{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,2,10]],"date-time":"2026-02-10T15:47:22Z","timestamp":1770738442464,"version":"3.49.0"},"reference-count":50,"publisher":"Rockefeller University Press","issue":"11","content-domain":{"domain":["rupress.org"],"crossmark-restriction":true},"short-container-title":[],"published-print":{"date-parts":[[1998,6,1]]},"abstract":"<jats:p>4-1BB ligand (4-1BBL) is a member of the tumor necrosis factor (TNF) family expressed on activated antigen-presenting cells. Its receptor, 4-1BB, is a member of the TNF receptor family expressed on activated CD4 and CD8 T cells. We have produced a soluble form of 4-1BBL using the baculovirus expression system. When coimmobilized on plastic with anti-CD3, soluble 4-1BBL induces interleukin (IL)-2 production by resting CD28+ or CD28\u2212 T cells, indicating that 4-1BBL can function independently of other cell surface molecules, including CD28, in costimulation of resting T cell activation. At low concentrations of anti-CD3, 4-1BBL is inferior to anti-CD28 in T cell activation. However, when 4-1BB ligand is provided together with strong TCR signals, then 4-1BBL and anti-CD28 are equally potent in stimulation of IL-2 production by resting T cells. We find that TNF receptor\u2013associated factor (TRAF)1 or TRAF2 associate with a glutathione S-transferase\u20134-1BB cytoplasmic domain fusion protein in vitro. In T cells, we find that association of TRAF1 and TRAF2 with 4-1BB requires 4-1BB cross-linking. In support of a functional role for TRAF2 in 4-1BB signaling, we find that resting T cells isolated from TRAF2-deficient mice or from mice expressing a dominant negative form of TRAF2 fail to augment IL-2 production in response to soluble 4-1BBL. Thus 4-1BB, via the TRAF2 molecule, can provide CD28-independent costimulatory signals to resting T cells.<\/jats:p>","DOI":"10.1084\/jem.187.11.1849","type":"journal-article","created":{"date-parts":[[2002,7,26]],"date-time":"2002-07-26T16:49:30Z","timestamp":1027702170000},"page":"1849-1862","update-policy":"https:\/\/doi.org\/10.1084\/jem.crossmarkpolicy","source":"Crossref","is-referenced-by-count":223,"title":["CD28-independent, TRAF2-dependent Costimulation of Resting T Cells by 4-1BB Ligand"],"prefix":"10.1084","volume":"187","author":[{"given":"Katina","family":"Saoulli","sequence":"first","affiliation":[{"name":"From the *Department of Immunology, the \u2021Department of Medical Biophysics, and Amgen Institute, University of Toronto, Toronto, Ontario M5S 1A8, Canada; and the \u00a7Howard Hughes Medical Institute and \u2016The Rockefeller University, New York 10021"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Soo Young","family":"Lee","sequence":"additional","affiliation":[{"name":"From the *Department of Immunology, the \u2021Department of Medical Biophysics, and Amgen Institute, University of Toronto, Toronto, Ontario M5S 1A8, Canada; and the \u00a7Howard Hughes Medical Institute and \u2016The Rockefeller University, New York 10021"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Jennifer L.","family":"Cannons","sequence":"additional","affiliation":[{"name":"From the *Department of Immunology, the \u2021Department of Medical Biophysics, and Amgen Institute, University of Toronto, Toronto, Ontario M5S 1A8, Canada; and the \u00a7Howard Hughes Medical Institute and \u2016The Rockefeller University, New York 10021"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Wen Chen","family":"Yeh","sequence":"additional","affiliation":[{"name":"From the *Department of Immunology, the \u2021Department of Medical Biophysics, and Amgen Institute, University of Toronto, Toronto, Ontario M5S 1A8, Canada; and the \u00a7Howard Hughes Medical Institute and \u2016The Rockefeller University, New York 10021"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Angela","family":"Santana","sequence":"additional","affiliation":[{"name":"From the *Department of Immunology, the \u2021Department of Medical Biophysics, and Amgen Institute, University of Toronto, Toronto, Ontario M5S 1A8, Canada; and the \u00a7Howard Hughes Medical Institute and \u2016The Rockefeller University, New York 10021"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Marni D.","family":"Goldstein","sequence":"additional","affiliation":[{"name":"From the *Department of Immunology, the \u2021Department of Medical Biophysics, and Amgen Institute, University of Toronto, Toronto, Ontario M5S 1A8, Canada; and the \u00a7Howard Hughes Medical Institute and \u2016The Rockefeller University, New York 10021"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Naveen","family":"Bangia","sequence":"additional","affiliation":[{"name":"From the *Department of Immunology, the \u2021Department of Medical Biophysics, and Amgen Institute, University of Toronto, Toronto, Ontario M5S 1A8, Canada; and the \u00a7Howard Hughes Medical Institute and \u2016The Rockefeller University, New York 10021"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Mark A.","family":"DeBenedette","sequence":"additional","affiliation":[{"name":"From the *Department of Immunology, the \u2021Department of Medical Biophysics, and Amgen Institute, University of Toronto, Toronto, Ontario M5S 1A8, Canada; and the \u00a7Howard Hughes Medical Institute and \u2016The Rockefeller University, New York 10021"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Tak W.","family":"Mak","sequence":"additional","affiliation":[{"name":"From the *Department of Immunology, the \u2021Department of Medical Biophysics, and Amgen Institute, University of Toronto, Toronto, Ontario M5S 1A8, Canada; and the \u00a7Howard Hughes Medical Institute and \u2016The Rockefeller University, New York 10021"},{"name":"From the *Department of Immunology, the \u2021Department of Medical Biophysics, and Amgen Institute, University of Toronto, Toronto, Ontario M5S 1A8, Canada; and the \u00a7Howard Hughes Medical Institute and \u2016The Rockefeller University, New York 10021"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Yongwon","family":"Choi","sequence":"additional","affiliation":[{"name":"From the *Department of Immunology, the \u2021Department of Medical Biophysics, and Amgen Institute, University of Toronto, Toronto, Ontario M5S 1A8, Canada; and the \u00a7Howard Hughes Medical Institute and \u2016The Rockefeller University, New York 10021"},{"name":"From the *Department of Immunology, the \u2021Department of Medical Biophysics, and Amgen Institute, University of Toronto, 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