Indian Journal of Biotechnology Vol 2, July 2003, pp 378-381 Effect of Lactic Acid Bacteria on Animal Performance Zwi G Weinberg' Forage Preservation and By-Products Research Unit, Department of Food Science, the Volcani Center, Bet Dagan, 50250, Israel Received 1 January 2003; accepted 20 February 2003 Lactic acid bacteria(LAB) which are included in inoculants for silage, sometimes impart beneficial effects, similar to when fed directly, on cattle performance. LABs might interact with rumen microorganisms in such a way that their activity is enhanced and fibre degradability is improved. Another possibility is that LABs produce bacteriocins in the silage which might inhibit detrimental microorganisms, both in the silage and in the rumen. Key words: lactic acid bacteria, ruminants, animal performance, probiotic effects Introduction Ensiling, a method of preserving moist forage, is widely used in North America, Europe, Israel and elsewhere. It is based on natural fermentation, where lactic acid bacteria (LAB) ferment water-soluble carbohydrates to organic acids, mainly lactic (LA), under anaerobic conditions. As a result, the pH decreases, inhibiting detrimental anaerobes and preserving the nutritional value and palatability of the moist forage. Inoculants including LABs are often used as silage additives to enhance LA fermentation and, hence, better to preserve the ensiled crops. Most commercially available inoculants contain homofermentative LABs which are fast and efficient producers of LA and thus improve the silage fermentation. Among the homofermentative LABs, most frequently used are Lactobacillus plantarum, Enterococcus faecium, Pediococcus acidilactici, P. pentosaceus and L. acidophilus. Heterofermentative LABs are also sometimes included in inoculants for silage, because they produce volatile fatty acids (VFAs), which inhibit the yeasts and moulds, which are activated upon aerobic exposure of the silage. The recommended application rates (105_106 viable cells/g) of these products are usually, sufficient to enable the inoculant LAB population to overtake those of the epiphytic LABs and become the predominant population in the silage. Many studies involving silage inoculants have been carried out (McDonald et al, 1991; Weinberg & Muck, 1996). *Tel: 972-968-3549, 968-3558; Fax: 972-3-960-4428 E-mail: [email protected] Effects of LAB Inoculant-treated Silage on Animal Performance The studies indicated that feeding cattle with silages treated with certain LABs improved ruminant performance. Certain LABs are believed to induce probiotic effects on humans. Improvements in animal performance are in many cases the principal economic justification for inoculant use. In several trials (Spoelstra, 1991; Muck, 1993), inoculants exhibited substantial effect on performance as follows: average increases in intake, 4-11; live weight gain, 7-11; milk production, 3-5; and feed efficiency, 9%. Inoculants have improved animal performance in part, which might be related to the inoculation rate. In alfalfa silages (Satter et al, 1987), an inoculation factor >10 (a more than ten-fold increase in LABs over the natural population) was necessary for increased milk production, a factor five times higher than that needed to affect fermentation consistently (Pahlow, 1991). A considerable number of animal experiments with low dry matter (DM) grass silage inoculated with. L plantarum MTD1, performed in Northern Ireland, showed improvements in both silage fermentation and animal performance. However, some studies showed improvement in animal performance with no apparent effect on fermentation (in some cases both the control and treated silages were of good quality and in some cases clostridial fermentation occurred in both) and vice versa suggesting that effect may be probiotic. Rooke & Kafilzadeh (1994) compared MTD 1 with two other LAB strains. All three strains improved fermentation similarly, but only MTDI resulted in a significant increase in DM intake in weathers, which WEINBERG: LACTIC ACID BACTERIA AND ANIMAL PERFORMANCE indicates that this phenomenon might be strain specific. In the rumen liquor of calves fed on MT01inoculated grass silage (Keady & Steen, 1994a,b, 1995), inoculation increased nitrogen degradability, which might be related to the liberation of slowly degradable nitrogen because of modification of the cellwall structure. The inoculant treatment increased the total ruminal VF A concentration and tended to decrease the butyrate and to increase the propionate contents. Similar trends in low OM grass silage were also observed (Sharp et ai, 1994) with a different inoculant, which contained strains of L. piantarum and E. faecalis applied at 105 CFUs per gram. Studies with other LAB inoculants had mixed results (Mayne, 1993; Smith et al, 1993). The effects on animal performance of LAB inoculants in silages other than grass are variable. Stokes (1992) found reduced OM intake by cattle fed on inoculated grass-legume silages and related this to the poor aerobic stability of such silages. Kung et al (1993), who used two different inoculants in corn silage, observed tendencies to higher fat-corrected milk yields and higher OM intake only with MTOl, which agrees with the results obtained in low OM grass silage. Sanderson (1993), who used corn silage inoculated with L. piantarum and E. Faecium, did not obtain any enhancement of fibre degradability. Salawu et al (2001) observed increased rates of in situ degradation of nitrogen and fibre in bi-crop pea/wheat silage inoculated with L. piantarum and L. buchneri. Effect of Direct-fed LABs on Animal Performance Several trials used direct-fed LABs in ruminant feeding. Wallace & Newbold (1993) used lactobacilli or enterococci as probiotics in calves and lambs. Feeding with live bacteria resulted in decreased counts of coliforms and improvements in intake and liveweight gain. Keady & Steen (1996) applied L. piantarum MTO 1 to well preserved grass silage at 109 colony-forming units (CFUs) per gram and immediately fed that silage to male cattle, no significant difference was observed in OM, nitrogen and fibre degradability, rumen pH, ammonia or VFA concentrations or feed intake. Malik & Sharma (1998) observed improved in vitro dry matter and organic matter digestibility and higher microbial protein nitrogen following addition of L. acidophilus to rumen liquor taken by male cattle which were fed wheat straw and concentrate (60:40) 379 rations. Addition of an LAB culture containing L. acidophiius, L. casei and L. jugarti to grain free diet of growing male crossbred cattle also had beneficial effects on weight gain and feed intake, and resulted in lower incidence of diarrhoea (Khuntia & Chaudhary, 2002). Ghorbani et al (2002), who fed Propionibacterium and Enterococcus faecium directly to steers, observed that the directly fed microbes might have decreased the risk of acidosis in feedlot cattle. Possible Mechanisms of the Probiotic Effect The cause of improved animal performance is unclear. The shifts in fermentation products and small improvements in silage OM recovery would enhance animal performance but not to the degree observed. Muck (1993) reported a high degree of correlation between effects on OM digestibility and on animal performance, and also noted that fibre digestibility was improved in 30% of the trials. This is certainly unexpected, in that the LABs used in inoculants are not known to have any direct activity on polysaccharides. LABs could interact with rumen microorganisms in such a way that their activity is enhanced and fibre degradability is improved. Weinberg et al (2003) observed higher pH in rumen fluid, which was inoculated with the LABs used in inoculants for silage. This trend might be associated in changes in VF A concentration in the rumen and might explain the observation that directly fed LAB might alleviate acidosis (Ghorbani et al, 2002). The sheep, immunized with live Streptococcus bovis vaccine, had higher rumen pH, higher feed intake and alleviated symptoms of acidosis (Gill et ai, 2000). LABs, that produce bacteriocins in the silage, might inhibit detrimental microorganisms, both in the silage and in the rumen. It is well known that certain LAB strains produce antimicrobial substances that inhibit other LABs or other microorganisms (Muller et al, 1996; Odenyo, 1994a; Vandenbergh, 1993). Conclusions and Research Needs Silage inoculant LABs has a probiotic effect on ruminant performance. However, the mechanism of this probiotic effect is not as yet clear. LABs might change the rumen environment and interact with rumen microorganisms in such a way as to enhance feed utilization. Therefore, helpful information might be obtained through measurements of fibre degradability and nitrogen metabolism or through studies of the interactions of LABs with selected rumen micro or- 380 INDIAN J BIOTECHNOL, ganisms by means of 16S rRNA probes (Odenyo et al, 1994a, b; Chen & Weimer, 2001). Evaluation of the bacteriocins production by the LAB strains used in inoculants for silage for bacteriocin production might be another useful research direction. A relevant question concerns whether the probiotic effect is already active in the inoculated silage or whether it is triggered in the rumen? The rumen system is complex and is affected by many factors including the feed (Weimer & Waghorn, 1999), the season, the number of days in milk, and animal variations. The probiotic effects endowed by the LABs might be affected by such factors, and research should address this point as well. JULY 2003 Mayne C S, 1993. The effect of formic acid, sulphuric acid and a bacterial inoculant on silage fermentation and the food intake and milk production of lactating dairy cows. Anim Prod, 56, 29-42. McDonald P et al, 1991. The Biochemistry of Silage, 2nd edn. ChaJcombe Publications, Aberystwyth, UK. Pp 184-236. Muck R E, 1993. The role of silage additives in making high quality silage. in Silage Production from Seed to Animal. Northeast Regional Agric. Engng. Service, NRAES-67, Syracuse, NY, February 23-25. Pp 106-116. Muller T et al, 1996. Antagonistic activity in plant-associated lactic acid bacteria. Microbiol Res, 151, 63-70. Odenyo A A et al, 1994a. The use of 16S rRNA- targeted oligonucleotide probes to study competition between ruminal fibrolytic bacteria: development of probes for Ruminococcus species and evidence for bacteriocin production. 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