{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,26]],"date-time":"2026-08-26T07:23:41Z","timestamp":1787729021793,"version":"build-2784847793"},"reference-count":62,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2026,8,26]],"date-time":"2026-08-26T00:00:00Z","timestamp":1787702400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":["frontiersin.org"],"crossmark-restriction":true},"short-container-title":["Front. Cell Dev. Biol."],"abstract":"<jats:p>Neurodegenerative, cardiovascular, and regenerative disorders remain major causes of disability and mortality worldwide, necessitating the development of multimodal therapeutic strategies. Tanshinone IIA (Tan IIA), a bioactive diterpenoid isolated from Salvia miltiorrhiza, possesses potent antioxidant, anti-inflammatory, anti-apoptotic, and pro-angiogenic properties. Mesenchymal stem cells (MSCs) have emerged as promising regenerative therapeutics owing to their differentiation potential, immunomodulatory capacity, and paracrine effects. Growing evidence indicates that the combination of Tan IIA and MSCs exerts synergistic therapeutic effects by enhancing MSC survival, proliferation, migration, and regenerative function while mitigating oxidative stress and inflammatory injury. This review summarizes current evidence regarding the combined effects of Tan IIA and MSCs on neuroprotection, cardiovascular repair, tissue regeneration, muscle protein synthesis, and gut microbiota modulation across neurodegenerative, cardiovascular, and regenerative disease models. The underlying molecular mechanisms involve the regulation of PI3K\/Akt, Wnt\/\u03b2-catenin, TGF-\u03b2\/Smad, and Nrf2 signaling pathways, together with improvements in angiogenesis, immune homeostasis, mitochondrial function, and cellular metabolism. Emerging evidence also suggests that modulation of the gut microbiota may contribute to enhanced systemic immune regulation and regenerative outcomes, whereas improvements in muscle protein synthesis may support tissue recovery and functional restoration. Although preclinical findings are promising, further mechanistic investigations and well-designed clinical trials are required to optimize treatment protocols and establish the efficacy, safety, and translational potential of this combinational therapeutic strategy.<\/jats:p>","DOI":"10.3389\/fcell.2026.1883952","type":"journal-article","created":{"date-parts":[[2026,8,26]],"date-time":"2026-08-26T06:54:19Z","timestamp":1787727259000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":0,"title":["Synergistic effects of tanshinone IIA and mesenchymal stem cells on neuroprotection, cardiovascular repair, tissue regeneration, muscle protein synthesis, and gut microbiota modulation across neurodegenerative, cardiovascular, and regenerative disease models"],"prefix":"10.3389","volume":"14","author":[{"given":"Wenzhe","family":"Wang","sequence":"first","affiliation":[{"name":"School of Medicine, Pingdingshan University","place":["Pingdingshan, Henan, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Hongyan","family":"Gao","sequence":"additional","affiliation":[{"name":"School of Medicine, Pingdingshan University","place":["Pingdingshan, Henan, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yuan","family":"Wang","sequence":"additional","affiliation":[{"name":"Internal Medicine, Brookdale University Hospital Medical Center","place":["New York, NY, United States"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1965","published-online":{"date-parts":[[2026,8,26]]},"reference":[{"key":"B1","doi-asserted-by":"publisher","first-page":"41","DOI":"10.1016\/j.diff.2016.02.005","article-title":"Key transcription factors in the differentiation of mesenchymal stem cells","volume":"92","author":"Almalki","year":"2016","journal-title":"Differentiation"},{"key":"B2","doi-asserted-by":"publisher","first-page":"98","DOI":"10.3389\/fncel.2016.00098","article-title":"Inflammogenesis of secondary spinal cord injury","volume":"10","author":"Anwar","year":"2016","journal-title":"Front. Cell Neurosci."},{"key":"B3","doi-asserted-by":"publisher","first-page":"534","DOI":"10.1007\/s12035-026-05819-7","article-title":"Reprogramming the inflammatory response to promote neural stem cell function after spinal cord injury","volume":"63","author":"Ardah","year":"2026","journal-title":"Mol. Neurobiol."},{"key":"B4","doi-asserted-by":"publisher","first-page":"61","DOI":"10.1097\/wnr.0000000000001755","article-title":"Mesenchymal stem cells after the proprocessing of tanshinone IIA attenuate cognitive deficits and oxidative stress injury in an amyloid \u03b2-peptide (25-35)-induced rodent model of alzheimer's disease","volume":"33","author":"Ba","year":"2022","journal-title":"Neuroreport"},{"key":"B5","doi-asserted-by":"publisher","first-page":"53","DOI":"10.62347\/YGXA7976","article-title":"Mesenchymal stem cells in injury repair of vital organs: from mechanism to clinical application","volume":"14","author":"Bao","year":"2025","journal-title":"Am. J. Stem Cells"},{"key":"B6","doi-asserted-by":"publisher","first-page":"831","DOI":"10.1007\/s12026-016-8798-6","article-title":"Immunomodulatory effects of mesenchymal stromal cells-derived exosome","volume":"64","author":"Chen","year":"2016","journal-title":"Immunol. Res."},{"key":"B7","doi-asserted-by":"publisher","first-page":"3239","DOI":"10.2147\/ndt.s265478","article-title":"Tanshinone IIA promotes M2 microglia by ER\u03b2\/IL-10 pathway and attenuates neuronal loss in mouse TBI model","volume":"16","author":"Chen","year":"2020","journal-title":"Neuropsychiatr. Dis. Treat."},{"key":"B8","doi-asserted-by":"publisher","first-page":"309","DOI":"10.1186\/s13018-024-04793-x","article-title":"Local application of tanshinone IIA protects mesenchymal stem cells from apoptosis and promotes fracture healing in ovariectomized mice","volume":"19","author":"Cheng","year":"2024","journal-title":"J. Orthop. Surg. Res."},{"key":"B9","doi-asserted-by":"publisher","first-page":"591065","DOI":"10.3389\/fimmu.2020.591065","article-title":"Immunomodulatory and regenerative effects of mesenchymal stem cells and extracellular vesicles: therapeutic outlook for inflammatory and degenerative diseases","volume":"11","author":"Dabrowska","year":"2020","journal-title":"Front. Immunol."},{"key":"B10","doi-asserted-by":"publisher","first-page":"270","DOI":"10.1097\/TP.0b013e3181c6ffbe","article-title":"Mesenchymal stem-cell immunosuppressive capabilities: therapeutic implications in islet transplantation","volume":"89","author":"Ding","year":"2010","journal-title":"Transplantation"},{"key":"B11","doi-asserted-by":"publisher","first-page":"1620152","DOI":"10.3389\/fphar.2025.1620152","article-title":"Harnessing the therapeutic value of tanshinone IIA: a breakthrough therapy in cardiovascular diseases","volume":"16","author":"Du","year":"2025","journal-title":"Front. Pharmacol."},{"key":"B12","doi-asserted-by":"publisher","first-page":"100841","DOI":"10.1016\/j.apmt.2020.100841","article-title":"Tanshinone IIA-Loaded aligned microfibers facilitate stem cell recruitment and capillary formation by inducing M2 macrophage polarization","volume":"21","author":"Gao","year":"2020","journal-title":"Appl. Mater. Today"},{"key":"B13","doi-asserted-by":"publisher","first-page":"7791","DOI":"10.3390\/ijms25147791","article-title":"Applications of modified mesenchymal stem cells as targeted systems against tumor cells","volume":"25","author":"Garza Trevi\u00f1o","year":"2024","journal-title":"Int. J. Mol. Sci."},{"key":"B14","doi-asserted-by":"publisher","first-page":"526","DOI":"10.4252\/wjsc.v6.i5.526","article-title":"Mesenchymal stem cells: emerging mechanisms of immunomodulation and therapy","volume":"6","author":"Glenn","year":"2014","journal-title":"World J. Stem Cells"},{"key":"B15","doi-asserted-by":"publisher","first-page":"92","DOI":"10.1038\/s41392-022-00932-0","article-title":"The secretion profile of mesenchymal stem cells and potential applications in treating human diseases","volume":"7","author":"Han","year":"2022","journal-title":"Signal Transduct. Target Ther."},{"key":"B16","doi-asserted-by":"publisher","first-page":"262","DOI":"10.1038\/s41392-025-02313-9","article-title":"Mesenchymal stem cells in treating human diseases: molecular mechanisms and clinical studies","volume":"10","author":"Han","year":"2025","journal-title":"Signal Transduct. Target Ther."},{"key":"B17","first-page":"8786","article-title":"Acute myocardial infarction and myocardial ischemia-reperfusion injury: a comparison","volume":"8","author":"Hashmi","year":"2015","journal-title":"Int. J. Clin. Exp. Pathol."},{"key":"B18","doi-asserted-by":"publisher","first-page":"284","DOI":"10.1186\/s12974-021-02337-2","article-title":"Inflammation after spinal cord injury: a review of the critical timeline of signaling cues and cellular infiltration","volume":"18","author":"Hellenbrand","year":"2021","journal-title":"J. Neuroinflammation"},{"key":"B19","doi-asserted-by":"publisher","first-page":"1338","DOI":"10.3390\/ph18091338","article-title":"Pharmacological effects and mechanisms of tanshinone IIA in bone injury repair","volume":"18","author":"Hu","year":"2025","journal-title":"Pharm. (Basel)"},{"key":"B20","doi-asserted-by":"publisher","first-page":"465","DOI":"10.1089\/cell.2011.0045","article-title":"Developmental definition of MSCs: new insights into pending questions","volume":"13","author":"Huang","year":"2011","journal-title":"Cell Reprogr."},{"key":"B21","doi-asserted-by":"publisher","first-page":"48","DOI":"10.1016\/j.bbr.2019.03.001","article-title":"Neuroprotective effect of tanshinone IIA-Incubated mesenchymal stem cells on A\u03b2(25-35)-induced neuroinflammation","volume":"365","author":"Huang","year":"2019","journal-title":"Behav. Brain Res."},{"key":"B22","doi-asserted-by":"publisher","first-page":"9977610","DOI":"10.1155\/2022\/9977610","article-title":"Tanshinone \u0399\u0399A\u2010Incubated mesenchymal stem cells inhibit lipopolysaccharide\u2010induced inflammation of N9 cells through TREM2 signaling pathway","volume":"2022","author":"Huang","year":"2022","journal-title":"Stem Cells Int."},{"key":"B23","doi-asserted-by":"publisher","first-page":"2520","DOI":"10.1038\/s41598-023-29282-9","article-title":"Tanshinone IIA-Loaded nanoparticles and neural stem cell combination therapy improves gut homeostasis and recovery in a pig ischemic stroke model","volume":"13","author":"Jeon","year":"2023","journal-title":"Sci. Rep."},{"key":"B24","doi-asserted-by":"publisher","first-page":"5334","DOI":"10.2174\/1381612825666190119130441","article-title":"Human mesenchymal stem cell-derived exosomes reduce ischemia\/reperfusion injury by the inhibitions of apoptosis and autophagy","volume":"24","author":"Jiang","year":"2018","journal-title":"Curr. Pharm. Des."},{"key":"B25","doi-asserted-by":"publisher","first-page":"202","DOI":"10.3389\/fphar.2019.00202","article-title":"Tanshinones, critical pharmacological components in Salvia miltiorrhiza","volume":"10","author":"Jiang","year":"2019","journal-title":"Front. Pharmacol."},{"key":"B26","doi-asserted-by":"publisher","first-page":"155","DOI":"10.1186\/s13287-023-03374-9","article-title":"Recent advances in pre-conditioned mesenchymal stem\/stromal cell (MSCs) therapy in organ failure; a comprehensive review of preclinical studies","volume":"14","author":"Kahrizi","year":"2023","journal-title":"Stem Cell Res. Ther."},{"key":"B27","doi-asserted-by":"publisher","first-page":"1061","DOI":"10.1093\/stcltm\/szac062","article-title":"Tanshinone IIA-loaded nanoparticle and neural stem cell therapy enhances recovery in a pig ischemic stroke model","volume":"11","author":"Kaiser","year":"2022","journal-title":"Stem Cells Transl. Med."},{"key":"B28","doi-asserted-by":"publisher","first-page":"175","DOI":"10.1016\/j.neulet.2016.11.024","article-title":"Synergistic effect of tanshinone IIA and mesenchymal stem cells on preventing learning and memory deficits via anti-apoptosis, attenuating tau phosphorylation and enhancing the activity of central cholinergic system in vascular dementia","volume":"637","author":"Kong","year":"2017","journal-title":"Neurosci. Lett."},{"key":"B29","doi-asserted-by":"publisher","first-page":"13","DOI":"10.1016\/j.jconrel.2023.04.014","article-title":"Tanshinone IIA enhances the therapeutic efficacy of mesenchymal stem cells derived exosomes in myocardial ischemia\/reperfusion injury via up-regulating miR-223-5p","volume":"358","author":"Li","year":"2023","journal-title":"J. Control Release"},{"key":"B30","doi-asserted-by":"publisher","first-page":"109904","DOI":"10.1016\/j.abb.2024.109904","article-title":"Tanshinone IIA attenuates osteoarthritis via inhibiting aberrant angiogenesis in subchondral bone","volume":"753","author":"Li","year":"2024","journal-title":"Archives Biochem. Biophysics"},{"key":"B31","doi-asserted-by":"publisher","first-page":"335","DOI":"10.1186\/s13287-024-03952-5","article-title":"Exosomes as promising bioactive materials in the treatment of spinal cord injury","volume":"15","author":"Li","year":"2024","journal-title":"Stem Cell Res. Ther."},{"key":"B32","doi-asserted-by":"publisher","first-page":"149235","DOI":"10.1016\/j.bbrc.2023.149235","article-title":"Tanshinone IIA promotes the proliferation and differentiation ability of primary muscle stem cells via MAPK and akt signaling","volume":"689","author":"Liu","year":"2023","journal-title":"Biochem. Biophys. Res. Commun."},{"key":"B33","doi-asserted-by":"publisher","first-page":"259","DOI":"10.20517\/evcna.2025.142","article-title":"Extracellular vesicle-based therapeutic strategies for spinal cord injury","volume":"7","author":"Liu","year":"2026","journal-title":"Extracell. Vesicles Circ. Nucl. Acids"},{"key":"B34","first-page":"3425","article-title":"Tanshinone II combined with bone marrow mesenchymal stem cell transplantation promotes nerve regeneration following cerebral infarction","volume":"20","author":"Lu","year":"2016","journal-title":"Chin. J. Tissue Eng. Res."},{"key":"B35","doi-asserted-by":"publisher","first-page":"96","DOI":"10.1016\/j.bioactmat.2026.02.024","article-title":"Targeting the osteoporotic bone microenvironment: mechanistic insight and therapeutic biomaterials for accelerating bone regeneration","volume":"62","author":"Lu","year":"2026","journal-title":"Bioact. Mater"},{"key":"B36","doi-asserted-by":"publisher","first-page":"814","DOI":"10.4252\/wjsc.v12.i8.814","article-title":"Mesenchymal stem cell-derived exosomes: toward cell-free therapeutic strategies in regenerative medicine","volume":"12","author":"Ma","year":"2020","journal-title":"World J. Stem Cells"},{"key":"B37","doi-asserted-by":"publisher","first-page":"4281","DOI":"10.2741\/3528","article-title":"Methodology, biology and clinical applications of mesenchymal stem cells","volume":"14","author":"Meirelles Lda","year":"2009","journal-title":"Front. Biosci. Landmark Ed."},{"key":"B38","doi-asserted-by":"publisher","first-page":"461718","DOI":"10.1155\/2012\/461718","article-title":"Mesenchymal stem cells isolated from adipose and other tissues: basic biological properties and clinical applications","volume":"2012","author":"Orbay","year":"2012","journal-title":"Stem Cells Int."},{"key":"B39","doi-asserted-by":"publisher","DOI":"10.3390\/pharmaceutics18030346","article-title":"Mechanistic insights into the cardioprotective effects of mesenchymal stem cell-derived exosomes in myocardial ischemic injury: a systematic review","volume":"18","author":"Othman","year":"2026","journal-title":"Pharmaceutics"},{"key":"B40","doi-asserted-by":"publisher","first-page":"3084","DOI":"10.3390\/ijms26073084","article-title":"The therapeutic use and potential of MSCs: advances in regenerative medicine","volume":"26","author":"P\u00eenzariu","year":"2025","journal-title":"Int. J. Mol. Sci."},{"key":"B41","doi-asserted-by":"publisher","first-page":"114011","DOI":"10.1016\/j.biopha.2022.114011","article-title":"Multiple strategies enhance the efficacy of MSCs transplantation for spinal cord injury","volume":"157","author":"Pang","year":"2023","journal-title":"Biomed. and Pharmacother."},{"key":"B42","doi-asserted-by":"publisher","first-page":"419","DOI":"10.1007\/s13346-022-01222-6","article-title":"Bone regeneration in osteoporosis: opportunities and challenges","volume":"13","author":"Patel","year":"2023","journal-title":"Drug Deliv. Transl. Res."},{"key":"B43","doi-asserted-by":"publisher","first-page":"51","DOI":"10.1080\/08923970903120997","article-title":"Tanshinone IIA protects against immune-mediated liver injury through activation of T-cell subsets and regulation of cytokines","volume":"32","author":"Qin","year":"2010","journal-title":"Immunopharmacol. Immunotoxicol."},{"key":"B44","doi-asserted-by":"publisher","first-page":"10","DOI":"10.3390\/life16010010","article-title":"Cell surface markers of mesenchymal stem cells: current knowledge and advances in characterization technologies","volume":"16","author":"Santoro","year":"2025","journal-title":"Life (Basel)"},{"key":"B45","doi-asserted-by":"publisher","first-page":"3924858","DOI":"10.1155\/2016\/3924858","article-title":"Preconditioning of human mesenchymal stem cells to enhance their regulation of the immune response","volume":"2016","author":"Saparov","year":"2016","journal-title":"Stem Cells Int."},{"key":"B46","doi-asserted-by":"publisher","first-page":"45511","DOI":"10.1021\/acsami.5c09766","article-title":"Exosomes derived from tanshinone IIA-pretreated umbilical cord mesenchymal stem cells repair traumatic spinal cord injury by miR-223-5p\/USP8\/NLRP3 axis","volume":"17","author":"Shen","year":"2025","journal-title":"ACS Appl. Mater Interfaces"},{"key":"B47","doi-asserted-by":"publisher","first-page":"3036","DOI":"10.3892\/mmr.2016.4886","article-title":"Effects of tanshinone IIA on fibrosis in a rat model of cirrhosis through heme oxygenase-1, inflammation, oxidative stress and apoptosis","volume":"13","author":"Shu","year":"2016","journal-title":"Mol. Med. Rep."},{"key":"B48","doi-asserted-by":"publisher","first-page":"1509418","DOI":"10.3389\/fphar.2025.1509418","article-title":"Optimizing therapeutic outcomes: preconditioning strategies for MSC-Derived extracellular vesicles","volume":"16","author":"Song","year":"2025","journal-title":"Front. Pharmacol."},{"key":"B49","doi-asserted-by":"publisher","first-page":"443","DOI":"10.1016\/j.phymed.2010.10.009","article-title":"Tanshinone IIA increases recruitment of bone marrow mesenchymal stem cells to infarct region via up-regulating stromal cell-derived factor-1\/CXC chemokine receptor 4 axis in a myocardial ischemia model","volume":"18","author":"Tong","year":"2011","journal-title":"Phytomedicine"},{"key":"B50","doi-asserted-by":"publisher","first-page":"26","DOI":"10.1186\/s40659-026-00687-w","article-title":"The progress and challenges of mesenchymal stromal cell-based therapy for diabetes and its complications","volume":"59","author":"Tu","year":"2026","journal-title":"Biol. Res."},{"key":"B51","doi-asserted-by":"publisher","first-page":"131","DOI":"10.1007\/s12013-025-01895-z","article-title":"Mesenchymal stem cell-based therapies: challenges and enhancement strategies","volume":"84","author":"Wong","year":"2026","journal-title":"Cell Biochem. Biophys."},{"key":"B52","doi-asserted-by":"publisher","first-page":"e29424","DOI":"10.1016\/j.heliyon.2024.e29424","article-title":"Neuroprotective effect of tanshinone IIA-Modified mesenchymal stem cells in a lipopolysaccharide-induced neuroinflammation model","volume":"10","author":"Wu","year":"2024","journal-title":"Heliyon"},{"key":"B53","doi-asserted-by":"publisher","first-page":"1141601","DOI":"10.3389\/fimmu.2023.1141601","article-title":"Mesenchymal stem cells in the treatment of spinal cord injury: mechanisms, current advances and future challenges","volume":"14","author":"Xia","year":"2023","journal-title":"Front. Immunol."},{"key":"B54","first-page":"3986","article-title":"First-trimester human decidual mesenchymal stem cells combined with tanshinone IIA in the treatment of intrauterine adhesions in rats","volume":"26","author":"Xu","year":"2022","journal-title":"Chin. J. Tissue Eng. Res."},{"key":"B55","doi-asserted-by":"publisher","first-page":"5769350","DOI":"10.1155\/2022\/5769350","article-title":"Tanshinone IIA ameliorates nonalcoholic steatohepatitis in mice by modulating neutrophil extracellular traps and hepatocyte apoptosis","volume":"2022","author":"Xu","year":"2022","journal-title":"Evid. Based Complement. Altern. Med."},{"key":"B56","doi-asserted-by":"publisher","first-page":"207","DOI":"10.1186\/s10020-024-00967-9","article-title":"Mesenchymal stem cells lineage and their role in disease development","volume":"30","author":"Xu","year":"2024","journal-title":"Mol. Med."},{"key":"B57","doi-asserted-by":"publisher","first-page":"110815","DOI":"10.1016\/j.biopha.2020.110815","article-title":"Tanshinone IIA exerts therapeutic effects by acting on endogenous stem cells in rats with liver cirrhosis","volume":"132","author":"Yang","year":"2020","journal-title":"Biomed. Pharmacother."},{"key":"B58","doi-asserted-by":"publisher","first-page":"103560","DOI":"10.4252\/wjsc.v17.i3.103560","article-title":"Immunomodulatory effects and clinical application of exosomes derived from mesenchymal stem cells","volume":"17","author":"Yi","year":"2025","journal-title":"World J. Stem Cells"},{"key":"B59","doi-asserted-by":"publisher","first-page":"2667","DOI":"10.1007\/s12015-025-10950-4","article-title":"A comparative study of mesenchymal stem cells from various sources in endometrial repair: the significant advantages of decidual mesenchymal stem cells","volume":"21","author":"Yu","year":"2025","journal-title":"Stem Cell Rev. Rep."},{"key":"B60","doi-asserted-by":"publisher","first-page":"963689720908500","DOI":"10.1177\/0963689720908500","article-title":"Mesenchymal stem cell-derived extracellular vesicles in tissue regeneration","volume":"29","author":"Zhang","year":"2020","journal-title":"Cell Transpl."},{"key":"B61","doi-asserted-by":"publisher","first-page":"193","DOI":"10.1186\/s12967-018-1571-y","article-title":"Tanshinone IIA promotes the differentiation of bone marrow mesenchymal stem cells into neuronal-like cells in a spinal cord injury model","volume":"16","author":"Zhang","year":"2018","journal-title":"J. Transl. Med."},{"key":"B62","doi-asserted-by":"publisher","DOI":"10.3390\/jcm8071025","article-title":"The immunomodulatory functions of mesenchymal stromal\/stem cells mediated via paracrine activity","volume":"8","author":"Zhou","year":"2019","journal-title":"J. Clin. Med."}],"container-title":["Frontiers in Cell and Developmental Biology"],"original-title":[],"link":[{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fcell.2026.1883952\/full","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,8,26]],"date-time":"2026-08-26T06:54:21Z","timestamp":1787727261000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fcell.2026.1883952\/full"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,8,26]]},"references-count":62,"alternative-id":["10.3389\/fcell.2026.1883952"],"URL":"https:\/\/doi.org\/10.3389\/fcell.2026.1883952","relation":{},"ISSN":["2296-634X"],"issn-type":[{"value":"2296-634X","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,8,26]]},"article-number":"1883952"}}