Trends in Food Science & Technology 114 (2021) 25–59

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Trends in Food Science & Technology

journal homepage: www.elsevier.com/locate/tifs

Understanding the potential of fruits, flowers, and ethnic beverages as

valuable sources of techno-functional and probiotics strains: Current
scenario and main challenges
Tatiana Colombo Pimentel a, Louise Iara Gomes de Oliveira b,
Elvira de Lourdes Chaves Macedo b, Giselle Nobre Costa b, Disney Ribeiro Dias c,
Rosane Freitas Schwan c, Marciane Magnani b, *
a
Federal Institute of Paraná, Campus Paranavaí, Paranavaí, Brazil
b
Department of Food Engineering, University of Paraíba, João Pessoa, PB, Brazil
c
Department of Food Science, Federal University of Lavras, Lavras, Minas Gerais, Brazil

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

Keywords: Background: Fruit, vegetables, flowers, and ethnic beverages may be a source of microbial species with techno-
Probiotic culture functional and health-promoting properties.
Tchapalo Scope and approach: This review explored the added-value microorganisms isolated from unconventional sources
Chicha
and their techno-functional and probiotic properties. Fruits (strawberry, guava, apple, peach, grape, and
Caxiri
Kombuha
papaya), vegetables (peppers, corn, zucchini, lettuce, cucumber, coffee beans, and olives), flowers (narcissus,
Cauim pink rose, red rose, yellow rose, and sunflower), and ethnic fermented beverages (tchapalo, tarubá, cauim,
chicha, caxiri, kombucha, and water kefir) are source of lactic acid bacteria (Lactobacillus and amended genera,
Leuconostoc, Enterococcus, Pediococcus, Fructobacillus, and Weissella) and yeasts (Saccharomyces, Candida, Pichia,
Rhodotorula, Torulaspora, Cryptococcus, Hansenula, and Debaromyces).
Key findings and conclusions: Strains isolated from unconventional sources showed antimicrobial capacity, pro­
duction of bioactive metabolites, and technological properties, suggesting their utilization as biopreservatives in
food products or against phytopathogens, and for improving the nutritional value and sensory characteristics of
food products. Their utilization as starter cultures in fermented foods may decrease the fermentation time and
improve the products’ characteristics. Some strains showed probiotic potential, presenting important adhesion
and auto and co-aggregation properties, cell surface hydrophobicity, safety, and resistance to the gastrointestinal
tract. These probiotic cultures showed anti-hypertensive, antilipidemic, immunomodulatory, and anti-diabetic
properties in in vitro assessments or animal models. However, clinical studies are necessary to demonstrate
the health effects in humans. In conclusion, cultures isolated from unconventional sources have a high potential
for use in processing and functionalization of foods and can be alternative tools for developing vegan probiotic
products.

1. Introduction et al., 2017).

The microbial composition of fruits, raw vegetables, and naturally
More attention has been devoted to fruit, raw vegetables, and flowers fermented beverages depends on the intrinsic (carbohydrate and protein
as source of microbial species with techno-functional and health- contents and pH values) and extrinsic (climatic conditions and har­
promoting properties in the last decade. Besides, naturally fermented vesting) parameters (Fessard & Remize, 2019; Garcia et al., 2016).
foods have been explored as niches for the isolation of bacteria and yeast Furthermore, it may be associated with pollinators that visit flowers and
strains with potential for biotechnological application and/or func­ fruits, birds fed with them, or insects (Rodríguez et al., 2019). Some of
tionalization of foods (Menezes et al., 2020; Pei et al., 2020; Semjonovs these strains can antagonize other microorganisms and show

* Corresponding author.
E-mail address: [email protected] (M. Magnani).

https://doi.org/10.1016/j.tifs.2021.05.024
Received 15 February 2021; Received in revised form 12 April 2021; Accepted 15 May 2021
Available online 27 May 2021
0924-2244/© 2021 Elsevier Ltd. All rights reserved.
T.C. Pimentel et al. Trends in Food Science & Technology 114 (2021) 25–59

biotechnological applications (Costa et al., 2018; Fessard et al., 2017; (Grijalva-Vallejos et al., 2020; Ramos et al., 2015). Several strains of
Linares-Morales et al., 2020), while others can present important health yeasts and LAB with techno-functional properties have been isolated
effects associated with their probiotic properties, such as microbiota from ethnic beverages over the last years, such as Limosilactobacillus
modulation (Costa et al., 2019), hypocholesterolemic, fermentum S6 (former Lactobacillus fermentum S6) and Pediococcus acid­
anti-hypertensive, and hypoglicemic properties (Verón et al., 2019), and ilactici S7 from tchapalo (Aka et al., 2020), S. cerevisiae EYS5 and Tor­
immunomodulatory effects (Fakruddin et al., 2017). ulaspora delbrueckii EGT1 from chicha (Grijalva-Vallejos et al., 2020),
In general, the microbial population of fruits and raw vegetables is L. plantarum SLG10, P. occidentalis, Candida sorboxylosa, Hanseniaspora
around 105 - 107 cfu/g. Yeasts are the dominant group (102–106 CFU/g), opuntiae, Komagataeibacter rhaeticus P 1463, and K. hansenii B22 from
and Saccharomyces, Candida, Pichia, Rhodotorula, Torulaspora, Crypto­ kombucha (Pei et al., 2020; Semjonovs et al., 2017; Taheur et al., 2020),
coccus, Hansenula, and Debaromyces are the most commonly found L. plantarum CCMA0743 and T. delbrueckii CCMA0235 from cauim
genera (Barros et al., 2019; Di Cagno et al., 2013; Leff & Fierer, 2013; (Freire et al., 2017), and L. plantarum, Levilactobacillus brevis (former
Martins et al., 2019; Riesute et al., 2021). Otherwise, Lactic Acid Bac­ Lactobacillus brevis), L. mesenteroides, Bacillus subtilis, and T. delbrueckii
teria (LAB) comprises between 102 - 104 cfu/g (Di Cagno et al., 2013; from tarubá (Ramos et al., 2015).
Leff & Fierer, 2013), and the most found genera are Lactobacillus, Leu­ Previous reviews have mainly explored the isolation of autochtho­
conostoc, Enterococcus, Pediococcus, Fructobacillus, and Weissella (Fessard nous microorganisms from vegetables, fruits, or fermented foods and
et al., 2017; Rodríguez et al., 2019). beverages and their use as starters, without evidencing the production of
Yeasts have stood out for their favorable performance in biotech­ metabolites, application of these cultures as biopreservatives or in the
nological applications, such as fermentation and metabolite production inhibition of phytopathogens (Pessôa et al., 2019; Torres et al., 2020;
(Karim et al., 2020; Semjonovs et al., 2017; Taheur et al., 2020). In Pérez-Armendáriz & Cardoso-Ugarte, 2020). Furthermore, the in vitro
addition, recent studies have demonstrated the probiotic potential of and in vivo probiotic effects of these isolated strains have not been
several yeast strains isolated from fruits and vegetables (Hsiung et al., extensively reported. None of the reviews discussed the flower micro­
2020; Menezes et al., 2020). At the same time, LAB can have probiotic biota as a source of added-value species. Therefore, this review aimed to
properties and can produce organic acids (mainly lactic acid) and other explore the added-value microorganisms isolated from unconventional
compounds contributing to increase the shelf life of food products and sources (fruit, raw vegetables, ethnic fermented beverages, and flowers)
improving their physicochemical and sensory characteristics (Costa and their associated techno-functional and potentially probiotic prop­
et al., 2018; Di Cagno et al., 2016; Oliveira, Araújo, et al., 2020). Various erties. The main challenges in characterizing new probiotic strains and
LAB strains have been isolated from fruit and vegetables, such as Lac­ functionalizing foods with these new strains are discussed. A focused
tiplantibacillus plantarum CM-3 (former Lactobacillus plantarum CM-3) discussion of the main reported effects of these strains as biotechno­
from strawberry (Fragaria × ananassa Duch. Cv. “Hongyan”) (Chen logical tools and the known in vitro/in vivo health-promoting effects is
et al., 2020), Leuconostoc mesenteroides, Enterococcus mundtii and provided. Furthermore, the antagonistic effects, the active microbial
Enterococcus faecium from fruits (guava, green apple, and orange) and metabolites, and the effects on products’ technological and sensory as­
vegetables (corn and chilaca and jalapeno peppers) (Linares-Morales pects are presented. Finally, the field’s future perspectives close this
et al., 2020), Weissella paramesenteroides FX1, FX2, FX5, FX9, FX12 from review, making available information to help researchers in further
sapota, cherry, banana, orange, and plum smashed fruits (Pabari et al., studies.
2020), L. plantarum 49, Lacticaseibacillus paracasei 108 (former Lacto­
bacillus. paracasei 108) and L. plantarum 201 from fruits (mango and 2. Probiotic potential of strains isolated from unconventional
acerola) and fruit by-products (mango, acerola, and soursop) (Garcia sources
et al., 2016), L. mesenteroides (OP4, OP9, OP21, OP18 and OP23) from
fresh fruits of Opuntia ficus-indica (L.) Mill. (Sanguigna genotype) (Di 2.1. Selection of probiotic cultures from unconventional sources
Cagno et al., 2016), and L. plantarum LPBR01 from coffee fruits (Pereira
et al., 2016). Probiotics are viable microorganisms that confer health effects to the
Flowers have been attracting attention due to the presence of bac­ individuals when consumed in adequate amounts (Hill et al., 2014). The
teria of the genus Fructobacillus that comprise species with fructophilic strains mostly used as probiotics are LAB, represented by species of
characteristics, i.e., preference for D-fructose over D-glucose, producing Lactobacillus (L. acidophilus), Lactiplantibacillus (L. plantarum), Ligilacto­
mannitol and acetate instead of ethanol (Behare et al., 2020; Maeno bacillus (L. salivarius), Lacticaseibacillus (L. rhamnosus and L. casei), and
et al., 2019; Patil et al., 2020; Sakandar et al., 2019). Mannitol is a Limosilactobacillus (L. reuteri and L. fermentum). Furthermore, strains of
low-calorie polyol widely used in the food industry, suggesting that the Bifidobacterium genus (B. longum, B. animalis, B. bifidum, B. lactis,
these strains can serve as tools for biotechnological application (Behare B. breve, and B. infantis) and yeasts (Saccharomyces cerevisiae) are
et al., 2020; Maeno et al., 2019). Recent studies have isolated fructo­ commonly used (Pimentel et al., 2021).
philic strains from various flowers and fruits such as F. fructosus MCC Dairy products are the main probiotic products available in the
3996 from nectar of Butea monosperma flower (Patil et al., 2020), and market. However, there is a demand for the development of non-dairy
F. pseudoficulneus JNGBKS, JNGBKS3, F. fructosus JNGBKS2, JNGBKS4, probiotic products based on cereals, fruits, and vegetables because of
and F. durionis JNGBKS5 from fresh fruits (apple, banana, Chinese lifestyle choices (e.g., vegetarianism and veganism) and the increased
peach, plum, cantaloupe, kiwi fruit, and lychee) and flowers (narcissus, rates of milk allergy and lactose intolerance in the population (Vera-­
pink rose, red rose, yellow rose, and sunflower) (Sakandar et al., 2019). Pingitore et al., 2016). However, most commercial probiotic strains are
Traditional fermented foods obtained from spontaneous fermenta­ derived from human gastrointestinal tract or dairy products (Pimentel
tion, are exploited as rich microbial niches, including fermented cereal- et al., 2021). Probiotic cultures used in plant-based fermented products
based beverages such as rice, corn, wheat, rye, cassava, or sorghum. should preferentially be isolated from plant materials because
Among these beverages stand out tchapalo, tarubá, cauim, chicha, caxiri, plant-derived substrates present significant technological and physio­
and fermented drinks based on teas and fruit juices such as kombucha logical challenges to which strains of the intestinal origin or dairy
and water kefir, which are produced by back-slopping from a consortium products might not be adapted (Gupta & Abu-Ghannam, 2012). Fig. 1
of bacteria and yeasts (Aka et al., 2020; Freire et al., 2017; Grijalva-­ presents the main probiotic properties observed for microorganisms
Vallejos et al., 2020; Pei et al., 2020; Ramos et al., 2015; Taheur et al., isolated from fruit, vegetables, and ethnic fermented beverages.
2020). In naturally fermented beverages, the main microbial groups Table 1 presents the main studies with potentially probiotic strains
comprise LAB, yeasts, and acetic acid bacteria, simultaneously found or isolated from unconventional sources and their main functional fea­
as predominant microbial groups during the fermentative process tures. The selection of probiotic strains considers several aspects,

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Fig. 1. Main genera of yeast and bacterial isolates and assays for probiotic potential.

including safety for the consumer (antibiotic susceptibility, hemolytic (Pabari et al., 2020), Saccharomyces cerevisiae IFST062013 isolated from
activity, and mucin degradation), and physiological functionalities (acid fruit (unspecified) (Fakruddin et al., 2017), L. fermentum 296 isolated
and bile salt tolerance, bile salt deconjugation, cell surface hydropho­ from fruit by-products (Albuquerque et al., 2018), W. koreensis FK121
bicity, auto-aggregation, co-aggregation with pathogens, antagonistic and Lactobacillus crispatus G19 isolated from south Indian fermented
activity against pathogens, and capability of surviving during exposure koozh and guerkins (fermented cucumber), L. pentosus 129, L. brevis 59,
to gastrointestinal conditions). Furthermore, the technological aspects and L. fermentum 111 isolated from fruit by-products (Garcia et al.,
(proteolytic and lipolytic activity, tolerance to NaCl, and EPS and 2016), L. buchneri SS50.4, L. plantarum SBR64.7, L. fermentum SS50.10
bioactive compounds production) may also be considered (Albuquerque isolated from silage, artisanal salami, and cocoa beans (Leandro et al.,
et al., 2018; Torres et al., 2020). Thus, the potentially probiotic strain 2020), and F. fructosus MCC 3996 isolated from nectar of flowers
should consider all these aspects. (B. monosperma) (Patil et al., 2020). The results suggest that acid and
The growth temperature is a primary characteristic for selecting bile salts tolerance are, generally, strain-specific and some strains iso­
probiotic strains since they must remain viable in the temperature found lated from fruits and fruits by-products may not withstand the harsh
in the host. Di Cagno et al. (2020) evaluated ten yeast strains belonging condition of the gastrointestinal tract (Albuquerque et al., 2018; Garcia
to different species and previously isolated from plant matrices. Only et al., 2016). Furthermore, it seems that strains isolated from sources
Saccharomyces cerevisiae DDNd10, Pichia kudriavzevii DCNa1, and with high acidity may better tolerate the stomach’s acidic conditions
Wickerhamomyces subpelliculosus DFNb6 were able to propagate at the (Cao et al., 2019).
temperature of 37 ◦ C, which is the common temperature found in the Co-aggregation and auto-aggregation are important probiotic char­
human body. Other previous studies also reported isolates from fruits acteristics, as they allow the entrapment of bacteria in an aggregated
and vegetables with growth at 37 ◦ C, which may be a differential for form, increasing the stability of microbial strains in the gastrointestinal
human applications (Fakruddin et al., 2017; Oliveira et al., 2017). The tract (Oliveira et al., 2017). Studies have described important
results suggest that it is important to evaluate the resistance of probiotic auto-aggregation performances among yeasts (Bonatsou et al., 2018; Di
cultures of vegetal origin to the target host’s temperatures, as the Cagno et al., 2020; Hsiung et al., 2020; Menezes et al., 2020) and bac­
adaptability to the host temperature may improve its survival and terial strains (Fernández-Pacheco et al., 2020; Leandro et al., 2020; Patil
health-related aspects (Gil-Rodríguez et al., 2015). et al., 2020) isolated from fruits, flowers, and traditional fermented
High viability and survival to adverse conditions are required for foods. Albuquerque et al. (2018) reported that LAB isolated from fruit
probiotics since the microbial population, and its metabolism directly by-products (L. plantarum 53, L. fermentum 60 and 296, and L. paracasei
relates to the effects on the environment or the host. Therefore, probiotic 106) showed high auto-aggregation capacity, resulting in a higher bar­
cultures should survive the food environment (Fiocco et al., 2020). rier effect, preventing the colonization by pathogens, and increasing the
Furthermore, the ability to survive in the upper digestive tract and reach competition for sites with pathogens. The authors also reported impor­
the large intestine demands tolerance to acidity and bile. Thus, the tant co-aggregation properties of the probiotics against pathogens (Lis­
microorganism’s survival to simulated gastrointestinal conditions is teria monocytogenes and Escherichia coli). The auto and co-aggregation
predictive of its survival in the gastrointestinal tract (Xu et al., 2020). capacities were stated as strain-specific and dependent on the pathogen
Studies have shown that microorganisms isolated from fruits, vegeta­ and incubation time (Albuquerque et al., 2018; Anandharaj et al., 2015;
bles, and flowers were able to withstand the simulated gastrointestinal Patil et al., 2020).
conditions, such as W. paramesenteroides isolated from citrus fruits Gut colonization by microorganisms is usually preceded by adhesion

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Table 1
Relevant studies approaching potential probiotics, isolation sources and main functional features of strain isolated from fruits, flowers, and traditional fermented
foods/beverages.
Strains Isolation source Performed test Findings References

Pichia kudriavzevii DCNa1 and Different fruits Simulated The cells remain viable in high Di Cagno et al. (2020)
Wickerhamomyces subpelliculosus gastrointestinal tract concentration, as well as after TGI
DFNb6 (GIT) conditions simulated conditions.
Cell surface All strains presented high hydrophobicity
hydrophobicity and faster auto-aggregation, higher
Aggregation properties biofilm formation and antioxidant activity
Biofilm formation at different levels.
Antioxidant activity None of the strains was hemolytic and
Hemolytic activity both presented sensitivity to fluconazole
Antibiotic susceptibility and nystatin.
Thirteen yeast strains Pistachio fruits (Pistacia vera) Simulated GIT Six from thirteen strains had a high Fernández-Pacheco
conditions survival rate to the GIT conditions et al. (2020)
Antioxidant activity Different auto-aggregation and
Aggregation properties hydrophobicity rates.
Cell surface Diutina rugosa 14 followed by Diutina
hydrophobicity rugosa 8 were the best wild yeast as
Biofilm formation potential probiotic and Hanseniaspora
Molecular guilliermondii 6 and Aureobasidium proteae
identification presented notable biocontrol and
antioxidant capabilities.
Lachancea dasiensis JYC2615MN648701, Fermented food and beverages in Tolerance to acid and The strains were acid tolerant and had Hsiung et al. (2020)
Candida metapsilosis JYC2616, Taiwan bile salts good auto-aggregation and cell-surface
Rhodotorula mucilaginosa JYC2617, Cell surface hydrophobicity.
Saccharomyces cerevisiae JYC2618, hydrophobicity The strains grew well under the bile salt
JYC2619, Brettanomyces bruxellensis Aggregation properties and acid conditions.
JYC2620, JYC2621 and Antioxidant activity The antioxidant capacities varied, and
S. boulardii I-3799 Enzymatic activity majority had a good performance
Molecular regarding cell surface hydrophobicity and
identification β-galactosidase activity.
Weissella confusa MD1 and Weissella Fermented batter Tolerance to acid and The strains had high survivability in Lakra et al. (2020)
cibaria MD2 bile salts gastrointestinal conditions, tolerance to
Simulated GIT lysozyme and phenol, adhesion in
conditions intestinal epithelial cells (HT-29).
Tolerance to phenol They also showed high bile salt hydrolase
Hemolytic activity activity towards sodium taurocholate and
Cell surface bile salt mixture.
hydrophobicity The intact cells showed strong antioxidant
Aggregation properties activity and inhibition of linoleic acid
Antioxidant activity peroxidation, and biofilm formation.
Adhesion properties Both strains showed sensitivity towards
Antibiotic susceptibility conventionally used antibiotics, no
hemolytic activity and exhibited strong
auto-aggregation property and co-
aggregation with Listeria monocytogenes.
Lactiplantibacillus plantarum SBR64.7 Forage plants, cocoa beans Tolerance to acid and Eighty-two isolates were screened, Leandro et al. (2020)
and S180.7 (former Lactobacillus fermented bile salts nineteen strains genotyping.
plantarum SBR64.7 and S180.7), Simulated GIT The species L. plantarum was prevalent.
Lentilactobacillus buchneri SS50.4 conditions The strains presented tolerance to acid
(former Lactobacillus buchneri SS50.4), Cell surface and capacity to grow in bile salts,
Limosilactobacillus fermentum SS50.10 hydrophobicity antimicrobial activity, susceptibility to
(former Lactobacillus fermentum Antibiotic susceptibility antibiotics, adhesion capacity and
SS50.10) Aggregation properties negative virulence factors.
Adhesion properties No biogenic amine production
Bile salt deconjugation The strain L. plantarum SBR64.71 was the
Enzymatic activity most promising for probiotic use.
Biogenic amine
production
Hemolytic activity
Molecular
identification
Nineteen LAB strains Theobroma cacao fermented fruit Tolerance to acid and All the tested LAB isolates showed at least Mabeku et al. (2020)
juice bile salts 50% cells survival at pH 2 after incubation
Acid and bacteriocin up to 3 h.
production They were also susceptible to antibiotics
Antibiotic susceptibility amoxicillin (30 μg), erythromycin (15 μg),
Molecular chloramphenicol (30 μg) and imipenem
identification (10 μg).
Weissella paramesenteroides FX1, FX2, Sapota, cherry, banana, orange, Survival at pH 2.5 and Ability to survive acid pH and sodium Pabari et al. (2020)
FX5, FX9, FX12 and plum smashed fruits salt 1.0% taurocholate.
Enzymatic activity Suitable adhesion to mucin, ability to
Hemolytic activity utilize low molecular weight
Adhesion properties galactooligosaccharides (GOS) and
Antimicrobial activity fructooligosaccharides (FOS), organic
(continued on next page)

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Table 1 (continued )
Strains Isolation source Performed test Findings References

Ability to prebiotic acids and SCFAs production.

utilization Biofilm formation, antimicrobial effect,
Short-chain fatty acids and not harbor any virulent traits.
(SCFAs) production The strains FX5 and FX9 were the best
Biogenic amine performance.
production No biogenic amine production
Molecular
identification
Fructobacillus fructosus MCC 3996 Nectar of Butea monosperma flower Tolerance to acid and Survival rate 67–68% at pH 2 and 3 and 52 Patil et al. (2020)
bile salts to 8% from bile, besides 81% at synthetic
Adhesion properties gastric juice.
Hemolytic activity Resistance to antibiotics amoxiclav,
Antibiotic susceptibility carbenicillin, cefotaxime, ciprofloxacin,
colistin, co-trimoxazole, co-trimazine,
gatifloxacin, nitrofurantoin, norfloxacin,
oxacillin and streptomycin.
GRAS status.
Levilactobacillus brevis B13-2 (former Chinese cabbage kimchi Tolerance to acid and L. brevis B13-2 exhibited tolerance against Song et al. (2020)
Lactobacillus brevis B13-2) bile salts artificial gastric acid and bile salts and did
Simulated GIT not produce β-glucuronidase.
conditions High DPPH radical scavenging activity,
Antioxidant activity Heat-
Enzymatic activity
Latilactobacillus sakei AM2, AM8, AM32, Kimchi Tolerance to acid and Two hundred and twenty-five lactic acid Won et al. (2020)
ADM14, ADM19, YRM13, YRM20, bile salts bacteria were isolated, fifteen strains were
RGM12, GP32, GP48, GN57, YG19 Intracellular tolerant to acid, and bile salts.
(former Lactobacillus sakei AM2, AM8, triglyceride level The strains showed excellent intestinal
AM32, ADM14, ADM19, YRM13, Antibiotic susceptibility cell adhesion, most of them did not
YRM20, RGM12, GP32, GP48, GN57, Enzymatic activity produce β-glucuronidase.
YG19), Molecular The susceptibility to ten tested antibiotics
L. plantarum S-2AM31 and Leuconostoc identification was varied between the strains.
pseudomesenteroides S-YRWM27, S- L. sakei ADM14 had the potential to be
YRWM28 used as a probiotic.
Multiple LAB strains Fermented broccoli, cherry, Tolerance to acid and In broccoli and ginger juices, the genus Xu et al. (2020)
ginger, white radish, and white- bile salts Lactobacillus occupied the dominant
fleshed pitaya juice Simulated GIT position (79.0 and 30.3%, respectively); in
conditions cherry and radish juices, Weissella
Adhesion properties occupied the dominant position (78.3 and
Antibiotic susceptibility 83.2%); and in pitaya juice, Streptococcus
Molecular and Lactococcus occupied the dominant
identification positions (52.2 and 37.0%).
The antibiotic resistance profile, adhesion
properties and survival rates at simulated
gastrointestinal transit were distinct
among all the isolates indicating a high
level of diversity.
Sixteen strains: Table olives biofilms Tolerance to acid and Five hundred and fifty-four lactic acid Benítez-Cabello et al.
13 Lactiplantibacillus pentosus (former bile salts bacteria were isolated, sixteen of which (2019)
Lactobacillus pentosus and) 3 Antibiotic susceptibility had predominant genotypes and were
L. plantarum Lpl15 Lactic acid production identified.
Aggregation properties All sixteen strains showed different levels
Simulated GIT of survival to simulated GIT, resistance to
conditions pH and bile, susceptibility to antibiotics,
Presence of bsh genes and ability to auto and co-aggregation.
Hemolytic activity β-hemolysis activity was detected for all
Enzymatic activity L. plantarum strains, but not for any of the
Molecular L. pentosus
identification Most strains do not exhibit α-glucosidase.
L. plantarum De’ang pickled tea Tolerance to acid and Twenty-six LAB strains were isolated. Cao et al. (2019)
(25 different strains), bile salts Eighteen strains showed a higher
Enterococcus casseliflavus S4b Simulated GIT tolerance to simulated GIT
conditions The strains showed adhesive abilities and
Antioxidant activity were auto-aggregative dependent on
Adherence capacity species and even strains.
Antibiotic susceptibility Two L. plantarum strains, ST and STDA10
Molecular exhibited good probiotic properties and
identification showed a good ability of scavenging DPPH
and ABTS+.
All tested LAB strains were resistant to
kanamycin, streptomycin, gentamicin,
and vancomycin and sensitive to
tetracycline and chloramphenicol.
Ten out of the eighteen strains are
resistant to ampicillin, and the remaining
strains are sensitive to ampicillin.
(continued on next page)

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Table 1 (continued )
Strains Isolation source Performed test Findings References

Four out of the eighteen strains showed

resistance to erythromycin.
L. plantarum P24-1, P24-2 P24-3, P24-4, Pulque and aguamiel Simulated GIT All the strains were susceptible to Cervantes-Elizarrarás
P24-, P24-6, P24-8 and Pediococcus conditions amoxicillin, amikacin, chloramphenicol, et al., (2019)
acidilactici A1-2, A2-1 Antibiotic susceptibility gentamicin, levofloxacin, spectinomycin,
Molecular tetracycline, except to vancomycin.
identification
Twenty-one LAB isolates Fresh vegetables (cauliflower, Tolerance to acid and Two hundred and sixty-six LAB were Junnarkar et al., (2019)
gherkins, cluster beans, fenugreek, bile salts isolated, of which twenty-one had a
cow pea, bitter gourd, french Simulated GIT potential probiotic.
beans, tomato, ridged gourd, conditions At different levels, the strains showed
cucumber, and bottle gourd) Antioxidant activity tolerance to bile salt, acidic pH and
Antibiotic susceptibility pancreatin.
They had the ability to survive in artificial
intestinal condition, presented resistance
to antibiotics and showed antioxidant
activity.
Saccharomyces cerevisiae (KU200270, Cucumber jangajji Tolerance to acid and High resistance to artificial gastric and Lee et al. (2019)
KU200280, and KU200284) bile salts bile juices
Adherence capacity High epithelial cell adhesion
Antibiotic susceptibility
Molecular
identification
103 bacterial isolates most of which Panchamirtham, an Indian ethnic Tolerance to acid and The bacterial isolates exhibited pH, bile Maheshwari et al.
were Bacillus sp., Enterobacter sp., fermented fruit mix bile salts salt tolerance, produced biofilm and (2019)
Acinetobacter sp., Pantoea sp., Biofilm formation exopolysaccharide, but exhibited no
Microbacterium sp., Klebsiella sp., Hemolytic activity hemolytic activity.
Sphingomonas sp.,Ornithinibacillus sp., Enzymatic activity The strains E. xiangfangensis M5S2B6,
Myroides sp., Kocuria sp., Alcaligenes Exopolysaccharides B. safensis M5S2B8, E. hormaechei subsp.
sp., and Brevundimonas sp. (EPSs) production oharae M6S1B2, B. velezensis M4S1B1 and
Cytotoxicity P. terrae M7S2B1 displayed essential
Antibiotic susceptibility characteristics of potential probiotics and
Molecular may be used as starter culture to produce
identification quality panchamirtham.
A hundred and sixteen yeasts isolated Brazilian indigenous fermented Simulated GIT Thirty-six strains were tolerant to Menezes et al. (2019)
food, cocoa fermentation conditions gastrointestinal conditions evaluated by
Cell surface tolerance to pH 2.0, bile salts (0.3% w/v),
hydrophobicity and 37 ◦ C.
Aggregation properties Fifteen isolates had a similar or higher
Antioxidant activity percentage of hydrophobicity, auto-
Phytate hydrolysis aggregation, and coaggregation with
E. coli than the reference strain
S. boulardii.
Fructobacillus pseudoficulneus JNGBKS, Fresh fruits (apple, banana, Tolerance to acid and All the strain exhibited some degree of Sakandar et al. (2019)
JNGBKS3, F. fructosus JNGBKS2, Chinese peach, plum, cantaloupe, bile salts auto-aggregation, inhibition of E. coli, S.
JNGBKS4, F. durionis JNGBKS5, kiwi fruit, and lychee) flowers Simulated GIT typhimurium, and S. aureus and presented
Apilactobacillus kunkeei JNGBKS6, (narcissus, pink rose, red rose, conditions resistance to ciprofloxacin, ceftriaxone,
JNGBKS7, JNGBKS8 (former yellow rose, and sunflower) and Aggregation properties gentamicin, novobiocin, vancomycin
Lactobacillus kunkeei JNGBKS6, honey, and rose petals jam Antimicrobial activity antibiotics.
JNGBKS7, JNGBKS8) Cholesterol assimilation L. kunkeei, F. durionis JNGBKS presented
Antibiotic susceptibility cholesterol assimilation.
Molecular
identification
L. plantarum, Enterococcus faecalis, Leaves of papaya, cassava, Growth at different Exhibited strong survival properties, Samedi and Charles
Lactiplantibacillus paraplantarum sugarcane, yam, and taro temperatures tolerance to simulated intestinal juices, (2019)
(former Lactobacillus paraplantarum), Tolerance to acid and susceptible to streptomycin, penicillin,
and Weissella paramesenteroides bile salts chloramphenicol, ampicillin, and
Simulated GIT tetracycline.
conditions
Antibiotic susceptibility
Molecular
identification
L. pentosus CHIG, NAG1 L. fermentum Fermented foods, fruits, and Tolerance to acid and Twenty-eight Lactobacillus strains were Shekh et al. (2020)
PRS1 vegetables bile salts isolated; six strains were selected to
Simulated GIT evaluate.
conditions The strains survive and grow in simulated
Tolerance to NaCl and oro-gastrointestinal tract, showed
phenol adhesion to mucin, biofilm formation, bile
Adhesion properties salt hydrolase and GABA production, also
Gamma amino butyric had β-galactosidase activities.
acid (GABA) production The strains do not degrade mucin, do not
Antibiotic susceptibility produce DNAse, and are non-hemolytic
Biogenic amine activity.
production
Enzymatic activity
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Table 1 (continued )
Strains Isolation source Performed test Findings References

Mucin degradation
Antimicrobial activity
LAB Strains: RG2A, RG3A, RG3B, RG6A, Cardinal and Red Globe grape Tolerance to acid and Eighteen isolates. Taroub et al. (2019)
RG6B, RG6C, RG7A, RG8B, C9C, fruits bile salts Six strains had an identification of species.
C11A, C11C, C11D, Pediococcus Simulated GIT Strains showed probiotic potential and
pentosaceus RG4A, RG7B, RG8A, RG8C conditions P. pentosaceus RG7B was more promising.
and C11C, L. plantarum RG8A Cell surface
hydrophobicity
Molecular
identification
Fifty yeasts strains Pineapple (Ananas comosus (L.) Tolerance to acid and A hundred and fifty yeasts were isolated. Amorin, Piccoli &
Merril) peel, fresh juice, and bile salts Fifty of which were evaluated, thirteen Duarte, (2018)
spontaneous fermentation of Simulated GIT strains were not able to grow at pH 2.0 but
pineapple conditions maintained high populations (>90%).
Aggregation properties Five strains resisted pepsin e four resisted
Hydrophobicity to bile salt.
properties All isolates showed high level of auto-
Antibiotic susceptibility aggregation and five of them, had a
Molecular hydrophobicity property and resistance to
identification ampicillin, chloramphenicol,
erythromycin, G penicillin, streptomycin,
and tetracycline.
Forty-nine yeast strains Kalamata table olive fermentation Simulated GIT Forty-two out of the forty-nine yeast Bonatsou et al. (2018)
conditions strains presented a survival rate higher
Aggregation properties than 50%, and 24 strains showed survival
Hemolytic activity higher than 70% at the end of the
digestions.
The majority strains showed
hydrophobicity higher than 75%, while
the auto-aggregation ability ranged
between 72 and 91%.
None of the strains showed hemolytic
activity.
L. fermentum 56, 60, 250, 263, 139, 141 Different fruit processing Tolerance to acid and The resistance to antibiotics varied among Albuquerque et al.
and 296, L. plantarum 53, byproducts bile salts the strains with potentially transferable (2018)
Lacticaseibacillus paracasei 106 (former Simulated GIT resistance to tetracycline.
Lactobacillus paracasei 106) conditions Had no resistance to ampicillin and
Cell surface chloramphenicol, as well as presented no
hydrophobicity hemolytic and mucinolytic activity or
Hemolytic activity ability to bile deconjugation.
Mucin degradation
Bile salt deconjugation
Antibiotic susceptibility
Liquorilactobacillus mali K8 Water kefir grains Antibiotic resistance L. mali K8 had tolerance to pH 2.5, and Koh et al. (2018)
(former Lactobacillus mali K8) Hemolytic activity resisted bile salts, pepsin and pancreatin,
Adaptation to food No hemolytic activity
matrix Susceptibility to the standard antibiotics
L. plantarum 3701, 3725, 3739, 3735, Mature Cornus officinalis fruits Tolerance to acid and The strains possess desirable probiotic Tang et al. (2018)
25234, 25294, 2519, 3736, 3711, bile salts properties, similar or superior to the strain
Lacticaseibacillus casei 3734 (former Cell surface Lacticaseibacillus rhamnosus GG (former
Lactobacillus casei 3734) hydrophobicity Lactobacillus rhamnosus GG) used as
Antioxidant activity reference.
Antibiotic susceptibility
Molecular
identification
Thirty different strains comprise Horreh, a traditional Iranian Tolerance to acid and A hundred and forty isolates were Vasiee et al. (2018)
Lactobacillus (fermentum, plantarum, fermented food bile salts identified as LAB and thirty of which were
and brevis) Weissella cibaria, Antibacterial activity evaluated.
Enterococcus (faecium and faecalis), Antibiotic susceptibility L. plantarum and L. mesenteroides subsp.
Leuconostoc (citreum and mesenteroides Molecular mesenteroides isolates were able to grow at
subsp. mesenteroides) and Pediococcus identification pH 2.5 and 3.5.
pentosaceus Some of L. fermentum and L. plantarum
isolates showed quite high hydrophobicity
and antimicrobial activity.
All strains were sensitive to ampicillin,
chloramphenicol, cycloheximide,
erythromycin, neomycin, streptomycin,
tetracycline, and rifampicin, but resistant
to vancomycin.
Leuconostoc mesenteroides E14, M67 Silage and honey Simulated GIT The strains E14 and M67 showed Zarour et al. (2018)
conditions resistance to gastrointestinal tract.
Antibiotic susceptibility The antibiogram presented resistance and
Cell surface sensitivity variables to several antibiotics.
hydrophobicity Both strains had a low EPS production and
Exopolysaccharides low rate of hydrophobicity.
(EPS) production
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Table 1 (continued )
Strains Isolation source Performed test Findings References

Saccharomyces cerevisiae IFST062013 Fruit (unspecified) Simulated GIT The strain was tolerant to a wide range of Fakruddin et al. (2017)
conditions temperature and pH, high concentration
Antioxidant activity of bile salt, NaCl, and presents resistance
Production of bioactive to gastric juice, α-amylase, trypsin, and
compounds lysozyme.
Tolerance to stress The strain produced killer toxin, vitamin
Antibiotic susceptibility B12, glutathione, siderophore, enzymes
(amylase, protease, lipase, cellulase),
biofilm, and strong antioxidant activity.
Also, it showed resistance to tetracycline,
ampicillin, gentamicin, penicillin,
polymyxin B and nalidixic acid.
L. plantarum CCMA0743, Torulaspora Cauim (blended of cassava and Tolerance to acid and The beverage in co-culture improved the Freire et al. (2017)
delbrueckii CCMA0235 rice) bile salts products digestibility by reducing starch,
Antioxidant activity for single and co-culture
Lactic acid production The assays containing yeast showed the
Enzymatic activity highest antioxidant activity (around 10%
by DPPH and ABTS methods).
Pediococcus acidilactici Ch-2 Chuli (a fermented apricot Tolerance to acid and The strain Ch-2 was resistance to low pH, Gupta and Sharma
product) bile salts bile salts and simulated gastric and (2017)
Simulated GIT intestinal conditions. Also, susceptible to
conditions selected eleven antibiotics, unable to
Aggregation properties produce gelatinase and DNase and non-
Metabolic profiling hemolytic nature.
The strain produced squalene – a rare and
therapeutic anticancer compound.
L. plantarum E1/18, E5/6, S0/3, S0/7 Fermented Tolerance to acid and The strain was susceptible to almost all Saelim et al. (2017)
and S3/16 stinky bean (Sa Taw Dong) bile salts antibiotics tested (ampicillin, kanamycin,
Simulated GIT streptomycin, erythromycin, clindamycin,
conditions tetracycline, chloramphenicol,
Cell surface vancomycin, and ciprofloxacin). It had
hydrophobicity neither hemolytic activity nor virulence
Growth under factor genes.
microaerobic and
anaerobic conditions
Hydrolase activity
Antimicrobial activity
Antibiotic susceptibility
Molecular
identification
Candida tropicalis 1A and 3A; Negrinha de Freixo cv. olives Simulated GIT S. cerevisiae strains showed high ability to Oliveira et al. (2017)
Debaryomyces hansenii 8A; conditions grow at 37 ◦ C, lipase activity. The strain
Galactomyces reessii 33A and 34A; Aggregation properties 15A showed higher auto- aggregation
Pichia guilliermondii 25A; P. manshurica Antioxidant activity percentage.
2A; P.membranifaciens 29A and 3B; Enzymatic activity P. guilliermondii and C. norvegica exhibited
Rhodotorula glutinis 27A; R. graminis Capacity to grow at the ability to survive human
20A; Saccharomyces cerevisiae 15A and 37 ◦ C gastrointestinal tract digestion.
15B; Candida boidinii 32A and 37A and S. cerevisiae strains showed high
C. norvegica 7A. antioxidant capacity.
L. casei 24, L. fermentum 38, 47, 62 and Fermentation Process of Simulated GIT The strains produced diffusible inhibitory Ornellas et al. (2017)
L. plantarum 81,90, 96, 100 and 105 “Cupuaçu” Theobroma grandiflorum conditions compounds and promoted co-aggregation
Aggregation properties with S. Typhimurium ATCC 6538.
L. brevis HAC06, HAC08, HAC09, White kimchi Simulated GIT The strains were resistant to adverse Park et al. (2016)
L. plantarum HAC01, HAC02, HAC03, conditions conditions and were able to grow in the
HAC07 and Latilactobacillus sakei Tolerance to phenol cabbage juice in which addition of 2% salt
HAC04, HAC05, HAC10, HAC011 Bile salt deconjugation favorably influenced.
(former Lactobacillus sakei HAC04, Hemolytic activity None of the strains showed hemolysis or
HAC05, HAC10, HAC011) Biogenic amine gelatin hydrolysis activity.
production L. brevis strains produced precursors of
Antibiotic susceptibility biogenic amines, and no other strain
Molecular showed this ability.
identification The susceptibility to erythromycin,
gentamicin, ampicillin, tetracycline,
chloramphenicol, streptomycin,
ciprofloxacin was variable among strains.
Four strains: L. plantarum S-811, S-TF2, Fresh and spoiled cactus (Opuntia Tolerance to acid and Seventeen isolated of LAB. Verón et al. (2017)
Fructobacillus fructosus S-22 and S-TF7 ficus-indica) pears of the green bile salts Four strains potentially probiotic selected.
cultivar Simulated GIT The strains showed starter potential in
conditions cactus pear juice fermentation.
Cell surface All showed resistance to bile salts and
hydrophobicity most of them were resistant to pH 3, but
Aggregation properties not to a pH 2 gastric juice.
Enzymatic activity Five strains (S-02, S-22, S-24, S-TF3 and S-
Antioxidant capacity TF7) showed hydrophobicity values above
Adaptation to food 40%.
matrix Related to auto-aggregation, had values
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Table 1 (continued )
Strains Isolation source Performed test Findings References

Screening of ferulic acid between 5.1 and 62.5%. 3 isolates, S-811,

Production of bioactive S-TF1 and S-TF2, showed feruloyl esterase
compounds activity, related to antioxidant activity.
Molecular
identification
L. brevis 59 Byproducts of fruit pulp process Tolerance to acid and The strains showed tolerance to different Garcia et al. (2016)
L. pentosus 129 Malpighia glabra L., Mangifera bile salts acidic conditions and bile salt
L. paracasei 108 indica L., Annona muricata L., and Ability to growth on concentrations.
L. plantarum 49 Fragaria vesca L. two different Suitable growth in two laboratory and
L. fermentum 111 cultivation media edible growth media, and variable
Molecular susceptibility to different antibiotics.
identification
L. plantarum F22 Traditional inoculum Phab used in Acid production The strain was resistant to bile salt, able to Handa and Sharma
chhang beverage Tolerance to acid and survive in simulated low gastric pH. (2016)
bile salts It showed broad antagonism against a
Cell surface wide range of foodborne/spoilage causing
hydrophobicity bacteria.
Aggregation properties It had a strong auto-aggregation,
Bacteriocin production hydrophobicity, and sensitivity to most of
Antibiotic susceptibility the clinical antibiotics.
Molecular
identification
L. plantarum Kimchi Tolerance to acid and Nutritionally improved DGK-17 was able Khan and Kang (2016)
DGK-17 bile salts to survive under low pH and high bile salt
Survival to GIT conditions, simulated gastric juice
conditions environment.
Aggregation properties Auto-aggregation, co-aggregation and
Adhesion properties adhesion indicated that DGK-17 prevents
colonization of pathogenic bacteria.
L. plantarum 277, 281, 286, 289, 291 and Brazilian cocoa fermentation Simulated GIT The strains showed variation on tolerance Santos et al. (2016)
L. fermentum 260, 265, 266 conditions to heat shock with reduction in viability
Cell surface ranged from 1.9 to 3.4 log orders.
hydrophobicity The resistance to GIT simulation varied
Heat tolerance among strains and L. plantarum 286
Antibiotic susceptibility showed the most promising result,
Molecular followed by L. plantarum 289.
identification L. plantarum 281 displayed higher
hydrophobicity (16.9%).
All strains showed sensitivity to both
amoxicillin and chloramphenicol but
resistance to both vancomycin and
nalidixic acid.
L. casei Q11, L. sakei Q82, L. plantarum Different quinoa varieties and Tolerance to acid and Eighteen LAB isolated were selected based Vera-Pingitore et al.
Q823, Q825 and Leuconostoc lactis amaranth seeds bile salts on the genomic profiling, five presented (2016)
Q615 Simulated GIT no resistance to gentamicin, kanamycin,
conditions streptomycin, erythromycin and
Antibiotic susceptibility clindamycin and were selected for their
potential probiotic.
The strains were tolerant to lysozyme, bile
salts, and had similar adhesion.
L. plantarum Q823 was selected for in vitro
and in vivo studies.
Genera Candida (2 strains), Strains from different sources of Capacity to grow at A hundred and thirty yeast strains isolated Gil-Rodríguez et al.
Kluyveromyces (4 strains), Lachancea which 66 strains were from grape 37 ◦ C from food. (2015)
(1strain), Metschnikowia (5 strains), must Survival the GIT About 50% of the yeast developed at 37%
Pichia (10 strains), Saccharomyces (68 conditions with growth kinetic parameters higher at
strains), Schizosaccharomyces (5 Antioxidant activity 37 ◦ C than at 25 ◦ C.
strains), Torulaspora (23 strains), Molecular A total of 95% of the strains survive to the
Wickerhamomyces (9 strains), Yarrowia identification exposition to conditions simulating to the
(2 strains) and Zygosaccharomyces (1 GIT.
strain) The strains exhibit auto-aggregation
phenotype, antioxidant activity and better
growth capacity at 37 ◦ C than at 25 ◦ C.
Good tolerance to GIT stress conditions,
and high auto-aggregation percentage.
Levilactobacillus spicheri G2 Gundruk a non-salted, fermented Acid and bacteriocin The strain produced lactic acid, Gautam and Sharma
(former Lactobacillus spicheri G2) acidic vegetable production bacteriocin and showed 53.65% of auto- (2015)
Aggregation properties aggregation capacity.
Adhesion to solvents Isolate showed some degree of bile salt
Tolerance to acid and tolerance and presented sensitive to 14
bile salts clinical antibiotics.
Antibiotic susceptibility
L. plantarum 0103, 0123, 0140, 0147, Malaysian Tolerance to acid and Based on acid tolerance thirteen LAB Ng et al. (2015)
0157, 0611, 0612, 0825, 1002; fermented Bambangan bile salts strains were selected.
L. brevis 0808, 08771, (21%), (Mangifera pajang) Aggregation properties Around 36% of the isolated LAB strains
L. rhamnosus 0504, Lactobacillus Enzymatic activity displayed a high survival rate at pH 3.0
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T.C. Pimentel et al. Trends in Food Science & Technology 114 (2021) 25–59

Table 1 (continued )
Strains Isolation source Performed test Findings References

delbrueckii, High temperature and with

L. paracasei and Pediococcus osmotic pressure at least 4 log CFU/mL after 24 h at 2.0%
pentosaceus 1001 Antimicrobial activity bile salt.
Adhesion to solvents Five L. plantarum strains showed at least
Molecular 70% viability at 60 ◦ C/10 min while one
identification L. brevis and three L. plantarum strains
were tolerant to 6% sodium chloride.
They presented aggregation activity
(>20%) in most of the LAB strains.
The maximum β-galactosidase activity
was found in four L. plantarum strains.

to intestinal mucosa leading to biofilm formation, and this feature dis­ antibiotic resistance and, mainly, the possible transmittance of genes
favors the binding of enteropathogens (Pabari et al., 2020). Moreover, should be evaluated for each potentially probiotic strain.
the biofilms formed around these bacteria increase their survival to Safety aspects should also consider the production of biogenic
adverse conditions. Within the used approaches, in vitro adhesion to the amines. Biogenic amines are recognized as undesirable metabolic
Caco-2 cell is the most frequent assay to evaluate that property. Yeast products of some starter cultures or probiotic strains that could be found
and bacteria from fruits, vegetables and traditional fermented foods in fermented foods. The consumption of high concentrations of biogenic
have been showed important adhesion properties (Bonatsou et al., 2018; amines may result in risk health effects for consumers, such as head­
Leandro et al., 2020; Menezes et al., 2019; Vera-Pingitore et al., 2016), aches, neurological disorders, tachycardia, hypotension, among others
which was associated to the production of enzymes and EPS that pro­ (Ku et al., 2020). Previous studies have reported no biogenic amine
mote interactions between probiotics and host-specific receptors and act production by probiotics (Lactiplantibacillus plantarum SBR64.7 and
as a capsule bound to the cell surface protecting against toxic agents and S180.7, Lentilactobacillus buchneri SS50.4, Limosilactobacillus fermentum
stressing conditions encountered during the gastrointestinal tract pas­ SS50.10) isolated from silage, cocoa beans, and artisanal salami (Lean­
sage or by food preservation technologies (García-Ruiz et al., 2014). The dro et al., 2020), and Weissella paramesenteroides isolated from fruits
adhesion properties are dependent on the source of the microorganisms, (Pabari et al., 2020). However, Park et al. (2016) observed that all
and some strains with low adhesion capacity may still present important L. brevis strains isolated from Korean white kimchi could produce
health effects. Therefore, this characteristic may not be primordial for biogenic amines (tyramine and/or histamine). Therefore, the biogenic
probiotic selection (Santos et al., 2016). amine production should be evaluated for each potentially probiotic
To provide health benefits to hosts by improving the nutrient con­ strain.
tent, a probiotic should have the ability to produce related enzymes Probiotic cultures could not present hemolytic activity or the ability
(Fakruddin et al., 2017). Fakruddin et al. (2017) reported that Saccha­ to degrade mucin. Hemolysins are toxins that can cause erythrocytes
romyces cerevisiae IFST062013 isolated from fruit could produce en­ lysis, while the production of enzymes capable of degrading mucin is a
zymes, such as lipase, cellulase, and protease, but it could not produce virulence factor in enteropathogens. Mucins are important as they cover
amylase, DNase, and gelatinase. The production of enzymes (amylase, the epithelial cells of the intestine and prevent mucosal penetration and
protease, lipase, cellulase, and galactosidase) is a positive characteristic translocation by pathogens (Albuquerque et al., 2018). Several LAB
for probiotics (Benitez-Cabello et al., 2019; Fakruddin et al., 2017; isolated from fruit by-products (L. plantarum 53, L. fermentum 56, 60,
Shekh et al., 2019; Song et al., 2020; Won et al., 2020; Xu et al., 2020), 250, 263, 139, 141 and 296, and L. paracasei 106) had no hemolytic
while the absence of production of gelatinase and DNase suggests their activity or ability to degrade mucin. Lack of ability to degrade mucin
safety to be used by human as most of the pathogenic microorganisms was observed for L. plantarum, P. acidilactici, L. fermentum, L. zeae, and
produce these enzymes as part of their pathogenesis. In a general view, L. buchneri isolated from cocoa beans, artisanal salami, and silage
the microorganisms isolated from fruits and vegetables produce bene­ (Leandro et al., 2020), Weisella strains isolated from fruits (banana, or­
ficial enzymes. ange, cherry, sapota, and plum (Pabari et al., 2020), and F. fructosus
The food industries may carefully evaluate new probiotic species’ MCC 3996 isolated from nectar of flowers (B. monosperma) (Patil et al.,
efficacy and safety before incorporating them into food products 2020). However, six bacteria isolated from panchamirtham (Indian
(Maheshiwari et al., 2019). Probiotic strains can be resistant to antibi­ ethnic fermented fruit mix) showed β-hemolytic activity (Maheshwari
otics, remaining in the environment where they are inserted and et al., 2019). Therefore, the hemolytic activity should be evaluated for
allowing their utilization in patients undergoing treatments with anti­ each potentially probiotic strain.
biotics. However, they should not transmit this resistance to other mi­ The reducing power measures the antioxidant activity, nitric oxide
croorganisms to be considered safe for human and animal consumption and hydroxyl radical scavenging activity, cell cytotoxicity and toxicity,
(Fakruddin et al., 2017). Previous studies reported that LAB isolated and metal ion chelating activity. Previous studies reported increased
from fruits and vegetables were sensitive to all the evaluated antibiotics antioxidant activity when microbial isolates of fruits, vegetables, and
(Leandro et al., 2020; Sakandar et al., 2019). Other studies found traditional fermented foods were used (Cao et al., 2019; Di Cagno et al.,
Lactobacillus strains with resistance to antibiotics, such as gentamicin, 2020; Fakruddin et al., 2017; Gil-Rodríguez et al., 2015; Hsiung et al.,
cefuroxime, kanamycin, vancomycin, and erythromycin, but it was 2020; Junnarkar et al., 2019; Lakra et al., 2020; Menezes et al., 2019;
natural or intrinsic and non-transmissible (Anandharaj et al., 2015; Oliveira et al., 2017; Song et al., 2020; Tang et al., 2018). The antioxi­
Pabari et al., 2020; Tang et al., 2018). However, a previous study dant activity was associated with the type of matrix and was
showed that strains isolated from fruit by-products (L. plantarum 53, and strain-specific (Cao et al., 2019; Di Cagno et al., 2020). Yeasts may show
L. fermentum 60) showed potentially transferable resistance to antibi­ higher antioxidant activity than LAB, which has been associated with
otics (tetracycline) (Albuquerque et al., 2018). This characteristic is of the high content of (1/3)-β-D glucan and other β-glucans found in their
paramount importance to guarantee the safety of potentially probiotic cell wall, among other cellular compounds (Menezes et al., 2020).
cultures, but many studies obtained strains with resistance to antibiotics The utilization of prebiotic compounds by probiotic cultures is an
and did not evaluate their possible transmittance of genes (Fakruddin important feature in the development of synbiotic products. Probiotics
et al., 2017; Garcia et al., 2016; Gautam & Sharma, 2015). Therefore, the may utilize prebiotic compounds and increase their viability and/or

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T.C. Pimentel et al. Trends in Food Science & Technology 114 (2021) 25–59

activity in the product. The utilization of prebiotic compounds seems to isolated from cactus pear (109 CFU/mL, 7 weeks) by obese mice resulted
be strain-dependent, as Weissella strains were able to grow in the pres­ in improvements in insulin resistance, hyperlipemia, and hyperglycemia
ence of FOS and GOS, but not on XOS (Pabari et al., 2020). However, (Verón et al., 2019). Finally, the consumption of L. plantarum 49 and
future studies are needed to better evaluate prebiotic compounds’ uti­ L. plantarum isolated from fruits (109 CFU/mL, 28 days) by male healthy
lization by probiotic cultures isolated from unconventional sources. Wistar rats resulted in a reduction in blood glucose levels and total
cholesterol. The effect of long-lasting glycemic control was dependent
on the probiotic strain, with L. plantarum 49 presenting a better per­
2.2. Health effects of probiotics strains from unconventional sources formance (Costa et al., 2019). The results suggest that the hypo­
cholesterolemic and anti-hypertensive effects were frequently observed
Fig. 2 presents the main health effects associated with probiotics after probiotic consumption, while the hypoglycemic properties were
isolated from unconventional sources. The main studies assessing the in dependent on the studied group (healthy, dyslipidemic, obese), pro­
vitro and/or in vivo (human and animal model) physiological effects of biotic strain, and vehiculation form (probiotic strain isolated or in a food
the probiotic strains isolated from unconventional sources are provided product). The reduction of cholesterol may be associated with decon­
in Table 2. The beneficial effects of probiotics on the host are related to jugation of bile salts, resulting in more demand of cholesterol for de novo
one or more mechanisms, such as modulation of the intestinal micro­ synthesis of bile salts and the reduction of its solubility and absorption in
biota, block of pathogen adhesion sites, modulation of the host immune the lumen of the intestine (Saelim et al., 2017). Furthermore, the
responses, and competition by nutrients (Vera-Pingitore et al., 2016). anti-hypertensive properties were related to decreases in systolic arterial
pressure and sympathetic hyperactivity (Cavalcante et al., 2019). The
2.2.1. Hypocholesterolemic, anti-hypertensive and hypoglycemic effects anti-diabetic properties may be influenced by changes in the microbiota
In vitro tests have demonstrated that probiotic cultures from uncon­ and production of short-chain fatty acids (Costa et al., 2019).
ventional sources could assimilate cholesterol. Weissella cibaria MD2 and
Weissella confusa MD1 isolated from Mudakathan dosai (fermented rice 2.2.2. Immunomodulatory and anti-inflammatory properties and antitumor
with leaves of herbaceous vine) could assimilate 67 and 78% of the activity
cholesterol medium (Lakra et al., 2020), while L. pentosus and In vitro tests have also demonstrated that probiotic cultures from
L. plantarum isolated from table olives biofilm assimilated unconventional sources show immunomodulatory properties. Levi­
13.09–38.42% (Benitez-Cabello et al., 2019). The capacity of cholesterol lactobacillus brevis B13-2 (former Lactobacillus brevis B13-2) isolated
assimilation was strain-dependent (Benitez-Cabello et al., 2019). from kimchi induced the expression of some cytokines (IL-1β, TNF-α,
In an in vivo test, the consumption of L. fermentum 296 isolated from and IL-6) and iNOS, indicating immunomodulatory activity (Song et al.,
fruit (109 CFU/mL, 4 weeks) by Wistar rats subjected to a high-fat diet 2020).
resulted in alleviation of hyperlipidemia and blood pressure but no In an in vivo test, the consumption of S. cerevisiae IFST 062013 iso­
impact on glucose tolerance and insulin resistance (Cavalcante et al., lated from fruits (109 CFU/mL, single dose) by male swiss albino mice
2019). At the same time, the consumption of L. fermentum 263, resulted in stimulation of T-lymphocyte specific proliferative response,
L. fermentum 139, and L. fermentum 296 isolated from fruit by-products potentiation of cell-mediated and humoral immunity, and antitumor
(109 CFU/mL, 1:1:1. 8 weeks) by Wistar rats subjected to a dyslipidemic activity. Furthermore, it induced anti and pro-inflammatory mediators
diet resulted in reductions of total cholesterol, increases in and maintained the balance of Th1 and Th2 cytokines, improving host
HDL-cholesterol, and improvements in blood pressure, but no impact on immunity (Fakruddin et al., 2017). The consumption of L. plantarum 81
insulin resistance (Oliveira, Cavalcante, et al., 2020). On the other hand, and L. plantarum 90 isolated from cupuaçu fermentation (108 CFU/mL,
the consumption of cactus pear juice fermented with L. plantarum S-811

Fig. 2. Most relevant in vivo and in vitro effects.

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T.C. Pimentel et al. Trends in Food Science & Technology 114 (2021) 25–59

Table 2
Main studies assessing the effects of probiotic isolated from unconventional sources.
Strains Type Isolation Source Assay Main Effects References
of
study

Hypocholesterolemic, anti-hypertensive and hypoglycemic effects

Weissella confusa MD1 and Weissella In Fermented batter Cholesterol assimilation High reduction in the Lakra et al. (2020)
cibaria MD2 vitro cholesterol levels (>67%)
Sixteen strains: 13 L. pentosus and 3 In Table olives biofilms Cholesterol assimilation All the strains showed the Benítez-Cabello et al.
L. plantarum vitro ability to reduce the cholesterol, (2020)
but with differences among
them.
L. plantarum E1/18, E5/6, S0/3, S0/ In Fermented Cholesterol assimilation All five strains exhibited the Saelim et al. (2017)
7 and S3/16 vitro stinky bean (Sa Taw Dong) ability to assimilate cholesterol.
The strains E5/6 and S0/7 were
more effective.
L. fermentum 296 In vivo Fruit pulp processing Male Wistar rats The administration of Cavalcante et al.
byproducts Control (CTL):commercial diet L. fermentum 296 for 4 weeks (2019)
and saline solution; alleviated hyperlipidemia,
(HF) received a High-fat sympathetic hyperactivity, and
commercial diet and saline reduced systolic blood pressure
solution in HF rats without affecting
(HF + Lf 296) treated with High baroreflex sensibility.
Fat (HF) diet + L. fermentum 296
in a solution of about 1x109 CFU/
mL.
Administration was performed
daily for 4 weeks by oral gavage.
Body weight, biochemical
measurements, glucose and
insulin tolerance tests, arterial
pressure and heart rate records,
spectral analysis of systolic
arterial pressure and cardiac
interval, baroreflex sensitivity,
sympathetic-vagal balance and
sympathetic vascular tone were
evaluated in each group.
L. fermentum 139, 263 and 296 In vivo Fruit by-products Wistar rats (n = 14) were fed with The strains of L. fermentum Oliveira et al. (2020)
a control diet (CTL = 7) or a reduced total cholesterol and
dyslipidemic diet (DLP = 7) increased HDL-c, but did not
during pregnancy and lactation. affect the insulin resistance
After weaning, male and female induced by maternal
offspring received a standard diet dyslipidemia in male and female
up to 90 days of life. offspring and reduced the
Rats were allocated groups: systolic and mean arterial
CTL group + saline solution (n = pressure, only in male.
14); The group that received the
DLP group + saline solution (n = probiotic showed improvement
14) in blood pressure and
DLP group receiving a probiotic sympathetic tone, without
cocktail (n = 14). affecting baroreflex modulation.
A CLT or probiotic formulation The probiotic intervention did
containing L. fermentum 139, 263 not cause significant changes in
and 296 at 1 × 109 CFU/mL) was the body weight, triglycerides,
administered daily by oral VLDL, TBARS, OGTT, UTT.
gavage/8 weeks.
The lipid and glucose profile,
insulin tolerance test,
malondialdehyde (MDA) as
indicator of oxidative stress,
arterial blood pressure,
peripheral chemoreflex,
sympathetic vascular tone were
evaluated.
L. plantarum S-811 In vivo Cactus Pear (Opuntia Ficus- Obese e lean mice were randomly Administration of fermented Verón et al. (2019)
Indica) Juice Fermented in two groups (n = 24): lean juice to obese mice ameliorated
group (L) that was fed a standard the insulin resistance,
diet (SD); an obese group (Ob) hyperglycemia, and
that was fed a high-fat diet hyperlipemia that characterize
(HFD). So, the animals were obesity.
subdivided in four groups (n = 6) O. ficus-indica fruits can reduce
received: 1) water ad libitum. 2) a the percentage of body fat, total
daily dose of L. plantarum S-811 cholesterol, and glycaemia.
as a suspension in water; 3)
pasteurized pear juice; 4) Cactus
pear juice fermented by
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36
T.C. Pimentel et al. Trends in Food Science & Technology 114 (2021) 25–59

Table 2 (continued )
Strains Type Isolation Source Assay Main Effects References
of
study

L. plantarum S-811 at 1.2 × 109

CFU/mL.
The intervention was 7 weeks.
Plasma leptin, insulin, glucose,
cholesterol, triglycerides, glucose
tolerance, homeostatic model
assessment (HOMA-IR) and
intestinal cytokines were
measured.
L. plantarum 49, and L. plantarum In vivo Pulp of Mangifera indica L., or Sixteen L. plantarum strains were Daily administration of Costa et al. (2019)
201 from industrial fruit used as growth promoting effects L. plantarum 49 and L. plantarum
pulp processing byproducts of models in Drosophila 201 did not affect food intake or
Malphigia glabra L., M. indica melanogaster. morphometric parameters.
L., Annona muricata L. and Two strains were selected to Both strains were associated
Fragaria ananassa L. observe the safety aspects and with reduction in blood glucose
beneficial effects of strains using levels and total cholesterol
Wistar rats (n = 48)/21 days. levels.
The animals were randomly The maintenance of the effects
distributed into three groups (n varied with the strain and time
= 16): group Lp49, which of ingestion.
received 9 log CFU/mL of strain The strains were detected in the
L. plantarum 49; group Lp201, intestine and did not cause
which received 9 log CFU/mL of alteration or translocate to
strain and control group, which spleen, kidneys, or liver during
received PBS daily by gavage/4 the experimental or wash-out
weeks. period.
The murinometric parameters,
levels of aspartate transaminase
(AST), alanine transaminase
(ALT) and lipids profile were
measured.
To evaluate the strains’ safety,
histopathological analysis was
performed to observe
translocation.
Immunomodulatory and anti-inflammatory properties, and antitumor activity
Lactiplantibacillus pentosus B13-2 In Chinese cabbage kimchi Expression of cytokines related to Induction of the expression of Song et al. (2020)
(former Lactobacillus pentosus vitro immunomodulation several cytokines (TNF-α, IL-1β,
B13-2) and IL-6) and iNOS by
activation of RAW 264.7 murine
macrophages.
Saccharomyces cerevisiae In vivo Fruit (unspecified) Dose of 150 μL (~109 CFU) were The isolate produced organic Fakruddin et al. (2017)
IFST062013 administered orally on albino acid, improved lymphocyte
mice to safety and immuno- proliferation and cytokine
modulatory activity evaluation. production in treated mice.
L. casei 24, L. fermentum 38, 47, 62 In vivo Fermentation Process of Germ-free Swiss mice of both The strains showed higher Ornellas et al. (2017)
and L. plantarum 81,90, 96, 100 “Cupuaçu” sexes with 6 weeks-old, received survival after enteropathogen
and 105. Theobroma grandiflorum a single dose of 0,1 mL of challenge, lower hepatic
Lactobacillus suspension translocation of
containing about 8.0 log of CFU enteropathogen, lower
by intragastric intubation. histopathological lesions in
The intestinal colonization was ileum and liver and anti-
evaluated in fresh faeces. inflammatory pattern of
Ex vivo antagonism in germ-free immunological response.
mice, cumulative mortality, L. plantarum 81 and L. plantarum
translocation to liver and spleen, 90 were selected as potential
histopathological examination of probiotics.
liver and ileum and mRNA
cytokine gene expression during
an experimental infection with
0,1 mL suspension
S. Typhimurium containing 5.0 log
CFU to enteropathogen
challenge.
Adipogenesis inhibition
Latilactobacillus sakei AM2, AM8, In Kimchi Cell culture and adipocyte Various strains presented Won et al. (2020)
AM32, ADM14, ADM19, YRM13, vitro differentiation test distinct inhibitory effects under
YRM20, RGM12, GP32, GP48, lipid accumulation through
GN57, YG19 (former Lactobacillus inhibition of lipid transcription
sakei AM2, AM8, AM32, ADM14, factors and gene expression.
ADM19, YRM13, YRM20, L. sakei ADM14 was very
RGM12, GP32, GP48, GN57, efficient to inhibit adipogenesis
YG19), L. plantarum S-2AM31 and in adipocytes.
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37
T.C. Pimentel et al. Trends in Food Science & Technology 114 (2021) 25–59

Table 2 (continued )
Strains Type Isolation Source Assay Main Effects References
of
study

Leuconostoc pseudomesenteroides
S-YRWM27, S-YRWM28
L. plantarum S-811 In vivo Cactus Pear (Opuntia Ficus- Obese e lean mice were randomly Administration of fermented Verón et al. (2019)
Indica) Juice Fermented in two groups (n = 24): lean juice to obese mice caused a
group (L) that was fed a standard significant decrease in the body
diet (SD); an obese group (Ob) weight gain.
that was fed a high-fat diet O. ficus-indica fruits can reduce
(HFD). So, the animals were the percentage of body fat.
subdivided in four groups (n = 6)
received: 1) water ad libitum. 2) a
daily dose of L. plantarum S-811
as a suspension in water; 3)
pasteurized pear juice; 4) Cactus
pear juice fermented by
L. plantarum S-811 at 1.2 × 109
CFU/mL.
The intervention was 7 weeks.
Body weight gain, adipose tissue
weight, plasma leptin, insulin,
glucose, cholesterol,
triglycerides, glucose tolerance,
homeostatic model assessment
(HOMA-IR) and intestinal
cytokines were measured.
L. brevis HAC06, HAC08, HAC09, In vivo White kimchi Seven-week-old C57BL/6J male The strain mixture showed a Park et al. (2016)
L. plantarum HAC01, HAC02, mice were housed and provided noticeable decrease in
HAC03, HAC07 and L. sakei with filtered water and rodent mesenteric adipose depot as
HAC04, HAC05, HAC10, diet, ad libitum for 10 weeks. compared to LGG and the
HAC011. After adaptation, each group (n positive control.
= 5) received 1 × 108 CFU viable The intervention resulted in a
cells of a strain mixture (50:50 of significantly lower body weight
L. plantarum HAC01 and L. sakei and total weight gain for eight
HAC13 suspension were given to weeks compared to the high-fat
the mice once/day for 10 more control group.
weeks.
Sterile suspension was
administered to the control
groups.
The L. rhamnosus GG, was used as
reference strain and administered
in the same way.
Antimicrobial activity
Lactiplantibacillus plantarum In Forage plants, cocoa beans Antimicrobial activity Inhibition of pathogenic and Leandro et al. (2020)
SBR64.7 and S180.7 (former vitro fermented spoilage microorganisms
Lactobacillus plantarum SBR64.7 against Staphylococcus sp.
and S180.7), Lentilactobacillus (ATCC® 27,626), Shigella sp.
buchneri SS50.4 (former (ATCC® 23,354), Salmonella
Lactobacillus buchneri SS50.4), sp. (ATCC® 700,623),
Limosilactobacillus fermentum Escherichia coli (ATCC®
SS50.10 (former Lactobacillus 25,922), Pseudomonas sp.
fermentum SS50.10) (ATCC® 19,151), Proteus
mirabilis (ATCC® 21,100), and
Klebsiella sp. (ATCC® 700,834)
and all strains had a high
percentage of CaCo2 cell
adhesion.
Weissella confusa MD1 and Weissella In Fermented batter Antimicrobial activity The strains exhibited Lakra et al. (2020)
cibaria MD2 vitro antimicrobial activity against
foodborne pathogens
(Escherichia coli, Salmonella
enterica, Listeria monocytogenes,
Salmonella typhi and
Staphylococcus aureus).
P. pentosaceus RG7B and C11C, and In Cardinal and Red Globe grape Antimicrobial activity P. pentosaceus RG7B and C11C Taroub et al. (2019)
L. plantarum RG7B vitro fruits and L. plantarum RG8A showed
antifungal activities against
Aspergillus niger aggrégats and
A. carbonarius. Pediococcus
pentosaceus RG7B
Weissella paramesenteroides FX5, In Sapota, cherry, banana, Antimicrobial activity The strains inhibited E. coli and Pabari et al. (2020)
FX9 vitro orange, and plum smashed S. aureus.
fruits
Twenty-one LAB isolates In Fresh vegetables (cauliflower, Antimicrobial activity The cellular extract produced Junnarkar et al. (2019)
vitro gherkins, cluster beans, bacteriocin and bacteriocin like
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38
T.C. Pimentel et al. Trends in Food Science & Technology 114 (2021) 25–59

Table 2 (continued )
Strains Type Isolation Source Assay Main Effects References
of
study

fenugreek, cow pea, bitter substances.

gourd, french beans, tomato, It had antibacterial potential
ridged gourd, cucumber, and against selected plant and
bottle gourd) human pathogens.
Fructobacillus pseudoficulneus In Fresh fruits (apple, banana, Antimicrobial activity The strains inhibited E. coli, S. Sakandar et al. (2019)
JNGBKS, JNGBKS3, F. fructosus vitro Chinese peach, plum, typhimurium, and S. aureus, L.
JNGBKS2, JNGBKS4, cantaloupe, kiwi fruit, and kunkeei strains
F. durionis JNGBKS5, and lychee) flowers (narcissus,
Apilactobacillus kunkeei JNGBKS6, pink rose, red rose, yellow
JNGBKS7, JNGBKS8 (former rose, and sunflower) and
Lactobacillus kunkeei JNGBKS6, honey, and rose petals jam
JNGBKS7, JNGBKS8)
Fructobacillus fructosus MCC 3996 In Nectar of Butea monosperma Antimicrobial activity Antagonistic activity against Patil et al. (2020)
vitro flower B. pumilus (NCIM 2327), E. coli
(NCIM 2109), S. typhimurium
(NCIM 2501), S. aureus (NCIM
2079), Proteus vulgaris (NCIM
2172) and P. aeruginosa (NCIM
2036).
L. plantarum, Enterococcus faecalis, In Leaves of papaya, cassava, Antimicrobial activity Exhibited antimicrobial activity Samedi and Charles
Lactiplantibacillus paraplantarum vitro sugarcane, yam, and taro against Gram-positive Bacillus (2019)
(former Lactobacillus cereus, Listeria monocytogenes,
paraplantarum), and Weissella Staphylococcus aureus,
paramesenteroides and the Gram-negative
Escherichia coli pathogen.
L. plantarum (twenty-five different In De’ang pickled tea Antimicrobial activity The strains had antimicrobial Cao et al. (2019)
strains), and Enterococcus vitro activity against Salmonella
casseliflavus S4b Typhimurium, Escherichia coli
and eight out of the eighteen
strains suppressed growth of
Shigella flexneri.
L. plantarum P24-1, P24-2, P24-3, In Pulque and aguamiel Antimicrobial activity Sixty percent of the isolates Cervantes-Elizarrarás
P24-4, P24-4, P24-5, P24-6, P24-8 vitro exhibited antimicrobial effect et al., (2019)
and Pediococcus acidilactici A1-2, against Escherichia coli,
A2-1 Staphylococcus aureus.
The growth of H. pylori ATCC
43504 was suppressed by all the
LAB.
A hundred and three bacterial In Panchamirtham, an Indian Antimicrobial activity Seventeen isolates exhibited Maheshwari et al.
isolates most of which were vitro ethnic fermented fruit mix antimicrobial activity against (2019)
Bacillus sp., Enterobacter sp., human pathogens: Listeria
acinetobacter sp., Pantoea sp., monocytogenes, Staphylococcus
Microbacterium sp., Klebsiella sp., aureus, Escherichia coli and
sphingomonas sp., Ornithinibacillus Pseudomonas aeruginosa.
sp., Myroides sp., Kocuria sp.,
Alcaligenes sp., and Brevundimonas
sp.
Nineteen LAB strains In Theobroma cacao fermented Antimicrobial activity The strain showed inhibitory Mabeku et al. (2020)
vitro fruit juice effect against 8 H. pylori clinical
strains (Hp0011, Hp0012,
Hp0013, Hp0014, Hp0015,
Hp0016, Hp00116 and
HP00117).
Antagonistic effect was
observed in 65.52% of isolated
LAB.
The LAB19 produced
bacteriocins to control H. pylori
and that LAB4′ , LAB8, LAB11′ ,
LAB12, LAB13′ , LAB15, LAB16
and LAB17 were through
organic acids.
The overall inhibitory activity
was two to three-fold reduced
when CFSs were used instead of
LAB isolates themselves.
Multiple lactic acid bacteria strains In Fermented broccoli, cherry, Antimicrobial activity L. plantarum 445 exhibited the Xu et al. (2020)
vitro ginger, white radish, and highest antagonistic activity
white-fleshed pitaya juice against E. coli, S. aureus, and
L. monocytogenes EGD-e, L.
Already, L. plantarum 430
exhibited the highest
antagonistic activities against
S. Enteritidis and S. Typhimurium.
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39
T.C. Pimentel et al. Trends in Food Science & Technology 114 (2021) 25–59

Table 2 (continued )
Strains Type Isolation Source Assay Main Effects References
of
study

L. pentosus CHIG, NAG1, and In Fermented foods, fruits, and Antimicrobial activity Exhibited inhibition against Shekh et al. (2020)
L. fermentum PRS1. vitro vegetables food spoilage organisms and
intestinal pathogens as
Escherichia coli, Enterobacter
aerogenes, Salmonella typhi
MTCC 98, Serratia marcescens
MTCC 97, Shigella sp.,
Pseudomonas aeruginosa
MTCC2587, Proteus vulgaris,
Klebsiella pneumoniae, Yersinia
enterocolitica MTCC4858,
Enterococcus faecalis,
Micrococcus luteus, Bacillus sp,
Listeria monocytogenes
MTCC1143, Staphylococcus
aureus MTCC1144, Aspergillus
niger MTCC3496, Aspergillus
flavus MTCC2798, Rhizoctonia
solani MTCC633, Penicillium
roqueforti MTCC933 and
Candida strains.
L. fermentum 56, 60, 250, 263, In Different fruit processing Antagonistic activity Inhibition of S. aureus INCQS Albuquerque et al.
139,141 and 296, L. plantarum 53, vitro byproducts 00015, S. typhimurium INCQS (2018)
Lacticaseibacillus paracasei 106 00150, S. enteritidis INCQS
(former Lactobacillus paracasei 00258, L. monocytogenes INCQS
106) 00266, and E. coli INCQS 00219.
L. fermentum (DUR18), L. plantarum In Tempoyak (Malaysian Antimicrobial activity The strains had inhibitory Khalil et al. (2018)
(DUR2, DUR5, DUR8), vitro fermented condiment) activities against the tested
Limosilactobacillus reuteri DUR12 enteric pathogens
(former Lactobacillus reuteri (Staphylococcus aureus, Listeria
DUR12), Lactobacillus crispatus monocytogenes, Salmonella
(DUR4), L. pentosus (DUR20). Typhimurium, E. coli and
Pseudomonas aeruginosa) with
some variations between the
strains and inhibition zone from
3.0 to 16.8 mm.
L. plantarum 3701, 3725, 3739, In Mature Cornus officinalis fruits Antimicrobial activity The strains present inhibition Tang et al. (2018)
3735, 25234, 25294, 2519, 3736, vitro capacity to E. coli ATCC 25922,
3711, L. casei 3734 Salmonella enteritidis ATCC
13076 and S. typhimurium ATCC
14028, Listeria monocytogenes
EGD-e and S.aureus ATCC
29213.
Thirty different strains comprise In Horreh, a traditional Iranian Antimicrobial activity The strains had an important Vasiee et al. (2018)
Lactobacillus (fermentum, vitro fermented food antimicrobial activity,
plantarum, and brevis), Weissella L. innocua and S. aureus were the
cibaria, Enterococcus (faecium and most sensitive indicator bacteria
faecalis), Leuconostoc (citreum and against the selected LAB species,
mesenteroides subsp. while E. coli and B. cereus
mesenteroides) and Pediococcus showed greater resistance.
pentosaceus.
Leuconostoc mesenteroides E14, M67 In Fermented Silage (oat and Antimicrobial activity Both strains had an inhibitory Zarour et al. (2018)
vitro barley) and honey effect against Staphylococcus
aureus and Escherichia coli
Saccharomyces cerevisiae In Fruit (unspecified) Antimicrobial activity Showed moderate antimicrobial Fakruddin et al. (2017)
IFST062013 vitro activity against bacteria and
fungi.
L. plantarum CCMA0743, In Cauim (blended of cassava Antimicrobial activity The strains inhibited Salmonella Freire et al. (2017)
Torulaspora delbrueckii vitro and rice) enterica subsp. enterica serovar
CCMA0235 Typhimurium ATCC 6538,
Escherichia coli ATCC 11229,
and Listeria monocytogenes ATCC
15313.
Pediococcus acidilactici Ch-2 In Chuli (a fermented apricot Antimicrobial activity The strain was able to produce Gupta and Sharma
vitro product) bacteriocin and lactic acid (2017)
against serious food borne and
spoilage microorganisms.
Saccharomyces cerevisiae 15A and In Negrinha de Freixo cv. olives Antimicrobial activity C. norvegica had antimicrobial Oliveira et al. (2017)
15B, Candida norvegica 7A vitro activity against C. neoformans.
Lacticaseibacillus casei 24 (former In Fermentation Process of Antimicrobial activity The strains produced diffusible Ornellas et al. (2017)
Lactobacillus casei 24), vitro “Cupuaçu” Theobroma inhibitory compounds
L. fermentum 38, 47, 62, grandiflorum
and L. plantarum 81,90, 96, 100
and 105
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T.C. Pimentel et al. Trends in Food Science & Technology 114 (2021) 25–59

Table 2 (continued )
Strains Type Isolation Source Assay Main Effects References
of
study

Four strains: L. plantarum S-811 and In Fresh and spoiled cactus Antimicrobial activity Only S-22, S-811, S-TF1, S-TF2 Verón et al. (2017)
S-TF2, Fructobacillus fructosus S- vitro (Opuntia ficus-indica) pears of and S-TF7 isolates showed some
22, and F. fructosus S-TF7 from the green cultivar antibacterial effect against juice
seventeen isolated of LAB contaminant Bacillus sp.
Sixteen strains: 13 L. pentosus and 3 In Table olives biofilms Antimicrobial activity The genotypes assayed were Benítez-Cabello et al.
L. plantarum vitro able to produce the inhibition of (2020)
the food-borne pathogens as E.
coli and L. monocytogenes
Levilactobacillus brevis In Byproducts of fruit pulp Antimicrobial activity The strains showed capacity to Garcia et al. (2016)
59 (former Lactobacillus brevis vitro process Malpighia glabra L., inhibit Staphylococcus aureus,
59), L. pentosus 129, L. paracasei Mangifera indica L., Annona Salmonella Typhimurium,
108, L. plantarum 49, and muricata L., and Fragaria vesca Salmonella Enteritidis, Listeria
L. fermentum 111 L. monocytogenes and Escherichia
coli.
L. plantarum F22 In Traditional inoculum Phab Antimicrobial activity The strain showed antagonistic Handa and Sharma
vitro used in chhang beverage effect against Staphylococcus (2016)
aureus IGMC, Enterococcus
faecalis MTCC 2729, Listeria
monocytogenes MTCC 839,
Clostridium perfringens MTCC
1739, Leucononstoc
mesenteroids, MTCC 107, and
Bacillus cereus CRI.
L. plantarum E1/18, E5/6, S0/3, S0/ In Fermented Antimicrobial activity The strain inhibited Bacillus Saelim et al. (2017)
7 and S3/16 vitro stinky bean (Sa Taw Dong) cereus MST5040, E. coli
DMST4212, Listeria
monocytogenes DMST17303,
Staphylococcus aureus
DMST8840 and Candida
sorphophila SM01 and was able
to inhibit the growth of coliform
during the fermentation.
L. plantarum In Kimchi Antimicrobial activity Nutritionally improved DGK-17 Khan and Kang (2016)
DGK-17 vitro showed antimicrobial activity
against several pathogenic
microorganisms with enhanced
antimicrobial activity and
bactericidal effect against
Pseudomonas aeruginosa 03K711
and Klebsiella pneumoniae
05K279.
Also, DGK-17 protects colon
cells against P. aeruginosa
(03K711) and K. pneumoniae
(05K279).
Endophytic Bacillus sp CaB5 In Surface of young fruits of Antimicrobial activity Identification of non-ribosomal Jasim et al. (2016)
vitro Capsicum annuum Screening for presence of peptide synthases, PKS Type I,
potential biosynthetic gene iturin, surfactin, DAPG and
clusters, the ability to influence gacA genes in the isolate CaB 5
in plant growth. from Capsicum annuum.
The strain proved growth
enhancement on Vigna radiata.
L. plantarum (277, 281, 286, 289, In Brazilian cocoa fermentation Antimicrobial activity All strains showed varying Santos et al. (2016)
291) L. fermentum (260, 265, 266) vitro degrees of pathogen inhibition
in the four tests evaluated.
Levilactobacillus spicheri G2 In Gundruk a non-salted, Antimicrobial activity The strain showed antagonistic Gautam and Sharma
(former Lactobacillus spicheri G2). vitro fermented acidic vegetable spectrum against foodborne (2015)
pathogens/food spoilage
bacteria as Listeria
monocytogenes MTCC 839,
Leuconostoc mesenteroides MTCC
107, Enterococcus faecalis MTCC
2729, Lactobacillus plantarum
CRI, Bacillus cereus CRI,
Clostridium perfringens MTCC
1739, Pectobacterium
carotovorum MTCC 1428,
Escherichiacoli IGMC,
Staphylococcus aureus IGMC and
Aeromonas hydrophila IGMC.
Strains: Candida (2 strains), In Strains from different sources Antimicrobial activity Only S. bayanus IFI-702, Gil-Rodríguez et al.
Kluyveromyces (4 strains), vitro of which 66 strains were from Killer phenotype S. cerevisiae IFI-716 and CYC- (2015)
Lachancea (1strain), grape must SGA, M. pulcherrima IFI-1242
Metschnikowia (5 strains), Pichia and W. subpelliculosa IFI-1253
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41
T.C. Pimentel et al. Trends in Food Science & Technology 114 (2021) 25–59

Table 2 (continued )
Strains Type Isolation Source Assay Main Effects References
of
study

(10 strains), Saccharomyces (68 (8.5% of the 59 strains studied)

strains), Schizosaccharomyces (5 displayed killer activity in at
strains), Torulaspora (23 strains), least one of the conditions
Wickerhamomyces tested.
(9 strains), Yarrowia (2 strains) Overall, the killer phenotype
and Zygosaccharomyces (1 strain) was not widely distributed
amongst the yeasts tested, as it
only appeared in 8.5% of the
strains.
Interestingly, all positive yeasts
to killer exhibited activity
against strains belonging to
other species, and sometimes,
even to different genera.
L. plantarum 0103, 0123, 0140, In Malaysian Antimicrobial activity The strains showed antibacterial Ng et al. (2015)
0147, 0157, 0611, 0612, 0825, vitro fermented Bambangan activity to pathogens
1002; L. brevis 0808, 08771, (Mangifera pajang) Staphylococcus aureus ATCC
(21%), L. rhamnosus 0504, 25,923, Listeria monocytogenes
Lactobacillus delbrueckii, ATCC 13,932, Salmonella
L. paracasei and Pediococcus Typhimurium ATCC 13,311,
pentosaceus 1001 S. Enteritidis ATCC 13,076 and
Yersinia enterocolitica ATCC
23,715.
L. In Italian rye-grass Antimicrobial activity The strain showed antifungal Vijayakumar et al.
plantarum KCC-24 vitro (L. multiflorum) forage activity against (2015)
strains of Aspergillus fumigatus,
Penicillium chrysogenum,
P. roqueforti, Botrytis elliptica,
Fusarium oxysporum
Pichia kudriavzevii DCNa1 and In Different fruits Culture-based bacteriological Yeasts modulated the intestinal Di Cagno et al. (2020)
Wickerhamomyces subpelliculosus vitro analyses of faecal batches microbiota increasing beneficial
DFNb6 Chemical detection of acids and microbes.
volatile compounds
L. fermentum 296 In vivo Fruit pulp processing Male Wistar rats divide in 3 The administration of Cavalcante et al.
byproducts groups: Control (CTL) that L. fermentum 296 for 4 weeks (2019)
received a commercial diet and recovered fecal Lactobacillus sp.
saline solution; (HF) received a counts
High-fat commercial diet and
saline solution and (HF + Lf 296)
treated with High Fat (HF) diet +
L. fermentum 296 in a solution of
about 1x109 CFU/mL.
Administration was performed
daily for 4 weeks by oral gavage.
Fecal microbiota
L. fermentum 139, 263 and 296. In vivo Fruit by-products Wistar rats (n = 14) were fed with The group that received the Oliveira et al. (2020)
a control diet (CTL = 7) or a probiotic showed improvement
dyslipidemic diet (DLP = 7) in fecal Lactobacillus sp. counts
during pregnancy and lactation. The microbial count varied
After weaning, male and female among male and female
offspring received a standard diet offspring and the probiotic
up to 90 days of life. intervention for eight weeks
Rats were allocated groups: CTL resulted in increase in fecal
group + saline solution (n = 14); counts of Lactobacillus spp. in
DLP group + saline solution (n = the DLP male and female
14) and DLP group receiving a offspring and decreased the
probiotic cocktail (n = 14). fecal counts of
A CLT or probiotic formulation Enterobacteriaceae when
containing L. fermentum 139, 263 compared to the CTL and DLP
and 296 at 1 × 109 CFU/mL) was groups.
administered daily by oral
gavage/8 weeks.
Microbial count on fecal samples
were evaluated.
L. plantarum 49, and L. plantarum In vivo Pulp of Mangifera indica L., or Sixteen L. plantarum strains were The strains were detected in the Costa et al. (2019)
201 from industrial fruit used as growth promoting effects intestine.
pulp processing byproducts of models in Drosophila
Malphigia glabra L., M. indica melanogaster.
L., Annona muricata L. and Two strains were selected to
Fragaria ananassa L. observe the safety aspects and
beneficial effects of strains using
Wistar rats (n = 48)/21 days.
The animals were randomly
distributed into three groups (n
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T.C. Pimentel et al. Trends in Food Science & Technology 114 (2021) 25–59

Table 2 (continued )
Strains Type Isolation Source Assay Main Effects References
of
study

= 16): group Lp49, which

received 9 log CFU/mL of strain
L. plantarum 49; group Lp201,
which received 9 log CFU/mL of
strain and control group, which
received PBS daily by gavage/4
weeks.
Fecal microbiota
L. plantarum Q823 In vivo Different quinoa varieties and Seven health individual Fecal counts of 5–7 Log10 CFU/ Vera-Pingitore et al.
amaranth seeds consumed 20 mL of the quinoa- g from the third day onwards. (2016)
based beverage containing 9.19 The counts started to decrease
Log10 CFU/mL once a day for 7 after the final dose. L. plantarum
day and for the subsequent Q823 was still present in all
period (washout) for another 7 subjects four days after the end
days of administration and remained
Fecal samples were analyzed to present after seven days in three
observe colonization. of the subjects.

single dose) by mice resulted in decreases in the production of IFN-γ and Probiotic cultures may modulate the intestinal microbiota. Anaer­
IL-6 (pro-inflammatory cytokines) and increases in the production of obes are the most found microorganisms in the small intestine, but, in
regulatory cytokine IL-10. Therefore, an anti-inflammatory activity was cases of dysbiosis they can be replaced by other microorganisms, mainly
observed (Ornellas et al., 2017). Probiotic cultures can metabolize hemolytic bacteria (Staphylococcus spp.). This can result in inflammation
non-digestible compounds and produce short-chain fatty acids, which in the intestine and/or dysfunction of the gut epithelial cells (Di Cagno
contribute to immunity (Song et al., 2020). et al., 2020). Probiotic cultures isolated from fruits, vegetables, flowers,
and traditional fermented beverages and included in food products may
2.2.3. Adipogenesis inhibition and body weight control improve the intestinal microbiota. Wickerhamomyces subpelliculosus
In vitro tests have demonstrated that probiotic cultures from uncon­ DFNb6 and Pichia kudriavzevii DCNa1 were used as starter cultures in
ventional sources could inhibit adipogenesis in adipocytes. Lat­ cornelian cherry beverage. In vitro tests showed that the functional
ilactobacillus sakei ADM14 (former Lactobacillus sakei ADM14) isolated beverage improved the number of total anaerobes bacteria and
from kimchi decreased the intracellular content of triglycerides on 3T3- decreased the number of Staphylococcus spp., with better results than
L1 adipocytes and reduced the expression of adipogenic marker genes FOS, a recognized prebiotic compound (Di Cagno et al., 2020).
(aP2, PPARγ, FAS, C/EBPα, and CD36), resulting in inhibition of adi­ In in vivo tests, the consumption of L. fermentum 296 isolated from
pogenesis (Won et al., 2020). In an in vivo test, the consumption of cactus fruit (Cavalcante et al., 2019), L. fermentum 263, L. fermentum 139, and
pear juice fermented with L. plantarum S-811 isolated from cactus pear L. fermentum 296 isolated from fruit by-products (Oliveira, Cavalcante,
(109 CFU/mL, 7 weeks) by obese mice resulted in decreases in body et al., 2020), and L. plantarum 49 and L. plantarum isolated from fruits
weight gain (Verón et al., 2019). In another study, the consumption of (Costa et al., 2019) by rats increased the counts of fecal Lactobacillus spp.
L. sakei HAC10 and L. plantarum HAC01 isolated from Kimchi (Korean and decreased Enterobacteriaceae counts (Oliveira, Cavalcante, et al.,
traditional fermented food, 108 CFU/mL, 10 weeks) by obese mice 2020). In a clinical trial, the consumption of a probiotic beverage based
resulted in lower total weight gain and body weight (Park et al., 2016). on quinoa (L. plantarum Q823 isolated from quinoa and amaranth seeds,
EPS produced by probiotic cultures can have inhibitory effects on adi­ 20 mL, 109 CFU/mL, 7 days) by healthy female individuals resulted in
pogenesis and body weight gain (Won et al., 2020). the survival of the probiotic culture to the gastrointestinal tract and
persistence for 5–7 days after the period of washout (Vera-Pingitore
2.2.4. Antimicrobial activity and improvement of intestinal microbiota et al., 2016). The results indicate that probiotic cultures could survive
The production of antimicrobial compounds, such as organic acids, and colonize the gastrointestinal tract of both animals and humans
short-chain fatty acids, and bacteriocins is a characteristic of probiotics (Costa et al., 2019; Vera-Pingitore et al., 2016). However, further studies
(Lakra et al., 2020; Leandro et al., 2020). The inhibition of the classic are needed, as clinical trials only evaluated females and with health
foodborne pathogens (E. coli, S. aureus, and L. monocytogenes, S. Enter­ status.
itidis and S. Typhimurium) by probiotic cultures is advantageous, is
considered as an appropriate alternative to antibiotic treatment, and 3. Techno-functional potential of strains isolated from
could increase the shelf life of food products (Xu et al., 2020). Antimi­ unconventional sources
crobial properties against classical foodborne pathogens have been re­
ported in several studies with LAB isolated from unconventional sources, 3.1. Fruits, vegetables, and flowers
such as forage plants, artisanal salami, and cocoa beans (Leandro et al.,
2020), fruits (Pabari et al., 2020; Taroub et al., 2019), vegetables Fruits, vegetables, and flowers are considered carriers of a wide va­
(Junnarkar et al., 2019), flowers (Patil et al., 2020; Sakandar et al., riety of microorganisms, belonging to the group of bacteria, yeasts, and
2019), plant leaves (Samedi & Charles, 2019), and fermented foods (Cao molds. The diversity and concentration of the microorganisms are
et al., 2019; Cervantes-Elizarrarás et al., 2019; Maheshiwari et al., associated with several factors such as water quality, environmental
2019). In another study, LAB strains isolated from Theobroma cacao conditions, maturation stage, among others (Fessard & Remize, 2019;
fermented fruit juice were able to inhibit Helicobacter pylori by produc­ Rodríguez et al., 2019). In recent years, a series of studies have
ing bacteriocins or organic acids. This is important, as Helicobacter pylori discovered new microorganisms isolated from fruits and vegetables with
is recognized as an infectious agent, causing gastritis, ulcers, and tumors high potential for use in food processing, to improve their nutritional
(Mabeku et al., 2020). In a general view, the antimicrobial effect of and sensory quality, or acting as natural biopreservatives (Barros et al.,
probiotic cultures was more pronounced for Gram-negative bacteria 2019; Costa et al., 2018; Di Cagno et al., 2016; Linares-Morales et al.,
than Gram-positive bacteria (Xu et al., 2020). 2020). However, studies involving the isolation of microorganisms from

43
T.C. Pimentel et al. Trends in Food Science & Technology 114 (2021) 25–59

flowers are still scarce (Patil et al., 2020; Sakandar et al., 2019). Fig. 3 agents, based on in vitro tests, which open opportunities to increase food
presents the main techno-functional properties of microorganisms iso­ products’ shelf life.
lated from fruit, vegetables, flowers, and ethnic fermented beverages. The increase of shelf life of perishable products using bio­
Table 3 presents the main studies that evaluated the techno- preservatives is of industrial interest, mainly due to consumers’ demand
functional properties of strains isolated from fruits, vegetables, and for products with no synthetic preservatives. L. plantarum 49,
flowers. Lactobacillus, Leuconostoc, Weissella, Enterococcus, and Ped­ L. paracasei 108 and L. plantarum 201 isolated from fruits (mango and
iococcus are the LAB genera most frequently isolated from fruits, vege­ acerola) and industrial fruit pulp processing by-products (mango,
tables, and flowers (Di Cagno et al., 2016). However, LAB represent a acerola, and soursop) were able to antagonize the growth of food-related
minority part of the indigenous microbiota of these products, and bacteria in vitro (5 log cfu/mL, Staphylococcus aureus ATCC 952806,
several yeast strains have also been isolated, such as Saccharomyces Pseudomonas aeruginosa ATCC 27853, Listeria monocytogenes ATCC
boulardii, Saccharomyces cerevisiae, Pichia anomala, Rhodotorula mucila­ 1915, Salmonella Enteritidis PT4 and S. Typhimurium PT4). Furthermore,
ginosa, Candida spp., and Ochrobactrum spp. (Barros et al., 2019; Habiba these strains showed a high capacity to produce organic acids and were
et al., 2019; Souza et al., 2017). effective as biopreservatives in Minas Frescal cheese and ground chicken
The LAB and yeast isolated from fruit and vegetables have shown the breast (Costa et al., 2018). The results suggest that LAB and yeasts’
ability to inactivate pathogenic and contaminant microorganisms antimicrobial effect isolated from fruits and vegetables are
(Martins et al., 2019; Peng et al., 2020; Pereira et al., 2015; Souza et al., strain-specific and may be a potentially replacer for synthetic pre­
2017). In in vitro tests, the growth of foodborne pathogens (Listeria servatives (Souza et al., 2017).
monocytogenes) and contaminants (Fusarium oxysporum) was inhibited Fruit and vegetables are very perishable products, resulting in
by Leuconostoc mesenteroides, Enterococcus mundtii and Enterococcus physiological deterioration and changes in the nutritional and quality
faecium isolated from pepper (chilaca and jalapeno), guava, green apple, characteristics due to consumption of internal water and use of reserve
corn, and orange (Linares-Morales et al., 2020). Saccharomyces cerevisiae substances by microorganisms. In products sensitive to fungal infection,
IFST 062013 (9 log cfu/mL) isolated from fruits exhibited moderate it is demanded alternatives that may act against phytopathogens, which
antibacterial (mainly against Gram Negative bacteria) and antifungal opens opportunities for LAB and yeasts isolated from fruits and vege­
(Aspergillus, Penicillium and Rhizopus) activities when compared to con­ tables (Habiba et al., 2019). The endophytic strain L. plantarum CM-3
trols (doxycycline and fluconazole, respectively) (Fakruddin et al., isolated from strawberry (Fragaria × ananassa Duch. Cv. “Hongyan”)
2017). Pichia anomala CCMA0148, Saccharomyces cerevisiae CCMA0159 exhibited inhibitory activity against Botrytis cinerea, which is a fungus
and Rhodotorula mucilaginosa (4–7 log cfu/mL) isolated from coffee and recognized for causing grey mold and a severely damage disease in
cocoa beans exhibited high potential as biocontrol agents, inhibiting the strawberries. This microorganism decreased mycelial growth (in vitro),
growth of the toxigenic fungi Aspergillus carbonarius (CCDCA 10608 and and inhibited spore germination and the incidence and diameter of the
CCDCA 10408) and Aspergillus ochraceus (CCDCA 10612) after cultiva­ lesions (in vivo) (Chen et al., 2020). Saccharomyces boulardii and
tion at 28 ◦ C for seven days, with a reduction of up to 53% in the Saccharomyces cerevisiae (7 log cfu/mL) obtained from the surfaces of
mycelial growth (A. ochraceus CCDCA10612). Furthermore, there was fruits (mango, orange, grapefruit, and lemon) and fresh vegetables
less production of ochratoxin (OTA) (Souza et al., 2017), which is (green pepper and tomato) exhibited antagonistic activity against phy­
important as OTA is recognized as a substance with hepatotoxic, neph­ topathogens, being the maximum activity observed against Penicillium
rotoxic, immunosuppressive, and teratogenic effects, and it can cause digitatum. In the in vivo tests, the yeast delayed natural postharvest
kidney and liver tumors (Souza et al., 2017). The results demonstrate the infection by P. digitatum in kinnow fruits, preserving their quality (lower
potential of using fruit and vegetable-derived strains as bioprotective loss of soluble solids and weight and higher total phenolic content)

Fig. 3. Unconventional sources of microbial isolation and their techno-functional properties.

44
T.C. Pimentel et al. Trends in Food Science & Technology 114 (2021) 25–59

Table 3
Main studies that evaluated the technological properties of strains isolated from fruits and vegetables.
Microorganisms Isolation source Experiments Analysis performed Insightful findings References

LAB strains Strawberry fruit It isolated endophytic LAB Evaluation of the inhibitory L. plantarum CM-3 demonstrated Chen et al.
Lactiplantibacillus (Fragaria × ananassa strains and evaluated their effect of isolates against the highest potential for in vitro (2020)
plantarum CM-3 Duch. cv. “Hongyan”) potential in relation to in vitro B. cinerea in vitro. and in vivo control of B. cinerea
(former Lactobacillus and in vivo biocontrol against Identification and L. plantarum CM-3 decreased the
plantarum CM-3) Botrytis cinerea. characterization of the most mycelial growth of B. cinerea in
(selected) potent antagonist vitro (between 55.27% and
Analysis of mycelial growth, 79.80%)
spore germination and Inhibition of spore germination
elongation of the germ tube in the presence of living cell
Biocontrol efficacy of CM-3 suspensions and reduction of
against B. cinerea in vivo decay incidence and lesion
(Storage at 20 ◦ C for 6 d) diameter of grey mold decay in in
Evaluation of colonization of vivo biocontrol assays
the selected antagonist in Concentrations of 8 and, 9 log
strawberry wounds cfu/mL− 1, decreased the
Concentration: incidence of grey mold in
L. plantarum CM-3: (6, 7, 8 48.23% and 75% respectively;
and, 9 log cfu/mL− 1) Cell-free filtrates of the strain
B. cinerea: (1 × 105 spores failed to provide any protection
mL− 1) against B. Cinerea
LAB strains Fresh fruits and It isolated strains of LAB from Isolation of Lactic Acid 76 LAB isolates were obtained. Linares-Morales
vegetables different fruits and vegetables Bacteria (LAB) Largest amounts of LAB were et al. (2020)
Chilaca chili, and evaluated the antimicrobial Antimicrobial Activity isolated from chilaca chilis (10),
Guava, activity and the enzymatic and (against Gram-positive, guava (8), green apple, jalapeno
Green apple, adhesion activity of these Gram-negative, LAB, mold, chilis, corn (6) and orange (5).
Jalapeño chili, isolates. and yeast strains) LAB strains of Enterococcus
Corn, Protease, lipase, amylase and faecium PIM4, Enterococcus
Orange, citrate metabolism mundtii ELO8, and Leuconostoc
Zucchini, Adhesion Capacity mesenteroides (PIM5, CAL14)
Peach, Biofilm Formation strongly inhibited the growth of
Red apple, Molecular Identification L. monocytogenes
pear, Enterococcus mundtii (JAV15,
Green tomato, TOV9) strongly inhibited the
Pomegranate, growth of F. oxysporum.
Lettuce, Seven isolates hydrolyzed starch,
Mandarin, 46 proteins, 14 lipids, and 36
Cucumber, metabolized citrate.
Bell pepper, LAB isolates with the best
Grape, activities were molecularly
Soybean sprout, identified as Leuconostoc
Nopal flower mesenteroides, Enterococcus
(Obtained from mundtii and Enterococcus faecium.
Farmlands, Street
markets, and
Supermarkets
in Chihuahua City,
Mexico)
L. plantarum 53, Pulp fruit of Malphigia It evaluated the effects of mixed Enumeration of viable counts The viable counts of LAB strains Oliveira et al.
Lacticaseibacillus glabra L. and culture of four potentially (zero - right after after the 120-h fermentation of (2020)
paracasei 106 (former Mangifera indica L. or probiotic Lactobacillus isolates homogenization and after 8, the by-products were ≥5.5 log
Lactobacillus paracasei Byproducts of on the physicochemical 24, 48, 72 and 120 h of cfu/mL
106), M. glabra L., M. indica characteristics, contents of fermentation). The fermentation increased the
Limosilactobacillus L. and bioactive compounds and Physico-chemical analysis luminosity, titratable acidity,
fermentum 56 Annona muricata L. antioxidant activity (color, pH, titratable acidity total flavonoids, total phenolics
(former Lactobacillus of acerola and guava industrial and total soluble solids). and antioxidant activity and
fermentum 56) and by-products submitted to a Determination of bioactive decreased the content of total
Lacticaseibacillus casei submerged fermentation process compounds soluble solids and the pH in the
L-26 (former for 120 h. Analysis of antioxidant two by-products tested.
Lactobacillus casei L-26) activity by FRAP and ABTS. Ascorbic acid content decreased
Determination of the content in the fermentation of the acerola
of ascorbic acid, carotenoids, by-product and increased in the
total flavonoids and total guava.
phenolics. There were no significant
Concentration: changes in the carotenoid
LAB: (10 log cfu/mL) content.
Mix of strains (1:1:1:1 ratio). Different phenolic compounds
(mainly Kaempferol and
quercetin) were found during the
fermentation of the by-products.
Yeasts strains Fresh fruits and seeds It isolated endophytic strains of Identification of Main genera identified were Barros et al.
of: yeasts present in the fruits of microorganisms Candida, Rhodotorula and (2019)
Achachairu (Garcinia achachairu, araçá-boi and Fermentation Ochrobactrum.
humilis), bacaba, and evaluated their Determination of total The largest number of strains
Araçá-boi (Eugenia potential phenolic compounds detected was in araçá-boi (3),
(continued on next page)

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T.C. Pimentel et al. Trends in Food Science & Technology 114 (2021) 25–59

Table 3 (continued )
Microorganisms Isolation source Experiments Analysis performed Insightful findings References

stipitata) and in the production of bioactive Identification and followed by bacaba (2) and
Aacaba (Oenocarpus compounds and their quantification of phenolic and achachairu (1).
bacaba) fruits antioxidant activity. flavonoid compounds Rhodotorula glutinis produced the
by UHPLC-QqQ-MS/MS highest amount of total phenolics
Analysis of antioxidant among all 3 fruit residues and
activity. several species of isolated
Concentration: microorganisms.
Yeasts: (7 log cfu/mL) The use of bacaba waste as a
medium resulted in a higher
production of total phenolics.
Higher concentrations
were found for chlorogenic acid
(460.00%), catechin (94.75%)
and rutin (27.53%).
The antioxidant activity
increased [ORAC (118.62%),
FRAP (90.32%) and ABTS
(39.25%)] in fermented
products.
LAB strains Pickled white Identified and characterized Ability to produce 77 isolated strains belonging to Fessard and
cabbage (Brassica LAB strains isolated from fruits exopolysaccharides (EPS) the species Lactobacillus Remize (2019)
oleracea and vegetables, as well as Identification of species and plantarum (3), Lactobacillus spp.,
var. capitata), evaluated the ability of isolates typing (3), Lactococcus lactis (13),
Papaya (Carica to produce exopolysaccharides Analysis of tolerance to acid, Leuconostoc pseudomesenteroides
papaya) and (EPS), the influence of oxidative and osmotic stress (25), Leuconostoc lactis (1),
Tomatoes growth temperature and and bile salts Leuconostoc mesenteroides (7),
(Lycopersicon tolerance to acid, salt, bile salts Analysis of growth parameters Leuconostoc citreum (14),
esculentum) or hydrogen peroxide. Weissella cibaria (4), Weissella
confusa (4), other Weissella
species (2) and Fructobacillus
tropaeoli (1).
Several isolates from Weissella
and Lactobacillus were
particularly tolerant to acid and
osmotic stress.
Lc. pseudomesenteroides 60 was
highly tolerant to oxidative
stress.
Weissella 30, 64 and 58,
Leuconostoc 60 and 12b,
Lactobacillus 75 and
Fructobacillus 77 were the
isolates that have the best
characteristics for their use as
starters or as preservatives.
Isolates of Saccharomyces Fresh fruit and Isolation and identification of Isolation and identification of 25 strains of epiphytic yeasts Habiba et al.
boulardii e vegetables surfaces: epiphytic yeasts from healthy epiphytic yeast were isolated, 20 of these (2019)
Saccharomyces Mango (Mangifera fruit and vegetables, and their In vitro antifungal assay showed maximum in vitro
cerevisiae indica L.) role in delaying the natural against Penicillium digitatum activity against P. digitatum.
Green chili (Capsicum postharvest P. digitatum rot and Evaluation of yeast in delaying Eight selected isolates showed
annuum L.) physiochemical properties in natural postharvest infection less infection of P. digitatum with
Orange (Citrus sinensis kinnow at room temperature to of P. digitatum in kinnow fruits minimal loss of quality compared
(L.) Osbeck cv. 15 days. Evaluation of yeast in delaying to the control and positive
Valencia) post-harvest decomposition in control (1% K sorbate).
Grape fruit (Citrus kinnow HAB-31 and HAB-53 isolates
paradise Macfad.) Physiochemical Analysis showed better characteristics.
Lemon (Citrus limon Concentration: Decreased weight loss (18.39%
(L.) Burm.) and Yeasts: (7 log cfu/mL) and 18.87%, respectively),
Tomato (Lycopersicon P. digitatum spores: (6 log cfu/ soluble solids and improved
esculentum Mill.) mL) phenolic content.
No effects on firmness.
S. cerevisiae CCMA 0200 Sugar cane Evaluated the performance of Microorganisms quantification T. delbrueckii CCMA 0684 Martins et al.
(former UFLA CA11) Coffea arabica L. var. yeasts Saccharomyces cerevisiae Analysis of acids and sugars showed better adaptation to the (2019)
T. delbrueckii CCMA Bourbon5 CCMA 0200 e Torulaspora Volatile compounds process (72 h of drying).
0684 delbrueckii Sensory analysis Eighteen volatile compounds
CCMA 0684 as a starter in coffee Concentration: were detected in green coffee
varieties processed by wet Yeasts: (7 log cfu/mL) and 75 in roasted coffee.
method fermentation and the 2-Furanmethanol propanoate
impact on sensory quality and and 2-ethyl-3,5-dimethylpyra­
compound profile zine were identified only in the
inoculated treatments.
All treatments received scores
greater than 80 in the sensory
analysis.
T. delbrueckii CCMA 0684
exhibited better results in
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46
T.C. Pimentel et al. Trends in Food Science & Technology 114 (2021) 25–59

Table 3 (continued )
Microorganisms Isolation source Experiments Analysis performed Insightful findings References

sensory analysis, being

preferable for use as a starter.
LAB strains Ripe wild fruits of Evaluated the lactic microbiota Microbiological Analyses and Bacteria of the genera Rodríguez et al.
guava (pink and of these tropical fruits and wild Isolation of LAB Enterococcus, Fructobacillus, (2019)
yellow varieties), flowers and their technological Analysis of metabolic Lactobacillus, Lactococcus,
Papaya, properties of metabolite properties (mannitol and Leuconostoc and Weissella were
Passion fruit, production. organic acids production) observed, and 21 different
Custard apple, Analysis of aroma compounds species were identified
Medlar, (diacetyl production) Most of the strains found were
Mulberry, Analysis of technological capable of producing mannitol,
Fig, properties (Acidifying capacity organic acids and aromatic
Khaki, and and growth rate bacterial, compounds; as well, some had
Flowers of Medlar, pectinolytic activity, activities of cinnamoyl esterase,
Passion fruit, cinnamoyl esterase activity, pectinase and esterase.
Custard apple, biogenic amine production)
Papaya
Sixteen LAB strains Pulp fruit of Malphigia Evaluated the antagonistic Antagonistic activity assays All LAB strains tested were able Costa et al.
glabra L. and activity of sixteen LAB strains Assays with cell-free culture to antagonize the growth of food- (2018)
Mangifera indica L. or derived from fruits against food- supernatants related bacteria in vitro.
Byproducts of related bacteria and the effects Analysis of organic acids The highest production of
M. glabra L., M. indica on the survival of production organic acids was detected for
L. and Annona L. monocytogenes and Bioconservation analyzes in L. plantarum 49, L. paracasei 108
muricata L. S. Enteritidis PT4 in fresh cheese ground chicken and cheese and L. plantarum 201.
and ground chicken breast "Minas Frescal" L. plantarum 49 and L. paracasei
Analysis of production of 108 decreased L. monocytogenes
organic acids in the matrices. counts in cheese.
Concentration: L. paracasei 108 decreased the
LAB strains and indicators: (5 count of S. Enteritidis PT4 in
log cfu/mL) chicken breast.
Saccharomyces cerevisiae Fruit (non-specified) Evaluated the antimicrobial and Analysis of antibacterial and Saccharomyces cerevisiae IFST Fakruddin et al.
IFST 062013 antioxidant activities of whole anti-fungal activity 062013 showed moderate (2017)
cells, culture supernatant and Analysis of antioxidant antibacterial and antifungal
cell lysate of the isolated activity. activity when compared to the
Saccharomyces cerevisiae IFST Concentration: control (doxycycline and
062013 Saccharomyces cerevisiae IFST fluconazole)
062013: (9 log cfu/mL) Cell lysate showed more effective
antibacterial and antifungal
effects.
Antibacterial effect of the isolate
was better against gram negative
pathogens
Saccharomyces cerevisiae IFST
062013 showed strong
antioxidant activity, reducing
power, sequestering activity of
nitric oxide and hydroxyl radical,
significant cytotoxicity of
artemia and acute toxicity and
chelating activity of metal ions.
Twenty-eight LAB strains Tomatoes Evaluated the production of Microbiological analyses and 34 strains of the genera Fessard et al.
(Lycopersicon exopolysaccharides (EPS) and isolation of LAB Lactobacillus, Leuconostoc, (2017)
esculantum), the acidification kinetics of LAB Analysis of EPS production Weissella, Lactococcus and
Papaya (Carica isolated from fruit and its and acidification kinetics Fructobacillus were isolated.
papaya) and potential as a starter for Growth in apple juice Among these, 13 species were
Sliced cabbage fermentation in pineapple juice Production of fermented represented.
(Brassica oleacera var. (40 mL), teas (green and black) Physico-chemical analyzes and EPS formation was observed for
capitata) (40 mL) or fruit puree (papaya viability all strains of Leuconostoc spp.,
or mango) (40 g) with views for Determination of sugar and W. confusa and W. cibaria.
technological use. phenolic content The kinetic parameters of
Sensory analysis acidification showed different
Analysis of antioxidant behavior within the group of
activity strains. Fermented juices were
Concentration: sensorially appreciated.
LAB: (5 log cfu/mL) W. cibaria 64 produced little
grew in apple or
pineapple juice, however,
significantly increased the
phenolic content and antioxidant
activity.
Lc. pseudomesenteroides 12b
increased antioxidant activity in
fermented papaya pulp,
however,
decreased phenolic content in
infusion of green tea and mango
pulp.
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T.C. Pimentel et al. Trends in Food Science & Technology 114 (2021) 25–59

Table 3 (continued )
Microorganisms Isolation source Experiments Analysis performed Insightful findings References

LAB strains Potato tocosh Evaluated the diversity of LAB Microbiological analyses and The genus Lactobacillus was Jiménez et al.
peruvian present in tocosh, in addition to isolation of LAB (both culture- predominant in LAB populations (2017)
(freshly harvested the safety aspects and the dependent and -independent at any stage of tocosh
potatoes, one-month biotechnological potential of approaches) fermentation
and eight-months these isolates for future Analysis of exopolysaccharides Latilactobacillus sakei (former
production) commercial applications (EPS) production Lactobacillus sakei) and Lc.
Analysis of phytate mesenteroides were present in all
degradation activity tocosh samples.
and amylolytic activity Of the twenty-four LAB species
Analysis of group B vitamin detected, six were recovered by
production and biogenic culture.
amines production Lb. sakei and Ln. mesenteroides
Analysis of antimicrobial strains exhibited the ability to
activity break down amylase and
phytate, as well as the
production of EPS and B vitamins
(riboflavin and folate)
Tocosh LAB species showed
antibacterial activities, as well as
the ability to produce biogenic
amines.
Thirty-two yeasts strains Coffee and Evaluated the antagonistic Toxigenic fungi biocontrol in Yeasts showed greater inhibitory Souza et al.
Cocoa beans potential of yeast isolates vitro assays effects (53%) on the mycelial (2017)
against Aspergillus carbonarius Evaluation of vegetative growth of the isolate A. ochraceus
(CCDCA 10608 and CCDCA growth (CCDCA10612).
10408) and Aspergillus ochraceus Evaluation of Aspergillus spore Pichia anomala CCMA0148 and
(CCDCA 10612) after cultivation production Saccharomyces cerevisiae
at 28 ◦ C for seven days. Antagonistic activity of CCMA0159 provided the greatest
different water activity over growth inhibition of all fungal
ochratoxin (OTA) production strains.
Analysis of antagonistic Rhodotorula mucilaginosa was
activity by potential yeast effective in inhibiting the
strain at cellular level. production of OTA by the three
Concentration: isolates of Aspergillus.
Yeasts: (4–7 log cfu/mL) S. cerevisiae CCMA0159 and
Aspergillus: (105 spores mL− 1) Pichia anomala CCMA0148
showed high potential as
biocontrol agents under the
conditions tested
Leuconostoc mesenteroides Fresh fruits of Opuntia It evaluated the technological Isolation, identification and Strains showed ability to degrade Di Cagno et al.
strains ficus-indica (L.) Mill., potential of autochthonous starter selection mucilage in vitro. (2016)
(genotype Sanguigna) lactic acid bacteria isolated from Fermentation of prickly pear Leuc. mesenteroides OP4, OP9,
prickly pear (Opuntia ficus-indica fruit puree OP21, OP18 and OP23 showed
L.) fruits as a starter for Microbiological analysis the best growth and acidification
fermentation, as well as Quality parameters performance, and synthesis of
improvement in shelf life, Analysis of carbohydrates, exopolysaccharides (EPS) during
rheological, sensory and organic acids, volatile fermentation.
functional characteristics in components, and free amino Strains improved their shelf life,
prickly pear fruit puree acids color parameters, darkening
Antimicrobial assays index, sensory attributes,
Analysis of determination of antimicrobial activity, vitamin C
total phenols and DPPH and betaine levels.
radical scavenging activity The fruit puree initiated
Vitamin C and betalains significantly inhibited the
analysis inflammatory state of Caco-2
Functionality analysis using cells in addition to improving the
Caco-2/Tc7 cells immunomodulatory activity of
Sensory analysis the fruit puree.
Concentration:
(4%, v/v) (7 log cfu/g− 1)
LAB strains Coffee fruits Evaluated the potential use of Microbiological analyses for L. plantarum LPBR01 showed Pereira et al.
lactic acid bacteria (LAB) LAB selection better performance as a starter (2016)
isolated from coffee fruits and Fermentation assays for the fermentation of coffee
selected to conduct the moist Analysis of determination of beans.
fermentation process of coffee sugars and organic acids by L. plantarum LPBR01 had no
beans in order to improve the HPLC significant influence on yeast
quality attributes of the Headspace solid phase micro growth.
beverages produced with these extraction and gas L. plantarum LPBR01 improved
beans chromatography analysis the production of volatile
Coffee sensorial quality aromatic compounds during the
evaluation fermentation process.
Concentration: Beverages produced with coffee
(7 log cfu/g− 1) beans inoculated with
L. plantarum LPBR01 presented
better aroma, flavor, acidity,
body, balance, aftertaste, and
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T.C. Pimentel et al. Trends in Food Science & Technology 114 (2021) 25–59

Table 3 (continued )
Microorganisms Isolation source Experiments Analysis performed Insightful findings References

overall quality in the quality test

compared with the conventional
process.
Lactiplantibacillus Industrially It evaluated the viability of Microbiological analyses Strains showed good survival Argyri et al.
pentosus B281 fermented olives potentially probiotic LAB during Physicochemical analyses (pH, rates. (2015)
(former Lactobacillus. the shelf life of fermented green water activity, colour, acidity, The initial and final LAB
pentosus B281) olives cv. Halkidiki stored (4 and salt content and firmness) population was 5.6–5.9 and
L. plantarum B282 20 ◦ C for 12 months) in modified Sensory Analyses 5.8–6.4 log CFU/g respectively.
atmosphere (70% N2–30% Concentration: Both strains were recovered in
CO2), as (~7–8 log cfu/mL) high percentages (64.7% for
well as the quality attributes. B281 and 94.1% for B282) after
6 months and in lower survival
rates after one year (35.0% for
B281 and 13.3% for B282) at
4 ◦ C.
Lower survival rates were
observed after one year.
Olives showed better
acceptability after 6 months than
12 months of storage at both
temperatures.
Pichia fermentans YC5.2 Spontaneous coffee Evaluated the use of Fermentation assays Sucrose did not interfere with the Pereira et al.
fermentation P. fermentans YC5.2 as an initial Microbiological analyses growth or frequency of P. (2015)
culture in controlled Metabolite target analysis fermentans YC5.2
fermentations of coffee beans of Determination of volatile P. fermentans YC5.2 increased the
varieties Mundo Novo and compounds and sugar production of specific volatile
Catuaí Vermelho with or composition aroma compounds (e.g., ethanol,
without the addition of 2% (w/ Quality evaluation acetaldehyde, ethyl acetate and
v) sucrose during wet isoamyl acetate) and decreased
processing, and its impact on the the production of lactic acid
metabolite profile. during the fermentation proce.ss.
Concentration: The strain improved the quality
(7 log cfu/mL) of the coffee by intensifying the
’vanilla’ flavor and ’floral’
aromas

compared to control and positive control (potassium sorbate at 1%) such as the capacity to hydrolyze starch, proteins, lipids and metabolize
(Habiba et al., 2019). Chemical fungicides are the cheapest approach to citrate (Linares-Morales et al., 2020). Enterococcus, Fructobacillus,
minimize postharvest spoilage of fruits; however, they can result in the Lactobacillus, Lactococcus, Leuconostoc, and Weissella isolated from ripe
development of chemical residues, pathogens resistance, phytotoxicity fruits and wildflowers presented enzymatic activity related to cinnamoyl
to the environment or other microorganisms, and public health issues esterase, pectinase, and esterase enzymes (Rodríguez et al., 2019).
(Chen et al., 2020). Therefore, the studies suggest that LAB and yeasts Therefore, these cultures may be used for producing enzymes. The
isolated from fruits and vegetables could be used as potential alterna­ production of enzymes by microorganisms is more desired than those
tives to synthetic fungicides, being eco-friendly, safe, and renewable, from other sources because they can show improved catalytic activities,
and resulting in lower postharvest losses and products with improved increased yields, and be produced using culture media of low costs
quality parameters. (Linares-Morales et al., 2020).
The food industry is interested in low-cost processes for obtaining The utilization of LAB from fruit and vegetables in food products
ingredients and compounds of interest. LAB and yeasts can contribute processing may improve their sensory properties and increase bioactive
with food quality due to their fermentative activity, producing various compounds’ content. Lactobacillus, Leuconostoc, Weissella, Lactococcus,
desirable compounds, such as organic acids, enzymes, bacteriocins, and and Fructobacillus isolated from tomato, papaya, and cabbage were able
EPS (Peng et al., 2020). Enterococcus, Fructobacillus, Lactobacillus, Lac­ to form EPS and showed good acidification properties. Therefore, they
tococcus, Leuconostoc and Weissella were isolated from ripe fruits (pink were used as starter cultures (5 log cfu/mL) in fermentation of apple
and yellow guava, papaya, passion fruit, pinecone, loquat, blackberry, juice, pineapple juice, and papaya pulp, improving the products’ anti­
fig, khaki) and wildflowers (flower loquat, passion fruit flower, pine­ oxidant activity and sensory profile (Fessard et al., 2017). Candida,
cone, papaya). Most of the strains showed good production capacity for Rhodotorula and Ochrobactrum isolated from Brazilian fruits (araçá-boi,
mannitol, organic acids, and aromatic compounds (Rodríguez et al., bacaba and achachairu) proved to be effective as starter cultures (7 log
2019). Lactilactobacillus sakei (former Lactobacillus sakei) and Leuconos­ cfu/mL) in fermentation of the residue of these fruits, increasing total
toc mesenteroides isolated from Peruvian tocosh potato demonstrated phenolic contents and antioxidant activity (Barros et al., 2019). A mix
ability to metabolize starch and phytate, and produce EPS, as well as (1:1:1:1 ratio) of the potentially probiotic strains L. plantarum 53,
synthesizing B vitamins (riboflavin and folate). Therefore, these LAB L. paracasei 106, L. fermentum 56, and Lacticaseibacillus casei L-26
could be used to improve the nutritional value of food products by (former Lactobacillus casei L-26) (10 log cfu/mL) isolated from fruits
increasing the concentrations of important compounds (vitamins) and (mango and acerola) and fruit by-products (mango, acerola, and sour­
decreasing antinutrients (phytate) (Jiménez et al., 2017). sop) positively affected the physicochemical characteristics of
The utilization of starter cultures of vegetal origin is important for agro-industrial by-products of acerola (Malpighia emarginata DC) and
vegan products, as most of the starter cultures commercially available guava (Psidium guajava L.) during submerged fermentation for 120 h.
are of dairy origin (Pimentel et al., 2021). Leuconostoc mesenteroides, They improved brightness, titratable acidity, flavonoid content, bioac­
Enterococcus mundtii and Enterococcus faecium isolated from pepper tive compounds content (ascorbic acid and phenolic compounds) and
(chilaca and jalapeno), guava, green apple, corn, and orange presented antioxidant activity (Oliveira, Araújo, et al., 2020). Finally, Leuconostoc
important characteristics to be used as starter cultures in food products, mesenteroides (OP4, OP9, OP21, OP18 and OP23) isolated from fresh

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T.C. Pimentel et al. Trends in Food Science & Technology 114 (2021) 25–59

fruits of Opuntia ficus-indica (L.) Mill. (Sanguigna genotype) showed in cell viability (even without glucose) for up to 15 days, good adaptability
vitro ability to degrade mucilage, and excellent performance of acidifi­ to high concentrations of salt (up to 10% NaCl), low temperature (7 ◦ C)
cation and synthesis of EPS during fermentation of mashed figs from and low pH (about 3.6), good tolerance to phenolic content of common
India (7 log cfu/g). Therefore, they increased shelf life, improved color olives (0.3% of total phenol), and high degradability of oleuropein and
parameters, darkening index, sensory attributes, increased antimicro­ β-glucosidase activity (Iorizzo et al., 2016). In another study, the yeast
bial activity, and vitamin C and betaine (Di Cagno et al., 2016). Overall, strain Wickerhamomyces anomalous Y12 and a combination of LAB
the studies demonstrate that the utilization of LAB and yeasts isolated strains (L. pentosus LPG1, L. pentosus Lp13, L. plantarum) resulted in
from fruit and vegetables as starter cultures may enhance safety, shelf lower pH values and higher acid production and sugar consumption,
life, texture, sensory properties, and health-promoting characteristics of LAB multiplication, biofilm formation, and production of auto-inducer-2
the products. (AI-2) in Spanish green olives. Furthermore, the sensorial analysis did
Coffee fruit is commonly processed by fermentation using the not show differences between the inoculated treatments and uninocu­
indigenous species, eliminating the mucilage of the beans and forming lated (spontaneous) control (Benítez-Cabello et al., 2020). L. pentosus
volatile compounds (Pereira et al., 2015). B281 and L. plantarum B282 (~7–8 log cfu/mL) isolated from industrial
It is of interest for the industry to obtain consistent quality products, fermented olives showed good viability (64.7% and 94.1%, respectively)
which may be achieved by controlling the fermentation step (Pereira after 6 months and 1 year (35.0% and 13.3% respectively) in fermented
et al., 2015). LAB and yeasts isolated from fruit and vegetables may be green olives (cv. Halkidiki) stored (4 and 20 ◦ C) in modified atmosphere
used in the processing of coffee. L. plantarum LPBR01 (7 log cfu/g) (70% N2–30% CO2). There were no significant changes in pH, acidity,
isolated from coffee fruits were used as starter culture for the wet or salt content, but the olives treated with the strains showed better
fermentation of coffee beans, resulting in improved production of vol­ results for color and firmness, reflecting better sensory acceptance
atile aromatic compounds during fermentation process. The produced (Argyri et al., 2015). The results suggest that microorganisms isolated
beverages showed better aroma, flavor, acidity, body, balance, residual from olives may be used as starter cultures in their fermentation,
flavor, and overall quality compared to that obtained by the sponta­ resulting in an accelerated fermentation process and products with
neous fermentation (Pereira et al., 2016). Saccharomyces cerevisiae similar or improved characteristics compared to those obtained by
CCMA 0200 isolated from sugar cane and Torulaspora delbrueckii CCMA spontaneous fermentation.
0684 isolated from Coffea arabica L. var. Bourbon5 were used in the wet LAB and yeasts with technological potential have also been isolated
fermentation of coffee varieties (Mundo Novo and Catuaí Vermelho). It from different traditional fermented foods and beverages worldwide.
was observed improvements on volatile compounds’ profile, resulting in The main technological properties reported were efficacy as starter
better sensory quality of the coffee beverages (Martins et al., 2019). In cultures, antimicrobial and antifungal activity, capacity for degradation
another study, Pichia fermentans YC5.2 (7 log cfu/mL) isolated from of toxins, and production of desirable compounds (Aka et al., 2020; Pei
spontaneous coffee fermentation increased the production of et al., 2020; Ramos et al., 2015; Taheur et al., 2020). Limosilactobacillus
aroma-specific volatile compounds (e.g. ethanol, acetaldehyde, ethyl fermentum S6 (former Lactobacillus fermentum S6) and Pediococcus acid­
acetate and isoamyl acetate) and decreased lactic acid production dur­ ilactici S7 were isolated from fermentation of tchapalo (traditional
ing the coffee bean fermentation process, resulting in a coffee beverage Ivoirian beverage produced from fermented sorghum) and showed good
with improved sensory quality due to the more intense “vanilla” and ability to act as starter cultures in the production of sorghum-based
“floral” aromas (Pereira et al., 2015). The results indicate that the use of beverages and to produce bacteriocins (Aka et al., 2020). Torulaspora
starter cultures isolated from fruits and vegetables can reduce the delbrueckii, Pichia exigua and Bacillus spp. isolated from tarubá (tradi­
fermentation time and improve the aroma and flavor of the coffee tional fermented beverage from cassava) demonstrated ability to
beverages, resulting in a product with a distinctive flavor and added degrade starch, proteolytic activity and thermotolerance. These char­
value (Martins et al., 2019; Pereira et al., 2016). acteristics are desired for cassava fermented foods production; there­
fore, the isolated microorganisms could be used as starter culture
3.2. Ethnic fermented beverages (Ramos et al., 2015). A diversity of microorganisms was isolated from
chichas (indigenous Andean beer) produced with rice, oats, grapes, and a
Fermented foods are good sources for the isolation of LAB and yeasts mixture of seven varieties of corn. Most microorganisms showed good
with tecno-functional properties (Menezes et al., 2020; Taheur et al., tolerance to stress conditions (concentrations of ethanol, low and high
2020). Fermented dairy products, such as fermented milks, yogurts, and temperature and osmotic stress). Cryptococcus sp. showed the highest
cheeses, have been the main source of microorganisms over the years, as amylase activity, while Saccharomyces cerevisiae EYS5 presented the
they are products produced on a large scale and with great commercial highest invertase enzyme activity and tolerance to dehydration. The
appeal (Rezac et al., 2018). However, several studies have shown that results suggested that the isolated yeasts could be used as dry starters to
the behavior and efficacy of a strain is related to its source, which has produce chichas or in other alcoholic fermentations (Grijalva-Vallejos
driven the interest in isolation of microorganisms from unconventional et al., 2020). Therefore, interesting results have been reported for the
sources, allowing their application in most diverse products and tech­ isolation of microorganisms from ethnic beverages and application as
nological processes (Maidana et al., 2020; Wuyts et al., 2020). In this starter cultures in the production of the same beverages, aiming to
context, the microbiota of plant origin’s fermented food still represents a replace spontaneous fermentation. This is important, as a previous study
source to be explored and with great potential for new biotechnological has observed that species of opportunistic pathogens (Candida tropicalis,
applications. Table 4 shows the main studies that evaluated the Candida inconspicua, and Kluyveromyces marxianus) could secrete hy­
techno-functional properties of strains isolated from ethnic fermented drolytic enzymes (that contributed to their pathogenicity) in ethnic
beverages and other fermented foods. fermented beverages such as tchapalo and bangui. Therefore, these re­
Olives represent one of the most consumed fermented vegetables sults open discussion about safety of traditional spontaneously fer­
around the world. They are produced through fermentation by their mented beverages (Egue et al., 2018).
native microbiota, which improves the sensory characteristics of the LAB and yeasts from fermented beverages or their metabolites may
products. In this way, studies have investigated the potential of LAB and show antimicrobial activities. L. plantarum SLG10 isolated from kom­
yeasts isolated from table olives for application and improvement of the bucha (traditional fermented beverage based on sugared tea) produced a
fermentation process (Benítez-Cabello et al., 2020; Iorizzo et al., 2016). bacteriocin that was thermostable (storage at 37 ◦ C for 14 days or 2
L. plantarum (B3 and B11) and Lactiplantibacillus pentosus (B4) (former months at 4 ◦ C), tolerant to a pH range of 2.0–7.0, and sensitive to most
Lactobacillus pentosus (B4)) isolated from fermented green olives cv. proteases (except trypsin and pepsin). Furthermore, it showed good
"Nocellara del Belice" have shown positive in vitro results, such as high antimicrobial activity against Gram-positive and Gram-negative

50
T.C. Pimentel et al. Trends in Food Science & Technology 114 (2021) 25–59

Table 4
Main studies that evaluated the techno-functional properties of strains isolated from traditional fermented foods/beverages.
Strain Isolation source Experiments Analysis performed Insightful findings References

LAB strains Tchapalo Identified and Tchapalo Processing Dominant LAB species in Aka et al. (2020)
traditional Ivoirian characterized LAB involved LAB isolation and tchapalo fermentation were
fermented beverage in the tchapalo enumeration L. fermentum (64%), followed
produced from sorghum fermentation process, in Analysis of antimicrobial by Pediococcus acidilactici
order to use them as activity (14%).
potential starter cultures, as Identification of LAB and Of the 465 isolates, 27 were
well as the production of analysis of genomic and capable of producing
bacteriocins by these general phylogenetic bacteriocins.
bacteria characteristics No antibiotic resistance genes
Detection of antibiotic or any potential virulence
resistance genes and factors have been detected.
potential virulence factors L. fermentum S6 and
P. acidilactici S7, and two
potential bacteriocin
producers, Weissella confusa
AB3E41 and Enterococcus
faecium AT1E22, were
selected for characterization.
L. fermentum S6 and
P. acidilactici S7 proved to be
good candidates for starter
cultures
Lactiplantibacillus pentosus Fermented table olives Evaluated the use of strains Inoculum treatment Wickerhanomyces anomalous Benítez-Cabello
LPG1 (former Lactobacillus biofilms (alone or in combination) Physicochemical analysis. Y12 followed by a et al. (2020)
pentosus LPG1), L. pentosus as multifunctional starters Microbiological analysis combination of the LAB
Lp13, Lactiplantibacillus for Manzanilla Spanish- Observation of the strains showed the best
plantarum Lp115 (former style green table olive presence of biofilm in the results (faster drop in pH,
Lactobacillus plantarum fermentations, as well as its fruits higher acid production, sugar
Lpl15), and impact on physical- Self-inducing bioassay-2 consumption, LAB growth,
Wickerhanomyces chemical, Genotyping of the LAB biofilm formation and AI-2
anomalous Y12 microbiological and population and production mainly in the first
sensory parameters. metataxonomic analysis weeks).
Sensory analysis LPG1 strain and, particularly
Concentration: Lp13, were excellent biofilms
LAB: (~7 log cfu/mL) former
Y12: (~5 log cfu/mL) Besides Lactobacillus, high
proportions of sequences
from other genera such as
Marinilactobacillus,
Alkalibacterium,
Halolactobacillus and low
levels of Halomonas and
Aerococcus were observed.
There were no differences
between the inoculated
treatments and the non-
inoculated (spontaneous)
control in the sensory
analysis.
Yeasts strains Ecuadorian Chichas It isolated, identified and Molecular yeast Eleven yeast genera and 16 Grijalva-Vallejos
(indigenous Andean beer) characterized yeast strains identification species were identified et al. (2020)
produced with: in order to select initiators Stress tolerance analysis Stress tolerance varied
Rice (RC), for application in the Detection of amylase between species
Oats (OC), fermentation process of activity One Cryptococcus sp. isolate
Grape (GC) and Chicha or other fermented Growth in molasses and had the highest amylase
A mixture of seven varieties foods, and biochemical and tolerance to dehydration activity.
of corn (yamor, YC) molecular aspects during Analysis of fermentative Ecuadorian strain of
fermentation were also capacity S. cerevisiae EYS5 exhibited
evaluated. Determination of invertase excellent fermentative
activity performance, invertase
activity (1.33 times greater
than the control) and showed
high viability after
dehydration.
T. delbrueckii EGT1 also
showed high potential as
non-Saccharomyces yeast for
alcoholic fermentations.
Thirty-four LAB strains Kimchi It selected LAB strains Identification of selected Six BAL identified as Hossain et al.
isolated from kimchi with anti-listerial LAB isolates L. plantarum (I.60, M.2 and (2020)
anti-listeria capacity and Analysis of cell surface M.21), L. curvatus (B.67),
investigated the cell aggregation and L. sakei (D.7) and Leuconostoc
surface properties (self- hydrophobicity activity mesenteroides
aggregation and Inhibition evaluation of (J.27) have shown anti-
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T.C. Pimentel et al. Trends in Food Science & Technology 114 (2021) 25–59

Table 4 (continued )
Strain Isolation source Experiments Analysis performed Insightful findings References

hydrophobicity) as well as Listeria monocytogenes listerial capability.

the inhibitory effect of biofilm activity in vitro, in All isolates exhibited
isolates against biofilm stainless steel coupons remarkable auto-aggregation
formation of (SS), in lettuce and in the and hydrophobicity
L. monocytogenes in lettuce MBEC ™ biofilm device. potential.
and MBEC™ biofilm Characterization analyzes LAB strains suppressed
device. of biofilm formation L. monocytogenes biofilms by
Concentration: up to 2.17 log cfu/cm2, 1.62
LAB: (8 log cfu/mL) log cfu/cm2 and 1.09 log cfu/
L. monocytogenes: (5 log peg in SS, lettuce and MBEC
cfu/mL) ™, respectively.
L. curvatus B.67 isolate was
the one with the highest
inhibition efficiency.
13 yeast strains: Kefir and Caxiri (indigenous It evaluated the Co-aggregation evaluation All strains were able to co- Menezes et al.
12 S. cerevisiae beverage) antimicrobial effect of Analysis of Scanning aggregate with the pathogens (2020)
P. kluyveri Cocoa fermentation potentially probiotic yeast electron microscopy (SEM) tested, however, this activity
strains, previously isolated of yeasts co-aggregated was dependent on the strain.
from fermented foods, on with bacterial pathogens All yeasts were effective in
adhesion of foodborne Analysis of inhibition of reducing adhesion and
pathogens to Caco-2 cells. pathogenic bacteria bacterial infection (over
adhesion to Caco-2 50%) in caco-2 cells.
Analysis of Antimicrobial As observed for P. kluyveri
activity (CCMA 0615) for EPEC in an
Concentration: exclusion test; S. cerevisiae
Yeast: (7 log cfu/mL) (CCMA 0731, CCMA 0732)
L. monocytogenes, EPEC e S. and P. kluyveri (CCMA 0615)
Enteritidis: (9 log cfu/mL) for L. monocytogenes in
exclusion and competition
tests; and S. cerevisiae (CCMA
0731) in exclusion and
S. cerevisiae (CCMA 0731,
CCMA 0732), P. kluyveri
(CCMA 0615) in a
competition trial for
S. Enteritidis.
Yeast antimicrobial
compounds have not been
produced
L. plantarum 332, Spontaneously fermented It selected LAB isolates as Curly kale juice L. paraplantarum G2114, Lb. Michalak et al.
Lactiplantibacillus curly kale starter cultures for the fermentation plantarum 332 and (2020)
paraplantarum G2114 fermentation of kale juice, Technological and Pediococcus pentosaceus 2211
(former Lactobacillus evaluating the functional functional potential showed better performance
paraplantarum G2114), and technological analysis of fermented curly in the evaluation of
Levilactobacillus brevis R413 characteristics of the kale juices (pH and total functional and technological
(former Lactobacillus brevis fermented juice from acidity, determination of properties by PCA and were
R413), Latilactobacillus multistrain formulations. viable cells, neutral sugar selected as starter cultures.
curvatus 154 (former and protein Kale juice fermented by the
Lactobacillus curvatus 154), content, antioxidant combined use of these strains
Weissella hellenica 152, activity (ABTS), gentisic showed rapid acidification
W. cibaria G44, and with a high number of viable
Pediococcus pentosaceus salicylic acid (HPLC) cells, increased antioxidant
2211, P. acidilactici 45AN, content) activity and antimicrobial
Leuconostoc mesenteroides Antibacterial activity test effects against some
153, and Lactococcus lactis PCA analysis for starter pathogenic bacteria tested
37BN culture selection (mainly Gram negative).
Analysis of technological
and functional
characteristics of
fermented juice with
starter culture
Analysis of the total
phenolic content (TPC),
Cell viability in cold
storage (4 ◦ C for 4 weeks)
Concentration:
LAB: (~7.5 log cfu/mL)
(5% v/v)
Salmonella enterica,
Staphylococcus aureus and
Escherichia coli: Briefly,
0.5 mL of inoculum
overnight (OD600 = 1.0)
L. plantarum SLG10 Traditional kombucha of It isolated an antimicrobial Isolation and identification SLG10 strain was selected Pei et al. (2020)
South China strain of Lactobacillus that of bacteriocinogenic LAB because it is the only strain
produces bacteriocin from Evaluation of bacteriocin capable of acting on both
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T.C. Pimentel et al. Trends in Food Science & Technology 114 (2021) 25–59

Table 4 (continued )
Strain Isolation source Experiments Analysis performed Insightful findings References

kombucha, in addition to production by the SLG10 Gram-positive and Gram-

tracking and purifying the strain negative bacteria (S. aureus
bacteriocin Bacteriocin purification CICC10384 and E. coli
produced, demonstrating analyzes CICC10302).
its mechanisms of action. Structural characterization The maximum production of
analyzes of bacteriocin bacteriocin SLG10 was
Bacteriocin stability recorded after 30 h of growth
analysis in MRS broth.
Antimicrobial activity of Bacteriocin SLG10 showed
bacteriocin SLG10 thermostability after storage
Analysis of inhibition of at 37 ◦ C for 14 days or even
biofilm formation after 2 months at 4 ◦ C, pH
Concentration: tolerance characteristics in
Indicator bacteria: (6 log the range of 2.0–7.0, and was
cfu/mL) sensitive to most proteases,
S. aureus CICC 10384: (8 but not to trypsin or pepsin.
log cfu/mL) Bacteriocin SLG10 proved to
be bactericidal, increased the
permeability of the cell
membrane, causing the
release of potassium ion and
was also able to inhibit the
formation of biofilms.
Six-teen strains: Kombucha (pellicle and It evaluated the capacity of Microbiological analysis Kombucha showed a Taheur et al.
LAB (7) and Yeasts (9) fermented tea) bacteria and/or yeasts Ability of kombucha for degradation capacity of 97% (2020)
isolated from Kombucha to degrading and adsorbing of AFB1 in black tea after 7
biodegrade Aflatoxin B1 mycotoxin from days of fermentation.
(AFB1), as well, as the contaminated tea In vitro degradation tests
cytotoxicity of the by- Evaluation of AFB showed that LAB degraded
products in Hep2 cells and biodegradation properties 20–45% of AFB1, while
in brine shrimp (Artemia of strains isolated from isolated yeasts showed no
salina). kombucha degradation capacity.
Identification of active In black tea, yeasts showed
strains degrading AFB1 better degradation potential
Mycotoxins contents of AFB1 (37%).
analysis (HPLC). Pichia occidentalis, Candida
Cytotoxicity test for AFB1 sorboxylosa and
on Hep2 cells and brine Hanseniaspora opuntiae have
shrimp. been identified as yeasts
Concentration: effective in degrading AFB1
LAB: (~9 log cfu/mL) P. occidentalis (59%) showed
Yeasts: (~7 log cfu/mL) the greatest degradation
capacity of AFB1 when grown
in black tea.
Biodegraded products were
less toxic than pure AFB1,
both in the cytotoxicity test in
Hep2 cells and in brine
shrimp.
LAB strains Fruits and vegetables: It isolated and identified Isolation of LAB 54 strains of LAB were Barrios-Roblero
Ilama (Annona LAB with antifungal Antifungal assays isolated, of these, 50 were et al. (2019)
macroprophyllata), papaya activity against Identification of LAB from fermented foods.
(Carica papaya L.), mango Colletotrichum Concentration: 18% of the isolates showed
(Mangifera indica L. var. gloeosporioides from LAB: not specified strong antifungal activity
Ataulfo), cocoa (Theobroma vegetables and fermented C. gloeosporioides Penz: (1 against C. gloeosporioides,
cacao L.), cabbage (Brassica foods. × 106 spores/mL.) inhibiting both spore
oleracea var. Capitata), germination and mycelial
melon (Cucumis melo L.), growth.
sapodilla (Pouteria sapota L.) The inhibition of spore
and sabila (Aloe vera L.) germination varied between
Fermented foods: 50 and 100% (inhibition
Pozol (traditional drink made range between 30 and 40
from mm)
corn and ground cocoa mass) All isolates presented were of
Tepache (fermented drink the genus Lactobacillus
made with fimxi), The isolated strains
Tejuino (fermented corn L. pentosus TEJ10, L. paracasei
drink) and TEP6 and L. plantarum TEP15
Sauerkraut (fermented showed the greatest
cabbage). inhibition of spore
germination of the
phytopathogen.
All strains inhibited the
growth of the mycelium of
C. gloeosporioides, surpassing
the
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T.C. Pimentel et al. Trends in Food Science & Technology 114 (2021) 25–59

Table 4 (continued )
Strain Isolation source Experiments Analysis performed Insightful findings References

positive controls (natamycin

and mancozebe)
L. plantarum LUHS135 and Spontaneously fermented It isolated strains of LAB Purification, isolation, L. plantarum LUHS135 and Bartkiene et al.
Lacticaseibacillus paracasei Rye flour from spontaneous identification and L. paracasei LUHS244 were (2018)
LUHS244 (former fermentation of rye flour characterization analyzes able to metabolize a wide
Lactobacillus. paracasei and evaluated their Analysis of antimicrobial spectrum of carbohydrates,
LUHS244) technological potential activity and antibiotic showed tolerance to acidity
regarding carbohydrate resistance (viable cells: 7.69 and 7.55
metabolism, gas Analysis of mycotoxin log10 cfu/mL - 1) and
production, low pH reducing properties susceptibility to antibiotics
survival, growth at Evaluation of growth and (gentamycin, tetracycline,
different temperatures, encapsulation in whey erythromycin, amoxicillin
antimicrobial properties, substrate and trimethoprim).
antibiotic resistance, Concentration: L. plantarum LUHS135 and
mycotoxin reducing LAB: (~7–8 log cfu/mL) L. paracasei LUHS24 inhibited
properties, as well as the growth of a wide range of
application in encapsulated pathogenic bacteria and
dairy by-products. showed no gas production.
The highest antimicrobial
activity was found for
L. plantarum LUHS135
against B. cereus
Candida strains Tchapalo and bangui (or palm It evaluated the diversity of Microbiological Analysis A total number of 136 yeasts Egue et al. (2018)
. wine) traditional beverages Candida species in popular Molecular identification (64 of tchapalo 72 of bangui)
alcoholic drinks from Côte tests of yeast isolates were identified, belonging to
d’Ivoire, as well as the Analysis of enzyme seven species: Candida
enzymatic activity of these activity tropicalis, Candida
strains. inconspicua, Candida rugosa,
Saccharomyces
cerevisiae, Kluyveromyces
marxianus, Hanseniaspora
guilliermondii, Trichosporon
asahii (7 log cfu/mL).
Three species of Candida (C.
tropicalis, C. inconspicua and
C. rugosa). were isolated from
beverages.
C. rugosa was specific to
bangui while T. asahii was
specific to tchapalo.
Four species were found to be
opportunistic pathogens
(C. tropicalis, C. inconspicua
and K. marxianus).
The pathogens evaluated
showed a variable amount of
secreted hydrolytic enzymes,
such as esterase, esterase
lipase, valine and cystine
arylamidase, alpha
chymotrypsin, alkaline
phosphatase and naphthol
phosphohydrolase.
Komagataeibacter strains Kombucha beverages Evaluated four different Isolation analysis and The best cellulose producers Semjonovs et al.
commercial Kombucha for identification of cellulose- were the B17 and B22 isolates (2017)
isolating strains with the producing strains belonging to the genus
potential to produce Characterization of BC Komagataeibacter (former
bacterial cellulose (BC) (FT-IR) Gluconacetobacter) and
under static cultivation BC production and classified as K. rhaeticus P
conditions using different quantification tests 1463 and K. hansenii B22,
sugars and sugar-alcohol Determination of sugars respectively.
mannitol as carbon sources, and acidic organics K. rhaeticus P 1463 was more
as well such as apple juice Determination of the effective with BC yield of
and cheese whey as low- degree of cellulose ~25% in Hestrin and
cost substrates. In addition, polymerization Schramm (HS) medium and
it also evaluated the Evaluation of the ~37% in fermented apple
physical and mechanical mechanical properties of juice for 5 days.
properties of the cellulose BC Synthesized BC showed good
produced. physical and mechanical
properties
L. plantarum (B3, B11) and Fermented Green Olives cv. It evaluated the Determination of bacterial All strains showed high cell Iorizzo et al.
L. pentosus (B4) “Nocellara del technological potential in viability and fermentative viability even with no glucose (2016)
Belice” vitro of Lactobacillus strains metabolism up to 15 days.
previously isolated from Determination of the Acidification capacity
fermented olives in terms of content of glucose, total (degradation of all glucose in
multiplication, phenolics, Oleuropein and 24 h, final pH: ~4.1),
(continued on next page)

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T.C. Pimentel et al. Trends in Food Science & Technology 114 (2021) 25–59

Table 4 (continued )
Strain Isolation source Experiments Analysis performed Insightful findings References

acidification capacity, hydroxytyrosol L. plantarum B3 and B11

enzymatic activities and Analysis of enzyme exhibited good adaptability
oleuropein degradation activity to high concentrations of salt
capacity under different Analysis of oleuropein- (up to 10% NaCl), low
conditions. degrading capability and temperature (7 ◦ C) and pH
β-Glucosidase (about 3.6).
activity Good tolerance to the phenol
content of common olives
(0.3% of total phenol) and
high degradability of
oleuropein.
Lesser action of degradation
of oleuropein was observed in
a medium rich in nutrients
All strains evaluated
produced β-glucosidase
activity, but without esterase
Komagataeibacter intermedius Fermented fruit juices of It isolated and identified a Preparation of fermented The isolated strain was Lin et al. (2016)
FST213-1 pineapple, apple and guava new strain producing fruit juices identified as
bacterial cellulose (BC) in Isolation of BC producing Komagataeibacter intermedius
fermented fruit juices, still strain FST213-1.
evaluated the production of Identification tests of BC K. intermedius FST213-1 was
BC at different pHs and producing strain able to produce BC at pH
characterized the BC Material property analysis values between 4 and 9,
produced. exhibiting maximum BC
production (1.2 g/L) at pH 8
in short term cultivation (4
days).
BC produced from
K. intermedius FST213-1
exhibited higher water
content (99.5%), lower
thermostability (315 ◦ C),
lower crystallinity (79.3%)
and similar mechanical
properties compared to the
producer BC,
Gluconacetobacter xylinus
23769.
LAB and Tarubá (traditional Identified and Microbial population and LAB and Bacillus species such Ramos et al.
yeasts strains fermented beverage characterized the characterization (by as L. plantarum, L. brevis, (2015)
produced from cassava) microbiota present during culture-dependent and Leuconostoc mesenteroides and
solid state fermentation in -independent methods) Bacillus subtilis were
the final stage of tarubá Analysis of amylolytic and predominant during
production, as well as the proteolytic activity fermentation.
chemical characteristics Thermotolerance analysis Torulaspora delbrueckii
throughout and at the end Analysis of chemical species was the dominant
of the fermentation of parameters yeast during fermentation.
beverage. Determination of sugars, Amylolytic, proteolytic and
organic acids and ethanol thermotolerance properties
by HPLC were observed for some
isolates.
T. delbrueckii, P. exigua and
Bacillus spp. exhibited all of
these properties.

bacteria (S. aureus CICC10384 and E. coli CICC10302) and inhibited the life.
formation of biofilms. Therefore, this bacteriocin could be used as an The capacity of microorganisms isolated from traditional beverages
antimicrobial compound in the food industry (Pei et al., 2020). Lacti­ for degradation of toxins has also been reported. Kombucha culture and
plantibacillus pentosus TEJ10 (former Lactobacillus pentosus TEJ10), isolates (Pichia occidentalis, Candida sorboxylosa and Hanseniaspora
L. paracasei TEP6 and L. plantarum TEP15 isolated from fermented opuntiae) could degrade aflatoxin B1 in black tea, decreasing its toxicity
foods/beverages (pozol, tepache, tejuino and sauerkraut) showed in vitro in both Hep2 and artemia cells. Aflatoxin B1 is recognized as the most
antifungal activity against phytopathogenic strain Colletotrichum gloeo­ harmful mycotoxin and its presence in tea makes it unsuitable for con­
sporioides Penz, being able to inhibit both spore germination (between sumption due to the risks to human health. Therefore, kombucha and its
50 and 100%) and mycelial growth (surpassing the positive controls strains can be applied in feed and food industries to detoxify aflatoxin B1
natamycin and mancozebe) (Barrios-Roblero et al., 2019). LAB isolated (Taheur et al., 2020).
from “chicha” (a traditional maize-based fermented beverage) and LAB and yeasts from fermented beverages may also be used to pro­
“tocosh” fermented potatoes exhibited antimicrobial capacity against duce desirable compounds. Four different commercial kombuchas and
fungi (A. oryzae and M. guilliermondii) and food pathogens (E. coli O157: the isolated strains (Komagataeibacter rhaeticus P 1463 and Komaga­
H7, L. innocua, and Salmonella Typhi) (Yépez et al., 2017). Therefore, the taeibacter hansenii B22) showed a strong capacity to produce cellulose
antimicrobial activity of LAB and yeasts isolated from fermented foods with good physical and mechanical properties both in vitro (~25%) and
may increase the plant resistance to pathogens and food products’ shelf in fermented apple juice for 5 days (~37%) (Semjonovs et al., 2017).

55
T.C. Pimentel et al. Trends in Food Science & Technology 114 (2021) 25–59

Komagataeibacter intermedius FST213-1 isolated from fermented fruit individuals, suggesting that studies covering new probiotic species
juices (pineapple, apple, and guava) exhibited ability to produce bac­ (yeasts and other LAB) from other sources (flowers, vegetables, and
terial cellulose (BC) (at pH between 4 and 9) with desirable techno­ ethnic beverages) and different types of hosts and diets (diabetics,
logical properties (water content capacity (99.5%), low thermostability healthy, elderly) are also needed. This highlights the range of study
(315 ◦ C), less crystallinity (79.3%) and good mechanical properties) (Lin opportunities in the area. The probiotic efficacy was strain-specific,
et al., 2016). Bacterial cellulose has many advantages compared to plant emphasizing the importance of having a catalog of probiotic strains
cellulose, such as higher crystallinity, mechanical strength, biocompat­ with information about their health effects, which would assist the in­
ibility, biodegradability, and purity. It has many applications, such as dustries in developing new functional products aiming specific human
biosensor, paper production, and tissue engineering (Lin et al., 2016). health promotion (Oliveira, Cavalcante, et al., 2020). Furthermore, it is
LAB and yeast from fermented foods may also be used to improve important to carry out the complete phenotypic and genotypic identi­
food products’ nutritional value (Fekri et al., 2020). Yépez et al. (2019) fication of the isolated probiotic cultures and not only demonstrate the
observed that LAB (L. plantarum M5MA1-B2) isolated from Andean health effects (Mabeku et al., 2020).
improved riboflavin content in kefir-like cereal-based beverages (ob­ In conclusion, fruits, vegetables, flowers, and ethnic fermented
tained from maize, oat, and barley flours). beverages are important sources of techno-functional and probiotic
Previous studies also reported the isolation of microorganisms from cultures with high potential for use in food processing to increase the
fermented foods that presented multiple functions. L. plantarum and nutritional and sensory quality, acting as natural biopreservatives, and
L. paracasei LUHS244 isolated from spontaneous fermentation of rye promote health effects. The utilization of these cultures may enhance the
flour were able to metabolize a wide spectrum of carbohydrates, development of vegan probiotic products and non-dairy fermented
inhibited the growth of a wide range of pathogenic bacteria, and beverages, allowing processing at lower times and resulting in products
reduced mycotoxins (3% of each inoculum) in in vitro tests. In addition, with more consistent characteristics. Furthermore, it opens opportu­
they showed good stability and multiplication when encapsulated in nities to isolate important microorganisms from available sources in
whey (enriched with 2.5% glucose, 2.0% yeast extract and 0.5% su­ each country, such as ethnic beverages or traditionally fermented foods.
crose) (Bartkiene et al., 2018). Lactiplantibacillus paraplantarum G2114
(former Lactobacillus paraplantarum G2114), L. plantarum 332 and Ped­ Declaration of competing interest
iococcus pentosaceus 2211 isolated from spontaneously fermented kale
showed rapid acidification, and high viability, antioxidant, and anti­ Authors declare no conflict of interest.
microbial activity (against Gram positive and Gram-negative bacteria).
These characteristics were maintained in fermented cabbage juice dur­ Acknowledgments
ing storage (4 ◦ C) for 4 weeks when the combination of these strains was
used as starter cultures (~7.5 log cfu/mL) (5% v/v) for fermentation This work was supported by the “Coordenação de Aconselhamento
(Michalak et al., 2020). de Pessoal de Nível Superior” (CAPES-Brazil; Finance code 001),
“Fundação de Apoio à Pesquisa do Estado da Paraíba” (FAPESq) and
4. Conclusions and future perspectives Conselho Nacional de Desenvolvimento Científico e Tecnológico
(CNPq).
This was the first review to explore the added-value microorganisms
isolated from unconventional sources (fruit, raw vegetables, ethnic References
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