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Challenges of the Non-Conventional Yeast Wickerhamomyces Anomalus in Winemaking

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Challenges of the Non-Conventional Yeast Wickerhamomyces Anomalus in Winemaking fermentation Review Challenges of the Non-Conventional Yeast Wickerhamomyces anomalus in Winemaking Beatriz Padilla 1, Jose V. Gil 2,3 and Paloma Manzanares 2,* 1 INCLIVA Health Research Institute, 46010 Valencia, Spain; [email protected] 2 Department of Biotechnology, Instituto de Agroquímica y Tecnología de Alimentos (IATA), Consejo Superior de Investigaciones Científicas (CSIC), Paterna, 46980 Valencia, Spain; [email protected] 3 Departamento de Medicina Preventiva y Salud Pública, Ciencias de la Alimentación, Toxicología y Medicina Legal, Facultad de Farmacia, Universitat de València, Burjassot, 46100 Valencia, Spain * Correspondence: [email protected]; Tel.: +34-96 390-0022 Received: 27 July 2018; Accepted: 18 August 2018; Published: 20 August 2018 Abstract: Nowadays it is widely accepted that non-Saccharomyces yeasts, which prevail during the early stages of alcoholic fermentation, contribute significantly to the character and quality of the final wine. Among these yeasts, Wickerhamomyces anomalus (formerly Pichia anomala, Hansenula anomala, Candida pelliculosa) has gained considerable importance for the wine industry since it exhibits interesting and potentially exploitable physiological and metabolic characteristics, although its growth along fermentation can still be seen as an uncontrollable risk. This species is widespread in nature and has been isolated from different environments including grapes and wines. Its use together with Saccharomyces cerevisiae in mixed culture fermentations has been proposed to increase wine particular characteristics. Here, we review the ability of W. anomalus to produce enzymes and metabolites of oenological relevance and we discuss its potential as a biocontrol agent in winemaking. Finally, biotechnological applications of W. anomalus beyond wine fermentation are briefly described. Keywords: non-Saccharomyces yeasts; Wickerhamomyces anomalus; Pichia anomala; enzymes; glycosidases; acetate esters; biocontrol; mixed starters; wine 1. Introduction Saccharomyces cerevisiae is the main microorganism involved in the alcoholic fermentation of grape must. Moreover, the use of selected S. cerevisiae strains has provided an improvement in the control and homogeneity of fermentations. However, winemaking is a non-sterile process, and many other species of yeasts belonging to various non-Saccharomyces genera prevail during the early stages of alcoholic fermentation and contribute significantly to the character and quality of the final wine [1]. In the past, non-Saccharomyces yeasts were considered of secondary significance or as undesirable spoilage yeasts. Nowadays, the role of non-Saccharomyces has been re-evaluated, and it is widely accepted that selected strains can positively influence the winemaking process [2]. Beyond the contribution of non-Saccharomyces yeasts to wine aroma complexity [3], these yeasts can help address some of the modern challenges in winemaking, including the reduction of the ethanol content of wine [4–7] or the control of wine spoilage [8,9]. Ecological studies have shown that species of mainly Hanseniaspora (Kloeckera), Candida, and Metschnikowia initiate the fermentation together with species of Pichia, Issatchenkia, and Kluyveromyces. Occasionally, representatives of Brettanomyces, Schizosaccharomyces, Torulaspora, Rhodotorula, Zygosaccharomyces, and Cryptococcus genera are also present. These yeasts decline by mid-fermentation, and then, S. cerevisiae becomes predominant and continues the fermentation [10]. Based on the capability of some of these non-Saccharomyces yeasts to produce flavor-enhancing enzymes or to modify the concentration Fermentation 2018, 4, 68; doi:10.3390/fermentation4030068 www.mdpi.com/journal/fermentation Fermentation 2018, 4, x FOR PEER REVIEW 2 of 13 mid-fermentation, and then, S. cerevisiae becomes predominant and continues the fermentation [10]. FermentationBased on 2018the , capability4, 68 of some of these non-Saccharomyces yeasts to produce flavor-enhancing2 of 14 enzymes or to modify the concentration of secondary metabolites, different mixed starters have been designed and proposed as a tool to enhance wine quality [3,11]. Moreover, several species including Lachanceaof secondary thermotolerans metabolites,, differentMetschnikowia mixed starterspulcherrima have, been Torulaspora designed delbrueckii and proposed, Pichia as a toolkluyvery to enhance, and wineSchizosaccharomyces quality [3,11]. Moreover, pombe are several already species commercially including available.Lachancea thermotolerans , Metschnikowia pulcherrima, TorulasporaWickerhamomyces delbrueckii, Pichia anomalus kluyvery, formerly, and Schizosaccharomyces known as Pichia pombe anomala,are already Hansenula commercially anomala, available. Candida pelliculosaWickerhamomyces was recently anomalus assigned, formerly to the knowngenus asWickerhamomycesPichia anomala, Hansenulabased on anomala,phylogenetic Candida analysis pelliculosa of wasgene recently sequences, assigned which to has the caused genus Wickerhamomycesmajor changes inbased the classification on phylogenetic of yeasts. analysis [12]. of This gene species sequences, has whichbeen frequently has caused isolated major changesfrom grapes in the and classification wines. Although of yeasts. traditionally [12]. This species W. anomalus has been is associated frequently withisolated excessive from grapes production and wines. of ethyl Although acetate, whic traditionallyh representsW. anomalus a seriousis handicap associated for with their excessive use in productionwinemaking, of ethylthis species acetate, has which gained represents considerable a serious importance handicap for for their the wine use in industry winemaking, since this it exhibits species interestinghas gained and considerable potentially importance exploitable for physiological the wine industry and metabolic since it exhibits characteristics interesting as summarized and potentially in Figureexploitable 1. Here, physiological we revisit andthe metaboliccontribution characteristics of W. anomalus assummarized in wine production. in Figure First,1. Here, we review we revisit the ecologythe contribution and prevalence of W. anomalus of this inyeast wine in production.winemaking, First, and we we review discuss the its ecology ability andto produce prevalence enzymes, of this yeastkiller intoxins, winemaking, and metabolites and we of discuss enological its ability relevanc to producee. Second, enzymes, we review killer the toxins, design and of mixed metabolites starters of ofenological W. anomalus relevance. with Second, S. cerevisiae we review to theimprove design ofwine mixed aroma starters complexity. of W. anomalus Finally,with S.we cerevisiae discussto biotechnologicalimprove wine aroma applications complexity. of Finally, W. anomalus we discuss beyond biotechnological wine fermentation. applications ofWhenW. anomalus citingbeyond older wineliterature, fermentation. the original When yeast citing species older name literature, will be the kept. original yeast species name will be kept. Floral and fruity notes Acetate esters mixed starter W. anomalus Glycosidases External addition or mixed starter Primary Control of aroma Proteases wine spoilage Haze prevention Figure 1. BenefitsBenefits of Wickerhamomyces anomalus in winemaking. 2. W. anomalus Is a Ubiquitous Yeast GenerallyGenerally AssociatedAssociated withwith WinemakingWinemaking W. anomalus is aa heterothallic,heterothallic, ascomycetous ascomycetous yeast, yeast, forming forming one one to fourto four hat-shaped hat-shaped ascospores ascospores[13, 14[13,14].]. The The placement placement of P. anomalaof P. anomalain the genusin theWickerhamomyces genus Wickerhamomyceswas due to was multigene due to phylogenetic multigene phylogeneticanalysis [11]. W.analysis anomalus [11].is W. a widely anomalus used is a name widely and used a proposal name and to conserve a proposal the to species conserve name the anomala species (-us).name W.anomala anomalus (-us).is a biotechnologicallyW. anomalus is relevanta biotechnologically yeast species withrelevant food, yeast environmental, species with industrial, food, environmental,and medical applications. industrial, Natural and medical habitats applications. of W. anomalus Naturalare very habitats diverse of and W. include anomalus tree exudates,are very plantsdiverse and and fruit include skins, tree insects, exudates, human plants tissues, and and fruit faeces, skins, and insects, also wastewaters human tissues, and marineand faeces, environments. and also Thewastewaters versatility and of thismarine species environments. is encouraged The by versatility its ability of to this tolerate species extreme is encouraged environmental by its conditions ability to toleratelike oxidative, extreme salt, environmental and osmotic conditions stress, as like well oxid as pHative, and salt, temperature and osmotic shocks stress,[15 as]. well Due as topH these and temperaturecharacteristics, shocks this yeast [15]. can Due be ato spoilage these characterist organism, forics, instance, this yeast inhigh-sugar can be a spoilage food products organism, [16,17 ]for or instance,silage [18 ].inIts high-sugar genome sequence food products is already [16,17] available, or silage providing [18]. Its the genome basis to sequence analyse metabolic is already capabilities, available, phylogeneticproviding the relationships, basis to andanalyse biotechnologically metabolic importantcapabilities, features phylogenetic [19,20]. The relationships, main physiological and biotechnologicallyand
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