{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,5,30]],"date-time":"2026-05-30T05:54:15Z","timestamp":1780120455545,"version":"3.54.0"},"reference-count":64,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2025,12,30]],"date-time":"2025-12-30T00:00:00Z","timestamp":1767052800000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0"},{"start":{"date-parts":[[2026,1,3]],"date-time":"2026-01-03T00:00:00Z","timestamp":1767398400000},"content-version":"vor","delay-in-days":4,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0"}],"funder":[{"name":"Science Achievement Scholarship of Thailand"},{"name":"Program Management Unit for Human Resources & Institutional Development, Research and Innovation","award":["B38G670011]"],"award-info":[{"award-number":["B38G670011]"]}]},{"name":"Program Management Unit for Human Resources & Institutional Development, Research and Innovation","award":["B38G670011]"],"award-info":[{"award-number":["B38G670011]"]}]}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Sci Rep"],"DOI":"10.1038\/s41598-025-29207-8","type":"journal-article","created":{"date-parts":[[2025,12,30]],"date-time":"2025-12-30T03:49:42Z","timestamp":1767066582000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":1,"title":["Riboflavin-PLGA nanoparticles enhance the effectiveness of the pyruvate carboxylase inhibitor ZY-444 in targeting lipogenesis in colon and breast cancer"],"prefix":"10.1038","volume":"16","author":[{"given":"Siraprapa","family":"Siritutsoontorn","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Preeyanuch","family":"Manohong","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Natthapat","family":"Sawektreeratana","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Chutima","family":"Kuhakarn","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Kanlaya Prapainop","family":"Katewongsa","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sarawut","family":"Jitrapakdee","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2025,12,30]]},"reference":[{"key":"29207_CR1","doi-asserted-by":"publisher","first-page":"85","DOI":"10.1038\/nrc2981","volume":"11","author":"RA Cairns","year":"2011","unstructured":"Cairns, R. A., Harris, I. S. & Mak, T. W. Regulation of cancer cell metabolism. Nat. Rev. Cancer. 11, 85\u201395. https:\/\/doi.org\/10.1038\/nrc2981 (2011).","journal-title":"Nat. Rev. Cancer"},{"key":"29207_CR2","doi-asserted-by":"publisher","first-page":"297","DOI":"10.1016\/j.ccr.2012.02.014","volume":"21","author":"PS Ward","year":"2012","unstructured":"Ward, P. S. & Thompson, C. B. Metabolic reprogramming: a cancer hallmark even Warburg did not anticipate. Cancer Cell. 21, 297\u2013308. https:\/\/doi.org\/10.1016\/j.ccr.2012.02.014 (2012).","journal-title":"Cancer Cell."},{"key":"29207_CR3","doi-asserted-by":"publisher","first-page":"1029","DOI":"10.1126\/science.1160809","volume":"324","author":"MG Vander Heiden","year":"2009","unstructured":"Vander Heiden, M. G., Cantley, L. C. & Thompson, C. B. Understanding the Warburg effect: the metabolic requirements of cell proliferation. Science 324, 1029\u20131033. https:\/\/doi.org\/10.1126\/science.1160809 (2009).","journal-title":"Science"},{"key":"29207_CR4","doi-asserted-by":"publisher","first-page":"66","DOI":"10.1111\/j.1749-6632.2009.05039.x","volume":"1177","author":"F Weinberg","year":"2009","unstructured":"Weinberg, F. & Chandel, N. S. Mitochondrial metabolism and cancer. Ann. N Y Acad. Sci. 1177, 66\u201373. https:\/\/doi.org\/10.1111\/j.1749-6632.2009.05039.x (2009).","journal-title":"Ann. N Y Acad. Sci."},{"key":"29207_CR5","doi-asserted-by":"publisher","first-page":"141","DOI":"10.1038\/s41573-021-00339-6","volume":"21","author":"ZE Stine","year":"2022","unstructured":"Stine, Z. E., Schug, Z. T., Salvino, J. M. & Dang, C. V. Targeting cancer metabolism in the era of precision oncology. Nat. Rev. Drug Discov. 21, 141\u2013162. https:\/\/doi.org\/10.1038\/s41573-021-00339-6 (2022).","journal-title":"Nat. Rev. Drug Discov"},{"key":"29207_CR6","doi-asserted-by":"publisher","first-page":"115064","DOI":"10.1016\/j.celrep.2024.115064","volume":"43","author":"Y Mao","year":"2024","unstructured":"Mao, Y., Xia, Z., Xia, W. & Jiang, P. Metabolic reprogramming, sensing, and cancer therapy. Cell. Rep. 43, 115064. https:\/\/doi.org\/10.1016\/j.celrep.2024.115064 (2024).","journal-title":"Cell. Rep."},{"key":"29207_CR7","doi-asserted-by":"publisher","first-page":"e0129848","DOI":"10.1371\/journal.pone.0129848","volume":"10","author":"P Phannasil","year":"2015","unstructured":"Phannasil, P. et al. Pyruvate carboxylase is Up-Regulated in breast cancer and essential to support growth and invasion of MDA-MB-231 cells. PLoS One. 10, e0129848. https:\/\/doi.org\/10.1371\/journal.pone.0129848 (2015).","journal-title":"PLoS One"},{"key":"29207_CR8","doi-asserted-by":"publisher","first-page":"6285","DOI":"10.21873\/anticanres.14649","volume":"40","author":"J Ngamkham","year":"2020","unstructured":"Ngamkham, J. et al. Overexpression of pyruvate carboxylase is correlated with colorectal cancer progression and supports growth of invasive colon cancer HT-29 cell line. Anticancer Res. 40, 6285\u20136293. https:\/\/doi.org\/10.21873\/anticanres.14649 (2020).","journal-title":"Anticancer Res."},{"key":"29207_CR9","doi-asserted-by":"publisher","first-page":"722","DOI":"10.1186\/s12885-021-08499-9","volume":"21","author":"C Liu","year":"2021","unstructured":"Liu, C., Zhou, X., Pan, Y., Liu, Y. & Zhang, Y. Pyruvate carboxylase promotes thyroid cancer aggressiveness through fatty acid synthesis. BMC Cancer. 21, 722. https:\/\/doi.org\/10.1186\/s12885-021-08499-9 (2021).","journal-title":"BMC Cancer"},{"key":"29207_CR10","doi-asserted-by":"publisher","first-page":"21874","DOI":"10.18632\/aging.104004","volume":"12","author":"H Shang","year":"2020","unstructured":"Shang, H., Zheng, J. & Tong, J. Integrated analysis of transcriptomic and metabolomic data demonstrates the significant role of pyruvate carboxylase in the progression of ovarian cancer. Aging (Albany NY). 12, 21874\u201321889. https:\/\/doi.org\/10.18632\/aging.104004 (2020).","journal-title":"Aging (Albany NY)"},{"key":"29207_CR11","doi-asserted-by":"publisher","first-page":"100903","DOI":"10.1016\/j.bbrep.2020.100903","volume":"25","author":"K Rattanapornsompong","year":"2021","unstructured":"Rattanapornsompong, K. et al. Impaired G2\/M cell cycle arrest induces apoptosis in pyruvate carboxylase knockdown MDA-MB-231 cells. Biochem. Biophys. Rep. 25, 100903. https:\/\/doi.org\/10.1016\/j.bbrep.2020.100903 (2021).","journal-title":"Biochem. Biophys. Rep."},{"key":"29207_CR12","doi-asserted-by":"publisher","unstructured":"Fan, T. W. et al. Altered regulation of metabolic pathways in human lung cancer discerned by (13)C stable isotope-resolved metabolomics (SIRM). Mol. Cancer. 8 https:\/\/doi.org\/10.1186\/1476-4598-8-41 (2009).","DOI":"10.1186\/1476-4598-8-41"},{"key":"29207_CR13","doi-asserted-by":"publisher","first-page":"687","DOI":"10.1172\/JCI72873","volume":"125","author":"K Sellers","year":"2015","unstructured":"Sellers, K. et al. Pyruvate carboxylase is critical for non-small-cell lung cancer proliferation. J. Clin. Invest. 125, 687\u2013698. https:\/\/doi.org\/10.1172\/JCI72873 (2015).","journal-title":"J. Clin. Invest."},{"key":"29207_CR14","doi-asserted-by":"publisher","first-page":"966089","DOI":"10.3389\/fonc.2022.966089","volume":"12","author":"J Ngamkham","year":"2022","unstructured":"Ngamkham, J., Siritutsoontorn, S., Saisomboon, S., Vaeteewoottacharn, K. & Jitrapakdee, S. Cas9-mediated ablation of pyruvate carboxylase gene in colon cancer cell line HT-29 inhibits growth and migration, induces apoptosis and increases sensitivity to 5-fluorouracil and glutaminase inhibitor. Front. Oncol. 12, 966089. https:\/\/doi.org\/10.3389\/fonc.2022.966089 (2022).","journal-title":"Front. Oncol."},{"key":"29207_CR15","doi-asserted-by":"publisher","first-page":"487","DOI":"10.4149\/BLL_2022_077","volume":"123","author":"E Gondas","year":"2022","unstructured":"Gondas, E. et al. The ubiquitous expression of pyruvate carboxylase among human prostate tumors. Bratisl Lek Listy. 123, 487\u2013490. https:\/\/doi.org\/10.4149\/BLL_2022_077 (2022).","journal-title":"Bratisl Lek Listy"},{"key":"29207_CR16","doi-asserted-by":"publisher","first-page":"3848","DOI":"10.1016\/j.molcel.2021.08.028","volume":"81","author":"S Igelmann","year":"2021","unstructured":"Igelmann, S. et al. A hydride transfer complex reprograms NAD metabolism and bypasses senescence. Mol. Cell. 81, 3848\u20133865. https:\/\/doi.org\/10.1016\/j.molcel.2021.08.028 (2021). e3819.","journal-title":"Mol. Cell."},{"key":"29207_CR17","first-page":"1533","volume":"13","author":"MT Han","year":"2023","unstructured":"Han, M. T. et al. ZY-444 inhibits the growth and metastasis of prostate cancer by targeting TNFAIP3 through TNF signaling pathway. Am. J. Cancer Res. 13, 1533\u20131546 (2023).","journal-title":"Am. J. Cancer Res."},{"key":"29207_CR18","doi-asserted-by":"publisher","first-page":"845","DOI":"10.1089\/thy.2018.0435","volume":"29","author":"A Strickaert","year":"2019","unstructured":"Strickaert, A. et al. Reprogramming of energy metabolism: increased expression and roles of pyruvate carboxylase in papillary thyroid cancer. Thyroid 29, 845\u2013857. https:\/\/doi.org\/10.1089\/thy.2018.0435 (2019).","journal-title":"Thyroid"},{"key":"29207_CR19","doi-asserted-by":"publisher","first-page":"423","DOI":"10.1038\/s41420-022-01214-y","volume":"8","author":"Y Liu","year":"2022","unstructured":"Liu, Y. et al. Pyruvate carboxylase promotes malignant transformation of papillary thyroid carcinoma and reduces iodine uptake. Cell. Death Discov. 8, 423. https:\/\/doi.org\/10.1038\/s41420-022-01214-y (2022).","journal-title":"Cell. Death Discov"},{"key":"29207_CR20","doi-asserted-by":"publisher","first-page":"643416","DOI":"10.3389\/fonc.2021.643416","volume":"11","author":"C Liu","year":"2021","unstructured":"Liu, C. et al. Value of pyruvate carboxylase in thyroid Fine-Needle aspiration Wash-Out fluid for predicting papillary thyroid cancer lymph node metastasis. Front. Oncol. 11, 643416. https:\/\/doi.org\/10.3389\/fonc.2021.643416 (2021).","journal-title":"Front. Oncol."},{"key":"29207_CR21","doi-asserted-by":"publisher","first-page":"8674","DOI":"10.1073\/pnas.1016627108","volume":"108","author":"T Cheng","year":"2011","unstructured":"Cheng, T. et al. Pyruvate carboxylase is required for glutamine-independent growth of tumor cells. Proc. Natl. Acad. Sci. U S A. 108, 8674\u20138679. https:\/\/doi.org\/10.1073\/pnas.1016627108 (2011).","journal-title":"Proc. Natl. Acad. Sci. U S A"},{"key":"29207_CR22","doi-asserted-by":"publisher","first-page":"127","DOI":"10.4149\/gpb_2021003","volume":"40","author":"E Gondas","year":"2021","unstructured":"Gondas, E. et al. Expression of pyruvate carboxylase in cultured human astrocytoma, glioblastoma and neuroblastoma cells. Gen. Physiol. Biophys. 40, 127\u2013135. https:\/\/doi.org\/10.4149\/gpb_2021003 (2021).","journal-title":"Gen. Physiol. Biophys."},{"key":"29207_CR23","doi-asserted-by":"publisher","first-page":"e108289","DOI":"10.1371\/journal.pone.0108289","volume":"9","author":"JL Izquierdo-Garcia","year":"2014","unstructured":"Izquierdo-Garcia, J. L. et al. Glioma cells with the IDH1 mutation modulate metabolic fractional flux through pyruvate carboxylase. PLoS One. 9, e108289. https:\/\/doi.org\/10.1371\/journal.pone.0108289 (2014).","journal-title":"PLoS One"},{"key":"29207_CR24","doi-asserted-by":"publisher","first-page":"e2202104","DOI":"10.1002\/advs.202202104","volume":"9","author":"C Yang","year":"2022","unstructured":"Yang, C. et al. Genome-Wide CRISPR\/Cas9 library screening revealed dietary restriction of glutamine in combination with Inhibition of pyruvate metabolism as effective liver cancer treatment. Adv. Sci. (Weinh). 9, e2202104. https:\/\/doi.org\/10.1002\/advs.202202104 (2022).","journal-title":"Adv. Sci. (Weinh)"},{"key":"29207_CR25","doi-asserted-by":"publisher","first-page":"1903483","DOI":"10.1002\/advs.201903483","volume":"7","author":"Q Lin","year":"2020","unstructured":"Lin, Q. et al. Targeting pyruvate carboxylase by a small molecule suppresses breast cancer progression. Adv. Sci. (Weinh). 7, 1903483. https:\/\/doi.org\/10.1002\/advs.201903483 (2020).","journal-title":"Adv. Sci. (Weinh)"},{"key":"29207_CR26","doi-asserted-by":"publisher","first-page":"101","DOI":"10.1038\/s41573-020-0090-8","volume":"20","author":"MJ Mitchell","year":"2021","unstructured":"Mitchell, M. J. et al. Engineering precision nanoparticles for drug delivery. Nat. Rev. Drug Discov. 20, 101\u2013124. https:\/\/doi.org\/10.1038\/s41573-020-0090-8 (2021).","journal-title":"Nat. Rev. Drug Discov"},{"key":"29207_CR27","doi-asserted-by":"publisher","first-page":"18","DOI":"10.1016\/j.jconrel.2007.03.012","volume":"120","author":"P Kocbek","year":"2007","unstructured":"Kocbek, P., Obermajer, N., Cegnar, M., Kos, J. & Kristl, J. Targeting cancer cells using PLGA nanoparticles surface modified with monoclonal antibody. J. Control Release. 120, 18\u201326. https:\/\/doi.org\/10.1016\/j.jconrel.2007.03.012 (2007).","journal-title":"J. Control Release"},{"key":"29207_CR28","doi-asserted-by":"publisher","first-page":"36092","DOI":"10.1021\/acsomega.2c03974","volume":"7","author":"R Sharma","year":"2022","unstructured":"Sharma, R. et al. Functionalized Peptide-Based nanoparticles for targeted cancer nanotherapeutics: A State-of-the-Art review. ACS Omega. 7, 36092\u201336107. https:\/\/doi.org\/10.1021\/acsomega.2c03974 (2022).","journal-title":"ACS Omega"},{"key":"29207_CR29","doi-asserted-by":"publisher","first-page":"1625","DOI":"10.1016\/j.jinorgbio.2004.07.009","volume":"98","author":"G Russell-Jones","year":"2004","unstructured":"Russell-Jones, G., McTavish, K., McEwan, J., Rice, J. & Nowotnik, D. Vitamin-mediated targeting as a potential mechanism to increase drug uptake by tumours. J. Inorg. Biochem. 98, 1625\u20131633. https:\/\/doi.org\/10.1016\/j.jinorgbio.2004.07.009 (2004).","journal-title":"J. Inorg. Biochem."},{"key":"29207_CR30","doi-asserted-by":"publisher","first-page":"2048","DOI":"10.1021\/acs.bioconjchem.6b00317","volume":"27","author":"N Beztsinna","year":"2016","unstructured":"Beztsinna, N. et al. Amphiphilic Phospholipid-Based riboflavin derivatives for tumor targeting nanomedicines. Bioconjug. Chem. 27, 2048\u20132061. https:\/\/doi.org\/10.1021\/acs.bioconjchem.6b00317 (2016).","journal-title":"Bioconjug. Chem."},{"key":"29207_CR31","doi-asserted-by":"publisher","first-page":"1548","DOI":"10.1111\/php.13464","volume":"97","author":"W Mekseriwattana","year":"2021","unstructured":"Mekseriwattana, W. et al. Dual functions of Riboflavin-functionalized Poly(lactic-co-glycolic acid) nanoparticles for enhanced drug delivery efficiency and photodynamic therapy in Triple-negative breast cancer cells. Photochem. Photobiol. 97, 1548\u20131557. https:\/\/doi.org\/10.1111\/php.13464 (2021).","journal-title":"Photochem. Photobiol"},{"key":"29207_CR32","doi-asserted-by":"publisher","first-page":"210","DOI":"10.1080\/21691401.2018.1548473","volume":"47","author":"CY Wu","year":"2019","unstructured":"Wu, C. Y. & Chen, Y. C. Riboflavin immobilized Fe(3)O(4) magnetic nanoparticles carried with n-butylidenephthalide as targeting-based anticancer agents. Artif. Cells Nanomed. Biotechnol. 47, 210\u2013220. https:\/\/doi.org\/10.1080\/21691401.2018.1548473 (2019).","journal-title":"Artif. Cells Nanomed. Biotechnol."},{"key":"29207_CR33","doi-asserted-by":"publisher","first-page":"1988","DOI":"10.1039\/d2na00015f","volume":"4","author":"W Mekseriwattana","year":"2022","unstructured":"Mekseriwattana, W. et al. Riboflavin-citrate conjugate multicore spions with enhanced magnetic responses and cellular uptake in breast cancer cells. Nanoscale Adv. 4, 1988\u20131998. https:\/\/doi.org\/10.1039\/d2na00015f (2022).","journal-title":"Nanoscale Adv."},{"key":"29207_CR34","doi-asserted-by":"publisher","first-page":"507","DOI":"10.1021\/bm201566g","volume":"13","author":"AB Witte","year":"2012","unstructured":"Witte, A. B. et al. Biophysical characterization of a riboflavin-conjugated dendrimer platform for targeted drug delivery. Biomacromolecules 13, 507\u2013516. https:\/\/doi.org\/10.1021\/bm201566g (2012).","journal-title":"Biomacromolecules"},{"key":"29207_CR35","doi-asserted-by":"publisher","first-page":"112141","DOI":"10.1016\/j.ejmech.2020.112141","volume":"190","author":"T Fan","year":"2020","unstructured":"Fan, T. et al. Design, synthesis and evaluation of Phenylthiazole and phenylthiophene pyrimidindiamine derivatives targeting the bacterial membrane. Eur. J. Med. Chem. 190, 112141. https:\/\/doi.org\/10.1016\/j.ejmech.2020.112141 (2020).","journal-title":"Eur. J. Med. Chem."},{"key":"29207_CR36","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1021\/jm801525s","volume":"52","author":"PS Portoghese","year":"2009","unstructured":"Portoghese, P. S. Revision of purity criteria for tested compounds. J. Med. Chem. 52, 1. https:\/\/doi.org\/10.1021\/jm801525s (2009).","journal-title":"J. Med. Chem."},{"key":"29207_CR37","doi-asserted-by":"publisher","first-page":"3131","DOI":"10.7150\/jca.18457","volume":"8","author":"X Dai","year":"2017","unstructured":"Dai, X., Cheng, H., Bai, Z. & Li, J. Breast cancer cell line classification and its relevance with breast tumor subtyping. J. Cancer. 8, 3131\u20133141. https:\/\/doi.org\/10.7150\/jca.18457 (2017).","journal-title":"J. Cancer"},{"key":"29207_CR38","doi-asserted-by":"publisher","first-page":"599","DOI":"10.3892\/ijo.2017.4206","volume":"52","author":"A Olejniczak","year":"2018","unstructured":"Olejniczak, A., Szarynska, M. & Kmiec, Z. In vitro characterization of spheres derived from colorectal cancer cell lines. Int. J. Oncol. 52, 599\u2013612. https:\/\/doi.org\/10.3892\/ijo.2017.4206 (2018).","journal-title":"Int. J. Oncol."},{"key":"29207_CR39","doi-asserted-by":"publisher","first-page":"162","DOI":"10.1111\/j.1365-2125.2010.03892.x","volume":"72","author":"B Pardini","year":"2011","unstructured":"Pardini, B. et al. Lorenzo Bermejo, 5-Fluorouracil-based chemotherapy for colorectal cancer and MTHFR\/MTRR genotypes. Br. J. Clin. Pharmacol. 72, 162\u2013163. https:\/\/doi.org\/10.1111\/j.1365-2125.2010.03892.x (2011).","journal-title":"Br. J. Clin. Pharmacol."},{"key":"29207_CR40","first-page":"19","volume":"11","author":"JF Bishop","year":"1997","unstructured":"Bishop, J. F. et al. Paclitaxel as first-line treatment for metastatic breast cancer. Oncol. (Williston Park). 11, 19\u201323 (1997).","journal-title":"Oncol. (Williston Park)"},{"key":"29207_CR41","doi-asserted-by":"publisher","first-page":"79","DOI":"10.3389\/fphar.2019.00079","volume":"10","author":"L Bartmann","year":"2019","unstructured":"Bartmann, L. et al. Evaluation of riboflavin transporters as targets for drug delivery and theranostics. Front. Pharmacol. 10, 79. https:\/\/doi.org\/10.3389\/fphar.2019.00079 (2019).","journal-title":"Front. Pharmacol."},{"key":"29207_CR42","doi-asserted-by":"publisher","first-page":"8","DOI":"10.1186\/s12935-021-02432-7","volume":"22","author":"L Zhang","year":"2022","unstructured":"Zhang, L. et al. Systematic analysis of the role of SLC52A2 in multiple human cancers. Cancer Cell. Int. 22, 8. https:\/\/doi.org\/10.1186\/s12935-021-02432-7 (2022).","journal-title":"Cancer Cell. Int."},{"key":"29207_CR43","doi-asserted-by":"publisher","first-page":"102342","DOI":"10.1016\/j.canep.2023.102342","volume":"83","author":"Y Ma","year":"2023","unstructured":"Ma, Y. et al. High serum riboflavin is associated with the risk of sporadic colorectal cancer. Cancer Epidemiol. 83, 102342. https:\/\/doi.org\/10.1016\/j.canep.2023.102342 (2023).","journal-title":"Cancer Epidemiol."},{"key":"29207_CR44","doi-asserted-by":"publisher","first-page":"83","DOI":"10.1016\/j.hepres.2006.03.009","volume":"35","author":"PN Rao","year":"2006","unstructured":"Rao, P. N. et al. Elevation of serum riboflavin carrier protein in hepatocellular carcinoma. Hepatol. Res. 35, 83\u201387. https:\/\/doi.org\/10.1016\/j.hepres.2006.03.009 (2006).","journal-title":"Hepatol. Res."},{"key":"29207_CR45","first-page":"985","volume":"8","author":"PN Rao","year":"1999","unstructured":"Rao, P. N. et al. Elevation of serum riboflavin carrier protein in breast cancer. Cancer Epidemiol. Biomarkers Prev. 8, 985\u2013990 (1999).","journal-title":"Cancer Epidemiol. Biomarkers Prev."},{"key":"29207_CR46","doi-asserted-by":"publisher","first-page":"3634","DOI":"10.2741\/3477","volume":"14","author":"T Johnson","year":"2009","unstructured":"Johnson, T. et al. Biochemical characterization of riboflavin carrier protein (RCP) in prostate cancer. Front. Biosci. (Landmark Ed). 14, 3634\u20133640. https:\/\/doi.org\/10.2741\/3477 (2009).","journal-title":"Front. Biosci. (Landmark Ed)"},{"key":"29207_CR47","doi-asserted-by":"publisher","first-page":"2659","DOI":"10.21873\/anticanres.12508","volume":"38","author":"V Tutino","year":"2018","unstructured":"Tutino, V. et al. The expression of riboflavin transporters in human colorectal cancer. Anticancer Res. 38, 2659\u20132667. https:\/\/doi.org\/10.21873\/anticanres.12508 (2018).","journal-title":"Anticancer Res."},{"key":"29207_CR48","doi-asserted-by":"publisher","first-page":"7530","DOI":"10.1038\/ncomms8530","volume":"6","author":"L Phan","year":"2015","unstructured":"Phan, L. et al. The cell cycle regulator 14-3-3sigma opposes and reverses cancer metabolic reprogramming. Nat. Commun. 6, 7530. https:\/\/doi.org\/10.1038\/ncomms8530 (2015).","journal-title":"Nat. Commun."},{"key":"29207_CR49","doi-asserted-by":"publisher","unstructured":"Boztepe, T. et al. Design, Synthesis, Characterization, and evaluation of the Anti-HT-29 colorectal cell line activity of novel 8-Oxyquinolinate-Platinum(II)-Loaded nanostructured lipid carriers targeted with riboflavin. Pharmaceutics 15 https:\/\/doi.org\/10.3390\/pharmaceutics15031021 (2023).","DOI":"10.3390\/pharmaceutics15031021"},{"key":"29207_CR50","doi-asserted-by":"publisher","first-page":"20180153","DOI":"10.1098\/rstb.2018.0153","volume":"374","author":"C Commisso","year":"2019","unstructured":"Commisso, C. The pervasiveness of macropinocytosis in oncological malignancies. Philos. Trans. R Soc. Lond. B Biol. Sci. 374, 20180153. https:\/\/doi.org\/10.1098\/rstb.2018.0153 (2019).","journal-title":"Philos. Trans. R Soc. Lond. B Biol. Sci."},{"key":"29207_CR51","doi-asserted-by":"publisher","first-page":"7868","DOI":"10.1021\/acsomega.7b01315","volume":"2","author":"S Palvai","year":"2017","unstructured":"Palvai, S., Kuman, M. M., Sengupta, P. & Basu, S. Hyaluronic acid layered chimeric nanoparticles: targeting MAPK-PI3K signaling hub in colon cancer cells. ACS Omega. 2, 7868\u20137880. https:\/\/doi.org\/10.1021\/acsomega.7b01315 (2017).","journal-title":"ACS Omega"},{"key":"29207_CR52","doi-asserted-by":"publisher","first-page":"2607","DOI":"10.1038\/s41467-018-05061-3","volume":"9","author":"M Yu","year":"2018","unstructured":"Yu, M. et al. Rapid transport of deformation-tuned nanoparticles across biological hydrogels and cellular barriers. Nat. Commun. 9, 2607. https:\/\/doi.org\/10.1038\/s41467-018-05061-3 (2018).","journal-title":"Nat. Commun."},{"key":"29207_CR53","doi-asserted-by":"publisher","unstructured":"Franchi, M., Piperigkou, Z., Karamanos, K. A., Franchi, L. & Masola, V. Extracellular matrix-mediated breast cancer cells morphological alterations, invasiveness, and microvesicles\/exosomes release. Cells. https:\/\/doi.org\/10.3390\/cells9092031 (2020).","DOI":"10.3390\/cells9092031"},{"key":"29207_CR54","doi-asserted-by":"publisher","first-page":"61","DOI":"10.1016\/j.ijpharm.2016.11.012","volume":"516","author":"MM El-Hammadi","year":"2017","unstructured":"El-Hammadi, M. M., Delgado, A. V., Melguizo, C., Prados, J. C. & Arias, J. L. Folic acid-decorated and pegylated PLGA nanoparticles for improving the antitumour activity of 5-fluorouracil. Int. J. Pharm. 516, 61\u201370. https:\/\/doi.org\/10.1016\/j.ijpharm.2016.11.012 (2017).","journal-title":"Int. J. Pharm."},{"key":"29207_CR55","doi-asserted-by":"publisher","first-page":"1271","DOI":"10.2147\/DDDT.S195493","volume":"13","author":"S Li","year":"2019","unstructured":"Li, S., Li, X., Ding, J., Han, L. & Guo, X. Anti-tumor efficacy of folate modified PLGA-based nanoparticles for the co-delivery of drugs in ovarian cancer. Drug Des. Devel Ther. 13, 1271\u20131280. https:\/\/doi.org\/10.2147\/DDDT.S195493 (2019).","journal-title":"Drug Des. Devel Ther."},{"key":"29207_CR56","doi-asserted-by":"publisher","first-page":"2749","DOI":"10.1021\/acs.molpharmaceut.7b00310","volume":"14","author":"Y Singh","year":"2017","unstructured":"Singh, Y. et al. Click biotinylation of PLGA template for biotin receptor oriented delivery of doxorubicin hydrochloride in 4T1 Cell-Induced breast cancer. Mol. Pharm. 14, 2749\u20132765. https:\/\/doi.org\/10.1021\/acs.molpharmaceut.7b00310 (2017).","journal-title":"Mol. Pharm."},{"key":"29207_CR57","doi-asserted-by":"publisher","unstructured":"Cheng, H. et al. Lipid metabolism and cancer. Life (Basel). https:\/\/doi.org\/10.3390\/life12060784. (2022).","DOI":"10.3390\/life12060784"},{"key":"29207_CR58","doi-asserted-by":"publisher","first-page":"7","DOI":"10.1038\/s44324-024-00047-w","volume":"3","author":"M Hornisch","year":"2025","unstructured":"Hornisch, M. & Piazza, I. Regulation of gene expression through protein-metabolite interactions. NPJ Metab. Health Dis. 3, 7. https:\/\/doi.org\/10.1038\/s44324-024-00047-w (2025).","journal-title":"NPJ Metab. Health Dis."},{"key":"29207_CR59","doi-asserted-by":"publisher","first-page":"102","DOI":"10.1038\/s41467-019-13668-3","volume":"11","author":"I Martinez-Reyes","year":"2020","unstructured":"Martinez-Reyes, I. & Chandel, N. S. Mitochondrial TCA cycle metabolites control physiology and disease. Nat. Commun. 11, 102. https:\/\/doi.org\/10.1038\/s41467-019-13668-3 (2020).","journal-title":"Nat. Commun."},{"key":"29207_CR60","doi-asserted-by":"publisher","first-page":"107","DOI":"10.1016\/j.cmet.2006.06.008","volume":"4","author":"K Uyeda","year":"2006","unstructured":"Uyeda, K. & Repa, J. J. Carbohydrate response element binding protein, ChREBP, a transcription factor coupling hepatic glucose utilization and lipid synthesis. Cell. Metab. 4, 107\u2013110. https:\/\/doi.org\/10.1016\/j.cmet.2006.06.008 (2006).","journal-title":"Cell. Metab."},{"key":"29207_CR61","doi-asserted-by":"publisher","first-page":"26122","DOI":"10.1074\/jbc.M702854200","volume":"282","author":"S Yoon","year":"2007","unstructured":"Yoon, S. et al. Up-regulation of acetyl-CoA carboxylase alpha and fatty acid synthase by human epidermal growth factor receptor 2 at the translational level in breast cancer cells. J. Biol. Chem. 282, 26122\u201326131. https:\/\/doi.org\/10.1074\/jbc.M702854200 (2007).","journal-title":"J. Biol. Chem."},{"key":"29207_CR62","doi-asserted-by":"publisher","first-page":"611","DOI":"10.1194\/jlr.TR119000439","volume":"61","author":"F Mollinedo","year":"2020","unstructured":"Mollinedo, F. & Gajate, C. Lipid rafts as signaling hubs in cancer cell survival\/death and invasion: implications in tumor progression and therapy: thematic review series: biology of lipid rafts. J. Lipid Res. 61, 611\u2013635. https:\/\/doi.org\/10.1194\/jlr.TR119000439 (2020).","journal-title":"J. Lipid Res."},{"key":"29207_CR63","doi-asserted-by":"publisher","first-page":"423","DOI":"10.1049\/iet-nbt.2017.0189","volume":"12","author":"K Sawasdee","year":"2018","unstructured":"Sawasdee, K. et al. Encapsulation of the reductase component of p-hydroxyphenylacetate hydroxylase in poly(lactide-co-glycolide) nanoparticles by three different emulsification techniques. IET Nanobiotechnol. 12, 423\u2013428. https:\/\/doi.org\/10.1049\/iet-nbt.2017.0189 (2018).","journal-title":"IET Nanobiotechnol."},{"key":"29207_CR64","doi-asserted-by":"publisher","first-page":"197","DOI":"10.1016\/0003-9861(91)90608-l","volume":"290","author":"M Rohde","year":"1991","unstructured":"Rohde, M., Lim, F. & Wallace, J. C. Electron microscopic localization of pyruvate carboxylase in rat liver and Saccharomyces cerevisiae by Immunogold procedures. Arch. Biochem. Biophys. 290, 197\u2013201. https:\/\/doi.org\/10.1016\/0003-9861(91)90608-l (1991).","journal-title":"Arch. Biochem. Biophys."}],"container-title":["Scientific Reports"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.nature.com\/articles\/s41598-025-29207-8","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41598-025-29207-8.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41598-025-29207-8.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,1,3]],"date-time":"2026-01-03T22:04:11Z","timestamp":1767477851000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.nature.com\/articles\/s41598-025-29207-8"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,12,30]]},"references-count":64,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2026,12]]}},"alternative-id":["29207"],"URL":"https:\/\/doi.org\/10.1038\/s41598-025-29207-8","relation":{},"ISSN":["2045-2322"],"issn-type":[{"value":"2045-2322","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,12,30]]},"assertion":[{"value":"8 April 2025","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"14 November 2025","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"30 December 2025","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Declarations"}},{"value":"The authors declare no competing interests.","order":2,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing interests"}}],"article-number":"125"}}