


































Food Science and Nutrition Studies 

ISSN 2573-1661 (Print) ISSN 2573-167X (Online) 

Vol. 3, No. 2, 2019 

www.scholink.org/ojs/index.php/fsns 

73 
 

Original Paper 

Selection and Characterization Criteria of Probiotics Intended 

for Human Use from the Past to the Future 

Valeria Sagheddu1*, Elena Guidesi1, Serena Galletti1 & Marina Elli1 

1 AAT-Advanced Analytical Technologies, Fiorenzuola d’Arda, Piacenza, Italy 

* Valeria Sagheddu, AAT-Advanced Analytical Technologies, Fiorenzuola d’Arda, Piacenza, Italy 

 

Received: April 28, 2019         Accepted: May 10, 2019        Online Published: May 20, 2019 

doi:10.22158/fsns.v3n2p73        URL: http://dx.doi.org/10.22158/fsns.v3n2p73 

 

Abstract 

The probiotic product consumption has recently increased with the prevalent intent to promote human 

and animal wellbeing. The complex selection process dealing with new-isolated probiotic candidates is 

the first challenge that has to be faced. From the isolation to the launch on the market, information 

about safety, tolerance to host physiological conditions, adhesion properties, genetics and interaction 

with the host has to be collected. Probiotics must be safe, survive to the exposition to bile salts and to 

gut transit, adhere to intestinal cells lining and colonize the lumen of the tract. The evaluation process 

of the possible probiotic health benefits is widely supported by in-vitro assays simulating the in-vivo 

conditions. The aim of this work is to summarize the classical models usually employed for the 

probiotic screening by underlying strengths and weaknesses of all models and to present some more 

recent analysis tools used in the probiotic field. The long term goal in new probiotic candidate 

selection experiencing these combined essays together would lead to the hypothetical assignment 

acknowledged as one strain-one function.  

Keywords 

screening, platform, in-vitro, in-vivo, model 

 

1. Introduction  

International Scientific Association for Probiotics and Prebiotics defined probiotics as “live 

microorganisms that, when administered in adequate amounts, confer a health benefit on the host” (Hill 

et al., 2014). In the recent years the global probiotic market has faced a massive growth, this expansion 

should be related to a robust and univocal legislation (Baldi & Arora, 2015). Nevertheless, focusing the 

attention on the European regulatory framework, it has to be noticed the absence of harmonized 

regulation. The Food Products Directive and Regulation (Regulation 178/2002/EC; Directive 



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2000/13/EU) and Herbal Medicinal Products Directive (2004/24/EC) are the main laws supervising 

probiotic products (Coppens et al., 2006). Furthermore, the European regulatory pressure in the field of 

probiotics led to a double negative impact by limiting the communication of their beneficial effects to 

consumers and by adversely impacting the market. After ten years from the enter into force of the 

Regulation on health claims made on food (EU 1924/2006), the market of probiotics shows a renewed 

interest about the health improving potential of beneficial microbes. The restored enthusiasm is mainly 

driven by the big pool of information derived from the omics approach for the study of the gut 

microbiota (Baugher & Klaenhammer, 2011). Unfortunately, the legislative framework seems to not 

run with the innovation associated with new analytical approaches. Actually, of the about 300 calls for 

health claims submitted to EFSA experts’ panel on the food area none has been accepted. The only 

exception was represented to the use of yoghurt for lactose intolerance (Papadimitriou et al., 2015). The 

implementation of new and comprehensive models, exploiting the newest omics technology could 

ameliorate the understanding of the probiotics mechanism of action and assessing the beneficial impact 

on health exerted by the administration of probiotics at a quantifiable level (Rebollar et al., 2016). The 

integration of the traditional in-vitro and ex-vivo screening methods of microorganisms with potential 

probiotic features are associated with omics techniques with the intent to boost the selection process, to 

deeper understand their functionality and to examine the opportunity to employ cell fractions instead of 

viable probiotics in finished products (Franzosa et al., 2015; Lahtinen, 2012). Recent market trends 

concerning probiotic are interestingly targeted to the zootechnic field. For instance, it has been 

proposed the probiotic use to produce safe and high-quality poultry meat (Park et al., 2016). Other 

application fields could be retrieved in cereals, dairy products, fermented meat, baked food, and dry 

food productions (Competitive Market Share & Forecast, 2016). For the application related to human 

consumption, the validated approach applied to the selection of probiotics firstly includes the safety 

assessment intended to verify the absence of pathological and virulence traits (Hill et al., 2014). The 

next step evaluates the potential application of the isolated strain by considering the prophylactic, 

therapeutic use or its ability to reduce the risk to develop a certain pathological condition. Regrettably, 

the implication of this validation results strictly dependent on the availability of measurable biomarkers 

related to a certain physiological condition.  

 

2. Classical in-vitro Approaches for New Probiotics Screening 

Guidelines for the evaluation of the probiotic use in food have been published several years ago 

(FAO/WHO, 2002) The principal focus identified was defined as a selection scheme for the 

conventional assessment of probiotic properties and for the characterization of functional bacterial 

strains as probiotics in food. Despite their time-consuming application, the in-vitro assays are routinely 

used due to their presumed predictive value to predetermine, the putative association between the 

isolated probiotic candidate strain and a specific health claim, targeting its potential general or specific 

application (Kumar et al., 2013). The major limitation of these assays is represented by the high 



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variability in terms of reproducibility even if a certain degree of standardization has been suggested. 

The natural continuation of these essays would be theoretically represented by in-vivo tests that are 

laborious, expensive and not feasible for ethical reasons.  

  

3. Persistence and Survival in the Human Gut 

A peculiar required probiotic feature is represented by the ability to survive the passage through the 

gastrointestinal tract remaining in a viable state and with the sufficient number of cells necessary to 

exert the probiotic effect. To verify this ability simulated gastric and pancreatic juices (Charteris et al., 

1998; Lavermicocca et al., 2008) and porcine or bovine bile have been extensively used, in fact these 

assays are easy and cheap if compared to more recent approaches including GIT simulators. 

Unfortunately, those in-vitro approaches are affected of some biases dealing with the unrealistic 

conditions simulated during benchtop assays. Moreover, potential probiotic candidates are tested 

against severe environmental parameters not really miming the stomach acidity and lower intestine 

conditions during food consumption (Papadimitriou et al., 2015).  

 

4. Safety for the Use in Human Nutrition 

The Joint Working Group FAO/WHO (2002) underlined the relevance of some safety issues for the use 

of probiotics in food. The EFSA panel of experts suggested the precise determination of the minimal 

inhibitory concentrations (MICs) towards 9 listed antibiotics and also stated the microbiological cutoff 

values to evaluate a new probiotic candidate as susceptible or resistant to antibiotics and the reference 

methodology to assess it (EFSA; 2012). Safety aspects usually assessed by in-vitro tests also include 

aggregation to platelets, to fibronectin and to fibrinogen, hemolytic activity, the synthesis of certain 

enzymes and the production of biogenic amines. The major concern related to these essays is based on 

the fact that the expression levels of the in-vivo traits could significantly vary from the in-vitro 

conditions inducing to an over- or an under-estimation of some vexing features of probiotics 

(Papadimitriou et al., 2015). 

 

5. Colonization Ability 

The colonization ability is an indicator of the probiotic cellular features to interact with the host. The 

adhesion of bacteria to mucus and epithelial cells is an important parameter evaluated with several 

in-vitro assays; this trait could both represent positive connotations associated with pathogens 

displacement and negative implications for instance the increasing risk of translocation (Sanders et al., 

2010). Furthermore, Caco-2, HT-29 epithelial cell adhesion assays often produce weak results due to 

the low reproducibility maintenance of the tested conditions. In order to circumvent the problem, 

resected gut tissue or whole tissue models including the mucus layer have been taken into consideration 

but reliability bias are still present (Vesterlund et al., 2005; van Tassell & Miller, 2011). To evaluate the 

ability of a strain to interact with the host, some physical features of probiotics have been considered, 



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for instance, the hydrophobicity of the cell surface (Yadav et al., 2014). Unfortunately, the scientific 

significance of these tests is only considered partially because of the lack of the peculiarity of this trait. 

The enzymatic activity expression such as GAPDH (glyceraldeyde-3-phosphate dehydrogenase) 

promoting the probiotic attachment to the mucus has been evaluated as a potential screening parameter 

even if with scarce results (Kinoshita et al., 2008). 

The assay conditions, even if optimized could not reproduce the complexity of the bacterial community 

structure and the possible interactions among different microorganisms. The adhesion assays are 

performed on a single and purified strain without considering different microbial species leading to 

doubts about the results, significance and their possible extensive transposition to the gut microbiota. 

 

6. Antimicrobial Potential 

It is known that probiotics could exert competitive inhibition towards pathogens exploiting several 

mechanisms, for instance the production of metabolites showing antimicrobial activity. This feature has 

been proven influence other bacteria, viruses and fungi at a large spectrum, taking advantage of 

different compounds like organic acids or bacteriocins (Denkova et al., 2013). Several in-vitro models 

have been experienced, from the agar diffusion test to more advanced systems targeting the release of 

antimicrobial substances using indicator strains (Szweda & Szweda, 2016). Probiotics exhibit other 

features as co-aggregation with pathogens and/or by reinforcing the integrity and functions of the gut 

inner barrier. Briefly, probiotic cells bind pathogens causing a sort of clumping effect limiting the 

pathogen interaction with the surfaces of the host and facilitating the excretion of pathogens trough 

biological fluids (e.g., saliva, feces) (Janković et al., 2012). The gut barrier could exert its function only 

if the tight junctions integrity is maintained, this condition represents a key issue to partially prevent 

the bacterial translocation, pathogens included. It has been proved that probiotics may improve the 

barrier integrity as measured by specific enzymatic activities (Putaala et al., 2008).  

The evaluation of probiotics antimicrobial potential is affected by some biases since the laboratory 

conditions only partially reproduce the in-vivo situation. Similarly to what has been observed for the 

described above in-vitro models, the level of expression of the real antimicrobial potential of tested 

probiotic strains is not completely defined. Substances as exopolysaccharides (EPS), biosurfactants, 

bacteriocins and organic acids represent the main antimicrobial compounds produced by probiotics 

(Evivie et al., 2017).  

 

7. Immunomodulatory Actions 

Probiotics could stimulate the secretion of antibodies by host cells and activate cell-mediated immune 

responses (Haghighi et al., 2005). Usually, immunomodulatory effects are promoted by M cells from 

the Payer’s patches or dendritic cells (DCs) and the response to the occurrence of bacterial cells is 

assessed by co-culture of probiotics and immune cells and by the detection and the quantification of 

cytokines (Gad et al., 2011). These co-incubation models are frequently applied to the selection of 



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probiotics showing anti-inflammatory features useful in the management of chronic inflammatory 

diseases (e.g., IBD) (Rutella and Locatelli, 2011) or to be employed as adjuvant in the treatment of 

allergic symptoms (Ozdemir, 2010). However, the center of the debate still remains the in-vivo 

modality of action of probiotics and on the possible solution to translate experimental findings to the 

concrete evaluation of the probiotic strains efficacy in the modulation of the gastrointestinal and 

systemic immunity. 

 

8. In-vitro Models of Cardiovascular Diseases 

Another trait that is required from probiotic candidates is represented by the expression of the bile salt 

hydrolase activity (BSH). This enzymatic activity allows the deconjugation of bile acids and it is pretty 

easy and quick to obtain qualitative and quantitative results (Cani & Van Hul, 2015; Zheng et al., 2013; 

Tomaro-Duchesneau et al., 2014). However, these routinely used models are sometimes considered 

relatively insufficient to estimate the real probiotics contribution to reduce cardiovascular diseases risks 

associated (Papadimitriou et al., 2015), albeit the beneficial effects seem to be highlighted by scientific 

evidences (Ebel et al., 2014).  

 

9. In-vitro Anti-cancer Models 

It has been proven that probiotics reinforce natural host body protection functions by improving 

metabolic and immunological parameters (Ashraf & Shah, 2014). Furthermore, these beneficial 

bacteria could exert antigenotoxic and antimutagenic activities and promote nitrosamines and 

heterocyclic amine degradation whose accumulation within the intestinal ecosystem could lead to 

potential negative effects (Duangjitcharoen et al., 2014; Faridnia et al., 2010). Excluding the ability of 

withdrawing dangerous metabolites, the data support the probiotics ability of releasing short chain fatty 

acids (SCFA) within the intestinal lumen (Table 1). These substances have been proven to exert some 

anticarcinogenic effects in-vivo (Burns & Rowland, 2004; Castro et al., 2010). Nevertheless, despite a 

supportive background is already available, researchers consider these in-vitro tests as not sufficient to 

provide consistent probiotic anticancer activity information and, for this reason, the in-vivo validation 

is compelled (Chong, 2014). 

To easily complete the probiotic profile of beneficial bacteria, a pool of other in-vitro model has been 

proposed, among them the measurement of B-galactosidase activity to monitor lactose intolerance 

symptoms (Vonk et al., 2012), B-group vitamin production (Leblanc et al., 2011), oxalate-degrading 

features (Abratt & Reid, 2010) and inhibition of pathogen oral strains responsible for volatile sulfur 

compounds (VSCs) production (Lee & Baek, 2014). 

 

 

 

 



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Table 1. Short Chain Fatty Acids (SCFA) Produced by Probiotics 

Probiotic microorganisms  Produced SCFA Reference 

L. salivarius spp salcinius JCM 1230, L. 

agilis JCM 1048 
Propionate/butyrate Meimandipour et al., 2010 

L. acidophilus CRL 1014 Acetate/butyrate/propionate Sivieri et al., 2013 

L. rhamnosus LGG Propionate Le Blanc et al., 2017 

L. gasseri PA 16/8 Propionate Le Blanc et al., 2017 

B. longum SP 07/3 Propionate/acetate Le Blanc et al., 2017 

B bifidum MF 20/5 Propionate/acetate Le Blanc et al., 2017 

Bifidobacteria Acetate/lactate Pessione et al., 2012 

 

10. Development of Innovative Screening Platforms 

The rationale underlying standardized in-vitro assays to study several aspects of probiotic effectiveness 

is driven by the necessity of set reproducible experimental protocols and consequently decrease the 

percentage of false positive and negative results. 

In-vivo models should be considered integrative for in-vitro ones with the purpose of choosing the 

appropriate probiotic for a potential application.  

Several animal models including invertebrates (e.g., insect, worm) and vertebrates (e.g., rodents, dogs, 

monkeys and swine), have been tested even though the predictivity for the situation in humans seems to 

be powerless and for this reason, a critical approach should be applied when results are transposed to 

humans. In the latest years the conventional screening pipeline for probiotics (in-vitro assays—animal 

model—clinical trial) including mice or rat models has strongly been debated and there was the 

tendency to directly check the findings from in-vitro evidences on small number human cohorts. 

However, animal models will certainly be exploited, particularly for studying specific pathological 

conditions, both induced by chemicals administration or using animal with genetic predispositions, or 

pathogenic infection conditions. Some probiotic positive modulation effects of intestinal inflammation 

have been successfully studied thanks to animal models. For instance, the co-occurrence of probiotics 

and carcinogenesis prevention, metabolic disorders control or auto-immunological conditions have 

been experienced. On the contrary, immune-deficient animals carrying human tissue or genes also 

known as humanized mice or rat models, are basically used more for pharmaceuticals purpose instead 

of probiotic validation (Papadimitriou et al., 2015).  

The combination of in-vitro and in-vivo test represents the new frontier, Europe coupled the two tests in 

some in-silico models to examine from a mathematical point of view the interaction between nutrients, 

epithelial cells, gut communities and host (Tan & Liong, 2014). These applications are supposed to 

mimic the complex gut environment processes by proposing predictive functions and taking in 

consideration the possible interactions between probiotics with multiple factors. 



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The omics approaches with the advent of new molecular methods greatly contributed to the 

comprehension of the usefulness of probiotic towards hosts, mainly quantifying the expression 

variation of certain housekeeping genes of probiotic cells in response to specific environmental 

conditions. The resistance to stressful conditions, modulation of the immune system, adhesion to host 

tissues, production of antimicrobial or noble nutrient compounds, degradation of prebiotics and quorum 

sensing are areas of interest investigated with special consideration (Papadimitriou et al., 2015).  

Omics technologies are assumed to explicate the complexity of the communication dynamics inside 

microbiome or between it and host tissues and to provide new insights on the impact of a foreign 

probiotic strain meeting those structured communities (De Melo Pereira et al., 2018). 

 

11. Conclusions  

The number of available assays to validate probiotic efficacy in-vitro and in-vivo is rather complex. The 

major limitations linked to those approaches are that they consider variables explanatory and not easily 

reproducible. As a consequence, often the efficacy of new-isolated probiotic candidates could be 

accomplished only by combining several assays.  

At the state of the art, on the one hand, there is an urging need of defining a clear workflow for the 

probiotics selection in order to obtain reliable and unquestionable proof of efficacy, and on the other, 

the design of appropriately designed clinical trials. Human studies are often associated with significant 

costs, consequently probiotics are still screened following the “funnel-like” approach, firstly using 

in-vitro tests and subsequently by expensive and appropriate validations on selected strains.  

The objective sought in new probiotic candidate selection would lead to the development of screening 

platforms, reducing from a large pool of strains to a restricted group intentionally selected for a specific 

human need to achieve the ideal aim of one strain-one function.   

 

References 

Abratt, V. R., & Reid, S. J. (2010). Oxalate-Degrading Bacteria of the Human Gut as Probiotics in the 

Management of Kidney Stone Disease (pp. 63-87).  

Ashraf, R., & Shah, N. P. (2014). Immune System Stimulation by Probiotic Microorganisms. Crit. Rev. 

Food Sci. Nutr., 54, 938-956. https://doi.org/10.1080/10408398.2011.619671 

Baldi, A., & Arora, M. (2015). Regulatory categories of probiotics across the globe: A review 

representing existing and recommended categorization. Indian J. Med. Microbiol., 33, 2. 

https://doi.org/10.4103/0255-0857.150868 

Baugher, J. L., & Klaenhammer, T. R. (2011). Invited review: Application of omics tools to 

understanding probiotic functionality. J. Dairy Sci., 94, 4753-4765. 

https://doi.org/10.3168/jds.2011-4384 

Burns, A. J., & Rowland, I. R. (2004). Antigenotoxicity of probiotics and prebiotics on faecal 

water-induced DNA damage in human colon adenocarcinoma cells. Mutat. Res., 551, 233-243. 



www.scholink.org/ojs/index.php/fsns                Food Science and Nutrition Studies                     Vol. 3, No. 2, 2019 

80 
Published by SCHOLINK INC. 

https://doi.org/10.1016/j.mrfmmm.2004.03.010 

Cani, P. D., & van Hul, M. (2015). Novel opportunities for next-generation probiotics targeting 

metabolic syndrome. Curr. Opin. Biotechnol. https://doi.org/10.1016/j.copbio.2014.10.006 

Castro, M. S., Molina, M. A., Di Sciullo, P., Azpiroz, M. B., Leocata Nieto, F., Sterín-Speziale, N. B., ... 

Manghi, M. A. (2010). Beneficial activity of Enterococcus faecalis CECT7121 in the 

anti-lymphoma protective response. J. Appl. Microbiol., 109, 1234-1243. 

https://doi.org/10.1111/j.1365-2672.2010.04747.x 

Charteris, Kelly, Morelli, & Collins. (1998). Development and application of an in vitro methodology 

to determine the transit tolerance of potentially probiotic Lactobacillus and Bifidobacterium 

species in the upper human gastrointestinal tract. J. Appl. Microbiol., 84, 759-768. 

https://doi.org/10.1046/j.1365-2672.1998.00407.x 

Chong, E. S. L. (2014). A potential role of probiotics in colorectal cancer prevention: Review of 

possible mechanisms of action. World J. Microbiol. Biotechnol., 30, 351-374. 

https://doi.org/10.1007/s11274-013-1499-6 

Coppens, P., Da Silva, M. F., & Pettman, S. (2006). European regulations on nutraceuticals, dietary 

supplements and functional foods: A framework based on safety Toxicology.  

De Melo Pereira, G. V., de Oliveira Coelho, B., Magalhães Júnior, A. I., Thomaz-Soccol, V., & Soccol, 

C. R. (2018). How to select a probiotic? A review and update of methods and criteria. Biotechnol. 

Adv. https://doi.org/10.1016/j.biotechadv.2018.09.003 

Denkova, R., Denkova, Z., Yanakieva, V., & Blazheva, D. (2013). Antimicrobial activity of probiotic 

lactobacilli, bifidobacteria and propionic acid bacteria, isolated from different sources (pp. 

857-864). 

Duangjitcharoen, Y., Kantachote, D., Prasitpuripreecha, C., Peerajan, S., & Chaiyasut, C. (2014). 

Selection and characterization of probiotic lactic acid bacteria with heterocyclic amine binding 

and nitrosamine degradation properties. J. Appl. Pharm. Sci., 4, 14-23. 

https://doi.org/10.7324/JAPS.2014.40703 

Ebel, B., Lemetais, G., Beney, L., Cachon, R., Sokol, H., Langella, P., & Gervais, P. (2014). Impact of 

Probiotics on Risk Factors for Cardiovascular Diseases. A Review. Crit. Rev. Food Sci. Nutr., 54, 

175-189. https://doi.org/10.1080/10408398.2011.579361 

EFSA. (2012). Guidance on the assessment of bacterial susceptibility to antimicrobials of human and 

veterinary importance.  

Evivie, S. E., Huo, G. C., Igene, J. O., & Bian, X. (2017). Some current applications, limitations and 

future perspectives of lactic acid bacteria as probiotics. Food Nutr. Res., 61(1), 1318034. 

https://doi.org/10.1080/16546628.2017.1318034 

FAO/WHO. (2002). Guidelines for the Evaluation of Probiotics in Food. Geneva, Switzerland: Food 

and Agriculture Organization (FAO), World Health Organization (WHO). 

Faridnia, F., Hussin, A. S. M., Saari, N., Mustafa, S., Yee, L. Y., & Manap, M. Y. A. (2010). In vitro 



www.scholink.org/ojs/index.php/fsns                Food Science and Nutrition Studies                     Vol. 3, No. 2, 2019 

81 
Published by SCHOLINK INC. 

binding of mutagenic heterocyclic aromatic amines by Bifidobacterium pseudocatenulatum G4. 

Benef. Microbes, 1, 149-154. https://doi.org/10.3920/BM2009.0035 

Franzosa, E. A., Hsu, T., Sirota-Madi, A., Shafquat, A., Abu-Ali, G., Morgan, X. C., & Huttenhower, C. 

(2015). Sequencing and beyond: Integrating molecular “omics” for microbial community profiling. 

Nat. Rev. Microbiol., 13, 360-372. https://doi.org/10.1038/nrmicro3451 

Gad, M., Ravn, P., Søborg, D. A., Lund-Jensen, K., Ouwehand, A. C., & Jensen, S. S. (2011). 

Regulation of the IL-10/IL-12 axis in human dendritic cells with probiotic bacteria. FEMS 

Immunol. Med. Microbiol., 63, 93-107. https://doi.org/10.1111/j.1574-695X.2011.00835.x 

Haghighi, H. R., Gong, J., Gyles, C. L., Hayes, M. A., Sanei, B., Parvizi, P., ... Sharif, S. (2005). 

Modulation of Antibody-Mediated Immune Response by Probiotics in Chickens. Clin. Vaccine 

Immunol., 12, 1387-1392. https://doi.org/10.1128/CDLI.12.12.1387-1392.2005 

Hill, C., Guarner, F., Reid, G., Gibson, G. R., Merenstein, D. J., Pot, B., ... Sanders, M. E. (2014). 

Expert consensus document: The International Scientific Association for Probiotics and Prebiotics 

consensus statement on the scope and appropriate use of the term probiotic. Nat. Rev. 

Gastroenterol. Hepatol., 11, 506-514. https://doi.org/10.1038/nrgastro.2014.66 

Janković, T., Frece, J., Abram, M., & Gobin, I. (2012). Aggregation ability of potential probiotic 

Lactobacillus plantarum strains. Int. J. Sanit. Eng. Res., 6, 19-24. 

Kinoshita, H., Wakahara, N., Watanabe, M., Kawasaki, T., Matsuo, H., Kawai, Y., ... Saito, T. (2008). 

Cell surface glyceraldehyde-3-phosphate dehydrogenase (GAPDH) of Lactobacillus plantarum LA 

318 recognizes human A and B blood group antigens. Res. Microbiol., 159, 685-691. 

https://doi.org/10.1016/j.resmic.2008.07.005 

Kumar, A., Kumar, M., Ghosh, M., & Ganguli, A. (2013). Modeling in vitro cholesterol reduction in 

relation to growth of probiotic Lactobacillus casei. Microbiol. Immunol., 57, 100-110. 

https://doi.org/10.1111/1348-0421.12008 

Lahtinen, S. J. (2012). Probiotic viability—Does it matter? Microb. Ecol. Heal. Dis., 23, 10-14. 

https://doi.org/10.3402/mehd.v23i0.18567 

Lavermicocca, P., Valerio, F., Lonigro, S. L., Di Leo, A., & Visconti, A. (2008). Antagonistic activity of 

potential probiotic lactobacilli against the ureolytic pathogen Yersinia enterocolitica. Curr. 

Microbiol., 56, 175-181. https://doi.org/10.1007/s00284-007-9069-5 

LeBlanc, J. G., Chain, F., Martín, R., Bermúdez-Humarán, L., Courau, S., & Langella, P. (2017). 

Beneficial effects on host energy metabolism of short-chain fatty acids and vitamins produced by 

commensal and probiotic bacteria. Microb. Cell Fact. https://doi.org/10.1186/s12934-017-0691-z 

Leblanc, J. G., Laiño, J. E., del Valle, M. J., Vannini, V., van Sinderen, D., Taranto, M. P., ... Sesma, F. 

(2011). B-Group vitamin production by lactic acid bacteria—Current knowledge and potential 

applications. J. Appl. Microbiol., 111, 1297-1309. 

https://doi.org/10.1111/j.1365-2672.2011.05157.x 

Lee, S.-H., & Baek, D.-H. (2014). Effects of Streptococcus thermophilus on volatile sulfur compounds 



www.scholink.org/ojs/index.php/fsns                Food Science and Nutrition Studies                     Vol. 3, No. 2, 2019 

82 
Published by SCHOLINK INC. 

produced by Porphyromonas gingivalis. Arch. Oral Biol., 59, 1205-1210. 

https://doi.org/10.1016/j.archoralbio.2014.07.006 

Meimandipour, A., Hair-Bejo, M., Shuhaimi, M., Azhar, K., Soleimani, A. F., Rasti, B., & Yazid, A. M. 

(2010). Gastrointestinal tract morphological alteration by unpleasant physical treatment and 

modulating role of Lactobacillus in broilers. Br. Poult. Sci. 

https://doi.org/10.1080/00071660903394455 

Ozdemir, O. (2010). Various effects of different probiotic strains in allergic disorders: An update from 

laboratory and clinical data. Clin. Exp. Immunol., 160, 295-304. 

https://doi.org/10.1111/j.1365-2249.2010.04109.x 

Papadimitriou, K., Zoumpopoulou, G., Foligné, B., Alexandraki, V., Kazou, M., Pot, B., & Tsakalidou, 

E. (2015). Discovering probiotic microorganisms: In vitro, in vivo, genetic and omics approaches. 

Front. Microbiol. https://doi.org/10.3389/fmicb.2015.00058 

Park, Y. H., Hamidon, F., Rajangan, C., Soh, K. P., Gan, C. Y., Lim, T. S., ... Liong, M. T. (2016). 

Application of Probiotics for the Production of Safe and High-quality Poultry Meat. Korean J. 

Food Sci. An, 36, 567-576. https://doi.org/10.5851/kosfa.2016.36.5.567 

Pessione, E. (2012). Lactic acid bacteria contribution to gut microbiota complexity: Lights and 

shadows. Front. Cell. Infect. Microbiol. https://doi.org/10.3389/fcimb.2012.00086 

Probiotics Market Size By End Use (Human, Animal). (n.d.). By Application (Functional Foods & 

Beverages [Dairy, Non-dairy, Cereals, Baked Goods, Fermented Meat Products, Dry Foods], 

Dietary Supplements [Food, Nutritional, Specialty, Infant Formula], Animal Feed Probiotics), 

Industry Analysis Report, Regional Outlook (U.S., Germany, UK, China, Japan, India, Brazil), 

Application Potential, Price Trends, Competitive Market Share & Forecast, 2016-2023. 

Putaala, H., Salusjärvi, T., Nordström, M., Saarinen, M., Ouwehand, A. C., Bech Hansen, E., & 

Rautonen, N. (2008). Effect of four probiotic strains and Escherichia coli O157:H7 on tight 

junction integrity and cyclo-oxygenase expression. Res. Microbiol., 159, 692-698. 

https://doi.org/10.1016/j.resmic.2008.08.002 

Rebollar, E. A., Antwis, R. E., Becker, M. H., Belden, L. K., Bletz, M. C., Brucker, R. M., ... Harris, R. 

N. (2016). Using “omics” and integrated multi-omics approaches to guide probiotic selection to 

mitigate chytridiomycosis and other emerging infectious diseases. Front. Microbiol., 7, 1-19. 

https://doi.org/10.3389/fmicb.2016.00068 

Rutella, S., & Locatelli, F. (2011). Intestinal dendritic cells in the pathogenesis of inflammatory bowel 

disease. World J. Gastroenterol, 17, 3761-3775. https://doi.org/10.3748/wjg.v17.i33.3761 

Sanders, M. E., Akkermans, L. M. A., Haller, D., Hammerman, C., Heimbach, J., Hörmannsperger, 

G., ... Vaughan, E. (2010). Safety assessment of probiotics for human use. Gut Microbes. 

https://doi.org/10.4161/gmic.1.3.12127 

Sivieri, K., Morales, M. L. V., Adorno, M. A. T., Sakamoto, I. K., Saad, S. M. I., & Rossi, E. A. (2013). 

Lactobacillus acidophilus CRL 1014 improved “gut health” in the SHIME® reactor. BMC 



www.scholink.org/ojs/index.php/fsns                Food Science and Nutrition Studies                     Vol. 3, No. 2, 2019 

83 
Published by SCHOLINK INC. 

Gastroenterol. https://doi.org/10.1186/1471-230X-13-100 

Szweda, P., & Szweda, P. (2016). World’s largest Science, Technology & Medicine Open Access book 

publisher Antimicrobial Activity of Honey Antimicrobial Activity of Honey.  

Tan, P. L., & Liong, M. T. (2014). In silico approaches for probiotic-derived bioactives. Trends 

Biotechnol, 32(12), 599-601. https://doi.org/10.1016/j.tibtech.2014.09.011 

Tomaro-Duchesneau, C., Jones, M. L., Shah, D., Jain, P., Saha, S., & Prakash, S. (2014). Cholesterol 

assimilation by Lactobacillus probiotic bacteria: An in vitro investigation. Biomed Res. Int., 2014, 

380316. https://doi.org/10.1155/2014/380316 

van Tassell, M. L., & Miller, M. J. (2011). Lactobacillus adhesion to mucus. Nutrients, 3(5), 613-636. 

https://doi.org/10.3390/nu3050613 

Vesterlund, S., Paltta, J., Karp, M., & Ouwehand, A. C. (2005). Adhesion of bacteria to resected human 

colonic tissue: Quantitative analysis of bacterial adhesion and viability. Res. Microbiol, 156, 

238-244. https://doi.org/10.1016/j.resmic.2004.08.012 

Vonk, R. J., Reckman, G. A. R., Harmsen, H. J. M., & Priebe, M. G. (2012). Probiotics and lactose 

intolerance. Probiotics, 149-160.  

Yadav, A. K., Tyagi, A., Kumar, A., Saklani, A. C., Grover, S., & Batish, V. K. (2014). Adhesion of 

indigenous Lactobacillus plantarum to gut extracellular matrix and its physicochemical 

characterization. Arch. Microbiol, 197, 155-164. https://doi.org/10.1007/s00203-014-1034-7 

Zheng, Y., Lu, Y., Wang, J., Yang, L., Pan, C., & Huang, Y. (2013). Probiotic Properties of 

Lactobacillus Strains Isolated from Tibetan Kefir Grains. PLoS One, 22;8(7), e69868. 

https://doi.org/10.1371/journal.pone.0069868 

 


