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BackgroundUse of plant ingredients in aquaculture feeds is impeded by high contents of antinutritional factors such as saponins, which may cause various pharmacological and biological effects. In this study, transcriptome changes were analyzed using a 21 k oligonucleotide microarray and qPCR in the distal intestine of Atlantic salmon fed diets based on five plant protein sources combined with soybean saponins.ResultsDiets with corn gluten, sunflower, rapeseed or horsebean produced minor effects while the combination of saponins with pea protein concentrate caused enteritis and major transcriptome changes. Acute inflammation was characterised by up-regulation of cytokines, NFkB and TNFalpha related genes and regulators of T-cell function, while the IFN-axis was suppressed. Induction of lectins, complement, metalloproteinases and the respiratory burst complex parallelled a down-regulation of genes for free radical scavengers and iron binding proteins. Marked down-regulation of xenobiotic metabolism was also observed, possibly increasing vulnerability of the intestinal tissue. A hallmark of metabolic changes was dramatic down-regulation of lipid, bile and steroid metabolism. Impairment of digestion was further suggested by expression changes of nutrient transporters and regulators of water balance (e.g. aquaporin, guanylin). On the other hand, microarray profiling revealed activation of multiple mucosal defence processes. Annexin-1, with important anti-inflammatory and gastroprotective properties, was markedly up-regulated. Furthermore, augmented synthesis of polyamines needed for cellular proliferation (up-regulation of arginase and ornithine decarboxylase) and increased mucus production (down-regulation of glycan turnover and goblet cell hyperplasia) could participate in mucosal healing and restoration of normal tissue function.ConclusionThe current study promoted understanding of salmon intestinal pathology and establishment of a model for feed induced enteritis. Multiple gene expression profiling further characterised the inflammation and described the intestinal pathology at the molecular level.Ethical approvalThe present experiment was approved by the Norwegian Animal Research Authority and conducted according to prevailing animal welfare regulations: FOR-1996-01-15-23 (Norway), European Convention for the Protection of Vertebrate Animals used for Experimental and Other Scientific Purposes (Strasbourg, 18.III.1986) and COUNCIL DIRECTIVE of 24 November 1986 on the approximation of laws, regulations and administrative provisions of the Member States regarding the protection of animals used for experimental and other scientific purposes (86/609/EEC).

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The effects of combining soyasaponins with plant ingredients on intestinal function and fish health were investigated in an 80 d study with Atlantic salmon (270 g) distributed thirty each into twenty-four tanks with seawater. Soyasaponins were supplemented (2 g/kg) to diets with maize gluten (MG), pea protein concentrate (PPC) and sunflower (SFM), rapeseed (RSM) or horsebean meals. A diet with soyabean meal (SBM) and another with wheat gluten and soyasaponins served as reference diets. Marked soyasaponin effects were observed when combined with PPC. This combination induced inflammation in the distal intestine (DI) similar to SBM, reduced feed intake, apparent digestibility of lipid, most amino acids and ash, decreased bile salt levels in intestinal chyme and decreased leucine aminopeptidase (LAP) activity but increased trypsin activity in the DI. No enteritis was observed in other diet groups, but small consistent negative soyasaponin effects were seen on lipid and fatty acid digestibility, faecal DM and LAP activity of the DI. Soyasaponin combination with RSM reduced digestibility of all nutrients including minerals. The mineral effect was also seen for SFM, whereas with MG and SFM a positive soyasaponin effect on feed intake was observed. Caution should be exercised to avoid ingredient combinations giving high saponin levels, a condition that appears to be a key factor in diet-induced enteritis together with certain plant ingredients.Key words: Soyasaponins: Plant protein ingredients: Antinutritional factors: Fish feed: Gastrointestinal tract Alternative dietary protein sources to supplement and replace limited marine ingredients in fish feeds are important for the future of the fish farming industry. Plant ingredients such as soyabeans hold promise with their good amino acid profiles that can easily be improved for fish requirements by supplementation with deficient amino acids. However, soyabean meal (SBM) inclusion has been demonstrated to induce enteritis and reduce performance in salmonids (1 -3) and carp (4) . The factors responsible for the disorders have not been conclusively identified, but soyasaponins, and possibly other bioactive antinutritional factors (ANF) in SBM, are implicated in the aetiology (5 -7) .Saponins are heat-stable glycosides present in soyabean and other legumes such as pea and lupin (8,9) . Saponins, with their membrane-active nature and affinity to cholesterol and bile salts (10,11) , possess a number of potential biological effects compatible with the negative effects observed in fish fed diets containing SBM. There have been conflicting findings from studies on dietary effects of saponins to teleost fish. In one study, a saponin-rich extract from SBM and Quillaja saponins, both at a 0·3 % dietary inclusion rate of saponin, greatly reduced feed intake and growth in Chinook salmon and depressed growth in rainbow trout (12) . Furthermore, the Quillaja saponin diets (0·15 and 0·3 % dietary inclusion) both induced substantial damage to the intestinal mucosa for both Chino...

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Rainbow trout with initial body weight 1144 g were fed two diets with high (feed A) or low (feed B) water stability for 6 weeks. During the last 2 weeks either stable or fluctuating oxygen saturation, salinity and temperature was introduced. High water stability of the feed was associated with harder pellets, less dust formation and less broken pellets compared to feed with low water stability. During the first 4 weeks, feed intake was 23% higher in trout fed diet B than in those fed diet A. Fluctuating environment resulted in a severe drop in feed intake for both dietary groups. At termination of the trial, stomachs of trout fed feed B contained mashed pellets and free water and oil. Stomachs of trout fed feed A contained more intact pellets and little fluid. The apparent digestibility of protein, starch, dry matter and energy was highest in feed A. The apparent digestibility of P and Zn was highest at stable environment, and the absorption of P was highest from feed A. In conclusion, physical quality affected the nutritional value of the feeds, and differences in nutritional value at stable or fluctuating environment appeared to be related to feed intake.

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