{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,23]],"date-time":"2026-09-23T01:23:32Z","timestamp":1790126612569,"version":"4.0.1"},"reference-count":69,"publisher":"MDPI AG","issue":"15","license":[{"start":{"date-parts":[[2026,7,23]],"date-time":"2026-07-23T00:00:00Z","timestamp":1784764800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"National Science Fund at the Ministry of Education and Science","award":["KP-06-H91\/13"],"award-info":[{"award-number":["KP-06-H91\/13"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Plants"],"abstract":"<jats:p>The comparative UHPLC-MS\/MS analysis of the underground and aerial parts of Primula veris subsp. columnae (Ten.) L\u0215di revealed a diverse range of secondary metabolites, including flavonoids and phenolic compounds and their glycosides, bisbibenzyl compounds and triterpene saponins. Phytochemical study of the aerial parts led to the isolation of 13 surface flavonoids and 6 flavonoid glycosides, of which 8,2\u2032-dimethoxy-5-hydroxyflavone and 8,3\u2032-dimethoxy-5-hydroxyflavone are new natural compounds. From the underground parts, a new triterpene saponin, primulasaponin VI, was isolated together with the known primulasaponins I, III and V; priverosaponin B 22-acetate; primulaverin; primverin; gaultherin; and 3-methoxy-4-primeverosylacetophenone. The structures of the new compounds were elucidated by NMR and HR-ESI-MS. The acetone extract obtained from the aerial parts and methanol extracts obtained from both aerial and underground parts were evaluated for their antibiofilm activity against Escherichia coli, Pseudomonas aeruginosa, and Staphylococcus aureus. The methanol extract obtained from underground parts possessed the strongest overall antibiofilm activity, particularly against E. coli and S. aureus. While the acetone extract inhibited the biofilm formation by E. coli and S. aureus, it promoted biofilm formation by P. aeruginosa. All tested extracts exhibited low cytotoxicity toward human keratinocytes and melanoma cells.<\/jats:p>","DOI":"10.3390\/plants15152259","type":"journal-article","created":{"date-parts":[[2026,7,23]],"date-time":"2026-07-23T15:05:42Z","timestamp":1784819142000},"page":"2259","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":1,"title":["Comparative Phytochemical Profiling and In Vitro Antibiofilm Activity of the Aerial and Underground Parts of Primula veris subsp. columnae (Ten.) L\u0215di"],"prefix":"10.3390","volume":"15","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-5273-176X","authenticated-orcid":false,"given":"Antoaneta","family":"Trendafilova","sequence":"first","affiliation":[{"name":"Institute of Organic Chemistry with Centre of Phytochemistry, Bulgarian Academy of Sciences, 1113 Sofia, Bulgaria"},{"name":"Centre of Competence \u201cSustainable Utilization of Bio-Resources and Waste of Medicinal and Aromatic Plants for Innovative Bioactive Products\u201d (BIORESOURCES BG), 1000 Sofia, Bulgaria"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0007-9056-0333","authenticated-orcid":false,"given":"Viktoria","family":"Ivanova","sequence":"additional","affiliation":[{"name":"Institute of Organic Chemistry with Centre of Phytochemistry, Bulgarian Academy of Sciences, 1113 Sofia, Bulgaria"},{"name":"Centre of Competence \u201cSustainable Utilization of Bio-Resources and Waste of Medicinal and Aromatic Plants for Innovative Bioactive Products\u201d (BIORESOURCES BG), 1000 Sofia, Bulgaria"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4984-041X","authenticated-orcid":false,"given":"Miroslav","family":"Novakovic","sequence":"additional","affiliation":[{"name":"Department of Chemistry, University of Belgrade\u2014Institute of Chemistry, Technology and Metallurgy, 11000 Belgrade, Serbia"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4235-3504","authenticated-orcid":false,"given":"Milka","family":"Jadranin","sequence":"additional","affiliation":[{"name":"Department of Chemistry, University of Belgrade\u2014Institute of Chemistry, Technology and Metallurgy, 11000 Belgrade, Serbia"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Plamena","family":"Staleva","sequence":"additional","affiliation":[{"name":"Institute of Organic Chemistry with Centre of Phytochemistry, Bulgarian Academy of Sciences, 1113 Sofia, Bulgaria"},{"name":"Research and Development and Innovation Consortium, Sofia Tech Park, 1784 Sofia, Bulgaria"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Petya","family":"Dimitrova","sequence":"additional","affiliation":[{"name":"The Stephan Angeloff Institute of Microbiology, Bulgarian Academy of Sciences, 1113 Sofia, Bulgaria"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ralitsa","family":"Veleva","sequence":"additional","affiliation":[{"name":"Centre of Competence \u201cSustainable Utilization of Bio-Resources and Waste of Medicinal and Aromatic Plants for Innovative Bioactive Products\u201d (BIORESOURCES BG), 1000 Sofia, Bulgaria"},{"name":"Department of Cellular and Developmental Biology, Faculty of Biology, Sofia University St. Kliment Ohridski, 1164 Sofia, Bulgaria"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1575-0790","authenticated-orcid":false,"given":"Tanya","family":"Topouzova-Hristova","sequence":"additional","affiliation":[{"name":"Centre of Competence \u201cSustainable Utilization of Bio-Resources and Waste of Medicinal and Aromatic Plants for Innovative Bioactive Products\u201d (BIORESOURCES BG), 1000 Sofia, Bulgaria"},{"name":"Department of Cellular and Developmental Biology, Faculty of Biology, Sofia University St. Kliment Ohridski, 1164 Sofia, Bulgaria"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8376-8917","authenticated-orcid":false,"given":"Tsvetelina","family":"Paunova-Krasteva","sequence":"additional","affiliation":[{"name":"The Stephan Angeloff Institute of Microbiology, Bulgarian Academy of Sciences, 1113 Sofia, Bulgaria"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2026,7,23]]},"reference":[{"key":"ref_1","unstructured":"(2026, June 06). Primulae Radix\u2014Herbal Medicinal Product|European Medicines Agency (EMA). Available online: https:\/\/www.ema.europa.eu\/en\/medicines\/herbal\/primulae-radix."},{"key":"ref_2","unstructured":"(2026, June 06). Primulae Flos\u2014Herbal Medicinal Product|European Medicines Agency (EMA). Available online: https:\/\/www.ema.europa.eu\/en\/medicines\/herbal\/primulae-flos."},{"key":"ref_3","doi-asserted-by":"crossref","unstructured":"Tsioutsiou, E.E., Amountzias, V., Vontzalidou, A., Dina, E., Stevanovi\u0107, Z.D., Cheilari, A., and Aligiannis, N. (2022). Medicinal Plants Used Traditionally for Skin Related Problems in the South Balkan and East Mediterranean Region\u2014A Review. Front. Pharmacol., 13.","DOI":"10.3389\/fphar.2022.936047"},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"17","DOI":"10.1016\/j.phymed.2018.09.015","article-title":"Composition and Cardioprotective Effects of Primula veris L. Solid Herbal Extract in Experimental Chronic Heart Failure","volume":"54","author":"Latypova","year":"2019","journal-title":"Phytomedicine"},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"2871579","DOI":"10.1155\/2017\/2871579","article-title":"Phenolics in Primula veris L. and P. elatior (L.) Hill Raw Materials","volume":"2017","author":"Baczek","year":"2017","journal-title":"Int. J. Anal. Chem."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"563","DOI":"10.1038\/nrmicro.2016.94","article-title":"Biofilms: An Emergent Form of Bacterial Life","volume":"14","author":"Flemming","year":"2016","journal-title":"Nat. Rev. Microbiol."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"276","DOI":"10.1093\/femsre\/fux010","article-title":"Molecular Mechanisms of Biofilm-Based Antibiotic Resistance and Tolerance in Pathogenic Bacteria","volume":"41","author":"Hall","year":"2017","journal-title":"FEMS Microbiol. Rev."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"1318","DOI":"10.1126\/science.284.5418.1318","article-title":"Bacterial Biofilms: A Common Cause of Persistent Infections","volume":"284","author":"Costerton","year":"1999","journal-title":"Science"},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1111\/apm.12099","article-title":"The Role of Bacterial Biofilms in Chronic Infections","volume":"121","author":"Bjarnsholt","year":"2013","journal-title":"Apmis"},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"88","DOI":"10.1080\/00087114.2008.10589614","article-title":"Antimitotic and Antibacterial Effects of the Primula veris L. Flower Extracts","volume":"61","author":"Biyik","year":"2008","journal-title":"Caryologia"},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"4475","DOI":"10.15376\/biores.18.3.4475-4491","article-title":"Chemical Content of Volatile Oil of Primula veris subsp. Columnae, Obtaining the Methanol Extracts and Their Biological Activities","volume":"18","year":"2023","journal-title":"Bioresources"},{"key":"ref_12","doi-asserted-by":"crossref","unstructured":"Ivani\u0161ov\u00e1, E., Hor\u0148\u00e1k, M., \u010cech, M., Harangozo, \u013d., Ka\u010d\u00e1niov\u00e1, M., Grygorieva, O., and Kowalczewski, P.\u0141. (2023). Polyphenol Content, Mineral Compounds Composition, Antimicrobial and Antioxidant Activities of Selected Medicinal Herbs from Slovak Republic. Appl. Sci., 13.","DOI":"10.3390\/app13031918"},{"key":"ref_13","doi-asserted-by":"crossref","unstructured":"Trendafilova, A., Raykova, D., Ivanova, V., Novakovic, M., Nedialkov, P., Paunova-Krasteva, T., Veleva, R., and Topouzova-Hristova, T. (2025). Phytochemical Characterization and Anti-Biofilm Activity of Primula veris L. Roots. Molecules, 30.","DOI":"10.3390\/molecules30081702"},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"1219","DOI":"10.1055\/a-1578-8731","article-title":"Development and Validation of an HPLC Method for the Analysis of Flowers of Wild-Growing Primula veris from Epirus, Greece","volume":"87","author":"Chintiroglou","year":"2021","journal-title":"Planta Med."},{"key":"ref_15","doi-asserted-by":"crossref","unstructured":"Apel, L., Kammerer, D.R., Stintzing, F.C., and Spring, O. (2017). Comparative Metabolite Profiling of Triterpenoid Saponins and Flavonoids in Flower Color Mutations of Primula veris L.. Int. J. Mol. Sci., 18.","DOI":"10.3390\/ijms18010153"},{"key":"ref_16","doi-asserted-by":"crossref","unstructured":"Stefanis, I., Chatzopoulou, P., Krigas, N., and Karioti, A. (2023). Exploring the Chemical Content of Primula veris L. subsp. veris Wild-Growing Populations along a Climate Gradient: An HPLC-PDA-MS Quality Assessment of Flowers, Leaves and Roots for Sustainable Exploitation. Horticulturae, 9.","DOI":"10.3390\/horticulturae9101120"},{"key":"ref_17","doi-asserted-by":"crossref","unstructured":"Graikou, K., Mpishinioti, A., Tsafantakis, N., Maloupa, E., Grigoriadou, K., and Chinou, I. (2023). Comparative Phytochemical Analyses of Flowers from Primula veris subsp. veris Growing Wild and from Ex Situ Cultivation in Greece. Foods, 12.","DOI":"10.3390\/foods12132623"},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"132","DOI":"10.1016\/j.phytochem.2017.07.005","article-title":"Phytochemistry of European Primula Species","volume":"143","author":"Colombo","year":"2017","journal-title":"Phytochemistry"},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"100510","DOI":"10.1016\/j.phyplu.2023.100510","article-title":"Genus Primula and Its Role in Phytomedicine; a Systematic Review","volume":"4","author":"Alam","year":"2024","journal-title":"Phytomed. Plus"},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"113719","DOI":"10.1016\/j.phytochem.2023.113719","article-title":"Bisbibenzyls from Serbian Primula veris subsp. Columnae (Ten.) L\u0215di and P. acaulis (L.) L.","volume":"212","author":"Novakovic","year":"2023","journal-title":"Phytochemistry"},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"712","DOI":"10.1007\/s10600-007-0241-8","article-title":"Riccardin C, a Bisbibenzyl Compound from Primula Macrocalyx","volume":"43","author":"Kosenkova","year":"2007","journal-title":"Chem. Nat. Compd."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"218","DOI":"10.1016\/j.chroma.2005.10.067","article-title":"Analysis of Phenolic Glycosides and Saponins in Primula elatior and Primula veris (Primula Root) by Liquid Chromatography, Evaporative Light Scattering Detection and Mass Spectrometry","volume":"1112","author":"Ganzera","year":"2006","journal-title":"J. Chromatogr. A"},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"90","DOI":"10.1016\/j.phytol.2021.06.014","article-title":"Hunting for Bis-Bibenzyls in Primula veris subsp. macrocalyx (Bunge) L\u00fcdi: Organ-Specific Accumulation and Cytotoxic Activity","volume":"44","author":"Bukvicki","year":"2021","journal-title":"Phytochem. Lett."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"1299","DOI":"10.1021\/np50087a019","article-title":"Triterpene Saponins from Primula veris subsp. Macrocalyx and Primula elatior subsp. Meyeri","volume":"55","author":"Wright","year":"1992","journal-title":"J. Nat. Prod."},{"key":"ref_25","doi-asserted-by":"crossref","unstructured":"W\u0142odarczyk, M., Pasikowski, P., Osiewa\u0142a, K., Frankiewicz, A., Dry\u015b, A., and Gle\u0144sk, M. (2020). In Search of High-Yielding and Single-Compound-Yielding Plants: New Sources of Pharmaceutically Important Saponins from the Primulaceae Family. Biomolecules, 10.","DOI":"10.3390\/biom10030376"},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"1033","DOI":"10.1016\/j.phytochem.2005.03.024","article-title":"Lipophilic Flavones of Primula veris L. from Field Cultivation and in Vitro Cultures","volume":"66","author":"Budzianowski","year":"2005","journal-title":"Phytochemistry"},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"665","DOI":"10.1016\/S0015-3796(86)80072-5","article-title":"Neue Flati Onoide Aus Primelmehl: New Flavonoids from Farinose Primula Exudates","volume":"181","author":"Wollenweber","year":"1986","journal-title":"Biochem. Und Physiol. Der Pflanz."},{"key":"ref_28","first-page":"587","article-title":"Diversification of Exudate Flavonoid Profiles in Further Primula spp.","volume":"7","author":"Bhutia","year":"2012","journal-title":"Nat. Prod. Commun."},{"key":"ref_29","first-page":"365","article-title":"Exudate Flavonoids of Primula spp: Structural and Biogenetic Chemodiversity","volume":"4","author":"Bhutia","year":"2009","journal-title":"Nat. Prod. Commun."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"419","DOI":"10.1016\/S0015-3796(17)30077-X","article-title":"Die Verbreitung Spezifischer Flavone in Der GattungPrimula","volume":"166","year":"1974","journal-title":"Biochem. Und Physiol. Der Pflanz."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"937","DOI":"10.1016\/j.phytochem.2010.03.004","article-title":"Chemodiversity of Exudate Flavonoids in Dionysia (Primulaceae): A Comparative Study","volume":"71","author":"Bhutia","year":"2010","journal-title":"Phytochemistry"},{"key":"ref_32","doi-asserted-by":"crossref","unstructured":"Ansari, P., Flatt, P.R., Harriott, P., and Abdel-Wahab, Y.H.A. (2020). Evaluation of the Antidiabetic and Insulin Releasing Effects of A. Squamosa, Including Isolation and Characterization of Active Phytochemicals. Plants, 9.","DOI":"10.3390\/plants9101348"},{"key":"ref_33","doi-asserted-by":"crossref","unstructured":"Ma, T., Sun, Y., Wang, L., Wang, J., Wu, B., Yan, T., and Jia, Y. (2022). An Investigation of the Anti-Depressive Properties of Phenylpropanoids and Flavonoids in Hemerocallis citrina Baroni. Molecules, 27.","DOI":"10.3390\/molecules27185809"},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"272","DOI":"10.1055\/s-2006-957425","article-title":"Structures of the Two Saponins Isolated from Commercially Available Root Extract of Primula sp.","volume":"64","author":"Siems","year":"1998","journal-title":"Planta Med."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"629","DOI":"10.2174\/138527208784577376","article-title":"Chemistry, Distribution and Biological Activities of 13,28-Epoxy-Oleanane Saponins from the Plant Families Myrsinaceae and Primulaceae","volume":"12","author":"Foubert","year":"2008","journal-title":"Curr. Org. Chem."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"817","DOI":"10.1080\/10286020.2011.596830","article-title":"Synthesis and Anti-Nociceptive and Anti-Inflammatory Effects of Gaultherin and Its Analogs","volume":"13","author":"Wang","year":"2011","journal-title":"J. Asian Nat. Prod. Res."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"4191","DOI":"10.1016\/0031-9422(92)80441-G","article-title":"Iridoid Glycosides from Penstemon Species: A C-5, C-9 Trans Iridoid and C-8 Epimeric Pairs","volume":"31","author":"Foderaro","year":"1992","journal-title":"Phytochemistry"},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"329","DOI":"10.1002\/fft2.473","article-title":"From Small-Scale Studies to an Encompassing View: Inhibiting Inflammation and Clinically Relevant Enzymes with Various Extracts of Primula Vulgaris Using in Vitro and in Silico Techniques","volume":"6","author":"Nilofar","year":"2025","journal-title":"Food Front."},{"key":"ref_39","unstructured":"(2026, June 05). MassBank of North America. Available online: https:\/\/mona.fiehnlab.ucdavis.edu\/spectra\/search."},{"key":"ref_40","doi-asserted-by":"crossref","unstructured":"Kurt-Celep, I., Zheleva-Dimitrova, D., Gevrenova, R., Uba, A.I., Zengin, G., Y\u0131ld\u0131ztugay, E., Picot-Allain, C.M.N., Lorenzo, J.M., Mahomoodally, M.F., and Montesano, D. (2022). An In-Depth Study on the Metabolite Profile and Biological Properties of Primula Auriculata Extracts: A Fascinating Sparkle on the Way from Nature to Functional Applications. Antioxidants, 11.","DOI":"10.3390\/antiox11071377"},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"518","DOI":"10.1016\/j.jfda.2017.06.009","article-title":"Systematic Study on Active Compounds as Antibacterial and Antibiofilm Agent in Aging Onions","volume":"26","author":"Sharma","year":"2018","journal-title":"J. Food Drug Anal."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"e01604","DOI":"10.1016\/j.heliyon.2019.e01604","article-title":"A Comparative Study on Chemical Composition, Antibiofilm and Biological Activities of Leaves Extracts of Four Tunisian Olive Cultivars","volume":"5","author":"Edziri","year":"2019","journal-title":"Heliyon"},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"e18114","DOI":"10.1590\/s2175-97902019000218114","article-title":"Antimicrobial and Antibiofilm Activities of Phenolic Compounds Extracted from Populus nigra and Populus alba Buds (Algeria)","volume":"55","author":"Nassima","year":"2019","journal-title":"Braz. J. Pharm. Sci."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"96","DOI":"10.1186\/s13020-025-01147-5","article-title":"Natural Phytochemical-Based Strategies for Antibiofilm Applications","volume":"20","author":"Zhou","year":"2025","journal-title":"Chin. Med."},{"key":"ref_45","doi-asserted-by":"crossref","unstructured":"Slobodn\u00edkov\u00e1, L., Fialov\u00e1, S., Rendekov\u00e1, K., Kov\u00e1\u010d, J., and Mu\u010daji, P. (2016). Antibiofilm Activity of Plant Polyphenols. Molecules, 21.","DOI":"10.3390\/molecules21121717"},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"299","DOI":"10.1016\/S0031-9422(99)00525-7","article-title":"Two Flavonoid Glycosides from Chenopodium murale","volume":"53","author":"Gohar","year":"2000","journal-title":"Phytochemistry"},{"key":"ref_47","doi-asserted-by":"crossref","unstructured":"Bouyahya, A., El Omari, N., El Menyiy, N., Guaouguaou, F.E., Balahbib, A., and Chamkhi, I. (2022). Anti-Quorum Sensing Agents from Natural Sources. Antimicrobial Resistance: Underlying Mechanisms and Therapeutic Approaches, Springer Nature.","DOI":"10.1007\/978-981-16-3120-7_17"},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"143","DOI":"10.1007\/s11274-019-2719-5","article-title":"An Overview on Anti-Biofilm Properties of Quercetin against Bacterial Pathogens","volume":"35","author":"Memariani","year":"2019","journal-title":"World J. Microbiol. Biotechnol."},{"key":"ref_49","doi-asserted-by":"crossref","unstructured":"Ming, D., Wang, D., Cao, F., Xiang, H., Mu, D., Cao, J., Li, B., Zhong, L., Dong, X., and Zhong, X. (2017). Kaempferol Inhibits the Primary Attachment Phase of Biofilm Formation in Staphylococcus aureus. Front. Microbiol., 8.","DOI":"10.3389\/fmicb.2017.02263"},{"key":"ref_50","doi-asserted-by":"crossref","unstructured":"Nazzaro, F., Coppola, F., Fratianni, F., Abdalrazeq, M., Ombra, M.N., De Giulio, B., Coppola, R., and Zengin, G. (2025). Polyphenols Bioactive Metabolites, and Their Anti-Biofilm and Neuroprotective Potential. Foods, 14.","DOI":"10.3390\/foods14223976"},{"key":"ref_51","first-page":"439","article-title":"Tradescantia Pallida Extract Inhibits Biofilm Formation in Pseudomonas aeruginosa","volume":"81","author":"Kamiya","year":"2019","journal-title":"Nagoya J. Med. Sci."},{"key":"ref_52","doi-asserted-by":"crossref","unstructured":"Dudek-Wicher, R., Junka, A.F., Migda\u0142, P., Korzeniowska-Kowal, A., Wzorek, A., and Bartoszewicz, M. (2022). The Antibiofilm Activity of Selected Substances Used in Oral Health Prophylaxis. BMC Oral Health, 22.","DOI":"10.1186\/s12903-022-02532-4"},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"1073","DOI":"10.1111\/apm.12083","article-title":"Effect of Plant Phenolic Compounds on Biofilm Formation by Pseudomonas aeruginosa","volume":"121","author":"Plyuta","year":"2013","journal-title":"Apmis"},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"519","DOI":"10.1099\/jmm.0.46804-0","article-title":"Effects of Oregano, Carvacrol and Thymol on Staphylococcus aureus and Staphylococcus epidermidis Biofilms","volume":"56","author":"Nostro","year":"2007","journal-title":"J. Med. Microbiol."},{"key":"ref_55","doi-asserted-by":"crossref","unstructured":"Hamion, G., Aucher, W., Tardif, C., Miranda, J., Rouger, C., Imbert, C., and Girardot, M. (2022). Valorization of Invasive Plant Extracts against the Bispecies Biofilm Staphylococcus aureus\u2013Candida albicans by a Bioguided Molecular Networking Screening. Antibiotics, 11.","DOI":"10.3390\/antibiotics11111595"},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"19434","DOI":"10.3390\/ijms141019434","article-title":"Systematic Exploration of Natural and Synthetic Flavonoids for the Inhibition of Staphylococcus aureus Biofilms","volume":"14","author":"Manner","year":"2013","journal-title":"Int. J. Mol. Sci."},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"193","DOI":"10.1016\/j.micpath.2017.03.033","article-title":"Inhibitory Effects of Flavonoids on Biofilm Formation by Staphylococcus aureus That Overexpresses Efflux Protein Genes","volume":"107","author":"Lopes","year":"2017","journal-title":"Microb. Pathog."},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"587","DOI":"10.1079\/BJN2002725","article-title":"The Biological Action of Saponins in Animal Systems: A Review","volume":"88","author":"Francis","year":"2002","journal-title":"Br. J. Nutr."},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"219","DOI":"10.1016\/j.jep.2004.05.016","article-title":"Biological Activities and Distribution of Plant Saponins","volume":"94","author":"Sparg","year":"2004","journal-title":"J. Ethnopharmacol."},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"343","DOI":"10.1016\/j.ijantimicag.2005.09.002","article-title":"Antimicrobial Activity of Flavonoids","volume":"26","author":"Cushnie","year":"2005","journal-title":"Int. J. Antimicrob. Agents"},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"21988","DOI":"10.1038\/s41598-022-26399-1","article-title":"Antimicrobial and Anti-Biofilm Activity of Polygonum Chinense L. Aqueous Extract against Staphylococcus aureus","volume":"12","author":"Zeng","year":"2022","journal-title":"Sci. Rep."},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"21416","DOI":"10.1038\/s41598-020-78379-y","article-title":"Mechanism and Antibacterial Activity of Vine Tea Extract and Dihydromyricetin against Staphylococcus aureus","volume":"10","author":"Liang","year":"2020","journal-title":"Sci. Rep."},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"3132","DOI":"10.1128\/AAC.00124-10","article-title":"Damage of the Bacterial Cell Envelope by Antimicrobial Peptides Gramicidin S and PGLa as Revealed by Transmission and Scanning Electron Microscopy","volume":"54","author":"Hartmann","year":"2010","journal-title":"Antimicrob. Agents Chemother."},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"99","DOI":"10.1016\/j.ijantimicag.2011.02.014","article-title":"Recent Advances in Understanding the Antibacterial Properties of Flavonoids","volume":"38","author":"Cushnie","year":"2011","journal-title":"Int. J. Antimicrob. Agents"},{"key":"ref_65","doi-asserted-by":"crossref","unstructured":"Juang, Y.-P., and Liang, P.-H. (2020). Biological and Pharmacological Effects of Synthetic Saponins. Molecules, 25.","DOI":"10.3390\/molecules25214974"},{"key":"ref_66","doi-asserted-by":"crossref","unstructured":"Stachelska, M.A., Karpi\u0144ski, P., and Kruszewski, B. (2025). A Comprehensive Review of Biological Properties of Flavonoids and Their Role in the Prevention of Metabolic, Cancer and Neurodegenerative Diseases. Appl. Sci., 15.","DOI":"10.3390\/app151910840"},{"key":"ref_67","doi-asserted-by":"crossref","unstructured":"Boyadzhieva, S.S., Tsvetanova, F.V., Coelho, J.A.P., Staleva, P., Kamenova-Nacheva, M., Taneva, S., and Stateva, R.P. (2026). Boosting the Sustainable Transformation of Cornus mas L. Stones Using a Hybrid Strategy Involving Microwave-Assisted Extraction. Molecules, 31.","DOI":"10.3390\/molecules31030525"},{"key":"ref_68","doi-asserted-by":"crossref","unstructured":"Paunova-Krasteva, T., Dimitrova, P.D., Damyanova, T., Borisova, D., Leseva, M., Uzunova, I., Dimitrova, P.A., Ivanova, V., Trendafilova, A., and Veleva, R. (2025). Interference of Pseudomonas Aeruginosa Virulence Factors by Different Extracts from Inula Species. Pharmaceuticals, 18.","DOI":"10.3390\/ph18121824"},{"key":"ref_69","doi-asserted-by":"crossref","unstructured":"Trendafilova, A., Staleva, P., Petkova, Z., Ivanova, V., Evstatieva, Y., Nikolova, D., Rasheva, I., Atanasov, N., Topouzova-Hristova, T., and Veleva, R. (2023). Phytochemical Profile, Antioxidant Potential, Antimicrobial Activity, and Cytotoxicity of Dry Extract from Rosa damascena Mill. Molecules, 28.","DOI":"10.3390\/molecules28227666"}],"container-title":["Plants"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2223-7747\/15\/15\/2259\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,7,23]],"date-time":"2026-07-23T15:27:51Z","timestamp":1784820471000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2223-7747\/15\/15\/2259"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,7,23]]},"references-count":69,"journal-issue":{"issue":"15","published-online":{"date-parts":[[2026,8]]}},"alternative-id":["plants15152259"],"URL":"https:\/\/doi.org\/10.3390\/plants15152259","relation":{},"ISSN":["2223-7747"],"issn-type":[{"value":"2223-7747","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,7,23]]}}}