ReviewThe use of probiotics in aquaculture
Introduction
Long before their discovery, microbes have been unawarely used to preserve food, and these empirical methods contributed to improve human health (Bengmark, 1998, Metchnikoff, 1908). Early in the century, Metchnikoff (1907), Metchnikoff (1908)(cited by Tannock, 1997) proposed to implant lactic acid bacteria into the human intestine, with a view to suppressing the detrimental activity of other microbes. The modern concept of probiotics was formulated only 25 years ago (Parker, 1974), then its pertinence was challenged for many years among the scientific community. Several definitions of probiotics were successively proposed. Parker (1974)originally referred to “organisms and substances which contribute to intestinal microbial balance” as probiotics. The definition was then restricted to “a live microbial feed supplement which beneficially affects the host animal by improving its intestinal microbial balance” (Fuller, 1989). Tannock (1997)noted that the effect on the “intestinal microbial balance” has not been demonstrated in most cases, and he proposed to speak of “living microbial cells administered as dietary supplements with the aim of improving health”.
To my knowledge, the first application of probiotics in aquaculture seems relatively recent (Kozasa, 1986), but the interest in such environment-friendly treatments is increasing rapidly. Scientific evaluation corroborated seldom the first empirical trials, and the information was mainly spread by “grey literature”. However, a growing number of scientific papers have dealt explicitly with probiotics, and it is now possible to survey the state of the art, from the empirical use to the scientific approach. A previous review was devoted to lactic acid bacteria in finfish (Ringø and Gatesoupe, 1998), but many other microbes have been tested as probiotics for various aquatic animals, appealing for a general overview.
It is essential to remind some definitions of ecological concepts to clear away the treatments termed improperly “probiotic”. The goals of this paper are (1) to examine the pertinence of such terminology applied to the aquatic environment, (2) to draw the different trends of applications, and (3) to point out needs for further research.
Section snippets
Is the intestinal environment of aquatic animals favourable to probiotics?
Aquatic animals are quite different from the land animals for which the probiotic concept was developed, and a preliminary question is the pertinence of probiotic applications to aquaculture.
Man and terrestrial livestock undergo embryonic development within an amnion, whereas the larval forms of most fish and shellfish are released in the external environment at an early ontogenetic stage. These larvae are highly exposed to gastrointestinal microbiota-associated disorders, because they start
Applications of the “probiotic” concept in a broad sense
The transience of aquatic microbes may legitimate the extension of the probiotic concept to living microbial preparations used to treat aquaculture ponds. Moriarty (1998)proposed to extend the definition of probiotics to microbial “water additives”. However, this extension would make too vague definition of Tannock (1997). I suggest an alternative definition of probiotics as: microbial cells that are administered in such a way as to enter the gastrointestinal tract and to be kept alive, with
Application of commercial products for terrestrial livestock
The first trials of incorporation of probiotics into aquaculture feeds used commercial preparations designed for land animals. Spores of Bacillus toyoi isolated from soil reduced the mortality of Japanese eel which were infected by Edwardsiella sp., (Kozasa, 1986). The same feed additive increased the growth rate of yellowtail (Kozasa, 1986). Spores may be easily incorporated into compound food, but their fate in the gastrointestinal tract of fish was not followed in these experiments. It would
Isolation and characterization of autochthonous microbes
In juvenile fish and shellfish, the autochthonous microbes may be isolated from the digestive tract after dissection, while distinguishing stomach and intestine regions. The microbes adherent to epithelial cells can be separated from those adherent to mucus, and from those transient in the lumen (Westerdahl et al., 1991). These methods are not applicable to larvae and live food organisms, but the external surface of larval fish may be washed with a 0.1% benzalkonium chloride saline solution to
Perspectives of development
The advantage of probiotics over antibiotics was discussed by Moriarty (1998), but most attention has been hitherto directed towards the production of inhibitory substances by the probiotics. The risk to select probiotic-resistant pathogens must not be underestimated, and it is particularly important to search for diversified antagonistic properties, which may lower the risk of multi-resistance. For example, the ability of some probiotics to adhere to intestinal mucus may block the intestinal
Conclusion
The application of probiotics in aquaculture shows promise, but needs considerable efforts of research. The first question, unanswered in many cases, is the fate of the probiotic in rearing medium and in gastrointestinal tract. Immunological and molecular probes will be useful tools to trace the probiotic cells (Ringø et al., 1996; Austin, 1998; O'Sullivan, 1999). It is essential to investigate the best way of introduction and the optimal dose, and technical solutions are required, especially
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