Available online at www.sciencedirect.
com
ScienceDirect
Contribution of non-conventional yeasts in alcoholic
beverages
Anne Gschaedler
Non-conventional yeasts includes a wide array of different
genera of non-Saccharomyces. In the past considered as
undesired or spoilage yeasts now they play an important role in
enhancing the composition and aroma profile of wine and
alcoholic beverages. The first publications dealing with non-
Saccharomyces describes their role in spontaneous
fermentations and their ability to influence the varietal flavors.
Recent publications highlights the role of non-Saccharomyces
as a selected strain for controlled fermentations mainly in co-
fermentations with selected Saccharomyces principally in wine.
In other beverages like beer, fruits wines, and distilled
beverages the positive contribution of non-Saccharomyces is
also discussed. Another interesting and new application of
non-Saccharomyces is as a tool for reducing the potential
ethanol content in wines.
Address
Centro de Investigación y Asistencia en Tecnologı́a y Diseño del Estado
de Jalisco, A.C., Camino Arenero 1227, El Bajio, Zapopan, Jalisco C.P.
45019, Mexico
Corresponding author: Gschaedler, Anne (agschaedler@ciatej.mx)
Current Opinion in Food Science 2017, 13:73–77
This review comes from a themed issue on Food bioprocessing
Edited by Rosane Freitas Schwan
For a complete overview see the Issue and the Editorial
Available online 16th February 2017
http://dx.doi.org/10.1016/j.cofs.2017.02.004
2214-7993/ã 2017 Published by Elsevier Ltd.
Introduction
The elaboration of alcoholic beverages (distilled or not)
involves a fermentation process that allows the conversion
of fermentable sugars into ethanol as well as a high
number of flavor-active compounds that give each of
the beverages-specific characteristics. The history of
the human being has always been accompanied in almost
all the civilizations with the elaboration of alcoholic
beverages. For centuries, the fermentation was carried
out spontaneously with a mixture of native microorgan-
isms including a large number of non-conventional yeasts
[1]. Since the 70s the concept of controlled fermenta-
tions inoculated with a specific selected yeast was intro-
duced and for decades the only species of yeast that
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was intentionally inoculated to make an alcoholic fermen-
tation was a specific strain of Saccharomyces cerevisiae and
consequently the most studied specie. However, in the 80s,
works were published demonstrating the positive effect of
some non-Saccharomyces mainly improving some character-
istics, such as complexity, texture and integration of flavors
in spontaneous fermentations. After three decades, the
non-Saccharomyces yeasts went from being an undesirable
microorganism associated with spoiled wines to yeasts that
can bring interesting characteristics [2]. Recently 37 yeast
species were mentioned as the main non-Saccharomyces
involved in wine, beer, tequila, mezcal and cachaça spon-
taneous fermentations. They belong to 20 genera men-
tioned below: Aureobasidium, Brettanomyces, Candida,
Clavispora, Cryptococcus, Debaryomyces, Hanseniaspora,
Hanseluna, Issatchenkia, Kluyveromyces, Lachancea, Metsch-
nikowia, Meyerozyma, Pichia, Rhodotorula, Saccharomy-
codes, Starmerella, Wickerhamomyces, Torulaspora and
Zygosaccharomyces [1].
Next, a summary of the information generated recently
focused on the characterization of spontaneous fermenta-
tions will be presented, and on the other hand, works in
which a selected non-conventional yeast is inoculated to
generate specific characteristics in the final product.
Non-Saccharomyces yeasts in grape juice
fermentation
There are numerous studies of the diversity of yeasts
present in the spontaneous fermentations of wines. Jolly
et al. [2] mention that within the nearly 1500 reported
species of yeasts more than 40 species have been isolated
from grape must in spontaneous fermentations. Reported
genera are Citeromyces, Debaryomyces, Dekkera, Hansenias-
pora, Lachancea, Metschnikowia, Meyerozyma, Millerozyma,
Pichia, Saccharomycodes, Starmerela, Torulaspora, Wickerha-
momyces, Zygoascus, Zygosaccharomyces. The great majority
of these strains came directly from the grape berry sur-
faces, cellar equipment surfaces and cellar environment.
Generally during the spontaneous fermentation process, a
dynamics of different yeast species occur: the non-Sac-
charomyces yeasts (mainly belonging to Hanseniaspora/
Kloeckera, Candida, Pichia, Zygosaccharomyces, Schizosac-
charomyces, Torulaspora, Kluyveromyces and Metschnikowia
genera) dominate the beginning of the alcoholic fermen-
tation and then they are replaced by Saccharomyces cere-
visiae that complete sugars conversion into ethanol [2–4].
Normally the non-Saccharomyces die soon after the begin-
ning of the fermentation and the initial belief was that
these strains die due to the rising concentration of ethanol
Current Opinion in Food Science 2017, 13:73–77
74 Food bioprocessing
and SO2 added. Recent investigations demonstrate that
the reasons of the disappearance are more complex and
different factors are involved: slow growth, inhibition by
the combined effects of SO2, high ethanol concentration,
low pH and oxygen deficiency [5,6] and certainly is strain
dependent.
In the thematic of the study of indigenous microbiota
recent investigations allowed to isolate non-conventional
yeasts associated with different types of vineyards looking
for new autochthonous strains with specific and interest-
ing characteristics [7–9]. Another important field of
research in this thematic are the methods of characteri-
zation and mainly the culture-independent methods
based on the nucleic acid sequences used to study the
microbial diversity of grapes and wines [10]. These tech-
niques open up new study perspectives, allowing the
massive analysis of samples. A recent study used the
high-throughput barcoding sequencing to analyze the
microbial diversity and dynamics during wine fermenta-
tion of Grenache grape variety evidencing a great varia-
tion of the relative abundance of the yeasts showing a
dominance of the genera Hanseniaspora at the beginning
of the fermentation accompanied by Issatchenkia and
Candida. Up to 10 days of fermentation there is a decrease
in the populations of non-Saccharomyces (except Candida)
and a much larger proportion of Saccharomyces.
In this specific area of investigation of autochthonous
microbiota, there is still an important field of research
since it has been demonstrated the great differences
existing between strains within the same species. An
example illustrating this last point is the phenotypic
characterization of yeasts strains (including 29 non-con-
ventional yeasts) to stresses inherent to wine fermenta-
tion in warm climates that allowed selecting some strains
with interesting biological characteristics resistant to
higher temperatures [11].
Regarding the process of elaboration, the non-conven-
tional yeasts are now use for developing specific inocula to
enhance sensorial complexity and aroma profiles of the
wines. Mixed starters designed to improve primary and
secondary wine aroma, using to types of inoculation (co-
inoculation or sequential) are tested. The species studied
in mixed starters with S. cerevisiae are Candida zemplinina
(Starmerella bacillaris), Debaryomyces pseudopolymorphus, D.
vanrijiae, Hanseniaspora guillermondii, H. uvarum, H. vinae,
Issatchenkia orientalis, Kluyveromyces gamospora, Lachancea
thermotolerans, Metschnikowia pulcherrima, Picha fermentans,
P. kluyveri, Schizosaccharomyces pombe, Torulaspora del-
brueckii, Wickerhamomyces anomalus, Williopsis. saturnus,
Zygosaccharomyces bailii [12]. In the investigations carried
out, the predominant strain studied is T. delbrueckii [13–
15] including an application in sparkling wine [16]. There
are also few trials of pure fermentation with non-Saccha-
romyces; one of them developed a killer strain of T.
Current Opinion in Food Science 2017, 13:73–77
delbrueckii. Wines obtained with this commercial strain
had dried food/pastry aromas but lower intensities in
fresh-fruit aromas comparing with a S. cerevisiae wine [17].
Another interesting and new application of non-Saccharo-
myces is as a tool for reducing the potential ethanol content
in wines. The global warming induces changes in the
chemical composition of the grapes mainly in an increase
in the sugar contents and a decrease in acids, anthocya-
nins and methoxypyrazine. These changes in the compo-
sition of the grapes have as consequence an increase of
the concentration of ethanol in the wine and alter mouth-
feel and flavor [18]. Röcker et al. [19] selected five
different non-Saccharomyces and performed fermentations
in different conditions with a sequential inoculation
(using always a selected commercial S. cerevisae in a
second phase of fermentation). The most promising strain
was the Metschnikovia pulcherrima which allows to reduce
the alcohol concentration. In another work the most
promising strain was a Candida sake and the authors
defined an interesting integrative parameter, ‘Efficacy’
which reflect the efficacy for alcohol level reduction [20].
A recent review presents the research on non-Saccharo-
myces aiming to produce wines with lower alcohol content.
The important conclusion is that the feasibility at indus-
trial level will require an improved understanding of the
metabolism of these non-conventional yeasts [21].
Non-Saccharomyces yeasts in fruit juice
fermentation
In many subtropical countries wine obtained with differ-
ent fruits has recently become more common. An exam-
ple is the production of Pineapple (Ananas comosus L.
Merr) wine in Angola. The characterization of native flora
permits the detection of different non-Saccharomyces:
Hanseniaspora guillermondi, H. uvarum, H. opuntiae, Pichia
anomala, Meyerozyma guillermondi, Wickerhamomyces
anomalus. Pure fermentations at laboratory scale demon-
strate the contribution of the non-Saccharomyces to the
aroma profile [22].
In Asia, another interesting wine is obtain by fermenta-
tion of a famous tropical fruit, the durian (Durio zibethinus
Murr). Normally the fermentation is carry out by S.
cerevisiae but the wine produced normally lacks flavor
complexity. Therefore, in recent investigations different
commercial non-Saccharomyces have been test: Torulas-
pora delbrueckii, Metschnikowia pulcherrima, Pichia kluyveri
and Kluyveromyces thermotolerans in pure or co-inoculated
fermentations. Like in the classic wine, an interesting
result is obtain with T. delbrueckii [23–25]. In Lychee wine
a sequential fermentation with T. delbrueckii first and then
S. cerevisiae in a second phase permit the production of a
wine with an acceptable ethanol concentration and a
complex aromatic profile [26].
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In Europe and in minor dimension in America the fer-
mentation of apple juice to obtain cider belongs to this
category. The indigenous yeast profile of traditional cider
fermentation is similar to that observed in spontaneous
wine fermentation. Few researches have been published
on the specific use of a non-Saccharomyces yeast for the
fermentation of cider. Nevertheless a recent work pro-
poses a controlled co-fermentation with S. cerevisiae and
Williopsis saturnus. Which is characterized by a high
production of fruity acetate esters. The co-fermentation
carried out with these two strains resulted in a more
complex volatile profile [27].
Non-Saccharomyces yeasts in brewing
In the case of brewing the raw material is starch coming
mostly from barley although others grains such as wheat,
rice oats, rye and corn can also be used. During decades,
beers were classified into two major types, ale and lager,
according to the yeast used and fermentations conditions.
In the two types of beers, different strains of Saccharomyces
are used. S. cerevisiae for the ale type and mainly
S. pastorianus for lager. Nowadays with the development
and success of craft beers the brewers are looking for new
alternatives to ferment and logically start to use non-
conventional yeast in order to impact on aroma and flavor
and generate differentiated products [28]. An exception
to the recent interest in non-Saccharomyces yeasts is the
use of Dekkera/Brettanomyces for the production of sour
beers. In particular, lambics are produced under a com-
plex succession of spontaneous fermentations with an
important participation of Dekkera/Brettanomyces in the
acidification-maturation stage [29]. Nowadays in the case
of craft beers the fermentation of wort is carried out by
different non-Saccharomyces pure starter culture. Each
strain is selected according to the characteristics sought
in the final product. The principal species studied
are: Torulaspora delbrueckii, Candida shehatae, Candida
tropicalis, Saccharomyces ludwigii, Zygosaccharomyces rouxii,
Pichia kluyveri, Cyberlindnera saturnus, Hanseniaspora
uvarum, Wickerhamomyces subpelliculosus, Wickerhamomyces
anomalus. The principal applications of these yeasts are
mainly bioflavoring, the generation of low calorie beer
and low alcohol beer [28,30,31,32]. Recently Candida
zemplinina was also evaluated for craft beer fermentation
with interesting results [33]. Finally is important to
mention that in Africa different types of beers are elabo-
rated and one of them is the ‘Tchoukoutou’ a Beninese
sorghum beer. The microbial ecology of the starter used
to produce this beer pointed out the presence of different
species of non-Saccharomyces: Pichia kudriavzevii, Candida
ethanolica, Debaryomyces hansenii [34].
Non-Saccharomyces in the production of
distillates
Like in other fermented beverages yeasts and the fer-
mentation stage play an essential role in the final sensory
profile of distilled spirits. In spite of the importance of
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Non-conventional yeasts in alcoholic beverages Gschaedler 75
fermentation there has been limited recent research on
the roles of non-Saccharomyces yeasts in this process. Some
interesting results have been generated in the case of
three distillates produced in America: tequila, mezcal and
cachaça. Tequila and mezcal are typical Mexican spirits
obtained from the distillation of the fermented juice of
different agave species cooked previously. In the case of
tequila only one agave specie is used, Agave tequilana
Weber var. azul and is always cultivated. Regarding the
development of fermentation, industry is divided into two
groups, the first still use ancestral spontaneous fermenta-
tions (the vast majority of small distilleries), and the latter,
mainly large industrial companies employ selected spe-
cific yeasts (S. cerevisiae). In the case of spontaneous
fermentations there is only one comprehensive study
of microorganisms present in the fermentation [35]. At
the early fermentation (up to 2 days) the samples contains
a rich mixture of yeast species: Candida intermedia,
Candida krusei, Brettanomyces anomalus, Kluyveromyces
marxianus, Pichia membranifaciens, Torulaspora delbrueckii,
Hanseniaspora spp., Candida spp., Saccharomyces cerevisisae,
Pichia anomala, Saccharomycodes ludwigii, Zygosaccharo-
myces bailii, Candida milleri, Brettanomyces bruxellensis,
Zygosaccharomyces rouxii. During fermentation, the popu-
lations of non-Saccharomyces yeasts tended to decrease,
with S. cerevisiae becoming dominant by the end of
fermentation. It is interesting to emphasize that in
the final phase of fermentation (more than 5 days of
process) as well as S. cerevisiae another four species were
detected in the fermentation: Pichia membranaefaciens,
Zygosaccharomyces bailii, Candida milleri and Brettanomyces
bruxellensis. This result shows that non-Saccharomyces are
able to survive throughout fermentation, contrary to what
is observed commonly in the fermentation of grapes.
Recent work has focused mainly on the use of non-
Saccharomyces yeasts in co-culture with Saccharomyces or
alone. The first work compared the fermenting behavior
in agave juice of five different yeasts at laboratory level
including two strains of Kloeckera (Hanseniaspora) [36].
The study highlights major differences between the three
S. cerevisiae especially from the point of view of produc-
tion of volatile compounds, in this case Kloeckera strains
did not achieved the alcoholic fermentation however
produce large amounts of esters. Another study explores
the possibility of using mixed cultures Saccharomyces/
Kloeckera with interesting conclusions especially on the
contribution of non-Saccharomyces yeasts on the aromatic
profile [37]. Kluyveromyces marxianus and Pichia kluyveri
are two interesting candidates to carry out the fer-
mentation of tequila. A study conducted at laboratory
level demonstrates that these two strains are able to carry
out alcoholic fermentation almost like a S. cerevisiae and
also produce a very interesting specific flavor profile with
high concentrations of esters [38]. These two strains were
tested at industrial level with very promising results [39].
Another strain of K. marxianus was tested and similar
results were obtained [40]. The big challenge for these
Current Opinion in Food Science 2017, 13:73–77
76 Food bioprocessing
non-Saccharomyces strains is to find the ideal conservation
conditions to obtain active dry yeast or lyophilized strains.
With classic conservation processes employed with S.
cerevisiae is not possible to generate a culture of these
non-Saccharomyces preserved with high viability.
In the case of mezcal the general process of production
of the spirit is similar to tequila, the big difference is the
specie of agave used as raw material. About 40 species of
agave (some of them wild species) are used, depending
of the region of production and availability. Native
strains of T. delbrueckii and K. marxianus were tested
as single or mixed starter (with a native strain of S.
cerevisiae) for mezcal fermentation. Like in the case of
tequila non-Saccharomyces demonstrate to have a positive
influence on the final sensory profile of the distillate
[41].
Cachaça, a spirit produced in Brazil is made from the fresh
sugar cane juice fermented and distillated. Its production
is in both artisanal and industrial scales, in sequential,
short duration fermentation cycles lasting about 18–
30 hours with successive filling of fresh sugarcane. The
principal fermentative strains are natives Saccharomyces
cerevisiae but recent studies investigate the use of non-
Saccharomyces in order to improve the aroma profile. For
example, 20 non-Saccharomyces yeast strains were evalu-
ated during co-inoculation with S. cerevisiae, but only two
yeast strains W. anomala and P. caribbica showed
improved fermentation kinetics, and in the last case
the fermentation showed increased concentrations of
desirable volatile compounds [42]. Recently a study of
native yeasts demonstrated that acting on a cluster con-
dition these yeasts may greatly contribute to the flavor
complexity of artisanal cachaça [43].
Conclusion and perspectives
The abundance of recent studies dealing with the use of
non-Saccharomyces demonstrate that is an important topic
in the elaboration of alcoholic beverages. The first and
clear contribution of non-Saccharomyces in all the bev-
erages is the positive impact on the sensory profiles by
increasing the complexity and organoleptic richness. The
majority of non-Saccharomyces are used in co-fermenta-
tion, so in this case the detailed study of the interaction
between Saccharomyces and non-Saccharomyces is a com-
plex key study in order to understand the relationship
between the strains including the molecular level [44,45].
Another important point is the strain specificity, so new
methods for high throughput screening has to be devel-
oped in order to screen a large number of strains for
selecting a specific trait [46]. The new applications of
non-Saccharomyces to generate low alcoholic wines or low
calories beers open interesting perspectives for these
strains. Finally in the case of some strains in order to
obtain practical commercial non-Saccharomyces yeasts the
principal challenge is to find the ideal conservation
Current Opinion in Food Science 2017, 13:73–77
conditions to obtain active dry yeast or lyophilized
strains.
Acknowledgement
Author thanks for financial support the Mexican Council of Science and
Technology (CONACYT), project # 252465.
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This article presents an interesting exploration of yeast biodiversity in
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