{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,23]],"date-time":"2026-08-23T17:56:44Z","timestamp":1787507804185,"version":"build-2736575974"},"reference-count":60,"publisher":"MDPI AG","issue":"9","license":[{"start":{"date-parts":[[2025,4,22]],"date-time":"2025-04-22T00:00:00Z","timestamp":1745280000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"State Plan for Scientific and Technical Research and Innovation; co-funded by the ISCIII Sub-Directorate General for Research Assessment and Promotion and the European Regional Development Fund (ERDF)","award":["RD16\/0025\/0013"],"award-info":[{"award-number":["RD16\/0025\/0013"]}]},{"name":"State Plan for Scientific and Technical Research and Innovation; co-funded by the ISCIII Sub-Directorate General for Research Assessment and Promotion and the European Regional Development Fund (ERDF)","award":["PI16\/01769"],"award-info":[{"award-number":["PI16\/01769"]}]},{"name":"State Plan for Scientific and Technical Research and Innovation; co-funded by the ISCIII Sub-Directorate General for Research Assessment and Promotion and the European Regional Development Fund (ERDF)","award":["PI19\/01237"],"award-info":[{"award-number":["PI19\/01237"]}]},{"name":"State Plan for Scientific and Technical Research and Innovation; co-funded by the ISCIII Sub-Directorate General for Research Assessment and Promotion and the European Regional Development Fund (ERDF)","award":["PI19\/00973"],"award-info":[{"award-number":["PI19\/00973"]}]},{"name":"State Plan for Scientific and Technical Research and Innovation; co-funded by the ISCIII Sub-Directorate General for Research Assessment and Promotion and the European Regional Development Fund (ERDF)","award":["RD16\/0025\/0019"],"award-info":[{"award-number":["RD16\/0025\/0019"]}]},{"name":"ISCIII Sub-Directorate General for Research Assessment and Promotion and the European Regional Development Fund (ERDF)","award":["RD16\/0025\/0013"],"award-info":[{"award-number":["RD16\/0025\/0013"]}]},{"name":"ISCIII Sub-Directorate General for Research Assessment and Promotion and the European Regional Development Fund (ERDF)","award":["PI16\/01769"],"award-info":[{"award-number":["PI16\/01769"]}]},{"name":"ISCIII Sub-Directorate General for Research Assessment and Promotion and the European Regional Development Fund (ERDF)","award":["PI19\/01237"],"award-info":[{"award-number":["PI19\/01237"]}]},{"name":"ISCIII Sub-Directorate General for Research Assessment and Promotion and the European Regional Development Fund (ERDF)","award":["PI19\/00973"],"award-info":[{"award-number":["PI19\/00973"]}]},{"name":"ISCIII Sub-Directorate General for Research Assessment and Promotion and the European Regional Development Fund (ERDF)","award":["RD16\/0025\/0019"],"award-info":[{"award-number":["RD16\/0025\/0019"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["IJMS"],"abstract":"<jats:p>Elite controllers (ECs) represent a unique subset of people living with HIV (PLWHs), who can suppress viral replication without requiring antiretroviral therapy (ART). However, despite this viral control, ECs exhibit increased incidences of various comorbid conditions and heightened systemic inflammation, which has been linked to monocyte activation. In this study, we performed an in-depth phenotypic analysis of monocytes in a cohort of long-term ECs (LTECs) and compared them to non-controller patients with ART-mediated control of HIV replication and to non-controller patients with uncontrolled viral replication. A total of 67 participants were included: 22 LTECs, 15 non-controllers on ART (onART), 10 non-controllers without ART (offART), and 20 uninfected controls (UCs) as a reference group. Monocyte phenotypes were analyzed using spectral flow cytometry with a 13-marker panel. The data were analyzed using two approaches: (a) FCS Express software v.7 to define different subsets of monocytes and assess the levels of expression of eight different monocyte functional markers and (b) R software v.4.1.1 for unsupervised multidimensional analysis, including batch correction, dimensionality reduction, and clustering analysis. Monocyte phenotypic profiling was conducted using three different approaches: (1) assessment of monocyte subsets (classical, intermediate, and non-classical monocytes); (2) evaluation of the levels of expression of eight monocyte functional markers, and (3) characterization of monocyte clusters defined through the dimensionality reduction of flow cytometry data (56 different clusters). The monocyte phenotype of the onART group closely resembled that of the UC group. In contrast, LTECs exhibited important alterations in the monocyte phenotype compared to that of the UCs, including (a) an increased proportion of intermediate monocytes and a decreased proportion of classical monocytes (p &lt; 0.01), (b) altered expressions of functional markers across monocyte subsets (p &lt; 0.05), and (c) alterations in sixteen different monocyte clusters (twelve decreased and four increased, p &lt; 0.05). Many of these alterations were also observed when comparing the LTEC and onART groups. Our findings suggest that monocyte-driven mechanisms may contribute to HIV control in LTECs; however, some of these alterations could also promote systemic inflammation and immune activation. These observations provide a compelling rationale for considering therapeutic interventions in this unique population of PLWHs.<\/jats:p>","DOI":"10.3390\/ijms26093926","type":"journal-article","created":{"date-parts":[[2025,4,22]],"date-time":"2025-04-22T03:17:32Z","timestamp":1745291852000},"page":"3926","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":4,"title":["Long-Term Elite Controllers of HIV-1 Infection Exhibit a Deep Perturbation of Monocyte Homeostasis"],"prefix":"10.3390","volume":"26","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-7172-049X","authenticated-orcid":false,"given":"Jos\u00e9 M.","family":"Benito","sequence":"first","affiliation":[{"name":"HIV and Viral Hepatitis Research Laboratory, Instituto de Investigaci\u00f3n Sanitaria Fundaci\u00f3n Jim\u00e9nez D\u00edaz, Universidad Aut\u00f3noma de Madrid (IIS-FJD, UAM), 28003 Madrid, Spain"},{"name":"Hospital Universitario Rey Juan Carlos, 28933 M\u00f3stoles, Spain"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Daniel","family":"Jim\u00e9nez-Carretero","sequence":"additional","affiliation":[{"name":"Unidad de Bioinform\u00e1tica, Centro Nacional de Investigaciones Cardiovasculares (CNIC), 28029 Madrid, Spain"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jaime","family":"Valent\u00edn-Quiroga","sequence":"additional","affiliation":[{"name":"Grupo de Respuesta Inmune Innata, IdiPAZ, Hospital Universitario La Paz, 28046 Madrid, Spain"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0558-6541","authenticated-orcid":false,"given":"Ignacio","family":"Mahillo","sequence":"additional","affiliation":[{"name":"Department of Statistics, Instituto de Investigaci\u00f3n Sanitaria Fundaci\u00f3n Jim\u00e9nez D\u00edaz, Universidad Aut\u00f3noma de Madrid (IIS-FJD, UAM), 28049 Madrid, Spain"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jos\u00e9 M.","family":"Ligos","sequence":"additional","affiliation":[{"name":"Cytek Biosciences, Inc., Fremont, CA 94538, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Clara","family":"Restrepo","sequence":"additional","affiliation":[{"name":"HIV and Viral Hepatitis Research Laboratory, Instituto de Investigaci\u00f3n Sanitaria Fundaci\u00f3n Jim\u00e9nez D\u00edaz, Universidad Aut\u00f3noma de Madrid (IIS-FJD, UAM), 28003 Madrid, Spain"},{"name":"Hospital Universitario Rey Juan Carlos, 28933 M\u00f3stoles, Spain"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-8429-8358","authenticated-orcid":false,"given":"Alfonso","family":"Cabello","sequence":"additional","affiliation":[{"name":"Hospital Universitario Fundaci\u00f3n Jim\u00e9nez D\u00edaz, 28040 Madrid, Spain"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Eduardo","family":"L\u00f3pez-Collazo","sequence":"additional","affiliation":[{"name":"Grupo de Respuesta Inmune Innata, IdiPAZ, Hospital Universitario La Paz, 28046 Madrid, Spain"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-1881-1664","authenticated-orcid":false,"given":"F\u00e1tima","family":"S\u00e1nchez-Cabo","sequence":"additional","affiliation":[{"name":"Unidad de Bioinform\u00e1tica, Centro Nacional de Investigaciones Cardiovasculares (CNIC), 28029 Madrid, Spain"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Miguel","family":"G\u00f3rgolas","sequence":"additional","affiliation":[{"name":"Hospital Universitario Fundaci\u00f3n Jim\u00e9nez D\u00edaz, 28040 Madrid, Spain"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Norma","family":"Rall\u00f3n","sequence":"additional","affiliation":[{"name":"HIV and Viral Hepatitis Research Laboratory, Instituto de Investigaci\u00f3n Sanitaria Fundaci\u00f3n Jim\u00e9nez D\u00edaz, Universidad Aut\u00f3noma de Madrid (IIS-FJD, UAM), 28003 Madrid, Spain"},{"name":"Hospital Universitario Rey Juan Carlos, 28933 M\u00f3stoles, Spain"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2025,4,22]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"181","DOI":"10.1097\/COH.0b013e328345a328","article-title":"Elite controllers as a model of functional cure","volume":"6","author":"Autran","year":"2011","journal-title":"Curr. Opin. HIV AIDS"},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"839","DOI":"10.1038\/nm.4108","article-title":"International AIDS Society global scientific strategy: Towards an HIV cure 2016","volume":"22","author":"Deeks","year":"2016","journal-title":"Nat. Med."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"889","DOI":"10.1080\/21505594.2020.1788887","article-title":"Elite controllers: A heterogeneous group of HIV-infected patients","volume":"11","author":"Restrepo","year":"2020","journal-title":"Virulence"},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"1163","DOI":"10.1097\/QAD.0b013e32832b44c8","article-title":"Prevalence and comparative characteristics of long-term nonprogressors and HIV controller patients in the French Hospital Database on HIV","volume":"23","author":"Grabar","year":"2009","journal-title":"AIDS"},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"2409","DOI":"10.1097\/QAD.0b013e32835a9950","article-title":"Increased Coronary Atherosclerosis and Immune Activation in HIV-1 Elite Controllers","volume":"26","author":"Pereyra","year":"2012","journal-title":"AIDS"},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"1692","DOI":"10.1093\/infdis\/jiu809","article-title":"Hospitalization Rates and Reasons Among HIV Elite Controllers and Persons with Medically Controlled HIV Infection","volume":"211","author":"Crowell","year":"2015","journal-title":"J. Infect. Dis."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"126","DOI":"10.1086\/524143","article-title":"Relationship Between T Cell Activation and CD4+ T Cell Count in HIV-Seropositive Individuals with Undetectable Plasma HIV RNA Levels in the Absence of Therapy","volume":"197","author":"Hunt","year":"2008","journal-title":"J. Infect. Dis."},{"key":"ref_8","doi-asserted-by":"crossref","unstructured":"Hunt, P.W., Landay, A.L., Sinclair, E., Martinson, J.A., Hatano, H., Emu, B., Norris, P.J., Busch, M., Martin, J.N., and Brooks, C. (2011). A Low T Regulatory Cell Response May Contribute to Both Viral Control and Generalized Immune Activation in HIV Controllers. PLoS ONE, 6.","DOI":"10.1371\/journal.pone.0015924"},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"931","DOI":"10.1093\/infdis\/jit581","article-title":"Evidence for Innate Immune System Activation in HIV Type 1-Infected Elite Controllers","volume":"209","author":"Krishnan","year":"2014","journal-title":"J. Infect. Dis."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"579","DOI":"10.1089\/aid.2015.0226","article-title":"Cytokine and Chemokine Signature in Elite Versus Viremic Controllers Infected with HIV","volume":"32","author":"Platten","year":"2016","journal-title":"AIDS Res. Human. Retroviruses"},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"184","DOI":"10.1016\/j.imlet.2014.01.008","article-title":"Persistent inflammation in HIV infection: Established concepts, new perspectives","volume":"161","author":"Nasi","year":"2014","journal-title":"Immunol. Lett"},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"1118","DOI":"10.1007\/s12026-016-8862-2","article-title":"HIV infection: Focus on the innate immune cells","volume":"64","author":"Soares","year":"2016","journal-title":"Immunol. Res."},{"key":"ref_13","doi-asserted-by":"crossref","unstructured":"Wacleche, V.S., Tremblay, C.L., Routy, J.P., and Ancuta, P. (2018). The Biology of Monocytes and Dendritic Cells: Contribution to HIV Pathogenesis. Viruses, 10.","DOI":"10.3390\/v10020065"},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"2","DOI":"10.1038\/icb.2010.130","article-title":"Toward a functional characterization of blood monocytes","volume":"89","author":"Saha","year":"2011","journal-title":"Immunol. Cell Biol."},{"key":"ref_15","doi-asserted-by":"crossref","unstructured":"Boyette, L.B., Macedo, C., Hadi, K., Elinoff, B.D., Walters, J.T., Ramaswami, B., Chalasani, G., Taboas, J.M., Lakkis, F.G., and Metes, D.M. (2017). Phenotype, function, and differentiation potential of human monocyte subsets. PLoS ONE, 12.","DOI":"10.1371\/journal.pone.0176460"},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"499","DOI":"10.1111\/j.1365-2249.2011.04526.x","article-title":"Impaired phagocytosis among patients infected by the human immunodeficiency virus: Implication for a role of highly active anti-retroviral therapy","volume":"167","author":"Michailidis","year":"2012","journal-title":"Clin. Exp. Immunol."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"4109","DOI":"10.1128\/JVI.79.7.4109-4119.2005","article-title":"Impaired monocyte maturation in response to CpG oligodeoxynucleotide is related to viral RNA levels in human immunodeficiency virus disease and is at least partially mediated by deficiencies in alpha\/beta interferon responsiveness and production","volume":"79","author":"Jiang","year":"2005","journal-title":"JVI"},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"803","DOI":"10.1189\/jlb.1111579","article-title":"Expansion of a subset of CD14highCD16negCCR2low\/neg monocytes functionally similar to myeloid-derived suppressor cells during SIV and HIV infection","volume":"91","author":"Gama","year":"2012","journal-title":"J. Leucoc. Biol."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"569819","DOI":"10.1155\/2014\/569819","article-title":"Monocytes as regulators of inflammation and HIV-related comorbidities during cART","volume":"2014","author":"Anzinger","year":"2014","journal-title":"J. Immunol. Res."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"302","DOI":"10.1007\/s10875-012-9790-0","article-title":"Selective expansion of pro-inflammatory chemokine CCL2-loaded CD14+CD16+ monocytes subset in HIV-infected therapy na\u00efve individuals","volume":"33","author":"Ansari","year":"2013","journal-title":"J. Clin. Immunol."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"2259","DOI":"10.1182\/blood-2012-03-418681","article-title":"Pivotal role of M-DC8+ monocytes from viremic HIV-infected patients in TNF overproduction in response to microbial products","volume":"120","author":"Dutertre","year":"2012","journal-title":"Blood"},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"1396","DOI":"10.1093\/infdis\/jiu275","article-title":"Monocyte-activation phenotypes are associated with biomarkers of inflammation and coagulation in chronic HIV infection","volume":"210","author":"Wilson","year":"2014","journal-title":"J. Infect. Dis."},{"key":"ref_23","doi-asserted-by":"crossref","unstructured":"Chen, P., Su, B., Zhang, T., Zhu, X., Xia, W., Fu, Y., Zhao, G., Xia, H., Dai, L., and Sun, L. (2017). Perturbations of Monocyte subsets and Their association with T helper cell Differentiation in acute and chronic HIV-1-infected Patients. Front. Immunol., 8.","DOI":"10.3389\/fimmu.2017.00272"},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"e17202","DOI":"10.1016\/j.heliyon.2023.e17202","article-title":"Differential expression of chemokine receptors on monocytes in TB and HIV S","volume":"9","author":"Tamene","year":"2023","journal-title":"Heliyon"},{"key":"ref_25","doi-asserted-by":"crossref","unstructured":"Prabhu, V.M., Singh, A.K., Padwal, V., Nagar, V., Patil, P., and Patel, V. (2019). Monocyte Based Correlates of Immune Activation and Viremia in HIV-Infected Long-Term Non-Progressors. Front. Immunol., 10.","DOI":"10.3389\/fimmu.2019.02849"},{"key":"ref_26","doi-asserted-by":"crossref","unstructured":"McCausland, M.R., Juchnowski, S.M., Zidar, D.A., Kuritzkes, D.R., Andrade, A., Sieg, S.F., Lederman, M.M., and Funderburg, N.T. (2015). Altered Monocyte Phenotype in HIV-1 Infection Tends to Normalize with Integrase- Inhibitor-Based Antiretroviral Therapy. PLoS ONE, 10.","DOI":"10.1371\/journal.pone.0139474"},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"447","DOI":"10.1097\/QAD.0000000000002766","article-title":"Monocyte activation in persons living with HIV and tuberculosis coinfection","volume":"35","author":"Huamana","year":"2021","journal-title":"AIDS"},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"146","DOI":"10.1097\/00042560-200206010-00002","article-title":"Increased HLA-DR expression on peripheral blood monocytes in subsets of subjects with primary HIV infection is associated with elevated CD4 T-cell apoptosis and CD4 T-cell depletion","volume":"30","author":"Gascon","year":"2002","journal-title":"J. Acquir. Immune Defic. Syndr."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"1029","DOI":"10.1093\/cvr\/cvy292","article-title":"Loss of CXCR4 on non-classical monocytes in participants of theWomen\u2019s Interagency HIV Study (WIHS) with subclinical atherosclerosis","volume":"115","author":"Mueller","year":"2019","journal-title":"Cardiovasc. Res."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"ofx108","DOI":"10.1093\/ofid\/ofx108","article-title":"High Cellular Monocyte Activation in People Living With Human Immunodeficiency Virus on Combination Antiretroviral Therapy and Lifestyle-Matched Controls Is Associated with Greater Inflammation in Cerebrospinal Fluid","volume":"4","author":"Booiman","year":"2017","journal-title":"Open Forum Infect. Dis."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"565","DOI":"10.1007\/s12250-020-00332-0","article-title":"Monocyte\/Macrophage-Mediated Innate Immunity in HIV-1 Infection: From Early Response to Late Dysregulation and Links to Cardiovascular Diseases Onset","volume":"36","author":"Teer","year":"2021","journal-title":"Virol. Sin."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"51","DOI":"10.1016\/j.jim.2012.07.006","article-title":"Role of monocytes in mediating HIV-specific antibody-dependent cellular cytotoxicity","volume":"384","author":"Kramski","year":"2012","journal-title":"J. Immunol. Methods"},{"key":"ref_33","doi-asserted-by":"crossref","unstructured":"Van den Bergh, R., Morin, S., Hans Sass, H.J., Grzesiek, S., Vekemans, M., Florence, E., Tran, H.T.T., Imiru, R.G., Heyndrickx, L., and Vanham, G. (2012). Monocytes Contribute to Differential Immune Pressure on R5 Versus X4 HIV Through the Adipocytokine Visfatin\/ NAMPT. PLoS ONE, 7.","DOI":"10.1371\/journal.pone.0035074"},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"e50","DOI":"10.1182\/blood-2011-01-326827","article-title":"SuperSAGE evidence for CD14++CD16+ monocytes as a third monocyte subset","volume":"118","author":"Zawada","year":"2011","journal-title":"Blood"},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"21","DOI":"10.1146\/annurev-immunol-083122-035233","article-title":"Immune Responses in Controllers of HIV Infection","volume":"42","author":"Kwaa","year":"2024","journal-title":"Annu. Rev. Immunol."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"9463","DOI":"10.1128\/JVI.01009-10","article-title":"Leukocyte immunoglobulin-like receptors maintain unique antigen-presenting properties of circulating myeloid dendritic cells in HIV-1-infected elite controllers","volume":"84","author":"Huang","year":"2010","journal-title":"J. Virol."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"165","DOI":"10.1093\/infdis\/jiae460","article-title":"Intact HIV reservoir in monocytes is associated with cognitive function in virally suppressed women with HIV","volume":"231","author":"Rubin","year":"2025","journal-title":"J. Infect. Dis."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"1177","DOI":"10.1097\/QAD.0000000000003534","article-title":"Increased transmigration of intermediate monocytes associated with atherosclerotic burden in people with HIV on antiretroviral therapy","volume":"37","author":"Chow","year":"2023","journal-title":"AIDS"},{"key":"ref_39","first-page":"e146215","article-title":"Higher circulating intermediate monocytes are associated with cognitive function in women with HIV","volume":"6","author":"Veenhuis","year":"2021","journal-title":"JCI Insight"},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"52","DOI":"10.1016\/j.atherosclerosis.2013.10.021","article-title":"Reduced CD14 expression on classical monocytes and vascular endothelial adhesion markers independently associate with carotid artery intima media thickness in chronically HIV-1 infected adults on virologically suppressive anti-retroviral therapy","volume":"232","author":"Barbour","year":"2014","journal-title":"Atherosclerosis"},{"key":"ref_41","doi-asserted-by":"crossref","unstructured":"Schittenhelm, L., Hilkens, C.M., and Morrison, V.L. (2017). \u03b22 Integrins as Regulators of Dendritic Cell, Monocyte, and Macrophage Function. Front. Immunol., 8.","DOI":"10.3389\/fimmu.2017.01866"},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"106","DOI":"10.1111\/j.1365-2249.1993.tb05955.x","article-title":"CR1 (CD35) and CR3 (CD11b\/CD18) mediate infection of human monocytes and monocytic cell lines with complement-opsonized HIV independently of CD4","volume":"92","author":"Thieblemont","year":"1993","journal-title":"Clin. Exp. Immunol."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"2932","DOI":"10.1182\/blood.V97.10.2932","article-title":"Ligation of CD11b and CD11c beta(2) integrins by antibodies or soluble CD23 induces macrophage inflammatory protein 1alpha (MIP-1alpha) and MIP-1beta production in primary human monocytes through a pathway dependent on nuclear factor-kappaB","volume":"97","author":"Rezzonico","year":"2001","journal-title":"Blood"},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"843","DOI":"10.1097\/QAD.0b013e328351f756","article-title":"HIV infection induces age-related changes to monocytes and innate immune activation in young men that persist despite combination antiretroviral therapy","volume":"26","author":"Hearps","year":"2012","journal-title":"AIDS"},{"key":"ref_45","doi-asserted-by":"crossref","unstructured":"Hofer, T.P., van de Loosdrecht, A.A., Stahl-Hennig, C., Cassatella, M.A., and Ziegler-Heitbrock, L. (2019). 6-Sulfo LacNAc (Slan) as a Marker for Non-Classical Monocytes. Front. Immunol., 10.","DOI":"10.3389\/fimmu.2019.02052"},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"76","DOI":"10.2174\/138945011701151217110917","article-title":"The CCL2\/CCR2 Axis in the Pathogenesis of HIV-1 Infection: A New Cellular Target for Therapy?","volume":"17","author":"Covino","year":"2016","journal-title":"Curr. Drug Targets"},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"39","DOI":"10.1002\/rmv.369","article-title":"The influence of cytokines, chemokines and their receptors on HIV-1 replication in monocytes and macrophages","volume":"13","author":"Kedzierska","year":"2003","journal-title":"Rev. Med. Virol."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"1235","DOI":"10.1007\/s10238-022-00927-9","article-title":"Intermediate monocytes expansion and homing markers expression in COVID-19 patients associate with kidney dysfunction","volume":"23","author":"Dentone","year":"2023","journal-title":"Clin. Exp. Med."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"1215","DOI":"10.1042\/CS20170009","article-title":"Monocyte subtypes and the CCR2 chemokine receptor in cardiovascular disease","volume":"131","author":"Izar","year":"2017","journal-title":"Clin. Sci."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"e36","DOI":"10.1212\/NXI.0000000000000036","article-title":"CCR2 on CD14(+)CD16(+) monocytes is a biomarker of HIV-associated neurocognitive disorders","volume":"1","author":"Williams","year":"2014","journal-title":"Neurol. Neuroimmunol. Neuroinflamm."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"75S","DOI":"10.1016\/S0002-9610(05)81210-3","article-title":"The human leukocyte antigens and their relationship to infection","volume":"165","author":"Cheadle","year":"1993","journal-title":"Am. J. Surg."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"236","DOI":"10.1086\/599837","article-title":"Differential HIV epitope processing in monocytes and CD4 T cells affects cytotoxic T lymphocyte recognition","volume":"200","author":"Lazaro","year":"2009","journal-title":"J. Infect. Dis."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"154","DOI":"10.1093\/infdis\/jir214","article-title":"Soluble CD163 Made by Monocyte\/Macrophages Is a Novel Marker of HIV Activity in Early and Chronic Infection Prior to and After Anti-Retroviral Therapy","volume":"204","author":"Burdo","year":"2011","journal-title":"J. Infect. Dis."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"1227","DOI":"10.1093\/infdis\/jir520","article-title":"Soluble CD163, a novel marker of activated macrophages, is elevated and associated with noncalcified coronary plaque in HIV-infected patients","volume":"204","author":"Burdo","year":"2011","journal-title":"J. Infect. Dis."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"1387","DOI":"10.1097\/QAD.0b013e32836010bd","article-title":"Elevated sCD163 in plasma but not cerebrospinal fluid is amarker of neurocognitive impairment in HIV infection","volume":"27","author":"Burdo","year":"2013","journal-title":"AIDS"},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"1198","DOI":"10.1093\/infdis\/jiw263","article-title":"Plasma Soluble CD163 Level Independently Predicts All-Cause Mortality in HIV-1-Infected Individuals","volume":"214","author":"Knudsen","year":"2016","journal-title":"J. Infect. Dis."},{"key":"ref_57","doi-asserted-by":"crossref","unstructured":"Ancuta, P., Kamat, A., Kunstman, K.J., Kim, E.Y., Autissier, P., and Wurcel, A. (2008). Microbial translocation is associated with increased monocyte activation and dementia in AIDS patients. PLoS ONE, 3.","DOI":"10.1371\/journal.pone.0002516"},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"347","DOI":"10.1080\/07853890410033171","article-title":"CD163: A regulated hemoglobin scavenger receptor with a role in the anti-inflammatory response","volume":"36","author":"Moestrup","year":"2004","journal-title":"Ann. Med."},{"key":"ref_59","doi-asserted-by":"crossref","unstructured":"Tippett, E., Cheng, W.J., Westhorpe, C., Cameron, P.U., Brew, B.J., Lewin, S.R., Jaworowski, A., and Crowe, S.M. (2006). Differential Expression of CD163 on Monocyte Subsets in Healthy and HIV-1 Infected Individuals. 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