Citation: Nasuti, C.; Ruffini, J.; Sola,
L.; Di Bacco, M.; Raimondi, S.;
Candeliere, F.; Solieri, L. Sour Beer as
Bioreservoir of Novel Craft Ale Yeast
Cultures. Microorganisms 2023, 11,
2138. https://doi.org/10.3390/
microorganisms11092138
Academic Editor: Maurizio Ciani
Received: 12 July 2023
Revised: 8 August 2023
Accepted: 20 August 2023
Published: 23 August 2023
Copyright: © 2023 by the authors.
Licensee MDPI, Basel, Switzerland.
This article is an open access article
distributed under the terms and
conditions of the Creative Commons
Attribution (CC BY) license (https://
creativecommons.org/licenses/by/
4.0/).
microorganisms
Article
Sour Beer as Bioreservoir of Novel Craft Ale Yeast Cultures
Chiara Nasuti
1
, Jennifer Ruffini
1
, Laura Sola
2
, Mario Di Bacco
3
, Stefano Raimondi
2
,
Francesco Candeliere
2
and Lisa Solieri
1,4,
*
1
Department of Life Sciences, University of Modena and Reggio Emilia, Via Amendola, 2-Pad. Besta,
42122 Reggio Emilia, Italy; chiara.nasuti@unimore.it (C.N.); 227818@studenti.unimore.it (J.R.)
2
Department of Life Sciences, University of Modena and Reggio Emilia, Via Campi 51, 41125 Modena, Italy;
laura.sola@unimore.it (L.S.); stefano.raimondi@unimore.it (S.R.); francesco.candeliere@unimore.it (F.C.)
3
Ca’ Del Brado Brewery, Via Andrea Costa, 146/2, 40065 Rastignano, Italy; quark@cadelbrado.it
4
National Biodiversity Future Center (NBFC), 90133 Palermo, Italy
* Correspondence: lisa.solieri@unimore.it; Tel.: +39-0522522026
Abstract: The increasing demand for craft beer is driving the search for novel ale yeast cultures from
brewing-related wild environments. The focus of bioprospecting for craft cultures is to identify feral
yeasts suitable to imprint unique sensorial attributes onto the final product. Here, we integrated phy-
logenetic, genotypic, genetic, and metabolomic techniques to demonstrate that sour beer during aging
in wooden barrels is a source of suitable craft ale yeast candidates. In contrast to the traditional lambic
beer maturation phase, during the aging of sour-matured production-style beer, different biotypes of
Saccharomyces cerevisiae dominated the cultivable in-house mycobiota, which were followed by Pichia
membranifaciens, Brettanomyces bruxellensis, and Brettanomyces anomalus. In addition, three putative
S. cerevisiae × Saccharomyces uvarum hybrids were identified. S. cerevisiae feral strains sporulated,
produced viable monosporic progenies, and had the STA1 gene downstream as a full-length promoter.
During hopped wort fermentation, four S. cerevisiae strains and the S. cerevisiae × S. uvarum hybrid
WY213 exceeded non-Saccharomyces strains in fermentative rate and ethanol production except for
P. membranifaciens WY122. This strain consumed maltose after a long lag phase, in contrast to the
phenotypic profile described for the species. According to the STA1+ genotype, S. cerevisiae partially
consumed dextrin. Among the volatile organic compounds (VOCs) produced by S. cerevisiae and
the S. cerevisiae × S. uvarum hybrid, phenylethyl alcohol, which has a fruit-like aroma, was the most
prevalent. In conclusion, the strains characterized here have relevant brewing properties and are
exploitable as indigenous craft beer starters.
Keywords: sour beer; craft brewing; STA1 gene; dextrin; hybrid; Saccharomyces cerevisiae; Saccharomyces
uvarum; Pichia membranifaciens
1. Introduction
Craft beer production has gained increasing interest in many countries as an alternative
to mainstream beer [1]. Craft brewing refers to relatively small breweries that aim to
produce beers with unique and original sensory characteristics based on quality and
diversity [2]. Special flavors increase perceived quality compared to commercial beer [3,4].
The sensory profile of beer is determined by the choice of malt or adjunct, hop variety,
water quality and yeast, which catalyzes metabolic reactions during wort fermentation,
giving the beer its flavor profile. Yeasts are responsible not only for sugar consumption and
ethanol and CO
2
production but also for releasing aroma-active secondary side-products
of alcoholic fermentation—higher alcohols and ethyl and acetate esters—that contribute
to solvent-like and fruity notes and a sweet taste [5]. They also can transform neutral
precursors in the wort and hops—glycosides, polyfunctional thiols, and terpenes—into
sensorial active compounds, as reviewed by Svedlund et al. [6]. This bio-flavoring aptitude
qualitatively and quantitatively varies depending on the strain and species employed [7].
Microorganisms 2023, 11, 2138. https://doi.org/10.3390/microorganisms11092138 https://www.mdpi.com/journal/microorganisms