{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,23]],"date-time":"2026-08-23T18:31:48Z","timestamp":1787509908520,"version":"build-2736575974"},"reference-count":62,"publisher":"MDPI AG","issue":"5","license":[{"start":{"date-parts":[[2025,2,28]],"date-time":"2025-02-28T00:00:00Z","timestamp":1740700800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"the European Union\u2014NextGenerationEU","award":["CN_00000033"],"award-info":[{"award-number":["CN_00000033"]}]},{"name":"the European Union\u2014NextGenerationEU","award":["CUP, H43C22000530001"],"award-info":[{"award-number":["CUP, H43C22000530001"]}]},{"name":"the Italian Ministry of University and Research","award":["CN_00000033"],"award-info":[{"award-number":["CN_00000033"]}]},{"name":"the Italian Ministry of University and Research","award":["CUP, H43C22000530001"],"award-info":[{"award-number":["CUP, H43C22000530001"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["IJMS"],"abstract":"<jats:p>Colorectal cancer (CRC) is the second deadliest cancer in the Western world. Increased body weight, a diet rich in red meat and alcohol, as well as a sedentary lifestyle, are all involved in sporadic CRC pathogenesis. Since current CRC therapies show several side effects, there is a need to find new and more effective therapeutic approaches, allowing conventional drug dosages and toxicity to be reduced. Gratiola officinalis alcoholic extract was characterized by LC-MS and its effect investigated on a healthy colon mucosa cell line and on different colorectal cancer cell lines. Cell viability, apoptosis and cell cycle progression were evaluated through flow cytometry; energy production and glycolysis were investigated using Seahorse technology, while cancer markers were analyzed through Western blotting. The untargeted metabolomics analysis of G. officinalis alcoholic extract revealed glycosides of different polyphenols and glycosides of cucurbitane-type triterpenes. This extract showed a stronger impact on CRC cell line viability compared to healthy colon cells. In the E705 CRC cell line, it induced cell apoptosis and caused the downregulation of glycolysis, inhibiting cell proliferation. On the other hand, SW480 CRC cells treated with G. officinalis extract showed G2\/M cell cycle arrest. This work shows that G. officinalis extract can reduce glycolysis and promote cell cycle arrest in CRC cells, suggesting that G. officinalis could represent a novel player in the prevention and treatment of CRC.<\/jats:p>","DOI":"10.3390\/ijms26052220","type":"journal-article","created":{"date-parts":[[2025,3,3]],"date-time":"2025-03-03T04:10:42Z","timestamp":1740975042000},"page":"2220","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":6,"title":["Gratiola officinalis Alcoholic Extract Targets Warburg Effect, Apoptosis and Cell Cycle Progression in Colorectal Cancer Cell Lines"],"prefix":"10.3390","volume":"26","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-2953-1852","authenticated-orcid":false,"given":"Stefano","family":"Bianchini","sequence":"first","affiliation":[{"name":"Department of Biotechnology and Biosciences, University of Milano-Bicocca, Piazza della Scienza 2, 20126 Milano, Italy"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-9603-7294","authenticated-orcid":false,"given":"Federica","family":"Bovio","sequence":"additional","affiliation":[{"name":"Department of Biotechnology and Biosciences, University of Milano-Bicocca, Piazza della Scienza 2, 20126 Milano, Italy"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4842-6831","authenticated-orcid":false,"given":"Stefano","family":"Negri","sequence":"additional","affiliation":[{"name":"Department of Biotechnology, University of Verona, Strada Le Grazie 15, 37134 Verona, Italy"},{"name":"National Biodiversity Future Center (NBFC), 90133 Palermo, Italy"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Flavia","family":"Guzzo","sequence":"additional","affiliation":[{"name":"Department of Biotechnology, University of Verona, Strada Le Grazie 15, 37134 Verona, Italy"},{"name":"National Biodiversity Future Center (NBFC), 90133 Palermo, Italy"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9706-4654","authenticated-orcid":false,"given":"Matilde","family":"Forcella","sequence":"additional","affiliation":[{"name":"Department of Biotechnology and Biosciences, University of Milano-Bicocca, Piazza della Scienza 2, 20126 Milano, Italy"},{"name":"National Biodiversity Future Center (NBFC), 90133 Palermo, Italy"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2746-1742","authenticated-orcid":false,"given":"Paola","family":"Fusi","sequence":"additional","affiliation":[{"name":"Department of Biotechnology and Biosciences, University of Milano-Bicocca, Piazza della Scienza 2, 20126 Milano, Italy"},{"name":"National Biodiversity Future Center (NBFC), 90133 Palermo, Italy"},{"name":"Integrated Models for Prevention and Protection in Environmental and Occupational Health (MISTRAL), Interuniversity Research Center, 25121 Brescia, Italy"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2025,2,28]]},"reference":[{"key":"ref_1","first-page":"34","article-title":"AntI-Inflammatory, Antipyretic and Antimicrobial Activity of Flavonoid-Containing Extract of Gratiola officinalis L","volume":"78","author":"Polukonova","year":"2015","journal-title":"Eksp. Klin. Farmakol."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"15","DOI":"10.1186\/s13002-020-00364-6","article-title":"Historical Ethnobotanical Review of Medicinal Plants Used to Treat Children Diseases in Romania (1860s\u20131970s)","volume":"16","author":"Petran","year":"2020","journal-title":"J. Ethnobiol. Ethnomed."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"110815","DOI":"10.1016\/j.fct.2019.110815","article-title":"Genotoxic Properties of Betonica Officinalis, Gratiola officinalis, Vincetoxicum Luteum and Vincetoxicum Hirundinaria Extracts","volume":"134","author":"Lazutka","year":"2019","journal-title":"Food Chem. Toxicol."},{"key":"ref_4","doi-asserted-by":"crossref","unstructured":"Navolokin, N., Lomova, M., Bucharskaya, A., Godage, O., Polukonova, N., Shirokov, A., Grinev, V., and Maslyakova, G. (2023). Antitumor Effects of Microencapsulated Gratiola officinalis Extract on Breast Carcinoma and Human Cervical Cancer Cells In Vitro. Materials, 16.","DOI":"10.3390\/ma16041470"},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"415","DOI":"10.15275\/rusomj.2022.0415","article-title":"Examining Apoptotic Activity of Gratiola officinalis L. (Scrophulariaceae) Extract On Cultured Human Tumor Cell Lines","volume":"11","author":"Polukonova","year":"2022","journal-title":"Russ. Open Med. J."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"e14264","DOI":"10.1111\/jfbc.14264","article-title":"Health Benefits of Polyphenols: A Concise Review","volume":"46","author":"Rana","year":"2022","journal-title":"J. Food Biochem."},{"key":"ref_7","doi-asserted-by":"crossref","unstructured":"Sammarco, G., Gallo, G., Vescio, G., Picciariello, A., De Paola, G., Trompetto, M., Curr\u00f2, G., and Ammendola, M. (2020). Mast Cells, MicroRNAs and Others: The Role of Translational Research on Colorectal Cancer in the Forthcoming Era of Precision Medicine. J. Clin. Med., 9.","DOI":"10.3390\/jcm9092852"},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"188699","DOI":"10.1016\/j.bbcan.2022.188699","article-title":"A Synopsis of Modern\u2014Day Colorectal Cancer: Where We Stand","volume":"1877","author":"Ranasinghe","year":"2022","journal-title":"Biochim. Biophys. Acta (BBA) Rev. Cancer"},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"1467","DOI":"10.1016\/S0140-6736(19)32319-0","article-title":"Colorectal Cancer","volume":"394","author":"Dekker","year":"2019","journal-title":"Lancet"},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"464","DOI":"10.1001\/jamaoncol.2016.5194","article-title":"Prevalence and Spectrum of Germline Cancer Susceptibility Gene Mutations Among Patients With Early-Onset Colorectal Cancer","volume":"3","author":"Pearlman","year":"2017","journal-title":"JAMA Oncol."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"897","DOI":"10.1053\/j.gastro.2017.11.004","article-title":"Germline Genetic Features of Young Individuals With Colorectal Cancer","volume":"154","author":"Stoffel","year":"2018","journal-title":"Gastroenterology"},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"1247","DOI":"10.1002\/jso.27320","article-title":"Colorectal Cancer in Young Adults","volume":"127","author":"Kim","year":"2023","journal-title":"J. Surg. Oncol."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"570","DOI":"10.1126\/science.aam9949","article-title":"Microbiota-Activated PPAR-\u03b3 Signaling Inhibits Dysbiotic Enterobacteriaceae Expansion","volume":"357","author":"Byndloss","year":"2017","journal-title":"Science"},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"273","DOI":"10.1016\/j.chom.2020.06.001","article-title":"High-Fat Diet and Antibiotics Cooperatively Impair Mitochondrial Bioenergetics to Trigger Dysbiosis That Exacerbates Pre-Inflammatory Bowel Disease","volume":"28","author":"Lee","year":"2020","journal-title":"Cell Host Microbe"},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"158","DOI":"10.1093\/jnci\/djy087","article-title":"Circulating Vitamin D and Colorectal Cancer Risk: An International Pooling Project of 17 Cohorts","volume":"111","author":"McCullough","year":"2019","journal-title":"J. Natl. Cancer Inst."},{"key":"ref_16","doi-asserted-by":"crossref","unstructured":"Thanikachalam, K., and Khan, G. (2019). Colorectal Cancer and Nutrition. Nutrients, 11.","DOI":"10.3390\/nu11010164"},{"key":"ref_17","doi-asserted-by":"crossref","unstructured":"Masci, D., Puxeddu, M., Silvestri, R., and La Regina, G. (2024). Metabolic Rewiring in Cancer: Small Molecule Inhibitors in Colorectal Cancer Therapy. Molecules, 29.","DOI":"10.3390\/molecules29092110"},{"key":"ref_18","doi-asserted-by":"crossref","unstructured":"R\u00edos-Hoyo, A., Monzon\u00eds, X., Vidal, J., Linares, J., and Montagut, C. (2024). Unveiling Acquired Resistance to Anti-EGFR Therapies in Colorectal Cancer: A Long and Winding Road. Front. Pharmacol., 15.","DOI":"10.3389\/fphar.2024.1398419"},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"113384","DOI":"10.1016\/j.biopha.2022.113384","article-title":"Plant-Derived Bioactive Compounds in Colon Cancer Treatment: An Updated Review","volume":"153","author":"Esmeeta","year":"2022","journal-title":"Biomed. Pharmacother."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"e4033","DOI":"10.1002\/cbf.4033","article-title":"Berberine: Potential Preventive and Therapeutic Strategies for Human Colorectal Cancer","volume":"42","author":"Feng","year":"2024","journal-title":"Cell Biochem. Funct."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"1927","DOI":"10.1016\/j.jff.2013.09.014","article-title":"Antioxidant Properties and Polyphenolics Composition of Common Hedge Hyssop (Gratiola officinalis L.)","volume":"5","author":"Venskutonis","year":"2013","journal-title":"J. Funct. Foods"},{"key":"ref_22","doi-asserted-by":"crossref","unstructured":"Plamada, D., and Vodnar, D.C. (2021). Polyphenols\u2014Gut Microbiota Interrelationship: A Transition to a New Generation of Prebiotics. Nutrients, 14.","DOI":"10.3390\/nu14010137"},{"key":"ref_23","doi-asserted-by":"crossref","unstructured":"Fabbrini, M., D\u2019Amico, F., Barone, M., Conti, G., Mengoli, M., Brigidi, P., and Turroni, S. (2022). Polyphenol and Tannin Nutraceuticals and Their Metabolites: How the Human Gut Microbiota Influences Their Properties. Biomolecules, 12.","DOI":"10.3390\/biom12070875"},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"1331","DOI":"10.1007\/BF00807062","article-title":"New Cucurbitacine Glycosides from Gratiola officinalis L.","volume":"126","author":"Rothenburger","year":"1995","journal-title":"Monatshefte Chem. Chem. Mon."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"121","DOI":"10.1002\/(SICI)1099-1565(200003\/04)11:2<121::AID-PCA493>3.0.CO;2-9","article-title":"Analysis of Cucurbitacins in Medicinal Plants by High-Pressure Liquid Chromatography\u2013Mass Spectrometry","volume":"11","author":"Sturm","year":"2000","journal-title":"Phytochem. Anal."},{"key":"ref_26","doi-asserted-by":"crossref","unstructured":"Frezza, C., Venditti, A., Bianco, A., Serafini, M., Pitorri, M., Sciubba, F., Di Cocco, M.E., Spinozzi, E., Cappellacci, L., and Hofer, A. (2020). Phytochemical Analysis and Trypanocidal Activity of Marrubium Incanum Desr. Molecules, 25.","DOI":"10.3390\/molecules25143140"},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"1344","DOI":"10.1021\/np900086y","article-title":"Phenylethanoid and Lignan Glycosides from Polar Extracts of Lantana, a Genus of Verbenaceous Plants Widely Used in Traditional Herbal Therapies","volume":"72","author":"Nimmo","year":"2009","journal-title":"J. Nat. Prod."},{"key":"ref_28","doi-asserted-by":"crossref","unstructured":"Daneshforouz, A., Nazemi, S., Gholami, O., Kafami, M., and Amin, B. (2021). The Cytotoxicity and Apoptotic Effects of Verbascoside on Breast Cancer 4T1 Cell Line. BMC Pharmacol. Toxicol., 22.","DOI":"10.1186\/s40360-021-00540-8"},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"2149","DOI":"10.2174\/1871520618666180611112125","article-title":"Verbascoside Attenuates Rac-1 and HIF-1\u03b1 Signaling Cascade in Colorectal Cancer Cells","volume":"18","author":"Seyfi","year":"2019","journal-title":"Anticancer Agents Med. Chem."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"16939","DOI":"10.1038\/s41598-018-35083-2","article-title":"Verbascoside: Identification, Quantification, and Potential Sensitization of Colorectal Cancer Cells to 5-FU by Targeting PI3K\/AKT Pathway","volume":"8","author":"Attia","year":"2018","journal-title":"Sci. Rep."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"264","DOI":"10.3906\/biy-2003-18","article-title":"Caffeic Acid Phenethyl Ester Induces Apoptosis in Colorectal Cancer Cells via Inhibition of Survivin","volume":"44","author":"Sari","year":"2020","journal-title":"Turk. J. Biol."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"120500","DOI":"10.1016\/j.lfs.2022.120500","article-title":"Ferulic Acid Nanocapsules as a Promising Treatment Modality for Colorectal Cancer: Preparation and in Vitro\/in Vivo Appraisal","volume":"298","author":"Nasr","year":"2022","journal-title":"Life Sci."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"e70095","DOI":"10.1002\/jbt.70095","article-title":"Targeting the SIRT3\/MnSOD and JNK\/HMGB1\/Beclin 1 Axes: Role of Apigenin in Multifaceted Metabolic Intervention in Colorectal Cancer","volume":"39","author":"Abdelmaksoud","year":"2025","journal-title":"J. Biochem. Mol. Toxicol."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"e0402","DOI":"10.15275\/rusomj.2018.0402","article-title":"The Apoptotic Activity of Flavonoid-Containing Gratiola officinalis Extract in Cell Cultures of Human Kidney Cancer","volume":"7","author":"Polukonova","year":"2018","journal-title":"Russ. Open Med. J."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"108948","DOI":"10.1016\/j.carres.2023.108948","article-title":"Isolation and Structural Elucidation of Prebiotic Oligosaccharides from Ziziphi Spinosae Semen","volume":"534","author":"Song","year":"2023","journal-title":"Carbohydr. Res."},{"key":"ref_36","doi-asserted-by":"crossref","unstructured":"Peixoto, J.A.B., \u00c1lvarez-Rivera, G., Costa, A.S.G., Machado, S., Cifuentes, A., Ib\u00e1\u00f1ez, E., Oliveira, M.B.P.P., and Alves, R.C. (2023). Contribution of Phenolics and Free Amino Acids on the Antioxidant Profile of Commercial Lemon Verbena Infusions. Antioxidants, 12.","DOI":"10.3390\/antiox12020251"},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"755","DOI":"10.1016\/j.foodres.2017.06.025","article-title":"Profiling Hydroxycinnamic Acid Glycosides, Iridoid Glycosides, and Phenylethanoid Glycosides in Baobab Fruit Pulp (Adansonia digitata)","volume":"99","author":"Li","year":"2017","journal-title":"Food Res. Int."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"1010","DOI":"10.1002\/jms.3413","article-title":"Approach to the Study of Flavone Di-C-glycosides by High Performance Liquid Chromatography-tandem Ion Trap Mass Spectrometry and Its Application to Characterization of Flavonoid Composition in Viola Yedoensis","volume":"49","author":"Cao","year":"2014","journal-title":"J. Mass Spectrom."},{"key":"ref_39","doi-asserted-by":"crossref","unstructured":"Formato, M., Piccolella, S., Zidorn, C., and Pacifico, S. (2021). UHPLC-HRMS Analysis of Fagus Sylvatica (Fagaceae) Leaves: A Renewable Source of Antioxidant Polyphenols. Antioxidants, 10.","DOI":"10.3390\/antiox10071140"},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"7976","DOI":"10.1021\/jf401937b","article-title":"Phytochemical Profiles of Black, Red, Brown, and White Rice from the Camargue Region of France","volume":"61","author":"Cros","year":"2013","journal-title":"J. Agric. Food Chem."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"1211","DOI":"10.1002\/jms.3844","article-title":"LC-PDA-ESI-MSn Analysis of Phenolic and Iridoid Compounds from Globularia Spp.","volume":"51","author":"Bucar","year":"2016","journal-title":"J. Mass Spectrom."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1002\/pca.802","article-title":"Identification by HPLC-PAD-MS and Quantification by HPLC-PAD of Phenylethanoid Glycosides of Five Phlomis Species","volume":"16","author":"Montoro","year":"2005","journal-title":"Phytochem. Anal."},{"key":"ref_43","doi-asserted-by":"crossref","unstructured":"Gao, H., Liu, Z., Song, F., Xing, J., Zheng, Z., and Liu, S. (2022). A Strategy for Identification and Structural Characterization of Compounds from Plantago asiatica L. by Liquid Chromatography-Mass Spectrometry Combined with Ion Mobility Spectrometry. Molecules, 27.","DOI":"10.3390\/molecules27134302"},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"237","DOI":"10.1002\/pca.2809","article-title":"Chemical Fingerprint and Bioactivity Evaluation of Globularia orientalis L. and Globularia trichosantha Fisch. & C. A. Mey. Using Non-targeted HPLC-ESI-QTOF-MS Approach","volume":"30","author":"Zengin","year":"2019","journal-title":"Phytochem. Anal."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"1084","DOI":"10.1248\/cpb.48.1084","article-title":"Isolation of Lignan Glucosides and Neolignan Sulfate from the Leaves of Glochidion zeylanicum (Gaertn) A. Juss","volume":"48","author":"Otsuka","year":"2000","journal-title":"Chem. Pharm. Bull."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"15992","DOI":"10.1038\/s41598-019-52404-1","article-title":"Metabolite Profiling and Quantitation of Cucurbitacins in Cucurbitaceae Plants by Liquid Chromatography Coupled to Tandem Mass Spectrometry","volume":"9","author":"Haq","year":"2019","journal-title":"Sci. Rep."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"e2300019","DOI":"10.1002\/jssc.202300019","article-title":"Identification and Quantification of Cucurbitacin in Watermelon Frost Using Molecular Networking Integrated with Ultra-high-performance Liquid Chromatography-tandem Mass Spectrometry","volume":"46","author":"Hu","year":"2023","journal-title":"J. Sep. Sci."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"1191","DOI":"10.1080\/10286020.2012.734299","article-title":"New Iridoid Glycoside from Gratiola officinalis","volume":"14","author":"Ali","year":"2012","journal-title":"J. Asian Nat. Prod. Res."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"259","DOI":"10.1515\/bmc-2016-0015","article-title":"Bcl-2 Family Proteins: Master Regulators of Cell Survival","volume":"7","author":"Hatok","year":"2016","journal-title":"Biomol. Concepts"},{"key":"ref_50","doi-asserted-by":"crossref","unstructured":"Uribe, M.L., Marrocco, I., and Yarden, Y. (2021). EGFR in Cancer: Signaling Mechanisms, Drugs, and Acquired Resistance. Cancers, 13.","DOI":"10.3390\/cancers13112748"},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"1299","DOI":"10.1016\/j.bbamcr.2006.11.010","article-title":"ERK Implication in Cell Cycle Regulation","volume":"1773","author":"Chambard","year":"2007","journal-title":"Biochim. Biophys. Acta (BBA) Mol. Cell Res."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"e952","DOI":"10.1038\/cddis.2013.484","article-title":"Wentilactone A as a Novel Potential Antitumor Agent Induces Apoptosis and G2\/M Arrest of Human Lung Carcinoma Cells, and Is Mediated by HRas-GTP Accumulation to Excessively Activate the Ras\/Raf\/ERK\/P53-P21 Pathway","volume":"4","author":"Lv","year":"2013","journal-title":"Cell Death Dis."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"10","DOI":"10.1016\/j.biocel.2014.01.025","article-title":"Who Controls the ATP Supply in Cancer Cells? Biochemistry Lessons to Understand Cancer Energy Metabolism","volume":"50","author":"Saavedra","year":"2014","journal-title":"Int. J. Biochem. Cell Biol."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"144158","DOI":"10.1016\/j.gene.2019.144158","article-title":"Crucial Players in Glycolysis: Cancer Progress","volume":"726","author":"Abbaszadeh","year":"2020","journal-title":"Gene"},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"342666","DOI":"10.1016\/j.aca.2024.342666","article-title":"Development of an Analytical Platform for the Affinity Screening of Natural Extracts by SEC-MS towards PPAR\u03b1 and PPAR\u03b3 Receptors","volume":"1309","author":"Rinaldi","year":"2024","journal-title":"Anal. Chim. Acta"},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"76","DOI":"10.1016\/0003-2697(85)90442-7","article-title":"Measurement of Protein Using Bicinchoninic Acid","volume":"150","author":"Smith","year":"1985","journal-title":"Anal. Biochem."},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"680","DOI":"10.1038\/227680a0","article-title":"Cleavage of Structural Proteins during the Assembly of the Head of Bacteriophage T4","volume":"227","author":"Laemmli","year":"1970","journal-title":"Nature"},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"248","DOI":"10.1016\/0003-2697(76)90527-3","article-title":"A Rapid and Sensitive Method for the Quantitation of Microgram Quantities of Protein Utilizing the Principle of Protein-Dye Binding","volume":"72","author":"Bradford","year":"1976","journal-title":"Anal. Biochem."},{"key":"ref_59","first-page":"473","article-title":"Enzymes as Biochemical Reagents","volume":"1","author":"Bergmeyer","year":"1974","journal-title":"Methods Enzym. Anal."},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"124","DOI":"10.9734\/EJMP\/2015\/15015","article-title":"Phytochemicals of Markhamia Species (Bignoniaceae) and Their Therapeutic Value: A Review","volume":"6","author":"Ali","year":"2015","journal-title":"Eur. J. Med. Plants"},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"1113","DOI":"10.1002\/jlac.199419941110","article-title":"Caffeic Acid Glycoside Esters from Gratiola officinalis L.","volume":"1994","author":"Rothenburger","year":"1994","journal-title":"Liebigs Ann. Chem."},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"237","DOI":"10.1016\/j.fitote.2012.12.014","article-title":"Phenylethanoid glycosides from the stems of Callicarpa peii (hemostatic drug)","volume":"84","author":"Wu","year":"2013","journal-title":"Fitoterapia"}],"container-title":["International Journal of Molecular Sciences"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/1422-0067\/26\/5\/2220\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,9]],"date-time":"2025-10-09T16:45:05Z","timestamp":1760028305000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/1422-0067\/26\/5\/2220"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,2,28]]},"references-count":62,"journal-issue":{"issue":"5","published-online":{"date-parts":[[2025,3]]}},"alternative-id":["ijms26052220"],"URL":"https:\/\/doi.org\/10.3390\/ijms26052220","relation":{},"ISSN":["1422-0067"],"issn-type":[{"value":"1422-0067","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,2,28]]}}}