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Metabolism of Short Chain Fatty Acids in the Colon and Faeces of Mice After a Supplementation of Diets With Agave FructansAlicia Huazano-Garcia1 and Mercedes G. Variations on the Efficacy of Probiotics in PoultryLuciana Kazue Otutumi1, Marcelo Biondaro Gois1, Elis Regina de Moraes Garcia2 and Maria Marta Loddi3[1] Universidade Paranaense,, Brazil[2] Universidade Estadual do Mato Grosso do Sul,, Brazil[3] Universidade Estadual de Ponta Grossa,, Brazil1.
From there, several studies have been made and continue being developed with the use of probiotics. Dairy Propionibacteria: Less Conventional Probiotics to Improve the Human and Animal HealthGabriela Zarate1[1] Centro de Referencias para Lactobacilos (CERELA)-CONICET, San Miguel de Tucuman, Argentina1. Propionibacteria are heterotrophic microorganisms that mean they need an organic carbon source to grow and posses a fermentative metabolism [41-43]. A total of 800 one-day-old chickens (Cobb 500) were obtained from a local hatchery and allocated randomly to five dietary treatments. Antibiotics have been extensively used as feed additives and growth promoters in animal feed industry. A total of 800 one-day-old chickens of the Cobb500 strain were obtained from a local hatchery.
Two Iranian indigenous herbal extracts, garlic (Allium Sativum) and thyme (Thymus Vulgaris) were added to a diet based on corn and soybean meal (CON diet). Non-significant effect of herb extracts of garlic and thyme on the performance of broiler chickens may be due to the hygienic situation of experiment.
There is little information available about the effect of garlic and thyme extracts on triglyceride, HDL and hematocrit concentrations. The results in this study showed that application of two herb extracts, garlic and thyme, had no significant effect on performance and percentage of carcass components (thigh, breast, abdominal fat, and carcass weight).
Daily consumption of an apple or apple polyphenols could help to ‘keep the doctor away’ by lowering levels of LDL cholesterol by up to 40%, suggest researchers. The disparity resulted from the differences in polyphenol composition differences between the extract, which came from a number of apple varieties, and the whole apples, which were just two varieties. Get FREE access to authoritative breaking news, videos, podcasts, webinars and white papers. Roundtable discussion: A lighter world: whose fault is globesity and what is being done to combat it? Concentration of short chain fatty acids generated by bifidobacteria and lactobacilli from the fermentation of Dasylirion sp. Lopez1[1] Departamento de Biotecnologia y Bioquimica, Centro de Investigacion y de Estudios Avanzados del IPN, Mexico1. Proposed interactions between competitive exclusion products, probiotics or immunostimulants, and avian intestinal immunity.
IntroductionIn face of the current debate about the use of antibiotics as growth promoters, due to the probable relationship with resistance to antibiotics used in human medicine, the presence of antibiotic residues in products of animal origin intended for human consumption and the emergent demand from consumer market for products free from additive residues, it was necessary to search for alternative products that could replace antibiotics used as promoters, without causing losses to productivity or product quality. Inconsistent results from the use of probiotics in animal production have been a constraint for the promotion of their use. Influence of bacterial surface components on lectins removal (a) and adhesion property (b). Transmission electron microscopy photomicrographs of the microvillous surface of the small bowel of mice fed with Con A (Group 2) (panels a-b) and those that consumed lectin plus propionibacteria (Group 4) (Panels c-d). IntroductionProbiotics are live microorganisms that confer health benefits to the host when administered in adequate amounts.
From a safety point of view, classical species have a long history of safe application on industrial processes whereas members of the cutaneous group are commonly considered opportunistic pathogens in compromised hosts.
The use of antibiotics as feed additives is hazardous due to crossresistance and multiple resistances of pathogens (Schwarz et al., 2001). Some herbs or herbal extracts can beneficially affect feed intake, secretion of digestive tract juices and immune system of animals (Mikulski et al., 2008).
One ml of blood was sampled in an EDTA-tube, mixed vigorously, and centrifuged for 10 minutes at 2000 rpm (Hettich EPA30, Germany) to obtain plasma. A general linear model was used to analyze the effect of diets on the broilers performance. In this study birds were kept in clean disinfected environment following all hygiene regulations.
Dietary treatments did not induce any significant effect on the carcass composition parameters including carcass yield (%), breast (%), thigh (%), and abdominal fat pad (%).
Although, this experiment was performed in a disinfected condition that may have resulted in a decreased performance of these extracts. Schematic representation of the crosstalk between probiotic bacteria and the intestinal mucosa. Effectiveness of probiotics in the prevention of Salmonella colonization in broiler chicken. An alternative is the use of probiotics, which are products made from living micro-organisms or their L-forms (without cell wall).
Variations in the efficacy of probiotics can be due to the difference in microbial species or micro-organism strains used, or with the additive preparation methods (Jin et al., 1998a). Reproduced from Zarate and Perez Chaia, Food Research International (2012), 47(1): 13-22 [145].
In the last decades there has been a great interest from food and pharmaceutical industries to develop products containing probiotics due to the great demands of healthy foods and alternatives to conventional chemotherapy. In consequence, the economic relevance of propionibacteria derives mainly from the industrial application of dairy species as cheese starters and as biological producers of propionic acid and other metabolites with a more recent interest on their usage as health promoters.“Dairy or classical” propionibacteria “Cutaneous” propionibacteriaP. The production of propionic acid by these bacteria involves a complex metabolic cycle with several reactions in which substrates are metabolized to pyruvate via glycolysis, pentose phosphate or the Entner-Doudoroff pathways, generating ATP and reduced co-enzymes. Therefore, European Union has banned the application of most of antibiotics in poultry diets.
Essential oils are derived mainly from herbs and spices and their purified compounds have antimicrobial properties (Faleiro et al., 2003).
The erythrocyte count was performed on EDTA mixed blood (Sysmex K21, UK-Japan) and plasma total lipid was obtained using an auto analyzer (Mindrey PM 200, China). The effects of diet and sex on the carcass composition, blood lipids and hematocrit were analyzed by exploiting a factorial design. In addition to the dietary treatments, the effect of sex was also not meaningful on hematocrit and plasma lipid profile. Sex of chickens did not have any significant effect on the carcass composition (data not shown).
More effective influences of these extracts could probably be seen in chickens rearing in the non-hygienic situation. FructansMost plants store starch or sucrose as reserve carbohydrates, but approximately 12-15% of higher plants (representing more than 40,000 species) synthesizes fructans as their main source of carbohydrates [1]. The micro-organisms included as probiotics are usually assumed to be non-pathogenic components of the normal microflora, such as the lactic acid bacteria. However, other factors can justify the variations in the results of probiotic use in poultry, such as origin species, probiotic preparation method, survival of colonizing micro-organisms to the gastrointestinal tract conditions, environment where the birds are raised, management (including the application time and application route of the probiotic), the immunologic status of the animals, the lineage of the poultry evaluated, as well as age and concomitant use or not of antibiotics.
Although the great bulk of evidence concerns lactobacilli and bifidobacteria, since they are members of the resident microbiota in the gastrointestinal tract, other less conventional genera like Saccharomyces, Streptococcus, Enterococcus, Pediococcus, Leuconostoc and Propionibacterium have also been considered.
Bassett (2000) reported that adding herbal essential oils to the feed or water increases feed conversion and weight of broiler chickens. Our results also showed no significant effect of garlic and thyme extracts on performance and carcass traits of broilers. Therefore, more studies with different growth facility need to be performed to analyze the effect of herb extracts on the performance, carcass yield, and hematology of poultry. The effects of an essential oil combination derived from selected herbs growing wild in turkey on broiler performance.
Effects of organic acids or natural plant extracts added to diets for turkeys on growth performance, gastrointestinal tract metabolism and carcass characteristics. Use of antibiotic growth promoter and two herbal natural feed additives with and without exogenous enzymes in wheat based broiler diets. Fructans are found naturally in plants as a heterogeneous mixture of different polymerization degrees, they are a polydisperse mixture. Probiotic bacteria can also enhance intestinal barrier function by (5) increasing mucus production (Adapted Ng et al., 2009). However, there is good evidence that non-pathogenic variants of pathogenic species can operate in much the same way as traditional probiotics. Thus, the aim of this review is to discuss the use of probiotics in poultry, with emphasis on the type of probiotic and micro-organisms used, action mechanism and its relation with the variations on the results of poultry survey.2. The latter transformation occurs via the Wood-Werkman cycle or transcarboxilase cycle which represents the key component of the central carbon metabolic pathway in propionibacteria [41]. There were no significant effects among the treatments on weight gain, feed intake and feed conversion ratio over the entire trial.
Birds were killed by cervical dislocation on day 42 of the experiment, and carcasses were weighed and dissected. This was in accordance with some other investigations showing that garlic powder and oils did not decrease blood cholesterol (Berthold et al., 1998).
Effect of dietary thyme oil and vitamin E on growth, lipid oxidation, meat fatty acid composition and serum lipoproteins of broilers.
The use of natural feed additives as alternatives for an antibiotic growth promoter in broiler diets. Influence of two plants extracts on broiler performance, digestibility and digestive organ size.
Production of short chain fatty acids in the caecum, colon and faeces in rats associated with human faecal contents and fed with inulins of different chain lengths.
Among plants that store fructans, many are economically important, due to its content of fructans, as it is the case of chicory (Cichorium intybus), agave (Agave spp.), artichoke (Cynara scolymus), dahlia (Dahlia variabilis), garlic (Allium sativum) and wheat (Triticum asetivum) [2, 3]. For example, avirulent mutants of Escherichia coli, Clostridium difficile, and Salmonella Typhimurium can also protect against infection by the respective virulent parent strain (Fuller, 1995).
Type of probiotic and micro-organisms usedThere are several types of probiotics available in the market to be used in poultry, with a range of micro-organisms present and, therefore, with different metabolic activities and action modes. Dairy propionibacteria are generally recognized as safe microorganisms whereas members of the cutaneous group have shown to be opportunistic pathogens in compromised hosts.
The most important reaction of this cycle is transcarboxylation that transfers a carboxyl group from methyllmalonyl-CoA to pyruvate to form oxaloacetate and propionyl-CoA, without ATP consumption. Also, no significant differences were observed between CON group and other treatments in plasma parameters (cholesterol, triglyceride and HDL) and hematocrit.
To the best of author's knowledge there is no report on the effect of indigenous Iranian herbal extracts on the broiler performance. In this trial, the use of growth promoter combinations may have had desirable effect on bird's performance.
The effect of a mixture of herbal essential oils or a-tocopheryl acetate on performance parameters and oxidation of body lipid in broiler.
Concentration of total short chain fatty acids production in the caecum and the three sections of the colon in mice fed a standard (STD) diet or diet supplemented with Raftiline (RNE-inulin) or Agave angustifolia Oax. Glucose, triglycerides and cholesterol levels in plasma of mice fed with a standard (STD) diet or diet supplemented with Raftiline (RNE) and A. Five different groups of fructans have been found in nature and distinguished according to the type of linkage between fructose units and the position of the glucose moiety within the structure. Also, they present variations as to the capacity of colonizing the intestine or not, which justifies variations on the results of their use.Bacillus, Bifidobacterium, Enterococcus, E. In consequence, the economic relevance of propionibacteria derives mainly from the industrial application of dairy species as cheese starters and as biological producers of propionic acid and other metabolites like exopolysaccharides and bacteriocins to be used as thickeners and foods preservers, respectively.
The enzyme catalyzing this reaction is a methylmalonyl-CoA carboxytransferase that has been fully characterized and its structure resolved [34; 40].

Therefore, the aim of the present study was to investigate the effect of two indigenous herbal extracts: garlic (Allium Sativum) and thyme (Thymus Vulgaris) on the broilers performance, carcass composition, and plasma parameters. Effect of dietary oregano essential oil on performance of chickens and on iron induced lipid oxidation of breast, thigh and abdominal fat tissues. The effect of dietary inclusions of herbs and their volatile oils on the performance of growing chickens. Effect of dietary garlic (Allium Sativum) on performance, carcass composition and blood chemistry changes in broiler chickens. These groups consist of inulins, neoseries inulins, levans, neoseries levans and graminans. In their experiments, the authors observed that the intestinal contents of normal adult birds, orally administered to chicks with one day of age, altered their sensitivity to infection by Salmonella spp. Cytotoxic effects of lectins, and protection of colonic cells by lectin removal by propionibacteria. However, the ability of dairy propionibacteria to improve the health of humans and animals by being used as dietary microbial adjuncts has been extensively investigated. Then, oxaloacetate is reduced to succinate, via malate and fumarate in two NADH requiring reactions. Supplementation of herb extracts did not affect carcass characteristics including carcass yield, breast, thigh and abdominal fat. Effects of dietary fat source and subtherapeatic levels of antibiotics on the bacterial community in the ileum of broiler chickens at various ages. However, even those belonging to the same species can have different strains and even these different strains from the same species can have different metabolic activities.
In this sense, our research group has been studying for the last two decades the probiotic properties of dairy propionibacteria isolated from different ecological niches. There were no significant differences observed in carcass characteristics between males and females fed with different levels of herb extracts. Jamroz and Kamel (2002) showed that broiler chickens fed with herb extract had higher breast weight compared to CON group.
Currently, inulins are extracted from chicory roots, containing fructose chains having a degree of polymerization (DP) from 3 to 60 [2] (Figure 1a). In the present article the current evidences supporting the potential of dairy propionibacteria to be used as probiotics are reviewed focusing in a less studied mechanism such as the protection of the intestinal mucosa by the binding of dietary toxic compounds.
Methylmalonyl-CoA is also regenerated from succinyl-CoA during propionate production, thus creating the second of the two transcarboxylase cycles, and can react with a new molecule of pyruvate. In conclusion, supplementation of garlic and thyme extracts as feed additives at two levels of 3 and 6% in the present study did not improve performance and carcass traits of broiler chicks. Antimicrobial activity of essential oils isolated from Portuguese endemic species of Thymus. The chemical or enzymatic (endoinulinases) hydrolysis of inulins produces inulins of shorter DP (DP<10), these are called fructooligosaccharides (FOS) [5, 6]. Viability was assessed by counting cells under the fluorescence microscope after propidium iodide ? fluorescein diacetate ? Hoescht staining.
Nowadays there are clear evidences that propionibacteria used alone or combined with other microorganisms can exert beneficial effects in the host. Department of Animal Science Virginia Polytechnic Institute and State University, Blacksburg, Virginia USA.
Mexico is considered the origin center of evolution and diversification of the Agave genus, since a large number of agave species are found in its territory. It must be emphasized that the Wood Werkman cycle used by propionibacteria to produce propionate is coupled to oxidative phosphorylation and yields more ATP than in the other bacteria producing propionic acid [42, 43].Depending on the strains, the substrate used, and the environmental conditions, propionibacteria modulate the proportions of pyruvate either reduced to propionate, or oxidised to acetate and CO2, to maintain the redox balance [43]. The Agave genus includes approximately 166 species and is the largest genus among the Agavaceae family that consists of 9 genera and approximately 293 species [7, 8]. The most documented probiotic effects for propionibacteria within these categories include: bifidogenic effect in the human gut, improvement of nutrients utilization, hypocholesterolemic effect and anticarcinogenic potential immune system stimulation. The agave plants have the ability to grow in extremely dry-hot environments, where sometimes this plant is the predominant or exclusive flora in that type of a geo-climatic zone, however, they can also be found in diverse ecosystems, such as productive highlands and elevated humidity [9]. Different studies have also described the ability of dairy propionibacteria to bind and remove toxic compounds from different environments such as the gut and food. These plants present a crassulacean acid metabolism (CAM) and their principal photosynthetic products are fructans [10], fructans are synthesized and stored in the stems of agave plants. During lactate fermentation, aspartate is deaminated to fumarate by an aspartate ammonia lyase; fumarate is then converted to succinate, with a concomitant production of NAD and ATP.
Agave is the most exploited genus and economically important as the raw materials are used on the production of alcoholic beverages such tequila (A. It has been proposed that probiotic microorganisms may reduce by binding, the availability of free toxic compounds within the intestinal tract which reduces in turn, their negative effects. Various agarized media with different degrees of selectivity have been used for detection and enumeration of classical propionibacteria in dairy environments, animal and human fecal samples. In this respect, in recent years we have been investigating the potential of dairy propionibacteria to protect the intestinal mucosa from the toxic and antinutritional effects of some common dietary substances like the plant lectins from the Leguminosae family. Among them it could be mentioned YELA [20], Pal Propiobac® medium, which contains glycerol, lithium lactate and antibiotics [21] or others including lithium chloride and sodium lactate in concentrations high enough to limit the growth of accompanying bacteria [22]. By in vitro and in vivo studies we have determined that certain strains are able to bind and remove different dietary lectins from media, preventing their cytotoxic effects on intestinal epithelial cells.
Although these media may be successful for the isolation of classical and cutaneous strains of Propionibacterium, they have limitations for selective enumeration of bacteria in very complex ecosystems like intestinal microbiota. They can grow in a minimal medium containing ammonium as the sole nitrogen source, but a higher growth is observed in media containing amino acids [45].Although P. Propionibacteria reduced the incidence of colonic lesions, the enlargement of organs, the disruption of brush border membranes and the decrease of their disaccharidase activities.
Since consumption of suitable propionibacteria may be an effective tool to avoid lectin-epithelia interactions, further investigations on their potential as probiotic detoxifying agents are actually ongoing With regard to animals’ health it has been reported that dairy propionibacteria directly fed to farm animals increased weight gain, food efficiency and health of many animals like chickens, laying hens, piglets and cows. Genus and species-specific primers targeted to the genes encoding 16S rRNA for PCR-based assays were also designed for detection of dairy propionibacteria [29].
Some proteinases have been described for Propionibacterium, one cell wall associated and one intracellular or membrane bound but their activities are weak.
With a wider insight, propionibacteria may be assayed as probiotics for other ruminants like goats and sheep since their milk-derived products are highly appreciated by consumers.
By contrast, different peptidases such as aminopeptidases, proline iminopeptidase, proline imidopeptidase, X-prolyl-dipeptidyl-amino-peptidase, endopeptidases and carboxypeptidase, have been described. It should be emphasized that much of the health benefits described above could be related to the ability of propionibacteria to remain in high numbers in the gastrointestinal tract by surviving the adverse environmental conditions and adhering to the intestinal mucosa.On the basis of the GRAS status of dairy propionibacteria and the positive results obtained by us and other authors, further studies are encouraged in order to select the appropriate strains for developing new functional foods that include these bacteria for human and animal nutrition. A FISH protocol and oligonucleotide probes targeting the 16S rRNA of dairy propionibacteria were developed in our laboratory [32] and successfully used for enumeration of P.
Agave fructans posses a molecular structure compose of a complex mixture containing highly branched molecules with ?(2-1) and ?(2-6) linkages, as well as internal and external glucose units, due to the existence of both types of glucose, agave fructans have been classified as graminans (external glucose) and agavins (internal glucose) [11] (Figure 1b). Amino acids, especially aspartic acid, alanine, serine and glycine, are degraded by Propionibacterium, with variations among species and strain [47].
Finally, a real-time PCR method, based on the transcription of the enzyme transcarboxylase involved in propionic fermentation, was successfully used to detect a strain of P. On the other side, cutaneous propionibacteria, have the ability to hydrolyze different proteins, like gelatin and fibronectin, and to promote damages and inflammation of the host tissues.Regarding vitamins, all propionibacteria strains require pantothenate (vitamin B5) and biotin (vitamin H).
General features and taxonomyPropionibacteria are Gram positive, catalase positive, high G+C%, non spore forming and non motile pleomorphic bacteria [1, 2]. In general, microorganisms of the genus Propionibacterium are anaerobic to slightly aerotolerant and morphologically heterogeneous including rod-shaped and filamentous branched cells that may occur singly, in pairs forming a V or a Y shape, or arranged in “Chinese characters”. They have a peculiar metabolism leading to the formation of propionic acid as main end-product of fermentation.
The G+C content in their DNA is in the range of 53-68 mol% and although they generally do not possess plasmids their existence has been reported in strains of P. Long term and stress survival of Propionibacteria It is known that propionibacteria are able to adapt and survive to different stresses like industrial processes and the gastrointestinal transit, as well as to remain active for long periods of time in such adverse environments [43]. Short chain fatty acids (SCFAs)The gastrointestinal tract is an extremely complex ecosystem containing about 1011 CFU (colony forming units) of bacteria per gram of intestinal content. Although in 1861, Louis Pasteur demonstrated that propionic fermentation was due to the biochemical activity of microorganisms, the first studies about the morphology and physiology of propionibacteria were carried out by Albert Fitz (1879) [3], who observed that organisms from cheeses with “eyes” ferment lactate to propionic and acetic acids and liberate carbon dioxide. In this sense, the manufacture of a swiss type cheese represents for microbial starters successive stresses like acidification of the curd, heating during cooking, osmotic stress due to brining, and low temperature (4 to 12 °C) during cheese ripening. This large population of bacteria plays a key role in the nutrition and health of the host [18]. The transit through the digestive tract also suppose stressful conditions for bacteria such as gastric acidity and the presence of other aggressive intestinal fluids like bile and pancreatic enzymes.
The colonic microbiota ferments organic material that cannot be digested otherwise by the host in the upper gut. Von Freudenreich and Sigurd Orla-Jensen (1906) [4] isolated the bacteria responsible for the “eyes” formation in Emmental cheese and some years later, the name Propionibacterium was suggested by Orla-Jensen [5] for referring to bacteria that produced large amounts of propionic acid.
These include resistant starch, non-digestible carbohydrates (fructans) as well as some proteins and amino acids [19]. Although several strains were isolated during the following years these microorganisms were not included in the Bergey’s Manual of Determinative Bacteriology till the third edition published in 1930. Since then, new species were described on the basis of their morphological and biochemical characteristics such as their typical pattern of Chinese characters, propionic acid production, and carbohydrate fermentation profile. The redundancy and inducibility of this chaperone and protease machinery is in agreement with the ability of P. In 1972, Johnson and Cummins [6], classified strains with several common features into eight homology groups based on DNA-DNA hybridization and peptidoglycan characteristics. Fructans fermentation also produces a few gases as CO2, CH4, H2 and additionally heat [19, 23]. This study was the basis for the classification of propionibacteria into “dairy or classical” and “cutaneous” groups included in the 8th edition of Bergey’s Manual of Determinative Bacteriology (1974). The presence of both, non-digestible carbohydrates and SCFAs in the colon can positively alter the colonic physiology drastically [24]. Acid and bile stresses, induce the synthesis of the following proteins: pyruvate-flavodoxin oxidoreductase and succinate dehydrogenase which are involved in electron transport and ATP synthesis, as well as glutamate decarboxylase and aspartate ammonia-lyase, which are involved in intracellular pH homeostasis. Bile also induces oxidative stress so that survival and activity within the gut depend on remediation of oxidative damages. An increased in SCFAs synthesis also creates a more acidic environment in the gut, which is important in vivo in terms of colonization resistance against pathogens [18, 20]. The production of SCFAs is affected by many factors, including the source of substrate [26], in particular, the chemical composition of the fermentable substrate, the amount of substrate available, its physical form (e.g. The gut of mice comprises four sections: caecum, proximal, transverse (medial) and distal colon. The caecum and proximal colon are the main sites where fermentation is carried out, given the number of bacteria and the availability of substrate, because as it moves through the intestine toward the distal colon, there is a lower concentration of water as well as a depletion of carbohydrates and increased pH [22]. SCFAs are rapidly absorbed in the caecum and colon being excreted in the faeces only from 5% to 10% of them [24]. The same scheme was followed in the first edition of Bergey’s Manual of Systematic Bacteriology [1]. Other inducible proteins involved in protection and repair of DNA damages include Ssb protein which is involved in DNA recombination and repair, as well as Dps which protects DNA against oxidative stress are stress-induced in P. In 1988, on the basis of 16S rRNA sequences, the species Arachnia propionica was reclassified as Propionibacterium propionicus [7]. Then, in Bergey’s Manual 9th edition (1994), the classification of previous edition was maintained but the subspecies P. Stress tolerance and cross-protection in strains of Propionibacterium freudenreichii were examined after exposure to heat, acid, bile and osmotic stresses. The presence of acetyl-CoA synthetase in the cytosol of adipose and mammary glands allows the use of C2:0 for lipogenesis once it enters the systemic circulation [24].

By contrast, some other heterologous pretreatments (hypothermic and hyperosmotic) had no effect on tolerance to bile salts.
Heat and acid responses did not present significant cross-protection and no cross-protection of salt-adapted cells against heat stress was observed for these propionibacteria [48-50].In addition, long term survival of propionibacteria on adverse environments could be due to the accumulation of storage compounds, compatible solutes, and the induction of a multi-tolerance response under carbon starvation [40]. C3:0 is also a substrate for hepatic gluconeogenesis and it has been reported that this acid inhibits cholesterol synthesis in hepatic tissue [31, 32]. Short chains of PolyP are synthesized when bacteria grow on glucose whereas long chains are accumulated when the main carbon source is lactate. It is the most important SCFA in colonocytes metabolism, where 70% to 90% of C4:0 is metabolized by the colonocytes.
The synthesis of PolyP is catalysed by polyphosphate kinase (PPK) that transfers the terminal phosphate of ATP to polyP. It is proposed that PolyPs enable microorganisms to tolerate adverse conditions since ppk mutants are unable to survive during stationary phase [51]. Approximately 95% of the C4:0 produced by colonic bacteria is transported across the epithelium, but concentrations in portal blood are usually undetectable as a result of a rapid utilization [33]. At present, the genus Propionibacterium is classified as Actinobacteria with a high G+C content, that make them more related to corynebacteria and mycobacteria than lactic acid bacteria. Classical propionibacteria include among their main habitats: raw milk and cheese [1, 2] but have been obtained also from silages and vegetables for human consumption [15], and from ruminal content and feces of cows and calves [16]. These enzymes seem to be involved in intracellular accumulation and hydrolysis of glycogen as neither P.
Furthermore, they are not limited to the gastrointestinal tract of ruminants being also isolated from the intestine of pigs and laying hens [17]. On the other side, cutaneous species are found mainly in the human skin, but have been isolated also from the intestine of humans, chicken and pigs [1, 2, 18], being best represented by the acne bacillus, Propionibacterium acnes. Trehalose is a non-reducing disaccharide that can be used by bacteria as a carbon and energy source and also can be accumulated as a compatible solute. The authors concluded that these differences correlated well with the structural differences, FOS has a short DP and inulin a long DP.
All dairy propionibacteria are able, in a strain dependent manner, to synthesize and accumulate trehalose from glucose and pyruvate [53]. Both processes are enhanced at stationary phase and under oxidative, osmotic, and acid stress conditions [54].
These researchers found that short inulins were rapidly fermented and produced higher concentrations of C4:0 compared with other inulins, hence, chain length is an important factor on the fermentation patterns of SCFAs (Table 1).
This long-term survival in stationary phase or dormant phase could be the consequence of a multi-tolerance response that involves the synthesis and accumulation of polyP, glycogen, trehalose and the over-expression of molecular protein chaperones.
Besides, a gene encoding an Rpf (resuscitation promoting factor) protein which is essential for the growth of dormant cells from actinobacteria has been described in the genome of P freudenreichii and is probably involved in long-term survival of propionibacteria [40]. The differences in SCFAs patterns in these studies may be also attributed to differences in bacteria species present in the faecal inocula or fermentation process used.
Dairy starters for Swiss-type cheeses and other productsThe main industrial application of the genus Propionibacterium is the usage of “classical propionibacteria” as dairy starters for the manufacture or Swiss type cheeses. These researchers found that branched fructans from Dasylirion (DSC) with a DP range from 3 to 20 and A. However, propionibacteria can also be used in the manufacture of various cheeses without eyes just to enhance flavour formation [58].In swiss type cheeses, propionibacteria may be present either as contaminants of raw milk or as components of starter cultures. The typical starter for this variety includes Streptococcus thermophilus, Lactobacillus helveticus, Lactobacillus delbrueckii subsps. Moreover, the major SCFAs fermentation product, were acetic, formic and lactic acids, wherein the proportions of the acids varied depending on the prebiotic type used by the different bacteria. Figure 4 shows the fermentation products only for the two agave fructans (DSC and ATG) and commercial inulins (RSE and RNE) that better stimulated the growth of bacteria. During manufacture and early stages of ripening, the thermophilic bacteria develop at expense of lactose of milk being responsible for lactic acid production, and also contributing to casein hydrolysis during pressing of the cheese. Interactions between microbiota and milk throughout ripening lead to biochemical changes that result in the development of the typical texture and flavor.
During maturation in the cold room (15 °C) most of lactic starter lyse and release peptidases that produce free amino acids, which are precursors of many flavor compounds. The subsequent period of warm room ripening is characterized by a marked growth of propionibacteria that metabolize the lactate produced by the lactic acid bacteria into propionate, acetate and CO2. The authors observed that agave fructans stimulated the growth of bifidobacteria and lactobacilli more efficiently (2-fold) that commercial inulins, either long- or short-DP. They also reported that short-DP fructans in the mixtures highly influenced the rate of fermentation by probiotc bacteria.
As described above, the end-products of propionic fermentation are considered as flavour compounds in cheese whereas the co-metabolism of aspartate leads to additional CO2 production. However, strains with a high ability to metabolise aspartate can be associated with undesirable slits and cracks [60].Propionibacteria degrade branched-chain amino acids to branched-chain volatile compounds mainly 2-methylbutanoic acid and 3-methylbutanoic acid, which derive from isoleucine and leucine degradation, respectively [61].
These important flavour compounds are almost entirely produced in cheese by propionibacteria that synthesize them in closely related manner to that of cell membrane fatty acids [62]. Their results indicated no significant differences among the growth of both bacteria genera with the different fructans used. Two esterases, one extracellular and other surface-exposed seem to be involved in lipolysis of milk glycerides [63, 64].
Other dairy products such as yogurt and fermented milks seem to be less appropriated for delivery of propionibacteria due to their weak proteolitic activity, the presence of inhibitory substances and the low pH attained by lactic acid fermentation that do not allow their development.
Currently, yogurt is used to deliver probiotic propionibacteria to the host’s intestine or to produce nutraceuticals, but in both cases inoculums higher than those used for cheese manufacturing are necessary. Antimicrobials production: Propionic acid and bacteriocinsPropionic acid and its salts, as well as Propionibacterium spp strains, are widely used as food and grain preservatives due to their antimicrobial activity at low pH.
They are commonly incorporated in the food industry to prolong the shelf-life of many products by suppressing the growth of mold and spoilage microorganisms in bread and cakes, on the surface of cheeses, meats, fruits, vegetables, and tobacco. Most commercial propionic acid is produced by petrochemical processes since biosynthesis by microbial fermentation is limited by low productivity, low conversion efficiency, by-product formation (acetic acid and succinic acid) and end-product inhibition.
However, different attempts have been made to improve biological production of propionic acid for industrial applications [68]. In this sense, it has been determined that the most appropriated species for bioproduction of propionic acid from carbohydrate-based feedstock, including glucose and whey lactose, is P.acidipropionici [69, 70].
Since the use of glycerol as the principal carbon source enables the production of propionic acid without acetic acid, recent investigations have focused on the optimization of this particular homopropionic fermentation by propionibacteria [71, 72].
Two commercial products that include propionibacteria or their metabolites aimed for controlling spoilage microorganisms are currently available at market. Microgard™ is a food grade biopreservative obtained by fermentation of skim milk with Propionibacterium shermanii that is active against some fungi and Gram negative bacteria, but not against Gram positive ones [73]. Production of SCFAs in vivoInulinsNilsson and Nyman [40] evaluated the formation of SCFAs in the hindgut of rats fed with lactulose, lactitol, FOS and inulins of different DP and solubility. Besides, they exert anticarcinogenic effects by inducing apoptosis of neoplastic cells but not of healthy mucosa [77].
Finally, SCFA may exert hypocholesterolemic effects, since propionate lowers blood glucose and alters lipid metabolism by suppressing cholesterol synthesis in the liver and intestine [78].Bacteriocins are antimicrobial peptides or proteins encoded by plasmid or chromosomal DNA of a wide range of Gram positive and negative bacteria.
They have an antagonistic activity against species genetically related to the producer strain, but many of them exhibit a rather wide spectrum of activity and inhibit the growth of spoilage and pathogenic bacteria belonging to other genera [79].Both starters and naturally occurring bacteria on food have the ability to produce bacteriocins. Hence, they may have potentially important applications as food biopreservatives or bacteriocin-producer probiotics to inhibit intestinal pathogens [80].
Moreover, FOS, inulins and the mixture of FOS-inulins increased the amount of C4:0 acid in the caecum and colon of the rats fed with the mixture regard to animals fed with standard diet. The authors concluded that the type of diet and the fermentation site in the colon affected the concentration of SCFAs (Table 2). Nutraceuticals production: CLA, vitamins, EPS and trehalosePropionibacteria are able to produce many biological compounds that enhance the human health so they can be used as “nutraceuticals cell factories” for food enrichment.
In this regard, propionibacteria have already been considered as rich sources of conjugated linoleic acid, vitamins, exopolysaccharides and trehalose.Many health benefits have been attributed to consumption of CLA-containing foods such as anticarcinogenic, antiatherogenic, antidiabetogenic and antioxidative properties, immune system modulation and reduction of body fat gain [94].
CLA-isomers are formed by biohydrogenation of LA in the rumen and through conversion of vaccenic acid by ?9-destaurase in the mammary gland so that ruminant meats and milk-derived products are main dietary sources of CLA.
In another study, the same authors fed rats with 10% of inulin, they found a higher concentration of C3:0 acid in the portal vein as well as a significant decrease in plasma cholesterol levels of the rats fed with this diet with regard to animals that consumed the standard diet [44].
However, some microorganisms like Bifidobacterium, Lactobacillus, Enterococcus and Propionibacterium posses a linoleic acid isomerase that allow them to form CLA as a detoxification mechanism [95]. On the other hand, a study carried out using obese rats that received a diet supplement with inulin, a two-fold greater C3:0 concentration in the portal vein and a decrement on triglyceride accumulation in the liver of these animals was observed [45]. In consequence, they have been intended, either as starter or adjunct cultures, to increase the CLA level and nutritional value of some fermented products like yoghurt and cheese. A similar result was seen in hamsters fed with different percentages of inulins (8, 12 and 16%). In this regard, several studies have shown the potential of propionibacteria for producing CLA enriched products. Plasma cholesterol and triglyceride concentrations were significantly lower with all the percentages of inulins studied with respect to hamsters fed with the standard diet [46]. By varying the source of LA for conjugation and the fermentation conditions it has been observed that P. Besides, it has been observed that CLA formation and growth of dairy propionibacteria in fermented milks were enhanced in the presence of yogurt microorganisms whereas organoleptic attributes obtained with yogurt starter cultures were not affected by co-cultures with the propionibacteria [100]. Vitamin B12 also called cobalamin, is an essential nutrient for the human body that plays a key role in the normal functioning of the brain and nervous system, the formation of blood and also the metabolism of every cell, especially affecting DNA synthesis and regulation, fatty acid synthesis and energy production. The pathway of vitamin B12 synthesis in Propionibacterium freudenreichii has been completely elucidated [40, 102]. This microorganism synthesizes cobalamin as a cofactor for propionic acid fermentation [41] and is the only bacteria, among B12 producers that possess the GRAS status of the United States Food and Drug Administration.In consequence dairy propionibacteria are the preferred microorganisms for the industrial production of this vitamin and many efforts have been made to improve the production process by using genetic engineering [102, 103] and other biotechnological strategies like fermentation manipulations [104, 105].
Vitamin B2, also known as riboflavin, is the central component of the cofactors FAD and FMN, and is therefore required by all flavoproteins.
As such, vitamin B2 is required for a wide variety of cellular reactions and is involved in vital metabolic processes in the body. Vitamin K (a group of 2-methyl-1,4-naphthoquinone derivatives), is an essential cofactor for the formation of ?-carboxyglutamic acid-containing proteins that bind calcium ions and are involved in blood coagulation and tissue calcification. Its deficiency has been associated with low bone density and increased risk of fractures from osteoporosis and intracranial hemorrhage in newborns [109]. Vitamin K1 or phylloquinone is present in plants, and vitamin K2, also called menaquinone, is produced in animals and bacteria that live in the intestine.
It has been reported that Propionibacterium freudenreichii produces large amounts of tetrahydromenaquinone-9 (MK-9 (4H)) and the precursor 1,4-dyhidroxy-2-naphtoicacid (DHNA) which is a known bifidogenic factor [110-112]. In order to improve the production of these metabolites, different laboratory culture protocols that could be applied to an industrial scale have been assayed finding out that DHNA production is markedly influenced by carbon source limitation and the oxygen supply. An improvement in DHNA production could be obtained by a cultivation method that combines anaerobic fed-batch and aerobic batch cultures [112, 113]. Both homopolysaccharide [116, 117] and heteropolymers [118] were described and it has been reported that production of EPS by propionibacteria is a strain-dependent property (due to an IS element in the gtf promoting sequence) that is influenced by the medium composition and the fermentation conditions [119, 120].
Further studies are needed to elucidate the role of these polymers and their potential applications.Trehalose has been proposed as a healthy sugar substitute in foods because of its anticariogenic and dietetic properties. As described in paragraphs above, propionibacteria synthesize trehalose as a reserve compound and as a stress-response metabolite [52-55].
With respect to the production of this sugar in situ in food products, it has been observed that P. It contributes to the typical flavor and the development of characteristic “eyes”[56, 57], [59].AntimicrobialsP.

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