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Probiotics: Production, Evaluation and Uses in Animal Feed, 2009: ISBN: 978-81-308-0323-4 Editors: Nelson Prez Guerra and Lorenzo Pastrana Castro 4 The use of probiotic in fish and shrimp aquaculture. A review
Maurilio Lara-Flores 1 and Gabriel Aguirre-Guzmn 2 1 Centro de Ecologa, Pesqueras y Oceanografa del Golfo de Mxico de la Universidad Autnoma de Campeche, Av. Agustn Melgar y Juan de la Barrera S/N, C. P. 240230, Campeche, Campeche Mxico; 2 Facultad de Medicina Veterinaria y Zootecnia de la Universidad Autnoma de Tamaulipas, Km. 5 Carr. Cd. Victoria a Mante, C.P. 87000, Cd. Victoria, Tamaulipas, Mxico E-mail: gabaguirre@uat.edu.mx, gaguirre_guzman@hotmail.com
Abstract In last decade, the total world fishery production decreased slightly and the human consumption for aquatic product increased. The reduction in capture fisheries was partly compensated for the fast growth of aquaculture industry. The need for enhanced disease resistance, feed efficiency, and growth performance of cultured organisms is substantial for various sectors of this industry. If growth performance
Correspondence/Reprint request: Dr. Maurilio Lara-Flores, Centro de Ecologa, Pesqueras y Oceanografa del Golfo de Mxico de la Universidad Autnoma de Campeche, Av. Agustn Melgar y Juan de la Barrera S/N C. P. 240230 Campeche, Campeche Mxico. E-mail: maurlara@uacam.mx, maurilio_lara@yahoo.com.mx Maurilio Lara-Flores & Gabriel Aguirre-Guzmn 2 and feed efficiency are increased in commercial aquaculture, the costs productions are likely to be reduced. Also if more aquatic organisms are able to resist diseases and survive the subsequent cost of medication and overall production costs would be reduced. Hormones, antibiotics, ionophers and some salts compounds have been used at some extent to prevent disease and as growth promoters; however, their inadequate application can produce adverse disorders, such as hormone imbalance, poisoning and predisposition to disease development. In the search of new options, several studies have been carried out to test new compounds, from which the aquaculture industry has developed the concept of functional additives. Among these additives, the additions of microorganisms to the diets, named probiotics, has shown to improve the energy expenditure derived from other sources such as carbohydrates and increase the incorporations of protein for growth; increase the immunity and disease resistance of the host organism. The use of probiotics in aquaculture activity is in the beginnings, since that the gastrointestinal microbiota of aquatic organisms has been poorly characterized; and their effects not be study extensive. This review summarizes and evaluates current knowledge of the use and the action of the probiotic in fish and shrimp culture; and the potential for further application of this in aquaculture production.
1. Introduction The human requirement for aquatics products for consumption and aquarist activity is higher and this is not satisfied for the world fishery production, those open an extensive range of opportunities to aquaculture industry for composes this requirement [1]. Aquaculture has become an important economic activity in many countries [2]. However, this aquaculture develop show many problems as widespread epizootics, feed efficiency and growth performance [3-5]. This is principal caused by the large-scale production facilities, where aquatic animals are exposed to stressful conditions, problems related to disease, inadequate balance of nutrient in the artificial diets an deterioration of environmental conditions, since the physiological stress is one of the primary contributing factors of aquatic organisms disease, poor growth and mortality in aquaculture [6-8]. In the present, the principal aims of the aquaculture industry are increase the growth or survival performance, feed efficiency, and resistance of aquatic organisms which show a positive effect on production costs [9]. Hormones, antibiotics, ionophers and some salts compounds have been used at some extent to prevent disease and as growth promoters; however, their inadequate application show a negative effects on aquaculture production and environment [10]. Probiotic in aquaculture 3 Functional additive, as probiotics, is a new concept on aquaculture [11]; where the additions of microorganisms on diets show a positive effect on growth caused by the best use of carbohydrates, protein, and energy [12-16], diminishing mortality by disease, antagonism to pathogen, better microbial intestinal balance and in the environment [3,12,17]. The concept of probiotics was originally used by Lilley and Stillwell [18] to mean a substance (s) that stimulates growth of other microorganisms [19]. Parker in 1974 modified the definition to organisms and substances which contribute to intestinal balance. Fuller [20] revised the definitions as A live microbial feed supplement which beneficially affects the host animal by improving its intestinal microbial balance. This revised definition has put forward the importance of live cells as the essential component of a potential probiotic and it clears the confusion created by the use of term substances. However, an effect on intestinal microbial balance has been defined and demonstrated only in a few cases. This was noted by Tannock [21], and he proposed the definition living microbial cells administered as dietary supplements with the aim of improving health. According to the accepted definition about probiotic this product is a live microorganism food supplement which improves the microbial balance of the host intestinal flora [22-24]. Other definitions in aquaculture show that probiotic is a live microbial food supplements that are consumed with the aim of providing health benefit to the host by contributing to an improved microbial balance within the intestinal microbiota [25,26], are biologically active components or single or mixed cultures of microorganisms capable of improving the health of the host [27,28], live microorganisms and/or disease resistance [29], live microorganisms administered in adequate amounts that confer a health effect on the host [30]. The components of these definitions reflect the use microorganism or their products (microbial cells element or cell free supernatant factors) to tanks and ponds in which animals live, as biological control or for their capacity of modified the bacterial composition of aquatic animals intestine, water and sediment, or used with feed as health supplement and/or biological control.
2. Criteria of probiotic selection in aquaculture The initial, major, purpose of using probiotics is to maintain or reestablish a favorable relationship between friendly and pathogenic microorganisms that constitute the flora of intestinal o skin mucus of aquatic animals. Since, successful probiotic is expected to have a few specific properties in order to certify a beneficial effect [31]. Maurilio Lara-Flores & Gabriel Aguirre-Guzmn 4 Generally, probiotic strains have been isolated from indigenous and exogenous microbiota of aquatic animals. Gram-negative facultative anaerobic bacteria such as Vibrio and Pseudomonas constitute the predominant indigenous microbiota of a variety of species of marine animals [32]. In contrast to saltwater organisms, the indigenous microbiota of freshwater animals tends to be dominated by members of the genera Aeromonas, Plesiomonas, representatives of the family Enterobacteriaceae, and obligate anaerobic bacteria of the genera Bacteroides, Fusubacterium, and Eubacterium [33]. Lactic acid producing bacteria, which are prevalent in the mammal or bird gut, are generally sub-dominant in fishes and represented essentially by the genus Carnobacterium [34]. Ideally, microbial probiotics should have a beneficial effect and not cause any harm to the host. Therefore, all strains have to be non-pathogenic and non-toxic in order to avoid undesirable side-effects when administrated to aquatic animals. Some research and products talk about the multifactorial action of the probiotics [30,35] on aquatic animals. However, the multifactorial effect is not agreed with evidence or is overestimate. Sometimes, this type of publicity about the potential of those products really affects the perspective of real probiotic designed for aquaculture industry. Different modes of action or properties are desire on the potential probiotic like antagonism to pathogens [34,36], ability of cells to produce metabolities (like vitamins) and enzymes [31], colonization or adhesion properties [37], enhance the immune systems [38] and others.
Competitive exclusion Competitive exclusion as it applies to the gastrointestinal tract is a phenomenon whereby an established microflora prevents or reduces the colonization of a competing bacterial challenge for the same location in the intestine. This microflora begins to form in the gut of aquatic animals during the hatching process and shortly thereafter from bacteria in the environment. The aim of probiotic products designed under competitive exclusion is obtain a stable, agree and controlled microbiota on culture based on competition for attachment sites on the mucosa, competition for nutrients, and production of inhibitory substance by the microflora which prevents replication and/or destroys the challenging bacteria and with this reduce its colonization [12,39]. Different strategies are displayed in the adhesion of microorganism to those attachment sites as passive forces, electrostatic interactions, hydrophobic, steric forces, lipoteichoic acids, adhesins and specific structures of adhesion [40]. Probiotic in aquaculture 5 The aquaculture industry display some probiotics products designed to adhesion on mucosal surfaces by a collection of microorganisms based on the competitive exclusion factors [39,41]. Those factors are important for adhesion to intestinal epithelial cells or in the activation of immune system, and help to the health of the organisms, intestinal homeostasis, and digestion [41,42]. These types of probiotic are extensively study in fish since this products were initialed development for vertebrates animals, but when are used in shrimp some problems were observed. For example, Beseres et al. [43] show that Litopenaeus vannamei, L. stiferus, Farfantepenaeus aztecus display a gut residence time of consumed feed from 45 to 90 min, which represent a short time to bacteria adhesion or colonization. Ziaei-Nejad et al. [24] show a low colonization rates of Bacillus sp. on nauplius, zoea, mysis, postlarvae (1-14) of Fenneropenaus indicus, and any significantly different in total bacterial counts between treatments and controls. The positive effects of this type of probiotics are designed for shrimp culture condition and using the competitive exclusion properties against microflora of aquatic organisms, these products show interesting results.
Antagonisms Control of microbial communities with high diversity has been regarded as difficult [44]. Such types of microbial communities can disperse the effect caused by the invasion or addition of certain extrinsic pathogenic organisms. Bacterial antagonism is a common phenomenon in nature; therefore, microbial interactions play a major role in the equilibrium between competing beneficial and potentially pathogenic microorganisms [6]. In addition, microorganisms can be sources of a variety of bioactive natural products of basic research and commercial interest that have inhibitory effects on microbial growth [45]. The antagonism is used in different probiotic products, this is focus to negative effect to pathogens of aquatic organisms (bacteria principally) with their positive effect on the health of host [30,45-47]. Vibrio species are part of the autochthonous flora of marine organisms and caused mortalities in shrimp larvae and juvenile [48-50]. Arthrobacter sp. (strain XE-7), Bacillus sp. (strains 11, P64, BT23), B. pumilus, B. sphaericus, B. subtilis, Lactobacillus bulgaricus (strains NCIM 2056, NCIM 2057), L. acidophilus (strain NCIM 2285), Nitrosomonas sp., Nitrobacter sp., Pseudomonas sp. (strain PS-102), Saccharomyces cerevisiae, S. exigus, Phaffia rodozoma, Streptococcus cremoris (strain NCIM 2285), Streptomyces sp., Tetraselmis suecica (strain CS-187), Vibrio sp. (strain P62, P63), V. alginolyticus (strain Ili) had been used as probiotics for their antagonism effect to Vibrio species as V. harveyi and V. parahaemolyticus (Table 1) [51-61]. Maurilio Lara-Flores & Gabriel Aguirre-Guzmn 6
T a b l e
1 .
P r o b i o t i c
u s e d
i n
a q u a c u l t u r e
i n d u s t r y
o f
s h r i m p .
Probiotic in aquaculture 7
T a b l e
1 .
C o n t i n u e d
Maurilio Lara-Flores & Gabriel Aguirre-Guzmn 8 In the other hand, in fish the probiotics are more extensively used for prevent and/or combat bacteria disease for example the vertebral column compression syndrome caused by Flavobacterium psychrophilum where the use of lactic acid bacteria [62,63] or yeast [64-67] caused a decrease of this bacteria. Vibrio sp. and Aeromonas sp. have become the most pathogenic microorganisms in fish aquaculture for control this pathogens pseudomonads have receive special attention as disease-protecting microorganisms and have been used as plant biocontrol [68]. In recent years, there has been great interest in the use of lactic acid bacteria as disinfection treatment [69] and for control the populations of the native microflora such as Aeromonas and Vibrio species [70]. In vitro antagonism test is based on the nature phenomenon of antimicrobial metabolites production of some bacteria strains and is a frequent way for screening probiotics. For example, antagonism of B. subtilis (strain BT23) against V. harveyi did confer protection to Penaeus monodon [56]. Antigenic components of diverse species of Pseudomonas demonstrated different levels of antagonism against Aeromonas hydrophila [45]. Cell free extracts of L. acidophilus, S. cremoris, L. bulgaricus (strains 56, 57) show negative effect on V. alginolyticus growth in agar plate test [57]. The origin of probiotic strain is an important element in the antagonisms test. The microorganisms present different physiologies or biochemical activities along their growth develop and based on their environments (fresh, seawater) and original source. These characteristics affect the probiotic potential for attachment sites [79] and may create a false impression of the ability of probiotics to inhibit pathogens in vivo test. The probiotics screening preferably requires different strategy of selection as antagonism, production of beneficial compounds, attachment and growth on various environments [80].
Immunity stimulation This is relations with the potential characteristic of some probiotic to stimulate the immune system of host against pathogen agents [81]. Some studies tried to explain the different actions mechanisms of probiotics stimulation of immune system in fish, (immune cell, antibodies, acid phosphatase, lysozyme, antimicrobial peptides, etc). Panigrahi et al. [82,83] demonstrated the increase of immune parameters as lysozyme action, phagocyte activity with the use of L. rhamnosus (strain JCM 1136) or their cell wall components on O. mykiss. Song et al., [84] observed an increase in acid phosphatase activity in Miichtys miiuy feeding with C. butyricum indicated a stimulate immune response. Probiotic in aquaculture 9 In the other hand, the shrimp not display an acquired immune system as in vertebrates but some studies demonstrated an immune response. Table 1 show that B. pumilus, B. sphaericus, B. subtilis, V. fluvialis (strain PM 17), Pseudomonas sp. (strain PM 11), S. cerevisiae, S. exigus, P. rodozoma are used as probiotic with immune response on P. monodon and L. vannamei [71,76,77]. Rengpipat et al. [71] showed that Bacillus sp. (strain S11) can be provided disease protection for activating of P. monodon immune defenses. Balcazar et al. [6] show that administration of a mixture of Bacillus sp. and Vibrio sp. promotes the resistance of juveniles of L. vannamei against V. harveyi. Vibrio cells, lipopolysaccharide (LPS), peptidoglycan (PG), 1-3 glucan, fucoidan, laminaria, yeast glucans have been experimentally tested in small-scale culture and their results suggest that can be used as an important element in the control of disease for their effect on shrimp or crustacean immunostimulation [85,86].
Adhesion Probiotics make up part of the resident microflora and contribute to the health or well-being of their host [87]. The ability of some strain of adhesion to mucus, gastrointestinal tract, epithelial cell and other tissues is a common characteristic used in the probiotic selection because it is associated with bacteria colonization [26,39,41]. The principal objective of adhesion design is obtaining a high and significant level of bacteria in the host and prevents them from being flushed out by the movement of food through the digestive tract. By attaching to the intestinal mucosa, probiotics can extend their time within the gut thereby influencing the gastrointestinal microflora of their host [53,76,88,89]. The attachment ability of some bacteria have been tested in vitro and in vivo and their results suggest that the pathogen was displaced by the potential probiotic, this is based on the ability of probiotics to attach to the mucus, where the growth of the pathogen in the digestive tract might be suppressed by the candidate probiotic presence [39,41,42,80]. This characteristic is higher associated with the competition for essential nutrients, space, etc. [39]. Different strains of acid lactic bacteria, like Enterococcus faecium and Lactobacillus sp.; and other groups of bacteria Gram-positive and Gram- negative as Bacillus sp., Vibrio sp., have been tested and posteriori used as probiotic for the ability of adhesion [15,16,53,57,80].
Digestive process Many studies on probiotics in aquaculture have used in vitro models of specific bacteria as antagonists of pathogens [80,90], measured the survival Maurilio Lara-Flores & Gabriel Aguirre-Guzmn 10 of probiotics in the fish gut [88,91] or evaluated the beneficial effect of probiotic on health management, disease resistance and immune response of fish [11,92]. But other important effect of the use of probiotic that its not extensively study but demonstrated an important effect is the feed efficiency and the growth promotion of aquatic animals by probiotic supplements [69,93]. The probiotic after transit through the stomach, they attach in the intestine and use a large number of carbohydrates for their growth and produce a range of relevant digestive enzymes (amylase, protease and lipase) that increase the digestibility of organic matter and protein, produce a higher growth, prevent the intestinal disorders and produce or/and stimulate a pre- digestion of secondary compounds present principal in plant protein sources [7,93]. In finfish the use of probiotics demonstrated beneficial effects on the growth performance, feed efficiency and digestibility of organic matter and protein, when used acid lactic bacteria and yeast [93-98]. In some case this beneficial effect had attribute to the capacity of the probiotic to stimulate and/or produce some enzymes on the intestinal tract. For example, Lara et al [99] observed a high activity of alkaline phosphatase in Nile tilapia (Oreochromis niloticus) when administered probiotics in the diet, the result show a high activity reflected a possible development of brush border membranes of enterocytes that can be stimulated by the probiotic and this it can be a indicator of carbohydrate and lipid absorption and explain the higher weight gain and the best feed conversion. In shrimp, some studies on the beneficial effects of probiotics in the nutritional and digestive process have been reported [24,41,72], where Bacillus sp. and spores of B. subtilis, B. licheniformis, B. polymyxa, B. laterosporus, B. circulans have been used in these works. Ziaei-Nead et al. [24] examined the effects of Bacillus sp. on F. indicus at different shrimp stages (Nauplius, Zoea, Mysis and postlarvae). In pond treated with probiotic show a significantly higher activity of amylase, total protease, and lipase. The increase of Bacillus sp. in L. vannamei diet shows a significantly higher apparent digestibility of some essential nutrients as phosphorus [72]. Some research about the application of probiotic strains in shrimp; display and apparent increase of some enzymes, but the short gut-residence time of feed on shrimp (45 to 90 min)[43] difficult the evaluation of this parameter.
3. Author opinion Initially, the efficient of probiotics was related with the strain multiplications and/or their presence on environment after application, and Probiotic in aquaculture 11 this attribute was associated with strain colonization on host and some benefic effect on health. Those are not agree with all probiotics products and help to obtain contradictories results about their effect on aquatic organisms. The evolution of probiotic is associated with the better understanding of the intestinal ecology application of this type of products, properties, and the specific strain-host relations. Actually, the specific use of probiotic is researched and associated with the effect on specific host, strain source and their active doses, effect on the activation immune system by RT-PCR studies, specific mechanism of adhesion to host and environment with use of gfp (green fluorescent protein) plasmid, symbiosis, and particularly environment condition of host and their effect in the probiotic strain. The direct use of probiotic on water (from fresh to seawater of farms and laboratories) is a special point of environment research consideration. Those products (probiotics) are commonly foreign or exogenous strain, and represent a possible risk of microorganism pollution, especially with the use of strain with genetic modification, specific adhesions or colonization niche, antibiotic production, synergistic action. The use and environment effect of those new probiotics generation its necessary to agree understand before massive application on aquaculture. However, a number of probiotic products have been thoroughly researched, and agree evidenced their efficacy a possible use on aquaculture. Beneficial bacterial preparations that are species-specific probiotics have become more widely available to the aquaculture community. These preparation show specific benefic effect as disease prevention and offer a natural element to obtain a stable healthy gut environment and immune system. The establishing of strong disease prevention program, which includes probiotic and good management practices can be hope to raise aquatic organisms production.
4. Acknowledgments This work was supported by grants from the Mexican Council of Science and Technology or CONACYT [Project FOMIX-CAMPECHE (CAMP- 2004-C02-1 and CAM-2005-C01-041)].
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