Tatsumi K, Kuriyama T, Tagawa M (2004) Expression of interleukin-22 in murine carcinoma cells did not influence tumour growth in vivo but did improve survival of the inoculated hosts. Scand J Immunol 60:449–454
74. Nagem RA, Colau D, Dumoutier L, Renauld JC, Ogata C, Polikarpov I (2002) Crystal structure of recombinant human interleukin-22. Structure 10:1051–1062
75. Nakagome K, Imamura M, Kawahata K, Harada H, Okunishi K, Matsumoto T, Sasaki O, Tanaka R, Kano MR, Chang H, Hanawa H, Miyazaki J, Yamamoto K, Dohi M (2011) High expression of IL-22 suppresses antigen-induced immune responses and eosinophilic airway inflammation via an IL-10-associated mechanism. J Immunol. 187:89–5077
76. Nie Y, Erion DM, Yuan Z, Dietrich M, Shulman GI, Horvath TL, Gao Q (2009) STAT3 inhibition of gluconeogenesis is downregulated by SirT1. Nat Cell Biol 11:492–500
77. Nograles KE, Zaba LC, Guttman-Yassky E, Fuentes-Duculan J, Suarez-Farinas M, Cardinale I, Khatcherian A, Gonzalez J, Pierson KC, White TR et al (2008) Th17 cytokines interleukin (IL)-17 and IL-22 modulate distinct inflammatory and keratinocyte-response pathways. Br J Dermatol 159:1092–1102
78. Nograles KE, Zaba LC, Shemer A, Fuentes-Duculan J, Cardinale I, Kikuchi T, Ramon M, Bergman R, Krueger JG, Guttman-Yassky E (2009) IL-22-producing “T22” T cells account for upregulated IL-22 in atopic dermatitis despite reduced IL-17-producing TH17 T cells. J Allergy ClinImmunol 123:1244–1252
79. Paget C, Ivanov S, Fontaine J, Renneson J, Blanc F, Pichavant M, Dumoutier L, Ryffel B, Renauld JC, Gosset P et al (2012) Interleukin-22 is produced by invariant natural killer T lymphocytes during influenza A virus infection: potential role in protection against lung epithelial damages. J BiolChem 287:8816–8829
80. Pan CX, Tang J, Wang XY, Wu FR, Ge JF, Chen FH (2014) Role of Interleukin-22 in liver diseases. Inflamm Res
81. Pan H, Hong F, Radaeva S, Gao B (2004) Hydrodynamic gene delivery of interleukin-22 protects the mouse liver from concanavalin A-, carbon tetrachloride-, and Fas ligand-induced injury via activation of STAT3. Cell Mol Immunol 1:43–49
82. Pan HF, Zhao XF, Yuan H, Zhang WH, Li XP, Wang GH, Wu GC, Tang XW, Li WX, Li LH et al (2009) Decreased serum IL-22 levels in patients with systemic lupus erythematosus. Clin Chim Acta 401:179–180
83. Park O, Wang H, Weng H, Feigenbaum L, Li H, Yin S, Ki SH, Yoo SH, Dooley S, Wang FS et al (2011) In vivo consequences of liver-specific interleukin-22 expression in mice: implications for human liver disease progression. Hepatology 54:252–261
84. Pickert G, Neufert C, Leppkes M, Zheng Y, Wittkopf N, Warntjen M, Lehr HA, Hirth S, Weigmann B, Wirtz S et al (2009) STAT3 links IL-22 signaling in intestinal epithelial cells to mucosal wound healing. J Exp Med 206:1465–1472
85. Radaeva S, Sun R, Pan HN, Hong F, Gao B (2004) Interleukin 22 (IL-22) plays a protective role in T cell-mediated murine hepatitis: IL-22 is a survival factor for hepatocytes via STAT3 activation. Hepatology 39:1332–1342
86. Ramirez JM, Brembilla NC, Sorg O, Chicheportiche R, Matthes T, Dayer JM, Saurat JH, Roosnek E, Chizzolini C (2010) Activation of the aryl hydrocarbon receptor reveals distinct requirements for IL-22 and IL-17 production by human T helper cells. Eur J Immunol 40:2450–2459
87. Rankin LC, Groom JR, Chopin M, Herold MJ, Walker JA, Mielke LA, McKenzie AN, Carotta S, Nutt SL, Belz GT (2013) The transcription factor T-bet is essential for the development of NKp46+ innate lymphocytes via the Notch pathway. Nat Immunol 14:389–395
88. Res PC, Piskin G, de Boer OJ, van der Loos CM, Teeling P, Bos JD, Teunissen MB (2010) Overrepresentation of IL-17A and IL-22 producing CD8 T cells in lesional skin suggests their involvement in the pathogenesis of psoriasis. PLoS ONE 5:e14108
89. Ricciardi L, Minciullo PL, Saitta S, Trombetta D, Saija A, Gangemi S (2009) Increased serum levels of IL-22 in patients with nickel contact dermatitis. Contact Dermatitis 60:57–58
90. Sabat R (2010) IL-10 family of cytokines. Cytokine Growth Factor Rev 21:315–324
91. Sabat R, Ouyang W, Wolk K (2014) Therapeutic opportunities of the IL-22-IL-22R1 system. Nat Rev Drug Discov 13:21–38
92. Sarkar S, Zhou X, Justa S, Bommireddy SR (2013) Interleukin-22 reduces the severity of collagen-induced arthritis in association with increased levels of interleukin-10. Arthritis Rheum 65:960–971
93. Satoh-Takayama N, Vosshenrich CA, Lesjean-Pottier S, Sawa S, Lochner M, Rattis F, Mention JJ, Thiam K, Cerf-Bensussan N, Mandelboim O et al (2008) Microbial flora drives interleukin 22 production in intestinal NKp46+ cells that provide innate mucosal immune defense. Immunity 29:958–970
94. Schulz SM, Kohler G, Schutze N, Knauer J, Straubinger RK, Chackerian AA, Witte E, Wolk K, Sabat R, Iwakura Y et al (2008) Protective immunity to systemic infection with attenuated Salmonella enterica serovar enteritidis in the absence of IL-12 is associated with IL-23- dependent IL-22, but not IL-17. J Immunol 181:7891–7901
95. Sekikawa A, Fukui H, Suzuki K, Karibe T, Fujii S, Ichikawa K, Tomita S, Imura J, Shiratori K, Chiba T et al (2010) Involvement of the IL-22/REG Ialpha axis in ulcerative colitis. Lab Invest 90:496–505
96. Sestito R, Madonna S, Scarponi C, Cianfarani F, Failla CM, Cavani A, Girolomoni G, Albanesi C (2011) STAT3-dependent effects of IL-22 in human keratinocytes are counter regulated by sirtuin 1 through a direct inhibition of STAT3 acetylation. Faseb J 25:916–927
97. Simonian PL, Wehrmann F, Roark CL, Born WK, O’Brien RL, Fontenot AP (2010) gammadelta T cells protect against lung fibrosis via IL-22. J Exp Med 207:2239–2253
98. Sonnenberg GF, Fouser LA, Artis D (2010) Functional biology of the IL-22-IL-22R pathway in regulating immunity and inflammation at barrier surfaces. Adv Immunol 107:1–29
99. Sonnenberg GF, Fouser LA, Artis D (2011) Border patrol: regulation of immunity, inflammation and tissue homeostasis at barrier surfaces by IL-22. Nat Immunol 12:383–390
100. Sonnenberg GF, Nair MG, Kirn TJ, Zaph C, Fouser LA, Artis D (2010) Pathological versus protective functions of IL-22 in airway inflammation are regulated by IL-17A. J Exp Med 207:1293–1305
101. Sugimoto K, Ogawa A, Mizoguchi E, Shimomura Y, Andoh A, Bhan AK, Blumberg RS, Xavier RJ, Mizoguchi A (2008) IL-22 ameliorates intestinal inflammation in a mouse model of ulcerative colitis. J Clin Invest 118:534–544
102. Tachiiri A, Imamura R, Wang Y, Fukui M, Umemura M, Suda T (2003) Genomic structure and inducible expression of the IL-22 receptor alpha chain in mice. Genes Immun 4:153–159
103. Takahashi K, Hirose K, Kawashima S, Niwa Y, Wakashin H, Iwata A, Tokoyoda K, Renauld JC, Iwamoto I, Nakayama T et al (2011) IL-22 attenuates IL-25 production by lung epithelial cells and inhibits antigen-induced eosinophilic airway inflammation. J Allergy Clin Immunol 128:1067–1076
104. Taube C, Tertilt C, Gyulveszi G, Dehzad N, Kreymborg K, Schneeweiss K, Michel E, Reuter S, Renauld JC, Arnold-Schild D et al (2011) IL-22 is produced by innate lymphoid cells and limits inflammation in allergic airway disease. PLoS ONE 6:e21799
105. Teraki Y, Sakurai A, Izaki S (2013) IL-13/IL-22-coproducing T cells, a novel subset, are increased in atopic dermatitis. J Allergy ClinImmunol 132:971–974
106. Trifari S, Kaplan CD, Tran EH, Crellin NK, Spits H (2009) Identification of a human helper T cell population that has abundant production of interleukin 22 and is distinct from T(H)-17, T(H)1 and T(H)2 cells. Nat Immunol 10:864–871
107. Truchetet ME, Brembilla NC, Montanari E, Allanore Y, Chizzolini C (2011) Increased frequency of circulating Th22 in addition to Th17 and Th2 lymphocytes in systemic sclerosis: association with interstitial lung disease. Arthritis Res Ther 13:R166
108. Veldhoen M, Hirota K, Westendorf AM, Buer J, Dumoutier L, Renauld JC, Stockinger B (2008) The aryl hydrocarbon receptor links TH17-cell-mediated autoimmunity to environmental toxins. Nature 453:106–109
109. Wang B, Zhao XP, Fan YC, Zhang JJ, Zhao J, Wang K (2013) IL-17A but not IL-22 suppresses the replication of hepatitis B virus mediated by over-expression of MxA and OAS mRNA in the HepG2.2.15 cell line. Antiviral Res 97:285–292
110. Wang M, Tan Z, Zhang R, Kotenko SV, Liang P (2002) Interleukin 24 (MDA-7/MOB-5) signals through two heterodimeric receptors, IL-22R1/IL-20R2 and IL-20R1/IL-20R2. J BiolChem 277:7341–7347
111. Wehkamp J, Fellermann K, Herrlinger KR, Baxmann S, Schmidt K, Schwind B, Duchrow M, Wohlschlager C, Feller AC, Stange EF (2002) Human beta-defensin 2 but not betadefensin 1 is expressed preferentially in colonic mucosa of inflammatory bowel disease. Eur J GastroenterolHepatol 14:745–752
112. Wei CC, Ho TW, Liang WG, Chen GY, Chang MS (2003) Cloning and characterization of mouse IL-22 binding protein. Genes Immun 4:204–211
113. Weiss B, Wolk K, Grunberg BH, Volk HD, Sterry W, Asadullah K, Sabat R (2004) Cloning of murine IL-22 receptor alpha 2 and comparison with its human counterpart. Genes Immun 5:330–336
114. Wilson MS, Feng CG, Barber DL, Yarovinsky F, Cheever AW, Sher A, Grigg M, Collins M, Fouser L, Wynn TA (2010) Redundant and pathogenic roles for IL-22 in mycobacterial, protozoan, and helminth infections. J Immunol 184:4378–4390
115. Wolk K, Haugen HS, Xu W, Witte E, Waggie K, Anderson M, Vom BE, Witte K, Warszawska K, Philipp S et al (2009) IL-22 and IL-20 are key mediators of the epidermal alterations in psoriasis while IL-17 and IFN-gamma are not. J Mol Med (Berl) 87:523–536
116. Wolk K, Kunz S, Witte E, Friedrich M, Asadullah K, Sabat R (2004) IL-22 increases the innate immunity of tissues. Immunity 21:241–254
117. Wolk K, Witte E, Hoffmann U, Doecke WD, Endesfelder S, Asadullah K, Sterry W, Volk HD, Wittig BM, Sabat R (2007) IL-22 induces lipopolysaccharide-binding protein in hepatocytes: a potential systemic role of IL-22 in Crohn’s disease. J Immunol 178:5973–5981
118. Wolk K, Witte E, Reineke U, Witte K, Friedrich M, Sterry W, Asadullah K, Volk HD, Sabat R (2005) Is there an interaction between interleukin-10 and interleukin-22? Genes Immun 6:8–18
119. Wolk K, Witte E, Wallace E, Docke WD, Kunz S, Asadullah K, Volk HD, Sterry W, Sabat R (2006) IL-22 regulates the expression of genes responsible for antimicrobial defense, cellular differentiation, and mobility in keratinocytes: a potential role in psoriasis. Eur J Immunol 36:1309–1323
120. Wolk K, Sabat R (2006) Interleukin-22: a novel T- and NK-cell derived cytokine that regulates the biology of tissue cells. Cytokine Growth Factor Rev 17:367–380
121. Wu PW, Li J, Kodangattil SR, Luxenberg DP, Bennett F, Martino M, Collins M, Dunussi- Joannopoulos K, Gill DS, Wolfman NM et al (2008) IL-22R, IL-10R2, and IL-22BP binding sites are topologically juxtaposed on adjacent and overlapping surfaces of IL-22. J MolBiol 382:1168–1183
122. Xie MH, Aggarwal S, Ho WH, Foster J, Zhang Z, Stinson J, Wood WI, Goddard AD, Gurney AL (2000) Interleukin (IL)-22, a novel human cytokine that signals through the interferon receptor-related proteins CRF2-4 and IL-22R. J BiolChem 275:31335–31339
123. Xing WW, Zou MJ, Liu S, Xu T, Wang JX, Xu DG (2011) Interleukin-22 protects against acute alcohol-induced hepatotoxicity in mice. BiosciBiotechnolBiochem 75:1290–1294
124. Xu T, Logsdon NJ, Walter MR (2005) Structure of insect-cell-derived IL-22. Acta Crystallogr D Biol Crystallogr 61:942–950
125. Xu W, Presnell SR, Parrish-Novak J, Kindsvogel W, Jaspers S, Chen Z, Dillon SR, Gao Z, Gilbert T, Madden K et al (2001) A soluble class II cytokine receptor, IL-22RA2, is a naturally occurring IL-22 antagonist. Proc Natl Acad Sci USA 98:9511–9516
126. Xue J, Nguyen DT, Habtezion A (2012) Aryl hydrocarbon receptor regulates pancreatic IL-22 production and protects mice from acute pancreatitis. Gastroenterology 143:1670–1680
127. Yang XO, Pappu BP, Nurieva R, Akimzhanov A, Kang HS, Chung Y, Ma L, Shah B, Panopoulos AD, Schluns KS et al (2008) T helper 17 lineage differentiation is programmed by orphan nuclear receptors ROR alpha and ROR gamma. Immunity 28:29–39
128. Yoon SI, Jones BC, Logsdon NJ, Harris BD, Deshpande A, Radaeva S, Halloran BA, Gao B, Walter MR (2010) Structure and mechanism of receptor sharing by the IL-10R2 common chain. Structure 18:638–648
129. Zenewicz LA, Yancopoulos GD, Valenzuela DM, Murphy AJ, Karow M, Flavell RA (2007) Interleukin-22 but not interleukin-17 provides protection to hepatocytes during acute liver inflammation. Immunity 27:647–659
130. Zhang L, Li JM, Liu XG, Ma DX, Hu NW, Li YG, Li W, Hu Y, Yu S, Qu X et al (2011) Elevated Th22 cells correlated with Th17 cells in patients with rheumatoid arthritis. J Clin Immunol 31:606–614
131. Zhang Y, Cobleigh MA, Lian JQ, Huang CX, Booth CJ, Bai XF, Robek MD (2011) A proinflammatory role for interleukin-22 in the immune response to hepatitis B virus. Gastroenterology 141:1897–1906
132. Zheng Y, Valdez PA, Danilenko DM, Hu Y, Sa SM, Gong Q, Abbas AR, Modrusan Z, Ghilardi N, de Sauvage FJ et al (2008) Interleukin-22 mediates early host defense against attaching and effacing bacterial pathogens. Nat Med 14:282–289
133. Zhuang Y, Peng LS, Zhao YL, Shi Y, Mao XH, Guo G, Chen W, Liu XF, Zhang JY, Liu T et al (2012) Increased intratumoral IL-22-producing CD4(+) T cells and Th22 cells correlate with gastric cancer progression and predict poor patient survival. Cancer ImmunolImmunother 61:1965–1975