{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2025,10,10]],"date-time":"2025-10-10T01:36:30Z","timestamp":1760060190516,"version":"build-2065373602"},"reference-count":93,"publisher":"MDPI AG","issue":"16","license":[{"start":{"date-parts":[[2025,8,9]],"date-time":"2025-08-09T00:00:00Z","timestamp":1754697600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"National Agency for Scientific Research, Technology and Innovation (NASRI)"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Molecules"],"abstract":"<jats:p>The growing demand for natural preservatives has driven interest in essential oils (EOs) from medicinal and aromatic plants. This study examines the potential of EOs from six wild populations of Albanian Lamiaceae, specifically Origanum vulgare subsp. hirtum, Thymbra capitata, and Satureja montana species, to be utilized for food conservation, among other possible uses. The EOs were extracted by hydrodistillation, and their chemical profiles were analyzed through GC-MS. DPPH and ABTS assays were performed to evaluate antioxidant activity. The antimicrobial efficacy of the oils was assessed using the broth microdilution method against six common foodborne pathogens: Salmonella enterica serovar Enteritidis, Escherichia coli, Pseudomonas aeruginosa, Stenotrophomonas maltophilia, Micrococcus luteus, and one fungus, Candida albicans. The most potent EOs in terms of yield and biological activity, resulting from O. vulgare subsp. hirtum and T. capitata, were encapsulated in oil-in-water emulsions, which were characterized for particle size and zeta potential. The results show that the populations of O.vulgare subsp. hirtum and T. capitata taken in the study belong to carvacrol chemotypes, and their EOs show strong antioxidant activity and are effective against all tested microorganisms. Nanoemulsions prepared with these EOs showed promising stability, indicating their potential as natural preservatives in food applications.<\/jats:p>","DOI":"10.3390\/molecules30163329","type":"journal-article","created":{"date-parts":[[2025,8,11]],"date-time":"2025-08-11T09:04:53Z","timestamp":1754903093000},"page":"3329","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Essential Oils from Wild Albanian Lamiaceae: GC-MS Profiling, Biological Activity, and Enhanced Delivery via Nanoencapsulation"],"prefix":"10.3390","volume":"30","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-7616-172X","authenticated-orcid":false,"given":"Elton","family":"Basha","sequence":"first","affiliation":[{"name":"Department of Agri-Food Technology, Agricultural University of Tirana, 1025 Tirana, Albania"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Erjon","family":"Mamo\u00e7i","sequence":"additional","affiliation":[{"name":"Department of Food Science and Biotechnology, Agricultural University of Tirana, 1025 Tirana, Albania"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8447-8495","authenticated-orcid":false,"given":"Aniket","family":"Sharma","sequence":"additional","affiliation":[{"name":"Department of Animal Science, University of Wyoming, Wyoming, WY 82071, USA"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Entela","family":"Hodaj-\u00c7eliku","sequence":"additional","affiliation":[{"name":"Department of Chemistry, Agricultural University of Tirana, 1025 Tirana, Albania"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-4782-8393","authenticated-orcid":false,"given":"Sanije","family":"Zejnelhoxha","sequence":"additional","affiliation":[{"name":"Department of Food Science and Biotechnology, Agricultural University of Tirana, 1025 Tirana, Albania"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3621-9632","authenticated-orcid":false,"given":"M\u0103d\u0103lina L.","family":"Medeleanu","sequence":"additional","affiliation":[{"name":"Department of Food Science, Faculty of Food Science and Technology, University of Agricultural Sciences and Veterinary Medicine of Cluj-Napoca, 400372 Cluj-Napoca, Romania"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0594-4628","authenticated-orcid":false,"given":"Sonia A.","family":"Socaci","sequence":"additional","affiliation":[{"name":"Department of Food Science, Faculty of Food Science and Technology, University of Agricultural Sciences and Veterinary Medicine of Cluj-Napoca, 400372 Cluj-Napoca, Romania"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3145-8025","authenticated-orcid":false,"given":"Bledar","family":"Bisha","sequence":"additional","affiliation":[{"name":"Department of Animal Science, University of Wyoming, Wyoming, WY 82071, USA"}],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"1968","published-online":{"date-parts":[[2025,8,9]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"9901018","DOI":"10.1155\/2022\/9901018","article-title":"Potentials of Natural Preservatives to Enhance Food Safety and Shelf Life: A Review","volume":"2022","author":"Teshome","year":"2022","journal-title":"Sci. World J."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"114548","DOI":"10.1016\/j.foodres.2024.114548","article-title":"Activity and Safety Evaluation of Natural Preservatives","volume":"190","author":"Dong","year":"2024","journal-title":"Food Res. Int."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"446","DOI":"10.1016\/j.fct.2007.09.106","article-title":"Biological Effects of Essential Oils\u2014A Review","volume":"46","author":"Bakkali","year":"2008","journal-title":"Food Chem. Toxicol."},{"key":"ref_4","doi-asserted-by":"crossref","unstructured":"Sun, J., Sun, P., Kang, C., Zhang, L., Guo, L., and Kou, Y. (2022). Chemical Composition and Biological Activities of Essential Oils from Six Lamiaceae Folk Medicinal Plants. Front. Plant Sci., 13.","DOI":"10.3389\/fpls.2022.919294"},{"key":"ref_5","doi-asserted-by":"crossref","unstructured":"Ietswaart, J.H. (1980). A Taxonomic Revision of the Genus Origanum (Labiatae), Leiden University Press.","DOI":"10.1007\/978-94-009-9156-9"},{"key":"ref_6","unstructured":"Vangjeli, J., Ruci, B., and Mullaj, A. (1996). Red Book of Albanian Flora, Institute of Biological Research, Academy of Sciences of Albania."},{"key":"ref_7","unstructured":"Padulosi, S. (1996, January 8\u201312). Taxonomy, Diversity and Distribution of Origanum Species. Proceedings of the IPGRI International Workshop on Oregano, Bari, Italy."},{"key":"ref_8","first-page":"449","article-title":"Genetic Variation of Oregano (Origanum vulgare L.) for Etheric Oil in Albania","volume":"4","author":"Elezi","year":"2013","journal-title":"Agric. Sci."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"5388","DOI":"10.1021\/jf061477i","article-title":"Polar Constituents from the Aerial Parts of Origanum vulgare L. ssp. hirtum Growing Wild in Greece","volume":"54","author":"Koukoulitsa","year":"2006","journal-title":"J. Agric. Food Chem."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"3120","DOI":"10.2174\/138161208786404218","article-title":"Portuguese Thymbra and Thymus Species Volatiles: Chemical Composition and Biological Activities","volume":"14","author":"Figueiredo","year":"2008","journal-title":"Curr. Pharm. Des."},{"key":"ref_11","doi-asserted-by":"crossref","unstructured":"Pirintsos, S.A., Bariotakis, M., Kampa, M., Sourvinos, G., Lionis, C., and Castanas, E. (2020). The Therapeutic Potential of the Essential Oil of Thymbra capitata (L.) Cav., Origanum dictamnus L. and Salvia fruticosa Mill. and a Case of Plant-Based Pharmaceutical Development. Front. Pharmacol., 11.","DOI":"10.3389\/fphar.2020.522213"},{"key":"ref_12","first-page":"58","article-title":"Analysis of Essential Oils of Three Wild Medicinal Plants in Albania","volume":"5","author":"Ibraliu","year":"2011","journal-title":"J. Med. Plants Res."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"463","DOI":"10.1016\/j.tifs.2022.10.014","article-title":"Phytochemistry, Pharmacological Investigations, Industrial Applications, and Encapsulation of Thymbra capitata L.: A Review","volume":"129","author":"Saoulajan","year":"2022","journal-title":"Trends Food Sci. Technol."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"8261","DOI":"10.1021\/jf049113i","article-title":"Essential Oils of Satureja, Origanum, and Thymbra Species: Chemical Composition and Antibacterial Activities Against Foodborne Pathogens","volume":"52","author":"Chorianopoulos","year":"2004","journal-title":"J. Agric. Food Chem."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"5377","DOI":"10.1007\/s11947-025-03759-z","article-title":"Enhanced Pathogen Control Through Thymol and Carvacrol Nanoemulsions: A Microfluidization Approach","volume":"18","author":"Ozogul","year":"2025","journal-title":"Food Bioprocess Technol."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"568","DOI":"10.1089\/jmf.2007.0520","article-title":"Antimicrobial Activities of the Essential Oils of Origanum onites L., Origanum vulgare L. Subspecies hirtum (Link) Ietswaart, Satureja thymbra L., and Thymbra cilicicus Boiss. & Bal. Growing Wild in Turkey","volume":"11","author":"Sarac","year":"2008","journal-title":"J. Med. Food"},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"235","DOI":"10.1002\/(SICI)1099-1026(1998070)13:4<235::AID-FFJ733>3.0.CO;2-T","article-title":"Antimicrobial and Antioxidant Properties of Some Commercial Essential Oils","volume":"13","author":"Baratta","year":"1998","journal-title":"Flavour Fragr. J."},{"key":"ref_18","first-page":"1035","article-title":"Chemical Composition, Olfactory Evaluation and Antioxidant Effects of the Essential Oil of Satureja montana L.","volume":"3","author":"Stoilova","year":"2008","journal-title":"Nat. Prod. Commun."},{"key":"ref_19","doi-asserted-by":"crossref","unstructured":"Chroho, M., Rouphael, Y., Petropoulos, S.A., and Bouissane, L. (2024). Carvacrol and Thymol Content Affects the Antioxidant and Antibacterial Activity of Origanum compactum and Thymbra zygis Essential Oils. Antibiotics, 13.","DOI":"10.3390\/antibiotics13020139"},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"1451","DOI":"10.3390\/ph6121451","article-title":"Effect of Essential Oils on Pathogenic Bacteria","volume":"6","author":"Nazzaro","year":"2013","journal-title":"Pharmaceuticals"},{"key":"ref_21","doi-asserted-by":"crossref","unstructured":"D\u2019Amato, S., Rossi, C., Maggio, F., Valbonetti, L., Savini, V., Paparella, A., and Serio, A. (2024). Antilisterial Effectiveness of Origanum vulgare var. hirtum and CoridoThymbra capitata Essential Oils and Hydrolates Alone and in Combination. Foods, 13.","DOI":"10.3390\/foods13060860"},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"223","DOI":"10.1016\/j.ijfoodmicro.2004.03.022","article-title":"Essential Oils: Their Antibacterial Properties and Potential Applications in Foods\u2014A Review","volume":"94","author":"Burt","year":"2004","journal-title":"Int. J. Food Microbiol."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"1545","DOI":"10.4315\/0362-028X-65.10.1545","article-title":"Bactericidal Activities of Plant Essential Oils and Some of Their Isolated Constituents Against Campylobacter jejuni, Escherichia coli, Listeria monocytogenes, and Salmonella enterica","volume":"65","author":"Friedman","year":"2002","journal-title":"J. Food Prot."},{"key":"ref_24","doi-asserted-by":"crossref","unstructured":"de Castro, R.D., de Souza, T.M., Bezerra, L.M., Ferreira, G.L., Costa, E.M., and Cavalcanti, A.L. (2015). Antifungal Activity and Mode of Action of Thymol and Its Synergism with Nystatin Against Candida Species Involved with Infections in the Oral Cavity: An In Vitro Study. BMC Complement. Altern. Med., 15.","DOI":"10.1186\/s12906-015-0947-2"},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"1719","DOI":"10.1039\/C2SM06903B","article-title":"Nanoemulsions versus Microemulsions: Terminology, Differences, and Similarities","volume":"8","author":"McClements","year":"2012","journal-title":"Soft Matter"},{"key":"ref_26","doi-asserted-by":"crossref","unstructured":"Maurya, A., Singh, V.K., Das, S., Prasad, J., Kedia, A., Upadhyay, N., Dubey, N.K., and Dwivedy, A.K. (2021). Essential Oil Nanoemulsion as Eco-Friendly and Safe Preservative: Bioefficacy Against Microbial Food Deterioration and Toxin Secretion, Mode of Action, and Future Opportunities. Front. Microbiol., 12.","DOI":"10.3389\/fmicb.2021.751062"},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"100066","DOI":"10.1016\/j.onano.2022.100066","article-title":"Preparation, Stability and Biological Activity of Essential Oil-Based Nanoemulsions: A Comprehensive Review","volume":"8","author":"Singh","year":"2022","journal-title":"OpenNano"},{"key":"ref_28","first-page":"327","article-title":"AnTC-Microbial Potential of Nano-Emulsion Form of Essential Oil Obtained from Aerial Parts of Origanum vulgare L. as Food Additive","volume":"11","author":"Enayatifard","year":"2021","journal-title":"Adv. Pharm. Bull."},{"key":"ref_29","doi-asserted-by":"crossref","unstructured":"Jayari, A., Dons\u00ec, F., Ferrari, G., and Maaroufi, A. (2022). Nanoencapsulation of Thyme Essential Oils: Formulation, Characterization, Storage Stability, and Biological Activity. Foods, 11.","DOI":"10.3390\/foods11131858"},{"key":"ref_30","doi-asserted-by":"crossref","unstructured":"Rinaldi, F., Maurizi, L., Conte, A.L., Marazzato, M., Maccelli, A., Crestoni, M.E., Hanieh, P.N., Forte, J., Conte, M.P., and Zagaglia, C. (2021). Nanoemulsions of Satureja montana Essential Oil: Antimicrobial and Antibiofilm Activity Against Avian Escherichia coli Strains. Pharmaceutics, 13.","DOI":"10.3390\/pharmaceutics13020134"},{"key":"ref_31","first-page":"785","article-title":"Antioxidant Activity and Chemical Composition of Essential Oils of Some Aromatic and Medicinal Plants from Albania","volume":"12","author":"Tsiftsoglou","year":"2017","journal-title":"Nat. Prod. Commun."},{"key":"ref_32","doi-asserted-by":"crossref","unstructured":"Giannoulis, K.D., Kamvoukou, C.-A., Gougoulias, N., and Wogiatzi, E. (2020). Irrigation and Nitrogen Application Affect Greek Oregano (Origanum vulgare ssp. hirtum) Dry Biomass, Essential Oil Yield and Composition. Ind. Crops Prod., 150.","DOI":"10.1016\/j.indcrop.2020.112392"},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"527","DOI":"10.1351\/pac200274040527","article-title":"Aromatic biodiversity among the flowering plant taxa of Turkey","volume":"74","author":"Baser","year":"2002","journal-title":"Pure Appl. Chem."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"1738","DOI":"10.1021\/jf9708296","article-title":"Antifungal activities of Origanum vulgare subsp. hirtum, Mentha spicata, Lavandula angustifolia and Salvia fruticosa Essential Oils against Human Pathogenic Fungi","volume":"46","author":"Adam","year":"1998","journal-title":"J. Agric. Food Chem."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"169","DOI":"10.54660\/.IJMRGE.2024.5.4.169-178","article-title":"Factors Influencing Yield, Chemical Composition and Efficacy of Essential Oils","volume":"5","author":"Mugao","year":"2024","journal-title":"Int. J. Multidiscip. Res. Growth Eval."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"26","DOI":"10.1002\/ffj.1029","article-title":"Variability in essential oil composition of Tunisian Thymbra capitata (L.) Hoffmanns. et Link","volume":"17","author":"Hedhili","year":"2002","journal-title":"Flavour Fragr. J."},{"key":"ref_37","doi-asserted-by":"crossref","unstructured":"Tagnaout, I., Zerkani, H., Hadi, N., El Moumen, B., El Makhoukhi, F., Bouhrim, M., Al-Salahi, R., Nasr, F.A., Mechchate, H., and Zair, T. (2022). Chemical Composition, Antioxidant and Antibacterial Activities of Thymbra broussonetii Boiss and Thymbra capitata (L.) Hoffmann and Link Essential Oils. Plants, 11.","DOI":"10.3390\/plants11070954"},{"key":"ref_38","first-page":"750","article-title":"Chemical Composition of the Essential Oil, Total Phenolics, Total Flavonoids and Antioxidant Activity of Methanolic Extracts of Satureja montana L.","volume":"10","author":"Hajdari","year":"2016","journal-title":"Rec. Nat. Prod."},{"key":"ref_39","doi-asserted-by":"crossref","unstructured":"Said-Al Ahl, H.A.H., Ka\u010d\u00e1niova, M., Mahmoud, A.A., Hikal, W.M., \u010cmikov\u00e1, N., Szczepanek, M., B\u0142aszczyk, K., Al-Balawi, S.M., Bianchi, A., and Smaoui, S. (2024). Phytochemical Characterization and Biological Activities of Essential Oil from Satureja montana L., a Medicinal Plant Grown under the Influence of Fertilization and Planting Dates. Biology, 13.","DOI":"10.3390\/biology13050328"},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"159","DOI":"10.24326\/asphc.2017.5.16","article-title":"Variation in winter savory (Satureja montana L.) yield and essential oil production as affected by different plant density and number of harvests","volume":"16","year":"2017","journal-title":"Acta Sci. Pol. Hortorum Cultus"},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"1554","DOI":"10.1002\/jsfa.4347","article-title":"Antioxidant and Antimicrobial Activity of Satureja montana L. Extracts","volume":"91","author":"Serrano","year":"2011","journal-title":"J. Sci. Food Agric."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"517","DOI":"10.1016\/0305-1978(94)90046-9","article-title":"Pattern of Geographic Variations of Origanum vulgare Trichomes and Essential Oil Content in Greece","volume":"22","author":"Kokkini","year":"1994","journal-title":"Biochem. Syst. Ecol."},{"key":"ref_43","doi-asserted-by":"crossref","unstructured":"M\u0105czka, W., Twardawska, M., Grabarczyk, M., and Wi\u0144ska, K. (2023). Carvacrol\u2014A Natural Phenolic Compound with Antimicrobial Properties. Antibiotics, 12.","DOI":"10.3390\/antibiotics12050824"},{"key":"ref_44","doi-asserted-by":"crossref","unstructured":"Maniki, E., Kostoglou, D., Paterakis, N., Nikolaou, A., Kourkoutas, Y., Papachristoforou, A., and Giaouris, E. (2023). Chemical Composition, Antioxidant, and Antibiofilm Properties of Essential Oil from Thymbra capitata Plants Organically Cultured on the Greek Island of Lemnos. Molecules, 28.","DOI":"10.3390\/molecules28031154"},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"572","DOI":"10.1055\/s-2004-827162","article-title":"Chemical Composition and Antifungal Activity of the Essential Oil of Thymbra capitata","volume":"70","author":"Salgueiro","year":"2004","journal-title":"Planta Med."},{"key":"ref_46","unstructured":"(2008). Oil of Origanum, Spanish Type [Coridothymus capitatus (L.) Rchb.f.] (Standard No. ISO 14717:2008)."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"88","DOI":"10.18689\/mjai-1000117","article-title":"Chemical Characterisation of Essential Oil for Satureja montana Population from Burreli Area, Albania","volume":"4","author":"Salihila","year":"2018","journal-title":"Madridge J. Anal. Sci. Instrum."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"402","DOI":"10.1016\/j.foodchem.2016.04.111","article-title":"Antibacterial and Antifungal Activities of Thymol: A Brief Review of the Literature","volume":"210","author":"Marchese","year":"2016","journal-title":"Food Chem."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"e2110092118","DOI":"10.1073\/pnas.2110092118","article-title":"The Biosynthesis of Thymol, Carvacrol, and Thymohydroquinone in Lamiaceae Proceeds via Cytochrome P450s and a Short-Chain Dehydrogenase","volume":"118","author":"Krause","year":"2021","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"3042","DOI":"10.1080\/10408398.2019.1675585","article-title":"The Antibacterial Properties of Phenolic Isomers, Carvacrol and Thymol","volume":"60","author":"Kachur","year":"2020","journal-title":"Crit. Rev. Food Sci. Nutr."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"59","DOI":"10.1016\/S0308-8146(98)00086-7","article-title":"Antioxidant Activity and Mechanism of Action of Thymol and Carvacrol in Two Lipid Systems","volume":"64","author":"Yanishlieva","year":"1999","journal-title":"Food Chem."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"3989","DOI":"10.1021\/jf073272v","article-title":"Comparative Study on the Antioxidant and Biological Activities of Carvacrol, Thymol, and Eugenol Derivatives","volume":"56","year":"2008","journal-title":"J. Agric. Food Chem."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"2083","DOI":"10.1007\/s00706-016-1837-0","article-title":"Essential Oils as Antioxidants: Their Evaluation by DPPH, ABTS, FRAP, CUPRAC, and \u03b2-Carotene Bleaching Methods","volume":"147","author":"Olszowy","year":"2016","journal-title":"Monatsh. Chem."},{"key":"ref_54","doi-asserted-by":"crossref","unstructured":"Ebhohimen, I.E., Okolie, N.P., Okpeku, M., Unweator, M., Adeleke, V.T., and Edemhanria, L. (2023). Evaluation of the Antioxidant Properties of Carvacrol as a Prospective Replacement for Crude Essential Oils and Synthetic Antioxidants in Food Storage. Molecules, 28.","DOI":"10.3390\/molecules28031315"},{"key":"ref_55","unstructured":"(2024). Performance Standards for Antimicrobial Susceptibility Testing, 34th Ed (Standard No. M100 Ed34)."},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"386","DOI":"10.1093\/jac\/dkh359","article-title":"Tolerance of Pseudomonas aeruginosa to Melaleuca alternifolia (Tea Tree) Oil Is Associated with the Outer Membrane and Energy-Dependent Cellular Processes","volume":"54","author":"Longbottom","year":"2004","journal-title":"J. Antimicrob. Chemother."},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"1013","DOI":"10.1080\/0972060X.2014.971069","article-title":"Chemical Profile, Antioxidant and Antibacterial Activity of Thyme and Oregano Essential Oils, Thymol and Carvacrol and Their Possible Synergism","volume":"18","author":"Gavaric","year":"2015","journal-title":"J. Essent. Oil Bear. Plants"},{"key":"ref_58","first-page":"1","article-title":"Comprehensive Review of the Combined Antimicrobial Activity of Essential Oil Mixtures and Synergism with Conventional Antimicrobials","volume":"20","author":"Soulaimani","year":"2025","journal-title":"Nat. Prod. Commun."},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"489","DOI":"10.1007\/s10863-021-09906-3","article-title":"Effect of Carvacrol and Thymol on NorA Efflux Pump Inhibition in Multidrug-Resistant (MDR) Staphylococcus aureus Strains","volume":"53","author":"Scherf","year":"2021","journal-title":"J. Bioenerg. Biomembr."},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"443","DOI":"10.1016\/j.phymed.2013.10.016","article-title":"Synergistic Interactions in Two-Drug and Three-Drug Combinations (Thymol, EDTA and Vancomycin) Against Multidrug-Resistant Bacteria Including E. coli","volume":"21","author":"Hamoud","year":"2014","journal-title":"Phytomedicine"},{"key":"ref_61","doi-asserted-by":"crossref","unstructured":"Wijesundara, N., Lee, S., Cheng, Z., Davidson, R., and Rupasinghe, H. (2021). Carvacrol Exhibits Rapid Bactericidal Activity Against Streptococcus pyogenes Through Cell Membrane Damage. Sci. Rep., 11.","DOI":"10.1038\/s41598-020-79713-0"},{"key":"ref_62","doi-asserted-by":"crossref","unstructured":"Gan, C., Langa, E., Valenzuela, A., Ballestero, D., and Pino-Ot\u00edn, M. (2023). Synergistic Activity of Thymol with Commercial Antibiotics Against Critical and High WHO Priority Pathogenic Bacteria. Plants, 12.","DOI":"10.3390\/plants12091868"},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"5964","DOI":"10.1080\/14786419.2020.1808635","article-title":"Anti-Acinetobacter baumannii Activity of Selected Phytochemicals Alone, in Binary Combinations and in Combinations with Conventional Antibiotics","volume":"35","author":"Nikolic","year":"2021","journal-title":"Nat. Prod. Res."},{"key":"ref_64","doi-asserted-by":"crossref","unstructured":"Bisso Ndezo, B., Tokam Kuate, C.R., and Dzoyem, J.P. (2021). Synergistic Antibiofilm Efficacy of Thymol and Piperine in Combination with Three Aminoglycoside Antibiotics Against Klebsiella pneumoniae Biofilms. Can. J. Infect. Dis. Med. Microbiol., 2021.","DOI":"10.1155\/2021\/7029944"},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"164","DOI":"10.1016\/j.ijfoodmicro.2010.04.001","article-title":"Use of Natural Antimicrobials to Increase Antibiotic Susceptibility of Drug Resistant Bacteria","volume":"140","author":"Palaniappan","year":"2010","journal-title":"Int. J. Food Microbiol."},{"key":"ref_66","doi-asserted-by":"crossref","unstructured":"Gan, C., Langa, E., Wang, G., Valenzuela, A., Ballestero, D., and Pino-Ot\u00edn, M. (2025). Mechanisms of Action and Resistance Prevention of Synergistic Thymol and Carvacrol Combinations with Antibiotics in Staphylococcus aureus and Acinetobacter baumannii. Nat. Prod. Bioprospect., 15.","DOI":"10.1007\/s13659-025-00518-7"},{"key":"ref_67","doi-asserted-by":"crossref","unstructured":"Farhadi, K., Rajabi, E., Varpaei, H.A., Iranzadasl, M., Khodaparast, S., and Salehi, M. (2024). Thymol and Carvacrol Against Klebsiella: Anti-Bacterial, Anti-Biofilm, and Synergistic Activities\u2014A Systematic Review. Front. Pharmacol., 15.","DOI":"10.3389\/fphar.2024.1487083"},{"key":"ref_68","doi-asserted-by":"crossref","unstructured":"McEwen, S.A., and Collignon, P.J. (2018). Antimicrobial Resistance: A One Health Perspective. Microbiol. Spectr., 6.","DOI":"10.1128\/microbiolspec.ARBA-0009-2017"},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"84","DOI":"10.1111\/jam.13755","article-title":"Combination of Essential Oil and Ciprofloxacin to Inhibit\/Eradicate Biofilms in Multidrug-Resistant Klebsiella pneumoniae","volume":"125","author":"Mohamed","year":"2018","journal-title":"J. Appl. Microbiol."},{"key":"ref_70","doi-asserted-by":"crossref","unstructured":"Touati, A., Mairi, A., Ibrahim, N.A., and Idres, T. (2025). Essential Oils for Biofilm Control: Mechanisms, Synergies, and Translational Challenges in the Era of Antimicrobial Resistance. Antibiotics, 14.","DOI":"10.3390\/antibiotics14050503"},{"key":"ref_71","doi-asserted-by":"crossref","unstructured":"Jayakumar, D., Mini, M., Kumar, P., Vaikkathillam, P., Mohan, A., and Khan, S. (2023). Synergistic Effect of Thymol-Ciprofloxacin Combination on Planktonic Cells and Biofilm of Pseudomonas aeruginosa. Curr. Microbiol., 81.","DOI":"10.1007\/s00284-023-03546-z"},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"56","DOI":"10.1016\/j.micpath.2017.01.012","article-title":"Use of Carvacrol, Thymol, and Eugenol for Biofilm Eradication and Resistance Modifying Susceptibility of Salmonella enterica Serovar Typhimurium Strains to Nalidixic Acid","volume":"104","author":"Miladi","year":"2017","journal-title":"Microb. Pathog."},{"key":"ref_73","doi-asserted-by":"crossref","unstructured":"Santos, M.I.S., Marques, C., Mota, J., Pedroso, L., and Lima, A. (2022). Applications of Essential Oils as Antibacterial Agents in Minimally Processed Fruits and Vegetables\u2014A Review. Microorganisms, 10.","DOI":"10.3390\/microorganisms10040760"},{"key":"ref_74","doi-asserted-by":"crossref","first-page":"273","DOI":"10.1016\/j.fm.2004.08.006","article-title":"Improvement in Shelf-Life and Safety of Perishable Foods by Plant Essential Oils and Smoke Antimicrobials","volume":"22","author":"Holley","year":"2005","journal-title":"Food Microbiol."},{"key":"ref_75","doi-asserted-by":"crossref","unstructured":"Kabir, M.N., Aras, S., Allison, A., Adhikari, J., Chowdhury, S., and Fouladkhah, A. (2019). Interactions of Carvacrol, Caprylic Acid, Habituation, and Mild Heat for Pressure-Based Inactivation of O157 and Non-O157 Serogroups of Shiga Toxin-Producing Escherichia coli in Acidic Environment. Microorganisms, 7.","DOI":"10.3390\/microorganisms7050145"},{"key":"ref_76","doi-asserted-by":"crossref","unstructured":"Zaharioudakis, K., Salmas, C.E., Andritsos, N.D., Kollia, E., Leontiou, A., Karabagias, V.K., Karydis-Messinis, A., Moschovas, D., Zafeiropoulos, N.E., and Avgeropoulos, A. (2024). Carvacrol, Citral, Eugenol and Cinnamaldehyde Casein-Based Edible Nanoemulsions as Novel Sustainable Active Coatings for Fresh Pork Tenderloin Meat Preservation. Front. Food Sci. Technol., 4.","DOI":"10.3389\/frfst.2024.1400224"},{"key":"ref_77","doi-asserted-by":"crossref","first-page":"103834","DOI":"10.1016\/j.jddst.2022.103834","article-title":"Nanotechnology as a Tool to Improve the Biological Activity of Carvacrol: A Review","volume":"76","author":"Dantas","year":"2022","journal-title":"J. Drug Deliv. Sci. Technol."},{"key":"ref_78","doi-asserted-by":"crossref","unstructured":"EFSA FEEDAP Panel (EFSA Panel on Additives and Products or Substances used in Animal Feed) (2016). Scientific opinion on the safety and efficacy of polyoxyethylene (20) sorbitan monooleate as a feed additive for all animal species. EFSA J., 14, 4443.","DOI":"10.2903\/j.efsa.2016.4443"},{"key":"ref_79","doi-asserted-by":"crossref","first-page":"285","DOI":"10.1080\/10408398.2011.559558","article-title":"Food-Grade Nanoemulsions: Formulation, Fabrication, Properties, Performance, Biological Fate, and Potential Toxicity","volume":"51","author":"McClements","year":"2011","journal-title":"Crit. Rev. Food Sci. Nutr."},{"key":"ref_80","doi-asserted-by":"crossref","first-page":"112935","DOI":"10.1016\/j.indcrop.2020.112935","article-title":"Essential Oil-Based Nano-Emulsions: Effect of Different Surfactants, Sonication and Plant Species on Physicochemical Characteristics","volume":"157","author":"Campolo","year":"2020","journal-title":"Ind. Crops Prod."},{"key":"ref_81","first-page":"e140923221094","article-title":"Antimicrobial Activity of Formulated Origanum and Thyme Essential Oil Nanoemulsions\u2014A Comparative Study","volume":"20","author":"Koirala","year":"2024","journal-title":"Curr. Nutr. Food Sci."},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"170","DOI":"10.1016\/j.ijpharm.2009.08.035","article-title":"Self-Nanoemulsifying Drug Delivery System (SNEDDS) for Oral Delivery of Zedoary Essential Oil: Formulation and Bioavailability Studies","volume":"383","author":"Zhao","year":"2010","journal-title":"Int. J. Pharm."},{"key":"ref_83","doi-asserted-by":"crossref","first-page":"155","DOI":"10.1016\/j.foodchem.2016.03.008","article-title":"Stability Evaluation of Lutein Nanodispersions Prepared via Solvent Displacement Method: The Effect of Emulsifiers with Different Stabilizing Mechanisms","volume":"205","author":"Tan","year":"2016","journal-title":"Food Chem."},{"key":"ref_84","doi-asserted-by":"crossref","first-page":"1516","DOI":"10.1016\/j.indcrop.2011.05.008","article-title":"Growing Conditions and Postharvest Management Can Affect the Essential Oil of Origanum vulgare L. ssp. hirtum (Link) Ietswaart","volume":"34","author":"Tibaldi","year":"2011","journal-title":"Ind. Crops Prod."},{"key":"ref_85","doi-asserted-by":"crossref","first-page":"118167","DOI":"10.1016\/j.indcrop.2024.118167","article-title":"Geographic Variability of Winter Savory Essential Oil","volume":"210","author":"Marin","year":"2024","journal-title":"Ind. Crops Prod."},{"key":"ref_86","unstructured":"The European Directorate for the Quality of Medicines & HealthCare (2020). European Pharmacopoeia, Council of Europe. [10th ed.]. Method 2.8.12\u2014Essential Oils in Herbal Drugs."},{"key":"ref_87","doi-asserted-by":"crossref","first-page":"463","DOI":"10.1016\/S0021-9673(01)80947-X","article-title":"A Generalization of the Retention Index System Including Linear Temperature Programmed Gas\u2013Liquid Partition Chromatography","volume":"11","author":"Kratz","year":"1963","journal-title":"J. Chromatogr."},{"key":"ref_88","unstructured":"Massada, Y. (1976). Analysis of Essential Oils by Gas Chromatography and Mass Spectrometry, Halsted Press: John Wiley & Sons."},{"key":"ref_89","unstructured":"Adams, R.P. (2007). Identification of Essential Oil Components by Gas Chromatography\/Mass Spectrometry, Allured Publishing Corporation. [4th ed.]."},{"key":"ref_90","doi-asserted-by":"crossref","first-page":"25","DOI":"10.1016\/S0023-6438(95)80008-5","article-title":"Use of a Free Radical Method to Evaluate Antioxidant Activity","volume":"28","author":"Cuvelier","year":"1995","journal-title":"LWT Food Sci. Technol."},{"key":"ref_91","doi-asserted-by":"crossref","first-page":"1231","DOI":"10.1016\/S0891-5849(98)00315-3","article-title":"Antioxidant Activity Applying an Improved ABTS Radical Cation Decolorization Assay","volume":"26","author":"Re","year":"1999","journal-title":"Free Radic. Biol. Med."},{"key":"ref_92","doi-asserted-by":"crossref","unstructured":"Pu\u0161k\u00e1rov\u00e1, A., Bu\u010dkov\u00e1, M., Krakov\u00e1, L., Pangallo, D., and Kozics, K. (2017). The Antibacterial and Antifungal Activity of Six Essential Oils and Their Cyto\/Genotoxicity to Human HEL 12469 Cells. Sci. Rep., 7.","DOI":"10.1038\/s41598-017-08673-9"},{"key":"ref_93","doi-asserted-by":"crossref","first-page":"115549","DOI":"10.1016\/j.lwt.2023.115549","article-title":"Washing Solutions Containing Nanoemulsified Essential Oils as a Potential Substitute of Commercial Washes for the Decontamination of Escherichia coli O157:H7 on Cherry Tomatoes","volume":"190","author":"Bodea","year":"2023","journal-title":"LWT"}],"container-title":["Molecules"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/1420-3049\/30\/16\/3329\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,9]],"date-time":"2025-10-09T18:27:05Z","timestamp":1760034425000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/1420-3049\/30\/16\/3329"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,8,9]]},"references-count":93,"journal-issue":{"issue":"16","published-online":{"date-parts":[[2025,8]]}},"alternative-id":["molecules30163329"],"URL":"https:\/\/doi.org\/10.3390\/molecules30163329","relation":{},"ISSN":["1420-3049"],"issn-type":[{"type":"electronic","value":"1420-3049"}],"subject":[],"published":{"date-parts":[[2025,8,9]]}}}