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Biology and regulation of IL-2: from molecular mechanisms to human therapy.

Spolski R, Li P, Leonard WJ.

Nat Rev Immunol. 2018 Aug 8. doi: 10.1038/s41577-018-0046-y. [Epub ahead of print] Review.


Defective IgA response to atypical intestinal commensals in IL-21 receptor deficiency reshapes immune cell homeostasis and mucosal immunity.

Cho H, Jaime H, de Oliveira RP, Kang B, Spolski R, Vaziri T, Myers TG, Thovarai V, Shen Z, Fox JG, Leonard WJ, Kelsall BL.

Mucosal Immunol. 2018 Aug 7. doi: 10.1038/s41385-018-0056-x. [Epub ahead of print]


The γ c family of cytokines: fine-tuning signals from IL-2 and IL-21 in the regulation of the immune response.

Spolski R, Gromer D, Leonard WJ.

F1000Res. 2017 Oct 23;6:1872. doi: 10.12688/f1000research.12202.1. eCollection 2017. Review.


Critical functions for STAT5 tetramers in the maturation and survival of natural killer cells.

Lin JX, Du N, Li P, Kazemian M, Gebregiorgis T, Spolski R, Leonard WJ.

Nat Commun. 2017 Nov 6;8(1):1320. doi: 10.1038/s41467-017-01477-5.


STAT5-mediated chromatin interactions in superenhancers activate IL-2 highly inducible genes: Functional dissection of the Il2ra gene locus.

Li P, Mitra S, Spolski R, Oh J, Liao W, Tang Z, Mo F, Li X, West EE, Gromer D, Lin JX, Liu C, Ruan Y, Leonard WJ.

Proc Natl Acad Sci U S A. 2017 Nov 14;114(46):12111-12119. doi: 10.1073/pnas.1714019114. Epub 2017 Oct 24.


A TSLP-complement axis mediates neutrophil killing of methicillin-resistant Staphylococcus aureus.

West EE, Spolski R, Kazemian M, Yu ZX, Kemper C, Leonard WJ.

Sci Immunol. 2016 Nov 18;1(5). pii: eaaf8471. doi: 10.1126/sciimmunol.aaf8471.


Interleukin-21 signaling in B cells, but not in T cells, is indispensable for the development of collagen-induced arthritis in mice.

Sakuraba K, Oyamada A, Fujimura K, Spolski R, Iwamoto Y, Leonard WJ, Yoshikai Y, Yamada H.

Arthritis Res Ther. 2016 Aug 17;18:188. doi: 10.1186/s13075-016-1086-y.


Cutting Edge: IL-4, IL-21, and IFN-γ Interact To Govern T-bet and CD11c Expression in TLR-Activated B Cells.

Naradikian MS, Myles A, Beiting DP, Roberts KJ, Dawson L, Herati RS, Bengsch B, Linderman SL, Stelekati E, Spolski R, Wherry EJ, Hunter C, Hensley SE, Leonard WJ, Cancro MP.

J Immunol. 2016 Aug 15;197(4):1023-8. doi: 10.4049/jimmunol.1600522. Epub 2016 Jul 18.


T helper 1 immunity requires complement-driven NLRP3 inflammasome activity in CD4⁺ T cells.

Arbore G, West EE, Spolski R, Robertson AAB, Klos A, Rheinheimer C, Dutow P, Woodruff TM, Yu ZX, O'Neill LA, Coll RC, Sher A, Leonard WJ, Köhl J, Monk P, Cooper MA, Arno M, Afzali B, Lachmann HJ, Cope AP, Mayer-Barber KD, Kemper C.

Science. 2016 Jun 17;352(6292):aad1210. doi: 10.1126/science.aad1210.


BACH2 regulates CD8(+) T cell differentiation by controlling access of AP-1 factors to enhancers.

Roychoudhuri R, Clever D, Li P, Wakabayashi Y, Quinn KM, Klebanoff CA, Ji Y, Sukumar M, Eil RL, Yu Z, Spolski R, Palmer DC, Pan JH, Patel SJ, Macallan DC, Fabozzi G, Shih HY, Kanno Y, Muto A, Zhu J, Gattinoni L, O'Shea JJ, Okkenhaug K, Igarashi K, Leonard WJ, Restifo NP.

Nat Immunol. 2016 Jul;17(7):851-860. doi: 10.1038/ni.3441. Epub 2016 May 9.


T Follicular Helper Cell Plasticity Shapes Pathogenic T Helper 2 Cell-Mediated Immunity to Inhaled House Dust Mite.

Ballesteros-Tato A, Randall TD, Lund FE, Spolski R, Leonard WJ, León B.

Immunity. 2016 Feb 16;44(2):259-73. doi: 10.1016/j.immuni.2015.11.017. Epub 2016 Jan 26.


Endothelial interleukin-21 receptor up-regulation in peripheral artery disease.

Wang T, Cunningham A, Houston K, Sharma AM, Chen L, Dokun AO, Lye RJ, Spolski R, Leonard WJ, Annex BH.

Vasc Med. 2016 Apr;21(2):99-104. doi: 10.1177/1358863X15621798. Epub 2015 Dec 24.


IL-21-mediated non-canonical pathway for IL-1β production in conventional dendritic cells.

Wan CK, Li P, Spolski R, Oh J, Andraski AB, Du N, Yu ZX, Dillon CP, Green DR, Leonard WJ.

Nat Commun. 2015 Aug 13;6:7988. doi: 10.1038/ncomms8988.


Comprehensive assembly of novel transcripts from unmapped human RNA-Seq data and their association with cancer.

Kazemian M, Ren M, Lin JX, Liao W, Spolski R, Leonard WJ.

Mol Syst Biol. 2015 Aug 7;11(8):826. doi: 10.15252/msb.156172.


Opposing roles of STAT1 and STAT3 in IL-21 function in CD4+ T cells.

Wan CK, Andraski AB, Spolski R, Li P, Kazemian M, Oh J, Samsel L, Swanson PA 2nd, McGavern DB, Sampaio EP, Freeman AF, Milner JD, Holland SM, Leonard WJ.

Proc Natl Acad Sci U S A. 2015 Jul 28;112(30):9394-9. doi: 10.1073/pnas.1511711112. Epub 2015 Jul 13.


Possible Human Papillomavirus 38 Contamination of Endometrial Cancer RNA Sequencing Samples in The Cancer Genome Atlas Database.

Kazemian M, Ren M, Lin JX, Liao W, Spolski R, Leonard WJ.

J Virol. 2015 Sep;89(17):8967-73. doi: 10.1128/JVI.00822-15. Epub 2015 Jun 17.


Interleukin-2 activity can be fine tuned with engineered receptor signaling clamps.

Mitra S, Ring AM, Amarnath S, Spangler JB, Li P, Ju W, Fischer S, Oh J, Spolski R, Weiskopf K, Kohrt H, Foley JE, Rajagopalan S, Long EO, Fowler DH, Waldmann TA, Garcia KC, Leonard WJ.

Immunity. 2015 May 19;42(5):826-38. doi: 10.1016/j.immuni.2015.04.018.


Loss of interleukin-21 receptor activation in hypoxic endothelial cells impairs perfusion recovery after hindlimb ischemia.

Wang T, Cunningham A, Dokun AO, Hazarika S, Houston K, Chen L, Lye RJ, Spolski R, Leonard WJ, Annex BH.

Arterioscler Thromb Vasc Biol. 2015 May;35(5):1218-25. doi: 10.1161/ATVBAHA.115.305476. Epub 2015 Apr 2.


Complex interactions of transcription factors in mediating cytokine biology in T cells.

Li P, Spolski R, Liao W, Leonard WJ.

Immunol Rev. 2014 Sep;261(1):141-56. doi: 10.1111/imr.12199. Review.


Interleukin-21: a double-edged sword with therapeutic potential.

Spolski R, Leonard WJ.

Nat Rev Drug Discov. 2014 May;13(5):379-95. doi: 10.1038/nrd4296. Epub 2014 Apr 22. Review.


Thymic stromal lymphopoietin and interleukin-4 mediate the pathogenesis of halothane-induced liver injury in mice.

Proctor WR, Chakraborty M, Fullerton AM, Korrapati MC, Ryan PM, Semple K, Morrison JC, Berkson JD, Chea LS, Yang Q, Li AP, Spolski R, West EE, Rochman Y, Leonard WJ, Bourdi M, Pohl LR.

Hepatology. 2014 Nov;60(5):1741-52. doi: 10.1002/hep.27169. Epub 2014 May 20.


Opposing actions of IL-2 and IL-21 on Th9 differentiation correlate with their differential regulation of BCL6 expression.

Liao W, Spolski R, Li P, Du N, West EE, Ren M, Mitra S, Leonard WJ.

Proc Natl Acad Sci U S A. 2014 Mar 4;111(9):3508-13. doi: 10.1073/pnas.1301138111. Epub 2014 Feb 18.


Interleukin-21 is a critical cytokine for the generation of virus-specific long-lived plasma cells.

Rasheed MA, Latner DR, Aubert RD, Gourley T, Spolski R, Davis CW, Langley WA, Ha SJ, Ye L, Sarkar S, Kalia V, Konieczny BT, Leonard WJ, Ahmed R.

J Virol. 2013 Jul;87(13):7737-46. doi: 10.1128/JVI.00063-13. Epub 2013 May 1.


The cytokines IL-21 and GM-CSF have opposing regulatory roles in the apoptosis of conventional dendritic cells.

Wan CK, Oh J, Li P, West EE, Wong EA, Andraski AB, Spolski R, Yu ZX, He J, Kelsall BL, Leonard WJ.

Immunity. 2013 Mar 21;38(3):514-27. doi: 10.1016/j.immuni.2013.02.011. Epub 2013 Feb 28.


Regulatory B cell (B10 Cell) expansion during Listeria infection governs innate and cellular immune responses in mice.

Horikawa M, Weimer ET, DiLillo DJ, Venturi GM, Spolski R, Leonard WJ, Heise MT, Tedder TF.

J Immunol. 2013 Feb 1;190(3):1158-68. doi: 10.4049/jimmunol.1201427. Epub 2012 Dec 28.


Mechanistic and structural insight into the functional dichotomy between IL-2 and IL-15.

Ring AM, Lin JX, Feng D, Mitra S, Rickert M, Bowman GR, Pande VS, Li P, Moraga I, Spolski R, Ozkan E, Leonard WJ, Garcia KC.

Nat Immunol. 2012 Dec;13(12):1187-95. doi: 10.1038/ni.2449. Epub 2012 Oct 28.


Regulatory B cells control T-cell autoimmunity through IL-21-dependent cognate interactions.

Yoshizaki A, Miyagaki T, DiLillo DJ, Matsushita T, Horikawa M, Kountikov EI, Spolski R, Poe JC, Leonard WJ, Tedder TF.

Nature. 2012 Nov 8;491(7423):264-8. doi: 10.1038/nature11501. Epub 2012 Oct 14.


BATF-JUN is critical for IRF4-mediated transcription in T cells.

Li P, Spolski R, Liao W, Wang L, Murphy TL, Murphy KM, Leonard WJ.

Nature. 2012 Oct 25;490(7421):543-6. doi: 10.1038/nature11530. Epub 2012 Sep 19.


Cd14 SNPs regulate the innate immune response.

Liu HH, Hu Y, Zheng M, Suhoski MM, Engleman EG, Dill DL, Hudnall M, Wang J, Spolski R, Leonard WJ, Peltz G.

Mol Immunol. 2012 Jun;51(2):112-27. doi: 10.1016/j.molimm.2012.02.112. Epub 2012 Mar 23.


IL-21 promotes the pathologic immune response to pneumovirus infection.

Spolski R, Wang L, Wan CK, Bonville CA, Domachowske JB, Kim HP, Yu Z, Leonard WJ.

J Immunol. 2012 Feb 15;188(4):1924-32. doi: 10.4049/jimmunol.1100767. Epub 2012 Jan 11.


Thymic stromal lymphopoietin is produced by dendritic cells.

Kashyap M, Rochman Y, Spolski R, Samsel L, Leonard WJ.

J Immunol. 2011 Aug 1;187(3):1207-11. doi: 10.4049/jimmunol.1100355. Epub 2011 Jun 20.


IL-21 is pivotal in determining age-dependent effectiveness of immune responses in a mouse model of human hepatitis B.

Publicover J, Goodsell A, Nishimura S, Vilarinho S, Wang ZE, Avanesyan L, Spolski R, Leonard WJ, Cooper S, Baron JL.

J Clin Invest. 2011 Mar;121(3):1154-62. doi: 10.1172/JCI44198.


IL-21 is an immune activator that also mediates suppression via IL-10.

Spolski R, Leonard WJ.

Crit Rev Immunol. 2010;30(6):559-70. Review.


Cutting edge: spontaneous development of IL-17-producing gamma delta T cells in the thymus occurs via a TGF-beta 1-dependent mechanism.

Do JS, Fink PJ, Li L, Spolski R, Robinson J, Leonard WJ, Letterio JJ, Min B.

J Immunol. 2010 Feb 15;184(4):1675-9. doi: 10.4049/jimmunol.0903539. Epub 2010 Jan 8.


IL-21 and T follicular helper cells.

Spolski R, Leonard WJ.

Int Immunol. 2010 Jan;22(1):7-12. doi: 10.1093/intimm/dxp112. Epub 2009 Nov 23. Review.


Adoptively transferred effector cells derived from naive rather than central memory CD8+ T cells mediate superior antitumor immunity.

Hinrichs CS, Borman ZA, Cassard L, Gattinoni L, Spolski R, Yu Z, Sanchez-Perez L, Muranski P, Kern SJ, Logun C, Palmer DC, Ji Y, Reger RN, Leonard WJ, Danner RL, Rosenberg SA, Restifo NP.

Proc Natl Acad Sci U S A. 2009 Oct 13;106(41):17469-74. doi: 10.1073/pnas.0907448106. Epub 2009 Sep 24.


New insights into the regulation of T cells by gamma(c) family cytokines.

Rochman Y, Spolski R, Leonard WJ.

Nat Rev Immunol. 2009 Jul;9(7):480-90. doi: 10.1038/nri2580. Review.


Cytokine mediators of Th17 function.

Spolski R, Leonard WJ.

Eur J Immunol. 2009 Mar;39(3):658-61. doi: 10.1002/eji.200839066.


IL-21 mediates suppressive effects via its induction of IL-10.

Spolski R, Kim HP, Zhu W, Levy DE, Leonard WJ.

J Immunol. 2009 Mar 1;182(5):2859-67. doi: 10.4049/jimmunol.0802978.


A critical role for IL-21 receptor signaling in the pathogenesis of systemic lupus erythematosus in BXSB-Yaa mice.

Bubier JA, Sproule TJ, Foreman O, Spolski R, Shaffer DJ, Morse HC 3rd, Leonard WJ, Roopenian DC.

Proc Natl Acad Sci U S A. 2009 Feb 3;106(5):1518-23. doi: 10.1073/pnas.0807309106. Epub 2009 Jan 21.


Interleukin-6/STAT3 signaling regulates the ability of naive T cells to acquire B-cell help capacities.

Eddahri F, Denanglaire S, Bureau F, Spolski R, Leonard WJ, Leo O, Andris F.

Blood. 2009 Mar 12;113(11):2426-33. doi: 10.1182/blood-2008-04-154682. Epub 2008 Nov 19.


IL-21 signaling is critical for the development of type I diabetes in the NOD mouse.

Spolski R, Kashyap M, Robinson C, Yu Z, Leonard WJ.

Proc Natl Acad Sci U S A. 2008 Sep 16;105(37):14028-33. doi: 10.1073/pnas.0804358105. Epub 2008 Sep 8.


Development of regulatory T cells requires IL-7Ralpha stimulation by IL-7 or TSLP.

Mazzucchelli R, Hixon JA, Spolski R, Chen X, Li WQ, Hall VL, Willette-Brown J, Hurwitz AA, Leonard WJ, Durum SK.

Blood. 2008 Oct 15;112(8):3283-92. doi: 10.1182/blood-2008-02-137414. Epub 2008 Jul 29.


The Yin and Yang of interleukin-21 in allergy, autoimmunity and cancer.

Spolski R, Leonard WJ.

Curr Opin Immunol. 2008 Jun;20(3):295-301. doi: 10.1016/j.coi.2008.02.004. Epub 2008 Jun 12. Review.


Interleukin 21: a cytokine/cytokine receptor system that has come of age.

Leonard WJ, Zeng R, Spolski R.

J Leukoc Biol. 2008 Aug;84(2):348-56. doi: 10.1189/jlb.0308149. Epub 2008 May 8. Review.


Measurement of interleukin-21.

Zeng R, Spolski R, Leonard WJ.

Curr Protoc Immunol. 2007 Aug;Chapter 6:Unit 6.30. doi: 10.1002/0471142735.im0630s78.


T-cell development and function are modulated by dual specificity phosphatase DUSP5.

Kovanen PE, Bernard J, Al-Shami A, Liu C, Bollenbacher-Reilley J, Young L, Pise-Masison C, Spolski R, Leonard WJ.

J Biol Chem. 2008 Jun 20;283(25):17362-9. doi: 10.1074/jbc.M709887200. Epub 2008 Apr 21.


IL-2 and IL-21 confer opposing differentiation programs to CD8+ T cells for adoptive immunotherapy.

Hinrichs CS, Spolski R, Paulos CM, Gattinoni L, Kerstann KW, Palmer DC, Klebanoff CA, Rosenberg SA, Leonard WJ, Restifo NP.

Blood. 2008 Jun 1;111(11):5326-33. doi: 10.1182/blood-2007-09-113050. Epub 2008 Feb 14.


Thymic stromal lymphopoietin is not necessary or sufficient to mediate the thymopoietic effects of keratinocyte growth factor.

Guimond M, Leonard WJ, Spolski R, Rossi SW, Veenstra RG, Hollander GA, Mackall CL, Blazar BR.

Blood. 2008 Jan 15;111(2):969-70. doi: 10.1182/blood-2007-09-113316. No abstract available.


Interleukin-21: basic biology and implications for cancer and autoimmunity.

Spolski R, Leonard WJ.

Annu Rev Immunol. 2008;26:57-79. Review.


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