{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,2]],"date-time":"2026-09-02T07:30:30Z","timestamp":1788334230771,"version":"build-2803163510"},"reference-count":78,"publisher":"American Chemical Society (ACS)","issue":"23","license":[{"start":{"date-parts":[[2026,6,2]],"date-time":"2026-06-02T00:00:00Z","timestamp":1780358400000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-029"},{"start":{"date-parts":[[2026,6,2]],"date-time":"2026-06-02T00:00:00Z","timestamp":1780358400000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-037"},{"start":{"date-parts":[[2026,6,2]],"date-time":"2026-06-02T00:00:00Z","timestamp":1780358400000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-045"}],"funder":[{"DOI":"10.13039\/100000057","name":"National Institute of General Medical Sciences","doi-asserted-by":"publisher","award":["R01GM121493-6"],"award-info":[{"award-number":["R01GM121493-6"]}],"id":[{"id":"10.13039\/100000057","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001691","name":"Japan Society for the Promotion of Science","doi-asserted-by":"publisher","award":["NA"],"award-info":[{"award-number":["NA"]}],"id":[{"id":"10.13039\/501100001691","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/na","name":"Ministerstvo \u0160kolstv\u00ed, Ml\u00e1de\u017ee a Telov\u00fdchovy","doi-asserted-by":"publisher","award":["MSMT-355\/2025-16"],"award-info":[{"award-number":["MSMT-355\/2025-16"]}],"id":[{"id":"10.13039\/na","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2026,6,17]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>Transmembrane (TM) asymmetry of glycerophospholipids in the plasma membrane is well established, yet still a complex and evolving area of study regarding the exact mechanisms, magnitude, and composition. The emerging controversy challenges \u201cFluid Mosaic Sandwich\u201d model simplicity by combining compositional (different types of lipids on each side) and numerical (different total amounts of phospholipids) asymmetries. While Gram-negative Escherichia coli phosphatidylethanolamine (PE) distribution is asymmetric (mostly cytoplasmic), the TM distribution of phosphatidylglycerol (PG) and cardiolipin (CL) across the inner (cytoplasmic) membrane (IM) is unknown. By integrating novel in vivo mapping techniques for PE, PG, and CL, we made the unexpected observation that their TM distribution does not correspond to the distribution of their total mass between the two IM leaflets. Instead, the underpopulated periplasmic leaflet can be occupied by lipid A, diacylglycerol (DAG), lipoproteins, and glycolipid MPase, whereas CL is found predominantly in the cytoplasmic leaflet of the IM. While PG is significantly enriched in the cytoplasmic leaflet of the IM in logarithmically growing E. coli, its asymmetrical distribution is inverted in stationary-grown cells, cells lacking CL, or cells subjected to growth in a hypotonic medium. These data suggest that cells can minimize the numerical differences in TM distribution by triggering a biosynthetic TM \u201ccleanup\u201d response, preventing membrane collapse. The numerical asymmetric interleaflet distribution of phospholipids may serve specific functional roles, potentially governed by critical threshold levels of lipid A and DAG that determine their compatibility for colocalization within the same leaflet. The overabundance of glycerophospholipids in the cytoplasmic leaflet of the IM may be driven by packing and localization constraints imposed during cell division and impacted by the rate of phospholipid biosynthesis and intermembrane flows. This imbalance may generate or result from the lipid concentration gradients and downhill flows across IM and between IM and outer membrane (OM), and the necessity to maintain appropriate electrostatic balance across the IM.<\/jats:p>","DOI":"10.1021\/jacs.6c06591","type":"journal-article","created":{"date-parts":[[2026,6,2]],"date-time":"2026-06-02T16:09:24Z","timestamp":1780416564000},"page":"24455-24476","source":"Crossref","is-referenced-by-count":1,"title":["Dark\nSide of\n                    <i>Escherichia coli<\/i>\n                    Biogenic Inner\nMembrane: Overabundance of Three Main Phospholipids\non Cytoplasmic Leaflet"],"prefix":"10.1021","volume":"148","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-9965-970X","authenticated-orcid":true,"given":"Katsuhiro","family":"Sawasato","sequence":"first","affiliation":[{"name":"The University of Texas Health Science Center, McGovern Medical School , , 6431 Fannin St., Suite 6.204 , , ,","place":["Houston, Texas, United States, 77030"]},{"name":"Iwate University , , , ,","place":["Morioka, Iwate, Japan, 020-8550"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Nataliia","family":"Bogdanova","sequence":"additional","affiliation":[{"name":"The University of Texas Health Science Center, McGovern Medical School , , 6431 Fannin St., Suite 6.204 , , ,","place":["Houston, Texas, United States, 77030"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"William","family":"Dowhan","sequence":"additional","affiliation":[{"name":"The University of Texas Health Science Center, McGovern Medical School , , 6431 Fannin St., Suite 6.204 , , ,","place":["Houston, Texas, United States, 77030"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6748-8931","authenticated-orcid":true,"given":"Semen","family":"Yesylevskyy","sequence":"additional","affiliation":[{"name":"Czech Academy of Sciences , , Flemingovo na\u0301me\u030csti\u0301 542\/2, 160 00 Prague 6 , ,","place":["Prague 6, Czech Republic, CZ-166 10"]},{"name":"Receptor.AI Inc ., 20-22 Wenlock Road , ,","place":["London, United Kingdom, N1 7GU"]},{"name":"Palacky\u0301 University Olomouc , , 17. listopadu 12 , ,","place":["Olomouc, Czech Republic, 771 46"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-7176-8127","authenticated-orcid":true,"given":"Mikhail","family":"Bogdanov","sequence":"additional","affiliation":[{"name":"The University of Texas Health Science Center, McGovern Medical School , , 6431 Fannin St., Suite 6.204 , , ,","place":["Houston, Texas, United States, 77030"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"316","published-online":{"date-parts":[[2026,6,2]]},"reference":[{"issue":"1","key":"2026073023165363300_cit1","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1042\/ETLS20220088","article-title":"The Power\nand Challenge of Lipid (a)Symmetry across the Membrane and Cell","volume":"7","author":"Bogdanov","year":"2023","journal-title":"Emerging Top. Life Sci."},{"issue":"23","key":"2026073023165363300_cit2","doi-asserted-by":"publisher","DOI":"10.1126\/sciadv.aaz6333","article-title":"Phospholipid\nDistribution in the\nCytoplasmic Membrane of Gram-Negative Bacteria is Highly Asymmetric,\nDynamic, and Cell Shape-Dependent","volume":"6","author":"Bogdanov","year":"2020","journal-title":"Sci. Adv."},{"issue":"1","key":"2026073023165363300_cit3","doi-asserted-by":"publisher","first-page":"137","DOI":"10.1042\/ETLS20230042","article-title":"Renovating\na Double Fence with or without Notifying the next Door and across\nthe Street Neighbors: Why the Biogenic Cytoplasmic Membrane of Gram-Negative\nBacteria Display Asymmetry?","volume":"7","author":"Bogdanov","year":"2023","journal-title":"Emerging Top. Life\nSci."},{"issue":"1","key":"2026073023165363300_cit4","doi-asserted-by":"publisher","first-page":"255","DOI":"10.1146\/annurev-micro-102215-095308","article-title":"The Power of Asymmetry: Architecture\nand Assembly of the Gram-Negative Outer Membrane Lipid Bilayer","volume":"70","author":"Henderson","year":"2016","journal-title":"Annu. Rev. Microbiol."},{"issue":"6178","key":"2026073023165363300_cit5","doi-asserted-by":"publisher","first-page":"173","DOI":"10.1038\/334173a0","article-title":"Phosphatidylglycerol is Involved in Protein Translocation\nacross Escherichia coli Inner Membranes","volume":"334","author":"de Vrije","year":"1988","journal-title":"Nature"},{"issue":"3","key":"2026073023165363300_cit6","doi-asserted-by":"publisher","first-page":"783","DOI":"10.1073\/pnas.92.3.783","article-title":"In vivo Evidence for the Involvement\nof Anionic Phospholipids in Initiation of DNA Replication in Escherichia coli","volume":"92","author":"Xia","year":"1995","journal-title":"Proc. Natl.\nAcad. Sci. U.S.A."},{"issue":"15","key":"2026073023165363300_cit7","doi-asserted-by":"publisher","first-page":"7131","DOI":"10.1016\/S0021-9258(18)68615-6","article-title":"Cardiolipin Activation of DnaA Protein, the Initiation\nProtein of Replication in Escherichia coli","volume":"263","author":"Sekimizu","year":"1988","journal-title":"J. Biol. Chem."},{"issue":"25","key":"2026073023165363300_cit8","doi-asserted-by":"publisher","first-page":"22193","DOI":"10.1074\/jbc.M302603200","article-title":"Effects of Phospholipid\nComposition on MinD-Membrane\nInteractions in vitro and in vivo","volume":"278","author":"Mileykovskaya","year":"2003","journal-title":"J. Biol. Chem."},{"issue":"2","key":"2026073023165363300_cit9","doi-asserted-by":"publisher","first-page":"135","DOI":"10.1016\/j.mib.2005.02.012","article-title":"Role of Membrane Lipids\nin Bacterial Division-Site\nSelection","volume":"8","author":"Mileykovskaya","year":"2005","journal-title":"Curr. Opin. Microbiol."},{"issue":"46","key":"2026073023165363300_cit10","doi-asserted-by":"publisher","first-page":"38835","DOI":"10.1074\/jbc.M112.407817","article-title":"MinD and MinE Interact\nwith Anionic Phospholipids and\nRegulate Division Plane Formation in Escherichia coli","volume":"287","author":"Renner","year":"2012","journal-title":"J. Biol. Chem."},{"key":"2026073023165363300_cit11","doi-asserted-by":"publisher","first-page":"189","DOI":"10.1016\/0006-3002(58)90308-1","article-title":"Plant Phospholipids\nI. Identification of the Phosphatidyl Glycerols","volume":"27","author":"Benson","year":"1958","journal-title":"Biochim. Biophys. Acta"},{"key":"2026073023165363300_cit12","doi-asserted-by":"publisher","first-page":"447","DOI":"10.1016\/S0006-3002(89)80039-3","article-title":"A \u201cnova\u201d in Phosphate\nMetabolism, GPG, and Discovery\nof Phosphatidylglycerol: A Commentary on \u201cPlant Phospholipids.\nI. Identification of the Phosphatidyl Glycerols\u201d","volume":"1000","author":"Benson","year":"1989","journal-title":"Biochim. Biophys. Acta"},{"issue":"1","key":"2026073023165363300_cit13","doi-asserted-by":"publisher","first-page":"210","DOI":"10.1128\/jb.116.1.210-214.1973","article-title":"Turnover of\nPhosphatidylglycerol\nin Escherichia coli","volume":"116","author":"Ballesta","year":"1973","journal-title":"J. Bacteriol."},{"issue":"41","key":"2026073023165363300_cit14","doi-asserted-by":"publisher","first-page":"16504","DOI":"10.1073\/pnas.1212797109","article-title":"Discovery of a Cardiolipin Synthase\nUtilizing Phosphatidylethanolamine\nand Phosphatidylglycerol as Substrates","volume":"109","author":"Tan","year":"2012","journal-title":"Proc.\nNatl. Acad. Sci. U.S.A."},{"issue":"48","key":"2026073023165363300_cit15","doi-asserted-by":"publisher","first-page":"25144","DOI":"10.1074\/jbc.M116.762070","article-title":"In vivo and in vitro Synthesis\nof Phosphatidylglycerol\nby an Escherichia coli Cardiolipin\nSynthase","volume":"291","author":"Li","year":"2016","journal-title":"J. Biol. Chem."},{"issue":"5","key":"2026073023165363300_cit16","doi-asserted-by":"publisher","DOI":"10.1016\/j.bbalip.2024.159483","article-title":"The Unusual Substrate\nSpecificity of Escherichia coli Cardiolipin\nSynthase C Does Not\nRequire the Product of the Transcriptionally Engaged ymdB Gene","volume":"1869","author":"Sawasato","year":"2024","journal-title":"Biochim. Biophys. Acta, Mol. Cell Biol.\nLipids"},{"issue":"9","key":"2026073023165363300_cit17","doi-asserted-by":"publisher","first-page":"2142","DOI":"10.1128\/JB.06641-11","article-title":"Phosphatidylglycerol:Prolipoprotein\nDiacylglyceryl Transferase (Lgt) of Escherichia coli Has Seven Transmembrane Segments, and its Essential Residues are\nEmbedded in the Membrane","volume":"194","author":"Pailler","year":"2012","journal-title":"J. Bacteriol."},{"issue":"5","key":"2026073023165363300_cit18","doi-asserted-by":"publisher","first-page":"1368","DOI":"10.1073\/pnas.70.5.1368","article-title":"Metabolism of Membrane\nPhospholipids and its Relation to a Novel Class of Oligosaccharides\nin Escherichia coli","volume":"70","author":"Van Golde","year":"1973","journal-title":"Proc. Natl. Acad. Sci. U.S.A."},{"issue":"4","key":"2026073023165363300_cit19","doi-asserted-by":"publisher","first-page":"1092","DOI":"10.1073\/pnas.79.4.1092","article-title":"Osmotic\nRegulation and the Biosynthesis of Membrane-Derived Oligosaccharides\nin Escherichia coli","volume":"79","author":"Kennedy","year":"1982","journal-title":"Proc. Natl. Acad. Sci. U.S.A."},{"issue":"5","key":"2026073023165363300_cit20","doi-asserted-by":"publisher","first-page":"690","DOI":"10.1111\/1758-2229.12325","article-title":"New Insights into the\nBiological Role of the Osmoregulated\nPeriplasmic Glucans in Pathogenic and Symbiotic Bacteria","volume":"7","author":"Bontemps-Gallo","year":"2015","journal-title":"Environ. Microbiol. Rep."},{"issue":"1","key":"2026073023165363300_cit21","doi-asserted-by":"publisher","DOI":"10.1128\/jb.00403-22","article-title":"Phosphatidylglycerol\nis the Lipid Donor for Synthesis\nof Phospholipid-Linked Enterobacterial Common Antigen","volume":"205","author":"Morris","year":"2023","journal-title":"J. Bacteriol."},{"issue":"9","key":"2026073023165363300_cit22","doi-asserted-by":"publisher","DOI":"10.1128\/JB.00804-18","article-title":"Rcs Phosphorelay Activation in Cardiolipin-Deficient Escherichia coli Reduces Biofilm Formation","volume":"201","author":"Nepper","year":"2019","journal-title":"J. Bacteriol."},{"issue":"1","key":"2026073023165363300_cit23","doi-asserted-by":"publisher","DOI":"10.1038\/s41598-020-63280-5","article-title":"Cardiolipin is Required in vivo for\nthe Stability of Bacterial Translocon and Optimal Membrane Protein\nTranslocation and Insertion","volume":"10","author":"Ryabichko","year":"2020","journal-title":"Sci. Rep."},{"issue":"1","key":"2026073023165363300_cit24","doi-asserted-by":"publisher","DOI":"10.1038\/s41598-019-54198-8","article-title":"Cardiolipin is an Optimal\nPhospholipid for the Assembly, Stability, and Proper Functionality\nof the Dimeric Form of NhaA Na+\/H+ Antiporter","volume":"9","author":"Rimon","year":"2019","journal-title":"Sci. Rep."},{"issue":"13","key":"2026073023165363300_cit25","doi-asserted-by":"publisher","DOI":"10.1128\/JB.00849-16","article-title":"Impact of Membrane Phospholipid Alterations\nin Escherichia coli on Cellular Function\nand Bacterial Stress Adaptation","volume":"199","author":"Rowlett","year":"2017","journal-title":"J. Bacteriol."},{"issue":"15","key":"2026073023165363300_cit26","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.2018329118","article-title":"Cardiolipin Aids in Lipopolysaccharide\nTransport to the Gram-Negative Outer Membrane","volume":"118","author":"Douglass","year":"2021","journal-title":"Proc. Natl. Acad. Sci. U.S.A."},{"issue":"10","key":"2026073023165363300_cit27","doi-asserted-by":"publisher","first-page":"5099","DOI":"10.3390\/ijms22105099","article-title":"Molecular Basis of Essentiality of\nEarly Critical Steps in the Lipopolysaccharide Biogenesis in Escherichia coli K-12: Requirement of MsbA, Cardiolipin,\nLpxL, LpxM and GcvB","volume":"22","author":"Gorzelak","year":"2021","journal-title":"Int. J. Mol. Sci."},{"issue":"19","key":"2026073023165363300_cit28","doi-asserted-by":"publisher","first-page":"3386","DOI":"10.1128\/JB.01877-14","article-title":"Localization of\nAnionic Phospholipids in Escherichia coli Cells","volume":"196","author":"Oliver","year":"2014","journal-title":"J.\nBacteriol."},{"issue":"2","key":"2026073023165363300_cit29","doi-asserted-by":"publisher","first-page":"1263","DOI":"10.1021\/ac403616c","article-title":"Superior Fluorescent Probe for Detection\nof Cardiolipin","volume":"86","author":"Leung","year":"2014","journal-title":"Anal. Chem."},{"issue":"43","key":"2026073023165363300_cit30","doi-asserted-by":"publisher","first-page":"9111","DOI":"10.1021\/acs.jpcb.9b07690","article-title":"Cardiolipin\nPreferentially Partitions to the Inner Leaflet of Mixed Lipid Large\nUnilamellar Vesicles","volume":"123","author":"Elmer-Dixon","year":"2019","journal-title":"J. Phys.Chem. B"},{"issue":"8","key":"2026073023165363300_cit31","doi-asserted-by":"publisher","DOI":"10.1016\/j.bbamem.2020.183241","article-title":"Lipid Asymmetry\nof a Model Mitochondrial Outer Membrane Affects Bax-Dependent Permeabilization","volume":"1862","author":"Bozelli","year":"2020","journal-title":"Biochim. Biophys. Acta, Biomembr."},{"issue":"17","key":"2026073023165363300_cit32","doi-asserted-by":"publisher","first-page":"3776","DOI":"10.1016\/j.bpj.2021.06.039","article-title":"TTAPE-Me Dye is Not Selective to\nCardiolipin and Binds to Common Anionic Phospholipids Nonspecifically","volume":"120","author":"Pyrshev","year":"2021","journal-title":"Biophys. J."},{"issue":"1","key":"2026073023165363300_cit33","doi-asserted-by":"publisher","DOI":"10.1126\/sciadv.ads0244","article-title":"Its Own Architect:\nFlipping Cardiolipin\nSynthase","volume":"11","author":"Sawasato","year":"2025","journal-title":"Sci. Adv."},{"issue":"2","key":"2026073023165363300_cit34","doi-asserted-by":"publisher","DOI":"10.1002\/bies.70114","article-title":"Critique of a Radically\nNew Model for Plasma Membrane\nBilayer Organization","volume":"48","author":"Lange","year":"2026","journal-title":"BioEssays"},{"issue":"22","key":"2026073023165363300_cit35","doi-asserted-by":"publisher","first-page":"3923","DOI":"10.1016\/j.bpj.2024.10.004","article-title":"Unexpected Asymmetric Distribution of Cholesterol and\nPhospholipids in Equilibrium Model Membranes","volume":"123","author":"Zhu","year":"2024","journal-title":"Biophys. J."},{"issue":"10","key":"2026073023165363300_cit36","doi-asserted-by":"publisher","first-page":"2586","DOI":"10.1016\/j.cell.2025.02.034","article-title":"Cell Membranes Sustain\nPhospholipid\nImbalance via Cholesterol Asymmetry","volume":"188","author":"Doktorova","year":"2025","journal-title":"Cell"},{"issue":"43","key":"2026073023165363300_cit37","doi-asserted-by":"publisher","first-page":"45102","DOI":"10.1074\/jbc.M408106200","article-title":"MsbA-Dependent\nTranslocation of Lipids\nacross the Inner Membrane of Escherichia coli","volume":"279","author":"Doerrler","year":"2004","journal-title":"J. Biol. Chem."},{"issue":"1","key":"2026073023165363300_cit38","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-026-68743-3","article-title":"Phospholipid\nComposition Strongly Affects the Assembly of \u03b2 Barrel Proteins\ninto Purified Bacterial Outer Membranes","volume":"17","author":"Nilaweera","year":"2026","journal-title":"Nat.\nCommun."},{"issue":"12","key":"2026073023165363300_cit39","doi-asserted-by":"publisher","first-page":"2555","DOI":"10.1021\/bi00553a003","article-title":"Determination\nof Phosphatidylglycerol\nAsymmetry in Small, Unilamellar Vesicles by Chemical Modification","volume":"19","author":"Lentz","year":"1980","journal-title":"Biochemistry"},{"key":"2026073023165363300_cit40","doi-asserted-by":"publisher","first-page":"1627","DOI":"10.1021\/bi00436a009","article-title":"Phospholipid Asymmetry\nin Large Unilamellar Vesicles\nInduced by Transmembrane pH Gradients","volume":"28","author":"Hope","year":"1989","journal-title":"Biochemistry"},{"issue":"1","key":"2026073023165363300_cit41","doi-asserted-by":"publisher","first-page":"35","DOI":"10.3109\/09687689709048168","article-title":"On the Accessibility\nof Phosphatidylglycerol to Periodate in Escherichia coli","volume":"14","author":"Huijbregts","year":"1997","journal-title":"Mol. Membr. Biol."},{"key":"2026073023165363300_cit42","doi-asserted-by":"crossref","unstructured":"Eisenhuth, A. G.; Adamson, L. S. R.; Zhang, C.; Abbas, G. S. K.; North, R. A.; Leyton, D. L.; Bernstein, H. D.; Brown, S. H. J.; Stewart, A. G.; Bailey, C. B.; Don, A. S.; Grosas, A. B.; Doyle, M. T.\n          Mechanism of\nPhospholipid Transport to the Bacterial Outer Membrane by TAM\n          bioRxiv\n          2026\n          10.64898\/2026.03.22.713439.","DOI":"10.64898\/2026.03.22.713439"},{"issue":"4","key":"2026073023165363300_cit43","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.2524588123","article-title":"A Surface-Exposed\nCardiolipin Synthase Provides an Unexpected Paradigm for Maintaining\nthe Gram-Negative Outer Membrane","volume":"123","author":"Herrera","year":"2026","journal-title":"Proc. Natl.\nAcad. Sci. U.S.A."},{"issue":"4","key":"2026073023165363300_cit44","doi-asserted-by":"publisher","first-page":"441","DOI":"10.1016\/j.chom.2015.03.003","article-title":"Delivery\nof Cardiolipins to the Salmonella\nOuter Membrane Is Necessary for Survival within Host Tissues and Virulence","volume":"17","author":"Dalebroux","year":"2015","journal-title":"Cell Host Microbe"},{"key":"2026073023165363300_cit45","doi-asserted-by":"publisher","first-page":"103","DOI":"10.1016\/j.biochi.2016.02.004","article-title":"Effects of\nElevated Growth Temperature and Heat Shock\non the Lipid Composition of the Inner and Outer Membranes of Yersinia pseudotuberculosis","volume":"123","author":"Davydova","year":"2016","journal-title":"Biochimie"},{"issue":"3","key":"2026073023165363300_cit46","doi-asserted-by":"publisher","first-page":"369","DOI":"10.1093\/jb\/mvab052","article-title":"Erylysin A\nInhibits Cytokinesis in Escherichia coli by Binding with Cardiolipin","volume":"170","author":"Sakihara","year":"2021","journal-title":"J. Biochem."},{"issue":"12","key":"2026073023165363300_cit47","doi-asserted-by":"publisher","first-page":"2008","DOI":"10.1091\/mbc.e13-03-0121","article-title":"Deacylation\non the Matrix Side of\nthe Mitochondrial Inner Membrane Regulates Cardiolipin Remodeling","volume":"24","author":"Baile","year":"2013","journal-title":"Mol. Biol. Cell"},{"issue":"9","key":"2026073023165363300_cit48","doi-asserted-by":"publisher","first-page":"3386","DOI":"10.1074\/jbc.RA117.001231","article-title":"The Phospholipid-Repair\nSystem LplT\/Aas\nin Gram-Negative Bacteria Protects the Bacterial Membrane Envelope\nfrom Host Phospholipase A2 Attack","volume":"293","author":"Lin","year":"2018","journal-title":"J. Biol.\nChem."},{"issue":"5","key":"2026073023165363300_cit49","doi-asserted-by":"publisher","first-page":"2136","DOI":"10.1074\/jbc.M115.700419","article-title":"Substrate\nSelectivity of Lysophospholipid Transporter LplT Involved in Membrane\nPhospholipid Remodeling in Escherichia coli","volume":"291","author":"Lin","year":"2016","journal-title":"J. Biol. Chem."},{"issue":"34","key":"2026073023165363300_cit50","doi-asserted-by":"publisher","DOI":"10.1126\/sciadv.abh2217","article-title":"Identification\nand Assessment of Cardiolipin Interactions\nwith E. coli Inner Membrane Proteins","volume":"7","author":"Corey","year":"2021","journal-title":"Sci. Adv."},{"issue":"38","key":"2026073023165363300_cit51","doi-asserted-by":"publisher","first-page":"8325","DOI":"10.1021\/bi101253s","article-title":"Escherichia coli Mutants That Synthesize Dephosphorylated Lipid A Molecules","volume":"49","author":"Ingram","year":"2010","journal-title":"Biochemistry"},{"key":"2026073023165363300_cit52","doi-asserted-by":"crossref","unstructured":"Denic, M.; Delprat, M.; Espinosa, L.; Barani, A. E.; Seduk, F.; Yamaryo-Botte, Y.; Botte, C. Y.; Audebert, S.; Camoin, L.; Magalon, A.\n          Cardiolipin Constrains\nLipid Unsaturation during Anaerobic Adaptation in Escherichia coli.  bioRxiv, 202610.64898\/2026.03.04.709520.","DOI":"10.64898\/2026.03.04.709520"},{"key":"2026073023165363300_cit53","doi-asserted-by":"publisher","first-page":"159","DOI":"10.1007\/978-1-0716-3445-5_10","article-title":"Preparation\nof Uniformly Oriented Inverted Inner (Cytoplasmic) Membrane Vesicles\nfrom Gram-Negative Bacterial Cells","volume":"2715","author":"Bogdanov","year":"2024","journal-title":"Methods\nMol. Biol."},{"issue":"9","key":"2026073023165363300_cit54","doi-asserted-by":"publisher","first-page":"3391","DOI":"10.1111\/1462-2920.12814","article-title":"Fatty Acid-releasing\nActivities in Sinorhizobium meliloti Include Unusual\nDiacylglycerol Lipase","volume":"17","author":"Sahonero-Canavesi","year":"2015","journal-title":"Environ. Microbiol."},{"key":"2026073023165363300_cit55","doi-asserted-by":"crossref","unstructured":"Bontemps-Gallo, S.; Bohin, J.-P.; Lacroix, J.-M.\n          Osmoregulated\nPeriplasmic Glucans\n          EcoSal Plus\n          2017; Vol. 7(2), 10.1128\/ecosalplus.esp-0001-2017.","DOI":"10.1128\/ecosalplus.esp-0001-2017"},{"issue":"18","key":"2026073023165363300_cit56","doi-asserted-by":"publisher","first-page":"4802","DOI":"10.1128\/JB.00576-12","article-title":"The MscS and MscL Families\nof Mechanosensitive Channels\nAct as Microbial Emergency Release Valves","volume":"194","author":"Booth","year":"2012","journal-title":"J.\nBacteriol."},{"issue":"34","key":"2026073023165363300_cit57","doi-asserted-by":"publisher","first-page":"4742","DOI":"10.1021\/acs.biochem.6b00541","article-title":"Three-Dimensional Distribution of Phospholipids in\nGram Negative Bacteria","volume":"55","author":"Furse","year":"2016","journal-title":"Biochemistry"},{"issue":"1","key":"2026073023165363300_cit58","doi-asserted-by":"publisher","first-page":"184","DOI":"10.1021\/acs.jpcb.1c07332","article-title":"Leaflet Asymmetry Modeling in the Lipid Composition\nof Escherichia coli Cytoplasmic Membranes","volume":"126","author":"Hsieh","year":"2022","journal-title":"J. Phys. Chem. B"},{"issue":"6","key":"2026073023165363300_cit59","doi-asserted-by":"publisher","first-page":"644","DOI":"10.1038\/s41589-020-0529-6","article-title":"Plasma Membranes\nare Asymmetric in Lipid Unsaturation, Packing and Protein Shape","volume":"16","author":"Lorent","year":"2020","journal-title":"Nat. Chem. Biol."},{"issue":"12","key":"2026073023165363300_cit60","doi-asserted-by":"publisher","first-page":"1321","DOI":"10.1038\/s41589-020-00688-0","article-title":"Structural\nand Functional Consequences\nof Reversible Lipid Asymmetry in Living Membranes","volume":"16","author":"Doktorova","year":"2020","journal-title":"Nat. Chem. Biol."},{"issue":"9","key":"2026073023165363300_cit61","doi-asserted-by":"publisher","first-page":"5098","DOI":"10.1021\/acs.chemrev.0c00587","article-title":"Assembly and\nMaintenance of Lipids\nat the Bacterial Outer Membrane","volume":"121","author":"Lundstedt","year":"2021","journal-title":"Chem. Rev."},{"issue":"1","key":"2026073023165363300_cit62","doi-asserted-by":"publisher","first-page":"553","DOI":"10.1146\/annurev-micro-032521-014507","article-title":"How Bacteria\nEstablish and Maintain Outer Membrane\nLipid Asymmetry","volume":"78","author":"Tan","year":"2024","journal-title":"Annu. Rev. Microbiol."},{"issue":"1","key":"2026073023165363300_cit63","doi-asserted-by":"publisher","DOI":"10.1038\/ncomms2267","article-title":"MPIase is a Glycolipozyme Essential for Membrane Protein\nIntegration","volume":"3","author":"Nishiyama","year":"2012","journal-title":"Nat. Commun."},{"issue":"1","key":"2026073023165363300_cit64","doi-asserted-by":"publisher","DOI":"10.1038\/s41598-018-37809-8","article-title":"CdsA is Involved in Biosynthesis of Glycolipid MPIase\nEssential for Membrane Protein Integration in Vivo","volume":"9","author":"Sawasato","year":"2019","journal-title":"Sci. Rep."},{"key":"2026073023165363300_cit65","doi-asserted-by":"publisher","DOI":"10.3389\/fmolb.2022.986602","article-title":"Interaction\nbetween Glycolipid MPIase\nand Proteinaceous Factors during Protein Integration into the Cytoplasmic\nMembrane of E. coli","volume":"9","author":"Nishikawa","year":"2022","journal-title":"Front. Mol. Biosci."},{"issue":"10","key":"2026073023165363300_cit66","doi-asserted-by":"publisher","first-page":"889","DOI":"10.3390\/pathogens13100889","article-title":"Breaking Barriers:\nExploiting Envelope Biogenesis and Stress Responses to Develop Novel\nAntimicrobial Strategies in Gram-Negative Bacteria","volume":"13","author":"Bisht","year":"2024","journal-title":"Pathogens"},{"key":"2026073023165363300_cit67","first-page":"121","article-title":"Exploring Uniform,\nDual, and Dynamic Topologies of Membrane Proteins by Substituted Cysteine\nAccessibility Method (SCAMTM)","volume-title":"Methods Mol. Biol.","author":"Bogdanov","year":"2024"},{"issue":"7","key":"2026073023165363300_cit68","first-page":"5595","article-title":"Extraction and Purification of Phosphatidic\nAcid of C18 Fatty Acids from Powdered Soybean Phospholipids","volume":"20","author":"Liu","year":"2008","journal-title":"Asian J. Chem."},{"issue":"11","key":"2026073023165363300_cit69","doi-asserted-by":"publisher","first-page":"1859","DOI":"10.1002\/jcc.20945","article-title":"CHARMM-GUI: A Web-based\nGraphical\nUser Interface for CHARMM","volume":"29","author":"Jo","year":"2008","journal-title":"J. Comput. Chem."},{"issue":"1","key":"2026073023165363300_cit70","doi-asserted-by":"publisher","first-page":"405","DOI":"10.1021\/acs.jctc.5b00935","article-title":"CHARMM-GUI Input Generator\nfor NAMD, GROMACS, AMBER, OpenMM, and CHARMM\/OpenMM Simulations Using\nthe CHARMM36 Additive Force Field","volume":"12","author":"Lee","year":"2016","journal-title":"J. Chem.\nTheory Comput."},{"key":"2026073023165363300_cit71","doi-asserted-by":"publisher","first-page":"19","DOI":"10.1016\/j.softx.2015.06.001","article-title":"GROMACS: High Performance\nMolecular Simulations through Multi-Level Parallelism from Laptops\nto Supercomputers","volume":"1-2","author":"Abraham","year":"2015","journal-title":"SoftwareX"},{"issue":"12","key":"2026073023165363300_cit72","doi-asserted-by":"publisher","first-page":"7182","DOI":"10.1063\/1.328693","article-title":"Polymorphic Transitions in Single\nCrystals: A New Molecular\nDynamics Method","volume":"52","author":"Parrinello","year":"1981","journal-title":"J. Appl. Phys."},{"issue":"1","key":"2026073023165363300_cit73","doi-asserted-by":"publisher","DOI":"10.1063\/1.2408420","article-title":"Canonical\nSampling through Velocity\nRescaling","volume":"126","author":"Bussi","year":"2007","journal-title":"J. Chem. Phys."},{"issue":"1","key":"2026073023165363300_cit74","doi-asserted-by":"publisher","first-page":"98","DOI":"10.1103\/PhysRev.159.98","article-title":"Computer \u201cExperiments\u201d\non Classical Fluids. I. Thermodynamical Properties of Lennard-Jones\nMolecules","volume":"159","author":"Verlet","year":"1967","journal-title":"Phys. Rev."},{"issue":"12","key":"2026073023165363300_cit75","doi-asserted-by":"publisher","first-page":"1463","DOI":"10.1002\/(SICI)1096-987X(199709)18:12&lt;1463::AID-JCC4&gt;3.0.CO;2-H","article-title":"LINCS: A Linear\nConstraint Solver for Molecular Simulations","volume":"18","author":"Hess","year":"1997","journal-title":"J. Comput. Chem."},{"issue":"23","key":"2026073023165363300_cit76","doi-asserted-by":"publisher","first-page":"7830","DOI":"10.1021\/jp101759q","article-title":"Update of the CHARMM\nAll-Atom Additive Force Field\nfor Lipids: Validation on Six Lipid Types","volume":"114","author":"Klauda","year":"2010","journal-title":"J.\nPhys. Chem. B"},{"issue":"2","key":"2026073023165363300_cit77","doi-asserted-by":"publisher","DOI":"10.1016\/j.bpr.2021.100034","article-title":"Recharging Your Fats:\nCHARMM36 Parameters for Neutral Lipids Triacylglycerol and Diacylglycerol","volume":"1","author":"Campomanes","year":"2021","journal-title":"Biophys. Rep."},{"issue":"1","key":"2026073023165363300_cit78","doi-asserted-by":"publisher","DOI":"10.1002\/jcc.27536","article-title":"MolAR: Memory-Safe\nLibrary for Analysis of MD Simulations Written in Rust","volume":"46","author":"Yesylevskyy","year":"2025","journal-title":"J. Comput. Chem."}],"container-title":["Journal of the American\nChemical Society"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/pubs.acs.org\/doi\/pdf\/10.1021\/jacs.6c06591","content-type":"application\/pdf","content-version":"vor","intended-application":"unspecified"},{"URL":"https:\/\/pubs.acs.org\/jacsat\/article-pdf\/148\/23\/24455\/66272245\/jacs.6c06591.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/pubs.acs.org\/jacsat\/article-pdf\/148\/23\/24455\/66272245\/jacs.6c06591.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,7,31]],"date-time":"2026-07-31T03:17:04Z","timestamp":1785467824000},"score":1,"resource":{"primary":{"URL":"https:\/\/pubs.acs.org\/jacsat\/article\/148\/23\/24455\/5233506\/Dark-Side-of-Escherichia-coli-Biogenic-Inner"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,6,2]]},"references-count":78,"journal-issue":{"issue":"23","published-online":{"date-parts":[[2026,6,2]]},"published-print":{"date-parts":[[2026,6,17]]}},"URL":"https:\/\/doi.org\/10.1021\/jacs.6c06591","relation":{},"ISSN":["0002-7863","1520-5126"],"issn-type":[{"value":"0002-7863","type":"print"},{"value":"1520-5126","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,6,2]]}}}