{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,5]],"date-time":"2026-10-05T07:56:05Z","timestamp":1791186965024,"version":"4.1.0"},"reference-count":71,"publisher":"Springer Science and Business Media LLC","issue":"9","license":[{"start":{"date-parts":[[2026,8,27]],"date-time":"2026-08-27T00:00:00Z","timestamp":1787788800000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.springernature.com\/gp\/researchers\/text-and-data-mining"},{"start":{"date-parts":[[2026,8,27]],"date-time":"2026-08-27T00:00:00Z","timestamp":1787788800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/www.springernature.com\/gp\/researchers\/text-and-data-mining"}],"funder":[{"name":"Joint TCM Science & Technology Projects of National Demonstration Zones for Comprehensive TCM Reform","award":["GZY-KJS-ZJ-2025\u2013054"],"award-info":[{"award-number":["GZY-KJS-ZJ-2025\u2013054"]}]},{"name":"Zhoushan \"Future via the Sea\" Haina Program Innovation Project"},{"name":"Medical Young Talents Program of Zhejiang Province"}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Mol Cell Biochem"],"DOI":"10.1007\/s11010-026-05706-8","type":"journal-article","created":{"date-parts":[[2026,8,27]],"date-time":"2026-08-27T08:37:11Z","timestamp":1787819831000},"page":"4987-5007","update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Actin polymerization drives endogenous MMP-9 upregulation and blood\u2013brain barrier disruption in ischemic brain endothelial cells"],"prefix":"10.1007","volume":"481","author":[{"given":"Yiyin","family":"Zhao","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Songbin","family":"He","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yiming","family":"Wang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Meng","family":"Jin","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yichao","family":"Fu","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xiaojing","family":"Zhou","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2026,8,27]]},"reference":[{"issue":"1","key":"5706_CR1","doi-asserted-by":"publisher","first-page":"125","DOI":"10.1007\/s00281-023-00984-6","volume":"45","author":"B Ma\u00efer","year":"2023","unstructured":"Ma\u00efer B, Tsai AS, Einhaus JF et al (2023) Neuroimaging is the new \u201cspatial omic\u201d: multi-omic approaches to neuro-inflammation and immuno-thrombosis in acute ischemic stroke. Semin Immunopathol 45(1):125\u2013143. https:\/\/doi.org\/10.1007\/s00281-023-00984-6","journal-title":"Semin Immunopathol"},{"issue":"1","key":"5706_CR2","doi-asserted-by":"publisher","first-page":"53","DOI":"10.1161\/strokeaha.118.022913","volume":"50","author":"S Sennf\u00e4lt","year":"2019","unstructured":"Sennf\u00e4lt S, Norrving B, Petersson J et al (2019) Long-term survival and function after stroke: a longitudinal observational study from the swedish stroke register. Stroke 50(1):53\u201361. https:\/\/doi.org\/10.1161\/strokeaha.118.022913","journal-title":"Stroke"},{"key":"5706_CR3","doi-asserted-by":"publisher","first-page":"1716041","DOI":"10.3389\/fimmu.2025.1716041","volume":"16","author":"H Wang","year":"2025","unstructured":"Wang H, Yiqiang Z, Cai J et al (2025) Immune-mediated blood-brain barrier disruption after ischemic stroke: mechanisms and therapeutic targets. Front Immunol 16:1716041. https:\/\/doi.org\/10.3389\/fimmu.2025.1716041","journal-title":"Front Immunol"},{"key":"5706_CR4","doi-asserted-by":"publisher","DOI":"10.1242\/jcs.188185","author":"MA Odenwald","year":"2017","unstructured":"Odenwald MA, Choi W, Buckley A et al (2017) ZO-1 interactions with F-actin and occludin direct epithelial polarization and single lumen specification in 3D culture. J Cell Sci. https:\/\/doi.org\/10.1242\/jcs.188185","journal-title":"J Cell Sci"},{"issue":"45","key":"5706_CR5","doi-asserted-by":"publisher","first-page":"29745","DOI":"10.1074\/jbc.273.45.29745","volume":"273","author":"AS Fanning","year":"1998","unstructured":"Fanning AS, Jameson BJ, Jesaitis LA et al (1998) The tight junction protein ZO-1 establishes a link between the transmembrane protein occludin and the actin cytoskeleton. J Biol Chem 273(45):29745\u201329753. https:\/\/doi.org\/10.1074\/jbc.273.45.29745","journal-title":"J Biol Chem"},{"issue":"24","key":"5706_CR6","doi-asserted-by":"publisher","first-page":"3783","DOI":"10.1016\/j.cub.2017.11.014","volume":"27","author":"D Spadaro","year":"2017","unstructured":"Spadaro D, Le S, Laroche T et al (2017) Tension-dependent stretching activates ZO-1 to control the junctional localization of its interactors. Curr Biol 27(24):3783-3795.e8. https:\/\/doi.org\/10.1016\/j.cub.2017.11.014","journal-title":"Curr Biol"},{"issue":"2","key":"5706_CR7","doi-asserted-by":"publisher","first-page":"324","DOI":"10.1016\/s0006-291x(03)00167-0","volume":"302","author":"G Kale","year":"2003","unstructured":"Kale G, Naren AP, Sheth P et al (2003) Tyrosine phosphorylation of occludin attenuates its interactions with ZO-1, ZO-2, and ZO-3. Biochem Biophys Res Commun 302(2):324\u2013329. https:\/\/doi.org\/10.1016\/s0006-291x(03)00167-0","journal-title":"Biochem Biophys Res Commun"},{"issue":"2","key":"5706_CR8","doi-asserted-by":"publisher","first-page":"279","DOI":"10.1007\/s00018-016-2325-8","volume":"74","author":"YL Dorland","year":"2017","unstructured":"Dorland YL, Huveneers S (2017) Cell-cell junctional mechanotransduction in endothelial remodeling. Cell Mol Life Sci 74(2):279\u2013292. https:\/\/doi.org\/10.1007\/s00018-016-2325-8","journal-title":"Cell Mol Life Sci"},{"key":"5706_CR9","doi-asserted-by":"publisher","DOI":"10.1083\/jcb.201909081","author":"CM Grimsley-Myers","year":"2020","unstructured":"Grimsley-Myers CM, Isaacson RH, Cadwell CM et al (2020) VE-cadherin endocytosis controls vascular integrity and patterning during development. J Cell Biol. https:\/\/doi.org\/10.1083\/jcb.201909081","journal-title":"J Cell Biol"},{"key":"5706_CR10","doi-asserted-by":"publisher","first-page":"257","DOI":"10.1007\/978-1-0716-3429-5_21","volume":"2711","author":"H Alluri","year":"2024","unstructured":"Alluri H, Peddaboina CS, Tharakan B (2024) Evaluation of tight junction integrity in brain endothelial cells using confocal microscopy. Methods Mol Biol 2711:257\u2013262. https:\/\/doi.org\/10.1007\/978-1-0716-3429-5_21","journal-title":"Methods Mol Biol"},{"key":"5706_CR11","doi-asserted-by":"publisher","DOI":"10.3390\/cells13191635","author":"Y Deng","year":"2024","unstructured":"Deng Y, Qiao Z, Zhou C et al (2024) Endothelial myosin IIA is required for the maintenance of blood-brain barrier integrity. Cells. https:\/\/doi.org\/10.3390\/cells13191635","journal-title":"Cells"},{"key":"5706_CR12","doi-asserted-by":"publisher","first-page":"256","DOI":"10.1016\/j.neuroscience.2020.10.034","volume":"453","author":"AS Douglas","year":"2021","unstructured":"Douglas AS, Shearer JA, Okolo A et al (2021) The relationship between cerebral reperfusion and regional expression of matrix metalloproteinase-9 in rat brain following focal cerebral ischemia. Neuroscience 453:256\u2013265. https:\/\/doi.org\/10.1016\/j.neuroscience.2020.10.034","journal-title":"Neuroscience"},{"key":"5706_CR13","doi-asserted-by":"publisher","first-page":"153505","DOI":"10.1016\/j.phymed.2021.153505","volume":"84","author":"Y Jiao-Yan","year":"2021","unstructured":"Jiao-Yan Y, Qing-Qing L, Xi L et al (2021) Oxymatrine improves blood-brain barrier integrity after cerebral ischemia-reperfusion injury by downregulating CAV1 and MMP9 expression. Phytomedicine 84:153505. https:\/\/doi.org\/10.1016\/j.phymed.2021.153505","journal-title":"Phytomedicine"},{"issue":"12","key":"5706_CR14","doi-asserted-by":"publisher","first-page":"1547","DOI":"10.1038\/s41401-020-0409-3","volume":"41","author":"MB Liu","year":"2020","unstructured":"Liu MB, Wang W, Gao JM et al (2020) Icariside II attenuates cerebral ischemia\/reperfusion-induced blood-brain barrier dysfunction in rats via regulating the balance of MMP9\/TIMP1. Acta Pharmacol Sin 41(12):1547\u20131556. https:\/\/doi.org\/10.1038\/s41401-020-0409-3","journal-title":"Acta Pharmacol Sin"},{"issue":"7","key":"5706_CR15","doi-asserted-by":"publisher","first-page":"1944","DOI":"10.4103\/nrr.Nrr-d-24-00392","volume":"20","author":"J Castillo-Gonz\u00e1lez","year":"2025","unstructured":"Castillo-Gonz\u00e1lez J, Gonz\u00e1lez-Rey E (2025) Beyond wrecking a wall: revisiting the concept of blood-brain barrier breakdown in ischemic stroke. Neural Regen Res 20(7):1944\u20131956. https:\/\/doi.org\/10.4103\/nrr.Nrr-d-24-00392","journal-title":"Neural Regen Res"},{"key":"5706_CR16","doi-asserted-by":"publisher","DOI":"10.1016\/j.phrs.2023.106720","volume":"190","author":"Y Ji","year":"2023","unstructured":"Ji Y, Gao Q, Ma Y et al (2023) An MMP-9 exclusive neutralizing antibody attenuates blood-brain barrier breakdown in mice with stroke and reduces stroke patient-derived MMP-9 activity. Pharmacol Res 190:106720. https:\/\/doi.org\/10.1016\/j.phrs.2023.106720","journal-title":"Pharmacol Res"},{"key":"5706_CR17","doi-asserted-by":"publisher","DOI":"10.3892\/mmr.2025.13590","author":"P Guo","year":"2025","unstructured":"Guo P, Li H, Zhang X et al (2025) Matrix metalloproteinase-9 in hemorrhagic transformation after acute ischemic stroke (Review). Mol Med Rep. https:\/\/doi.org\/10.3892\/mmr.2025.13590","journal-title":"Mol Med Rep"},{"issue":"33","key":"5706_CR18","doi-asserted-by":"publisher","first-page":"3169","DOI":"10.1093\/eurheartj\/ehaa207","volume":"41","author":"B Liao","year":"2020","unstructured":"Liao B, Geng L, Zhang F et al (2020) Adipocyte fatty acid-binding protein exacerbates cerebral ischaemia injury by disrupting the blood-brain barrier. Eur Heart J 41(33):3169\u20133180. https:\/\/doi.org\/10.1093\/eurheartj\/ehaa207","journal-title":"Eur Heart J"},{"issue":"38","key":"5706_CR19","doi-asserted-by":"publisher","first-page":"9451","DOI":"10.1523\/jneurosci.2674-08.2008","volume":"28","author":"BW McColl","year":"2008","unstructured":"McColl BW, Rothwell NJ, Allan SM (2008) Systemic inflammation alters the kinetics of cerebrovascular tight junction disruption after experimental stroke in mice. J Neurosci 28(38):9451\u20139462. https:\/\/doi.org\/10.1523\/jneurosci.2674-08.2008","journal-title":"J Neurosci"},{"issue":"8","key":"5706_CR20","doi-asserted-by":"publisher","first-page":"2120","DOI":"10.1038\/s41401-025-01505-x","volume":"46","author":"LD Du","year":"2025","unstructured":"Du LD, Fang C, Wang YQ et al (2025) MMP-9 inhibitor SB-3CT improves neurological outcomes in ischemic stroke mice by modulation of astrocytic lipid metabolism. Acta Pharmacol Sin 46(8):2120\u20132135. https:\/\/doi.org\/10.1038\/s41401-025-01505-x","journal-title":"Acta Pharmacol Sin"},{"issue":"12","key":"5706_CR21","doi-asserted-by":"publisher","first-page":"1430","DOI":"10.1097\/01.Wcb.0000090680.07515.C8","volume":"23","author":"C Justicia","year":"2003","unstructured":"Justicia C, Pan\u00e9s J, Sol\u00e9 S et al (2003) Neutrophil infiltration increases matrix metalloproteinase-9 in the ischemic brain after occlusion\/reperfusion of the middle cerebral artery in rats. J Cereb Blood Flow Metab 23(12):1430\u20131440. https:\/\/doi.org\/10.1097\/01.Wcb.0000090680.07515.C8","journal-title":"J Cereb Blood Flow Metab"},{"issue":"4","key":"5706_CR22","doi-asserted-by":"publisher","first-page":"313","DOI":"10.1136\/thx.2003.008854","volume":"59","author":"O Prause","year":"2004","unstructured":"Prause O, Bozinovski S, Anderson GP et al (2004) Increased matrix metalloproteinase-9 concentration and activity after stimulation with interleukin-17 in mouse airways. Thorax 59(4):313\u2013317. https:\/\/doi.org\/10.1136\/thx.2003.008854","journal-title":"Thorax"},{"issue":"1","key":"5706_CR23","doi-asserted-by":"publisher","first-page":"563","DOI":"10.1007\/s12035-013-8538-z","volume":"49","author":"M Chaturvedi","year":"2014","unstructured":"Chaturvedi M, Kaczmarek L (2014) Mmp-9 inhibition: a therapeutic strategy in ischemic stroke. Mol Neurobiol 49(1):563\u2013573. https:\/\/doi.org\/10.1007\/s12035-013-8538-z","journal-title":"Mol Neurobiol"},{"issue":"5","key":"5706_CR24","doi-asserted-by":"publisher","first-page":"1169","DOI":"10.1161\/01.STR.0000125861.55804.f2","volume":"35","author":"CM Maier","year":"2004","unstructured":"Maier CM, Hsieh L, Yu F et al (2004) Matrix metalloproteinase-9 and myeloperoxidase expression: quantitative analysis by antigen immunohistochemistry in a model of transient focal cerebral ischemia. Stroke 35(5):1169\u20131174. https:\/\/doi.org\/10.1161\/01.STR.0000125861.55804.f2","journal-title":"Stroke"},{"issue":"6","key":"5706_CR25","doi-asserted-by":"publisher","first-page":"1399","DOI":"10.1161\/01.STR.0000223001.06264.af","volume":"37","author":"A Rosell","year":"2006","unstructured":"Rosell A, Ortega-Aznar A, Alvarez-Sab\u00edn J et al (2006) Increased brain expression of matrix metalloproteinase-9 after ischemic and hemorrhagic human stroke. Stroke 37(6):1399\u20131406. https:\/\/doi.org\/10.1161\/01.STR.0000223001.06264.af","journal-title":"Stroke"},{"issue":"6","key":"5706_CR26","doi-asserted-by":"publisher","first-page":"107727","DOI":"10.1016\/j.jstrokecerebrovasdis.2024.107727","volume":"33","author":"J Li","year":"2024","unstructured":"Li J, Wang B, Dai F et al (2024) The predictive value of serum F-actin on the severity and early neurological deterioration of acute ischemic stroke: Predictive value of F-actin in stroke. J Stroke Cerebrovasc Dis 33(6):107727. https:\/\/doi.org\/10.1016\/j.jstrokecerebrovasdis.2024.107727","journal-title":"J Stroke Cerebrovasc Dis"},{"key":"5706_CR27","doi-asserted-by":"publisher","DOI":"10.1016\/j.tice.2025.102946","volume":"95","author":"Y Zhao","year":"2025","unstructured":"Zhao Y, Zhou X, He S et al (2025) Oxygen-glucose deprivation induces actin spillover in brain endothelial cells. Tissue Cell 95:102946. https:\/\/doi.org\/10.1016\/j.tice.2025.102946","journal-title":"Tissue Cell"},{"key":"5706_CR28","doi-asserted-by":"publisher","first-page":"1100938","DOI":"10.3389\/fcell.2023.1100938","volume":"11","author":"T Mgrditchian","year":"2023","unstructured":"Mgrditchian T, Brown-Clay J, Hoffmann C et al (2023) Actin cytoskeleton depolymerization increases matrix metalloproteinase gene expression in breast cancer cells by promoting translocation of cysteine-rich protein 2 to the nucleus. Front Cell Dev Biol 11:1100938. https:\/\/doi.org\/10.3389\/fcell.2023.1100938","journal-title":"Front Cell Dev Biol"},{"key":"5706_CR29","doi-asserted-by":"publisher","first-page":"115141","DOI":"10.1016\/j.biopha.2023.115141","volume":"165","author":"W Lu","year":"2023","unstructured":"Lu W, Chen Z, Wen J (2023) The role of RhoA\/ROCK pathway in the ischemic stroke-induced neuroinflammation. Biomed Pharmacother 165:115141. https:\/\/doi.org\/10.1016\/j.biopha.2023.115141","journal-title":"Biomed Pharmacother"},{"key":"5706_CR30","doi-asserted-by":"publisher","DOI":"10.3390\/cells10010166","author":"I Figiel","year":"2021","unstructured":"Figiel I, Kruk PK, Zar\u0119ba-Kozio\u0142 M et al (2021) MMP-9 signaling pathways that engage Rho GTPases in brain plasticity. Cells. https:\/\/doi.org\/10.3390\/cells10010166","journal-title":"Cells"},{"issue":"5","key":"5706_CR31","doi-asserted-by":"publisher","first-page":"891","DOI":"10.1097\/MIB.0b013e3182813297","volume":"19","author":"LA Johnson","year":"2013","unstructured":"Johnson LA, Rodansky ES, Sauder KL et al (2013) Matrix stiffness corresponding to strictured bowel induces a fibrogenic response in human colonic fibroblasts. Inflamm Bowel Dis 19(5):891\u2013903. https:\/\/doi.org\/10.1097\/MIB.0b013e3182813297","journal-title":"Inflamm Bowel Dis"},{"issue":"10","key":"5706_CR32","doi-asserted-by":"publisher","first-page":"2029","DOI":"10.1002\/term.2737","volume":"12","author":"L Zheng","year":"2018","unstructured":"Zheng L, Zhang L, Chen L et al (2018) Static magnetic field regulates proliferation, migration, differentiation, and YAP\/TAZ activation of human dental pulp stem cells. J Tissue Eng Regen Med 12(10):2029\u20132040. https:\/\/doi.org\/10.1002\/term.2737","journal-title":"J Tissue Eng Regen Med"},{"issue":"1","key":"5706_CR33","doi-asserted-by":"publisher","first-page":"449","DOI":"10.1186\/s12964-025-02452-0","volume":"23","author":"A Simiczyjew","year":"2025","unstructured":"Simiczyjew A, Kot M, Majkowski M et al (2025) Phenotype switching in highly invasive resistant to vemurafenib and cobimetinib melanoma cells. Cell Commun Signal 23(1):449. https:\/\/doi.org\/10.1186\/s12964-025-02452-0","journal-title":"Cell Commun Signal"},{"issue":"16","key":"5706_CR34","doi-asserted-by":"publisher","DOI":"10.1111\/jcmm.70008","volume":"28","author":"C Zhou","year":"2024","unstructured":"Zhou C, Zhou Y, Vong CT et al (2024) 3,3\u2019,4,5\u2019-Tetramethoxy-trans-stilbene and 3,4\u2019,5-trimethoxy-trans-stilbene prevent oxygen-glucose deprivation-induced injury in brain endothelial cell. J Cell Mol Med 28(16):e70008. https:\/\/doi.org\/10.1111\/jcmm.70008","journal-title":"J Cell Mol Med"},{"key":"5706_CR35","doi-asserted-by":"publisher","DOI":"10.3389\/fnins.2026.1754128","volume":"20","author":"M Dong","year":"2026","unstructured":"Dong M, Huang L, Zhang Q et al (2026) Suppression of miR-195 attenuates oxygen-glucose deprivation\/reperfusion-induced BBB destruction, possibly via targeting BCL2L2. Front Neurosci 20:1754128. https:\/\/doi.org\/10.3389\/fnins.2026.1754128","journal-title":"Front Neurosci"},{"issue":"4","key":"5706_CR36","doi-asserted-by":"publisher","first-page":"907","DOI":"10.1007\/s11064-021-03492-x","volume":"47","author":"C Xu","year":"2022","unstructured":"Xu C, Yu H, Chen B et al (2022) Serum exosomal mir-340-5p promotes angiogenesis in brain microvascular endothelial cells during oxygen-glucose deprivation. Neurochem Res 47(4):907\u2013920. https:\/\/doi.org\/10.1007\/s11064-021-03492-x","journal-title":"Neurochem Res"},{"key":"5706_CR37","doi-asserted-by":"publisher","DOI":"10.1186\/1756-6606-6-14","volume":"6","author":"KE Hawkins","year":"2013","unstructured":"Hawkins KE, DeMars KM, Yang C et al (2013) Fluorometric immunocapture assay for the specific measurement of matrix metalloproteinase-9 activity in biological samples: application to brain and plasma from rats with ischemic stroke. Mol Brain 6:14. https:\/\/doi.org\/10.1186\/1756-6606-6-14","journal-title":"Mol Brain"},{"key":"5706_CR38","doi-asserted-by":"publisher","DOI":"10.1186\/s13104-015-1284-8","volume":"8","author":"J Christensen","year":"2015","unstructured":"Christensen J, Shastri VP (2015) Matrix-metalloproteinase-9 is cleaved and activated by cathepsin K. BMC Res Notes 8:322. https:\/\/doi.org\/10.1186\/s13104-015-1284-8","journal-title":"BMC Res Notes"},{"issue":"43","key":"5706_CR39","doi-asserted-by":"publisher","first-page":"17963","DOI":"10.1074\/jbc.M117.806075","volume":"292","author":"RH Scannevin","year":"2017","unstructured":"Scannevin RH, Alexander R, Haarlander TM et al (2017) Discovery of a highly selective chemical inhibitor of matrix metalloproteinase-9 (MMP-9) that allosterically inhibits zymogen activation. J Biol Chem 292(43):17963\u201317974. https:\/\/doi.org\/10.1074\/jbc.M117.806075","journal-title":"J Biol Chem"},{"issue":"2","key":"5706_CR40","doi-asserted-by":"publisher","first-page":"1222","DOI":"10.3390\/ijms12021222","volume":"12","author":"K Liu","year":"2011","unstructured":"Liu K, Li Z, Wu T et al (2011) Role of rho kinase in microvascular damage following cerebral ischemia reperfusion in rats. Int J Mol Sci 12(2):1222\u20131231. https:\/\/doi.org\/10.3390\/ijms12021222","journal-title":"Int J Mol Sci"},{"key":"5706_CR41","doi-asserted-by":"publisher","DOI":"10.3389\/fphys.2018.00824","volume":"9","author":"F Martino","year":"2018","unstructured":"Martino F, Perestrelo AR, Vinarsk\u00fd V et al (2018) Cellular mechanotransduction: from tension to function. Front Physiol 9:824. https:\/\/doi.org\/10.3389\/fphys.2018.00824","journal-title":"Front Physiol"},{"issue":"4","key":"5706_CR42","doi-asserted-by":"publisher","first-page":"445","DOI":"10.1083\/jcb.201609037","volume":"215","author":"Z Sun","year":"2016","unstructured":"Sun Z, Guo SS, F\u00e4ssler R (2016) Integrin-mediated mechanotransduction. J Cell Biol 215(4):445\u2013456. https:\/\/doi.org\/10.1083\/jcb.201609037","journal-title":"J Cell Biol"},{"key":"5706_CR43","doi-asserted-by":"publisher","first-page":"347","DOI":"10.1016\/j.brainres.2010.10.002","volume":"1367","author":"Q Hu","year":"2011","unstructured":"Hu Q, Chen C, Khatibi NH et al (2011) Lentivirus-mediated transfer of MMP-9 shRNA provides neuroprotection following focal ischemic brain injury in rats. Brain Res 1367:347\u2013359. https:\/\/doi.org\/10.1016\/j.brainres.2010.10.002","journal-title":"Brain Res"},{"issue":"15","key":"5706_CR44","doi-asserted-by":"publisher","first-page":"2769","DOI":"10.1091\/mbc.E15-04-0232","volume":"26","author":"CM Van Itallie","year":"2015","unstructured":"Van Itallie CM, Tietgens AJ, Krystofiak E et al (2015) A complex of ZO-1 and the BAR-domain protein TOCA-1 regulates actin assembly at the tight junction. Mol Biol Cell 26(15):2769\u20132787. https:\/\/doi.org\/10.1091\/mbc.E15-04-0232","journal-title":"Mol Biol Cell"},{"issue":"5","key":"5706_CR45","doi-asserted-by":"publisher","first-page":"3747","DOI":"10.1074\/jbc.M411365200","volume":"280","author":"SL M\u00fcller","year":"2005","unstructured":"M\u00fcller SL, Portwich M, Schmidt A et al (2005) The tight junction protein occludin and the adherens junction protein alpha-catenin share a common interaction mechanism with ZO-1. J Biol Chem 280(5):3747\u20133756. https:\/\/doi.org\/10.1074\/jbc.M411365200","journal-title":"J Biol Chem"},{"issue":"2","key":"5706_CR46","doi-asserted-by":"publisher","first-page":"691","DOI":"10.1074\/jbc.M115.648386","volume":"291","author":"CM Cadwell","year":"2016","unstructured":"Cadwell CM, Jenkins PM, Bennett V et al (2016) Ankyrin-G inhibits endocytosis of cadherin dimers. J Biol Chem 291(2):691\u2013704. https:\/\/doi.org\/10.1074\/jbc.M115.648386","journal-title":"J Biol Chem"},{"key":"5706_CR47","doi-asserted-by":"publisher","DOI":"10.1242\/jcs.259012","author":"CN Duong","year":"2021","unstructured":"Duong CN, Br\u00fcckner R, Schmitt M et al (2021) Force-induced changes of \u03b1-catenin conformation stabilize vascular junctions independently of vinculin. J Cell Sci. https:\/\/doi.org\/10.1242\/jcs.259012","journal-title":"J Cell Sci"},{"key":"5706_CR48","doi-asserted-by":"publisher","DOI":"10.1172\/jci164199","author":"J Tang","year":"2023","unstructured":"Tang J, Kang Y, Zhou Y et al (2023) TIMP2 ameliorates blood-brain barrier disruption in traumatic brain injury by inhibiting Src-dependent VE-cadherin internalization. J Clin Invest. https:\/\/doi.org\/10.1172\/jci164199","journal-title":"J Clin Invest"},{"issue":"12","key":"5706_CR49","doi-asserted-by":"publisher","first-page":"2075","DOI":"10.1039\/d0mt00187b","volume":"12","author":"J Wu","year":"2020","unstructured":"Wu J, Yang J, Yu M et al (2020) Lanthanum chloride causes blood-brain barrier disruption through intracellular calcium-mediated RhoA\/Rho kinase signaling and myosin light chain kinase. Metallomics 12(12):2075\u20132083. https:\/\/doi.org\/10.1039\/d0mt00187b","journal-title":"Metallomics"},{"issue":"7","key":"5706_CR50","doi-asserted-by":"publisher","DOI":"10.1590\/1414-431x20186201","volume":"51","author":"PL Luo","year":"2018","unstructured":"Luo PL, Wang YJ, Yang YY et al (2018) Hypoxia-induced hyperpermeability of rat glomerular endothelial cells involves HIF-2\u03b1 mediated changes in the expression of occludin and ZO-1. Braz J Med Biol Res 51(7):e6201. https:\/\/doi.org\/10.1590\/1414-431x20186201","journal-title":"Braz J Med Biol Res"},{"key":"5706_CR51","doi-asserted-by":"publisher","first-page":"126","DOI":"10.1016\/j.neuroscience.2018.02.035","volume":"377","author":"S Zhang","year":"2018","unstructured":"Zhang S, An Q, Wang T et al (2018) Autophagy- and MMP-2\/9-mediated reduction and redistribution of ZO-1 contribute to hyperglycemia-increased blood-brain barrier permeability during early reperfusion in stroke. Neuroscience 377:126\u2013137. https:\/\/doi.org\/10.1016\/j.neuroscience.2018.02.035","journal-title":"Neuroscience"},{"issue":"1","key":"5706_CR52","doi-asserted-by":"publisher","first-page":"597","DOI":"10.1038\/s41598-022-27308-2","volume":"13","author":"C Ivaldo","year":"2023","unstructured":"Ivaldo C, Passalacqua M, Furfaro AL et al (2023) Oxidative stress-induced MMP- and \u03b3-secretase-dependent VE-cadherin processing is modulated by the proteasome and BMP9\/10. Sci Rep 13(1):597. https:\/\/doi.org\/10.1038\/s41598-022-27308-2","journal-title":"Sci Rep"},{"key":"5706_CR53","doi-asserted-by":"publisher","first-page":"12","DOI":"10.1016\/j.expneurol.2019.03.017","volume":"316","author":"X Zuo","year":"2019","unstructured":"Zuo X, Lu J, Manaenko A et al (2019) MicroRNA-132 attenuates cerebral injury by protecting blood-brain-barrier in MCAO mice. Exp Neurol 316:12\u201319. https:\/\/doi.org\/10.1016\/j.expneurol.2019.03.017","journal-title":"Exp Neurol"},{"issue":"4","key":"5706_CR54","doi-asserted-by":"publisher","first-page":"C951","DOI":"10.1152\/ajpcell.00342.2023","volume":"325","author":"TS Wendt","year":"2023","unstructured":"Wendt TS, Gonzales RJ (2023) Ozanimod differentially preserves human cerebrovascular endothelial barrier proteins and attenuates matrix metalloproteinase-9 activity following in vitro acute ischemic injury. Am J Physiol Cell Physiol 325(4):C951-c971. https:\/\/doi.org\/10.1152\/ajpcell.00342.2023","journal-title":"Am J Physiol Cell Physiol"},{"key":"5706_CR55","doi-asserted-by":"publisher","first-page":"68","DOI":"10.1016\/j.expneurol.2017.12.016","volume":"302","author":"XF Li","year":"2018","unstructured":"Li XF, Zhang XJ, Zhang C et al (2018) Ulinastatin protects brain against cerebral ischemia\/reperfusion injury through inhibiting MMP-9 and alleviating loss of ZO-1 and occludin proteins in mice. Exp Neurol 302:68\u201374. https:\/\/doi.org\/10.1016\/j.expneurol.2017.12.016","journal-title":"Exp Neurol"},{"issue":"1","key":"5706_CR56","doi-asserted-by":"publisher","first-page":"273","DOI":"10.1007\/s12035-025-05592-z","volume":"63","author":"SS Gopal","year":"2025","unstructured":"Gopal SS, Kaur M, Lanzkron S et al (2025) The breakdown of neurovascular barriers: molecular mechanisms of tight junction dysfunction. Mol Neurobiol 63(1):273. https:\/\/doi.org\/10.1007\/s12035-025-05592-z","journal-title":"Mol Neurobiol"},{"issue":"49","key":"5706_CR57","doi-asserted-by":"publisher","first-page":"19520","DOI":"10.1073\/pnas.0807248105","volume":"105","author":"XB Wang","year":"2008","unstructured":"Wang XB, Bozdagi O, Nikitczuk JS et al (2008) Extracellular proteolysis by matrix metalloproteinase-9 drives dendritic spine enlargement and long-term potentiation coordinately. Proc Natl Acad Sci U S A 105(49):19520\u201319525. https:\/\/doi.org\/10.1073\/pnas.0807248105","journal-title":"Proc Natl Acad Sci U S A"},{"issue":"5\u20136","key":"5706_CR58","doi-asserted-by":"publisher","first-page":"321","DOI":"10.1515\/bmc-2016-0022","volume":"7","author":"N Bildyug","year":"2016","unstructured":"Bildyug N (2016) Matrix metalloproteinases: an emerging role in regulation of actin microfilament system. Biomol Concepts 7(5\u20136):321\u2013329. https:\/\/doi.org\/10.1515\/bmc-2016-0022","journal-title":"Biomol Concepts"},{"key":"5706_CR59","doi-asserted-by":"publisher","first-page":"8669","DOI":"10.12659\/msm.911394","volume":"24","author":"X Dai","year":"2018","unstructured":"Dai X, Geng F, Dai J et al (2018) Rho GTPase activating protein 24 (ARHGAP24) regulates the anti-cancer activity of sorafenib against breast cancer MDA-MB-231 cells via the signal transducer and activator of transcription 3 (STAT3) signaling pathway. Med Sci Monit 24:8669\u20138677. https:\/\/doi.org\/10.12659\/msm.911394","journal-title":"Med Sci Monit"},{"issue":"4","key":"5706_CR60","doi-asserted-by":"publisher","first-page":"1209","DOI":"10.3892\/ijo.2017.4100","volume":"51","author":"D Luo","year":"2017","unstructured":"Luo D, Chen H, Li X et al (2017) Activation of the ROCK1\/MMP-9 pathway is associated with the invasion and poor prognosis in papillary thyroid carcinoma. Int J Oncol 51(4):1209\u20131218. https:\/\/doi.org\/10.3892\/ijo.2017.4100","journal-title":"Int J Oncol"},{"key":"5706_CR61","doi-asserted-by":"publisher","first-page":"146297","DOI":"10.1016\/j.brainres.2019.06.016","volume":"1720","author":"CY Guo","year":"2019","unstructured":"Guo CY, Xiong TQ, Tan BH et al (2019) The temporal and spatial changes of actin cytoskeleton in the hippocampal CA1 neurons following transient global ischemia. Brain Res 1720:146297. https:\/\/doi.org\/10.1016\/j.brainres.2019.06.016","journal-title":"Brain Res"},{"key":"5706_CR62","doi-asserted-by":"publisher","first-page":"92","DOI":"10.1016\/j.brainres.2009.11.087","volume":"1316","author":"L Gisselsson","year":"2010","unstructured":"Gisselsson L, Toresson H, Ruscher K et al (2010) Rho kinase inhibition protects CA1 cells in organotypic hippocampal slices during in vitro ischemia. Brain Res 1316:92\u2013100. https:\/\/doi.org\/10.1016\/j.brainres.2009.11.087","journal-title":"Brain Res"},{"issue":"5","key":"5706_CR63","doi-asserted-by":"publisher","first-page":"H1011","DOI":"10.1152\/ajpheart.00664.2017","volume":"314","author":"L Weise-Cross","year":"2018","unstructured":"Weise-Cross L, Sands MA, Sheak JR et al (2018) Actin polymerization contributes to enhanced pulmonary vasoconstrictor reactivity after chronic hypoxia. Am J Physiol Heart Circ Physiol 314(5):H1011\u2013H1021. https:\/\/doi.org\/10.1152\/ajpheart.00664.2017","journal-title":"Am J Physiol Heart Circ Physiol"},{"issue":"2","key":"5706_CR64","doi-asserted-by":"publisher","first-page":"181","DOI":"10.1016\/s0014-4835(03)00118-0","volume":"77","author":"X Liu","year":"2003","unstructured":"Liu X, Wu Z, Sheibani N et al (2003) Low dose latrunculin-A inhibits dexamethasone-induced changes in the actin cytoskeleton and alters extracellular matrix protein expression in cultured human trabecular meshwork cells. Exp Eye Res 77(2):181\u2013188. https:\/\/doi.org\/10.1016\/s0014-4835(03)00118-0","journal-title":"Exp Eye Res"},{"issue":"4","key":"5706_CR65","doi-asserted-by":"publisher","first-page":"3395","DOI":"10.3892\/mmr.2019.10584","volume":"20","author":"L Li","year":"2019","unstructured":"Li L, Liu B (2019) ROCK inhibitor Y-27632 protects rats against cerebral ischemia\/reperfusion-induced behavioral deficits and hippocampal damage. Mol Med Rep 20(4):3395\u20133405. https:\/\/doi.org\/10.3892\/mmr.2019.10584","journal-title":"Mol Med Rep"},{"issue":"2","key":"5706_CR66","doi-asserted-by":"publisher","first-page":"201","DOI":"10.1016\/j.mcn.2009.11.001","volume":"43","author":"J Ding","year":"2010","unstructured":"Ding J, Li QY, Yu JZ et al (2010) Fasudil, a Rho kinase inhibitor, drives mobilization of adult neural stem cells after hypoxia\/reoxygenation injury in mice. Mol Cell Neurosci 43(2):201\u2013208. https:\/\/doi.org\/10.1016\/j.mcn.2009.11.001","journal-title":"Mol Cell Neurosci"},{"issue":"1","key":"5706_CR67","doi-asserted-by":"publisher","first-page":"337","DOI":"10.1124\/jpet.110.177675","volume":"338","author":"A Koumura","year":"2011","unstructured":"Koumura A, Hamanaka J, Kawasaki K et al (2011) Fasudil and ozagrel in combination show neuroprotective effects on cerebral infarction after murine middle cerebral artery occlusion. J Pharmacol Exp Ther 338(1):337\u2013344. https:\/\/doi.org\/10.1124\/jpet.110.177675","journal-title":"J Pharmacol Exp Ther"},{"issue":"1","key":"5706_CR68","doi-asserted-by":"publisher","first-page":"90","DOI":"10.1186\/s12987-025-00701-5","volume":"22","author":"A Blase","year":"2025","unstructured":"Blase A, di Girasole CG, Benjamin L et al (2025) Phased blood-brain barrier disruption in ischaemic stroke: implications for therapy? Fluids Barriers CNS 22(1):90. https:\/\/doi.org\/10.1186\/s12987-025-00701-5","journal-title":"Fluids Barriers CNS"},{"issue":"11","key":"5706_CR69","doi-asserted-by":"publisher","first-page":"3590","DOI":"10.1038\/s41380-024-02626-1","volume":"29","author":"W Du","year":"2024","unstructured":"Du W, Chen H, Gr\u00f3f I et al (2024) Antidepressant-induced membrane trafficking regulates blood-brain barrier permeability. Mol Psychiatry 29(11):3590\u20133598. https:\/\/doi.org\/10.1038\/s41380-024-02626-1","journal-title":"Mol Psychiatry"},{"issue":"6","key":"5706_CR70","doi-asserted-by":"publisher","first-page":"1099","DOI":"10.33549\/physiolres.935432","volume":"73","author":"P Kozler","year":"2024","unstructured":"Kozler P, Mare\u0161ov\u00e1 D, Pokorn\u00fd J (2024) Assessment of blood-brain barrier permeability in a cerebral ischemia-reperfusion model in rats; a pilot study. Physiol Res 73(6):1099\u20131105. https:\/\/doi.org\/10.33549\/physiolres.935432","journal-title":"Physiol Res"},{"key":"5706_CR71","doi-asserted-by":"publisher","first-page":"165","DOI":"10.1016\/j.jconrel.2017.03.020","volume":"253","author":"K Shiraishi","year":"2017","unstructured":"Shiraishi K, Wang Z, Kokuryo D et al (2017) A polymeric micelle magnetic resonance imaging (MRI) contrast agent reveals blood-brain barrier (BBB) permeability for macromolecules in cerebral ischemia-reperfusion injury. J Control Release 253:165\u2013171. https:\/\/doi.org\/10.1016\/j.jconrel.2017.03.020","journal-title":"J Control Release"}],"container-title":["Molecular and Cellular Biochemistry"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1007\/s11010-026-05706-8.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/article\/10.1007\/s11010-026-05706-8","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1007\/s11010-026-05706-8.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,10,5]],"date-time":"2026-10-05T07:45:19Z","timestamp":1791186319000},"score":1,"resource":{"primary":{"URL":"https:\/\/link.springer.com\/10.1007\/s11010-026-05706-8"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,8,27]]},"references-count":71,"journal-issue":{"issue":"9","published-online":{"date-parts":[[2026,9]]}},"alternative-id":["5706"],"URL":"https:\/\/doi.org\/10.1007\/s11010-026-05706-8","relation":{"has-preprint":[{"id-type":"doi","id":"10.21203\/rs.3.rs-9032259\/v1","asserted-by":"object"}]},"ISSN":["1573-4919"],"issn-type":[{"value":"1573-4919","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,8,27]]},"assertion":[{"value":"4 March 2026","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"11 August 2026","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"27 August 2026","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"The authors declared no potential conflicts of interest with respect to the research, authorship, and\/or publication of this article.","order":1,"name":"Ethics","label":"Competing interests","group":{"name":"EthicsHeading","label":"Declarations"}}]}}