{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,7]],"date-time":"2026-08-07T23:44:19Z","timestamp":1786146259183,"version":"build-2736575974"},"reference-count":69,"publisher":"Elsevier BV","issue":"7","license":[{"start":{"date-parts":[[2026,7,1]],"date-time":"2026-07-01T00:00:00Z","timestamp":1782864000000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/tdm\/userlicense\/1.0\/"},{"start":{"date-parts":[[2026,7,1]],"date-time":"2026-07-01T00:00:00Z","timestamp":1782864000000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/legal\/tdmrep-license"},{"start":{"date-parts":[[2026,5,13]],"date-time":"2026-05-13T00:00:00Z","timestamp":1778630400000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0\/"}],"funder":[{"DOI":"10.13039\/100008496","name":"The Catholic University of America","doi-asserted-by":"publisher","id":[{"id":"10.13039\/100008496","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000054","name":"National Institutes of Health National Cancer Institute","doi-asserted-by":"publisher","award":["R15CA267890-01"],"award-info":[{"award-number":["R15CA267890-01"]}],"id":[{"id":"10.13039\/100000054","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100019077","name":"Mount Sinai Children's Center Foundation","doi-asserted-by":"publisher","award":["2200138"],"award-info":[{"award-number":["2200138"]}],"id":[{"id":"10.13039\/100019077","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100009633","name":"National Institute of Child Health and Human Development","doi-asserted-by":"publisher","award":["1ZIA-HD008955"],"award-info":[{"award-number":["1ZIA-HD008955"]}],"id":[{"id":"10.13039\/100009633","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100009633","name":"National Institute of Child Health and Human Development","doi-asserted-by":"publisher","award":["ZIA HD000072"],"award-info":[{"award-number":["ZIA HD000072"]}],"id":[{"id":"10.13039\/100009633","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000001","name":"National Science Foundation","doi-asserted-by":"publisher","id":[{"id":"10.13039\/100000001","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"publisher","id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["jbc.org","elsevier.com","sciencedirect.com"],"crossmark-restriction":true},"short-container-title":["Journal of Biological Chemistry"],"published-print":{"date-parts":[[2026,7]]},"DOI":"10.1016\/j.jbc.2026.113164","type":"journal-article","created":{"date-parts":[[2026,5,15]],"date-time":"2026-05-15T23:56:25Z","timestamp":1778889385000},"page":"113164","update-policy":"https:\/\/doi.org\/10.1016\/elsevier_cm_policy","source":"Crossref","is-referenced-by-count":0,"title":["The multiple sclerosis drug fingolimod induces pore-like membrane defects, contributing to its antimicrobial activity"],"prefix":"10.1016","volume":"302","author":[{"given":"Antara","family":"Syam","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Benjamin","family":"Rees","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sebastian","family":"Cuervo","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Fengtian","family":"Xue","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Alexander J.","family":"Sodt","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ekaterina M.","family":"Nestorovich","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Tatiana K.","family":"Rostovtseva","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sergey M.","family":"Bezrukov","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-9196-3280","authenticated-orcid":false,"given":"John S.","family":"Choy","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"78","reference":[{"key":"10.1016\/j.jbc.2026.113164_bib1","doi-asserted-by":"crossref","first-page":"373","DOI":"10.1038\/nrneurol.2010.76","article-title":"Fingolimod is a potential novel therapy for multiple sclerosis","volume":"6","author":"Aktas","year":"2010","journal-title":"Nat. Rev. Neurol."},{"key":"10.1016\/j.jbc.2026.113164_bib2","doi-asserted-by":"crossref","first-page":"2071","DOI":"10.1016\/0006-2952(91)90340-B","article-title":"Sphingoid bases as endogenous cationic amphiphilic \u201cdrugs\u201d","volume":"42","author":"Liscovitch","year":"1991","journal-title":"Biochem. Pharmacol."},{"key":"10.1016\/j.jbc.2026.113164_bib3","article-title":"New insights into the intracellular distribution pattern of cationic amphiphilic drugs","volume":"7","author":"Vater","year":"2017","journal-title":"Sci. Rep."},{"key":"10.1016\/j.jbc.2026.113164_bib4","doi-asserted-by":"crossref","first-page":"53","DOI":"10.1177\/019262339702500111","article-title":"Cationic amphiphilic drug-induced phospholipidosis","volume":"25","author":"Halliwell","year":"1997","journal-title":"Toxicol. Pathol."},{"key":"10.1016\/j.jbc.2026.113164_bib5","doi-asserted-by":"crossref","DOI":"10.1128\/mbio.02191-22","article-title":"TCA and SSRI antidepressants exert selection pressure for efflux-dependent antibiotic resistance mechanisms in Escherichia coli","volume":"13","author":"Ou","year":"2022","journal-title":"MBio"},{"key":"10.1016\/j.jbc.2026.113164_bib6","doi-asserted-by":"crossref","DOI":"10.1038\/s41598-020-74934-9","article-title":"Unravelling the antimicrobial action of antidepressants on gut commensal microbes","volume":"10","author":"Ait Chait","year":"2020","journal-title":"Sci. Rep."},{"key":"10.1016\/j.jbc.2026.113164_bib8","doi-asserted-by":"crossref","first-page":"299","DOI":"10.1186\/s12866-025-03973-x","article-title":"Fingolimod as a potent anti-Staphylococcus aureus: pH-dependent cell envelope damage and eradication of biofilms\/persisters","volume":"25","author":"Shang","year":"2025","journal-title":"BMC Microbiol."},{"key":"10.1016\/j.jbc.2026.113164_bib9","doi-asserted-by":"crossref","first-page":"1834","DOI":"10.3390\/microorganisms8111834","article-title":"Screening of FDA-approved drugs using a 384-well plate-based biofilm platform: the case of fingolimod","volume":"8","author":"Gilbert-Girard","year":"2020","journal-title":"Microorganisms"},{"key":"10.1016\/j.jbc.2026.113164_bib10","doi-asserted-by":"crossref","first-page":"18465","DOI":"10.1021\/acsomega.1c02591","article-title":"Synthesis and biological evaluation of fingolimod derivatives as antibacterial agents","volume":"6","author":"Zore","year":"2021","journal-title":"ACS Omega"},{"key":"10.1016\/j.jbc.2026.113164_bib11","doi-asserted-by":"crossref","first-page":"1043","DOI":"10.3390\/antibiotics11081043","article-title":"Fingolimod promotes antibacterial effect of doripenem against carbapenem-resistant Escherichia coli","volume":"11","author":"Jin","year":"2022","journal-title":"Antibiotics"},{"key":"10.1016\/j.jbc.2026.113164_bib12","doi-asserted-by":"crossref","DOI":"10.3389\/fmicb.2024.1396663","article-title":"Fingolimod synergizes and reverses K. pneumoniae resistance to colistin","volume":"15","author":"Geng","year":"2024","journal-title":"Front. Microbiol."},{"key":"10.1016\/j.jbc.2026.113164_bib13","doi-asserted-by":"crossref","first-page":"11","DOI":"10.3389\/fcimb.2017.00011","article-title":"Oral multiple sclerosis drugs inhibit the in vitro growth of epsilon toxin producing gut bacterium, Clostridium perfringens","volume":"7","author":"Rumah","year":"2017","journal-title":"Front. Cell. Infect. Microbiol."},{"key":"10.1016\/j.jbc.2026.113164_bib14","doi-asserted-by":"crossref","first-page":"373","DOI":"10.1098\/rsif.2009.0443","article-title":"Lipid bilayer regulation of membrane protein function: gramicidin channels as molecular force probes","volume":"7","author":"Lundb\u00e6k","year":"2010","journal-title":"J. R. Soc. Interf."},{"key":"10.1016\/j.jbc.2026.113164_bib15","doi-asserted-by":"crossref","first-page":"1256","DOI":"10.1126\/science.1700867","article-title":"Kinetics of gramicidin channel formation in lipid bilayers: Transmembrane monomer association","volume":"250","author":"O'Connell","year":"1990","journal-title":"Science"},{"key":"10.1016\/j.jbc.2026.113164_bib16","doi-asserted-by":"crossref","first-page":"95","DOI":"10.1016\/0005-2736(83)90264-X","article-title":"The effects of bilayer thickness and tension on gramicidin single-channel lifetime","volume":"735","author":"Elliott","year":"1983","journal-title":"Biochim. Biophys. Acta"},{"key":"10.1016\/j.jbc.2026.113164_bib17","doi-asserted-by":"crossref","first-page":"159","DOI":"10.1016\/S0304-4157(00)00016-2","article-title":"Structure of lipid bilayers","volume":"1469","author":"Nagle","year":"2000","journal-title":"Biochim. Biophys. Acta"},{"key":"10.1016\/j.jbc.2026.113164_bib18","doi-asserted-by":"crossref","first-page":"937","DOI":"10.1016\/S0006-3495(99)77257-7","article-title":"Experimental evidence for hydrophobic matching and membrane-mediated interactions in lipid bilayers containing gramicidin","volume":"76","author":"Harroun","year":"1999","journal-title":"Biophys. J."},{"key":"10.1016\/j.jbc.2026.113164_bib19","doi-asserted-by":"crossref","first-page":"9333","DOI":"10.1021\/bi980233r","article-title":"Influence of lipid\/peptide hydrophobic mismatch on the thickness of diacylphosphatidylcholine bilayers. A 2H NMR and ESR study using designed transmembrane a-helical peptides and gramicidin A","volume":"37","author":"Planque","year":"1998","journal-title":"Biochemistry"},{"key":"10.1016\/j.jbc.2026.113164_bib20","doi-asserted-by":"crossref","first-page":"328","DOI":"10.1016\/j.bbamem.2013.10.005","article-title":"Ion channel stability of Gramicidin A in lipid bilayers: effect of hydrophobic mismatch","volume":"1838","author":"Basu","year":"2014","journal-title":"Biochim. Biophys. Acta"},{"key":"10.1016\/j.jbc.2026.113164_bib21","doi-asserted-by":"crossref","first-page":"1551","DOI":"10.1016\/j.bpj.2012.03.014","article-title":"Influence of hydrophobic mismatch on structures and dynamics of gramicidin a and lipid bilayers","volume":"102","author":"Kim","year":"2012","journal-title":"Biophysical J."},{"key":"10.1016\/j.jbc.2026.113164_bib22","doi-asserted-by":"crossref","first-page":"1061","DOI":"10.1016\/S0006-3495(86)83550-0","article-title":"Deformation free energy of bilayer membrane and its effect on gramicidin channel lifetime","volume":"50","author":"Huang","year":"1986","journal-title":"Biophysical J."},{"key":"10.1016\/j.jbc.2026.113164_bib23","doi-asserted-by":"crossref","first-page":"1783","DOI":"10.1016\/S0006-3495(98)77620-9","article-title":"Membrane surface-charge titration probed by gramicidin A channel conductance","volume":"75","author":"Rostovtseva","year":"1998","journal-title":"Biophysical J."},{"key":"10.1016\/j.jbc.2026.113164_bib24","doi-asserted-by":"crossref","first-page":"342","DOI":"10.1085\/jgp.201812263","article-title":"Antidepressants are modifiers of lipid bilayer properties","volume":"151","author":"Kapoor","year":"2019","journal-title":"J. Gen. Physiol."},{"key":"10.1016\/j.jbc.2026.113164_bib25","doi-asserted-by":"crossref","first-page":"237","DOI":"10.1016\/S1359-0294(00)00061-3","article-title":"Functional consequences of lipid packing stress","volume":"5","author":"Bezrukov","year":"2000","journal-title":"Curr. Opin. Colloid Interf. Sci."},{"key":"10.1016\/j.jbc.2026.113164_bib26","doi-asserted-by":"crossref","first-page":"15427","DOI":"10.1073\/pnas.1007455107","article-title":"Amphiphile regulation of ion channel function by changes in the bilayer spring constant","volume":"107","author":"Lundb\u00e6k","year":"2010","journal-title":"Proc. Natl. Acad. Sci. U. S. A."},{"key":"10.1016\/j.jbc.2026.113164_bib27","doi-asserted-by":"crossref","first-page":"889","DOI":"10.1016\/S0006-3495(99)77252-8","article-title":"Spring constants for channel-induced lipid bilayer deformations estimates using gramicidin channels","volume":"76","author":"Lundb\u00e6k","year":"1999","journal-title":"Biophysical J."},{"key":"10.1016\/j.jbc.2026.113164_bib28","article-title":"Measuring cell death by propidium iodide uptake and flow cytometry","volume":"2016","author":"Crowley","year":"2016","journal-title":"Cold Spring Harbor. Protoc."},{"key":"10.1016\/j.jbc.2026.113164_bib29","doi-asserted-by":"crossref","first-page":"1829","DOI":"10.1016\/S0006-3495(01)76153-X","article-title":"Voltage-induced nonconductive pre-pores and metastable single pores in unmodified planar lipid bilayer","volume":"80","author":"Melikov","year":"2001","journal-title":"Biophysical J."},{"key":"10.1016\/j.jbc.2026.113164_bib30","doi-asserted-by":"crossref","first-page":"992","DOI":"10.1111\/cbdd.12060","article-title":"Chemotherapy drugs form ion pores in membranes due to physical interactions with lipids","volume":"80","author":"Ashrafuzzaman","year":"2012","journal-title":"Chem. Biol. Drug Des."},{"key":"10.1016\/j.jbc.2026.113164_bib31","doi-asserted-by":"crossref","first-page":"1985","DOI":"10.1016\/S0006-3495(02)75547-1","article-title":"Syringomycin E channel: a lipidic pore stabilized by lipopeptide?","volume":"82","author":"Malev","year":"2002","journal-title":"Biophysical J."},{"key":"10.1016\/j.jbc.2026.113164_bib32","doi-asserted-by":"crossref","first-page":"E3622","DOI":"10.1073\/pnas.1619806114","article-title":"Structural features and lipid binding domain of tubulin on biomimetic mitochondrial membranes","volume":"114","author":"Hoogerheide","year":"2017","journal-title":"Proc. Natl. Acad. Sci. U. S. A."},{"key":"10.1016\/j.jbc.2026.113164_bib33","first-page":"170","article-title":"Measurement of differences in the surface potentials of bilayer membranes according to the second harmonic of a capacitance current","volume":"25","author":"Sokolov","year":"1980","journal-title":"Biofizika"},{"key":"10.1016\/j.jbc.2026.113164_bib34","doi-asserted-by":"crossref","first-page":"19","DOI":"10.1016\/S0009-3084(02)00013-0","article-title":"Origin of membrane dipole potential: contribution of the phospholipid fatty acid chains","volume":"117","author":"Peterson","year":"2002","journal-title":"Chem. Phys. Lipids"},{"key":"10.1016\/j.jbc.2026.113164_bib35","doi-asserted-by":"crossref","first-page":"301","DOI":"10.1016\/0021-9797(76)90205-8","article-title":"Voltage dependence of bilayer membrane capacitance: harmonic response to ac excitation with dc bias","volume":"57","author":"Carius","year":"1976","journal-title":"J. Colloid Interf. Sci."},{"key":"10.1016\/j.jbc.2026.113164_bib36","doi-asserted-by":"crossref","first-page":"ar25","DOI":"10.1091\/mbc.E23-06-0263","article-title":"Cationic amphiphilic drugs induce accumulation of cytolytic lysoglycerophospholipids in the lysosomes of cancer cells and block their recycling into common membrane glycerophospholipids","volume":"35","author":"Nielsen","year":"2024","journal-title":"Mol. Biol. Cell"},{"key":"10.1016\/j.jbc.2026.113164_bib37","doi-asserted-by":"crossref","first-page":"725","DOI":"10.1016\/S0006-3495(01)75737-2","article-title":"Mesoscopic simulation of cell membrane damage, morphology change and rupture by nonionic surfactants","volume":"81","author":"Groot","year":"2001","journal-title":"Biophysical J."},{"key":"10.1016\/j.jbc.2026.113164_bib38","doi-asserted-by":"crossref","first-page":"2156","DOI":"10.1016\/S0006-3495(04)74275-7","article-title":"Molecular dynamics simulations of hydrophilic pores in lipid bilayers","volume":"86","author":"Leontiadou","year":"2004","journal-title":"Biophysical J."},{"key":"10.1016\/j.jbc.2026.113164_bib39","doi-asserted-by":"crossref","first-page":"2647","DOI":"10.1529\/biophysj.108.131409","article-title":"Bilayer edge and curvature effects on partitioning of lipids by tail length: atomistic simulations","volume":"95","author":"Wang","year":"2008","journal-title":"Biophysical J."},{"key":"10.1016\/j.jbc.2026.113164_bib40","doi-asserted-by":"crossref","DOI":"10.1103\/PhysRevE.103.042413","article-title":"Spatial extent of a single lipid's influence on bilayer mechanics","volume":"103","author":"Sapp","year":"2021","journal-title":"Phys. Rev. E"},{"key":"10.1016\/j.jbc.2026.113164_bib41","doi-asserted-by":"crossref","first-page":"3188","DOI":"10.1016\/j.bpj.2022.07.036","article-title":"Molecular mechanisms of spontaneous curvature and softening in complex lipid bilayer mixtures","volume":"121","author":"Lessen","year":"2022","journal-title":"Biophysical J."},{"key":"10.1016\/j.jbc.2026.113164_bib42","doi-asserted-by":"crossref","first-page":"5380","DOI":"10.21105\/joss.05380","article-title":"MembraneAnalysis. jl: A Julia package for analyzing molecular dynamics simulations of lipid membranes","volume":"8","author":"Hossein","year":"2023","journal-title":"J. Open Source Softw."},{"key":"10.1016\/j.jbc.2026.113164_bib43","doi-asserted-by":"crossref","DOI":"10.3389\/fonc.2020.605361","article-title":"Repurposing cationic amphiphilic drugs and derivatives to engage lysosomal cell death in cancer treatment","volume":"10","author":"Hu","year":"2020","journal-title":"Front. Oncol."},{"key":"10.1016\/j.jbc.2026.113164_bib44","doi-asserted-by":"crossref","first-page":"897","DOI":"10.1124\/dmd.112.050054","article-title":"Lysosomal sequestration (trapping) of lipophilic amine (cationic amphiphilic) drugs in immortalized human hepatocytes (Fa2N-4 cells)","volume":"41","author":"Kazmi","year":"2013","journal-title":"Drug Metab. Disposition"},{"key":"10.1016\/j.jbc.2026.113164_bib45","doi-asserted-by":"crossref","first-page":"1384","DOI":"10.1021\/mp200641e","article-title":"Cationic amphiphilic drugs cause a marked expansion of apparent lysosomal volume: implications for an intracellular distribution-based drug interaction","volume":"9","author":"Funk","year":"2012","journal-title":"Mol. pharmaceutics"},{"key":"10.1016\/j.jbc.2026.113164_bib46","doi-asserted-by":"crossref","first-page":"321","DOI":"10.1016\/j.bbrc.2010.11.115","article-title":"Gilenya (FTY720) inhibits acid sphingomyelinase by a mechanism similar to tricyclic antidepressants","volume":"404","author":"Dawson","year":"2011","journal-title":"Biochem. biophys. Res. Commun."},{"key":"10.1016\/j.jbc.2026.113164_bib47","doi-asserted-by":"crossref","first-page":"807","DOI":"10.1016\/j.it.2023.08.008","article-title":"pH sensing at the intersection of tissue homeostasis and inflammation","volume":"44","author":"Hajjar","year":"2023","journal-title":"Trends Immunol."},{"key":"10.1016\/j.jbc.2026.113164_bib48","doi-asserted-by":"crossref","first-page":"5087","DOI":"10.1073\/pnas.0710625105","article-title":"Mechanism and kinetics of pore formation in membranes by water-soluble amphipathic peptides","volume":"105","author":"Lee","year":"2008","journal-title":"Proc. Natl. Acad. Sci. U. S. A."},{"key":"10.1016\/j.jbc.2026.113164_bib49","doi-asserted-by":"crossref","first-page":"7231","DOI":"10.1038\/s41467-025-62604-1","article-title":"Antimicrobial peptide class that forms discrete \u03b2-barrel stable pores anchored by transmembrane helices","volume":"16","author":"Dickey","year":"2025","journal-title":"Nat. Commun."},{"key":"10.1016\/j.jbc.2026.113164_bib50","doi-asserted-by":"crossref","first-page":"2323","DOI":"10.1016\/S0006-3495(95)80102-5","article-title":"Two classes of alamethicin transmembrane channels: molecular models from single-channel properties","volume":"69","author":"Mak","year":"1995","journal-title":"Biophysical J."},{"key":"10.1016\/j.jbc.2026.113164_bib51","doi-asserted-by":"crossref","DOI":"10.1016\/j.mib.2023.102315","article-title":"Bacterial lipid biophysics and membrane organization","volume":"74","author":"Mitchison-Field","year":"2023","journal-title":"Curr. Opin. Microbiol."},{"key":"10.1016\/j.jbc.2026.113164_bib52","doi-asserted-by":"crossref","first-page":"76","DOI":"10.1016\/j.mib.2017.02.001","article-title":"Exploring functional membrane microdomains in bacteria: an overview","volume":"36","author":"Lopez","year":"2017","journal-title":"Curr. Opin. Microbiol."},{"key":"10.1016\/j.jbc.2026.113164_bib53","doi-asserted-by":"crossref","DOI":"10.1098\/rsif.2013.0810","article-title":"Asymmetric phospholipid: lipopolysaccharide bilayers; a Gram-negative bacterial outer membrane mimic","volume":"10","author":"Clifton","year":"2013","journal-title":"J. R. Soc. Interf."},{"key":"10.1016\/j.jbc.2026.113164_bib54","doi-asserted-by":"crossref","first-page":"33","DOI":"10.5483\/BMBRep.2024-0179","article-title":"Advancing membrane biology: single-molecule approaches meet model membrane systems","volume":"58","author":"Shin","year":"2025","journal-title":"BMB Rep."},{"key":"10.1016\/j.jbc.2026.113164_bib55","doi-asserted-by":"crossref","first-page":"813","DOI":"10.1016\/0019-2791(76)90181-6","article-title":"Binding of polymyxin B to the lipid A portion of bacterial lipopolysaccharides","volume":"13","author":"Morrison","year":"1976","journal-title":"Immunochemistry"},{"key":"10.1016\/j.jbc.2026.113164_bib56","doi-asserted-by":"crossref","first-page":"3485","DOI":"10.1021\/acs.langmuir.6b00240","article-title":"The effect of lipopolysaccharide core oligosaccharide size on the electrostatic binding of antimicrobial proteins to models of the gram negative bacterial outer membrane","volume":"32","author":"Clifton","year":"2016","journal-title":"Langmuir."},{"key":"10.1016\/j.jbc.2026.113164_bib57","doi-asserted-by":"crossref","first-page":"8009","DOI":"10.1073\/pnas.0903229106","article-title":"An ABC transport system that maintains lipid asymmetry in the gram-negative outer membrane","volume":"106","author":"Malinverni","year":"2009","journal-title":"Proc. Natl. Acad. Sci. U. S. A."},{"key":"10.1016\/j.jbc.2026.113164_bib58","doi-asserted-by":"crossref","first-page":"586","DOI":"10.3389\/fcell.2020.00586","article-title":"The beauty of asymmetric membranes: reconstitution of the outer membrane of gram-negative bacteria","volume":"8","author":"Paulowski","year":"2020","journal-title":"Front. Cell. Dev. Biol."},{"key":"10.1016\/j.jbc.2026.113164_bib59","doi-asserted-by":"crossref","first-page":"5195","DOI":"10.1021\/acs.langmuir.5b04562","article-title":"Subnanometer structure of an asymmetric model membrane: interleaflet coupling influences domain properties","volume":"32","author":"Heberle","year":"2016","journal-title":"Langmuir"},{"key":"10.1016\/j.jbc.2026.113164_bib60","doi-asserted-by":"crossref","DOI":"10.1073\/pnas.2308723120","article-title":"Experimentally determined leaflet\u2013leaflet phase diagram of an asymmetric lipid bilayer","volume":"120","author":"Enoki","year":"2023","journal-title":"Proc. Natl. Acad. Sci. U. S. A."},{"key":"10.1016\/j.jbc.2026.113164_bib61","doi-asserted-by":"crossref","first-page":"361","DOI":"10.1038\/nrm.2017.16","article-title":"The mystery of membrane organization: composition, regulation and roles of lipid rafts","volume":"18","author":"Sezgin","year":"2017","journal-title":"Nat. Rev. Mol. Cell. Biol."},{"key":"10.1016\/j.jbc.2026.113164_bib62","doi-asserted-by":"crossref","first-page":"289","DOI":"10.1016\/j.bbamem.2008.08.023","article-title":"Lipid domains in bacterial membranes and the action of antimicrobial agents","volume":"1788","author":"Epand","year":"2009","journal-title":"Biochim. Biophys. Acta"},{"key":"10.1016\/j.jbc.2026.113164_bib63","doi-asserted-by":"crossref","first-page":"532","DOI":"10.1177\/0091270004264165","article-title":"Multiple-dose FTY720: tolerability, pharmacokinetics, and lymphocyte responses in healthy subjects","volume":"44","author":"Kovarik","year":"2004","journal-title":"J. Clin. Pharmacol."},{"key":"10.1016\/j.jbc.2026.113164_bib64","doi-asserted-by":"crossref","DOI":"10.1016\/j.msard.2024.106246","article-title":"Whole blood concentrations of fingolimod and its pharmacologically active metabolite fingolimod phosphate obtained during routine health care of patients with multiple sclerosis","volume":"93","author":"Kacirova","year":"2025","journal-title":"Mult. Scler. Relat. Disord."},{"key":"10.1016\/j.jbc.2026.113164_bib65","doi-asserted-by":"crossref","first-page":"654","DOI":"10.1093\/toxsci\/kfs310","article-title":"Multiple NSAID-induced hits injure the small intestine: underlying mechanisms and novel strategies","volume":"131","author":"Boelsterli","year":"2013","journal-title":"Toxicol. Sci."},{"key":"10.1016\/j.jbc.2026.113164_bib66","doi-asserted-by":"crossref","first-page":"37496","DOI":"10.1074\/jbc.M602548200","article-title":"Voltage gating of VDAC is regulated by nonlamellar lipids of mitochondrial membranes","volume":"281","author":"Rostovtseva","year":"2006","journal-title":"J. Biol. Chem."},{"key":"10.1016\/j.jbc.2026.113164_bib67","doi-asserted-by":"crossref","first-page":"3561","DOI":"10.1073\/pnas.69.12.3561","article-title":"Formation of bimolecular membranes from lipid monolayers and a study of their electrical properties","volume":"69","author":"Montal","year":"1972","journal-title":"Proc. Natl. Acad. Sci. U. S. A."},{"key":"10.1016\/j.jbc.2026.113164_bib7","doi-asserted-by":"crossref","first-page":"13291","DOI":"10.1039\/C7NR03926C","article-title":"Lipid nanodomains change ion channel function","volume":"9","author":"Weinrich","year":"2017","journal-title":"Nanoscale"},{"key":"10.1016\/j.jbc.2026.113164_bib68","doi-asserted-by":"crossref","first-page":"1668","DOI":"10.1002\/jcc.20290","article-title":"The Amber biomolecular simulation programs","volume":"26","author":"Case","year":"2005","journal-title":"J. Comput. Chem."},{"key":"10.1016\/j.jbc.2026.113164_bib69","doi-asserted-by":"crossref","DOI":"10.1103\/PhysRevE.107.054403","article-title":"Kinetic relaxation of giant vesicles validates diffusional softening in a binary lipid mixture","volume":"107","author":"Sapp","year":"2023","journal-title":"Phys. Rev. E"}],"container-title":["Journal of Biological Chemistry"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S0021925826020363?httpAccept=text\/xml","content-type":"text\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S0021925826020363?httpAccept=text\/plain","content-type":"text\/plain","content-version":"vor","intended-application":"text-mining"}],"deposited":{"date-parts":[[2026,8,7]],"date-time":"2026-08-07T23:28:32Z","timestamp":1786145312000},"score":1,"resource":{"primary":{"URL":"https:\/\/linkinghub.elsevier.com\/retrieve\/pii\/S0021925826020363"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,7]]},"references-count":69,"journal-issue":{"issue":"7","published-print":{"date-parts":[[2026,7]]}},"alternative-id":["S0021925826020363"],"URL":"https:\/\/doi.org\/10.1016\/j.jbc.2026.113164","relation":{},"ISSN":["0021-9258"],"issn-type":[{"value":"0021-9258","type":"print"}],"subject":[],"published":{"date-parts":[[2026,7]]},"assertion":[{"value":"Elsevier","name":"publisher","label":"This article is maintained by"},{"value":"The multiple sclerosis drug fingolimod induces pore-like membrane defects, contributing to its antimicrobial activity","name":"articletitle","label":"Article Title"},{"value":"Journal of Biological Chemistry","name":"journaltitle","label":"Journal Title"},{"value":"https:\/\/doi.org\/10.1016\/j.jbc.2026.113164","name":"articlelink","label":"CrossRef DOI link to publisher maintained version"},{"value":"article","name":"content_type","label":"Content Type"},{"value":"\u00a9 2026 The Authors. Published by Elsevier Inc on behalf of American Society for Biochemistry and Molecular Biology.","name":"copyright","label":"Copyright"}],"article-number":"113164"}}