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In germinal centers, T follicular helper (TFH) cells provide important help to B-cell antibody production but also contribute to HIV persistence. T follicular regulatory (TFR) cells, which inhibit the function of TFH cells, express similar surface markers. Since FOXP3 is the only marker that distinguishes TFR from TFH cells it is unknown whether the increase in TFH cells observed in HIV infection and HIV persistence may be partly due to an increase in TFR cells. Using multicolor flow cytometry to detect TFH and TFR cells in cryopreserved peripheral blood mononuclear cells from HIV-infected and non-infected participants in the UCLA Multicenter AIDS Cohort Study (MACS), we identified CD3+CXCR5+CD4+CD8\u2212BCL6+ peripheral blood TFH (pTFH) cells and CD3+CXCR5+CD4+CD8\u2212FOXP3+ peripheral blood TFR (pTFR) cells. Unlike TFR cells in germinal centers, pTFR cells do not express B cell lymphoma 6 (BCL6), a TFH cell master transcriptional regulator. Our major findings are that the frequency of pTFH cells, but not pTFR cells was higher in HIV-infected participants of the MACS and that pTFH cells expressed less CCR5 in HIV-infected MACS participants. Constitutive expression of CCR5 in TFR cells supports their potential to contribute to HIV persistence.<\/jats:p>","DOI":"10.3390\/cells12020296","type":"journal-article","created":{"date-parts":[[2023,1,12]],"date-time":"2023-01-12T05:03:03Z","timestamp":1673499783000},"page":"296","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":3,"title":["Characterization of T Follicular Helper Cells and T Follicular Regulatory Cells in HIV-Infected and Sero-Negative Individuals"],"prefix":"10.3390","volume":"12","author":[{"given":"Bradley","family":"Salvatore","sequence":"first","affiliation":[{"name":"Department of Microbiology, Immunology and Molecular Genetics, University of California Los Angeles (UCLA), Los Angeles, CA 90095, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Rachel","family":"Resop","sequence":"additional","affiliation":[{"name":"Department of Microbiology, Immunology and Molecular Genetics, University of California Los Angeles (UCLA), Los Angeles, CA 90095, USA"},{"name":"UCLA AIDS Institute, David Geffen School of Medicine, University of California Los Angeles (UCLA), Los Angeles, CA 90095, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Brent","family":"Gordon","sequence":"additional","affiliation":[{"name":"Department of Microbiology, Immunology and Molecular Genetics, University of California Los Angeles (UCLA), Los Angeles, CA 90095, USA"},{"name":"UCLA AIDS Institute, David Geffen School of Medicine, University of California Los Angeles (UCLA), Los Angeles, CA 90095, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-7605-5994","authenticated-orcid":false,"given":"Marta","family":"Epeldegui","sequence":"additional","affiliation":[{"name":"UCLA AIDS Institute, David Geffen School of Medicine, University of California Los Angeles (UCLA), Los Angeles, CA 90095, USA"},{"name":"Department of Obstetrics 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USA"},{"name":"Department of Surgical-Oncology, University of California Los Angeles (UCLA), Los Angeles, CA 90095, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2879-719X","authenticated-orcid":false,"given":"Alex","family":"Lam","sequence":"additional","affiliation":[{"name":"Department of Molecular Pharmacology, University of California Los Angeles (UCLA), Los Angeles, CA 90095, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ting-Ting","family":"Wu","sequence":"additional","affiliation":[{"name":"UCLA AIDS Institute, David Geffen School of Medicine, University of California Los Angeles (UCLA), Los Angeles, CA 90095, USA"},{"name":"Jonsson Comprehensive Cancer Center (JCCC), University of California Los Angeles (UCLA), Los Angeles, CA 90095, USA"},{"name":"Department of Molecular Pharmacology, University of California Los Angeles (UCLA), Los Angeles, CA 90095, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0825-6521","authenticated-orcid":false,"given":"Christel","family":"Uittenbogaart","sequence":"additional","affiliation":[{"name":"Department of Microbiology, Immunology and Molecular Genetics, University of California Los Angeles (UCLA), Los Angeles, CA 90095, USA"},{"name":"UCLA AIDS Institute, David Geffen School of Medicine, University of California Los Angeles (UCLA), Los Angeles, CA 90095, USA"},{"name":"Jonsson Comprehensive Cancer Center (JCCC), University of California Los Angeles (UCLA), Los Angeles, CA 90095, USA"},{"name":"Department of Pediatrics, University of California Los Angeles (UCLA), Los Angeles, CA 90095, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2023,1,12]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"868","DOI":"10.1126\/science.6189183","article-title":"Isolation of a T-lymphotropic retrovirus from a patient at risk for acquired immune deficiency syndrome (AIDS)","volume":"220","author":"Chermann","year":"1983","journal-title":"Science"},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"500","DOI":"10.1126\/science.6200936","article-title":"Frequent detection and isolation of cytopathic retroviruses (HTLV-III) from patients with AIDS and at risk for AIDS","volume":"224","author":"Gallo","year":"1984","journal-title":"Science"},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"12","DOI":"10.1016\/j.jaci.2014.05.026","article-title":"Recent developments in the search for a cure for HIV-1 infection: Targeting the latent reservoir for HIV-1","volume":"134","author":"Siliciano","year":"2014","journal-title":"J. Allergy Clin. Immunol."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"377","DOI":"10.1016\/j.immuni.2012.08.010","article-title":"Redefining the viral reservoirs that prevent HIV-1 eradication","volume":"37","author":"Eisele","year":"2012","journal-title":"Immunity"},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"1284","DOI":"10.1038\/nm1295-1284","article-title":"In vivo fate of HIV-1-infected T cells: Quantitative analysis of the transition to stable latency","volume":"1","author":"Chun","year":"1995","journal-title":"Nat. Med."},{"key":"ref_6","doi-asserted-by":"crossref","unstructured":"Tran, T.A., de Goer de Herve, M.G., Hendel-Chavez, H., Dembele, B., Le Nevot, E., Abbed, K., Pallier, C., Goujard, C., Gasnault, J., and Delfraissy, J.F. (2008). Resting regulatory CD4 T cells: A site of HIV persistence in patients on long-term effective antiretroviral therapy. PLoS ONE, 3.","DOI":"10.1371\/journal.pone.0003305"},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"143","DOI":"10.1084\/jem.20121932","article-title":"Follicular helper T cells serve as the major CD4 T cell compartment for HIV-1 infection, replication, and production","volume":"210","author":"Perreau","year":"2013","journal-title":"J. Exp. Med."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"621","DOI":"10.1146\/annurev-immunol-031210-101400","article-title":"Follicular helper CD4 T cells (TFH)","volume":"29","author":"Crotty","year":"2011","journal-title":"Annu. Rev. Immunol."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"1006","DOI":"10.1126\/science.1175870","article-title":"Bcl6 and Blimp-1 are reciprocal and antagonistic regulators of T follicular helper cell differentiation","volume":"325","author":"Johnston","year":"2009","journal-title":"Science"},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"377","DOI":"10.1016\/j.it.2010.07.001","article-title":"The elusive identity of T follicular helper cells","volume":"31","author":"Yu","year":"2010","journal-title":"Trends Immunol."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"1001","DOI":"10.1126\/science.1176676","article-title":"Bcl6 mediates the development of T follicular helper cells","volume":"325","author":"Nurieva","year":"2009","journal-title":"Science"},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"529","DOI":"10.1016\/j.immuni.2014.10.004","article-title":"T follicular helper cell differentiation, function, and roles in disease","volume":"41","author":"Crotty","year":"2014","journal-title":"Immunity"},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"983","DOI":"10.1038\/nm.2426","article-title":"Follicular regulatory T cells expressing Foxp3 and Bcl-6 suppress germinal center reactions","volume":"17","author":"Chung","year":"2011","journal-title":"Nat. Med."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"975","DOI":"10.1038\/nm.2425","article-title":"Foxp3+ follicular regulatory T cells control the germinal center response","volume":"17","author":"Linterman","year":"2011","journal-title":"Nat. Med."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"4553","DOI":"10.4049\/jimmunol.1101328","article-title":"Regulation of the germinal center reaction by Foxp3+ follicular regulatory T cells","volume":"187","author":"Wollenberg","year":"2011","journal-title":"J. Immunol."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"1151","DOI":"10.4049\/jimmunol.155.3.1151","article-title":"Immunologic self-tolerance maintained by activated T cells expressing IL-2 receptor alpha-chains (CD25). Breakdown of a single mechanism of self-tolerance causes various autoimmune diseases","volume":"155","author":"Sakaguchi","year":"1995","journal-title":"J. Immunol."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"61","DOI":"10.1093\/intimm\/dxt060","article-title":"Foxp3(+) T(reg) cells in humoral immunity","volume":"26","author":"Wing","year":"2014","journal-title":"Int. Immunol."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"2666","DOI":"10.1182\/blood-2014-07-585976","article-title":"Human T-follicular helper and T-follicular regulatory cell maintenance is independent of germinal centers","volume":"124","author":"Wallin","year":"2014","journal-title":"Blood"},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"227","DOI":"10.4110\/in.2014.14.5.227","article-title":"Regulatory T cells in B cell follicles","volume":"14","author":"Chang","year":"2014","journal-title":"Immune Netw."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"eaan1487","DOI":"10.1126\/sciimmunol.aan1487","article-title":"Human blood Tfr cells are indicators of ongoing humoral activity not fully licensed with suppressive function","volume":"2","author":"Fonseca","year":"2017","journal-title":"Sci. Immunol."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"659","DOI":"10.3389\/fimmu.2016.00659","article-title":"CD4 T Follicular Helper and Regulatory Cell Dynamics and Function in HIV Infection","volume":"7","author":"Miles","year":"2016","journal-title":"Front. Immunol."},{"key":"ref_22","doi-asserted-by":"crossref","unstructured":"Colantonio, A.D., Epeldegui, M., Jesiak, M., Jachimowski, L., Blom, B., and Uittenbogaart, C.H. (2011). IFN-alpha is constitutively expressed in the human thymus, but not in peripheral lymphoid organs. PLoS ONE, 6.","DOI":"10.1371\/journal.pone.0024252"},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"47","DOI":"10.1097\/PPO.0000000000000303","article-title":"A Pilot Trial of the Combination of Transgenic NY-ESO-1 -reactive Adoptive Cellular Therapy with Dendritic Cell vaccination with or without ipilimumab","volume":"24","author":"Nowicki","year":"2018","journal-title":"Cancer J."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"51","DOI":"10.2174\/1570162043485086","article-title":"Role of CD4 receptor down-regulation during HIV-1 infection","volume":"2","author":"Levesque","year":"2004","journal-title":"Curr. HIV Res."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"3271","DOI":"10.1172\/JCI64314","article-title":"Expansion of HIV-specific T follicular helper cells in chronic HIV infection","volume":"122","author":"Lindqvist","year":"2012","journal-title":"J. Clin. Investig."},{"key":"ref_26","doi-asserted-by":"crossref","unstructured":"Colineau, L., Rouers, A., Yamamoto, T., Xu, Y., Urrutia, A., Pham, H.P., Cardinaud, S., Samri, A., Dorgham, K., and Coulon, P.G. (2015). HIV-Infected Spleens Present Altered Follicular Helper T Cell (Tfh) Subsets and Skewed B Cell Maturation. PLoS ONE, 10.","DOI":"10.1371\/journal.pone.0140978"},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"8608","DOI":"10.1038\/ncomms9608","article-title":"Follicular regulatory T cells impair follicular T helper cells in HIV and SIV infection","volume":"6","author":"Miles","year":"2015","journal-title":"Nat. Commun."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"949","DOI":"10.1084\/jem.187.6.949","article-title":"The unenlarged lymph nodes of HIV-1-infected, asymptomatic patients with high CD4 T cell counts are sites for virus replication and CD4 T cell proliferation. The impact of highly active antiretroviral therapy","volume":"187","author":"Stellbrink","year":"1998","journal-title":"J. Exp. Med."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"1152","DOI":"10.1002\/eji.201546094","article-title":"Follicular regulatory T cells repress cytokine production by follicular helper T cells and optimize IgG responses in mice","volume":"46","author":"Wu","year":"2016","journal-title":"Eur. J. Immunol."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"25","DOI":"10.1111\/imm.12774","article-title":"T follicular regulatory cells in mice and men","volume":"152","author":"Maceiras","year":"2017","journal-title":"Immunology"},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"1373","DOI":"10.1084\/jem.193.12.1373","article-title":"Subspecialization of CXCR5+ T cells: B helper activity is focused in a germinal center-localized subset of CXCR5+ T cells","volume":"193","author":"Kim","year":"2001","journal-title":"J. Exp. Med."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"754","DOI":"10.1038\/nm.4113","article-title":"PD-1(+) and follicular helper T cells are responsible for persistent HIV-1 transcription in treated aviremic individuals","volume":"22","author":"Banga","year":"2016","journal-title":"Nat. Med."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"e1000766","DOI":"10.1371\/journal.ppat.1000766","article-title":"IFN-alpha-induced upregulation of CCR5 leads to expanded HIV tropism in vivo","volume":"6","author":"Stoddart","year":"2010","journal-title":"PLoS Pathog."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"e00430-17","DOI":"10.1128\/JVI.00430-17","article-title":"Follicular Regulatory T Cells Are Highly Permissive to R5-Tropic HIV-1","volume":"91","author":"Miller","year":"2017","journal-title":"J. Virol."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"152","DOI":"10.1038\/ni.2496","article-title":"The receptor PD-1 controls follicular regulatory T cells in the lymph nodes and blood","volume":"14","author":"Sage","year":"2013","journal-title":"Nat. Immunol."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"1249","DOI":"10.1038\/ni.3837","article-title":"Dynamic regulation of T follicular regulatory cell responses by interleukin 2 during influenza infection","volume":"18","author":"Botta","year":"2017","journal-title":"Nat. Immunol."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"269","DOI":"10.1093\/intimm\/dxh206","article-title":"A distinct role for ICOS-mediated co-stimulatory signaling in CD4+ and CD8+ T cell subsets","volume":"17","author":"Watanabe","year":"2005","journal-title":"Int. Immunol."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"304","DOI":"10.3389\/fimmu.2016.00304","article-title":"ICOS Co-Stimulation: Friend or Foe?","volume":"7","author":"Wikenheiser","year":"2016","journal-title":"Front. Immunol."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"1415","DOI":"10.1002\/eji.201747407","article-title":"An optimized method to measure human FOXP3(+) regulatory T cells from multiple tissue types using mass cytometry","volume":"48","author":"Dawson","year":"2018","journal-title":"Eur. J. Immunol."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"776","DOI":"10.1016\/j.jinf.2020.09.014","article-title":"Alterations of the frequency and functions of follicular regulatory T cells and related mechanisms in HIV infection","volume":"81","author":"Zhao","year":"2020","journal-title":"J. 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