Critical Reviews in Food Science and Nutrition

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Bio-functional properties of probiotic Lactobacillus:

current applications and research perspectives

Jagrani Minj, Priyanka Chandra, Catherine Paul & Rakesh Kumar Sharma

To cite this article: Jagrani Minj, Priyanka Chandra, Catherine Paul & Rakesh Kumar
Sharma (2020): Bio-functional properties of probiotic Lactobacillus: current applications
and research perspectives, Critical Reviews in Food Science and Nutrition, DOI:
10.1080/10408398.2020.1774496

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Published online: 10 Jun 2020.

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CRITICAL REVIEWS IN FOOD SCIENCE AND NUTRITION
https://doi.org/10.1080/10408398.2020.1774496

REVIEW

Bio-functional properties of probiotic Lactobacillus: current applications and

research perspectives
Jagrani Minja, Priyanka Chandrab, Catherine Paula, and Rakesh Kumar Sharmac
a
Department of Food Science and Technology, Nebraska Innovation Campus (NIC), University of Nebraska, Lincoln, Nebraska, USA; bICAR-
Central Soil Salinity Research Institute, Karnal, India; cDepartment of Biosciences, Manipal University Jaipur, Jaipur, Rajasthan, India

ABSTRACT KEYWORDS
Lactic acid bacteria as a starter culture are very important component in the fermentation process Bioactive metabolites; lactic
of dairy and food industry. Application of lactic acid bacteria as probiotic bacteria adds more func- acid bacteria;
tionality to the developed product. Gut colonizing bacteria have attractive benefits related to Lactobacillus; probiotics
human health. Bio-functional properties such as antimicrobial activity, anti-inflammatory, ACE-
inhibitory, antioxidant, antidiarrheal, antiviral, immunomodulatory, hypocholesterolemic, anti-dia-
betic and anti-cancer activities are the most applicable research areas of lactic acid bacteria.
Different strains of Lactobacillus are generally consumed as probiotics and colonize the gastro-
intestinal tract. Sometimes these bacteria may possess antimicrobial activity and may positively
influence the effect of antibiotics. Use of Lactobacillus spp. for the development of functional
foods is one of the promising areas of current research and applications. Individual bacterial spe-
cies have unique biological activity, which may vary from strains to strains and identification of
this uniqueness could be helpful in the development of functional and therapeutic food products.

Introduction generation probiotics (O’Toole, Marchesi, and Hill 2017). As

Lactic acid bacteria (LAB) group are known to play an
important role in the fermentation processes in small house-
holds, food and dairy industry. Among different LAB,
Lactobacillus are mainly used for the production of lactic
acid but these are also responsible for the production
organic acids, gas, flavor, aroma and texture development in
the food and dairy products. These bacteria are also known
to boost therapeutic and health benefits to the consumers.
The health promoting properties of any LAB is related to its
probiotic characteristics. Definition of probiotics is “live
microorganisms which when administered in adequate
amounts confer a health benefit on the host”. Probiotic bac-
teria enters into the human gastrointestinal tract and adhere
to the intestinal epithelial cells and shows various beneficial
biological activities, like competitive adhesion to the epithe-
lial layer and production of small antimicrobial molecules
and peptides that suppress the harmful bacteria, stimulation
of various immunomodulatory cytokines for the balance of
pro-inflammatory and anti-inflammatory cytokines, produc-
tion of anti-tumor compounds, bacteriocins (glycoproteins,
which possesses antimicrobial and bactericidal activity) etc.
Positive health effects of probiotics in the combination of
prebiotics (collectively known as synbiotics) have also been
reported (Markowiak and
Slizewska 2017). Such formula-
tions may positively influence the growth of bacteria in
human intestine. Probiotic bacteria with some additional
properties have also been explored and developed as next
the research is progressing, different related concepts includ-
ing paraprobiotics (inactivated probiotics), postbiotics (bio-
active soluble factors: produced by bacterial metabolism or
released from cell rupture) and psychobiotics (probiotics,
which provide mental health benefits) have also been intro-
duced (Barros et al. 2020). However, the role of lactobacilli
in all these preparations may be well correlated.
Lactobacillus species are widely used in the dairy prepara-
tions because of its efficient fermentation ability and related
health benefits (Tachedjian et al. 2017). On food processing
point of view, the role of Lactobacillus is not limited to fer-
mentation but also deals with therapeutic aspects.
Consumers’ awareness is steadily increasing about the health
benefits of fermented food, thus, numerous functional foods
with probiotics (i.e., fermented foods containing probiotic
microbes as starters or adjuncts culture) are readily available
in the market. Therapeutic properties of Lactobacillus are
regulated by the metabolites produced during fermentation,
which posses unique functional properties (Plaza-Diaz et al.
2019). Some bioactive peptides having a positive influence
on human health are either produced through degradation
of food proteins during food processing or food digestion.
Applications of enzymes e.g. phytases from LAB in food
processing industries to enhance the micronutrient uptake
have been well explored (Sharma, Chandel, et al. 2020).
Numerous studies have been conducted targeting the pro-
duction and physiological actions of bioactive peptides, i.e.

CONTACT R. K. Sharma rakeshkumar.sharma@jaipur.manipal.edu Department of Biosciences, Manipal University Jaipur, Jaipur 303007, Rajasthan, India;
Priyanka Chandra priyanka.chandra921@gmail.com ICAR-Central Soil Salinity Research Institute, Karnal, India
ß 2020 Taylor & Francis Group, LLC
2 J. MINJ ET AL.
antioxidants, antimicrobial, antithrombotic and immuno-
modulatory activities (Chandra and Vij 2018; Chandra,
Sharma, and Singh Arora 2020). Thus, this comprehensive
review includes the compilation of bio-functional properties
of common Lactobacillus spp., which are used as probiotics
or as starter cultures in the dairy and other food products.
Some of the Lactobacillus species used as probiotics in dairy
products are discussed in detail and enlisted in Table 1.
Lactobacillus acidophilus
Lactobacillus acidophilus is a normal flora of intestine,
mouth and vagina in humans. According to several research
findings, it provides various health benefits to the host by
the production of a variety of vitamins and enzymes
(Akolkar, Sajgure, and Lele 2005). Enzyme lactase has been
known for the metabolic breakdown of lactose sugar into
glucose and galactose. These broken smaller sugar molecules
are easily absorbed by the human intestinal cells.
Additionally, the enzyme lactase may acts as a therapeutic
agent for the lactose intolerance people. L. acidophilus is
used in the making of yoghurt (as an additional culture),
dahi, soy based fermented products like miso and tempeh
and some fermented dairy products in food and dairy indus-
try. It can either be used as a single organism or in the com-
bination with one or more other microbes as per the
requirement of the final products. Gut LAB L. acidophilus
Pul13_14 may breakdown the starchy branched a-glucan
oligomers into short a-1, 6-branches. These starch glycans,
branched a-1, 6-glucans and glycogen act as a pool of meta-
bolic resources (prebiotics) for the intestinal microorganisms
because these are not degraded by the human enzymes
(Møller et al. 2017).
Probiotic L. acidophilus NCFM exhibited adherence prop-
erty to the intestinal epithelial cells (Celebioglu et al. 2017).
Therapeutic application of L. acidophilus has been mostly
studied as antimicrobial, antiviral and toward eradication of
diarrhea. L. acidophilus has also been very active against
rotaviruses, enterobacteriaceae and candida infection.
Rotaviruses are the main cause of acute gastroenteritis and
diarrheal conditions mainly in children under the age of
5 years. Research work has been carried out on the preven-
tion of rotavirus diarrhea using many probiotic bacteria,
among them L. acidophilus has demonstrated its potential to
inhibit infection of rotavirus under in vitro conditions. It
also decreased the duration of diarrhea in the pediatric
patients. The synergistic inhibitory action of probiotic bac-
teria L. acidophilus and Bifidobacterium longum against rota-
virus infections has also been reported (Lee et al. 2015).
Antimicrobial activity of L. acidophilus is governed by the
production of bacteriocins like acidocin, acidophilin, acido-
philicin, acidophilucin, lactacins B, F and some organic acids
(Dinev et al. 2018). The antimicrobial activity of L. acidoph-
ilus against Listeria monocytogenes, Helicobacter pylori,
Escherichia coli, Staphylococcus aureus, Pseudomonas aerugi-
nosa, Salmonella, Shigella and Bacillus species has already
been confirmed. Helicobacter pylori infection has been
treated with the probiotic supplementations but some of the
contradictory results were also reported while using a com-
mercial probiotic preparation including L. acidophilus, L.
rhamnosus and L. sporogenes, which revealed that these pro-
biotics were not as effective as the antibiotics used for eradi-
cation of H. pylori infection (Lee, Shih, et al. 2017). On the
other hand, inhibitory action of L. acidophilus and L. planta-
rum against multidrug-resistant enteroaggregative
Escherichia coli (MDR-EAEC) increased when all these
microbes were co-cultured. The synergistic antimicrobial
effect of both the probiotic bacteria was significantly higher
as compared to the individual probiotic bacteria (Kumar
et al. 2016). Probiotic L. acidophilus ATCC 4357 played a
key role in mediating the electro-neutral NaCl absorption in
the intestinal epithelial luminal membrane NHE3 (Naþ/Hþ
exchanger 3) and DRA (Down-regulated in Adenoma; Cl/
HCO3 exchanger). L. acidophilus ATCC 4357 thus counter-
acted the inhibition of NHE3 and DRA expression in the
mice infected with Citrobacter rodentium (Kumar
et al., 2016).
Production of short chain fatty acids by L. acidophilus
ATCC 4357 also enhanced its functional property. A short-
chain fatty acid butyrate was produced by this bacterium,
which served as a major substrate for colonocytes and
reduced mucosal inflammation, and also stimulated NaCl
absorption. L. acidophilus ATCC 4357 significantly increased
Monocarboxylate transporter 1 (MCT1) mediated butyrate
uptake in Caco-2 cells. Furthermore, the culture supernatant
of L. acidophilus ATCC 4357 enhanced the apical membrane
levels of MCT1 protein by decreasing its basal endocytosis.
It also attenuated enteropathogenic E. coli (EPEC) mediated
endocytosis of MCT1 and inhibition of butyrate uptake
(Kumar et al. 2015).
Anti-candidal effects of probiotic L. acidophilus have
been reported earlier that could be a good choice for the
treatment of candidal infections (Matsubara et al. 2016).
EPS producing L. acidophilus strains are particularly useful
in the yoghurt production. Co-culture of Bifidobacterium
animalis subsp. lactis and L. acidophilus La5 produced EPS,
which may be directly used as natural antioxidants in the
food industry (Amiri et al. 2019). Anti-diabetic potential of
probiotic yoghurt containing L. acidophilus La5 and
Bifidobacterium lactis Bb12 have been reported to improve
fasting blood glucose in patients (300 g/day) with type 2 dia-
betes and it also improved the antioxidant enzyme level
(Ejtahed et al. 2012).
Lactobacillus casei
Lactobacillus casei is commonly found in human intestine
and considered as safe probiotic bacteria for the consump-
tion. L. casei is a dominant species of nonstarter LAB gener-
ally used during the ripening process of cheddar cheese. L.
casei also possesses numerous health promotional and pro-
tective activities. Recently, a significant increase in different
biological activities such as anti-cancer, antioxidant and
angiotensin converting enzyme inhibitory activities were
reported during the storage of probiotic L. casei ATCC 393
(Abdel-Hamid et al. 2019). Cytotoxic activity against various
 CRITICAL REVIEWS IN FOOD SCIENCE AND NUTRITION 3

Table 1. Bio-functional properties of different Lactobacillus strains.

Starter culture /Used in Used dose/initial
dairy products/ concentrations and
S. No. Lactobacillus strain Food Products Biological activities time duration References
1. L. acidophilus NA Antibacterial activity, 5  107 CFU (48 h of (Shao et al. 2017)
ATCC 4356 Cholesterol lowering ability fermentation)
2. L. acidophilus NA Anti-tumor and immune 108 CFU (72 h of fermentation) (Hsieh et al. 2016)
BCRC 14079 modulatory activity
3. L. acidophilus W37 NA Antimicrobial activity 109 CFU (24–48 h of (Campana et al. 2017)
fermentation)
4. L. acidophilus La NA Antioxidant activity 3 days of fermentation (Lin and Chen 2017)
5. L. acidophilus NCFM NA Antioxidant activity 5  108 CFU/mL (6 h of (Hougaard et al. 2016)
fermentation)
6. L. acidophilus NCDC-15 Fermented milk (Lassi) ACE-inhibitory activity Cell free supernatant (12 h of (Padghan et al. 2017)
fermentation)
7. L. acidophilus LA-5 Iranian fermented Decontamination of 107 CFU/mL (28 d) (Sarlak et al. 2017)
milk drink Aflatoxins M1
8. L. acidophilus NA Maintenance of 103–106 CFU (3-9 d) (Azevedo et al. 2012)
ATCC NCFM immunological homeostasis
9. L. acidophilus NCC90 Synbiotic yoghurt Effect on bone mineralization 1–5  106 CFU/g (7 d) (Scholz-Ahrens
et al. 2016)
10. L. delbrueckii ssp. NA Antioxidant property EPS fraction (48 h of (Tang et al. 2017)
bulgaricus SRFM-1 fermentation)
11. L. delbrueckii ssp. NA Anti-biofilm Activity 107–108 CFU/mL (48 h of (Sarikaya, Aslim, and
bulgaricus A-12 fermentation) Yuksekdag 2017)
12. L. delbrueckii ssp. NA Antiproliferative activity 108 CFU/mL (28 d) (Sah et al. 2016)
bulgaricus Lb1466
13. L. bulgaricus Yogurt Immune stimulating activity 109 CFU (18 h of fermentation) (Makino et al. 2016)
OLL1073R-1
14. Lactobacillius casei SB27 NA Antitumor activity 1.5  109 CFU/mL (EPS (Di et al. 2017)
fraction,36 h of
fermentation)
15. L. casei KNE-1 Fermented milk Thiamine and 0to 24 d
et al. 2015)
(Drywien
Riboflavin producers
16. L. casei CRL 431 Fermented milk Anti-tumor activity 1.2  10 CFU/day (2 d)
9
(Bonet et al. 2005)
17. L. casei strain Shirota Fermented milk Anti-hypertensive activity 0.9–40  109CFU (3 times (Aoyagi et al. 2017)
per week)
18. L. casei PRA205 Parmigiano Probiotic property 109 CFU/mL (10 months of (Solieri et al. 2014)
Reggiano cheese fermentation)
19. L. fermentum LB-69 NA Anti-biofilm activity 107–108 CFU/mL (48 h of (Sarikaya, Aslim, and
fermentation) Yuksekdag 2017)
20. L. fermentum KJ23 Fermented stinky bean Cell surface 109 CFU/mL (24 h of (Jampaphaeng, Cocolin,
(Sataw-Dong) hydrophobicity activity fermentation) and Maneerat 2017)
21. L. fermentum Fermented melinjo flour Antioxidant and DNA damage 108 cells/mL (48 h of (Siswoyo, Ardyati, and
protection activity fermentation) Hosokawa 2017)
22. L. fermentum I5007 Fermented milk Alteration of intestinal 6  109 CFU/day (14 days) (Liu et al. 2014)
microbial composition,
especially reduction in
Clostridium spp.
23. L. fermentum I5007 NA Antioxidant and anti- 5  109 CFU/day (7 days) (Hou et al. 2014)
inflammatory activity
24. L. fermentum I5007 NA Alleviation of oxidative stress 20 ml of 108 CFU/mL (21 d) (Wang et al. 2013)
25. L. helveticus MTCC5463 Swiss cheese, Parmesan Proteolytic activity 107 CFU/mL (12 h of (Hati, Sakure, and
cheese, Romano fermentation) Mandal 2017)
cheese, Provolone
cheese and
Mozzarella cheese
26. L. helveticus IMAU80851 Traditional fermented ACE-Inhibitory activity 106 CFU/mL (18 h of (Chen et al. 2015)
dairy product fermentation)
27. L. helveticus KLDS1.8701 Fermented Antifungal activity 107 CFU/mL (24 h of (Bian et al. 2016)
soybean milk fermentation)
28. L. helveticus MB2-1 Fermented whey Antioxidant activity EPS fractions (Li et al. 2014)
29. L. plantarum GD-2 NA Anti-biofilm activity 10 –108 CFU/mL (48 h of
7
(Sarikaya, Aslim, and
fermentation) Yuksekdag 2017)
30. L. plantarum 55 NA Anti-inflammatory, 108 CFU/mL (Peptide (Aguilar-Toal
a
antihemolytic, fractions,48 h of et al. 2017)
antioxidant activity fermentation)
31. L. plantarumLAT3 Fermented milk ACE-inhibitory activity 107 CFU/mL (Peptide fractions, (Chaves-L
opez
36 h of fermentation) et al. 2014)
32. L. plantarum WHE92 Munster cheese Pathogen inhibitor 105 CFU/ml (7 d) (Ennahar, Assobhei, and
Hasselmann 1998)
33. L. plantarum YW11 Kefir Improvement of microbiota 4  108 CFU/mL (82 d) (Zhang et al. 2017)
composition
34. L. plantarum CRL 2130 Fermented milk Vitamin producers 2  107 CFU/mL (4 d) (Levit et al. 2017)
35. L. plantarum NA Antimicrobial activity 107 CFU/mL (24–48 h of (Tremonte et al. 2017)
fermentation)
(continued)
4 J. MINJ ET AL.

Table 1. Continued.
Starter culture /Used in Used dose/initial
dairy products/ concentrations and
S. No. Lactobacillus strain Food Products Biological activities time duration References
36. L. plantarum ZDY2013 NA Radical scavenging activity EPS fraction (24 h of (Z. Zhang, Guo,
fermentation) et al. 2016)
37. L. plantarum KJ03 Fermented stinky bean Cholesterol lowering activity 109 CFU/mL (24 h of (Jampaphaeng, Cocolin,
(Sataw-Dong) fermentation) and Maneerat 2017)
38. L. plantarum LP3 Fermented sarshir Radical scavenging activity 109 CFU/g (10 d) (Hashemi et al. 2017)
39. L. paracasei FT700 Goat cheese Proteolytic activity 107 CFU/mL (24 h of (Tulini et al. 2015)
fermentation)
40. L. paracasei FT700 Fermented milk Cytotoxicity and 107 CFU/mL (24 h of (Tulini et al. 2015)
immunomodulatory activity fermentation)
41. L. paracasei CBA L74 Fermented milk Anti-inflammatory activity 109 CFU (14 d) (Zagato et al. 2014)
42. L. paracasei subsp. NA Bactericidal activity Peptide fraction (Miao et al. 2016)
tolerans FX-6
43. L. paracasei subsp. NA Reducing effect on 108 CFU/day (3,6,9 weeks) (Chiang and Pan 2012)
paracasei NTU 101 hyperlipidemia,
atherosclerosis and
hypertension
44. L. paracasei subsp. Fermented Neuroprotective effect 108 CFU/day (3,6,9 weeks) (Chiang and Pan 2012)
paracasei NTU 101 skimmed milk
45. L. reuteri DSM 17938 NA Membrane vesicles (MVs) 8.38  106 CFU/mL (24 h) (Grande et al. 2017)
formation which is useful
for communication
between beneficial bacteria
and intestinal mucosa cells
46. L. reuteri NA Improves the integrity of 2  109 CFU/mL (16 d) (Dicksved et al. 2012)
mucus layer and or
prevention from distortion
during colitis
47. L. reuteri BR11 NA Reduction in inflammatory 109 CFU/mL/day(12 d) (Atkins et al. 2012)
bowel disease
48. L. reuteri ATCC 23272 NA Maintenance of 103– 106 CFU (3–9 d) (Azevedo et al. 2012)
immunological homeostasis
49. L. reuteri NCIMB 30242 NA Improvement of vitamin 2.9  10 CFU/capsule
9
(Jones et al. 2013)
D status (9 weeks)
50. L. reuteri GMNL-263 NA Improvement in insulin 2  109 CFU/day (14 weeks) (Hsieh et al. 2013)
resistance and ameliorates
hepatic steatosis
51. L. reuteri NA Managing immune response 109 CFU/0.1 mL (43 d) (Huang, Lin, and
against food allergy Jan 2017)
52. L. rhamnosus GD-11 NA Anti-biofilm activity 107–108 CFU/mL (48 h of (Sarikaya, Aslim, and
fermentation) Yuksekdag 2017)
53. L. rhamnosus PTCC 1637 Fermented camel milk ACE-inhibitory activity Peptide fraction (21 d of (Moslehishad
fermentation) et al. 2013)
54. L. rhamnosus ZY NA Anti-oxidant and anti- EPS fraction (24 h of (Ng and Xue 2017)
pathogenic activity fermentation)
55. L. rhamnosusA238 Cottage cheese Antifungal activity 106 CFU/mL (up to 21 d) (Fernandez et al. 2017)
56. L. rhamnosus PRA172 Parmigiano Probiotic property 109 CFU/mL (10 months of (Solieri et al. 2014)
Reggiano cheese fermentation)
NA: Not known.

tumor cells has been well demonstrated by Lactobacillus
casei ATCC 25180. Antitumoural activity of this bacterium
was mediated by peptidoglycan. It was observed that the
peptidoglycan fragments enriched with D-configurations of
the amino acids Ala, Gln and Asn demonstrated a signifi-
cant inhibitory effect on the mitochondrial enzyme activity.
Further, it was also observed that only D-amino acids could
enter complexes with hexokinase. So, the cytotoxicity of the
peptide was not only affected by animo acids but their con-
figuration also governed the specific activity of a peptide
(Fichera, Fichera, and Milone 2016).
Immunomodulatory activity of L. casei ATCC 27139 has
been demonstrated by enhancing the host innate immunity
by helping the asparagine synthetase (asnH), which contrib-
uted to peptide synthesis in the peptidoglycan layer. Some
cell wall structures of L. casei were responsible for the
improvement in host innate immunity (Ito et al. 2014). L.
casei BL23 promoted the functional changes in bacteria by
regulating the T cells and FoxP3 and also produced anti-
inflammatory cytokine interleukin. It may potentially modify
the immunomodulatory properties of bacteria by cross talk
with the host cells (Jacouton et al. 2019). A recombinant L.
casei secreted fusion protein CTB (cholera nontoxic B sub-
unit) with YVAD (tetra peptide composed of alanine,
aspartic acid, tyrosine, valine), which can translocate into
intestinal epithelial cells, exerted anti-inflammatory effects in
the intestine (Hiramatsu et al. 2014).
L. casei may produce many antibacterial peptides during
fermentation but their ability to produce biological active
peptides varies among different strains. An aproline rich
antimicrobial peptide PR39 of 4.7 kDa was produced by L.
Casei 393, which was active against Escherichia coli,
Staphylococcus aureus and Salmonella sp. (Zhang, Guo, et al.
2016). The antagonistic property against Salmonella

typhimurium has also been demonstrated by L. casei GG.
Colonization of L. casei GG significantly affected the mesen-
teric lymph nodes, liver and the spleen and delayed the
occurrences of mortality in the animals by 100% (Hudault
et al. 1997). L. casei ATCC 334 demonstrated the ability to
combat with Mycobacterium avium subsp. paratuberculosis,
a causative agent of Johne’s disease, one of the most infec-
tious diseases worldwide in dairy cattle. The oral doses of L.
casei ATCC 334 significantly altered the cytokine responses
and their histology indicated a trend in immunomodulation
and reduction of pathogenesis (Cooney et al. 2014).
Antifungal potential of L. casei has suggested its use to
increase the quality and safety of foods. Antifungal activity
of five L. casei strains has been established against different
food spoiling molds (Aspergillus flavus, Botrytis cinerea,
Colletotrichum gloeosporioides, Penicillium expansum).
Significant antifungal activity was demonstrated by anti-
microbial compound 3-phenyllactic acid, which was pro-
duced by L. casei 21/1 (Cort
es-Zavaleta et al. 2014). The
probiotic strains L. casei strain B and Lactococcus latics
subsp. lactis strain X extracts exhibited fungistatic activity.
The antifungal compounds were identified as lactic acid and
6-O-(a-D-glucopyranosyl)-1, 6-di-O-pentadecanoyl-a-D-glu-
copyranose, a novel fatty acids derivative having MIC of
10 mg/ml (Nyanzi et al. 2014).
Apart from antibacterial and antifungal activities, the
metabolites of L. casei CMPUJ 415 demonstrated the ability
to reduce rotavirus infection by decreasing the damage to
cells, which further prevented the loss of electrolyte.
Produced metabolites interfered with the final amount of
intracellular NSP4 protein and controlled Ca2þ regulation,
which was suggested as an approach to the mechanism used
by probiotic bacteria against rotavirus infection (Olaya
Gal
an et al. 2016).
Reduction in total cholesterol and triglycerides levels by
L. casei Shirota is one of the most important health benefits
to the heart related patients. It effectively decreased the
cholesterol level and also reduced the chance of atheroscler-
otic plaque development in aortic sinus in apo E-deficient
mice. The cardio protective effect mediated through several
mechanisms that correspond to improving serum total chol-
esterol concentration and decreasing cholesterol accumula-
tion in the liver and aortic arterial tissue. Simultaneously, it
improved the level of high density lipoprotein (HDL) chol-
esterol concentration and leading the inhibition of the for-
mation of atherosclerotic lesion (Tang et al. 2014). Effects of
probiotic strain L. casei Shirota on the glucose intolerance,
insulin resistance and lipid metabolisms in rats, which were
fed with a high fat diet, were studied. The results revealed
that it might affect the tissue-specific autonomic nerves
through the vagal nerve pathway to modulate glucose and
lipid metabolism. Suppression of WAT-SNA, ASNA and
Liv- SNA has also been observed in bacterial treated rats
(Tanida et al. 2014). The health benefits of L. casei Shirota
has also been studied for its protection against nonalcoholic
fatty liver disease. Probiotic treatment decreased hepatic
steatosis and plasma alanine-amino transferase levels and
also activated the Toll like receptor 4 signaling cascade in
CRITICAL REVIEWS IN FOOD SCIENCE AND NUTRITION 5
the liver of mice (Wagnerberger et al. 2013). On the other
hand, successful development of vaccine has been done
using marker free strain L. casei ATCC 393. This recombin-
ant bacterium has been found to be a safe tool for the
effective food grade vaccine production (Song et al. 2014).
Lactobacillus delbrueckii subsp. bulgaricus
Two specific yoghurt bacteria i. e., Streptococcus thermophi-
lus and Lactobacillus delbrueckii subsp. bulgaricus in 1:1
ratio commonly is used in the yoghurt production. These
bacteria act in a synergistic way to promote the fermentation
process and develop typical yoghurt flavor. Apart from syn-
ergistic action in yoghurt itself, each bacterium also contrib-
utes to the promotion of bio-functional properties of the
product. Probiotic L. delbrueckii subsp. bulgaricus boosts the
health benefits of yoghurt, which has been reported to pre-
vent inflammatory bowel disease (IBD) and colitis. The anti-
microbial activity of L. delbrueckii subsp. bulgaricus has been
reported against many pathogenic bacteria including
Escherichia coli, Listeria monocytogenes, Shigella etc.
Production of lactic acid along with the organic acids, bio-
active peptides and bacteriocins may govern the antimicro-
bial activity of L. bulgaricus. L. bulgaricus K41 showed
antagonistic activity against L. monocytogenes and also
exhibited the anti-listerial activity by producing bacteriocins
(Zaeim, Soleimanian-Zad, and Sheikh-Zeinoddin 2014).
Similarly, Arabian yoghurt isolate L. bulgaricus Z55 exhib-
ited antimicrobial activity against other LAB and food borne
pathogenic bacteria. L. bulgaricus Z55 produced bacteriocin
like compound which was stable at acidic and neutral pH
conditions and responsible for its inhibitory action against
other bacteria (Abdel-Shafi et al. 2014). Two simultaneous
health benefits such as anti-inflammatory and antibacterial
effects of L. bulgaricus have been analyzed in the chronic
wound healing. The wound dressing of polyvinyl alcohol/
Carrageenan hydrogel was loaded with L. bulgaricus extract
and exhibited promising result on healing process and also
down regulated the expression of TNF-a gene (El-Fawal,
Yassin, and El-Deeb 2017).
Immune stimulating and anti-inflammatory activity of L.
delbrueckii subsp. bulgaricus have also been reported. L. del-
brueckii subsp. lactis CNRZ327 isolated from cheese demon-
strated the anti- inflammatory effect, in vitro as well as
in vivo. This strain showed protective effects in a mouse
dextran sodium sulfate colitis model by modulating the pro-
duction of TGF-b, IL-6 and IL-12 in colonic tissue, TGF- b
and IL-6 in the spleen and it was also responsible for an
expansion of CD4þ Foxp3þ regulatory T cells in the cecal
lymph nodes (Santos Rocha et al. 2014). Anti-inflammatory
potential of cheese starter culture L. delbrueckii subsp. lactis
CNRZ327 and Propionibacterium freundenreichii have also
been reported to protect epithelial cell damage (Pl
e
et al. 2016).
Antihypertensive activity of LAB is one of the most
important biofunctional properties. Angiotensin-converting
enzyme (ACE) has been known for regulating the blood
pressure by conversion of angiotensin-I to angiotensin-II,
6 J. MINJ ET AL.
which is a potent vasoconstrictor. Blood pressure and
sodium retention increases due to the release of aldosterone,
which was induced by enzyme angiotensin-II (Muro Urista
et al. 2011). Thus, inhibition of angiotensin-I to angiotensin-
II could be the best mechanism to reduce of higher blood
pressure and sodium retention as well. To overcome the
side effects associated with antihypertensive drugs, naturally
produced antihypertensive enzyme by LAB may work as an
alternative. Antihypertensive and ACE inhibitor activity of
L. delbrueckii subsp. bulgaricus ACA-DC 87, Enterococcus
spp. and Streptococcus thermophilus, which were isolated
from artisanal Greek yoghurt and fermented milk exhibited
the probiotic properties. It was observed that N- or C- ter-
minal is racidin peptide region of a S1-casein and two
internal peptide fragments j-casein and b-casein were
responsible for the ACE-I inhibitory activity (Georgalaki
et al. 2017). Production of ACE inhibitor peptides by L. bul-
garicus LB6 during the fermentation of goat milk has also
been demonstrated (Shu et al. 2014).
Toxin neutralizing potential has also been reported in
some L. delbrueckii subsp. Bulgaricus strains. Mutant of L.
delbrueckii subsp. Bulgaricus DUC3-17 has demonstrated the
decomposition of uremic toxins such as creatinine, homo-
cysteine, phosphorus, parathyroid hormone urea nitrogen
and uric acid. These toxins decomposition property may be
beneficial for the treatment of chronic renal failure (Bai,
Jiang, and Jiang 2014). Some of the contradictory results of
L. bulgaricus have also been demonstrated in the case of
Clostridium difficile associated diarrhea, as the use of pro-
biotic ACTIMEL (L. bulgaricus, L. casei and S. thermophilus)
did not reduce C. difficile associated diarrhea in elderly
orthogerietric patients (Mallina et al. 2018).
Lactobacillus fermentum
Lactobacillus fermentum is mainly found in the fermented
food and dairy products, which provides several health ben-
efits to gastrointestinal tract. L. fermentum AGR1487 was
able to reduce the expression of intestinal cell tight junction
proteins and genes, which resulted into reduced barrier
integrity in the epithelial cell lines and increased the expres-
sion level of seven tubulin genes and three microtubule
associated proteins. These genes were linked to disassemble
of tight junctions, present in the epithelial cell lines. The
adherence ability of L. fermentum AGR1487 to caco-2 cell
lines has also been improved by facilitating cell to cell inter-
action. However contradictory results have also been found
as L. fermentum AGR1485 negatively affected the barrier
integrity in vitro. Later on its effect on pro-inflammatory
response in germ-free rats colon have been observed in vitro
and the results showed that this bacterium activated TLR
signaling and induced pro-inflammatory cytokines in
immune cells (Anderson et al. 2013). This indicated the
potential of a strain of the same species to differently elicit
inflammatory responses to the host and highlighted the
importance of strain characterization in probiotics
approaches to cure inflammatory conditions (Anderson
et al. 2016). Recently, L. fermentum CNU1969 and L.
salivarius CNU1334 demonstrated the inhibitory activity
against methicillin resistant Staphylococcus aureus (MRSA)
(Kang et al. 2017). EPS produced by probiotic different L.
fermentum strains may also have an active role in its func-
tional properties (Ale et al. 2020).
L. fermentum ATCC 11976 significantly lowered the con-
centration of esterified cholesterol, hepatic free cholesterol,
phospholipids and triglycerides in animals suffering from
nonalcoholic fatty liver disease. Fat deposition in liver,
serum total cholesterol, uric acid, triglycerides and insulin
resistance also decreased in the L. fermentum ATCC 11976
treated animals. All these significant changes were linked to
the expression of key enzymes in the liver and significantly
decreased in the marker enzymes for lipid peroxidation
(Bhathena et al. 2013). Human milk isolate L. fermentum
HM3 and fermented dates isolate L. buchneri FD2 strains
were investigated for their probiotic properties in terms of
safety using sub-acute and acute oral toxicity tests in the
rats as animal model. A significant reduction in b-glucosi-
dase and b-glucuronidase enzymes was observed in the L.
fermentum HM3 and L. buchneri FD2 fed group of rats. It
indicated toward the significant reduction in harmful patho-
genic bacteria, as these are the marker enzymes of harmful
pathogenic bacteria. The effective and safe dose of these
Lactobacillus strains have been noted up to 1010 CFU/kg
body weight per day in a 14 days or 28 days of treatment
period (Shokryazdan et al. 2016). Recently, abomasum
driven rennet cheese for the presence of lactobacilli have
been screened for their potential probiotic properties.
Parameters such as bile and acid resistance, antimicrobial
activity against pathogenic bacteria and antibiotic suscepti-
bility were evaluated (Maleki Kakelar et al. 2019).
Lactobacillus helveticus
Lactobacillus helveticus is generally found in cheese milk and
mainly used in the production of Swiss cheese. The main
purpose of this culture is to develop acid and ripening of
cheese. Apart from Swiss cheese, L. helveticus has also been
used as a starter culture for various cheeses such as
Parmesan, Romano, Provolone and Mozzarella cheese. The
proteolytic nature of L. helveticus strain produces many bio-
logically active peptides which are responsible for a typical
cheese flavor. Many L. helveticus strains possess key pro-
biotic properties, such as ability to survive in the gastro-
intestinal tract, adherance to epithelial cells, and antagonistic
activity against pathogens. L. helveticus DPC4571 prevented
the gastrointestinal infections, improved protection against
pathogens, modulated host immune responses and enriched
intestinal microbiota (Slattery et al. 2010).
L. helveticus containing cheese was found to possess
immuno-regulatory activities like increase in regulatory T
cell and reduction of pro-inflammatory cytokines in mice.
In an in vitro analysis, L. helveticus SBT2171 significantly
repressed the proliferation of mesenteric lymph nodes
(MLN) immune cells and pro-inflammatory cytokines such
as IL-6 and IL-b stimulated with anti-CD3/CD28. The anti-
proliferative potency of L. helveticus was also effective on

peyer’s patches and spleens stimulated with anti-Cd3/CD28
or lipopolysaccarides (LPS). L. helveticus SBT2171 decreased
the IL-1b production from spleens and LPS-stimulated IL-6
production from MLN, peyer’s patches and spleen
(Yamashita et al. 2014). A probiotic mixture including
Bifidobacterium longum Bar33 and L. helveticus Bar13 ame-
liorated immune response in older adults by increasing
naive, activated memory, regulatory T cells, B cells, and nat-
ural killer activity and decreasing memory T cells.
Characterization of S-layer protein (SlpA) and functional
role of L. helveticus M92 have been demonstrated as purified
SlpA from probiotic L. helveticus M92 showed immunomo-
dulatory effects in orally immunized mice. The SlpA was
also involved in the autoaggregation of L. helveticus M92
cells and co-aggregation of L. helveticus M92 with
Salmonella typhimurium. Probiotic L. helveticus M92 also
reduced the infection of Salmonella typhimurium in mice,
which was associated with competitive exclusion in the
intestinal tract and thus enhanced the immune protection
(Beganovi
c et al. 2011). L. helveticus has been studied for
several other health benefits such as inhibition of potential
pathogens, antimutagenic and anti-tumorogenic activities,
anti-hypertensive activity, stimulation of immune and
digestive system, control of diarrhea and reduction of lactose
intolerances (Taverniti and Guglielmetti 2012). Milk fer-
mented by L. helveticus effectively increased calcium absorp-
tion, bone mineral density and bone mineral contents. It has
also been shown to support the bone mineral density in ani-
mal models by increasing calcium absorption in postmeno-
pausal women (Narva et al. 2004). In a study, it was
demonstrated that L. helveticus LBK-16H fermented milk
contains certain tripeptides that showed a positive effect on
arterial stiffness. This tripeptide isoleucine-proline-proline
demonstrated a significant reduction in high blood pressure
in rats (Seppo et al. 2003). Each peptide has unique func-
tional characteristics and this can be identified with various
available techniques. Genome sequencing of L. helveticus
CNRZ 32 has revealed the presence of four proteinase genes
i.e. prtH, prtH2, prtH3 and prtH4 in the proteolytic system.
During the study of proteolytic systems in 51 strains, several
strains of L. helveticus exhibited highly intra specific diver-
sity of genes, which encoded the cell envelope-associated
proteinases (CEPs). These CEPs were responsible for the
functional characteristics of this bacteria (Broadbent et al.
2011). Several L. helveticus strains have been isolated from
traditional dairy products and nine of them confirmed the
presence of two CEPs i.e. prtH and prtH2 (Nejati, Babaei,
and Taghi-Zadeh 2016). L. helveticus SBT2171 contributed
to the many properties including anti-allergic effects
(Yamashita et al. 2019). As L. helveticus has been reported
to exhibit various techno-functional properties, it may be
used to promote the bio-functional and therapeutic proper-
ties of the products.
Lactobacillus paracasei
Lactobacillus paracasei is commonly present in the human
intestinal tract, which comes from fermented milk,
CRITICAL REVIEWS IN FOOD SCIENCE AND NUTRITION 7
vegetables and meat products. L. paracasei along with L.
casei are used as a probiotic and dietary supplements for
many clinical aspects. Changes in the gut microbial popula-
tion may change the host health and its physiology.
Probiotic gut bacteria follow some specific metabolic path-
ways such as degradation of dietary carbohydrates. Similarly,
L. paracasei subsp. paracasei W20 has been reported to
degrade prebiotic inulin in the human gut and could be
used as a commercial symbiotic (Boger, van Bueren, and
Dijkhuizen 2018).
Probiotic L. paracasei K5 isolated from dairy products
has been used for the Feta type cheese production. The use
of immobilized biocatalyst which was composed of wheat
bran prebiotic and L. paracasei K5 cells improved its aroma,
acceptability, quality and shelf life of the feta type cheese
(Terpou et al. 2018). L. paracasei subsp. tolerans JG22 was
isolated from pepper-leaf and reported to have high survival
rate during its exposed to pepsin containing gastric juice
and intestinal fluid amended with oxgall. This strain showed
adherence to the caco-2 cell lines and resistance to rifampi-
cin, erythromycin, cephalothin and ciprofloxacin. Its anti-
mutagenicity and mutagen binding ability demonstrated
highest antimutagenicity against 2-nitroflourene (2-NF) with
51.37% inhibition, and relatively lower against 4-nitroquino-
line-1-oxide (4-NQO). Therefore, it was suggested that this
potential probiotic strain may have an important role in
decreasing the risk of cancer by suppressing the mutagenesis
and absorbing mutagens (Lim 2014). b-Glucosidase and
b-Glucuronidase have been known as potential carcinogenic
biomarkers and L. paracasei 0919 reduced this enzyme activ-
ity by 76% in the children and 82% in elder people (Nowak
and
Sli_zewska 2014). L. paracasei DTA93 and L. paracasei
DTA81 could survive in the simulated gastric conditions
and showed antimicrobial activity, bile salt hydrolase activ-
ity, hemolytic activity, and inhibitory activity against biofilm
formation. It also exhibited anti-cancerous activity by attach-
ing with HT-29 human cancer cell lines (Tarrah et al. 2019).
Different LAB including L. paracasei were able to reduce the
level of cholesterol and also produced bacteriocins like
inhibitory substances, which showed antagonistic activity
against Listeria monocytogens, Salmonella enteritidis and
Staphylococcus aureus (Iranmanesh, Ezzatpanah, and
Mojgani 2014).
Immune modulation property of L. paracasei was con-
firmed by preventing the weight loss up to 50% in the colitis
mice. L. paracasei strains KBL382, 384 and 385 have been
used for the treatment of inflammatory bowel diseases via
immune modulation and positive impact on gut microbial
population. These strains of L. paracasei exhibited significant
improvement in the dextran sodium sulfate-induced colitis.
It significantly reduced the tumor necrosis factor, interleu-
kin-4, IL-6, IL-17A and interferon gamma; whereas anti-
inflammatory cytokines such as IL-10, CD4þ, CD25þ,
Foxp3þ and Tregulatory cell populations were significantly
higher (Kim et al. 2019). Fermented milk containing L. par-
acasei strain CIDCA8339 has been given to the mice and it
did not changed the IL-10, TNF-a and TGF-b mRNA
expression as well as sIgA levels in the ileum. However, the
8 J. MINJ ET AL.
decreased level of anti-inflammatory cytokine INF-c has
been reported in the ileum (Bengoa et al. 2019). Any alter-
ation in the intestinal microbiota and immunosenescence is
generally associated with the aging also. Probiotics and fer-
mented food might be helpful in the stabilization of altered
intestinal microbiota. L. paracasei strain Lpc-37 evaluated to
modulate the immune markers including natural killer cell
activity, cytokine profiles and phagocytic activity. Activity
and composition of the intestinal microbiota in a random-
ized, double-blind and placebo-controlled group, showed a
very limited effect on the intestinal microbiota and immune
markers of healthy elderly population (Forssten et al. 2011).
Different metabolites produced by L. paracasei CBA L74
during fermentation of milk for infant formula and culture
media demonstrated significant anti-inflammatory properties
on dendritic cells (DCs) in response to the inflammatory
enteric pathogenic bacterium Salmonella typhimurium
(Zagato et al. 2014). Allergic rhinitis (AR) generally occurs
due to an imbalance between Th1 and Th2 cells. Subjects
with persistent AR who are treated with an oral H1-antihis-
tamine showed improvement in the quality of life due to
probiotic L. paracasei LP-33. This treatment also improved
the ocular symptoms consistently, whereas no change was
observed in nasal symptoms (Costa et al. 2014). Insulin
resistance (IR) is another crucial pathophysiological param-
eter in the development of nonalcoholic fatty liver diseases.
IR and steatosis induced rats were treated with L. paracasei
B21060 along with arabinogalactan and fructo-oligosacchar-
ides. Liver inflammatory markers have been shown to be
down regulated and the expression of nuclear peroxisome
proliferator activated receptors was increased in the rats
who received the synbiotic treatment. Synbiotic treatment
also reduced the cytokines synthesis and deregulated the
TLR 2, 4 and 9 mRNAs. Among all these functions, synbi-
otic feeding effectively protected the gut barrier integrity
and reduced the population of harmful bacteria in the
colonic mucosa (Mattace Raso et al. 2014). Synbiotic com-
prised of probiotic L. paracasei DSM 46331 and
Bifidobacterium animalis subsp. lactis along with prebiotic
oat b-glucan has significantly alleviated the high fat diet
induced obesity and weight gain. This synbiotic has also
been played a significant role in the homeostasis of gut
microbiota (Ke et al. 2019).
Lactobacillus plantarum
Functional properties of L. plantarum are regulated by the
production of EPS, bioactive peptides and some organic
acids. The metabolites produced by L. plantarum I-UL4
exhibited selective cytotoxic effect on some cancer cells in
dose and time dependent manners. On the other hand, the
same metabolites did not cause any toxic effect or hemolysis
on normal cells. This selective toxicity was mediated by anti-
proliferative effect and induction of apoptosis against malig-
nant cancer cells. On the basis of these results, postbiotic
metabolites produced by Lactobacillus plantarum I-UL4
were suggested for their potential use as adjunctive treat-
ment for cancer (Chuah et al. 2019).
Milk fermented by probiotic L. plantarum K25 demon-
strated the production of volatiles such as acetoin and 2,3-
butanediol. Among different metabolites (glyceric acid, malic
acid, succinic acid, glycine, alanine, ribose, and 1,3-dihy-
droxyaceton), c-aminobutyric acid was the most prominent.
This fermented milk also showed a significant increase in
ACE inhibitory activity (Yang et al. 2020). Cheeses made
with different cultures exhibited different bio-functional
activities e.g. cheese made with camel milk isolate EPS pro-
ducing culture exhibited higher a-amylase activity than com-
mercial EPS producing culture; whereas traditional culture
or non-EPS producing culture exhibited good radical scav-
enging activity. The ACE-inhibitory activity has been
reported significantly higher in the cheese made with camel
milk EPS producing culture (Al-Dhaheri et al. 2017). The
purified EPS from L. plantarum MYU 74 composed of glu-
cose and galactose with a molar ratio of 2:1 demonstrated
scavenging ability on hydroxyl radicals and protective effects
on H2O2-induced Caco-2 cells oxidative injury in vitro. This
EPS demonstrated the scavenging of reactive oxygen species
(ROS), reduction of lipid peroxidation and up-regulation of
enzymatic & non-enzymatic antioxidant activities
(Yamamoto et al. 2019). An acidic EPS (12.4 kDa), consisted
of many sugars with different composition, which exhibited
a very good antioxidant against ABTS (65.5%), DPPH
(60.5%) and hydroxyl radicals (80.4%) at a concentration of
10 mg/mL (Min et al. 2019). Similarly, the amylolytic poten-
tial of L. plantarum Bom2 could resist the acids, bile salts
and antibiotics (Gotcheva et al. 2018). Among 63 isolated
LAB, L. plantarum OF101 demonstrated good hydrophobi-
city in the presence of xylene (46.2%) and n-hexadecane
(43.6%), and the bacterium was resistant to low pH, bile salt
and simulated gastric transit (Adesulu-Dahunsi, Jeyaram,
and Sanni 2018).
Fermented soy milk with L. plantarum had more agly-
cones, which helped in decreasing the hepatic cholesterol
and serum triglyceride contents. It also reduced the hepatic
triglycerides, serum free fatty acids and simultaneously ele-
vated the serum HDL-cholesterol and fecal cholesterol con-
tent. This fermented soy milk down regulated the expression
of SREBP-1c and SREBP-2 by targeting the genes FAS,
SCD1, LDLR and HMGCR. Fermented soy milk elevated the
adipose tissues gene expression such as genes involved in
triglyceride rich lipoprotein uptake, fatty acid oxidation,
HDL-biogenesis and adiponectin signaling as well as the lev-
els of ACC and phosphorylated AMPK (Kim et al. 2014).
The oral dose of probiotics significantly improved the
immunological parameters, reduced the pancreatic and plas-
matic a-amylase activities, and protected the pancreatic tis-
sues and plasma glucose levels in the diabetic rats. L.
plantarum TN627 demonstrated efficient protective effects
to the liver and kidney by decreasing serum aspartate trans-
aminase, gamma-glutamyl transpeptidase activities, alanine
transaminase, lactate dehydrogenase, urea and creatinine
content. Thus, it was suggested that L. plantarum TN627
has the potential to combat with anti-diabetic effects and its
complications (Bejar et al. 2013). Any change in gut micro-
bial population directly affects the host immune system in

many ways. Probiotic treatment in ovariectomy mice signifi-
cantly altered the immune system associated bone loss and
bone mineral density. In ovariectomy condition, the level of
resorption marker C-terminal telopeptide and the urinary
fractional excretion of calcium were improved by probiotic
treatment. But the probiotic administration suppressed the
expression of two main inflammatory cytokines TNFa and
IL-1b, and simultaneously it also increased the expression of
OPG, a possible inhibitor of osteoclastogenesis, in cortical
bone of ovariectomy mice (Ohlsson et al. 2014). Similarly
bone health and age related osteoporosis studies have been
done on mouse model. Dietary enriched powdered whole
grape and probiotics (equal ratio of Bifidobacterium bifidum,
B. breve, L. bulgaricus, L. casei and L. plantarum) had a
positive effect on bone health and did not alter the bone
micro architecture (Blanton 2018).
Fermented soshiho-tang (SST) made with L. plantarum
KFRI-144 exhibited anti-proliferative activity in vascular
smooth muscle cell (VSMC). The SST fermented with this
strain enhanced the inhibition of platelet derived growth fac-
tors (PDGF)-BB-induced VSMC proliferation. These inhibi-
tions were mediated through down regulation of Akt
phosphorylation (Lee et al. 2014). Applications of L. planta-
rum KCCM 11322 have also been studied for the promotion
of vascular endothelial growth factor (VEGF) on vascular
and hair growth. The main angiogenic functions of VEGF
includes: to increase the vascular permeability, stimulation
of capillary formation, induction of proliferation of endothe-
lial cells etc. The hydrolysate of L. plantarum KCCM 11322
with enzyme proteases, treated with MG-63 osteoblasts
showed the excellent VEGF production in the C57BL/6 male
mice (Woo et al. 2019).
Milk allergy with protein b-lactoglobulin can also be
treated with the oral probiotic supplementation. Probiotic L.
plantarum ZDY2013 and L. rhamnosus GG have been
reported to suppress the allergic response, decrease the
serum IgE concentration and were able to relieve the ana-
phylaxis symptoms (Fu et al. 2019). It was reported that the
fermented ginseng extract significantly inhibited the secre-
tion of interleukin-4 (IL-4) and b-hexosaminidase from the
mast cells, which are known as allergic causing agents
(Hwang et al. 2018).
A study reported that the patients produced a higher
inflammatory response toward L. plantarum WCFS1 as
compared to APCs from healthy and HIV negative controls.
APC expression of TLR2 and CD36 increased the signaling
through p38-MAPK, and on the other side the expression of
MAP Kinase phosphatase-1 (MKP-1) decreased in chronic
HIV infection. However, an enhancement in responsiveness
of APCs to the commensal lactobacilli might have contrib-
uted to the immune activation (Nagy et al. 2013). L. planta-
rum strain AYA also provided the protection against
Influenza Virus (IFV) infection. IFV infection has been
known to infect the local immunity and it continuously
weakens the immune system. L. plantarum strain AYA
induced the anti-inflammatory cytokines by the promotion
of IgA and protected from IFV. Moreover, L. plantarum
strain AYA produced the IL-6 in Peyer’s patch dendritic
CRITICAL REVIEWS IN FOOD SCIENCE AND NUTRITION 9
cells and simultaneously it enhanced the production of IgA
in the small intestine and lung. Therefore, L. plantarum
strain AYA may be used for the enhancement of local
immunity by the production of mucosal IgA and to reduce
the risk of IFV infection (Kikuchi et al. 2014).
Antibacterial property of L. plantarum might be due to
its adherence ability to the intestinal epithelial cells and then
production of antibacterial compounds, which may inhibit
the growth of harmful and pathogenic bacteria.
Antimicrobial activity of L. plantarum ATCC 8014 against
Clostridium perfringens, C. butyricum and C. difficile has
already been established. This inhibitory property of L. plan-
tarum ATCC 8014 was facilitated by easy colonization abil-
ity and survival rate under the harsh acidic and bile salt
concentrations (Monteiro et al. 2019). Self-life of food prod-
ucts might be increased during long term preservation using
antifungal strains and their produced metabolites.
Antifungal potential of L. plantarum UFG 121 has been
used for the production of oat-based beverage. This product
was artificially contaminated with Fusarium culmorum, while
L. plantarum UFG 121 produced metabolites acted as a
good bio-preservative against fungal contamination and
inhibited fungal growth (Russo et al. 2017). L. plantarum
HS produced different bioactive peptides sequences, but
only one peptide sequence SGADTTFLTK significantly
reduced the growth of Aspergillus parasiticus (73%) and
Penicillium expansum (58%) (Luz et al. 2017). Similarly,
antifungal activity against Candida albicans and Candida
krusei has also been observed (Bulgasem et al. 2016).
Aflatoxin produced by certain molds can be degraded by
L. plantarum. L. plantarum MYS44 was highly useful in the
degradation of toxic aflatoxin- B by growth inhibition as
well as suppression of aflatoxin-B production
(Poornachandra Rao et al. 2019). The antifungal substances
produced by L. plantarum K35 were resistant to physical
sterilization as well as to some proteolytic enzymes (trypsin
& proteinase K). L. plantarum K35 supernatant caused
severe damages to the cell wall and cytoplasmic membrane
along with the complete destruction of cell organelles
including mitochondria and nucleus of fungal strains
(Sangmanee and Hongpattarakere 2014). Thus, antifungal
agents from L. plantarum supernatant could be effectively
used for the prevention of food and food products from
spoilage and aflatoxin.
Lactobacillus reuteri
Lactobacillus reuteri has also been widely studied for its dif-
ferent aspects of bio-functional activities such as antimicro-
bial, antidiarrheal activity, improvement of gastrointestinal
microflora, adhesion to the intestinal epithelial surface and
balancing the composition of healthy bacteria in the gut.
Antibacterial activity of L. reuteri can be positively corre-
lated with a metabolite, reuterin, which has demonstrated
bactericidal and bacteriostatic activities against a large num-
ber of gram positive and gram negative bacteria and even
though to some fungi. Reuterin produced by L. reuteri
ATCC 55730 demonstrated bactericidal activity at a
10 J. MINJ ET AL.
concentration of 0.08 to 20.48 mM against H. pylori. This
antibacterial activity of reuterin did not alter the gut micro-
bial populations. Simultaneously, H. pylori virulence genes
flaA and vacA were also down regulated by reuterin
(Urrutia-Baca et al. 2018). Preventive impact of L. reuteri
ATCC 6475 on fast food induced obesity and age associated
weight gain in mice was studied using oral administration.
This therapy altered the pro-inflammatory immune cell pro-
file and prevented the abdominal fat pathology and age
associated weight gain in mice. L. reuteri ATCC 6475 sig-
nificantly restored the CD4þ T cell balance and also helped
in the weight management (Poutahidis et al. 2013). Anti-
inflammatory effects of L. reuteri in the trinitrobenzene sul-
fonic acid (TNBS)-induced colitis in mouse model was
observed, which revealed that L. reuteri possess a chromo-
somal hdc gene cluster (genes hdcA, hdcB, and hdcP) and
could also repressed human TNF production. Suppression of
acute colitis in mice have been documented by colonic
injury, serum amyloid A (SAA) protein concentration,
diminished weight loss and lowering the uptake of [18F] flu-
orodeoxyglucose by positron emission tomography (Gao
et al. 2015).
Antibacterial activity and gastrointestinal symptoms such
as gastrointestinal disturbances and gastritis in the case of
diarrhea caused by H. pylori was cured and eradicated by L.
reuteri DSM 17938 and L. reuteri ATCC PTA 6475
(Francavilla et al. 2014). Similarly, antimicrobial activity of
probiotic L. reuteri has been studied in 104 strains isolated
from porcine feces. Out of which, 16 strains exhibited higher
and 8 strains showed lower anti pathogenic effect against
two pathogens related to swine, Escherichia coli and
Salmonella enteric subsp. enteric serovar typhimurium.
Furthermore, these strains have been selected for genomic
comparison and results demonstrated the group differences
in the pdu-cbi-cob-hem gene cluster, involved in the reu-
terin and cobalamin biosynthesis (Lee, Shih, et al. 2017).
Thus, it could be concluded that bacteriocins like molecule
reuterin was responsible for the inhibitory action against
swine-related pathogens. A 47 kDa, putative sulfated-galacto-
sylceramide (sulfatide)-binding protein purified from the cell
surface of L. reuteri JCM1081 also exhibited the potential
binding ability to the mucosal surfaces. This purified protein
has been identified as elongation factor-Tu (EF-Tu) and EF-
Tu (Nishiyama et al. 2013).
Lactobacillus rhamnosus
Lactobacillus rhamnosus may easily adhere to the mucous
membrane of intestinal epithelial cells and it plays a signifi-
cant role in the homeostasis of intestinal microbiota. This
adherence property of L. rhamnosus further promotes the
innate immune response and enhances the adaptive immun-
ity. It also produces antibacterial compounds and some
other health promoting biologically active compounds.
However, all these properties of L. rhamnosus cumulatively
promote the gut barrier functions. Probiotic strain L. rham-
nosus DR20 has shown good adherences to the intestinal
mucous in vitro, resistant to bile salt, low pH and sensitive
to antibiotics. Immunomodulatory effects have also been
observed by significant increase in IgA content (Mart
ınez-
Abad et al. 2016). Similarly, the immune modulatory effects
of probiotics on the neonatal gnotobiotic piglets infected
with virulent human rotavirus revealed that total IgA levels
in the post and pre human rotavirus challenged piglets
serum were significantly higher in Escherichia coli Nissle
colonized piglets rather than L. rhamnosus GG colonized
piglets. This study have suggested that different strains have
difference in adherence properties in the intestinal micro-
environment, which further contributed differentially in the
modulation of rotavirus infection and immune cell
responses (Kandasamy et al. 2016). The immune mediated
diseases such as allergy could also be treated with the long
term perinatal probiotic administration and have been con-
sidered as safe for the reduction of allergy prevalence in the
children (Lundelin et al. 2017).
Cranberry proanthocyanidins fermented by Lactobacillus
rhamnosus ATCC 9595 under anaerobic conditions pro-
duced different metabolites including: 4-hydroxyphenylacetic
acid, 3-(4-hydroxyphenyl)propionic acid, hydrocinnamic
acid, catechol, and pyrogallol. The extract containing these
metabolites completely inhibited human hepatocellular car-
cinoma HepG2 cells proliferation. The cytotoxic effect of the
extract was found to be mitochondria-controlled and not by
proapoptotic caspase-3/7 dependent (Rupasinghe, Parmar,
and Neir 2019). Extract containing the metabolites produced
by L. rhamnosus MD 14 exhibited both antigenotoxic and
cytotoxic potential against colon cancer. These metabolites
contained both high and low molecular weight heat sensitive
organic acids and proteins along with the chemical com-
pounds such as ethers, amides, esters, etc (Sharma, Chandel,
et al. 2020).
The antibacterial potential of L. rhamnosus GG was
mediated by the production of low molecular weight non-
proteinaceous, heat stable bactericidal substance, which has
been active at acidic pH conditions against pathogens.
Another therapeutic application of probiotic L. rhamnosus
GG has also been observed on antibiotic associated diarrhea
and Clostridium difficile associated diarrhea, however, the
effectiveness of this probiotic was dose depended
(Mantegazza et al. 2018). Synbiotic containing B. infantis, B.
breve, L. acidophilus, L. casei, L. delbrueckii subsp. bulgari-
cus, L. rhamnosus, S. thermophilus and a prebiotic fructo-
oligosaccharide was able to reduce colic in breastfed infants
(Mugambi et al. 2012). Similarly, the biological activities of
L. rhamnosus NCDC17 have been reported in the manage-
ment of obesity and different types of diabetes. L. rhamnosus
NCDC17 improved the oral glucose tolerance test and many
other biochemical parameters such as fasting blood glucose,
free fatty acids, glycosylated hemoglobin and cholesterol
level. Improvement in the antioxidant enzymes such as cata-
lase, glutathione peroxidase, superoxide dismutase, and thio-
barbituric acid reactive substances, expression of glucagon
like peptide-1 gene, tumor necrosis factor-a, and interlukin-
6 incresed in L. rhamnosus NCDC17 fed group of rats
(Singh et al. 2017). Freeze-dried strains of L. acidophilus, L.
rhamnosus, L. casei, L. bulgaricus, Streptococcus thermophiles,

Figure 1. Possible mechanisms of resistant lactobacillus to transfer the antibiotic resistance and production of bioactive metabolites for health benefits.
Bifidobacterium longum and Bifidobacterium breve and
100 mg of prebiotic fructo-oligosaccharide with lactose as
carrier substance have been supplemented in the form of
capsules to see its effects on the type 2 diabetes patients.
The regular consumption of this probiotic supplements for
six weeks significantly decreased fasting plasma glucose and
increased the HDL cholesterol concentrations (Razmpoosh
et al. 2019). Antioxidant activity of a human breast milk iso-
late, EPS producing six L. rhamnosus strains has been
reported to chelate the metal ions, which is considered as
one of the antioxidant mechanisms (Riaz Rajoka et al.
2018). L. rhamnosus IMC 501 accelerated the bone depos-
ition through stimulation of the expression of gene Mpk1/3,
involved in ossifications. Thus, this probiotic strain helped
in the skeleton development and treatment of bone disorder
(Maradonna et al. 2013). Probiotic containing foods have
also been used to prevent respiratory tract and gastrointes-
tinal infections in adult and pediatric patients. Treatment
with probiotic L. rhamnosus GG and Bifidobacterium lon-
gum reduced the lung injury, infection and sepsis (Khailova
et al. 2014).
Influence of antibiotics on gut colonizing probiotic
Lactobacillus strains
Besides preventing antibiotic associated diarrhea, probiotic
Lactobacillus may help to maintain the gut microorganisms
and restore the micro-flora during antibiotic treatment. It
also helps in fighting off the pathogenic bacteria even in
children (Mohseni et al. 2013; Wu et al. 2018). However,
taking probiotic lactobacilli along with antibiotics may
decrease its effectiveness as most of the bacterial probiotics
are generally sensitive to generally prescribed antibiotics.
Such antibiotics may disturb the diversity of native gut colo-
nizing microbiota (Mousavi Khaneghah et al. 2020). To
avoid this interaction and alteration in the diversity of gut
microbiota, a time interval is required. Alternatively, use of
resistant lactobacillus as probiotic was also suggested (Neut,
Mahieux, and Dubreuil 2017).
Various bioactive metabolites and antibiotic producing
lactobacillus are considered as second generation probiotics.
CRITICAL REVIEWS IN FOOD SCIENCE AND NUTRITION 11
However, there is a risk associated with the use of probiotic
strains having antibiotic resistance genes, as these resistant
bacteria may potentially pass these genes to other probiotic
or pathogenic bacteria through horizontal gene transfer,
while retain the gene through vertical gene transfer
(Figure 1). Commercially available probiotic formulations
may possess antibiotic resistance genes (Rozman et al. 2020).
These genes are generally present in mobile genetic elements
e.g. plasmid and transposons. On the other hand, some spe-
cies of LAB may have intrinsic resistance to a number of
antibiotics i.e. bacitracin, beta-lactams, kanamycin, teico-
planin, and vancomycin (Imperial and Ibana 2016). The
influence of resistant or susceptible probiotic lactobacillus
during the administration of antibiotics on gut colonization
has been illustrated in Figure 2. Probiotic containing resist-
ant microbes may lead to a case with higher gut coloniza-
tion by resistant bacteria (G2), while the probiotic dose
containing susceptible or native bacteria will result in nor-
mal gut colonization (E1). Thus, it may be recommended to
provide a susceptible probiotic strain for longer duration
even after finishing the antibiotic dose.
Role of Lactobacillus as a supportive treatment in
emerging disease
Lactobacillus has shown its potential toward the treatment
for various infectious diseases. Recent outbreak of SARS
Covid 19 has been declared as pandemic by WHO. There
are several reports being published on this deadly disease
but none of them could establish a significant role of pro-
biotic doses with the treatment of Covid 19. Lower gut
microbiota diversity in old age can be positively correlated
with lesser immunity. It was suggested that higher gut
microbiota colonization and diversity may be achieved by
personalized nutrition and supplementation, which may
improve immunity and the impact of this disease can be
minimized in elderly and immune-compromised patients
(Dhar and Mohanty 2020). Recently, a case study of recov-
ered Covid 10 patient Dr. Jaime F. Flor (ENT specialist)
communicated by himself, clears some of the points related
to the supporting role of Covid 19. He consumed two
12 J. MINJ ET AL.

Figure 2. Schematic presentation shows the administration of susceptible or resistant lactobacillus during antibiotic treatment. A: Pathogen attack on the gut colo-
nizing lactobacillus; B: Pathogen – Lactobacilli interaction; C: Antibiotic administration; D1: Supplementation of susceptible probiotic lactobacillus increases the com-
petition; E1: Continued probiotic lactobacillus result in higher colonization of susceptible with a few resistant lactobacillus; In the second case, D2:Supplementation
of resistant probiotic lactobacillus; E2: Interaction between resistant lactobacillus and other microbes will increase the chances of horizontal gene transfer; F2:
Continued probiotic lactobacillus result in higher colonization of resistant bacteria; G2: Colonization by resistant lactobacillus.

consortium pills daily containing Lactobacillus acidophilus
20 billion CFU along with antibiotics and other medicines.
L. acidophilus stimulated the Gut Associated Lymphoid
Tissue to produce antibodies against virus and bacteria. This
dose of L. acidophilus stimulated sufficient production of the
IgG and IgM and the bacterium produced vitamin B12,
which also played a positive role along with vitamin C and
Zinc (Flor 2020).
On the other hand, in a study, patients with Covid 19
were given antibiotics, and diarrhea occurred in some of the
patients. To reinforce the colonic microbiota a probiotic
combination consisted of Lactobacilli and Bifidobacteria was
prescribed to the patients. These probiotics demonstrated
modest efficacy in reducing the antibiotic-associated diar-
rhea. On the other hand, when a probiotics preparation con-
sists of Lactobacillus rhamnosus GG, Bacillus subtilis, and
Enterococcus faecalis was given to critically ill patients on
mechanical ventilation, they developed significantly less ven-
tilator-associated pneumonia as compared to placebo (Mak,
Chan, and Ng 2020). Thus, on the basis of this study the
efficacy of probiotics in the treatment of Covid 19 may not
be established, which needs further investigation, but a diet
plant with probiotics may be suggested as a supportive treat-
ment. It further highlights the importance of Lactobacillus
containing probiotics and provides a direction toeards fur-
ther research dealing with its metabolites, immunomodela-
tory effects and molecular mechanisms involved to combat
such emerging diseases.
Concluding remarks and future outlook
Several research works have demonstrated a number of bio-
functional properties of Lactobacillus spp. which can be used
in the treatment of concerned physiological disorder. It is
recommend that a proper diet plan including suitable dose
of probiotics may be prescribed to the patients suffering
from different emerging or well recognized diseases. Many
research studies have been done in vitro which needs
in vivo confirmation using an animal model as well as
human trials. Deciding optimum dosages of probiotics
should be well defined during antibiotic treatment. A cross
validation and regular quality assurance is also important in
the case of probiotic containing foods. Further, studying the
effect of probiotic microbes on gene regulations may reveal
molecular level interactions. Application of non traditional
lactobacilli with different bio-functional properties can be
explored for their use in probiotic formulation. These pro-
biotic microbes may be explored for postbiotic properties,
which will increase the scope of their use in a wide variety
of food including bakery items.
Disclosure statement
No potential conflict of interest was reported by the authors.
ORCID
Rakesh Kumar Sharma http://orcid.org/0000-0002-2826-1291
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