{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,5,1]],"date-time":"2026-05-01T20:57:15Z","timestamp":1777669035679,"version":"3.51.4"},"reference-count":57,"publisher":"MDPI AG","issue":"2","license":[{"start":{"date-parts":[[2018,4,10]],"date-time":"2018-04-10T00:00:00Z","timestamp":1523318400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Beverages"],"abstract":"<jats:p>The unwanted modification of wine sensory attributes by yeasts of the species Brettanomyces bruxellensis due to the production of volatile phenols is presently the main microbiological threat to red wine quality. The effects of ethylphenols and other metabolites on wine flavor is now recognized worldwide and the object of lively debate. The focus of this review is to provide an update of the present knowledge and practice on the prevention of this problem in the wine industry. Brettanomyces bruxellensis, or its teleomorph, Dekkera bruxellensis, are rarely found in the natural environment and, although frequently isolated from fermenting substrates, their numbers are relatively low when compared with other fermenting species. Despite this rarity, they have long been studied for their unusual metabolical features (e.g., the Custers effect). Rising interest over the last decades is mostly due to volatile phenol production affecting high quality red wines worldwide. The challenges have been dealt with together by researchers and winemakers in an effective way and this has enabled a state where, presently, knowledge and prevention of the problem at the winery level is readily accessible. Today, the main issues have shifted from technological to sensory science concerning the effects of metabolites other than ethylphenols and the over estimation of the detrimental impact by ethylphenols on flavor. Hopefully, these questions will continue to be tackled together by science and industry for the benefit of wine enjoyment.<\/jats:p>","DOI":"10.3390\/beverages4020032","type":"journal-article","created":{"date-parts":[[2018,4,10]],"date-time":"2018-04-10T13:06:08Z","timestamp":1523365568000},"page":"32","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":44,"title":["Two Decades of \u201cHorse Sweat\u201d Taint and Brettanomyces Yeasts in Wine: Where do We Stand Now?"],"prefix":"10.3390","volume":"4","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-7985-963X","authenticated-orcid":false,"given":"Manuel","family":"Malfeito-Ferreira","sequence":"first","affiliation":[{"name":"Linking Landscape, Environment, Agriculture and Food Research Centre (LEAF), Instituto Superior de Agronomia, University of Lisbon, Tapada da Ajuda, 1349-017 Lisboa, Portugal"}]}],"member":"1968","published-online":{"date-parts":[[2018,4,10]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"323","DOI":"10.1002\/yea.3023","article-title":"The wine and beer yeast Dekkera bruxellensis","volume":"31","author":"Schifferdecker","year":"2014","journal-title":"Yeast"},{"key":"ref_2","first-page":"A31","article-title":"The Custers effect (negative Pasteur effect) as a diagnostic criterion for the genus Brettanomyces","volume":"35","author":"Scheffers","year":"1969","journal-title":"Antonie Leeuwenhoek"},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"165","DOI":"10.1002\/jsfa.2740600205","article-title":"The origin of ethylphenols in wines","volume":"60","author":"Chatonnet","year":"1992","journal-title":"J. Sci. Food Agric."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"680","DOI":"10.1111\/ajgw.12200","article-title":"Harnessing improved understanding of Brettanomyces bruxellensis biology to mitigate the risk of wine spoilage","volume":"21","author":"Curtin","year":"2015","journal-title":"Aust. J. Grape Wine Res."},{"key":"ref_5","doi-asserted-by":"crossref","unstructured":"Blackburn, C. (2006). Spoilage activities of Dekkera\/Brettanomyces spp.. Food Spoilage Microorganisms, Woodhead Publishers. Chapter 13.","DOI":"10.1201\/9781439824573"},{"key":"ref_6","first-page":"161","article-title":"The origin of Brettanomyces bruxellensis in wines: A review","volume":"41","author":"Renouf","year":"2007","journal-title":"J. Int. Sci. Vigne Vin"},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"10","DOI":"10.1016\/j.foodchem.2006.03.030","article-title":"The production of ethylphenols in wine by yeasts of the genera Brettanomyces and Dekkera: A review","volume":"102","author":"Morata","year":"2007","journal-title":"Food Chem."},{"key":"ref_8","first-page":"128","article-title":"Significance of Brettanomyces and Dekkera during Winemaking: A Synoptic Review","volume":"29","author":"Oelofse","year":"2008","journal-title":"S. Afr. J. Enol. Vitic."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"1474","DOI":"10.1016\/j.lwt.2010.06.010","article-title":"The challenge of Brettanomyces in wine","volume":"43","author":"Wedral","year":"2010","journal-title":"LWT Food Sci. Technol."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"57","DOI":"10.1146\/annurev-food-030212-182533","article-title":"Advances in the control of wine spoilage by Zygosaccharomyces and Dekkera\/Brettanomyces","volume":"4","author":"Zuehlke","year":"2013","journal-title":"Annu. Rev. Food Sci. Technol."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"24","DOI":"10.1016\/j.ijfoodmicro.2015.04.005","article-title":"Brettanomyces yeasts\u2014From spoilage organisms to valuable contributors to industrial fermentations","volume":"206","author":"Steensels","year":"2015","journal-title":"Int. J. Food Microbiol."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"161","DOI":"10.1016\/j.fm.2016.06.008","article-title":"Brettanomyces bruxellensis, a survivalist prepared for the wine apocalypse and other beverages","volume":"59","author":"Smith","year":"2016","journal-title":"Food Microbiol."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"180","DOI":"10.1007\/s11274-017-2345-z","article-title":"Brettanomyces bruxellensis yeasts: Impact on wine and winemaking","volume":"33","author":"Agnolucci","year":"2017","journal-title":"World J. Microbiol. Biotechnol."},{"key":"ref_14","unstructured":"Goode, J., and Harrop, S. (2008, January 15). Wine Faults and Their Prevalence: Data from the World\u2019s Largest Blind Tasting. Proceedings of the 20th Entretiens Scientifiques Lallemand, Horsens, Denmark."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"161","DOI":"10.1016\/j.foodres.2017.06.057","article-title":"Influence of Brettanomyces ethylphenols on red wine aroma evaluated by consumers in the United States and Portugal","volume":"100","author":"Schumaker","year":"2017","journal-title":"Food Res. Int."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"9401","DOI":"10.1021\/acs.jafc.6b04074","article-title":"Hydroxycinnamoyl Glucose and Tartrate Esters and Their Role in the Formation of Ethylphenols in Wine","volume":"64","author":"Hixson","year":"2016","journal-title":"J. Agric. Food Chem."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"11610","DOI":"10.1021\/jf403440k","article-title":"Metabolism of Nonesterified and Esterified Hydroxycinnamic Acids in Red Wines by Brettanomyces bruxellensis","volume":"61","author":"Schopp","year":"2013","journal-title":"J. Agric. Food Chem."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"2293","DOI":"10.1021\/jf204908s","article-title":"Hydroxycinnamic Acid Ethyl Esters as Precursors to Ethylphenols in Wine","volume":"60","author":"Hixson","year":"2012","journal-title":"J. Agric. Food Chem."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"1281","DOI":"10.1016\/j.talanta.2007.08.045","article-title":"Impact of malolactic fermentation on low molecular weight phenolic compounds","volume":"74","author":"Cabrita","year":"2008","journal-title":"Talanta"},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"357","DOI":"10.5344\/ajev.2015.14108","article-title":"Impact of Oenococcus oeni on wine hydroxycinnamic acids and volatile phenol production by Brettanomyces bruxellensis","volume":"66","author":"Chescheir","year":"2015","journal-title":"Am. J. Enol. Vitic."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"23","DOI":"10.5344\/ajev.2016.16015","article-title":"Influence of Oenococcus oeni and Brettanomyces bruxellensis on hydroxycinnamic acids and volatile phenols of aged wine","volume":"68","author":"Madsen","year":"2017","journal-title":"Am. J. Enol. Vitic."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"377","DOI":"10.1016\/S0740-0020(03)00023-6","article-title":"Factors affecting the production of 4-ethylphenol by the yeast Dekkera bruxellensis in enological conditions","volume":"20","author":"Dias","year":"2003","journal-title":"Food Microbiol."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"487","DOI":"10.1007\/s00217-015-2559-y","article-title":"Growth and volatile phenol production by Brettanomyces bruxellensis in different grapevine varieties during fermentation and in finished wine","volume":"242","author":"Chandra","year":"2015","journal-title":"Eur. Food Res. Technol."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"201","DOI":"10.1016\/j.ijfoodmicro.2007.11.020","article-title":"Survival patterns of Dekkera bruxellensis in wines and inhibitory effect of sulphur dioxide","volume":"121","author":"Barata","year":"2008","journal-title":"Int. J. Food Microbiol."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"133","DOI":"10.5344\/ajev.2006.57.2.133","article-title":"Growth and 4-ethylphenol production by the yeast Pichia guilliermondii in grape juices","volume":"57","author":"Barata","year":"2006","journal-title":"Am. J. Enol. Vitic."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"567","DOI":"10.1016\/S0740-0020(02)00152-1","article-title":"Identification of yeasts isolated from wine related environments and capable of producing 4-ethylphenol","volume":"20","author":"Dias","year":"2003","journal-title":"Food Microbiol."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"79","DOI":"10.1016\/j.ijfoodmicro.2005.05.014","article-title":"Molecular typing of the yeast species Dekkera bruxellensis and Pichia guilliermondii recovered from wine related sources","volume":"106","author":"Martorell","year":"2006","journal-title":"Int. J. Food Microbiol."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"11","DOI":"10.1111\/j.1472-765X.1987.tb01633.x","article-title":"Determination of the carbonation tolerance of yeasts","volume":"5","author":"Ison","year":"1987","journal-title":"Lett. Appl. Microbiol."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"588","DOI":"10.1046\/j.1365-2672.2001.01275.x","article-title":"Development and use of a new medium to detect yeasts of the genera Dekkera\/Brettanomyces spp.","volume":"90","author":"Rodrigues","year":"2001","journal-title":"J. Appl. Microbiol."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"154","DOI":"10.1016\/j.micres.2006.02.006","article-title":"Development of an enrichment medium to detect Dekkera\/Brettanomyces bruxellensis a spoilage yeast, on the surface of grape berries","volume":"162","author":"Renouf","year":"2007","journal-title":"Microbiol. Res."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"571","DOI":"10.1128\/aem.53.3.571-576.1987","article-title":"Analysis of yeast flora associated with grape sour rot and of the chemical disease markers","volume":"53","author":"Guerzoni","year":"1987","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"322","DOI":"10.5344\/ajev.2002.53.4.322","article-title":"Rapid detection and identification of Brettanomyces from winery air samples based on peptide nucleic acid analysis","volume":"53","author":"Connell","year":"2002","journal-title":"Am. J. Enol. Vitic."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"379","DOI":"10.5344\/ajev.2007.58.3.379","article-title":"Survival of wine microorganisms in the bottle during storage","volume":"58","author":"Renouf","year":"2007","journal-title":"Am. J. Enol. Vitic."},{"key":"ref_34","first-page":"34","article-title":"The influence of oxygen on the \u201chorse sweat taint\u201d in red wines","volume":"24","author":"Rodrigues","year":"2001","journal-title":"Ital. Food Beverage Technol."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"95","DOI":"10.1007\/s13213-010-0098-0","article-title":"Yeasts and wine off-flavours: A technological perspective","volume":"61","year":"2011","journal-title":"Ann. Microbiol."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"196","DOI":"10.1016\/j.fm.2016.06.007","article-title":"Viable But Not Culturable (VBNC) state of Brettanomyces bruxellensis in wine: New insights on molecular basis of VBNC behaviour using a transcriptomic approach","volume":"59","author":"Capozzi","year":"2016","journal-title":"Food Microbiol."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"965","DOI":"10.1007\/s00253-009-2409-6","article-title":"Saccharomyces cerevisiae CCMI 885 secretes peptides that inhibit the growth of some non-Saccharomyces wine-related strains","volume":"86","author":"Albergaria","year":"2010","journal-title":"Appl. Microbiol. Biotechnol."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"207","DOI":"10.1111\/jam.13121","article-title":"Evaluation of damage induced by Kwkt and Pikt zymocins against Brettanomyces\/Dekkera spoilage yeast, as compared to sulphur dioxide","volume":"121","author":"Oro","year":"2016","journal-title":"J. Appl. Microbiol."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"422","DOI":"10.1016\/j.fm.2007.10.003","article-title":"Evaluation of the inhibitory effect of dimethyl dicarbonate (DMDC) against wine microorganisms","volume":"25","author":"Costa","year":"2008","journal-title":"Food Microbiol."},{"key":"ref_40","first-page":"49","article-title":"Effectiveness of chitosan preparations against Brettanomyces bruxellensis grown in culture media and red wines","volume":"50","author":"Petrova","year":"2016","journal-title":"J. Int. Sci. Vigne Vin"},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"186","DOI":"10.1111\/lam.12444","article-title":"Heat inactivation of wine spoilage yeast Dekkera bruxellensis by hot water treatment","volume":"61","author":"Fabrizio","year":"2015","journal-title":"Lett. Appl. Microbiol."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"152","DOI":"10.1111\/jam.12981","article-title":"Impact of pulsed-electric field and high-voltage electrical discharges on red wine microbial stabilization and quality characteristics","volume":"120","author":"Delsart","year":"2016","journal-title":"J. Appl. Microbiol."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/j.foodcont.2017.05.028","article-title":"High pressure processing inactivation of Brettanomyces bruxellensis in seven different table wines","volume":"81","author":"Silva","year":"2017","journal-title":"Food Control"},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"1013","DOI":"10.1111\/j.1567-1364.2007.00267.x","article-title":"Species-specific identification of Dekkera\/Brettanomyces yeasts by fluorescently labeled DNA probes targeting the 26S rRNA","volume":"7","year":"2007","journal-title":"FEMS Yeast Res."},{"key":"ref_45","first-page":"153","article-title":"Detection of Dekkera\/Brettanomyces bruxellensis in pressed Sangiovese grapes by real time PCR","volume":"19","author":"Agnolucci","year":"2007","journal-title":"Ital. J. Food Sci."},{"key":"ref_46","first-page":"223","article-title":"Evaluation of three Brettanomyces qPCR commercial kits: Results from an interlaboratory study","volume":"50","author":"Longin","year":"2016","journal-title":"J. Int. Sci. Vigne Vin"},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"196","DOI":"10.1016\/j.fm.2014.03.010","article-title":"Use of propidium monoazide for the enumeration of viable Brettanomyces bruxellensis in wine and beer by quantitative PCR","volume":"42","author":"Vendrame","year":"2014","journal-title":"Food Microbiol."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"549","DOI":"10.1007\/s13213-010-0085-5","article-title":"Susceptibility of wine spoilage yeasts and bacteria in the planktonic state and in biofilms to disinfectants","volume":"60","author":"Tristezza","year":"2010","journal-title":"Ann. Microbiol."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"140","DOI":"10.5539\/jfr.v2n1p140","article-title":"Enumeration and identification of 4-ethylphenol producing yeasts recovered from the wood of wine ageing barriques after different sanitation treatments","volume":"2","author":"Barata","year":"2013","journal-title":"J. Food Res."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"799","DOI":"10.1111\/j.1745-4549.2012.00702.x","article-title":"Removal of Brettanomyces bruxellensis from red wine using membrane filtration","volume":"37","author":"Umiker","year":"2013","journal-title":"J. Food Process. Preserv."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"1209","DOI":"10.1111\/jam.12446","article-title":"Antimicrobial activity of Metschnikowia pulcherrima on wine yeasts","volume":"116","author":"Oro","year":"2014","journal-title":"J. Appl. Microbiol."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"40","DOI":"10.1016\/j.fm.2014.03.002","article-title":"A response surface methodology study on the role of factors affecting growth and volatile phenol production by Brettanomyces bruxellensis ISA 2211 in wine","volume":"42","author":"Chandra","year":"2014","journal-title":"Food Microbiol."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"1097","DOI":"10.1111\/j.1567-1364.2008.00415.x","article-title":"The effect of sugar concentration and temperature on growth and volatile phenol production by Dekkera bruxellensis in wine","volume":"8","author":"Barata","year":"2008","journal-title":"FEMS Yeast Res."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"183","DOI":"10.1016\/j.jchromb.2016.10.036","article-title":"A simple, cheap and reliable method for control of 4-ethylphenol and4-ethylguaiacol in red wines. Screening of fining agents for reducing volatile phenols levels in red wines","volume":"1041\u20131042","author":"Milheiro","year":"2017","journal-title":"J. Chromatogr. B"},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"2126","DOI":"10.1016\/j.foodchem.2011.12.012","article-title":"4-Ethylphenol and 4-ethylguaiacol depletion in wine using esterified cellulose","volume":"132","author":"Larcher","year":"2012","journal-title":"Food Chem."},{"key":"ref_56","first-page":"12","article-title":"Creation and Use of a Brettanomyces Aroma Wheel","volume":"1","author":"Joseph","year":"2017","journal-title":"Catalyst"},{"key":"ref_57","doi-asserted-by":"crossref","unstructured":"Smith, C. (2013). Postmodern Winemaking, Rethinking the Modern Science of an Ancient Craft, University California Press.","DOI":"10.1525\/9780520958548"}],"container-title":["Beverages"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2306-5710\/4\/2\/32\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,11]],"date-time":"2025-10-11T15:00:08Z","timestamp":1760194808000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2306-5710\/4\/2\/32"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2018,4,10]]},"references-count":57,"journal-issue":{"issue":"2","published-online":{"date-parts":[[2018,6]]}},"alternative-id":["beverages4020032"],"URL":"https:\/\/doi.org\/10.3390\/beverages4020032","relation":{},"ISSN":["2306-5710"],"issn-type":[{"value":"2306-5710","type":"electronic"}],"subject":[],"published":{"date-parts":[[2018,4,10]]}}}