Grain & Oil Science and Technology 7 (2024) 124–132

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Grain & Oil Science and Technology

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Review article

Impact of sourdough fermentation on nutrient transformations in

cereal-based foods: Mechanisms, practical applications, and
health implications
Zhen Wang, Luyang Wang *
State Key Laboratory of Crop Stress Adaptation and Improvement, College of Agriculture, Henan University, Kaifeng 475004, China

A R T I C L E I N F O A B S T R A C T

Keywords: Sourdough is often considered a healthy choice and quality improver for use in cereal production due to its
Sourdough fermentation unique microbial composition and fermentation properties. During sourdough fermentation of cereals,
Lactic acid bacteria biotransformation of nutrients occurs, resulting in notable changes to proteins, carbohydrates, fats, vitamins, and
Proteolysis
minerals. Each nutrient undergoes specific transformations, providing various advantages for human health.
Starch hydrolysis
Proteins undergo hydrolysis to produce small molecular weight peptides and amino acids that are more easily
Low-GI
Nutrients digested and absorbed by the human body. Carbohydrates break down to improve the digestibility and ab­
sorption of cereals and lower the glycemic index. Fatty acids experience oxidation to produce new substances
with health benefits. Additionally, the application of sourdough fermentation can enhance the texture, flavor,
and nutritional value of cereal foods while also extending their shelf life and improving food safety. In
conclusion, sourdough fermentation has a broad range of applications in cereal food processing. Further research
is encouraged to investigate the mechanisms and processes of sourdough fermentation to develop even more
nutritious, healthy, and flavorful cereal-based foods.

1. Introduction
Fermentation has been a crucial technique and approach employed
in the manufacturing of cereal-based foods for almost a millennium.
Sourdough fermentation technology is an ancient and primitive food
manufacturing biotechnology used worldwide [1]. Scientific research
on sourdough biotechnology began in the 1990s. Initially, the focus was
on the impact of sourdough on taste, texture, and shelf life, such as
slowing staling and preventing spoilage, as well as studying the micro­
bial relationships within this complex environment [2]. Recently, sci­
entific research has also centered on the functional and nutritional
properties of sourdough fermentation. According to research, sourdough
fermentation can improve the taste, consistency, and nutritional value of
grain foods [3]. It has also been found to lower the glycaemic index [4],
and enhance baking when compared to fermentation with commercial
dry yeast [5].
The fermentation of cereal has shown promise in enhancing and
formulating the nutritional value of food and ingredients, as well as
influencing their effects on health [6]. Sourdough can enhance the
sensory characteristics of whole grain, fiber-rich, or gluten-free items,
and can also effectively inhibit the digestion of starch, resulting in a
decrease in glycaemic response. Sourdough fermentation has been
shown to regulate the levels and accessibility of bioactive substances, as
well as enhance the bioavailability of minerals [6,7]. Meanwhile, this
process can generate polysaccharides that are difficult to digest or
modify the availability of grain fiber complexes to intestinal bacteria
[8]. It has been suggested that the breakdown of gluten in sourdough
bread may make it more suitable for individuals with celiac disease [9].
Modifications to the grain structure may enhance its nutritional value.
This can be achieved by generating acids and adjusting pH levels to
promote the activity of specific internal enzymes, which can modify the
availability of minerals and phytochemicals [10]. Furthermore, the
microbial ecology of sourdough depends on grain enzymes. There exists
a mutual relationship between bacterial metabolism and grain enzyme
activity in sourdough [11]. Cereal enzyme activity is regulated by
acidification, oxygen utilization, and the accumulation of thiols in mi­
crobial metabolism. In turn, grain enzymes provide substrates for mi­
croorganisms to grow. During fermentation, enzymes break down large
molecules in grains, including proteins and polysaccharides found in cell
walls. Sourdough fermentation alters the characteristics of cereal-based

* Corresponding author.
E-mail address: luyangwang@henu.edu.cn (L. Wang).

https://doi.org/10.1016/j.gaost.2024.03.001
Received 7 October 2023; Received in revised form 4 March 2024; Accepted 6 March 2024
Available online 11 March 2024
2590-2598/© 2023 Henan University of Technology. Publishing services by Elsevier B.V. on behalf of KeAi Communications Co. Ltd. CC BY-NC-ND 4.0 This is an
open access article under the CC BY-NC-ND license (http://creativecommons.org/licenses/by-nc-nd/4.0/).
Z. Wang and L. Wang
foods, which may affect the assimilation of essential and non-essential
components derived from grains [12]. Also, sourdough fermentation
can produce new bioactive substances, such as prebiotics, oligosaccha­
rides, and other compounds, which can improve the nutritional prop­
erties and functionality of grains [7].
The aim of this paper is to analyze recent researches on the use of
sourdough fermentation technology to improve the quality of cereal
products. The paper provides a detailed description of the changes in
nutrient composition of cereal products, particularly in bakery matrices,
resulting from the sourdough fermentation process, and illustrates the
associated mechanisms. The utilization of sourdough fermentation
technology may extend to the manufacturing of premium staple foods as
well as innovative functional foods.
2. Impact of sourdough fermentation on the macronutrient
composition of cereal-based foods
2.1. Sourdough and its classification
Sourdough fermentation, mastered by humans, is the oldest
biotechnology used to produce cereal-based products, especially baked
goods. Fermented grain products, enjoyed worldwide, provide a crucial
solution to future food production challenges. Sourdoughs can be clas­
sified into three groups based on the method and process of fermenta­
tion. The type-I involves natural sourdough fermentation using yeast
and lactic acid bacteria (LAB) found in the flour [13]. The type-II un­
dergoes sourdough fermentation after precise inoculation with a suit­
able leavening agent [14]. The type-III is a dehydrated version of the
second group, and the type-IV is a combination of the first and second
groups, currently manufactured only in laboratories [15]. The survey
shows that fermented foods, which are commonly found in 47 countries
worldwide (excluding Antarctica), exhibit a wide range of microbio­
logical, biochemical, and/or technological traits [3]. Italy and France
have an advantage in the production of sourdough baked goods due to
their rich historical background and cultural heritage in using sour­
dough. A stable microbial community structure typically emerges in
sourdough after prolonged and continuous colonization, despite the
intricate composition of fermenting organisms: This occurs after more
than 10 cycles of colonization [16]. The dominant LAB species include
Lactobacillus sanfranciscensis (heterofermentative), Lactobacillus planta­
rum (homofermentative), Lactobacillus brevis (heterofermentative), as
well as homofermentative species like Pentosaccharomyces cerevisiae and
Saccharomyces cerevisiae [17–19]. Furthermore, there are additional
secondary communities comprising of various LAB species like Lacto­
bacillus reuteri, Leuconostoc, and Weissella, along with other yeast species
belonging to the Kazachstania branch [20,21].
2.2. Impact of sourdough fermentation on the protein in grain foods
The proteolytic activity of LAB in sourdough processing enhances the
production of cereal products with proteins that are easier to digest.
Sourdough fermentation resulted in a significant increase in the di­
gestibility of wheat-derived protein and the content of soluble protein
[22,23]. This could be attributed to the fact that sourdough, which is a
co-fermentation system of LAB and yeast, produces a higher number of
organic acids. These acids enhance the activity of proteases in the
sourdough fermentation system, as shown in Fig. 1. This, in turn, pro­
motes protein degradation by proteases [24]. The hydrolysis of proteins
in sourdough relies on the presence of proteases found in cereals. The
enzymatic function of LAB in sourdough influences the formation of
biologically potent peptides, amino acids, and metabolites derived from
amino acids in dough and bread. Initially, when wheat and rye flour
were used in the sourdough system, enzyme activation was detected in
the co-fermentation systems of LAB and yeast. Both flours demonstrated
significant protein hydrolysis activity, which ensured the sustainability
of the yeast population [25]. On the other hand, Lactobacillus
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Grain & Oil Science and Technology 7 (2024) 124–132
Fig. 1. The protein hydrolysis process under the fermentation conditions of
lactic acid bacteria (LAB) is carried out in the acid dough system.
sanfranciscensis contains glutathione reductase, while organisms
belonging to the Lactobacillus reuteri group do not have it [26]. Both type
I and type II yeasts have strain-specific characteristics, including the
presence of peptidases involved in the production of bioactive peptides
and the conversion of free fatty acids into antifungal hydroxyl fatty acids
[15]. In the co-fermentation system, a higher abundance of L-lactate
dehydrogenase and UDP glucose 4-differential isomerase, which
enhance the quality of sourdough fermentation, was observed compared
to sourdoughs produced from monocultures (only LAB or yeast present)
[27]. Additionally, the sourdough fermentation system showed an in­
crease in calreticulin, enolase, serine tRNA synthetase, ribosomal pro­
tein L23, ribosomal protein L16, and ribosomal protein L5, all of which
play a crucial role in protein stability [27].
During research on sourdough fermentation for making flour prod­
ucts, it was discovered that every type of LAB had distinct impacts on the
composition of gluten proteins. These effects included breaking down
gluten macromolecules into fibrous and lamellar microstructures, as
well as creating protofibril networks linked to antiparallel β-sheet [28].
Gluten proteins undergo depolymerization to form microstructures such
as fibrous networks and lamellar structures, which are associated with
an increase in β-sheet structure. These changes depend on the unique
acidification kinetics of each strain [29]: For example, four strains of
LAB were used in fermentations, Lactobacillus delbrueckii subsp. bulgar­
icus, Lactobacillus fermentum, Lactobacillus plantarum, and Pediococcus
pentosaceus. Lactobacillus fermentum and Lactobacillus plantarum
fermented-gluten proteins were better arranged into sheet-like struc­
tures and both were dominated by parallel β-sheets conformation;
Lactobacillus delbrueckii subsp. bulgaricus and Pediococcus pentosaceus
fermented-gluten proteins were dominated by the development of a
three-dimensional fibrous network accompanied by an increase in
antiparallel β-sheets. This is another indication that the fermentation
activity of LAB also leads to changes in the microenvironment associated
with disulphide bonds and aromatic amino acids [30].
Proteomics provides a powerful tool for studying changes in protein
metabolism during sourdough fermentation. Reale et al. [31] demon­
strated that sourdoughs containing different fermenting Lactobacillus
spp. exhibited distinct peptide profiles; simultaneously, β-amylase and
serpin serve as the main proteolytic targets in sourdough. This partly
exhibits why fermentative sugar are reduced in sourdough fermentation
systems. Through the utilization of proteomics as a method of evalua­
tion, it has been discovered that Lactobacillus paracasei is the foremost
significant LAB for gliadin hydrolysis, with Lactobacillus plantarum
coming in second place [23].
Z. Wang and L. Wang
2.3. Impact of sourdough fermentation on carbohydrates in cereal-based
products
The consumption of sourdough bread can decrease the glycemic and
insulin responses after a meal [32]. This can be attributed to the organic
acids, such as lactic and acetic acids, present in sourdough, which lower
its estimated glycaemic index (eGI). There are two potential mechanisms
that may be involved in lowering the glycemic index of foods in sour­
dough fermentation: (i) Acidity regulation: acid produced during sour­
dough fermentation lowers the pH of food, which slows down the rate of
digestion and absorption of carbohydrates in food [33]. This means that
the glucose in food is released into the bloodstream at a slower rate, thus
reducing the rate of rise in blood glucose. (ii) Inhibition of saccharifi­
cation: organic acids (e.g. lactic acid) produced during sourdough
fermentation may have the effect of inhibiting saccharification reactions
in food [34]. Glycation is the process by which carbohydrates combine
with proteins or fats to form high glycemic reactive substances. By
inhibiting the saccharification reaction, sourdough fermented foods may
reduce the production of hyperglycemic reactive substances, thus
lowering the glycemic index of the food.
In our previous study, we discovered a fascinating occurrence where
intact wheat starch granules were only broken down during synergistic
fermentation with the presence of LAB and yeasts, which contrasts
greatly with the scenario of yeast being the sole participant in the
fermentation process [35]. According to the agreement in the realm of
cereal science, it is widely accepted that yeast can only consume
damaged starch (which are cereal starch granules that have been broken
by mechanical forces during milling) as a source of sugar, whereas it
cannot utilize whole starch granules (as shown in Fig. 2). The combi­
nation of lactic acid-induced pH decrease and activation of cereal pro­
teases during fermentation may influence the change in starch surface
morphology, which could have a significant impact [6]. Deckardt et al.
[36] investigated the modifying effect of lactic acid on starch and found
that the surface morphology of starch changed slightly with decreasing
acidity. Additionally, both acetic and lactic acids induce the deforma­
tion of certain wheat starch granules, along with the presence of cracks
and spots on the surface [37]. However, it is important to note that the
crystalline arrangement of starch cannot be fully broken down by the co-
fermentation of LAB and yeast, also known as type II sourdough [35].
The preservation of the crystal arrangement will directly affect the
ability of starch in sourdough fermentation products to resist digestion.
It was demonstrated that sourdough fermentation is a beneficial
Fig. 2. A new perspective on the hydrolysis of starch in the co-fermentation
process of lactic acid bacteria (LAB) and yeast.
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Grain & Oil Science and Technology 7 (2024) 124–132
process for regulating starch digestibility, which reduces the eGI in
bread products. The most significant decrease in eGI was observed in
whole-wheat sourdough samples obtained at a temperature of 30 ◦ C
using the type II fermentation method. The study found that the type II
sourdough samples derived from whole-wheat flours showed the most
efficient enhancement in RS (resistant starch) content when compared
to the type I sourdough fermentation methods [38]. Additionally, the
study included Tef (Eragrostis tef (Zucc.) Trotter), an ancient gluten-free
cereal that is processed into whole flour and contains high levels of fiber
and minerals. Over time, the glycemic index of fresh Tef bread slightly
increased due to the implementation of sourdough fermentation. How­
ever, the fast-digestible starch decreased by over two times, while the
slow-digestible starch and resistant starch (RS) increased by two-fold
and three-fold, respectively. This was observed in a study where the
bread was stored [39]. The ratio of RS may increase as a result of
α-amylase breaking down branched-chain starches and forming more
limited dextrins. The phenomenon of coagulation during storage can be
explained by the changes in the interaction between the protein matrix
and starch granules that occur in the process [34]. Additionally, freezing
bread further enhances the RS content due to the retrogradation char­
acteristics of starch [39].
Furthermore, sourdough fermentation not only helps reduce the
glycemic index (GI) values through starch retrogradation, but it also
offers another way to achieve this. The fermentation process of sour­
dough decreases the levels of fermentable sugars and free glucose in the
system [40]. Wheat flour contains a type of indigestible carbohydrates,
specifically fermentable oligosaccharides, disaccharides, mono­
saccharides, and polyols (FODMAPs). However, in certain cases of in­
testinal disorders, these sugars can directly or indirectly worsen the
inflammatory condition of the gastrointestinal tract [8]. Initial research
suggests that during the initial fermentation of dough, there is a sig­
nificant reduction in all FODMAPs except for polyols like sorbitol and
mannitol. Additionally, sucrose, fructose, and glucose are completely
broken down during the first fermentation stage and by the end of the
baking process [41]. As a result, the natural fermentation process re­
duces FODMAPs by at least 30%, leading to sourdough bread with lower
levels of fermentable carbohydrates and unbound glucose [42].
2.4. Impact of sourdough fermentation on fats
The oxidation of lipids during dough mixing produces volatile flavors
and affects dough rheology by oxidizing flour components [43]. The
metabolic process of Lactobacillus species may promote lipid oxidation
or exhibit a potent antioxidant impact during sourdough fermentation.
Homofermentative LAB enhances lipid oxidation and the production of
nonenal and decenal throughout the sourdough fermentation process
[44]. The phenomenon’s occurrence is attributed to the production of
hydrogen peroxide during glucose metabolism through homo­
fermentation. In contrast, heterofermentative LAB reduces the sour­
dough’s redox potential and specifically synthesizes glutathione or
similar low molecular weight thiol compounds [45]. LAB accumulates
thiols through heterofermentative metabolism, which generates
reducing agents in the pentose phosphate pathway. This enables a
potent capacity for converting lipid peroxides into hydroxyl groups.
Additionally, during sourdough growth, the alcohol dehydrogenases of
heterofermentative Lactobacillus spp. convert (E)-2-nonenal and (E,E)-
2,4-decadienal to their corresponding alcohols [7]. Additionally, the
sourdough fermentation system showed a reduction in the proportion of
flavor-active compounds, including heptenal, nonenal, nonbutenal, and
dodecenal [12].
Z. Wang and L. Wang Grain & Oil Science and Technology 7 (2024) 124–132

3. Impact of sourdough fermentation on micronutrient content
in cereal-based products
3.1. Vitamins
Whole-grain products are rich in vitamins E, thiamin, and folate,
primarily found in the germ and bran [46]. The fermentation process in
sourdough enhances the vitamin content, especially when specific types
of LABs are involved, as they can produce vitamins like riboflavin,
thiamin, and folate [47]. Furthermore, the activity of microorganisms,
such as LAB that prefer fructose, contributes to the production of nico­
tinamide mononucleotides during sourdough fermentation [48].
Therefore, incorporating LAB in sourdough is an alternative solution
to prevent vitamin deficiencies, both clinical and subclinical. To in­
crease the bioavailability of vitamins, it is recommended to select strains
that produce an excess of vitamins [5]. Additionally, there is a hy­
pothesis that brewer’s yeast could enhance the growth of lactic acid
Lactobacilli by providing vitamins (such as B12, C, or D) that are not
present in wheat flour [5]. Using flour made from germinated grains can
also improve the accessibility of the vitamins found in them, as they are
produced along with other bioactive substances during the transition
from dormancy [49].
However, it is important to note that baking leads to a decrease in
vitamin content, including vitamin B12 in its various forms, such as
cyanocobalamin, hydroxocobalamin, and the form synthesized during
baking [50]. Therefore, it is necessary to develop new strategies to
maintain the bioavailability and vitamin levels in sourdough fermen­
tation products.
3.2. Minerals
Cereal-based foods primarily contain minerals such as K, P, Mg, and
Zn. However, they also contain phytic acid, which can decrease mineral
absorption. Phytic acid is an anti-nutrient found in varying quantities in
cereals and cereal-based foods. It forms phytates with minerals such as
calcium, iron, potassium, magnesium, manganese, and zinc, rendering
them insoluble [51]. Phytates are present in high concentrations in the
outer layers of cereals and can be broken down by the phytase enzyme
found within them [52].
Inositol hexaphosphate esterase (EC 3.1.3.8/EC 3.1.3.26) is the
enzyme that hydrolyzes phytase, releasing soluble inorganic phosphate,
low-grade inositol phosphates, and myo-inositol in sequence. Sourdough
fermentation increases phytase, amylase, and xylanase activities, while
decreasing pH [53,54]. Therefore, proper hydration enables the break­
down of most phytic acid and ensures optimal mineral absorption during
the pre-fermentation process of whole grains or bran [6]. In sourdough
fermentation, the degradation of polysaccharides is typically more
pronounced compared to yeast fermentation. However, to successfully
execute this procedure, it is necessary to maintain the appropriate pH
level, precise timing, and optimal temperature conditions. The use of
sourdough, along with the activity of phytase and the fermentation
process, collectively improve the availability of minerals. Studies have
shown that certain strains of Lactobacillus spp. can enhance the avail­
ability of calcium, zinc, and magnesium to some extent [54,55]. Uti­
lizing a combination of specific LAB and yeast strains, the phytic acid
content was reduced by more than 12% [56]. Table 1
[5,23,24,37,40,43,45,50,52,55,57,58] summarizes the impact of sour­
dough fermentation on the key nutrients in cereal-based foods, including
their biotransformation and health benefits.
4. Interaction between LABs and yeasts is an important
mechanism for nutrient changes in cereal foods fermented by
sourdough
During sourdough fermentation, LABs and yeasts engage in various
interactions that include nutrient competition, mutualistic relationships,
acidity regulation, flavor formation, and nutrient utilization [59]. These
interactions have a significant impact on substance metabolism and
flavor development during the fermentation process, ultimately leading
to changes in the nutrient composition of the grain matrix. The mech­
anisms underlying the interaction between LABs and yeasts in sour­
dough fermentation systems are not fully understood. However, it is
believed that LABs and yeasts communicate through the release of
various compounds, such as organic acids, enzymes, and signaling
molecules [60]. These compounds may act as signals or nutrients for the
growth and activity of the other organism.
Firstly, LABs and yeasts work together in a symbiotic relationship,
where each organism benefits from the activities of the other [61]. The
LABs are responsible for producing lactic acid, while yeast produces
carbon dioxide and ethanol. The interaction between LABs and yeasts
begins with the fermentation of carbohydrates present in the flour by
LABs. LABs convert these carbohydrates into organic acids, primarily
lactic acid, through a process called lactic acid fermentation [62]. This
acidification of the dough creates an acidic environment that is

Table 1
Biotransformation and health benefits of the major nutrients in cereals due to sourdough fermentation.
Nutrients Nutritional changes Correlation with quality of cereal foods Health benefits References

Proteins Soluble protein ↑ Texture Absorbability ↑ [23,24]

Molecular weight ↓ Specific volume Texture Anti-cancer function ↑
Gliadin ↓ Flavor Sensitization↓
Bioactive peptide ↑ Color
Bioavailability ↑
Secondary structural changes
Carbohydrates Starch hydrolysis ↑ Texture eGI ↓ [37,40]
Soluble dietary fiber ↑ Specific volume Texture Intestinal homeostasis ↑
Fermentable sugar ↓ Flavor Immune ↑
Free glucose ↓ Shelf life
FODMAPs ↑
Lipids Unsaturated fatty acid ↑ Flavor Preventing cardiovascular diseases ↑ [43,45]
Hydroxy fatty acids ↑ Shelf life Immune ↑
Short-chain fatty acid ↑ Functionality
Lipid oxidation ↓
Vitamins Riboflavin ↑ Functionality anti-inflammatory ↑ [5,50]
Thiamin ↑ Antioxidant ↑
Folic acid ↑
Minerals Bioavailability ↑ Functionality Mineral absorption rate ↑ [52,55]
Phytate ↓ Antinutritional factor ↓
Others Exopolysaccharides ↑ Flavor Antibacterial properties ↑ [57,58]
Tannin ↓ Shelf life Antioxidant ↑
Functionality

127
Z. Wang and L. Wang
favorable for the growth of LABs and inhibitory for the growth of un­
desirable microorganisms [63]. The presence of lactic acid produced by
LABs also plays a crucial role in yeasts activity. Yeasts are able to tolerate
relatively high levels of lactic acid compared to other microorganisms.
The acidic environment created by LABs promotes the growth and ac­
tivity of yeasts, allowing it to ferment carbohydrates and produce carbon
dioxide and ethanol [64]. The carbon dioxide gas produced by yeasts is
responsible for the leavening effect in sourdough bread, creating the
characteristic airy texture. In addition to creating an acidic environ­
ment, LABs also produce various metabolites that contribute to the fla­
vor and aroma of sourdough bread [65]. These metabolites include
acetic acid, ethanol, diacetyl, and various flavor compounds. The pres­
ence of LABs in sourdough fermentation systems is therefore crucial for
the development of the unique taste and aroma associated with sour­
dough bread. Furthermore, LABs and yeasts can also interact physically
through cell-to-cell contact. This physical interaction may facilitate the
exchange of metabolites and enhance the overall fermentation process
[66]. It is also believed that LABs may provide protective effects to
yeasts, increasing its tolerance to environmental stresses such as high
acidity [67]. Overall, the interactions between LABs and yeasts in
sourdough fermentation systems are complex and multifaceted. They
involve metabolic, physical, and potentially signaling interactions that
contribute to the production of sourdough bread with its characteristic
taste, texture, and aroma, as shown in Table 2 [68–87].
Table 2
LAB synergistic fermentation with yeast versus only yeast fermentation.
Items LAB co-fermentation with yeast
(Sourdough fermentation)
Fermentation Multiple microbial communities, including yeast and LAB, work together
mechanism in a symbiotic relationship. Yeast ferments sugars into alcohol and
carbon dioxide, while LAB produce lactic acid and acetic acid through
the metabolism of carbohydrates.
Lactobacillus Synergistic fermentation in sourdough is characterized by the presence of
subspecies various LAB subspecies, which may include Lactobacillus sanfranciscensis,
Lactobacillus plantarum, Lactobacillus fermentum, Lactobacillus brevis, and
others.
Enzymes Synergistic fermentation in sourdough involves the production of a wide
range of enzymes by both yeast and Lactobacilli. Yeast enzymes, such as
amylases, proteases, and lipases, break down complex carbohydrates,
proteins, and lipids into simpler forms for fermentation. LAB also
produce enzymes that aid in the breakdown of carbohydrates.
Extracellular One notable aspect of sourdough fermentation is the production of
polysaccharides extracellular polysaccharides by LAB. These polysaccharides, such as
dextran and fructan, contribute to the viscosity and texture of the dough,
influencing the overall quality of the bread.
Acid production Synergistic fermentation in sourdough leads to the production of lactic
acid and acetic acid by Lactobacilli. These organic acids contribute to the
sour taste and tangy flavor of sourdough bread.
Flavor complexity The combination of multiple microbial communities in sourdough
fermentation contributes to a complex flavor profile. The presence of
yeast and Lactobacilli leads to the development of fruity, tangy, and nutty
flavors, providing a rich sensory experience.
Fermentation time Synergistic fermentation in sourdough typically requires a longer
fermentation time compared to fermentation using only yeast. The
involvement of multiple microbial communities and the complex
metabolic processes involved necessitate a more extended period for the
fermentation to reach its desired stage.
Bread texture The presence of multiple microbial communities in sourdough
fermentation contributes to a softer and more elastic bread texture. The
extracellular polysaccharides produced by LAB, along with the gas
produced by yeast, result in a lighter and more open crumb structure.
Shelf life Sourdough bread often exhibits a longer shelf life compared to bread
made through fermentation using only yeast. The presence of lactic acid
and acetic acid acts as natural preservatives, inhibiting the growth of
spoilage microorganisms and extending the bread’s freshness.
Health benefits The Lactobacilli present in sourdough fermentation produce lactic acid,
which promotes gut health and aids in digestion. Additionally, the
fermentation process breaks down certain anti-nutrients, making
nutrients more bioavailable.
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Grain & Oil Science and Technology 7 (2024) 124–132
5. Other advantages of sourdough fermentation in relation to
cereal products
5.1. Increasing the content of bioactive substances
Grains contain several classes of phytochemicals, including flavo­
noids (which are relatively rare) and non-flavonoid phenols. The most
abundant non-flavonoid phenols are predominantly alkylresorcinols,
lignans, and phenolic acids. Other phytochemicals found in grains
include benzoxazines, betaines, folic acid, phytates, and sterols [88].
These substances possess inherent functional properties, primarily
acting as antioxidants. However, processing conditions such as milling,
malting, and fermentation greatly influence their levels and bioavail­
ability [89]. The enzymatic transformation and microbial metabolic
efficiency of phenolic compounds in sourdough fermentation have been
linked to the presence of high levels of phenolic compounds, as seen in
sorghum and millet [90].
Peptides, amino acids, and microbial metabolites of amino acids play
a crucial role in determining the quality of bread. These compounds act
as taste-active compounds, flavor precursors, and bioactives, contrib­
uting to the accumulation of antioxidant, antihypertensive, and cancer-
preventive peptide, and amino acid metabolites in foods [1,40]. The
presence of specific peptidases from the Lactobacillus strain in sourdough
made from rye malt significantly affects the formation of bioactive
peptides. In cases where the starter cultures have low PepO and high
PepN activity, higher concentrations of antihypertensive tripeptides like
Fermentation using yeast only References
Involves only yeast converting sugars into alcohol and carbon [68,69]
dioxide.
Fermentation using only yeast does not involve the presence of [70,71]
Lactobacilli subspecies.
Fermentation using only yeast relies primarily on yeast-produced [72,73]
enzymes, such as invertase and zymase.
Fermentation using only yeast does not involve the production of [74,75]
extracellular polysaccharides.
Fermentation using only yeast primarily produces ethanol [76,77]
(alcohol), resulting in an alcoholic taste in the final product.
Fermentation using only yeast results in a simpler flavor profile, [78,79]
with the dominant taste being that of alcohol.
Fermentation using only yeast is generally faster, as yeast is a [80,81]
more efficient starter of sugars.
Fermentation using only yeast tends to produce a denser bread [82,83]
texture due to the absence of extracellular polysaccharides and the
dominance of alcohol production.
Fermentation using only yeast does not provide the same [84,85]
preservation effects, resulting in a shorter shelf life for the bread.
Yeast fermentation helps to the production of B vitamins and [86,87]
minerals in the bread.
Z. Wang and L. Wang
LQP and LLP were observed [91]. A study demonstrated that sourdough
fermentation effectively enriches baked goods with γ-Aminobutyrate (a
biologically active byproduct of glutamate) and concentrations of anti­
hypertensive tripeptide activity [92]. Additionally, the production of
flavor peptides can help mask the reduction of salt content and enrich
baked products with functional anti-hypertensive compounds. The
synthesis of flavored FAA (free amino acids) and amino acid derivatives
during sourdough fermentation compensates for the reduction in salt,
which is influenced by the type and strain of LAB. By incorporating rye
malt sourdough fermented with Lactobacillus reuteri, which accumulates
glutamate, the salt level in bread can be decreased from 1.5% to 1.0%
while maintaining the same flavor and quality characteristics [93].
Sourdough-fermented wheat germ bread has a saltier taste compared to
conventional bread [94]. This higher salinity is a result of acidification,
protein breakdown, and the chemistry of wheat germ. Furthermore, the
presence of microorganisms during sourdough fermentation can lead to
the development of specific nutritional components. For example,
glutamine provides energy for the mucosa of the small intestine, while
short-chain fatty acids serve as energy sources for the mucosa of the
colon, along with linoleic acid and arginine.
5.2. Extending the shelf life of cereal-based products
Extensive experience has demonstrated that sourdough bread pos­
sesses a considerably extended shelf life compared to white bread.
Considering this, the baking sector has recently started reevaluating this
conventional fermentation technique as a potential substitute for
chemical additives in order to guarantee a clean label. Consequently, the
scientific investigation and emphasis on sourdough fermentation tech­
nology have extensively examined its function as a biological agent and
inhibitor of bread spoilage.
Scientific research has established that sourdough possesses the ca­
pacity to slow down staling and safeguard bread against staling, thereby
aiding in prolonging the bread’s shelf life, albeit with certain limitations.
LAB, such as Lactobacillus acidophilus, Lactobacillus amylovorus, Lacto­
bacillus brevis, Lactobacillus fermentum, Lactobacillus gasseri, Lactobacillus
plantarum, Lactobacillus reuteri, Lactobacillus rhamnosus, Lactococcus,
Lactobacillus pentosus, and other bacteria participating in sourdough
fermentation, primarily provide this capability, demonstrating a certain
preservative impact on bread [95].
The ability of sourdough LAB to combat bacteria, microorganisms,
and fungi is linked to their production and/or release of active sub­
stances. These substances can supplement or potentially replace the
need for chemical preservatives. Certain small molecules, including
cyclic dipeptides, hydroxy fatty acids, phenyl, and substituted phenyl
derivatives (such as 3-phenyl lactic acid, 4-hydroxyvaleric acid, and
benzoic acid), diacetyl groups, hydrogen peroxide, caproate esters,
reuterin, and fungicidal peptides [96,97], are the primary substances
that demonstrate antifungal effects. LAB that is heterofermentative
specifically releases organic acids with antifungal properties. Lactoba­
cillus sanfranciscensis can generate a range of organic acids, such as
acetic, butyric, hexanoic, formic, n-pentanoic, and propionic acids [95].
These organic acids exhibit effective antifungal properties against
Fusarium, Penicillium, and Aspergillus. The production of inhibitory
compounds like 4-hydroxyphenolactic acid and phenylactic acid by
Lactobacillus plantarum has demonstrated its extensive antifungal capa­
bilities [98]. Active reuterin, a compound in the form of a hydrated
monomer and cyclic dimer of 3-hydroxypropionaldehyde, is released by
Lactobacillus reuteri. This low molecular weight antibiotic effectively
targets Gram-positive LAB and yeasts, exhibiting antimicrobial proper­
ties against various food spoilage organisms such as Gram-positive and
negative bacteria, yeasts, and molds [99].
The antimicrobial activity of sourdough fermentation is linked to the
production of organic acids, extracellular polysaccharides, and antimi­
crobial substances such as lactic acid. Lactic acid also aids in the
transformation of phenolic compounds and lipids. Heterofermentative
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Grain & Oil Science and Technology 7 (2024) 124–132
LAB, commonly found in wheat and rye sourdough, prefer to metabolize
sucrose and maltose. Maltose is produced by cereal enzymes during the
fermentation process. Sucrose aids in the metabolism of hetero­
fermentative LAB, resulting in the production of acetate and extracel­
lular polysaccharides [100]. However, research on the transcriptional
regulation of bacterial EPS remains limited. Most identified regulators
have been focused on harmful pathogenic bacteria, such as Streptococcus
pneumoniae, Staphylococcus aureus, and Pseudomonas aeruginosa. In
contrast, there is a lack of reported regulators in beneficial probiotic
bacteria, such as Lactobacillus, Bacillus licheniformis, and Bacillus subtilis.
Additionally, most of the identified regulators are for capsule poly­
saccharides, with fewer reported for mucopolysaccharides. Lipid meta­
bolism also contributes to shelf life. As previously mentioned, LAB uses
hydroxy fatty acids to synthesize oleic, linoleic, and linolenic acids. The
conversion of linoleic acid to 13-hydroxy-9-octadecenoic acid by fatty
acid hydratase results in the production of the antifungal 10-hydroxy-
12-octadecene [95]. The growth of Lactobacillus hammesii in sour­
dough results in the production of hydroxy fatty acids, which signifi­
cantly increases the shelf life of bread [101]. Additional research is
required to elucidate the precise mechanisms involved in the relation­
ship between fatty acid conversion by LAB and membrane homeostasis
[102], as the physiological role remains unclear.
5.3. Sourdough fermentation and celiac disease
Celiac disease (CD) is a lifelong systemic autoimmune disorder that
occurs in genetically susceptible individuals who ingest gluten proteins.
In patients with CD, gluten proteins induce chronic inflammation in the
intestinal epithelium, leading to CD-related pathological alterations
such as increased crypt size, reduced villi length, and infiltration of
lymphocytes within the epithelial layer. Dietary advice for people with
celiac disease is to refrain from eating foods containing gluten for life.
The sourdough fermentation process is thought to alter the surface
structure of gluten and to be able to reduce antigen-antibody reactions
that occur in vivo after fermented food ingestion. Celiac peptides
generate α- and γ-gliadin proteins which comprise of an unorganized
initial structural section that is elongated by a C-terminal structural
section folded into an α-helix. The celiac peptides are primarily found in
the unorganized section of the α- and γ-gliadin proteins molecules and
are easily accessible to proteolytic hydrolases in the stomach (pepsin)
and intestines (trypsin, chymotrypsin) fluids, while also being in direct
proximity to them. So far, over 50% of the possible peritoneal peptides
discovered in alcohol-soluble proteins exhibit pepsin cleavage sites. The
sourdough fermentation caused the breakdown of α- and γ-gliadin pro­
teins in the N-terminal structural domain, while the core protein region
known as the crypto-celiac epitope remained unaffected [31]. None­
theless, during docking trials, celiac peptides that were shortened by
proteolytic hydrolysis of either one or two amino acid residues displayed
considerable efficacy when interacting with the human leukocyte anti­
gen HLA-DQ2 and HLA-DQ8 receptors. On the other hand, the process of
sourdough fermentation includes additional protein hydrolyzing en­
zymes that are easily broken down by the celiac peptide. However, since
sourdough-treated breads and foods may still contain small quantities of
celiac peptides that can harm individuals with CD, it is advisable for
celiac patients to refrain from consuming such products. By utilizing the
supplementary sourdough technique on genetically engineered wheat
varieties with reduced gluten content or deemed safe for individuals
with celiac disease, it is possible to create food products that are more
secure for those who are vulnerable to gluten and have celiac disease
[103]. However, it should be noted that sourdough-fermented foods are
not yet licensed in large-scale clinical settings.
Z. Wang and L. Wang
6. Challenges and future perspectives
6.1. Challenges in sourdough research
Sourdough research faces challenges related to the complex nature of
the microbial ecosystem, variability, lack of standardization, limited
understanding of microbial interactions, industrial-scale applications,
bread quality evaluation, and preservation methods etc.:
(i) Complexity: Sourdough is a complex microbial ecosystem con­
sisting of a diverse community of yeasts and LAB. Understanding the
dynamics and interactions of these microorganisms is a challenging task.
(ii) Variability: Sourdoughs can vary significantly in terms of mi­
crobial composition and activity, depending on factors such as
geographical location, ingredients used, and fermentation conditions.
This variability makes it difficult to draw general conclusions or estab­
lish standardized protocols.
(iii) Lack of standardized methods: There is a lack of standardized
methods for sourdough research, making it challenging to compare re­
sults across studies. This hinders the progress and collaboration within
the field.
(iv) Limited knowledge of microbial interactions: The interactions
between yeasts and LAB in sourdough fermentation are not yet fully
understood. Investigating these interactions and their impact on bread
quality is a complex task.
(v) Industrial-scale applications: Scaling up sourdough production
for industrial applications while maintaining the desired microbial
composition and fermentation characteristics is a challenge. Developing
efficient and consistent sourdough production methods on a larger scale
is an ongoing area of research.
(vi) Preservation and storage: Developing methods to preserve and
store sourdough starters or cultures for extended periods without losing
their functionality or altering their microbial composition is a challenge.
This is particularly important for commercial applications.
6.2. Future perspectives
Several intriguing scientific inquiries remain unanswered, such as
the potential for measurable synergistic or antagonistic interactions
among various strains of LAB, and the feasibility of preserving the me­
tabolites of LAB during the manufacturing of cereal foods. Meanwhile, it
is advisable to conduct further research on the procedure and mecha­
nism of sourdough fermentation in order to enhance its significance in
the realm of cereal food processing. Utilizing Foodomics as a tool shows
great potential in connecting nutrient transformations and microbial
metabolism in sourdough fermentation. Exploring the application of
omics tools to tackle present challenges will be a highly promising
avenue for research. Exploring the impact of sourdough fermentation on
human health is a crucial area for future research, particularly in rela­
tion to alterations in nutrient composition. Exploring the integration of
sourdough fermentation with other food processing methods may lead
to the development of cereal products that have increased nutritional
value.
7. Conclusion
Sourdough fermentation plays an important role in cereal-based
foods and has a significant impact on nutrient retention and trans­
formation. An in-depth understanding of its mechanism of action could
provide a more effective means of food processing, leading to better
retention of nutrients in foods, improved bioavailability of foods, and
potentially customized foods for populations with specific health needs.
For example, sourdough fermentation increases the solubility of iron
and zinc in cereal foods, thereby improving the bioavailability of these
key minerals. This has important implications for preventing and
addressing micronutrient deficiencies. Second, sourdough fermentation
also improves the digestibility of fiber in cereals. The enzymes produced
130
Grain & Oil Science and Technology 7 (2024) 124–132
during the fermentation process break down the cellulose, making it
easier for the body to digest and absorb. This not only improves the
digestibility of foods, but also helps maintain intestinal health. In
addition, sourdough fermentation modifies the interference of nutrient
absorption by anti-nutritional factors in cereals, such as phytates. In
terms of practical applications, by controlling the conditions of sour­
dough fermentation, food products can be customized to suit the needs
of different populations. For example, for people who need to control
their blood sugar, low-sugar foods can be produced by selecting the
fermentation process; for people who need to increase their mineral
intake, mineral-rich fermented foods can be developed.
Overall, the use of sourdough fermentation in cereal-based foods has
important health implications. By gaining a deeper understanding of its
mechanism of action, we can better utilize this natural food processing
method to provide more possibilities for healthy food production.
CRediT authorship contribution statement
Zhen Wang: Writing – original draft. luyang Wang: Writing – re­
view & editing, Funding acquisition.
Ethical statement
There are no human or animal subjects in this article.
Declaration of Competing Interest
The authors declare no competing interests.
Acknowledgements
This work was supported by the Graduate Education Innovation and
Quality Improvement Project of Henan University (No. SYLYC2023185).
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