Skip to main content

Advertisement

Springer Nature Link
Log in

The transcription factor network in Th9 cells

  • Review
  • Published:
Seminars in Immunopathology Aims and scope Submit manuscript

Abstract

The development of T helper cell subsets requires activated T cells that respond to a polarizing cytokine environment resulting in the activation and expression of transcription factors. The subset-specific transcription factors bind and induce the production of specific effector cytokines. Th9 cells express IL-9 and develop in the presence of TGFβ, IL-4, and IL-2. Each of these cytokines activates signaling pathways that are required for Th9 differentiation and IL-9 production. In this review, I summarize what is currently understood about the signaling pathways and transcription factors that promote the Th9 genetic program, providing some perspective for the integration of the signals in regulating the Il9 gene and dictating the expression of other Th9-associated genes. I highlight how experiments in mouse cells have established a transcriptional network that is conserved in human T cells and set the stage toward defining the next important questions for a more detailed understanding of Th9 cell development and function.

This is a preview of subscription content, log in via an institution to check access.

Access this article

Log in via an institution

Subscribe and save

Springer+ Basic
$34.99 /Month
  • Get 10 units per month
  • Download Article/Chapter or eBook
  • 1 Unit = 1 Article or 1 Chapter
  • Cancel anytime
Subscribe now

Buy Now

Price excludes VAT (USA)
Tax calculation will be finalised during checkout.

Instant access to the full article PDF.

Institutional subscriptions

Fig. 1
Fig. 2

Similar content being viewed by others

References

  1. Kaplan MH, Hufford MM, Olson MR (2015) The development and in vivo function of T helper 9 cells. Nat Rev Immunol 15:295–307

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  2. Agarwal S, Rao A (1998) Modulation of chromatin structure regulates cytokine gene expression during T cell differentiation. Immunity 9:765–775

    Article  CAS  PubMed  Google Scholar 

  3. Avni O, Lee D, Macian F, Szabo SJ, Glimcher LH et al (2002) T(H) cell differentiation is accompanied by dynamic changes in histone acetylation of cytokine genes. Nat Immunol 3:643–651

    CAS  PubMed  Google Scholar 

  4. Bird JJ, Brown DR, Mullen AC, Moskowitz NH, Mahowald MA et al (1998) Helper T cell differentiation is controlled by the cell cycle. Immunity 9:229–237

    Article  CAS  PubMed  Google Scholar 

  5. Murphy KM, Reiner SL (2002) The lineage decisions of helper T cells. Nat Rev Immunol 2:933–944

    Article  CAS  PubMed  Google Scholar 

  6. Grogan JL, Mohrs M, Harmon B, Lacy DA, Sedat JW et al (2001) Early transcription and silencing of cytokine genes underlie polarization of T helper cell subsets. Immunity 14:205–215

    Article  CAS  PubMed  Google Scholar 

  7. Stritesky GL, Yeh N, Kaplan MH (2008) IL-23 mediates stability but not commitment to the Th17 lineage. J Immunol 181:5948–5955

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  8. Ghoreschi K, Laurence A, Yang XP, Tato CM, McGeachy MJ et al (2010) Generation of pathogenic T(H)17 cells in the absence of TGF-beta signalling. Nature 467:967–971

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  9. McGeachy MJ, Bak-Jensen KS, Chen Y, Tato CM, Blumenschein W et al (2007) TGF-beta and IL-6 drive the production of IL-17 and IL-10 by T cells and restrain T(H)-17 cell-mediated pathology. Nat Immunol 8:1390–1397

    Article  CAS  PubMed  Google Scholar 

  10. Tan C, Aziz MK, Lovaas JD, Vistica BP, Shi G et al (2010) Antigen-specific Th9 cells exhibit uniqueness in their kinetics of cytokine production and short retention at the inflammatory site. J Immunol 185:6795–6801

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  11. Perumal NB, Kaplan MH (2011) Regulating Il9 transcription in T helper cells. Trends Immunol 32:146–150

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  12. Chang HC, Sehra S, Goswami R, Yao W, Yu Q et al (2010) The transcription factor PU.1 is required for the development of IL-9-producing T cells and allergic inflammation. Nat Immunol 11:527–534

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  13. O'Shea JJ, Lahesmaa R, Vahedi G, Laurence A, Kanno Y (2011) Genomic views of STAT function in CD4+ T helper cell differentiation. Nat Rev Immunol 11:239–250

    Article  PubMed  PubMed Central  Google Scholar 

  14. O'Shea JJ, Plenge R (2012) JAK and STAT signaling molecules in immunoregulation and immune-mediated disease. Immunity 36:542–550

    Article  PubMed  PubMed Central  Google Scholar 

  15. Wei G, Wei L, Zhu J, Zang C, Hu-Li J et al (2009) Global mapping of H3K4me3 and H3K27me3 reveals specificity and plasticity in lineage fate determination of differentiating CD4+ T cells. Immunity 30:155–167

    Article  PubMed  PubMed Central  Google Scholar 

  16. Wei L, Vahedi G, Sun HW, Watford WT, Takatori H et al (2010) Discrete roles of STAT4 and STAT6 transcription factors in tuning epigenetic modifications and transcription during T helper cell differentiation. Immunity 32:840–851

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  17. Vahedi G, Takahashi H, Nakayamada S, Sun HW, Sartorelli V et al (2012) STATs shape the active enhancer landscape of T cell populations. Cell 151:981–993

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  18. Pham D, Yu Q, Walline CC, Muthukrishnan R, Blum JS et al (2013) Opposing roles of STAT4 and Dnmt3a in Th1 gene regulation. J Immunol 191:902–911

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  19. Yu Q, Thieu VT, Kaplan MH (2007) Stat4 limits DNA methyltransferase recruitment and DNA methylation of the IL-18Ralpha gene during Th1 differentiation. EMBO J 26:2052–2060

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  20. Yu Q, Zhou B, Zhang Y, Nguyen ET, Du J et al (2012) DNA methyltransferase 3a limits the expression of interleukin-13 in T helper 2 cells and allergic airway inflammation. Proc Natl Acad Sci U S A 109:541–546

    Article  CAS  PubMed  Google Scholar 

  21. Elo LL, Jarvenpaa H, Tuomela S, Raghav S, Ahlfors H et al (2010) Genome-wide profiling of interleukin-4 and STAT6 transcription factor regulation of human Th2 cell programming. Immunity 32:852–862

    Article  CAS  PubMed  Google Scholar 

  22. Lee DU, Agarwal S, Rao A (2002) Th2 lineage commitment and efficient IL-4 production involves extended demethylation of the IL-4 gene. Immunity 16:649–660

    Article  CAS  PubMed  Google Scholar 

  23. Dardalhon V, Awasthi A, Kwon H, Galileos G, Gao W et al (2008) IL-4 inhibits TGF-beta-induced Foxp3+ T cells and, together with TGF-beta, generates IL-9+ IL-10+ Foxp3(−) effector T cells. Nat Immunol 9:1347–1355

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  24. Goswami R, Jabeen R, Yagi R, Pham D, Zhu J et al (2012) STAT6-dependent regulation of Th9 development. J Immunol 188:968–975

    Article  CAS  PubMed  Google Scholar 

  25. Veldhoen M, Uyttenhove C, van Snick J, Helmby H, Westendorf A et al (2008) Transforming growth factor-beta ‘reprograms’ the differentiation of T helper 2 cells and promotes an interleukin 9-producing subset. Nat Immunol 9:1341–1346

    Article  CAS  PubMed  Google Scholar 

  26. Angkasekwinai P, Chang SH, Thapa M, Watarai H, Dong C (2010) Regulation of IL-9 expression by IL-25 signaling. Nat Immunol 11:250–256

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  27. Jabeen R, Goswami R, Awe O, Kulkarni A, Nguyen ET et al (2013) Th9 cell development requires a BATF-regulated transcriptional network. J Clin Invest 123:4641–4653

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  28. Schmitt E, Germann T, Goedert S, Hoehn P, Huels C et al (1994) IL-9 production of naive CD4+ T cells depends on IL-2, is synergistically enhanced by a combination of TGF-beta and IL-4, and is inhibited by IFN-gamma. J Immunol 153:3989–3996

    CAS  PubMed  Google Scholar 

  29. Gomez-Rodriguez J, Meylan F, Handon R, Hayes ET, Anderson SM et al (2016) Itk is required for Th9 differentiation via TCR-mediated induction of IL-2 and IRF4. Nat Commun 7:10857

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  30. Niedbala W, Besnard AG, Nascimento DC, Donate PB, Sonego F et al (2014) Nitric oxide enhances Th9 cell differentiation and airway inflammation. Nat Commun 5:4575

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  31. Richard AC, Tan C, Hawley ET, Gomez-Rodriguez J, Goswami R et al (2015) The TNF-family ligand TL1A and its receptor DR3 promote T cell-mediated allergic immunopathology by enhancing differentiation and pathogenicity of IL-9-producing T cells. J Immunol 194:3567–3582

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  32. Bassil R, Orent W, Olah M, Kurdi AT, Frangieh M et al (2014) BCL6 controls Th9 cell development by repressing Il9 transcription. J Immunol 193:198–207

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  33. Liao W, Spolski R, Li P, Du N, West EE et al (2014) Opposing actions of IL-2 and IL-21 on Th9 differentiation correlate with their differential regulation of BCL6 expression. Proc Natl Acad Sci U S A 111:3508–3513

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  34. Yao W, Zhang Y, Jabeen R, Nguyen ET, Wilkes DS et al (2013) Interleukin-9 is required for allergic airway inflammation mediated by the cytokine TSLP. Immunity 38:360–372

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  35. Liao W, Schones DE, Oh J, Cui Y, Cui K et al (2008) Priming for T helper type 2 differentiation by interleukin 2-mediated induction of interleukin 4 receptor alpha-chain expression. Nat Immunol 9:1288–1296

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  36. Olson MR, Verdan FF, Hufford MM, Dent AL, Kaplan MH (2016) STAT3 impairs STAT5 activation in the development of IL-9-secreting T cells. J Immunol 196:3297–3304

    Article  CAS  PubMed  Google Scholar 

  37. Murugaiyan G, Beynon V, Pires Da Cunha A, Joller N, Weiner HL (2012) IFN-gamma limits Th9-mediated autoimmune inflammation through dendritic cell modulation of IL-27. J Immunol 189:5277–5283

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  38. Vegran F, Berger H, Boidot R, Mignot G, Bruchard M et al (2014) The transcription factor IRF1 dictates the IL-21-dependent anticancer functions of TH9 cells. Nat Immunol 15:758–766

    Article  CAS  PubMed  Google Scholar 

  39. Wang A, Pan D, Lee YH, Martinez GJ, Feng XH et al (2013) Cutting edge: Smad2 and Smad4 regulate TGF-beta-mediated Il9 gene expression via EZH2 displacement. J Immunol 191:4908–4912

    Article  CAS  PubMed  Google Scholar 

  40. Tamiya T, Ichiyama K, Kotani H, Fukaya T, Sekiya T et al (2013) Smad2/3 and IRF4 play a cooperative role in IL-9-producing T cell induction. J Immunol 191:2360–2371

    Article  CAS  PubMed  Google Scholar 

  41. Ebel ME, Kansas GS (2016) Functions of Smad transcription factors in TGF-beta1-induced selectin ligand expression on murine CD4 Th cells. J Immunol 197:2627–2634

  42. Elyaman W, Bassil R, Bradshaw EM, Orent W, Lahoud Y et al (2012) Notch receptors and Smad3 signaling cooperate in the induction of interleukin-9-producing T cells. Immunity 36:623–634

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  43. Nakatsukasa H, Zhang D, Maruyama T, Chen H, Cui K et al (2015) The DNA-binding inhibitor Id3 regulates IL-9 production in CD4(+) T cells. Nat Immunol 16:1077–1084

    Article  CAS  PubMed  Google Scholar 

  44. Pham D, Walline CC, Hollister K, Dent AL, Blum JS et al (2013) The transcription factor Twist1 limits T helper 17 and T follicular helper cell development by repressing the gene encoding the interleukin-6 receptor alpha chain. J Biol Chem 288:27423–27433

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  45. Wang Y, Bi Y, Chen X, Li C, Li Y et al (2016) Histone deacetylase SIRT1 negatively regulates the differentiation of interleukin-9-producing CD4(+) T cells. Immunity 44:1337–1349

    Article  CAS  PubMed  Google Scholar 

  46. Singh Y, Garden OA, Lang F, Cobb BS (2016) MicroRNAs regulate T-cell production of interleukin-9 and identify hypoxia-inducible factor-2alpha as an important regulator of T helper 9 and regulatory T-cell differentiation. Immunology 149:74–86

    Article  CAS  PubMed  Google Scholar 

  47. Waickman AT, Powell JD (2012) mTOR, metabolism, and the regulation of T-cell differentiation and function. Immunol Rev 249:43–58

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  48. Goswami R, Kaplan MH (2012) Gcn5 is required for PU.1-dependent interleukin-9 (IL-9) induction in Th9 cells. J Immunol 189:3026–3033

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  49. Ramming A, Druzd D, Leipe J, Schulze-Koops H, Skapenko A (2012) Maturation-related histone modifications in the PU.1 promoter regulate Th9-cell development. Blood 119:4665–4674

    Article  CAS  PubMed  Google Scholar 

  50. Gerlach K, Hwang Y, Nikolaev A, Atreya R, Dornhoff H et al (2014) TH9 cells that express the transcription factor PU.1 drive T cell-mediated colitis via IL-9 receptor signaling in intestinal epithelial cells. Nat Immunol 15:676–686

    Article  CAS  PubMed  Google Scholar 

  51. Sehra S, Yao W, Nguyen ET, Glosson-Byers NL, Akhtar N et al (2015) TH9 cells are required for tissue mast cell accumulation during allergic inflammation. J Allergy Clin Immunol 136:433–440

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  52. Ahyi AN, Chang HC, Dent AL, Nutt SL, Kaplan MH (2009) IFN regulatory factor 4 regulates the expression of a subset of Th2 cytokines. J Immunol 183:1598–1606

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  53. Chang HC, Han L, Jabeen R, Carotta S, Nutt SL et al (2009) PU.1 regulates TCR expression by modulating GATA-3 activity. J Immunol 183:4887–4894

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  54. Chang HC, Zhang S, Thieu VT, Slee RB, Bruns HA et al (2005) PU.1 expression delineates heterogeneity in primary Th2 cells. Immunity 22:693–703

    Article  CAS  PubMed  Google Scholar 

  55. Awe O, Hufford MM, Wu H, Pham D, Chang HC et al (2015) PU.1 expression in T follicular helper cells limits CD40L-dependent germinal center B cell development. J Immunol 195:3705–3715

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  56. Koh B, Hufford MM, Pham D, Olson MR, Wu T et al (2016) The ETS family transcription factors Etv5 and PU.1 function in parallel to promote Th9 cell development. J Immunol 197:2465–2472

    Article  CAS  PubMed  Google Scholar 

  57. Pham D, Sehra S, Sun X, Kaplan MH (2014) The transcription factor Etv5 controls TH17 cell development and allergic airway inflammation. J Allergy Clin Immunol 134:204–214

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  58. Hollenhorst PC, McIntosh LP, Graves BJ (2011) Genomic and biochemical insights into the specificity of ETS transcription factors. Annu Rev Biochem 80:437–471

    Article  CAS  PubMed  Google Scholar 

  59. Staudt V, Bothur E, Klein M, Lingnau K, Reuter S et al (2010) Interferon-regulatory factor 4 is essential for the developmental program of T helper 9 cells. Immunity 33:192–202

    Article  CAS  PubMed  Google Scholar 

  60. Ubel C, Sopel N, Graser A, Hildner K, Reinhardt C et al (2014) The activating protein 1 transcription factor basic leucine zipper transcription factor, ATF-like (BATF), regulates lymphocyte- and mast cell-driven immune responses in the setting of allergic asthma. J Allergy Clin Immunol 133(198–206):e191–e199

    Google Scholar 

  61. Brustle A, Heink S, Huber M, Rosenplanter C, Stadelmann C et al (2007) The development of inflammatory T(H)-17 cells requires interferon-regulatory factor 4. Nat Immunol 8:958–966

    Article  PubMed  Google Scholar 

  62. Lohoff M, Mittrucker HW, Prechtl S, Bischof S, Sommer F et al (2002) Dysregulated T helper cell differentiation in the absence of interferon regulatory factor 4. Proc Natl Acad Sci U S A 99:11808–11812

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  63. Ise W, Kohyama M, Schraml BU, Zhang T, Schwer B et al (2011) The transcription factor BATF controls the global regulators of class-switch recombination in both B cells and T cells. Nat Immunol 12:536–543

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  64. Schraml BU, Hildner K, Ise W, Lee WL, Smith WA et al (2009) The AP-1 transcription factor Batf controls T(H)17 differentiation. Nature 460:405–409

    CAS  PubMed  PubMed Central  Google Scholar 

  65. Betz BC, Jordan-Williams KL, Wang C, Kang SG, Liao J et al (2010) Batf coordinates multiple aspects of B and T cell function required for normal antibody responses. J Exp Med 207:933–942

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  66. Kwon H, Thierry-Mieg D, Thierry-Mieg J, Kim HP, Oh J et al (2009) Analysis of interleukin-21-induced Prdm1 gene regulation reveals functional cooperation of STAT3 and IRF4 transcription factors. Immunity 31:941–952

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  67. Li P, Spolski R, Liao W, Wang L, Murphy TL et al (2012) BATF-JUN is critical for IRF4-mediated transcription in T cells. Nature 490:543–546

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  68. Glasmacher E, Agrawal S, Chang AB, Murphy TL, Zeng W et al (2012) A genomic regulatory element that directs assembly and function of immune-specific AP-1-IRF complexes. Science 338:975–980

    Article  CAS  PubMed  Google Scholar 

  69. Ciofani M, Madar A, Galan C, Sellars M, Mace K et al (2012) A validated regulatory network for Th17 cell specification. Cell 151:289–303

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  70. Jash A, Sahoo A, Kim GC, Chae CS, Hwang JS et al (2012) Nuclear factor of activated T cells 1 (NFAT1)-induced permissive chromatin modification facilitates nuclear factor-kappaB (NF-kappaB)-mediated interleukin-9 (IL-9) transactivation. J Biol Chem 287:15445–15457

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  71. Koch S, Graser A, Mirzakhani H, Zimmermann T, Melichar VO et al (2016) Increased expression of nuclear factor of activated T cells 1 drives IL-9-mediated allergic asthma. J Allergy Clin Immunol 137:1898–1902

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  72. Xiao X, Balasubramanian S, Liu W, Chu X, Wang H et al (2012) OX40 signaling favors the induction of T(H)9 cells and airway inflammation. Nat Immunol 13:981–990

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  73. Kim IK, Kim BS, Koh CH, Seok JW, Park JS et al (2015) Glucocorticoid-induced tumor necrosis factor receptor-related protein co-stimulation facilitates tumor regression by inducing IL-9-producing helper T cells. Nat Med 21:1010–1017

    Article  CAS  PubMed  Google Scholar 

  74. Xiao X, Shi X, Fan Y, Zhang X, Wu M et al (2015) GITR subverts Foxp3(+) Tregs to boost Th9 immunity through regulation of histone acetylation. Nat Commun 6:8266

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  75. O'Shea JJ, Paul WE (2010) Mechanisms underlying lineage commitment and plasticity of helper CD4+ T cells. Science 327:1098–1102

    Article  PubMed  PubMed Central  Google Scholar 

  76. Zhou L, Chong MM, Littman DR (2009) Plasticity of CD4+ T cell lineage differentiation. Immunity 30:646–655

    Article  CAS  PubMed  Google Scholar 

  77. Stritesky GL, Muthukrishnan R, Sehra S, Goswami R, Pham D et al (2011) The transcription factor STAT3 is required for Th2 cell development. Immunity 34:39–49

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  78. Glosson-Byers NL, Sehra S, Stritesky GL, Yu Q, Awe O et al (2014) Th17 cells demonstrate stable cytokine production in a proallergic environment. J Immunol 193:2631–2640

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  79. Xiao X, Shi X, Fan Y, Wu C, Zhang X et al (2016) The costimulatory receptor OX40 inhibits interleukin-17 expression through activation of repressive chromatin remodeling pathways. Immunity 44:1271–1283

    Article  CAS  PubMed  Google Scholar 

  80. Laurence A, Tato CM, Davidson TS, Kanno Y, Chen Z et al (2007) Interleukin-2 signaling via STAT5 constrains T helper 17 cell generation. Immunity 26:371–381

    Article  CAS  PubMed  Google Scholar 

  81. Lee SH, Chang HS, Jang AS, Park SW, Park JS et al (2011) The association of a single-nucleotide polymorphism of the IL-2 inducible T-cell kinase gene with asthma. Ann Hum Genet 75:359–369

    Article  CAS  PubMed  Google Scholar 

  82. Becker KL, Rosler B, Wang X, Lachmandas E, Kamsteeg M, et al. (2016) Th2 and Th9 responses in patients with chronic mucocutaneous candidiasis and hyper IgE syndrome. Clin Exp Allergy

  83. Yao W, Tepper RS, Kaplan MH (2011) Predisposition to the development of IL-9-secreting T cells in atopic infants. J Allergy Clin Immunol 128:1357–1360

    Article  CAS  PubMed  PubMed Central  Google Scholar 

Download references

Acknowledgments

This work was supported by Public Health Service grants R01 AI057459 and R03 AI101628 to MHK. Support provided by the Herman B Wells Center was in part from the Riley Children’s Foundation. I thank Drs. R. Nicholas Laribee, Geoffrey Kansas, and Matthew Olson for comments on this review.

Author information

Authors and Affiliations

  1. Department of Pediatrics and Herman B Wells Center for Pediatric Research, Department of Microbiology and Immunology, Indiana University School of Medicine, 1044 West Walnut St #202, Indianapolis, IN, 46202, USA

    Mark H. Kaplan

Authors
  1. Mark H. Kaplan
    View author publications

    You can also search for this author inPubMed Google Scholar

Corresponding author

Correspondence to Mark H. Kaplan.

Additional information

This article is a contribution to the special issue on Th9 Cells in Immunity and Immunopathological Diseases - Guest Editors: Mark Kaplan and Markus Neurath

Rights and permissions

Reprints and permissions

About this article

Check for updates. Verify currency and authenticity via CrossMark

Cite this article

Kaplan, M.H. The transcription factor network in Th9 cells. Semin Immunopathol 39, 11–20 (2017). https://doi.org/10.1007/s00281-016-0600-2

Download citation

  • Received:

  • Accepted:

  • Published:

  • Issue Date:

  • DOI: https://doi.org/10.1007/s00281-016-0600-2

Keywords