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Yeasts and molds in fermented food production: an
ancient bioprocess
Marina Venturini Copetti
The fermentation of carbon sources by fungi leading to the
production of foods and beverages represents one of the
oldest and most economically significant of all biotechnologies.
Also, some traditional fermented foods require a ripening
period to develop their best sensorial characteristics. On
ancient times, fermentation was carried out spontaneously.
With scientific development, single strains were selected and
adapted for use as starters in some large-scale food producing.
Currently an apparent revival of interest in complex
fermentation is occurring, as well in the use of mixed strains
wondering production of food and beverages with more diverse
and peculiar flavors impressed by a more complex micro-
organisms interaction and the associated health functionalities.
Address
Federal University of Santa Maria, Rural Sciences Center, Department of
Food Technology and Science, 1000 Roraima Avenue, Santa Maria, RS,
97105-900, Brazil
Corresponding author: Venturini Copetti, Marina (mvc@smail.ufsm.br)
Current Opinion in Food Science 2019, 25:57–61
This review comes from a themed issue on Food bioprocessing
Edited by Cristiano Ragagnin de Menezes
For a complete overview see the Issue and the Editorial
Available online 28th February 2019
https://doi.org/10.1016/j.cofs.2019.02.014
2214-7993/ã 2019 Elsevier Ltd. All rights reserved.
Introduction
Fungi, both yeast and filamentous, by virtue of their
metabolic versatility have important role in food industry
processes. The role of yeasts on the production of fer-
mented foods is well established. A study revealed that
industrial yeasts are genetically and phenotypically dis-
tinct from wild strains and stem from a limited set of
ancestral strains adapted to man-made environments
(clades of: Asian strains such as sake yeasts, one mostly
containing wine yeasts, a mixed clade containing bread
and other yeasts, and two separate families of beer yeasts)
[1]. In contrast, foods showing presence of molds are, in
general, considered inadequate for consumption, but
selected species of molds are useful for the manufacture
of certain foods or ingredients (Figure 1).
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Fungal biota plays an important role in the development
of peculiar sensorial characteristics of some traditional
fermented foods, participating on metabolic processes
through production of many enzymes and metabolites
during the ripening period [2]. These biotas sometimes
are diverse and only spontaneously inoculated in the
products, depending on the maturation environment.
In other cases, fungal strains can be selected and used
as starter, accelerating the fermentation and standardizing
the food produced [3].
This review will focus on the use of yeasts and molds as
bioprocessing agents for the manufacture of foods
through fermentation, highlighting ancient processes cur-
rently developed at industrial level.
Alcoholic beverages
Saccharomyces sp. is the major yeast genus involved in the
production of alcoholic beverages. Probably the best
understood processes are wine and beer [4]. Besides
the well-established use of Saccharomyces as starters,
non-Saccharomyces yeasts used in the co-fermentation of
beverages impact on the sensory profiles by increasing the
complexity and organoleptic richness [5].
Wine
Winemaking is an ancient biotechnology, and the basic
activities of modern wineries are fundamentally the same
as those undertaken traditionally: Extraction of sugars
from grapes and their fermentation by yeasts to produce
an alcoholic beverage. It can be carried out spontaneously
throughout endogenous micro-organisms from grape and
winery surfaces or using exogenous micro-organisms from
starter culture of single or mixed yeast strains [6–8,9].
Modern, large-scale wineries generally use specially
selected starter cultures of Saccharomyces cerevisiae [10].
Spontaneous fermentation of grape juice is driven by a
complex community of microorganisms. A yeast succes-
sion from Hanseniaspora to Saccharomyces has been estab-
lished, due differences on species sensitivity to the
produced ethanol, and so, population driven on selectiv-
ity [7,9,11,12].
The technological diffusion of the use of selected strains
of S. cerevisiae simplified the fermentation process,
although, more recently there is an increasing interest
toward controlled mixed fermentations using more than
one selected yeast strain [6–8]. The micro-organisms
selected and even the sequence and time of inoculation
Current Opinion in Food Science 2019, 25:57–61
58 Food bioprocessing

Figure 1

Alcoholic
beverages
Pigments Asian Foods

F
O F
O O
Organic acids Bread
D O
I D
N P
G R
R
E
D
I
Oils and Fatty
acids Fungi Cheese
ripening
O
D
U
C
E T
N I
T O
S Mycoprotein Cocoa N

Flavoring Cured meats

Enzymes

Current Opinion in Food Science

Fungal importance in some industrial food and ingredients production bioprocesses.

can exert strong influence in the sensorial profile of wines
[13–15].
The use of more complex and controlled inoculum (co-
inoculation or sequential inoculation with non-Saccharo-
myces and S. cerevisiae) [13,16–19] and the revival of
spontaneous fermentation, labeled as ‘wild ferments’
[15], are recent trends in wine market. These kind of
products meet the consumers claim for more natural and/
or peculiar food; the diversity of microbial population of
natural (or spontaneous) and sequential fermentation can
offer the desired complexity and diversification to these
wines [20].
Beer
Beer brewing is one of most ancient human pursuits,
representing one of the first forays in biotechnology.
Differently from other alcoholic beverages which are
originated from fermentation of simple fruit sugars, beer
is obtained through fermentation of starch derivative
sources. The starch must be first solubilized in the wort
using enzymes to generate more simple sugars, ferment-
able into ethyl alcohol, carbon dioxide, and other fermen-
tation by-products, including higher alcohols and esters,
through the action of brewer’s yeasts, mostly Saccharomy-
ces sp. [21].
Most of the peculiar flavor and aroma of any beer are
determined by the yeast strain and fermentation condi-
tions [1]. The majority of beer consumed around the
world is the bottom-fermented lager beer and the top-
fermented ale beer, fermented with strains of Saccharo-
myces pastorianus [Saccharomyces uvarum (carlsbergensis)] at
7–10  C and S. cerevisiae at 18–25 C, respectively [21].
In recent years, with the development and success of craft
beers [22,23,24], the brewers are also starting to use non-
conventional yeast in order to impact on aroma and flavor
and generate differentiated products [25–29]. Brettano-
myces and other non-conventional yeasts would be highly
attractive for the development of specialty beers, since
yield natural flavors out of scope of conventional Saccha-
romyces species, resulting in products with novel attractive
flavor compositions [27,29,30].
Craft beers are also unpasteurized and unfiltered, so, have
the potential for delivering probiotic yeasts. Inclusion of
S. cerevisiae var. boulardii in mixed starter increased the
antioxidant activity and polyphenols content in beer
without negative influence in beer aroma [31].
Asian foods
Many fermented Asian foods employ fungi for their
manufacture [32]. The molds Aspergillus oryzae and

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Aspergillus sojae play a pivotal role in some Asian food
processing, through their use as the starter for prepara-
tion of koji, a source of fungal enzymes (both amylolytic,
lipolytic and proteolytic) necessary for assimilation of
nutrients by other microorganisms in subsequent fer-
mentations [33]. The koji also contributes to color,
flavor, and aroma, which are important for the overall
character of the fermented products, such as soy sauce
and sake [33]. In recent times, efforts have been made
to explore the health-promoting effects of Koji and its
derived products consumption, including their antioxi-
dant, antimutagenic, antitumoral, and stress regulation
activities, capability on elimination of gluten allergen
and lowering cholesterol [33].
Sake or rice wine
Sake is a Japanese alcoholic beverage made from rice and
manufactured by means of a mixed fermentation of molds
and yeasts in two stages [34]. At the first stage of
fermentation, rice is steamed, cooled down, inoculated
with A. oryzae, and kept at 30 C for two days for koji
production. Then koji is mixed with steamed rice, water
and inoculated with S. cerevisiae for alcoholic fermenta-
tion. Approximately two months is needed to complete all
the fermentation steps and obtaining the sake [32].
The development of high-class sake based on acidity has
begun in Japan in more recent years; thus, the sake
industry has entered a new era, and the use of diverse
yeasts is becoming relevant and, like some wines, can
drive on certification of geographical indication [35].
Shoyu or soy sauce
Shoyu or soy sauce is a traditional Japanese liquid
seasoning also manufactured by a two-stage fermenta-
tion process in which a consortium of filamentous fungi
and yeasts species occurs [34]. The filamentous fungi
A. oryzae or A. sojae are grown on a mixture of steamed
soybeans, cracked wheat and wheat bran to koji pro-
duction. The koji is then mixed with water and salt,
following 6–12 months fermentation. The fermented
product is then pressed to produce the fermented juice,
shoyu [32,33].
The flavor of soy sauce is chiefly determined by protei-
nases, amylases, and other enzymes synthesized by the
mold species used. In contrast, yeasts such as Hansenula
spp. and Zygosaccharomyces rouxii have an important role
on the development of a tangy taste, and bacteria such as
Tetragonococcus halophila on acidic aroma and clear
appearance [34].
Soy sauce contains, in addition to taste and aroma com-
pounds, certain bioactive components that have anti-
hypertensive, anti-carcinogenic, anti-microbial, anti-oxi-
dative and anti-platelet activities [33,36].
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Yeasts and molds in food production Venturini Copetti 59
Bread
Bread is a food prepared by baking fermented dough,
essentially made from flour, water, and leavening agent
[37]. The fermentation occurs through the metabolism
of complex fermentable carbohydrates (such as sucrose,
maltose, and starch) from the dough into glucose and
fructose by the yeast S. cerevisiae. During baking, protein
denaturation leads to a hardening of the glutinic struc-
ture, determining the conservation of bread shape and
volume [37].
Besides the action as leavening agent, S. cerevisiae impress
subtle flavors, mainly through the synthesis of organic
acids and volatile products. In contrast, the association of
S. cerevisiae to other species of yeasts and lactobacilli can
lead to production of a more acid product and with
peculiar flavor, the sourdough.
Sourdough bread
Sourdough is a mixture of cereal flour and water that is
spontaneously fermented by consortia covering hetero-
fermentative lactic acid bacteria and yeasts [38].
The use of sourdough in baking is an ancient technology
that is currently undergoing a revival of interest. Bread
production relied on the use of sourdough as leavening
agent for most of human history; the use of baker’s yeast
dates back less than 150 years [39]. The sourdough
technology remained in European countries mainly
because rye flour requires acidification to achieve an
optimal bread quality. Some well-known sourdough
breads are the sweet breads from Mediterranean coun-
tries, such as panettone and colomba [40], as well the San
Francisco bread.
Sourdough promotes an improvement in dough structure,
aroma, palatability, nutritional value and prolonged shelf-
life, mainly due its stability against fungal spoilage [41].
Besides, in more recent years, functional/nutritional fea-
tures of sourdough fermentation (such as potential to
lower glycemic index, increase mineral bioavailability
and decrease the gluten content, among others) have
been highlighted [42].
Mold-ripened cheeses
Mold-ripened cheeses are of two major types: surface
mold-ripened cheeses (such as Camembert and Brie,
generally ripened by Penicillium camemberti, forming a
velvety white rind) and internal mold-ripened cheeses
(or blueveined cheeses, ripened by Penicillium roqueforti,
as Danablu, Roquefort, Stilton, and Gorgonzola) [43].
Basically, the making and ripening of mold-ripened
cheeses involve lactic acid bacteria as primary starters,
besides yeasts and molds. The fungal biota is notably
involved in the consumption of lactic acid and lactate
produced by lactic acid bacteria, raising the pH, as well as
Current Opinion in Food Science 2019, 25:57–61
60 Food bioprocessing
in proteolysis and lipolysis, fundamental processes for
ripening. The type and concentration of acids, primary
and secondary alcohols, carbonyl compounds, esters, and
hydrocarbons determine the peculiar aroma/flavor of
these cheeses [34]. Interactions between microorgan-
isms and their environmental factors and between micro-
organisms themselves are determinant in controlling rip-
ening and sensorial properties of cheeses [43].
Cocoa
Cocoa is the main raw material for chocolate production.
The primary objectives of cocoa fermentation are the
removal of mucilage and the development of chocolate
flavor precursors. The sugar rich bean mucilage is fer-
mented by yeasts; the violet cotyledons of beans become
brown during the week-long fermentation, are dried and
roasted [44].
Microbial populations involved in spontaneous cocoa
fermentations demonstrate yeast–bacterial interactions,
with well-defined and inter-dependent role. These fer-
mentations are dominated by yeasts and both lactic acid
and acetic acid bacteria, in a definite succession. Yeasts
(most Saccharomyces sp. and Candida sp.) proliferate early,
leading to production of ethanol, secretion of pectinolytic
enzymes and metabolization of citric acid, raising the pH.
This is followed by a phase in which bacteria predomi-
nate; principally the microaerophilic lactic-acid bacteria
and the ethanol oxidative acetic-acid bacteria [45].
Cured meats
In the twenty-first century, sausage manufacturing is still
made from the ancient recipe: comminuted meat, sea-
soned with salt and spices, stuffed into casings, fermen-
ted, and ripened. The industrial development led to the
use of starters but some manufacturers are still producing
artisanal fermented sausages with spontaneous fermenta-
tion [46].
The ecology of fermented sausages is complex and
includes different bacteria, yeasts, and molds, with a
specific microbiota that is typical of the region or area
where they are produced [47]. During maturation period,
meat enzymes, assisted mostly by lactic acid bacteria,
contribute to the fermentation, improving taste, and
acidifying the product. Fungi have also an important role,
because they produce many lipolytic and proteolytic
enzymes involved in the development of specific flavors
in dry meats. Debaryomyces hansenii is one of the predomi-
nant yeasts in meat products and Penicillium species are
the most common filamentous fungi colonizing both
industrially and manufactured dry-fermented meat pro-
ducts [47–50].
Conclusion
This review summarized some of the most ancient, and
industrially adapted, bioprocessing of foods through
Current Opinion in Food Science 2019, 25:57–61
fungal development. Even then starters are applied in
modern food processing to improve fermentations and
standardize the product, spontaneous fermentations
remain and, in a certain extent, are showing a revival
of interest due to the diverse and peculiar flavors
impressed by different micro-organisms interacting dur-
ing the food fermentation, as well due some health
benefits associated.
Conflict of interest statement
Nothing declared.
Acknowledgement
To Conselho Nacional de Desenvolvimento Cientı́fico e Tecnológico -
CNPq for a research grant to M.V.C. (Process 309691/2015-0).
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