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Lactic Acid Bacteria as Probiotics:Characteristics, Selection Criteria and Role in Immunomodulation of Human GI Muccosal BarrierDaoud Harzallah and Hani Belhadj[1] Laboratory of Applied Microbiology, Faculty of Natural and Life Sciences, University Ferhat Abbas, Setif, Algeria1. The gut is colonized with different types of microorganisms which have dynamic and diverse symbiotic relationships. The capacity to ferment complex polysaccharides to short-chain fatty acids (SCFAs) by intestinal microflora has a profound effect on energy homeostasis, providing as much as ~70% of energy in ruminants and 20–30% for several monogastric animals (8).
During the last 15 years, our laboratories have worked toward the identification of probiotic candidates for use in poultry as a defined LAB-based probiotic culture (FloraMax® B11). In spite of the success shown by the development of the LAB probiotic for use in commercial poultry (above), there is still an urgent need for commercial probiotics that are shelf-stable, cost-effective, and feed-stable (tolerance to the heat pelletization process) to increase compliance and widespread utilization. At the present time, our laboratory’s aim is to develop a novel, cost-effective, feed-stable probiotic with widespread utilization, simple delivery, and clinical efficacy for human and animal use. Other isolates or combinations of isolates with increased potency and efficacy may be identified with continued research. Digestive tract of a hen: (a) beak and mouth, (b) esophagus, (c) crop, (d) proventriculus, (e) gizzard, (f) duodenal loop, (g) pancreas, (h) liver, (i) gallbladder, (j) jejunum, (k) ileum, (l) ceca, (m) rectum, (n) cloaca, (o) vent. View from right of digestive tract of a cow: (a) mouth, (b) esophagus, (c) rumen, (d) reticulum, (e) omasum, (f) abomasum, (g) duodenum, (h) liver, (i) gallbladder, (j) pancreas, (k) small intestine, (l) large intestine, (m) ceca, (n) rectum. The mutualism of the ruminant is dependent upon the pre-gastric ruminal fermentation which converts feedstuffs to SCFAs, and microbial cells (microbial crude protein), which are washed out of the rumen and are degraded by the host gastric stomach and absorbed by the intestinal tract (71). Ruminant diets are comprised of many variable feedstuffs that contain complex carbohydrates (e.g.
The advent of next generation sequencing has produced a bloom of information available about the composition of the resident gastrointestinal microbiome under different conditions; and metagenomics, proteomics, and KEGG analyses (Kyoto Encyclopedia of Genes and Genomes) have led to an unprecedented understanding of the metabolic actions of and interactions between ruminal microorganisms and their host. Most of our knowledge based on the microbial ecology of the rumen and GIT is derived from culture-based methodologies; however, over the past 5 years, more than 3,000 known bacterial species have been detected in cattle rumens and GITs, and estimates of the unidentified species in the rumen range from 10- to 100-fold higher.
As an example of changes in the gastrointestinal microbiome caused by diet, starch can significantly alter the population composition of the lower GIT. While bacteria are the best understood members of the microbial consortium, protozoa (Eucarya) comprise as much as 50% of the biomass of the rumen, yet relatively little is known of the types of protozoa found in the rumen.
Although much work remains to be done to determine their exact mechanism of action, bacteriocins and bacteriocin-producing bacteria have the potential to improve animal health and by extension, improve the safety of animal products consumed by humans. The interest in digestive physiology and the role of microorganisms has generated data whereby human and animal wellbeing can be enhanced and the risk of disease reduced. Metchnikoff founded the research field of probiotics, aimed at modulating the intestinal microflora (115).
The authors acknowledge the support of the European Science Foundation (ESF), in the framework of the Research Networking Programe, The European Network for Gastrointestinal Health Research.
The authors have not received any funding or benefits from industry or elsewhere to conduct this study. Probiotics, prebiotics, energy balance, and obesity: mechanistic insights and therapeutic implications.
Probiotics and prebiotics for severe acute malnutrition (PRONUT study): a double-blind efficacy randomised controlled trial in Malawi.
Host-microbial symbiosis in the vertebrate gastrointestinal tract and the Lactobacillus reuteri paradigm. Antibiotic resistance in bacteria associated with food animals: a United States perspective of livestock production.
Leukocytic responses and intestinal mucin dynamics of broilers protected with Enterococcus faecium EF55 and challenged with Salmonella Enteritidis. Comparison of the immune responses induced by local immunizations with recombinant Lactobacillus plantarum producing tetanus toxin fragment C in different cellular locations.
Upregulation of oxidative burst and degranulation in chicken heterophils stimulated with probiotic bacteria. Spherical lactic acid bacteria activate plasmacytoid dendritic cells immunomodulatory function via TLR9-dependent crosstalk with myeloid dendritic cells. Prophylactic Lactobacillus GG reduces antibiotic-associated diarrhea in children with respiratory infections: A randomized study.
The role of the intestinal microflora for the development of the immune system in early childhood. Effect of lactic acid bacteria probiotic culture treatment timing on Salmonella Enteritidis in neonatal broilers. Transcriptional profiling of cecal gene expression in probiotic-and Salmonella-challenged neonatal chicks. Effect of probiotic treatment in broiler chicks on intestinal macrophage numbers and phagocytosis of Salmonella Enteritidis by abdominal exudate cells. Evaluation of spray application of a Lactobacillus-based probiotic on Salmonella Enteritidis colonization in broiler chickens.
Effect of poultry guard litter amendment on horizontal transmission of Salmonella Enteritidis in broiler chicks.
Effect of a selected Lactobacillus spp.-based probiotic on Salmonella enterica serovar Enteritidis-infected broiler chicks. Effect of lactic acid bacteria probiotic culture for the treatment of Salmonella enterica serovar Heidelberg in neonatal broiler chickens and turkey poults.
Evaluation of intervention strategies for idiopathic diarrhea in commercial turkey brooding houses. Identification and characterization of lactic acid bacteria in a commercial probiotic culture. A probiotic strain of Bacillus polyfermenticus reduces DMH induced precancerous lesions in F344 male rat. Probiotic impact on microbial flora, inflammation and tumour development in IL-10 knockout mice. Evaluation of germination, distribution and persistence of Bacillus subtilis spores through the gastrointestinal tract of chickens. Germination of the spore in the gastrointestinal tract provides a novel route for heterologous antigen delivery. Evaluation of a screening and selection method for bacillus isolates for use as effective direct-fed microbials in commercial poultry. Evaluation of Bacillus species as potential candidates for direct-fed microbials in commercial poultry. Regulation of lactate production in Streptococcus bovis: a spiraling effect that contributes to rumen acidosis. Comparative studies of microbial populations in the rumen, duodenum, ileum and faeces of lactating dairy cows. Comparison of bacterial communities in faeces of beef cattle fed diets containing corn and wet distillers’ grain with solubles. Impact of reducing the level of wet distillers grains fed to cattle prior to harvest on prevalence and levels of Escherichia coli O157:H7 in feces and on hides.
Bacterial community analysis of beef cattle feedlots reveals pen surface is distinct from feces. Evaluation of the bacterial diversity in the feces of cattle using bacterial tag-encoded FLX amplicon pyrosequencing (bTEFAP). Ruminal bacterial, archaeal, and fungal diversity of dairy cows with normal and reduced ruminal fauna.
Evaluation of bacterial diversity in the rumen and feces of cattle fed different levels of dried distillers grains plus solubles using bacterial tag-encoded FLX amplicon pyrosequencing. Impact of high-concentrate feeding and low ruminal pH on methanogens and protozoa in the rumen of dairy cows. Distribution of anaerobic fungi in the digestive tract of cattle and their survival in faeces.
Commercial probiotics are not effective for short-term control of enterohemorrhagic Escherichia coli O157 infection in beef cattle. Use of a trial probiotic product in calves experimentally infected with Escherichia coli O157. Reduction of fecal shedding of enterohemorrhagic Escherichia coli O157:H7 in lambs by feeding microbial feed supplement.
Evaluation of a direct-fed microbial product effect on the prevalence and load of Escherichia coli O157:H7 in feedlot cattle. Isolation of bovine intestinal Lactobacillus plantarum and Pediococcus acidilactici with inhibitory activity against Escherichia coli O157 and F5. Extracts from bitter orange on its own or combined with caffeine do not pose any risks at doses commonly used by humans, says a new review of published and un-published clinical data.
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Vitamins and minerals in foods are bound to natural food complexes with proteins, carbohydrates and lipids. Also, some vitamins and minerals are extracted from food sources, leaving behind toxic residues if chemical solvents were used to remove them. This powder tends to curb cravings and leaves you feeling cleansed - although the smell isn't the greatest. IntroductionAs it was reported by Chow (2002), the notion that food could serve as medicine was first conceived thousands of years ago by the Greek philosopher and father of medicine, Hippocrates, who once wrote: 'Let food be thy medicine, and let medicine be thy food'. Epithelial cells lining the gastrointestinal tract are able to respond to infection by initiating either nonspecific or specific host-defence response (Kagnoff and Eckmann 1997, Strober 1998). Dietary ingredients have a profound effect on the composition of the gut microflora, which in turn regulates the physiology of metazoans.
The gut microbiome serves multiple functions including the conversion of several nutrients that the host cannot digest to end products, a process which has a direct impact on digestive physiology and immunology. In this way, prebiotics selectively modify the colonic microflora and can potentially influence gut metabolism (4). Alternatively, different studies have showed that the administration of a lactic acid based probiotic 2 hours after Salmonella challenge, had no effect during the first 12 hours on increasing cecal colonization by this pathogen, although marked and rapid decreases were observed between 12 and 24 hours post-challenge (40). This has demonstrated an accelerated development of normal microflora in chickens and turkeys, providing increased resistance to Salmonella spp. Among the large number of probiotic products in use today some are bacterial spore formers, mostly of the genus Bacillus. Together, these studies not only show that spores are not transient passengers in the gut, but that they have an intimate interaction with the host cells or microflora that can enhance their potential probiotic effect. We have demonstrated that one Bacillus subtilis spores isolate is as effective as our LAB-based probiotic (FloraMax™) for Salmonella reduction (68, 69), and equal to bacitracin (an antibiotic) for the prevention of necrotic enteritis in experimental and commercial field trials. Some of these environmental Bacillus isolates have been evaluated in vitro for antimicrobial activity against selected bacterial pathogens, heat stability, and the ability to grow to high numbers. The primary SCFAs of interest in the ruminant are acetate, propionate, and butyrate, which when absorbed by the host, can provide up to 80% of the animals energy requirement (71). An additional microbial consortium occupies the cecum and colon of cattle that is distinct from the ruminal population (74). However, our methods for understanding the biology and ecology of these consortia are still limiting.
Based on our new insights into the composition of the ruminal microbiota, the ruminant GIT is dominated by two major bacterial phyla Firmicutes and Bacteroidetes, which comprise approximately 80–92% of the intestinal microbiota in the human and bovine gut (77, 78). Firmicutes populations decreased with increasing fecal starch concentrations, whereas Bacteroidetes increased as starch concentration increased (78). Their role in starch sequestration and conversion of bacterial protein to high-quality protein has been noted but knowledge about the diversity of these important ruminal components has largely been limited to microscopy (84). The mechanisms of their antimicrobial activity can vary from bactericidal modulation of enzymatic activity, through to the inhibition of anionic transporters, or to form pores in the cytoplasmic membrane.
New molecular techniques allow an accurate assessment of the floral composition, resulting in improved strategies to elucidate the different mechanism of action that probiotics have to enhance performance of livestock animal species. However, other parts of the body containing endogenous microflora or problems relating to the immune system may also be candidates for probiotic therapy.
Lactobacillus rhamnosus induces peripheral hyporesponsiveness in stimulated CD4+ T cells via modulation of dendritic cell function. Both of which will support, guide, and inspire you toward the best possible health outcomes for you and your family. It does not have a hard cell wall making it harder to digest, but a soft, easily digested cell wall made of a mucopolysaccharide. Farming practices have depleted our soils of minerals, overuse of chemical fertilizers is killing valuable microorganisms that contribute to the mineral content in soils. Since many conventional foods are nutrient depleted, more people are taking spirulina and other green superfoods.
Potential and established health benefits associated with the usage of probiotics (Leroy et al., 2008).
However, during recent times, the concept of food having medicinal value has been reborn as 'functional foods'.
Bacterial adhesion to the host cell or recognition by the host cell is often an essential first stage in the disease process. As such, nutritional components of the diet are of critical importance not only for meeting the nutrient requirements of the host, but also for the microbiome. Although age, diet, environment, and ethnicity of the subjects have crucial impacts on the microbial gut composition, the improvement of high-throughput technologies has made it possible to better understand the contribution of microbiota to the human health status (1). Bacteria fed with a preferential food substrate have a proliferative advantage over other bacteria (2). However, the bacterial nutrient package will not be advantageous without the presence of the specific beneficial bacteria that use it, and similarly the live microbial product will not succeed if the environment into which it is introduced is unfavorable (6).

Later, using the same challenge model and microarray analysis of gut mRNA expression, gene expression differences were found in birds treated with probiotic compared to saline treated birds, especially those associated with the NFκB complex and apoptosis, suggesting that increased apoptosis may be a mechanism by which this probiotic reduces Salmonella infection (41).
Used primarily in their spore form, some (though not all) have been shown to prevent selected gastrointestinal disorders and the diversity of species used and their applications are astonishing. Several commercial spore-forming Bacillus cultures have been shown to reduce food-borne pathogens (67).
Our preliminary data suggests that these isolates could be an effective alternative to antibiotic growth promoters for commercial poultry.
Ruminants, such as cattle, exist in a unique symbiotic relationship with a pre-gastric microbial consortium of bacteria, protozoa, archaea, viruses, and fungi that ferment relatively low-quality feedstuffs and in turn supply the ruminant with nutrients to produce high-quality meat, milk, and fiber (70). Although acetate is the primary SCFA produced in the ruminal fermentation, propionate is glucogenic and is more desirable in animals growing rapidly (e.g. Thus, as dietary components change, the nutrient niches and bacterial species occupying them also change.
This microbial population performs a secondary fermentation in the lower gut that captures more dietary energy that bypasses ruminal fermentation.
The proportions and diversity of organisms representing this community are strongly influenced by diet as postulated in the nutrient-niche theory (72, 75, 78). Prevotellaceae and Lachnospiraceae were the dominant members of Bacteroidetes and Firmicutes, respectively in grain-fed cattle; however in forage-fed cattle, the Ruminococcaceae and Bacteroidales became dominant. Furthermore, while it has been long known that members of Kingdom Archaea, most notably methanogens, inhabit the rumen, only with the advent of molecular techniques have details of the Euryarchaeota population composition been reported (85, 86).
Given the recent international legislation and domestic consumer pressures to withdraw growth-promoting antibiotics and limit antibiotics available for treatment of bacterial infections, probiotics can offer alternative options. Much research has been completed in an effort to understand and apply the natural benefits of non-pathogenic bacteria, but there is much still to do.
Spirulina is a non nitrogen-fixing blue-green alga, which makes it safe for human consumption.
Most vitamin and mineral supplements, however, are synthetic combinations of isolated USP vitamins and minerals. These wholefoods offer functional nutrients and phytochemicals, new frontiers for disease prevention research, way beyond isolated vitamins and mineral supplements.
This productivity breakthrough yields over 20 times more protein than soybeans on the same area, 40 times corn and 400 times beef. Key and desirable criteria for the selection of probiotics in commercial applications (Vasiljevic and Shah, 2008).
The list of health benefits accredited to functional food continues to increase, and the gut is an obvious target for the development of functional foods, because it acts as an interface between the diet and all other body functions. A wide range of gastrointestinal cell surface constituents, such as several glygoconjucates, can serve as receptors for bacterial adherence (Servin and Coconnier 2003, Pretzer et al., 2005).
During their coevolution, bacterial microbiota has established multiple mechanisms to influence the eukaryotic host, generally in a beneficial fashion.
Some prebiotics have shown to selectively stimulate the growth of endogenous lactic acid bacteria (LAB) and Bifidobacteria in the gut to improve the health of the host (3, 4).
The concept of symbiotic has been proposed recently to characterize foods with both prebiotic and probiotic properties as health-enhancing functional foods. In addition, Toll-like receptors (TLRs) represent evolutionary conserved pathogen recognition receptors which are necessary components in the protection against invading microorganisms (42). Published experimental and commercial studies have shown that these selected probiotic organisms are able to reduce idiopathic diarrhea in commercial turkey brooding houses (48).
While not all Bacillus spores are highly heat tolerant, some specific isolates are the toughest life form known on earth (51) and can be used under extreme heat conditions. However, cost issues associated with achieving necessary concentrations of spores in feed have greatly limited commercial acceptance in the animal industry (57).
Importantly, improved efficiency of amplification and sporulation is absolutely essential to gain widespread industry acceptance of a feed-based probiotic for ante mortem food-borne pathogen intervention.
The intestinal tract of ruminants is different from monogastrics because it contains a large (up to 100 liters) anaerobic fermentation chamber, the reticulo-rumen, located prior to feedstuff entry into the gastric stomach, the abomasum (Figs. For instance, starch that reaches the hindgut is fermented, which causes higher fecal SCFA and lactate concentrations and a lower pH than that found in forage-based diets. Grain feeding decreased Ruminococcaceae populations but increased Lachnospiraceae, Clostridiaceae, and Succinivibrionaceae populations (78). While microscopy studies have previously demonstrated that protozoa and methanogens exist in symbiosis, this relationship can now be quantified and specific relationships between members of these understudied kingdoms can be determined. New advances in the application of probiotics are directed to produce significant changes in gut physiology and provide even higher levels of health as well as increasing performance parameters in different animals. One of the most promising areas for the development of functional food components lies in the use of probiotics and prebiotics which scientific researches have demonstrated therapeutic evidence. Furthermore, epithelial cells express constitutively host pattern recognition receptors (PRRS), such as Toll-like receptors (TLR). The microbiome encrypts a variety of metabolic functions that complements the physiology of their hosts. Nevertheless, probiotics can only be effective if the requirements for their growth are present in the gastrointestinal tract (GIT). Certain probiotic strains may modulate TLR expression as a mechanism of their protective effect against pathogens.
Large-scale commercial trials indicated that appropriate administration of this probiotic mixture to turkeys and chickens increased performance and reduced costs of production (45, 49). Several studies have shown that either live vegetative cells or endospores of some isolates can prevent colon carcinogenesis (52) or discharge antimicrobial substances against Gram-positive bacteria, such as Staphylococcus aureus, E. Recently, both vegetative growth and sporulation rate have been optimized in our laboratory, which may lead to new efficiencies for commercial amplification and manufacture of a cost-effective product at very high spore counts (68). The acetate:propionate ratio, an indicator of the energetic efficiency of the fermentation to the host animal, can be altered by simply increasing the amount of cereal grain in the ration.
For example, the primary driver of the shift in ruminal propionate proportions by cereal grain feeding is starch. This shift in the ruminal population could be explained by the hypothesized role Ruminococci play in degradation of hemicellulose and xylan. Fungi have also been commonly isolated from the rumen, yet the role of fungi is still unclear (87). But these vitamins and minerals are not bound to anything, and may have an entirely different chemical structure than those found in foods.
This way, it can augment the food supply not by clearing the disappearing rainforests, but by cultivating the expanding deserts.
Nowadays, consumers are aware of the link among lifestyle, diet and good health, which explains the emerging demand for products that are able to enhance health beyond providing basic nutrition. Over a century ago Eli Metchnikoff proposed the revolutionary idea to consume viable bacteria to promote health by modulating the intestinal microflora.
In order to select even more effective isolates, we are currently focused on the mechanistic action of new Bacillus candidates. The mutualistic nature of the ruminant animal and the resident microbial population is critical to the success of ruminant animals in a variety of environmental niches around the world. Grain feeding decreases the acetate:propionate ratio, meaning more energy is available to the host for growth.
Ruminal starch is fermented by Streptococci and Lactococci to produce SCFAs and lactic acid which results in a lowering of ruminal pH (73).
Prevotellae and Bacteroides, on the other hand, utilize hemicellulose but also degrade starch and pectin rapidly, yet Bacteroides are more commonly found in cattle feces than Prevotella. Clostridium, Enterococcus, Staphylococcus, and Peptostreptococcus) have displayed a marked ability to ferment peptides and amino acids, indicating that diets rich in available protein would provide a competitive advantage for these genera.
Bacteriophages (bacterial viruses) have been frequently isolated from the rumen, and it has been hypothesized that they play a role in nutrient recycling in the rumen as well as diurnal variation of the ruminal populations (88). Different researchers have compared several of the commercially available growth enhancement probiotics and yeast products and found that feeding these probiotics provided no effect in regards to pathogen levels in cattle (90, 91). Vitamin and mineral formulas may ignore antagonistic and synergistic effects of vitamins and minerals both in regard to absorption and metabolic reactions once absorbed. The idea is more applicable now than ever, since bacterial antimicrobial resistance has become a serious worldwide problem both in medical and agricultural fields. Tolerance and resistance to acidic pH, high osmotic concentration of NaCl, and bile salts of these isolates may have contributed to the efficacy of these isolates to survive and provide beneficial effects. These results provide evidence of colonization and antimicrobial activity of probiotic bacteria, thus, products containing Bacillus spores are used commercially as probiotics, and they offer potential advantages over the more common LAB products since they can be used as direct feed microbials. Preliminary studies conducted in our laboratory indicate a potential mechanistic action of these new Bacillus candidates at least partially involves the rapid activation of innate host immune mechanisms in chickens and turkeys (unpublished data). Further benefits to ruminants of this mutualistic relationship include the fact that B-complex vitamins are produced by the ruminal and gastrointestinal microbiome which eliminates the need for dietary B-vitamin supplementation. As a result of this increase in lactic acid concentrations, populations of the lactate-utilizing ruminal bacteria Megasphaera elsdenii and Selenomonas ruminantium increase due to their ability to secondarily ferment lactic acid to form propionate (70). Collectively, these population shifts illustrate the elasticity of the ruminant microbiome response to environmental (dietary) shifts.
However, a probiotic culture comprised of Streptococcus bovis and Lactobacillus gallinarum from the rumen of cattle resulted in the reduction of E. It is well known that many vitamin and mineral supplements, especially calcium and iron are not well absorbed. TLRs are also found on innate immune cells, such as dendritic cells and macrophages (Vinderola et al., 2005).
The impending ban of antibiotics in animal feed due to the current concern over the spread of antibiotic resistance genes makes a compelling case for the development of alternative prophylactics.
Both strains also have in vitro antibacterial activity against Salmonella enterica serovar Enteritidis, Escherichia coli (O157:H7), and Campylobacter jejuni (50). This data provides an exciting possibility for the identification of vastly superior and more potent probiotics in the near future. Feeding wet distiller’s grains to cattle changes the microbial community structure (most notably Prevotellae) in the feces (75) which significantly increased E. They can be found in fermented products as meat, milk products, vegetables, beverages and bakery products. Nutritional approaches to counteract the debilitating effects of stress and infection may provide producers with useful alternatives to antibiotics. In addition, previous research from our laboratory indicates very rapid induction of specific host-gene expression pathways, which are associated with reductions in enteric colonization with Salmonella (41). The aforementioned characteristics, may contribute to the efficacy previously reported in laboratory and field conditions (24). Interestingly, while pens where cattle are housed are often covered with fresh feces, recent molecular studies have indicated that the bacterial communities of feedlot surfaces (and by extension other animal housing environments) are distinctive from fecal bacterial populations (81). They are part of the microbiota on mucous membranes, such as the intestines, mouth, skin, urinary and genital organs of both humans and animals, and may have a beneficial influence on these ecosystems. Other known recognition receptors are nucleotide-binding oligomerization domain proteins, which recognize both gram-positive and gram-negative bacteria.
While many mechanisms of action have been proposed for the observed efficacy, precise modalities have not been completely described for this highly effective culture. LAB that grow as the adventitious microflora of foods or that are added to foods as cultures are generally considered to be harmless or even an advantage for human health. This review presents some of the alternatives currently used in food-producing animals to influence their health in relation to human health.
Since their discovery, LAB has been gained mush interest in various applications, as starter cultures in food and feed fermentations, pharmaceuticals, probiotics and as biological control agents. Increased epithelial barrier permeability is frequently associated with gastrointestinal disorders contributing to both disease onset and persistence (Lu and Walker 2001, Berkes 2003). In food industry, LAB are widely used as starters to achieve favorable changes in texture, aroma, flavor and acidity (Leory and De Vuyst, 2004). The gatekeeper of the paracellular pathway is the tight junction, which is an apically located cell-cell junction between epithelial cells. The tight junction permits the passage of small molecules, such as ions, while restricting the movement of large molecules, such as antigens and microorganisms, which can cause inflammation. Du to their antimicrobial and antioxidant activities some LAB strains are used in food biopreservation.
Origine and safety of probiotics An old dogma of probiotic selection has been that the probiotic strains should be of “human origin”. Many of the indications for probiotic activity have been obtained from effects observed in various clinical situations.
One may argue that from evolutionary point of view, describing bacteria to be of human origin does not make much sense at all.
The requirement for probiotics to be of human origin relates actually to the isolation of the strain rather than the “origin” itself. A DFM comprised of Bacillus subtilis did not affect the fecal prevalence or concentration of E. Usually, the strains claimed to be “of human origin” have been isolated from faecal samples of healthy human subjects, and have therefore been considered to be “part of normal healthy human gut microbiota”. Overview of probioticsThe most tried and tested manner in which the gut microbiota composition may be influenced is through the use of live microbial dietary additions, as probiotics. In reality the recovery of a strain from a faecal sample does not necessarily mean that this strain is part of the normal microbiota of this individual, since microbes passing the GI tract transiently can also be recovered from the faecal samples (Forssten et al., 2011). Studies have also indicated that cultures of Lactobacillus acidilactici and Pediococcus could directly inhibit E.
In practice it is impossible to know the actual origin of the probiotic strains, regardless of whether they have been isolated from faecal samples, fermented dairy products or any other source for that matter.
Isolation of a strain from faeces of a healthy individual is also not a guarantee of the safety of the strain—such a sample will also always contain commensal microbes which can act as opportunistic pathogens, or even low levels of true pathogens, which are present in the individual at sub-clinical levels.
However, at the beginning of this century probiotics were first put onto a scientific basis by the work of Metchnikoff (1908). He hypothesised that the normal gut microflora could exert adverse effects on the host and that consumption of ‘soured milks’ reversed this effect.
However, many species of the genera Lactobacillus, Leuconostoc, Pediococcus, Enterococcus, and Bifidobacterium were isolated frequently from various types of infective lesions. The origin of the first use can be traced back to Kollath (1953), who used it to describe the restoration of the health of malnourished patients by different organic and inorganic supplements.
Later, Vergin (1954) proposed that the microbial imbalance in the body caused by antibiotic treatment could have been restored by a probiotic rich diet; a suggestion cited by many as the first reference to probiotics as they are defined nowadays.
Similarly, Kolb recognized detrimental effects of antibiotic therapy and proposed the prevention by probiotics (Vasiljevic and Shah, 2008) Later on, Lilly and Stillwell (1965) defined probiotics as “…microorganisms promoting the growth of other microorganisms”.
The idea of health-promoting effects of LAB is by no means new, as Metchnikoff proposed that lactobacilli may fight against intestinal putrefaction and contribute to long life. Although minor side effects of the use of probiotics have been reported, infections with probiotic bacteria occur and invariably only in immunocompromised patients or those with intestinal bleeding (Leroy et al., 2008). Other definitions advanced through the years have been restrictive by specification of mechanisms, site of action, delivery format, method, or host.

An issue of concern regarding the use of probiotics is the presence of chromosomal, transposon, or plasmid-located antibiotic resistance genes amongst the probiotic microorganisms.
At this moment, insufficient information is available on situations in which these genetic elements could be mobilised, and it is not known if situations could arise where this would become a clinical problem (Leroy et al., 2008). The mechanism of action of probiotics (e.g, having an impact on the intestinal microbiota or enhancing immune function) was dropped from the definition to encompass health effects due to novel mechanisms and to allow application of the term before the mechanism is confirmed. Furthermore, certain mechanisms of action (such as delivery of certain enzymes to the intestine) may not require live cells. In vitro safety screenings of probiotics may include, among others, antibiotic resistance assays, screenings for virulence factors, resistance to host defence mechanisms and induction of haemolysis. In relation to food, probiotics are considered as “viable preparations in foods or dietary supplements to improve the health of humans and animals”. According to these definitions, an impressive number of microbial species are considered as probiotics. ConclusionThe individual diversity of the intestinal microflora underscores the difficulty of identifying the entire human microbiota and poses barriers to this ?eld of research. Selection of probioticsMany in vitro tests are performed when screening for potential probiotic strains.
The first step in the selection of a probiotic LAB strain is the determination of its taxonomic classification, which may give an indication of the origin, habitat and physiology of the strain.
It is also apparent that even a single strain of probiotic may exert its actions via multiple, concomitant pathways.
All these characteristics have important consequences on the selection of the novel strains (Morelli, 2007). Probiotics have long been used as an alternative to traditional medicine with the goal of maintaining enteric homeostasis and preventing disease. This conclusion was brought forward due to uncertainty of the origin of the human intestinal microflora since the infants are borne with virtually sterile intestine.
Clinical trials have shown that probiotic treatment can reduce the risk of some diseases, especially antibiotic-associated diarrhea, but conclusive evidence is impeded owing to the wide range of doses and strains of bacteria used.
However, the panel also underlined a need for improvement of in vitro tests to predict the performance of probiotics in humans. While many probiotics meet criteria such as acid and bile resistance and survival during gastrointestinal transit, an ideal probiotic strain remains to be identified for any given indication. Many studies, as discussed above, have shown that probiotics increase barrier function in terms of increased mucus, antimicrobial peptides, and sIgA production, competitive adherence for pathogens, and increased TJ integrity of epithelial cells. Furthermore, it seems unlikely that a single probiotic will be equally suited to all indications; selection of strains for disease-specific indications will be required (Shanahan, 2003). Current investigation into the mechanism of action of speci?c probiotics has focused on probiotic-induced changes in the innate immune functions involvingTLRs and its downstream systems Like NF-?B, and other pathways (Yoon and Sun, 2011). Although the immunomodulatory effects of probiotics have been demonstrated in experimental animal models of allergy, autoimmunity, and IBD, information from clinical trials in humans is scarce. The ability to adhere to the intestinal mucosa is one of the more important selection criteria for probiotics because adhesion to the intestinal mucosa is considered to be a prerequisite for colonization (Tuomola et al., 2001). The table below (Table 2) indicates key creteria for sellecting probiotic candidat for commercial application, and figure 1 presents major and cardinal steps for sellecting probiotic candidats.It is of high importance that the probiotic strain can survive the location where it is presumed to be active. Therefore, more research, especially in the form of well-designed clinical trials, is needed to evaluate the ef?cacy and safety of probiotics (Ezendam and Van Loveren, 2008). For a longer and perhaps higher activity, it is necessary that the strain can proliferate and colonise at this specific location.
Probably only host-specific microbial strains are able to compete with the indigenous microflora and to colonise the niches. Besides, the probiotic strain must be tolerated by the immune system and not provoke the formation of antibodies against the probiotic strain. On the other hand, the probiotic strain can act as an adjuvant and stimulate the immune system against pathogenic microorganisms. Basic initial characterization of strain identity and taxonomy should be conducted, followed by evaluation with validated assays both in studies of animal models and in controlled studies in the target host.
In vitro assays are frequently conducted that have not been proved to be predictive of in vivo function.
Technological robustness must also be determined, such as the strain’s ability to be grown to high numbers, concentrated, stabilized, and incorporated into a ?nal product with good sensory properties, if applicable, and to be stable, both physiologically and genetically, through the end of the shelf life of the product and at the active site in the host. Assessment of stability can also be a challenge, since factors such as chain length and injury may challenge the typical assessment of colony-forming units, as well as in vivo function (Sanders, 2008). Dose levels of probiotics should be based on levels found to be ef?cacious in human studies.
Furthermore, the impact of product format on Figure 1.Scheme of the Guidelines for the Evaluation of Probiotics for Food Use. The common quality-control parameter of colony-forming units per gram may not be the only parameter indicative of the ef?cacy of the ?nal product. Other factors, such as probiotic growth during product manufacture, coating, preservation technology, metabolic state of the probiotic, and the presence of other functional ingredients in the ?nal product, may play a role in the effectiveness of a product. Potential mechanisms of action of probioticsA wide variety of potential beneficial health effects have been attributed to probiotics (Table 3).
Claimed effects range from the alleviation of constipation to the prevention of major life-threatening diseases such as inflammatory bowel disease, cancer, and cardiovascular incidents. Some of these claims, such as the effects of probiotics on the shortening of intestinal transit time or the relief from lactose maldigestion, are considered well-established, while others, such as cancer prevention or the effect on blood cholesterol levels, need further scientific backup (Leroy et al., 2008). The mechanisms of action may vary from one probiotic strain to another and are, in most cases, probably a combination of activities, thus making the investigation of the responsible mechanisms a very difficult and complex task.
In general, three levels of action can be distinguished: probiotics can influence human Probiotic organisms can provide a beneficial effect on intestinal epithelial cells in numerous ways. Gut microbiotaThe human gastrointestinal tract is inhabited by a complex and dynamic population of around 500-1000 of different microbial species which remain in a complex equilibrium. It has been estimated that bacteria account for 35–50% of the volume content of the human colon.
These include Bacteroides, Lactobacillus, Clostridium, Fusobacterium, Bifidobacterium, Eubacterium, Peptococcus, Peptostreptococcus, Escherichia and Veillonella. The bacterial strains with identified beneficial properties include mainly Bifidobacterium and Lactobacillus species. The dominant microbial composition of the intestine have been shown to be stable over time during adulthood, and the microbial patterns are unique for each individual. However, there are numerous external factors that have potential to influence the microbial composition in the gut as host genetics, birth delivery mode, diet, age, antibiotic treatments and also, other microorganisms as probiotics. The intestine is one of the main surfaces of contact with exogenous agents (viruses, bacteria, allergens) in the human body. It has a primary role in the host defense against external aggressions by means of the intestinal mucosa, the local immune system, and the interactions with the intestinal microbiota (resident and in transitbacteria). Gut microbiota influences human health through an impact on the gut defense barrier, immune function, nutrient utilization and potentially by direct signaling with the gastrointestinal epithelium (Collado et al., 2009). In healthy adults, 80% of phylotypes belong to four major phylogenetic groups, which are the Clostiridium leptum, Clostridium coccoides, Bacteroides and Bifidobacteria groups. Also, studies have found that mucosal microbiota is stable along the distal gastrointestinal tract from ileum to rectum, but mucosa-associated microbiota is different from fecal microbiota. The number of bacterial cells present in the mammalian gut shows a continuum that goes from 101 to 103 bacteria per gram of contents in the stomach and duodenum, progressing to 104 to 107 bacteria per gram in the jejunum and ileum and culminating in 1011 to 1012 cells per gram in the colon (Figure 3a). In addition to the longitudinal heterogeneity displayed by the intestinal microbiota, there is also a great deal of latitudinal variation in the microbiota composition (Figure 3b). The intestinal epithelium is separated from the lumen by a thick and physicochemically complex mucus layer. The microbiota present in the intestinal lumen differs significantly from the microbiota attached and embedded in this mucus layer as well as the microbiota present in the immediate a: variations in microbial numbers and composition across the length of the gastrointestinal tract. For instance, Bacteroides, Bifidobacterium, Streptococcus, members of Enterobacteriacea, Enterococcus, Clostridium, Lactobacillus, and Ruminococcus were all found in feces, whereas only Clostridium, Lactobacillus, and Enterococcus were detected in the mucus layer and epithelial crypts of the small intestine (Sekirov et al., 2010).
Upon passage through the birth canal, infants are exposed to a complex microbial population.
After the initial establishment of the intestinal microbiota and during the first year of life, the microbial composition of the mammalian intestine is relatively simple and varies widely between different individuals and also with time. Survival and antagonism effects of probiotics in the gutThe intestinal epithelium is the largest mucosal surface in the human body, provides an interface between the external environment and the host.
The gut epithelium is constantly exposed to foreign microbes and antigens derived from digested foods. Thus, the gut epithelium acts as a physical barrier against microbial invaders and is equipped with various elements of the innate defense system. In the gut, two key elements govern the interplay between environmental triggers and the host: intestinal permeability and intestinal mucosal defense. Resident bacteria can interact with pathogenic microorganisms and external antigens to protect the gut using various strategies.According to the generally accepted de?nition of a probiotic, the probiotic microorganism should be viable at the time of ingestion to confer a health bene?t. Although not explicitly stated, this de?nition implies that a probiotic should survive GI tract passage and, colonize the host epithelium. A variety of traits are believed to be relevant for surviving GI tract passage, the most important of which is tolerance both to the highly acidic conditions present in the stomach and to concentrations of bile salts found in the small intestine. These properties have consequently become important selection criteria for new probiotic functionality. One of the mechanisms by which the gut ?ora resists colonization by pathogenic bacteria is by the production of a physiologically restrictive environment, with respect to pH, redox potential, and hydrogen sul?de production. Probiotic bacteria decrease the luminal pH, as has been demonstrated in patients with ulcerative colitis (UC) following ingestion of the probiotic preparation VSL#3. Several bacteriocins produced by different species from the genus Lactobacillus have been described. The inhibitory activity of these bacteriocins varies; some inhibit taxonomically related Gram-positive bacteria, and some are active against a much wider range of Gram-positive and Gram-negative bacteria as well as yeasts and molds. Lacticin 3147, a broad-spectrum bacteriocin produced by Lactococcus lactis, inhibits a range of genetically distinct Clostridium dif?cile isolates from healthy subjects and patients with IBD. A further example is the antimicrobial effect of Lactobacillus species on Helicobacter pylori infection of gastric mucosa, achieved by the release of bacteriocins and the ability to decrease adherence of this pathogen to epithelial cells (Gotteland et al., 2006). The pretreatment of intestinal (T84) cells with lactic acid-producing bacteria reduced the ability of pathogenic E. Adhesion and invasion of an intestinal epithelial cell line (Intestine 407) by adherent invasive E. Probiotics and the mucous layerMost mucosal surfaces are covered by a hydrated gel formed by mucins.
Mucins are secreted by specialized epithelial cells, such as gastric foveolar mucous cells and intestinal goblet cells, Goblet cells are found along the entire length of the intestinal tract, as well as other mucosal surfaces. Of the 18 mucin-type glycoproteins expressed by humans, MUC2 is the predominant glycoprotein found in the small and large bowel mucus.
The NH2- and COOH-termini are not glycosylated to the same extent, but are rich in cysteine residues that form intra- and inter-molecular disul?de bonds.
These glycan groups confer proteolytic resistance and hydrophilicity to the mucins, whereas the disul?de linkages form a matrix of glycoproteins that is the backbone of the mucous layer (Ohland and MacNaughton, 2010).
Although small molecules pass through the heavily glycosylated mucus layer with relative ease, bulk fluid flow is limited and thereby contributes to the development of an unstirred layer of fluid at the epithelial cell surface. As the unstirred layer is protected from convective mixing forces, the diffusion of ions and small solutes is slowed (Turner, 2009).
This gel layer provides protection by shielding the epithelium from potentially harmful antigens and molecules including bacteria from directly contacting the epithelial cell layer, while acting as a lubricant for intestinal motility. Mucins can also bind the epithelial cell surface carbohydrates and form the bottom layer, which is ?rmly attached to the mucosa, whereas the upper layer is loosely adherent.
The mucus is the ?rst barrier that intestinal bacteria meet, and pathogens must penetrate it to reach the epithelial cells during infection (Ohland and MacNaughton, 2010).Probiotics may promote mucus secretion as one mechanism to improve barrier function and exclusion of pathogens. In support of this concept, probiotics have been shown to increase mucin expression in vitro, contributing to barrier function and exclusion of pathogens. Several studies showed that increased mucin expression in the human intestinal cell lines Caco-2 (MUC2) and HT29 (MUC2 and 3), thus blocking pathogenic E. However, healthy rats did not display increased colonic TFF3 expression after stimulation by VSL#3 probiotics (Caballero-Franco et al., 2007).
Furthermore, mice treated with 1% dextran sodium sulfate (DSS) to induce chronic colitis did not exhibit increased TFF3 expression or wound healing when subsequently treated with VSL#3. This observation indicates that probiotics do not enhance barrier function by up-regulation of TFF3, nor are they effective at healing established in?ammation. Therefore, use of current probiotics is likely to be effective only in preventing in?ammation as shown by studies in animal models (Ohland and MacNaughton, 2010).5. Interaction of probiotic bacteria with gut epitheliumThe composition of the commensal gut microbiota is probably influenced by the combination of food practices and other factors like the geographical localization, various levels of hygiene or various climates. The establishment of a normal microbiota provides the most substantial antigenic challenge to the immune system, thus helping the gut associated lymphoid tissus (GALT) maturation.
The intestinal microbiota contributes to the anti-inflammatory character of the intestinal immune system. Several immunoregulatory mechanisms, including regulatory cells, cytokines, apoptosis among others, participate in the control of immune responses by preventing the pathological processes associated with excessive reactivity.
An interesting premise for probiotic physiological action is their capacity to modulate the immune system. Consequently, many studies have focused on the effects of probiotics on diverse aspects of the immune response. Following consumption of probiotic products, the interaction of these bacteria with intestinal enterocytes initiates a host response, since intestinal cells produce various immunomodulatory molecules when stimulated by bacteria (Delcenseri et al., 2009).
Furthermore, The indigenous microbiota is a natural resistance factor against potential pathogenic microorganisms and provides colonization resistance, also known as gut barrier, by controlling the growth of opportunistic microorganisms. It has been suggested that commensal bacteria protect their host against microbial pathogens by interfering with their adhesion and toxic effects (Myllyluoma, 2007).A fraction of ingested probiotics are able to interact with intestinal epithelial cells (IECs) and dendritic cells (DCs), depending on the presence of a dynamic mucus layer.
Probiotics can occasionally encounter DCs through two routes: DCs residing in the lamina propria sample luminal bacterial antigens by passing their dendrites between IECs into the gut lumen, and DCs can also interact directly with bacteria that have gained access to the dome region of the gut-associated lymphoid tissue (GALT) through specialized epithelial cells, termed microfold or M cells.
The interaction of the host cells with microorganism-associated molecular patterns (MAMPs) that are present on the surface macromolecules of probiotic bacteria will induce a certain molecular response.
The host pattern recognition receptors (PRRs) that can perceive probiotic signals include Toll-like receptors (TLRs) and the C type lectin DC-specific intercellular adhesion molecule 3-grabbing non-integrin (DC-SIGN).
Some molecular responses of IECs depend on the subtype of cell, for example, Paneth cells produce defensins and goblet cells produce mucus.
Important responses of DCs against probiotics include the production of cytokines, major histocompatibility complex molecules for antigen presentation, and co-stimulatory molecules that polarize T cells into T helper or CD4+CD25+ regulatory T cells in the mesenteric lymph nodes (MLNs) or subepithelial dome of the GALT. Host interactions of probiotic bacterial surface molecules: comparison with commensals and pathogens.

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