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Chances are you’ve heard the term probiotics and know that these good bacteria that live in your gut promote healthy digestion. While researchers are still unraveling exactly how these bacteria accomplish all that they do, it’s understood that probiotics help establish a healthy environment in your digestive tract, which is important considering 70 percent of your body’s immune system resides there. For thousands of years, many cultures relied on fermented foods like yogurt, sauerkraut and kimchi to obtain probiotics. The secret to these new probiotic foods and beverages is a unique probiotic strain called GanedenBC30, which is one of the only strains that can withstand extreme heat and cold, plus survive extended periods on grocery store shelves, with science and research backing its benefits. The next time you’re at the grocery store, take a closer look and you’ll find more than 500 foods and beverages on the shelves with this strain in a variety of surprising categories that your entire family will love.
Finished products often have health benefits, but probiotics can now be found in the ingredients used for baking, as well.
Whether you prefer tea and coffee or juices and waters, there are probiotic options on the shelves for cold and hot beverage drinkers alike. In the past, frozen products weren’t a well-known option for probiotics, but these days you can find healthy bacteria added to a variety of items, including breakfast burritos, popsicles and even ice cream. AbstractUnderstanding of probiotic-induced regulatory gene expression and networking is critical to further explore their roles in controlling infection. The human normal flora was determined thousands of years before humans ever consumed these agricultural products. Each harvest is third party tested for potency, purity and safety and is stored in a refrigerated warehouse to maximize freshness. It is not uncommon to experience what is referred to as excessive drainage syndrome when ridding the intestine of pathogenic bacteria and yeast and re-establishing normal flora. This is really a positive sign that the intestine is being cleansed of these toxin releasing and illness-promoting bacteria and fungi but it can be uncomfortable for a few days. Did you know that probiotics are also key to a healthy immune system and protein absorption? Another thing scientists have found is that probiotic benefits are strain-specific, which means it matters which probiotic is used in the products you buy.
Then, pasteurization became popular and most of the bacteria in fermented food was killed – even the beneficial kind. This means that you can now get your probiotics from many of your favorite foods and drinks without taking pills.
From sugar and buttery spreads to baking mixes themselves, a variety of probiotic options can easily be added to your next baking project. Look for well-known brands like Suja or Bigelow that are leading the way in innovative probiotic products.
Probiotics are taking over the snacking category, and companies have found ways to add them to options including chocolate, granola, trail mix, nut butters, kale chips, cookies and even Mariani fruit snacks. Transcriptional profile of selected innate immune genes in primary bovine intestinal epithelial cells was assessed over a time course of incubation with the probiotic Lactobacillus plantarum 299v. The more science reveals about them, the clearer it becomes that probiotics are something you want to consume on a daily basis.
Since then, people have relied on supplements to get probiotics, but today an increasing number of food and beverage companies are introducing everyday offerings that include the healthy bacteria and can easily be added to daily routines. Based on gene expression results, a time point was chosen to prime epithelial cells with the probiotic prior to infection with rotavirus. Plaque assays and genomic analysis provided the basis for establishing the efficacy of probiotics in preventing a rotaviral infection. Plaque assays revealed that the probiotic is capable of decreasing (at least by 100-fold) the levels of live virus when the cells were primed with the probiotic. There are over 1,000 species of commensals, composing approximately 1012 organisms per milliliter of content in the lumen of the large intestine of humans [9], which comprised of beneficial bacteria such as Bifidobacterium and Lactobacillus [3].
The intestinal environment provides nutrients to the commensals, and in turn the commensals benefit the host by aiding in the absorption of otherwise indigestible nutrients and preventing overgrowth of potential pathogens in the intestinal lumen [4]. This can occur through several different mechanisms such as secreting antimicrobial substances, providing competition at adhesion sites on the intestinal epithelial cells or enhancing the mucosal immune system [4].Of the known commensals, there are two species of established probiotics, Bifidobacterium and Lactobacillus. Probiotics are bacteria that can be consumed or found naturally in the intestine, in an active form, which confer beneficial properties to the host [7].
Recently, it has been established that there are four different mechanisms by which probiotics benefit the host.
The first is the ability of the probiotic to normalize or alter gut permeability, which decreases pathogen translocation into the intestinal epithelial cell and thus creates a defense barrier within the gut [10, 17].
Secondly, the probiotic may reduce the ability of pathogens to survive, and as such infection is reduced or eliminated.


Thirdly, the probiotic can modulate signal transduction, which can regulate gut homeostasis.
Lastly, innate and adaptive immune modulation through altered gene expression can occur due to probiotic presence in the gut [2, 17].Given that the intestinal epithelial lining is the first line of defense for intestinal infections, exploration of enhanced innate immunity through prophylactic probiotic presence is of major interest [8]. Upon pathogen entry into the epithelial cell, the hosta€™s innate immune response is activated via germline-encoded pattern recognition receptors (PRR) present on and within organelles in the epithelial cell [20]. This recognition of PAMPs by PRRs leads to a change in the gene expression of cytokines and chemokines [17]. These chemokines and cytokines are pro-inflammatory or anti-inflammatory and ultimately regulate the non-specific killing of epithelial cells and pathogens and aid in the recruitment of cells and host defense peptides [20].
Over a period of days, following induction of certain chemokines and cytokines, the hosta€™s adaptive immune response is activated, which regulates T and B cell responses [5]. Commensals are known to provide a form of continuous activation of the host epithelial cells, which leads to the continual production of factors for repairing tissue, thus maintaining homeostasis [9].
This maintenance of homeostasis or increase in the expression of certain genes involved in innate immunity is one mechanism of alleviating the clinical symptoms of pathogen infection [14].Lactobacillus plantarum 299v (Lp299v) is a probiotic that has been shown to aid in passive immunity and minimization of effects of viral infection in humans [6]. Prophylactic therapy in humans has been used with many lactobacillus strains, and success has been found in the alleviation of several diarrheagenic infections [15].
Lp299va€™s possibilities for therapeutic use have been initiated through the current study using a bovine model of rotaviral infection that can be translated to humans.Of particular interest is the potential beneficial effect probiotics have when the gastrointestinal tract is presented with a rotavirus infection.
A rotaviral challenge in vitro has, until recently, been impossible due to a lack of a stable bovine intestinal epithelial cell line. Over 25A years have passed since it was first proposed that a stable intestinal epithelial line should be developed and the effects of passive immunity in preventing a rotavirus infection should be researched [16]. We have developed this in-house stable line of bovine intestinal epithelial cells (BIEC) that is susceptible to rotavirus infection [2, 11]. Bovine rotavirus (BRV) causes an enteric infection in newborn calves up to approximately 3A weeks of age [16].
The clinical manifestations are diarrhea lasting several days, and viral shedding occurring in the feces, facilitating the spread of the disease to herds of cattle [19]. Its proposed mechanism of infection is through the luminal side of the intestine, making this virus a good candidate for in vitro study with intestinal epithelial cells from the small intestine [16].In this report, we have presented the effect Lp299v has on BIEC in vitro through a time course study.
The transcriptional profile of selected genes involved in innate immunity was analyzed using qRT-PCR to determine the best incubation time of the BIEC with Lp299v.
Once an optimal time point was chosen for priming of the BIEC with Lp299v, a time course infection with BRV was performed with and without the priming of the BIEC with Lp299v.
Adaptation of the primary cell line yielded a more sustainable culture, known as EVM07-BIEC. The supplemented Dulbeccoa€™s Modified Eagles Medium will now be referred to as Complete medium. To split the cells, the media was aspirated from the flask, and the monolayer was rinsed once with versene.
David Mack (Department of Pediatrics, University of Ottawa and Chief of Gastroenterology, Childrena€™s Hospital of Eastern Ontario, ON) and was stored in a?’80A°C.
A streak of the frozen stock was plated on sterile MRS agar (VWR, Mississauga, ON), prepared according to manufacturera€™s instructions. A single colony was picked from the plateA 16-h prior to the experiment and placed into 5A ml of pre-warmed MRS broth. Confirmation of BRV infection and count number was obtained through a plaque assay with the supernatant from each flask. The plaque assay was performed as described previously [1].Adaption of Rotavirus for Growth in BIECThe EVM07-BIECs were seeded in Corning six-well cell-binding tissue culture plates obtained from Sigmaa€“Aldrich (Oakville, ON), placed in a 37A°C incubator with 5% CO2 and grown for 24 h to the point of confluency. Rotavirus was added at a multiplicity of infection (MOI) of 1 to the plate and was allowed to absorb for 1-h in the 37A°C incubator with 5% CO2. Complete medium containing no FBS was added in and the plate was placed again in the incubator until cytopathic effect was observed.
The following morning, the Complete medium was removed and replaced with Complete medium that did not contain gentamycin or fetal bovine serum. Following incubation points, lysis of the cells and isolation of RNA were performed.BIEC Time Course with Lp299v and BRV InfectionFollowing priming the intestinal epithelial cells with Lp299v for 6-h, BRV strain C486 was added to the cells at a multiplicity of infection of 1. RNA was isolated following 6-h priming of cells with and without Lp299v and 6-h priming plus 2, 4, 6, 12 and 18-h BRV infection.
To determine RNA quantity following isolation, a spectrophotometer was used to read absorbances at A260, A280 and A230. The 44 (Table S1) oligonucleotide primers for qRT-PCR using cDNA were obtained from Invitrogen, Carlsbad, CA. All reactions were carried out using an iCycler IQ real-time PCR system (BioRad, Hercules, CA). Results were analyzed using the 2a?’a?†a?†Ct method as described by Livak and Schmittgen [13].Clustering and Pathway AnalysisK-means Clustering based on gene expression values of innate immune genes were calculated using Genespring software (Agilent Technologies, USA).
The primers of 44 genes (Table S1) coding for PRRs, cytokines or chemokines were selected to understand the mechanism of the innate immune response in BIEC. FigureA 1 shows the kinetics of fold change values of 12 selected innate immune genes with respect to time-matched media control following priming of BIEC with Lp299v. Results revealed that the cells are stimulated even at 2-h following priming, and this stimulation is relatively stable throughout the time course until 12-h post-priming.
Il-6 gene expression was the strongest at 2 and 4-h post-priming but stayed at an average constant level of a 3.6-fold increase at the 6 and 12-h time points.
MyD88, which forms a complex used to increase Type I IFN and cytokine responses, showed the most significant increase in gene expression at the 6 and 12-h time points [12].


TLR3, TLR7 and TLR9 are all responsible for viral recognition and the induction of Type I, or IFNI± and IFNI? responses [12]. Expression in terms of fold change values of selected innate immune genes in BIEC by qRT-PCR following treatment with Lp299v is shown. Error bars indicate A±1-SD of the dataAfter analysis of genes at all time points, we selected 6-h as the optimum time to prime the BIEC prior to infection with BRV to study the roles of transcriptional gene regulation in controlling BRV infection.
This point provided the best balance of gene expression, along with a logistical point for further experimentation. Once the time point of priming with Lp299v was established, BRV was added into the flasks following priming.
Thirty-two genes were selected based on information they provided in the probiotic-priming experiment. The change in these genes was analyzed, and expression revealed that homeostasis is maintained, or gene expression is moderately increased in important cytokines and chemokines.In order to better view the innate immune response with respect to primed versus non-primed BRV-infected cells, a classification analysis was performed using Gene Tree cluster analysis. The result displays gene expression changes at each time point being reflected as a color change (Fig.A 2). In looking at selected genes with Lp299v priming, with the exception of TLR4, all genes show steady expression throughout all time points, with slight upregulation in most of the genes. Genes that are deregulated when the cells are not first primed, such as IL-10 and TLR9, show no change in expression or increases in expression when the cells are first primed with Lp299v. Time course of transcriptional expression profile clustering by K-means of selected innate immune genes following BRV infection in Lp299v-treated and untreated BIEC is shown.
Blue on the color code indicates downregulation, red upregulation and yellow no change with respect to time-matched media controlIn analyzing gene expression, Fig.A 2, promising results were found when looking at the Type I interferons. IFN production leads to the activation of macrophages, cells that can induce TNFI± and IL-6 (which have antiviral effects).
IFN1I± shows slight upregulation when BIECs are primed with the probiotic prior to infection. This theme is present throughout the time course, with cytokine and chemokine expression either maintaining homeostasis, or being slightly upregulated when cells are primed with Lp299v.
Given there is no IFNI? in BIEC, IFNI± and IFNI? are responsible for the antiviral effects they produce in cells that express these. Interferons are not constitutively expressed in cells, so cellular induction with probiotic priming prior to rotaviral infection could mean that the BIECs are in a more protected state. On the system level, increased interferons prior to infection could lead to faster clearing of the virus from the system.Further analysis of the differentially expressed genes and their ontology-based classifications revealed some important physiological information to understand how probiotics or commensal microbiota can prepare the hosta€™s immune system to combat infection. TableA 1 shows various gene ontology (GO) terms and descriptions with a number of genes as extracted from our selected innate immune genes studied for the current report following 6-h of incubation of BIEC with Lp299v.
Similarly, TableA 2 reveals the list of differentially expressed genes with GO classifications at 18-h post-BRV infection of BIEC.
By comparing the two tables, it is obvious that Lp299v priming of BIEC enriches 19 different GO classes while BRV affects 11 different classes. FigureA 3 demonstrates the gene expression profile of selected innate immune genes following BRV infection in the presence and absence of Lp299v priming of the BIEC. This pathway analysis shows all time points with and without priming using Lp299v within the same pathway for comparative analysis.
Results from the pathway analysis in controlling BRV infection in the presence of Lp299v were consistent with the plaque assay. KEGG description of TLR pathway with expression profile for Lp299v-primed and non-primed BIEC with BRV infection. The left half of each gene box contains non-Lp299v-primed BIEC at 2, 4, 6, 12 and 18-h post-BRV infection, respectively. The right half of each gene box contains Lp299v-primed BIEC at 2, 4, 6, 12 and 18-h post-BRV infection, respectively. Color bar on the right indicates the expression level as a fold increase or decreaseWith the advent of establishing in vitro BIEC that are susceptible to infection by BRV, it has become possible to study the gene expression of the cells in a wide array of conditions effectively and efficiently.
The use of the probiotic Lp299v as a possible mechanism of inhibiting the BRV infection cycle has shown promising results in limiting growth of the virus, along with inducing the expression of selected genes involved in the innate immune response to prime the immune system prior to infection.Results of the rotaviral infection with and without Lp299v priming show that the probiotic is capable of maintaining consistent gene expression over time and thus homeostasis, or in some cases a boosted immune response in the presence of rotavirus. This indicates that symptoms caused by the deregulation of innate immune gene expression could be alleviated through probiotic supplementation.
With probiotic priming, transcriptional increases in genes responsible for the recognition of the virus were observed.
This is a sign that the clearing of the virus through recognition by the innate immune system and subsequent stimulation of the adaptive immune system are enhanced.Use of the plaque assay to assess the change in live virus over time showed that the probiotic may be capable of decreasing the levels of virus, thus preventing entry into the BIEC altogether. This is an exciting possibility, as the prevention of infection is an optimal solution.This experiment has furthered current understanding on the mechanism of immune enhancement by gut microbiota.
It has both qualitatively and quantitatively shown in vitro that immune stimulation and protection from viral infection are possible with probiotic supplementation.
On a larger scale, these findings may be applied to our general understanding of the pathways of infection in the bovine mucosal model.
The ultimate endpoint to this experiment is testing an in vivo model of probiotic priming followed by a viral infection.AcknowledgmentsWe thank Elaine van Moorlehem for supplying and adapting the EVM07-BIEC. We thank Jamille Heer for maintaining the BIEC and Wayne Connor for technical assistance in running the qRT-PCR analysis. We would also like to thank Neil Rawlyk for technical assistance in growing the bacterial culture.



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