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Lactic Acid Bacteria: Probiotic Characteristic, Selection Criteria, and its Role in
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Lactic Acid Bacteria: Probiotic Characteristic,

Selection Criteria, and its Role in Human Health
(A Review)
1
Rahul Gupta, 1*Kadirvelu Jeevaratnam, 2Amrin Fatima
1
PhD scholar, 1*Professor (Corresponding Author), 2Post Graduate
1
Department of Biochemistry and Molecular Biology,
1
Pondicherry University, Pondicherry, India.
2
Department of Biotechnology,
2
Central University of South Bihar, India.

Abstract: The traditional usage of antibiotics as a therapeutic agent has engendered severe side effects on Human health. Therefore, the demand
for the natural alternatives has been enticed by both the manufacturer and the consumers. This natural alternative is the usage of “Probiotics”.
According to FAO/WHO- “Probiotics are the living microorganisms that are administered/consumed in an ample amount which produces
beneficial effects on the host”. Lactic Acid Bacteria (LABs) are considered to be the utmost studied probiotic microorganisms because of their
dynamic nutritional value; confer several health benefits, modulating host immune responses, inhibiting the growth of the food-borne and other
harmful pathogen. LABs are commonly present in fermented milk products, fermented foods, and beverages. However, for the assortment of
probiotic LAB strains for commercialization, a number of aspects have to be measured which includes: Safety, Stability and Feasibility, Functional
and Technological aspects, and proper Quality control. Hence, the current review comprehends the scenario of LABs and its Probiotic
characteristics, selection criteria and their beneficial impacts on Human health along with the present guidelines and code of practice.

Keywords— Lactic Acid Bacteria, Probiotics, Lactobacillus, Fermented Food, Immunomodulation, Selection Criteria, Human
Health, Gastrointestinal Tract, and Microbiota.

Abbreviations-LABs/LAB, Lactic Acid Bacteria; GRAS, Generally Recognized As Safe; BSH, Bile Salt Hydrolases; ACE,
Angiotensin Converting Enzyme; FAO, Food and Agriculture Organization; WHO, World Health Organization; and AIDS,
Acquired Immunodeficiency Syndrome.

I. INTRODUCTION

The traditional treatment of antibiotics has engendered multidrug-resistant pathogens which has been a severe threat to human health
and therefore this has led to the development of a new method of preventing various bacterial and fungal diseases of the digestive
tract in humans by the usage of the safe and natural antimicrobial biomolecules (low molecular weight compounds and ribosomally
synthesized peptides/proteins) extracted from the Lactic acid bacteria (LABs) [16, 26]. LABs are reflected in a group of gram-positive
bacteria (cocci or rods) which does not form spores, consider as non-pathogenic, and typically are anaerobic or facultative aerobic
bacteria which has the capability to make use of carbohydrates and metabolically yields lactic acid as the key product during
fermentation [9, 43]. They are typically found in decayed plants and animal matter, fecal substances, in numerous raw materials that
are used to produce fermented foods like milk, meat, and flour and even in the gut of herbivorous animals and humans as a symbiotic
[29, 31, 46]. LABs are reported for being used in alcoholic and food fermentations since 6000 of years [30]. It has been well
documented that probiotic LAB strains show very good anti-bacterial, anti-fungal, and anti-viral properties by producing numerous
anti-microbial compounds which include Lactic Acid, Acetic Acid, Propionic Acid, Alcohol, Diacetyl, and Bacteriocin (ribosomally
synthesized proteinaceous toxins which inhibit the growth of similar or closely related bacteria). These antimicrobial compounds
produced by the LAB, interact with the cell membranes of the harmful pathogens (Listeria monocytogens, Pseudomonas aeruginosa,
Staphylococcus aureus, Escherichia coli, Helicobacter pylori, Candida albicans, etc.) and inhibit their growth by intracellular
acidification (lowering the pH) and denaturing the proteins of the membrane which leads to the dysfunction of the membrane
permeability [1, 7, 12, 43, 78, 82]. This ability of LAB led to its use as a starter culture in many food fermentation processes because
it protects the lifespan of many foods by inhibiting the growth of other harmful pathogens and also help in maintaining the palatability
of the food [2, 5, 16].

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II. LACTIC ACID BACTERIA

Lactic acid bacteria are Gram-positive, non-spore-forming Rods, Cocci, and Cocco-bacilli, non-respiring but aerotolerant, fastidious,
acid tolerant and catalase negative devoid of cytochromes, which produces lactic acid as one of the key fermentation products by
utilizing carbohydrates during fermentation. These bacteria not only produces lactic acid as an end product of carbohydrate catabolism
but also produces organic substances that contribute to the flavor, texture, and aroma and thus aiding unique organoleptic
characteristics to the products [1, 8, 51, 52, 64].

2.1 Brief History

From more than 4000 years lactic acid bacteria have been used to extend the shelf life of various foods through fermentation processes.
Carl Wilhelm Scheele was the first to discover the lactic acid bacteria while isolating lactic acid from sour milk. In 1857, Pasteur
discovered that microorganisms were responsible for the souring of milk and can be heat killed by boiling the milk and further he
found that this microorganism was involved in the lactic acid production. In 1881, M/s Clinton processing company (USA) was the
first to produce lactic acid commercially through fermentation processes. The first commercial starter cultures were prepared in
Denmark from the raw milk around the end of the 19th century and it was unknown mixes of microbes. The idea of using pure single-
strain starter cultures started evolving in the 1940s and 1950s comprising lactic acid bacteria and became commercially available.
After 1980s researchers observed the widespread application of lactic acid bacteria in the field of biomedicine, food preservative,
food processing, and fermentation and animal husbandry [27, 40, 83].

2.2 Classification of Lactic Acid Bacteria

LAB have also been classified based on acid production into different genera/species by fermenting sugars and its growth at specific
temperatures [38, 53]. The conventional approach of LAB classification was based on physiological and biochemical characteristics
but recently, molecular characterization has become an important tool for classification and identification of LAB. Molecular
characterization includes: random amplified polymorphic DNA profiling, 16S rRNA gene sequencing, PCR-based fingerprinting and
soluble protein patterns [54] and differentiation of species by multiplex PCR assay by using specific recA derived primers [55].

2.2.1 Homofermentative and Heterofermentative

The lactic acid bacteria are classified into homofermentative and heterofermentative organisms based on their ability to ferment
carbohydrates [56]. The homofermentative lactic acid bacteria such as Lactococcus and Streptococcus yield two molecules of lactates
from one glucose molecule whereas heterofermentative lactic acid bacteria such as Leuconostoc, Wiessella and some lactobacilli
generates lactate, ethanol and carbon dioxide from one molecule of glucose [54].

2.2.2 Taxonomical Classification

In recent taxonomic classification, lactic acid bacteria comes under the Phylum of Firmicutes, class Bacilli, and order Lactobacillales
which comprised of various different genera but the major genera includes Lactobacillus, Lactococcus, Weissella, Pediococcus,
Enterococcus, Streptococcus, Melissococcus, Lactosphaera, Leuconostoc, Carnobacterium, Oenococcus, Vagococcus, and
Tetragenococcus (Fig.1). Other genera include Aerococcus, Microbacterium, Propionibacterium, and Bifidobacterium [22, 55].
However, the largest genus in this group is the lactobacillus and it consists of more than 80 recognized species such as Lactobacillus
acidophilus, Lactobacillus plantarum, Lactococcus lactis cremoris, Bifidobacterium bifidum, Lactobacillus Casei, Lactobacillus
rhamnosus, Lactobacillus delbrueckii bulgaricus, Lactobacillus fermentum, Lactobacillus reuteri, Lactococcus lactis lactis,
Bifidobacterium infantis, Bifidobacterium breve, Enterococcus faecalis, Enterococcus faecium, Bifidobacterium adolecentis, and
Bifidobacterium longum [54, 57].

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Figure 1. Differentiation of Species according to the recent Taxonomy

2.3 Lactic Acid Bacteria in Fermented Foods

Lactic acid bacteria are the imperative groups of microbes in food fermentation and processing. Lactic acid bacteria have been
extensively used for the processing of many fermented food products such as kefir, cheese, butter, yogurt, sauerkraut, buttermilk,
brined vegetables, sourdough, soya curd, koumiss, idly batter, uttapam, fermented meat, and beverages (TABLE I). It is now well
established that microbes, especially lactic acid bacteria (LAB) which contribute more to the properties of foods, mediate the
fermentation process. LAB produces lactic acid and some other organic acid during sugar fermentation, which results in the reduction
of pH of the environment and thereby inhibiting the growth of undesired microbial agents. They also contribute for the preservation
of the food by producing various secondary metabolites like lactic acid, fatty acid, and bacteriocin by inhibiting the growth of spoilage
and pathogenic bacteria [47, 52, 56-60, 81].

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TABLE I. Types of Fermented Food products and Beverages and their associated Lactic Acid Bacteria
Category of fermented products Different species of lactic acid bacteria
Dairy based products
Butter and buttermilk Lactococcus lactis ssp. lactis, Lactococcus lactis ssp. Lactis var. diacetylactis,
Lactococcus lactis ssp. cremoris, Leuconostoc mesenteroides ssp. Cremoris

Yogurt Lactobacillus delbrueckii ssp. bulgaricus, Streptococcus thermophiles

Ice cream Lactobacillus johnsonii, Lactobacillus acidophilus, Lactobacillus delbrueckii subsp.

bulgaricus and Bifidobacterium bifidum
Fermented probiotic milk, Dahi (Curd), Lactobacillus casei, Lactobacillus acidophilus, Lactobacillus rhamnosus, Lactobacillus
and Lassi johnsonii, Bifidobacterium lactis, Bifidobacterium bifidum, Bifidobacterium breve
Kefir Lactobacillus kefir, Lactobacillus kefiranofacies, Lactobacillus brevi
Cheeses Lactococcus lactis ssp. lactis, Lactococcus lactis ssp. cremoris, Lactococcus lactis ssp.
lactis, Lactococcus lactis ssp. Lactis var. diacetylactis, Lactococcus lactis ssp. cremoris,
Leuconostoc mesenteroides ssp. cremoris, Lactobacillus delbrueckii ssp. lactis,
Lactobacillus helveticus, Lactobacillus casei, Lactobacillus delbrueckii ssp. bulgaricus,
Streptococcus thermophiles

Fermented vegetables
Pickles Leuconostoc mesenteroides, Pediococcus cerevisiae, Lactobacillus brevis, Lactobacillus
plantarum
Fermented olives Leuconostoc mesenteroides, Lactobacillus pentosus, Lactobacillus plantarum
Sauerkraut Leuconostoc mesenteroides, Lactobacillus plantarum, Pediococcus acidilactici

Fermented cereals
Sourdough, Idli and Dosa Lactobacillus sanfransiscensis, Lactobacillus farciminis, Lactobacillus fermentum,
Lactobacillus brevis, Lactobacillus plantarum, Lactobacillus amylovorus, Lactobacillus
reuteri, Lactobacillus pontis, Lactobacillus panis, Lactobacillus alimentarius, Weissella
cibaria
Alcoholic beverages
Wine (malolactic fermentation) Oenococcus oeni
Rice wine Lactobacillus sakei

Other fermented products

Soysauce Tegragenococcus halophilus
Soymilk Lactobacillus acidophilus, Lactobacillus casei, Lactobacillus gasseri, Lactobacillus
plantarum , Bifidobacterium
Fish Lactobacillus alimentarius, Carnobacterium piscicola

Fermented sausage Pediococcus acidilactici, Lactobacillus sakei, Lactobacillus curvatus, Pediococcus

pentosaceus
Tomato, orange, and grape juice Lactobacillus plantarum, Lactobacillus acidophilus, Lactobacillus casei, Lactobacillus
rhamnosus GG, Lactobacillus paracasei

III. PROBIOTICS: CHARACTERISTIC AND SELECTION CRITERIA

Recently, LABs have enticed much consideration to be used as a biopreservative constituent mainly in dairy and food industries and
also used as an analog of antibiotic in human therapeutics treatment by modulating the human host system against the various food-
borne pathogens and thus leading to the coining of the term “Probiotics”. Probiotics are the living microorganisms that are consumed
in an ample quantity as a part of nutrition (directly or indirectly), producing a beneficial effect by hindering the growth of detrimental
microbes and refining the microbiota balance in the gastrointestinal system of the host (human and animal). LABs are the most
extensively studied microorganism for human health as probiotics and further, most of the LABs have been considered as GRAS
(Generally Recognized as Safe) microorganism (8, 28, 50).

Good probiotics must fulfill the subsequent criteria and functional properties (Fig.2) for producing a beneficial effect on the human
health which includes:
 It must be safe for human use i.e., non-pathogenic and non-toxic;
 It must survive and show resistance to gastric juices and bile toxicity;
 It must adhere to and colonize the Gastrointestinal tract and Urinary tract;
 It must remain stable and viable for a long period of time during storage and food fermentation process;

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 It must produce antimicrobial substances or antimicrobial peptides; and

 It must modulate immune responses in the host.

Figure 2. Shows important Paradigms for the selection of Probiotic LABs

Some important selection criteria are discussed below:

3.1 Origin

The probiotic LAB can be isolated from different sources such as fermented foods, animal and human but to use as a probiotic strain
for the human purpose than it should be isolated from human microflora system which will have the ability to adhere more in the
human intestinal cell walls than the others and also must be safe. However, there are several potent probiotic LAB which has been
used to develop commercial probiotic products such as Lactobacillus casei and Lactobacillus plantarum isolated from the cheese and
have shown good and prominent impact human health [4, 9, 35, 58].

3.2 Biosafety

The assortment of potent probiotics needs to endure a safety assessment which includes taxonomic classification (phenotypic and
genetic methods) which will lead to the identification of the origin, habitat, and physiology of the strain. However, according to
WHO, all the strains which are considered to be probiotic should be identified and must be deposited in recognized culture collection
(National and International) before use [4, 27]. Further, well-designed protocol and experiments must be designed to administer
probiotic into human and animal model; must endure host specific interaction studies between probiotic and the host (in vivo and in-
vitro) must be done; and finally, it must not show any side effects while taking orally or administer along with foods [10, 28, 32, 39,
40, 44]. In 2002, FAO and WHO clarify that the source of the microorganism is not so essential criteria instead of that action of the
probiotic strain on the human health is more important criteria for the selection of the probiotic strains. Further, they collaborated
together and developed guidelines for the selection and methods to identify probiotics which are outlined in Fig. 2 and Fig. 3.

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Figure 3. FAO/WHO (2002)-Guidelines for the Assortment of Probiotic strains

3.3 Stability and Viability

Another important attribute for the selection of the potent probiotics is to determine the efficacy of the probiotics strain. Here, efficacy
relates to the stability (both physiologically and genetically); high growth in number; good sensory properties; and viability of the
strain throughout the food processing step, storage, and till the end of the shelf life of the products; and must survive and colonize
while passing through the gastrointestinal tract. However, dose-response studies must be done on human or animal models. Probiotics
products are usually recommended to store at 4℃ to 5℃ and must be used before the date of expiration of the product [27, 40, 58].

3.4 Resistance to bile salt and low pH environment

According to definition, probiotic isolates must survive, prevail and tolerate the stress environment such as low pH (acidic condition)
in the stomach and high bile salt concentration in the small intestine including various hydrolytic enzymes and thereby passing the
Gastro-Intestinal Tract (GIT) and colonizing the gut epithelial tissue of the host by competing with the other bacteria and thus,
produces antagonistic effect against harmful pathogens [62, 61]. Grosu-Tudor and Zamfir (2012) have isolated six potent probiotic
LABs strain from fermented vegetables and evaluated each strain for acid and high bile salt concentration tolerance. All the six strains
found to be resistant to pH 3.0 and pH 4.0 with viability rates of 109 CFU/ml and 1012 CFU/ml after 24 hours of incubation,
respectively. Likewise, all the six isolates also revealed a high resistance to bile salt concentration of 0.3% (w/v) with the viability
rates of 105 -108 CFU/ml after 24 hours of incubation [63]. Moreover, it has been reported that the survival rate of ingested probiotics
against bile salt and low pH tolerance varies with strain to strain for example Bifidobacteria and Lactobacillus acidophilus can
efficiently pass through the gut region without getting diminished in their concentration and LABs strains which produce high
exopolysaccharide such as Streptococcus thermophiles, have higher tolerance to bile salts as compared to Lactobacillus delbrueckii
subsp. Bulgaricus strains [6, 8, 79, 80].

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3.5 Adhesion and Colonization to Gastro-Intestinal Tract (GIT)

The capability to adhere to intestinal surfaces is one of the most vital criteria for selection of the potent probiotics isolates because
adhesion to the intestinal epithelial tissue is considered to be a criterion for colonization and thus producing health benefits by killing
the detrimental pathogens and balancing the microbiota balance in GIT. The adhesion of LAB in the intestinal epithelial tissue is
related to a good adherence ability, flow-rate of the LAB through the gut and presence of mucins to trap, protect and lubricate the
intestinal surfaces [64]. Furthermore, adhesion of LAB may involve the binding of LABs to a specific cell surface receptor or through
hydrophobic interaction between cell surface of LABs and intestinal cell surface or it may bind to extracellular matrix such as
fibronectin, collagen, and vitronectin [65] and this adhesion properties can be measured by using adhesion to hydrocarbons, salt
aggregation test (SAT) and the contact angle method [51, 65]. The colonization of probiotics must attribute to health-promoting traits
to the host like the production of vitamins, amino acids, enzymes, lactic acids, and natural antibiotics or antimicrobial
peptides/compounds [46]. The LAB which has genes responsible for the degradation and utilization of simple sugars and complex
carbohydrates are able to better survive and multiply in the gut [67, 68]. Most of the article has suggested that the LABs residing in
the intestine of the host are able to restrict the adhesion of enteropathogenic bacteria to intestinal epithelial cell surfaces by producing
more intestinal mucins such as Lactobacillus plantarum, found to inhibit the attachment of enteropathogenic Escherichia coli by
increasing the level of mRNA expression of some mucins [69-71]. Some prominent probiotic strains have been isolated from
fermented foods shows substantial adherent property such as Lactococcus lactis IS-16183 and Lactobacillus rhamnosus IS-7257
from dadih, which has restrained the adhesion of E. coli O157: H7 from adhering to mucus layers and Caco-2 cells. Furthermore,
these two strains have shown significant and relatively comparable adhesion properties than the commercial probiotics strains such
as Lactobacillus casei Shirota and Lactobacillus rhamnosus GG [6]. Another article suggested that Lactobacillus plantarum MB427
along with Lactobacillus plantarum IS-10506 isolated from a fermented fruit called as “mandai” also shown a good adhesive
properties on HCT-116 cells and also found to inhibit the adhesion of enteropathogens like E. coli, Salmonella enterica ATCC 14028,
and Listeria monocytogenes ATCC 13932 to mucus, significantly [72].

3.6 Immunomodulatory effects

Usually, foreign microorganisms invade the host by the process of translocation through the epithelial layer or through the Peyer's
patches and finally, penetrates into the GUT wall. LAB from the genera Lactobacillus is mostly present in the probiotic foods and
beverages and they are able to cross the intestinal mucous layer where they survive and elucidate the immune response of the host.
Most studies have revealed that the potent probiotic strains are able to modulate and regulate the innate and acquired immune
responses against a various infectious pathogen, some allergic reaction, inhibiting the pro-inflammatory mediators, and also helps in
increasing the immunogenicity of vaccine which is orally administered such as Polio, Cholera, Influenza, etc [4, 9, 28].
Probiotics can influence human health in three stages:
i. By way of Colonizing, interacting and eliminating microorganisms present on the site of action;
ii. By way of increasing the synthesis of mucus layer; and
iii. Finally, modulating the immune system of the host.
There are numerous species of Lactobacillus which have been identified to colonize in a GUT and modulate the immune
system of the host and these are Lactobacillus casei, Lactobacillus acidophilus, Lactobacillus rhamnous, Lactobacillus salivarius,
and Lactobacillus brevis. These LAB can modulate the innate and acquired immune responses by binding to specific receptors present
on the immune cells and other tissues such as intestinal epithelial tissue and induces production of chemokines, cytokines, regulatory
T cells, activation of dendritic cells and macrophages and also stimulate the production of specific antibodies and mucous [6, 8, 41,
46].
A number of studies have reported that LAB significantly modulates the host immune responses and also produces
antimicrobial substances against various foodborne pathogen and also prevent from causing diarrhea. As most of the fermented foods
are enrich in LAB which provides good texture to the food, therefore, consumption of these fermented food results in the production
of lactoferrin, bioactive peptides, flavonoids and many dietary compounds which help in maintaining the GUT microflora [35, 46].
Other studies suggested that potent probiotic LAB present in the fermented foods aids in the production of bioactive compounds such
as antioxidants and anti-inflammatory, neuropeptides, agmatine, and polyamines that modulates and also benefits brain health [6,
27]. However, all these studies highlight the roles and impacts of probiotic foods and beverages on human health but there is a major
gap in understanding of the interaction of the probiotic in in vivo (human and animal) modal and this is because of the complexity of
the GUT ecosystem, generation of varieties of probiotic strains from different sources and also each strain may modulate the host
immune system in different mode, and further, the immune modulation mechanism is not fully understood and explicated. Therefore,
more concise and precise research is required to reveal the immuno-modulatory effect of the probiotic strains on the human
physiology [32, 40, 67, 69, 71].

3.7 Antibiotic Resistance Pattern and Stability/ Transferability

The probiotic LAB can be used to kill the opportunistic harmful antibiotic-resistant pathogen by proliferating and colonizing into
GUT but to proliferate in the GUT it must be resistant to the common antibiotics by having a wide variety of antibiotic-resistant
genes. This antibiotic-resistant genes must be stable and must not get a transfer to the GUT pathogens as it will lead to a catastrophic
effect on the human health. Therefore, it is important to evaluate the stability or transferability rate of the antibiotic resistance genes
of the probiotic strains. Numerous antibiotic resistant LAB strains have been isolated and reported from different sources [73-77].
Safety assessment and sorting of the antibiotic resistant probiotic LAB can be done by following methods:

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• Determining the Antibiotic Susceptibility Test with the commonly used antibiotics for each LAB strains;
• Using standard PCR (Polymerase Chain Reaction) method;
• Determining the expression of the target gene by using Real-Time Quantitative PCR (RTqPCR); and
• Using Genome sequencing and Microarray methods.

3.8 Antimicrobial activity

It has been well documented that probiotic LAB shows very good anti-bacterial, anti-fungal, and anti-viral properties by producing
numerous anti-microbial compounds which include lactic acid, acetic acid, propionic acid, alcohol, and diacetyl. These antimicrobial
compounds produced by the LAB, interacts with the cell membranes of the harmful pathogens and inhibits their growth by
intracellular acidification (lowering the pH) and denaturing the proteins of the membrane which leads to the dysfunction of the
membrane permeability. In addition to these antimicrobial compounds, LAB also produces “Bacteriocins” which are ribosomally
synthesized proteinaceous toxins which inhibit the growth of similar or closely related bacteria by making pores in the membrane of
the pathogenic bacteria and thus, disrupting the permeability of the membrane (TABLE II) [2, 16, 33, 43, 50].

TABLE II. Antimicrobial Substances produced by Lactic Acid Bacteria

Antimicrobial Mechanism and Target Microorganisms
compounds/metabolites

Lactic acid, Acetic acid, It inhibits a wide range of both gram-positive and gram-negative bacteria, clostridia, molds,
Propionic acid, Reuterin, some fungi as well as yeast by interacting with the membrane, denaturing membrane protein,
Diacetyl, Fatty acid causing intracellular acidification, and disrupting membrane integrity.

Hydrogen peroxide It inhibits Pathogens and spoilage organism especially in protein-rich food by peroxidation
of membrane lipid and thus causes membrane dysfunction.
Antimicrobial Enzymes Mechanism and Target Microorganisms

Lactoperoxidase system It is a peroxidase enzyme which shows strong antimicrobial properties against pathogens
and spoilage-causing bacteria (milk and dairy products) by the addition of hydrogen
peroxide and thiocyanate.
Bacteriocins Mechanism and Target Microorganisms

Class I bacteriocins Ribosomally synthesized bacteriocins that consist of one or two peptides which include nisin
(Lantibiotics) and other lantibiotics that go through post-translational modification to have lanthionine, β-
methyllanthionine, dehydrated amino acid and some of them also have D-alanine. They are
small peptides (<5kDa) which are further alienated into two subgroups:
Group Ia: small cationic peptide which is flexible, amphipathic in nature, and having a
screw-shaped structure that makes pores in the membrane by unspecific interaction and thus
disrupting membrane permeability.
Group Ib: globular shape peptides which are usually neutral or anionic in nature.

Class II bacteriocins (Non-
lantibiotics)
Ribosomally synthesized low molecular weight peptide (<10kDa) which does not have any
modified amino acid. They are heat stable and are generally produced as an inactive form.
They become active when its N-terminal leader peptide is post-translationally cleaved. They
are further sub-classified into different sub-groups:
Group IIa: It consists of single peptide with strong anti-listerial activity and have
consensus sequence YGNGGVXC near N-termini. It shows a broad range of anti-bacterial
activity.
Group IIb: This class of bacteriocins is called two-peptide bacteriocin because it requires
two different peptides for anti-bacterial activity.
Group IIc: These are circular bacteriocins that have a broad range of effects on target
bacterial cells such as disruption of membrane permeability and cell wall formation.

Class III bacteriocins This class of bacteriocins are large, heat labile, antibiotic in nature-proteins and are least
(Non-lantibiotics) characterized with a molecular mass greater than 30kDa.

Class IV bacteriocins This class of bacteriocins is complexed with lipids or carbohydrates moieties which is
essential for their antimicrobial activity and are comparatively hydrophobic in nature and
heat stable.

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3.9 Quality Control Aspects

Before claiming the probiotic strains to be beneficial for the human health, it must go through the common quality parameter designed
by the FAO and WHO which includes: genetic identification of the claimed probiotic strains must be verified; high growth in number
i.e., Colony Forming Units (CFU) per gram of the final products must be verified of each strain; stability and viability must be
verified, purity of the selected strain must be verified; and proper labeling of the probiotic products must be done such as detail
information of the nutrition supplement; manufacturing date-Lot number-expiration date must be specified; genus, species, strain
information must be specified; proper storage condition and shelf life of the strain must be specified; minimum viability of the stain
at the end of the shelf life of the product must be specified; and complete contact and address details of manufacturer must be
mentioned in the probiotic products [9, 10, 13, 28, 40]. Some commercialized probiotic strains are outlined in TABLE III.
TABLE III. List of commercialized Probiotic strains explored by various Industries/Institutes

Probiotic Commercialized Strains Country Company/ Institute Name

Lactobacillus rhamnosus GG Helsinki, Finland Valio Dairy
Lactobacillus johnsonii Lal Lausanne, Switzerland Nestle
Lactobacillus casei Shirota Tokyo, Japan Yakult
Bifidobacterium breve Yakult
Lactobacillus acidophilus NCFM Madison, USA Rhodia
Lactobacillus casei CRL-43i Gilliland(La- Wisconsin, USA Chr. Hansens
Mo)
Lactobacillus reuteri SD 2112 Wisconsin, USA BioGaia
Lactobacillus plantarum 299V Lund, Sweden Probi
Lactobacillus Rhamnosus 271
Lactobacillus caseri DN01-4001 (Immunitas) Paris, France Danone Le PlessisRobinson
Streptococcus thermophiles 1131, Tokyo, Japan Meiji milk products
Lactobacillus acidophilus SBT-2062,
Lactobacillus delbrueckii subsp.,
Bulgaricus 2038
Saccharomyces boulardii Seattle, USA Biocodex
Bifidobacterium longum BB536 Zama-City, Japan Moringa milk industry Co., Ltd

Lactobacillus acidophilus LA-1, Horsholm, Denmark Chr. Hansen

Lactobacillus paracasei CRL 431,
Bifidobacterium lactis Bb-12
Lactobacillus acidophilus SBT-2062, Tokyo, Japan Snow Brand Milk Products Co.,
Bifidobacterium longum SBT-2928, Ltd
Bifidobacterium longum SBT-2928
Lactobacillus acidophilus R0011, Montreal, Canada Institute Rosell
Lactobacillus rhamnosus ROO52
Lactobacillus acidophilus DDS-1 Lincoln, NE Nebraska
Lactobacillus fermentum RC-14, London, Ontario, Urex Biotech Inc
Lactobacillus rhamnosus GR-1 Canada

Lactobacillus johnsonsii La1 Lausanne, Switzerland Nestle

Lactobacillus rhamnosus LB21, Umea, Sweden Essum AB
Lactococcus lactis L1A
Lactobacillus salivarius UCC118 Cork, Ireland University College Cork
Lactobacillus acidophilus LB Houdan, France Lacteol Laboratory
Lactobacillus paracasei F19 Stockholm, Sweden Arla Dairy
Lactobacillus crispatus CTV05, Seattle, USA Gynelogix, Boulder, Co
Lactobacillus casei DN114,
Saccharomyces boulardii

Other selection criteria for assessment of potent probiotic LABs includes: production of organic acids and hydrogen peroxide, anti-
mutagenic, enhancing metabolic activities such production of vitamins, reducing cholesterol, anti-carcinogenic, anti-hemolytic
activity, mucin degradation, and reducing lactose intolerance [8, 40, 43, 51].

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IV. BENEFICIAL EFFECTS OF THE PROBIOTIC LAB ON HUMAN HEALTH

Trillions of microorganisms (“microbiota/microflora”) have been found to be colonized in the intestine and the gut of the mammalian
system, which are vital for the human health. Lactobacillus, Pediococcus, Bifdobacterium, Lactococcus, Streptococcus, and
Leuconostoc are the most extensively isolated organisms from the fermented foods, beverages and also from the human and animal
GUT [8, 32].
LABs exhibit potential human health benefits such as reducing serum cholesterol [25], alleviate lactose intolerance [32,
40], possess antitumor activity [3, 19, 23, 32] and have an immune stimulatory effect [6]. Due to these beneficial properties, the
research in isolation and characterization of the LAB from several food sources and their applicability in the food industry have
exaggerated in recent years. Numerous health promoting LAB strains (such as Bifidobacterium sp., Lactococcus sp. and
Lactobacillus sp.) have been found that have shown optimistic consequences upon human health [47]. Thus, consumption of probiotic
LAB will have the following advantages on human health: Assist in maintaining the healthy microbiota balance of the intestine by
competing and excluding harmful pathogen by adhering to the active site [4, 35]; Enhancing the immune responses by increasing the
release of serum antibodies (IgM, IgG and IgA) and balancing pro-inflammatory and anti-inflammatory cytokines [27, 34, 40, 41];
Decreasing the duration and preventing intestinal diseases such as Inflammatory Bowel Disease, diarrhea and constipation by
colonizing and modulating GUT microflora, synthesizing antimicrobial compounds, Enhancing secretion of mucus and immune
responses [9, 13]; Helps in prevention of metabolic disorder such diabetes, obesity and cardiovascular diseases by enhancing gut
microbiota, restoring antioxidant system, decreasing insulin resistant and inflammation [34, 38]; Inhibit the growth of Candida and
Helicobacter pylori to prevent intestinal infection by competing and adhering to the mucosal surface and enhancing immune
responses [27]; Preventing cancer growth by acting as an therapeutic agent for cancer treatment by detoxification of chemical
carcinogens, decreasing the release of toxic metabolites, enhancing antioxidant system, modulating immune response to inhibit self-
proliferation of cancer, generating metabolites like butyrate which improves cancer cell death (apoptosis) and producing anti-
mutagenic effect [11, 14, 18, 20, 34, 36, 51]; Assist in maintaining and alleviating lactose intolerance by providing β-galactosidase
(Lactase) enzyme which breakdown the lactose into its constituents [27, 38, 45]; Benefits in lowering the cholesterol level by
precipitating cholesterol with free bile salts into bile acids and thereby reducing cholesterol absorption [20, 37, 42]; Enhances
absorption of minerals such as calcium to prevent mineral deficiency diseases such as osteoporosis [40]; Enhances nutritional value
by synthesizing cofactors and vitamins (K and B); and also produces several enzymes which are helpful for digestion of the food [28,
32, 38, 46].
Further, probiotic LAB act as bio-therapeutic agent for the treatment of Ulcer [17]; Allergic Disorders [15]; Urogenital
Infection [29]; Reducing Blood Pressure [48]; Kidney Stones [24]; AIDS and Leukemia [38]; and fungal diseases [21, 32].

V. FUTURE PERSPECTIVES

LAB from the genera Lactobacillus is mostly present in the probiotic foods and beverages and they are able to cross the intestinal
mucous layer where they survive and elucidate the immune response of the host. Most studies have revealed that the potent probiotic
strains are able to modulate and regulate the innate and acquired immune responses against a various infectious pathogen, some
allergic reaction, inhibiting the pro-inflammatory mediators, and also helps in increasing the immunogenicity of vaccine which is
orally administered such as Polio, Cholera, Influenza, etc. As consumers are becoming more aware of the usage of probiotics as a
therapeutic agent, there is an increasing demand for the probiotic supplements in the last few decades. Further, research on this area
is growing rapidly that leads to the evolving of new probiotic LAB strains from different sources which must be regulated more
stringent manner for the safety of patients and to avoid consumer misleading of the probiotic products globally. Therefore, there is a
need for well-designed code of practice regarding selection of the probiotic LAB strains such as origin, strain identification, and
safety properties must be studied; stability and viability during the product manufacturing must be studied; Functional and
Technological properties must be tested (in vitro and in vivo); Mode of action must be studied; Optimal dosage and duration of the
specific treatment must be known; and well documented evidence of the health benefit claimed by the usage of specific probiotic
strain must be generated. However, in addition to this traditional selection criteria, novel methods are needed to isolate and
characterize target or site-specific probiotic strains with help of a Genomic database. These target or site-specific probiotics are called
as third generation (micro-encapsulated) probiotics and fourth generation (dual coated – pH-dependent release system, moisture,
temperature, and pressure protection) probiotics. Therefore, there is a need for new selection paradigms and guidelines to evaluate
these target or site-specific probiotics which may emerge in the near future.

VI. ACKNOWLEDGMENT

Mr. Rahul Gupta is grateful to UGC for providing UGC Research Fellowship and I am also grateful to UGC-SAP for funding the
research facilities to the Department of Biochemistry & Molecular Biology, Pondicherry University, India.

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