{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,7]],"date-time":"2026-08-07T17:17:04Z","timestamp":1786123024685,"version":"build-2736575974"},"reference-count":69,"publisher":"American Chemical Society (ACS)","issue":"8","license":[{"start":{"date-parts":[[2027,7,6]],"date-time":"2027-07-06T00:00:00Z","timestamp":1814832000000},"content-version":"vor","delay-in-days":365,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0\/"},{"start":{"date-parts":[[2026,7,6]],"date-time":"2026-07-06T00:00:00Z","timestamp":1783296000000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-029"},{"start":{"date-parts":[[2026,7,6]],"date-time":"2026-07-06T00:00:00Z","timestamp":1783296000000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-037"},{"start":{"date-parts":[[2026,7,6]],"date-time":"2026-07-06T00:00:00Z","timestamp":1783296000000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-045"}],"funder":[{"DOI":"10.13039\/501100002915","name":"Fondation pour la Recherche M?dicale","doi-asserted-by":"publisher","award":["FDT202204014720"],"award-info":[{"award-number":["FDT202204014720"]}],"id":[{"id":"10.13039\/501100002915","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2026,8,7]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>Endothelial cells (ECs) are key regulators in vascular homeostasis, and their physiological state depends on hemodynamic forces produced by the blood flow. Laminar shear stress (LSS) maintains endothelial integrity, whereas oscillatory shear stress (OSS) leads to endothelial dysfunction and subsequently to cardiovascular disease. Here, we delineated the proteomic signatures of human umbilical vein endothelial cells (HUVECs) exposed to a protective flow (LSS) or a disturbed flow (OSS) in comparison to static conditions, using trapped ion mobility spectrometry coupled with parallel accumulation-serial fragmentation. OSS induced limited changes in cell morphology and the proteomic profile, whereas LSS triggered major modifications in cell shape and a proteomic signature related to the cell surface and extracellular matrix. Membrane subproteomic profiling confirmed the identification of membrane effectors involved in flow-mediated responses and in silico integrating promoter analysis suggested Kru\u0308ppel-like Factor 4 (KLF4) as an important transcriptional regulator of this effector cluster. Using a novel EC model, TeloHAECs, we demonstrated that KLF4 likely contributes to LSS-induced alignment and elongation by increasing the expression of CD34, HEG1, PI16, and ITGB4 and that KLF4 is involved in PI16 and ITGB4 expression. Collectively, these findings provide a robust discovery data set on EC proteomic profiles related to flow and a refined view of KLF4-associated regulatory signatures at the level of membrane protein\u2013coding genes in response to LSS.<\/jats:p>","DOI":"10.1021\/acs.jproteome.5c01254","type":"journal-article","created":{"date-parts":[[2026,7,6]],"date-time":"2026-07-06T14:08:44Z","timestamp":1783346924000},"page":"3987-4004","source":"Crossref","is-referenced-by-count":0,"title":["Proteomic\nProfiling of Endothelial Cells Under Laminar\nShear Stress Confirms the Importance of KLF4 in the Regulation of\nMembrane Protein Expression Compared to Oscillatory Flow"],"prefix":"10.1021","volume":"25","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-7143-5966","authenticated-orcid":true,"given":"Le\u0301a","family":"Re\u0301thore\u0301","sequence":"first","affiliation":[{"name":"Univ Angers, INSERM, CNRS, MITOVASC, Equipe CarME, SFR ICAT , ,","place":["Angers, France, F-49000"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Benjamin","family":"Barre\u0301","sequence":"additional","affiliation":[{"name":"Univ Angers, INSERM, CNRS, MITOVASC, Equipe CarME, SFR ICAT , ,","place":["Angers, France, F-49000"]},{"name":"Institut de Cance\u0301rologie de l\u2019Ouest (ICO), Prot\u2019ICO (ICO Proteomic Core Facility) , ,","place":["Angers, France, F-49055"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Anne-Laure","family":"Guihot","sequence":"additional","affiliation":[{"name":"Univ Angers, INSERM, CNRS, MITOVASC, Equipe CarME, SFR ICAT , ,","place":["Angers, France, F-49000"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Linda","family":"Grimaud","sequence":"additional","affiliation":[{"name":"Univ Angers, INSERM, CNRS, MITOVASC, Equipe CarME, SFR ICAT , ,","place":["Angers, France, F-49000"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Coralyne","family":"Proux","sequence":"additional","affiliation":[{"name":"Univ Angers, INSERM, CNRS, MITOVASC, Equipe CarME, SFR ICAT , ,","place":["Angers, France, F-49000"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Alice","family":"Boissard","sequence":"additional","affiliation":[{"name":"Institut de Cance\u0301rologie de l\u2019Ouest (ICO), Prot\u2019ICO (ICO Proteomic Core Facility) , ,","place":["Angers, France, F-49055"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ce\u0301cile","family":"Henry","sequence":"additional","affiliation":[{"name":"Institut de Cance\u0301rologie de l\u2019Ouest (ICO), Prot\u2019ICO (ICO Proteomic Core Facility) , ,","place":["Angers, France, F-49055"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Catherine","family":"Guette","sequence":"additional","affiliation":[{"name":"Institut de Cance\u0301rologie de l\u2019Ouest (ICO), Prot\u2019ICO (ICO Proteomic Core Facility) , ,","place":["Angers, France, F-49055"]},{"name":"Univ Angers, Nantes Universite\u0301, INSERM, CNRS, CRCI2NA , ,","place":["Angers, France, F-49000"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Franc\u0327ois","family":"Guillonneau","sequence":"additional","affiliation":[{"name":"Institut de Cance\u0301rologie de l\u2019Ouest (ICO), Prot\u2019ICO (ICO Proteomic Core Facility) , ,","place":["Angers, France, F-49055"]},{"name":"Univ Angers, Nantes Universite\u0301, INSERM, CNRS, CRCI2NA , ,","place":["Angers, France, F-49000"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Daniel","family":"Henrion","sequence":"additional","affiliation":[{"name":"Univ Angers, INSERM, CNRS, MITOVASC, Equipe CarME, SFR ICAT , ,","place":["Angers, France, F-49000"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Christian","family":"Legros","sequence":"additional","affiliation":[{"name":"Univ Angers, INSERM, CNRS, MITOVASC, Equipe CarME, SFR ICAT , ,","place":["Angers, France, F-49000"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0248-1111","authenticated-orcid":true,"given":"Claire","family":"Legendre","sequence":"additional","affiliation":[{"name":"Univ Angers, INSERM, CNRS, MITOVASC, Equipe CarME, SFR ICAT , ,","place":["Angers, France, F-49000"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"316","published-online":{"date-parts":[[2026,7,6]]},"reference":[{"issue":"10","key":"2026080709075693600_cit1","doi-asserted-by":"publisher","first-page":"1285","DOI":"10.1161\/CIRCULATIONAHA.106.652859","article-title":"Endothelial Function and Dysfunction:\nTesting and Clinical Relevance","volume":"115","author":"Deanfield","year":"2007","journal-title":"Circulation"},{"issue":"5","key":"2026080709075693600_cit2","doi-asserted-by":"publisher","first-page":"1425","DOI":"10.1161\/HYPERTENSIONAHA.120.16128","article-title":"\u201cMan Is as\nOld as His Arteries\u201d Taken\nLiterally: In Search of the Best Metric","volume":"76","author":"Weber","year":"2020","journal-title":"Hypertension"},{"key":"2026080709075693600_cit3","doi-asserted-by":"publisher","first-page":"738","DOI":"10.1038\/s41569-023-00883-1","article-title":"Flow-Induced\nReprogramming of Endothelial Cells in Atherosclerosis","volume":"20","author":"Tamargo","year":"2023","journal-title":"Nat. Rev. Cardiol."},{"issue":"2","key":"2026080709075693600_cit4","doi-asserted-by":"publisher","first-page":"1247","DOI":"10.1152\/physrev.00053.2021","article-title":"Vascular Mechanotransduction","volume":"103","author":"Davis","year":"2023","journal-title":"Physiol. Rev."},{"key":"2026080709075693600_cit5","doi-asserted-by":"publisher","first-page":"524","DOI":"10.3389\/fphys.2018.00524","article-title":"Endothelial\nMechanotransduction, Redox Signaling and the Regulation of Vascular\nInflammatory Pathways","volume":"9","author":"Chatterjee","year":"2018","journal-title":"Front. Physiol."},{"key":"2026080709075693600_cit6","doi-asserted-by":"publisher","first-page":"65","DOI":"10.1159\/000512618","article-title":"Polarized\nProteins\nin Endothelium and Their Contribution to Function","volume":"58","author":"Wolpe","year":"2021","journal-title":"J. Vasc. Res."},{"issue":"7","key":"2026080709075693600_cit7","doi-asserted-by":"publisher","first-page":"752","DOI":"10.1161\/CIRCRESAHA.124.325685","article-title":"Biophysical and Biochemical\nRoles of Shear Stress on Endothelium: A Revisit and New Insights","volume":"136","author":"Cheng","year":"2025","journal-title":"Circ. Res."},{"issue":"13","key":"2026080709075693600_cit8","doi-asserted-by":"publisher","first-page":"1745","DOI":"10.1091\/mbc.e16-06-0389","article-title":"Flow-Induced Endothelial\nCell Alignment Requires the\nRhoGEF Trio as a Scaffold Protein to Polarize Active Rac1 Distribution","volume":"28","author":"Kroon","year":"2017","journal-title":"MBoC"},{"issue":"4","key":"2026080709075693600_cit9","doi-asserted-by":"publisher","first-page":"253","DOI":"10.1016\/j.tips.2019.02.004","article-title":"Targeting Mechanosensitive\nTranscription Factors in Atherosclerosis","volume":"40","author":"Niu","year":"2019","journal-title":"Trends\nPharmacol. Sci."},{"issue":"18","key":"2026080709075693600_cit10","doi-asserted-by":"publisher","first-page":"13769","DOI":"10.1074\/jbc.M700078200","article-title":"Kruppel-like\nFactor 4 Regulates Endothelial Inflammation","volume":"282","author":"Hamik","year":"2007","journal-title":"J. Biol. Chem."},{"issue":"1","key":"2026080709075693600_cit11","doi-asserted-by":"publisher","first-page":"49","DOI":"10.1172\/JCI24787","article-title":"Integration\nof Flow-dependent Endothelial Phenotypes by Kruppel-like Factor 2","volume":"116","author":"Parmar","year":"2005","journal-title":"J. Clin. Invest."},{"issue":"5","key":"2026080709075693600_cit12","doi-asserted-by":"publisher","first-page":"1689","DOI":"10.1182\/blood-2002-01-0046","article-title":"Prolonged Fluid Shear Stress Induces a Distinct Set\nof Endothelial Cell Genes, Most Specifically Lung Kru\u0308ppel-like\nFactor (KLF2)","volume":"100","author":"Dekker","year":"2002","journal-title":"Blood"},{"issue":"1","key":"2026080709075693600_cit13","doi-asserted-by":"publisher","first-page":"4795","DOI":"10.1038\/s41598-022-08645-8","article-title":"Shear Stress Switches the Association\nof Endothelial Enhancers from ETV\/ETS to KLF Transcription Factor\nBinding Sites","volume":"12","author":"Tsaryk","year":"2022","journal-title":"Sci. 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Physiol."},{"issue":"7","key":"2026080709075693600_cit20","doi-asserted-by":"publisher","first-page":"1179","DOI":"10.1074\/mcp.RA120.001964","article-title":"Flow-Induced\nReorganization of Laminin-Integrin Networks\nWithin the Endothelial Basement Membrane Uncovered by Proteomics","volume":"19","author":"Be\u0301guin","year":"2020","journal-title":"Mol. Cell. Proteomics"},{"issue":"5","key":"2026080709075693600_cit21","doi-asserted-by":"publisher","first-page":"551","DOI":"10.1165\/rcmb.2022-0340OC","article-title":"Shear Stress\nMarkedly Alters the Proteomic Response to Hypoxia in Human Pulmonary\nEndothelial Cells","volume":"68","author":"Kostyunina","year":"2023","journal-title":"Am. J. Respir. Cell Mol.\nBiol."},{"issue":"8","key":"2026080709075693600_cit22","doi-asserted-by":"publisher","first-page":"1012","DOI":"10.1093\/clinchem\/hvaa125","article-title":"The Digital\nMIQE Guidelines Update: Minimum Information for Publication of Quantitative\nDigital PCR Experiments for 2020","volume":"66","author":"Huggett","year":"2020","journal-title":"Clin Chem."},{"issue":"4","key":"2026080709075693600_cit23","doi-asserted-by":"publisher","first-page":"611","DOI":"10.1373\/clinchem.2008.112797","article-title":"The MIQE Guidelines: Minimum Information for Publication\nof Quantitative Real-Time PCR Experiments","volume":"55","author":"Bustin","year":"2009","journal-title":"Clin\nChem."},{"issue":"5","key":"2026080709075693600_cit24","doi-asserted-by":"publisher","first-page":"430","DOI":"10.1038\/nmeth.1584","article-title":"mProphet: Automated Data Processing and Statistical\nValidation for Large-Scale SRM Experiments","volume":"8","author":"Reiter","year":"2011","journal-title":"Nat. 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Protoc."},{"key":"2026080709075693600_cit27","doi-asserted-by":"publisher","first-page":"49","DOI":"10.1016\/j.ddtec.2021.06.006","article-title":"Data-independent Acquisition (DIA):\nAn Emerging Proteomics Technology for Analysis of Drug-Metabolizing\nEnzymes and Transporters","volume":"39","author":"Li","year":"2021","journal-title":"Drug Discov Today\nTechnol."},{"issue":"8","key":"2026080709075693600_cit28","doi-asserted-by":"publisher","first-page":"1179","DOI":"10.1136\/annrheumdis-2021-221867","article-title":"Kru\u0308ppel-like\nFactors-4 and \u22122 Are Important Regulators of Joint Tissue Cells\nand Protect against Tissue Destruction and Inflammation in Osteoarthritis","volume":"81","author":"Kawata","year":"2022","journal-title":"Ann. Rheum. Dis."},{"issue":"14","key":"2026080709075693600_cit29","doi-asserted-by":"publisher","first-page":"1315","DOI":"10.1161\/CIRCULATIONAHA.117.027462","article-title":"KLF4 Regulation\nof Ch25h and LXR Mitigates Atherosclerosis Susceptibility","volume":"136","author":"Li","year":"2017","journal-title":"Circulation"},{"issue":"D1","key":"2026080709075693600_cit30","doi-asserted-by":"publisher","first-page":"D87","DOI":"10.1093\/nar\/gkz1001","article-title":"JASPAR 2020: Update of the Open-Access\nDatabase of\nTranscription Factor Binding Profiles","volume":"48","author":"Fornes","year":"2019","journal-title":"Nucleic\nAcids Res."},{"issue":"2","key":"2026080709075693600_cit31","doi-asserted-by":"publisher","first-page":"026122","DOI":"10.1063\/5.0263936","article-title":"Pulsatile\nLow Shear Stress Increases\nSusceptibility to Endothelial Inflammation via Upregulation of IFT\nand Activation of YAP","volume":"9","author":"Hou","year":"2025","journal-title":"APL Bioeng."},{"issue":"1","key":"2026080709075693600_cit32","doi-asserted-by":"publisher","first-page":"114410","DOI":"10.1016\/j.yexcr.2025.114410","article-title":"Low Shear\nStress Induces Vascular Endothelial Cells Apoptosis via miR-330\/SOD2\/HSP70\nSignaling Pathway","volume":"445","author":"Li","year":"2025","journal-title":"Exp. Cell Res."},{"issue":"1","key":"2026080709075693600_cit33","doi-asserted-by":"publisher","first-page":"39553","DOI":"10.1038\/srep39553","article-title":"PI16 Is\na Shear\nStress and Inflammation-Regulated Inhibitor of MMP2","volume":"6","author":"Hazell","year":"2016","journal-title":"Sci. Rep"},{"key":"2026080709075693600_cit34","doi-asserted-by":"publisher","first-page":"189","DOI":"10.1016\/B978-0-12-405943-6.00005-1","article-title":"Id Proteins:\nSmall Molecules, Mighty\nRegulators","volume":"110","author":"Ling","year":"2014","journal-title":"Curr. Top. Dev. Biol."},{"issue":"16","key":"2026080709075693600_cit35","doi-asserted-by":"publisher","first-page":"5623","DOI":"10.3390\/ijms21165623","article-title":"Ephs and Ephrins\nin Adult Endothelial Biology","volume":"21","author":"Vreeken","year":"2020","journal-title":"Int. J. Mol. Sci."},{"issue":"D1","key":"2026080709075693600_cit36","doi-asserted-by":"publisher","first-page":"D174","DOI":"10.1093\/nar\/gkad1059","article-title":"JASPAR 2024:20th\nAnniversary of the Open-Access Database of Transcription Factor Binding\nProfiles","volume":"52","author":"Rauluseviciute","year":"2024","journal-title":"Nucleic Acids Res."},{"issue":"4","key":"2026080709075693600_cit37","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.2023236118","article-title":"Longitudinal\nShear Stress Response in Human Endothelial Cells to Atheroprone and\nAtheroprotective Conditions","volume":"118","author":"Maurya","year":"2021","journal-title":"Proc. Natl. Acad.\nSci. U. S. A."},{"issue":"11","key":"2026080709075693600_cit38","doi-asserted-by":"publisher","DOI":"10.1096\/fj.202401558RR","article-title":"A Novel Function of NaV Channel \u03923 Subunit in\nEndothelial Cell Alignment Through Autophagy Modulation","volume":"39","author":"Re\u0301thore\u0301","year":"2025","journal-title":"FASEB J."},{"key":"2026080709075693600_cit39","doi-asserted-by":"publisher","first-page":"133","DOI":"10.1186\/s13059-019-1749-5","article-title":"Integrative Analysis of Vascular\nEndothelial Cell Genomic Features Identifies AIDA as a Coronary Artery\nDisease Candidate Gene","volume":"20","author":"Lalonde","year":"2019","journal-title":"Genome Biol."},{"issue":"4","key":"2026080709075693600_cit40","doi-asserted-by":"publisher","first-page":"2536","DOI":"10.1074\/jbc.M110.180489","article-title":"Endothelial\nArgininosuccinate\nSynthetase 1 Regulates Nitric Oxide Production and Monocyte Adhesion\nunder Static and Laminar Shear Stress Conditions","volume":"286","author":"Mun","year":"2011","journal-title":"J. Biol. Chem."},{"issue":"13","key":"2026080709075693600_cit41","doi-asserted-by":"publisher","first-page":"1217","DOI":"10.1161\/CIRCULATIONAHA.116.026361","article-title":"Disturbed Flow Promotes\nArterial Stiffening Through Thrombospondin-1","volume":"136","author":"Kim","year":"2017","journal-title":"Circulation"},{"issue":"1","key":"2026080709075693600_cit42","doi-asserted-by":"publisher","first-page":"7394","DOI":"10.1038\/s41467-025-62857-w","article-title":"Molecular\nBasis of Human Taurine Transporter Uptake and Inhibition","volume":"16","author":"Du","year":"2025","journal-title":"Nat. Commun."},{"key":"2026080709075693600_cit43","doi-asserted-by":"publisher","first-page":"823","DOI":"10.3389\/fimmu.2019.00823","article-title":"Pentraxin 3 in Cardiovascular\nDisease","volume":"10","author":"Ristagno","year":"2019","journal-title":"Front. Immunol."},{"issue":"7","key":"2026080709075693600_cit44","doi-asserted-by":"publisher","DOI":"10.1161\/CIRCRESAHA.125.327592","article-title":"FOXO1 Integrates Endothelial Hemodynamic, Inflammatory,\nand Metabolic Pathways in Atherosclerosis","volume":"138","author":"Deng","year":"2026","journal-title":"Circ.\nRes."},{"issue":"18","key":"2026080709075693600_cit45","doi-asserted-by":"publisher","DOI":"10.1172\/jci.insight.159286","article-title":"Flow Pattern\u2013dependent\nMitochondrial Dynamics Regulates the Metabolic Profile and Inflammatory\nState of Endothelial Cells","volume":"7","author":"Hong","year":"2022","journal-title":"JCI Insight"},{"issue":"16","key":"2026080709075693600_cit46","doi-asserted-by":"publisher","first-page":"8653","DOI":"10.1074\/jbc.M115.713149","article-title":"Differential\nRoles of Protein Complexes NOX1-NOXO1\nand NOX2-P47phox in Mediating Endothelial Redox Responses to Oscillatory\nand Unidirectional Laminar Shear Stress","volume":"291","author":"Siu","year":"2016","journal-title":"J.\nBiol. Chem."},{"issue":"2","key":"2026080709075693600_cit47","doi-asserted-by":"publisher","first-page":"429","DOI":"10.1083\/jcb.200210135","article-title":"Shear Stress\u2013Induced Endothelial Cell Polarization\nIs Mediated by Rho and Rac but Not Cdc42 or PI 3-Kinases","volume":"161","author":"Wojciak-Stothard","year":"2003","journal-title":"J. Cell Biol."},{"issue":"4","key":"2026080709075693600_cit48","doi-asserted-by":"publisher","first-page":"142","DOI":"10.1002\/cm.21679","article-title":"Posttranslational Modifications of the Cytoskeleton","volume":"78","author":"MacTaggart","year":"2021","journal-title":"Cytoskeleton (Hoboken)"},{"key":"2026080709075693600_cit49","doi-asserted-by":"publisher","first-page":"925923","DOI":"10.3389\/fcvm.2022.925923","article-title":"Mechanisms of Oxidized LDL-Mediated\nEndothelial Dysfunction and Its Consequences for the Development of\nAtherosclerosis","volume":"9","author":"Jiang","year":"2022","journal-title":"Front. Cardiovasc. 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