For centuries, lactic acid bacteria (LAB) have been used for the preservation of food for human consumption. LAB are a large group of fermentative, anaerobe facultative, aerotolerant microorganisms which are usually present in the gut of humans and other animals, raw vegetables, meat and meat products, and cereals (Carr et al., 2002). In animals, their numbers may vary with the species, the age of the host, or the location within the gut (De Vries et al., 2006). In the food industry, lactic acid bacterial strains are widely employed either as starter cultures or as non-starter lactic acid bacteria. Furthermore, owing to their probiotic properties, several LAB strains are used as adjunctive cultures in foods and feed (Sanders, 2000; Leroy & de Vuyst, 2004).
The term “probiotic” originated from the Greek word “probios” meaning “for life” (as opposed to “antibiotic,” which means “against life”) (Longdet et al., 2011). Probiotics are microbial food supplements which, when administered in adequate amounts, confer health benefits to consumers by maintaining or improving their intestinal microbial flora (Salminen et al., 1998; Reid et al., 2003). The US Food and Drug Administration uses other terms for live microbes for regulatory purposes (Sanders, 2008); live microbes used in animal feeds are called “direct-fed microbials” (FDA, 1995), and, when intended for use as human drugs, they are classiﬁed as “live biotherapeutics” (Vaillancourt, 2006). Probiotics are mainly members of the genera
The research of novel probiotic strains is important in order to satisfy the increasing request of the market and to obtain functional products in which the probiotic cultures are more active and with better probiotic characteristics than those already present on the market (Verdenelli, et al., 2009). According to a recent market research report ‘Probiotics Market (2009-2014)’, the global probiotics market generated US $15.9 billion in 2008 and is expected to be worth US $ 32.6 billion by 2014 with a compound annual growth rate of 12.6 percent from 2009 to 2014 (FB 1046, 2009).
Several aspects, including general, functional and technological characteristics, have to be taken into consideration while selecting probiotic strains (Sanders & Huis in’t Veld 1999; Šušković et al., 2001). This chapter includes selection criteria of bacteria as probiotics, technological usage of probiotics, new approaches for enhancing the performance of probiotics, and health effects of probiotic bacteria.
2. Selection of probiotic bacteria
Probiotics are living, health-promoting microorganisms that are incorporated into various kinds of foods. Although there has been a growing interest in using LAB isolated both from naturally fermented products and humans for health benefits (Lim & Im, 2009), the strains should preferably be of human origin and possess a Generally-Recognized-As-Safe status (Rönkä et al., 2003).
In order to exhibit their beneﬁcial effects, probiotic bacteria need to survive during the food-manufacturing process and in human ecosystem conditions; therefore it is important to investigate bacterial behavior under conditions which mimic the GIT (Zago et al., 2011; Lo Curto et al., 2011). Stresses to microorganisms begin in the mouth, with the lysozyme-containing saliva; continue in the stomach, which has a pH between 1.5 and 3.0; and go on to the upper intestine, which contains bile (Corzo & Gilliland, 1999). Acid and bile tolerances are two fundamental properties that indicate the ability of a probiotic microorganism to survive the passage through the GIT, resisting the acidic conditions in the stomach and the bile acids at the beginning of the small intestine (Prasad et al., 1998; Park et al., 2002). To evaluate the probiotic survival in the GIT, several
Effects of probiotics are strain specific. Strain identity is important in order to link a strain with a specific health effect, as well as to enable accurate surveillance and epidemiological studies (Ganguly et al., 2011). It is very important to be able to identify speciﬁcally and unambiguously the particular probiotic LAB strains from clinical fecal and intestinal biopsy specimens and from food samples (Tilsala-Timisjärvi & Tapanialtossava, 1998). Identiﬁcation of bacterial species and strains from commercialized probiotics has been conducted mostly using molecular methods (Holzapfel et al., 2001; Schillinger et al., 2003; Huys et al., 2006; Sheu et al., 2009).
Verdenelli et al. (2009) investigated the probiotic potential of 11
Başyiğit Kılıç & Karahan (2010) isolated one hundred seven strains of human originated LAB identified by 16S rRNA analysis and examined them for resistance to acidic pH, bile salts and antibiotic susceptibility. They found that
Lo Curto et al. (2011) investigated the survival of three commercial probiotic strains (
The safety of probiotic bacteria must be carefully assessed, with particular attention to transferable antibiotic resistance (Mathur & Singh, 2005). In the last decade, increasing concern has arisen about the safe use of LAB cultures for food and feed applications, in light of the latest knowledge about their possible role as an antibiotic-resistant gene reservoir. Particular concern is due to evidence of widespread occurrence in this bacterial group of conjugative plasmids and transposons (Clementi & Aquilanti, 2011). It is known that lactobacilli have a high natural resistance to bacitracin, cefoxitin, ciprofloxacin, fusidic acid, kanamycin, gentamicin, metronidazole, nitrofurantoin, norfloxacin, streptomycin, sulphadiazine, teicoplanin, trimethoprim/sulphamethoxazole, and vancomycin (Danielsen & Wind, 2003).
One of the primary beneﬁts associated with probiotic bacterial cultures is that they can exclude pathogenic bacteria from the small and large intestine (Kos et al., 2008). Another benefit is that in food products, antimicrobial activity of probiotic bacteria may contribute to an improvement in the quality of fermented foods. This may result from control of spoilage and pathogenic bacteria, extension of shelf life, and improvement of sensory quality (Wei et al., 2006; Siripatrawan & Harte, 2007). Kos et al. (2008) used overnight cultures and cell-free supernatants of the three probiotic strains
Production of antimicrobial compounds, which may take part in the inhibition of intestinal pathogens, is another criterion for classifying a potentially probiotic bacteria (Hutt et al., 2006). The inhibition of pathogenic microorganisms by selected probiotic strains may occur via a) production of antibiotic-like substances, b) bacteriocins and bacteriocin-like inhibitory substances such as acidophilin and reuterin, c) lowering of pH by producing organic acids such as acetic, lactic and phenyllactic acid, d) production of hydrogen peroxide and short chain fatty acids, e) decreasing the redox potential, and f) consumption of available nutrients (Holzapfel et al., 1995;Ouwehand, 1998; Tharmaraj & Shah, 2009).
The ability of LAB to adhere to epithelial cells and mucosal surfaces is thought to be an important property of many bacterial strains used as probiotics (FAO/WHO, 2001). Cell adhesion is a complex process involving contact between the bacterial cell membrane and interacting surfaces. Difficulties experienced in studying bacterial adhesion
The ability of probiotic bacteria to adhere to Caco-2 cells can be determined by plate counting or real time PCR (Matijasic et al., 2003; Candela et al., 2005). Nawaz et al. (2011) used both of these methods and did not find a statistically significant difference. Gaudana et al. (2010) investigated the ability of four different isolates (
3. Technological usage of probiotics
The use of starter cultures in the production of fermented food is necessary for guaranteeing safety and standardizing properties. LAB functions primarily to drop the pH of the batter; lower pH a) promotes product safety by inactivating pathogens, b) creates the biochemical conditions to attain the ﬁnal sensory properties through modiﬁcation of the raw materials, and c) improves the product stability and shelf life by inhibiting undesirable changes brought about by spoilage microorganisms or abiotic reactions (Ammor & Mayo, 2007).
Functional starter cultures are deﬁned as microbes that possess at least one inherently functional property aimed at improving the quality of the end product (De Vuyst, 2000). The use of probiotics in food has reinforced the acclaimed healthy properties and given rise to an increased consumption of these products in Europe and the USA (Kristo et al., 2003). Probiotics have been evaluated as functional starter cultures in various types of fermented food products such as yoghurt, cheese, dry sausage, salami, and sourdough. They have also been studied in therapeutic preparations to assess their positive effects on physico-chemical properties of foods and their impact on the nutritional quality and functional performance of the raw material (Knorr, 1998; Rodgers, 2008).
Fermented dairy products are widely-accepted, healthy food products and valued components of diets. The incorporation of probiotic bacteria as adjuncts in various fermented milk products is currently an important topic with industrial and commercial consequences. A number of dairy products containing probiotic bacteria are currently on the market. Fermented milk and cheeses have been described as the most suitable carriers, because they enhance the transit tolerance of bacteria (Saarela et al., 2000; Lourens-Hattingh & Viljoen, 2001). Some strains of
Although the number of cells required to produce therapeutic beneﬁts is not known and might vary as a function of the strain and the health effect desired, in general a minimum level of more than 106 viable probiotic bacteria per mililitre or gram of food product is accepted (Ouwehand & Salminen, 1998). The study of new probiotic strains for their technological relevance and use in food products is important for trade and industry. The search for strains which show resistance to biological barriers of the human GIT, and which possess physiological characteristics compatible with probiotic properties among LAB isolated from food, may eventually lead to the discovery of new probiotic strains for functional food products (Bude-Ugarte et al., 2006).
Studies of fermented food products as a source of new isolates are rapidly accumulating. For example, a mixture of human-derived probiotic strains was tested in the manufacture of ice cream; some of the ice cream was sweetened with sucrose and some was sweetened with aspartame (Başyiğit et al. 2006). The results showed that neither frozen conditions during the storage period nor the type of sweeteners used had any undesired effect on the survival of the probiotic cultures. Georgieva et al. (2009) studied technologically relevant properties of eight candidate probiotic
Essid et al. (2009) characterized 17 strains of
Floros et al. (2012) tested 19 facultatively heterofermentative lactobacilli from Feta, Kasseri, and Graviera cheeses for potential probiotic strains. Data from this study revealed that isolates B1, G16, G22, E22, E35, and H30 from Feta; PB2.2 from Kasseri; and 631 from Graviera have promising probiotic properties
Wang et al. (2010) identified and established the functional and technological characteristics of potential probiotic
3.1. Methods to increase survival and viability of probiotics
Researchers have long been encouraged to find new, efficient methods of improving the viability of probiotics in food products (especially fermented types), since viability can be affected by the acidic-bile conditions of the gastrointestinal tract (Mortazavian et al., 2007). The latest developments focus on fermentation technologies for producing probiotic bacteria; new approaches for enhancing the performance of these fastidious organisms during fermentation, downstream processing, and utilization in commercial products; and improving functionality in the gut. Processes to optimize survival and functionality in the gut include sublethal stress applications during cell production and new fermentation technologies, such as immobilized cell bioﬁlm-type fermentations, are promising in this respect (Lacroix & Yildirim, 2007).
3.1.1. Immobilized cell biofilm
Cell immobilization in fermentations is an attractive and rapidly expanding research area because of its technical and economic advantages, compared to a free cell system (Stewart & Russell, 1986). The immobilization method is cheap, simple and easy (Kourkoutas et al., 2006). The technology of cell immobilization allows an increase in cell stability and a decrease of the lethal effect on the microbial cells, providing protection from the conditions of the environment (Champagne et al., 1994; Grosso & Fávaro-Trindade, 2004). Thus immobilization techniques could provide protection to acid-sensitive LAB and increase their survival rate during the shelf life of the yoghurt and during their passage through the gastrointestinal tract (Cui et al., 2000; Fávaro-Trindade & Grosso, 2002). Kushal et al. (2006) determined that the process of co-immobilization of probiotic strains of
Encapsulation is the process of forming a continuous coating around an inner matrix that is wholly contained within the capsule wall as a core of encapsulated material (Kailasapathy, 2002). Encapsulation occurs naturally when bacterial cells grow and produce exo-polysaccharides. The microbial cells are entrapped within their own secretions that act as a protective structure or a capsule, reducing the permeability of material through the capsule, and making it less exposed to adverse environmental factors. Many LAB synthesise exo-polysaccharides, but they produce insufficient amounts to encapsulate themselves fully (Shah, 2002). Encapsulating probiotics in hydrocolloid beads has been investigated as a means of improving their viability and survival in food products and in the intestinal tract (Picot & Lacroix, 2004). Other benefits of encapsulation include reduction of cell injury, protection of probiotics from bacteriophages (Steenson et al., 1987), increased survival during freeze-drying and freezing (Kim & Yoon, 1995), and greater stability during storage (Kebary et al., 1998). Several methods of encapsulation have been used on probiotics in fermented milk products and biomass production: emulsion or two phase systems, the extrusion or droplet method, and spray drying and spray coating (Mortazavian et al., 2007). The common materials used for microencapsulation of probiotics are alginate and its derivatives, starch, mixtures of xanthan-gelan, carrageenan and its mixtures, gelatin, cellulose acetate phethalate, chitosan, and miscellaneous compounds such as whey proteins, soybean oil, gums, wax, and calcium chloride (Rao et al., 1989, Picot & Lacroix, 2004, Chandramouli et al., 2004).
Hou et al. (2003) demonstrated that encapsulation of
4. Effects of probiotics on human health
Probiotics have the potential for contributing greatly to human and animal health via a wide range of applications. Historically, probiotics have been used in food for humans and animals without any side effects, while providing for the balance of intestinal flora (Holzapfel & Wood, 1998). The health-promoting effects of probiotics have been widely explored and include stabilization of the indigenous microbial population, boosting of the immune system, inhibition of the growth of pathogenic organisms, prevention of diarrhea from various causes, alleviation of lactose intolerance, increased nutritional value of foods, reduction of serum cholesterol levels, antimutagenicity and anticarcinogenicity, reduction of the risk of inflammatory bowel conditions, improvement of digestion of proteins and fats, synthesis of vitamins, and detoxification and protection from toxins (Klaenhammer, 1998; Perdigon et al., 2002; Gaudana et al., 2010).
Anderson & Gilliland (1999) conducted two controlled clinical studies to test effects of yoghurt on heart-related health. They reported an average reduction of serum cholesterol by 2.9% with regular consumption of yoghurt containing
Isolauri et al. (1999) found significant improvement when a supplement of either
Can (2003) used an experimental animal model to study the effects of a probiotic mixture and
Ziarno et al. (2007) studied cholesterol assimilation by commercial starter cultures, reporting
Many probiotic species have been identified to be effective in children suffering from rotaviral diarrhea (Saavedra, 2000). Longdet et al. (2011) investigated the probiotic efficacy of
Senol et al. (2011a) investigated the protective effect of a probiotic mixture of 13 different bacteria and a-tocopherol on 98% ethanol-induced gastric mucosal injury. Levels of gastric mucosal pro-and anti-inﬂammatory cytokines, malondialdehyde, and secretory immunglobulin A were measured. Results showed that probiotic pretreatment signiﬁcantly suppressed the ethanol-induced increase of gastric mucosal interleukin-4 levels. Pretreatment with either probiotic or a–tocopherol inhibited the ethanol-induced increase of mucosal malondialdehyde concentration. Probiotic pretreatment enhanced the gastric mucosal secretory immunoglobulin A concentration. The researchers indicated that the probitic mixture and a-tocopherol reduced ethanol-induced gastric mucosal lipid peroxidation, suggesting that these probiotics may be beneﬁcial for helping heal gastric lesions induced by lower ethanol concentration. In another study, the role of a probiotic mixture, including 13 different bacteria, in the prevention of aspirin-induced gastric mucosal injury was investigated. Pretreatment with the probiotic mixture reduced aspirin-induced gastric damage and exerted a tendency toward downregulation of proinflammatory cytokines elicited by aspirin. Researchers also found that the probiotic mixture increased sIgA production approximately 7.5-fold in the stomach, and significantly reduced the malondialdehyde increase in the gastric mucosa elicited by aspirin. Additionally, pretreatment with the probiotic mixture alleviated aspirin-induced reduction of mast cell count in the gastric mucosa. Probiotic mixture pretreatment attenuates the aspirin-induced gastric lesions by reducing the lipid peroxidation, enhancing mucosal sIgA production, and stabilizing mucosal mast cell degranulation into the gastric mucosa (Senol et al., 2011b).
5. Final remarks
Significant data have been accumulated on probiotics and their beneficial health effects. Furthermore, more insights and key findings on the impact of processing and storage on probiotic viability and stability have been gained. A variety of microorganisms, typically food grade LAB, have been evaluated for their probiotic potential and are applied as adjunct cultures in various types of food products or in therapeutic preparations. In addition, further studies are needed to determine if preventive probiotic strategies are safe with regard to development of probiotic infections. Cooperation amongst food technologists, medical and nutrition scientists, and anticipation of future consumer demands are crucial for future success in probiotics.
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