{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,28]],"date-time":"2026-09-28T23:43:16Z","timestamp":1790638996388,"version":"4.1.0"},"reference-count":95,"publisher":"MDPI AG","issue":"5","license":[{"start":{"date-parts":[[2021,5,11]],"date-time":"2021-05-11T00:00:00Z","timestamp":1620691200000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Cells"],"abstract":"<jats:p>HIV-1 is a complex retrovirus that is adapted to replicate in cells of the immune system. To do so, HIV-1, like other viruses, developed strategies to use several cellular processes to its advantage, but had also to come to terms with an arsenal of cellular innate defense proteins, or antiviral factors, that target more or less efficiently, virtually every step of the virus replicative cycle. Among antiviral restriction factors, the family of interferon-induced transmembrane proteins (IFITMs) has emerged as a crucial component of cellular innate defenses for their ability to interfere with both early and late phases of viral replication by inhibiting cellular and viral membranes fusion. Here, we review the enormous advances made since the discovery of IFITMs as interferon-regulated genes more than thirty years ago, with a particular focus on HIV-1 and on the elements that modulate its susceptibility or resistance towards members of this family. Given the recent advances of the field in the elucidation of the mechanism of IFITM inhibition and on the mechanism(s) of viral resistance, we expect that future years will bring novel insights into the definition of the multiple facets of IFITMs and on their possible use for novel therapeutical approaches.<\/jats:p>","DOI":"10.3390\/cells10051171","type":"journal-article","created":{"date-parts":[[2021,5,11]],"date-time":"2021-05-11T22:53:40Z","timestamp":1620773620000},"page":"1171","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":21,"title":["Membrane Interference Against HIV-1 by Intrinsic Antiviral Factors: The Case of IFITMs"],"prefix":"10.3390","volume":"10","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-5614-3997","authenticated-orcid":false,"given":"Federico","family":"Marziali","sequence":"first","affiliation":[{"name":"Centre International de Recherche en Infectiologie (CIRI), Universit\u00e9 de Lyon, Inserm U1111, CNRS, UMR5308, ENS de Lyon, Universite\u0301 Claude Bernard Lyon 1, 46 All\u00e9e d\u2019Italie, 69007 Lyon, France"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0892-0488","authenticated-orcid":false,"given":"Andrea","family":"Cimarelli","sequence":"additional","affiliation":[{"name":"Centre International de Recherche en Infectiologie (CIRI), Universit\u00e9 de Lyon, Inserm U1111, CNRS, UMR5308, ENS de Lyon, Universite\u0301 Claude Bernard Lyon 1, 46 All\u00e9e d\u2019Italie, 69007 Lyon, France"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2021,5,11]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"a006940","DOI":"10.1101\/cshperspect.a006940","article-title":"HIV Restriction Factors and Mechanisms of Evasion","volume":"2","author":"Malim","year":"2012","journal-title":"Cold Spring Harb. Perspect. Med."},{"key":"ref_2","doi-asserted-by":"crossref","unstructured":"Ramirez, P.W., Sharma, S., Singh, R., Stoneham, C.A., Vollbrecht, T., and Guatelli, J. (2019). Plasma Membrane-Associated Restriction Factors and Their Counteraction by HIV-1 Accessory Proteins. Cells, 8.","DOI":"10.3390\/cells8091020"},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"211","DOI":"10.2174\/1570162X14666151216145521","article-title":"Cellular Antiviral Factors that Target Particle Infectivity of HIV-1","volume":"14","author":"Goffinet","year":"2016","journal-title":"Curr. HIV Res."},{"key":"ref_4","doi-asserted-by":"crossref","unstructured":"Alm\u00e9n, S.M., Bringeland, N., Fredriksson, R., and Schi\u00f6th, H.B. (2012). The Dispanins: A Novel Gene Family of Ancient Origin that Contains 14 Human Members. PLoS ONE, 7.","DOI":"10.1371\/journal.pone.0031961"},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"745","DOI":"10.1016\/0092-8674(84)90270-8","article-title":"Transcriptional and Posttranscriptional Regulation of Interferon-Induced Gene Expression in Human Cells","volume":"38","author":"Friedman","year":"1984","journal-title":"Cell"},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"183","DOI":"10.1089\/jir.2010.0112","article-title":"The Small Interferon-Induced Transmembrane Genes and Proteins","volume":"31","author":"Siegrist","year":"2011","journal-title":"J. Interferon Cytokine Res."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"840","DOI":"10.1073\/pnas.86.3.840","article-title":"A Single DNA Response Element Can Confer Inducibility by Both Alpha- and Gamma-Interferons","volume":"86","author":"Reid","year":"1989","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"1136","DOI":"10.1002\/ajmg.a.36409","article-title":"Two Mutations in IFITM5 Causing Distinct Forms of Osteogenesis Imperfecta","volume":"164","author":"Valencia","year":"2014","journal-title":"Am. J. Med. Genet. A"},{"key":"ref_9","doi-asserted-by":"crossref","unstructured":"Coomer, C.A., Rahman, K., and Compton, A.A. (2021). CD225 Proteins: A Family Portrait of Fusion Regulators. Trends Genet.","DOI":"10.1016\/j.tig.2021.01.004"},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"103","DOI":"10.1186\/s12977-014-0103-y","article-title":"IFITM Proteins Are Incorporated onto HIV-1 Virion Particles and Negatively Imprint Their Infectivity","volume":"11","author":"Tartour","year":"2014","journal-title":"Retrovirology"},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"261","DOI":"10.1146\/annurev-virology-031413-085537","article-title":"IFITM-Family Proteins: The Cell\u2019s First Line of Antiviral Defense","volume":"1","author":"Bailey","year":"2014","journal-title":"Annu. Rev. Virol."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"24029","DOI":"10.1038\/srep24029","article-title":"Combined Approaches of EPR and NMR Illustrate Only One Transmembrane Helix in the Human IFITM3","volume":"6","author":"Ling","year":"2016","journal-title":"Sci. Rep."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"745","DOI":"10.1016\/j.devcel.2005.10.010","article-title":"IFITM\/Mil\/Fragilis Family Proteins IFITM1 and IFITM3 Play Distinct Roles in Mouse Primordial Germ Cell Homing and Repulsion","volume":"9","author":"Tanaka","year":"2005","journal-title":"Dev. Cell"},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"4688","DOI":"10.1128\/MCB.00272-08","article-title":"Normal Germ Line Establishment in Mice Carrying a Deletion of the Ifitm\/Fragilis Gene Family Cluster","volume":"28","author":"Lange","year":"2008","journal-title":"Mol. Cell. Biol."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"266","DOI":"10.1002\/cncr.23551","article-title":"Gene-Expression Profiling in Chinese Patients with Colon Cancer by Coupling Experimental and Bioinformatic Genomewide Gene-Expression Analyses: Identification and Validation of IFITM3 as a Biomarker of Early Colon Carcinogenesis","volume":"113","author":"Fan","year":"2008","journal-title":"Cancer"},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"95","DOI":"10.3892\/ol.2010.197","article-title":"Detection of Fecal Interferon-Induced Transmembrane Protein Messenger RNA for Colorectal Cancer Screening","volume":"2","author":"Miyamoto","year":"2011","journal-title":"Oncol. Lett."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"618","DOI":"10.1093\/ajcp\/aqz205","article-title":"Expression and Prognostic Value of IFIT1 and IFITM3 in Head and Neck Squamous Cell Carcinoma","volume":"153","author":"Li","year":"2020","journal-title":"Am. J. Clin. Pathol."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"2648","DOI":"10.3892\/or.2014.3522","article-title":"Mechanism and Biological Significance of the Overexpression of IFITM3 in Gastric Cancer","volume":"32","author":"Hu","year":"2014","journal-title":"Oncol. Rep."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"38","DOI":"10.1038\/s41417-019-0093-y","article-title":"High IFITM3 Expression Predicts Adverse Prognosis in Acute Myeloid Leukemia","volume":"27","author":"Liu","year":"2020","journal-title":"Cancer Gene Ther."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"433","DOI":"10.1007\/s11033-019-05146-2","article-title":"IFITM3\/STAT3 Axis Promotes Glioma Cells Invasion and Is Modulated by TGF-\u03b2","volume":"47","author":"Wang","year":"2020","journal-title":"Mol. Biol. Rep."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"1299","DOI":"10.1002\/2211-5463.12479","article-title":"IFITM3 Promotes Hepatocellular Carcinoma Invasion and Metastasis by Regulating MMP9 through P38\/MAPK Signaling","volume":"8","author":"Min","year":"2018","journal-title":"FEBS Open Bio"},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"592","DOI":"10.1111\/1759-7714.12488","article-title":"Overexpression of IFITM3 Predicts Poor Prognosis in Stage IIA Esophageal Squamous Cell Carcinoma after Ivor Lewis Esophagectomy","volume":"8","author":"Jia","year":"2017","journal-title":"Thorac. Cancer"},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"337","DOI":"10.1111\/1759-7714.12451","article-title":"Interferon Induced Transmembrane Protein 3 Regulates the Growth and Invasion of Human Lung Adenocarcinoma","volume":"8","author":"Zhang","year":"2017","journal-title":"Thorac. Cancer"},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"1949","DOI":"10.1158\/0008-5472.CAN-05-2731","article-title":"Identification of the IFITM Family as a New Molecular Marker in Human Colorectal Tumors","volume":"66","author":"Andreu","year":"2006","journal-title":"Cancer Res."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"593245","DOI":"10.3389\/fonc.2020.593245","article-title":"Malignancy and IFITM3: Friend or Foe?","volume":"10","author":"Rajapaksa","year":"2020","journal-title":"Front. Oncol."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"375","DOI":"10.1089\/jir.1996.16.375","article-title":"Partial Inhibition of Vesicular Stomatitis Virus by the Interferon-Induced Human 9-27 Protein","volume":"16","author":"Alber","year":"1996","journal-title":"J. Interferon Cytokine Res."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"1243","DOI":"10.1016\/j.cell.2009.12.017","article-title":"The IFITM Proteins Mediate Cellular Resistance to Influenza A H1N1 Virus, West Nile Virus, and Dengue Virus","volume":"139","author":"Brass","year":"2009","journal-title":"Cell"},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"1255","DOI":"10.1016\/j.cell.2009.12.018","article-title":"A Physical and Regulatory Map of Host-Influenza Interactions Reveals Pathways in H1N1 Infection","volume":"139","author":"Shapira","year":"2009","journal-title":"Cell"},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"229","DOI":"10.1007\/s40588-018-0103-0","article-title":"Antiviral Protection by IFITM3 In Vivo","volume":"5","author":"Zani","year":"2018","journal-title":"Curr. Clin. Microbiol. Rep."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"4937","DOI":"10.1016\/j.jmb.2013.09.024","article-title":"IFITMs Restrict the Replication of Multiple Pathogenic Viruses","volume":"425","author":"Perreira","year":"2013","journal-title":"J. Mol. Biol."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"53","DOI":"10.1186\/s12977-017-0377-y","article-title":"More than Meets the I: The Diverse Antiviral and Cellular Functions of Interferon-Induced Transmembrane Proteins","volume":"14","author":"Shi","year":"2017","journal-title":"Retrovirology"},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"2126","DOI":"10.1128\/JVI.01531-10","article-title":"The IFITM Proteins Inhibit HIV-1 Infection","volume":"85","author":"Lu","year":"2011","journal-title":"J. Virol."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"736","DOI":"10.1016\/j.chom.2014.11.001","article-title":"IFITM Proteins Incorporated into HIV-1 Virions Impair Viral Fusion and Spread","volume":"16","author":"Compton","year":"2014","journal-title":"Cell Host Microbe"},{"key":"ref_34","doi-asserted-by":"crossref","unstructured":"Tartour, K., Nguyen, X.-N., Appourchaux, R., Assil, S., Barateau, V., Bloyet, L.-M., Burlaud Gaillard, J., Confort, M.-P., Escudero-Perez, B., and Gruffat, H. (2017). Interference with the Production of Infectious Viral Particles and Bimodal Inhibition of Replication Are Broadly Conserved Antiviral Properties of IFITMs. PLoS Pathog., 13.","DOI":"10.1371\/journal.ppat.1006610"},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"14551","DOI":"10.1038\/s41598-018-32785-5","article-title":"IFITM Proteins Inhibit HIV-1 Protein Synthesis","volume":"8","author":"Lee","year":"2018","journal-title":"Sci. Rep."},{"key":"ref_36","doi-asserted-by":"crossref","unstructured":"Li, K., Markosyan, R.M., Zheng, Y.-M., Golfetto, O., Bungart, B., Li, M., Ding, S., He, Y., Liang, C., and Lee, J.C. (2013). IFITM Proteins Restrict Viral Membrane Hemifusion. PLoS Pathog., 9.","DOI":"10.1371\/journal.ppat.1003124"},{"key":"ref_37","doi-asserted-by":"crossref","unstructured":"Desai, T.M., Marin, M., Chin, C.R., Savidis, G., Brass, A.L., and Melikyan, G.B. (2014). IFITM3 Restricts Influenza A Virus Entry by Blocking the Formation of Fusion PoRes. Following Virus-Endosome Hemifusion. PLoS Pathog., 10.","DOI":"10.1371\/journal.ppat.1004048"},{"key":"ref_38","doi-asserted-by":"crossref","unstructured":"Suddala, K.C., Lee, C.C., Meraner, P., Marin, M., Markosyan, R.M., Desai, T.M., Cohen, F.S., Brass, A.L., and Melikyan, G.B. (2019). Interferon-Induced Transmembrane Protein 3 Blocks Fusion of Sensitive but Not Resistant Viruses by Partitioning into Virus-Carrying Endosomes. PLoS Pathog., 15.","DOI":"10.1371\/journal.ppat.1007532"},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"259","DOI":"10.1038\/s41589-018-0213-2","article-title":"IFITM3 Directly Engages and Shuttles Incoming Virus Particles to Lysosomes","volume":"15","author":"Spence","year":"2019","journal-title":"Nat. Chem. Biol."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"360","DOI":"10.1016\/j.cub.2005.01.049","article-title":"Endocytic Delivery to Lysosomes Mediated by Concurrent Fusion and Kissing Events in Living Cells","volume":"15","author":"Bright","year":"2005","journal-title":"Curr. Biol."},{"key":"ref_41","first-page":"1289","article-title":"Virus Entry at a Glance","volume":"126","author":"Yamauchi","year":"2013","journal-title":"J. Cell. Sci."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"878","DOI":"10.1016\/j.tim.2019.06.002","article-title":"Molecular Mechanism of HIV-1 Entry","volume":"27","author":"Chen","year":"2019","journal-title":"Trends Microbiol."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"13697","DOI":"10.1128\/JVI.01828-12","article-title":"The N-Terminal Region of IFITM3 Modulates Its Antiviral Activity by Regulating IFITM3 Cellular Localization","volume":"86","author":"Jia","year":"2012","journal-title":"J. Virol."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"429","DOI":"10.1016\/j.chom.2016.08.006","article-title":"Resistance of Transmitted Founder HIV-1 to IFITM-Mediated Restriction","volume":"20","author":"Foster","year":"2016","journal-title":"Cell Host Microbe"},{"key":"ref_45","doi-asserted-by":"crossref","unstructured":"Appourchaux, R., Delpeuch, M., Zhong, L., Burlaud-Gaillard, J., Tartour, K., Savidis, G., Brass, A., Etienne, L., Roingeard, P., and Cimarelli, A. (2019). Functional Mapping of Regions Involved in the Negative Imprinting of Virion Particle Infectivity and in Target Cell Protection by Interferon-Induced Transmembrane Protein 3 against HIV-1. J. Virol., 93.","DOI":"10.1128\/JVI.01716-18"},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"433","DOI":"10.1016\/j.cell.2009.02.046","article-title":"HIV Enters Cells via Endocytosis and Dynamin-Dependent Fusion with Endosomes","volume":"137","author":"Miyauchi","year":"2009","journal-title":"Cell"},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/j.micpath.2014.11.003","article-title":"Endocytosis of Human Immunodeficiency Virus 1 (HIV-1) in Astrocytes: A Fiery Path to Its Destination","volume":"78","author":"Chauhan","year":"2015","journal-title":"Microb. Pathog."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"1005","DOI":"10.1016\/S0960-9822(00)00654-0","article-title":"Endocytic Entry of HIV-1","volume":"10","author":"Fackler","year":"2000","journal-title":"Curr. Biol."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"8110","DOI":"10.1128\/JVI.00815-13","article-title":"Productive Entry of HIV-1 during Cell-to-Cell Transmission via Dynamin-Dependent Endocytosis","volume":"87","author":"Sloan","year":"2013","journal-title":"J. Virol."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"13956","DOI":"10.1128\/JVI.01543-14","article-title":"HIV-1 Entry in SupT1-R5, CEM-Ss, and Primary CD4+ T Cells Occurs at the Plasma Membrane and Does Not Require Endocytosis","volume":"88","author":"Herold","year":"2014","journal-title":"J. Virol."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"659","DOI":"10.1016\/0092-8674(87)90542-3","article-title":"PH-Independent HIV Entry into CD4-Positive T Cells via Virus Envelope Fusion to the Plasma Membrane","volume":"49","author":"Stein","year":"1987","journal-title":"Cell"},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"513","DOI":"10.1002\/j.1460-2075.1988.tb02839.x","article-title":"Human Immunodeficiency Virus Infection of CD4-Bearing Cells Occurs by a PH-Independent Mechanism","volume":"7","author":"McClure","year":"1988","journal-title":"EMBO J."},{"key":"ref_53","doi-asserted-by":"crossref","unstructured":"Hulseberg, C.E., F\u00e9n\u00e9ant, L., Szyma\u0144ska, K.M., and White, J.M. (2018). Lamp1 Increases the Efficiency of Lassa Virus Infection by Promoting Fusion in Less Acidic Endosomal Compartments. mBio, 9.","DOI":"10.1128\/mBio.01818-17"},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"7677","DOI":"10.1128\/JVI.00560-08","article-title":"Different Mechanisms of Cell Entry by Human-Pathogenic Old World and New World Arenaviruses","volume":"82","author":"Rojek","year":"2008","journal-title":"J. Virol."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"e58537","DOI":"10.7554\/eLife.58537","article-title":"Homology-Guided Identification of a Conserved Motif Linking the Antiviral Functions of IFITM3 to Its Oligomeric State","volume":"9","author":"Rahman","year":"2020","journal-title":"Elife"},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"145","DOI":"10.1016\/j.celrep.2015.08.055","article-title":"IFITM Proteins Restrict HIV-1 Infection by Antagonizing the Envelope Glycoprotein","volume":"13","author":"Yu","year":"2015","journal-title":"Cell Rep."},{"key":"ref_57","doi-asserted-by":"crossref","unstructured":"Ahi, Y.S., Yimer, D., Shi, G., Majdoul, S., Rahman, K., Rein, A., and Compton, A.A. (2020). IFITM3 Reduces Retroviral Envelope Abundance and Function and Is Counteracted by GlycoGag. mBio, 11.","DOI":"10.1128\/mBio.03088-19"},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"218","DOI":"10.1038\/nature15400","article-title":"SERINC3 and SERINC5 Restrict HIV-1 Infectivity and Are Counteracted by Nef","volume":"526","author":"Usami","year":"2015","journal-title":"Nature"},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"7013","DOI":"10.1074\/jbc.RA119.007662","article-title":"The Retroviral Accessory Proteins S2, Nef, and GlycoMA Use Similar Mechanisms for Antagonizing the Host Restriction Factor SERINC5","volume":"294","author":"Ahmad","year":"2019","journal-title":"J. Biol. Chem."},{"key":"ref_60","doi-asserted-by":"crossref","unstructured":"Guo, X., Steink\u00fchler, J., Marin, M., Li, X., Lu, W., Dimova, R., and Melikyan, G.B. (2021). Interferon-Induced Transmembrane Protein 3 Blocks Fusion of Diverse Enveloped Viruses by Altering Mechanical Properties of Cell Membranes. ACS Nano.","DOI":"10.1101\/2020.06.25.171280"},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"838","DOI":"10.1016\/j.bbrc.2006.06.152","article-title":"Laurdan Fluorescence Senses Mechanical Strain in the Lipid Bilayer Membrane","volume":"347","author":"Zhang","year":"2006","journal-title":"Biochem. Biophys. Res. Commun."},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"895","DOI":"10.1016\/j.celrep.2013.10.033","article-title":"Amphotericin B Increases Influenza A Virus Infection by Preventing IFITM3-Mediated Restriction","volume":"5","author":"Lin","year":"2013","journal-title":"Cell Rep."},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"368","DOI":"10.1016\/j.tim.2011.03.007","article-title":"Multifaceted Roles for Lipids in Viral Infection","volume":"19","author":"Heaton","year":"2011","journal-title":"Trends Microbiol."},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"1740","DOI":"10.15252\/embr.201744100","article-title":"IFITM 3 RequiRes. An Amphipathic Helix for Antiviral Activity","volume":"18","author":"Chesarino","year":"2017","journal-title":"EMBO Rep."},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"452","DOI":"10.1016\/j.chom.2013.03.006","article-title":"The Antiviral Effector IFITM3 Disrupts Intracellular Cholesterol Homeostasis to Block Viral Entry","volume":"13","author":"Choi","year":"2013","journal-title":"Cell Host Microbe"},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"3683","DOI":"10.3390\/v6093683","article-title":"IFITM Proteins Inhibit Entry Driven by the MERS-Coronavirus Spike Protein: Evidence for Cholesterol-Independent Mechanisms","volume":"6","author":"Wrensch","year":"2014","journal-title":"Viruses"},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"952","DOI":"10.1016\/j.bbalip.2016.02.009","article-title":"VAP, a Versatile Access Point for the Endoplasmic Reticulum: Review and Analysis of FFAT-like Motifs in the VAPome","volume":"1861","author":"Murphy","year":"2016","journal-title":"Biochim. Biophys. Acta"},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"3052","DOI":"10.1242\/jcs.028696","article-title":"FFAT Rescues VAPA-Mediated Inhibition of ER-to-Golgi Transport and VAPB-Mediated ER Aggregation","volume":"121","author":"Prosser","year":"2008","journal-title":"J. Cell. Sci."},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"e104369","DOI":"10.15252\/embj.2019104369","article-title":"FFAT Motif Phosphorylation Controls Formation and Lipid Transfer Function of Inter-Organelle Contacts","volume":"39","author":"Martinet","year":"2020","journal-title":"EMBO J."},{"key":"ref_70","doi-asserted-by":"crossref","first-page":"1689","DOI":"10.1007\/s00018-020-03604-w","article-title":"ORP\/Osh Mediate Cross-Talk between ER-Plasma Membrane Contact Site Components and Plasma Membrane SNAREs","volume":"78","author":"Trimbuch","year":"2021","journal-title":"Cell Mol. Life Sci."},{"key":"ref_71","doi-asserted-by":"crossref","unstructured":"Quon, E., Sere, Y.Y., Chauhan, N., Johansen, J., Sullivan, D.P., Dittman, J.S., Rice, W.J., Chan, R.B., Di Paolo, G., and Beh, C.T. (2018). Endoplasmic Reticulum-Plasma Membrane Contact Sites Integrate Sterol and Phospholipid Regulation. PLoS Biol., 16.","DOI":"10.1371\/journal.pbio.2003864"},{"key":"ref_72","doi-asserted-by":"crossref","unstructured":"Zhang, S., Nguyen, H.T., Ding, H., Wang, J., Zou, S., Liu, L., Guha, D., Gabuzda, D., Ho, D.D., and Kappes, J.C. (2021). Dual Pathways of Human Immunodeficiency Virus Type 1 Envelope Glycoprotein Trafficking Modulate the Selective Exclusion of Uncleaved Oligomers from Virions. J. Virol., 95.","DOI":"10.1128\/JVI.01369-20"},{"key":"ref_73","doi-asserted-by":"crossref","unstructured":"Drouin, A., Migraine, J., Durand, M.-A., Moreau, A., Burlaud-Gaillard, J., Beretta, M., Roingeard, P., Bouvin-Pley, M., and Braibant, M. (2020). Escape of HIV-1 Envelope Glycoprotein from the Restriction of Infection by IFITM3. J. Virol.","DOI":"10.1128\/JVI.01994-20"},{"key":"ref_74","doi-asserted-by":"crossref","first-page":"6626","DOI":"10.1073\/pnas.1304288110","article-title":"Phenotypic Properties of Transmitted Founder HIV-1","volume":"110","author":"Parrish","year":"2013","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_75","doi-asserted-by":"crossref","unstructured":"Wang, Y., Pan, Q., Ding, S., Wang, Z., Yu, J., Finzi, A., Liu, S.-L., and Liang, C. (2017). The V3 Loop of HIV-1 Env Determines Viral Susceptibility to IFITM3 Impairment of Viral Infectivity. J. Virol., 91.","DOI":"10.1128\/JVI.02441-16"},{"key":"ref_76","doi-asserted-by":"crossref","unstructured":"Beitari, S., Pan, Q., Finzi, A., and Liang, C. (2020). Differential PressuRes. of SERINC5 and IFITM3 on HIV-1 Envelope Glycoprotein over the Course of HIV-1 Infection. J. Virol., 94.","DOI":"10.1128\/JVI.00514-20"},{"key":"ref_77","doi-asserted-by":"crossref","unstructured":"Magnus, C., and Regoes, R.R. (2010). Estimating the Stoichiometry of HIV Neutralization. PLoS Comput. Biol., 6.","DOI":"10.1371\/journal.pcbi.1000713"},{"key":"ref_78","doi-asserted-by":"crossref","first-page":"12132","DOI":"10.1128\/JVI.79.19.12132-12147.2005","article-title":"Stoichiometry of Envelope Glycoprotein Trimers in the Entry of Human Immunodeficiency Virus Type 1","volume":"79","author":"Yang","year":"2005","journal-title":"J. Virol."},{"key":"ref_79","doi-asserted-by":"crossref","unstructured":"Brandenberg, O.F., Magnus, C., Rusert, P., Regoes, R.R., and Trkola, A. (2015). Different Infectivity of HIV-1 Strains Is Linked to Number of Envelope Trimers Required for Entry. PLoS Pathog., 11.","DOI":"10.1371\/journal.ppat.1004595"},{"key":"ref_80","doi-asserted-by":"crossref","first-page":"5752","DOI":"10.1128\/JVI.03187-14","article-title":"Structure and Dynamics of the Native HIV-1 Env Trimer","volume":"89","author":"Munro","year":"2015","journal-title":"J. Virol."},{"key":"ref_81","doi-asserted-by":"crossref","first-page":"655","DOI":"10.1016\/j.tim.2020.03.007","article-title":"The Conformational States of the HIV-1 Envelope Glycoproteins","volume":"28","author":"Wang","year":"2020","journal-title":"Trends Microbiol."},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"635","DOI":"10.1016\/j.chembiol.2017.04.007","article-title":"A Versatile Tool for Live-Cell Imaging and Super-Resolution Nanoscopy Studies of HIV-1 Env Distribution and Mobility","volume":"24","author":"Sakin","year":"2017","journal-title":"Cell Chem. Biol."},{"key":"ref_83","doi-asserted-by":"crossref","first-page":"524","DOI":"10.1126\/science.1226359","article-title":"Maturation-Dependent HIV-1 Surface Protein Redistribution Revealed by Fluorescence Nanoscopy","volume":"338","author":"Chojnacki","year":"2012","journal-title":"Science"},{"key":"ref_84","doi-asserted-by":"crossref","first-page":"348","DOI":"10.1074\/jbc.RA119.009481","article-title":"The Lipid Membrane of HIV-1 Stabilizes the Viral Envelope Glycoproteins and Modulates Their Sensitivity to Antibody Neutralization","volume":"295","author":"Salimi","year":"2020","journal-title":"J. Biol. Chem."},{"key":"ref_85","doi-asserted-by":"crossref","unstructured":"Pacheco, B., Alsahafi, N., Debbeche, O., Pr\u00e9vost, J., Ding, S., Chapleau, J.-P., Herschhorn, A., Madani, N., Princiotto, A., and Melillo, B. (2017). Residues in the Gp41 Ectodomain Regulate HIV-1 Envelope Glycoprotein Conformational Transitions Induced by Gp120-Directed Inhibitors. J. Virol., 91.","DOI":"10.1128\/JVI.02219-16"},{"key":"ref_86","doi-asserted-by":"crossref","first-page":"191","DOI":"10.1126\/science.aaa9804","article-title":"HIV-1 ENVELOPE. Effect of the Cytoplasmic Domain on Antigenic Characteristics of HIV-1 Envelope Glycoprotein","volume":"349","author":"Chen","year":"2015","journal-title":"Science"},{"key":"ref_87","doi-asserted-by":"crossref","unstructured":"Alfadhli, A., Staubus, A.O., Tedbury, P.R., Novikova, M., Freed, E.O., and Barklis, E. (2019). Analysis of HIV-1 Matrix-Envelope Cytoplasmic Tail Interactions. J. Virol., 93.","DOI":"10.1128\/JVI.01079-19"},{"key":"ref_88","doi-asserted-by":"crossref","first-page":"25269","DOI":"10.1073\/pnas.1910008116","article-title":"Single-Molecule Imaging of HIV-1 Envelope Glycoprotein Dynamics and Gag Lattice Association Exposes Determinants Responsible for Virus Incorporation","volume":"116","author":"Pezeshkian","year":"2019","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_89","doi-asserted-by":"crossref","first-page":"32105","DOI":"10.1073\/pnas.2012197117","article-title":"Cholesterol 25-Hydroxylase Suppresses SARS-CoV-2 Replication by Blocking Membrane Fusion","volume":"117","author":"Zang","year":"2020","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_90","doi-asserted-by":"crossref","first-page":"274","DOI":"10.1016\/j.molmed.2015.02.007","article-title":"APOBEC3 Genes: Retroviral Restriction Factors to Cancer Drivers","volume":"21","author":"Henderson","year":"2015","journal-title":"Trends Mol. Med."},{"key":"ref_91","doi-asserted-by":"crossref","first-page":"519","DOI":"10.1016\/S0092-8674(03)00112-0","article-title":"Membrane Fusion","volume":"112","author":"Jahn","year":"2003","journal-title":"Cell"},{"key":"ref_92","doi-asserted-by":"crossref","unstructured":"Wee, Y.S., Weis, J.J., Gahring, L.C., Rogers, S.W., and Weis, J.H. (2015). Age-Related Onset of Obesity Corresponds with Metabolic Dysregulation and Altered Microglia Morphology in Mice Deficient for Ifitm Proteins. PLoS ONE, 10.","DOI":"10.1371\/journal.pone.0123218"},{"key":"ref_93","doi-asserted-by":"crossref","first-page":"491","DOI":"10.1038\/s41586-020-2884-6","article-title":"IFITM3 Functions as a PIP3 Scaffold to Amplify PI3K Signalling in B Cells","volume":"588","author":"Lee","year":"2020","journal-title":"Nature"},{"key":"ref_94","doi-asserted-by":"crossref","first-page":"176","DOI":"10.1126\/science.aaw7733","article-title":"IFITM Proteins Inhibit Placental Syncytiotrophoblast Formation and Promote Fetal Demise","volume":"365","author":"Buchrieser","year":"2019","journal-title":"Science"},{"key":"ref_95","doi-asserted-by":"crossref","first-page":"19844","DOI":"10.1074\/jbc.AC119.010611","article-title":"Interferon-Induced Transmembrane Proteins Inhibit Cell Fusion Mediated by Trophoblast Syncytins","volume":"294","author":"Zani","year":"2019","journal-title":"J. Biol. Chem."}],"container-title":["Cells"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2073-4409\/10\/5\/1171\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,11]],"date-time":"2025-10-11T05:59:27Z","timestamp":1760162367000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2073-4409\/10\/5\/1171"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2021,5,11]]},"references-count":95,"journal-issue":{"issue":"5","published-online":{"date-parts":[[2021,5]]}},"alternative-id":["cells10051171"],"URL":"https:\/\/doi.org\/10.3390\/cells10051171","relation":{},"ISSN":["2073-4409"],"issn-type":[{"value":"2073-4409","type":"electronic"}],"subject":[],"published":{"date-parts":[[2021,5,11]]}}}