TYPE Mini Review

PUBLISHED 01 December 2022

DOI 10.3389/fmicb.2022.1070773

Mechanism and application of

OPEN ACCESS fermentation to remove beany
flavor from plant-based meat
EDITED BY
Changyu Zhou,
Ningbo University,
China

REVIEWED BY
analogs: A mini review
Hao Dong,
Zhongkai University of Agriculture and
Engineering, China
Anqi Tao 1†, Hongyu Zhang 1†, Junnan Duan 1†, Ying Xiao 1,2, Yao
Zhang Jiamin, Liu 3, Jianwei Li 4, Jieyu Huang 4, Tian Zhong 1* and Xi Yu 1,2*
Chengdu University,
China 1
Faculty of Medicine, Macau University of Science and Technology, Macau, Macau SAR, China,
Liao Guozhou, 2
Guangdong-Hong Kong-Macao Joint Laboratory for Contaminants Exposure and Health, School
Yunnan Agricultural University, of Environmental Science and Engineering, Institute of Environmental Health and Pollution Control,
China Guangdong University of Technology, Guangzhou, China, 3 School of Pharmacy and Food Science,
Zhuhai College of Science and Technology, Zhuhai, China, 4 Macau Uni-Win Biotechnology Co.,
*CORRESPONDENCE
Ltd, Macau, Macau SAR, China
Tian Zhong
[email protected]
Xi Yu
[email protected] Over the past few decades, there has been a noticeable surge in the market
†
These authors have contributed equally to of plant-based meat analogs (PBMA). Such popularity stems from their
this work environmentally friendly production procedures as well as their positive
SPECIALTY SECTION health effects. In order to meet the market demand, it is necessary to look
This article was submitted to for plant protein processing techniques that can help them match the
Food Microbiology,
a section of the journal quality of conventional meat protein from the aspects of sensory, quality
Frontiers in Microbiology and functionality. Bean proteins are ideal options for PBMA with their easy
RECEIVED 15October 2022 accessibility, high nutrient-density and reasonable price. However, the high
ACCEPTED 11 November 2022 polyunsaturated lipids content of beans inevitably leads to the unpleasant
PUBLISHED 01 December 2022
beany flavor of soy protein products, which severely affects the promotion
CITATION
of soy protein-based PBMA. In order to solve this issue, various methods
Tao A, Zhang H, Duan J, Xiao Y, Liu Y, Li J,
Huang J, Zhong T and Yu X (2022) including bleaching, enzyme and fermentation etc. are developed. Among
Mechanism and application of fermentation these, fermentation is widely investigated due to its high efficiency, less harm to
to remove beany flavor from plant-based
meat analogs: A mini review. the protein matrix, targeted performance and low budget. In addition, proper
Front. Microbiol. 13:1070773. utilization of microbiome during the fermentation process not only reduces
doi: 10.3389/fmicb.2022.1070773
the unpleasant beany flavors, but also enhances the aroma profile of the final
COPYRIGHT product. In this review, we provide a thorough and succinct overview of the
© 2022 Tao, Zhang, Duan, Xiao, Liu, Li,
Huang, Zhong and Yu. This is an open-
mechanism underlying the formation and elimination of beany flavor with
access article distributed under the terms associated fermentation process. The pros and cons of typical fermentation
of the Creative Commons Attribution technologies for removing beany flavors are discussed in alongside with their
License (CC BY). The use, distribution or
reproduction in other forums is permitted, application scenarios. Additionally, the variations among different methods are
provided the original author(s) and the compared in terms of the strains, fermentation condition, target functionality,
copyright owner(s) are credited and that
the original publication in this journal is
matrix for application, sensory perception etc.
cited, in accordance with accepted
academic practice. No use, distribution or KEYWORDS
reproduction is permitted which does not
comply with these terms. beany-flavor, plant-based meat analogs, fermentation, soy proteins, pea proteins,
off-flavor

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Tao et al. 10.3389/fmicb.2022.1070773

Introduction to cover the off flavors as a current mainstream solution. However,

the addition of extra seasonings may also have an impact on the
The Food and Agriculture Organization of the United Nations overall taste and bring about adverse health issues for the
estimates that in 2019, humans consumed about 3.25 million tons consumers. Other methods to reduce the beany flavor include
of meat; demand for meat is expected to increase by another 12% temperature control, enzyme treatment, acid–base treatment,
by 2029; and by 2050, demand for meat will increase by about supercritical CO2 extraction, new cultivars breeding, genetic
70%. If traditional meat production and management patterns engineering, etc. (Zhang et al., 2012). However, some of these
remain unchanged, an additional 30%–50% of the land will methods have hidden risks for food safety and quality, while the
be needed for livestock and meat production by then (Chen et al., high cost and extra energy consumption also bring about new
2019). Moreover, such situation of undersupply can be further problems (Kumari et al., 2015; Nedele et al., 2021; Korma et al.,
intensified by natural disasters showing up every now and then. 2022). Therefore, searching for novel technology to remove beany
Animal diseases such as African swine fever and avian influenza off-flavors instead of covering them is significant to improve the
in 2018 caused an increasing number of small and medium-sized overall taste and future development of PBMA (Sun et al., 2022).
livestock companies to exit the market. During the Coronavirus Recently, through the effort of modern microbiologists, it is
Disease 2019 (COVID-19) pandemic, the meat, poultry, and more and more revealed that fermentation using microorganisms
animal product processing plants were the sectors most affected, can effectively remove the beany flavor from next-generation
which leads to imbalance between global supply and demand for plant-based food products such as PBMA. Moreover, a new
meat products (Chen and Yang, 2021). This situation has now aromatic taste can be developed during the fermentation to mask
triggered an increase in the price of traditional meat products. The or cover the beany flavor sometimes (Wang Z. et al., 2022). Other
global agriculture and meat industry is facing serious challenges than that, fermentation has many additional benefits, such as
considering factors such as increasing pressure on the natural adjusting the gut microbiota and remediating the detriment on the
environment, population growth, consumer trends in health and gut epithelium caused by food additives (Yu and Zuo, 2021). This
environmental protection, and food innovation (Kumar et al., review discusses the mechanism of the formation of beany odors,
2017). To relieve the supply pressure of meat products, the the mechanism of using fermentation to remove them, and the
development of plant-based meat analogs (PBMA) can be an applications and the differences of various traditional and newly
effective way to address the imbalance between meat production emerged techniques. In the end, summary and discussion are
and consumption (Taufik et al., 2019; Sun et al., 2022). made about the possibilities to utilize and improve current
Recent PBMA research and development has been focusing fermentation technique to better develop our food for future
on utilizing raw materials such as soy and pea proteins to mimic signatured by PBMA.
the flavor, smell, appearance and texture of traditional meats. It is
beneficial for the whole mankind not only in terms of promoting
a sustainable development, but also from a nutrition aspect The formation of beany flavor
(Kyriakopoulou et al., 2021). Compared to traditional meat, the
energy, land, and water consumption of plant-based meat is much Unsaturated fatty acids are the main cause of beany flavor
lowered. That is why this new star can effectively mitigate carbon formation in legume-based foods. The formation of off-flavor
emissions and moderate global environmental change (Pimentel compounds usually results from the oxidation of unsaturated fatty
and Pimentel, 2003). Many epidemiological studies have shown acids and the hydrolysis of lipids (Jelen, 2011). Legume seeds
that long-term meat consumption, especially red and processed contain a large amount of unsaturated fatty acids, the most
meats, increases the incidence of digestive cancer, cardiovascular abundant being oleic acid, linolenic acid, and linoleic acid
disease and hypercholesterolemia (Cordelle et al., 2022). (Khrisanapant et al., 2019). The oxidation mechanism generally
Conversely pieces of evidence support the health benefits of consists of three categories: automatic oxidation, photo-oxidation,
consuming a plant-based diet and increasing the intake of and enzymatic oxidation (Wang Y. Q. et al., 2022).
legumes. Significantly reduces the incidence of heart disease, high
blood pressure, stroke, and type 2 diabetes (Polak et al., 2015).
Nevertheless, despite the many benefits of PBMA, their Automatic oxidation
market share is still low at around 1% (Choudhury et al., 2020).
The major bottleneck for developing ideal products which can As shown in the yellow route in Figure 1, automatic oxidation
cater the public lies in the texture as well as the taste. Other is a free radical chain reaction involving oxygen and unsaturated
problems include high energy consumption of the production lipids (Wang Y. Q. et al., 2022). Principally, reactive singlet oxygen
line, rough finished products and premature control technology. attacks H, forming α-methylene near the double bonds, thus
Currently, the key problem with the flavor of PBMA is that soy forming alkyl radicals (R·). After that, further oxygen absorption
and pea proteins have an unpleasant beany-flavor which hinders leads to the formation of peroxyl radicals (ROO·) and hydrogen
consumer preference and acceptability (Mittermeier-Kleßinger peroxide, which ends up with a wide range of volatile and
et al., 2021). Flavorings are added during the production process non-volatile secondary products, odor compounds (Wang et al.,

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Tao et al. 10.3389/fmicb.2022.1070773

FIGURE 1
The generation and removal mechanism of beany-flavor.

2021). Once the chain reaction starts, it is very difficult to After the formation of hydrogen peroxide derivatives through
be stopped. Therefore, preventing chain reaction initiation is the three processes: automatic oxidation, photosensitized oxidation
most effective way to control autoxidation. and enzymatic oxidation. As shown in Figure 1, the main volatile
flavor compounds, and nonvolatile lipid oxidation products were
produced through three different oxidation mechanisms (Roland
Photosensitized oxidation et al., 2017; Zhang et al., 2020). Hydroperoxide lyase (HPL),
alcohol dehydrogenase (ADH), and peroxygenase (POX) will
The left green route in Figure 1 displays the photosensitized further catalyze the generation of products and aggregation,
oxidation, which differs from automatic oxidation in that during resulting in beany flavor. Hydroperoxide lyase (HPL) reacts with
photosensitized oxidation, highly reactive singlet oxygen directly hydroperoxyl derivative to form volatiles such as C6 aldehydes
attacks the double bonds of unsaturated fatty acids, resulting in and C9 aldehydes, and in the presence of ADH, these aldehydes
oxygen binding directly to the fatty acids in the formation of are reacted to the corresponding alcohols, such as alcohols and
hydrogen peroxide. Singlet oxygen is more significant for acids. And hydroperoxyl derivative forms non-volatile lipid
photosensitized oxidation in the oxidation process than automatic oxidation products such as hydroxy, epoxy, and fatty acids in the
oxidation (Hammond and White, 2011). presence of POX (Wang Y. Q. et al., 2022).
Therefore, in order to control the production of beany flavor,
we have to control the oxidation and degradation of unsaturated
Enzymatic oxidation fatty acids. In the automatic oxidation process, the concentration
of oxygen is extremely important, the higher the concentration of
As illustrated in the purple route in Figure 1, enzymatic oxygen, the faster the rate of automatic oxidation (Wang et al.,
oxidation is the main reason to produce most volatiles in legumes, 2021). On the contrary, in the photosensitized oxidation process,
which is the primary way in which off-flavor is generated. the oxygen concentration has little effect, and the number and
Lipoxygenase (LOX) is a major element of enzymatic oxidation distribution of unsaturated double bonds have a significant effect
(Roland et al., 2017). LOX belongs to the family of nonheme iron- (Wang et al., 2020). And in the enzymatic reaction process, lox is
containing enzymes that can effectively catalyze the deoxygenation its limiting factor (Roland et al., 2017; Wang et al., 2021).
of polyunsaturated fatty acids (PUFAs) such as linoleic acid (18:2)
and linolenic acid (18:3), to produce hydroperoxyl derivative
(Alhendi et al., 2018). Because oxidation reactions usually occur The mechanism underlying the
in specific regions and oxygen is generally bound within C9 and fermentation removal of beany
C13 (Gardner, 1991), Soy LOX isoenzymes are classified into three off-flavor
types: 9-, 13- and 9-/13-LOXs. For example, soy LOX-1 is a
13-LOX, LOX-2 is a 9-/13-LOX, and soy LOX-3 is a 9-LOX or There are currently two major types of methods to remove
9-/13-LOX (Wang Y. Q. et al., 2022). beany flavor from plants. The first way is to use enzymes to

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Tao et al. 10.3389/fmicb.2022.1070773

decompose the components that produce the beany flavor in Among them, the precursors of the flavor of legumes are mainly
plants and use enzymes to decompose the precursor substances phospholipids (PLs) and free fatty acids (FFAs).
of the plants that produce the beany flavor (Zhu and Phospholipase can be used to remove such precursors, among
Damodaran, 2018; Fischer et al., 2022). The second way is to which the combined phospholipase A2 (PLA2) and cyclodextrin
generate new aromatic profile which can mask the original mixed with soybean meal in a water bath and it is found that the
beany flavor. removal rate of phospholipids is more effective (Zhu and
Damodaran, 2018).
In previous studies, it was found that the principle of
Direct decomposition of compounds removing beany flavor was mainly divided into two steps:
with beany off-flavor hydrolyzing PLA 2 to decompose PL, and then removing the
hydrolyzed product by forming an inclusion complex through
It was found that ADH and aldehyde dehydrogenase (ALDH) β-cyclodextrin (β-CD). PLA2 selectively cleaves the ester bond of
had the most obvious effect on the removal of beany flavor. the acyl chain at the sn-2 position of PL and generates Lyso-PL
Because the selection of compounds involved in the beany flavor and FFA. The product is a non-polar material. β-CD is a cyclic
included nine aldehydes, one furan, four alcohols, four ketones, non-reducing oligosaccharide. Its special construction makes its
three sulfides, and five pyrazines, the compound with the greatest inner cavity hydrophobic. β-CD and other cyclodextrins can form
effect on “beany” are aldehydes (Xu et al., 2020). ADH converts water-soluble inclusion complexes with insoluble non-polar
aldehydes to primary alcohols and ketones to secondary alcohols. substances. The water-soluble inclusion compound can
ALDH converts aldehydes to carboxylic acids. Reduces beany be dissolved into the supernatant by a polar solvent such as
flavor by converting odorous compounds (aldehydes and ketones) deionized water, so as to achieve the effect of separation from
to less odorous products (alcohols and carboxylic acids; Shi soybean flour.
et al., 2021). Additional studies have shown that after the application of
As illustrated in the upper right of Figure 1, there are five main Alcalase, papain or a combination of enzymes, the acidic subunits
ways they remove the beany flavor: (a) The blue route illustrates of β-conglycinin and glycinin disappear completely, resulting in
the homofermentation: For the catabolism that occurs in this type the removal of some of the precursors of the beany flavor, thereby
of fermentation, ADH does not participate in basal metabolism, reducing the beany flavor (Meinlschmidt et al., 2016).
but plays a role in the mixed acid pathway. Such as Lactobacillus
acidophilus and Streptococcus thermophiles (Shi et al., 2021). (b)
Obligatory heterofermentative Lactic acid bacteria illustrated with Odor masking and transformation
the red route (LAB): such as L. fermentum. ADH is present in
metabolism as a constitutive enzyme as well as an aldehyde/ As shown in the low right of Figure 1, some strains can
alcohol dehydrogenase (Ortakci et al., 2015; Verce et al., 2020). (c) produce new aldehydes with aroma after fermentation while
Facultative heterofermentative LAB: Only the heterofermentative reducing the content of compounds with beany smell. In this way,
pathway is inducible. Such as L. plantarum. Its basic metabolism the new aroma can mask the original beany smell to further
is the homofermentative pathway (Prückler et al., 2015). (d) enhance the total aroma profile of the product. For example,
Alcoholic fermentation illustrated with the cyan route: ADH and phenylacetaldehyde produced by fermentation of L. rhamose L08
aldehyde dehydrogenase are present in the catabolism of the strain can bring floral and honey-like aromas (Pei et al., 2022). And the
as constitutive enzymes. For example Saccharomyces cerevisiae main contributors to the aroma profile of A. aegerita fermented
(Mizuno et al., 2006). (e) Oxidative metabolism. “Oxidative soy beverages remain soy-derived compounds. At the same time,
fermentation” is the incomplete oxidation of substrates, carried by ethyl esters and lactones were produced during fermentation,
the dehydrogenases of the respiratory chain, resulting in the which resulted in increased fruity, floral, and creamy/dairy aromas
extracellular accumulation of oxidized products. The “oxidative present in cheese aromas and altered the overall aroma of the
system” is located in the cytoplasmic membrane and is connected samples (Nedele et al., 2022a). Other scents include milky, nutty,
to the respiratory chain, for example, Gluconobacter suboxydans and more.
(Deppenmeier and Ehrenreich, 2009; Lynch et al., 2019).

Application of microorganisms to
Breaking down beany flavor precursors remove beany flavor
in plants
In recent years, fermentation has become the primary method
Previous studies have indicated that the beany flavor in employed in many studies to reduce the beany flavor due to its
legumes is primarily the result of interactions between lipid various advantages. Several strains are discussed below that have
oxidation products, proteins, and phytochemicals. Lipid-derived shown potential to remove or cover the beany flavor in bean
off-flavors are believed to be the main cause of beany flavor. protein-based products (Table 1).

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Tao et al.
TABLE 1 Application of fermentation methods for removing beany flavor.

Fermentation
Matrices Strains Method Function Sensory perception References
conditions
Lactobacillus Mung bean LAB (L. plantarum) 37 ± 1°C Biotransformation Transform aldehydes into esters. The esters give the fermented Yi et al. (2021)
48 h products a pleasant fruity odor.
Lupinus angustifolius L. Five lactic acid bacteria 28°C Decomposition and Reduce aldehydes, especially hexanal, Increase sourness and “vinegar” Laaksonen et al. (2021)
48 h masking which possesses “green” odor; Create odor; Reduce the “beany” flavor
new pleasant aromatic compounds. as well as the unpleasant off
flavor.
Pea flour (Pisum L. rhamnosus L08 37°C Decomposition and Increase the variety of acids and esters; Reduce the unpleasant beany Pei et al. (2022)
sativum L.) 2 days masking Reduce the unpleasant flavor compounds flavor; Produce floral and
such as nonanal, decanal, octanal, honey-like aromas.
1-hexanol and 2-ethyl-1-hexanol;
Produce phenylethyl aldehyde that could
bring pleasant aromas.
Pea protein isolates L. plantarum, L. casei and 37°C Decomposition Remove around 42% aldehyde and 64% Decrease the overall unpleasant Shi (2020)
mixed strains of probiotics 5, 10, 15, 20, 25, and 30 h ketone content; Produce a small amount aroma and flavor intensity.
Anaerobic conditions of alcohol.
05

Lupin protein extracts L. plantarum L1047 and − Decomposition and Decrease the concentration of n-hexanal The more pleasant odor of the Schindler et al. (2011)
(LPE) Pediococcus pentosaceus P113 masking and prevent its re-formation; Change the fermented protein extracts,
aroma profile which may mask off- compared to the unfermented
flavors. protein extracts is explained by
its different aroma profile.
Soymilk Lactobacilli and Streptococci 37°C Decomposition Reduce or even eliminate the The heat treatment applied to Blagden and Gilliland
12 h concentrations of volatile components the soymilk in the present (2005)
that have been associated with the beany study would certainly cause a
flavor of soymilk, such as methanol, severe cooked flavor. Thus, the
acetaldehyde, ethanol, and hexanal. resulting fermented product
would not have been suitable
for sensory analysis.

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Pea protein isolates L. plantarum 37°C Decomposition Eliminate aroma compounds that belong Reduce the off-flavor (“hay” Shi et al. (2021)
(PPI) 30 h to the aldehyde, ketone, and alcohol and “green” like aroma);
Anaerobic conditions groups. Improve the aroma profile.
Pea (Pisum sativum) L. plantarum L1047 or 37°C Decomposition and Decrease the n-hexanal content; Reduce Improve the aroma profile. Schindler et al. (2012)
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protein extract P. pentosaceus P113 48 h masking or mask undesirable green notes.

(Continued)
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Tao et al.
TABLE 1 (Continued)
Fermentation
Matrices Strains Method Function Sensory perception References
conditions

Lupin protein isolate Lactobacilli 30°C (L. parabuchneri, Decomposition and Decrease the n-hexanal content; Reduce Reduce the intensity of Schlegel et al. (2019)
L. brevis) masking or mask undesirable green notes. characteristic aroma impression
37°C (L. helveticus, (pea-like, green bell pepper-
L. delbrueckii, L. sakei sub sp. like) from 4.5 to 1.0.
carnosus, L. reuteri, S. xylosus)
42°C (L. amylolyticus)
36–48 h
Yeast Okara Yeast (Lindnera saturnus) 30°C Biotransformation Aldehydes are reduced into alcohols by Transform the aroma profile of Vong and Liu (2018)
48 h yeast alcohol dehydrogenase or oxidized okara from a green, grassy
Solid-state fermentation into acids by yeast aldehyde off-flavor into a markedly fruity
dehydrogenase, and finally form esters and sweet aroma.
via enzymatic reactions.
Soybean residue (okara) Four “dairy yeasts” (Geotrichum 30°C Biotransformation Oxidize the undesirable aldehydes into The final fermented okara had a Vong and Liu (2017)
candidum, Yarrowia lipolytica, 72 h fatty acids, or reduce them into alcohols, very strong fruity and estery
06

Debaryomyces hansenii and Solid-state fermentation and finally yield esters; Yeast proteinases character.
Kluyveromyces lactis) and six and peptidases break down the protein
“wine yeasts” (Saccharomyces in okara. Yeasts degrade the free amino
cerevisiae, Lachancea acids to yield higher alcohols and esters.
thermotolerans, Metschnikowia
pulcherrima, Pichia kluyveri,
Torulaspora delbrueckii, and
Williopsis saturnus)
Soy (tofu) whey commercial non-Saccharomyces 20°C Decomposition & Metabolize endogenous carbonyls and Each yeast produced different Chua et al. (2018)
yeasts (T. delbrueckii; 3 days Masking alcohols to low or trace levels; Produce levels of different volatile
L. thermotolerans; new alcohols, esters and acids to enrich compounds that can contribute
M. pulcherrima; P. kluyveri and aroma profiles that were unique to each to the different aroma profiles

10.3389/fmicb.2022.1070773
W. saturnus) non-Saccharomyces yeast. of the fermented whey.
Edible Soybean products (soy Lycoperdon pyriforme 24°C Biotransformation Saturated aldehydes were metabolized by Impart slightly bitter almond- Nedele et al. (2022b)
Basidiomycetes drink and soy protein 28 h the fungus to their corresponding like, fungal and nutty odor
isolate) In darkness alcohols; Di-unsaturated aldehydes were notes without recognition of
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synthesized to a non-volatile product. the soy product off-flavors.

(Continued)
 TABLE 1 (Continued)
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Tao et al.
Fermentation
Matrices Strains Method Function Sensory perception References
conditions

Okara Wolfiporia cocos CGMCC 5.55, 25 ± 1°C Decomposition & Decrease the content of off-flavor All fermented products had Wang Z. et al. (2022)
W. cocos CGMCC 5.528, 7 days Masking compounds like hexanal; New aromatic very little characteristics of
W. cocos CGMCC 5.78 and compounds were generated. beany flavor, and a fragrant,
Tremella fuciformis CGMCC floral, and sweet aroma was
5.466 present.
Soy drink Agrocybe aegerita 24°C Decomposition and Many typical soybean off-flavor Produce a natural and vegan Nedele et al. (2022a)
75 h masking contributors were reduced in their cheese aroma;
In darkness intensity such as green aldehydes; Decrease the off-flavor in
Synthesize ethyl esters and lactone, soybean-based products;
which change the overall aroma of the Produce a sweet, floral and
sample. fruity flavor impression.
Soy drink L. pyriforme 24°C Decomposition & Increase aroma compounds such as The aroma of soy drink turned Nedele et al. (2021)
28 h Masking 1-octen-3-one with a mushroom-like from green, beany, and oat-like
In darkness odor and benzyl alcohol with a sweetish to oat-like, mushroom-like, and
flavor; Decrease most of the key aroma almond-like.
compounds with a green off-flavor.
07

Co-fermentation Soybean (soymilk) Three isolated new yeasts 36°C Biotransformation Transform aldehydes into either acids, Remove the beany flavor; Korma et al. (2022)
(K. marxianus SP-1, Candida 5h alcohols and esters. Produce rich aromatic
ethanolica ATW-1, and LAB and yeast ratio components; Improved the
P. amenthionina Y) and a (5:2, v/v) flavor and taste of drinks.
commercial yeast
(K. marxianus K) along with
five strains of lactic acid
bacteria (LAB)
Okara (soybean residue) Probiotic (L. paracasei) and 30°C Biotransformation Produce large amounts of esters to give a Give a natural fruity aroma. Vong and Liu (2019)
yeast (L. saturnus) 48 h natural fruity aroma.
(viable cell count ratio of
probiotic:yeast was about
100:1)

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Pea protein-based Lactic acid bacteria and yeasts pH 7.1 ± 0.1 to 4.55 Decomposition & Degrade many off-flavor compounds; Reduce the concentration of El Youssef et al. (2020)
product 30°C to 4°C Masking Trigger the generation of esters pea off-notes; Generate new
compounds with fruity and floral notes. notes that could modify the
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perception of sensory defects;

Improve the aroma quality of
fermented beverages.

(Continued)
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Lactobacillus

Lactic fermentation might be a promising strategy to improve

Shi et al. (2020)

References
the aroma profile of plant-based food products as it results in a
reduction or covering of undesirable flavors. Some Lactobacillus
strains can reduce or eliminate aroma compounds that have been
linked with the beany flavor which are members of the aldehyde,
and taste. Sensory acceptability
Give the okara a pleasant smell

is greatly improved compared

ketone, and alcohol groups. Several authors demonstrated how
Sensory perception

fermentation with lactic acid bacteria in pea, lupin protein extract,

to unfermented okara.

and soymilk, respectively, reduced the concentration of hexanal,

which is primarily responsible for the greeny and beany off-flavor
(Blagden and Gilliland, 2005; Schindler et al., 2011, 2012; Shi,
2020). Other researches have shown that lactic fermentation can
produce higher aldehydes, alcohols, acids, and ester compounds
through further biotransformation to cover the beany flavors
Convert aldehydes into alcohols and

(Schindler et al., 2011; Shi et al., 2021; Yi et al., 2021). Lipid

esters, among other compounds to

degradation, which also contributes to the creation of the odors

improve the flavor of okara.

found in fermented foods, takes place concurrently with this

biotransformation. For instance, L. rhamnosus L08 fermentation
produced phenylethyl aldehyde that could bring floral and honey-
like aromas, phenylethyl alcohol that exhibits a fresh bread-like,
Function

rose-like aroma, and several esters with floral and fruity

fragrances, which had the effect of covering undesirable flavors
(Pei et al., 2022).
Biotransformation

Yeast
Method

Some yeasts are also used to modify the odor characteristics

of plant-based products by biotransformation. The products of
yeast fermentation often carry a pleasant flavor profile of alcohols
Solid-state fermentation
Anaerobic conditions

and esters. Chua et al. fermented soy whey samples using five
Fermentation

commercial non-Saccharomyces yeasts. Volatile compounds such

37°C for 2 days
conditions

28°C for 1 day

as ethanol and 2-phenylethanol were found in the fermented

products, giving them a rose-like aromatic character (Chua et al.,
2018). Other studies used yeasts to ferment okara and obtained a
very strong fruity and estery character since most of the
and B. longum, S. cerevisiae and
Hansenula sp.) and mixed yeast
(L. plantarum, L. acidophiluss,
Bifidobacterium lactis, L. casei

undesirable aldehydes were reduced into alcohols and esters by the

(S. cerevisiae and Hansenula

yeasts fermentation (Vong and Liu, 2017; Vong and Liu, 2018).
Compound probiotics

Unfortunately, although fermentation with lactic acid bacteria

or yeasts reduces the beany flavor substances, the fermentation
process of certain strains inevitably produces acids such as lactic
Strains

acid and hexanoic acid, causing strong sour and wine flavors,
sp.)

which to some extent aggravate the undesirable flavor of the bean

protein-based product (Laaksonen et al., 2021; Nedele et al., 2021;
Wang Z. et al., 2022). Thus, the optimization of fermentation
conditions is required to prevent the creation of unpleasant
Matrices

flavor compounds.
Okara
TABLE 1 (Continued)

Edible basidiomycetes

In the food processing industry, edible fungi are common

fermentation strains. Among them, basidiomycetes have garnered

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Tao et al. 10.3389/fmicb.2022.1070773

increasing interest due to their abundance, diversity, accessibility, high acidity, which may not be preferred by consumers. More
and nutritional benefits (Mahboubi et al., 2017; Sun et al., 2020; research is needed to select suitable strains and optimize
Bentil, 2021). While most conventional starter cultures mainly fermentation process to meet the preferences of consumers
produce primary metabolites, basidiomycetes are noted for their (Laaksonen et al., 2021). (b) Aside from legume proteins, other
sensory modification of various food products (Nedele et al., 2021; ingredients including microalgae, konjac, and edible
Rigling et al., 2021; Nedele et al., 2022a). They are able to modify mushrooms could be promising substitutes for meat analogs
the flavor because their highly sophisticated secretomes produce due to their superior production capabilities and high
abundant natural flavor molecules (Bouws et al., 2008). The nutritional content. Scientists can process these ingredients
scientists used four types of edible fungi to reduce the beany flavor using fermentation method to explore the next generation of
and gain new aromatic flavors of okara. These new flavors contain meat analogs (Jiménez-Colmenero et al., 2012; Fu et al., 2021;
benzene, ethanol, and linalool, which are probably byproducts of Yuan et al., 2022). (c) The fermentation process produces amino
enzymatic events occurring during the growth and metabolism of acids, sugars, and a series of precursors for Maillard reactions.
edible fungi. In addition to adding floral, sweet, and orange Therefore, if the fermentation process is properly designed and
fragrances, the presence of these compounds can disguise the oriented, it will effectively promote the Maillard reaction,
flavor characteristics of some unwanted components (Wang providing better flavor and taste to plant-based meat analogs
Z. et al., 2022). Apart from that, recent studies have confirmed that and eliminating the beany flavor. (d) The metabolomic pathway
by using a unique fermentation procedure with Lycoperdon of fermentation process should be further investigated to
pyriforme, the beany flavor of soy beverages was diminished while demonstrate how the beany flavor is decomposed with the aid
the nutritional profile was maintained. During this fermentation of mass spectrometry, nuclear magnetic resonance or isotopic
process, aldehydes, the main off-flavor contributors, were labeling experiments (Ran et al., 2022). And the influence of
decreased and some pleasant aroma components were created, more volatile compounds exist at low amounts should be further
imparting the finished product an almond- and nutty-like smell studied to confirm their effect on the aroma change of the
(Nedele et al., 2021, 2022b). Therefore, it can be expected that the fermented product. Improvements of current fermentation
edible basidiomycetes are likely to be a promising strain for flavor techniques will benefit from the results of such research. (e)
improvement in plant-based foods. Further research is required There is still a lack of comprehensive research on the application
to ascertain which kind of strains can successfully grow in the of fermentation to remove beany flavor from plant-based meat
various sources of legume dietary fiber systems as well as what analog. Not only the technological aspects, but also the safety
fermentation conditions can facilitate the aroma profile concerns should be taken into consideration. Possible safety
improvement. problems such as microbiological stability, allergenic potential
and heavy metal and toxic substance contamination during
fermentation should be investigated to develop safe and efficient
Conclusion and outlook fermentation protocols for the production of plant-based
meat analogs.
In summary, oxidation of unsaturated fatty acids and
hydrolysis of lipids are the primary causes of undesirable flavor in
a plant protein-base food. At present, masking, biotransformation Author contributions
and enzymatic degradation are the main mechanisms and
approaches for removing the beany flavor. Several species of XY and TZ contributed to conception and design of the study.
microorganisms including lactobacillus, yeast, and edible AT, HZ, and JD wrote the first draft of the manuscript. YX, YL, JL,
basidiomycetes were demonstrated to reduce the level of beany and JH helped review and revise the manuscript. All authors
flavor compounds by converting them into less impacting contributed to the article and approved the submitted version.
compounds or covering them with new pleasant compounds
formed during fermentation. However, different fermentation
approaches will produce different final flavor characteristics due Funding
to the different metabolic pathways and capacities of the strains.
Thus, the strains utilized have a significant impact on how the This work was supported by the Science and Technology
fermentation affects the fragrance profile of the plant proteins. It Development Funds, Macau SAR (0024/2022/A and 0004/2021/
should be noted that no technique is perfect, and each method has ITP), the Science and Technology Planning Project of Guangdong
its own inherent advantages and drawbacks. Province (2020B1212030008), Innovation Cultivation Project of
In order to further improve the flavor quality of plant-based Zhuhai College of Science and Technology (2019XJCQ006), and
meat analogs, future product development and application the Open Fund of Guangdong-Hong Kong-Macao Joint
research can mainly focus on the following aspects: (a) Laboratory for Contaminants Exposure and Health
Fermentation using some strains may produce products with (GHMJLCEH-05).

Frontiers in Microbiology 09 frontiersin.org

Tao et al. 10.3389/fmicb.2022.1070773

Conflict of interest Publisher’s note

JL and JH were employed by Macau Uni-Win Biotechnology All claims expressed in this article are solely those of the
Co., Ltd. authors and do not necessarily represent those of their affiliated
The remaining authors declare that the research was organizations, or those of the publisher, the editors and the
conducted in the absence of any commercial or financial reviewers. Any product that may be evaluated in this article, or
relationships that could be construed as a potential conflict claim that may be made by its manufacturer, is not guaranteed or
of interest. endorsed by the publisher.

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