Review

Exploring the Nutritional Impact of Sourdough Fermentation:

Its Mechanisms and Functional Potential
Zuhal Alkay 1, Fereshteh Falah 2, Hasan Cankurt 3 and Enes Dertli 4,*

1 Food Engineering Department, Faculty of Engineering, Necmettin Erbakan University,

Konya 42010, Türkiye; [email protected]
2 Department of Food Science and Technology, Faculty of Agriculture, Ferdowsi University of Mashhad,

Mashhad 9177948974, Iran; [email protected]

3 Food Technology Department, Safiye Cikrikcioglu Vocational School, Kayseri University,

Kayseri 38000, Türkiye; [email protected]

4 Department of Food Engineering, Faculty of Chemical and Metallurgical Engineering,

Yildiz Technical University, Davutpasa Campüs, Istanbul 34210, Türkiye

* Correspondence: [email protected] or [email protected]

Abstract: Sourdough fermentation is one of the oldest traditional methods in food technology and
occurs as a result of fermentation of flour prepared from grains. The nutritional role of sourdough
is related to the final composition of fermented foods prepared through sourdough fermentation,
and recently, sourdough has become an important application to improve nutrition characteristics
of bread. Thanks to lactic acid bacteria (LAB) presented in sourdough microflora and metabolites
partially produced by yeasts, technological and important nutritional features of the bread improve
and an increase in shelf life is achieved. In addition, sourdough bread has a low glycemic index
value, high protein digestibility, high mineral and antioxidant content, and improved dietary fiber
composition, making it more attractive for human nutrition compared to regular bread. When the
sourdough process is applied, the chemical and physical properties of fibers vary according to the
degree of fermentation, revealing the physiological importance of dietary fiber and its importance
to humans’ large intestine microbiota. Therefore, taking these approach frameworks into consider-
ation, this review highlights the benefits of sourdough fermentation in increasing nutrient availa-
bility and contributing positively to support human health.

Citation: Alkay, Z.; Falah, F.; Keywords: sourdough; microflora; nutritional role
Cankurt, H.; Dertli, E. Exploring the
Nutritional Impact of Sourdough
Fermentation: Its Mechanisms and
Functional Potential. Foods 2024, 13,
1. Introduction
1732. https://doi.org/10.3390/
foods13111732 Sourdough fermentation is a traditional method employed to enhance the nutri-
tional, functional, and workability characteristics of cereals [1]. Utilizing sourdough for
Received: 14 May 2024
leavening is an age-old technique in grain fermentation. The process involves grinding
Revised: 28 May 2024
cereals, pseudocereals, or legumes and adding water to create dough, which then trans-
Accepted: 29 May 2024
Published: 31 May 2024
forms into sourdough over time [2]. The sourdough transformation process used for cen-
turies is not only important for food technology but also represents an important evolu-
tion for mankind. Sourdough bread and similar products crafted from grains serve as
embodiments of diverse knowledge, encompassing agricultural techniques, technological
Copyright: © 2024 by the authors.
advancements, and cultural heritage. Bread holds a significant place in human suste-
Licensee MDPI, Basel, Switzerland.
nance, particularly in moderate climates, intertwining with tradition, societal practices,
This article is an open access article
distributed under the terms and
and religious beliefs. Various languages feature expressions highlighting the deep con-
conditions of the Creative Commons nection between life and bread, like “to earn one’s bread” or “remove bread from one’s
Attribution (CC BY) license mouth”, reflecting the enduring significance of this staple food. The etymology of words
(https://creativecommons.org/license further underscores this bond, as seen in the origin of “companio” from Latin and French
s/by/4.0/). roots, denoting someone with whom bread is shared. Notably, in 2020, “companio” was

Foods 2024, 13, 1732. https://doi.org/10.3390/foods13111732 www.mdpi.com/journal/foods

Foods 2024, 13, 1732 2 of 20

used to name a genus of lactobacilli, Companilactobacillus, commonly found in sourdough

cultures [3].
Sourdough, a traditional natural starter, has been utilized for centuries in the pro-
duction of fermented baked goods as a substitute for baker’s yeast and chemical leavening
agents. Over nearly three decades, research has gathered evidence demonstrating its ef-
fectiveness [4]. Recent advancements have seen a resurgence in the use of sourdough as
the primary method for producing bread. The fermentation process of sourdough enables
the creation of bread using just two ingredients—flour and water. This simplicity is highly
appealing in markets where customers prefer “clean label” products [3].
Nutritional properties of food products are typically associated with their final chem-
ical composition and their bioactive constituents [5]. The presence of lactic acid bacteria
(LAB) in sourdough contributes to the increased nutritional value of bread [6]. Sourdough
not only alters the flavor of bread but also enhances its texture and provides beneficial
effects on health [7]. As a result, sourdough fermentation is recognized as one of the ear-
liest food biotechnologies utilized to ferment grain matrices, primarily valued for its in-
fluence on the sensory, structural, and shelf-life attributes of leavened bakery products
[8]. A diagrammatic representation illustrating the nutritional properties of sourdough
fermentation is depicted in Figure 1, which summarizes the alterations in the main dough
and bread characteristics following the sourdough fermentation process. For instance, a
decrement in the glycemic index of sourdough bread can be observed which is mainly
associated with alterations in the starch characteristics. Similarly, sourdough fermentation
results in a decrement in the phytic acid content of dough and bread that increases mineral
bioavailability. Sourdough fermentation also leads to an increment in the phenolic com-
pounds, protein digestibility, and dietary fiber content of the final bread, all of which are
important nutritional properties discussed in detail below. Importantly, the positive ef-
fects of sourdough fermentation on the reduction in fermentable oligosaccharides, disac-
charides, monosaccharides, and polyols (FODMAPs) as well as the salt content of sour-
dough have recently gained special interest, especially from the consumer’s side.

Figure 1. The effect of sourdough fermentation on the nutritional properties of sourdough bread.
The direction of each arrow demonstrates the alteration in each component or characteristics de-
pending on the sourdough fermentation system.
Foods 2024, 13, 1732 3 of 20

The presence of well-adapted LAB initiates the lactic acid fermentation process of
sourdough, which eventually matures into a stable culture over an extended period. This
culture can be obtained through three established methods, leading to the categorization
of sourdough into types I, II, and III. Type I sourdough represents the traditional form
that requires continuous propagation through regular refreshing with fresh flour and wa-
ter. Type II sourdough involves the industrial inoculation of adapted cultures as dough
acidifiers, while type III sourdough is typically dried for convenient storage and use. Type
I sourdough can further be classified into subtypes Ia, Ib, and Ic based on different origins
and fermentation methods. Additionally, there is a type 0 sourdough, which includes pre-
doughs or sponge doughs with the addition of baker’s yeast (Saccharomyces cerevisiae).
Manufacturers strive to identify and develop types II and III sourdough to ensure opti-
mized and consistent product manufacturing [2]. The type of sourdough together with
other process parameters can play crucial roles in determining the final nutritional char-
acteristics of sourdough bread, and in this review, the effect of sourdough technology on
the nutritional characteristics of sourdough bread will be discussed in a broad context.

2. Nutritional Functionality of Sourdough

2.1. Mineral Bioavailability and Phytic Acid
It is widely recognized that cereals, pseudo cereals, and legumes serve as significant
sources of minerals such as potassium, phosphate, magnesium, and zinc. However, the
utilization of these minerals is hindered by the presence of phytic acid [9]. Phytic acid
possesses the ability to bind to minerals such as calcium, manganese, potassium, iron,
zinc, magnesium, selenium, and copper in the digestive system, preventing their dissolu-
tion and absorption, which results in its classification as an antinutrient [10]. Phytic acid,
also known as myo-inositol hexaphosphate, is present in cereals, seeds, nuts, legumes,
and beans. Serving as the storage form of phosphorus in many plants (rich in six phos-
phate groups), phytic acid is found in varying concentrations in grains [9–11]. It is pre-
dominantly located in the bran portion of grains (within the aleurone layers of cereals)
and exhibits strong chelating properties [10,12]. It is estimated that phytic acid constitutes
approximately 1–5% of the dry mass of cereals [10].
Phytic acid functions as a proton donor, leading to the acquisition of a negative
charge by phosphate groups due to the release of hydrogen ions [13]. This transformation
results in the conversion of phytic acid to phytate, which contains six negatively charged
phosphate groups [13]. Phytates are predominantly found in the outer layers of cereals
and can be hydrolyzed by the phytase enzyme present within them [14]. Due to their
strong negative charge over a wide pH range, the presence of phytates in the diet nega-
tively impacts the bioavailability of divalent and trivalent mineral ions such as Zn2+, Mg2+,
Ca2+, Mn2+, Cu2+, Fe2+, and Fe3+ [11]. Several enzymes are responsible for the hydrolysis of
phytic acid, including soluble inorganic PO43⁻, low-molecular-weight inositol phosphate,
and myoinositol-releasing phytases [15]. Furthermore, phytic acid can hinder the activity
of enzymes like trypsin, pepsin, α-amylase, and β-galactosidase [16]. Phytases, enzymes
that degrade phytic acid, can cleave the phosphomonoester bonds of phytic acid and con-
vert it into lower inositol phosphate esters and inorganic phosphate [1]. Only myo-inositol
hexakisphosphate (IP6) and inositol pentaphosphate (IP5) negatively affect mineral bioa-
vailability [17]. Phytase, an enzyme naturally present in grains, seeds, and microbes, in-
cluding LAB, is responsible for phytic acid breakdown [1]. Aside from LAB, phytase ac-
tivity is also observed in certain parts of the intestinal mucosa and in bacteria residing in
the gastrointestinal tract. Phytates evade digestion as ingested phytate remains largely
intact during its passage through the gastrointestinal tract, thereby impairing mineral bi-
oavailability [10]. Additionally, the phytate–mineral complex undergoes partial hydroly-
sis in the human intestine due to the absence of phytate-degrading enzymes in the small
Foods 2024, 13, 1732 4 of 20

intestine and the limited microbial population in the upper digestive tract [18]. Consider-
ing these factors, excessive consumption of phytic acid is believed to cause gastrointestinal
issues and lead to the binding of essential minerals in food [19].
Numerous studies have demonstrated that LAB generally exhibit low extracellular
phytase activity [20,21]. However, only a few studies have identified LAB strains with
notable extracellular phytase activity [16,22]. Intracellular phytase enzymes are unable to
access phytic acid unless the cell is disrupted. Therefore, using LAB strains with extracel-
lular phytase activity as a starter culture in bread production has been reported to be more
effective than those with intracellular phytase activity [1]. Phytase activity is influenced
by various factors, including temperature, pH, proteolytic stability, and substrate speci-
ficity [16]. Recent research has suggested that the addition of sourdough yeast can en-
hance phytate hydrolysis [1]. It has been proposed that the significant improvement in
mineral bioavailability is attributed to acidification by sourdough yeast, which indirectly
stimulates the inherent phytases in the flour and enhances microbial enzyme efficacy [15].
Fekri et al. [23] reported in their study that Kluyveromyces marxianus, Kluyveromyces
lactis, and Kluyveromyces aestuarri as yeasts, as well as Enterococcus faecium, Pedostiocococus,
and Leuconostoc citreum as LAB, found in the sourdough yeast microbiota exhibit phytase
activity and are resistant to low pH and bile effects. They observed that K. marxianus
strains showed higher phytase efficiency compared to S. cerevisiae strains but had lower
phytic acid content than whole wheat flour [23]. Another study conducted by Yıldırım
and Arıcı [24] identified Levilactobacillus brevis HEB33 and Lactiplantibacillus plantarum
ELB78 strains among the LAB isolates from sourdough samples with the highest phytase
activity [24]. Furthermore, studies have shown that combining LAB and yeast strains can
lead to a more than 40% reduction in phytic acid content [25]. Researchers attribute this
reduction to the synergistic action of LAB and exogenous phytases produced by the yeast
in the sourdough microbiota [21,26,27]. It was also noted that the endogenous phytase
activity in wheat flour and sourdough mixture influences the phytase activity of microor-
ganisms in sourdough [12,28]. The activity of endogenous cereal phytase is typically en-
hanced under acidic conditions during sourdough fermentation. The optimal pH range
for achieving over 70% phytic acid degradation is usually between 4.3 and 4.6 [9]. In the
acidic environment created during sourdough fermentation, phytase activity is usually
rapid. While the optimal pH for wheat phytase is 5.0, yeast phytase functions optimally
at pH 3.5 [29]. Although phytase enzymes are present in rye and wheat, they are often
insufficient to significantly improve mineral bioavailability in whole-grain breads. The
transformation of flour into bread leads to a decrease in phytate content due to flour
phytase activity, but this reduction may not be substantial enough to significantly enhance
mineral bioavailability. The decrease in phytate content during bread production is at-
tributed to phytase activity, which is influenced by various factors such as the flour extrac-
tion rate, fermentation duration and temperature, dough acidity, and the addition of yeast
and enzymes to the dough [30]. Phytate can bind directly to starch through H+ bonding with
a PO43⁻ group or indirectly through proteins, resulting in reduced starch solubility and di-
gestibility [31,32]. Using sourdough technology can directly or indirectly lead to the reduc-
tion in phytate content during bread production and enhance mineral bioavailability.

2.2. Effect on Starch Digestibility and Glycemic Index of Sourdough Fermentation

Another benefit of incorporating sourdough is its ability to slow down the digestibil-
ity of starch in wheat flour-based products like bread [33]. This slowing down of starch
digestibility is a significant nutritional concern, particularly for populations that rely on
commercially leavened wheat bread in their diets. When starch content is rapidly digested
and absorbed, it can lead to hyperglycemia in individuals with diabetes [34,35]. However,
by utilizing sourdough, the organic acids generated during the fermentation process (such
as acetic acid, which reduces gastric emptying, and lactic acid, which hinders the availa-
bility of starch through interactions with gluten) can lower the postprandial glycemic re-
Foods 2024, 13, 1732 5 of 20

sponse in human blood and delay starch digestion [36–38]. Following sourdough fermen-
tation, starch digestibility likely becomes a fundamental aspect in starch, grains, and ce-
reals [39].
The macro and microstructure of grain nutrition has a significant effect on the digest-
ibility of starch. In particular, the properties of starch are important for the glucose re-
sponse [12]. Starch retrogradation, which is the recombination of amylose and amylopec-
tin to create dual helices and likely crystal structures, favors the slow digestibility of
starch. Moreover, the high branch intensity of amylopectin decelerates starch digestibility
[8]. Starch digestibility may be influenced by many agents such as α-amylase binding to
the substrate, gastric unloading, enzyme inhibitors, physiological parts, and the features
and viscosities of digestive enzymes [40]. Amylose-rich starches have been reported to be
more durable than amylolysis waxy or typical starches. It is noted that inherent starches
are hydrolyzed heavily and to a restricted degree by amylases [41]. With the occurrence
of gelatinization during processing, the in vitro proportion of amylolysis rises signifi-
cantly [42], because the more starch gelatinizes, the faster it is digested [43]. For example,
high gelatinization of starch is noted in many prevalent starchy foods like normal white
wheat bread, emphasizing that the product structure is very porous. These properties
cause starch to be broken down rapidly in the small intestine. In addition, it makes the
blood sugar level rise very quickly [12]. However, at the product level, the glycemic re-
sponse is affected by the texture integrity, porosity, and structure of starch [12], because
starch digestion and the glycemic index of foods are directly related [44].
The glycemic index (GI) is defined as a method that classifies foods based on their
carbohydrate content and their effect on blood sugar levels. The classification and GI val-
ues of grain-based products are shown in Figure 2 [45,46]. The GI concept is considered
clinically inadequate. The glycemic index varies between foods, and these differences are
not fully accounted for when consumed as part of a complex meal [47]. This is because
when a complex meal is composed of a few carbohydrate sources, the impact of the lower
GI component may be overshadowed by the quantity of carbohydrates from other
sources. Accurate calculation of the GI of complex meals is crucial for this purpose [48].
The mechanism of action of the GI is as follows [49]:
 Its metabolic effects are generally related to the ratio of glucose that is absorbed from
the small bowel.
 The ratio of glucose decreases after consumption of low-GI carbohydrate foods. For
example, intestinal hormones such as incretins and insulin contribute to the reduced
rate of absorption of increased postprandial glucose. Prolonged absorption of carbo-
hydrates over time maintains the repression of free fatty acids (FFAs) and counter-
regulatory reactions, resulting in lower blood glucose concentrations.
 A decrease in FFA concentrations over time and an increase in tissue insulinization
and respiratory coefficients lead to faster withdrawal of glucose from the circulation.
As a result, glucose absorption from the small intestine continues, but blood glucose
concentrations return to baseline. Thus, the increase in postprandial blood glucose
decreases with increasing blood glucose area above baseline [49].
The GI can be influenced not only by the raw ingredients but also by the baking
methods [37]. It is suggested that products with a low or medium GI (70) are considered
beneficial for health, particularly in preventing cardiovascular diseases, obesity, and type
1 or 2 diabetes [50]. Therefore, understanding the techniques behind high-GI products like
bread is of primary importance [37]. One reason for which white wheat breads have a
relatively high GI is that the starch becomes highly gelatinized during baking at a temper-
ature of 250 °C. This makes it more accessible to salivary and pancreatic α-amylases [37].
It has been noted that the glycemic response of bread, classified as a high-GI product, can
only be reduced through fermentation. It is observed that if the pH values of the dough
are between 3.5 and 4.0, the formation of resistant starch decreases starch digestibility and,
consequently, blood sugar levels [51–53].
Foods 2024, 13, 1732 6 of 20

Resistant starch is characterized as the sum of products that are not absorbed in the
small intestine of healthy individuals, are resistant to enzymatic breakdown, and possess
an exceptionally regular molecular structure [54,55]. The presence of organic acids during
baking leads to the separation of branches in the amylopectin components, facilitating the
formation of resistant starch [56]. This newly formed resistant starch can limit the extent
of gelatinization by encapsulating the starch granules and also provides a physical barrier
against the enzymatic action of α-amylases [57].
It is highlighted that the consumption of wholemeal bread, in comparison to white
flour breads, leads to a reduction in GI. This reduction can be further enhanced by using
sourdough [14,58,59]. Sourdough fermentation is particularly effective within low pH
ranges (3.5–4.0) and contributes to decreased soluble fiber and GI [37]. Consuming sour-
dough bread results in lower postprandial blood sugar and insulin responses [40]. Fur-
thermore, the glycemic response to bread can vary among individuals [60].

Low GI (GI ≤ 55)

Starch/Grains

Medium GI (56–69)
Starch/Grains

High GI (GI ≥ 70)

Starch/Grains
Foods 2024, 13, 1732 7 of 20

Figure 2. Glycemic index (GI) classification and GI values of grain-based products.

In both in vitro and in vivo studies, it has been shown that sourdough fermentation
has the potential to reduce the GI of bread from high to medium. When sourdough fer-
mentation is combined with the addition of dietary fiber (5–10%), the GI can drop below
55 [61]. Research indicates that sourdough fermentation using starter cultures like Levl.
brevis, Lpb. plantarum, and baker’s yeast results in a bread with a higher resistant starch
content compared to spontaneous sourdough fermentation. The impact of fermentation
type on the estimated GI is particularly noticeable in wheat-grain bread [40]. The process
of gelatinization and puffing of starch particles during sourdough fermentation is be-
lieved to increase the GI, making starch more digestible [62,63]. On the other hand, it is
suggested that starch hydrolysis is hindered in the sourdough fermentation process of rye
products, potentially reducing the postprandial glucose response due to the formation of
the amylose layer [63]. Factors such as leavening, baking time and temperature, and flour
type can also influence the starch structure and glycemic response of bread [64–66]. Ad-
ditionally, the glycemic response is influenced by factors like starch sensitivity, protein
and lipid content, and the overall structure of the food [37]. When considering the impact
on the GI of sourdough breads, the type of flour used is crucial. While a decrease in GI is
observed when sourdough is used in wheat breads, the use of sourdough in gluten-free
breads made with buckwheat flour, quinoa, and teff may increase the estimated GI [62,65].
Researchers have noted that the use of sourdough or unleavened pseudocereal flours does
not always result in a rise in heavily digestible starch. Some studies suggest that flours
with smaller starch particle sizes may lead to a higher GI [61]. Various strategies have
been explored to lower the GI, such as incorporating high-fiber flours like rye, oat, and
barley, or adding dietary fiber supplements like β-glucans [67].
In a study conducted by Gil-Cardoso et al. [68], rats were fed different types of bread
containing sourdough. These included a traditional refined wheat bread without sour-
dough (C-WhB), a spelt bread leavened with Rebola sourdough (Re-SpB), a durum wheat
bread with Rebola sourdough (Re-DuB), and a multi-grain bread leavened with Rebola
(Re-MGB), Carla (Ca-MGB), or San Francisco (SF-MGB) sourdoughs. The results indicated
that rats fed with Re-SpB, Re-DuB, and Re-MGB had lower postprandial blood glucose
levels compared to the control group. Additionally, rats fed with SF-MGB showed a re-
duced postprandial blood insulin response.

2.3. Effect on Protein Digestibility

In addition to the impact of sourdough on the GI, the role of LAB in processing or-
ganic nitrogen has been extensively researched [69,70]. LAB from sourdough exhibit pro-
Foods 2024, 13, 1732 8 of 20

teolytic efficiency through cell wall proteinase, intracellular peptidases, and specific mem-
brane transporters [8]. Simultaneous acidification also activates endogenous flour prote-
ases, leading to progressive hydrolysis of natural proteins and an increase in the concen-
tration of peptides and essential free amino acids (FAAs) during LAB fermentation [5].
Sourdough fermentation is considered a pre-digestion process driven by the enzymatic
activity of LAB. Products containing sourdough are generally associated with improved
digestibility and a higher nutritional profile in the protein fraction [71]. Proteolysis is uti-
lized to enhance the digestibility of protein-rich flours such as seeds, reducing the amount
of gluten that can trigger allergic and intolerant reactions in susceptible individuals [8],
while also fortifying grain-based products [40,72].
Recent empirical and scientific studies have highlighted the ability of sourdough fer-
mentation to enhance protein digestibility. In vitro research has demonstrated that sour-
dough fermentation improves the digestibility of protein, measured as the percentage of
total protein dissolved after enzyme hydrolysis. A study by Rizzello et al. [73] investigated
the consumption response to different types of sourdough bread among 36 healthy vol-
unteers. The study found that sourdough bread with medium acidification increased ap-
petite and reduced feelings of fullness, while bread with a more intense acidic flavor led
to a perception of quick and high satiety. Also, they compared the digestibility of bread
made with S. cerevisiae E10 and sourdough bread. In the results, they monitored a rise of
16% in the digestibility of sourdough bread and a rise of 18.7% in the biological amount
of the proteins. Although gallbladder response did not differ between the breads, it was
observed that gastric emptying was faster with sourdough bread compared to commercial
leavened bread. Furthermore, it was noted that levels of essential FAAs in the blood-
stream remained elevated for an extended period after consuming sourdough bread [74].
The literature suggests that sourdough bread enhances protein digestibility compared to
commercial leavened bread, attributed to the proteolysis occurring during fermentation.
Proteolysis breaks down proteins into bioactive peptides and amino acids, facilitating
their absorption by enterocytes [53].
In recent years, there has been a growing trend of adding dried fruits to sourdough
to enhance the essential amino acid content. A study found that incorporating peanut dust
into flour or semolina significantly increased the lysine content in sourdough products
[75]. Another study reported a notable increase in the condensation of essential amino
acids like valine and methionine through sourdough fermentation with the addition of
dried pears and oranges [76]. Notably, there was a significant elevation in gamma-amino-
butyric acid (GABA) among the essential amino acids. GABA, a nonprotein amino acid
primarily derived from the decarboxylation of L-glutamic acid, is known for its roles in
neurotransmission, inducing hypotension, and exhibiting diuretic and sedative effects; so,
we can prove the effects of sourdough on protein digestibility [77,78].

2.4. Salt Reduction Feature

Numerous studies have demonstrated that fermenting essential amino acids can help
reduce the need for salt supplementation without compromising food quality [52]. High
dietary sodium intake, around 10 g/day, is associated with various health risks such as
cardiovascular diseases, hypertension, paralysis, stomach cancer, kidney issues, and bone
demineralization [10]. Therefore, controlling blood pressure by reducing sodium intake is
crucial [10]. However, reducing salt can be challenging as sodium chloride (NaCl) impacts
the taste and physical characteristics of baked goods. To address this, researchers have
explored various technological options [79], including salt substitutes like KCl, season-
ings, flavor enhancers, malt, barley, amino acids, monosodium glutamate (MSG), and
yeast products. However, it has been found that none of these alternatives provide a com-
pletely effective or convenient salt replacement strategy [52]. In this regard, sourdough
fermentation is utilized to mask reduced salt content or enhance bakery products with
functional anti-hypertensive components. During sourdough fermentation, sweetener
FAAs and amino acid derivatives are produced. The extent of salt reduction depends on
Foods 2024, 13, 1732 9 of 20

the type and strain of LAB used, with sourdough exerting stabilizing effects on them. A
study revealed that incorporating rye malt sour yeast leavened with glutamate-accumu-
lating Limosilactobacillus reuteri strains reduced the salt content of bread from 1.5% to 1%
without compromising flavor or other quality aspects [80,81].

2.5. Sourdough Fermentation and Dietary Fiber

Dietary fiber consists of polysaccharides or lignins that are primarily the edible parts
of plants or similar carbohydrates resistant to digestion and absorption in the human
small intestine, undergoing whole or partial fermentation in the gut [8,82]. As per the
American Association of Cereal Chemists (AACC), DFs are plant carbohydrates that can
be fermented by colon microbes but resist hydrolysis by human enzymes, categorized into
soluble and insoluble types [44]. The composition of DF varies based on factors like chem-
ical and physical properties, including physiological effects, the polymerization rate of
polysaccharides, the presence of side chains, crosslinking rate, granule size, and cell wall
integrity [83]. These properties can be influenced by acidification and activation of endog-
enous enzymes [8]. Dietary fiber is recognized as a dietary component that significantly
contributes to health improvement, with varying evidence supporting this claim [84]. It
aids in lowering cholesterol and glucose levels in the blood and plays a crucial role in
promoting health. There is a strong association between obesity, chronic diseases, and
dietary fiber intake, with impacts varying depending on the type of fiber consumed.
Whole-cereal dietary fiber is a valuable source of beneficial molecules like β-glucans, fruc-
tans, resistant starch, and arabinoxylans for the host. The bioavailability of these compo-
nents in whole-cereal products often depends on specific technological factors [85]. Many
consumers adhere to various processing practices, such as fortifying bread with dietary
fiber, to ensure high-quality products [86]. Modern diets recommend a daily intake of 25
g or less of dietary fiber, as per the World Health Organization (WHO). Consequently,
there is a focus on strategies to enhance the fiber content of foods to address this defi-
ciency. Common sources of dietary fiber include cereals like hemicellulose, resistant
starch, β-glucans, arabinoxylans, and other cereal components. Additionally, bran, wheat
germ, by-products from enriching bakery products through grinding, and legume flour
are gaining popularity. However, supplementing these ingredients in high quantities can
alter the sensory and rheological properties of bakery products. One solution to mitigate
these challenges is the use of sourdough, as it has the potential to be incorporated into
bakery items, overcoming the technological drawbacks of high-fiber flours [9].
When utilizing the sourdough process, the chemical and physical characteristics of
the fibers may change based on the level of fermentation [85]. Moreover, the enzymatic
impact can alter the ratio of insoluble fibers to soluble fibers during bread production [87].
It has been observed that in sourdough breads, fibers can undergo two types of enzymatic
hydrolysis. Specifically, different grain-specific enzymes are activated when the flour is
moist, including the hemicellulase enzyme responsible for breaking down hemicelluloses
[88]. Additionally, LAB can secrete enzymes with glycolytic activity that target the fibers
present in the dough [14].
An in vitro study conducted by Mihhalevski et al. [89] explored the enrichment of
sourdough breads with dietary fiber, specifically focusing on rye sourdough bread. The
study revealed an increase in both soluble and insoluble DF ratios during the processing
of rye sourdough, attributed to the biochemical and microbiological processes involved
in sourdough bread production. The rise in total dietary fiber content was linked to the
formation of resistant starch, while the increase in soluble dietary fiber content was asso-
ciated with the conversion of insoluble fibers into soluble fibers during rye flour fermen-
tation [89].
Another study highlighted that fermenting sourdough breads with barley enhanced
their capacity, color, and sensory characteristics. This fermentation process was found to
elevate the levels of dietary fiber, arabinoxylans, and β-glucans [90]. Sourdough fermen-
tation is recognized as a suitable biotechnology for producing wholemeal rye and wheat
Foods 2024, 13, 1732 10 of 20

flour products rich in dietary fiber [2,39]. Sourdough contributes to enhance the flavor,
texture, and shelf life of whole-grain rye breads, as the production of sourdough whole-
meal rye or wheat–rye flour blends without sourdough technology can be challenging [8].
This is because sourdough plays a significant role in traditional rye bread baking by im-
proving workability, flavor, and structure. It is important to note that wholemeal rye
bread relies on fermentation for its production [12].
Specifically, the aspartic protease enzyme, one of the endogenous rye proteases,
breaks down rye proteins and secalins during rye–sourdough fermentation, generating
amino acids and peptides that contribute to the bread’s flavor profile [91]. Recent studies
have focused on the dietary fiber ratios in various sourdough bread varieties, with their
findings summarized in Table 1.

Table 1. Studies on dietary fiber ratios in sourdough bread varieties in recent years and total dietary
fiber amounts in results.

Reference Flour Type Microorganism Total Dietary Fiber Ratio Determined as a Result
Khorasan flour at immature phase using sour yeast
fermentation at high temperatures resulted in (10.26
Lpb. plantarum, mg/100 g). Khorasan flour at entirely mature phase
Kamut khorasan and
Fructilactobacillus using sour yeast fermentation at high temperature re-
Saa et al. [85] durum wheat-grain flour
sanfranciscensis, Levl. Brevis, sulted in (19.25 mg/100 g). Breads acquired with durum
(milky and entirely mature)
and S. cerevisiae wheat flour at entirely mature phase using industrial
fermentation at high temperatures resulted in (15.48
mg/100 g).
Barley sour yeast ended in a higher condensation of
Saccharomyces chevalieri,
Wheat flour and a wheat– both dietary fiber and the arabinoxylans and β-glucan
Pejcz et al. [90] Lacticaseibacillus casei, and
barley blen fractions compared to barley whole wheat. Total dietary
Levl. brevis
fiber was 10%.
The highest total dietary fiber amount (17.2%) was
Pediococcus pentosaceus SA8
Olojede et al. [92] Sorghum flour found in sour yeast bread made with P. pentosaceus SA8
and S. cerevisiae YC1
and S. cerevisiae YC1 strains.
Liml. reuteri, Levl. brevis,
Sourdough breads are Immature wheat sourdough bread (26 day) resulted
Çetin-Babaoğlu Lbp. plantarum, Liml. Fermen-
prepared from immature inm2.18%. Immature wheat sourdough bread (36 day)
et al. [93] tum, and Lacticaseibacillus
wheat flour (26 and 36 days) resulted in 2.10%
rhamnosus
Millet flours such as kodo,
barn, small, and foxtail Sourdough starter culture Foxtail 20–50%, refined flour 65–35%, chickpea flour
Das et al. [94]
were used for sourdough mix 10%, and tapioca flour 5%.
bread production
The highest total dietary fiber amount (15.9%) was
Sorghum flour and corn P. pentosaceus and Weissella found in sourdough bread with P. pentosaceus, while the
Olojede et al. [95]
starch were used confusa lowest total dietary fiber amount was observed in the
control bread without sour yeast (13.25%).
Whole wheat flour
Subaşı and Ercan Lpb. Plantarum and Fruc. Whole wheat bread of Bezostaja-1 had the highest total
(Tosunbey, Kenanbey,
[96] sanfranciscensis dietary fiber amount (15.94%).
İkizce-96, Bezostaja-1)

2.6. Sourdough Fermentation and Gut Microbiota

The human gastrointestinal system is an extremely complex tract. This system, with
its physiological ways which are highly dynamic, appears to be closely coordinated with
diverse microbiota [97]. The challenge lies in the composition and functionality of the
bowel microbiota, with this mutual connection persisting throughout life [98]. There is
now ample evidence that compounds within the bowel microbiota help to combat entero-
pathogens, ensure energy and essential nutrients, improve nutrient digestibility, and
maintain optimal intestinal motility. The intestinal ecosystem also releases chemicals that
Foods 2024, 13, 1732 11 of 20

play a role in immune system activity and potentially cancer prevention [99]. The intesti-
nal microbiota seems to play a role in supplementing human nutrient metabolism and
significantly contributes to the maintenance of a robust and highly active immune system
[12]. The intestinal microbiota is a common term for the microbes residing in the gastro-
intestinal system of all vertebrates. In humans, the bowel serves as the primary habitat for
a person’s microbiota. The intestinal microbiota comprises a variety of bacterial and yeast
species, with its diversity and composition varying at different life stages. This variation
is evident in both healthy and unhealthy individuals [100]. Furthermore, the microbial
composition of the intestinal microbiota varies across different parts of the gastrointestinal
system. The colon harbors a dense microbial population, while the stomach and small
bowel typically have fewer bacteria. Anaerobic microorganisms make up over 99% of the
bacteria in the intestine. The predominant bacterial strains found in the human intestine
belong to the phyla Firmicutes, Bacteroidetes, Actinobacteria, Proteobacteria, and Verrucomi-
crobia [100,101]. It is worth noting that fermented products, particularly grain-based foods,
contain numerous components that reach the gastrointestinal system and are accessible to
the host’s intestinal microbes [44].
It is understood that sourdough leavening, a grain-based fermented product, along
with grain fibers, increases the production of non-digestible polysaccharides in the small
bowel and has the ability to slow down the digestibility of starch, which can then be fer-
mented by the gut microbiota [102]. The impact of sourdough fermentation on bowel
health operates through various mechanisms, as illustrated in Figure 3 [12,14].

Figure 3. Mechanism of action of sourdough fermentation on intestinal health.

Some LAB appear to produce EPSs like glucans, fructans, glucooligosaccharides, and
fructooligosaccharides, which have the potential to improve intestinal health [14]. For ex-
ample, levans produced by Fruc. sanfranciscensis have prebiotic characteristics [103]. In
addition, studies have shown the formation of oligosaccharides and polysaccharides with
prebiotic properties by Liml. reuteri LTH5448 and Weissella cibaria 10M in sorghum sour-
dough [104]. Intestinal microorganisms show the potential to metabolize these compo-
nents, which have been indicated to have prebiotic features [14]. Thanks to the metabolism
of these components, cholesterol and triglyceride levels decrease and insulin susceptibil-
ity is increased. In addition, propionic acid with various beneficial effects can be produced
[105]. However, studies on how the consumption of sourdough bread affects not only the
gut microbiota but also the leave of health-supporting metabolites along with the transi-
tion and continuation of the human gut microbiota profile are limited. In a related study,
Abbondio et al. [106] fed rats a diet supplemented with sourdough breads to test the ef-
fects of sourdough on the composition and function of the bowel microbiota. It was found
in this study that adding sourdough bread to the diet resulted in a reduction in low-pro-
tein diets which are desired by bowel pathogens. Moreover, some studies have indicated
that the cell wall compounds of Lpb. plantarum found in sourdough have the capability to
stimulate the immune response in the gut even when the bacteria are not viable [107].
Another significant benefit of sourdough to host health through the gut system can be the
potential of the sourdough LAB as probiotics and their products as postbiotics [14]. An-
other study was conducted by Korem et al. [108] and in this study, volunteers were fed
Foods 2024, 13, 1732 12 of 20

bread made from refined white wheat and whole wheat sourdough as dietary interven-
tions. The effectiveness of these two bread types was compared and it was concluded that
no significant differences were found between the two 1-week dietary interventions clin-
ically, one involving the consumption of commercial white bread and the other including
the consumption of sourdough bread made from whole cereals. However, person-specific
factors were found to be effective for the role of bread type on clinical parameters, sug-
gesting the role of the gut microbiota and other genetic factors as part of the response to
different breads [108]. In another in vivo study, Wistar rats were fed a control diet, recon-
stituted whole wheat flour (white flour plus bran), commercial bread, and sourdough
bread [109]. The sum cecum pond of short-chain fatty acids (SCFAs), especially the butyr-
ate pond, has been noted to increase significantly with the consumption of unrefined
products. Recent in vivo studies on the effectiveness of sourdough bread varieties on the
gut microbiota are given in Table 2.

Table 2. Recent in vivo studies on the influence of sourdough bread varieties on the gut microbiota.

Human Studies
Type of Flour, Type of Sourdough Bread Made
Reference Results for Gut Microbiota
with LAB, and Number of Volunteers
Wheat flour, Lpb. plantarum CR1, Furl. rossiae CR5, The amount of SCFAs and isovaleric and 2-methylbutyric
Da Ros et al. [98]
and S. cerevisiae E10, 40 healthy volunteers acids increased.
Animal Studies
Type of Sourdough Bread and Type of Animal
Reference Results for Gut Microbiota
Used
In general, after bread ingestion, there was a significant re-
duction in the F. phylum relative to the baseline. There was
Celta bread, S. cerevisiae, S. pastorianus, C. sakei, Lpb.
a substantial rise in Bacteroidetes bacteria in the intestinal
Arias et al. [110] paralimentarius, P. parvulus, Levl. Brevis, and Leu.
microbiota of the mice fed with the commercial bread. In
citreum, 10 female 8-week-old C57BL/6 J mice.
the group fed with sour yeast bread in the Celta class, the
main change was related to Verrucomicrobiaphylum.
It has been noted that diets enriched with bran produce a
Whole-grain wheat and whole-grain rye, C. milleri,
Koistinen et al. rise in the relative amount of a few bacterial taxa, like
Levl. Brevis, and Lpb. plantarum, C57BL/6 J male mice
[111] Akkermansia, Bifidobacterium, Coriobacteriaceae, Lactobacillus,
(n = 74)
Parasutterella, and Ruminococcus.
Mice fed with sourdough bread showed a diabetes-
lowering effect by decreasing the GI, owing to the existence
Yeast-leavened white bread and sourdough bread,
Kwon et al. [112] of dietary fiber and SCFAs. Some useful bowel bacteria like
male C57BL/6 mice
Akkermansia, Bifidobacterium, and Lactobacillus were increased
in mice in the sourdough bread-fed group.

2.7. Sourdough Fermentation and FODMAPs

Another benefit of sourdough fermentation is its ability to reduce FODMAPs. Recent
research has highlighted the importance of FODMAPs, a type of carbohydrates, in relation
to overall health [14]. FODMAPs can impact a person’s health based on the amount of
sugar consumed through food. When consumed in appropriate quantities, FODMAPs can
have a positive effect on health (some FODMAP sugars even have a prebiotic effect)
[113,114]. However, consuming FODMAP-rich foods in excess, over 20 g per day, can lead
to sugar accumulation in the intestines, potentially causing various digestive issues, espe-
cially in individuals with irritable bowel syndrome (IBS) [115,116]. Since FODMAPs are
not easily digestible carbohydrates, they can contribute to the development of conditions
like IBS or non-celiac gluten sensitivity (NCGS) [14].
IBS is a gastrointestinal disorder that affects a significant portion of the population,
with symptoms typically appearing after consuming foods high in FODMAPs. These
symptoms include abdominal discomfort, stomach pain, bloating, constipation, or diar-
Foods 2024, 13, 1732 13 of 20

rhea [117,118]. While simple sugars and polyols have a strong osmotic effect, carbohy-
drates like fructans, fructooligosaccharides, and galactooligosaccharides are more prone
to fermentation by the gut microbiota [116,119]. Some studies suggest that FODMAP com-
pounds can trigger symptoms in individuals with IBS [120]. A low-FODMAP diet is often
recommended by dietitians to help manage IBS symptoms [121], and the sourdough fer-
mentation process plays a crucial role in achieving this. By utilizing the sourdough
method, it is possible to reduce the levels of indigestible oligosaccharides, such as fructans
and α-galactooligosaccharides, making products more suitable for individuals with
NCGS and IBS [122].
During sourdough fermentation, bacterial activity can lead to the production of pol-
ysaccharides, oligosaccharides, and polyols. The breakdown of fructans during bread
making is partially due to the efficiency of invertase in yeast [123]. While cereal fructans
break down to some extent during regular bread making, the process is different with
sourdough. The fructose produced is converted to mannitol by lactobacilli in sourdough,
which is then fermented by the gut microbiota [123]. The extended fermentation period
required for sourdough bread production allows for significant changes in carbohydrate
composition [121].
To further reduce FODMAP content in bread, selecting specific microbes responsible
for fermenting and breaking down sugars that cause gastrointestinal issues is crucial [116].
However, the extent of FODMAP reduction can vary based on the fermentation process,
the type of grain used, and the amount of sour yeast incorporated into the bread dough
[123]. There is limited research available on the production of grain-based products with
low-FODMAP contents [44]. In a study by Struyf et al. [124], the FODMAP content of
whole wheat bread was reduced by over 90% by adding K. marxianus and S. cerevisiae to
the dough. It was noted that using both yeast strains together was essential to achieve a
product with desirable sensory properties and a low-FODMAP content, as the control
bread made with only K. marxianus did not produce enough carbon dioxide for the desired
volume.
In their study, Laurent et al. [125] compared the impact of K. marxianus CBS6014
strain and S. cerevisiae commercial yeast on FODMAP levels. The CBS6014 strain of K.
marxianus has been found to exhibit greater fructan hydrolysis and lower fructan content
in rye sourdough bread, as well as white and whole-grain toast bread. While the presence
of K. marxianus in sourdough rye bread led to a reduction in fructan levels, it also posi-
tively influenced the size of the bread. This highlights the importance of the K. marxianus
strain in producing high-fiber products with a reduced FODMAP content.
In a recent study, Menezes et al. [126] examined the effects of sourdough fermenta-
tion on FODMAPs and organic acids during the production of sourdough and bread. The
levels of organic acids were initially higher during the early stages of fermentation, but
stabilized in the later stages. It was observed that all FODMAPs, except polyols, were sig-
nificantly reduced during the process. During the initial fermentation step, sucrose was
completely converted into fructose and glucose, while the concentrations of other carbo-
hydrates decreased after the fourth step. The study highlighted that sourdough bread con-
tained higher levels of organic acids and polyols, and lower levels of fructans, sucrose,
fructose, and glucose compared to bread leavened with commercial yeast. The study sug-
gests that sourdough fermentation could be utilized to create low-FODMAP wheat flour
products, as the reduction in fructan content increased from 69% to 75%.
Schmidt and Sciurba [127] analyzed the impact of extended fermentation and the ad-
dition of sourdough on FODMAPs in final products. Their findings indicate that supple-
menting sourdough alters the FODMAP composition by reducing fructan content and in-
creasing mannitol content. While breads made from refined wheat flour adhere to low-
FODMAP standards, those made from rye and whole wheat flour are considered high
FODMAP regardless of the fermentation process. Rye breads exceed the limits for fructans
and mannitol, while whole wheat breads surpass the threshold for excessive fructose con-
tent [127].
Foods 2024, 13, 1732 14 of 20

Another study conducted by Boakye et al. [128] aimed to investigate the FODMAP
contents of twenty-two different wheat cultivars grown in Minnesota, USA. The study
also examined the impact of type I sour yeast fermentation (4 and 12 h fermentation peri-
ods) on FODMAP levels in these cultivars. It was found that the levels of fructans and
raffinose decreased by 69% and 69%, respectively, in the sourdoughs produced with a 12
h fermentation period. However, there was a significant increase (550%) in mannitol levels
observed after sourdough fermentation. These results highlight the importance of moni-
toring mannitol and other FODMAPs during type I sourdough fermentation, and strate-
gies to reduce mannitol levels during fermentation and in the final product should be
considered [128].

2.8. The Impact of Sourdough Fermentation on Vitamins

Another advantage of sourdough fermentation is its ability to enhance the vitamin
content. Cereal crops contain tocopherol, vitamin B1, and folates concentrated in the seed
and bran [12,39], while pseudocereals include retinol, tocopherol, and folate [129]. Certain
strains of LAB are known to synthesize water-soluble vitamins from group B, such as fo-
lates B9, riboflavin B2, and vitamin B12 [14,130]. In addition to LAB, specific sourdough
yeast strains have shown a significant capability to increase folate levels in rye sourdough
[8]. The fermentation process with sourdough has been reported to boost folate content
[131], decrease vitamin E and tocotrienol content [131], and alter vitamin B1 content de-
pending on the processing method [132]. Therefore, the fermentation stage can impact the
overall vitamin content during the baking process [132]. It is also believed that S. cerevisiae
can promote LAB growth by producing vitamins like cobalamin, ascorbic acid, or calcif-
erol, which are not naturally present in wheat flour [133]. While sourdough is known to
enhance vitamin levels, factors such as the baking process can lead to a reduction in vita-
min content [14,134].

2.9. Sourdough Fermentation and Phenolic Compounds

Phenolic compounds are secondary plant metabolites found in significant amounts
in grains [135]. These components are known for their bitter flavor and their ability to
hinder the digestion of starch and proteins [136]. They are synthesized through various
metabolic pathways, including shikimate, phenylpropanoid, acetyl CoA, malonyl CoA,
pyruvate, acetate, and amino acids like phenylalanine and tyrosine [44]. Phenolic com-
pounds also act as antioxidants, providing various health benefits such as antidiabetic,
anticancer, anti-inflammatory, antimicrobial, antioxidant, neuroprotective, cardioprotec-
tive, and hepatoprotective functions [135,137,138]. Their chemical structure allows them
to scavenge free radicals and chelate metals by facilitating electron transfer or H+ donation
from the OH- groups of their aromatic rings [44]. Solubility is crucial for phenolic com-
pounds to exert their antioxidant effects in the human bloodstream [44]. Wheat and rye
contain primary phenolic components like phenolic acids and alkylresorcinols, with
lignans being abundant in wheat [139]. While lipophilic alkyl resorcinols in cereal bran
layers do not affect bread quality, they serve as a biomarker for whole-cereal intake [140].
Ferulic acid is the main phenolic acid in rye, constituting around 50% of total phenolic
acids, alongside caffeic acid, dihydrobenzoic acid, and sinapic acids [136].
Phenolic acids in wheat and rye are mostly found in connected figures and as dimers
[136]. The increase in the phenolic content of cereals is provided by certain factors. These
are particle size decrease, germination, supplementation of hydrolytic enzymes, and fer-
mentation [44]. In addition, it has been found that fermentation increases the antioxidant
efficiency of the phenolic part [141]. Parameters such as temperature during fermentation,
final pH, time, microorganisms added, grain kind, and cereal texture play a significant
part in the results related to the leave of bound phenolics [44]. Among the added micro-
organisms, LAB phenolic acid metabolism is achieved through reductases and decarbox-
ylases. Subsequently, decarboxylating hydroxybenzoic and hydroxycinnamic acids to
Foods 2024, 13, 1732 15 of 20

phenol or vinyl derivatives takes place. Hydroxycinnamic acids and their vinyl deriva-
tives are transformed by reductases that hydrogenate the double bond. The bioconver-
sions of phenolic acids are generally species-specific [136].
There are limited studies on the addition of grain enzymes and specific starter cul-
tures to facilitate the transformation of phenolic compounds in sourdough fermentation
[136]. The significance of phenolic components in sourdoughs stems from the presence of
these compounds in wheat flour [139]. Both whole- and white wheat flour contain some
phenolics that may not be readily bioavailable [139]. However, natural fermentation can
enhance the bioavailability of these phenolics. This is because yeast microbes produce en-
zymes that break down insoluble phenolics, making them more accessible [139]. It has
been observed that phenolic compounds in flour undergo changes during fermentation.
In previous studies, the antioxidant potential of a wheat bran–flour mixture and rye
flour was compared with sourdough fermentation. It has been reported that the antioxi-
dant efficiency following sourdough fermentation surpasses that of traditional products
[142]. However, research indicates that in addition to enhancing antioxidant capacity,
sourdough fermentation also hinders the performance of α-amylase and α-glucosidases,
impacting their activity in starch hydrolysis [143,144].

3. Conclusions
In conclusion, this comprehensive evaluation of the nutritional role of sourdough
fermentation has shed light on its diverse mechanisms and potential functional aspects.
Through this study, the significant impact of sourdough fermentation on various nutri-
tional factors, including mineral bioavailability and phytic acid, starch digestibility and
glycemic index, protein digestibility, salt reduction, dietary fiber content, gut microbiota
modulation, FODMAPs, vitamins, and phenolic compounds, has been explored. The find-
ings highlight the multifaceted benefits of sourdough fermentation in enhancing nutrient
availability, promoting digestive health, and potentially contributing to overall well-be-
ing. Further research in this area could provide valuable insights into optimizing the nu-
tritional quality of fermented foods for improved dietary outcomes.

Author Contributions: Conceptualization, methodology, investigation, and writing—original draft

preparation, Z.A.; writing—original draft preparation, F.F.; conceptualization and writing—review and
editing, H.C. and E.D. All authors have read and agreed to the published version of the manuscript.
Funding: This research received no external funding.
Institutional Review Board Statement: Not applicable.
Informed Consent Statement: Not applicable.
Data Availability Statement: The original contributions presented in the study are included in the
article, further inquiries can be directed to the corresponding author.
Conflicts of Interest: The authors declare no conflict of interest.

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