{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,7,30]],"date-time":"2026-07-30T13:18:36Z","timestamp":1785417516236,"version":"3.56.0"},"reference-count":52,"publisher":"MDPI AG","issue":"14","license":[{"start":{"date-parts":[[2026,7,9]],"date-time":"2026-07-09T00:00:00Z","timestamp":1783555200000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"crossref","award":["32272965"],"award-info":[{"award-number":["32272965"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"crossref"}]},{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"crossref","award":["32472982"],"award-info":[{"award-number":["32472982"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"crossref"}]},{"award":["32272965"],"award-info":[{"award-number":["32272965"]}],"id":[{"id":"https:\/\/ror.org\/01h0zpd94","id-type":"ROR","asserted-by":"publisher"}]},{"award":["32472982"],"award-info":[{"award-number":["32472982"]}],"id":[{"id":"https:\/\/ror.org\/01h0zpd94","id-type":"ROR","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100012131","name":"Science and Technology Program Project of Liaoning Province","doi-asserted-by":"publisher","award":["2023011979-JH3\/4600"],"award-info":[{"award-number":["2023011979-JH3\/4600"]}],"id":[{"id":"10.13039\/501100012131","id-type":"DOI","asserted-by":"publisher"}]},{"award":["2023011979-JH3\/4600"],"award-info":[{"award-number":["2023011979-JH3\/4600"]}],"id":[{"id":"https:\/\/ror.org\/02f09cx53","id-type":"ROR","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Cells"],"abstract":"<jats:p>Background: Metabolic dysfunction-associated steatotic liver disease (MASLD) is now the most prevalent chronic liver disease worldwide, imposing a severe public health burden. Its core pathological hallmark is excessive hepatic lipid accumulation driven by systemic metabolic dysregulation. Concomitant hepatocellular injury impairs hepatic lactate clearance, leading to aberrant lactate buildup in the liver microenvironment. However, the causal role of lactate in exacerbating liver lipid metabolism dysfunction and driving the progression of MASLD remains unclear. Methods: First, we performed a comprehensive bioinformatic analysis of publicly available transcriptomic datasets. Mining of the Gene Expression Omnibus (GEO) database showed that lactate dehydrogenase (LDH) expression was significantly upregulated in liver tissues from both metabolic dysfunction-associated fatty liver disease (MASLD) patients and MASLD mouse models. Next, network pharmacology approaches were employed to predict putative molecular targets that could mediate lactate\u2019s biological effects. Gene Ontology (GO) and Kyoto Encyclopedia of Genes and Genomes (KEGG) enrichment analyses indicated that these candidate targets were predominantly enriched in pathways governing fatty acid metabolism and long-chain fatty acid transport. Molecular docking and molecular dynamics simulations further suggested possible interactions and supported the prioritization of cluster of differentiation 36 (CD36) as candidate lipid metabolism regulators potentially involved in lactate-mediated effects. Finally, liver-specific Ldha knockdown mice (AAV8-TBG-shRNA) and free fatty acid-induced steatotic AML12 hepatocytes were used to investigate the functional relevance of these findings in vivo and in vitro. Results: Network pharmacology analyses preliminarily identified the PPAR signaling pathway as a candidate pathway potentially linking lactate to MASLD. Experimental results showed that exogenous lactate administration was associated with significantly increased lipid accumulation in steatotic AML12 hepatocytes and the livers of MASLD mice, manifested as elevated triglyceride levels and enhanced lipid droplet formation, accompanied by upregulated expression of PPAR\u03b3 and CD36. Conversely, inhibiting endogenous lactate production or silencing PPAR\u03b3 or CD36 attenuated this lipid-accumulation phenotype and significantly reduced intracellular triglyceride levels. Conclusions: In conclusion, these findings indicate that lactate exposure is associated with hepatic lipid accumulation and upregulation of the PPAR\u03b3\/CD36 axis. Pharmacological inhibition or silencing of PPAR\u03b3 or CD36 attenuates this phenotype, suggesting that this pathway may contribute to lactate-associated hepatic steatosis and potentially accelerate MASLD progression.<\/jats:p>","DOI":"10.3390\/cells15141240","type":"journal-article","created":{"date-parts":[[2026,7,9]],"date-time":"2026-07-09T10:16:22Z","timestamp":1783592182000},"page":"1240","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Lactate Aggravates MASLD via PPAR\u03b3\/CD36-Mediated Hepatocellular Fatty Acid Uptake"],"prefix":"10.3390","volume":"15","author":[{"ORCID":"https:\/\/orcid.org\/0009-0001-5197-5911","authenticated-orcid":false,"given":"Wenke","family":"Sun","sequence":"first","affiliation":[{"name":"College of Animal Science and Medicine, Shenyang Agricultural University, No. 120, Dongling Road, Shenyang 110866, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0002-5079-1878","authenticated-orcid":false,"given":"Weiwei","family":"Li","sequence":"additional","affiliation":[{"name":"College of Animal Science and Medicine, Shenyang Agricultural University, No. 120, Dongling Road, Shenyang 110866, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Guangyi","family":"Ouyang","sequence":"additional","affiliation":[{"name":"College of Animal Science and Medicine, Shenyang Agricultural University, No. 120, Dongling Road, Shenyang 110866, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jishuang","family":"San","sequence":"additional","affiliation":[{"name":"College of Animal Science and Medicine, Shenyang Agricultural University, No. 120, Dongling Road, Shenyang 110866, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yue","family":"Zhu","sequence":"additional","affiliation":[{"name":"College of Animal Science and Medicine, Shenyang Agricultural University, No. 120, Dongling Road, Shenyang 110866, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yunheng","family":"Liu","sequence":"additional","affiliation":[{"name":"College of Animal Science and Medicine, Shenyang Agricultural University, No. 120, Dongling Road, Shenyang 110866, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4509-0893","authenticated-orcid":false,"given":"Jiancheng","family":"Yang","sequence":"additional","affiliation":[{"name":"College of Animal Science and Medicine, Shenyang Agricultural University, No. 120, Dongling Road, Shenyang 110866, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Gaofeng","family":"Wu","sequence":"additional","affiliation":[{"name":"College of Animal Science and Medicine, Shenyang Agricultural University, No. 120, Dongling Road, Shenyang 110866, China"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2026,7,9]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"106925","DOI":"10.1016\/j.phrs.2023.106925","article-title":"Natural products in non-alcoholic fatty liver disease (NAFLD): Novel lead discovery for drug development","volume":"196","author":"Wang","year":"2023","journal-title":"Pharmacol. Res."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"4020249","DOI":"10.1155\/2020\/4020249","article-title":"An Overview of Lipid Metabolism and Nonalcoholic Fatty Liver Disease","volume":"2020","author":"Pei","year":"2020","journal-title":"BioMed Res. Int."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"122714","DOI":"10.1016\/j.lfs.2024.122714","article-title":"PIM1 alleviated liver oxidative stress and NAFLD by regulating the NRF2\/HO-1\/NQO1 pathway","volume":"349","author":"Yang","year":"2024","journal-title":"Life Sci."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"2263","DOI":"10.1001\/jama.2015.5370","article-title":"Nonalcoholic fatty liver disease: A systematic review","volume":"313","author":"Rinella","year":"2015","journal-title":"JAMA"},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"1250","DOI":"10.1038\/cr.2014.120","article-title":"Liver Med23 ablation improves glucose and lipid metabolism through modulating FOXO1 activity","volume":"24","author":"Chu","year":"2014","journal-title":"Cell Res."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"757","DOI":"10.1016\/j.cmet.2018.03.008","article-title":"The Science and Translation of Lactate Shuttle Theory","volume":"27","author":"Brooks","year":"2018","journal-title":"Cell Metab."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"149","DOI":"10.1007\/s10555-019-09787-4","article-title":"Acidosis and cancer: From mechanism to neutralization","volume":"38","author":"Estrella","year":"2019","journal-title":"Cancer Metastasis Rev."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"1127","DOI":"10.1016\/j.mayocp.2013.06.012","article-title":"Etiology and therapeutic approach to elevated lactate levels","volume":"88","author":"Andersen","year":"2013","journal-title":"Mayo Clin. Proc."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"E534","DOI":"10.1152\/ajpendo.00462.2004","article-title":"Sodium lactate increases LPS-stimulated MMP and cytokine expression in U937 histiocytes by enhancing AP-1 and NF-kappaB transcriptional activities","volume":"289","author":"Nareika","year":"2005","journal-title":"Am. J. Physiol. Endocrinol. Metab."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"1101","DOI":"10.1016\/j.bbrc.2020.06.126","article-title":"Secreted phosphoglucose isomerase is a novel biomarker of nonalcoholic fatty liver in mice and humans","volume":"529","author":"Lin","year":"2020","journal-title":"Biochem. Biophys. Res. Commun."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"112","DOI":"10.1016\/j.bbrc.2016.08.118","article-title":"Metabolic biomarkers for non-alcoholic fatty liver disease induced by high-fat diet: In vivo magnetic resonance spectroscopy of hyperpolarized [1-13C] pyruvate","volume":"482","author":"Moon","year":"2017","journal-title":"Biochem. Biophys. Res. Commun."},{"key":"ref_12","doi-asserted-by":"crossref","unstructured":"Huang, W., Su, J., Chen, X., Li, Y., Xing, Z., Guo, L., Li, S., and Zhang, J. (2023). High-Intensity Interval Training Induces Protein Lactylation in Different Tissues of Mice with Specificity and Time Dependence. Metabolites, 13.","DOI":"10.3390\/metabo13050647"},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"59","DOI":"10.3892\/ijmm.2024.5383","article-title":"Role of lactate and lactate metabolism in liver diseases (Review)","volume":"54","author":"Yao","year":"2024","journal-title":"Int. J. Mol. Med."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"3925","DOI":"10.12659\/MSM.902491","article-title":"Kaempferol Alleviates the Interleukin-1\u03b2-Induced Inflammation in Rat Osteoarthritis Chondrocytes via Suppression of NF-\u03baB","volume":"23","author":"Zhuang","year":"2017","journal-title":"Med. Sci. Monit."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"166","DOI":"10.3892\/ijmm.2021.4999","article-title":"Role of the S100 protein family in liver disease (Review)","volume":"48","author":"Yao","year":"2021","journal-title":"Int. J. Mol. Med."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"1605","DOI":"10.1002\/hep.31173","article-title":"Changes in the Global Burden of Chronic Liver Diseases From 2012 to 2017: The Growing Impact of NAFLD","volume":"72","author":"Paik","year":"2020","journal-title":"Hepatology"},{"key":"ref_17","unstructured":"Saxena, R. (2011). Practical Hepatic Pathology: A Diagnostic Approach, Elsevier."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"6","DOI":"10.21037\/jlpm-20-93","article-title":"Aspartate aminotransferase and cardiovascular disease\u2014A narrative review","volume":"6","author":"Ndrepepa","year":"2021","journal-title":"J. Lab. Precis. Med."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"25920","DOI":"10.18632\/aging.203780","article-title":"LDH-A inhibitors as remedies to enhance the anticancer effects of PARP inhibitors in ovarian cancer cells","volume":"13","author":"Xiang","year":"2021","journal-title":"Aging"},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"132","DOI":"10.3109\/07388551.2014.930017","article-title":"Liver cell-targeted delivery of therapeutic molecules","volume":"36","author":"Kang","year":"2016","journal-title":"Crit. Rev. Biotechnol."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"1723184","DOI":"10.1155\/2018\/1723184","article-title":"Prognostic Value of Lactate Dehydrogenase in Patients with Hepatocellular Carcinoma: A Meta-Analysis","volume":"2018","author":"Kong","year":"2018","journal-title":"BioMed Res. Int."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"1939","DOI":"10.1038\/s42255-024-01134-4","article-title":"Inhibition of hepatic oxalate overproduction ameliorates metabolic dysfunction-associated steatohepatitis","volume":"6","author":"Das","year":"2024","journal-title":"Nat. Metab."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"566","DOI":"10.1038\/s42255-020-0243-4","article-title":"Lactate: The ugly duckling of energy metabolism","volume":"2","author":"Rabinowitz","year":"2020","journal-title":"Nat. Metab."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"268","DOI":"10.1038\/s41580-020-0227-y","article-title":"Cellular adaptation to hypoxia through hypoxia inducible factors and beyond","volume":"21","author":"Lee","year":"2020","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"106524","DOI":"10.1016\/j.isci.2023.106524","article-title":"CD36 regulates diurnal glucose metabolism and hepatic clock to maintain glucose homeostasis in mice","volume":"26","author":"Chen","year":"2023","journal-title":"iScience"},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"1695","DOI":"10.1093\/hmg\/ddn060","article-title":"Variants in the CD36 gene associate with the metabolic syndrome and high-density lipoprotein cholesterol","volume":"17","author":"Sherva","year":"2008","journal-title":"Hum. Mol. Genet."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"389","DOI":"10.1016\/j.ymgme.2010.12.012","article-title":"The macrophage Ox-LDL receptor, CD36 and its association with type II diabetes mellitus","volume":"102","author":"Gautam","year":"2011","journal-title":"Mol. Genet. Metab."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"562","DOI":"10.1097\/FJC.0000000000001634","article-title":"Peroxisome Proliferator-Activated Receptor Gamma Regulates Interleukin-6-Induced Lipoprotein (a) Gene Expression in Human HepG2 Cells","volume":"84","author":"Harb","year":"2024","journal-title":"J. Cardiovasc. Pharmacol."},{"key":"ref_29","doi-asserted-by":"crossref","unstructured":"Karunakaran, U., Elumalai, S., Moon, J.S., and Won, K.C. (2021). CD36 Signal Transduction in Metabolic Diseases: Novel Insights and Therapeutic Targeting. Cells, 10.","DOI":"10.3390\/cells10071833"},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"575","DOI":"10.1038\/s41586-019-1678-1","article-title":"Metabolic regulation of gene expression by histone lactylation","volume":"574","author":"Zhang","year":"2019","journal-title":"Nature"},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"150","DOI":"10.1186\/s10020-023-00740-4","article-title":"Swimming exercise ameliorates insulin resistance and nonalcoholic fatty liver by negatively regulating PPAR\u03b3 transcriptional network in mice fed high fat diet","volume":"29","author":"Zhang","year":"2023","journal-title":"Mol. Med."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"3829","DOI":"10.2147\/DDDT.S386982","article-title":"Biological Mechanisms and Related Natural Inhibitors of CD36 in Nonalcoholic Fatty Liver","volume":"16","author":"Feng","year":"2022","journal-title":"Drug Des. Dev. Ther."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"106786","DOI":"10.1016\/j.phrs.2023.106786","article-title":"Roles of the peroxisome proliferator-activated receptors (PPARs) in the pathogenesis of nonalcoholic fatty liver disease (NAFLD)","volume":"192","author":"Qiu","year":"2023","journal-title":"Pharmacol. Res."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"2140","DOI":"10.1194\/jlr.M088419","article-title":"Extrahepatic PPAR\u03b1 modulates fatty acid oxidation and attenuates fasting-induced hepatosteatosis in mice","volume":"59","author":"Brocker","year":"2018","journal-title":"J. Lipid Res."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"557","DOI":"10.1038\/nm.3159","article-title":"PPAR\u03b3 signaling and metabolism: The good, the bad and the future","volume":"19","author":"Ahmadian","year":"2013","journal-title":"Nat. Med."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"289","DOI":"10.1146\/annurev.biochem.77.061307.091829","article-title":"Fat and beyond: The diverse biology of PPARgamma","volume":"77","author":"Tontonoz","year":"2008","journal-title":"Annu. Rev. Biochem."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"72","DOI":"10.1016\/j.tem.2008.11.001","article-title":"Cellular fatty acid uptake: A pathway under construction","volume":"20","author":"Su","year":"2009","journal-title":"Trends Endocrinol. Metab."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"556","DOI":"10.1053\/j.gastro.2007.11.037","article-title":"Hepatic fatty acid transporter Cd36 is a common target of LXR, PXR, and PPARgamma in promoting steatosis","volume":"134","author":"Zhou","year":"2008","journal-title":"Gastroenterology"},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"726","DOI":"10.1053\/j.gastro.2013.11.049","article-title":"Increased de novo lipogenesis is a distinct characteristic of individuals with nonalcoholic fatty liver disease","volume":"146","author":"Lambert","year":"2014","journal-title":"Gastroenterology"},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"924","DOI":"10.1038\/nsmb.1474","article-title":"Structural basis for the activation of PPARgamma by oxidized fatty acids","volume":"15","author":"Itoh","year":"2008","journal-title":"Nat. Struct. Mol. Biol."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"1362","DOI":"10.1126\/science.aaa1299","article-title":"Targeting LDH enzymes with a stiripentol analog to treat epilepsy","volume":"347","author":"Sada","year":"2015","journal-title":"Science"},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"1102","DOI":"10.1016\/j.bbagen.2012.04.016","article-title":"Mitochondrial dysfunction promotes cell migration via reactive oxygen species-enhanced \u03b25-integrin expression in human gastric cancer SC-M1 cells","volume":"1820","author":"Hung","year":"2012","journal-title":"Biochim. Biophys. Acta"},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"139852","DOI":"10.1016\/j.jhazmat.2025.139852","article-title":"PDHA1-mediated H3K18 lactylation is involved in arsenic-induced nonalcoholic fatty liver disease by activating the CD36-NLRP3 inflammasome axis","volume":"498","author":"Li","year":"2025","journal-title":"J. Hazard. Mater."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"137","DOI":"10.1016\/j.omtm.2022.03.006","article-title":"In vivo CRISPR-Cas9 inhibition of hepatic LDH as treatment of primary hyperoxaluria","volume":"25","author":"Salido","year":"2022","journal-title":"Mol. Ther. Methods Clin. Dev."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"574","DOI":"10.1053\/j.gastro.2015.11.043","article-title":"Circadian Clock Control of Liver Metabolic Functions","volume":"150","author":"Reinke","year":"2016","journal-title":"Gastroenterology"},{"key":"ref_46","first-page":"3","article-title":"Circadian clock genes: Their influence on liver metabolism, disease development and treatment (Review)","volume":"33","author":"Wang","year":"2026","journal-title":"Mol. Med. Rep."},{"key":"ref_47","doi-asserted-by":"crossref","unstructured":"Guo, X., Zheng, J., Zhang, S., Jiang, X., Chen, T., Yu, J., Wang, S., Ma, X., and Wu, C. (2021). Advances in Unhealthy Nutrition and Circadian Dysregulation in Pathophysiology of NAFLD. Front. Clin. Diabetes Healthc., 2.","DOI":"10.3389\/fcdhc.2021.691828"},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"120","DOI":"10.1016\/j.jhep.2024.07.034","article-title":"Targeting the liver clock improves fibrosis by restoring TGF-\u03b2 signaling","volume":"82","author":"Crouchet","year":"2025","journal-title":"J. Hepatol."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"305","DOI":"10.1038\/s41392-022-01151-3","article-title":"Lactate metabolism in human health and disease","volume":"7","author":"Li","year":"2022","journal-title":"Signal Transduct. Target. Ther."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"1011","DOI":"10.1016\/j.ajem.2016.02.053","article-title":"Lactate clearance and mortality in septic patients with hepatic dysfunction","volume":"34","author":"Ha","year":"2016","journal-title":"Am. J. Emerg. Med."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"61","DOI":"10.1038\/s42255-022-00710-w","article-title":"Lactylome analysis suggests lactylation-dependent mechanisms of metabolic adaptation in hepatocellular carcinoma","volume":"5","author":"Yang","year":"2023","journal-title":"Nat. Metab."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"850","DOI":"10.1093\/toxres\/tfab068","article-title":"Lactate and blood ammonia on admission as biomarkers to predict the prognosis of patients with acute mushroom poisoning and liver failure: A retrospective study","volume":"10","author":"Gao","year":"2021","journal-title":"Toxicol. Res."}],"container-title":["Cells"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2073-4409\/15\/14\/1240\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,7,11]],"date-time":"2026-07-11T04:17:39Z","timestamp":1783743459000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2073-4409\/15\/14\/1240"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,7,9]]},"references-count":52,"journal-issue":{"issue":"14","published-online":{"date-parts":[[2026,7]]}},"alternative-id":["cells15141240"],"URL":"https:\/\/doi.org\/10.3390\/cells15141240","relation":{},"ISSN":["2073-4409"],"issn-type":[{"value":"2073-4409","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,7,9]]}}}