Saccharomyces Cerevisiae in the Production of Fermented Beverages
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beverages Review Saccharomyces cerevisiae in the Production of Fermented Beverages Graeme M Walker 1,* and Graham G Stewart 2 1 Abertay University, Dundee, Scotland DD1 1HG, UK 2 Heriot-Watt University, Edinburgh, Scotland EH14 4AS, UK; [email protected] * Correspondence: [email protected]; Tel.: +44-1382-308658 Academic Editor: Edgar Chambers IV Received: 20 October 2016; Accepted: 11 November 2016; Published: 17 November 2016 Abstract: Alcoholic beverages are produced following the fermentation of sugars by yeasts, mainly (but not exclusively) strains of the species, Saccharomyces cerevisiae. The sugary starting materials may emanate from cereal starches (which require enzymatic pre-hydrolysis) in the case of beers and whiskies, sucrose-rich plants (molasses or sugar juice from sugarcane) in the case of rums, or from fruits (which do not require pre-hydrolysis) in the case of wines and brandies. In the presence of sugars, together with other essential nutrients such as amino acids, minerals and vitamins, S. cerevisiae will conduct fermentative metabolism to ethanol and carbon dioxide (as the primary fermentation metabolites) as the cells strive to make energy and regenerate the coenzyme NAD+ under anaerobic conditions. Yeasts will also produce numerous secondary metabolites which act as important beverage flavour congeners, including higher alcohols, esters, carbonyls and sulphur compounds. These are very important in dictating the final flavour and aroma characteristics of beverages such as beer and wine, but also in distilled beverages such as whisky, rum and brandy. Therefore, yeasts are of vital importance in providing the alcohol content and the sensory profiles of such beverages. This Introductory Chapter reviews, in general, the growth, physiology and metabolism of S. cerevisiae in alcoholic beverage fermentations. Keywords: Saccharomyces cerevisiae; fermented beverages 1. Introduction Yeasts in Alcoholic Beverage Fermentations The production of alcoholic beverages from fermentable carbon sources by yeast is the oldest and most economically important of all biotechnologies. Yeast plays a vital role in the production of all alcoholic beverages (see Figure1) and the selection of suitable yeast strains is essential not only to maximise alcohol yield, but also to maintain beverage sensory quality. The yeast species that dominates in the production of alcoholic beverages worldwide is Saccharomyces cerevisiae, and the particular strains of this species employed in fermentation exert a profound influence on the flavour and aroma characteristics of different beverages. For large-scale beverage fermentations, as in brewing, winemaking and distilled spirit production, pure cultures of selected strains of S. cerevisiae are usually used. These strains are either sourced in house or supplied from yeast producing companies. In smaller-scale (artisanal) processes, spontaneous fermentations may be allowed to occur that rely on indigenous microbiological flora (wild yeasts and bacteria) present in the raw material and in the production facility. For example, this would be typical in small distilleries in Mexico (for Tequila and Mezcal production) and in Brazil (for Cachaça production). In some types of alcoholic beverage fermentations, non-S. cerevisiae yeasts may be employed either as starter cultures, or occur naturally. For example, in winemaking the S. cerevisiae yeast strain used Beverages 2016, 2, 30; doi:10.3390/beverages2040030 www.mdpi.com/journal/beverages Beverages 2016, 2, 30 2 of 12 Beverages 2016, 2, 30 2 of 12 tocommence commence fermentation fermentation may may be be overrun overrun by by the the indigenous indigenous yeast yeast flora flora associated associated with with the the grapes. grapes. Table1 1 summarises summarises different different yeast yeast species species encountered encountered in in alcoholic alcoholic beverage beverage fermentations. fermentations. Raw Materials (Cereals, Sugar Cane, Molasses, Fruit, Agave, Whey, etc.) Fermentable Sugars (Maltose, Maltotriose, Sucrose, Fructose, Glucose) Fermentation S. cerevisiae Maturation Distillation (e.g., Beer, Wine, Cider, Mead) Matured Spirits Non‐Matured Spirits (e.g., Whisky, Cognac, Rum) Non‐Flavoured Flavoured (e.g., Vodka, Grappa, Tsiporo) (e.g., Gin) Figure 1. The key roles of Saccharomyces cerevisiae in production of fermentedfermented beverages. Table 1. The main types of yeast used in the production of selected alcoholic beverages. Table 1. The main types of yeast used in the production of selected alcoholic beverages. Beverage Yeast Involved Comments Beverage Yeast InvolvedLager yeasts are likely a natural hybrid Comments (S. cerevisiae & S. eubayanus). Relatively fewLager strains yeasts employed are likely a naturalin lager hybrid fermentations. (S. cerevisiae Lager& S. eubayanus strains ). Lager beer: Saccharomyces utilise maltotrioseRelatively more few strainsefficiently employed than ale in lager strains, fermentations. and they ferment Lager strains at pastorianus Lager beer: Saccharomyces pastorianuscooler temperatures.utilise maltotriose Ale yeasts more are efficiently polyploid than strains. ale strains, Numerous and they fermentstrains at Beer Ale: SaccharomycesAle: Saccharomyces cerevisiae cerevisiae cooler temperatures. Ale yeasts are polyploid strains. Numerous strains Beer employed in ale brewing. Ale yeasts ferment at warmer temperatures LambicLambic beer: beer:BrettanomycesBrettanomyces bruxellensis employed in ale brewing. Ale yeasts ferment at warmer temperatures compared with lager yeasts. Lambic beer traditionally produced via bruxellensisand other and yeasts other yeasts compared with lager yeasts. Lambic beer traditionally produced via spontaneousspontaneous fermentation, fermentation, but some but lambic some lambicand Belgian and Belgian-style‐style beers beers employ pureemploy starter pure cultures starter culturesof Brettanomyces of Brettanomyces spp. spp. TraditionalTraditional winemaking winemaking is characterised is characterised by spontaneous by spontaneous fermentations fermentations of grape mustof with grape naturally must with occurring naturally microflora occurring microflora (the main (the yeast main genera yeast genera associated with grapes are: Kloeckera and Hanseniaspora, associated with lesser grapes representations are: Kloeckera of andCandida Hanseniaspora, Metchnikowia, with, Cryptococcus lesser , Pichia Saccharomyces cerevisiae, Saccharomyces cerevisiae, representationsand Kluyveromyces of Candida, Metchnikowiaand very low populations, Cryptococcus of Saccharomyces, Pichia and cerevisiae). Wine Saccharomyces bayanus (pure cultures) Saccharomyces bayanus (pure KluyveromycesModern, and very large-scale low populations wineries use speciallyof Saccharomyces selected starter cerevisiae cultures). of Wine and naturally-occurring yeasts cultures) and naturally‐ Modern, largeS. cerevisiae‐scale wineriesstrains available use specially in dried selected form (e.g., starter active cultures dry yeast, of ADY) from specialist yeast supply companies. Occasionally, secondary occurring yeasts S. cerevisiae strains available in dried form (e.g., active dry yeast, ADY) commercial non-Saccharomyces starter cultures (e.g., Candida stellata) from specialistmay beyeast employed supply to companies. impart specific Occasionally, flavour and secondary aroma to wine. commercialScotch non‐Saccharomyces whisky producers starter currently cultures use selected (e.g., Candida distilling stellata strains) may of be employedS. cerevisiae to impartin threespecific main flavour formats, and cream aroma yeast, to pressed wine. (cake) and dried Whisky Saccharomyces cerevisiae Scotch whiskyyeast. producers Malt whisky currently distilleries use traditionally selected distilling use pressed strains yeast, of but larger S. cerevisiaegrain in three distillers main have formats, now adopted cream yeast, cream pressed yeast. Dried (cake) yeasts and are dried not as prevalent as pressed and cream formats in whisky fermentations. Whisky Saccharomyces cerevisiae yeast. Malt whisky distilleries traditionally use pressed yeast, but larger grain distillersS. cerevisiae have nowstrains adopted in rum fermentationscream yeast. areDried developed yeasts are as starter not as cultures and provide faster fermentation with more higher alcohols and prevalent asfatty pressed acids, butand less cream esters formats resulting in inwhisky lighter fermentations. style rums. Saccharomyces cerevisiae and Rum S. cerevisiaeSchiz. strains pombe in rumin rum fermentations fermentations are provides developed slower as fermentations starter cultures Schizosaccharomyces pombe and provideleading faster to fermentation less higher alcohols with more and fatty higher acids, alcohols but more and esters fatty resulting acids, but less estersin heavy, resulting strong in aroma lighter rums. style Growth rums. of Schiz. pombe is favoured by low Saccharomyces cerevisiae and pH, higher sugar conc. Rum Schiz. pombe in rum fermentations provides slower fermentations leading Schizosaccharomyces pombe Various yeasts have been isolated from such processes: S. cerevisiae, Tequila, Mezcal, Natural yeasts in artisanal Agaveto less higher alcohols and fatty acids, but more esters resulting in heavy, Kluyveromyces marxianus, Pichia spp., Brettanomyces spp., Bacanora fermentations strong aromaRhodotorula rums. Growthspp., etc. of Schiz. pombe is favoured by low pH, higher sugar conc. For brandies, cognac, etc. the base wine is produced by pure starter Brandies, Gin, Tequila,