{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,25]],"date-time":"2026-10-25T03:57:58Z","timestamp":1792900678121,"version":"4.5.0"},"reference-count":47,"publisher":"Elsevier BV","issue":"12","license":[{"start":{"date-parts":[[2016,12,1]],"date-time":"2016-12-01T00:00:00Z","timestamp":1480550400000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/tdm\/userlicense\/1.0\/"},{"start":{"date-parts":[[2017,12,20]],"date-time":"2017-12-20T00:00:00Z","timestamp":1513728000000},"content-version":"vor","delay-in-days":384,"URL":"https:\/\/www.elsevier.com\/open-access\/userlicense\/1.0\/"}],"funder":[{"name":"NIH","award":["R01 GM106720"],"award-info":[{"award-number":["R01 GM106720"]}]},{"name":"Stanley Cohen Innovation Fund","award":["DK048106"],"award-info":[{"award-number":["DK048106"]}]},{"name":"Stanley Cohen Innovation Fund","award":["DK084424"],"award-info":[{"award-number":["DK084424"]}]},{"name":"Stanley Cohen Innovation Fund","award":["DK034854"],"award-info":[{"award-number":["DK034854"]}]},{"name":"Harvard Digestive Disease Center"}],"content-domain":{"domain":["biophysj.org","elsevier.com","sciencedirect.com"],"crossmark-restriction":true},"short-container-title":["Biophysical Journal"],"published-print":{"date-parts":[[2016,12]]},"DOI":"10.1016\/j.bpj.2016.11.008","type":"journal-article","created":{"date-parts":[[2016,11,30]],"date-time":"2016-11-30T07:30:18Z","timestamp":1480491018000},"page":"2547-2550","update-policy":"https:\/\/doi.org\/10.1016\/elsevier_cm_policy","source":"Crossref","is-referenced-by-count":39,"title":["Glycolipid Crosslinking Is Required for Cholera Toxin to Partition Into and Stabilize Ordered Domains"],"prefix":"10.1016","volume":"111","author":[{"given":"Krishnan","family":"Raghunathan","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Tiffany H.","family":"Wong","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Daniel J.","family":"Chinnapen","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Wayne I.","family":"Lencer","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Michael G.","family":"Jobling","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Anne K.","family":"Kenworthy","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"78","reference":[{"key":"10.1016\/j.bpj.2016.11.008_bib1","doi-asserted-by":"crossref","first-page":"46","DOI":"10.1126\/science.1174621","article-title":"Lipid rafts as a membrane-organizing principle","volume":"327","author":"Lingwood","year":"2010","journal-title":"Science"},{"key":"10.1016\/j.bpj.2016.11.008_bib2","doi-asserted-by":"crossref","first-page":"3165","DOI":"10.1073\/pnas.0611357104","article-title":"Large-scale fluid\/fluid phase separation of proteins and lipids in giant plasma membrane vesicles","volume":"104","author":"Baumgart","year":"2007","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"10.1016\/j.bpj.2016.11.008_bib3","doi-asserted-by":"crossref","first-page":"287","DOI":"10.1021\/cb800012x","article-title":"Critical fluctuations in plasma membrane vesicles","volume":"3","author":"Veatch","year":"2008","journal-title":"ACS Chem. Biol."},{"key":"10.1016\/j.bpj.2016.11.008_bib4","doi-asserted-by":"crossref","first-page":"20","DOI":"10.1016\/j.bbamem.2007.08.028","article-title":"Structural determinants for partitioning of lipids and proteins between coexisting fluid phases in giant plasma membrane vesicles","volume":"1778","author":"Sengupta","year":"2008","journal-title":"Biochim. Biophys. Acta"},{"key":"10.1016\/j.bpj.2016.11.008_bib5","doi-asserted-by":"crossref","first-page":"135","DOI":"10.1042\/bse0570135","article-title":"Functions of cholera toxin B-subunit as a raft cross-linker","volume":"57","author":"Day","year":"2015","journal-title":"Essays Biochem."},{"key":"10.1016\/j.bpj.2016.11.008_bib6","doi-asserted-by":"crossref","first-page":"10005","DOI":"10.1073\/pnas.0804374105","article-title":"Plasma membranes are poised for activation of raft phase coalescence at physiological temperature","volume":"105","author":"Lingwood","year":"2008","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"10.1016\/j.bpj.2016.11.008_bib7","doi-asserted-by":"crossref","first-page":"1427","DOI":"10.1016\/j.bbamem.2010.03.009","article-title":"Temperature-dependent phase behavior and protein partitioning in giant plasma membrane vesicles","volume":"1798","author":"Johnson","year":"2010","journal-title":"Biochim. Biophys. Acta."},{"key":"10.1016\/j.bpj.2016.11.008_bib8","doi-asserted-by":"crossref","first-page":"6320","DOI":"10.1073\/pnas.0405654102","article-title":"Crosslinking a lipid raft component triggers liquid ordered-liquid disordered phase separation in model plasma membranes","volume":"102","author":"Hammond","year":"2005","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"10.1016\/j.bpj.2016.11.008_bib9","doi-asserted-by":"crossref","first-page":"1777","DOI":"10.1016\/j.bbamem.2012.03.007","article-title":"Partitioning, diffusion, and ligand binding of raft lipid analogs in model and cellular plasma membranes","volume":"1818","author":"Sezgin","year":"2012","journal-title":"Biochim. Biophys. Acta."},{"key":"10.1016\/j.bpj.2016.11.008_bib10","doi-asserted-by":"crossref","first-page":"2104","DOI":"10.1016\/j.bpj.2012.03.054","article-title":"Dynamics and size of cross-linking-induced lipid nanodomains in model membranes","volume":"102","author":"\u0160tefl","year":"2012","journal-title":"Biophys. J."},{"key":"10.1016\/j.bpj.2016.11.008_bib11","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1128\/mBio.00401-12","article-title":"A single native\u00a0ganglioside GM1-binding site is sufficient for cholera toxin\u00a0to\u00a0bind to cells and complete the intoxication pathway","volume":"3","author":"Jobling","year":"2012","journal-title":"MBio"},{"key":"10.1016\/j.bpj.2016.11.008_bib12","doi-asserted-by":"crossref","first-page":"2751","DOI":"10.1016\/j.bpj.2013.11.005","article-title":"Liquid general anesthetics lower critical temperatures in plasma membrane vesicles","volume":"105","author":"Gray","year":"2013","journal-title":"Biophys. J."},{"key":"10.1016\/j.bpj.2016.11.008_bib13","doi-asserted-by":"crossref","first-page":"e0140925","DOI":"10.1371\/journal.pone.0140925","article-title":"Membrane transition temperature determines cisplatin response","volume":"10","author":"Raghunathan","year":"2015","journal-title":"PLoS One"},{"key":"10.1016\/j.bpj.2016.11.008_bib14","doi-asserted-by":"crossref","first-page":"3272","DOI":"10.1073\/pnas.0408215102","article-title":"Sterol structure determines the separation of phases and the curvature of the liquid-ordered phase in model membranes","volume":"102","author":"Bacia","year":"2005","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"10.1016\/j.bpj.2016.11.008_bib15","doi-asserted-by":"crossref","first-page":"104","DOI":"10.1007\/s13361-013-0751-5","article-title":"Measuring positive cooperativity using the direct ESI-MS assay. Cholera toxin B subunit homopentamer binding to GM1 pentasaccharide","volume":"25","author":"Lin","year":"2014","journal-title":"J.\u00a0Am. Soc. Mass Spectrom."},{"key":"10.1016\/j.bpj.2016.11.008_bib16","doi-asserted-by":"crossref","first-page":"3126","DOI":"10.1128\/IAI.55.12.3126-3130.1987","article-title":"Differential binding kinetics of cholera toxin to intestinal microvillus membrane during development","volume":"55","author":"Lencer","year":"1987","journal-title":"Infect. Immun."},{"key":"10.1016\/j.bpj.2016.11.008_bib17","doi-asserted-by":"crossref","first-page":"5954","DOI":"10.1021\/ja069375w","article-title":"GM1 clustering inhibits cholera toxin binding in supported phospholipid membranes","volume":"129","author":"Shi","year":"2007","journal-title":"J.\u00a0Am. Chem. Soc."},{"key":"10.1016\/j.bpj.2016.11.008_bib18","doi-asserted-by":"crossref","first-page":"36049","DOI":"10.1074\/jbc.M110.110189","article-title":"A major fraction of glycosphingolipids in model and cellular cholesterol-containing membranes is undetectable by their binding proteins","volume":"285","author":"Mahfoud","year":"2010","journal-title":"J.\u00a0Biol. Chem."},{"key":"10.1016\/j.bpj.2016.11.008_bib19","doi-asserted-by":"crossref","first-page":"260","DOI":"10.1038\/nchembio.551","article-title":"Cholesterol modulates glycolipid conformation and receptor activity","volume":"7","author":"Lingwood","year":"2011","journal-title":"Nat. Chem. Biol."},{"key":"10.1016\/j.bpj.2016.11.008_bib20","doi-asserted-by":"crossref","first-page":"850","DOI":"10.1016\/j.bbamcr.2014.07.016","article-title":"On multivalent receptor activity of GM1 in cholesterol containing membranes","volume":"1853","author":"\u0160achl","year":"2014","journal-title":"Biochim. Biophys. Acta."},{"key":"10.1016\/j.bpj.2016.11.008_bib21","doi-asserted-by":"crossref","first-page":"573","DOI":"10.1016\/j.devcel.2012.08.002","article-title":"Lipid\u00a0sorting by ceramide structure from plasma membrane to ER for\u00a0the cholera toxin receptor ganglioside GM1","volume":"23","author":"Chinnapen","year":"2012","journal-title":"Dev. Cell"},{"key":"10.1016\/j.bpj.2016.11.008_bib22","doi-asserted-by":"crossref","first-page":"1476","DOI":"10.1128\/IAI.01286-07","article-title":"Attenuated endocytosis and toxicity of a mutant cholera toxin with decreased ability to cluster ganglioside GM1 molecules","volume":"76","author":"Wolf","year":"2008","journal-title":"Infect. Immun."},{"key":"10.1016\/j.bpj.2016.11.008_bib23","doi-asserted-by":"crossref","first-page":"11","DOI":"10.1038\/ncb1999","article-title":"GM1 structure determines SV40-induced membrane invagination and infection","volume":"12","author":"Ewers","year":"2010","journal-title":"Nat. Cell Biol."},{"key":"10.1016\/j.bpj.2016.11.008_bib24","doi-asserted-by":"crossref","first-page":"670","DOI":"10.1038\/nature05996","article-title":"Shiga toxin induces tubular membrane invaginations for its uptake into cells","volume":"450","author":"R\u00f6mer","year":"2007","journal-title":"Nature"},{"key":"10.1016\/j.bpj.2016.11.008_bib25","doi-asserted-by":"crossref","first-page":"407","DOI":"10.1016\/j.tibs.2015.04.005","article-title":"The multi-tasked life of GM1 ganglioside, a true factotum of nature","volume":"40","author":"Ledeen","year":"2015","journal-title":"Trends Biochem. Sci."},{"key":"10.1016\/j.bpj.2016.11.008_bib26","doi-asserted-by":"crossref","first-page":"90","DOI":"10.1016\/j.bbalip.2014.06.009","article-title":"Organization and functions of glycolipid-enriched microdomains in phagocytes","volume":"1851","author":"Ekyalongo","year":"2015","journal-title":"Biochim. Biophys Acta."},{"key":"10.1016\/j.bpj.2016.11.008_bib27","doi-asserted-by":"crossref","first-page":"6433","DOI":"10.1021\/bi200771m","article-title":"Mechanism of amyloid \u03b2-protein aggregation mediated by GM1 ganglioside clusters","volume":"50","author":"Ikeda","year":"2011","journal-title":"Biochemistry"},{"key":"10.1016\/j.bpj.2016.11.008_bib28","doi-asserted-by":"crossref","first-page":"289","DOI":"10.1242\/jcs.030338","article-title":"Binding of laminin-1 to monosialoganglioside GM1 in lipid rafts is crucial for neurite outgrowth","volume":"122","author":"Ichikawa","year":"2009","journal-title":"J.\u00a0Cell Sci."},{"key":"10.1016\/j.bpj.2016.11.008_bib29","doi-asserted-by":"crossref","first-page":"203","DOI":"10.1016\/j.bbamcr.2005.09.002","article-title":"How principles of domain formation in model membranes may explain ambiguities concerning lipid raft formation in cells","volume":"1746","author":"London","year":"2005","journal-title":"Biochim. Biophys. Acta."},{"key":"10.1016\/j.bpj.2016.11.008_bib30","doi-asserted-by":"crossref","first-page":"14267","DOI":"10.1021\/bi035759c","article-title":"Steroid structural requirements for stabilizing or disrupting lipid domains","volume":"42","author":"Wenz","year":"2003","journal-title":"Biochemistry"},{"key":"10.1016\/j.bpj.2016.11.008_bib31","doi-asserted-by":"crossref","first-page":"1800","DOI":"10.1016\/j.bpj.2016.03.012","article-title":"Polyunsaturated lipids regulate membrane domain stability by tuning membrane order","volume":"110","author":"Levental","year":"2016","journal-title":"Biophys. J."},{"key":"10.1016\/j.bpj.2016.11.008_bib32","doi-asserted-by":"crossref","first-page":"4935","DOI":"10.1016\/j.bpj.2009.04.019","article-title":"Theory of raft formation by the cross-linking of saturated or unsaturated lipids in model lipid bilayers","volume":"96","author":"Putzel","year":"2009","journal-title":"Biophys. J."},{"key":"10.1016\/j.bpj.2016.11.008_bib33","doi-asserted-by":"crossref","first-page":"138101","DOI":"10.1103\/PhysRevLett.109.138101","article-title":"Critical Casimir forces in cellular membranes","volume":"109","author":"Machta","year":"2012","journal-title":"Phys. Rev. Lett."},{"key":"10.1016\/j.bpj.2016.11.008_bib34","doi-asserted-by":"crossref","first-page":"e01671","DOI":"10.7554\/eLife.01671","article-title":"A lipid bound actin meshwork organizes liquid phase separation in model membranes","volume":"3","author":"Honigmann","year":"2014","journal-title":"eLife"},{"key":"10.1016\/j.bpj.2016.11.008_bib35","doi-asserted-by":"crossref","first-page":"257","DOI":"10.1146\/annurev.biophys.32.110601.142439","article-title":"The state of lipid rafts: from model membranes to cells","volume":"32","author":"Edidin","year":"2003","journal-title":"Annu. Rev. Biophys. Biomol. Struct."},{"key":"10.1016\/j.bpj.2016.11.008_bib36","doi-asserted-by":"crossref","first-page":"269","DOI":"10.1146\/annurev.biophys.32.110601.141803","article-title":"Model systems, lipid rafts, and cell membranes","volume":"33","author":"Simons","year":"2004","journal-title":"Annu. Rev. Biophys."},{"key":"10.1016\/j.bpj.2016.11.008_bib37","doi-asserted-by":"crossref","first-page":"1109","DOI":"10.1016\/j.jmb.2004.12.026","article-title":"Lipid rafts have different sizes depending on membrane composition: a time-resolved fluorescence resonance energy transfer study","volume":"346","author":"de Almeida","year":"2005","journal-title":"J.\u00a0Mol. Biol."},{"key":"10.1016\/j.bpj.2016.11.008_bib38","doi-asserted-by":"crossref","first-page":"9105","DOI":"10.1021\/acs.langmuir.5b01866","article-title":"Nanodomain formation of ganglioside GM1 in lipid membrane: effects of Cholera toxin mediated crosslinking","volume":"31","author":"Sun","year":"2015","journal-title":"Langmuir"},{"key":"10.1016\/j.bpj.2016.11.008_bib39","doi-asserted-by":"crossref","first-page":"2676","DOI":"10.1016\/j.bpj.2008.11.067","article-title":"Sorting of lipids and proteins in membrane curvature gradients","volume":"96","author":"Tian","year":"2009","journal-title":"Biophys. J."},{"key":"10.1016\/j.bpj.2016.11.008_bib40","doi-asserted-by":"crossref","first-page":"1519","DOI":"10.1111\/j.1600-0854.2010.01116.x","article-title":"Lipid cosorting mediated by Shiga toxin induced tubulation","volume":"11","author":"Safouane","year":"2010","journal-title":"Traffic"},{"key":"10.1016\/j.bpj.2016.11.008_bib41","doi-asserted-by":"crossref","first-page":"572","DOI":"10.1111\/tra.12269","article-title":"Microtubule motors power plasma membrane tubulation in clathrin-independent endocytosis","volume":"16","author":"Day","year":"2015","journal-title":"Traffic"},{"key":"10.1016\/j.bpj.2016.11.008_bib42","doi-asserted-by":"crossref","first-page":"14662","DOI":"10.1073\/pnas.011442598","article-title":"Identification of motifs in cholera toxin A1 polypeptide that are required for its interaction with human ADP-ribosylation factor 6 in a bacterial two-hybrid system","volume":"97","author":"Jobling","year":"2000","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"10.1016\/j.bpj.2016.11.008_bib43","doi-asserted-by":"crossref","first-page":"1755","DOI":"10.1111\/j.1365-2958.1991.tb01925.x","article-title":"Analysis of structure and function of the B subunit of cholera toxin by the use of site-directed mutagenesis","volume":"5","author":"Jobling","year":"1991","journal-title":"Mol. Microbiol."},{"key":"10.1016\/j.bpj.2016.11.008_bib44","doi-asserted-by":"crossref","first-page":"1516","DOI":"10.1002\/pro.5560060716","article-title":"Structural studies of receptor binding by cholera toxin mutants","volume":"6","author":"Merritt","year":"1997","journal-title":"Protein Sci."},{"key":"10.1016\/j.bpj.2016.11.008_bib45","doi-asserted-by":"crossref","first-page":"4783","DOI":"10.1091\/mbc.e03-06-0354","article-title":"Gangliosides that associate with lipid rafts mediate transport of cholera and related toxins from the plasma membrane to endoplasmic reticulum","volume":"14","author":"Fujinaga","year":"2003","journal-title":"Mol. Biol. Cell"},{"key":"10.1016\/j.bpj.2016.11.008_bib46","doi-asserted-by":"crossref","first-page":"125","DOI":"10.1016\/S0022-1759(97)00170-1","article-title":"Fusions to the cholera toxin B subunit: influence on pentamerization and GM1 binding","volume":"210","author":"Liljeqvist","year":"1997","journal-title":"J.\u00a0Immunol. Methods"},{"key":"10.1016\/j.bpj.2016.11.008_bib47","doi-asserted-by":"crossref","first-page":"1511","DOI":"10.5012\/bkcs.2002.23.11.1511","article-title":"Highly sensitive and fast protein detection with Coomassie brilliant blue in sodium dodecyl sulfate-polyacrylamide gel electrophoresis","volume":"23","author":"Kang","year":"2002","journal-title":"Bull. Korean Chem. Soc."}],"container-title":["Biophysical Journal"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S0006349516310359?httpAccept=text\/xml","content-type":"text\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S0006349516310359?httpAccept=text\/plain","content-type":"text\/plain","content-version":"vor","intended-application":"text-mining"}],"deposited":{"date-parts":[[2020,9,27]],"date-time":"2020-09-27T12:31:29Z","timestamp":1601209889000},"score":1,"resource":{"primary":{"URL":"https:\/\/linkinghub.elsevier.com\/retrieve\/pii\/S0006349516310359"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2016,12]]},"references-count":47,"journal-issue":{"issue":"12","published-print":{"date-parts":[[2016,12]]}},"alternative-id":["S0006349516310359"],"URL":"https:\/\/doi.org\/10.1016\/j.bpj.2016.11.008","relation":{},"ISSN":["0006-3495"],"issn-type":[{"value":"0006-3495","type":"print"}],"subject":[],"published":{"date-parts":[[2016,12]]},"assertion":[{"value":"Elsevier","name":"publisher","label":"This article is maintained by"},{"value":"Glycolipid Crosslinking Is Required for Cholera Toxin to Partition Into and Stabilize Ordered Domains","name":"articletitle","label":"Article Title"},{"value":"Biophysical Journal","name":"journaltitle","label":"Journal Title"},{"value":"https:\/\/doi.org\/10.1016\/j.bpj.2016.11.008","name":"articlelink","label":"CrossRef DOI link to publisher maintained version"},{"value":"article","name":"content_type","label":"Content Type"},{"value":"\u00a9 2016 Biophysical Society.","name":"copyright","label":"Copyright"}]}}