{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,7]],"date-time":"2026-09-07T22:31:24Z","timestamp":1788820284716,"version":"build-2803163510"},"reference-count":80,"publisher":"MDPI AG","issue":"6","license":[{"start":{"date-parts":[[2026,3,20]],"date-time":"2026-03-20T00:00:00Z","timestamp":1773964800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Polymers"],"abstract":"<jats:p>Kombucha fermentation is driven by a Symbiotic Culture of Bacteria and Yeast (SCOBY), a cellulose-rich biofilm that hosts a complex microbial consortium. While most kombucha studies focus on the liquid beverage, the SCOBY pellicle itself remains underexplored, particularly with respect to species-level microbial resolution and its intrinsic biological activities. In this study, a commercial kombucha SCOBY was characterized using full-length 16S rRNA gene and ITS amplicon sequencing based on Oxford Nanopore Technology, enabling species-level taxonomic resolution. In parallel, hydroalcoholic and aqueous extracts of dried SCOBY biomass were evaluated for in vitro antioxidant activity (DPPH and ABTS assays), antidiabetic-related enzyme inhibition (\u03b1-glucosidase and dipeptidyl peptidase-4, DPP4), and anti-aging-related enzyme inhibition (tyrosinase and elastase). The SCOBY bacterial community was strongly dominated by acetic acid bacteria, with Komagataeibacter saccharivorans and Acetobacter tropicalis accounting for more than 60% of total reads, reflecting a biofilm structure optimized for cellulose production and oxidative metabolism. The yeast community showed marked unevenness, with Brettanomyces bruxellensis representing over 80% of reads, consistent with its known role in ethanol production and stress tolerance within kombucha systems. In vitro assays revealed that hydroalcoholic SCOBY extracts consistently exhibited higher biological activity than aqueous extracts across all tested assays. However, both extracts showed substantially lower potency than purified reference compounds, indicating moderate but measurable bioactivity typical of complex fermented matrices. These findings support the potential valorization of SCOBY as a fermentation-derived biomaterial and functional ingredient while underscoring the need for further chemical characterization, mechanistic studies, and biological validation beyond enzyme-based assays.<\/jats:p>","DOI":"10.3390\/polym18060764","type":"journal-article","created":{"date-parts":[[2026,3,20]],"date-time":"2026-03-20T17:02:43Z","timestamp":1774026163000},"page":"764","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":1,"title":["Kombucha SCOBY as a Fermentation-Derived Biofilm Matrix: Species-Resolved Microbial Communities and Multidimensional In Vitro Bioactivities"],"prefix":"10.3390","volume":"18","author":[{"given":"Anita","family":"Hartono","sequence":"first","affiliation":[{"name":"Mambucha Scobylogy International, Jakarta 14240, Indonesia"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0001-1367-785X","authenticated-orcid":false,"given":"Kyra Singgih","family":"Palupi","sequence":"additional","affiliation":[{"name":"Biology Department, Faculty of Science and Technology, State Islamic University of Sunan Kalijaga (UIN Sunan Kalijaga), Yogyakarta 55281, Indonesia"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1327-1200","authenticated-orcid":false,"given":"Riza-Arief","family":"Putranto","sequence":"additional","affiliation":[{"name":"Department of Biotechnology, Faculty of Medical Sciences, Esa Unggul University, Jakarta 11510, Indonesia"},{"name":"Indonesian Oil Palm Research Institute, Medan 20158, Indonesia"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5505-3327","authenticated-orcid":false,"given":"Antonello","family":"Santini","sequence":"additional","affiliation":[{"name":"Department of Pharmacy, University of Napoli Federico II, Via Domenico Montesano, 49, 80131 Napoli, Italy"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2151-0854","authenticated-orcid":false,"given":"Fahrul","family":"Nurkolis","sequence":"additional","affiliation":[{"name":"Institute for Research and Community Service, State Islamic University of Sunan Kalijaga (UIN Sunan Kalijaga), Yogyakarta 55281, Indonesia"},{"name":"Master of Basic Medical Science, Faculty of Medicine, Universitas Airlangga, Surabaya 60131, Indonesia"},{"name":"Medical Research Center of Indonesia, Surabaya 60281, Indonesia"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2026,3,20]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","unstructured":"Anantachoke, N., Duangrat, R., Sutthiphatkul, T., Ochaikul, D., and Mangmool, S. (2023). Kombucha Beverages Produced from Fruits, Vegetables, and Plants: A Review on Their Pharmacological Activities and Health Benefits. Foods, 12.","DOI":"10.3390\/foods12091818"},{"key":"ref_2","doi-asserted-by":"crossref","unstructured":"Ntovolou, I., Farkatsi, D., and Ellinas, K. (2025). The Quest towards Superhydrophobic Cellulose and Bacterial Cellulose Membranes and Their Perspective Applications. Micro, 5.","DOI":"10.3390\/micro5030037"},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"85","DOI":"10.17113\/ftb.61.01.23.7811","article-title":"Role of Acetic Acid Bacteria in Food and Beverages","volume":"61","author":"Surek","year":"2023","journal-title":"Food Technol. Biotechnol."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"7911","DOI":"10.1038\/s41598-017-06918-1","article-title":"Reconstruction of a Genome-Scale Metabolic Network of Komagataeibacter nataicola RZS01 for Cellulose Production","volume":"7","author":"Zhang","year":"2017","journal-title":"Sci. Rep."},{"key":"ref_5","doi-asserted-by":"crossref","unstructured":"Andrade, D.K.A., Wang, B., Lima, E.M.F., Shebeko, S.K., Ermakov, A.M., Khramova, V.N., Ivanova, I.V., Rocha, R.S., Vaz-Velho, M., and Mutukumira, A.N. (2025). Kombucha: An Old Tradition into a New Concept of a Beneficial, Health-Promoting Beverage. Foods, 14.","DOI":"10.3390\/foods14091547"},{"key":"ref_6","doi-asserted-by":"crossref","unstructured":"Alkan, G., Konar Erol, N.M., Y\u0131km\u0131\u015f, S., Er, H., \u00d6\u011f\u00fct, S., and Yinan\u00e7, A. (2025). Kombucha\u2019s Functional Features and Fermentation Dynamics: A Bibliometric Assessment in Sustainable Food Production. Front. Sustain. Food Syst., 9.","DOI":"10.3389\/fsufs.2025.1593348"},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"63","DOI":"10.1016\/j.ijfoodmicro.2015.12.015","article-title":"Kombucha Tea Fermentation: Microbial and Biochemical Dynamics","volume":"220","author":"Chakravorty","year":"2016","journal-title":"Int. J. Food Microbiol."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"580","DOI":"10.1111\/1750-3841.14068","article-title":"Understanding Kombucha Tea Fermentation: A Review","volume":"83","author":"Beaufort","year":"2018","journal-title":"J. Food Sci."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"538","DOI":"10.1111\/1541-4337.12073","article-title":"A Review on Kombucha Tea-Microbiology, Composition, Fermentation, Beneficial Effects, Toxicity, and Tea Fungus","volume":"13","author":"Jayabalan","year":"2014","journal-title":"Compr. Rev. Food Sci. Food Saf."},{"key":"ref_10","doi-asserted-by":"crossref","unstructured":"Antolak, H., Piechota, D., and Kucharska, A. (2021). Kombucha Tea-A Double Power of Bioactive Compounds from Tea and Symbiotic Culture of Bacteria and Yeasts (SCOBY). Antioxidants, 10.","DOI":"10.3390\/antiox10101541"},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"100025","DOI":"10.1016\/j.focha.2022.100025","article-title":"Kombucha: Biochemical and Microbiological Impacts on the Chemical and Flavor Profile","volume":"1","author":"Bishop","year":"2022","journal-title":"Food Chem. Adv."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"455","DOI":"10.1111\/1750-3841.14992","article-title":"Microbial Composition of Kombucha Determined Using Amplicon Sequencing and Shotgun Metagenomics","volume":"85","author":"Mitchell","year":"2020","journal-title":"J. Food Sci."},{"key":"ref_13","doi-asserted-by":"crossref","unstructured":"Tran, T., Grandvalet, C., Winckler, P., Verdier, F., Martin, A., Alexandre, H., and Tourdot-Mar\u00e9chal, R. (2021). Shedding Light on the Formation and Structure of Kombucha Biofilm Using Two-Photon Fluorescence Microscopy. Front. Microbiol., 12.","DOI":"10.3389\/fmicb.2021.725379"},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"124","DOI":"10.1186\/s13568-015-0124-5","article-title":"Metabarcoding of the Kombucha Microbial Community Grown in Different Microenvironments","volume":"5","author":"Reva","year":"2015","journal-title":"AMB Express"},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"73","DOI":"10.3390\/applmicrobiol2010005","article-title":"Advances in Kombucha Tea Fermentation: A Review","volume":"2","author":"Nyhan","year":"2022","journal-title":"Appl. Microbiol."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"41","DOI":"10.1016\/j.procbio.2020.07.022","article-title":"Conducting High Acetic Acid and Temperature Acetification Processes by Acetobacter pasteurianus UMCC 2951","volume":"98","author":"Pothimon","year":"2020","journal-title":"Process Biochem."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"e00606","DOI":"10.1016\/j.btre.2021.e00606","article-title":"Functional and Genomic Characterization of Komagataeibacter uvaceti FXV3, a Multiple Stress Resistant Bacterium Producing Increased Levels of Cellulose","volume":"30","author":"Nascimento","year":"2021","journal-title":"Biotechnol. Rep."},{"key":"ref_18","doi-asserted-by":"crossref","unstructured":"Karni\u010dnik, B., Accetto, T., Fanedl, L., Jugovi\u0107, I., and Tr\u010dek, J. (2025). Isolation and Characterization of Komagataeibacter piraceti sp. nov. and Novacetimonas labruscae sp. nov.: Two Novel Microaerobic Cellulose-Producing Acetic Acid Bacteria from Vinegars. Microorganisms, 13.","DOI":"10.3390\/microorganisms13020456"},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"14768","DOI":"10.1038\/s41598-024-65841-4","article-title":"Short-Read Full-Length 16S RRNA Amplicon Sequencing for Characterisation of the Respiratory Bacteriome of Captive and Free-Ranging African Elephants (Loxodonta africana)","volume":"14","author":"Martin","year":"2024","journal-title":"Sci. Rep."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"219","DOI":"10.1186\/s40168-025-02206-7","article-title":"High-Resolution Profiling of Bacterial and Fungal Communities Using Pangenome-Informed Taxon-Specific Long-Read Amplicons","volume":"13","author":"Stalder","year":"2025","journal-title":"Microbiome"},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"bpaa026","DOI":"10.1093\/biomethods\/bpaa026","article-title":"Rapid and Real-Time Identification of Fungi up to Species Level with Long Amplicon Nanopore Sequencing from Clinical Samples","volume":"6","author":"Francino","year":"2021","journal-title":"Biol. Methods Protoc."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"9785","DOI":"10.1038\/s41598-023-37016-0","article-title":"Using Nanopore Sequencing to Identify Fungi from Clinical Samples with High Phylogenetic Resolution","volume":"13","author":"Ohta","year":"2023","journal-title":"Sci. Rep."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"248","DOI":"10.1007\/s00203-024-03985-7","article-title":"A Comparison between Full-Length 16S RRNA Oxford Nanopore Sequencing and Illumina V3-V4 16S RRNA Sequencing in Head and Neck Cancer Tissues","volume":"206","author":"Yeo","year":"2024","journal-title":"Arch. Microbiol."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"e0015722","DOI":"10.1128\/msystems.00157-22","article-title":"Microbial Diversity and Interaction Specificity in Kombucha Tea Fermentations","volume":"7","author":"Landis","year":"2022","journal-title":"mSystems"},{"key":"ref_25","doi-asserted-by":"crossref","unstructured":"Harrison, K., and Curtin, C. (2021). Microbial Composition of SCOBY Starter Cultures Used by Commercial Kombucha Brewers in North America. Microorganisms, 9.","DOI":"10.3390\/microorganisms9051060"},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"e76401","DOI":"10.7554\/eLife.76401","article-title":"A Systematic, Complexity-Reduction Approach to Dissect the Kombucha Tea Microbiome","volume":"11","author":"Huang","year":"2022","journal-title":"Elife"},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"e0060523","DOI":"10.1128\/aem.00605-23","article-title":"The Newest Oxford Nanopore R10.4.1 Full-Length 16S RRNA Sequencing Enables the Accurate Resolution of Species-Level Microbial Community Profiling","volume":"89","author":"Zhang","year":"2023","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"1088","DOI":"10.1007\/s13197-022-05476-3","article-title":"Metagenomic and Physicochemical Analysis of Kombucha Beverage Produced from Tea Waste","volume":"60","author":"Pradhan","year":"2023","journal-title":"J. Food Sci. Technol."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"fix048","DOI":"10.1093\/femsec\/fix048","article-title":"Unraveling Microbial Ecology of Industrial-Scale Kombucha Fermentations by Metabarcoding and Culture-Based Methods","volume":"93","author":"Coton","year":"2017","journal-title":"FEMS Microbiol. Ecol."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"502","DOI":"10.3746\/pnf.2023.28.4.502","article-title":"Multi-Strain Probiotics Enhance the Bioactivity of Cascara Kombucha during Microbial Composition-Controlled Fermentation","volume":"28","author":"Van","year":"2023","journal-title":"Prev. Nutr. Food Sci."},{"key":"ref_31","doi-asserted-by":"crossref","unstructured":"Dechakhamphu, A., Wongchum, N., Chumroenphat, T., Tanomtong, A., Pinlaor, S., and Siriamornpun, S. (2023). In Vitro and In Vivo Evaluation for Antioxidant and Anti-Diabetic Properties of Cyperus rotundus L. Kombucha. Foods, 12.","DOI":"10.20944\/preprints202310.1201.v1"},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"2893","DOI":"10.1007\/s13197-023-05808-x","article-title":"Native Bacterial Cellulose Films Based on Kombucha Pellicle as a Potential Active Food Packaging","volume":"60","year":"2023","journal-title":"J. Food Sci. Technol."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"347","DOI":"10.3746\/pnf.2023.28.3.347","article-title":"Antioxidant Properties of Kombucha Made with Tartary Buckwheat Tea and Burdock Tea","volume":"28","author":"Lee","year":"2023","journal-title":"Prev. Nutr. Food Sci."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"69","DOI":"10.1007\/s13668-025-00658-9","article-title":"Kombucha Tea: A Functional Beverage and All Its Aspects","volume":"14","author":"Onsun","year":"2025","journal-title":"Curr. Nutr. Rep."},{"key":"ref_35","doi-asserted-by":"crossref","unstructured":"Wang, B., Rutherfurd-Markwick, K., Zhang, X.-X., and Mutukumira, A.N. (2022). Kombucha: Production and Microbiological Research. Foods, 11.","DOI":"10.3390\/foods11213456"},{"key":"ref_36","doi-asserted-by":"crossref","unstructured":"Cubas, A.L.V., Provin, A.P., Dutra, A.R.A., Mouro, C., and Gouveia, I.C. (2023). Advances in the Production of Biomaterials through Kombucha Using Food Waste: Concepts, Challenges, and Potential. Polymers, 15.","DOI":"10.3390\/polym15071701"},{"key":"ref_37","doi-asserted-by":"crossref","unstructured":"Vianna, N.M., Albagli, G., Pereira, A.S., and Amaral, P.F.F. (2025). From Tea Fermentation to New Technologies: Multisectoral Applications of Kombucha SCOBY Through the Lens of Methodi ordinatio. Fermentation, 11.","DOI":"10.3390\/fermentation11100589"},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"100421","DOI":"10.1016\/j.focha.2023.100421","article-title":"Kombucha Fermentation: Recent Trends in Process Dynamics, Functional Bioactivities, Toxicity Management, and Potential Applications","volume":"3","author":"Sanwal","year":"2023","journal-title":"Food Chem. Adv."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"105555","DOI":"10.1016\/j.jff.2023.105555","article-title":"In Silico and in Vitro Studies Reveal a Synergistic Potential Source of Novel Anti-Ageing from Two Indonesian Green Algae","volume":"104","author":"Nurkolis","year":"2023","journal-title":"J. Funct. Foods"},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"1251","DOI":"10.1016\/j.crfs.2022.08.005","article-title":"Modulation of Gut Microbiota and Markers of Metabolic Syndrome in Mice on Cholesterol and Fat Enriched Diet by Butterfly Pea Flower Kombucha","volume":"5","author":"Permatasari","year":"2022","journal-title":"Curr. Res. Food Sci."},{"key":"ref_41","doi-asserted-by":"crossref","unstructured":"He, C., Liu, X., Jiang, Z., Geng, S., Ma, H., and Liu, B. (2019). Interaction Mechanism of Flavonoids and \u03b1-Glucosidase: Experimental and Molecular Modelling Studies. Foods, 8.","DOI":"10.3390\/foods8090355"},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"1216","DOI":"10.1080\/14756366.2017.1368503","article-title":"\u03b1-Glucosidase Inhibition by Flavonoids: An In Vitro and In Silico Structure-Activity Relationship Study","volume":"32","author":"Freitas","year":"2017","journal-title":"J. Enzyme Inhib. Med. Chem."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"456","DOI":"10.1016\/j.cca.2011.10.031","article-title":"Method Comparison of Dipeptidyl Peptidase IV Activity Assays and Their Application in Biological Samples Containing Reversible Inhibitors","volume":"413","author":"Matheeussen","year":"2012","journal-title":"Clin. Chim. Acta"},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"e1305","DOI":"10.7717\/peerj.1305","article-title":"Antityrosinase Activity of Euphorbia characias Extracts","volume":"3","author":"Pintus","year":"2015","journal-title":"PeerJ"},{"key":"ref_45","doi-asserted-by":"crossref","unstructured":"Thring, T.S.A., Hili, P., and Naughton, D.P. (2009). Anti-Collagenase, Anti-Elastase and Anti-Oxidant Activities of Extracts from 21 Plants. BMC Complement. Altern. Med., 9.","DOI":"10.1186\/1472-6882-9-27"},{"key":"ref_46","doi-asserted-by":"crossref","unstructured":"Younis, M.M., Ayoub, I.M., Mostafa, N.M., El Hassab, M.A., Eldehna, W.M., Al-Rashood, S.T., and Eldahshan, O.A. (2022). GC\/MS Profiling, Anti-Collagenase, Anti-Elastase, Anti-Tyrosinase and Anti-Hyaluronidase Activities of a Stenocarpus sinuatus Leaves Extract. Plants, 11.","DOI":"10.3390\/plants11070918"},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"18","DOI":"10.1186\/s43008-023-00125-6","article-title":"Targeted Sequencing Analysis Pipeline for Species Identification of Human Pathogenic Fungi Using Long-Read Nanopore Sequencing","volume":"14","author":"Langsiri","year":"2023","journal-title":"IMA Fungus"},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"131346","DOI":"10.1016\/j.foodchem.2021.131346","article-title":"Bacterial Cellulose Films Production by Kombucha Symbiotic Community Cultured on Different Herbal Infusions","volume":"372","author":"Peltzer","year":"2022","journal-title":"Food Chem."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"e00731","DOI":"10.1002\/mbo3.731","article-title":"Comparative Genomics of the Komagataeibacter Strains-Efficient Bionanocellulose Producers","volume":"8","author":"Kubiak","year":"2019","journal-title":"Microbiologyopen"},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"4015","DOI":"10.1039\/D1FO00630D","article-title":"Transient in Situ Measurement of Kombucha Biofilm Growth and Mechanical Properties","volume":"12","author":"Bertsch","year":"2021","journal-title":"Food Funct."},{"key":"ref_51","doi-asserted-by":"crossref","unstructured":"Devanthi, P.V.P., Kho, K., Nurdiansyah, R., Briot, A., Taherzadeh, M.J., and Aslanzadeh, S. (2021). Do Kombucha Symbiotic Cultures of Bacteria and Yeast Affect Bacterial Cellulose Yield in Molasses?. J. Fungi, 7.","DOI":"10.3390\/jof7090705"},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"foaf005","DOI":"10.1093\/femsyr\/foaf005","article-title":"Comprehensive Survey of Kombucha Microbial Communities of Diverse Origins and Fermentation Practices","volume":"25","author":"Friedrich","year":"2025","journal-title":"FEMS Yeast Res."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"103379","DOI":"10.1016\/j.fm.2019.103379","article-title":"Brettanomyces bruxellensis Phenotypic Diversity, Tolerance to Wine Stress and Wine Spoilage Ability","volume":"87","author":"Cibrario","year":"2020","journal-title":"Food Microbiol."},{"key":"ref_54","doi-asserted-by":"crossref","unstructured":"Colomer, M.S., Chailyan, A., Fennessy, R.T., Olsson, K.F., Johnsen, L., Solodovnikova, N., and Forster, J. (2020). Assessing Population Diversity of Brettanomyces Yeast Species and Identification of Strains for Brewing Applications. Front. Microbiol., 11.","DOI":"10.3389\/fmicb.2020.00637"},{"key":"ref_55","doi-asserted-by":"crossref","unstructured":"Liao, T., Li, X.-R., Fan, L., Zhang, B., Zheng, W.-M., Hua, J.-J., Li, L., Mahror, N., and Cheng, L.-H. (2024). Nature of Back Slopping Kombucha Fermentation Process: Insights from the Microbial Succession, Metabolites Composition Changes and Their Correlations. Front. Microbiol., 15.","DOI":"10.3389\/fmicb.2024.1433127"},{"key":"ref_56","doi-asserted-by":"crossref","unstructured":"Andreson, M., Kazantseva, J., Malv, E., Kuldj\u00e4rv, R., Priidik, R., and K\u00fctt, M.-L. (2023). Evaluation of Microbial Dynamics of Kombucha Consortia upon Continuous Backslopping in Coffee and Orange Juice. Foods, 12.","DOI":"10.3390\/foods12193545"},{"key":"ref_57","first-page":"110","article-title":"Brettanomyces Bruxellensis, Essential Contributor in Spontaneous Beer Fermentations Providing Novel Opportunities for the Brewing Industry","volume":"68","author":"Crauwels","year":"2015","journal-title":"BrSc"},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"244","DOI":"10.1186\/s13068-016-0652-5","article-title":"Presence Does Not Imply Activity: DNA and RNA Patterns Differ in Response to Salt Perturbation in Anaerobic Digestion","volume":"9","author":"Regueiro","year":"2016","journal-title":"Biotechnol. Biofuels"},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"507","DOI":"10.1038\/nature24460","article-title":"Quantitative Microbiome Profiling Links Gut Community Variation to Microbial Load","volume":"551","author":"Vandeputte","year":"2017","journal-title":"Nature"},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"2590","DOI":"10.1038\/s41467-020-16224-6","article-title":"A Quantitative Sequencing Framework for Absolute Abundance Measurements of Mucosal and Lumenal Microbial Communities","volume":"11","author":"Barlow","year":"2020","journal-title":"Nat. Commun."},{"key":"ref_61","doi-asserted-by":"crossref","unstructured":"Wu, X., Zhang, Y., Zhang, B., Tian, H., Liang, Y., Dang, H., and Zhao, Y. (2023). Dynamic Changes in Microbial Communities, Physicochemical Properties, and Flavor of Kombucha Made from Fu-Brick Tea. Foods, 12.","DOI":"10.3390\/foods12234242"},{"key":"ref_62","doi-asserted-by":"crossref","unstructured":"Salter, S.J., Cox, M.J., Turek, E.M., Calus, S.T., Cookson, W.O., Moffatt, M.F., Turner, P., Parkhill, J., Loman, N.J., and Walker, A.W. (2014). Reagent and Laboratory Contamination Can Critically Impact Sequence-Based Microbiome Analyses. BMC Biol., 12.","DOI":"10.1186\/s12915-014-0087-z"},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"e00290-19","DOI":"10.1128\/mSystems.00290-19","article-title":"Controlling for Contaminants in Low-Biomass 16S RRNA Gene Sequencing Experiments","volume":"4","author":"Karstens","year":"2019","journal-title":"mSystems"},{"key":"ref_64","doi-asserted-by":"crossref","unstructured":"Jakubczyk, K., Ka\u0142du\u0144ska, J., Kochman, J., and Janda, K. (2020). Chemical Profile and Antioxidant Activity of the Kombucha Beverage Derived from White, Green, Black and Red Tea. Antioxidants, 9.","DOI":"10.3390\/antiox9050447"},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"108782","DOI":"10.1016\/j.foodres.2019.108782","article-title":"de Kombuchas from Green and Black Teas Have Different Phenolic Profile, Which Impacts Their Antioxidant Capacities, Antibacterial and Antiproliferative Activities","volume":"128","author":"Cardoso","year":"2020","journal-title":"Food Res. Int."},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"503","DOI":"10.1111\/1750-3841.16029","article-title":"Kombucha: A Review of Substrates, Regulations, Composition, and Biological Properties","volume":"87","author":"Ruiz","year":"2022","journal-title":"J. Food Sci."},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"18792","DOI":"10.1038\/s41598-021-98191-6","article-title":"Effect of Fermentation Time on the Content of Bioactive Compounds with Cosmetic and Dermatological Properties in Kombucha Yerba Mate Extracts","volume":"11","author":"Ziemlewska","year":"2021","journal-title":"Sci. Rep."},{"key":"ref_68","doi-asserted-by":"crossref","unstructured":"Arulmozhiraja, S., Matsuo, N., Ishitsubo, E., Okazaki, S., Shimano, H., and Tokiwa, H. (2016). Comparative Binding Analysis of Dipeptidyl Peptidase IV (DPP-4) with Antidiabetic Drugs\u2014An Ab Initio Fragment Molecular Orbital Study. PLoS ONE, 11.","DOI":"10.1371\/journal.pone.0166275"},{"key":"ref_69","doi-asserted-by":"crossref","unstructured":"Shaikh, S., Lee, E.-J., Ahmad, K., Ahmad, S.-S., Lim, J.-H., and Choi, I. (2021). A Comprehensive Review and Perspective on Natural Sources as Dipeptidyl Peptidase-4 Inhibitors for Management of Diabetes. Pharmaceuticals, 14.","DOI":"10.3390\/ph14060591"},{"key":"ref_70","doi-asserted-by":"crossref","first-page":"7880387","DOI":"10.1155\/2022\/7880387","article-title":"Enzymes Inhibition and Antioxidant Potential of Medicinal Plants Growing in Oman","volume":"2022","author":"Rehman","year":"2022","journal-title":"Biomed Res. Int."},{"key":"ref_71","doi-asserted-by":"crossref","first-page":"100288","DOI":"10.1016\/j.focha.2023.100288","article-title":"Kombucha Benefits, Risks and Regulatory Frameworks: A Review","volume":"2","author":"Ogeda","year":"2023","journal-title":"Food Chem. Adv."},{"key":"ref_72","doi-asserted-by":"crossref","unstructured":"Ojo, A.O., and de Smidt, O. (2023). Microbial Composition, Bioactive Compounds, Potential Benefits and Risks Associated with Kombucha: A Concise Review. Fermentation, 9.","DOI":"10.3390\/fermentation9050472"},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"66","DOI":"10.1016\/j.annepidem.2018.11.001","article-title":"Kombucha: A Systematic Review of the Empirical Evidence of Human Health Benefit","volume":"30","author":"Kapp","year":"2019","journal-title":"Ann. Epidemiol."},{"key":"ref_74","doi-asserted-by":"crossref","unstructured":"Puspitasari, Y.E., Tuenter, E., Breynaert, A., Foubert, K., Herawati, H., Hariati, A.M., Aulanni\u2019am, A., De Bruyne, T., and Hermans, N. (2024). \u03b1-Glucosidase Inhibitory Activity of Tea and Kombucha from Rhizophora mucronata Leaves. Beverages, 10.","DOI":"10.3390\/beverages10010022"},{"key":"ref_75","doi-asserted-by":"crossref","unstructured":"Mechqoq, H., Hourfane, S., El Yaagoubi, M., El Hamdaoui, A., da Silva Almeida, J.R.G., Rocha, J.M., and El Aouad, N. (2022). Molecular Docking, Tyrosinase, Collagenase, and Elastase Inhibition Activities of Argan by-Products. Cosmetics, 9.","DOI":"10.3390\/cosmetics9010024"},{"key":"ref_76","doi-asserted-by":"crossref","first-page":"3811","DOI":"10.3390\/ijms10093811","article-title":"Variations in IC(50) Values with Purity of Mushroom Tyrosinase","volume":"10","author":"Neeley","year":"2009","journal-title":"Int. J. Mol. Sci."},{"key":"ref_77","first-page":"1","article-title":"The Effect of Polyphenolic Extracts from 10 New Zealand Kiwifruit Cultivars in Inhibiting Tyrosinase Enzyme Activity In Vitro","volume":"19","author":"Carroll","year":"2024","journal-title":"Nat. Prod. Commun."},{"key":"ref_78","doi-asserted-by":"crossref","unstructured":"Di Lorenzo, R., Maisto, M., Ricci, L., Piccolo, V., Marzocchi, A., Greco, G., Tenore, G.C., and Laneri, S. (2024). Annurca Apple Oleolite as Functional Ingredient for the Formulation of Cosmetics with Skin-Antiaging Activity. Int. J. Mol. Sci., 25.","DOI":"10.3390\/ijms25031677"},{"key":"ref_79","first-page":"103","article-title":"The phenolic content and antioxidant activity of the aqueous and hydroalcoholic extracts of hops and their pellets","volume":"119","author":"Kowalczyk","year":"2013","journal-title":"J. Inst. Brew."},{"key":"ref_80","doi-asserted-by":"crossref","unstructured":"Roda, G., Marinello, C., Grassi, A., Picozzi, C., Aldini, G., Carini, M., and Regazzoni, L. (2019). Ripe and Raw Pu-Erh Tea: LC-MS Profiling, Antioxidant Capacity and Enzyme Inhibition Activities of Aqueous and Hydro-Alcoholic Extracts. Molecules, 24.","DOI":"10.3390\/molecules24030473"}],"container-title":["Polymers"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2073-4360\/18\/6\/764\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,3,24]],"date-time":"2026-03-24T05:51:59Z","timestamp":1774331519000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2073-4360\/18\/6\/764"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,3,20]]},"references-count":80,"journal-issue":{"issue":"6","published-online":{"date-parts":[[2026,3]]}},"alternative-id":["polym18060764"],"URL":"https:\/\/doi.org\/10.3390\/polym18060764","relation":{},"ISSN":["2073-4360"],"issn-type":[{"value":"2073-4360","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,3,20]]}}}