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- How tryptophan levels in plant-based aquafeeds affect fish physiology, metabolism and proteomePublication . Cerqueira, Marco; Schrama, Denise; Silva, Tome S.; Colen, R.; Engrola, Sofia; Conceicao, Luis E. C.; Rodrigues, Pedro; Farinha, Ana PaulaFish meal replacement by plant-protein sources is a priority in aquaculture feeds. Within this framework, dietary supplementation with essential amino acids (EAA), as tryptophan (TRP), is strategic to ensure that the individual nutritional needs are met, besides promoting enhanced immunological status. The purpose of this study was to examine the beneficial effects of TRP incorporation in plant-protein source diets on fish growth performance and nutritional status. We tested diets with 20% lower (LTRP) and 27% higher (HTRP) of the putative requirements of TRP for seabream (Sparus aurata) and assessed its impact on fish physiology and liver metabolism and proteome. After 12 weeks, growth performance, body proximate, hepatic composition and liver metabolic profiling were similar between diets. Nevertheless, liver proteome analysis indicated a higher accumulation of proteins involved in acute-phase responses, typically triggered by infection, inflammation or trauma, in fish fed with HTRP diet as compared with those fed with LTRP. The overall results obtained suggest a potential beneficial effect of TRP supplementation in terms of immune stimulation, without compromising growth or feed intake. Moreover, proteomics and metabolic profiling demonstrate to be valuable tools in this endeavour. Significance: Nutritional needs are hard to assess in aquaculture fisheries, and many times controversial depending on the methodology employed. The estimated amino acid requirements depend on both fish species and stage development, making it extremely hard to standardise. On the other hand, the substitution of fish-based to plant-based protein sources diets towards a sustainable aquaculture, may imbalance these requirements, being necessary further studies to assess the impact on fish growth and development. Finally, the incorporation of crystalized amino acids such as TRP into diets aims global better performance both at fish health/immune condition and growth development. This work focused on the potential beneficial effects of TRP supplementation into diets with a plant-based protein source, addressing the effects on the liver metabolism and proteome, and on growth performance of Gilthead seabream juveniles, a species with special relevance and economical importance in the Mediterranean region. The present study by employing proteomics together with metabolic profiling shows that TRP supplementation at the tested doses, does not compromise growth performance, and seems to stimulate the immune system. Our findings can contribute to the development of new feed formulations for Gilthead seabream species, therefore, reinforcing the resilience and competitiveness of the on-growing aquaculture industry and impact directly the sustainability of living resources with the decrease of the fisheries needs to fulfil the human search for quality proteins consume.
- The role of dietary methionine concentrations on growth, metabolism and N-retention in cobia (Rachycentron canadum) at elevated water temperaturesPublication . Nguyen, Minh Van; Espe, Marit; Conceição, Louis E. C.; Le, Hoang Minh; Yúfera, Manuel; Engrola, Sofia; Jordal, A. E. O.; Rønnestad, IvarThis study determined impacts of dietary methionine concentrations at two temperatures on growth, feeding efficiency and N-metabolites in juvenile cobia. Methionine concentrations of the experimental diets were deficient (M9; 9 g/kg), sufficient (M12; 12 g/kg) and surplus (M16, 16 g/kg). Water temperature was normal (30°C) or elevated (34°C). Twenty cobia in triplicate tanks were fed the experimental diets for 6 weeks. Both methionine and temperature affected cobia's growth and feeding efficiency. Cobia fed M9 performed lower than the fish fed M12 and M16 diets. Additionally, cobia reared at 34°C performed poorer than at 30°C, probably due to lower voluntary feed intake in the fish reared at 34°C. Protein efficiency ratio and protein productive value in cobia fed M9 diet were less than M12 or M16 diets. This was confirmed with the improved retentions of indispensable amino acids (AAs). No interactions between methionine and temperature were observed in growth and protein accretion. At 30°C, CF improved, while HSI and VSI declined upon methionine supplementation levels. Of which an interaction between temperature and methionine was present. Plasma, muscle and liver free AA and N-metabolites were affected by methionine and temperature. Furthermore, temperature affected cobia's lipid class composition, resulting in increased phospholipids and cholesterol at 34°C.
- Transition from endogenous to exogenous feeding in hatchery-cultured European eel larvaePublication . Benini, Elisa; Engrola, Sofia; Politis, Sebastian Nikitas; Sørensen, Sune Riis; Nielsen, Anders; Conceição, Luis E. C.; Santos, André; Tomkiewicz, JonnaThe transition from endogenous to exogenous feeding is critical during fish early life, where appropriate feed availability and timing of initiation of feeding influence survival. For European eel (Anguilla anguilla), establishing first feeding culture is at a pioneering state, where successful production of larvae has recently enabled feeding experiments. In the present study, three diets and potential benefits of early feeding during the transition from yolk-sac stage to feeding larvae were explored, including molecular analyses of genes involved in digestive functions and growth. Three consecutive trials were performed using hatchery produced eel offspring. In Feeding regimes 1 and 3, expression of npy and cck (appetite regulation) was higher, while expression of pomca (food intake) was lower in non-prefed larvae, indicating increased fasting and higher starvation risk. In contrast, Feeding regime 2 led to the highest survival ever registered for European eel larvae i.e. 20% at 20 dph, in spite that prefeeding resulted in reduced survival rate during the endogenous feeding stage. This was associated with initial hsp90 (stress/repair) upregulation in larvae receiving prefeeding, however, with subsequent downregulation during exogenous feeding. Notably, the growth related gh expression was higher in prefed larvae, indicating growth benefits of prefeeding. Likewise, prefeeding resulted in pomca as well as try, tgl, and amyl2a (digestion) upregulation, providing evidence of beneficial maturation of gut functionalities. Essentially, Feeding regime 2 demonstrated a continuous upregulation of growth, appetite and digestion related genes, which in combination with the highest survival suggest that dietary requirements were partially met. Moreover, in Feeding regime 2, gh and tgl were expressed at a higher level in prefeeding larvae than in the control, indicating that prefeeding might be advantageous in spite observed mortality, but further research is needed, including timing of feed application.
- Alternative proteins for fish diets: implications beyond growthPublication . Aragão, Cláudia; Gonçalves, Ana Teresa; Costas, Benjamín; Azeredo, Rita; Xavier, Maria João; Engrola, SofiaAquaculture has been challenged to find alternative ingredients to develop innovative feed formulations that foster a sustainable future growth. Given the most recent trends in fish feed formulation on the use of alternative protein sources to decrease the dependency of fishmeal, it is fundamental to evaluate the implications of this new paradigm for fish health and welfare. This work intends to comprehensively review the impacts of alternative and novel dietary protein sources on fish gut microbiota and health, stress and immune responses, disease resistance, and antioxidant capacity. The research results indicate that alternative protein sources, such as terrestrial plant proteins, rendered animal by-products, insect meals, micro- and macroalgae, and single cell proteins (e.g., yeasts), may negatively impact gut microbiota and health, thus affecting immune and stress responses. Nevertheless, some of the novel protein sources, such as insects and algae meals, have functional properties and may exert an immunostimulatory activity. Further research on the effects of novel protein sources, beyond growth, is clearly needed. The information gathered here is of utmost importance, in order to develop innovative diets that guarantee the production of healthy fish with high quality standards and optimised welfare conditions, thus contributing to a sustainable growth of the aquaculture industry.
- Dietary tryptophan supplementation induces a transient immune enhancement of gilthead seabream (Sparus aurata) juveniles fed fishmeal-free dietsPublication . Ramos-Pinto, Lourenco; Martos-Sitcha, Juan A.; Reis, Bruno; Azeredo, Rita; Fernandez-Boo, Sergio; Perez-Sanchez, Jaume; Alvar Calduch-Giner, Josep; Engrola, Sofia; Conceicao, Luis E. C.; Dias, Jorge; Silva, Tome S.; Costas, BenjaminEuropean aquaculture is an industry with a high sustainability profile contributing to the supply of safe seafood. However, several diseases can affect farmed fish and it is imperative to find alternatives for chemotherapeutic treatments when disease outbreaks occur. Maintenance of health through nutrition is well-establish in modern animal farming, and amino acids (AA) are promising candidates as functional additives to improve fish health. Therefore, the goal of this research is to provide a better understanding of the influence of tryptophan supplementation on nutritional condition and immune mechanisms in fish. Triplicate groups of fish (13.3 +/- 0.3g) previously fed with a fishmeal-based diet were either fed a control diet with an extreme formulation (0% fishmeal) but meeting the AA requirements (CTRL), or the SUP diet, formulated as the CTRL with an increase in tryptophan (TRP) content. After 2 and 13 weeks of feeding, head-kidney (HK), liver (L) and white skeletal muscle (WSM) were collected for gene expression, whereas plasma was suited for humoral immune parameters. A holistic approach using transcriptomic, humoral and zootechnical parameters was undertaken. The expression of 29-31 genes for WSM, L or HK confirms an effect due to the treatment across time. A two-way ANOVA analysis revealed that 15-24 genes varied significantly depending on the tissue, and the multivariate analysis by means of PLS-DA explained (R-2) and predicted (Q(2)) with four components up to 93% and 78% of total variance, respectively. Component 1 (R-2 = 50.06%) represented the time effects, whereas components 2 (24.36%) and 3 (13.89%) grouped fish on the basis of dietary treatment, at early sampling. The HK results in particular suggest that fish fed SUP diet displayed an immunostimulated state at 2 weeks. No major differences were observed in plasma humoral parameters, despite an increase in antiprotease and peroxidase activities after 13 weeks regardless of dietary treatment. These results suggest that tryptophan supplementation may improve the seabream immune status after 2 weeks. Hence, the use of functional feeds is especially relevant during a short-term feeding period before a predictable stressful event or disease outbreak, considering that these putative advantageous effects seem to disappear after a 13 weeks feeding period.
- Optimizing diets to decrease environmental impact of Nile tilapia (Oreochromis niloticus) productionPublication . Teodósio, Rita; Engrola, Sofia; Colen, R.; Masagounder, Karthik; Aragão, CláudiaAquaculture is one of the most thriving animal production sectors, and Nile tilapia (Oreochromis niloticus) farming represents 8% of total finfish culture. However, the industry sustainability depends on the development of cost‐effective and environmental friendly feeds. This study aimed to reduce dietary protein levels in diets for juvenile tilapia and to minimize diet environmental impact while maximizing biological efficiency. A growth trial was performed using five isoenergetic plant protein‐based diets with decreasing levels of crude protein: 360, 340, 320, 300 and 280 g/kg diet (D360, D340, D320, D300 and D280, respectively). Dietary protein utilization was assessed by metabolic trials using a radiolabelled amino acid mixture. Tilapia in all treatments showed similar growth performance and feed intake. Feed conversion ratio was significantly higher in fish fed the D280 than the D360 diet, while no differences were found for other treatments. Protein retention was significantly higher in tilapia fed the D300 than the D360 diet. Amino acid catabolism increased in fish fed the D360 diet, though without significant differences in muscle amino acid retention. This study demonstrates that dietary protein levels can be reduced to 300 g/kg diet without hindering tilapia growth and feed conversion ratio, while reducing environmental nitrogen losses.
- The effect of tank cover on welfare of farmed Nile tilapiaPublication . Saraiva, João L.; Nogueirinha, Margarida; Teodósio, Rita; Aragão, Cláudia; Engrola, Sofia; Arechavala Lopez, PabloWelfare in aquaculture is a pressing topic. One of the main measures to improve the life of farmed fish is implementing environmental enrichment, which can include the addition of structural covers to rearing tanks. Here we test the effects of tank covers (fully covered, 50 % covered and uncovered) in an all-male population (N = 75 tagged individuals: 5 individuals per tank, 5 tanks per treatment) of farmed Nile tilapia using behavioural and physiological welfare indicators. Video recordings were performed over 21 days to evaluate overall swimming (swim bursts, freezing on the bottom or hovering in the water column) and aggression events (bites and chases). At the end of the experiment, blood was collected to assess circulating cortisol. Results showed a clearly lower welfare in the partially covered tanks regarding behavioural and physiological variables. The present study highlights the need to correctly study 1) the ethology of the farmed species and 2) the enrichment strategy to be implemented, before any measure is taken. While environmental enrichment is undoubtedly a much needed area of research and a promising solution to improve the welfare of farmed fish, a proper ethological assessment must be made a priori to avoid detrimental effects of poorly devised approaches.
- Amino acid metabolism in gilthead seabream is affected by the dietary protein to energy ratiosPublication . Teodósio, Rita; Aragão, Cláudia; Conceicao, Luis; Dias, Jorge; Engrola, SofiaThe dietary protein to energy ratio (P/E) has proven to influence protein utilization and/or growth in several fish species. This study intended to unravel the bioavailability and metabolic fate of lysine and methionine in gilthead seabream (Sparus aurata) juveniles fed plant diets with different P/E ratios. Seabream juveniles were fed two isonitrogenous diets (45% crude protein) differing in crude lipids (20 and 14%): LowP/E (P/E ratio=20.0 mg protein kJ-1) and HighP/E (P/E ratio=21.4 mg protein kJ-1). After three weeks, fish (11.6 +/- 4.3 g) were tube-fed the respective diet labelled with C-14-protein (L-amino acid mixture), C-14-lysine, or C-14-methionine. Protein, lysine, and methionine utilization were determined based on the proportion of C-14-amino acid evacuated, retained in the free or protein-bound fraction of liver and muscle, or catabolized. This study revealed that a decrease in P/E ratio resulted in lower amino acid evacuation (p < 0.05), contributing to a more efficient amino acid uptake. Results indicate that amino acids are retained as protein in the liver and not only temporarily available in the free pool. The amount of free amino acids retained in the muscle of LowP/E fed fish was significantly higher than in HighP/E fish (p < 0.05) due to a simultaneous higher retention of lysine and methionine, without affecting the overall protein retention. Methionine catabolism was significantly lower than lysine or protein independently of the P/E ratio (p < 0.05), reinforcing that this amino acid is preferentially spared for metabolic functions and not used as energy source. In contrast, increasing the dietary P/E ratio decreased lysine catabolism and increased its availability for growth. The bioavailability and metabolism of individual amino acids should be considered when optimizing P/E ratios in diets for gilthead seabream juveniles. Formulating diets with optimum P/E ratios will improve diet utilization and fish performance.
- Improving growth potential in Senegalese sole (Solea senegalensis) through dietary proteinPublication . Canada, Paula; Engrola, Sofia; Conceicao, Luis E. C.; Valente, Luisa M. P.In the present work, recent studies that contributed for improving dietary protein in practical microdiets for Senegalese sole larvae are revised, in an attempt to overcome some of the current problems in this species larviculture, such as the difficult early adaptation to inert diets and highly variable growth rates. Different diet formulation strategies were employed in an attempt to improve Senegalese sole larvae capacity to utilize and deposit protein throughout metamorphosis, to further maximize growth potential: (1) increasing indispensable amino acids (IAA) content (2) meeting the ideal IAA profile by adjusting the dietary AA profile to the larval body AA profile and (3) decreasing the complexity (molecular weight) of dietary protein to increase its digestibility. Either manipulating the quality or the complexity of dietary protein had impact on the larvae capacity to utilize protein and direct it for growth. Sole larvae seem to be able to adapt their digestive functions and metabolic capacity to dietary protein. Moreover, increasing the dietary IAA/DAA ratio and changing the degree of hydrolysis of dietary protein affected the expression pattern of muscle growth related genes, with consequences on muscle cellularity and potential for growth. The expression of DNA methyltransferases was altered in response to changes in dietary protein. The novelty of such information in fish may trigger further studies on the effect of dietary protein on the epigenetic regulation of growth. In conclusion, optimal protein quality for Senegalese sole seems to change during larval development. Whereas the inclusion of a moderately hydrolysed protein comes up as a promising way to improve growth in early larval stages, larger peptides and intact protein seem to be more suitable to sole post-larvae and young juveniles. Therefore, these results suggest that dietary protein fraction formulation of microdiets for Senegalese sole should be adapted to each developmental stage, what has important consequences for practical larval microdiets formulation and feeding protocols.
- Metabolic and nutritional responses of Nile tilapia juveniles to dietary methionine sourcesPublication . Teodósio, Rita; Engrola, Sofia; Cabano, Miguel; Colen, R.; Masagounder, Karthik; Raquel Cêa de Aragão Teixeira, CláudiaCommercial diets for tilapia juveniles contain high levels of plant protein sources. Soybean meal has been utilised due to its high protein content; however, soy-based diets are limited in methionine (Met) and require its supplementation to fulfil fish requirements. DL-Methinone (DL-Met) and Ca bis-methionine hydroxyl analogue (MHA-Ca) are synthetic Met sources supplemented in aquafeeds, which may differ in biological efficiency due to structural differences. The present study evaluated the effect of both methionine sources on metabolism and growth of Nile tilapia. A growth trial was performed using three isonitrogenous and isoenergetic diets, containing plant ingredients as protein sources: DLM and MHA diets were supplemented on equimolar levels of Met, while REF diet was not supplemented. Hepatic free Met and one-carbon metabolites were determined in fish fed for 57 d. Metabolism of DL-Met and MHA was analysed by an in vivo time-course trial using 14C-labelled tracers. Only DLMet supplementation significantly increased final body weight and improved feed conversion and protein efficiency ratios compared with the REF diet. Our findings indicate that Met in DLM fed fish follows the transsulphuration pathway, while in fish fed MHA and REF diets it is reme thylated. The in vivo trial revealed that 14C-DL-Met is absorbed faster and more retained than 14C-MHA, resulting in a greater availability of free Met in the tissues when fish is fed with DLM diet. Our study indicates that dietary DL-Met supplementation improves growth performance and N retention, and that Met absorption and utilisation are influenced by the dietary source in tilapia juveniles.