{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,23]],"date-time":"2026-10-23T19:15:52Z","timestamp":1792782952147,"version":"4.5.0"},"reference-count":86,"publisher":"Springer Science and Business Media LLC","issue":"4","license":[{"start":{"date-parts":[[2026,10,15]],"date-time":"2026-10-15T00:00:00Z","timestamp":1792022400000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0"},{"start":{"date-parts":[[2026,10,15]],"date-time":"2026-10-15T00:00:00Z","timestamp":1792022400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0"}],"funder":[{"DOI":"10.13039\/100000968","name":"American Heart Association","doi-asserted-by":"publisher","id":[{"id":"10.13039\/100000968","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"publisher","id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000057","name":"National Institute of General Medical Sciences","doi-asserted-by":"publisher","award":["R35GM158040"],"award-info":[{"award-number":["R35GM158040"]}],"id":[{"id":"10.13039\/100000057","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000057","name":"National Institute of General Medical Sciences","doi-asserted-by":"publisher","award":["R35GM142944"],"award-info":[{"award-number":["R35GM142944"]}],"id":[{"id":"10.13039\/100000057","id-type":"DOI","asserted-by":"publisher"}]},{"name":"National Institutes of General Medical Sciences","award":["R35GM130312"],"award-info":[{"award-number":["R35GM130312"]}],"id":[{"id":"https:\/\/ror.org\/04q48ey07","id-type":"ROR","asserted-by":"crossref"}]},{"DOI":"10.13039\/100000050","name":"National Heart, Lung, and Blood Institute","doi-asserted-by":"publisher","award":["R35HL139950"],"award-info":[{"award-number":["R35HL139950"]}],"id":[{"id":"10.13039\/100000050","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Angiogenesis"],"DOI":"10.1007\/s10456-026-10098-w","type":"journal-article","created":{"date-parts":[[2026,10,15]],"date-time":"2026-10-15T20:51:24Z","timestamp":1792097484000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Nuclear SUN2 coordinates endothelial cell-matrix interactions to regulate blood vessel homeostasis and barrier function"],"prefix":"10.1007","volume":"29","author":[{"given":"Pauline","family":"Bougaran","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Danielle B.","family":"Buglak","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Alexandra","family":"Neal","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Mitesh","family":"Rathod","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Michaelanthony","family":"Gore","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Max A.","family":"Hockenberry","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Aryan A.","family":"Amin","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Natalie","family":"Tanke","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Morgan","family":"Oatley","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Wesley R.","family":"Legant","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ziqing","family":"Liu","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"James E.","family":"Bear","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"William J.","family":"Polacheck","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Victoria L.","family":"Bautch","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2026,10,15]]},"reference":[{"key":"10098_CR1","doi-asserted-by":"publisher","DOI":"10.3389\/fphys.2020.00861","author":"E Roux","year":"2020","unstructured":"Roux E, Bougaran P, Dufourcq P, Couffinhal T (2020) Fluid Shear Stress Sensing by the Endothelial Layer. Front Physiol. https:\/\/doi.org\/10.3389\/fphys.2020.00861","journal-title":"Front Physiol"},{"key":"10098_CR2","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1146\/annurev-bioeng-092419-060810","volume":"23","author":"JD Humphrey","year":"2021","unstructured":"Humphrey JD, Schwartz MA (2021) Vascular mechanobiology: homeostasis, adaptation, and disease. Annu Rev Biomed Eng 23:1\u201327. https:\/\/doi.org\/10.1146\/annurev-bioeng-092419-060810","journal-title":"Annu Rev Biomed Eng"},{"key":"10098_CR3","doi-asserted-by":"publisher","first-page":"882","DOI":"10.1161\/ATVBAHA.125.322557","volume":"45","author":"H Deng","year":"2025","unstructured":"Deng H, Eichmann A, Schwartz MA (2025) Fluid shear stress-regulated vascular remodeling: past, present, and future. Arterioscler Thromb Vasc Biol 45:882\u2013900. https:\/\/doi.org\/10.1161\/ATVBAHA.125.322557","journal-title":"Arterioscler Thromb Vasc Biol"},{"key":"10098_CR4","doi-asserted-by":"publisher","DOI":"10.3389\/fphys.2020.00684","volume":"11","author":"E Gordon","year":"2020","unstructured":"Gordon E, Schimmel L, Frye M (2020) The importance of mechanical forces for in vitro endothelial cell biology. Front Physiol 11:684. https:\/\/doi.org\/10.3389\/fphys.2020.00684","journal-title":"Front Physiol"},{"key":"10098_CR5","doi-asserted-by":"publisher","DOI":"10.1038\/s41392-023-01501-9","volume":"8","author":"X Di","year":"2023","unstructured":"Di X, Gao X, Peng L et al (2023) Cellular mechanotransduction in health and diseases: from molecular mechanism to therapeutic targets. Sig Transduct Target Ther 8:282. https:\/\/doi.org\/10.1038\/s41392-023-01501-9","journal-title":"Sig Transduct Target Ther"},{"key":"10098_CR6","doi-asserted-by":"publisher","first-page":"4639","DOI":"10.1093\/emboj\/20.17.4639","volume":"20","author":"E Tzima","year":"2001","unstructured":"Tzima E, del Pozo MA, Shattil SJ et al (2001) Activation of integrins in endothelial cells by fluid shear stress mediates Rho-dependent cytoskeletal alignment. EMBO J 20:4639\u20134647. https:\/\/doi.org\/10.1093\/emboj\/20.17.4639","journal-title":"EMBO J"},{"key":"10098_CR7","doi-asserted-by":"publisher","first-page":"426","DOI":"10.1038\/nature03952","volume":"437","author":"E Tzima","year":"2005","unstructured":"Tzima E, Irani-Tehrani M, Kiosses WB et al (2005) A mechanosensory complex that mediates the endothelial cell response to fluid shear stress. Nature 437:426\u2013431. https:\/\/doi.org\/10.1038\/nature03952","journal-title":"Nature"},{"key":"10098_CR8","doi-asserted-by":"publisher","DOI":"10.1242\/jcs.229542","author":"I Xanthis","year":"2019","unstructured":"Xanthis I, Souilhol C, Serbanovic-Canic J et al (2019) \u03b21 integrin is a sensor of blood flow direction. J Cell Sci. https:\/\/doi.org\/10.1242\/jcs.229542","journal-title":"J Cell Sci"},{"key":"10098_CR9","doi-asserted-by":"publisher","first-page":"217","DOI":"10.1002\/cm.21652","volume":"78","author":"K Tanaka","year":"2021","unstructured":"Tanaka K, Joshi D, Timalsina S, Schwartz MA (2021) Early events in endothelial flow sensing. Cytoskeleton 78:217\u2013231. https:\/\/doi.org\/10.1002\/cm.21652","journal-title":"Cytoskeleton"},{"key":"10098_CR10","doi-asserted-by":"publisher","first-page":"443","DOI":"10.1146\/annurev-bioeng-060418-052139","volume":"21","author":"M Maurer","year":"2019","unstructured":"Maurer M, Lammerding J (2019) The driving force: nuclear mechanotransduction in cellular function, fate, and disease. Annu Rev Biomed Eng 21:443\u2013468. https:\/\/doi.org\/10.1146\/annurev-bioeng-060418-052139","journal-title":"Annu Rev Biomed Eng"},{"key":"10098_CR11","doi-asserted-by":"publisher","first-page":"905927","DOI":"10.3389\/fcell.2022.905927","volume":"10","author":"J Salvador","year":"2022","unstructured":"Salvador J, Iruela-Arispe ML (2022) Nuclear mechanosensation and Mechanotransduction in Vascular Cells. Front Cell Dev Biol 10:905927","journal-title":"Front Cell Dev Biol"},{"key":"10098_CR12","doi-asserted-by":"publisher","DOI":"10.3389\/fphys.2024.1411995","author":"P Bougaran","year":"2024","unstructured":"Bougaran P, Bautch VL (2024) Life at the crossroads: the nuclear LINC complex and vascular mechanotransduction. Front Physiol. https:\/\/doi.org\/10.3389\/fphys.2024.1411995","journal-title":"Front Physiol"},{"key":"10098_CR13","doi-asserted-by":"publisher","first-page":"1007","DOI":"10.1242\/bio.20134622","volume":"2","author":"E Tkachenko","year":"2013","unstructured":"Tkachenko E, Gutierrez E, Saikin SK et al (2013) The nucleus of endothelial cell as a sensor of blood flow direction. Biol Open 2:1007\u20131012. https:\/\/doi.org\/10.1242\/bio.20134622","journal-title":"Biol Open"},{"key":"10098_CR14","doi-asserted-by":"publisher","first-page":"519","DOI":"10.1152\/physrev.1995.75.3.519","volume":"75","author":"PF Davies","year":"1995","unstructured":"Davies PF (1995) Flow-mediated endothelial mechanotransduction. Physiol Rev 75:519\u2013560. https:\/\/doi.org\/10.1152\/physrev.1995.75.3.519","journal-title":"Physiol Rev"},{"key":"10098_CR15","doi-asserted-by":"publisher","first-page":"939","DOI":"10.1161\/01.RES.0000216595.15868.55","volume":"98","author":"S McCue","year":"2006","unstructured":"McCue S, Dajnowiec D, Xu F et al (2006) Shear stress regulates forward and reverse planar cell polarity of vascular endothelium in vivo and in vitro. Circ Res 98:939\u2013946. https:\/\/doi.org\/10.1161\/01.RES.0000216595.15868.55","journal-title":"Circ Res"},{"key":"10098_CR16","doi-asserted-by":"publisher","first-page":"376","DOI":"10.1038\/ncb2927","volume":"16","author":"C Guilluy","year":"2014","unstructured":"Guilluy C, Osborne LD, Van Landeghem L et al (2014) Isolated nuclei adapt to force and reveal a mechanotransduction pathway within the nucleus. Nat Cell Biol 16:376\u2013381. https:\/\/doi.org\/10.1038\/ncb2927","journal-title":"Nat Cell Biol"},{"key":"10098_CR17","doi-asserted-by":"publisher","first-page":"895","DOI":"10.1083\/jcb.201706097","volume":"217","author":"DM Graham","year":"2018","unstructured":"Graham DM, Andersen T, Sharek L et al (2018) Enucleated cells reveal differential roles of the nucleus in cell migration, polarity, and mechanotransduction. J Cell Biol 217:895\u2013914. https:\/\/doi.org\/10.1083\/jcb.201706097","journal-title":"J Cell Biol"},{"key":"10098_CR18","doi-asserted-by":"publisher","first-page":"75","DOI":"10.1038\/nrm2594","volume":"10","author":"N Wang","year":"2009","unstructured":"Wang N, Tytell JD, Ingber DE (2009) Mechanotransduction at a distance: mechanically coupling the extracellular matrix with the nucleus. Nat Rev Mol Cell Biol 10:75\u201382. https:\/\/doi.org\/10.1038\/nrm2594","journal-title":"Nat Rev Mol Cell Biol"},{"key":"10098_CR19","doi-asserted-by":"publisher","first-page":"34","DOI":"10.1016\/j.molmed.2025.03.005","volume":"32","author":"I Benedicto","year":"2026","unstructured":"Benedicto I, Hamczyk MR, Dorado B, Andr\u00e9s V (2026) Vascular cell types in progeria: victims or villains? Trends Mol Med 32:34\u201348. https:\/\/doi.org\/10.1016\/j.molmed.2025.03.005","journal-title":"Trends Mol Med"},{"key":"10098_CR20","doi-asserted-by":"publisher","first-page":"295","DOI":"10.1515\/hsz-2014-0267","volume":"396","author":"DI Kim","year":"2015","unstructured":"Kim DI, Kc B, Roux KJ (2015) Making the LINC: SUN and KASH protein interactions. Biol Chem 396:295\u2013310. https:\/\/doi.org\/10.1515\/hsz-2014-0267","journal-title":"Biol Chem"},{"key":"10098_CR21","doi-asserted-by":"publisher","first-page":"208","DOI":"10.1186\/s12964-024-01593-y","volume":"22","author":"Z Zi-yi","year":"2024","unstructured":"Zi-yi Z, Qin Q, Fei Z et al (2024) Nesprin proteins: bridging nuclear envelope dynamics to muscular dysfunction. Cell Commun Signal 22:208. https:\/\/doi.org\/10.1186\/s12964-024-01593-y","journal-title":"Cell Commun Signal"},{"key":"10098_CR22","doi-asserted-by":"publisher","first-page":"218","DOI":"10.1242\/dev.114769","volume":"142","author":"MH Wilson","year":"2015","unstructured":"Wilson MH, Holzbaur ELF (2015) Nesprins anchor kinesin-1 motors to the nucleus to drive nuclear distribution in muscle cells. Development 142:218\u2013228. https:\/\/doi.org\/10.1242\/dev.114769","journal-title":"Development"},{"key":"10098_CR23","doi-asserted-by":"publisher","first-page":"1035","DOI":"10.1016\/j.cell.2012.03.046","volume":"149","author":"BA Sosa","year":"2012","unstructured":"Sosa BA, Rothballer A, Kutay U, Schwartz TU (2012) LINC complexes form by binding of three KASH peptides to domain interfaces of trimeric SUN proteins. Cell 149:1035\u20131047. https:\/\/doi.org\/10.1016\/j.cell.2012.03.046","journal-title":"Cell"},{"key":"10098_CR24","doi-asserted-by":"publisher","first-page":"1190","DOI":"10.1016\/j.bpj.2018.01.015","volume":"114","author":"Z Jahed","year":"2018","unstructured":"Jahed Z, Fadavi D, Vu UT et al (2018) Molecular insights into the mechanisms of SUN1 oligomerization in the nuclear envelope. Biophys J 114:1190\u20131203. https:\/\/doi.org\/10.1016\/j.bpj.2018.01.015","journal-title":"Biophys J"},{"key":"10098_CR25","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0197621","volume":"13","author":"CK May","year":"2018","unstructured":"May CK, Carroll CW (2018) Differential incorporation of SUN-domain proteins into LINC complexes is coupled to gene expression. PLoS ONE 13:e0197621. https:\/\/doi.org\/10.1371\/journal.pone.0197621","journal-title":"PLoS ONE"},{"key":"10098_CR26","doi-asserted-by":"publisher","DOI":"10.3389\/fphys.2024.1464678","volume":"15","author":"KL Shores","year":"2024","unstructured":"Shores KL, Truskey GA (2024) Mechanotransduction of the vasculature in Hutchinson-Gilford progeria syndrome. Front Physiol 15:1464678. https:\/\/doi.org\/10.3389\/fphys.2024.1464678","journal-title":"Front Physiol"},{"key":"10098_CR27","doi-asserted-by":"publisher","first-page":"115","DOI":"10.1016\/j.bpj.2010.04.011","volume":"99","author":"TJ Chancellor","year":"2010","unstructured":"Chancellor TJ, Lee J, Thodeti CK, Lele T (2010) Actomyosin tension exerted on the nucleus through Nesprin-1 connections influences endothelial cell adhesion, migration, and cyclic strain-induced reorientation. Biophys J 99:115\u2013123. https:\/\/doi.org\/10.1016\/j.bpj.2010.04.011","journal-title":"Biophys J"},{"key":"10098_CR28","doi-asserted-by":"publisher","first-page":"647","DOI":"10.3389\/fbioe.2020.00647","volume":"8","author":"F Yang","year":"2020","unstructured":"Yang F, Zhang Y, Zhu J et al (2020) Laminar flow Protects Vascular Endothelial Tight Junctions and Barrier Function via Maintaining the Expression of Long Non-coding RNA MALAT1. Front Bioeng Biotechnol 8:647","journal-title":"Front Bioeng Biotechnol"},{"key":"10098_CR29","doi-asserted-by":"publisher","first-page":"565","DOI":"10.1016\/j.cell.2012.01.059","volume":"149","author":"C-Y Chen","year":"2012","unstructured":"Chen C-Y, Chi Y-H, Mutalif RA et al (2012) Accumulation of the inner nuclear envelope protein Sun1 is pathogenic in progeric and dystrophic laminopathies. Cell 149:565\u2013577. https:\/\/doi.org\/10.1016\/j.cell.2012.01.059","journal-title":"Cell"},{"key":"10098_CR30","doi-asserted-by":"publisher","first-page":"531","DOI":"10.1172\/JCI121297","volume":"129","author":"S Osmanagic-Myers","year":"2019","unstructured":"Osmanagic-Myers S, Kiss A, Manakanatas C et al (2019) Endothelial progerin expression causes cardiovascular pathology through an impaired mechanoresponse. J Clin Invest 129:531\u2013545. https:\/\/doi.org\/10.1172\/JCI121297","journal-title":"J Clin Invest"},{"key":"10098_CR31","doi-asserted-by":"publisher","DOI":"10.1016\/j.celrep.2023.112534","volume":"42","author":"S Vidak","year":"2023","unstructured":"Vidak S, Serebryannyy LA, Pegoraro G, Misteli T (2023) Activation of endoplasmic reticulum stress in premature aging via the inner nuclear membrane protein SUN2. Cell Rep 42:112534. https:\/\/doi.org\/10.1016\/j.celrep.2023.112534","journal-title":"Cell Rep"},{"key":"10098_CR32","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.83652","volume":"12","author":"DB Buglak","year":"2023","unstructured":"Buglak DB, Bougaran P, Kulikauskas MR et al (2023) Nuclear SUN1 stabilizes endothelial cell junctions via microtubules to regulate blood vessel formation. Elife 12:e83652. https:\/\/doi.org\/10.7554\/eLife.83652","journal-title":"Elife"},{"key":"10098_CR33","doi-asserted-by":"publisher","first-page":"5680","DOI":"10.1182\/blood-2008-08-174508","volume":"113","author":"I S\u00f6rensen","year":"2009","unstructured":"S\u00f6rensen I, Adams RH, Gossler A (2009) DLL1-mediated Notch activation regulates endothelial identity in mouse fetal arteries. Blood 113:5680\u20135688. https:\/\/doi.org\/10.1182\/blood-2008-08-174508","journal-title":"Blood"},{"key":"10098_CR34","doi-asserted-by":"publisher","first-page":"183","DOI":"10.1038\/s41586-022-05027-y","volume":"609","author":"MWG Schneider","year":"2022","unstructured":"Schneider MWG, Gibson BA, Otsuka S et al (2022) A mitotic chromatin phase transition prevents perforation by microtubules. Nature 609:183\u2013190. https:\/\/doi.org\/10.1038\/s41586-022-05027-y","journal-title":"Nature"},{"key":"10098_CR35","doi-asserted-by":"publisher","first-page":"263","DOI":"10.1038\/nature09198","volume":"466","author":"C Grashoff","year":"2010","unstructured":"Grashoff C, Hoffman BD, Brenner MD et al (2010) Measuring mechanical tension across vinculin reveals regulation of focal adhesion dynamics. Nature 466:263\u2013266. https:\/\/doi.org\/10.1038\/nature09198","journal-title":"Nature"},{"key":"10098_CR36","doi-asserted-by":"publisher","first-page":"1265","DOI":"10.1161\/ATVBAHA.124.320671","volume":"44","author":"NT Tanke","year":"2024","unstructured":"Tanke NT, Liu Z, Gore MT et al (2024) Endothelial cell flow-mediated quiescence is temporally regulated and utilizes the cell cycle inhibitor p27. Arterioscler Thromb Vasc Biol 44:1265\u20131282. https:\/\/doi.org\/10.1161\/ATVBAHA.124.320671","journal-title":"Arterioscler Thromb Vasc Biol"},{"key":"10098_CR37","doi-asserted-by":"publisher","DOI":"10.1002\/smll.202307901","volume":"20","author":"ML Rathod","year":"2024","unstructured":"Rathod ML, Aw WY, Huang S et al (2024) Donor-derived engineered microvessels for cardiovascular risk stratification of patients with kidney failure. Small 20:2307901. https:\/\/doi.org\/10.1002\/smll.202307901","journal-title":"Small"},{"key":"10098_CR38","doi-asserted-by":"publisher","first-page":"1425","DOI":"10.1038\/s41596-019-0144-8","volume":"14","author":"WJ Polacheck","year":"2019","unstructured":"Polacheck WJ, Kutys ML, Tefft JB, Chen CS (2019) Microfabricated blood vessels for modeling the vascular transport barrier. Nat Protoc 14:1425\u20131454. https:\/\/doi.org\/10.1038\/s41596-019-0144-8","journal-title":"Nat Protoc"},{"key":"10098_CR39","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.2509535122","volume":"122","author":"MA Hockenberry","year":"2025","unstructured":"Hockenberry MA, Ulmer AJ, Rapp JL et al (2025) Measurement of cellular traction forces during confined migration. Proc Natl Acad Sci U S A 122:e2509535122. https:\/\/doi.org\/10.1073\/pnas.2509535122","journal-title":"Proc Natl Acad Sci U S A"},{"key":"10098_CR40","doi-asserted-by":"publisher","DOI":"10.1083\/jcb.202111126","volume":"221","author":"ZT King","year":"2022","unstructured":"King ZT, Butler MT, Hockenberry MA et al (2022) Coro1B and Coro1C regulate lamellipodia dynamics and cell motility by tuning branched actin turnover. J Cell Biol 221:e202111126. https:\/\/doi.org\/10.1083\/jcb.202111126","journal-title":"J Cell Biol"},{"key":"10098_CR41","doi-asserted-by":"publisher","first-page":"653","DOI":"10.1038\/nmeth.3430","volume":"12","author":"SJ Han","year":"2015","unstructured":"Han SJ, Oak Y, Groisman A, Danuser G (2015) Traction microscopy to identify force modulation in subresolution adhesions. Nat Methods 12:653\u2013656. https:\/\/doi.org\/10.1038\/nmeth.3430","journal-title":"Nat Methods"},{"key":"10098_CR42","doi-asserted-by":"publisher","first-page":"C595","DOI":"10.1152\/ajpcell.00270.2001","volume":"282","author":"JP Butler","year":"2002","unstructured":"Butler JP, Toli\u0107-N\u00f8rrelykke IM, Fabry B, Fredberg JJ (2002) Traction fields, moments, and strain energy that cells exert on their surroundings. Am J Physiol Cell Physiol 282:C595\u2013C605. https:\/\/doi.org\/10.1152\/ajpcell.00270.2001","journal-title":"Am J Physiol Cell Physiol"},{"key":"10098_CR43","doi-asserted-by":"publisher","first-page":"254","DOI":"10.1038\/labinvest.2012.159","volume":"93","author":"O Dubrovskyi","year":"2013","unstructured":"Dubrovskyi O, Birukova AA, Birukov KG (2013) Measurement of local permeability at subcellular level in cell models of agonist- and ventilator-induced lung injury. Lab Invest 93:254\u2013263. https:\/\/doi.org\/10.1038\/labinvest.2012.159","journal-title":"Lab Invest"},{"key":"10098_CR44","doi-asserted-by":"publisher","DOI":"10.1242\/dev.201811","volume":"150","author":"MR Kulikauskas","year":"2023","unstructured":"Kulikauskas MR, Oatley M, Yu T et al (2023) Endothelial cell SMAD6 balances Alk1 function to regulate adherens junctions and hepatic vascular development. Development 150:dev201811. https:\/\/doi.org\/10.1242\/dev.201811","journal-title":"Development"},{"key":"10098_CR45","doi-asserted-by":"publisher","first-page":"15","DOI":"10.1093\/bioinformatics\/bts635","volume":"29","author":"A Dobin","year":"2013","unstructured":"Dobin A, Davis CA, Schlesinger F et al (2013) STAR: ultrafast universal RNA-seq aligner. Bioinformatics 29:15\u201321. https:\/\/doi.org\/10.1093\/bioinformatics\/bts635","journal-title":"Bioinformatics"},{"key":"10098_CR46","doi-asserted-by":"publisher","first-page":"2943","DOI":"10.1093\/bioinformatics\/btac166","volume":"38","author":"GH Putri","year":"2022","unstructured":"Putri GH, Anders S, Pyl PT et al (2022) Analysing high-throughput sequencing data in Python with HTSeq 2.0. Bioinformatics 38:2943\u20132945. https:\/\/doi.org\/10.1093\/bioinformatics\/btac166","journal-title":"Bioinformatics"},{"key":"10098_CR47","doi-asserted-by":"publisher","first-page":"550","DOI":"10.1186\/s13059-014-0550-8","volume":"15","author":"MI Love","year":"2014","unstructured":"Love MI, Huber W, Anders S (2014) Moderated estimation of fold change and dispersion for RNA-seq data with DESeq2. Genome Biol 15:550. https:\/\/doi.org\/10.1186\/s13059-014-0550-8","journal-title":"Genome Biol"},{"key":"10098_CR48","doi-asserted-by":"publisher","first-page":"676","DOI":"10.1038\/nmeth.2019","volume":"9","author":"J Schindelin","year":"2012","unstructured":"Schindelin J, Arganda-Carreras I, Frise E et al (2012) Fiji: an open-source platform for biological-image analysis. Nat Methods 9:676\u2013682. https:\/\/doi.org\/10.1038\/nmeth.2019","journal-title":"Nat Methods"},{"key":"10098_CR49","doi-asserted-by":"publisher","DOI":"10.1038\/s41598-022-26662-5","volume":"12","author":"A Mieremet","year":"2022","unstructured":"Mieremet A, van der Stoel M, Li S et al (2022) Endothelial dysfunction in Marfan syndrome mice is restored by resveratrol. Sci Rep 12:22504. https:\/\/doi.org\/10.1038\/s41598-022-26662-5","journal-title":"Sci Rep"},{"key":"10098_CR50","doi-asserted-by":"publisher","first-page":"1654","DOI":"10.1091\/mbc.E20-11-0698","volume":"32","author":"KB Denis","year":"2021","unstructured":"Denis KB, Cabe JI, Danielsson BE et al (2021) The LINC complex is required for endothelial cell adhesion and adaptation to shear stress and cyclic stretch. Mol Biol Cell 32:1654\u20131663. https:\/\/doi.org\/10.1091\/mbc.E20-11-0698","journal-title":"Mol Biol Cell"},{"key":"10098_CR51","doi-asserted-by":"publisher","first-page":"702","DOI":"10.1016\/0142-9612(90)90031-k","volume":"11","author":"MJ Levesque","year":"1990","unstructured":"Levesque MJ, Nerem RM, Sprague EA (1990) Vascular endothelial cell proliferation in culture and the influence of flow. Biomaterials 11:702\u2013707. https:\/\/doi.org\/10.1016\/0142-9612(90)90031-k","journal-title":"Biomaterials"},{"key":"10098_CR52","doi-asserted-by":"publisher","first-page":"203","DOI":"10.1093\/cvr\/cvm102","volume":"78","author":"VWM van Hinsbergh","year":"2008","unstructured":"van Hinsbergh VWM, Koolwijk P (2008) Endothelial sprouting and angiogenesis: matrix metalloproteinases in the lead. Cardiovasc Res 78:203\u2013212. https:\/\/doi.org\/10.1093\/cvr\/cvm102","journal-title":"Cardiovasc Res"},{"key":"10098_CR53","doi-asserted-by":"publisher","DOI":"10.1101\/cshperspect.a004903","volume":"4","author":"RO Hynes","year":"2012","unstructured":"Hynes RO, Naba A (2012) Overview of the matrisome\u2014an inventory of extracellular matrix constituents and functions. Cold Spring Harb Perspect Biol 4:a004903. https:\/\/doi.org\/10.1101\/cshperspect.a004903","journal-title":"Cold Spring Harb Perspect Biol"},{"key":"10098_CR54","doi-asserted-by":"publisher","DOI":"10.1186\/s13059-015-0676-3","volume":"16","author":"R Kelwick","year":"2015","unstructured":"Kelwick R, Desanlis I, Wheeler GN, Edwards DR (2015) The ADAMTS (a disintegrin and metalloproteinase with thrombospondin motifs) family. Genome Biol 16:113. https:\/\/doi.org\/10.1186\/s13059-015-0676-3","journal-title":"Genome Biol"},{"key":"10098_CR55","doi-asserted-by":"publisher","first-page":"126","DOI":"10.1161\/ATVBAHA.118.310956","volume":"39","author":"DM Milewicz","year":"2019","unstructured":"Milewicz DM, Ramirez F (2019) Therapies for thoracic aortic aneurysms and acute aortic dissections. Arterioscler Thromb Vasc Biol 39:126\u2013136. https:\/\/doi.org\/10.1161\/ATVBAHA.118.310956","journal-title":"Arterioscler Thromb Vasc Biol"},{"key":"10098_CR56","doi-asserted-by":"publisher","DOI":"10.3390\/biom12010128","volume":"12","author":"K Asano","year":"2022","unstructured":"Asano K, Cantalupo A, Sedes L, Ramirez F (2022) Pathophysiology and therapeutics of thoracic aortic aneurysm in Marfan syndrome. Biomolecules 12:128. https:\/\/doi.org\/10.3390\/biom12010128","journal-title":"Biomolecules"},{"key":"10098_CR57","doi-asserted-by":"publisher","first-page":"346","DOI":"10.1016\/j.actbio.2023.05.015","volume":"166","author":"EL Doherty","year":"2023","unstructured":"Doherty EL, Aw WY, Warren EC et al (2023) Patient-derived extracellular matrix demonstrates role of COL3A1 in blood vessel mechanics. Acta Biomater 166:346\u2013359. https:\/\/doi.org\/10.1016\/j.actbio.2023.05.015","journal-title":"Acta Biomater"},{"key":"10098_CR58","doi-asserted-by":"publisher","DOI":"10.1101\/cshperspect.a005058","volume":"3","author":"P Lu","year":"2011","unstructured":"Lu P, Takai K, Weaver VM, Werb Z (2011) Extracellular matrix degradation and remodeling in development and disease. Cold Spring Harb Perspect Biol 3:a005058. https:\/\/doi.org\/10.1101\/cshperspect.a005058","journal-title":"Cold Spring Harb Perspect Biol"},{"key":"10098_CR59","doi-asserted-by":"publisher","first-page":"280","DOI":"10.1007\/s11033-024-09255-5","volume":"51","author":"KB Rypdal","year":"2024","unstructured":"Rypdal KB, Apte SS, Lunde IG (2024) Emerging roles for the ADAMTS-like family of matricellular proteins in cardiovascular disease through regulation of the extracellular microenvironment. Mol Biol Rep 51:280. https:\/\/doi.org\/10.1007\/s11033-024-09255-5","journal-title":"Mol Biol Rep"},{"key":"10098_CR60","doi-asserted-by":"publisher","first-page":"118","DOI":"10.3390\/ph12030118","volume":"12","author":"EM Maguire","year":"2019","unstructured":"Maguire EM, Pearce SWA, Xiao R et al (2019) Matrix metalloproteinase in abdominal aortic aneurysm and aortic dissection. Pharmaceuticals (Basel) 12:118. https:\/\/doi.org\/10.3390\/ph12030118","journal-title":"Pharmaceuticals (Basel)"},{"key":"10098_CR61","doi-asserted-by":"publisher","first-page":"391","DOI":"10.1152\/physiol.00029.2013","volume":"28","author":"A Yabluchanskiy","year":"2013","unstructured":"Yabluchanskiy A, Ma Y, Iyer RP et al (2013) Matrix metalloproteinase-9: many shades of function in cardiovascular disease. Physiology (Bethesda) 28:391\u2013403. https:\/\/doi.org\/10.1152\/physiol.00029.2013","journal-title":"Physiology (Bethesda)"},{"key":"10098_CR62","doi-asserted-by":"publisher","first-page":"355","DOI":"10.1161\/CIRCRESAHA.122.322332","volume":"132","author":"J Aman","year":"2023","unstructured":"Aman J, Margadant C (2023) Integrin-dependent cell-matrix adhesion in endothelial health and disease. Circ Res 132:355\u2013378. https:\/\/doi.org\/10.1161\/CIRCRESAHA.122.322332","journal-title":"Circ Res"},{"key":"10098_CR63","doi-asserted-by":"publisher","first-page":"58","DOI":"10.1164\/rccm.201108-1381OC","volume":"185","author":"G Su","year":"2012","unstructured":"Su G, Atakilit A, Li JT et al (2012) Absence of integrin \u03b1v\u03b23 enhances vascular leak in mice by inhibiting endothelial cortical actin formation. Am J Respir Crit Care Med 185:58\u201366. https:\/\/doi.org\/10.1164\/rccm.201108-1381OC","journal-title":"Am J Respir Crit Care Med"},{"key":"10098_CR64","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.71032","volume":"11","author":"AL Godeau","year":"2022","unstructured":"Godeau AL, Leoni M, Comelles J et al (2022) 3D single cell migration driven by temporal correlation between oscillating force dipoles. Elife 11:e71032. https:\/\/doi.org\/10.7554\/eLife.71032","journal-title":"Elife"},{"key":"10098_CR65","doi-asserted-by":"publisher","first-page":"1279","DOI":"10.1016\/s0960-9822(02)70714-8","volume":"6","author":"A Bershadsky","year":"1996","unstructured":"Bershadsky A, Chausovsky A, Becker E et al (1996) Involvement of microtubules in the control of adhesion-dependent signal transduction. Curr Biol 6:1279\u20131289. https:\/\/doi.org\/10.1016\/s0960-9822(02)70714-8","journal-title":"Curr Biol"},{"key":"10098_CR66","doi-asserted-by":"publisher","first-page":"481","DOI":"10.1083\/jcb.201206050","volume":"198","author":"S Stehbens","year":"2012","unstructured":"Stehbens S, Wittmann T (2012) Targeting and transport: how microtubules control focal adhesion dynamics. J Cell Biol 198:481\u2013489. https:\/\/doi.org\/10.1083\/jcb.201206050","journal-title":"J Cell Biol"},{"key":"10098_CR67","doi-asserted-by":"publisher","first-page":"2715","DOI":"10.1038\/s44318-024-00114-4","volume":"43","author":"J Aureille","year":"2024","unstructured":"Aureille J, Prabhu SS, Barnett SF et al (2024) Focal adhesions are controlled by microtubules through local contractility regulation. EMBO J 43:2715\u20132732. https:\/\/doi.org\/10.1038\/s44318-024-00114-4","journal-title":"EMBO J"},{"key":"10098_CR68","doi-asserted-by":"publisher","first-page":"1331","DOI":"10.1042\/BST20221541","volume":"51","author":"S De Silva","year":"2023","unstructured":"De Silva S, Fan Z, Kang B et al (2023) Nesprin-1: novel regulator of striated muscle nuclear positioning and mechanotransduction. Biochem Soc Trans 51:1331\u20131345. https:\/\/doi.org\/10.1042\/BST20221541","journal-title":"Biochem Soc Trans"},{"key":"10098_CR69","doi-asserted-by":"publisher","first-page":"585","DOI":"10.1016\/j.ijcard.2015.03.423","volume":"187","author":"CJA Ramachandra","year":"2015","unstructured":"Ramachandra CJA, Mehta A, Guo KWQ et al (2015) Molecular pathogenesis of Marfan syndrome. Int J Cardiol 187:585\u2013591. https:\/\/doi.org\/10.1016\/j.ijcard.2015.03.423","journal-title":"Int J Cardiol"},{"key":"10098_CR70","doi-asserted-by":"publisher","DOI":"10.1038\/s41572-020-0194-9","volume":"6","author":"F Malfait","year":"2020","unstructured":"Malfait F, Castori M, Francomano CA et al (2020) The Ehlers\u2013Danlos syndromes. Nat Rev Dis Primers 6:64. https:\/\/doi.org\/10.1038\/s41572-020-0194-9","journal-title":"Nat Rev Dis Primers"},{"key":"10098_CR71","doi-asserted-by":"publisher","DOI":"10.1016\/j.vph.2023.107215","volume":"153","author":"S Seeburun","year":"2023","unstructured":"Seeburun S, Wu S, Hemani D et al (2023) Insights into elastic fiber fragmentation: mechanisms and treatment of aortic aneurysm in Marfan syndrome. Vascul Pharmacol 153:107215. https:\/\/doi.org\/10.1016\/j.vph.2023.107215","journal-title":"Vascul Pharmacol"},{"key":"10098_CR72","doi-asserted-by":"publisher","DOI":"10.1038\/ncomms4984","volume":"5","author":"C Collins","year":"2014","unstructured":"Collins C, Osborne LD, Guilluy C et al (2014) Haemodynamic and extracellular matrix cues regulate the mechanical phenotype and stiffness of aortic endothelial cells. Nat Commun 5:3984. https:\/\/doi.org\/10.1038\/ncomms4984","journal-title":"Nat Commun"},{"key":"10098_CR73","doi-asserted-by":"publisher","DOI":"10.1016\/j.lfs.2023.122233","volume":"334","author":"L Cheng","year":"2023","unstructured":"Cheng L, Yue H, Zhang H et al (2023) The influence of microenvironment stiffness on endothelial cell fate: implication for occurrence and progression of atherosclerosis. Life Sci 334:122233. https:\/\/doi.org\/10.1016\/j.lfs.2023.122233","journal-title":"Life Sci"},{"key":"10098_CR74","doi-asserted-by":"publisher","first-page":"718","DOI":"10.1038\/s44161-023-00311-0","volume":"2","author":"D Wang","year":"2023","unstructured":"Wang D, Brady T, Santhanam L, Gerecht S (2023) The extracellular matrix mechanics in the vasculature. Nat Cardiovasc Res 2:718\u2013732. https:\/\/doi.org\/10.1038\/s44161-023-00311-0","journal-title":"Nat Cardiovasc Res"},{"key":"10098_CR75","doi-asserted-by":"publisher","first-page":"1664","DOI":"10.1091\/mbc.E18-07-0438","volume":"30","author":"RM Stewart","year":"2019","unstructured":"Stewart RM, Rodriguez EC, King MC (2019) Ablation of SUN2-containing LINC complexes drives cardiac hypertrophy without interstitial fibrosis. Mol Biol Cell 30:1664\u20131675. https:\/\/doi.org\/10.1091\/mbc.E18-07-0438","journal-title":"Mol Biol Cell"},{"key":"10098_CR76","doi-asserted-by":"publisher","DOI":"10.1242\/jcs.232843","volume":"132","author":"S Seetharaman","year":"2019","unstructured":"Seetharaman S, Etienne-Manneville S (2019) Microtubules at focal adhesions \u2013 a double-edged sword. J Cell Sci 132:jcs232843. https:\/\/doi.org\/10.1242\/jcs.232843","journal-title":"J Cell Sci"},{"key":"10098_CR77","doi-asserted-by":"publisher","DOI":"10.3389\/fphys.2024.1425620","author":"K Moise","year":"2024","unstructured":"Moise K, Arun KM, Pillai M et al (2024) Endothelial cell elongation and alignment in response to shear stress requires acetylation of microtubules. Front Physiol. https:\/\/doi.org\/10.3389\/fphys.2024.1425620","journal-title":"Front Physiol"},{"key":"10098_CR78","doi-asserted-by":"publisher","first-page":"549","DOI":"10.1016\/j.mcn.2008.08.004","volume":"39","author":"O Sbai","year":"2008","unstructured":"Sbai O, Ferhat L, Bernard A et al (2008) Vesicular trafficking and secretion of matrix metalloproteinases-2, -9 and tissue inhibitor of metalloproteinases-1 in neuronal cells. Mol Cell Neurosci 39:549\u2013568. https:\/\/doi.org\/10.1016\/j.mcn.2008.08.004","journal-title":"Mol Cell Neurosci"},{"key":"10098_CR79","doi-asserted-by":"publisher","first-page":"8468","DOI":"10.1074\/jbc.M111.290676","volume":"287","author":"R Hanania","year":"2012","unstructured":"Hanania R, Sun HS, Xu K et al (2012) Classically activated macrophages use stable microtubules for matrix metalloproteinase-9 (MMP-9) secretion. J Biol Chem 287:8468\u20138483. https:\/\/doi.org\/10.1074\/jbc.M111.290676","journal-title":"J Biol Chem"},{"key":"10098_CR80","doi-asserted-by":"publisher","first-page":"960","DOI":"10.1161\/ATVBAHA.114.304412","volume":"35","author":"E Crosas-Molist","year":"2015","unstructured":"Crosas-Molist E, Meirelles T, L\u00f3pez-Luque J et al (2015) Vascular smooth muscle cell phenotypic changes in patients with Marfan syndrome. Arterioscler Thromb Vasc Biol 35:960\u2013972. https:\/\/doi.org\/10.1161\/ATVBAHA.114.304412","journal-title":"Arterioscler Thromb Vasc Biol"},{"key":"10098_CR81","doi-asserted-by":"publisher","first-page":"H1037","DOI":"10.1152\/ajpheart.00391.2024","volume":"327","author":"KK Dwivedi","year":"2024","unstructured":"Dwivedi KK, Rother J, Wagenseil JE (2024) Age- and sex-specific biomechanics and extracellular matrix remodeling of the ascending aorta in a mouse model of severe Marfan syndrome. Am J Physiol Heart Circ Physiol 327:H1037\u2013H1051. https:\/\/doi.org\/10.1152\/ajpheart.00391.2024","journal-title":"Am J Physiol Heart Circ Physiol"},{"key":"10098_CR82","doi-asserted-by":"publisher","first-page":"182","DOI":"10.1091\/mbc.E16-06-0467","volume":"28","author":"K Thakar","year":"2017","unstructured":"Thakar K, May CK, Rogers A, Carroll CW (2017) Opposing roles for distinct LINC complexes in regulation of the small GTPase RhoA. Mol Biol Cell 28:182\u2013191. https:\/\/doi.org\/10.1091\/mbc.E16-06-0467","journal-title":"Mol Biol Cell"},{"key":"10098_CR83","doi-asserted-by":"publisher","first-page":"3097","DOI":"10.1016\/j.cub.2017.08.073","volume":"27","author":"R Zhu","year":"2017","unstructured":"Zhu R, Antoku S, Gundersen GG (2017) Centrifugal displacement of nuclei reveals multiple LINC complex mechanisms for homeostatic nuclear positioning. Curr Biol 27:3097-3110.e5. https:\/\/doi.org\/10.1016\/j.cub.2017.08.073","journal-title":"Curr Biol"},{"key":"10098_CR84","doi-asserted-by":"publisher","first-page":"2258","DOI":"10.1093\/hmg\/ddx116","volume":"26","author":"C Zhou","year":"2017","unstructured":"Zhou C, Li C, Zhou B et al (2017) Novel nesprin-1 mutations associated with dilated cardiomyopathy cause nuclear envelope disruption and defects in myogenesis. Hum Mol Genet 26:2258\u20132276. https:\/\/doi.org\/10.1093\/hmg\/ddx116","journal-title":"Hum Mol Genet"},{"key":"10098_CR85","doi-asserted-by":"publisher","DOI":"10.1038\/s41598-019-50728-6","volume":"9","author":"I Holt","year":"2019","unstructured":"Holt I, Fuller HR, Lam LT et al (2019) Nesprin-1-alpha2 associates with kinesin at myotube outer nuclear membranes, but is restricted to neuromuscular junction nuclei in adult muscle. Sci Rep 9:14202. https:\/\/doi.org\/10.1038\/s41598-019-50728-6","journal-title":"Sci Rep"},{"key":"10098_CR86","doi-asserted-by":"publisher","first-page":"86","DOI":"10.1038\/s41580-024-00780-6","volume":"26","author":"A Yildiz","year":"2025","unstructured":"Yildiz A (2025) Mechanism and regulation of kinesin motors. Nat Rev Mol Cell Biol 26:86\u2013103. https:\/\/doi.org\/10.1038\/s41580-024-00780-6","journal-title":"Nat Rev Mol Cell Biol"}],"container-title":["Angiogenesis"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1007\/s10456-026-10098-w.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/article\/10.1007\/s10456-026-10098-w","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1007\/s10456-026-10098-w.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,10,15]],"date-time":"2026-10-15T20:51:26Z","timestamp":1792097486000},"score":1,"resource":{"primary":{"URL":"https:\/\/link.springer.com\/article\/10.1007\/s10456-026-10098-w"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,10,15]]},"references-count":86,"journal-issue":{"issue":"4","published-online":{"date-parts":[[2026,11]]}},"alternative-id":["10098"],"URL":"https:\/\/doi.org\/10.1007\/s10456-026-10098-w","relation":{},"ISSN":["1573-7209"],"issn-type":[{"value":"1573-7209","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,10,15]]},"assertion":[{"value":"18 May 2026","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"26 September 2026","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"15 October 2026","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"The authors declare no competing interests.","order":1,"name":"Ethics","label":"Conflict of interest","group":{"name":"EthicsHeading","label":"Declarations"}}],"article-number":"78"}}