{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,5,27]],"date-time":"2026-05-27T15:48:32Z","timestamp":1779896912354,"version":"3.53.1"},"reference-count":49,"publisher":"Elsevier BV","issue":"4","license":[{"start":{"date-parts":[[2011,2,1]],"date-time":"2011-02-01T00:00:00Z","timestamp":1296518400000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/tdm\/userlicense\/1.0\/"},{"start":{"date-parts":[[2011,2,1]],"date-time":"2011-02-01T00:00:00Z","timestamp":1296518400000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/legal\/tdmrep-license"},{"start":{"date-parts":[[2013,7,16]],"date-time":"2013-07-16T00:00:00Z","timestamp":1373932800000},"content-version":"vor","delay-in-days":896,"URL":"http:\/\/www.elsevier.com\/open-access\/userlicense\/1.0\/"}],"funder":[{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"publisher","award":["GM55203"],"award-info":[{"award-number":["GM55203"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["cell.com","elsevier.com","sciencedirect.com"],"crossmark-restriction":true},"short-container-title":["Biophysical Journal"],"published-print":{"date-parts":[[2011,2]]},"DOI":"10.1016\/j.bpj.2011.01.013","type":"journal-article","created":{"date-parts":[[2011,2,16]],"date-time":"2011-02-16T11:35:13Z","timestamp":1297856113000},"page":"987-995","update-policy":"https:\/\/doi.org\/10.1016\/elsevier_cm_policy","source":"Crossref","is-referenced-by-count":24,"title":["Adhesion and Merging of Lipid Bilayers: A Method for Measuring the Free Energy of Adhesion and Hemifusion"],"prefix":"10.1016","volume":"100","author":[{"given":"Yen","family":"Sun","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Chang-Chun","family":"Lee","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Huey W.","family":"Huang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"78","reference":[{"key":"10.1016\/j.bpj.2011.01.013_bib1","doi-asserted-by":"crossref","first-page":"423","DOI":"10.1016\/S0006-3495(84)84039-4","article-title":"Free energy potential for aggregation of giant, neutral lipid bilayer vesicles by Van der Waals attraction","volume":"46","author":"Evans","year":"1984","journal-title":"Biophys.\u00a0J."},{"key":"10.1016\/j.bpj.2011.01.013_bib2","doi-asserted-by":"crossref","first-page":"3003","DOI":"10.1021\/la950802l","article-title":"Direct measurement of polyethylene glycol induced depletion attraction between lipid bilayers","volume":"12","author":"Kuhl","year":"1996","journal-title":"Langmuir"},{"key":"10.1016\/j.bpj.2011.01.013_bib3","doi-asserted-by":"crossref","first-page":"351","DOI":"10.1016\/0304-4157(89)90010-5","article-title":"Hydration forces between phospholipid bilayers","volume":"988","author":"Rand","year":"1989","journal-title":"Biochim. Biophys. Acta"},{"key":"10.1016\/j.bpj.2011.01.013_bib4","first-page":"305","article-title":"Steric interaction of fluid membranes in multilayer systems","volume":"33a","author":"Helfrich","year":"1978","journal-title":"Z.\u00a0Naturforsch.\u00a0C."},{"key":"10.1016\/j.bpj.2011.01.013_bib5","doi-asserted-by":"crossref","first-page":"203","DOI":"10.1021\/la047876u","article-title":"Distorted hexagonal phase studied by neutron diffraction: lipid components demixed in a bent monolayer","volume":"21","author":"Ding","year":"2005","journal-title":"Langmuir"},{"key":"10.1016\/j.bpj.2011.01.013_bib6","doi-asserted-by":"crossref","first-page":"2819","DOI":"10.1529\/biophysj.106.097923","article-title":"Evidence of cholesterol accumulated in high curvature regions: implication to the curvature elastic energy for lipid mixtures","volume":"92","author":"Wang","year":"2007","journal-title":"Biophys.\u00a0J."},{"key":"10.1016\/j.bpj.2011.01.013_bib7","first-page":"361","article-title":"On the theory of membrane fusion. The stalk mechanism","volume":"3","author":"Markin","year":"1984","journal-title":"Gen. Physiol. Biophys."},{"key":"10.1016\/j.bpj.2011.01.013_bib8","doi-asserted-by":"crossref","first-page":"2124","DOI":"10.1016\/S0006-3495(93)81256-6","article-title":"Energetics of intermediates in membrane fusion: comparison of stalk and inverted micellar intermediate mechanisms","volume":"65","author":"Siegel","year":"1993","journal-title":"Biophys.\u00a0J."},{"key":"10.1016\/j.bpj.2011.01.013_bib9","doi-asserted-by":"crossref","first-page":"7235","DOI":"10.1073\/pnas.121191898","article-title":"A quantitative model for membrane fusion based on low-energy intermediates","volume":"98","author":"Kuzmin","year":"2001","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"10.1016\/j.bpj.2011.01.013_bib10","doi-asserted-by":"crossref","first-page":"882","DOI":"10.1016\/S0006-3495(02)75450-7","article-title":"Stalk model of membrane fusion: solution of energy crisis","volume":"82","author":"Kozlovsky","year":"2002","journal-title":"Biophys.\u00a0J."},{"key":"10.1016\/j.bpj.2011.01.013_bib11","doi-asserted-by":"crossref","first-page":"693","DOI":"10.1016\/S0006-3495(02)75432-5","article-title":"Membrane fusion: stalk model revisited","volume":"82","author":"Markin","year":"2002","journal-title":"Biophys.\u00a0J."},{"key":"10.1016\/j.bpj.2011.01.013_bib12","doi-asserted-by":"crossref","first-page":"555","DOI":"10.1016\/S0006-3495(02)75420-9","article-title":"Filling potholes on the path to fusion pores","volume":"82","author":"Lentz","year":"2002","journal-title":"Biophys.\u00a0J."},{"key":"10.1016\/j.bpj.2011.01.013_bib13","doi-asserted-by":"crossref","first-page":"627","DOI":"10.1146\/annurev.cellbio.12.1.627","article-title":"Virus-cell and cell-cell fusion","volume":"12","author":"Hernandez","year":"1996","journal-title":"Annu. Rev. Cell Dev. Biol."},{"key":"10.1016\/j.bpj.2011.01.013_bib14","doi-asserted-by":"crossref","first-page":"81","DOI":"10.1083\/jcb.136.1.81","article-title":"An early stage of membrane fusion mediated by the low pH conformation of influenza hemagglutinin depends upon membrane lipids","volume":"136","author":"Chernomordik","year":"1997","journal-title":"J.\u00a0Cell Biol."},{"key":"10.1016\/j.bpj.2011.01.013_bib15","doi-asserted-by":"crossref","first-page":"2451","DOI":"10.1529\/biophysj.106.096362","article-title":"Solution pH alters mechanical and electrical properties of phosphatidylcholine membranes: relation between interfacial electrostatics, intramembrane potential, and bending elasticity","volume":"92","author":"Zhou","year":"2007","journal-title":"Biophys.\u00a0J."},{"key":"10.1016\/j.bpj.2011.01.013_bib16","doi-asserted-by":"crossref","first-page":"1822","DOI":"10.1021\/ma00184a049","article-title":"Attraction between lipid bilayer membranes in concentrated solutions of nonadsorbing polymers: comparison of mean-field theory with measurements of adhesion energy","volume":"21","author":"Evans","year":"1988","journal-title":"Macromolecules"},{"key":"10.1016\/j.bpj.2011.01.013_bib17","doi-asserted-by":"crossref","first-page":"35109","DOI":"10.1074\/jbc.M301726200","article-title":"Antennapedia and HIV transactivator of transcription (TAT) \u201cprotein transduction domains\u201d promote endocytosis of high molecular weight cargo upon binding to cell surface glycosaminoglycans","volume":"278","author":"Console","year":"2003","journal-title":"J.\u00a0Biol. Chem."},{"key":"10.1016\/j.bpj.2011.01.013_bib18","doi-asserted-by":"crossref","first-page":"15300","DOI":"10.1074\/jbc.M401604200","article-title":"Cellular uptake of unconjugated TAT peptide involves clathrin-dependent endocytosis and heparan sulfate receptors","volume":"280","author":"Richard","year":"2005","journal-title":"J.\u00a0Biol. Chem."},{"key":"10.1016\/j.bpj.2011.01.013_bib19","doi-asserted-by":"crossref","first-page":"310","DOI":"10.1038\/nm996","article-title":"Transducible TAT-HA fusogenic peptide enhances escape of TAT-fusion proteins after lipid raft macropinocytosis","volume":"10","author":"Wadia","year":"2004","journal-title":"Nat. Med."},{"key":"10.1016\/j.bpj.2011.01.013_bib20","doi-asserted-by":"crossref","first-page":"2525","DOI":"10.1016\/j.bpj.2010.08.029","article-title":"Cell-penetrating peptide induces leaky fusion of liposomes containing late endosome-specific anionic lipid","volume":"99","author":"Yang","year":"2010","journal-title":"Biophys.\u00a0J."},{"key":"10.1016\/j.bpj.2011.01.013_bib21","doi-asserted-by":"crossref","first-page":"425","DOI":"10.1016\/S0006-3495(80)85069-7","article-title":"Analysis of adhesion of large vesicles to surfaces","volume":"31","author":"Evans","year":"1980","journal-title":"Biophys.\u00a0J."},{"key":"10.1016\/j.bpj.2011.01.013_bib22","doi-asserted-by":"crossref","first-page":"715","DOI":"10.1016\/S0006-3495(84)84213-7","article-title":"Free energy potential for aggregation of mixed phosphatidylcholine\/phosphatidylserine lipid vesicles in glucose polymer (dextran) solutions","volume":"45","author":"Evans","year":"1984","journal-title":"Biophys.\u00a0J."},{"key":"10.1016\/j.bpj.2011.01.013_bib23","doi-asserted-by":"crossref","first-page":"4219","DOI":"10.1021\/j100300a003","article-title":"Physical properties of surfactant bilayer membranes: thermal transitions, elasticity, rigidity, cohesion, and colloidal interactions","volume":"91","author":"Evans","year":"1987","journal-title":"J.\u00a0Phys. Chem."},{"key":"10.1016\/j.bpj.2011.01.013_bib24","series-title":"Proteins at Interfaces","first-page":"88","article-title":"Nonspecific adhesion of phospholipid bilayer membranes in solutions of plasma proteins: measurement of free energy potentials and theoretical concepts","author":"Evans","year":"1987"},{"key":"10.1016\/j.bpj.2011.01.013_bib25","series-title":"Giant Vesicles","first-page":"27","article-title":"Liposome electroformation","author":"Angelova","year":"2000"},{"key":"10.1016\/j.bpj.2011.01.013_bib26","doi-asserted-by":"crossref","first-page":"2318","DOI":"10.1529\/biophysj.108.133736","article-title":"The bound states of amphipathic drugs in lipid bilayers: study of curcumin","volume":"95","author":"Sun","year":"2008","journal-title":"Biophys.\u00a0J."},{"key":"10.1016\/j.bpj.2011.01.013_bib27","doi-asserted-by":"crossref","first-page":"675","DOI":"10.1038\/nsmb.1455","article-title":"Mechanics of membrane fusion","volume":"15","author":"Chernomordik","year":"2008","journal-title":"Nat. Struct. Mol. Biol."},{"key":"10.1016\/j.bpj.2011.01.013_bib28","doi-asserted-by":"crossref","first-page":"269","DOI":"10.1140\/epje\/i2003-10151-2","article-title":"Hemifusion and fusion of giant vesicles induced by reduction of inter-membrane distance","volume":"14","author":"Heuvingh","year":"2004","journal-title":"Eur. Phys. J. E. Soft Matter"},{"key":"10.1016\/j.bpj.2011.01.013_bib29","doi-asserted-by":"crossref","first-page":"281","DOI":"10.1016\/j.bbagen.2005.04.028","article-title":"Indirect evidence of submicroscopic pores in giant unilamellar [correction of unilamelar] vesicles","volume":"1724","author":"Rodriguez","year":"2005","journal-title":"Biochim. Biophys. Acta"},{"key":"10.1016\/j.bpj.2011.01.013_bib30","doi-asserted-by":"crossref","first-page":"328","DOI":"10.1016\/S0006-3495(00)76295-3","article-title":"Effect of chain length and unsaturation on elasticity of lipid bilayers","volume":"79","author":"Rawicz","year":"2000","journal-title":"Biophys.\u00a0J."},{"key":"10.1016\/j.bpj.2011.01.013_bib31","doi-asserted-by":"crossref","first-page":"637","DOI":"10.1016\/S0006-3495(81)84817-5","article-title":"Thermoelasticity of large lecithin bilayer vesicles","volume":"35","author":"Kwok","year":"1981","journal-title":"Biophys.\u00a0J."},{"key":"10.1016\/j.bpj.2011.01.013_bib32","doi-asserted-by":"crossref","first-page":"1026","DOI":"10.1016\/j.bpj.2008.11.007","article-title":"Interaction of tea catechin (\u2212)-epigallocatechin gallate with lipid bilayers","volume":"96","author":"Sun","year":"2009","journal-title":"Biophys.\u00a0J."},{"key":"10.1016\/j.bpj.2011.01.013_bib33","doi-asserted-by":"crossref","first-page":"544","DOI":"10.1016\/j.bpj.2010.04.045","article-title":"Kinetic process of beta-amyloid formation via membrane binding","volume":"99","author":"Sun","year":"2010","journal-title":"Biophys.\u00a0J."},{"key":"10.1016\/j.bpj.2011.01.013_bib34","doi-asserted-by":"crossref","first-page":"89","DOI":"10.1016\/0005-2736(93)90319-U","article-title":"Tension-stabilized pores in giant vesicles: determination of pore size and pore line tension","volume":"1147","author":"Zhelev","year":"1993","journal-title":"Biochim. Biophys. Acta"},{"key":"10.1016\/j.bpj.2011.01.013_bib35","doi-asserted-by":"crossref","first-page":"109","DOI":"10.1016\/0006-3002(63)90990-9","article-title":"The denaturation of proteins. IV. Conalbumin and iron(III)-conalbumin in urea solution","volume":"71","author":"Glazer","year":"1963","journal-title":"Biochim. Biophys. Acta"},{"key":"10.1016\/j.bpj.2011.01.013_bib36","doi-asserted-by":"crossref","first-page":"922","DOI":"10.1016\/S0006-3495(95)79966-0","article-title":"The hemifusion intermediate and its conversion to complete fusion: regulation by membrane composition","volume":"69","author":"Chernomordik","year":"1995","journal-title":"Biophys.\u00a0J."},{"key":"10.1016\/j.bpj.2011.01.013_bib37","doi-asserted-by":"crossref","first-page":"5160","DOI":"10.1021\/la049681s","article-title":"Membrane fusion of giant unilamellar vesicles of neutral phospholipid membranes induced by La3+","volume":"20","author":"Tanaka","year":"2004","journal-title":"Langmuir"},{"key":"10.1016\/j.bpj.2011.01.013_bib38","doi-asserted-by":"crossref","first-page":"15841","DOI":"10.1073\/pnas.0602766103","article-title":"Time scales of membrane fusion revealed by direct imaging of vesicle fusion with high temporal resolution","volume":"103","author":"Haluska","year":"2006","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"10.1016\/j.bpj.2011.01.013_bib39","doi-asserted-by":"crossref","first-page":"318","DOI":"10.1038\/362318a0","article-title":"SNAP receptors implicated in vesicle targeting and fusion","volume":"362","author":"S\u00f6llner","year":"1993","journal-title":"Nature"},{"key":"10.1016\/j.bpj.2011.01.013_bib40","doi-asserted-by":"crossref","first-page":"474","DOI":"10.1126\/science.1161748","article-title":"Membrane fusion: grappling with SNARE and SM proteins","volume":"323","author":"S\u00fcdhof","year":"2009","journal-title":"Science"},{"key":"10.1016\/j.bpj.2011.01.013_bib41","doi-asserted-by":"crossref","first-page":"665","DOI":"10.1038\/nsmb.1450","article-title":"Synaptic vesicle fusion","volume":"15","author":"Rizo","year":"2008","journal-title":"Nat. Struct. Mol. Biol."},{"key":"10.1016\/j.bpj.2011.01.013_bib42","doi-asserted-by":"crossref","first-page":"658","DOI":"10.1038\/nsmb.1451","article-title":"Membrane fusion","volume":"15","author":"Wickner","year":"2008","journal-title":"Nat. Struct. Mol. Biol."},{"key":"10.1016\/j.bpj.2011.01.013_bib43","doi-asserted-by":"crossref","first-page":"543","DOI":"10.1038\/nrm2417","article-title":"Mechanisms of membrane fusion: disparate players and common principles","volume":"9","author":"Martens","year":"2008","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"10.1016\/j.bpj.2011.01.013_bib44","doi-asserted-by":"crossref","first-page":"512","DOI":"10.1126\/science.1166500","article-title":"Alternative zippering as an on-off switch for SNARE-mediated fusion","volume":"323","author":"Giraudo","year":"2009","journal-title":"Science"},{"key":"10.1016\/j.bpj.2011.01.013_bib45","doi-asserted-by":"crossref","first-page":"516","DOI":"10.1126\/science.1166505","article-title":"Complexin controls the force transfer from SNARE complexes to membranes in fusion","volume":"323","author":"Maximov","year":"2009","journal-title":"Science"},{"key":"10.1016\/j.bpj.2011.01.013_bib46","doi-asserted-by":"crossref","first-page":"105","DOI":"10.1083\/jcb.150.1.105","article-title":"Close is not enough: SNARE-dependent membrane fusion requires an active mechanism that transduces force to membrane anchors","volume":"150","author":"McNew","year":"2000","journal-title":"J.\u00a0Cell Biol."},{"key":"10.1016\/j.bpj.2011.01.013_bib47","doi-asserted-by":"crossref","first-page":"699","DOI":"10.1016\/j.tibs.2010.06.003","article-title":"Protein-driven membrane stresses in fusion and fission","volume":"35","author":"Kozlov","year":"2010","journal-title":"Trends Biochem. Sci."},{"key":"10.1016\/j.bpj.2011.01.013_bib48","doi-asserted-by":"crossref","first-page":"601","DOI":"10.1016\/j.cell.2010.02.017","article-title":"Membrane curvature in synaptic vesicle fusion and beyond","volume":"140","author":"McMahon","year":"2010","journal-title":"Cell"},{"key":"10.1016\/j.bpj.2011.01.013_bib49","doi-asserted-by":"crossref","first-page":"1877","DOI":"10.1126\/science.1074354","article-title":"Observation of a membrane fusion intermediate structure","volume":"297","author":"Yang","year":"2002","journal-title":"Science"}],"container-title":["Biophysical Journal"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S0006349511000592?httpAccept=text\/xml","content-type":"text\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S0006349511000592?httpAccept=text\/plain","content-type":"text\/plain","content-version":"vor","intended-application":"text-mining"}],"deposited":{"date-parts":[[2025,11,2]],"date-time":"2025-11-02T16:29:52Z","timestamp":1762100992000},"score":1,"resource":{"primary":{"URL":"https:\/\/linkinghub.elsevier.com\/retrieve\/pii\/S0006349511000592"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2011,2]]},"references-count":49,"journal-issue":{"issue":"4","published-print":{"date-parts":[[2011,2]]}},"alternative-id":["S0006349511000592"],"URL":"https:\/\/doi.org\/10.1016\/j.bpj.2011.01.013","relation":{},"ISSN":["0006-3495"],"issn-type":[{"value":"0006-3495","type":"print"}],"subject":[],"published":{"date-parts":[[2011,2]]},"assertion":[{"value":"Elsevier","name":"publisher","label":"This article is maintained by"},{"value":"Adhesion and Merging of Lipid Bilayers: A Method for Measuring the Free Energy of Adhesion and Hemifusion","name":"articletitle","label":"Article Title"},{"value":"Biophysical Journal","name":"journaltitle","label":"Journal Title"},{"value":"https:\/\/doi.org\/10.1016\/j.bpj.2011.01.013","name":"articlelink","label":"CrossRef DOI link to publisher maintained version"},{"value":"article","name":"content_type","label":"Content Type"},{"value":"Copyright \u00a9 2011 Biophysical Society. Published by Elsevier Inc. All rights reserved.","name":"copyright","label":"Copyright"}]}}