{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,11,5]],"date-time":"2026-11-05T05:29:32Z","timestamp":1793856572732,"version":"4.6.1"},"reference-count":65,"publisher":"Elsevier BV","license":[{"start":{"date-parts":[[2026,10,1]],"date-time":"2026-10-01T00:00:00Z","timestamp":1790812800000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/tdm\/userlicense\/1.0\/"},{"start":{"date-parts":[[2026,10,1]],"date-time":"2026-10-01T00:00:00Z","timestamp":1790812800000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/legal\/tdmrep-license"},{"start":{"date-parts":[[2026,10,22]],"date-time":"2026-10-22T00:00:00Z","timestamp":1792627200000},"content-version":"vor","delay-in-days":21,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/100006492","name":"National Institute of Allergy and Infectious Diseases Division of Intramural Research","doi-asserted-by":"publisher","id":[{"id":"10.13039\/100006492","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["clinicalkey.com","clinicalkey.com.au","clinicalkey.es","clinicalkey.fr","clinicalkey.jp","mucosalimmunology.org","elsevier.com","sciencedirect.com"],"crossmark-restriction":true},"short-container-title":["Mucosal Immunology"],"published-print":{"date-parts":[[2026,10]]},"DOI":"10.1016\/j.mucimm.2026.100422","type":"journal-article","created":{"date-parts":[[2026,10,23]],"date-time":"2026-10-23T23:14:07Z","timestamp":1792797247000},"page":"100422","update-policy":"https:\/\/doi.org\/10.1016\/elsevier_cm_policy","source":"Crossref","is-referenced-by-count":0,"title":["Control of M. tuberculosis infection requires CD28 to generate lung-homing T cells and CTLA-4 to limit immunopathology"],"prefix":"10.1016","author":[{"given":"Christine E.","family":"Nelson","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Shunsuke","family":"Sakai","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Keith D.","family":"Kauffman","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Taylor W.","family":"Foreman","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Melanie","family":"Cohen","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Julie","family":"Laux","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Eduardo P.","family":"Amaral","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Emily M.","family":"Grayson","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Anne I.","family":"Sperling","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Katrin D.","family":"Mayer-Barber","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Daniel L.","family":"Barber","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"78","reference":[{"key":"10.1016\/j.mucimm.2026.100422_bb0005","series-title":"Global Tuberculosis Report 2024","year":"2024"},{"key":"10.1016\/j.mucimm.2026.100422_bb0010","doi-asserted-by":"crossref","first-page":"2243","DOI":"10.1084\/jem.178.6.2243","article-title":"Disseminated tuberculosis in interferon gamma gene-disrupted mice","volume":"178","author":"Cooper","year":"1993","journal-title":"J. Exp. Med."},{"key":"10.1016\/j.mucimm.2026.100422_bb0015","doi-asserted-by":"crossref","first-page":"2249","DOI":"10.1084\/jem.178.6.2249","article-title":"An essential role for interferon gamma in resistance to Mycobacterium tuberculosis infection","volume":"178","author":"Flynn","year":"1993","journal-title":"J. Exp. Med."},{"key":"10.1016\/j.mucimm.2026.100422_bb0020","doi-asserted-by":"crossref","first-page":"347","DOI":"10.1084\/jem.192.3.347","article-title":"Depletion of CD4(+) T cells causes reactivation of murine persistent tuberculosis despite continued expression of interferon gamma and nitric oxide synthase 2","volume":"192","author":"Scanga","year":"2000","journal-title":"J Exp Med"},{"key":"10.1016\/j.mucimm.2026.100422_bb0025","doi-asserted-by":"crossref","first-page":"19408","DOI":"10.1073\/pnas.1006298107","article-title":"Distinct functions of antigen-specific CD4 T cells during murine Mycobacterium tuberculosis infection","volume":"107","author":"Reiley","year":"2010","journal-title":"Proc Natl Acad Sci"},{"key":"10.1016\/j.mucimm.2026.100422_bb0030","doi-asserted-by":"crossref","first-page":"2965","DOI":"10.4049\/jimmunol.1400019","article-title":"Control of Mycobacterium tuberculosis infection by a subset of lung parenchyma homing CD4 T cells","volume":"192","author":"Sakai","year":"2014","journal-title":"J Immunol"},{"key":"10.1016\/j.mucimm.2026.100422_bb0035","doi-asserted-by":"crossref","first-page":"715","DOI":"10.1084\/jem.20141518","article-title":"ICOS and Bcl6-dependent pathways maintain a CD4 T cell population with memory-like properties during tuberculosis","volume":"212","author":"Moguche","year":"2015","journal-title":"J Exp Med"},{"key":"10.1016\/j.mucimm.2026.100422_bb0040","doi-asserted-by":"crossref","first-page":"3091","DOI":"10.1016\/j.celrep.2017.03.007","article-title":"Th1 differentiation drives the accumulation of intravascular, non-protective CD4 T cells during tuberculosis","volume":"18","author":"Sallin","year":"2017","journal-title":"Cell Rep"},{"key":"10.1016\/j.mucimm.2026.100422_bb0045","doi-asserted-by":"crossref","first-page":"6311","DOI":"10.4049\/jimmunol.1300248","article-title":"Control of chronic Mycobacterium tuberculosis infection by CD4 KLRG1\u2212 IL-2\u2013secreting central memory cells","volume":"190","author":"Lindenstr\u00f8m","year":"2013","journal-title":"J. Immunol."},{"key":"10.1016\/j.mucimm.2026.100422_bb0050","doi-asserted-by":"crossref","first-page":"3247","DOI":"10.4049\/jimmunol.1300283","article-title":"Protective CD4 T cells targeting cryptic epitopes of Mycobacterium tuberculosis resist infection-driven terminal differentiation","volume":"192","author":"Woodworth","year":"2014","journal-title":"J. Immunol."},{"key":"10.1016\/j.mucimm.2026.100422_bb0055","doi-asserted-by":"crossref","first-page":"555","DOI":"10.1038\/mi.2016.70","article-title":"Subunit vaccine H56\/CAF01 induces a population of circulating CD4 T cells that traffic into the Mycobacterium tuberculosis-infected lung","volume":"10","author":"Woodworth","year":"2017","journal-title":"Mucosal Immunol"},{"key":"10.1016\/j.mucimm.2026.100422_bb0060","doi-asserted-by":"crossref","DOI":"10.1128\/mBio.00226-21","article-title":"In vivo antigen expression regulates CD4 T cell differentiation and vaccine efficacy against Mycobacterium tuberculosis infection","volume":"12","author":"Clemmensen","year":"2021","journal-title":"mBio"},{"key":"10.1016\/j.mucimm.2026.100422_bb0065","doi-asserted-by":"crossref","first-page":"1449","DOI":"10.1084\/jem.20141520","article-title":"Interleukin 27R regulates CD4+ T cell phenotype and impacts protective immunity during Mycobacterium tuberculosis infection","volume":"212","author":"Torrado","year":"2015","journal-title":"J Exp Med"},{"key":"10.1016\/j.mucimm.2026.100422_bb0070","doi-asserted-by":"crossref","first-page":"1198","DOI":"10.1038\/s41564-018-0231-6","article-title":"Host resistance to pulmonary Mycobacterium tuberculosis infection requires CD153 expression","volume":"3","author":"Sallin","year":"2018","journal-title":"Nat Microbiol"},{"key":"10.1016\/j.mucimm.2026.100422_bb0075","doi-asserted-by":"crossref","first-page":"491","DOI":"10.1038\/s41385-020-0322-6","article-title":"Mycobacterium tuberculosis-specific CD4 T cells expressing CD153 inversely associate with bacterial load and disease severity in human tuberculosis","volume":"14","author":"Du Bruyn","year":"2021","journal-title":"Mucosal Immunol"},{"key":"10.1016\/j.mucimm.2026.100422_bb0080","doi-asserted-by":"crossref","DOI":"10.1084\/jem.20222090","article-title":"CD30 co-stimulation drives differentiation of protective T cells during Mycobacterium tuberculosis infection","volume":"220","author":"Foreman","year":"2023","journal-title":"J Exp Med"},{"key":"10.1016\/j.mucimm.2026.100422_bb0085","doi-asserted-by":"crossref","first-page":"3793","DOI":"10.4049\/jimmunol.0802996","article-title":"B7 costimulation is critical for host control of chronic Mycobacterium tuberculosis infection","volume":"182","author":"Bhatt","year":"2009","journal-title":"J Immunol"},{"key":"10.1016\/j.mucimm.2026.100422_bb0090","doi-asserted-by":"crossref","first-page":"975","DOI":"10.1093\/infdis\/jir868","article-title":"OX40 ligand fusion protein delivered simultaneously with the BCG vaccine provides superior protection against murine Mycobacterium tuberculosis infection","volume":"205","author":"Snelgrove","year":"2012","journal-title":"J Infect Dis"},{"key":"10.1016\/j.mucimm.2026.100422_bb0095","doi-asserted-by":"crossref","first-page":"8423","DOI":"10.1038\/s41467-023-44152-8","article-title":"Recently activated CD4 T cells in tuberculosis express OX40 as a target for host-directed immunotherapy","volume":"14","author":"Gress","year":"2023","journal-title":"Nat Commun"},{"key":"10.1016\/j.mucimm.2026.100422_bb0100","doi-asserted-by":"crossref","unstructured":"Fern\u00e1ndez Do Porto, D. A. et al. CD137 differentially regulates innate and adaptive immunity against Mycobacterium tuberculosis. Immunol Cell Biol 90, 449\u2013456 (2012).","DOI":"10.1038\/icb.2011.63"},{"key":"10.1016\/j.mucimm.2026.100422_bb0105","doi-asserted-by":"crossref","first-page":"233","DOI":"10.1146\/annurev.immunol.14.1.233","article-title":"CD28\/B7 system of t cell COSTIMULATION","volume":"14","author":"Lenschow","year":"1996","journal-title":"Annu Rev Immunol"},{"key":"10.1016\/j.mucimm.2026.100422_bb0110","doi-asserted-by":"crossref","first-page":"180","DOI":"10.4049\/jimmunol.174.1.180","article-title":"CD28 regulates the translation of Bcl-xL via the phosphatidylinositol 3-kinase\/mammalian target of rapamycin pathway","volume":"174","author":"Wu","year":"2005","journal-title":"J Immunol"},{"key":"10.1016\/j.mucimm.2026.100422_bb0115","doi-asserted-by":"crossref","first-page":"1007","DOI":"10.4049\/jimmunol.154.3.1007","article-title":"CD28 costimulation up-regulates long-term IL-2R beta expression in human T cells through combined transcriptional and post-transcriptional regulation","volume":"154","author":"Cerdan","year":"1995","journal-title":"J Immunol"},{"key":"10.1016\/j.mucimm.2026.100422_bb0120","doi-asserted-by":"crossref","first-page":"385","DOI":"10.1016\/j.cell.2017.08.018","article-title":"Mitochondrial priming by CD28","volume":"171","author":"Klein Geltink","year":"2017","journal-title":"Cell"},{"key":"10.1016\/j.mucimm.2026.100422_bb0125","doi-asserted-by":"crossref","first-page":"1497","DOI":"10.1097\/00007890-199711270-00025","article-title":"Differential down-regulation of CD28 by B7-1 and B7-2 engagement","volume":"64","author":"Eck","year":"1997","journal-title":"Transplantation"},{"key":"10.1016\/j.mucimm.2026.100422_bb0130","doi-asserted-by":"crossref","first-page":"641","DOI":"10.1002\/jlb.65.5.641","article-title":"Generation of CD28- cells from long-term-stimulated CD8+CD28+ T cells: a possible mechanism accounting for the increased number of CD8+CD28- T cells in HIV-1-infected patients","volume":"65","author":"Fiorentini","year":"1999","journal-title":"J Leukoc Biol"},{"key":"10.1016\/j.mucimm.2026.100422_bb0135","doi-asserted-by":"crossref","first-page":"eadg0878","DOI":"10.1126\/sciimmunol.adg0878","article-title":"Sustained CD28 costimulation is required for self-renewal and differentiation of TCF-1+ PD-1+ CD8 T cells","volume":"8","author":"Humblin","year":"2023","journal-title":"Sci. Immunol."},{"key":"10.1016\/j.mucimm.2026.100422_bb0140","doi-asserted-by":"crossref","first-page":"e55","DOI":"10.1016\/S1470-2045(19)30674-6","article-title":"Immune checkpoint inhibitors and tuberculosis: an old disease in a new context","volume":"21","author":"Langan","year":"2020","journal-title":"Lancet Oncol"},{"key":"10.1016\/j.mucimm.2026.100422_bb0145","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pone.0006307","article-title":"CTLA4 autoimmunity-associated genotype contributes to severe pulmonary tuberculosis in an African population","volume":"4","author":"Thye","year":"2009","journal-title":"PloS One"},{"key":"10.1016\/j.mucimm.2026.100422_bb0150","doi-asserted-by":"crossref","first-page":"945","DOI":"10.7150\/ijbs.4390","article-title":"Association of CTLA4 gene polymorphisms with susceptibility and pathology correlation to pulmonary tuberculosis in southern Han Chinese","volume":"8","author":"Wang","year":"2012","journal-title":"Int J Biol Sci"},{"key":"10.1016\/j.mucimm.2026.100422_bb0155","article-title":"Association of tuberculosis risk with genetic polymorphisms of the immune checkpoint genes PDCD1, CTLA-4, and TIM3","volume":"19","author":"Liu","year":"2024","journal-title":"PloS One"},{"key":"10.1016\/j.mucimm.2026.100422_bb0160","doi-asserted-by":"crossref","first-page":"337","DOI":"10.47665\/tb.42.3.013","article-title":"Cytotoxic T-lymphocyte Antigen-4 (CTLA-4) gene polymorphism is associated with pulmonary tuberculosis susceptibility in a Thai population","volume":"42","author":"Tiwongsa","year":"2025","journal-title":"Trop Biomed"},{"key":"10.1016\/j.mucimm.2026.100422_bb0165","doi-asserted-by":"crossref","DOI":"10.1016\/j.celrep.2022.110983","article-title":"Mycobacterium tuberculosis infection drives a type I IFN signature in lung lymphocytes","volume":"39","author":"Akter","year":"2022","journal-title":"Cell Rep"},{"key":"10.1016\/j.mucimm.2026.100422_bb0170","doi-asserted-by":"crossref","first-page":"263","DOI":"10.1038\/16717","article-title":"ICOS is an inducible T-cell co-stimulator structurally and functionally related to CD28","volume":"397","author":"Hutloff","year":"1999","journal-title":"Nature"},{"key":"10.1016\/j.mucimm.2026.100422_bb0175","doi-asserted-by":"crossref","first-page":"459","DOI":"10.1084\/jem.182.2.459","article-title":"CD28 and CTLA-4 have opposing effects on the response of T cells to stimulation","volume":"182","author":"Krummel","year":"1995","journal-title":"J Exp Med"},{"key":"10.1016\/j.mucimm.2026.100422_bb0180","doi-asserted-by":"crossref","first-page":"1032","DOI":"10.4049\/jimmunol.155.3.1032","article-title":"Antigen-dependent clonal expansion of a trace population of antigen-specific CD4+ T cells in vivo is dependent on CD28 costimulation and inhibited by CTLA-4","volume":"155","author":"Kearney","year":"1995","journal-title":"J Immunol"},{"key":"10.1016\/j.mucimm.2026.100422_bb0185","doi-asserted-by":"crossref","first-page":"1052","DOI":"10.4049\/jimmunol.177.2.1052","article-title":"CTLA-4 overexpression inhibits T cell responses through a CD28-B7-dependent mechanism","volume":"177","author":"Engelhardt","year":"2006","journal-title":"J Immunol"},{"key":"10.1016\/j.mucimm.2026.100422_bb0190","doi-asserted-by":"crossref","first-page":"1423","DOI":"10.1126\/science.aaf0683","article-title":"Rescue of exhausted CD8 T cells by PD-1\u2013targeted therapies is CD28-dependent","volume":"355","author":"Kamphorst","year":"2017","journal-title":"Science"},{"key":"10.1016\/j.mucimm.2026.100422_bb0195","doi-asserted-by":"crossref","first-page":"1428","DOI":"10.1126\/science.aaf1292","article-title":"T cell costimulatory receptor CD28 is a primary target for PD-1\u2013mediated inhibition","volume":"355","author":"Hui","year":"2017","journal-title":"Science"},{"key":"10.1016\/j.mucimm.2026.100422_bb0200","doi-asserted-by":"crossref","first-page":"407","DOI":"10.1093\/intimm\/11.3.407","article-title":"Changes in the strength of co-stimulation through the B7\/CD28 pathway alter functional T cell responses to altered peptide ligands","volume":"11","author":"Murtaza","year":"1999","journal-title":"Int Immunol"},{"key":"10.1016\/j.mucimm.2026.100422_bb0205","doi-asserted-by":"crossref","first-page":"74","DOI":"10.1006\/cimm.1997.1137","article-title":"High concentrations of antigenic ligand activate and do not tolerize naive CD4 T cells in the absence of CD28\/B7 costimulation","volume":"179","author":"Teh","year":"1997","journal-title":"Cell Immunol"},{"key":"10.1016\/j.mucimm.2026.100422_bb0210","doi-asserted-by":"crossref","first-page":"1207","DOI":"10.4049\/jimmunol.169.3.1207","article-title":"Direct ex vivo analysis of human CD4(+) memory T cell activation requirements at the single clonotype level","volume":"169","author":"Bitmansour","year":"2002","journal-title":"J Immunol"},{"key":"10.1016\/j.mucimm.2026.100422_bb0215","doi-asserted-by":"crossref","first-page":"631","DOI":"10.1084\/jem.180.2.631","article-title":"Comparative analysis of B7-1 and B7-2 costimulatory ligands: expression and function","volume":"180","author":"Hathcock","year":"1994","journal-title":"J Exp Med"},{"key":"10.1016\/j.mucimm.2026.100422_bb0220","doi-asserted-by":"crossref","first-page":"5268","DOI":"10.4049\/jimmunol.161.10.5268","article-title":"B7.1 is a quantitatively stronger Costimulus than B7.2 in the activation of naive CD8+ TCR-transgenic T Cells1","volume":"161","author":"Fields","year":"1998","journal-title":"J. Immunol."},{"key":"10.1016\/j.mucimm.2026.100422_bb0225","doi-asserted-by":"crossref","first-page":"401","DOI":"10.1016\/j.immuni.2004.06.017","article-title":"B7\u20131 and B7\u20132 selectively recruit CTLA-4 and CD28 to the immunological synapse","volume":"21","author":"Pentcheva-Hoang","year":"2004","journal-title":"Immunity"},{"key":"10.1016\/j.mucimm.2026.100422_bb0230","doi-asserted-by":"crossref","first-page":"6264","DOI":"10.1038\/s41467-020-20070-x","article-title":"B7-CD28 co-stimulation modulates central tolerance via thymic clonal deletion and Treg generation through distinct mechanisms","volume":"11","author":"Watanabe","year":"2020","journal-title":"Nat Commun"},{"key":"10.1016\/j.mucimm.2026.100422_bb0235","doi-asserted-by":"crossref","first-page":"1016","DOI":"10.4049\/jimmunol.1301236","article-title":"Cutting edge: direct recognition of infected cells by CD4 T cells is required for control of intracellular mycobacterium tuberculosis in vivo","volume":"191","author":"Srivastava","year":"2013","journal-title":"J. Immunol."},{"key":"10.1016\/j.mucimm.2026.100422_bb0240","doi-asserted-by":"crossref","first-page":"439","DOI":"10.1016\/j.chom.2018.08.001","article-title":"Alveolar macrophages provide an early Mycobacterium tuberculosis niche and initiate dissemination","volume":"24","author":"Cohen","year":"2018","journal-title":"Cell Host Microbe"},{"key":"10.1016\/j.mucimm.2026.100422_bb0245","doi-asserted-by":"crossref","DOI":"10.1371\/journal.ppat.1008621","article-title":"CD11cHi monocyte-derived macrophages are a major cellular compartment infected by Mycobacterium tuberculosis","volume":"16","author":"Lee","year":"2020","journal-title":"PLoS Pathog"},{"key":"10.1016\/j.mucimm.2026.100422_bb0250","doi-asserted-by":"crossref","first-page":"2509","DOI":"10.4049\/jimmunol.179.4.2509","article-title":"Mycobacterium tuberculosis infects dendritic cells with high frequency and impairs their function in Vivo1","volume":"179","author":"Wolf","year":"2007","journal-title":"J Immunol"},{"key":"10.1016\/j.mucimm.2026.100422_bb0255","doi-asserted-by":"crossref","DOI":"10.1371\/journal.ppat.1012205","article-title":"Mycobacterium tuberculosis resides in lysosome-poor monocyte-derived lung cells during chronic infection","volume":"20","author":"Zheng","year":"2024","journal-title":"PLoS Pathog"},{"key":"10.1016\/j.mucimm.2026.100422_bb0260","doi-asserted-by":"crossref","first-page":"1603","DOI":"10.1084\/jem.20141030","article-title":"Deletion of CTLA-4 on regulatory T cells during adulthood leads to resistance to autoimmunity","volume":"212","author":"Paterson","year":"2015","journal-title":"J Exp Med"},{"key":"10.1016\/j.mucimm.2026.100422_bb0265","doi-asserted-by":"crossref","first-page":"295","DOI":"10.1084\/jem.192.2.295","article-title":"Cytotoxic T Lymphocyte\u2013Associated Antigen 4 Plays an Essential Role in the Function of Cd25+Cd4+ Regulatory Cells That Control Intestinal Inflammation","volume":"192","author":"Read","year":"2000","journal-title":"J. Exp. Med."},{"key":"10.1016\/j.mucimm.2026.100422_bb0270","doi-asserted-by":"crossref","first-page":"303","DOI":"10.1084\/jem.192.2.303","article-title":"Immunologic Self-Tolerance Maintained by Cd25+Cd4+Regulatory T Cells Constitutively Expressing Cytotoxic T Lymphocyte\u2013Associated Antigen 4","volume":"192","author":"Takahashi","year":"2000","journal-title":"J. Exp. Med."},{"key":"10.1016\/j.mucimm.2026.100422_bb0275","doi-asserted-by":"crossref","first-page":"324","DOI":"10.1038\/mi.2012.75","article-title":"CTLA-4 promotes Foxp3 induction and regulatory T cell accumulation in the intestinal lamina propria","volume":"6","author":"Barnes","year":"2013","journal-title":"Mucosal Immunol"},{"key":"10.1016\/j.mucimm.2026.100422_bb0280","doi-asserted-by":"crossref","first-page":"363","DOI":"10.1038\/s41385-018-0105-5","article-title":"GITRL on inflammatory antigen presenting cells in the lung parenchyma provides signal 4 for T-cell accumulation and tissue-resident memory T-cell formation","volume":"12","author":"Chu","year":"2019","journal-title":"Mucosal Immunol"},{"key":"10.1016\/j.mucimm.2026.100422_bb0285","doi-asserted-by":"crossref","first-page":"3344","DOI":"10.4049\/jimmunol.163.6.3344","article-title":"Role of CD28 in the generation of effector and memory responses required for resistance to toxoplasma gondii","volume":"163","author":"Villegas","year":"1999","journal-title":"J Immunol"},{"key":"10.1016\/j.mucimm.2026.100422_bb0290","doi-asserted-by":"crossref","first-page":"103","DOI":"10.1016\/j.cellimm.2006.02.004","article-title":"CD28 is required for induction and maintenance of immunological memory in toxin-reactive CD4+ T cells in vivo","volume":"238","author":"Fukada","year":"2005","journal-title":"Cell Immunol"},{"key":"10.1016\/j.mucimm.2026.100422_bb0295","doi-asserted-by":"crossref","first-page":"130","DOI":"10.1016\/j.cellimm.2010.09.008","article-title":"Differential requirements for CD80\/86-CD28 costimulation in primary and memory CD4 T cell responses to vaccinia virus","volume":"266","author":"Fuse","year":"2011","journal-title":"Cell Immunol"},{"key":"10.1016\/j.mucimm.2026.100422_bb0300","doi-asserted-by":"crossref","first-page":"2909","DOI":"10.4049\/jimmunol.1103231","article-title":"CD28 promotes CD4+ T cell clonal expansion during infection independently of its YMNM and PYAP motifs","volume":"189","author":"Pag\u00e1n","year":"2012","journal-title":"J Immunol"},{"key":"10.1016\/j.mucimm.2026.100422_bb0305","doi-asserted-by":"crossref","DOI":"10.1371\/journal.ppat.1005667","article-title":"CD4 T cell-derived IFN-\u03b3 plays a minimal role in control of pulmonary Mycobacterium tuberculosis infection and must be actively repressed by PD-1 to prevent lethal disease","volume":"12","author":"Sakai","year":"2016","journal-title":"PLoS Pathog"},{"key":"10.1016\/j.mucimm.2026.100422_bb0310","doi-asserted-by":"crossref","first-page":"1598","DOI":"10.4049\/jimmunol.1003304","article-title":"CD4 T cells promote rather than control tuberculosis in the absence of PD-1\u2013mediated inhibition","volume":"186","author":"Barber","year":"2011","journal-title":"J. Immunol."},{"key":"10.1016\/j.mucimm.2026.100422_bb0315","doi-asserted-by":"crossref","first-page":"1409","DOI":"10.1084\/jem.20091885","article-title":"Pathogen-specific regulatory T cells delay the arrival of effector T cells in the lung during early tuberculosis","volume":"207","author":"Shafiani","year":"2010","journal-title":"J Exp Med"},{"key":"10.1016\/j.mucimm.2026.100422_bb0320","doi-asserted-by":"crossref","DOI":"10.1371\/journal.ppat.1012339","article-title":"IL-10 suppresses T cell expansion while promoting tissue-resident memory cell formation during SARS-CoV-2 infection in rhesus macaques","volume":"20","author":"Nelson","year":"2024","journal-title":"PLoS Pathog"},{"key":"10.1016\/j.mucimm.2026.100422_bb0325","doi-asserted-by":"crossref","first-page":"3573","DOI":"10.1016\/j.cell.2021.04.048","article-title":"Integrated analysis of multimodal single-cell data","volume":"184","author":"Hao","year":"2021","journal-title":"Cell"}],"container-title":["Mucosal Immunology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S1933021926001248?httpAccept=text\/xml","content-type":"text\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S1933021926001248?httpAccept=text\/plain","content-type":"text\/plain","content-version":"vor","intended-application":"text-mining"}],"deposited":{"date-parts":[[2026,11,5]],"date-time":"2026-11-05T05:13:29Z","timestamp":1793855609000},"score":1,"resource":{"primary":{"URL":"https:\/\/linkinghub.elsevier.com\/retrieve\/pii\/S1933021926001248"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,10]]},"references-count":65,"alternative-id":["S1933021926001248"],"URL":"https:\/\/doi.org\/10.1016\/j.mucimm.2026.100422","relation":{},"ISSN":["1933-0219"],"issn-type":[{"value":"1933-0219","type":"print"}],"subject":[],"published":{"date-parts":[[2026,10]]},"assertion":[{"value":"Elsevier","name":"publisher","label":"This article is maintained by"},{"value":"Control of M. tuberculosis infection requires CD28 to generate lung-homing T cells and CTLA-4 to limit immunopathology","name":"articletitle","label":"Article Title"},{"value":"Mucosal Immunology","name":"journaltitle","label":"Journal Title"},{"value":"https:\/\/doi.org\/10.1016\/j.mucimm.2026.100422","name":"articlelink","label":"CrossRef DOI link to publisher maintained version"},{"value":"article","name":"content_type","label":"Content Type"},{"value":"Published by Elsevier Inc. on behalf of Society for Mucosal Immunology.","name":"copyright","label":"Copyright"}],"article-number":"100422"}}