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Smith with reported antimicrobial properties, alone or combined with Dextranase (DEX) and DNase I (DNase), against polymicrobial biofilms of fluconazole-susceptible and fluconazole-resistant Candida albicans and Streptococcus mutans. Mixed-species biofilms were formed with C. albicans ATCC 90028 (CaS), C. albicans ATCC 96901 (CaR), and S. mutans UA159 and assessed in inhibition and disruption assays. Treatments included ZER, DEX, and DNase, individually or in combination. Efficacy was analyzed by total biofilm mass, metabolic activity (CFU\/mm2), and live\/dead cell assessment using CLSM. Data were analyzed by one- and two-way ANOVA followed by Tukey\u2019s posthoc test (\u03b1 = 0.05). In the biofilm inhibition assay, ZER and its combinations, particularly ZER + DEX, demonstrated enhanced activity against both C. albicans and S. mutans strains. For CaS + Sm biofilms, ZER, ZER + DEX, and DNase + DEX achieved similar and the greatest reduction in CaS viability (&amp;gt;2.7 log10), whereas, for S. mutans, DNase + DEX, ZER, and ZER + DEX were the most effective (&amp;gt;2.1 log10). ZER + DEX also significantly reduced total microbial load in CaR + Sm biofilm. Notably, ZER + Dextranase was the only treatment that significantly reduced dry weight in susceptible biofilms and produced the highest biomass reduction in both susceptible and resistant biofilms as measured by crystal violet staining. In the biofilm disruption assay, ZER + DNase caused the greatest reduction in CaS viability (2.08 log10), whereas ZER and ZER + DNase were most effective against S. mutans (&amp;gt;2.5 log10). For CaR + Sm biofilms, ZER + DEX and ZER + DNase were among the most effective treatments. The confocal microscopy results corroborated these findings, showing that ZER and all treatments combined with ZER significantly increased the proportion of dead cells under both tested conditions. Within the limitations of this in vitro study, ZER alone or in combination with matrix-degrading enzymes (DEX and DNase), showed enhanced activity against both inhibition and disruption of mixed-species biofilms of C. albicans and S. mutans, including fluconazole-resistant strains, suggesting potential for further investigation as an adjunctive antimicrobial approach.<\/jats:p>","DOI":"10.1021\/acsomega.6c03152","type":"journal-article","created":{"date-parts":[[2026,8,19]],"date-time":"2026-08-19T18:02:02Z","timestamp":1787162522000},"page":"51080-51093","source":"Crossref","is-referenced-by-count":0,"title":["Zerumbone Combined\nwith Hydrolytic Enzymes: A Promising\nStrategy to Control Mixed Biofilms of\n                    <i>Streptococcus mutans<\/i>\n                    and Fluconazole-Resistant\n                    <i>Candida albicans<\/i>"],"prefix":"10.1021","volume":"11","author":[{"given":"Analu\u0301 Barros","family":"de Oliveira","sequence":"first","affiliation":[{"name":"Sa\u0303o Paulo State University (UNESP) , , , ,","place":["Araraquara, Sa\u0303o Paulo, Brazil, 14801-903"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2138-1223","authenticated-orcid":true,"given":"Beatriz H. D.","family":"Panariello","sequence":"additional","affiliation":[{"name":"LECOM School of Dental Medicine , 4800 Lakewood Ranch Blvd. , , ,","place":["Bradenton, Florida, United States, 34211"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Paula Aboud","family":"Barbugli","sequence":"additional","affiliation":[{"name":"Sa\u0303o Paulo State University (UNESP) , , , ,","place":["Araraquara, Sa\u0303o Paulo, Brazil, 14801-903"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9933-2819","authenticated-orcid":true,"given":"Tu\u0301lio Morandin","family":"Ferrisse","sequence":"additional","affiliation":[{"name":"Sa\u0303o Paulo State University (UNESP) , , , ,","place":["Araraquara, Sa\u0303o Paulo, Brazil, 14801-903"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9231-1994","authenticated-orcid":true,"given":"Ana Claudia","family":"Pavarina","sequence":"additional","affiliation":[{"name":"Sa\u0303o Paulo State University (UNESP) , , , ,","place":["Araraquara, Sa\u0303o Paulo, Brazil, 14801-903"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"316","published-online":{"date-parts":[[2026,8,19]]},"reference":[{"issue":"9","key":"2026090307164343300_cit1","doi-asserted-by":"publisher","first-page":"1797","DOI":"10.3390\/microorganisms12091797","article-title":"Oral Microbiome: A\nReview of Its Impact on Oral and Systemic Health","volume":"12","author":"Rajasekaran","year":"2024","journal-title":"Microorganisms"},{"issue":"3","key":"2026090307164343300_cit2","doi-asserted-by":"publisher","first-page":"335","DOI":"10.3390\/pathogens11030335","article-title":"Oral Cavity\nand Candida albicans: Colonisation to the Development\nof Infection","volume":"11","author":"Patel","year":"2022","journal-title":"Pathogens"},{"issue":"4","key":"2026090307164343300_cit3","doi-asserted-by":"publisher","DOI":"10.1128\/spectrum.00708-23","article-title":"Six-Year Retrospective\nAnalysis of Epidemiology, Risk\nFactors, and Antifungal Susceptibilities of Candidiasis from a Tertiary\nCare Hospital in South China","volume":"11","author":"Bilal","year":"2023","journal-title":"Microbiol. Spectr."},{"key":"2026090307164343300_cit4","first-page":"15","article-title":"Oral Candidiasis: A Disease of Opportunity","volume-title":"J. Fungi (Basel)","author":"Vila","year":"2020"},{"key":"2026090307164343300_cit5","doi-asserted-by":"publisher","first-page":"e280","DOI":"10.1016\/S1473-3099(24)00749-7","article-title":"Global guideline for\nthe diagnosis and management of candidiasis: an initiative of the\nECMM in cooperation with ISHAM and ASM","volume":"25","author":"Cornely","year":"2025","journal-title":"Lancet"},{"key":"2026090307164343300_cit6","doi-asserted-by":"publisher","first-page":"237","DOI":"10.2147\/IDR.S118892","article-title":"Fluconazole resistance\nin Candida species: a current\nperspective","volume":"10","author":"Berkow","year":"2017","journal-title":"Infect Drug Resist."},{"issue":"1","key":"2026090307164343300_cit7","doi-asserted-by":"publisher","DOI":"10.1128\/microbiolspec.gpp3-0051-2018","article-title":"The Biology of Streptococcus mutans","volume":"7","author":"Lemos","year":"2019","journal-title":"Microbiol\nSpectr."},{"key":"2026090307164343300_cit8","doi-asserted-by":"publisher","first-page":"1645","DOI":"10.12688\/f1000research.16275.2","article-title":"Relationship between Candida albicans and Streptococcus mutans in early childhood caries,\nevaluated by quantitative PCR","volume":"7","author":"Bachtiar","year":"2018","journal-title":"F1000Res."},{"issue":"5","key":"2026090307164343300_cit9","doi-asserted-by":"publisher","first-page":"1968","DOI":"10.1128\/IAI.00087-14","article-title":"Symbiotic\nrelationship between Streptococcus mutans and Candida albicans synergizes\nvirulence of plaque biofilms in vivo","volume":"82","author":"Falsetta","year":"2014","journal-title":"Infect.\nImmun."},{"issue":"1","key":"2026090307164343300_cit10","doi-asserted-by":"publisher","DOI":"10.1080\/20002297.2019.1581520","article-title":"Dual-species biofilms of Streptococcus mutans and Candida albicans exhibit more biomass and are mutually beneficial compared with single-species\nbiofilms","volume":"11","author":"Lobo","year":"2019","journal-title":"J. Oral Microbiol."},{"key":"2026090307164343300_cit11","first-page":"3","article-title":"Biofilms\nas Promoters of Bacterial Antibiotic Resistance and Tolerance","volume-title":"Antibiotics (Basel)","author":"Urue\u0301n","year":"2020"},{"key":"2026090307164343300_cit12","doi-asserted-by":"publisher","first-page":"124","DOI":"10.1016\/j.pdpdt.2019.05.038","article-title":"DNase increases\nthe efficacy of antimicrobial photodynamic therapy on Candida albicans\nbiofilms","volume":"27","author":"Panariello","year":"2019","journal-title":"Photodiagnosis Photodyn Ther"},{"issue":"4","key":"2026090307164343300_cit13","doi-asserted-by":"publisher","first-page":"983","DOI":"10.3390\/jcm9040983","article-title":"Combined DNase\nand Proteinase Treatment\nInterferes with Composition and Structural Integrity of Multispecies\nOral Biofilms","volume":"9","author":"Karygianni","year":"2020","journal-title":"J. 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Biochem."},{"issue":"10","key":"2026090307164343300_cit22","doi-asserted-by":"publisher","first-page":"2113","DOI":"10.1111\/pbi.13643","article-title":"Affordable oral health\ncare: dental biofilm disruption using chloroplast\nmade enzymes with chewing gum delivery","volume":"19","author":"Singh","year":"2021","journal-title":"Plant\nBiotechnol J."},{"issue":"2","key":"2026090307164343300_cit23","doi-asserted-by":"publisher","first-page":"223","DOI":"10.3390\/microorganisms10020223","article-title":"Modulation of Streptococcus mutans Adherence to Hydroxyapatite by Engineered\nSalivary Peptides","volume":"10","author":"Marin","year":"2022","journal-title":"Microorganisms"},{"issue":"1","key":"2026090307164343300_cit24","doi-asserted-by":"publisher","DOI":"10.1080\/20002297.2018.1476644","article-title":"Fluconazole impacts\nthe extracellular matrix of fluconazole-susceptible and -resistant Candida albicans and Candida glabrata biofilms","volume":"10","author":"Panariello","year":"2018","journal-title":"J. Oral Microbiol."},{"issue":"8","key":"2026090307164343300_cit25","doi-asserted-by":"publisher","first-page":"891","DOI":"10.1111\/j.1600-0463.2007.apm_630.x","article-title":"Quantification of biofilm\nin microtiter plates: overview of testing conditions and practical\nrecommendations for assessment of biofilm production by staphylococci","volume":"115","author":"Stepanovic\u0301","year":"2007","journal-title":"APMIS."},{"key":"2026090307164343300_cit26","doi-asserted-by":"publisher","DOI":"10.3389\/fmicb.2023.1274201","article-title":"DNase improves the\nefficacy of antimicrobial photodynamic therapy in the treatment of\ncandidiasis induced with Candida albicans","volume":"14","author":"Jorda\u0303o","year":"2023","journal-title":"Front Microbiol."},{"issue":"2","key":"2026090307164343300_cit27","doi-asserted-by":"publisher","first-page":"72","DOI":"10.1159\/000535980","article-title":"The Effect of Enzymatic Treatment\nwith Mutanase, Beta-Glucanase, and DNase on a Saliva-Derived Biofilm\nModel","volume":"58","author":"Dukanovic Rikvold","year":"2024","journal-title":"Caries Res."},{"issue":"1","key":"2026090307164343300_cit28","doi-asserted-by":"publisher","first-page":"5","DOI":"10.3390\/plants12010005","article-title":"Healthy\nZerumbone: From Natural Sources to Strategies to Improve Its Bioavailability\nand Oral Administration","volume":"12","author":"Iba\u0301n\u0303ez","year":"2023","journal-title":"Plants (Basel)"},{"issue":"4","key":"2026090307164343300_cit29","first-page":"163","article-title":"Characteristics\nof biofilm formation by Candida albicans","volume":"18","author":"Ramage","year":"2001","journal-title":"Rev.\nIberoam Micol."},{"issue":"1","key":"2026090307164343300_cit30","first-page":"162","article-title":"A novel assay for quantification\nof Candida albicans biofilm formation on plastic\nsurfaces","volume":"58","author":"Alem","year":"2006","journal-title":"J. Antimicrob. Chemother."},{"issue":"17","key":"2026090307164343300_cit31","first-page":"1888","article-title":"In vitro antifungal\nactivity of the sesquiterpene lactone anastatin against Candida albicans biofilms","volume":"77","author":"Marcos-Arias","year":"2011","journal-title":"Planta Medica"},{"issue":"1","key":"2026090307164343300_cit32","doi-asserted-by":"publisher","first-page":"32","DOI":"10.1111\/prd.12361","article-title":"The dental\nplaque biofilm matrix","volume":"86","author":"Jakubovics","year":"2021","journal-title":"Periodontol 2000"},{"key":"2026090307164343300_cit33","doi-asserted-by":"publisher","DOI":"10.1016\/j.bioflm.2024.100202","article-title":"Effect of mutanase and Dextranase on biofilms of cariogenic\nbacteria: A systematic review of in vitro studies","volume":"7","author":"Del Rey","year":"2024","journal-title":"Biofilm"},{"issue":"18","key":"2026090307164343300_cit34","doi-asserted-by":"publisher","first-page":"6357","DOI":"10.1128\/AEM.05203-11","article-title":"Role of glucosyltransferase\nB in\ninteractions of Candida albicans with Streptococcus mutans and with experimental pellicle on hydroxyapatite surfaces","volume":"77","author":"Gregoire","year":"2011","journal-title":"Appl. Environ. Microbiol."},{"issue":"1","key":"2026090307164343300_cit35","doi-asserted-by":"publisher","first-page":"299","DOI":"10.1007\/s10103-025-04552-2","article-title":"Efficiency\nof the extracellular polymeric\nmatrix disruptor Zerumbone in combination with Photodithazine in the\nphotodynamic inactivation of monospecies biofilms","volume":"40","author":"Abreu-Pereira","year":"2025","journal-title":"Lasers Med. Sci."},{"issue":"12","key":"2026090307164343300_cit36","doi-asserted-by":"publisher","DOI":"10.1371\/journal.ppat.1007342","article-title":"Candida\u2013streptococcal interactions in biofilm-associated oral diseases","volume":"14","author":"Koo","year":"2018","journal-title":"PLoS Pathogens"},{"issue":"19","key":"2026090307164343300_cit37","doi-asserted-by":"publisher","first-page":"6727","DOI":"10.3390\/ma15196727","article-title":"Quantitative Analyses of Biofilm by Using Crystal Violet\nStaining and Optical Reflection","volume":"15","author":"Kamimura","year":"2022","journal-title":"Materials (Basel)"},{"issue":"9","key":"2026090307164343300_cit38","doi-asserted-by":"publisher","first-page":"563","DOI":"10.1038\/nrmicro.2016.94","article-title":"Biofilms:\nan emergent form of bacterial\nlife","volume":"14","author":"Flemming","year":"2016","journal-title":"Nat. Rev. Microbiol."},{"issue":"3","key":"2026090307164343300_cit39","doi-asserted-by":"publisher","first-page":"212","DOI":"10.1111\/2049-632X.12118","article-title":"Biophysics\nof biofilm infection","volume":"70","author":"Stewart","year":"2014","journal-title":"Pathog Dis."},{"issue":"5","key":"2026090307164343300_cit40","doi-asserted-by":"publisher","first-page":"887","DOI":"10.1039\/C6BM00832A","article-title":"Modulation\nof the mechanical properties\nof bacterial biofilms in response to environmental challenges","volume":"5","author":"Tallawi","year":"2017","journal-title":"Biomater Sci."},{"issue":"3","key":"2026090307164343300_cit41","doi-asserted-by":"publisher","first-page":"205","DOI":"10.1177\/0022034509359403","article-title":"Biofilm\ndispersal: mechanisms, clinical implications, and potential therapeutic\nuses","volume":"89","author":"Kaplan","year":"2010","journal-title":"J. Dent Res."},{"key":"2026090307164343300_cit42","doi-asserted-by":"publisher","DOI":"10.1016\/j.mib.2022.102237","article-title":"Antifungal drug-resistance\nmechanisms in Candida biofilms","volume":"71","author":"Kaur","year":"2023","journal-title":"Curr. Opin Microbiol."},{"key":"2026090307164343300_cit43","doi-asserted-by":"publisher","first-page":"2173","DOI":"10.3389\/fmicb.2016.02173","article-title":"Azole antifungal resistance in Candida\nalbicans and emerging non-albicans Candida species","volume":"7","author":"Whaley","year":"2017","journal-title":"Front Microbiol."},{"issue":"6","key":"2026090307164343300_cit44","doi-asserted-by":"publisher","first-page":"1773","DOI":"10.1128\/AAC.46.6.1773-1780.2002","article-title":"Antifungal susceptibility of Candida biofilms: unique\nefficacy of amphotericin B lipid formulations and echinocandins","volume":"46","author":"Kuhn","year":"2002","journal-title":"Antimicrob. Agents Chemother."},{"key":"2026090307164343300_cit45","doi-asserted-by":"publisher","DOI":"10.3389\/fmicb.2021.737458","article-title":"DexA70, the\nTruncated Form of a Self-Produced Dextranase, Effectively Disrupts Streptococcus mutans Biofilm","volume":"12","author":"Liu","year":"2021","journal-title":"Front\nMicrobiol"},{"key":"2026090307164343300_cit46","doi-asserted-by":"publisher","DOI":"10.1016\/j.micpath.2021.104838","article-title":"Effects of extracellular\nDNA on dual-species biofilm formed by Streptococcus mutans and Candida albicans","volume":"154","author":"Guo","year":"2021","journal-title":"Microb.\nPathogen."},{"issue":"11","key":"2026090307164343300_cit47","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pcbi.1001000","article-title":"How cations\ncan assist DNase I in DNA binding and hydrolysis","volume":"6","author":"Gue\u0301roult","year":"2010","journal-title":"PLoS Comput. Biol."},{"issue":"3","key":"2026090307164343300_cit48","doi-asserted-by":"publisher","first-page":"665","DOI":"10.5051\/jkape.2002.32.3.665","article-title":"Effects of Lipomyces\nstarkeyi KSM 22 glucanhydrolase on human gingival fibroblasts","volume":"32","author":"Yun","year":"2002","journal-title":"J. Korean Acad. Periodontol."},{"issue":"1","key":"2026090307164343300_cit49","doi-asserted-by":"publisher","DOI":"10.1038\/s41598-025-06340-y","article-title":"Multifunctional nano-delivery\nsystem based on DNase I and photodynamic therapy for combatting enterococcus\nfaecalis biofilm infections","volume":"15","author":"Xia","year":"2025","journal-title":"Sci. Rep."},{"key":"2026090307164343300_cit50","doi-asserted-by":"publisher","first-page":"41717","DOI":"10.1021\/acsomega.6c01470","article-title":"Antifungal\nActivity, Biocompatibility, and Anti-Inflammatory Effects of Zerumbone-Enhanced\nAntimicrobial Photodynamic Inactivation in a Three-Dimensional Oral\nTissue Model","volume":"11","author":"Abreu-Pereira","year":"2026","journal-title":"ACS Omega"}],"container-title":["ACS Omega"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/pubs.acs.org\/doi\/pdf\/10.1021\/acsomega.6c03152","content-type":"application\/pdf","content-version":"vor","intended-application":"unspecified"},{"URL":"https:\/\/pubs.acs.org\/acsodf\/article-pdf\/11\/34\/51080\/66883461\/acsomega.6c03152.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/pubs.acs.org\/acsodf\/article-pdf\/11\/34\/51080\/66883461\/acsomega.6c03152.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,9,3]],"date-time":"2026-09-03T11:16:48Z","timestamp":1788434208000},"score":1,"resource":{"primary":{"URL":"https:\/\/pubs.acs.org\/acsodf\/article\/11\/34\/51080\/5276978\/Zerumbone-Combined-with-Hydrolytic-Enzymes-A"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,8,19]]},"references-count":50,"journal-issue":{"issue":"34","published-online":{"date-parts":[[2026,8,19]]},"published-print":{"date-parts":[[2026,9,1]]}},"URL":"https:\/\/doi.org\/10.1021\/acsomega.6c03152","relation":{},"ISSN":["2470-1343"],"issn-type":[{"value":"2470-1343","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,8,19]]}}}