Critical Reviews in Food Science and Nutrition

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Sourdough production: fermentation strategies,

microbial ecology, and use of non-flour ingredients

Luc De Vuyst, Andrea Comasio & Simon Van Kerrebroeck

To cite this article: Luc De Vuyst, Andrea Comasio & Simon Van Kerrebroeck (2021): Sourdough
production: fermentation strategies, microbial ecology, and use of non-flour ingredients, Critical
Reviews in Food Science and Nutrition, DOI: 10.1080/10408398.2021.1976100

To link to this article: https://doi.org/10.1080/10408398.2021.1976100

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Critical Reviews in Food Science and Nutrition
https://doi.org/10.1080/10408398.2021.1976100

Review

Sourdough production: fermentation strategies, microbial ecology, and use

of non-flour ingredients
Luc De Vuyst , Andrea Comasio and Simon Van Kerrebroeck
Research Group of Industrial Microbiology and Food Biotechnology (IMDO), Faculty of Sciences and Bioengineering Sciences, Vrije
Universiteit Brussel (VUB), Brussels, Belgium

ABSTRACT
Sourdough production is an ancient method to ferment flour from cereals for the manufacturing KEYWORDS
of baked goods. This review deals with the state-of-the-art of current fermentation strategies for Acetic acid bacteria;
sourdough production and the microbial ecology of mature sourdoughs, with a particular focus bread;
on the use of non-flour ingredients. Flour fermentation processes for sourdough production are lactic acid bacteria;
typically carried out by heterogeneous communities of lactic acid bacteria and yeasts. Acetic acid sourdough;
bacteria may also occur, although their presence and role in sourdough production can be criticized. yeasts
Based on the inoculum used, sourdough productions can be distinguished in fermentation processes
using backslopping procedures, originating from a spontaneously fermented flour-water mixture
(Type 1), starter culture-initiated fermentation processes (Type 2), and starter culture-initiated
fermentation processes that are followed by backslopping (Type 3). In traditional recipes for the
initiation and/or propagation of Type 1 sourdough productions, non-flour ingredients are often
added to the flour-water mixture. These ingredients may be the source of an additional microbial
inoculum and/or serve as (co-)substrates for fermentation. An example of the former is the addition
of yoghurt; an example of the latter is the use of fruit juices. The survival of microorganisms
transferred from the ingredients to the fermenting flour-water mixture depends on the
competitiveness toward particular strains of the microbial species present under the harsh conditions
of the sourdough ecosystem. Their survival and growth is also determined by the presence of the
appropriate substrates, whether or not carried over by the ingredients added.

Introduction
Fermentation of food raw materials is an old biotechnological
process that is applied to prolong the shelf-life of agricultural
and animal husbandry products (Leroy and De Vuyst 2004;
Bourdichon et al. 2012; Wolfe and Dutton 2015; Tamang,
Watanabe, and Holzapfel 2016; Marco et al. 2017, 2021; Rezac
et al. 2018). Diverse nutrients of these raw materials are used
as substrates by bacteria and fungi and, hence, these micro-
organisms convert these raw materials into fermented food
products with desirable, organoleptic, and durable properties.
Agricultural raw materials that are frequently used for fer-
mented food productions are cereals, in particular in the case
of beer, yeast-leavened bread, and sourdough productions.
The preparation of bread based on sourdough is a typical
example of an old and traditional fermentation process
(Cappelle et al. 2013; Gobbetti et al. 2019). In fact, it was
the first way of producing bread through natural leavening,
dating back thousands of years, making use of endogenous
lactic acid bacteria (LAB) and/or yeasts unconsciously, as
these microorganisms are present on the raw materials, until
commercial yeast (first brewer’s yeast starting in the Middle
Ages and later on baker’s yeast starting in the 19th century)
was used for leavening. For sourdough production, a mixture
of cereal flour(s) (fractions) and water is prepared that is
either fermented spontaneously and further subjected to a
backslopping process in the case of traditional sourdoughs,
or fermented in a single step for a couple of hours to several
days in the case of industrial sourdoughs (De Vuyst, Van
Kerrebroeck, and Leroy 2017). The sourdoughs obtained are
used as functional bread dough ingredients for the production
of leavened breads or to substitute baker’s yeast to produce
traditional sourdough breads. Nowadays, the use of sour-
doughs as source of flavor, nutritional, and health-promoting
compounds for breadmaking is increasingly popular (Poutanen,
Flander, and Katina 2009; Gobbetti et al. 2014, 2019; Poutanen,
Sozer, and Della Valle 2014; Rizzello et al. 2019).
This review deals with the state-of-the-art of current
fermentation strategies for sourdough production, seen from
the point of view of both the bakery industry and the
non-bakery food fermentation industry in general, and the
microbial ecology of mature sourdoughs, with a particular
focus on the use of non-flour ingredients.

CONTACT Luc De Vuyst luc.de.vuyst@vub.be

*These authors contributed equally to this study.
© 2021 The Author(s). Published with license by Taylor & Francis Group, LLC.
This is an Open Access article distributed under the terms of the Creative Commons Attribution-NonCommercial-NoDerivatives License (http://creativecommons.org/licenses/by-nc-nd/4.0/),
which permits non-commercial re-use, distribution, and reproduction in any medium, provided the original work is properly cited, and is not altered, transformed, or built upon in any way.
2 L. DE VUYST ET AL.
Flours used for sourdough production
Mainly flour made from kernels of cereals belonging to the
Poaceae is used for sourdough production. In particular,
flour from soft wheat (Triticum aestivum L.) and/or hard
wheat (Triticum aestivum L. and Triticum durum Desf.) is
used. A dry milling process on the wheat kernels yields the
flour, the baking quality of which is determined by the
gluten network that is formed during dough preparation.
However, the baking quality of cereal flour is influenced by
several factors.
Wheat flour is mainly composed of carbohydrates, in
particular starch (70–75 %, m/m), next to free saccharides
(1–2 %), and crude fiber (2–3 %) (Koehler and Wieser
2013). The starch absorbs water, is incorporated into the
gas-holding gluten network of the dough, whose viscosity
it contributes to, and serves as primary energy source (after
enzymatic hydrolysis) for fermentation by yeasts. The dietary
fiber mainly consists of fructans (1–2 %) and arabinoxylan
pentosans (1.5–2.5 %). Fructans contribute to the yeast activ-
ity and are mainly of nutritional importance. Arabinoxylans
contribute to the dough texture by enhancing the viscoelas-
ticity of the dough through water absorption and better gas
retention of the dough. The latter is of utmost importance,
as LAB metabolism (proteolytic and oxidoreductase activi-
ties) may partially destroy the gluten network during fer-
mentation. Further, wheat flour contains proteins (8–15 %),
in particular gliadins and glutenins, that provide the dough
with the actual viscoelasticity and gas retention capacity
through water-binding and the formation of disulfide bridges
via thiol/disulfide exchange reactions in the glutenin mac-
ropolymer of the gluten network during the mixing and
kneading steps of the dough preparation. LAB acidification
of the dough enables an increased bread loaf volume and
crumb softness, thanks to a higher solubility of the gluten
in an acidic environment, because of a positive net charge
of these proteins, as well as a delayed staling due to starch
retrogradation, because of the promotion of the swelling of
the starch granules. Besides the storage proteins mentioned
above, several enzymes are present in wheat flour, such as
α-amylase and β-amylase, proteases and oxidizing enzymes
(lipoxygenase, hydroperoxide lyase, and polyphenol oxidase),
which influence the baking quality of the flour too. In addi-
tion, lipids (1–2 %) are present in wheat flour. In particular,
the starch non-associated glycolipids have a positive influ-
ence on the baking quality of the flour by enhancing the
knead ability and gas retention capacity of the dough, pro-
longing the oven spring, increasing the bread volume and
crumb resilience, and retarding the starch retrogradation of
the bread crumb. Finally, wheat flour contains several other
compounds, such as phenolics, minerals, and vitamins, that
contribute to the organoleptic and/or nutritional quality
of bread.
Compared to wheat flour, flour from rye (Secale cereale
L.) contains lower amounts of starch (on average 65 %),
more fiber, in particular more arabinoxylan pentosans (6–8
%), approximately the same amount of proteins (on average
12 %) but less appropriate gluten proteins (secalins and
secalinins) for the dough texture (no real gluten network),
and more α-amylase and less β-amylase, requesting appro-
priate acidifying processing through fermentation to avoid
too much starch hydrolysis (Deleu et al. 2020). Further,
fermentation of a rye flour-water mixture increases the sol-
ubility of the pentosans. Together with their sensitivity to
oxidative gelatinization in an acidic environment, the arab-
inoxylan properties (water-binding and gas-holding capacity)
are responsible for the actual texture of rye doughs.
Other cereal flours used for sourdough production are,
for instance, barley (Zannini et al. 2009; Mariotti et al. 2014;
Harth, Van Kerrebroeck, and De Vuyst 2016; Sadeghi et al.
2016; Bartkiene et al. 2017; Pejcz et al. 2017; Montemurro
et al. 2019; Palla et al. 2020), millet (Vogelmann et al. 2009;
Coda, Rizzello, and Gobbetti 2010; Akinola and Osundahunsi
2017; Adisa et al. 2019; Nami et al. 2019), oat (Hüttner, Dal
Bello, and Arendt 2010; Bartkiene et al. 2017; Lu et al. 2019;
Hajinia, Sadeghi, and Mahoonak 2021), sorghum (Gassem
1999; Galle et al. 2010, 2011; Svensson et al. 2010;
Sekwati-Monang, Valcheva, and Gänzle 2012), and einkorn
(Çakır, Muhammet, and Durak 2020). Although the flour
is usually of cereal origin, also flours from pseudocereals
(e.g., amaranth, buckwheat, and quinoa), legumes (e.g.,
beans, chickpeas, lentils, and lupine), and alternative seeds
(e.g., acorn, chestnut, chia, flaxseed, hempseed, and sun-
flower) are used (Vogelmann et al. 2009; Coda, Rizzello,
and Gobbetti 2010; Bartkiene et al. 2011, 2013, 2014, 2016;
Moroni et al. 2012; Aponte et al. 2013, 2014; Torino et al.
2013; Rizzello et al. 2014, 2015; Curiel et al. 2015; Fritsch
et al. 2016; Coda et al. 2017b; Rinaldi et al. 2017; Sáez et
al. 2017, 2018; Nionelli et al. 2018; Montemurro et al. 2019;
Venturi et al. 2019; Franco et al. 2020; Galli et al. 2020;
Gunduz et al. 2020; Maidana et al. 2020; Nissen, Bordoni,
and Gianotti 2020; Purabdolah et al. 2020). All these studies
on alternative cereal flours refer mainly to laboratory-made
sourdoughs that are initiated with the flour and sterile water
and that may or may not relate or lead to bakery practice.
These flours have not only different compositions but also
often contain specific compounds, such as more β-glucans
in barley and oat flours and higher concentrations of phe-
nolic compounds in sorghum and millet flours (Svensson
et al. 2010; Sekwati-Monang, Valcheva, and Gänzle 2012;
Koehler and Wieser 2013). Moreover, certain compounds,
such as phenolics, tannins, and certain enzymes in oat and
buckwheat flours may display antimicrobial activity (De
Vuyst, Van Kerrebroeck, and Leroy 2017). It is interesting
to note that artisan and industrial fermentation processes
of barley, maize, millet, oat, sorghum, and pulses flours are
widely used for the production of cereal gruels, porridges,
and beverages; although the technological aim of these fer-
mentation processes is very different from sourdoughs (used
for baking), the microbial ecology is often very comparable
to flour-water fermentations for sourdough productions
(Gänzle and Zheng 2019).
Flour fermentation for sourdough production
Natural (spontaneous) flour fermentation is carried out by
a specific microbial consortium that consists of mainly wild

LAB (in particular belonging to the family Lactobacillaceae)
and wild yeasts (De Vuyst et al. 2009, 2014, 2016; Gobbetti
et al. 2014, 2016, 2019; Minervini et al. 2014; Gänzle and
Ripari 2016; De Vuyst, Van Kerrebroeck, and Leroy 2017;
Settanni 2017; Van Kerrebroeck, Maes, and De Vuyst 2017).
Here, the term wild refers to strains coming from the direct
environment. Typical microorganisms that occur mainly in
natural (traditional) sourdoughs are Fructilactobacillus san-
franciscensis (previously known as Lactobacillus sanfrancis-
censis; Zheng et al. 2020) as LAB species and Kazachstania
humilis (previously known as Candida humilis; Jacques et
al. 2016) as yeast species (De Vuyst and Neysens 2005;
Corsetti and Settanni 2007; De Vuyst et al. 2009, 2014, 2016;
De Vuyst, Van Kerrebroeck, and Leroy 2017; Van Kerrebroeck,
Maes, and De Vuyst 2017). The LAB are responsible for the
acidification of the flour-water mixture by the production
of lactic acid and/or acetic acid as well as for flavor for-
mation (mainly the acidic flavor), whereas the yeasts are
responsible for extended flavor formation [in particular
ascribed to higher alcohols (a key compound is
3-methyl-1-butanol) and esters (a key compound is ethyl
acetate)] and leavening by the production of carbon dioxide
(De Vuyst et al. 2009, 2014, 2016; Birch, Petersen, and
Hansen 2014; Gänzle 2015; Pico, Bernal, and Gómez 2015;
De Vuyst, Van Kerrebroeck, and Leroy 2017; Pétel, Onno,
and Prost 2017). Other microorganisms such as acetic acid
bacteria (AAB) can occur too; however, they do not belong
to the core microbiota of sourdoughs (Minervini et al.
2012a; Zhang and He 2013; Lhomme et al. 2015; Li, Li, and
Bian 2016; Liu et al. 2016; Ripari, Gänzle, et al. 2016; De
Vuyst, Van Kerrebroeck, and Leroy 2017; Comasio et al.
2019; Comasio et al. 2020a). Moreover, it can be questioned
if the presence of AAB is desirable during flour fermenta-
tion. Their oxygen requirement cannot be fulfilled through
aeration, as this would enhance mold growth and flour
compound oxidation as well. Further, their oxidation of
ethanol and lactic acid would increase the flavor-impacting
concentrations of acetic acid and acetoin to a too great
extent and hence result in variations in volatile organic
compound compositions. Finally, they may have a retarded
impact on the dough rise.
The ecosystem of a sourdough represents a stressful envi-
ronment for residing microorganisms, and hence it allows
growth through adaptation of a specific microbiota express-
ing the necessary physiological and stress responses for
metabolic activities and survival (De Vuyst et al. 2009, 2014,
2016; Vogelmann and Hertel 2011; Minervini et al. 2014;
Gobbetti et al. 2016; Van Kerrebroeck et al. 2016; De Vuyst,
Van Kerrebroeck, and Leroy 2017; Van Kerrebroeck, Maes,
and De Vuyst 2017; Oshiro, Zendo, and Nakayama 2021).
This is ascribed to the variable carbohydrate and other
nutrient concentrations that occur during backslopping and/
or as a function of fermentation time, in particular the
concentration of maltose, the acidic stress due to an envi-
ronmental pH of around 4.0 and even lower caused by the
LAB acidification, and the variable and limited oxygen con-
centration due to the high viscosity {depending on the
dough yield [DY or (dough mass/flour mass) x 100], which
reflects the flour to water ratio and thus the sourdough
Critical Reviews in Food Science and Nutrition 3
consistency} and limited mixing. Alternatively, the introduc-
tion of oxygen during mixing may cause stress as well, as
the oxygen tolerance of LAB in sourdoughs is rather low
(Gänzle and Zheng 2019). The LAB counts [typically 108
colony-forming units (CFU) per gram or higher] are at least
a factor ten higher than the yeast counts (typically 10 7
CFU/g or lower). A pH value of 4.0 has been shown to be
the borderline pH concerning prevalence of certain
sourdough-specific LAB species, such as Frul. sanfranciscen-
sis (Van Kerrebroeck, Maes, and De Vuyst 2017).
Backslopped spontaneous sourdough productions starting
from scratch go through a three-step fermentation cascade
with respect to the growth of the bacterial communities in
the flour-water mixture, in particular LAB (Corsetti et al.
2007; Van der Meulen et al. 2007; De Vuyst et al. 2009,
2014; Ercolini et al. 2013; Minervini et al. 2014; De Vuyst,
Van Kerrebroeck, and Leroy 2017; Oshiro, Zendo, and
Nakayama 2021). The microbial load of the flour [< 4 to
7 log (CFU/g)] and its nutrient availability, as well as the
physicochemical parameters of the flour-water mixture, such
as temperature, DY, pH, total titratable acidity and oxygen
tension, and the fermentation time, determine the start and
course of the microbiology of a backslopping process for
sourdough production (Corsetti 2013; De Vuyst, Van
Kerrebroeck, and Leroy 2017; Minervini et al. 2018; Oshiro
et al. 2020). Hence, fermentation of the flour-water mixture
starts with autochthonous, sourdough nonspecific bacteria,
such as proteobacteria, staphylococci and diverse LAB spe-
cies, belonging to the enterococci, lactococci, and strepto-
cocci. This is followed by the growth of autochthonous LAB
species, belonging to various species of the Lactobacillaceae
(encompassing Lactobacillus, various genera related to
Lactobacillus, Leuconostoc, Pediococcus, and Weissella; Zheng
et al. 2020). All these microorganisms cause a fast acidifi-
cation of the flour-water mixture. This is finally followed
by the growth (or stabilization) of mainly sourdough-specific
LAB species that prevail in the mature sourdoughs [such
as Levilactobacillus brevis (previously known as Lactobacillus
brevis), Limosilactobacillus fermentum (previously known as
Lactobacillus fermentum), Companilactobacillus paralimenta-
rius (previously known as Lactobacillus paralimentarius),
Lactiplantibacillus plantarum (previously known as
Lactobacillus plantarum), Frul. sanfranciscensis, and
Pediococcus pentosaceus] (Van der Meulen et al. 2007;
Vrancken et al. 2010, 2011; Weckx et al. 2010a, 2010b;
Weckx et al. 2011; Minervini et al. 2012a; Harth, Van
Kerrebroeck, and De Vuyst 2016; Harth, Van Kerrebroeck,
and Vuyst 2018; Oshiro et al. 2019). Although mainly
Saccharomyces cerevisiae grows during backslopped sour-
dough productions (Gatto and Torriani 2004; Ercolini et al.
2013; Zhang et al. 2015; Harth, Van Kerrebroeck, and De
Vuyst 2016), probably because of its natural occurrence in
the flour and/or environmental contamination with baker’s
yeast (Succi et al. 2003; Vrancken et al. 2010; Minervini et
al. 2012a; Minervini et al. 2015; Lahue et al. 2020; Bigey et
al. 2021), the yeast communities may vary as a function of
processing conditions and time too (Meroth, Hammes, and
Hertel 2003; Harth, Van Kerrebroeck, and De Vuyst 2016).
For instance, backslopped wheat sourdoughs produced in
4 L. DE VUYST ET AL.
the laboratory (high temperature, high DY) often carry
Candida glabrata and Wickerhamomyces anomalus (Vrancken
et al. 2010). Backslopped rye-based laboratory sourdoughs
can harbor different combinations of C. glabrata, Kazachstania
unispora, Pichia kudriavzevii, and S. cerevisiae (Bessmeltseva
et al. 2014). Finally, during backslopped barley sourdough
productions carried out in the laboratory, both under bakery
(low DY) and industrial (high DY) conditions, only S. cer-
evisiae has been retrieved (Harth, Van Kerrebroeck, and De
Vuyst 2016). Backslopped oat and buckwheat laboratory
sourdoughs lack yeasts, probably due to growth inhibition
by endogenous antifungal compounds (Vogelmann et al.
2009; Moroni et al. 2010; Sørensen et al. 2010; Moroni,
Arendt, and Dal Bello 2011). Unless for the preparation of
new sourdough starter cultures, these spontaneous sour-
dough productions (with or without backslopping) are rel-
evant mainly for laboratory model studies, and for many of
the cereal fermentation processes that produce gruels, por-
ridges, or beverages (Gänzle and Zheng 2019). In bakery
practice, sourdough productions are almost always controlled
by backslopping, by the use of sourdough starter cultures,
or by the addition of ingredients that shortcuts the typical
succession of the microbiota in backslopped sourdough pro-
ductions (Gänzle and Ripari 2016; Gänzle and Zheng 2019).
During fermentation of the flour-water mixture, endog-
enous flour enzymes are active (Brandt and Gänzle 2006;
Gänzle 2014). The starch is degraded by the amylases, releas-
ing maltodextrins and maltose. Endogenous flour proteases
are responsible for the primary proteolysis of the flour pro-
teins, which depolymerize the gluten network and elaborate
oligopeptides and free amino acids (Gänzle, Loponen, and
Gobbetti 2008). They optimally act at pH 4.2–5.5 and low
redox potential, which is reached during flour fermentation
as a result of the LAB acidification and carbon dioxide
production, respectively. The LAB ferment free maltose and
monosaccharides to lactic acid and/or acetic acid (Gänzle,
Vermeulen, and Vogel 2007; Gänzle and Gobbetti 2013).
Further, the LAB cause a secondary proteolysis, during
which intracellular peptidases with an optimal activity
around pH 6.0–8.0 hydrolyze the oligopeptides taken up
and contribute to an accumulation of amino acids in the
sourdough matrix as a result of a release of excess amino
acids from the cells (Di Cagno et al. 2002; Rollán et al.
2005; Gänzle, Loponen, and Gobbetti 2008). These amino
acids not only enrich sourdoughs nutritionally but also are
the precursors for further enzymatic conversions into
flavor-active volatile organic compounds (aldehydes, alcohols,
carboxylic acids, and/or esters) by LAB and/or yeasts
(Gänzle, Vermeulen, and Vogel 2007; Gänzle and Gobbetti
2013; Guerzoni et al. 2013). This has an impact on the
dough rheology and aroma and taste of the bread. Further,
glutathione reductase activity by LAB reduces the disulfide
bonds of the gluten network, which further weakens the
gluten network, thereby accumulating thiol compounds that
further lower the redox potential of the sourdough matrix.
In contrast, arabinoxylans exposed to the acidic environment
of the sourdough matrix and the production of oligosac-
charides (kestose, nystose, and kestopentaose) by LAB
enhance the gas-holding capacity of the dough, hence
increasing the bread volume, next to a decreased firmness
because of their water-binding capacity. Thanks to these
oligosaccharides, the improved shelf-life of the baked goods
can be ascribed to a slower evaporation of the bound water
and reduced staling. Shelf-life improvement is of course also
ascribed to the (modest) acidification of the flour-water
mixture and the production of dedicated organic acids and
peptides as antifungal and anti-rope compounds. Oxidation
of fatty acids present in the flour, catalyzed by lipoxygenase
and hydroperoxide lyase, causes the production of volatile
aldehydes, such as 4,5-epoxy-(E)-2-decenal and (E)-2-nonenal,
that influence the bread flavor (Czerny and Schieberle 2002;
Birch, Petersen, and Hansen 2014; Gänzle 2014; Pétel, Onno,
and Prost 2017).
Fermentation of a rye flour-water mixture for sourdough
production causes a desirable inhibition of the α-amylase
that is present abundantly in rye flour by limiting too exten-
sive starch degradation, good hydration of the arabinoxylans
yielding an airier and stronger dough (creating a
pseudo-gluten network), and converting undesirable volatile
organic compounds (Kirchhoff and Schieberle 2002; Brandt
2007; Cappelle et al. 2013; Koehler and Wieser 2013; Deleu
et al. 2020).
The technological and health-promoting properties of
sourdoughs have been reviewed before extensively (Arendt,
Ryan, and Dal Bello 2007; Corsetti and Settanni 2007;
Poutanen, Flander, and Katina 2009; Corsetti 2013; Galle
2013; Katina and Poutanen 2013; Gänzle 2014; Gobbetti et
al. 2014, 2019; Gänzle and Ripari 2016; Nionelli and Rizzello
2016; De Vuyst, Van Kerrebroeck, and Leroy 2017; Koistinen
et al. 2018; Loponen and Gänzle 2018; Menezes et al. 2018;
Sahin et al. 2019a; Fernández-Peláez, Paesani, and
Gómez 2020).
Classification of sourdough production processes
Sourdough production processes can be classified according
to the inoculum used to initiate the fermentation of the
flour-water mixture, to make them comparable with fer-
mentation processes carried out in the non-bakery food
fermentation industry, or based on the process technology
applied, a traditional classification approach (De Vuyst, Van
Kerrebroeck, and Leroy 2017). These classifications are inde-
pendent of their geographical origin (De Vuyst et al. 2014;
Van Kerrebroeck, Maes, and De Vuyst 2017; Landis et
al. 2021).
Inoculum-based classification
According to their inoculum, three types of sourdough pro-
duction processes (Types 1, 2, and 3) can be distinguished,
which influence the microbiology of the mature sourdoughs
(Figure 1; De Vuyst, Van Kerrebroeck, and Leroy 2017).
Type 1 sourdough production processes represent fermen-
tations of the flour-water mixture through traditional daily
backslopping to keep the sourdough LAB and yeasts met-
abolically active (De Vuyst et al. 2014; De Vuyst, Van
Kerrebroeck, and Leroy 2017). Backslopping is based on

Figure 1. Various types of sourdough production processes based on the way of inoculation of the flour-water mixture (based on De Vuyst, Van Kerrebroeck,
and Leroy 2017).
several refreshment steps of a part of the preceding sour-
dough with a fresh flour-water mixture, in the very first
step starting from the mother sourdough kept. However,
backslopping can also refer to refreshment of a leftover
portion of sourdough for the next breadmaking (Urien et
al. 2019). Sometimes a resting time is applied, for instance
refrigerated storage of the sourdough for several days or
weeks, to allow for less frequent backslopping and/or have
the microbial composition under control (Venturi, Guerrini,
and Vincenzini 2012). Type 1 sourdoughs, which are live
sourdoughs with leavening capacity, are generally made and
Critical Reviews in Food Science and Nutrition 5
maintained in artisan bakeries and on household level to
produce traditional breads. Further, artisan bakery practices
often encompass tight and efficient control of the fermen-
tation conditions (storage and environmental temperature,
length of the fermentation period between refreshments,
refreshment ratio, and number of refreshments) and thus
the fermentation microbiota, so that a vast majority of bak-
eries possesses sourdoughs characterized by the presence of
Frul. sanfranciscensis and K. humilis as LAB and yeast spe-
cies, respectively (Gänzle and Zheng 2019). Indeed, these
sourdough-specific microorganisms require well-controlled
6 L. DE VUYST ET AL.
production and maintenance conditions (Venturi, Guerrini,
and Vincenzini 2012; Van Kerrebroeck et al. 2018b). Other
microorganisms involved are, for instance, Coml. paralimen-
tarius, Levl. brevis, and Lacp. plantarum (Gänzle and Zheng
2019). In contrast, household sourdoughs can be subjected
to less strict conditions for fermentation and maintenance,
causing their fermentation microbiota more dependent on
the local environmental conditions, production practices,
and ingredients (Landis et al. 2021). Type 2 sourdough
production processes represent prolonged fermentations of
the flour-water mixture that are initiated through the addi-
tion of a common starter culture that targets specific prop-
erties in the end-products, such as acidification and flavor
formation. Type 2 sourdoughs are industrial products. The
sourdoughs thus obtained are commercialized as semi-liquid,
heat-treated, spray-dried, drum-dried, or freeze-dried prepa-
rations, to be used as flavor or technological ingredients
(dough acidifier or improver, bread shelf-life improver) in
(industrial) bakeries. Only in the case of semi-liquid sour-
doughs, the microorganisms are generally still metabolically
active. Type 2 sourdoughs are obtained through
temperature-controlled fermentation of the flour-water mix-
ture with specific strains of acid-tolerant LAB species [e.g.,
Lactobacillus amylovorus, Levl. brevis, Liml. fermentum, Lacp.
plantarum, Limosilactobacillus reuteri (previously known as
Lactobacillus reuteri), and Furfurilactobacillus rossiae (pre-
viously known as Lactobacillus rossiae)], causing a fast acid-
ification or the expression of desirable organoleptic
properties. Finally, Type 3 sourdough production processes
represent starter culture-initiated fermentations of the
flour-water mixture (as in Type 2) followed by daily back-
slopping (as in Type 1). The latter practice will determine
which microorganisms end up in the mature sourdoughs,
which depends on the competitiveness of the strain(s) pres-
ent in the common starter culture used for inoculation of
the initial flour-water mixture (Siragusa et al. 2009; Minervini
et al. 2010; Moroni et al. 2010; Lin and Gänzle 2014; Coda
et al. 2018). Type 3 sourdough production processes are
thus a combination of Type 1 and Type 2 sourdough pro-
duction processes and are applied by some bakeries, both
artisan and industrial ones.
Process technology-based classification
According to their process technology, four types of sour-
dough production processes (Types 0, I, II, and III) can be
distinguished (Figure 2; Corsetti 2013; De Vuyst, Van
Kerrebroeck, and Leroy 2017). Type 0 sourdough production
processes represent sponge doughs or pre-doughs, during
which fermentation of the flour-water mixture is allowed
for a limited time, so that the LAB species, which are nat-
urally present in the flour and as contaminants of the baker’s
yeast preparation, can grow (Reale et al. 2013). Although
the yeasts grow faster than the LAB, the LAB species cause
a limited acidification and flavor formation. Typical Type 0
sourdough products are French baguettes, Italian ciabattas,
and American crackers. They are often not considered as
sourdough products. Type I sourdough production processes
represent traditional sourdoughs that are daily backslopped
at room temperature (20–30 °C) and low DY (< 200), with
a short to moderate fermentation time (6–24 h) to reach a
pH of about 4.0. The maintenance of such original mother
doughs for years has built up a rich collection of bakery
sourdoughs of Type I. Some of them are preserved in the
sourdough library of Puratos (www.puratos.com/en/innova-
tion/Center-for-Bread-Flavor/Sourdough-library/). Typical
examples are traditional sourdoughs, such as San Francisco
sourdough and various Italian sourdoughs, which are often
legally protected as protected designation of origin (PDO)
or protected geographic indication (PGI) (e.g., Pane di
Altamura PDO and Pane di Matera PGI; Vernocchi et al.
2004; Ricciardi et al. 2005; Zotta et al. 2008; Minervini et
al. 2012b). They frequently harbor Frul. sanfranciscensis,
Lacp. plantarum, Levl. brevis, Companilactobacillus (previ-
ously known as Lactobacillus) species (e.g., Coml. crustorum,
Coml. mindensis, Coml. nantensis, and Coml. paralimentar-
ius), leuconostocs, and weissellas (Van Kerrebroeck, Maes,
and De Vuyst 2017; Gänzle and Zheng 2019). Type II sour-
dough production processes represent acid-tolerant starter
culture-initiated and thus accelerated, one-step fermentations
carried out at controlled temperatures above ambient tem-
perature (30–37 °C), which enhance acidification and/or
target other sourdough-specific traits, and at high DY (>
200), which enables a simple handling of the sourdoughs
(for instance mixing and pumping). These processes last
one to three days, thereby often achieving pH values lower
than 4.0. These sourdoughs are available in liquid or pasty
form or as stiff or crumbly products; dried powders obtained
through gentle drying processes (spray, fluidized bed, or
freeze-drying), which keeps the microorganisms alive, are
available too. They frequently harbor species of
Limosilactobacillus (previously known as Lactobacillus; e.g.,
Liml. pontis, Liml. fermentum, Liml. reuteri, Liml. frumenti,
and Liml. panis) and Lactobacillus (e.g., Lb. amylovorus, Lb.
acidophilus, and Lb. crispatus) (Van Kerrebroeck, Maes, and
De Vuyst 2017; Gänzle and Zheng 2019). Finally, Type III
sourdough production processes yield sourdoughs (produced
as in Type II) that are dried by means of a spray or drum
drier after the fermentation process. These ready-to-use
sourdoughs are not metabolically active, which differentiates
them from the Type II sourdoughs. Type II and Type III
sourdoughs are used as acidifying or flavor carriers in bread
doughs by bakeries, both artisan and industrial ones. They
require the addition of baker’s yeast for leavening during
dough preparation, because yeasts are either inactivated
through fast and extensive acidification (Type II) or dry
processing (Type III).
Starter cultures
Concerning starter cultures used in Type II and Type III
sourdough production processes, distinction can be made
between a common defined starter culture as it is used in
the non-bakery food fermentation industry (cf. Type 2 sour-
dough production processes) or a traditional sourdough
starter culture. Common defined starter cultures enable the

inoculation of (a) pure culture(s) of carefully selected strains
of LAB species, whether or not combined with strains of
yeast species, in defined ratios. These starter cultures are
commercially available as frozen and freeze-dried monocul-
tures or freeze-dried blended cultures. Also refrigerated or
freeze-dried common defined starter cultures that are pro-
duced in cereal substrates as well as refrigerated, undefined
starter cultures that the starter culture suppliers maintain
Figure 2. Various types of sourdough production processes based on their process technology applied (based on De Vuyst, Van Kerrebroeck, and Leroy 2017).
Critical Reviews in Food Science and Nutrition 7
by backslopping are available (Brandt 2014, 2019). The only
difference between the last type of starter cultures and a
backslopped sourdough (cf. Type 1/I) is that the backslop-
ping is done by two different companies, namely the
culture-supplying company and the bakery, respectively.
When no yeast is co-cultivated, sometimes baker’s yeast is
added to the sourdough matrix at the end of the sourdough
production process.
8 L. DE VUYST ET AL.
Traditional sourdough starter cultures (cf. Type 1/I) rep-
resent stable microbial consortia (alive and active) under
the form of a liquid/pasty batter or a stiff/crumbly dough,
depending on the DY (ratio of water to flour in the sour-
dough starter), maintained through backslopping. They are
derived from an original sourdough starter that was the
result of a spontaneously fermented flour-water mixture that
became stabilized upon backslopping and, hence, may be
many years old. As long as the sourdough starter culture
is regularly refreshed by the periodic addition of flour and
water, it will remain metabolically active. The regularity of
the refreshments is mainly determined by the acid tolerance
of the microorganisms involved (Gänzle and Zheng 2019).
It can be questioned if the addition of manure, fruits,
etc., either at the initiation of the sourdough build-up or
upon refreshment, can be considered as a type of starter
culture; yet, these materials carry microorganisms that start
and/or contribute to the fermentation of the flour-water
mixture (cf. Section on Sourdough Productions with Added
Non-flour Ingredients).
Sourdough microorganisms
The microbial species diversity of sourdoughs, in particular
the phylogenetic diversity, is limited. It concerns yeasts
(Ascomycota), LAB (Firmicutes, mainly Lactobacillaceae),
and sometimes AAB (α-proteobacteria).
Yeasts
Yeasts are unicellular fungi that are mainly facultatively
aerobic and reproduce either asexually (mainly budding) or,
under starvation conditions, sexually (spore formation)
(Neiman 2005; Kurtzman, Fell, and Boekhout 2011). They
play an important role in fermented food production by
causing desirable transformations of appropriate food raw
materials (e.g., bread, beer, and wine productions).
Alternatively, they are responsible for the spoilage of differ-
ent foods. Sourdoughs mainly contain yeasts that belong to
the Ascomycota, being fermentative, acid-tolerant, and
adapted to the sourdough ecosystem (Huys, Daniel, and De
Vuyst 2013; De Vuyst et al. 2016).
Examples of sourdough yeasts, in decreasing order of
relative abundance in 394 backslopped sourdoughs exam-
ined, are S. cerevisiae (68 % of the sourdoughs), K. humilis
(20 %), W. anomalus (6 %), Torulaspora delbrueckii (6 %),
Pi. kudriavzevii (6 %), and C. glabrata (4 %) (Van
Kerrebroeck, Maes, and De Vuyst 2017). The number of
different yeast species isolated from a single sourdough is
in general equal or lower than the number of LAB species
(De Vuyst, Van Kerrebroeck, and Leroy 2017; Van
Kerrebroeck, Maes, and De Vuyst 2017). Also, one or two
yeast species are often inhabiting a single bakery sourdough,
among which S. cerevisiae and K. humilis occur most fre-
quently (De Vuyst et al. 2009, 2014, 2016; De Vuyst, Van
Kerrebroeck, and Leroy 2017; Van Kerrebroeck, Maes, and
De Vuyst 2017). Saccharomyces cerevisiae represents common
baker’s yeast that is maltose-positive and relatively
acid-tolerant (De Vuyst et al. 2016; Lahue et al. 2020).
Thanks to its maltose-negative fermentation behavior and
acid-tolerant (no effect on growth between pH 3.5 and 7.0)
and relatively thermotolerant (up to 36 °C) properties, K.
humilis does not compete with the maltose-positive
sourdough-specific LAB species, such as Frul. sanfranciscen-
sis, during sourdough production (Gänzle, Ehrmann, and
Hammes 1998; Brandt, Hammes, and Gänzle 2004). In con-
trast, a trophic relationship occurs between K. humilis and
Frul. sanfranciscensis, making it a typical sourdough yeast
species in Type 1/I sourdough production processes (Stolz
et al. 1993; Corsetti and Settanni 2007; Huys, Daniel, and
De Vuyst 2013; De Vuyst et al. 2016; Van Kerrebroeck,
Maes, and De Vuyst 2017). However, this mutual relationship
seems to be strain-dependent for both species (Carbonetto
et al. 2020; Rogalski, Ehrmann, and Vogel 2020). The nutri-
tional relationship further encompasses the provision of
fructose (from glucofructans through invertase activity),
amino acids (complementing auxotrophies of Frul. sanfran-
ciscensis), and vitamins by K. humilis to Frul. sanfranciscensis
(Vogel et al. 2011; Vogel 2015). In addition, K. humilis
causes oxidative stress through thiol oxidation that is with-
stood by Frul. sanfranciscensis via its redox homeostasis
(e.g., glutathione reductase activity, NADH oxidase activity).
Fructilactobacillus sanfranciscensis causes acidic stress
through the production of lactic acid and acetic acid that
is withstood by K. humilis through its acetic acid tolerance
(Brandt, Hammes, and Gänzle 2004). However, it is not
excluded that other more complex interactions occur,
encompassing both mutualistic and antagonistic interactions
(Carbonetto et al. 2020; Rogalski, Ehrmann, and Vogel 2020).
Sourdough yeasts ferment the flour saccharides (malt-
os e, sucros e, g lucos e, and f r uc tos e) v ia the
Embden-Meyerhof-Parnas (EMP) pathway into pyruvate,
thereby generating both ATP and reducing power
(NADH + H+), and further convert pyruvate into ethanol
and carbon dioxide (alcoholic fermentation), thereby
regenerating the cofactor NAD+ consumed in the upper
part of the EMP pathway (Figure 3; De Vuyst et al. 2016;
De Vuyst, Van Kerrebroeck, and Leroy 2017). The forma-
tion of glycerol from dihydroxy acetone phosphate gen-
erated by the EMP pathway as well as the formation of
succinic acid through the reductive tricarboxylic acid
(TCA) cycle further contribute to redox balancing (NAD+
regeneration). In particular, the formation of carbon diox-
ide is important during dough fermentation and, hence,
for the production of bread, although ethanol (that evap-
orates during the baking process), glycerol (produced as
osmoprotectant and improving carbon dioxide retention),
and succinic acid (causing a pH decrease) contribute to
the formation of the gluten network and the rheological
properties of the dough too (Jayaram et al. 2014a, 2014b;
Aslankoohi et al. 2015; Rezaei, Verstrepen, and Courtin
2015; Rezaei et al. 2016; De Vuyst, Van Kerrebroeck, and
Leroy 2017).
Further, the yeasts contribute to the bread flavor by pro-
ducing different flavor-active metabolites, in particular
higher alcohols from amino acids, such as 3-methyl-1-butanol
(important contributor to the bread crumb flavor) from

Figure 3. Overview of the metabolism of yeasts in a sourdough matrix. Only the uptake of appropriate substrates and the most relevant pathways are repre-
sented. EMP, Embden-Meyerhof-Parnas pathway; PPP, pentose phosphate pathway; TCA, tricarboxylic acid cycle (from De Vuyst, Van Kerrebroeck, and Leroy
2017, adapted version).
L-leucine, either anabolically (side-product of amino acid
biosynthesis) or catabolically (amino acid conversion via the
Ehrlich pathway) (Hazelwood et al. 2008; Birch et al. 2013;
Birch, Petersen, and Hansen 2014; Pico, Bernal, and Gómez
2015; De Vuyst, Van Kerrebroeck, and Leroy 2017; Pétel,
Onno, and Prost 2017). Esters as condensation products of
ethanol and fatty acids (ethyl esters) or higher alcohols and
acetate (acetate esters) contribute mainly fruity and floral
notes to the bread flavor. Also the production of diacetyl
(butter aroma), produced either via pyruvate metabolism
during fermentation or by Strecker degradation during bak-
ing, contributes to the crumb flavor. The inclusion of air
during the mixing and kneading steps of dough preparation
enhances the production of the yeast metabolites thanks to
a better growth of the yeasts aerobically. Moreover, in the
absence of yeasts, air inclusion is adverse, as oxygen is then
used by the flour lipoxygenase that oxidizes the flour lipids
to aldehydes that are undesirable in bread doughs. Further,
yeasts produce acetic acid and succinic acid that not only
slightly acidify the dough and contribute to the dough tex-
ture but also affect the dough and bread flavor. Finally, the
yeasts degrade the flour fructans and contribute to acetic
acid formation and flavor indirectly, namely by elaborating
fructose that can be used by heterofermentative LAB species
as alternative external electron acceptor (instead of pyruvate)
Critical Reviews in Food Science and Nutrition 9
to produce mannitol (for NAD+ regeneration) and acetic
acid (instead of ethanol production for NAD+ regeneration,
thereby enabling extra ATP formation). This particularly
takes place during Type 1/I sourdough productions (De
Vuyst et al. 2016; De Vuyst, Van Kerrebroeck, and
Leroy 2017).
Lactic acid bacteria
LAB are Gram-positive, catalase-negative, non-sporulating,
anaerobic (aerotolerant) bacteria that belong to the Firmicutes
(Axelsson 2004; Holzapfel and Wood 2014). They occur in
various niches, ranging from (fermented) food products,
plant materials, and insects to the human body. As they are
fermentative, redox balancing is accomplished by the pro-
duction of lactic acid from pyruvate, produced via the EMP
pathway from flour saccharides, to generate both ATP and
reducing power in the case of homofermentative LAB spe-
cies, or by the production of lactic acid and ethanol or
acetic acid (produced from pyruvate and acetyl-phosphate,
respectively) and carbon dioxide, produced via the phos-
phogluconate pathway from flour saccharides, to generate
both ATP and reducing power in the case of heterofermen-
tative LAB species (Figure 4). They play an important role
10 L. DE VUYST ET AL.
Figure 4. Overview of the metabolism of heterofermentative LAB species in a sourdough matrix. Only the uptake of appropriate substrates and the most
relevant pathways are represented (from De Vuyst, Van Kerrebroeck, and Leroy 2017, adapted version).
in the production of many fermented foods and beverages
(e.g., fermented dairy, meat, cereal, and vegetable products).
Sourdoughs mainly contain heterofermentative LAB species
belonging to the Lactobacillaceae that are particularly
adapted to the sourdough ecosystem (De Vuyst and Neysens
2005; De Vuyst, Van Kerrebroeck, and Leroy 2017; Van
Kerrebroeck, Maes, and De Vuyst 2017). According to their
lifestyles, distinction can be made between insect-adapted
Frul. sanfranciscensis, typical for Type I sourdoughs (often
associated with Companilactobacillus species), vertrebate
host-adapted Lactobacillus and Limosilactobacillus species,
typical for Type II sourdoughs, and environmental or
nomadic Liml. fermentum, Lacp. plantarum, and Levl. brevis,
typical for laboratory sourdoughs, Type 0 sourdoughs, and
sourdoughs made from flours other than those of wheat or
rye (Gänzle and Zheng 2019).
Examples of heterofermentative sourdough LAB species,
in decreasing order of relative abundance in 527 backslopped
sourdoughs examined, are Frul. sanfranciscensis (47 %), Levl.
brevis (17 %), and Liml. fermentum (12 %) (Van Kerrebroeck,
Maes, and De Vuyst 2017). Further, homofermentative LAB
species occur, such as Lacp. plantarum (43 %), P. pentosaceus
(14 %), and Coml. paralimentarius (13 %). Heterofermentative
leuconostocs and weissellas are usually present in sour-
doughs with high pH (> 4.0) and fermented at low tem-
perature (< 30 °C) and low DY (< 200). In general, less than
three different LAB species inhabit a single bakery sour-
dough. The LAB species Frul. sanfranciscensis, with its small
genome of 1.3 Mbp and, hence, a restricted metabolism, is
typically associated with the sourdough ecosystem (Vogel et
al. 2011; Zheng et al. 2015; Gänzle and Zheng 2019).
Similarly, Coml. (par)alimentarius is rarely found in other
ecosystems than sourdough, albeit that this LAB species
harbors a genome twice as large (2.55 Mbp) as that of Frul.
sanfranciscensis (Gänzle and Zheng 2019; Zheng et al. 2020).
Thus, it is not always clear how a matrix specificity can be
explained genomically. This homofermentative,
maltose-negative LAB species often occurs together with
Frul. sanfranciscensis and, in contrast with the latter, fer-
ments fructose (De Vuyst et al. 2002; Gänzle and Zheng
2019). Hence, its co-occurrence behavior and broader sac-
charide metabolism spectrum may require a larger genome.
Adaptations of heterofermentative LAB species, such as
Frul. sanfranciscensis, Liml. fermentum, and Levl. brevis to
the sourdough ecosystem encompass several physiological
traits, depending on the environmental conditions (Figure
4; Gänzle 2015; De Vuyst, Van Kerrebroeck, and Leroy 2017;
Gänzle and Zheng 2019). In the first place, there is the
preferred fermentation of maltose by means of a constitu-
tively expressed, energy-efficient maltose phosphorylase.
Maltose fermentation is not subject to glucose repression
and does not interfere with yeast growth, thanks to the
mutual relationship between the maltose
phosphorylase-positive heterofermentative LAB species (e.g.,
Frul. sanfranciscensis, Liml. fermentum, and Liml. reuteri)
and the maltose-negative yeasts (in particular K. humilis)
in the sourdough matrix. Further, the use of alternative
external electron acceptors, such as fructose (continuously
elaborated by yeast activity) that is then reduced to mannitol
to recuperate NAD+, allows to improve the strains’ compet-
itiveness (thanks to extra ATP formation) and to enhance
the production of acetic acid by circumventing ethanol for-
mation and activating the acetate kinase branch of the pyru-
vate metabolism. This is also the case for the use of fructose
6-phosphate that is reduced to erythritol, the use of oxalo-
acetic acid that is converted into malate, which can be
further converted into fumarate that is in turn reduced to
succinate (via the reductive TCA cycle), the use of oxygen

gas that is reduced to hydrogen peroxide (by pyruvate oxi-
dase activity), the use of (lipid oxidation-derived) aldehydes
that are converted into alcohols (by alcohol dehydrogenases),
and the use of glutathione disulfide (oxidized glutathione
of the gluten network) that is reduced to glutathione (by
glutathione reductase). All these activities are LAB species-
and/or strain-dependent (Gänzle, Vermeulen, and Vogel
2007; Gänzle and Gobbetti 2013; Gänzle 2015; Gänzle and
Zheng 2019).
Further, there is the ability by LAB species to ferment
both hexoses and pentoses (via xylulose 5-phosphate)
present in the flour through the phosphogluconate path-
way, for instance by Coml. paralimentarius and Lacp. plan-
tarum. The use of sucrose by means of an energy-efficient
sucrose phosphorylase by, for instance, Liml. reuteri is
also advantageous. Alternatively, dedicated strains of Frul.
sanfranciscensis use sucrose by extracellular levansucrase
activity, thereby elaborating free glucose; however, glucose
(as energy source) and fructose (as alternative external
electron acceptor) are mostly provided through yeast
invertase activity, as the growth of Frul. sanfranciscensis
is commonly associated with that of yeasts and sometimes
sucrose-metabolizing LAB species that carry out extracel-
lular levansucrase activity (Gänzle and Zheng 2019). Also,
sucrose can be converted into dough texture-contributing
homooligosaccharides by glucan- and fructansucrase or
levansucrase activities by Limosilactobacillus species as well
as leuconostocs and weissellas. Furthermore, acid-tolerant
Limosilactobacillus species execute amino acid conversions
that enable a protection toward acidic stress. The arginine
deiminase (ADI) pathway converts arginine into ornithine,
thereby releasing ammonia that slightly increases the pH
(acidic stress response) and allows extra ATP formation,
which further enhances the strains’ competitiveness (Figure
4; Gänzle and Gobbetti 2013; Gänzle 2015; De Vuyst, Van
Kerrebroeck, and Leroy 2017). Further, deamidation of
glutamine (glutaminase activity) and glutamate (glutamate
dehydrogenase activity) consumes protons and releases
ammonia, whereas decarboxylation of dedicated amino
acids also consumes protons and may produce biogenic
amines (Bartkiene et al. 2011, 2014; Diana, Rafecas, and
Quílez 2014). Both metabolic activities again allow to
increase the intracellular pH. This is also the case for the
decarboxylation of glutamate through glutamate decarbox-
ylase activity that yields γ-aminobutyric acid (GABA), an
anti-hypertensive and thus health-promoting compound.
Cystathionine conversion and glutathione reduction are
dedicated responses of LAB species toward acidic and
oxidative stresses, respectively. However, glutaminase, glu-
tamate decarboxylase, and ADI pathway activity are absent
in Frul. sanfranciscensis, which makes this species sensitive
to low pH and thus highly dependent on regular sour-
dough refreshments (Vogel et al. 2011; Zheng et al. 2015;
Gänzle and Zheng 2019; Rogalski, Ehrmann, and
Vogel 2021).
Further, LAB are responsible for the formation of certain
flavor compounds (Gänzle 2014, 2015; De Vuyst, Van
Kerrebroeck, and Leroy 2017). First, lactic acid (homo- and
heterofermentative LAB species) and acetic acid
Critical Reviews in Food Science and Nutrition 11
(heterofermentative LAB species) cause the acidic taste of
sourdough-based breads. The ratio of lactic acid to acetic
acid is a determining factor for the acidic taste of sour-
doughs, often represented as the fermentation quotient (pref-
erably around 3.0–5.0 moles of lactic acid per mole of acetic
acid), as both lactic acid (delivering a fresh and soft acidity)
and acetic acid (delivering a sharp and strong vinegar-like
acidity) reduce the pH, but only acetic acid is volatile and
hence contributes to the aroma. Second, pyruvate metabo-
lism by homofermentative Lactobacillaceae allows the for-
mation of, for instance, diacetyl and acetoin (buttery notes).
Diacetyl can be further converted into pyrazines through
Maillard reactions during the bread baking process. Third,
the accumulation of amino acids, as a result of peptide
hydrolysis, may result in, for instance, an umami taste (accu-
mulation of glutamate). In addition, ornithine (elaborated
via the ADI pathway) can react with the carbonyl compound
2-oxopropanal (methylglyoxal) during Maillard reactions
taking place during the bread baking process, forming
2-acetyl-1-pyrroline that is responsible for the bread crust
aroma (Moskowitz et al. 2012; Gänzle 2014; Pico, Bernal,
and Gómez 2015; De Vuyst, Van Kerrebroeck, and Leroy
2017). Moreover, as sourdough decreases the pH of the
bread dough, Amadori rearrangements are favored during
Maillard reactions and, hence, the generation of various
Maillard compounds, such as furans, pyrazines, pyrroles,
pyrrolines, and sulfuric compounds, takes place during the
baking process (Cho and Peterson 2010; Pico, Bernal, and
Gómez 2015). Finally, the formation of taste-active peptides
through proteolysis, such as glutathione and some γ-glutamyl
dipeptides and tripeptides, cause a kokumi taste (Zhao and
Gänzle 2016; Zhao, Schieber, and Gänzle 2016). Also the
phenolic acids and flavonoids present in the flour may be
converted into diverse aroma precursors by strain-specific
phenolic acid decarboxylases and cinnamic acid reductases,
for instance in the case of Lacp. plantarum, albeit that sim-
ilar activity by other LAB species and/or yeasts cannot be
excluded (Czerny and Schieberle 2002; Gänzle 2014; De
Vuyst, Van Kerrebroeck, and Leroy 2017; Ripari, Yunpeng,
and Gänzle 2019; Boudaoud et al. 2021).
Acetic acid bacteria
A second group of bacteria that sometimes occurs in sour-
doughs are AAB (De Vuyst, Van Kerrebroeck, and Leroy
2017). AAB are Gram-negative, catalase-negative,
oxidase-positive, non-sporulating, aerobic bacteria that
belong to the α-proteobacteria (Matsushita et al. 2016; De
Roos and De Vuyst 2018). They mainly occur in ethanol-
and carbohydrate-rich niches, such as flowers and fruits.
Examples of AAB that have been found in sourdoughs
are Acetobacter and/or Gluconobacter species (not specified
in Scheirlinck et al. 2008; Vogelmann et al. 2009; and Zhang
and He 2013), Acetobacter cerevisiae (Li, Li, and Bian 2016;
Ripari, Gänzle, et al. 2016; Comasio et al. 2020a), Acetobacter
fabarum (Comasio et al. 2020a), Acetobacter indonesiensis
(Comasio et al. 2019), species of the Acetobacter lovaniensis
group (Landis et al. 2021), species of the Acetobacter
12 L. DE VUYST ET AL.
Figure 5. Overview of the metabolism of acetic acid bacteria (AAB; based on Acetobacter sp. and Gluconobacter sp.) of potential importance in a sourdough
matrix. Only the uptake of appropriate substrates and the most relevant pathways are represented (based on Matsushita et al. 2016). Dashed lines represent
multiple metabolic reactions. The dotted line represents a non-enzymatic reaction. EMP, Embden-Meyerhof-Parnas pathway; PPP, pentose phosphate pathway;
TCA, tricarboxylic acid cycle.
malorum group (Landis et al. 2021), Acetobacter orleanensis
(Minervini et al. 2012a; Li, Li, and Bian 2016), Acetobacter
oryzifermentans (Comasio et al. 2020a), Acetobacter pasteur-
ianus (Lhomme et al. 2015; Comasio et al. 2020a; Landis
et al. 2021), Acetobacter senegalensis (Comasio et al. 2020a),
Acetobacter sicerae (Comasio et al. 2020a), Acetobacter trop-
icalis (Li, Li, and Bian 2016), Gluconobacter cerinus (Ripari,
Gänzle, et al. 2016), species of the Gluconobacter frateurii
group (Landis et al. 2021), Komagataeibacter sucrofermentans
(Comasio et al. 2020a), and Komagataeibacter xylinus
(Comasio et al. 2020a).
AAB produce acetic acid from ethanol and/or gluconic
acid from glucose by partial oxidation of these substrates
in the cells’ periplasm, using dedicated cofactor-depending
dehydrogenases that are embedded in the cytoplasmic mem-
brane at the periplasmic site (Figure 5; Matsushita et al.
2016). The acetic acid produced may contribute to the
acidic taste of sourdough breads. Also, the aroma formation
potential (e.g., acetoin and diacetyl) of starter culture-initiated
sourdough productions with A. cerevisiae, Gluconobacter
albidus, and Kozakia baliensis has been tested (Ripari,
Cecchi, et al. 2016; Ua-Arak, Jakob, and Vogel 2017).
Further, AAB are able to produce fructans, which may
contribute to the texture of sourdough breads, which has
been tested through starter culture-initiated sourdough pro-
ductions, in the presence of sucrose or sugarcane molasses,
with G. albidus, K. baliensis, and Neoasaia chiangmaiensis
(Hermann, Petermeier, and Vogel 2015; Ua-Arak, Jakob,
and Vogel 2016, 2017). However, in all these cases, a con-
tinuous agitation was applied to provide oxygen for the
AAB growth. Also, AAB may contribute to the coloration
of the bread crumb, likely by the production of dihydroxy
acetone, hence enabling to produce brown wheat breads
(Comasio et al. 2020b).
Selection of functional starter cultures for
sourdough productions
As mentioned above, starter cultures composed of defined
strains can be used in Type 2, Type 3, and Type II/III sour-
dough production processes. Most of these commercial
starter cultures in use have been selected based on their
growth and acidification capacity; some of them are pro-
moted because of their contribution to organoleptic proper-
ties, either flavor generation or texture improvement (Decock
and Cappelle 2005; Brandt 2007, 2014; De Vuyst et al. 2009,
2014; De Vuyst, Van Kerrebroeck, and Leroy 2017). Nowadays,
much attention is paid to extra functional properties of both
LAB and yeast strains, for instance shelf-life extension and
health promotion; even AAB strains have been considered
as candidate members for the development of functional
starter cultures for sourdough production (De Vuyst et al.
2016; De Vuyst, Van Kerrebroeck, and Leroy 2017).

Microbial performance of a starter culture is of course
a first criterion that has been examined thoroughly (Meroth
et al. 2003; Siragusa et al. 2009; Vogelmann et al. 2009;
Moroni et al. 2010; Banu, Vasilean, and Aprodu 2011; Ravyts
and De Vuyst 2011; Banu and Aprodu 2012; Capuani et al.
2014; Nionelli et al. 2014; Picozzi et al. 2016; Harth, Van
Kerrebroeck, and Vuyst 2018; Van Kerrebroeck et al. 2018a;
Francesca et al. 2019; Galli et al. 2019; Gaglio et al. 2020;
Gunduz et al. 2020; Montemurro et al. 2020; Teleky et al.
2020). With regard to functional properties of LAB, it
mainly concerns flavor formation (Meignen et al. 2001;
Katina, Poutanen, and Autio 2004; Paramithiotis et al. 2005;
Katina et al. 2006; Settanni et al. 2013; Prückler et al. 2015;
Alfonzo et al. 2016; Corona et al. 2016; Ripari, Cecchi, et
al. 2016; Van Kerrebroeck et al. 2016, 2018a, 2018b; D. Xu
et al. 2019; Comasio et al. 2020b; Liu et al. 2020), texture
improvement (Katina et al. 2006, 2009; Robert et al. 2006;
Coda et al. 2010; Galle et al. 2010, 2011; Palomba et al.
2012; Bartkiene et al. 2013, 2016, 2017; Tamani, Goh, and
Brennan 2013; Pontonio et al. 2015; Chen, Levy, and Gänzle
2016; Kajala et al. 2016; Mamhoud et al. 2016, Y. Xu et al.
2017, 2019; D. Xu et al. 2018; Chen et al. 2018, Coda et
al. 2018; Nionelli et al. 2018; Bockwoldt et al. 2020; Comasio
et al. 2020b; Galli et al. 2020; Ispirli et al. 2020; Olojede,
Sanni, and Banwo 2020; Sun et al. 2020), antimicrobial
activity (Leroy et al. 2007; Choi et al. 2012; Axel et al. 2015;
Axel et al. 2016a, 2016b; Belz et al. 2019; Sadeghi et al.
2019; Bartkiene et al. 2020; Sun et al. 2020; Jin et al. 2021),
and health promotion (Rizzello et al. 2008; Rizzello et al.
2016; Banu, Vasilean, and Aprodu 2010; Coda, Rizzello, and
Gobbetti 2010; Bartkiene et al. 2011; Coda et al. 2012;
Rizzello et al. 2012a; Rizzello et al. 2012b; Wolter et al.
2014; Pallin et al. 2016; Picozzi et al. 2016; Rizzello,
Montemurro, and Gobbetti 2016; Ruiz Rodríguez et al.
2016a, 2016b; Coda et al. 2017a; Rizzello et al. 2017a;
Rizzello et al. 2017b; Galli et al. 2018, 2020; Sahin et al.
2019b, 2019c; Venturi et al. 2019; Fraberger et al. 2020;
Gaglio et al. 2020; Hajinia, Sadeghi, and Mahoonak 2021;
Luti et al. 2020; Pontonio et al. 2020; Purabdolah et al.
2020; Sun et al. 2020).
In the case of AAB strains, it concerns flavor formation
(Ripari, Cecchi, et al. 2016; Ua-Arak, Jakob, and Vogel 2017;
Van Kerrebroeck et al. 2018a, 2018b), texture improvement
(Hermann, Petermeier, and Vogel 2015; Ua-Arak, Jakob, and
Vogel 2016, 2017), and color improvement (Comasio et
al. 2020b).
Also Zymomonas mobilis has been used during flour
fermentation for sourdough production (in co-culture with
Frul. sanfranciscensis) and for dough making, thanks to its
leavening capacity through alcoholic fermentation (Musatti
et al. 2015, 2016, 2018). It turned out that Z. mobilis could
be used to substitute S. cerevisiae for the production of
baked goods.
The application of non-conventional yeasts as to their
capacity to dough leavening, metabolism of fermentable
oligosaccharides, disaccharides, monosaccharides, and polyols
(FODMAPs), and flavor formation during bread making is
of increasing interest (Aslankoohi et al. 2016; De Vuyst et
al. 2016; Struyf et al. 2017, 2018; Zhou et al. 2017; Fraberger
Critical Reviews in Food Science and Nutrition 13
et al. 2018). Yeast species belonging to the genera
Brettanomyces/Dekkera, Kazachstania, Kluyveromyces,
Lachancea, Meyerozyma, Pichia, Saccharomyces, Torulaspora,
and Wickerhamomyces have been tested. Further, it is pos-
sible to produce Panettone by means of a co-culture of Liml.
fermentum and W. anomalus strains (Stefanello et al. 2019).
Finally, if sourdoughs are used as leavening agent, carbon
dioxide production by the fermentation microbiota is the
determining selection criterion (Guerzoni et al. 2013; Gänzle
and Zheng 2019; Carbonetto et al. 2020; Palla et al. 2020;
Bigey et al. 2021). In general, yeasts have a greater leavening
capacity than heterofermentative LAB species. Further, func-
tional yeast starter cultures may contribute to nutritional
(e.g., production of vitamins) and functional traits (e.g.,
phytase activity to degrade phytate and anti-oxidant activity
by elaborating phenolic compounds) as well (Palla et al.
2020). Interestingly, sourdough strains of S. cerevisiae are
better adapted to the sourdough environment thanks to an
increased growth on maltose (Bigey et al. 2021).
Sourdough productions with added non-flour
ingredients
Since ancient times, many non-flour ingredients have been
used to produce doughs for baked goods. Already the
Egyptians used beer foam (unconsciously as a source of
yeasts) to leaven their doughs to make their breads soft and
light (Cappelle et al. 2013). A document of Pliny the Elder
(Historia Naturalis) from the first century A.D. refers to the
use of grapes for bread production.
Nowadays, as in the past, when started from scratch,
sourdough production processes of Type 1/I are usually
initiated with the spontaneous fermentation of a cereal
flour-water mixture, whether or not supplemented with
non-flour ingredients, which is then followed by a
long-lasting backslopping process. Indeed, these backslopping
steps are often carried out until the baker passes the sour-
dough on to the next owner of the bakery, who then
refreshes the sourdough through backslopping until it is
passed on again. Some sourdoughs have a documented age
of more than 100 years of backslopping, representing >
100,000 propagation steps (Gänzle and Ripari 2016; Gänzle
and Zheng 2019). The one and only spontaneous step that
may or may not have been used to start the backslopping
cycle is insignificant in this respect, as long as the mature
sourdough is robust enough to withstand new refreshments,
either to maintain the sourdough or to switch to other flour
and/or non-flour ingredients, DYs, etc. (Di Cagno et al.
2014; Lhomme et al. 2014; Comasio, Van Kerrebroeck, and
De Vuyst 2021). Yet, ingredients, environmental conditions,
fermentation strategies, sourdough maintenance practices,
and (multi)species interactions can impact the taxonomic
structure of the microbial communities involved (Van
Kerrebroeck, Maes, and De Vuyst 2017; Landis et al. 2021).
To improve the nutritional value of the breads produced,
wholemeal flours or bran fractions are sometimes used
(Rizzello et al. 2012b; Caputo, Visconti, and De Angelis
2015; Coda, Katina, and Rizzello 2015; Clément et al. 2018;
14 L. DE VUYST ET AL.
De Angelis et al. 2019). However, other, non-flour ingredi-
ents daily used in the kitchen can be added to the flour-water
mixture as well, such as fruits, herbs, honey, milk, salt,
sugar, and yoghurt (Table 1). Some of these ingredients may
be the source of certain microorganisms and, hence, serve
as an additional microbial inoculum, whereas other ingre-
dients may be the source of substrates and/or co-substrates
for microbial fermentation and conversion and, thus, acti-
vating or selecting microorganisms that are naturally present,
either in the flour or in the ingredients (Table 2).
Consequently, applying an appropriate functional starter
culture and/or dedicated ingredients in Type 2/II sourdough
production processes may have an influence on not only
the microbial composition but also the organoleptic prop-
erties of a sourdough, in particular its aroma and taste
profiles (Minervini et al. 2012b; Minervini et al. 2014, 2016).
The effect of certain compounds on the flavor of sour-
doughs is directly linked with the strains’ metabolism or
as a result of subprocesses taking place during flour fer-
mentation (e.g., cell lysis). Concerning the former, lactic
acid (soft acidity) and acetic acid (sharp acidity) as
end-metabolites of carbohydrate breakdown contribute to
the acidic taste of sourdoughs, whereas the volatile acetic
acid further contributes to their aroma (Corsetti 2013;
Gänzle 2014; Pico, Bernal, and Gómez 2015; De Vuyst, Van
Kerrebroeck, and Leroy 2017). Similarly, the accumulation
of certain amino acids, such as glutamate (e.g., by Liml.
reuteri as a response to acidic stress through glutaminase
activity on glutamine, an amino acid that is liberated at
high concentrations through wheat flour proteolysis), and
taste-active peptides (e.g., glutathione accumulation and
glutamyl dipeptide formation by Liml. reuteri, because of
the high abundance of oxidized glutathione and glutamate,
isoleucine and leucine in the flour, respectively) contribute
to the umami taste and kokumi taste, respectively (Thiele,
Gänzle, and Vogel 2002; Vermeulen, Gänzle, and Vogel
2007; Gänzle 2014; Pico, Bernal, and Gómez 2015; Zhao
et al. 2015; Zhao and Gänzle 2016; Cavallo et al. 2017).
The release of amino acids and other compounds (e.g.,
minerals and vitamins) can also be the result of expulsion
or cell lysis. For instance, amino acid expulsion by LAB
follows the enhanced proteolysis by the flour proteases as
a result of acidification and peptidase activity (Di Cagno
et al. 2002; Rollán et al. 2005; Gänzle, Vermeulen, and
Vogel 2007; Gänzle, Loponen, and Gobbetti 2008; Gänzle
et al. 2009; Gobbetti et al. 2014). Yeast cell lysis can sustain
LAB growth and enhance sourdough flavor thanks to the
release of amino acids, peptides, and vitamins (Gobbetti
1998). Moreover, lysed yeast cells excrete ornithine and
glutathione, influencing the bread taste and aroma (Vrancken
et al. 2011; Yan et al. 2018).
Sourdough productions with the addition of pure
chemical compounds
A first type of ingredients that can be added to the
flour-water mixtures are pure chemical compounds, which
has been studied mainly during starter culture-initiated
sourdough productions carried out in the laboratory (using
either strains or mature sourdoughs; Table 3) or making
use of simulation media (Gobbetti, Corsetti, and Rossi 1995;
Neysens, Messens, and De Vuyst 2003; Vermeulen, Gänzle,
and Vogel 2006). The starter culture strain used can either
consume or convert these chemical compounds. Alternatively,
its growth can be activated by these chemical compounds.
In all these cases, a desirable effect on the mature sour-
doughs can be generated. Typical examples, whether or not
in the research phase, are the addition of salt (NaCl), car-
bohydrates (fructose, sucrose, pentoses, and pentosans),
amino acids (arginine and glutamate), and other organic
acids (citrate) (Table 3). The concomitant starter culture
strains used belong to certain LAB, yeast, and even AAB
species.
Salt
Salt is a common ingredient used in the production of
bread. For instance, the production of Pugliese sourdough
bread includes the addition of salt to the flour-water mixture
at each refreshment step of the sourdough prior to bread-
making (Ottogalli, Galli, and Foschino 1996). Salt evidently
contributes to the salty taste of bread. Also, salt has a direct
effect on the microbial ecosystem of a sourdough matrix,
as too high concentrations of salt inhibit the growth of
many microorganisms. In contrast, the growth of certain
LAB species is often favored in the presence of salt, a com-
mon practice for the use of salt to control food fermentation
processes involving LAB, whereas that of, for instance,
enterobacteria is inhibited (Wouters et al. 2013). As an
example, it has been shown that the maximal specific growth
rate and cell yield of Lb. amylovorus DCE 471 increases
with increasing salt concentrations (up to a concentration
of 0.5 % NaCl) in a wheat sourdough simulation medium
(Neysens, Messens, and De Vuyst 2003). Similarly, LAB are
stimulated in a wheat sourdough initiated with a ripe rye
sourdough by low salt concentrations (up to 0.7 % NaCl),
whereas higher concentrations negatively affect LAB and,
to a lesser extent, yeast growth (Simonson, Salovaara, and
Korhola 2003). Alternatively, high salt concentrations (up
to 4.0 %) activate the ADI pathway of the heterofermentative
Liml. fermentum IMDO 130101 strain (Vrancken et al.
2009a; Verce, De Vuyst, and Weckx 2020). Yeasts generally
tolerate low water activity values or increased osmotic pres-
sure better than LAB species and, hence, display a higher
tolerance toward salt (Minervini et al. 2015; De Vuyst et
al. 2016). It may explain the better fitness of W. anomalus
compared to S. cerevisiae in spontaneous backslopped lab-
oratory sourdoughs (Vrancken et al. 2010; Daniel et al. 2011;
Landis et al. 2021). Also, K. humilis is not inhibited by the
addition of 4.0 % salt, whereas Frul. sanfranciscensis does
not tolerate high salt concentrations, which has been demon-
strated in a rye sourdough fermented with Frul. sanfrancis-
censis LTH 2581 and K. humilis LTH H198 (Brandt, Hammes,
and Gänzle 2004). Finally, salt impacts the properties of the
bread dough, through its impact on the positively charged
gluten proteins to form a stronger dough, and bread (Miller
and Hoseney 2008).
 Critical Reviews in Food Science and Nutrition 15

Table 1. Non-exhaustive overview of the lactic acid bacteria (LAB), acetic acid bacteria (AAB), and yeast species diversity of backslopped sourdough productions
(Type I), spontaneously initiated with flour and non-flour ingredients (and propagation in the case of NaCl addition). The data represent the last sampling point
of each backslopping process. The microorganisms were identified culture-independently (CI) and/or culture-dependently [molecular approach (CD) or pheno-
typical approach (CD*)].
Species diversity
AAB (A., Acetobacter; G., Gluconobacter);
LAB (Coml., Companilactobacillus; E.,
Enterococcus; Frul., Fructilactobacillus; Furl.,
Furfurilactobacillus; Lacc., Lacticaseibacillus;
Lacp., Lactiplantibacillus; Lc., Lactococcus;
Latl., Latilactobacillus; Leuc., Leuconostoc;
Levl., Levilactobacillus; Liml.,
Limosilactobacillus; P., Pediococcus; W.,
Ingredient Weissella);
(number of Dough Fermentation Fermentation Yeasts (K., Kazachstania; Pi., Pichia; S.,
sourdoughs if > yield temperature time Type of Saccharomyces; T., Torulaspora; Y.,
1) Cereal flour (DY) (°C) (h) producer Yarrowia). Reference
Apple Wheat na na na Bakery LAB: Levl. brevis (CD), Latl. graminis (CD), Ripari, Gänzle, et al.
Latl. sakei (CD) (2016)
AAB: A. cerevisiae (CD)
Yeasts: S. cerevisiae (CD)
Apple Wheat na na na Bakery LAB: Lactiplantibacillus species (CD) Ripari, Gänzle, et al.
AAB: A. cerevisiae (CD) (2016)
Yeasts: S. cerevisiae (CD), S. unisporus
(now K. unispora; CD)
Apple Wheat 150 29 4-4-16 Laboratory LAB: Lacp. plantarum (CD), Latl. sakei (CD), Gordún et al.
P. pentosaceus (CD) (2015)
Yeasts: Saccharomyces group
Apple flowers Wheat 200 20–25 48 Laboratory LAB: Lactiplantibacillus species (CD, CI), Furl. Ripari, Gänzle, et al.
rossiae (CD, CI) (2016)
Yeasts: S. cerevisiae (CI)
Apple flowers Wheat 200 30 24 Laboratory LAB: Lactiplantibacillus species (CD), Furl. Ripari, Cecchi, et al.
rossiae (CD) (2016)
Yeasts: S. cerevisiae (CD)
Apple pulp Wheat na na na Bakery LAB: Lactiplantibacillus species (CD), Latl. Ripari, Gänzle, et al.
sakei (CD), P. pentosaceus (CD) (2016)
AAB: A. cerevisiae (CD)
Yeasts: S. cerevisiae (CD)
Baker’s yeast Durum wheat 160 30 5 Laboratory LAB: Levl. brevis (CD), Latl. curvatus (CD), Minervini et al.
Liml. fermentum (CD), Lacp. plantarum (2016)
(CD)
Yeasts: S. cerevisiae (CD)
Basil (2) Wheat na RT 24 Household Yeasts: P. membranifaciens (CD), S. Paramithiotis et al.
cerevisiae (CD) (2000)
Basil Wheat na RT 24 Household Yeasts: S. cerevisiae (CD), Y. lipolytica (CD) Paramithiotis et al.
(2000)
Basil (2) Wheat na RT 24 Bakery Yeasts: S. cerevisiae (CD) Paramithiotis et al.
(2000)
Basil Wheat na RT na Household LAB: Lacp. plantarum (CD), P. pentosaceus Paramithiotis,
(CD) Tsiasiotou, and
Yeasts: T. delbrueckii (CD), S. cerevisiae Drosinos (2010)
(CD)
Basil (7) Wheat na RT na Household LAB: Lacp. plantarum (CD) Paramithiotis,
Yeasts: T. delbrueckii (CD) Tsiasiotou, and
Drosinos (2010)
Basil (2) Wheat na RT na Household LAB: Lacp. plantarum (CD), P. pentosaceus Paramithiotis,
(CD) Tsiasiotou, and
Yeasts: T. delbrueckii (CD) Drosinos (2010)
Blueberries Wheat na 25 6 Laboratory LAB: Leuc. mesenteroides group (CD*) Vilanova et al.
(2015)
Cow manure Wheat na na na Bakery LAB: Lacp. plantarum, Frul. sanfranciscensis Ripari, Gänzle, et al.
(CD) (2016)
Yeasts: S. cerevisiae (CD)
Crystal pear Wheat 200 RT 24 Laboratory LAB: Levl. brevis (CD), Lacp. plantarum (CD), Yu et al. (2018)
Furl. rossiae (CD)
Yeasts: S. cerevisiae (CD)
Dry apricot Wheat na 25 6 Laboratory LAB: Leuc. mesenteroides group (CD*) Vilanova et al.
(2015)
Fruits Wheat na na na Bakery LAB: Levl. brevis (CD), Lactiplantibacillus Ripari, Gänzle, et al.
species (CD), Furl. rossiae (CD), Leuc. (2016)
holzapfelii (CD), Leuc.
pseudomesenteroides (CD), Leuc. kimchii
(CD), W. cibaria (CD)
AAB: A. cerevisiae (CD)
Yeasts: S. cerevisiae (CD)

(Continued)
16 L. DE VUYST ET AL.

Table 1. (Continued ).
Species diversity
AAB (A., Acetobacter; G., Gluconobacter);
LAB (Coml., Companilactobacillus; E.,
Enterococcus; Frul., Fructilactobacillus; Furl.,
Furfurilactobacillus; Lacc., Lacticaseibacillus;
Lacp., Lactiplantibacillus; Lc., Lactococcus;
Latl., Latilactobacillus; Leuc., Leuconostoc;
Levl., Levilactobacillus; Liml.,
Limosilactobacillus; P., Pediococcus; W.,
Ingredient Weissella);
(number of Dough Fermentation Fermentation Yeasts (K., Kazachstania; Pi., Pichia; S.,
sourdoughs if > Cereal yield temperature time Type of Saccharomyces; T., Torulaspora; Y.,
1) flour (DY) (°C) (h) producer Yarrowia). Reference
Grape juice or Durum wheat na na 24 Bakery LAB: Lacp. pentosus (CD) Randazzo et al.
honey (2) (2005)
Grape juice or Durum wheat na na 24 Bakery LAB: Lacc. casei (CD), Coml. kimchii (CD), Randazzo et al.
honey Lacp. pentosus (CD) (2005)
Grape juice or Durum wheat na na 24 Bakery LAB: Lacp. pentosus (CD), Frul. Randazzo et al.
honey (3) sanfranciscensis (CD) (2005)
Grape juice or Durum wheat na na 24 Bakery LAB: Lacp. plantarum (CD) Randazzo et al.
honey (2005)
Grape juice or Durum wheat na na 24 Bakery LAB: Coml. kimchii (CD), Lacp. pentosus Randazzo et al.
honey (CD), Frul. sanfranciscensis (CD) (2005)
Grape juice or Durum wheat na na 24 Bakery LAB: Lacc. casei (CD), Lacp. plantarum (CD) Randazzo et al.
honey (2005)
Grape must Durum wheat 160 30 5 Laboratory LAB: Lacp. plantarum (CD), Leuc. Minervini et al.
mesenteroides (CD) (2016)
Yeasts: S. cerevisiae (CD)
Grapes Wheat 150 29 4-4-16 Laboratory LAB: Lacp. plantarum (CD), Latl. sakei (CD), Gordún et al.
P. pentosaceus (CD) (2015)
Yeasts: Saccharomyces group
Grapes Wheat + rye 167 5–25 24 Laboratory LAB: Lacc. casei/rhamnosus/paracasei (CD) Aplevicz et al.
Yeasts: S. cerevisiae (CD) (2014b)
Hawthorn Wheat 200 20–25 48 Laboratory LAB: Frul. sanfranciscensis (CD, CI) Ripari, Gänzle, et al.
berries Yeasts: unidentified species (CI) (2016)
Honey Durum wheat 160 30 5 Laboratory LAB: Lacp. plantarum (CD) Minervini et al.
Yeasts: S. cerevisiae (CD) (2016)
Honey Wheat na na na Bakery LAB: Lactiplantibacillus species (CD), P. Ripari, Gänzle, et al.
pentosaceus (CD), W. cibaria (CD) (2016)
Yeasts: S. cerevisiae (CD), W. anomalus
(CD)
Honey Wheat na na na Bakery LAB: Latl. graminis (CD), P. pentosaceus (CD) Ripari, Gänzle, et al.
Yeasts: S. cerevisiae (CD) (2016)
Honey Wheat na na na Bakery LAB: Levl. brevis (CD), P. pentosaceus (CD) Ripari, Gänzle, et al.
Yeasts: S. cerevisiae (CD) (2016)
Honey Wheat na na na Bakery LAB: Latl. graminis (CD), P. pentosaceus (CD) Ripari, Gänzle, et al.
Yeasts: S. cerevisiae (CD), S. barnettii (2016)
(now K. barnettii) (CD)
Honey Wheat na na na Bakery LAB: Frul. sanfranciscensis (CD), Leuc. Ripari, Gänzle, et al.
holzapfelii (CD), Leuc. mesenteroides (2016)
(CD), Leuc. kimchii (CD), W. confusa (CD)
Yeasts: S. cerevisiae (CD)
Macerated pears Durum wheat 160 30 5 Laboratory LAB: Lacp. pentosus (CD), Lacp. plantarum Minervini et al.
(CD) (2016)
Yeasts: S. cerevisiae (CD)
Mother of Wheat 200 20–25 48 Laboratory LAB: Frul. sanfranciscensis (CD, CI) Ripari, Gänzle, et al.
vinegar Yeasts: S. cerevisiae (CI) (2016)
Mother of Wheat 200 30 24 Laboratory LAB: Frul. sanfranciscensis (CD) Ripari, Cecchi, et al.
vinegar Yeasts: S. cerevisiae (CD) (2016)
Myrtle berries Wheat 200 20–25 48 Laboratory LAB: Frul. sanfranciscensis (CD, CI) Ripari, Gänzle, et al.
Yeasts: S. cerevisiae (CI) (2016)
Myrtle berries Wheat 200 30 24 Laboratory LAB: Frul. sanfranciscensis (CD) Ripari, Cecchi, et al.
Yeasts: S. cerevisiae (CD) (2016)
NaCl Durum wheat 150 25 13 Bakery LAB: Lacp. plantarum (CD), Latl. sakei (CD), Minervini et al.
Leuc. citreum (CD) (2012a)
Yeasts: S. cerevisiae (CI)
NaCl Durum wheat 160 25 6 Bakery LAB: Frul. sanfranciscensis (CD) Minervini et al.
Yeasts: K. humilis/barnettii (CI) (2012a)
NaCl Durum wheat 160 25 10 Bakery LAB: W. cibaria (CD), W. confusa (CD) Minervini et al.
Yeasts: S. cerevisiae (CD) (2012b)
NaCl Durum wheat 160 25 6 Bakery LAB: Frul. sanfranciscensis (CD), Leuc. Minervini et al.
citreum (CD) (2012b)
Yeasts: S. cerevisiae (CD)
 Critical Reviews in Food Science and Nutrition 17

Table 1. (Continued ).
Species diversity
AAB (A., Acetobacter; G., Gluconobacter);
LAB (Coml., Companilactobacillus; E.,
Enterococcus; Frul., Fructilactobacillus; Furl.,
Furfurilactobacillus; Lacc., Lacticaseibacillus;
Lacp., Lactiplantibacillus; Lc., Lactococcus;
Latl., Latilactobacillus; Leuc., Leuconostoc;
Levl., Levilactobacillus; Liml.,
Limosilactobacillus; P., Pediococcus; W.,
Ingredient Weissella);
(number of Dough Fermentation Fermentation Yeasts (K., Kazachstania; Pi., Pichia; S.,
sourdoughs if > Cereal yield temperature time Type of Saccharomyces; T., Torulaspora; Y.,
1) flour (DY) (°C) (h) producer Yarrowia). Reference
NaCl Durum wheat 160 25 6 Bakery LAB: Frul. sanfranciscensis (CD), Lacp. Minervini et al.
plantarum (CD), Leuc. citreum (CD) (2012b)
Yeasts: S. cerevisiae (CD)
NaCl Wheat + durum 160 25 6 Bakery Yeasts: S. cerevisiae (CD) Minervini et al.
wheat (2012b)
NaCl Wheat 160 22 6 Bakery LAB: Latl. sakei (CD), Frul. sanfranciscensis Minervini et al.
(CD), Leuc. mesenteroides subsp. (2012b)
mesenteroides (CD)
Yeasts: S. cerevisiae (CD)
NaCl Wheat 160 22 10 Bakery LAB: Frul. sanfranciscensis (CD) Minervini et al.
Yeasts: S. cerevisiae (CD) (2012b)
NaCl Wheat 160 22 12 Bakery LAB: Coml. paralimentarius (CD), Lacp. Minervini et al.
plantarum (CD) (2012b)
Yeasts: S. cerevisiae (CD)
NaCl Wheat 160 23 6 Bakery LAB: Lacc. casei (CD), Lacp. plantarum (CD), Minervini et al.
Frul. sanfranciscensis (CD), P. inopinatus (2012b)
(CD)
Yeasts: S. cerevisiae (CD)
NaCl Durum wheat 160 26 4 Bakery LAB: Lacp. plantarum (CD), Frul. Minervini et al.
sanfranciscensis (CD) (2012b)
Yeasts: K. humilis (CD)
NaCl Durum wheat 160 22 11 Bakery LAB: Levl. brevis (CD), Lacp. plantarum (CD), Minervini et al.
Furl. rossiae (CD), Frul. sanfranciscensis (2012b)
(CD), P. argentinicus (CD)
Yeasts: S. cerevisiae (CD)
NaCl Durum wheat 160 27 11 Bakery LAB: Levl. brevis (CD), Lacp. plantarum (CD), Minervini et al.
Frul. sanfranciscensis (CD), Lc. lactis (2012b)
subsp. lactis (CD), Leuc. mesenteroides
subsp. mesenteroides (CD), P.
pentosaceus (CD), W. paramesenteroides
(CD)
Yeasts: S. cerevisiae (CD)
NaCl Durum wheat 160 26 10 Bakery LAB: Coml. paralimentarius (CD), Levl. Minervini et al.
spicheri (CD) (2012b)
Yeasts: K. barnettii (CD), K. exigua (CD)
NaCl Durum wheat 160 25 11 Bakery LAB: Levl. namurensis (CD), Coml. Minervini et al.
paralimentarius (CD), Lacp. plantarum (2012b)
(CD), Frul. sanfranciscensis (CD)
Yeasts: S. cerevisiae (CD)
NaCl Durum wheat 160 22 5 Bakery LAB: E. durans (CD), Frul. sanfranciscensis Minervini et al.
(CD) (2012b)
Yeasts: K. humilis (CD)
NaCl + baker’s Wheat 160 30 6 Bakery LAB: Lacp. plantarum (CD), P. pentosaceus Minervini et al.
yeast (CD) (2012b)
Yeasts: S. cerevisiae (CD)
NaCl + baker’s Wheat 160 27 18 Bakery LAB: Coml. paralimentarius (CD), Lacp. Minervini et al.
yeast plantarum (CD), Leuc. mesenteroides (2012b)
subsp. mesenteroides (CD)
Yeasts: S. cerevisiae (CD)
Navel orange Wheat 200 RT 24 Laboratory LAB: Levl. brevis (CD), Lacp. plantarum (CD), Yu et al. (2018)
Furl. rossiae (CD)
Yeasts: S. cerevisiae (CD)
Pomegranate Wheat 200 20–25 48 Laboratory LAB: Lactiplantibacillus species (CD, CI), Frul. Ripari, Gänzle, et al.
sanfranciscensis (CI) (2016)
Yeasts: S. cerevisiae (CI)
Pomegranate Wheat 200 30 24 Laboratory LAB: Lactiplantibacillus species (CD) Ripari, Cecchi, et al.
Yeasts: S. cerevisiae (CD) (2016)
Rye cider Wheat na 25 6 Laboratory LAB: Leuc. mesenteroides group (CD*) Vilanova et al.
(2015)
Speedwell Wheat 200 20–25 48 Laboratory LAB: Frul. sanfranciscensis (CD, CI) Ripari, Gänzle, et al.
flowers AAB: G. cerinus (CD, CI) (2016)
(Continued)
18 L. DE VUYST ET AL.

Table 1. (Continued ).
Species diversity
AAB (A., Acetobacter; G., Gluconobacter);
LAB (Coml., Companilactobacillus; E.,
Enterococcus; Frul., Fructilactobacillus; Furl.,
Furfurilactobacillus; Lacc., Lacticaseibacillus;
Lacp., Lactiplantibacillus; Lc., Lactococcus;
Latl., Latilactobacillus; Leuc., Leuconostoc;
Levl., Levilactobacillus; Liml.,
Limosilactobacillus; P., Pediococcus; W.,
Ingredient Weissella);
(number of Dough Fermentation Fermentation Yeasts (K., Kazachstania; Pi., Pichia; S.,
sourdoughs if > Cereal yield temperature time Type of Saccharomyces; T., Torulaspora; Y.,
1) flour (DY) (°C) (h) producer Yarrowia). Reference
Speedwell Wheat 200 30 24 Laboratory LAB: Frul. sanfranciscensis (CD) Ripari, Cecchi, et al.
flowers Yeasts: S. cerevisiae (CD) (2016)
Vinegar Wheat na na na Bakery LAB: Lactiplantibacillus species (CD), Furl. Ripari, Gänzle, et al.
rossiae (CD), Leuc. mesenteroides (CD) (2016)
Yeasts: S. cerevisiae (CD)
Water from Durum wheat 160 30 5 Laboratory LAB: Lacp. plantarum (CD) Minervini et al.
macerate Yeasts: S. cerevisiae (CD) (2016)
pears
White cider Wheat na 25 6 Laboratory LAB: Leuc. mesenteroides group (CD*) Vilanova et al.
(2015)
White mustard Wheat 200 20–25 48 Laboratory LAB: Latl. graminis (CD, CI) Ripari, Gänzle, et al.
flowers Yeasts: unidentified species (CI) (2016)
White mustard Wheat 200 30 24 Laboratory LAB: Latl. graminis (CD) Ripari, Cecchi, et al.
flowers Yeasts: S. cerevisiae (CD) (2016)
Yoghurt Wheat na na na Bakery LAB: Leuc. holzapfelii (CD), Leuc. Ripari, Gänzle, et al.
mesenteroides (CD), P. pentosaceus (CD), (2016)
W. cibaria (CD)
Yeasts: S. cerevisiae (CD)
Yoghurt Wheat na na na Bakery LAB: Frul. sanfranciscensis (CD), Leuc. Ripari, Gänzle, et al.
holzapfelii (CD), W. cibaria (CD) (2016)
Yeasts: S. cerevisiae (CD)
Yoghurt Wheat na na na Bakery LAB: Lacc. paracasei (CD), Lactiplantibacillus Ripari, Gänzle, et al.
species (CD), Furl. rossiae (CD), Leuc. (2016)
holzapfelii (CD), Leuc. mesenteroides
(CD), P. pentosaceus (CD)
Yeasts: S. cerevisiae (CD)
Yoghurt Wheat 150 29 4-4-16 Laboratory LAB: Levl. brevis (CD), Lacp. plantarum (CD), Gordún et al.
Latl. sakei (CD), P. pentosaceus (CD), W. (2015)
cibaria (CD)
Yeasts: Saccharomyces group
na, not available; RT, room temperature.

Table 2. Non-exhaustive overview of laboratory sourdough productions (Type 2) initiated with a lactic acid bacterial starter culture to study the effects of the
addition of non-flour ingredients.
Dough Fermentation Fermentation
yield temperature time
Ingredient Starter culture Cereal flour (DY) (°C) (h) Effect Reference
Apple juice Companilactobacillus Wheat 200 30 72 or 10 days of Enhanced production of Comasio, Van
crustorum LMG 24 h lactic acid Kerrebroeck,
23699 backslopping Increased lactic acid and De Vuyst
bacterial population (2021)
Honey Pediococcus Wheat + rye 200 32 19 Influence on bread dough Nutter et al.
pentosaceus CRL development (only Liml. (2017)
922 and fermentum)
Limosilactobacillus
fermentum CRL
220
Lemon juice Coml. crustorum LMG Wheat 200 30 72 or 10 days of Enhanced production of Comasio, Van
23699 24 h acetic acid, lactic acid, Kerrebroeck,
backslopping acetoin, and diacetyl and De Vuyst
(2021)
Milk whey Lactobacillus Wheat or 200 30 24 More exopolysaccharides Galle et al. (2011)
buchneri FUA 3154 sorghum
 Critical Reviews in Food Science and Nutrition 19

Table 3. Non-exhaustive overview of laboratory sourdough productions (Type 2) initiated with a lactic acid bacterial and/or yeast starter culture or a mature
sourdough to study the effects of the addition of pure chemical compounds.
Dough Fermentation Fermentation
Cereal yield temperature time
Compound Starter culture flour (DY) (°C) (h) Effect Reference
Arginine Furfurilactobacillus Wheat 160 30 6 More ornithine De Angelis et al.
rossiae CB1 and 7 and 12 Increased growth (2002)
(originally rate and enhanced
identified as tolerance to acidic
Fructilactobacillus stress
sanfranciscensis Improved cell
CB1) survival during
storage
Citrate Companilactobacillus Wheat 200–400 30 72 More lactic acid, Van Kerrebroeck
crustorum LMG acetoin, and et al. (2018a,
23699 diacetyl 2018b);
Comasio et
al. (2019)
Citrate Furl. rossiae CB1 Wheat 148 28 8 Increased growth rate Gobbetti and
More lactic acid Corsetti
and acetic acid 1996)
Reduced
fermentation
quotient (ratio of
lactic acid to
acetic acid)
Less ethanol
Fructose Furl. rossiae CB1 Wheat 148 28 8 Higher TTA Gobbetti,
More acetic acid Corsetti, and
Reduced Rossi (1995)
fermentation
quotient
Fructose Levilactobacillus Wheat 180–320 25–40 20 More acetic acid Röcken, Rick,
brevis 25a Reduced and
fermentation Reinkemeier
quotient (1992)
α-Ketoglutaric acid Frul. sanfranciscensis Wheat 200 30–37 48 More lactic acid and Zhang and
DSM 20451 acetic acid Gänzle
Limosilactobacillus More (2010)
reuteri LTH 2584 2-hydroxyglutarate
NaCl Frul. sanfranciscensis Rye 200 28 24 Inhibited growth of Brandt,
LTH 2581 and Frul. Hammes,
Kazachstania sanfranciscensis and Gänzle
humilis LTH H198 and less lactic (2004)
acid with high
NaCl
concentrations
NaCl Rye sourdough Wheat 220 28 24 Enhanced LAB Simonson,
growth with low Salovaara,
NaCl and Korhola
concentrations (2003)
Inhibited growth
of yeasts and LAB
with high NaCl
concentrations
Pentosans and Lactobacillus hilgardii Wheat 135–220 30 24 More acetic acid Gobbetti et al.
endo-xylanase S32 Increased (1999)
acidification rate
and TTA
Pentosans and Lactiplantibacillus Wheat 170 30 8 More acetic acid Gobbetti et al.
α-L- plantarum 20B Increased (2000)
arabinofuranosidase acidification rate
and TTA
Sucrose Frul. sanfranciscensis Wheat-rye 200 30 24 More OS, acetate, Korakli et al.
LTH 2590 and mannitol (2001)
Sucrose Lactobacillaceae spp. Wheat 200 30–37 24 More OS Tieking et al.
(2003)
TTA, total titratable acidity; LAB, lactic acid bacteria; OS, oligosaccharides.

Fructose as alternative external electron acceptor by heterofermen-

The addition of fructose to the flour-water mixture allows tative LAB species, thereby producing mannitol (Figure 4;
the production of higher concentrations of acetic acid (and Röcken, Rick, and Reinkemeier 1992; De Vuyst et al. 2002).
lower concentrations of ethanol), as fructose can be used This has been shown during co-metabolism of maltose and
20 L. DE VUYST ET AL.

Figure 6.  Co-metabolism of citrate or malate and maltose in lactic acid bacteria (LAB): citrate metabolism starts with its conversion into acetate and oxaloac-
etate by a citrate lyase; oxaloacetate can be further converted into succinate (common route for heterofermentative LAB species) via the reductive branch of
the tricarboxylic acid cycle or to pyruvate by an oxaloacetate decarboxylase; pyruvate can be further converted into lactic acid or α-acetolactate with a lactate
dehydrogenase or an α-acetolactate synthase (releasing carbon dioxide), respectively, the latter being converted into acetoin enzymatically (acetolactate decar-
boxylase) or diacetyl non-enzymatically (oxidative decarboxylation; dashed arrow). Malate can be converted into lactate by a malolactic enzyme (releasing
carbon dioxide). The most relevant intermediates and end-products (shown in bold) as well as the use and regeneration of the cofactor NAD+ are represented
(based on Gänzle 2015).

fructose in heterofermentative LAB species (Gobbetti, Arginine

Corsetti, and Rossi 1995; Stolz, Böcker, et al. 1995; Stolz,
Vogel, et al. 1995). Acidity due to the presence of acetic
acid contributes to the aroma and taste of sourdough breads.
Sucrose
The addition of sucrose that activates levansucrase activity
for oligosaccharide production (Figure 4), thereby accumu-
lating glucose, also supports acetate formation by hetero-
fermentative LAB species, such as in the case of Frul.
sanfranciscensis (Korakli et al. 2001; Tieking et al. 2003;
Tieking and Gänzle 2005; Korakli and Vogel 2006; Galle et
al. 2010). However, high acetic acid concentrations accom-
panied with oligosaccharide production may have an adverse
effect on the bread texture (Kaditzky et al. 2008).
Pentoses and pentosans
The addition of pentoses also allows the enhancement of
the production of acetic acid. For instance, the
co-fermentation of arabinose, xylose, or ribose and maltose
by appropriate strains of LAB species makes this possible
(Gobbetti et al. 1999, 2000). Therefore, arabinoxylans (as
source of arabinose that is present in branches on the xylan
backbone chain) may be added to the wheat flour-water
mixture (as the arabinoxylan concentration is low in wheat
flour) together with α-L-arabinofuranosidase, an enzyme
that is capable to elaborate arabinose from the xylan back-
bone chain and that is rarely produced by LAB species
(Michlmayr et al. 2013; Linares-Pastén et al. 2017).
Heterofermentative LAB species can prefer arabinose in the
presence of maltose. Arabinose enables a higher specific
growth rate and cell yield, thereby causing a faster acidifi-
cation of the sourdough matrix and an enhanced production
of acetic acid. This can be ascribed to the higher production
of ATP (Figure 4; Gänzle 2015).
The addition of arginine to the flour-water mixture allows
the production of ornithine via the ADI pathway in mainly
heterofermentative LAB species (Figure 4; Gänzle and
Gobbetti 2013; De Vuyst, Van Kerrebroeck, and Leroy 2017;
Weckx, Van Kerrebroeck, and De Vuyst 2019). In particular,
this pathway enhances the acidic stress response (through
ammonia production) and the strains’ competitiveness
(through extra ATP formation) in species of Limosilactobacillus
(not in Frul. sanfranciscensis). Ornithine is a precursor for
the production of the crust aroma compound
2-acetyl-1-pyrroline that is formed through Maillard reaction
during the baking process (Schieberle 1990; Grosch and
Schieberle 1997; Thiele, Gänzle, and Vogel 2002; Fernández
and Zúñiga 2006; Pacyński, Zawirska-Wojtasiak, and
Mildner-Szkudlarz 2015). The ADI pathway in heterofer-
mentative LAB species has been studied in detail for several
strains, among which Liml. fermentum IMDO 130101, for
which it turned out that the environmental pH determines
the release of citrulline, an intermediate of the ADI pathway,
and ornithine, the end-product of the ADI pathway
(Vrancken et al. 2009a, 2009b). Addition of a small con-
centration of arginine (6 mM) to the flour-water mixture
enhances the growth and survival of a strain of Furl. rossiae
(described as Frul. sanfranciscensis CB1 in the original
paper), which produces more ornithine compared to a fer-
mentation without added arginine (De Angelis et al. 2002).
An increased roasty note has been found with the addition
of Liml. pontis DSM 8475 to a wheat flour-water mixture,
which is ascribed to ornithine production (Thiele, Gänzle,
and Vogel 2002).
Glutamate
Doughs chemically acidified with glutamate (47 mmol/kg)
significantly change the taste of breads. Moreover, during
starter culture-initiated fermentations of flour with high
proteolytic activity (obtained through fungal protease

addition), higher concentrations of glutamate are produced
(via deamination of glutamine) in a strain-dependent man-
ner, enhancing the acid tolerance of the LAB and improving
the taste of the breads produced thereof (Vermeulen, Gänzle,
and Vogel 2007). Heterofermentative LAB species, such as
Frul. sanfranciscensis and Liml. reuteri, can impact the pro-
duction of aroma compounds through their transamination
pathway (Vermeulen, Gänzle, and Vogel 2006; Zhang and
Gänzle 2010). For instance, the co-fermentation of maltose
as energy source and α-ketoglutarate as electron acceptor
instead of amino group acceptor leads to the production of
2-hydroxyglutarate instead of an α-keto acid and glutamate,
the latter being favored in the presence of an amino group
donor (e.g., phenylalanine) and citrate. Also, it enables a
faster growth of the appropriate strains as well as the pro-
duction of more acetic acid at the cost of ethanol.
Citrate
The addition of citrate in the presence of a citrate-positive
LAB strain, for instance the homofermentative Coml. crus-
torum LMG 23699, enables the production of not only acetic
acid and lactic acid, but also acetoin and diacetyl (Figures
4 and 6; Van Kerrebroeck et al. 2018a, 2018b; Comasio et
al. 2019; Comasio, Van Kerrebroeck, and De Vuyst 2021).
In particular, diacetyl, which is responsible for a buttery
aroma, may have interesting applications in bread products,
at least when it can be produced in concentrations that are
higher than those present naturally and above the flavor
threshold. The low concentrations of diacetyl commonly
present in breads are possibly due to its further conversion
into 2,3-butanediol by common baker’s yeast, the latter com-
pound having a higher flavor threshold (Birch, Petersen,
and Hansen 2013; Ripari, Cecchi, et al. 2016). Co-fermentation
of maltose and citrate by the heterofermentative Furl. rossiae
CB1 (described as citrate-negative Frul. sanfranciscensis CB1
in the original paper), which is actually a strain uncapable
to use citrate as the sole energy source, leads to a faster
growth, increased concentrations of lactic acid (via a pyru-
vate surplus from citrate, indicating co-fermentation) and
particularly acetic acid (made possible through the enhanced
NAD+ recuperation via the production of extra lactic acid)
compared to maltose solely (Gobbetti and Corsetti 1996).
Sourdough productions with the addition of daily
non-flour ingredients
In traditional bakery recipes, non-flour ingredients, such as
butter, eggs and raisins, are often added to the dough to
enrich the baked goods organoleptically, for instance in the
production of Italian sweet baked goods and biscuits such
as Panettone and Lagaccio, respectively (Venturi, Guerrini,
and Vincenzini 2012). Alternatively, extra ingredients can
be used to initiate and/or propagate sourdoughs. Non-flour
ingredients often added to flour-water mixtures for the pro-
duction of dedicated sourdoughs are alcoholic beverages,
bakers’ yeast, flowers, fruits, herbs, honey, milk, salt, vinegar,
and yoghurt (Table 1). Besides serving as inoculum of cer-
tain microorganisms, they may stimulate or hamper certain
Critical Reviews in Food Science and Nutrition 21
microbial communities already present in the raw materials,
such as the flour and water used.
The most abundant LAB species encountered in sour-
doughs supplemented with non-flour ingredients do not
differ from common sourdoughs, as inventoried before (De
Vuyst, Van Kerrebroeck, and Leroy 2017). In decreasing
order of relative abundance through the examination of 83
sourdoughs (Table 1), these are Lacp. plantarum (31 sour-
doughs) and other Lactiplantibacillus species (formerly
known as the Lb. plantarum group) (19 sourdoughs), Frul.
sanfranciscensis (38 sourdoughs), P. pentosaceus (15 sour-
doughs), and Levl. brevis (10 sourdoughs). Saccharomyces
cerevisiae is the main yeast species found (50 sourdoughs),
followed by T. delbrueckii (10 sourdoughs), K. humilis (3
sourdoughs), Kazachstania barnettii (3 sourdoughs), and
Pichia membranifaciens (2 sourdoughs). AAB species can
particularly occur when fruits or flowers are added to the
flour-water mixture (Ripari, Gänzle, et al. 2016) or when
AAB-carrying beers, such as lambic beer, are used (Comasio
et al. 2020a). Some sourdoughs harbor species that are not
frequently detected, such as the LAB species Pediococcus
inopinatus and Pediococcus argentinicus in wheat and durum
sourdoughs from an Italian artisan bakery, respectively
(Minervini et al. 2012b). Yet, P. argentinicus was originally
isolated from fermented wheat flour (De Bruyne et al. 2008).
Further, Leuconostoc kimchii has been found in Italian bak-
ery wheat sourdoughs, obtained through flour fermentation
in the presence of fruits or honey (Ripari, Gänzle, et al.
2016). Concerning AAB, G. cerinus has been found in Italian
laboratory sourdoughs, obtained through flour fermentation
in the presence of flowers (Ripari, Gänzle, et al. 2016).
Finally, the yeast Yarrowia lipolytica occurs in Greek house-
hold wheat sourdoughs obtained through flour fermentation
in the presence of basil (Paramithiotis et al. 2000).
Fruits
Fruits are often used in sourdough preparations. It concerns,
for instance, apples, apricots, berries, grapes, oranges, pears,
and pomegranates (Randazzo et al. 2005; Aplevicz et al.
2014a, 2014b; Gordún et al. 2015; Vilanova et al. 2015;
Minervini et al. 2016; Ripari, Cecchi, et al. 2016; Ripari,
Gänzle, et al. 2016; Yu et al. 2018). There is a possibility
that these fruits determine the microorganisms thriving in
the sourdoughs, either accidentally (cross-contamination) or
consciously (addition). For instance, retrieval of the yeast
species Metschnikowia pulcherrima from sweet baked prod-
ucts of an Italian artisan bakery has been connected with
the presence of fruits in that bakery (Palomba et al. 2011).
Similarly, Frul. sanfranciscensis can be linked with flowers
and insects, which may contaminate the bakery environment
(Wong, Ng, and Douglas 2011; Ripari, Gänzle, et al. 2016;
Gänzle and Zheng 2019).
The addition of oranges or pears to wheat flour-water
mixtures leads to a different LAB consortium (Levl. brevis,
Lacp. plantarum, and Furl. rossiae, albeit in different pro-
portions) after fermentation than that of the use of wheat
flour only [Leuconostoc mesenteroides, Latilactobacillus cur-
vatus (previously known as Lactobacillus curvatus), and P.
22 L. DE VUYST ET AL.
pentosaceus] for sourdough production (Yu et al. 2018).
Although several yeasts may be present through the addition
of fruits at the start of the flour fermentation processes,
yeasts belonging to the Saccharomyces group are often most
competitive (Gordún et al. 2015). In particular, S. cerevisiae
mostly thrives in the presence of fruits and flowers; other
yeasts, such as T. delbrueckii and Kazachstania spp., often
occur in the presence of herbs or honey. In addition, fruits
cause a low initial pH and enable a faster acidification of
the flour-water mixture, which not only steers the growth
of certain LAB and yeast species but also controls the
growth of undesirable microorganisms (Gordún et al. 2015;
Minervini et al. 2016; Yu et al. 2018). Furthermore, the
addition of fruits enhances the concentrations of certain
amino acids and thus the aroma formation during flour
fermentation, which is particularly pronounced in the con-
comitant breads produced (Minervini et al. 2016; Yu et al.
2018). This is the result of not only bacterial metabolic
activity (influenced by the fruits and pH, for instance
enhanced production of ornithine and GABA) but also
higher activity of the endogenous flour enzymes.
Fruit juices
The addition of lemon juice (as a source of citrate) and
apple juice (as a source of malate) to a wheat flour-water
mixture, fermented with the citrate- and malate-positive
Coml. crustorum LMG 23699, enables this starter culture
strain to prevail and co-ferment the extra carbon sources
during both Type 2 and Type 3 sourdough production pro-
cesses; yeast growth is favored by apple juice but retarded
by lemon juice (Comasio, Van Kerrebroeck, and De Vuyst
2021). The Type 3 fermentation strategy requires, however,
the addition of juice at each backslopping step. Lemon juice
enables citrate conversion and thus the production of lactic
acid, acetic acid, and the buttery flavor compounds acetoin
and diacetyl. Apple juice enables malolactate fermentation
and thus an enhanced production of lactic acid. These inno-
vative sourdoughs lead to the production of baked goods
with different flavor characteristics.
Honey
Honey is a carbohydrate-rich product and hence may influ-
ence the growth of microorganisms naturally present in a
flour-water mixture to which it is added (Table 1; Randazzo
et al. 2005; Minervini et al. 2016; Ripari, Gänzle, et al. 2016)
as well as after the addition of a starter culture (Table 2;
Nutter et al. 2017). Indeed, the high concentrations of glu-
cose (used as energy source) and fructose (used as energy
source or alternative external electron acceptor) present in
honey accelerate the growth and cell yield of strains of the
LAB species added, both homofermentative ones (such as
P. pentosaceus) and heterofermentative ones (such as Liml.
fermentum), during a Type 2 sourdough production process
(Nutter et al. 2017). In contrast, the antimicrobial com-
pounds present in honey inhibit the growth of yeasts that
are naturally present in the flour (Minervini et al. 2016).
Further, honey contains glucose oxidase, which oxidizes
glucose to glucono-δ-lactone and hydrogen peroxide, which
in turn oxidizes free thiol groups of the gluten proteins,
thereby forming disulfide bridges. Consequently, the addition
of honey may enforce the gluten network of sourdough.
However, the addition of honey retards the acidification of
the sourdough matrix by the homofermentative P. pentosa-
ceus through reduction of the activity of the endogenous
proteases, which influences the gluten network too. Yet, the
final pH is too low for glucose oxidase activity and, hence,
the addition of honey has no impact on the gluten network
with this LAB species. In contrast, in the presence of the
heterofermentative Liml. fermentum, a milder acidification
of the sourdough matrix occurs, which does not affect glu-
cose oxidase activity, thereby minimizing the effect of glu-
tathione reduction and, hence, forming a stronger gluten
network.
Milk and yoghurt
In traditional sourdoughs from the western region in Inner
Mongolia (China), sourdoughs produced in the presence of
milk contain Leuconostoc citreum that is naturally present
in the milk, next to Lacp. plantarum (Zhang et al. 2011).
Fermented dairy products are a source of dedicated LAB
species, for instance the homofermentative Lactobacillus del-
brueckii subsp. bulgaricus and Streptococcus thermophilus of
yoghurt. The addition of yoghurt to a flour-water mixture
increases the LAB counts, which are thus much higher than
the yeast counts, at the start of a sourdough production
process. However, upon backslopping, the yeast counts
increase and P. pentosaceus and S. cerevisiae become prev-
alent in a fermenting flour-water mixture inoculated with
yoghurt, indicating that the yoghurt LAB species are not
appropriate to adapt to and survive in the sourdough envi-
ronment (Gordún et al. 2015).
Alcoholic beverages
Alternative ingredients used in the production of
region-specific sourdoughs are, for instance, beer. Birote
sourdough is produced based on eggs, lemons, and beer
and is used for the production of Pan de Birote, a French
baguette-like baked product in Mexico (Puratos 2018).
In Greece, a small amount of ouzo (a strong
anise-flavoured distilled spirit produced from pomace) is
sometimes added to a mixture of crushed chickpeas, basil
leaves, and pelargonium, which is fermented overnight and
used in the production of wheat sourdoughs (De Vuyst et
al. 2002).
Also white and rye ciders have been used as alcoholic
non-flour ingredient (Vilanova et al. 2015). It has turned
out that the sourdoughs made with white cider have more
Leuc. mesenteroides than those made with rye cider.
Finally, lambic beer (a beer obtained through spontaneous
fermentation by means of yeasts, LAB, and AAB; Bogaerts,
De Roos, and De Vuyst 2021) has been used to produce
sourdough. It turned out that the sourdoughs carry yeast
species such as Dekkera anomala and P. membranifaciens as
well as Acetobacter species, reflecting the lambic beer eco-
system and brewery environment (Comasio et al. 2020a).

Conclusions
Addition of non-flour ingredients to the flour-water mixture
for sourdough production is a common practice in artisan
bakeries. However, the influence of these ingredients (and
the microorganisms they carry) on the microbial ecology
of the sourdough matrices and the breads produced thereof
has not been studied in detail yet. The survival of micro-
organisms transferred from the ingredients to the fermenting
flour-water mixture depends on the competitiveness toward
particular strains of the microbial species present under the
harsh conditions of the sourdough ecosystem. Their survival
and growth is also determined by the presence of the appro-
priate substrates, whether or not carried over by the ingre-
dients added. Yet, a thorough understanding of the behavior
of the microorganisms and the impact of certain substrates
from the ingredients used may lead to the production of
new, stable sourdoughs and concomitant innovative sour-
dough breads.
Disclosure statement
No potential conflict of interest was reported by the authors.
Funding
The authors acknowledge their financial support from the Research
Council of the Vrije Universiteit Brussel (SRP7, IRP2, IOF2442, and
IOF3017 projects), the Hercules Foundation (projects UABR09004 and
UAB13002), Flanders’ FOOD (projects Innocereal, Innocereal II,
SourDough, FibraxFun, and SourFun), and the Agency Flanders
Innovation and Entrepreneurship (VLAIO; projects SourDough and
SourFun).
ORCID
Luc De Vuyst http://orcid.org/0000-0002-7534-7186
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