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Effects of Dietary NEXT ENHANCE150.Pdf Accepted Manuscript Effects of dietary NEXT ENHANCE ®150 on growth performance and expression of immune and intestinal integrity related genes in gilthead sea bream (Sparus aurata L.) Jaume Pérez-Sánchez, Laura Benedito-Palos, Itziar Estensoro, Yiannis Petropoulos, Josep Alvar Calduch-Giner, Craig L. Browdy, Dr. Ariadna Sitjà-Bobadilla PII: S1050-4648(15)00057-1 DOI: 10.1016/j.fsi.2015.01.039 Reference: YFSIM 3318 To appear in: Fish and Shellfish Immunology Received Date: 31 October 2014 Revised Date: 30 January 2015 Accepted Date: 30 January 2015 Please cite this article as: Pérez-Sánchez J, Benedito-Palos L, Estensoro I, Petropoulos Y, Calduch- Giner JA, Browdy CL, Sitjà-Bobadilla A, Effects of dietary NEXT ENHANCE ®150 on growth performance and expression of immune and intestinal integrity related genes in gilthead sea bream (Sparus aurata L.), Fish and Shellfish Immunology (2015), doi: 10.1016/j.fsi.2015.01.039. This is a PDF file of an unedited manuscript that has been accepted for publication. As a service to our customers we are providing this early version of the manuscript. The manuscript will undergo copyediting, typesetting, and review of the resulting proof before it is published in its final form. Please note that during the production process errors may be discovered which could affect the content, and all legal disclaimers that apply to the journal pertain. ACCEPTED MANUSCRIPT 1 Effects of dietary NEXT ENHANCE ®150 on growth performance and expression of 2 immune and intestinal integrity related genes in gilthead sea bream ( Sparus aurata 3 L.). 4 5 Jaume Pérez-Sánchez 1, Laura Benedito-Palos 1, Itziar Estensoro 2, Yiannis Petropoulos 3, 6 Josep Alvar Calduch-Giner 1, Craig L. Browdy 4, Ariadna Sitjà-Bobadilla 2 7 8 1Nutrigenomics and Fish Growth Endocrinology Group, Instituto de Acuicultura Torre de 9 la Sal (IATS-CSIC), Castellón, Spain 10 2 Fish Pathology Group, Instituto de Acuicultura Torre de la Sal (IATS-CSIC), Castellón, 11 Spain 12 3Andromeda S.A., Rion Achaias, Greece 13 4Novus International, Inc., St. Charles, MO, USAMANUSCRIPT 14 15 16 Corresponding author: Dr. Ariadna Sitjà Bobadilla 17 Instituto de Acuicultura Torre de la Sal 18 Consejo Superior de Investigaciones Científicas 19 12595 Ribera de Cabanes, Castellón, Spain 20 e-mail: [email protected] 21 ACCEPTED phone: +34-964319500 ext. 38 22 fax: +34-964319509 23 24 1 ACCEPTED MANUSCRIPT 25 ABSTRACT 26 27 Gilthead sea bream juveniles were fed different doses (0, 50, 100, 200, 300 ppm) of NEXT 28 ENHANCE ®150 (NE) for 9 weeks. Feed gain ratio (FGR) was improved by a 10% with 29 all the doses, but feed intake decreased in a dose dependent manner. The optimum inclusion 30 level to achieve maximum growth was set at 100 ppm. The hepatosomatic index did not 31 vary and only at the highest dose, viscerosomatic and splenosomatic indexes were 32 significantly decreased. No significant changes were found in haematological parameters, 33 plasma biochemistry, total antioxidant capacity and respiratory burst. In a second trial, NE 34 was given at 100 ppm alone (D1) or in combination with the prebiotic PREVIDA® (0.5%) 35 (PRE) (D2) for 17 weeks. There were no differences in the growth rates, and FGR was 36 equally improved for D1 and D2. No significant changes in haematology and plasma 37 antioxidant capacity were detected. The histologicaMANUSCRIPTl examination of the liver and the 38 intestine showed no outstanding differences in the liver, but the number of mucosal 39 foldings appeared to be higher in D1 and D2 vs CTRL diet and the density of enterocytes 40 and goblet cells also appeared higher, particularly in the anterior intestine. A 87-gene PCR- 41 array was constructed based on our transcriptomic database (www.nutrigroup- 42 iats.org/seabreamdb) and applied to samples of anterior (AI) and posterior (PI) intestine. It 43 included 54 new gene sequences and other sequences as markers of cell differentiation and 44 proliferation, intestinal architecture and permeability, enterocyte mass and epithelial 45 damage, interleukinsACCEPTED and cytokines, pattern recognition receptors (PRR), and mitochondrial 46 function and biogenesis. More than half of the studied genes had significantly different 47 expression between AI and PI segments. The functional significance of this differential 48 tissue expression is discussed. The experimental diets induced significant changes in the 2 ACCEPTED MANUSCRIPT 49 expression of 26 genes. The intensity of these changes and the number of genes that were 50 significantly regulated were higher at PI than at AI. At PI, both diets invoked a clear down- 51 regulation of genes involved in cell differentiation and proliferation, some involved in cell 52 to cell communication, cytokines and several PRR. By contrast, up-regulation was mostly 53 found for genes related to enterocyte mass, cell epithelial damage and mitochondrial 54 activity at AI. The changes were of the same order for D1 and D2, except for fatty acid- 55 binding proteins 2 and 6 and the PRR fucolectin, which were higher in D2 and D1 fed fish, 56 respectively. Thus, NE alone or in combination with PRE seems to induce an anti- 57 inflammatory and anti-proliferative transcriptomic profile with probable improvement in 58 the absorptive capacity of the intestine that would explain the improved FGR. 59 60 Key words: Teleostei, prebiotics, carvacrol, thymol, aquafeeds, intestine, transcriptomics, 61 immunology MANUSCRIPT ACCEPTED 3 ACCEPTED MANUSCRIPT 62 1. Introduction 63 64 The role of dietary nutrients and additives on immune system function in fish has 65 been investigated for several decades as a means of reducing the presence of opportunistic 66 pathogens and simultaneously stimulating the host immunological responses [1, 2]. Among 67 the different nutraceutical products, prebiotics, probiotics and antioxidants are most 68 studied. Prebiotics are non-digestible selectively fermented feed ingredients that allow 69 specific changes in the composition and/or activity in the gut microbiota that confers 70 benefits upon host wealth and health, which mediate immunological development and 71 functionality, particularly at the mucosal interface within the gastrointestinal (GI) tract [3- 72 5]. The use of dietary prebiotics in farmed fish has been approached, sometimes with 73 contradictory effects [6-10]. 74 Phytoadditives obtained from different herbsMANUSCRIPT have also been tested in fish feeds for 75 effects on fish immune response and disease susceptibility [11-15]. They represent a 76 promising alternative to traditional drugs as they have low costs and some of them are 77 legally registered as food additives for human consumption [16]. Phytogenic extracts 78 containing phenolic and flavonoid chemical compounds are known to exert an array of 79 positive effects that include improvement of gut microbiota composition, enhanced immune 80 function and resistance to pathogens, and improved gut barrier structure and function in 81 humans and animals [17, 18]. Among these additives, carvacrol and thymol are essential 82 oils from oreganoACCEPTED ( Origanum vulgare ) that have been shown to improve growth 83 performance in different animal production systems [19, 20]. They are also known for their 84 antibacterial [21] and antioxidant [22] properties, for synergizing effects of antibiotics [23], 85 and for their potential applications in foods [24], including fish fillets [25, 26]. Carvacrol in 4 ACCEPTED MANUSCRIPT 86 particular exhibits a range of biological activities: antimicrobial, antitumoral, 87 antimutagenic, antigenotoxic, analgesic, antispasmodic, antiinflammatory, angiogenic, 88 antiparasitic, antiplatelet, AChe inhibitory, antielastase, insecticidal, antihepatotoxic and 89 hepatoprotective (reviewed by Baser [27]). However, its role as an immunostimulant is 90 rather controversial, with contradictory results in swine [28-30]. Carvacrol administration 91 in fish species is starting to be documented. Thus far, it has been tested alone in tilapia [31] 92 and European sea bass [32], or in combination with thymol in channel catfish [33] and 93 rainbow trout [34, 35]. In these previous experiments, these phytoadditives had a beneficial 94 effect on growth and disease resistance to bacterial challenges. However, the pathways 95 involved in their immunostimulant and/or antioxidant roles in fish are still poorly 96 understand. 97 Thus, in the present study we tested the effect of Next Enhance NE®150 (NE), an 98 encapsulated combination of carvacrol and thymol,MANUSCRIPT as a feed additive for gilthead sea 99 bream, alone or in combination with the prebiotic Previda® (PRE). In a first short-term 100 trial, the best dose of NE was established in terms of growth performance. Subsequently, a 101 longer trial was set up to assess the effect on gut and liver histomorphology, antioxidant 102 status, and the expression of immune and intestinal integrity related genes, using an 87- 103 gene PCR-array derived from the updated transcriptomic database of gilthead sea bream 104 [36] hosted at www.nutrigroup-iats.org/seabreamdb. The development of the array involved 105 the molecular definition of 54 new gilthead sea bream sequences, selected together with 106 other known sequencesACCEPTED as markers of intestine function, immunity and integrity, based on 107 the intestine gene expression profiling in fish challenged with different diets and the 108 parasite Enteromyxum leei [37-40]. 109 5 ACCEPTED MANUSCRIPT 110 2. Materials and Methods 111 112 2.1. Animal care, feeding trials design and sampling procedure 113 Clinically healthy juvenile gilthead sea bream (GSB) were obtained from a local 114 commercial hatchery and acclimatized to the IATS rearing facilities for four weeks. A first 115 feeding trial (trial 1) was set up to establish the best NE dose. Fish with an initial weight of 116 26-27 g (1 st May) were randomly distributed in triplicated 90L tanks (20 animals/tank) and 117 fed a basal diet (CTRL, see below) or the experimental diets with four inclusion levels of 118 the active compound NE: 50 ppm (D50), 100 ppm (D100), 200 ppm (D200) and 300 ppm 119 (D300).
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