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Alternative Saccharomyces Interspecies Hybrid Combinations and Their Potential for Low- Temperature Wort Fermentation

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Alternative Saccharomyces Interspecies Hybrid Combinations and Their Potential for Low- Temperature Wort Fermentation This is an electronic reprint of the original article. This reprint may differ from the original in pagination and typographic detail. Nikulin, Jarkko; Krogerus, Kristoffer; Gibson, Brian Alternative Saccharomyces interspecies hybrid combinations and their potential for low- temperature wort fermentation Published in: Yeast DOI: 10.1002/yea.3246 Published: 01/01/2018 Document Version Publisher's PDF, also known as Version of record Published under the following license: CC BY Please cite the original version: Nikulin, J., Krogerus, K., & Gibson, B. (2018). Alternative Saccharomyces interspecies hybrid combinations and their potential for low-temperature wort fermentation. Yeast, 35(1), 113-127. https://doi.org/10.1002/yea.3246 This material is protected by copyright and other intellectual property rights, and duplication or sale of all or part of any of the repository collections is not permitted, except that material may be duplicated by you for your research use or educational purposes in electronic or print form. You must obtain permission for any other use. Electronic or print copies may not be offered, whether for sale or otherwise to anyone who is not an authorised user. Powered by TCPDF (www.tcpdf.org) Yeast Yeast 2018; 35: 113–127. Published online 30 August 2017 in Wiley Online Library (wileyonlinelibrary.com) DOI: 10.1002/yea.3246 Yeast Interspecies Hybrids Alternative Saccharomyces interspecies hybrid combinations and their potential for low-temperature wort fermentation † † Jarkko Nikulin1,2 , Kristoffer Krogerus1,3 and Brian Gibson1* 1VTT Technical Research Centre of Finland Ltd, Tietotie 2, PO Box 1000FI-02044 VTT Espoo, Finland 2Chemical Process Engineering, Faculty of Technology, University of Oulu, PO Box 8000FI-90014 Oulun Yliopisto, Finland 3Department of Biotechnology and Chemical Technology, Aalto University, School of Chemical Technology, Kemistintie 1, Aalto, PO Box 16100 FI- 00076 Espoo, Finland *Correspondence to: Abstract Brian Gibson, VTT Technical Saccharomyces cerevisiae Saccharomyces eubayanus Research Centre of Finland Ltd, The lager yeast hybrid ( × ) possesses fi Tietotie 2, PO Box 1000, FI- two key characteristics that are essential for lager brewing: ef cient sugar utilization 02044 VTT, Espoo, Finland. and cold tolerance. Here we explore the possibility that the lager yeast phenotype E-mail: brian.gibson@vtt.fi can be recreated by hybridizing S. cerevisiae ale yeast with a number of cold-tolerant † Saccharomyces species including Saccharomyces arboricola, Saccharomyces eubayanus, Joint first authors. Saccharomyces mikatae and Saccharomyces uvarum. Interspecies hybrids performed better than parental strains in lager brewing conditions (12°C and 12°P wort), with the S. mikatae hybrid performing as well as the S. eubayanus hybrid. Where the S. cerevisiae parent was capable of utilizing maltotriose, this trait was inherited by the hybrids. A greater production of higher alcohols and esters by the hybrids resulted in the produc- tion of more aromatic beers relative to the parents. Strong fermentation performance relative to the parents was dependent on ploidy, with polyploid hybrids (3n, 4n) performing better than diploid hybrids. All hybrids produced 4-vinyl guaiacol, a smoke/clove aroma generally considered an off flavour in lager beer. This characteristic could however be eliminated by isolating spore clones from a fertile hybrid of S. cerevisiae and S. mikatae. The results suggest that S. eubayanus is dispensable when constructing yeast hybrids that express the typical lager yeast phenotype. © 2017 The Authors. Yeast published by John Wiley & Sons, Ltd. Keywords: saccharomyces pastorianus; lager beer; hybridization; cold tolerance; Received: 26 May 2017 maltotriose; aroma; phenolic off flavour Accepted: 18 July 2017 Introduction to its central role in the brewing industry is pre- sumably due to a fortuitous combination of cir- Traditionally, lager beers are brewed with Saccha- cumstance and its superior biological properties: romyces pastorianus (Gibson and Liti, 2015). This the ability to efficiently ferment wort sugars, bottom-fermenting yeast is an interspecies hybrid which has been obtained from a S. cerevisiae of Saccharomyces cerevisiae and Saccharomyces parent, and cold tolerance, which has been eubayanus (Libkind et al., 2011). This hybridiza- obtained from a S. eubayanus parent (Gibson tion presumably occurred in Bavaria during the et al., 2013). The recent creation of de novo lager fifteenth or sixteenth centuries, coincident with yeast hybrids has revealed that hybrid strains the rise of low-temperature brewing of lager beer typically outperform parent strains during wort (Meussdoerffer, 2009). The rise of S. pastorianus fermentation and that these hybrids grow well at © 2017 The Authors. Yeast published by John Wiley & Sons, Ltd. This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited. 114 J. Nikulin et al. the lower temperatures associated with lager attributable to the two parent species is not brewing (Krogerus et al., 2015; Mertens et al., completely understood. Studies on the maltose 2015). The lager yeasts that are used today in and maltotriose uptake capabilities of the non-S. the brewing industry are closely related and ex- cerevisiae species in the Saccharomyces complex hibit little genetic or phenotypic diversity (Dunn are limited, but in general maltose and particularly and Sherlock, 2008; Okuno et al., 2016). Genetic maltotriose use tends to be poor in non- and functional diversity could potentially be domesticated strains (Bell et al., 2001; Gallone increased through the creation of de novo et al., 2016). The distribution of sugar transporters lager yeast hybrids from diverse lineages in the in natural lager yeasts suggests that the majority of Saccharomyces genus, as long as the requisite them have been obtained from the S. cerevisiae characteristics are present. parent. While it has been hypothesized that the S. eubayanus is not the only cold tolerant species maltotriose-transporting Mtt1 transporter in lager in the Saccharomyces genus. Strains of Saccharo- yeast is derived from the S. eubayanus parent myces kudriavzevii and Saccharomyces uvarum (Dietvorst et al., 2005; Nakao et al., 2009), recent are also adept at growing and fermenting at low studies have found this transporter in S. cerevisiae temperatures (Gonçalves et al., 2011; González strains (Magalhães et al., 2016; Vidgren, 2010). et al., 2006; López-Malo et al., 2013; Masneuf- Hence, it is possible that the sugar-utilizing ability Pomarède et al., 2010; Paget et al., 2014). These of the non-S. cerevisiae parent of de novo lager species are usually associated with wine and cider yeast hybrids has little effect on the hybrid’s fermentation (Naumov et al., 2001; González sugar utilization abilities, as long as the S. et al., 2006). Hybrids of these, i.e. S. cerevisiae × cerevisiae parent is capable of efficient maltose S. kudriavzevii and S. cerevisiae × S. uvarum, have and maltotriose uptake. also been isolated from both brewing and wine- One important feature of lager yeast is their making environments (Pérez-Torrado et al., 2017; inability to produce 4-vinyl guaiacol (Mukai Sampaio and Gonçalves, 2008). As these ‘alterna- et al., 2014; Richard et al., 2015). This compound tive’ Saccharomyces species are also cold tolerant, contributes to so-called phenolic off-flavour (POF) they may be feasible alternatives to S. eubayanus and is considered undesirable in most beer styles in the creation of interspecific hybrids for the (Gallone et al., 2016; Gonçalves et al., 2016). 4- purpose of lager brewing. The potential of de novo Vinyl guaiacol is derived from ferulic acid and is interspecific hybrids created from other species in usually associated with non-domesticated yeasts. the Saccharomyces genus (i.e. Saccharomyces In nature such hydroxycinnamic acid transforma- arboricola, S. kudriavzevii, Saccharomyces tions serve a protective function (Stratford et al., mikatae, Saccharomyces paradoxus or S. uvarum) 2007), but are unlikely to be important for survival for brewing purposes has, to our knowledge, not in the brewing environment. The phenotype is been explored. However, their potential in attributed to the action of the adjacent PAD1 and winemaking conditions has been demonstrated in FDC1 genes, and we have found it to be a feature recent studies. The use of de novo S. cerevisiae of the type-strains of all non-S. cerevisiae Saccha- interspecific hybrids with S. kudriavzevii (Bellon romyces species, including S. eubayanus. De novo et al., 2011; Lopandic et al., 2016), S. mikatae lager yeast hybrids created in recent studies have (Bellon et al., 2013), S. paradoxus (Bellon et al., all produced 4-vinyl guaiacol owing to inheritance 2011) and S. uvarum (Bellon et al., 2011; of functional PAD1 and FDC1 genes from S. Lopandic et al., 2016) has revealed their potential eubayanus (Krogerus et al., 2015; Mertens et al., for increasing aromatic diversity and fermentation 2015). However, if one of the parent strains con- performance. tains loss-of-function mutations in either PAD1 or While cold tolerance probably played an impor- FDC1, and the hybrid is also fertile, the phenotype tant role in the emergence of lager yeast, its can be removed through meiotic recombination success can also be attributed to its fermentation and sporulation, as demonstrated by Krogerus properties. These
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