{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"institution":[{"name":"bioRxiv"}],"indexed":{"date-parts":[[2026,7,30]],"date-time":"2026-07-30T11:19:18Z","timestamp":1785410358178,"version":"3.56.0"},"posted":{"date-parts":[[2025,6,25]]},"group-title":"Molecular Biology","reference-count":115,"publisher":"openRxiv","license":[{"start":{"date-parts":[[2025,6,25]],"date-time":"2025-06-25T00:00:00Z","timestamp":1750809600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/www.biorxiv.org\/about\/FAQ#license"}],"funder":[{"award":["R01GM136975"],"award-info":[{"award-number":["R01GM136975"]}],"id":[{"id":"https:\/\/ror.org\/04q48ey07","id-type":"ROR","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"accepted":{"date-parts":[[2025,6,25]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                <jats:p>Styrene-maleic acid (SMA) copolymers are powerful tools for the detergent-free solubilization of biological membranes. Yet the influence of specific lipids on SMA activity remains an open question. Here, we examined the effects of the mitochondriaspecific phospholipid cardiolipin on SMA-mediated membrane solubilization and their ability to form SMA-bound nanodiscs. To this end, we prepared a series of model membranes with cardiolipin and other test lipids with comparable surface charge and lateral packing characteristics. Using multiple independent experimental approaches, we found that cardiolipin inhibited SMA solubilization. Our results indicate that this effect was not attributable to headgroup charge effects, but rather to cardiolipininduced increase in lateral packing pressure at the interfacial region. Reduction of this lateral packing pressure using bilayer-active alcohols partially restored SMA solubilization. Our results highlight the importance of lipid geometry and packing in SMA nanodisc formation and could help guide the design of copolymers tailored to specific membranes.<\/jats:p>","DOI":"10.1101\/2025.06.23.659572","type":"posted-content","created":{"date-parts":[[2025,6,26]],"date-time":"2025-06-26T01:20:14Z","timestamp":1750900814000},"source":"Crossref","is-referenced-by-count":0,"title":["Lateral lipid packing governs bilayer solubilization by styrene-maleic acid copolymers: a case study with cardiolipin-containing membranes"],"prefix":"10.64898","author":[{"given":"Joseph C.","family":"Iovine","sequence":"first","affiliation":[{"name":"Department of Molecular and Cell Biology, University of Connecticut, Storrs, CT, 06269 USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Nathan N.","family":"Alder","sequence":"additional","affiliation":[{"name":"Department of Molecular and Cell Biology, University of Connecticut, Storrs, CT, 06269 USA"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"54368","reference":[{"key":"2026041210101056000_2025.06.23.659572v1.1","doi-asserted-by":"publisher","DOI":"10.3390\/nano12030361"},{"key":"2026041210101056000_2025.06.23.659572v1.2","doi-asserted-by":"publisher","DOI":"10.1021\/acs.biomac.1c01166"},{"key":"2026041210101056000_2025.06.23.659572v1.3","doi-asserted-by":"publisher","DOI":"10.1021\/ja810046q"},{"key":"2026041210101056000_2025.06.23.659572v1.4","doi-asserted-by":"publisher","DOI":"10.1002\/anie.201201355"},{"key":"2026041210101056000_2025.06.23.659572v1.5","doi-asserted-by":"publisher","DOI":"10.1007\/s00249-015-1093-y"},{"key":"2026041210101056000_2025.06.23.659572v1.6","doi-asserted-by":"publisher","DOI":"10.1016\/j.chemphyslip.2019.03.008"},{"key":"2026041210101056000_2025.06.23.659572v1.7","doi-asserted-by":"publisher","DOI":"10.1007\/s12274-014-0560-6"},{"key":"2026041210101056000_2025.06.23.659572v1.8","doi-asserted-by":"publisher","DOI":"10.1039\/C6NR02089E"},{"key":"2026041210101056000_2025.06.23.659572v1.9","doi-asserted-by":"publisher","DOI":"10.1021\/acs.langmuir.1c00304"},{"key":"2026041210101056000_2025.06.23.659572v1.10","doi-asserted-by":"publisher","DOI":"10.1186\/1472-6750-13-41"},{"key":"2026041210101056000_2025.06.23.659572v1.11","doi-asserted-by":"publisher","DOI":"10.1016\/j.bbamem.2017.07.011"},{"key":"2026041210101056000_2025.06.23.659572v1.12","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-019-09425-1"},{"key":"2026041210101056000_2025.06.23.659572v1.13","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.1416205112"},{"key":"2026041210101056000_2025.06.23.659572v1.14","doi-asserted-by":"publisher","DOI":"10.1002\/anie.201406412"},{"key":"2026041210101056000_2025.06.23.659572v1.15","doi-asserted-by":"publisher","DOI":"10.1042\/BJ20131477"},{"key":"2026041210101056000_2025.06.23.659572v1.16","doi-asserted-by":"publisher","DOI":"10.1016\/j.bbamem.2015.06.024"},{"key":"2026041210101056000_2025.06.23.659572v1.17","doi-asserted-by":"publisher","DOI":"10.1080\/02648725.2014.921502"},{"key":"2026041210101056000_2025.06.23.659572v1.18","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0206692"},{"key":"2026041210101056000_2025.06.23.659572v1.19","doi-asserted-by":"publisher","DOI":"10.1016\/j.bbamem.2017.10.019"},{"key":"2026041210101056000_2025.06.23.659572v1.20","doi-asserted-by":"publisher","DOI":"10.1038\/s41586-018-0061-y"},{"key":"2026041210101056000_2025.06.23.659572v1.21","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.1712725115"},{"key":"2026041210101056000_2025.06.23.659572v1.22","doi-asserted-by":"publisher","DOI":"10.1016\/j.bbamem.2017.10.005"},{"key":"2026041210101056000_2025.06.23.659572v1.23","doi-asserted-by":"publisher","DOI":"10.1021\/jacs.8b08441"},{"key":"2026041210101056000_2025.06.23.659572v1.24","doi-asserted-by":"publisher","DOI":"10.1177\/2472555219857691"},{"key":"2026041210101056000_2025.06.23.659572v1.25","doi-asserted-by":"publisher","DOI":"10.2147\/IJN.S187089"},{"key":"2026041210101056000_2025.06.23.659572v1.26","doi-asserted-by":"publisher","DOI":"10.1016\/j.toxrep.2019.04.004"},{"key":"2026041210101056000_2025.06.23.659572v1.27","doi-asserted-by":"publisher","DOI":"10.1016\/j.ijpharm.2019.118719"},{"key":"2026041210101056000_2025.06.23.659572v1.28","doi-asserted-by":"publisher","DOI":"10.2174\/1574892814666190919163731"},{"key":"2026041210101056000_2025.06.23.659572v1.29","doi-asserted-by":"publisher","DOI":"10.1016\/j.bpj.2016.09.025"},{"key":"2026041210101056000_2025.06.23.659572v1.30","doi-asserted-by":"publisher","DOI":"10.1016\/j.chemphyslip.2017.08.010"},{"key":"2026041210101056000_2025.06.23.659572v1.31","doi-asserted-by":"publisher","DOI":"10.1016\/j.bbamem.2019.183125"},{"key":"2026041210101056000_2025.06.23.659572v1.32","doi-asserted-by":"publisher","DOI":"10.1016\/j.bpj.2014.11.3464"},{"key":"2026041210101056000_2025.06.23.659572v1.33","doi-asserted-by":"publisher","DOI":"10.1039\/C5NR06353A"},{"key":"2026041210101056000_2025.06.23.659572v1.34","doi-asserted-by":"publisher","DOI":"10.1007\/s00249-016-1181-7"},{"key":"2026041210101056000_2025.06.23.659572v1.35","doi-asserted-by":"publisher","DOI":"10.1016\/j.bbamem.2020.183209"},{"key":"2026041210101056000_2025.06.23.659572v1.36","doi-asserted-by":"publisher","DOI":"10.1016\/j.bbamem.2017.08.010"},{"key":"2026041210101056000_2025.06.23.659572v1.37","doi-asserted-by":"publisher","DOI":"10.1038\/s41598-019-51896-1"},{"key":"2026041210101056000_2025.06.23.659572v1.38","doi-asserted-by":"publisher","DOI":"10.1016\/j.bbamem.2021.183641"},{"key":"2026041210101056000_2025.06.23.659572v1.39","doi-asserted-by":"publisher","DOI":"10.1038\/s41598-019-54999-x"},{"key":"2026041210101056000_2025.06.23.659572v1.40","doi-asserted-by":"publisher","DOI":"10.1083\/jcb.202003131"},{"key":"2026041210101056000_2025.06.23.659572v1.41","doi-asserted-by":"publisher","DOI":"10.1016\/0304-4157(85)90002-4"},{"key":"2026041210101056000_2025.06.23.659572v1.42","doi-asserted-by":"publisher","DOI":"10.1038\/nrm2330"},{"key":"2026041210101056000_2025.06.23.659572v1.43","doi-asserted-by":"publisher","DOI":"10.1002\/1873-3468.12887"},{"key":"2026041210101056000_2025.06.23.659572v1.44","doi-asserted-by":"publisher","DOI":"10.1039\/c8sm02133c"},{"key":"2026041210101056000_2025.06.23.659572v1.45","doi-asserted-by":"publisher","DOI":"10.1016\/j.tibs.2021.06.003"},{"key":"2026041210101056000_2025.06.23.659572v1.46","doi-asserted-by":"publisher","DOI":"10.1038\/s42003-023-05657-6"},{"key":"2026041210101056000_2025.06.23.659572v1.47","doi-asserted-by":"publisher","DOI":"10.1369\/0022155415574818"},{"key":"2026041210101056000_2025.06.23.659572v1.48","doi-asserted-by":"publisher","DOI":"10.1126\/sciadv.1700532"},{"key":"2026041210101056000_2025.06.23.659572v1.49","doi-asserted-by":"publisher","DOI":"10.1016\/j.plipres.2022.101195"},{"key":"2026041210101056000_2025.06.23.659572v1.50","doi-asserted-by":"publisher","DOI":"10.1038\/s44318-024-00132-2"},{"key":"2026041210101056000_2025.06.23.659572v1.51","doi-asserted-by":"publisher","DOI":"10.1016\/j.bbamem.2017.03.013"},{"key":"2026041210101056000_2025.06.23.659572v1.52","doi-asserted-by":"publisher","DOI":"10.1021\/acs.biochem.9b00495"},{"key":"2026041210101056000_2025.06.23.659572v1.53","doi-asserted-by":"publisher","DOI":"10.1021\/bi00895a023"},{"key":"2026041210101056000_2025.06.23.659572v1.54","doi-asserted-by":"publisher","DOI":"10.1016\/j.bbamem.2021.183725"},{"key":"2026041210101056000_2025.06.23.659572v1.55","doi-asserted-by":"publisher","DOI":"10.1038\/nprot.2016.070"},{"key":"2026041210101056000_2025.06.23.659572v1.56","doi-asserted-by":"publisher","DOI":"10.1016\/B978-0-12-420067-8.00021-0"},{"key":"2026041210101056000_2025.06.23.659572v1.57","doi-asserted-by":"publisher","DOI":"10.1016\/0003-2697(80)90269-9"},{"key":"2026041210101056000_2025.06.23.659572v1.58","doi-asserted-by":"publisher","DOI":"10.3109\/10715769709065767"},{"key":"2026041210101056000_2025.06.23.659572v1.59","doi-asserted-by":"publisher","DOI":"10.1016\/s0301-4622(02)00111-4"},{"key":"2026041210101056000_2025.06.23.659572v1.60","doi-asserted-by":"publisher","DOI":"10.1016\/j.bpc.2005.06.007"},{"key":"2026041210101056000_2025.06.23.659572v1.61","doi-asserted-by":"publisher","DOI":"10.1016\/j.abb.2007.06.031"},{"key":"2026041210101056000_2025.06.23.659572v1.62","doi-asserted-by":"publisher","DOI":"10.1007\/s00249-011-0786-0"},{"key":"2026041210101056000_2025.06.23.659572v1.63","doi-asserted-by":"publisher","DOI":"10.1016\/j.bpc.2016.11.004"},{"key":"2026041210101056000_2025.06.23.659572v1.64","doi-asserted-by":"publisher","DOI":"10.3390\/toxins12110705"},{"key":"2026041210101056000_2025.06.23.659572v1.65","doi-asserted-by":"publisher","DOI":"10.1016\/S0006-3495(90)82637-0"},{"key":"2026041210101056000_2025.06.23.659572v1.66","doi-asserted-by":"publisher","DOI":"10.1562\/0031-8655(1999)070<0557:AMFTIO>2.3.CO;2"},{"key":"2026041210101056000_2025.06.23.659572v1.67","doi-asserted-by":"publisher","DOI":"10.3390\/ijms21196981"},{"key":"2026041210101056000_2025.06.23.659572v1.68","doi-asserted-by":"publisher","DOI":"10.1038\/s41598-019-40723-2"},{"key":"2026041210101056000_2025.06.23.659572v1.69","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0073040"},{"key":"2026041210101056000_2025.06.23.659572v1.70","doi-asserted-by":"publisher","DOI":"10.1016\/j.vibspec.2014.10.003"},{"key":"2026041210101056000_2025.06.23.659572v1.71","doi-asserted-by":"publisher","DOI":"10.1016\/j.bbamem.2016.03.007"},{"key":"2026041210101056000_2025.06.23.659572v1.72","doi-asserted-by":"publisher","DOI":"10.1016\/j.bbamem.2016.10.013"},{"key":"2026041210101056000_2025.06.23.659572v1.73","doi-asserted-by":"publisher","DOI":"10.1017\/s0033583500001645"},{"key":"2026041210101056000_2025.06.23.659572v1.74","doi-asserted-by":"publisher","DOI":"10.1016\/0009-3084(86)90067-8"},{"key":"2026041210101056000_2025.06.23.659572v1.75","doi-asserted-by":"publisher","DOI":"10.1016\/0005-2736(83)90134-7"},{"key":"2026041210101056000_2025.06.23.659572v1.76","doi-asserted-by":"publisher","DOI":"10.1016\/j.ultramic.2007.04.009"},{"key":"2026041210101056000_2025.06.23.659572v1.77","doi-asserted-by":"publisher","DOI":"10.1016\/j.bbamem.2009.03.014"},{"key":"2026041210101056000_2025.06.23.659572v1.78","doi-asserted-by":"publisher","DOI":"10.1016\/0005-2736(89)90331-3"},{"key":"2026041210101056000_2025.06.23.659572v1.79","doi-asserted-by":"publisher","DOI":"10.1016\/j.bpj.2018.04.001"},{"key":"2026041210101056000_2025.06.23.659572v1.80","doi-asserted-by":"publisher","DOI":"10.3390\/pharmaceutics8040036"},{"key":"2026041210101056000_2025.06.23.659572v1.81","doi-asserted-by":"publisher","DOI":"10.1007\/s13346-024-01584-z"},{"key":"2026041210101056000_2025.06.23.659572v1.82","doi-asserted-by":"publisher","DOI":"10.1006\/jcis.2002.8432"},{"key":"2026041210101056000_2025.06.23.659572v1.83","doi-asserted-by":"publisher","DOI":"10.1007\/s00216-017-0527-z"},{"key":"2026041210101056000_2025.06.23.659572v1.84","doi-asserted-by":"publisher","DOI":"10.1016\/j.cbi.2024.111215"},{"key":"2026041210101056000_2025.06.23.659572v1.85","doi-asserted-by":"publisher","DOI":"10.1529\/biophysj.108.136077"},{"key":"2026041210101056000_2025.06.23.659572v1.86","doi-asserted-by":"publisher","DOI":"10.1016\/j.biochi.2011.06.027"},{"key":"2026041210101056000_2025.06.23.659572v1.87","doi-asserted-by":"publisher","DOI":"10.1007\/s10895-016-1951-8"},{"key":"2026041210101056000_2025.06.23.659572v1.88","doi-asserted-by":"publisher","DOI":"10.1016\/S0006-3495(91)82041-0"},{"key":"2026041210101056000_2025.06.23.659572v1.89","doi-asserted-by":"publisher","DOI":"10.1023\/A:1020528716621"},{"key":"2026041210101056000_2025.06.23.659572v1.90","doi-asserted-by":"publisher","DOI":"10.1002\/1873-3468.13737"},{"key":"2026041210101056000_2025.06.23.659572v1.91","doi-asserted-by":"publisher","DOI":"10.1016\/j.chemphyslip.2012.02.002"},{"key":"2026041210101056000_2025.06.23.659572v1.92","doi-asserted-by":"publisher","DOI":"10.1039\/d1sm01591e"},{"key":"2026041210101056000_2025.06.23.659572v1.93","doi-asserted-by":"publisher","DOI":"10.1039\/a806472e"},{"key":"2026041210101056000_2025.06.23.659572v1.94","doi-asserted-by":"publisher","DOI":"10.1016\/j.bbamem.2004.06.010"},{"key":"2026041210101056000_2025.06.23.659572v1.95","doi-asserted-by":"publisher","DOI":"10.1021\/acs.jpcb.5b06604"},{"key":"2026041210101056000_2025.06.23.659572v1.96","doi-asserted-by":"publisher","DOI":"10.1016\/j.bpj.2019.06.023"},{"key":"2026041210101056000_2025.06.23.659572v1.97","doi-asserted-by":"publisher","DOI":"10.3390\/ijms242316570"},{"key":"2026041210101056000_2025.06.23.659572v1.98","doi-asserted-by":"publisher","DOI":"10.1016\/0304-4157(87)90017-7"},{"key":"2026041210101056000_2025.06.23.659572v1.99","doi-asserted-by":"publisher","DOI":"10.1074\/jbc.M115.706523"},{"key":"2026041210101056000_2025.06.23.659572v1.100","doi-asserted-by":"publisher","DOI":"10.1016\/j.bbamem.2011.07.013"},{"key":"2026041210101056000_2025.06.23.659572v1.101","doi-asserted-by":"publisher","DOI":"10.1016\/j.bpc.2018.04.001"},{"key":"2026041210101056000_2025.06.23.659572v1.102","doi-asserted-by":"publisher","DOI":"10.1099\/mic.0.28345-0"},{"key":"2026041210101056000_2025.06.23.659572v1.103","doi-asserted-by":"publisher","DOI":"10.1007\/s12551-016-0218-6"},{"key":"2026041210101056000_2025.06.23.659572v1.104","doi-asserted-by":"publisher","DOI":"10.1016\/j.bbamem.2011.12.010"},{"key":"2026041210101056000_2025.06.23.659572v1.105","doi-asserted-by":"publisher","DOI":"10.1016\/0304-4157(90)90002-T"},{"key":"2026041210101056000_2025.06.23.659572v1.106","doi-asserted-by":"publisher","DOI":"10.1016\/S0006-3495(99)77415-1"},{"key":"2026041210101056000_2025.06.23.659572v1.107","doi-asserted-by":"publisher","DOI":"10.1101\/cshperspect.a004747"},{"key":"2026041210101056000_2025.06.23.659572v1.108","doi-asserted-by":"publisher","DOI":"10.1016\/j.bbamem.2020.183450"},{"key":"2026041210101056000_2025.06.23.659572v1.109","doi-asserted-by":"publisher","DOI":"10.1021\/jacs.6b10615"},{"key":"2026041210101056000_2025.06.23.659572v1.110","doi-asserted-by":"publisher","DOI":"10.1016\/j.bpj.2011.07.013"},{"key":"2026041210101056000_2025.06.23.659572v1.111","doi-asserted-by":"publisher","DOI":"10.1016\/j.jlr.2022.100220"},{"key":"2026041210101056000_2025.06.23.659572v1.112","doi-asserted-by":"publisher","DOI":"10.1016\/j.bpj.2016.11.3205"},{"key":"2026041210101056000_2025.06.23.659572v1.113","doi-asserted-by":"publisher","DOI":"10.1021\/jp0750811"},{"key":"2026041210101056000_2025.06.23.659572v1.114","doi-asserted-by":"publisher","DOI":"10.1016\/j.bpc.2005.09.003"},{"key":"2026041210101056000_2025.06.23.659572v1.115","doi-asserted-by":"publisher","DOI":"10.1529\/biophysj.103.034280"}],"container-title":[],"original-title":[],"link":[{"URL":"https:\/\/syndication.highwire.org\/content\/doi\/10.1101\/2025.06.23.659572","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,6,24]],"date-time":"2026-06-24T04:57:05Z","timestamp":1782277025000},"score":1,"resource":{"primary":{"URL":"http:\/\/biorxiv.org\/lookup\/doi\/10.1101\/2025.06.23.659572"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,6,25]]},"references-count":115,"URL":"https:\/\/doi.org\/10.1101\/2025.06.23.659572","relation":{"is-preprint-of":[{"id-type":"doi","id":"10.1016\/j.bbamem.2025.184470","asserted-by":"subject"}]},"subject":[],"published":{"date-parts":[[2025,6,25]]},"subtype":"preprint"}}