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In this study, we define distinct patterns of surface marker and cytokine expression among the ILC subsets that may further delineate their migration and function. Most notably, we found that the subset previously defined as group 1 ILC (ILC1) contains CD4+ CD8\u2212, CD4\u2212 CD8+, and CD4\u2212 CD8\u2212 populations. Although all ILC1 subsets shared characteristics with Th1 cells, CD4+ ILC1 also demonstrated significant phenotypic and functional heterogeneity. We also show that the frequencies of CD4+ ILC1 and NKp44+ group 3 ILC, but not CD4\u2212 ILC1 or group 2 ILC, are increased in the peripheral blood of individuals with systemic sclerosis (SSc), a disease characterized by fibrotic and vascular pathology, as well as immune dysregulation. Furthermore, we demonstrate that CD4+ and CD4\u2212 ILC1 are functionally divergent based on their IL-6R\u03b1 expression and that the frequency of IL-6R\u03b1 expression on ILC is altered in SSc. The distinct phenotypic and functional features of CD4+ and CD4\u2212 ILC1 suggest that they may have differing roles in the pathogenesis of immune-mediated diseases, such as SSc.<\/jats:p>","DOI":"10.4049\/jimmunol.1501491","type":"journal-article","created":{"date-parts":[[2016,1,30]],"date-time":"2016-01-30T03:33:53Z","timestamp":1454124833000},"page":"2051-2062","source":"Crossref","is-referenced-by-count":122,"title":["CD4+ Group 1 Innate Lymphoid Cells (ILC) Form a Functionally Distinct ILC Subset That Is Increased in Systemic Sclerosis"],"prefix":"10.1093","volume":"196","author":[{"given":"Florence","family":"Roan","sequence":"first","affiliation":[{"name":"Immunology Program, Benaroya Research Institute at Virginia Mason , Seattle, WA 98101"},{"name":"Division of Allergy and Infectious Diseases, University of Washington , Seattle, WA 98109"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Thomas A","family":"Stoklasek","sequence":"additional","affiliation":[{"name":"Immunology Program, Benaroya Research Institute at Virginia Mason , Seattle, WA 98101"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Elizabeth","family":"Whalen","sequence":"additional","affiliation":[{"name":"Systems Immunology Division, Benaroya Research Institute at Virginia Mason , Seattle, WA 98101"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jerry A","family":"Molitor","sequence":"additional","affiliation":[{"name":"Division of Rheumatology, University of Minnesota , Minneapolis, MN 55455"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-8793-7848","authenticated-orcid":false,"given":"Jeffrey A","family":"Bluestone","sequence":"additional","affiliation":[{"name":"Diabetes Center, University of California San Francisco , San Francisco, CA 94143"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jane H","family":"Buckner","sequence":"additional","affiliation":[{"name":"Translational Research Program, Benaroya Research Institute at Virginia Mason , Seattle, WA 98101"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Steven F","family":"Ziegler","sequence":"additional","affiliation":[{"name":"Immunology Program, Benaroya Research Institute at Virginia Mason , Seattle, WA 98101"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"286","published-online":{"date-parts":[[2016,3,1]]},"reference":[{"key":"2025030702140770500_r1","doi-asserted-by":"crossref","first-page":"245","DOI":"10.1038\/nature12526","article-title":"Type 2 innate lymphoid cells control eosinophil homeostasis","volume":"502","author":"Nussbaum","year":"2013","journal-title":"Nature"},{"key":"2025030702140770500_r2","doi-asserted-by":"crossref","first-page":"74","DOI":"10.1016\/j.cell.2014.12.011","article-title":"Activated type 2 innate lymphoid cells regulate beige fat biogenesis","volume":"160","author":"Lee","year":"2015","journal-title":"Cell"},{"key":"2025030702140770500_r3","doi-asserted-by":"crossref","first-page":"242","DOI":"10.1038\/nature14115","article-title":"Group 2 innate lymphoid cells promote beiging of white adipose tissue and limit obesity","volume":"519","author":"Brestoff","year":"2015","journal-title":"Nature"},{"key":"2025030702140770500_r4","doi-asserted-by":"crossref","first-page":"3572","DOI":"10.1182\/blood-2014-07-587493","article-title":"Interleukin-5-producing group 2 innate lymphoid cells control eosinophilia induced by interleukin-2 therapy","volume":"124","author":"Van Gool","year":"2014","journal-title":"Blood"},{"key":"2025030702140770500_r5","doi-asserted-by":"crossref","first-page":"320","DOI":"10.1038\/ni.2002","article-title":"ROR\u03b3t+ innate lymphoid cells regulate intestinal homeostasis by integrating negative signals from the symbiotic microbiota","volume":"12","author":"Sawa","year":"2011","journal-title":"Nat. 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