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J. Mol. Sci.","IJMS"],"abstract":"<jats:p>Antiretroviral therapy (ART) has proven effective in suppressing HIV-1 replication, but further development of HIV-1 inhibitors is continually driven by the challenge of drug resistance and viral adaptation. The HIV-1 capsid is a promising target for treatment due to its high sequence conservation as well as its crucial role in the viral life cycle. Recently, we have developed a novel capsid-targeting biologic that prevents HIV-1 replication by efficient degradation of newly synthesized capsid. Here, we have investigated the sensitivity to viral escape as well as the breadth of this biologic against HIV-1 subtypes. The capsid-targeting biologic efficiently blocked replication of different primary HIV-1 isolates, and continuous exposure of these viruses to the biologic resulted in viral breakthrough of two out of ten primary HIV-1 isolates tested. Notably, the breakthrough variants did not have amino acid changes in the nanobody epitope but primarily in the matrix region. The breakthrough variants remained sensitive to the biologic albeit to a lesser extent. In the absence of the biologic, breakthrough variants showed increased replication kinetics when compared to their parental virus, suggesting that adaption to the biologic is likely due to the increased viral production and that the target area of the biologic is too conserved for actual escape. This is further underscored by the broad specificity of the biologic as importantly the biologic blocked infection of different HIV-1 subtypes that occur worldwide (A, B, C, D, CRF01_AE, CRF02_AG). These results demonstrate the broad neutralization potential of anti-capsid biologics with a high barrier to resistance, making capsid-targeting inhibitors important for novel antiretroviral drug strategies worldwide.<\/jats:p>","DOI":"10.3390\/ijms27156883","type":"journal-article","created":{"date-parts":[[2026,8,3]],"date-time":"2026-08-03T12:57:56Z","timestamp":1785761876000},"page":"6883","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Capsid-Targeting Biologic Achieves Broad HIV-1 Neutralization with a High Barrier to Resistance"],"prefix":"10.3390","volume":"27","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-3781-8223","authenticated-orcid":false,"given":"Florence M.","family":"Stel","sequence":"first","affiliation":[{"name":"Department of Experimental Immunology, Amsterdam UMC Location AMC, University of Amsterdam, 1105 AZ Amsterdam, The Netherlands"},{"name":"Amsterdam Institute for Immunology and Infectious Diseases, 1105 AZ Amsterdam, The Netherlands"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Esther M.","family":"Zijlstra-Willems","sequence":"additional","affiliation":[{"name":"Department of Experimental Immunology, Amsterdam UMC Location AMC, University of Amsterdam, 1105 AZ Amsterdam, The Netherlands"},{"name":"Amsterdam Institute for Immunology and Infectious Diseases, 1105 AZ Amsterdam, The Netherlands"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ad C. van","family":"Nuenen","sequence":"additional","affiliation":[{"name":"Department of Experimental Immunology, Amsterdam UMC Location AMC, University of Amsterdam, 1105 AZ Amsterdam, The Netherlands"},{"name":"Amsterdam Institute for Immunology and Infectious Diseases, 1105 AZ Amsterdam, The Netherlands"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Brigitte D. M.","family":"Boeser-Nunnink","sequence":"additional","affiliation":[{"name":"Department of Experimental Immunology, Amsterdam UMC Location AMC, University of Amsterdam, 1105 AZ Amsterdam, The Netherlands"},{"name":"Amsterdam Institute for Immunology and Infectious Diseases, 1105 AZ Amsterdam, The Netherlands"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-5710-2839","authenticated-orcid":false,"given":"Teunis B. H.","family":"Geijtenbeek","sequence":"additional","affiliation":[{"name":"Department of Experimental Immunology, Amsterdam UMC Location AMC, University of Amsterdam, 1105 AZ Amsterdam, The Netherlands"},{"name":"Amsterdam Institute for Immunology and Infectious Diseases, 1105 AZ Amsterdam, The Netherlands"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-9429-7754","authenticated-orcid":false,"given":"Neeltje A.","family":"Kootstra","sequence":"additional","affiliation":[{"name":"Department of Experimental Immunology, Amsterdam UMC Location AMC, University of Amsterdam, 1105 AZ Amsterdam, The Netherlands"},{"name":"Amsterdam Institute for Immunology and Infectious Diseases, 1105 AZ Amsterdam, The Netherlands"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2026,8,1]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"13596535231201162","DOI":"10.1177\/13596535231201162","article-title":"HIV drug resistance in the era of contemporary antiretroviral therapy: A clinical perspective","volume":"28","author":"Carr","year":"2023","journal-title":"Antivir. Ther."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"55","DOI":"10.1126\/science.288.5463.55d","article-title":"HIV-1 nomenclature proposal","volume":"288","author":"Robertson","year":"2000","journal-title":"Science"},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"100912","DOI":"10.1016\/S2666-5247(24)00151-4","article-title":"Global and regional genetic diversity of HIV-1 in 2010-21: Systematic review and analysis of prevalence","volume":"5","author":"Nair","year":"2024","journal-title":"Lancet Microbe"},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"fuac040","DOI":"10.1093\/femsre\/fuac040","article-title":"New antiretroviral inhibitors and HIV-1 drug resistance: More focus on 90% HIV-1 isolates?","volume":"47","author":"Ndashimye","year":"2023","journal-title":"FEMS Microbiol. Rev."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"35936","DOI":"10.1038\/s41598-025-04144-8","article-title":"In vitro comparison of the susceptibilities of the same drug resistance mutations to reverse-transcriptase inhibitors of subtype B and CRF01_AE HIV-1 strains","volume":"15","author":"Li","year":"2025","journal-title":"Sci. Rep."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"ofv158","DOI":"10.1093\/ofid\/ofv158","article-title":"Global Comparison of Drug Resistance Mutations After First-Line Antiretroviral Therapy Across Human Immunodeficiency Virus-1 Subtypes","volume":"3","author":"Huang","year":"2016","journal-title":"Open Forum Infect. Dis."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"636","DOI":"10.1016\/j.coviro.2012.08.006","article-title":"Are subtype differences important in HIV drug resistance?","volume":"2","author":"Lessells","year":"2012","journal-title":"Curr. Opin. Virol."},{"key":"ref_8","doi-asserted-by":"crossref","unstructured":"Troyano-Hernaez, P., Reinosa, R., and Holguin, A. (2022). HIV Capsid Protein Genetic Diversity Across HIV-1 Variants and Impact on New Capsid-Inhibitor Lenacapavir. Front. Microbiol., 13.","DOI":"10.3389\/fmicb.2022.854974"},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"126","DOI":"10.1186\/1742-4690-10-126","article-title":"Functional conservation of HIV-1 Gag: Implications for rational drug design","volume":"10","author":"Li","year":"2013","journal-title":"Retrovirology"},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"477","DOI":"10.1016\/j.chembiol.2024.02.012","article-title":"Identification of clickable HIV-1 capsid-targeting probes for viral replication inhibition","volume":"31","author":"McFadden","year":"2024","journal-title":"Cell Chem. Biol."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"41","DOI":"10.1186\/s12977-021-00583-z","article-title":"Rotten to the core: Antivirals targeting the HIV-1 capsid core","volume":"18","author":"McFadden","year":"2021","journal-title":"Retrovirology"},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"849","DOI":"10.1126\/science.278.5339.849","article-title":"Structure of the carboxyl-terminal dimerization domain of the HIV-1 capsid protein","volume":"278","author":"Gamble","year":"1997","journal-title":"Science"},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"1944","DOI":"10.1128\/JVI.02698-15","article-title":"Mutations of Conserved Residues in the Major Homology Region Arrest Assembling HIV-1 Gag as a Membrane-Targeted Intermediate Containing Genomic RNA and Cellular Proteins","volume":"90","author":"Tanaka","year":"2016","journal-title":"J. Virol."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"471","DOI":"10.1038\/nrmicro3503","article-title":"HIV-1 capsid: The multifaceted key player in HIV-1 infection","volume":"13","author":"Campbell","year":"2015","journal-title":"Nat. Rev. Microbiol."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"119","DOI":"10.1007\/s12250-019-00095-3","article-title":"Multiple Roles of HIV-1 Capsid during the Virus Replication Cycle","volume":"34","author":"Novikova","year":"2019","journal-title":"Virol. Sin."},{"key":"ref_16","doi-asserted-by":"crossref","unstructured":"Zila, V., Muller, T.G., Muller, B., and Krausslich, H.G. (2021). HIV-1 capsid is the key orchestrator of early viral replication. PLoS Pathog., 17.","DOI":"10.1371\/journal.ppat.1010109"},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"741","DOI":"10.1016\/j.tim.2017.04.004","article-title":"Capsid-Dependent Host Factors in HIV-1 Infection","volume":"25","author":"Yamashita","year":"2017","journal-title":"Trends Microbiol."},{"key":"ref_18","doi-asserted-by":"crossref","unstructured":"Dick, R.A., Mallery, D.L., Vogt, V.M., and James, L.C. (2018). IP6 Regulation of HIV Capsid Assembly, Stability, and Uncoating. Viruses, 10.","DOI":"10.3390\/v10110640"},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"1285","DOI":"10.1016\/S0092-8674(00)81823-1","article-title":"Crystal structure of human cyclophilin A bound to the amino-terminal domain of HIV-1 capsid","volume":"87","author":"Gamble","year":"1996","journal-title":"Cell"},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"836","DOI":"10.1038\/s41586-023-06969-7","article-title":"The HIV capsid mimics karyopherin engagement of FG-nucleoporins","volume":"626","author":"Dickson","year":"2024","journal-title":"Nature"},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"32","DOI":"10.1186\/s12977-021-00577-x","article-title":"HIV-1 capsid variability: Viral exploitation and evasion of capsid-binding molecules","volume":"18","author":"Saito","year":"2021","journal-title":"Retrovirology"},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"5439","DOI":"10.1128\/JVI.77.9.5439-5450.2003","article-title":"Functional surfaces of the human immunodeficiency virus type 1 capsid protein","volume":"77","author":"Stray","year":"2003","journal-title":"J. Virol."},{"key":"ref_23","doi-asserted-by":"crossref","unstructured":"Rihn, S.J., Wilson, S.J., Loman, N.J., Alim, M., Bakker, S.E., Bhella, D., Gifford, R.J., Rixon, F.J., and Bieniasz, P.D. (2013). Extreme genetic fragility of the HIV-1 capsid. PLoS Pathog., 9.","DOI":"10.1371\/journal.ppat.1003461"},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"10","DOI":"10.1128\/AAC.02057-20","article-title":"Absence of Lenacapavir (GS-6207) Phenotypic Resistance in HIV Gag Cleavage Site Mutants and in Isolates with Resistance to Existing Drug Classes","volume":"65","author":"Margot","year":"2021","journal-title":"Antimicrob. Agents Chemother."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"614","DOI":"10.1038\/s41586-020-2443-1","article-title":"Clinical targeting of HIV capsid protein with a long-acting small molecule","volume":"584","author":"Link","year":"2020","journal-title":"Nature"},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"1793","DOI":"10.1056\/NEJMoa2115542","article-title":"Capsid Inhibition with Lenacapavir in Multidrug-Resistant HIV-1 Infection","volume":"386","author":"DeJesus","year":"2022","journal-title":"N. Engl. J. Med."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"989","DOI":"10.1093\/jac\/dkab503","article-title":"Phenotypic resistance to lenacapavir and monotherapy efficacy in a proof-of-concept clinical study","volume":"77","author":"Margot","year":"2022","journal-title":"J. Antimicrob. Chemother."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"e15","DOI":"10.1016\/S2352-3018(22)00291-0","article-title":"Lenacapavir administered every 26 weeks or daily in combination with oral daily antiretroviral therapy for initial treatment of HIV: A randomised, open-label, active-controlled, phase 2 trial","volume":"10","author":"Gupta","year":"2023","journal-title":"Lancet HIV"},{"key":"ref_29","first-page":"1878","article-title":"Emergence of in-vitro resistance to lenacapavir is similar across HIV-1 subtypes","volume":"39","author":"Naik","year":"2025","journal-title":"AIDS"},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"360","DOI":"10.1126\/science.abb4808","article-title":"Structural and mechanistic bases for a potent HIV-1 capsid inhibitor","volume":"370","author":"Bester","year":"2020","journal-title":"Science"},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"e0180422","DOI":"10.1128\/mbio.01804-22","article-title":"Structural and Mechanistic Bases of Viral Resistance to HIV-1 Capsid Inhibitor Lenacapavir","volume":"13","author":"Bester","year":"2022","journal-title":"mBio"},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"1261","DOI":"10.1056\/NEJMoa2411858","article-title":"Twice-Yearly Lenacapavir for HIV Prevention in Men and Gender-Diverse Persons","volume":"392","author":"Kelley","year":"2025","journal-title":"N. Engl. J. Med."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"e497","DOI":"10.1016\/S2352-3018(23)00113-3","article-title":"Efficacy and safety of the novel capsid inhibitor lenacapavir to treat multidrug-resistant HIV: Week 52 results of a phase 2\/3 trial","volume":"10","author":"Ogbuagu","year":"2023","journal-title":"Lancet HIV"},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"e282","DOI":"10.1016\/S2352-3018(25)00329-7","article-title":"Standardisation of monitoring routines for new long-acting antiretrovirals in development","volume":"13","author":"Hoffmann","year":"2026","journal-title":"Lancet HIV"},{"key":"ref_35","doi-asserted-by":"crossref","unstructured":"Stel, F.M., Verkuijlen, C.E.J., Zijlstra-Willems, E.M., Dekkers, G., Heukers, R., Kootstra, N.A., and Geijtenbeek, T.B.H. (2026). Anti-capsid nanobodies function as HIV-1 capsid inhibitors by intracellular capsid degradation. Commun. Biol.","DOI":"10.1038\/s42003-026-10662-6"},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"6045","DOI":"10.1073\/pnas.0800159105","article-title":"TRIM21 is an IgG receptor that is structurally, thermodynamically, and kinetically conserved","volume":"105","author":"Keeble","year":"2008","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"19985","DOI":"10.1073\/pnas.1014074107","article-title":"Antibodies mediate intracellular immunity through tripartite motif-containing 21 (TRIM21)","volume":"107","author":"Mallery","year":"2010","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"7932","DOI":"10.1128\/JVI.00757-08","article-title":"Autologous neutralizing humoral immunity and evolution of the viral envelope in the course of subtype B human immunodeficiency virus type 1 infection","volume":"82","author":"Bunnik","year":"2008","journal-title":"J. Virol."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"1569","DOI":"10.1172\/JCI7953C1","article-title":"Differential coreceptor expression allows for independent evolution of non-syncytium-inducing and syncytium-inducing HIV-1","volume":"106","author":"Blaak","year":"2000","journal-title":"J. Clin. Investig."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"479","DOI":"10.1021\/acsinfecdis.6b00189","article-title":"Unravelling the Molecular Basis of High Affinity Nanobodies against HIV p24: In Vitro Functional, Structural, and in Silico Insights","volume":"3","author":"Gray","year":"2017","journal-title":"ACS Infect. Dis."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"1692","DOI":"10.1016\/j.cell.2017.10.033","article-title":"A Method for the Acute and Rapid Degradation of Endogenous Proteins","volume":"171","author":"Clift","year":"2017","journal-title":"Cell"},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"89","DOI":"10.1016\/j.virusres.2014.07.001","article-title":"How HIV-1 Gag assembles in cells: Putting together pieces of the puzzle","volume":"193","author":"Lingappa","year":"2014","journal-title":"Virus Res."},{"key":"ref_43","doi-asserted-by":"crossref","unstructured":"Troyer, R.M., McNevin, J., Liu, Y., Zhang, S.C., Krizan, R.W., Abraha, A., Tebit, D.M., Zhao, H., Avila, S., and Lobritz, M.A. (2009). Variable fitness impact of HIV-1 escape mutations to cytotoxic T lymphocyte (CTL) response. PLoS Pathog., 5.","DOI":"10.1371\/journal.ppat.1000365"},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"12179","DOI":"10.1128\/JVI.01277-07","article-title":"Evolution of human immunodeficiency virus type 1 cytotoxic T-lymphocyte epitopes: Fitness-balanced escape","volume":"81","author":"Liu","year":"2007","journal-title":"J. Virol."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"565","DOI":"10.1016\/j.jmb.2011.03.037","article-title":"Flexible nature and specific functions of the HIV-1 nucleocapsid protein","volume":"410","author":"Darlix","year":"2011","journal-title":"J. Mol. Biol."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"100321","DOI":"10.1016\/j.jbc.2021.100321","article-title":"Structural characterization of HIV-1 matrix mutants implicated in envelope incorporation","volume":"296","author":"Eastep","year":"2021","journal-title":"J. Biol. Chem."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"5311","DOI":"10.1128\/jvi.68.8.5311-5320.1994","article-title":"Single amino acid changes in the human immunodeficiency virus type 1 matrix protein block virus particle production","volume":"68","author":"Freed","year":"1994","journal-title":"J. Virol."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1006\/viro.1998.9398","article-title":"HIV-1 gag proteins: Diverse functions in the virus life cycle","volume":"251","author":"Freed","year":"1998","journal-title":"Virology"},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"4966","DOI":"10.1128\/jvi.66.8.4966-4971.1992","article-title":"The matrix protein of human immunodeficiency virus type 1 is required for incorporation of viral envelope protein into mature virions","volume":"66","author":"Yu","year":"1992","journal-title":"J. Virol."},{"key":"ref_50","doi-asserted-by":"crossref","unstructured":"Tedbury, P.R., Ablan, S.D., and Freed, E.O. (2013). Global rescue of defects in HIV-1 envelope glycoprotein incorporation: Implications for matrix structure. PLoS Pathog., 9.","DOI":"10.1371\/journal.ppat.1003739"},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"198","DOI":"10.1006\/jmbi.1994.1719","article-title":"Three-dimensional structure of the human immunodeficiency virus type 1 matrix protein","volume":"244","author":"Massiah","year":"1994","journal-title":"J. Mol. Biol."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"47","DOI":"10.1016\/j.virol.2007.06.046","article-title":"Characterization of replication defects induced by mutations in the basic domain and C-terminus of HIV-1 matrix","volume":"369","author":"Bhatia","year":"2007","journal-title":"Virology"},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"2855","DOI":"10.1128\/JVI.74.6.2855-2866.2000","article-title":"Role of the Gag matrix domain in targeting human immunodeficiency virus type 1 assembly","volume":"74","author":"Ono","year":"2000","journal-title":"J. Virol."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"512","DOI":"10.1016\/j.jmb.2011.04.015","article-title":"Molecular determinants that regulate plasma membrane association of HIV-1 Gag","volume":"410","author":"Chukkapalli","year":"2011","journal-title":"J. Mol. Biol."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"1637","DOI":"10.1016\/j.jmb.2016.03.005","article-title":"Structural and Molecular Determinants of Membrane Binding by the HIV-1 Matrix Protein","volume":"428","author":"Mercredi","year":"2016","journal-title":"J. Mol. Biol."},{"key":"ref_56","doi-asserted-by":"crossref","unstructured":"Prince, J.L., Claiborne, D.T., Carlson, J.M., Schaefer, M., Yu, T., Lahki, S., Prentice, H.A., Yue, L., Vishwanathan, S.A., and Kilembe, W. (2012). Role of transmitted Gag CTL polymorphisms in defining replicative capacity and early HIV-1 pathogenesis. PLoS Pathog., 8.","DOI":"10.1371\/journal.ppat.1003041"},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"e2200794119","DOI":"10.1073\/pnas.2200794119","article-title":"Atomic view of the HIV-1 matrix lattice; implications on virus assembly and envelope incorporation","volume":"119","author":"Samal","year":"2022","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"341","DOI":"10.1128\/jvi.70.1.341-351.1996","article-title":"Domains of the human immunodeficiency virus type 1 matrix and gp41 cytoplasmic tail required for envelope incorporation into virions","volume":"70","author":"Freed","year":"1996","journal-title":"J. Virol."},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"4116","DOI":"10.1128\/JVI.72.5.4116-4126.1998","article-title":"Role of matrix in an early postentry step in the human immunodeficiency virus type 1 life cycle","volume":"72","author":"Kiernan","year":"1998","journal-title":"J. Virol."},{"key":"ref_60","doi-asserted-by":"crossref","unstructured":"Takemura, T., and Murakami, T. (2012). Functional constraints on HIV-1 capsid: Their impacts on the viral immune escape potency. Front. Microbiol., 3.","DOI":"10.3389\/fmicb.2012.00369"},{"key":"ref_61","doi-asserted-by":"crossref","unstructured":"Sinay, Z., Tiggeler, A., Palstra, R.J., and Mahmoudi, T. (2026). Tools to Quantify and Characterize the Persistent Reservoir in People with HIV-1: Focus on Non-B Subtypes. Viruses, 18.","DOI":"10.3390\/v18010110"},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"4979","DOI":"10.1038\/s41467-025-60001-2","article-title":"Efficient mRNA delivery to resting T cells to reverse HIV latency","volume":"16","author":"Cevaal","year":"2025","journal-title":"Nat. Commun."},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"3273","DOI":"10.1074\/jbc.M806889200","article-title":"General strategy to humanize a camelid single-domain antibody and identification of a universal humanized nanobody scaffold","volume":"284","author":"Vincke","year":"2009","journal-title":"J. Biol. Chem."},{"key":"ref_64","doi-asserted-by":"crossref","unstructured":"Ali, S.A., Teow, S.Y., Omar, T.C., Khoo, A.S., Choon, T.S., and Yusoff, N.M. (2016). A Cell Internalizing Antibody Targeting Capsid Protein (p24) Inhibits the Replication of HIV-1 in T Cells Lines and PBMCs: A Proof of Concept Study. PLoS ONE, 11.","DOI":"10.1371\/journal.pone.0145986"},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"7478","DOI":"10.1128\/jvi.71.10.7478-7487.1997","article-title":"Coreceptor usage of primary human immunodeficiency virus type 1 isolates varies according to biological phenotype","volume":"71","author":"Bjorndal","year":"1997","journal-title":"J. Virol."},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"363","DOI":"10.1038\/nprot.2008.3","article-title":"Isolation and propagation of HIV-1 on peripheral blood mononuclear cells","volume":"3","author":"Schuitemaker","year":"2008","journal-title":"Nat. Protoc."},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"149","DOI":"10.1016\/0042-6822(89)90521-7","article-title":"Detection and Subtyping of Hiv-1 Isolates with a Panel of Characterized Monoclonal-Antibodies to Hiv P24gag","volume":"171","author":"Tersmette","year":"1989","journal-title":"Virology"},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"1298","DOI":"10.1073\/pnas.0337541100","article-title":"Abrogation of postentry restriction of HIV-1-based lentiviral vector transduction in simian cells","volume":"100","author":"Kootstra","year":"2003","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"269","DOI":"10.1016\/j.virol.2011.01.010","article-title":"Comparison of in vivo and in vitro evolution of CCR5 to CXCR4 coreceptor use of primary human immunodeficiency virus type 1 variants","volume":"412","author":"Setiawan","year":"2011","journal-title":"Virology"},{"key":"ref_70","unstructured":"Hall, T.A. (1999). BioEdit: A user-friendly biological sequence alignment editor and analysis program for Windows 95\/98\/NT. Nucleic Acids Symposium Series, Oxford University Press."}],"container-title":["International Journal of Molecular Sciences"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/1422-0067\/27\/15\/6883\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,8,5]],"date-time":"2026-08-05T09:17:01Z","timestamp":1785921421000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/1422-0067\/27\/15\/6883"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,8,1]]},"references-count":70,"journal-issue":{"issue":"15","published-online":{"date-parts":[[2026,8]]}},"alternative-id":["ijms27156883"],"URL":"https:\/\/doi.org\/10.3390\/ijms27156883","relation":{},"ISSN":["1422-0067"],"issn-type":[{"value":"1422-0067","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,8,1]]}}}