fermentation

Review
Enhancing Micronutrients Bioavailability through
Fermentation of Plant-Based Foods: A Concise Review
Mrinal Samtiya 1 , Rotimi E. Aluko 2, * , Anil Kumar Puniya 3 and Tejpal Dhewa 1, *

1 Department of Nutrition Biology, School of Interdisciplinary and Applied Sciences, Central University of
Haryana, Mahendergarh 123031, Haryana, India; [email protected]
2
Pwede dito mang-galing Department of Food and Human Nutritional Sciences, University of Manitoba,
ang explanation ko Winnipeg, MB R3T 2N2, Canada
3 Dairy Microbiology Division, ICAR-National Dairy Research Institute, Karnal 132001, Haryana, India;
[email protected]
* Correspondence: [email protected] (R.E.A.); [email protected] (T.D.)

Abstract: Plant-based foods are rich sources of vitamins and essential micronutrients. For the
proper functioning of the human body and their crucial role, trace minerals (iron, zinc, magnesium,
manganese, etc.) are required in appropriate amounts. Cereals and pulses are the chief sources of
these trace minerals. Despite these minerals, adequate consumption of plant foods cannot fulfill
the human body’s total nutrient requirement. Plant foods also contain ample amounts of anti-
nutritional factors such as phytate, tannins, phenols, oxalates, etc. These factors can compromise the
bioavailability of several essential micronutrients in plant foods. However, literature reports show
that fermentation and related processing methods can improve nutrient and mineral bioavailability



of plant foods. In this review, studies related to fermentation methods that can be used to improve


micronutrient bioavailability in plant foods are discussed.
Citation: Samtiya, M.; Aluko, R.E.;
Puniya, A.K.; Dhewa, T. Enhancing
Keywords: fermentation; antinutritional factors; micronutrients; bioaccessibility; bioavailability;
Micronutrients Bioavailability
plant foods; cereals; pulses
through Fermentation of Plant-Based
Foods: A Concise Review.
Fermentation 2021, 7, 63.
https://doi.org/10.3390/
fermentation7020063 1. Introduction
Health in general depends on food intake that meets the nutritional needs of con-
Academic Editor: Mohamed Koubaa sumers. Foods are also explored as an approach to inhibit the development of different
disorders, especially those that are due to improper, deficient or excessive intake [1]. Nutri-
Received: 9 April 2021 ents (e.g., minerals) carry out different vital functions in body like development of strong
Accepted: 16 April 2021 bones to conduct nerve impulses for sustained health [2]. Due to lack of proper nutrition,
Published: 20 April 2021 macronutrients and micronutrient deficiency occurs leading to undernutrition. Excessive
intake of these macro/ micronutrients also result in overnutrition leading to obesity and
Publisher’s Note: MDPI stays neutral associated metabolic disorders [3]. In the absence of proper consumption of Zn, Fe and
with regard to jurisdictional claims in
Ca severe malnutrition occurs (i.e., high risk of infection, anemia and osteoporosis, respec-
published maps and institutional affil-
tively). These micronutrients are adequately present in beverages and foods that fulfill
iations.
body’s daily needs; however, due to their low bioavailability, people face severe micronu-
trient deficiencies [4]. In the developing world, occurrence of micronutrient insufficiencies
adversely affects the health of vulnerable people. The main factor for deficiency of key
micronutrients (i.e., Zn, vitamin A, and Fe) is the limited bioavailability and poor diet
Copyright: © 2021 by the authors. quality [5]. To achieve sufficient intake of minerals, some countries recommend dietary
Licensee MDPI, Basel, Switzerland. guidelines to increase consumption of whole grains, as plant-based foods are excellent
This article is an open access article
sources of nearly all the vital nutrients [6]. However, the presence of different antinutri-
distributed under the terms and
tional factors such as polyphenolics and phytic acids could impede the immense nutritional
conditions of the Creative Commons
influence of whole grains [7,8]. Fermentation, cooking, soaking, puffing and germination
Attribution (CC BY) license (https://
are some of the traditional food processing methods used to reduce the antinutritional
creativecommons.org/licenses/by/
components and enhance protein digestibility [9–12]. Due to this, fermented foodstuffs
4.0/).

Fermentation 2021, 7, 63. https://doi.org/10.3390/fermentation7020063 https://www.mdpi.com/journal/fermentation

Fermentation 2021, 7, 63 2 of 13

have been a vital part of human nutrition and is essential in several developing nations
where these are consumed as part of indigenous traditions [13–15] Fermented foods are
produced by using lactic acid bacteria (LAB) comprised of a number of genera (e.g., Pedic-
occus, Lactobacillus, Streptococcus, Lactococcus, Leuconostoc, Enterococcus, etc.) that yield lactic
acid as a major metabolite. The fermentation is carried out by bacteria, yeast, filamentous
fungi or a combination of these [13,16]. During fermentation, the microbial metabolism
enhances the macromolecules’ digestibility and improves the bioavailability of macro/
micronutrients and phytochemicals. For the removal of antinutrients, allergens and toxins,
fermentation is considered as one of the most effective processing methods [17]. Phytic acid
salts, also known by the name phytate or myo-inositol hexakisdihydrogen phosphate, are
the central storage (1–5% by weight) form of both myo-inositol and phosphate in oilseeds,
cereal grains, legumes, plant seeds and nuts [18]. Phytic acid salts commonly found in
plant-based foods are shown in Table 1.

Table 1. Phytate content in common plant-based foods.

Plant Food Phytate Content (%), Dry Weight Basis

Almonds 0.35–9.42
Beans 0.61–2.38
Brazil nuts 0.29–6.34
Cashew nuts 0.19–4.98
Chickpeas 0.28–1.60
Corn 0.72–6.39
Eggplant seed 1.42
Kiwi fruits 1.34
Lentils 0.27–1.51
Peas 0.22–1.22
Rapeseed 2.50–7.50
Rice 0.06–8.70
Sesame seed 1.44–5.36
Soybean 1.00–10.7
Tomato seed 1.66
Walnuts 0.20–6.69
Source: [19–22].

Phytic acid chelates the multivalent positive cations (e.g., Mg2+ , Zn2+ , Ca2+ , Fe2+ , Mn2+
and amino group derivatives in protein moieties), which leads to reduced nutrient solubility,
bioavailability and absorption [16]. Fermentation also improves the mineral bioavailability
by producing a phytase enzyme that degrades the phytic acids in plant foods. Such a
phytic acid reduction may enhance the level of calcium, iron and zinc several-fold [23]. In
this review, we emphasize the fermentation used to improve the bioavailability of minerals
that could be further utilized to enhance the level of micronutrients in plant-based foods
alleviating micronutrient deficiency.

2. Fermentation
Fermentation is a type of metabolic processing that oxidizes carbohydrates to release
energy in absence of the external electron acceptors. LAB are acid-tolerant, catalase-
negative, non-spore-forming, Gram-positive, fastidious, cytochrome deficient, aerotolerant
and fermentative microorganisms [24,25]. As per the expert panel report of The Inter-
national Scientific Association for Probiotics and Prebiotics (ISAPP), a fermented food is
“food made through desired microbial growth and enzymatic conversions of food compo-
Fermentation 2021, 7, 63 3 of 13

nents” [26]. Primarily people consume processed foods; among those, fermented foods
have been the very first ones. Fermented foods are linked to the cultural relationships
between different nations and associated with its own population’s nutritional status.
Fermentation gained much interest of researchers to understand the mechanism and for
commercialization of food products [27]. It is a typical method applied to improve the
organoleptic, nutritional and shelf-life properties of food ingredients [28]. Fermented
foods have several functional attributes that provide full benefits to the consumers (i.e.,
antioxidative properties, production of enzymes, probiotic potentials, antimicrobial fea-
tures, bioactive peptides, etc.) [29]. Microorganisms used in fermentation can produce a
range of enzymes (i.e., phytase, protease, lipase, amylase, etc.) that can hydrolyze lipids,
carbohydrates and proteins into simple digestible constituents with desirable texture and
taste. Microbial enzymes may also degrade different antinutritional such as protease in-
hibitors, tannins and phytates. This has increased much interest in using fermentation for
the improvement of bioavailability and absorption of minerals present in plant seeds [30].
Use of microbial enzymes as human food ingredients remains a challenge from safety
point of view [31]. Efforts were made to isolate microorganisms that can produce phytase
and are safe for human consumption [32]. During food fermentation, endogenous enzymes
are activated due to lower pH that contribute to the reduction of phytic acid. Fermentation
improves mineral (i.e., Ca, Zn and Fe) bioavailability by producing phytases that reduce
phytic acid constituents in plant-based foods [23,33]. Aside from this, fermentation is used
as an enzymatic and microbial food processing method. Although it enhances the quality
of raw food components by improving safety, antinutrients elimination, extended shelf-
life, attractive flavor and nutritional enrichment [11], by lowering polyphenol and phytic
acids, fermentation may also improve the bioavailability of minerals through production
of organic acids [34]. There are different fermented foods (i.e., pearl millet fermented gruel,
lamtoro tempeh, corn steep liquor, soybean tempeh, shalgam, gembus tempeh, kimchi,
uttapam batter, sourdough, kara tempeh and other ethnic products) that can be used as a
source for the isolation of phytase-producing LAB [35–39]. Graphical abstract represents
that fermentation not only improve the plant food proteins and carbohydrates digestibility
but also enhances minerals bioavailability by increasing phytase production that catalyzes
removal of phosphate from phytic acid resulting into myo-inositol, phosphate and other
minerals and nutrients.

3. Fermentation Affects Enzyme Activation to Improve the Nutritional Value of

Plant-Based Food
In modern times, the food and nutrition safety processing of products derived from
agriculture is very important. Due to developments such as urbanization, food produced
and processed in the remote areas, for feeding purpose of the growing population, is
transported into cities and towns. Moreover, due to seasonality, agricultural products
should be processed so that foods can be accessible throughout the year [40].
In developing nations, legume and cereal starches provide major calories due to the
high contents of carbohydrates in these crops [41]. Previous evidence shows that the
fermentation process stimulates enzymes such as maltase and α-amylase that degrade
starch into simple sugars and maltodextrins. Furthermore, studies have emphasized
that, in the early fermentation phase, glucose is increased due to activated α-amylase
and maltase [42–44]. Moreover, fermentation subsequently enhances the ethanol and
carbon dioxide production and reduces the starch component in different varieties of
millet in addition to facilitating phytase enzyme production and considerable reductions
in pH [42,45,46].
Earlier literature reports show that the effect of fermentation on protein content
is unpredictable, which could be due to different study times, designs of experiments
and variations in the initial amino acid or protein profile of the foods. Some studies
proposed that fermentation could increase or decrease the amino acid and protein contents
of foods [47–49]. Due to the fermentation process, relative changes occur by dry matter
loss because of the action of microorganisms when they metabolize and hydrolyze fats and
Fermentation 2021, 7, 63 4 of 13

carbohydrates as energy sources. However, plant protein digestibility increases during the
fermentation process [40]. A previous study showed that, due to loss of carbohydrates,
protein content was increased after 24 h of fermentation but arginine, glycine and lysine
levels were reduced [43]. On the other hand, it has been reported that the use of amino
acids as nutrients by fermenting microbes reduced the quality and content of proteins
in certain fermented foods [50]. In general, the digestibility of plant proteins is much
lower when compared to animal-derived proteins. Therefore, plant-derived proteins
may affect the gastrointestinal environment, because of the increased protein excretion
in the fecal matter. Thus, the undigested protein level could be reduced by increasing
protein digestibility, which could reduce food allergies associated with decreased intestinal
absorption of proteins [51].
Fermentation can also reduce the content of several plant compounds such as phytic
acid, oxalate, tannins, carbohydrates and various protein complexes in addition to microbial
production of digestive enzymes [10,52]. An in vitro study by Pranoto et al. [50] assessed
the influence of 36 h natural fermentation and Lactobacillus plantarum on the digestibility
of sorghum flour proteins. They found that protein digestibility was enhanced 92% and
47% by L. plantarum and natural fermentation, respectively. These findings suggest that
L. plantarum increased the level of proteolytic enzymes better than natural fermentation,
which led to better degradation of the protein–tannin complexes as well as other protein
complexes to liberate amino acids and peptides [50].
Minerals present in plant products have very little bioavailability due to their forma-
tion of complexes with non-digestible matter. These minerals are bonded and entrapped in
the complex matrices, which is the main factor responsible for the lower bioavailability.
Fermentation is one of the suitable methods that could degrade these complexes to make
minerals free, easily assessable and bioavailable [40,50]. The level of certain fermented
food minerals such as zinc, calcium, iron and magnesium are increased by fermentation,
which is connected with the reduced concentration of phytates [50,53]. Nonetheless, during
fermentation, mineral content increases may also be due to the loss of dry matter, as fer-
menting microorganisms degrade proteins and carbohydrates [54]. By degrading phytates
and oxalates, factors that form complexes with minerals, fermentation also enhances iron,
phosphorous and calcium bioavailability [55,56]. This is because embedded minerals ions
are loosened from the complex matrix by the fermentation process. Both α-amylase and
phytase, which are produced during microbial fermentation, degrade starch and phytate
contents, respectively, to loosen the complexes. Additionally, certain fermenting microbes
can breakdown the dietary fiber materials to further loosen the food matrix [57]. Con-
sequently, fermentation effects are also influenced by food composition, as other food
constituents such as fibers could reduce specific mineral availability. To overcome these
difficulties, seed germination and incubation of foods with polyphenol oxidase or phytase
(through fermentation) could help enhance the availability of minerals by decreasing the
phytate or tannin levels [58,59]. During fermentation, pH is reduced, which enhances the
absorption of iron because of ferric iron conversion to the more readily absorbed ferrous
iron. Furthermore, fermentation leads to an optimal pH environment, which is necessary
for phytase activity to degrade phytate. Besides, if fermentation is preceded by grinding
the food materials, there is enhanced mineral bioavailability. The positive effect of grinding
is due to breaks in the cellular structure of the food material coupled with enlarged surface
that facilitates dispersion of phytase for a more effective breakdown of phytate [40,46].
For a very long time, the significance of phytochemicals to the health and nutrition
of humans was not well understood because they were considered as non-functional sec-
ondary metabolites of plants that are synthesized during plant development by shikimate
pathways and phenyl propanoid biosynthesis [60,61]. However, fermentation has shown
both adverse as well beneficial effects on phytonutrients. A previous study reported that
LAB fermentation of soybean germs for 48 h considerably reduced tocopherol, glycosy-
lated soyasaponin and phytosterol contents. For example, the phytosterol content was
reduced from 4.2 to 1.1 mg/g at the end of the experiment. The findings suggest that
Fermentation 2021, 7, 63 5 of 13

the reduction of glycosylated soya saponins maybe because of their transformation from
2,3-dihydroxy-2,5-dihydroxy-6-methyl-4H-pyran-4-one (DDMP) to non-DDMP forms [62].
Wang et al. [63] evaluated fermentation effect on the antioxidant attributes of four types
of cereals using L. plantarum and Bacillus subtilis. They concluded that, by using these
starter cultures, there was a considerable increase in the total flavonoid and total phenolic
acid contents [63]. Consequently, the capability of fermentation to enhance antioxidant
attributes of foods can be explored as a gainful approach to alleviate the oxidative stress
within the human body by consuming these foods [64–66].
Literature reports suggest dual effects of fermentation on the glycemic index (GI) of
foods. Certain studies have confirmed that fermentation increased the GI [67–69], although
some evidence indicates that GI decreased [70,71]. The low glycemic index of fermented
foods may be due to the microbial conversion of glucose to short-chain organic acids
such as propionic acid, acetic acid and lactic acid, which reduces the amount of glucose
that is absorbed into blood circulation [72,73]. Ostman et al. [74] also suggested another
mechanism for the lowering of glucose, which involves pH reduction by the lactic acid and
consequent reduction in the activity of starch-hydrolyzing enzymes [74]. The mechanism
of action for the acetic and propionic acids is different from the lactic acid mechanism;
these organic acids reduced the gastric emptying rate in addition to decreasing the activity
of enzymes [40]. This mechanism may be one of the reasons that sourdough bread reduces
the postprandial glucose concentration and enhances glucose homeostasis in a healthy
individual. Based on these data interpretations, it is reasonable to conclude that the natural
fermentation of sugars and starch lead to formation of propionic and lactic acids that
decrease glucose level in foods, which leads to low GI [71,73,75].

4. Influence of Fermentation on Micronutrients Bioavailability of Plant-Based Foods

In several countries, traditional fermented foods are a significant portion of their
routine dietary needs, although, in certain countries, these foods are recommended for
regular consumption [76]. Fermented foods consumptions are frequently associated with
several health benefits. Certain fermented foods that comprise probiotics bacteria enhance
nutritional attributes by improving mineral bioavailability and forming bioactive com-
pounds [17]. Plant-derived food micronutrient bioavailability is usually improved by
fermentation, which provides better shelf-life to foods and with enhanced gut microbial
ecology that promotes a healthy status. These health benefits are mainly due to the activities
of fermenting microorganisms known as probiotics, specifically the lactic acid producing
type [77]. Lactobacillus and Lactococcus are the most frequently used LAB culture, although
others, such as molds and yeast, are also used for fermentation purposes [78]. Due to
these health benefits provided by fermented foods/products, fermentation may become
an effective approach to mitigate the mineral deficiency concerns among the vulnerable
populations, particularly in the developing nations where unprocessed pulses/cereals are
significantly consumed [40].
Some evidence confirms that different fermentation processing methods improved
the mineral bioavailability in plant-originated foods. A previous study confirmed the
potential of five LAB strains in relation to mineral bioavailability improvement. The results
confirm that L. fermentum B4655, L. plantarum B4495, L. casei B1922, L. bulgaricus CFR2028
and L. acidophilus B4496 strains of LAB reduced the phytic acid content when used for
the soymilk fermentation at 37 ◦ C for 24 h. Moreover, the results show an increase in the
level of Mg and Ca of fermented soymilk when compared to the control [79]. Bahaciu
et al. [80] recently investigated the effects of germination and fermentation (Lactobacillus)
on the mineral bioavailability of soybean seeds. They detected that germination of soybean
seeds for four days at 25 ◦ C increased the level of Zn, Mg, Fe and Ca by 28.04%, 15.77%,
22.31% and 48.76%, respectively in comparison to control samples. Furthermore, they
confirmed that germination followed by fermentation led to 40.87%, 43.41%, 59.56% and
53.4% increases for Zn, Mg, Fe and Ca, respectively, which are higher than values obtained
for germination alone. In addition, optimal levels of mineral bioavailability were obtained
Fermentation 2021, 7, 63 6 of 13

in soybean seeds that germinated for four days at 25 ◦ C followed by 72 h of lactic acid
fermentation in a medium containing 3% saccharose [80]. Previous findings also confirm
the effect of natural fermentation on mineral bioavailability in sorghum grain and finger
millet. Results showed that natural fermentation reduced the phytic acid of sorghum
grain and finer millet by 64.8% and 72.3% after 96 h, respectively. Furthermore, 39.0%
and 54.3% of phytic acid levels decreased in sorghum grain and finger millet after 72 h of
natural fermentation, respectively. The results show that 72–96 h of natural fermentation
increased the bioavailability of calcium, manganese and iron in both finger millet and
sorghum grains [81]. There is evidence that fermentation with L. plantarum strains reduced
the phytic acid content and enhanced the availability of Ca, Zn and Fe of quinoa flour.
For instance, a study confirmed that fermentation (16–18 h, at 30 ◦ C) of quinoa flour
by L. plantarum improved the iron content significantly when compared to unfermented
samples in addition to substantial decrease in the phytic acid content. Furthermore, the
highest phytic acid reduction (98%) was observed after fermentation of the germinated
flour [82]. Moreover, Castro-Alba et al. [83] recently reported that fermentation (4 or
10 h, at 30 ◦ C) of milled quinoa seeds using L. plantarum 299v reduced the phytic acid
significantly and improved the bioavailability of minerals such as Ca, Fe, and Zn [83]. In
another previous study, the effect of natural sourdough fermentation of wheat showed a
much better improvement of copper and magnesium absorption in comparison to non-
fermented wheat flour, using male Wistar rats. The authors found that fermented wheat
flour samples by both natural fermentation as well as Saccharomyces cerevisiae enhanced Zn
and Fe absorption. The results suggest that the reduction of phytic acid could be the reason
for the enhanced mineral bioavailability [84]. The latest study by Chiş et al. [85] confirmed
that fermentation of rice sourdough for 24 h with L. Spicheri DSM 15429 enhanced the level
of manganese, zinc, copper, potassium, magnesium and calcium by 1.68, 1.92, 2.55, 1.98, 1.94
and 2.3 folds, respectively. Moreover, mineral levels improved by 0.7–1.52-fold after 24 h of
spontaneous sourdough fermentation, compared to starting level before fermentation [85].
In addition, earlier research reported that fermentation by Pediococcus pentosaceus KTU05-8
and P. pentosaceus KTU05-9 strains, which are phytase-producing organisms, improved
the whole meal wheat flour minerals (phosphorus, calcium, manganese, zinc, and iron)
solubility by 30% [86]. Furthermore, Anastasio et al. [87] found that the phytase-producing
LAB, (L. plantarum H5 and Enterococcus faecium A86) produced significant 98% increase
in the solubility of iron and manganese when used for brown dough fermentation. In
comparison, the non-phytase producing LAB strains produced lower mineral solubility
during bacterial fermentation [87]. A previous study confirmed the potential of Leuconostoc
mesenteroides FSC2 and Lactobacillus pentosus FSC1 in relation to the bioavailability of
minerals content. The authors found that fermentation of carrot juice by L. mesenteroides
FSC2 and L. pentosus FSC1 improved the solubility of minerals such as Cu, Zn, Fe and
Mn by 1-, 1.2-, 1.5–1.7- and 2-fold, respectively. Furthermore, they showed that LAB-
fermented juice improved the cellular (Caco-2 cell line) uptake of iron by 6–7-fold [88]. Giri
et al. [89] recently confirmed the capability of L. plantarum L7 in favor of improving the
bioavailability of minerals by preparing a rice-based fermented drink. The results show
that the phytase activity of the LAB strain was responsible for the considerable increases in
the iron, manganese, magnesium, calcium and sodium levels of the fermented drink [89].
Furthermore, other evidence confirms the ability of lactic acid fermentation of plant-based
diet (cassava) in relation to the enhancement of Zn bioavailability using the rat model.
Results have found absorption of Zn level was 40.2% in the animal model that consumed
LAB-fermented cassava when compared to 16.5% in those that consumed the unfermented
sample. In addition, higher zinc levels have been shown to be present in femur, serum
and liver of animals that consumed a LAB-fermented diet [90]. A previous clinical study
by Scheers et al. [91] confirmed that fermentation enhanced iron bioavailability in human
volunteers who consumed a L. plantarum fermented vegetable diet when compared to the
group who consumed the non-fermented diet [91]. A recent research study reported the
impacts of fermentation on the bioavailability of Zn in hydrothermally treated, malted and
Fermentation 2021, 7, 63 7 of 13

native finger millet seeds. The authors found that L. pentosus CFR3 fermentation reduced
the phytate content in the hydrothermally treated, malted, and native finger millet seed coat
by 87.8%, 66.6%, and 56.7%, respectively. Moreover, the findings confirm that zinc in vitro
bioaccessibility improved by 12.1%, 34.6% and 28.4% in hydrothermally treated, malted
and native finger millet seed coat, respectively, after 24 h of fermentation [92]. Chawla
et al. [93] evaluated the effect of fermentation on the in vitro Fe and Zn bioavailability in
black-eyed pea (cowpea) flour by using Caco-2 cell line and gastrointestinal digestion study.
They confirmed that 96 h of solid-state fermentation of black-eyed pea flour by Aspergillus
oryzae led to enhanced in vitro bioaccessibility of Zn and Fe from 14.3% to 29.6% and
17.2% to 30.2%, respectively. The improved mineral bioaccessibility was attributed to the
antinutritional content degradation during fermentation. Furthermore, after fermentation,
Zn and Fe uptakes by Caco-2 cells were also enhanced from 18% to 28% and 22% to 32%,
respectively [93]. Khodaii et al. [94] estimated the influence of lactic acid fermentation
of bread dough on iron absorption using serum ferritin in rats and a Caco-2 cell line
model. They showed that ferritin formation was considerably enhanced in the in vivo
(serum) and in vitro (cell line) models in comparison to the controls when L. acidophilus
was added into the dough [94]. Other previous evidence confirms that lactic fermentation
could reduce the antinutrients content such as phytic acid, tannins and hence improve the
mineral bioavailability of different cereals [95]. Carrizo et al. [96] confirmed the potential
of L. plantarum strains fermented quinoa sourdough to prevent minerals and vitamin
deficiency using an in vivo mouse model. The findings show that mice that consumed
the L. plantarum CRL 1964 and L. plantarum CRL 2107-fermented pasta (quinoa) had
enhanced blood levels of vitamins B9 and B2. Furthermore, supplementation of mice
diet with fermented pasta led to 4.85, 0.37, 10.70 and 18.75 mg/dL blood levels of Mg+2 ,
Fe+2 , Ca+2 and P, respectively, compared to 3.34, 0.26, 9.90 and 9.85 mg/dL for the group
that consumed the non-fermented diet [96]. Table 2 shows the studies related to the
enhancement of mineral bioavailability using different fermentation methods.

Table 2. Studies showing that fermentation influence the bioavailability of the minerals.

Types of Food Fermentation Organisms Results References

Lactobacillus acidophilus B4496,
Lactobacillus bulgaricus CFR2028,
Increased calcium and magnesium
Soymilk Lactobacillus casei B1922, Lactobacillus [79]
availability
plantarum B4495 and Lactobacillus
fermentum B4655
Increased calcium (53.4%), iron
Soybeans Lactobacilli (59.56%), magnesium (43.41%) and [80]
Zinc (40.87%) levels
Improved solubility of Fe, Ca, Mg
Sorghum and finger millet Natural fermentation [81]
and Mn.
Quinoa seeds L. plantarum Iron solubility increased [82]
Reduced phytic acid content and
Quinoa seeds L. plantarum [83]
improved Zn, Fe and Ca availability
Saccharomyces cerevisiae and natural Increased the apparent absorption
Wheat flour [84]
fermentation of Fe and Zn
Improved levels of calcium
(2.3-fold), magnesium (1.94-fold),
Rice Lactobacillus Spicheri DSM 15429 potassium (1.98-fold), copper [85]
(2.55-fold), zinc (1.92-fold), and
manganese (1.68-fold).
Increased solubility of iron, zinc,
Pediococcus pentosaceus strains KTU05-8
Wheat manganese and calcium and [86]
and KTU05-9
phosphorus (average of 30%).
Fermentation 2021, 7, 63 8 of 13

Table 2. Cont.

Types of Food Fermentation Organisms Results References

Improved mineral solubility:
manganese (2.2–2.5-fold); iron
(1.5–1.7-fold); Zinc (1.2-fold);
Lactobacillus pentosus FSC1 and
Carrot Copper (1-fold). Cellular (Caco-2) [88]
Leuconostoc mesenteroides FSC2
uptake of ferrous iron in LAB
fermented juice improved by 6 to
7-fold.
Increased in the levels of free
Rice L. plantarum L7 minerals like sodium, calcium, [89]
magnesium, manganese and iron
Cassava Spontaneous fermentation Increased bioavailability of zinc [90]
Vegetable mix (carrots,
turnips, white cabbage, L. plantarum Bioavailability of iron increased [91]
parsnip, celery and onion)
Increased in the bioavailability and
Finger millet seed L. pentosus CFR3 [92]
the in vitro bioaccessibility of zinc.
In vitro bioaccessibility of iron and
zinc after 96 h of fermentation
increased from 17.2% to 30.2% and
14.3% to 29.6%, respectively.
Black-eyed pea Aspergillus oryzae [93]
Uptake of iron and zinc by Caco-2
cells similarly improved from 22%
to 32% and 18% to 28%, respectively
after fermentation.
Increased iron absorption; ferritin
formation increased significantly
Whole meal flour (bread) L. acidophilus [94]
compared to controls in the
intestinal cells (in vitro)
Improved the nutritional status,
L. plantarum CRL 2017 and L. plantarum
Quinoa pasta improve calcium, iron and [96]
CRL 1964
magnesium levels in the blood

5. Future Perspectives
Fermentation is a processing method that involves microorganism and their enzymes
to carry out primary food matrix biochemical modifications. Fermentation not only im-
proves the shelf-life and organoleptic properties of food products but also increases their
nutrient bioavailability and bioaccessibility [40]. The microorganism that is most commonly
used for food fermentation is LAB, which produce lactic acid that contributes to enhanced
food preservation. Around 4000 years BC, LAB were used in the making of yogurt, when
the Thracians farmed sheep. There are several commercial microbial enzymes that have
been commonly used to enhance the functional value of plant phytochemicals, yet fermen-
tation (lactic acid) is favored to advance the foods nutraceutical significance because it is
not so expensive while increasing the overall nutritional and organoleptic properties [25].
Fermentation has gained much interest due to the considerable health benefits, which
it imparts on food products. Besides, fermentation does not require any complicated
equipment and needs very low energy input. Therefore, it could be a feasible solution for
the developing nations to valorize local raw ingredients for improved local economies,
in addition to being a suitable strategy to overcome malnutrition because it enhances the
nutritional value of food products [97]. Several methods could possibly be used to improve
product yield, reduce the cost and proficiency of fermentation procedures. Strategies that
have been considered include developing technologies to simplify fermentation along
with the development of new, more robust and effective cultures through old and current
Fermentation 2021, 7, 63 9 of 13

“omic” practices; the use of analytical technologies such as real-time monitoring systems
to measure processing factors such as dissolved oxygen, pressure, temperature, pH, etc.;
and automated fermenters with upgraded in-built controls to direct the fermentation pro-
cess [17,98]. An emerging trend is the use of bacterial probiotics culture and cultures that
could address the functionality and safety of the fermented product [99].

6. Conclusions
People all over the world are facing mineral deficiency concerns, especially in devel-
oping countries. This is because people consume ample amounts of cereals and pulses in
their diets, and these plant foods are rich sources of minerals such as iron, zinc, magnesium,
calcium, etc. However, these plant foods also contain high levels of antinutritional factors,
especially phytic acid that binds and chelates these mineral ions to reduce absorption and
bioavailability in the gut. Phytate hinders the absorption of micronutrients in the human
diet because of the absence of intestinal phytase enzymes. The presence of the phytate and
other antinutrient components makes micronutrients unavailable for human absorption
and is one of the causes of mineral malnutrition worldwide. Several approaches have been
used to increase the bioavailability of minerals ions from plant-based food. Fermentation
is one of the best strategies that can be used traditionally and on an industrial scale to
enhance mineral accessibility from plant foodstuffs. Fermentation processing methods
that improve minerals availability could be transferred to local communities to improve
minerals intake in the households. Studies have shown that natural fermentation and
strain-specific fermentation provide better mineral bioavailability through a more effective
degradation of the plant food’s phytic acid content. It also provides viable environmental
conditions (such as low pH) that help the growth of beneficial strain and enhance phytase
activity. Additionally, during fermentation, metabolic activities of microbes improve the
nutritional, physical and sensory properties of food products. Overall, the findings in this
review suggest that selective/appropriate fermentation processing methods may be useful
to produce functional foods with improved bioavailability of essential micronutrients,
which could contribute to alleviated micronutrient deficiency.

Author Contributions: M.S. prepared the original draft of the manuscript; M.S., R.E.A. and T.D.
were involved in substantial revision of the original draft of the manuscript; R.E.A., A.K.P. and T.D.
were involved in supervision, conceptualization and revision of the final draft of the manuscript; All
authors have read and agreed to the published version of the manuscript.
Funding: Not applicable.
Institutional Review Board Statement: Not applicable.
Informed Consent Statement: Not applicable.
Data Availability Statement: Not applicable.
Acknowledgments: Author (Mrinal Samtiya) is thankful to SERB-DST (File No. ECR/2016/001893)
India for the award of Junior Research Fellowship (Financial support).
Conflicts of Interest: The authors declare no conflict of interest.

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