{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,23]],"date-time":"2026-10-23T21:19:16Z","timestamp":1792790356363,"version":"4.5.0"},"reference-count":101,"publisher":"eLife Sciences Publications, Ltd","license":[{"start":{"date-parts":[[2026,10,9]],"date-time":"2026-10-09T00:00:00Z","timestamp":1791504000000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/publicdomain\/zero\/1.0\/"},{"start":{"date-parts":[[2026,10,9]],"date-time":"2026-10-09T00:00:00Z","timestamp":1791504000000},"content-version":"am","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/publicdomain\/zero\/1.0\/"},{"start":{"date-parts":[[2026,10,9]],"date-time":"2026-10-09T00:00:00Z","timestamp":1791504000000},"content-version":"tdm","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/publicdomain\/zero\/1.0\/"}],"funder":[{"name":"Aix-Marseille Universit\u00e9","award":["AMX-21-PEP-022"],"award-info":[{"award-number":["AMX-21-PEP-022"]}],"id":[{"id":"https:\/\/ror.org\/035xkbk20","id-type":"ROR","asserted-by":"crossref"}]},{"award":["AMX-21-PEP-022"],"award-info":[{"award-number":["AMX-21-PEP-022"]}],"id":[{"id":"https:\/\/ror.org\/035xkbk20","id-type":"ROR","asserted-by":"publisher"}]},{"name":"Agence Nationale de la Recherche","award":["ApoMito"],"award-info":[{"award-number":["ApoMito"]}],"id":[{"id":"https:\/\/ror.org\/00rbzpz17","id-type":"ROR","asserted-by":"crossref"}]},{"award":["ApoMito"],"award-info":[{"award-number":["ApoMito"]}],"id":[{"id":"https:\/\/ror.org\/00rbzpz17","id-type":"ROR","asserted-by":"publisher"}]},{"name":"National Institutes of Health","award":["ZIA HD000072-18"],"award-info":[{"award-number":["ZIA HD000072-18"]}]},{"award":["ZIA HD000072-18"],"award-info":[{"award-number":["ZIA HD000072-18"]}],"id":[{"id":"https:\/\/ror.org\/01cwqze88","id-type":"ROR","asserted-by":"publisher"}]}],"content-domain":{"domain":["elifesciences.org"],"crossmark-restriction":false},"short-container-title":[],"abstract":"<jats:p>Voltage-dependent anion channel (VDAC) is the most abundant protein of the mitochondrial outer membrane and a key regulator of metabolite exchange and mitochondrial physiology. Its oligomerization has been proposed to control processes such as mitochondrial DNA release and membrane remodeling, yet the underlying mechanisms remain poorly defined. VBIT-4 has been widely used as a putative inhibitor of VDAC1 oligomerization, despite limited mechanistic validation. Here, using high-speed atomic force microscopy (AFM), we visualized VDAC1 assemblies in lipid membranes and examined the effect of VBIT-4. Unexpectedly, VBIT-4 induced membrane defects and permeabilization at micromolar concentrations, independently of VDAC1. Quantitative AFM analysis further shows that VBIT-4 does not alter VDAC1 cluster organization. Complementary electrophysiology, microscale thermophoresis, and coarse-grained molecular dynamics demonstrate that VBIT-4 partitions into lipid bilayers, increases membrane permeability, and destabilizes membrane structure, without detectable effects on VDAC1 channel properties or assemblies. Consistent with this mechanism, VBIT-4 induces VDAC1-independent cytotoxicity in HeLa cells at concentrations above 10 \u00b5M. Together, these results demonstrate that VBIT-4 does not act as a specific inhibitor of VDAC1 oligomerization but instead functions as a membrane-active compound. This work provides a revised framework for interpreting studies using VBIT-4 and highlights the importance of systematically assessing drug-membrane interactions when targeting membrane proteins.<\/jats:p>","DOI":"10.7554\/elife.111980","type":"journal-article","created":{"date-parts":[[2026,7,13]],"date-time":"2026-07-13T14:30:15Z","timestamp":1783953015000},"update-policy":"https:\/\/doi.org\/10.7554\/elife.111980","source":"Crossref","is-referenced-by-count":0,"title":["A membrane-disruptive action of VBIT-4 challenges its role as a widely used VDAC1 oligomerization inhibitor"],"prefix":"10.7554","volume":"15","author":[{"ORCID":"https:\/\/orcid.org\/0009-0007-6890-6924","authenticated-orcid":true,"given":"Varun","family":"Ravishankar","sequence":"first","affiliation":[{"id":[{"id":"https:\/\/ror.org\/035xkbk20","id-type":"ROR","asserted-by":"publisher"}],"name":"Laboratoire d'Ing\u00e9nierie des Syst\u00e8mes Macromol\u00e9culaires, CNRS, UMR 7255 \u2013 Aix-Marseille Universit\u00e9","place":["Marseille, France"]}],"role":[{"role":"author","vocabulary":"crossref"},{"role":"formal-analysis","vocabulary":"credit"},{"role":"investigation","vocabulary":"credit"},{"role":"writing-original-draft","vocabulary":"credit"},{"role":"writing-review-editing","vocabulary":"credit"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3004-7750","authenticated-orcid":true,"given":"Lu\u00eds","family":"Borges-Ara\u00fajo","sequence":"first","affiliation":[{"id":[{"id":"https:\/\/ror.org\/029brtt94","id-type":"ROR","asserted-by":"publisher"}],"name":"Laboratoire de Biologie et Mod\u00e9lisation de la Cellule, CNRS, UMR 5239, Inserm, U1293, Universit\u00e9 Claude Bernard Lyon 1, Ecole Normale Sup\u00e9rieure de Lyon","place":["Lyon, France"]},{"id":[{"id":"https:\/\/ror.org\/04zmssz18","id-type":"ROR","asserted-by":"publisher"}],"name":"Centre Blaise Pascal de Simulation et de Mod\u00e9lisation Num\u00e9rique, Ecole Normale Sup\u00e9rieure de Lyon","place":["Lyon, France"]}],"role":[{"role":"author","vocabulary":"crossref"},{"role":"conceptualization","vocabulary":"credit"},{"role":"formal-analysis","vocabulary":"credit"},{"role":"investigation","vocabulary":"credit"},{"role":"writing-original-draft","vocabulary":"credit"},{"role":"writing-review-editing","vocabulary":"credit"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-8078-4738","authenticated-orcid":true,"given":"Megha","family":"Rajendran","sequence":"first","affiliation":[{"id":[{"id":"https:\/\/ror.org\/04byxyr05","id-type":"ROR","asserted-by":"publisher"}],"name":"Eunice Kennedy Shriver National Institute of Child Health and Human Development","place":["Bethesda, United States"]}],"role":[{"role":"author","vocabulary":"crossref"},{"role":"conceptualization","vocabulary":"credit"},{"role":"formal-analysis","vocabulary":"credit"},{"role":"investigation","vocabulary":"credit"},{"role":"supervision","vocabulary":"credit"},{"role":"writing-original-draft","vocabulary":"credit"},{"role":"writing-review-editing","vocabulary":"credit"}]},{"ORCID":"https:\/\/orcid.org\/0009-0003-1737-9723","authenticated-orcid":true,"given":"Elodie","family":"Lafargue","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/035xkbk20","id-type":"ROR","asserted-by":"publisher"}],"name":"U1325 Aix-Marseille Universit\u00e9, INSERM, DyNaMo, Turing Centre for Living System","place":["Marseille, France"]}],"role":[{"role":"author","vocabulary":"crossref"},{"role":"investigation","vocabulary":"credit"},{"role":"visualization","vocabulary":"credit"},{"role":"writing-review-editing","vocabulary":"credit"}]},{"given":"Deborah","family":"Byrne","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/035xkbk20","id-type":"ROR","asserted-by":"publisher"}],"name":"Biomass and Protein Engineering Aix-Marseille Universit\u00e9, CNRS, IMM UAR2044","place":["Marseille, France"]}],"role":[{"role":"author","vocabulary":"crossref"},{"role":"methodology","vocabulary":"credit"},{"role":"supervision","vocabulary":"credit"}]},{"given":"Nicolas","family":"Buzhinsky","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/035xkbk20","id-type":"ROR","asserted-by":"publisher"}],"name":"U1325 Aix-Marseille Universit\u00e9, INSERM, DyNaMo, Turing Centre for Living System","place":["Marseille, France"]}],"role":[{"role":"author","vocabulary":"crossref"},{"role":"methodology","vocabulary":"credit"},{"role":"supervision","vocabulary":"credit"}]},{"given":"Mya S","family":"Wolfe","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/04byxyr05","id-type":"ROR","asserted-by":"publisher"}],"name":"Eunice Kennedy Shriver National Institute of Child Health and Human Development","place":["Bethesda, United States"]}],"role":[{"role":"author","vocabulary":"crossref"},{"role":"investigation","vocabulary":"credit"}]},{"given":"Wendy","family":"Fitzgerald","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/04byxyr05","id-type":"ROR","asserted-by":"publisher"}],"name":"Eunice Kennedy Shriver National Institute of Child Health and Human Development","place":["Bethesda, United States"]}],"role":[{"role":"author","vocabulary":"crossref"},{"role":"investigation","vocabulary":"credit"}]},{"given":"Nina A","family":"Bautista","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/04byxyr05","id-type":"ROR","asserted-by":"publisher"}],"name":"Eunice Kennedy Shriver National Institute of Child Health and Human Development","place":["Bethesda, United States"]}],"role":[{"role":"author","vocabulary":"crossref"},{"role":"investigation","vocabulary":"credit"}]},{"given":"Bethel G","family":"Beyene","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/04byxyr05","id-type":"ROR","asserted-by":"publisher"}],"name":"Eunice Kennedy Shriver National Institute of Child Health and Human Development","place":["Bethesda, United States"]}],"role":[{"role":"author","vocabulary":"crossref"},{"role":"investigation","vocabulary":"credit"}]},{"given":"Motahareh G","family":"Larimi","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/04byxyr05","id-type":"ROR","asserted-by":"publisher"}],"name":"Eunice Kennedy Shriver National Institute of Child Health and Human Development","place":["Bethesda, United States"]}],"role":[{"role":"author","vocabulary":"crossref"},{"role":"investigation","vocabulary":"credit"}]},{"given":"Jean-Pierre","family":"Duneau","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/035xkbk20","id-type":"ROR","asserted-by":"publisher"}],"name":"Laboratoire d'Ing\u00e9nierie des Syst\u00e8mes Macromol\u00e9culaires, CNRS, UMR 7255 \u2013 Aix-Marseille Universit\u00e9","place":["Marseille, France"]}],"role":[{"role":"author","vocabulary":"crossref"},{"role":"formal-analysis","vocabulary":"credit"},{"role":"investigation","vocabulary":"credit"},{"role":"software","vocabulary":"credit"},{"role":"writing-review-editing","vocabulary":"credit"}]},{"given":"James","family":"Sturgis","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/035xkbk20","id-type":"ROR","asserted-by":"publisher"}],"name":"Laboratoire d'Ing\u00e9nierie des Syst\u00e8mes Macromol\u00e9culaires, CNRS, UMR 7255 \u2013 Aix-Marseille Universit\u00e9","place":["Marseille, France"]}],"role":[{"role":"author","vocabulary":"crossref"},{"role":"methodology","vocabulary":"credit"},{"role":"writing-review-editing","vocabulary":"credit"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8209-8050","authenticated-orcid":true,"given":"Sergey M","family":"Bezrukov","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/04byxyr05","id-type":"ROR","asserted-by":"publisher"}],"name":"Eunice Kennedy Shriver National Institute of Child Health and Human Development","place":["Bethesda, United States"]}],"role":[{"role":"author","vocabulary":"crossref"},{"role":"conceptualization","vocabulary":"credit"},{"role":"writing-review-editing","vocabulary":"credit"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7192-0136","authenticated-orcid":true,"given":"Ignacio","family":"Casuso","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/035xkbk20","id-type":"ROR","asserted-by":"publisher"}],"name":"U1325 Aix-Marseille Universit\u00e9, INSERM, DyNaMo, Turing Centre for Living System","place":["Marseille, France"]}],"role":[{"role":"author","vocabulary":"crossref"},{"role":"conceptualization","vocabulary":"credit"},{"role":"writing-review-editing","vocabulary":"credit"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7299-7362","authenticated-orcid":true,"given":"Tatiana K","family":"Rostovtseva","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/04byxyr05","id-type":"ROR","asserted-by":"publisher"}],"name":"Eunice Kennedy Shriver National Institute of Child Health and Human Development","place":["Bethesda, United States"]}],"role":[{"role":"author","vocabulary":"crossref"},{"role":"conceptualization","vocabulary":"credit"},{"role":"formal-analysis","vocabulary":"credit"},{"role":"funding-acquisition","vocabulary":"credit"},{"role":"supervision","vocabulary":"credit"},{"role":"writing-original-draft","vocabulary":"credit"},{"role":"writing-review-editing","vocabulary":"credit"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-6068-238X","authenticated-orcid":true,"given":"Lucie","family":"Bergdoll","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/035xkbk20","id-type":"ROR","asserted-by":"publisher"}],"name":"Laboratoire d'Ing\u00e9nierie des Syst\u00e8mes Macromol\u00e9culaires, CNRS, UMR 7255 \u2013 Aix-Marseille Universit\u00e9","place":["Marseille, France"]}],"role":[{"role":"author","vocabulary":"crossref"},{"role":"corresponding-author","vocabulary":"crossref"},{"role":"conceptualization","vocabulary":"credit"},{"role":"formal-analysis","vocabulary":"credit"},{"role":"investigation","vocabulary":"credit"},{"role":"supervision","vocabulary":"credit"},{"role":"visualization","vocabulary":"credit"},{"role":"writing-original-draft","vocabulary":"credit"},{"role":"writing-review-editing","vocabulary":"credit"}]}],"member":"4374","published-online":{"date-parts":[[2026,10,9]]},"reference":[{"key":"bib1","doi-asserted-by":"publisher","first-page":"12468","DOI":"10.1073\/pnas.211400898","type":"journal-article","article-title":"A high-speed atomic force microscope for studying biological macromolecules","volume":"98","author":"Ando","year":"2001","journal-title":"PNAS"},{"key":"bib2","doi-asserted-by":"publisher","DOI":"10.1101\/2024.07.23.604601","type":"preprint","article-title":"Creating coarse-grained systems with COBY: towards higher accuracy in membrane complexity","volume-title":"bioRxiv","author":"Andreasen","year":"2024"},{"key":"bib3","doi-asserted-by":"publisher","DOI":"10.1140\/epje\/i2020-11989-0","type":"journal-article","article-title":"Estimation of the parameters of the Smoluchowski equation describing the occurrence of pores in a bilayer lipid membrane under soft poration","volume":"43","author":"Anosov","year":"2020","journal-title":"The European Physical Journal. E, Soft Matter"},{"key":"bib4","doi-asserted-by":"publisher","first-page":"992","DOI":"10.1111\/cbdd.12060","type":"journal-article","article-title":"Chemotherapy drugs form ion pores in membranes due to physical interactions with lipids","volume":"80","author":"Ashrafuzzaman","year":"2012","journal-title":"Chemical Biology & Drug Design"},{"key":"bib5","doi-asserted-by":"publisher","first-page":"183","DOI":"10.1002\/jps.2600820214","type":"journal-article","article-title":"pH-metric log P. II: Refinement of partition coefficients and ionization constants of multiprotic substances","volume":"82","author":"Avdeef","year":"1993","journal-title":"Journal of Pharmaceutical Sciences"},{"key":"bib6","doi-asserted-by":"publisher","first-page":"109","DOI":"10.1201\/9781351060318-6","type":"book-chapter","article-title":"Free-energy calculations of pore formation in lipid membranes","volume-title":"Biomembrane Simulations","author":"Awasthi","year":"2019"},{"key":"bib7","doi-asserted-by":"publisher","DOI":"10.1126\/sciimmunol.ade7652","type":"journal-article","article-title":"Hexokinase dissociation from mitochondria promotes oligomerization of VDAC that facilitates NLRP3 inflammasome assembly and activation","volume":"8","author":"Baik","year":"2023","journal-title":"Science Immunology"},{"key":"bib8","doi-asserted-by":"publisher","DOI":"10.3390\/antiox12071459","type":"journal-article","article-title":"Pharmacological and genetic suppression of VDAC1 alleviates the development of mitochondrial dysfunction in endothelial and fibroblast cell cultures upon hyperglycemic conditions","volume":"12","author":"Belosludtsev","year":"2023","journal-title":"Antioxidants"},{"key":"bib9","doi-asserted-by":"publisher","DOI":"10.1016\/j.bbamem.2024.184329","type":"journal-article","article-title":"Effect of VBIT-4 on the functional activity of isolated mitochondria and cell viability","volume":"1866","author":"Belosludtsev","year":"2024","journal-title":"Biochimica et Biophysica Acta. Biomembranes"},{"key":"bib10","doi-asserted-by":"publisher","first-page":"24986","DOI":"10.1074\/jbc.M116.744284","type":"journal-article","article-title":"Novel compounds targeting the mitochondrial protein VDAC1 inhibit apoptosis and protect against mitochondrial dysfunction","volume":"291","author":"Ben-Hail","year":"2016","journal-title":"The Journal of Biological Chemistry"},{"key":"bib11","doi-asserted-by":"publisher","first-page":"3684","DOI":"10.1063\/1.448118","type":"journal-article","article-title":"Molecular dynamics with coupling to an external bath","volume":"81","author":"Berendsen","year":"1984","journal-title":"The Journal of Chemical Physics"},{"key":"bib12","doi-asserted-by":"publisher","first-page":"E172","DOI":"10.1073\/pnas.1715464115","type":"journal-article","article-title":"Protonation state of glutamate 73 regulates the formation of a specific dimeric association of mVDAC1","volume":"115","author":"Bergdoll","year":"2018","journal-title":"PNAS"},{"key":"bib13","doi-asserted-by":"publisher","DOI":"10.1063\/5.0020514","type":"journal-article","article-title":"Pressure control using stochastic cell rescaling","volume":"153","author":"Bernetti","year":"2020","journal-title":"The Journal of Chemical Physics"},{"key":"bib14","doi-asserted-by":"publisher","DOI":"10.1101\/2024.07.22.604557","type":"preprint","article-title":"Hexokinase-I directly binds to a charged membrane-buried glutamate of mitochondrial VDAC1 and VDAC2","volume-title":"bioRxiv","author":"Bieker","year":"2024"},{"key":"bib15","doi-asserted-by":"publisher","first-page":"206","DOI":"10.1016\/j.str.2019.11.015","type":"journal-article","article-title":"The structural basis for low conductance in the membrane protein VDAC upon \u03b2-NADH binding and voltage gating","volume":"28","author":"B\u00f6hm","year":"2020","journal-title":"Structure"},{"key":"bib16","doi-asserted-by":"publisher","DOI":"10.1126\/sciadv.adi2767","type":"journal-article","article-title":"ANT-dependent MPTP underlies necrotic myofiber death in muscular dystrophy","volume":"9","author":"Bround","year":"2023","journal-title":"Science Advances"},{"key":"bib17","doi-asserted-by":"publisher","DOI":"10.1063\/1.2408420","type":"journal-article","article-title":"Canonical sampling through velocity rescaling","volume":"126","author":"Bussi","year":"2007","journal-title":"The Journal of Chemical Physics"},{"key":"bib18","doi-asserted-by":"publisher","first-page":"303","DOI":"10.1126\/science.adu6445","type":"journal-article","article-title":"Structure of human PINK1 at a mitochondrial TOM-VDAC array","volume":"388","author":"Callegari","year":"2025","journal-title":"Science"},{"key":"bib19","doi-asserted-by":"publisher","DOI":"10.3389\/fphys.2017.00460","type":"journal-article","article-title":"Mitochondrial VDAC1: a key gatekeeper as potential therapeutic target","volume":"8","author":"Camara","year":"2017","journal-title":"Frontiers in Physiology"},{"key":"bib20","doi-asserted-by":"publisher","first-page":"429","DOI":"10.1016\/j.surfrep.2006.06.001","type":"journal-article","article-title":"Solid supported lipid bilayers: from biophysical studies to sensor design","volume":"61","author":"Castellana","year":"2006","journal-title":"Surface Science Reports"},{"key":"bib21","doi-asserted-by":"publisher","first-page":"525","DOI":"10.1038\/nnano.2012.109","type":"journal-article","article-title":"Characterization of the motion of membrane proteins using high-speed atomic force microscopy","volume":"7","author":"Casuso","year":"2012","journal-title":"Nature Nanotechnology"},{"key":"bib22","doi-asserted-by":"publisher","first-page":"2140","DOI":"10.1021\/ci700257y","type":"journal-article","article-title":"Computation of octanol-water partition coefficients by guiding an additive model with knowledge","volume":"47","author":"Cheng","year":"2007","journal-title":"Journal of Chemical Information and Modeling"},{"key":"bib23","doi-asserted-by":"publisher","first-page":"33","DOI":"10.1007\/978-3-030-21162-2_3","type":"book-chapter","article-title":"Ceramide channels","volume-title":"Bioactive Ceramides in Health and Disease","author":"Colombini","year":"2019"},{"key":"bib24","doi-asserted-by":"publisher","first-page":"485","DOI":"10.1016\/0040-6031(72)87040-0","type":"journal-article","article-title":"The free energy, enthalpy and entropy changes of the dissociation of diprotonated diamines","volume":"4","author":"Creyf","year":"1972","journal-title":"Thermochimica Acta"},{"key":"bib25","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-019-09654-4","type":"journal-article","article-title":"Ceramides bind VDAC2 to trigger mitochondrial apoptosis","volume":"10","author":"Dadsena","year":"2019","journal-title":"Nature Communications"},{"key":"bib26","doi-asserted-by":"publisher","first-page":"3284","DOI":"10.1021\/ci500467k","type":"journal-article","article-title":"iLOGP: a simple, robust, and efficient description of n-octanol\/water partition coefficient for drug design using the GB\/SA approach","volume":"54","author":"Daina","year":"2014","journal-title":"Journal of Chemical Information and Modeling"},{"key":"bib27","doi-asserted-by":"publisher","DOI":"10.1038\/srep42717","type":"journal-article","article-title":"SwissADME: a free web tool to evaluate pharmacokinetics, drug-likeness and medicinal chemistry friendliness of small molecules","volume":"7","author":"Daina","year":"2017","journal-title":"Scientific Reports"},{"key":"bib28","doi-asserted-by":"publisher","first-page":"10089","DOI":"10.1063\/1.464397","type":"journal-article","article-title":"Particle mesh Ewald: An N\u22c5log(N) method for Ewald sums in large systems","volume":"98","author":"Darden","year":"1993","journal-title":"Journal of Chemical Physics"},{"key":"bib29","doi-asserted-by":"publisher","DOI":"10.1016\/j.xpro.2024.103240","type":"journal-article","article-title":"Protocol for the production and reconstitution of VDAC1 for functional assays","volume":"5","author":"Dearden","year":"2024","journal-title":"STAR Protocols"},{"key":"bib30","type":"software","article-title":"ComBiol2025","volume-title":"Github","author":"deejy","year":"2025","unstructured":"deejy. 2025. ComBiol2025. Github. d0cdec8. https:\/\/github.com\/deejy\/ComBiol2025."},{"key":"bib31","doi-asserted-by":"publisher","first-page":"3477","DOI":"10.1021\/ja0393574","type":"journal-article","article-title":"Directed self-assembly of monodisperse phospholipid bilayer Nanodiscs with controlled size","volume":"126","author":"Denisov","year":"2004","journal-title":"Journal of the American Chemical Society"},{"key":"bib32","doi-asserted-by":"publisher","first-page":"W331","DOI":"10.1093\/nar\/gkx312","type":"journal-article","article-title":"LigParGen web server: an automatic OPLS-AA parameter generator for organic ligands","volume":"45","author":"Dodda","year":"2017","journal-title":"Nucleic Acids Research"},{"key":"bib33","doi-asserted-by":"publisher","first-page":"3864","DOI":"10.1021\/acs.jpcb.7b00272","type":"journal-article","article-title":"1.14*CM1A-LBCC: localized bond-charge corrected CM1A charges for condensed-phase simulations","volume":"121","author":"Dodda","year":"2017","journal-title":"The Journal of Physical Chemistry. B"},{"key":"bib34","first-page":"226","type":"conference-paper","article-title":"A density-based algorithm for discovering clusters in large spatial databases with noise","author":"Ester","year":"1996","unstructured":"Ester M, Kriegel HP, Sander J, Xu X. 1996. A density-based algorithm for discovering clusters in large spatial databases with noise. Proceedings of the Second International Conference on Knowledge Discovery and Data Mining."},{"key":"bib35","doi-asserted-by":"publisher","first-page":"569","DOI":"10.1021\/acs.jcim.9b00830","type":"journal-article","article-title":"Lipophilicity of coarse-grained cholesterol models","volume":"60","author":"Fornasier","year":"2020","journal-title":"Journal of Chemical Information and Modeling"},{"key":"bib36","doi-asserted-by":"publisher","first-page":"413","DOI":"10.1016\/j.jmb.2007.03.063","type":"journal-article","article-title":"Supramolecular assembly of VDAC in native mitochondrial outer membranes","volume":"369","author":"Gon\u00e7alves","year":"2007","journal-title":"Journal of Molecular Biology"},{"key":"bib37","type":"report","volume-title":"MDAnalysis: a python package for the rapid analysis of molecular dynamics simulations","author":"Gowers","year":"2019","unstructured":"Gowers RJ, Linke M, Barnoud J, Reddy TJE, Melo MN, Seyler SL, Domanski J, Dotson DL, Buchoux S, Kenney IM, Beckstein O. 2019. LA-UR-19-29136, Los Alamos National Laboratory (LANL). MDAnalysis: a python package for the rapid analysis of molecular dynamics simulations."},{"key":"bib38","doi-asserted-by":"publisher","first-page":"374","DOI":"10.1038\/s41586-023-06373-1","type":"journal-article","article-title":"cGAS-STING drives ageing-related inflammation and neurodegeneration","volume":"620","author":"Gulen","year":"2023","journal-title":"Nature"},{"key":"bib39","doi-asserted-by":"publisher","first-page":"1463","DOI":"10.1002\/(SICI)1096-987X(199709)18:12<1463::AID-JCC4>3.0.CO;2-H","type":"journal-article","article-title":"LINCS: A linear constraint solver for molecular simulations","volume":"18","author":"Hess","year":"1997","journal-title":"Journal of Computational Chemistry"},{"key":"bib40","doi-asserted-by":"publisher","first-page":"246","DOI":"10.1016\/j.jmb.2007.04.073","type":"journal-article","article-title":"The supramolecular assemblies of voltage-dependent anion channels in the native membrane","volume":"370","author":"Hoogenboom","year":"2007","journal-title":"Journal of Molecular Biology"},{"key":"bib41","doi-asserted-by":"publisher","DOI":"10.1063\/1.2432410","type":"journal-article","article-title":"WSXM: a software for scanning probe microscopy and a tool for nanotechnology","volume":"78","author":"Horcas","year":"2007","journal-title":"The Review of Scientific Instruments"},{"key":"bib42","doi-asserted-by":"publisher","first-page":"2352","DOI":"10.1021\/acs.jctc.7b00106","type":"journal-article","article-title":"Probing a continuous polar defect: a reaction coordinate for pore formation in lipid membranes","volume":"13","author":"Hub","year":"2017","journal-title":"Journal of Chemical Theory and Computation"},{"key":"bib43","doi-asserted-by":"publisher","first-page":"335","DOI":"10.1007\/s10822-020-00295-0","type":"journal-article","article-title":"Assessing the accuracy of octanol-water partition coefficient predictions in the SAMPL6 Part II log P Challenge","volume":"34","author":"I\u015f\u0131k","year":"2020","journal-title":"Journal of Computer-Aided Molecular Design"},{"key":"bib44","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-023-43570-y","type":"journal-article","article-title":"Phospholipids are imported into mitochondria by VDAC, a dimeric beta barrel scramblase","volume":"14","author":"Jahn","year":"2023","journal-title":"Nature Communications"},{"key":"bib45","doi-asserted-by":"publisher","first-page":"6665","DOI":"10.1073\/pnas.0408037102","type":"journal-article","article-title":"Potential energy functions for atomic-level simulations of water and organic and biomolecular systems","volume":"102","author":"Jorgensen","year":"2005","journal-title":"PNAS"},{"key":"bib46","doi-asserted-by":"publisher","first-page":"5698","DOI":"10.1128\/MCB.00165-10","type":"journal-article","article-title":"Oligomerization of the mitochondrial protein voltage-dependent anion channel is coupled to the induction of apoptosis","volume":"30","author":"Keinan","year":"2010","journal-title":"Molecular and Cellular Biology"},{"key":"bib47","doi-asserted-by":"publisher","first-page":"1531","DOI":"10.1126\/science.aav4011","type":"journal-article","article-title":"VDAC oligomers form mitochondrial pores to release mtDNA fragments and promote lupus-like disease","volume":"366","author":"Kim","year":"2019","journal-title":"Science"},{"key":"bib48","doi-asserted-by":"publisher","first-page":"8919","DOI":"10.1021\/acs.jctc.3c00777","type":"journal-article","article-title":"Neighbor list artifacts in molecular dynamics simulations","volume":"19","author":"Kim","year":"2023","journal-title":"Journal of Chemical Theory and Computation"},{"key":"bib49","doi-asserted-by":"publisher","DOI":"10.1038\/s41598-020-79056-w","type":"journal-article","article-title":"VDAC1 in the diseased myocardium and the effect of VDAC1-interacting compound on atrial fibrosis induced by hyperaldosteronism","volume":"10","author":"Klapper-Goldstein","year":"2020","journal-title":"Scientific Reports"},{"key":"bib50","doi-asserted-by":"publisher","first-page":"397","DOI":"10.1007\/s10822-015-9840-9","type":"journal-article","article-title":"Guidelines for the analysis of free energy calculations","volume":"29","author":"Klimovich","year":"2015","journal-title":"Journal of Computer-Aided Molecular Design"},{"key":"bib51","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.90627.4","type":"journal-article","article-title":"Martinize2 and Vermouth provide a unified framework for molecular topology generation","volume":"12","author":"Kroon","year":"2025","journal-title":"eLife"},{"key":"bib52","doi-asserted-by":"publisher","first-page":"1011","DOI":"10.1002\/jcc.540130812","type":"journal-article","article-title":"THE weighted histogram analysis method for free\u2010energy calculations on biomolecules. I. THE method","volume":"13","author":"Kumar","year":"1992","journal-title":"Journal of Computational Chemistry"},{"key":"bib53","doi-asserted-by":"publisher","DOI":"10.1038\/s42003-025-08311-5","type":"journal-article","article-title":"Membrane lipid composition modulates the organization of VDAC1, a mitochondrial gatekeeper","volume":"8","author":"Lafargue","year":"2025","journal-title":"Communications Biology"},{"key":"bib54","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.2110996118","type":"journal-article","article-title":"In-cell structures of conserved supramolecular protein arrays at the mitochondria-cytoskeleton interface in mammalian sperm","volume":"118","author":"Leung","year":"2021","journal-title":"PNAS"},{"key":"bib55","doi-asserted-by":"publisher","first-page":"3","DOI":"10.1016\/s0169-409x(00)00129-0","type":"journal-article","article-title":"Experimental and computational approaches to estimate solubility and permeability in drug discovery and development settings","volume":"46","author":"Lipinski","year":"2001","journal-title":"Advanced Drug Delivery Reviews"},{"key":"bib56","doi-asserted-by":"publisher","first-page":"5171","DOI":"10.1021\/acs.jmedchem.1c00251","type":"journal-article","article-title":"Interface-mediated mechanism of action-the root of the cytoprotective effect of immediate-release omeprazole","volume":"64","author":"Lopes-de-Campos","year":"2021","journal-title":"Journal of Medicinal Chemistry"},{"key":"bib57","doi-asserted-by":"publisher","DOI":"10.1007\/s11064-025-04366-2","type":"journal-article","article-title":"VDAC1 Inhibition mitigates inflammatory status and oxidative stress in epileptic mice treated with the ketogenic diet","volume":"50","author":"Luo","year":"2025","journal-title":"Neurochemical Research"},{"key":"bib58","doi-asserted-by":"publisher","first-page":"4470","DOI":"10.2174\/0929867324666170601073920","type":"journal-article","article-title":"Interactions of VDAC with proteins involved in neurodegenerative aggregation: An opportunity for advancement on therapeutic molecules","volume":"24","author":"Magri","year":"2017","journal-title":"Current Medicinal Chemistry"},{"key":"bib59","doi-asserted-by":"publisher","first-page":"2243","DOI":"10.1073\/pnas.80.8.2243","type":"journal-article","article-title":"Structural and functional evidence for multiple channel complexes in the outer membrane of Neurospora crassa mitochondria","volume":"80","author":"Mannella","year":"1983","journal-title":"PNAS"},{"key":"bib60","doi-asserted-by":"publisher","first-page":"861","DOI":"10.1002\/jps.21494","type":"journal-article","article-title":"Calculation of molecular lipophilicity: State-of-the-art and comparison of log P methods on more than 96,000 compounds","volume":"98","author":"Mannhold","year":"2009","journal-title":"Journal of Pharmaceutical Sciences"},{"key":"bib61","doi-asserted-by":"publisher","first-page":"1207","DOI":"10.1016\/j.carres.2004.12.038","type":"journal-article","article-title":"Octanol-water partition coefficient of glucose, sucrose, and trehalose","volume":"340","author":"Mazzobre","year":"2005","journal-title":"Carbohydrate Research"},{"key":"bib62","doi-asserted-by":"publisher","first-page":"2319","DOI":"10.1002\/jcc.21787","type":"journal-article","article-title":"MDAnalysis: a toolkit for the analysis of molecular dynamics simulations","volume":"32","author":"Michaud-Agrawal","year":"2011","journal-title":"Journal of Computational Chemistry"},{"key":"bib63","doi-asserted-by":"publisher","DOI":"10.3390\/biom13030557","type":"journal-article","article-title":"Ablation of GPR56 causes \u03b2-cell dysfunction by ATP loss through mistargeting of mitochondrial VDAC1 to the plasma membrane","volume":"13","author":"Mohammad Al-Amily","year":"2023","journal-title":"Biomolecules"},{"key":"bib64","doi-asserted-by":"publisher","first-page":"E5590","DOI":"10.1073\/pnas.1510574112","type":"journal-article","article-title":"Motifs of VDAC2 required for mitochondrial Bak import and tBid-induced apoptosis","volume":"112","author":"Naghdi","year":"2015","journal-title":"PNAS"},{"key":"bib65","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.64787","type":"journal-article","article-title":"ATP and large signaling metabolites flux through caspase-activated Pannexin 1 channels","volume":"10","author":"Narahari","year":"2021","journal-title":"eLife"},{"key":"bib66","type":"software","article-title":"Particle_Analysis","volume-title":"Github","author":"Ortega","year":"2024","unstructured":"Ortega A. 2024. Particle_Analysis. Github. 7484a16. https:\/\/github.com\/CodeCanSolveAll\/Particle_Analysis."},{"key":"bib67","doi-asserted-by":"publisher","first-page":"2641","DOI":"10.1016\/j.cpc.2013.06.003","type":"journal-article","article-title":"A flexible algorithm for calculating pair interactions on SIMD architectures","volume":"184","author":"P\u00e1ll","year":"2013","journal-title":"Computer Physics Communications"},{"key":"bib68","doi-asserted-by":"publisher","first-page":"3159","DOI":"10.1021\/acs.jcim.1c00075","type":"journal-article","article-title":"MolGpka: a web server for small molecule pKa prediction using a graph-convolutional neural network","volume":"61","author":"Pan","year":"2021","journal-title":"Journal of Chemical Information and Modeling"},{"key":"bib69","doi-asserted-by":"publisher","first-page":"7182","DOI":"10.1063\/1.328693","type":"journal-article","article-title":"Polymorphic transitions in single crystals: a new molecular dynamics method","volume":"52","author":"Parrinello","year":"1981","journal-title":"Journal of Applied Physics"},{"key":"bib70","doi-asserted-by":"publisher","first-page":"1598","DOI":"10.1021\/acscentsci.5c00755","type":"journal-article","article-title":"The martini 3 lipidome: expanded and refined parameters improve lipid phase behavior","volume":"11","author":"Pedersen","year":"2025","journal-title":"ACS Central Science"},{"key":"bib71","first-page":"2825","type":"journal-article","article-title":"Scikit-learn: machine learning in python","volume":"12","author":"Pedregosa","year":"2018","journal-title":"Journal of Machine Learning Research"},{"key":"bib72","doi-asserted-by":"publisher","DOI":"10.3390\/molecules24030516","type":"journal-article","article-title":"Licofelone-DPPC interactions: putting membrane lipids on the radar of drug development","volume":"24","author":"Pereira-Leite","year":"2019","journal-title":"Molecules"},{"key":"bib73","doi-asserted-by":"publisher","first-page":"230","DOI":"10.1016\/j.csbj.2021.12.021","type":"journal-article","article-title":"Dynorphin A induces membrane permeabilization by formation of proteolipidic pores","volume":"20","author":"Perini","year":"2022","journal-title":"Insights from Electrophysiology and Computational Simulations. Computational and Structural Biotechnology Journal"},{"key":"bib74","doi-asserted-by":"publisher","first-page":"22","DOI":"10.1016\/j.bbabio.2018.11.001","type":"journal-article","article-title":"Assessing the role of residue E73 and lipid headgroup charge in VDAC1 voltage gating","volume":"1860","author":"Queralt-Mart\u00edn","year":"2019","journal-title":"Biochimica et Biophysica Acta. Bioenergetics"},{"key":"bib75","doi-asserted-by":"publisher","DOI":"10.1085\/jgp.201912501","type":"journal-article","article-title":"A lower affinity to cytosolic proteins reveals VDAC3 isoform-specific role in mitochondrial biology","volume":"152","author":"Queralt-Mart\u00edn","year":"2020","journal-title":"The Journal of General Physiology"},{"key":"bib76","doi-asserted-by":"publisher","DOI":"10.1016\/j.jbc.2025.111009","type":"journal-article","article-title":"Three mammalian VDAC isoforms distinctly regulate mitochondrial function and proteome to maintain cell metabolism","volume":"302","author":"Rajendran","year":"2026","journal-title":"The Journal of Biological Chemistry"},{"key":"bib77","doi-asserted-by":"publisher","first-page":"465","DOI":"10.1007\/s00249-015-1049-2","type":"journal-article","article-title":"Conductance hysteresis in the voltage-dependent anion channel","volume":"44","author":"Rappaport","year":"2015","journal-title":"European Biophysics Journal"},{"key":"bib78","doi-asserted-by":"publisher","DOI":"10.5281\/zenodo.22285808","type":"software","article-title":"Repository of MD data for membrane-disruptive action of VBIT-4 challenges its role as a widely used VDAC1 oligomerization inhibitor","volume-title":"Zenodo","author":"Ravishankar","year":"2026","unstructured":"Ravishankar V, Borges-Ara\u00fajo L, Rajendran M, Lafargue E, Byrne D, Buzhinsky N, Wolfe MS, Fitzgerald W, Bautista NA, Beyene BG, Larimi MG, Duneau JP, Sturgis J, Bezrukov SM, Casuso I, Rostovtseva T, Bergdoll L. 2026. Repository of MD data for membrane-disruptive action of VBIT-4 challenges its role as a widely used VDAC1 oligomerization inhibitor. Zenodo. Version v1. https:\/\/doi.org\/10.5281\/zenodo.22285808."},{"key":"bib79","doi-asserted-by":"publisher","first-page":"4447","DOI":"10.2174\/0929867324666170530074039","type":"journal-article","article-title":"Anti-cancer compounds targeted to VDAC: potential and perspectives","volume":"24","author":"Reina","year":"2017","journal-title":"Current Medicinal Chemistry"},{"key":"bib80","doi-asserted-by":"publisher","first-page":"211","DOI":"10.1016\/S0076-6879(09)64011-8","type":"journal-article","article-title":"Chapter 11 - Reconstitution of membrane proteins in phospholipid bilayer nanodiscs","volume":"464","author":"Ritchie","year":"2009","journal-title":"Methods in Enzymology"},{"key":"bib81","doi-asserted-by":"publisher","DOI":"10.1016\/j.ceca.2021.102356","type":"journal-article","article-title":"VDAC regulation of mitochondrial calcium flux: from channel biophysics to disease","volume":"94","author":"Rosencrans","year":"2021","journal-title":"Cell Calcium"},{"key":"bib82","doi-asserted-by":"publisher","first-page":"37496","DOI":"10.1074\/jbc.M602548200","type":"journal-article","article-title":"Voltage gating of VDAC is regulated by nonlamellar lipids of mitochondrial membranes","volume":"281","author":"Rostovtseva","year":"2006","journal-title":"The Journal of Biological Chemistry"},{"key":"bib83","doi-asserted-by":"publisher","DOI":"10.3390\/ijms25042204","type":"journal-article","article-title":"Dimeric tubulin modifies mechanical properties of lipid bilayer, as probed using gramicidin a channel","volume":"25","author":"Rostovtseva","year":"2024","journal-title":"International Journal of Molecular Sciences"},{"key":"bib84","doi-asserted-by":"publisher","DOI":"10.3390\/ijms22020946","type":"journal-article","article-title":"A calcium guard in the outer membrane: is VDAC a regulated gatekeeper of mitochondrial calcium uptake?","volume":"22","author":"Sander","year":"2021","journal-title":"International Journal of Molecular Sciences"},{"key":"bib85","doi-asserted-by":"publisher","first-page":"671","DOI":"10.1038\/nmeth.2089","type":"journal-article","article-title":"NIH Image to ImageJ: 25 years of image analysis","volume":"9","author":"Schneider","year":"2012","journal-title":"Nature Methods"},{"key":"bib86","doi-asserted-by":"publisher","first-page":"12566","DOI":"10.1074\/jbc.M113.497438","type":"journal-article","article-title":"High resolution structure and double electron-electron resonance of the zebrafish voltage-dependent anion channel 2 reveal an oligomeric population","volume":"289","author":"Schredelseker","year":"2014","journal-title":"The Journal of Biological Chemistry"},{"key":"bib87","doi-asserted-by":"publisher","first-page":"10866","DOI":"10.1073\/pnas.1605104113","type":"journal-article","article-title":"A\u03b242 assembles into specific \u03b2-barrel pore-forming oligomers in membrane-mimicking environments","volume":"113","author":"Serra-Batiste","year":"2016","journal-title":"PNAS"},{"key":"bib88","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-021-24869-0","type":"journal-article","article-title":"Cardiac-specific deletion of voltage dependent anion channel 2 leads to dilated cardiomyopathy by altering calcium homeostasis","volume":"12","author":"Shankar","year":"2021","journal-title":"Nature Communications"},{"key":"bib89","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-020-17437-5","type":"journal-article","article-title":"Protein\u2013ligand binding with the coarse-grained Martini model","volume":"11","author":"Souza","year":"2020","journal-title":"Nature Communications"},{"key":"bib90","doi-asserted-by":"publisher","first-page":"382","DOI":"10.1038\/s41592-021-01098-3","type":"journal-article","article-title":"Martini 3: a general purpose force field for coarse-grained molecular dynamics","volume":"18","author":"Souza","year":"2021","journal-title":"Nature Methods"},{"key":"bib91","doi-asserted-by":"publisher","first-page":"27659","DOI":"10.1074\/jbc.M112.381939","type":"journal-article","article-title":"Phosphatidylinositol 4,5-bisphosphate (PI(4,5)P2)-dependent oligomerization of fibroblast growth factor 2 (FGF2) triggers the formation of a lipidic membrane pore implicated in unconventional secretion","volume":"287","author":"Steringer","year":"2012","journal-title":"The Journal of Biological Chemistry"},{"key":"bib92","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-025-62021-4","type":"journal-article","article-title":"Oligomer-based functions of mitochondrial porin","volume":"16","author":"Takeda","year":"2025","journal-title":"Nature Communications"},{"key":"bib93","doi-asserted-by":"publisher","DOI":"10.1126\/sciadv.ado4618","type":"journal-article","article-title":"ATR inhibition activates cancer cell cGAS\/STING-interferon signaling and promotes antitumor immunity in small-cell lung cancer","volume":"10","author":"Taniguchi","year":"2024","journal-title":"Science Advances"},{"key":"bib94","doi-asserted-by":"publisher","first-page":"23670","DOI":"10.1074\/jbc.M114.576314","type":"journal-article","article-title":"Acidification asymmetrically affects voltage-dependent anion channel implicating the involvement of salt bridges","volume":"289","author":"Teijido","year":"2014","journal-title":"The Journal of Biological Chemistry"},{"key":"bib95","doi-asserted-by":"publisher","first-page":"30081","DOI":"10.1074\/jbc.M313420200","type":"journal-article","article-title":"Lipidic pore formation by the concerted action of proapoptotic BAX and tBID","volume":"279","author":"Terrones","year":"2004","journal-title":"The Journal of Biological Chemistry"},{"key":"bib96","doi-asserted-by":"publisher","DOI":"10.1186\/s40035-022-00329-7","type":"journal-article","article-title":"Targeting the overexpressed mitochondrial protein VDAC1 in a mouse model of Alzheimer\u2019s disease protects against mitochondrial dysfunction and mitigates brain pathology","volume":"11","author":"Verma","year":"2022","journal-title":"Translational Neurodegeneration"},{"key":"bib97","doi-asserted-by":"publisher","DOI":"10.1093\/sleep\/zsad039","type":"journal-article","article-title":"Microglial voltage-dependent anion channel 1 signaling modulates sleep deprivation-induced transition to chronic postsurgical pain","volume":"46","author":"Wei","year":"2023","journal-title":"Sleep"},{"key":"bib98","doi-asserted-by":"publisher","first-page":"868","DOI":"10.1021\/ci990307l","type":"journal-article","article-title":"Prediction of physicochemical parameters by atomic contributions","volume":"39","author":"Wildman","year":"1999","journal-title":"Journal of Chemical Information and Computer Sciences"},{"key":"bib99","doi-asserted-by":"publisher","first-page":"89","DOI":"10.1007\/s002329900540","type":"journal-article","article-title":"Mouse VDAC isoforms expressed in yeast: channel properties and their roles in mitochondrial outer membrane permeability","volume":"170","author":"Xu","year":"1999","journal-title":"The Journal of Membrane Biology"},{"key":"bib100","doi-asserted-by":"publisher","first-page":"2787","DOI":"10.1042\/BST20210753","type":"journal-article","article-title":"VDAC2 and the BCL-2 family of proteins","volume":"49","author":"Yuan","year":"2021","journal-title":"Biochemical Society Transactions"},{"key":"bib101","doi-asserted-by":"publisher","first-page":"64","DOI":"10.1016\/j.cmet.2018.09.008","type":"journal-article","article-title":"Preserving insulin secretion in diabetes by inhibiting VDAC1 overexpression and surface translocation in \u03b2 Cells","volume":"29","author":"Zhang","year":"2019","journal-title":"Cell Metabolism"}],"container-title":["eLife"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/cdn.elifesciences.org\/articles\/111980\/elife-111980-v1.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/cdn.elifesciences.org\/articles\/111980\/elife-111980-v1.xml","content-type":"application\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/elifesciences.org\/articles\/111980","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,10,9]],"date-time":"2026-10-09T14:55:46Z","timestamp":1791557746000},"score":1,"resource":{"primary":{"URL":"https:\/\/elifesciences.org\/articles\/111980"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,10,9]]},"references-count":101,"alternative-id":["10.7554\/eLife.111980"],"URL":"https:\/\/doi.org\/10.7554\/elife.111980","archive":["CLOCKSS"],"relation":{"has-preprint":[{"id-type":"doi","id":"10.1101\/2025.06.30.661942","asserted-by":"subject"}],"is-same-as":[{"id-type":"doi","id":"10.7554\/eLife.111980.3","asserted-by":"subject"}],"is-version-of":[{"id-type":"doi","id":"10.7554\/eLife.111980.1","asserted-by":"subject"},{"id-type":"doi","id":"10.7554\/eLife.111980.2","asserted-by":"subject"}]},"ISSN":["2050-084X"],"issn-type":[{"value":"2050-084X","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,10,9]]},"assertion":[{"value":"single anonymised","name":"peer_review_transparency","label":"Peer review transparency","group":{"name":"peer_review_taxonomy"}},{"value":"other reviewer(s), editor","name":"peer_review_interaction","label":"Peer review interaction","group":{"name":"peer_review_taxonomy"}},{"value":"review summaries, review reports, author\/editor communication, reviewer identities reviewer opt in, editor identities","name":"peer_review_published","label":"Peer review published","group":{"name":"peer_review_taxonomy"}},{"value":"open (sign in with ORCID iD required)","name":"post_publication_commenting","label":"Post publication commenting","group":{"name":"post_publication_commenting"}}],"article-number":"RP111980"}}