{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,3,3]],"date-time":"2026-03-03T09:34:23Z","timestamp":1772530463599,"version":"3.50.1"},"reference-count":65,"publisher":"MDPI AG","issue":"6","license":[{"start":{"date-parts":[[2025,5,24]],"date-time":"2025-05-24T00:00:00Z","timestamp":1748044800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"European Union through FEDER funds","award":["NORTE-01-0145-FEDER-000052"],"award-info":[{"award-number":["NORTE-01-0145-FEDER-000052"]}]},{"name":"European Union through FEDER funds","award":["10.54499\/UIDB\/50006\/2020"],"award-info":[{"award-number":["10.54499\/UIDB\/50006\/2020"]}]},{"name":"European Union through FEDER funds","award":["10.54499\/UIDP\/50006\/2020"],"award-info":[{"award-number":["10.54499\/UIDP\/50006\/2020"]}]},{"name":"European Union through FEDER funds","award":["Norma Transit\u00f3ria\u2014DL 57\/2016\/CP1346\/CT0006"],"award-info":[{"award-number":["Norma Transit\u00f3ria\u2014DL 57\/2016\/CP1346\/CT0006"]}]},{"name":"European Union through FEDER funds","award":["892321"],"award-info":[{"award-number":["892321"]}]},{"name":"European Union through FEDER funds","award":["892321"],"award-info":[{"award-number":["892321"]}]},{"name":"European Union through FEDER funds","award":["CEECIND\/00098\/2018"],"award-info":[{"award-number":["CEECIND\/00098\/2018"]}]},{"name":"European Union through FEDER funds","award":["REQUIMTE 2019-34"],"award-info":[{"award-number":["REQUIMTE 2019-34"]}]},{"name":"FCT\/MCTES","award":["NORTE-01-0145-FEDER-000052"],"award-info":[{"award-number":["NORTE-01-0145-FEDER-000052"]}]},{"name":"FCT\/MCTES","award":["10.54499\/UIDB\/50006\/2020"],"award-info":[{"award-number":["10.54499\/UIDB\/50006\/2020"]}]},{"name":"FCT\/MCTES","award":["10.54499\/UIDP\/50006\/2020"],"award-info":[{"award-number":["10.54499\/UIDP\/50006\/2020"]}]},{"name":"FCT\/MCTES","award":["Norma Transit\u00f3ria\u2014DL 57\/2016\/CP1346\/CT0006"],"award-info":[{"award-number":["Norma Transit\u00f3ria\u2014DL 57\/2016\/CP1346\/CT0006"]}]},{"name":"FCT\/MCTES","award":["892321"],"award-info":[{"award-number":["892321"]}]},{"name":"FCT\/MCTES","award":["892321"],"award-info":[{"award-number":["892321"]}]},{"name":"FCT\/MCTES","award":["CEECIND\/00098\/2018"],"award-info":[{"award-number":["CEECIND\/00098\/2018"]}]},{"name":"FCT\/MCTES","award":["REQUIMTE 2019-34"],"award-info":[{"award-number":["REQUIMTE 2019-34"]}]},{"name":"FCT\u2014Funda\u00e7\u00e3o para a Ci\u00eancia e a Tecnologia","award":["NORTE-01-0145-FEDER-000052"],"award-info":[{"award-number":["NORTE-01-0145-FEDER-000052"]}]},{"name":"FCT\u2014Funda\u00e7\u00e3o para a Ci\u00eancia e a Tecnologia","award":["10.54499\/UIDB\/50006\/2020"],"award-info":[{"award-number":["10.54499\/UIDB\/50006\/2020"]}]},{"name":"FCT\u2014Funda\u00e7\u00e3o para a Ci\u00eancia e a Tecnologia","award":["10.54499\/UIDP\/50006\/2020"],"award-info":[{"award-number":["10.54499\/UIDP\/50006\/2020"]}]},{"name":"FCT\u2014Funda\u00e7\u00e3o para a Ci\u00eancia e a Tecnologia","award":["Norma Transit\u00f3ria\u2014DL 57\/2016\/CP1346\/CT0006"],"award-info":[{"award-number":["Norma Transit\u00f3ria\u2014DL 57\/2016\/CP1346\/CT0006"]}]},{"name":"FCT\u2014Funda\u00e7\u00e3o para a Ci\u00eancia e a Tecnologia","award":["892321"],"award-info":[{"award-number":["892321"]}]},{"name":"FCT\u2014Funda\u00e7\u00e3o para a Ci\u00eancia e a Tecnologia","award":["892321"],"award-info":[{"award-number":["892321"]}]},{"name":"FCT\u2014Funda\u00e7\u00e3o para a Ci\u00eancia e a Tecnologia","award":["CEECIND\/00098\/2018"],"award-info":[{"award-number":["CEECIND\/00098\/2018"]}]},{"name":"FCT\u2014Funda\u00e7\u00e3o para a Ci\u00eancia e a Tecnologia","award":["REQUIMTE 2019-34"],"award-info":[{"award-number":["REQUIMTE 2019-34"]}]},{"name":"European Union\u2019s Horizon 2020 Research and Innovation Programme","award":["NORTE-01-0145-FEDER-000052"],"award-info":[{"award-number":["NORTE-01-0145-FEDER-000052"]}]},{"name":"European Union\u2019s Horizon 2020 Research and Innovation Programme","award":["10.54499\/UIDB\/50006\/2020"],"award-info":[{"award-number":["10.54499\/UIDB\/50006\/2020"]}]},{"name":"European Union\u2019s Horizon 2020 Research and Innovation Programme","award":["10.54499\/UIDP\/50006\/2020"],"award-info":[{"award-number":["10.54499\/UIDP\/50006\/2020"]}]},{"name":"European Union\u2019s Horizon 2020 Research and Innovation Programme","award":["Norma Transit\u00f3ria\u2014DL 57\/2016\/CP1346\/CT0006"],"award-info":[{"award-number":["Norma Transit\u00f3ria\u2014DL 57\/2016\/CP1346\/CT0006"]}]},{"name":"European Union\u2019s Horizon 2020 Research and Innovation Programme","award":["892321"],"award-info":[{"award-number":["892321"]}]},{"name":"European Union\u2019s Horizon 2020 Research and Innovation Programme","award":["892321"],"award-info":[{"award-number":["892321"]}]},{"name":"European Union\u2019s Horizon 2020 Research and Innovation Programme","award":["CEECIND\/00098\/2018"],"award-info":[{"award-number":["CEECIND\/00098\/2018"]}]},{"name":"European Union\u2019s Horizon 2020 Research and Innovation Programme","award":["REQUIMTE 2019-34"],"award-info":[{"award-number":["REQUIMTE 2019-34"]}]},{"name":"FCT Scientific Employment Stimulus","award":["NORTE-01-0145-FEDER-000052"],"award-info":[{"award-number":["NORTE-01-0145-FEDER-000052"]}]},{"name":"FCT Scientific Employment Stimulus","award":["10.54499\/UIDB\/50006\/2020"],"award-info":[{"award-number":["10.54499\/UIDB\/50006\/2020"]}]},{"name":"FCT Scientific Employment Stimulus","award":["10.54499\/UIDP\/50006\/2020"],"award-info":[{"award-number":["10.54499\/UIDP\/50006\/2020"]}]},{"name":"FCT Scientific Employment Stimulus","award":["Norma Transit\u00f3ria\u2014DL 57\/2016\/CP1346\/CT0006"],"award-info":[{"award-number":["Norma Transit\u00f3ria\u2014DL 57\/2016\/CP1346\/CT0006"]}]},{"name":"FCT Scientific Employment Stimulus","award":["892321"],"award-info":[{"award-number":["892321"]}]},{"name":"FCT Scientific Employment Stimulus","award":["892321"],"award-info":[{"award-number":["892321"]}]},{"name":"FCT Scientific Employment Stimulus","award":["CEECIND\/00098\/2018"],"award-info":[{"award-number":["CEECIND\/00098\/2018"]}]},{"name":"FCT Scientific Employment Stimulus","award":["REQUIMTE 2019-34"],"award-info":[{"award-number":["REQUIMTE 2019-34"]}]},{"name":"FCT Scientific Employment Stimulus","award":["NORTE-01-0145-FEDER-000052"],"award-info":[{"award-number":["NORTE-01-0145-FEDER-000052"]}]},{"name":"FCT Scientific Employment Stimulus","award":["10.54499\/UIDB\/50006\/2020"],"award-info":[{"award-number":["10.54499\/UIDB\/50006\/2020"]}]},{"name":"FCT Scientific Employment Stimulus","award":["10.54499\/UIDP\/50006\/2020"],"award-info":[{"award-number":["10.54499\/UIDP\/50006\/2020"]}]},{"name":"FCT Scientific Employment Stimulus","award":["Norma Transit\u00f3ria\u2014DL 57\/2016\/CP1346\/CT0006"],"award-info":[{"award-number":["Norma Transit\u00f3ria\u2014DL 57\/2016\/CP1346\/CT0006"]}]},{"name":"FCT Scientific Employment Stimulus","award":["892321"],"award-info":[{"award-number":["892321"]}]},{"name":"FCT Scientific Employment Stimulus","award":["892321"],"award-info":[{"award-number":["892321"]}]},{"name":"FCT Scientific Employment Stimulus","award":["CEECIND\/00098\/2018"],"award-info":[{"award-number":["CEECIND\/00098\/2018"]}]},{"name":"FCT Scientific Employment Stimulus","award":["REQUIMTE 2019-34"],"award-info":[{"award-number":["REQUIMTE 2019-34"]}]},{"name":"FCT and LAQV\/REQUIMTE for a post-doc fellowship","award":["NORTE-01-0145-FEDER-000052"],"award-info":[{"award-number":["NORTE-01-0145-FEDER-000052"]}]},{"name":"FCT and LAQV\/REQUIMTE for a post-doc fellowship","award":["10.54499\/UIDB\/50006\/2020"],"award-info":[{"award-number":["10.54499\/UIDB\/50006\/2020"]}]},{"name":"FCT and LAQV\/REQUIMTE for a post-doc fellowship","award":["10.54499\/UIDP\/50006\/2020"],"award-info":[{"award-number":["10.54499\/UIDP\/50006\/2020"]}]},{"name":"FCT and LAQV\/REQUIMTE for a post-doc fellowship","award":["Norma Transit\u00f3ria\u2014DL 57\/2016\/CP1346\/CT0006"],"award-info":[{"award-number":["Norma Transit\u00f3ria\u2014DL 57\/2016\/CP1346\/CT0006"]}]},{"name":"FCT and LAQV\/REQUIMTE for a post-doc fellowship","award":["892321"],"award-info":[{"award-number":["892321"]}]},{"name":"FCT and LAQV\/REQUIMTE for a post-doc fellowship","award":["892321"],"award-info":[{"award-number":["892321"]}]},{"name":"FCT and LAQV\/REQUIMTE for a post-doc fellowship","award":["CEECIND\/00098\/2018"],"award-info":[{"award-number":["CEECIND\/00098\/2018"]}]},{"name":"FCT and LAQV\/REQUIMTE for a post-doc fellowship","award":["REQUIMTE 2019-34"],"award-info":[{"award-number":["REQUIMTE 2019-34"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Membranes"],"abstract":"<jats:p>As a surface-located molecule in biological membranes, cholesterol sulphate (CholS) plays a major role in membrane-driven cell\u2013cell processes and events including platelet-cell adhesion, T-cell receptor signalling, sperm\u2013egg interaction, membrane fusion, and skin differentiation. Despite this, little is known about the biophysical implications of CholS at the membrane in cells and organelles. In this study, we investigate the effect of increasing the content of CholS on the biophysical properties in cholesterol-poor and cholesterol-rich biomimetic models. Data obtained show that increasing amounts of CholS result in a slight increase in anisotropy, evidence for decreased membrane fluidity at higher CholS content (10 mol%) in cholesterol-poor systems but only negligible in rigidified epithelial-like cholesterol-rich systems. On the other hand, incorporation of CholS has an overall increasing ordering effect on membrane organisation and on-surface potential that is influenced by the lipid composition and cholesterol content. Though further research is needed to gain better insights on the (patho)physiological levels of CholS in cells and organelles, our findings are discussed in the context of diet\u2013microbiota\u2013host interactions in membrane-driven events in inflammatory-related disorders.<\/jats:p>","DOI":"10.3390\/membranes15060159","type":"journal-article","created":{"date-parts":[[2025,5,25]],"date-time":"2025-05-25T20:16:22Z","timestamp":1748204182000},"page":"159","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":2,"title":["Cholesterol Sulfate in Biological Membranes: A Biophysical Study in Cholesterol-Poor and Cholesterol-Rich Biomimetic Models"],"prefix":"10.3390","volume":"15","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-4537-9902","authenticated-orcid":false,"given":"Ana","family":"Reis","sequence":"first","affiliation":[{"name":"Rede de Quimica e Tecnologia\u2014Laboratorio Associado de Quimica Verde, Departamento de Qu\u00edmica e Bioqu\u00edmica, Faculdade de Ci\u00eancias, Universidade do Porto, Rua do Campo Alegre, 4169-007 Porto, Portugal"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-1765-4724","authenticated-orcid":false,"given":"Maria Jo\u00e3o","family":"Sarmento","sequence":"additional","affiliation":[{"name":"Instituto de Medicina Molecular, Faculdade de Medicina, Universidade de Lisboa, 1649-028 Lisboa, Portugal"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9907-9518","authenticated-orcid":false,"given":"Mariana","family":"Ferreira","sequence":"additional","affiliation":[{"name":"Rede de Quimica e Tecnologia\u2014Laboratorio Associado de Quimica Verde, Departamento de Qu\u00edmica e Bioqu\u00edmica, Faculdade de Ci\u00eancias, Universidade do Porto, Rua do Campo Alegre, 4169-007 Porto, Portugal"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4747-6853","authenticated-orcid":false,"given":"Paula","family":"Gameiro","sequence":"additional","affiliation":[{"name":"Rede de Quimica e Tecnologia\u2014Laboratorio Associado de Quimica Verde, Departamento de Qu\u00edmica e Bioqu\u00edmica, Faculdade de Ci\u00eancias, Universidade do Porto, Rua do Campo Alegre, 4169-007 Porto, Portugal"}]},{"given":"Victor","family":"de Freitas","sequence":"additional","affiliation":[{"name":"Rede de Quimica e Tecnologia\u2014Laboratorio Associado de Quimica Verde, Departamento de Qu\u00edmica e Bioqu\u00edmica, Faculdade de Ci\u00eancias, Universidade do Porto, Rua do Campo Alegre, 4169-007 Porto, Portugal"}]}],"member":"1968","published-online":{"date-parts":[[2025,5,24]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"1268","DOI":"10.1194\/jlr.R300005-JLR200","article-title":"Cholesterol sulfate in human physiology: What\u2019s it all about?","volume":"44","author":"Strott","year":"2003","journal-title":"J. Lipid Res."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"401","DOI":"10.1042\/EBC20230090","article-title":"Oxysterol sulfates in fluids, cells and tissues: How much do we know about their clinical significance, biological relevance and biophysical implications?","volume":"68","author":"Reis","year":"2024","journal-title":"Essays Biochem."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"53","DOI":"10.1016\/j.chemphyslip.2019.03.009","article-title":"Sulfate-based lipids: Analysis of healthy human fluids and cell extracts","volume":"221","author":"Dias","year":"2019","journal-title":"Chem. Phys. Lipids"},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"521","DOI":"10.1124\/pharmrev.122.000679","article-title":"Sulfotransferase 2B1b, Sterol Sulfonation, and Disease","volume":"75","author":"Cook","year":"2023","journal-title":"Pharmacol. Rev."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"T271","DOI":"10.1530\/JME-18-0086","article-title":"Insights into steroid sulfation and desulfation pathways","volume":"61","author":"Foster","year":"2018","journal-title":"J. Mol. Endocrinol."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"1390","DOI":"10.1038\/s41564-022-01195-9","article-title":"Characterization of interactions of dietary cholesterol with the murine and human gut microbiome","volume":"7","author":"Le","year":"2022","journal-title":"Nat. Microbiol."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"1404","DOI":"10.1172\/JCI110391","article-title":"Stratum corneum lipids in disorders of cornification. Increased cholesterol sulfate content of stratum corneum in recessive X-linked ichthyosis","volume":"68","author":"Williams","year":"1981","journal-title":"J. Clin. Investig."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"1414","DOI":"10.1172\/JCI111552","article-title":"Stratum Corneum Lipids in Disorders of Cornification","volume":"74","author":"Elias","year":"1984","journal-title":"JCI"},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"1085","DOI":"10.1016\/0003-9969(95)00091-7","article-title":"Regional variation in content, composition and organization of porcine epithelial barrier lipids revealed by thin-layer chromatography and transmission electron microscopy","volume":"40","author":"Law","year":"1995","journal-title":"Archs. Oral Biol."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"833","DOI":"10.1007\/BF02536238","article-title":"Higher levels of plasma cholesterol sulfate in patients with liver cirrhosis and hypercholesterolemia","volume":"28","author":"Tamasawa","year":"1993","journal-title":"Lipids"},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"3430","DOI":"10.1021\/ac9029482","article-title":"Cholesterol Sulfate Imaging in Human Prostate Cancer Tissue by Desorption Electrospray Ionization Mass Spectrometry","volume":"82","author":"Eberlin","year":"2010","journal-title":"Anal. Chem."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"353","DOI":"10.1016\/j.bbalip.2013.11.009","article-title":"Role of cholesterol sulfate in epidermal structure and function: Lessons from X-linked ichthyosis","volume":"1841","author":"Elias","year":"2014","journal-title":"Biochim. Biophys. Acta Mol. Cell Biol. Lipids"},{"key":"ref_13","doi-asserted-by":"crossref","unstructured":"Lopalco, P., Vitale, R., Cho, Y.S., Totaro, P., Corcelli, A., and Lobasso, S. (2019). Alteration of Cholesterol Sulfate\/Seminolipid Ratio in Semen Lipid Profile of Men With Oligoasthenozoospermia. Front. Physiol., 10.","DOI":"10.3389\/fphys.2019.01344"},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"4428","DOI":"10.1038\/s41467-022-32158-7","article-title":"Cholesterol sulfate alleviates ulcerative colitis by promoting cholesterol biosynthesis in colonic epithelial cells","volume":"13","author":"Xu","year":"2022","journal-title":"Nat. Commun."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"299","DOI":"10.1016\/j.bbamem.2006.08.022","article-title":"Visualizing the localization of sulfoglycolipids in lipid raft domains in model membranes and sperm membrane extracts","volume":"1768","author":"Weerachatyanukul","year":"2007","journal-title":"Biochim. Biophys. Acta Biomembr."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"844","DOI":"10.1038\/ni.3462","article-title":"Inhibition of T cell receptor signalling by cholesterol sulfate, a naturally occurring derivative of membrane cholesterol","volume":"17","author":"Wang","year":"2016","journal-title":"Nat. Immunol."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"2032","DOI":"10.1161\/01.CIR.103.16.2032","article-title":"Cholesterol sulfate: A new adhesive molecule for platelets","volume":"103","author":"Merten","year":"2001","journal-title":"Circulation"},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"e00299-22","DOI":"10.1128\/msphere.00299-22","article-title":"Pleiotropic Roles of Cholesteryl Sulfate during Entamoeba Encystation: Involvement in Cell Rounding and Development of Membrane Impermeability","volume":"7","author":"Tsugawa","year":"2022","journal-title":"mSphere"},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"16683","DOI":"10.1038\/s41598-018-34286-x","article-title":"Imaging the distribution of skin lipids and topically applied compounds in human skin using mass spectrometry","volume":"8","author":"Skedung","year":"2018","journal-title":"Sci. Rep."},{"key":"ref_20","doi-asserted-by":"crossref","unstructured":"Morino, K., Kunimura, K., Sugiura, Y., Izumi, Y., Matsubara, K., Akiyoshi, S., Maeda, R., Hirotani, K., Sakata, D., and Mizuno, S. (2023). Cholesterol sulfate limits neutrophil recruitment and gut inflammation during mucosal injury. Front. Immunol., 14.","DOI":"10.3389\/fimmu.2023.1131146"},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"1672","DOI":"10.1016\/S0006-3495(00)76719-1","article-title":"Molecular dynamics simulation of dipalmitoylphosphatidylcholine membrane with cholesterol sulfate","volume":"78","author":"Smondyrev","year":"2000","journal-title":"Biophys. J."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"100177","DOI":"10.1016\/j.jlr.2022.100177","article-title":"Cholesterol sulfate fluidizes the sterol fraction of the stratum corneum lipid phase and increases its permeability","volume":"63","author":"Fandrei","year":"2022","journal-title":"J. Lipid Res."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"73","DOI":"10.1016\/S0009-9120(97)00166-5","article-title":"Morphological changes of human erythrocytes induced by cholesterol sulphate","volume":"31","author":"Przybylska","year":"1998","journal-title":"Clin. Biochem."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"23","DOI":"10.1016\/S0009-3084(98)00065-6","article-title":"Cholesterol versus cholesterol sulfate: Effects on properties of phospholipid bilayers containing docosahexaenoic acid","volume":"95","author":"Schofield","year":"1998","journal-title":"Chem. Phys. Lipids"},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"1380","DOI":"10.1016\/S0006-3495(96)79696-0","article-title":"Comparative effects of cholesterol and cholesterol sulfate on hydration and ordering of dimyristoylphosphatidylcholine membranes","volume":"70","author":"Faure","year":"1996","journal-title":"Biophys. J."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"5","DOI":"10.1242\/jcs.071233","article-title":"Lipid map of the mammalian cell","volume":"124","year":"2011","journal-title":"J. Cell Sci."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"237","DOI":"10.1007\/s00018-023-04889-3","article-title":"The expanding organelle lipidomes: Current knowledge and challenges","volume":"80","author":"Sarmento","year":"2023","journal-title":"Cell. Mol. Life Sci."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"55","DOI":"10.1016\/0005-2736(85)90521-8","article-title":"Production of large unilamellar vesicles by a rapid extrusion procedure. Characterization of size distribution, trapped volume and ability to maintain a membrane potential","volume":"812","author":"Hope","year":"1985","journal-title":"Biochim. Biophys. Acta Biomembr."},{"key":"ref_29","doi-asserted-by":"crossref","unstructured":"Lakowicz, J. (2006). Principles of Fluorescence Spectroscopy, Springer. [3rd ed.].","DOI":"10.1007\/978-0-387-46312-4"},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"2413","DOI":"10.1016\/S0006-3495(97)78887-8","article-title":"Two-photon fluorescence microscopy of Laurdan generalized polarization domains in model and natural membranes","volume":"72","author":"Tiziana","year":"1997","journal-title":"Biophys. J."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"11","DOI":"10.1016\/j.chemphyslip.2005.09.003","article-title":"Determination of phase transition temperatures of lipids by light scattering","volume":"139","author":"Michel","year":"2006","journal-title":"Chem. Phys. Lipids"},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"2406","DOI":"10.1016\/S0006-3495(03)74664-5","article-title":"Sphingomyelin\/phosphatidylcholine\/cholesterol phase diagram: Boundaries and composition of lipid rafts","volume":"85","author":"Fedorov","year":"2003","journal-title":"Biophys. J."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"665","DOI":"10.1016\/j.bbalip.2008.09.002","article-title":"Phase diagrams of lipid mixtures relevant to the study of membrane rafts","volume":"1781","author":"Alonso","year":"2008","journal-title":"Biochim. Biophys. Acta"},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"172","DOI":"10.1016\/j.bbamcr.2005.06.010","article-title":"Seeing spots: Complex phase behavior in simple membranes","volume":"1746","author":"Veatch","year":"2005","journal-title":"Biochim. Biophys. Acta Mol. Cell Res."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"a004630","DOI":"10.1101\/cshperspect.a004630","article-title":"Phase separation in lipid membranes","volume":"3","author":"Heberle","year":"2011","journal-title":"Cold Spring Harb. Perspect. Biol."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"16645","DOI":"10.1073\/pnas.0908987106","article-title":"Order of lipid phases in model and plasma membranes","volume":"106","author":"Kaiser","year":"2009","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"590","DOI":"10.1016\/j.plipres.2013.07.002","article-title":"Lipids of mitochondria","volume":"52","author":"Horvath","year":"2013","journal-title":"Prog. Lipid Res."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"264","DOI":"10.1080\/10409230903193307","article-title":"The ins and outs of phospholipid asymmetry in the plasma membrane: Roles in health and disease","volume":"44","author":"Fadeel","year":"2009","journal-title":"Crit. Rev. Biochem. Mol. Biol."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"333","DOI":"10.1016\/j.tibs.2024.01.007","article-title":"Exploring membrane asymmetry and its effects on membrane proteins","volume":"49","author":"Pabst","year":"2024","journal-title":"Trends Biochem. Sci."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"825","DOI":"10.1038\/ki.1992.357","article-title":"Membrane fluidity and transport properties in epithelia","volume":"42","author":"Friedlander","year":"1992","journal-title":"Kidney Int."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"3230","DOI":"10.1016\/j.febslet.2008.08.016","article-title":"Cholesterol displacement by ceramide in sphingomyelin-containing liquid-ordered domains, and generation of gel regions in giant lipidic vesicles","volume":"582","author":"Sot","year":"2008","journal-title":"FEBS Lett."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"1357","DOI":"10.1007\/s00216-010-4028-6","article-title":"Cardiolipin, a key component to mimic the E. coli bacterial membrane in model systems revealed by dynamic light scattering and steady-state fluorescence anisotropy","volume":"398","author":"Lopes","year":"2010","journal-title":"Anal. Bioanal. Chem."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"1534","DOI":"10.3390\/biom12101534","article-title":"Modelling Hyperglycaemia in an Epithelial Membrane Model: Biophysical Characterisation","volume":"12","author":"Reis","year":"2022","journal-title":"Biomolecules"},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"1362","DOI":"10.1016\/j.bbamem.2017.04.026","article-title":"Interaction of procyanidin B3 with membrane lipids\u2013Fluorescence, DSC and FTIR studies","volume":"1859","author":"Jurkiewicz","year":"2017","journal-title":"Biochim. Biophys. Acta Biomembr."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"40","DOI":"10.1016\/j.fct.2018.01.017","article-title":"Cholesterol modulates the liposome membrane fluidity and permeability for a hydrophilic molecule","volume":"113","author":"Kaddah","year":"2018","journal-title":"Food Chem. Toxicol."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"12851","DOI":"10.1021\/la302620t","article-title":"Influence of cholesterol on the phase transition of lipid bilayers: A temperature-controlled force spectroscopy study","volume":"28","author":"Giannotti","year":"2012","journal-title":"Langmuir"},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"14815","DOI":"10.1038\/s41598-018-33107-5","article-title":"Determination of the Main Phase Transition Temperature of Phospholipids by Nanoplasmonic Sensing","volume":"8","author":"Chen","year":"2018","journal-title":"Sci. Rep."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"97","DOI":"10.1016\/j.bbamem.2008.08.022","article-title":"Ordering effects of cholesterol and its analogues","volume":"1788","author":"Vattulainen","year":"2009","journal-title":"Biochim. Biophys. Acta Biomembr."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"2647","DOI":"10.1016\/j.bbamem.2016.07.013","article-title":"Diphenylhexatriene membrane probes DPH and TMA-DPH: A comparative molecular dynamics simulation study","volume":"1858","author":"Robalo","year":"2016","journal-title":"Biochim. Biophys. Acta Biomembr."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"26","DOI":"10.1016\/j.biochi.2011.06.027","article-title":"Lipid hydration and mobility: An interplay between fluorescence solvent relaxation experiments and molecular dynamics simulations","volume":"94","author":"Jurkiewicz","year":"2012","journal-title":"Biochimie"},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"112404","DOI":"10.1016\/j.jphotobiol.2022.112404","article-title":"Laurdan in live cell imaging: Effect of acquisition settings, cell culture conditions and data analysis on generalized polarization measurements","volume":"228","author":"Pokorna","year":"2022","journal-title":"J. Photochem. Photobiol. B Biol."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"9417","DOI":"10.1039\/C6SM01777K","article-title":"Lipid bilayer thickness determines cholesterols location in model membranes","volume":"12","author":"Marquardt","year":"2016","journal-title":"Soft Matter"},{"key":"ref_53","doi-asserted-by":"crossref","unstructured":"Ma, Y., Poole, K., Goyette, J., and Gaus, K. (2017). Introducing membrane charge and membrane potential to T cell signaling. Front. Immunol., 8.","DOI":"10.3389\/fimmu.2017.01513"},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"579","DOI":"10.1038\/mi.2013.76","article-title":"Pro-inflammatory effects of matrix metalloproteinase 7 in acute inflammation","volume":"7","author":"Vandenbroucke","year":"2014","journal-title":"Mucosal Immunol."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"2099","DOI":"10.1016\/j.str.2015.08.013","article-title":"Charge-Triggered Membrane Insertion of Matrix Metalloproteinase-7, Supporter of Innate Immunity and Tumors","volume":"23","author":"Prior","year":"2015","journal-title":"Structure"},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"529","DOI":"10.1016\/0305-0491(86)90291-9","article-title":"Lipids of epidermis and keratinized and non-keratinized oral epithelia","volume":"83","author":"Wertz","year":"1986","journal-title":"Comp. Biochem. Physiol. Part B Biochem."},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"138","DOI":"10.1016\/0005-2736(84)90284-0","article-title":"Lipid composition and fluidity of rat enterocyte basolateral membranes regional differences","volume":"774","author":"Brasitus","year":"1984","journal-title":"Biochim. Biophys. Acta Biomembr."},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"403","DOI":"10.1007\/s10337-014-2787-5","article-title":"Lipid Analysis of Airway Epithelial Cells for Studying Respiratory Diseases","volume":"78","author":"Zehethofer","year":"2014","journal-title":"Chromatographia"},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"213","DOI":"10.1083\/jcb.201108175","article-title":"Quantitative analysis of the lipidomes of the influenza virus envelope and MDCK cell apical membrane","volume":"196","author":"Gerl","year":"2012","journal-title":"J. Cell Biol."},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"100084","DOI":"10.1016\/j.jlr.2021.100084","article-title":"Evaluation of the available cholesterol concentration in the inner leaflet of the plasma membrane of mammalian cells","volume":"62","author":"Buwaneka","year":"2021","journal-title":"J. Lipid Res."},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"14554","DOI":"10.1021\/ja507832e","article-title":"Lipid organization of the plasma membrane","volume":"136","author":"Melo","year":"2014","journal-title":"J. Am. Chem. Soc."},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"644","DOI":"10.1038\/s41589-020-0529-6","article-title":"Plasma membranes are asymmetric in lipid unsaturation, packing and protein shape","volume":"16","author":"Lorent","year":"2020","journal-title":"Nat. Chem. Biol."},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"1812","DOI":"10.1136\/gutjnl-2015-309957","article-title":"High-level adherence to a Mediterranean diet beneficially impacts the gut microbiota and associated metabolome","volume":"65","author":"Pellegrini","year":"2016","journal-title":"Gut"},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"4137","DOI":"10.1016\/j.cell.2021.06.019","article-title":"Gut-microbiota-targeted diets modulate human immune status","volume":"184","author":"Wastyk","year":"2021","journal-title":"Cell"},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"605","DOI":"10.1093\/ajcn\/nqab077","article-title":"Associations of healthy food choices with gut microbiota profiles","volume":"114","author":"Koponen","year":"2021","journal-title":"Am. J. Clin. Nutr."}],"container-title":["Membranes"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2077-0375\/15\/6\/159\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,9]],"date-time":"2025-10-09T17:39:44Z","timestamp":1760031584000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2077-0375\/15\/6\/159"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,5,24]]},"references-count":65,"journal-issue":{"issue":"6","published-online":{"date-parts":[[2025,6]]}},"alternative-id":["membranes15060159"],"URL":"https:\/\/doi.org\/10.3390\/membranes15060159","relation":{},"ISSN":["2077-0375"],"issn-type":[{"value":"2077-0375","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,5,24]]}}}