1.Connor, WE (2000) Importance of n-3 fatty acids in health and disease. Am J Clin Nutr 71, 171S–175S.
2.Simopoulos, AP (2008) The importance of the omega-6/omega-3 fatty acid ratio in cardiovascular disease and other chronic diseases. Exp Biol Med 233, 674–688.
3.Mozaffarian, D & Wu, JHY (2012) (n-3) Fatty acids and cardiovascular health: are effects of EPA and DHA shared or complementary? J Nutr 142, 614S–625S.
4.U.S. Department of Health and Human Services (2015) 2015–2020 Dietary Guidelines for Americans. Washington, DC: USDA.
5.FAO (2016) The State of World Fisheries and Aquaculture 2016. Contributing to Food Security and Nutrition for All. Rome, Italy: FAO.
6.Sargent, J, Tocher, D & Bell, J (2002) The lipids. In Fish Nutrition, pp. 181–257 [Halver, JE and RW Hardy, editors]. San Diego, CA: Academic Press.
7.Castro, LF, Tocher, DR & Monroig, O (2016) Long-chain polyunsaturated fatty acid biosynthesis in chordates: insights into the evolution of Fads and Elovl gene repertoire. Prog Lipid Res 62, 25–40.
8.Leaver, MJ, Villeneuve, LA, Obach, A, et al. (2008) Functional genomics reveals increases in cholesterol biosynthetic genes and highly unsaturated fatty acid biosynthesis after dietary substitution of fish oil with vegetable oils in Atlantic salmon (Salmo salar). BMC Genom 9, 299.
9.Sargent, J, Bell, G, McEvoy, L, et al. (1999) Recent developments in the essential fatty acid nutrition of fish. Aquaculture 177, 191–199.
10.Tocher, DR (2010) Fatty acid requirements in ontogeny of marine and freshwater fish. Aquac Res 41, 717–732.
11.Monroig, Ó, Tocher, DR & Navarro, JC (2013) Biosynthesis of polyunsaturated fatty acids in marine invertebrates: recent advances in molecular mechanisms. Mar Drugs 11, 3998–4018.
12.Monroig, Ó, Wang, S, Zhang, L, et al. (2012) Elongation of long-chain fatty acids in rabbitfish Siganus canaliculatus: cloning, functional characterisation and tissue distribution of Elovl5- and Elovl4-like elongases. Aquaculture 350–353, 63–70.
13.Navarro-Guillén, C, Engrola, S, Castanheira, F, et al. (2014) Effect of varying dietary levels of LC-PUFA and vegetable oil sources on performance and fatty acids of Senegalese sole post larvae: puzzling results suggest complete biosynthesis pathway from C18 PUFA to DHA. Comp Biochem Phys B: Biochem Mol Biol 167, 51–58.
14.Xu, H, Dong, X, Ai, Q, et al. (2014) Regulation of tissue LC-PUFA contents, Δ6 fatty acyl desaturase (FADS2) gene expression and the methylation of the putative FADS2 gene promoter by different dietary fatty acid profiles in Japanese seabass (Lateolabrax japonicus). PLOS ONE 9, e87726.
15.Tocher, DR & Ghioni, C (1999) Fatty acid metabolism in marine fish: low activity of fatty acyl Δ5 desaturation in gilthead sea bream (Sparus aurata) cells. Lipids 34, 433–440.
16.Seiliez, I, Panserat, S, Corraze, G, et al. (2003) Cloning and nutritional regulation of a Δ6-desaturase-like enzyme in the marine teleost gilthead seabream (Sparus aurata). Comp Biochem Physiol Part B: Biochem Mol Biol 135, 449–460.
17.Izquierdo, MS, Robaina, L, Juárez-Carrillo, E, et al. (2008) Regulation of growth, fatty acid composition and Δ6 desaturase expression by dietary lipids in gilthead seabream larvae (Sparus aurata). Fish Physiol Biochem 34, 117–127.
18.Izquierdo, MS, Turkmen, S, Montero, D, et al. (2015) Nutritional programming through broodstock diets to improve utilization of very low fishmeal and fish oil diets in gilthead sea bream. Aquaculture 449, 18–26.
19.Turkmen, S, Castro, PL, Caballero, MJ, et al. (2017) Nutritional stimuli of gilthead seabream (Sparus aurata) larvae by dietary fatty acids: effects on larval performance, gene expression and neurogenesis. Aquac Res 48, 202–213.
20.Turkmen, S, Zamorano, MJ, Fernández-Palacios, H, et al. (2017) Parental nutritional programming and a reminder during juvenile stage affect growth, lipid metabolism and utilisation in later developmental stages of a marine teleost, the gilthead sea bream (Sparus aurata). Br J Nutr 118, 500–512.
21.Ainge, H, Thompson, C, Ozanne, SE, et al. (2010) A systematic review on animal models of maternal high fat feeding and offspring glycaemic control. Int J Obes 35, 325–335.
22.Burdge, GC & Lillycrop, KA (2010) Nutrition, epigenetics, and developmental plasticity: implications for understanding human disease. Ann Rev Nutr 30, 315–339.
23.Lillycrop, KA & Burdge, GC (2018) Chapter 13 – Interactions between polyunsaturated fatty acids and the epigenome. In Polyunsaturated Fatty Acid Metabolism, pp. 225–239. Urbana, IL: AOCS Press.
24.Kelsall, CJ, Hoile, SP, Irvine, NA, et al. (2012) Vascular dysfunction induced in offspring by maternal dietary fat involves altered arterial polyunsaturated fatty acid biosynthesis. PLOS ONE 7, e34492.
25.Hoile, SP, Irvine, NA, Kelsall, CJ, et al. (2013) Maternal fat intake in rats alters 20 : 4n-6 and 22 : 6n-3 status and the epigenetic regulation of Fads2 in offspring liver. J Nutr Biochem 24, 1213–1220.
26.Calder, PC (2006) n-3 Polyunsaturated fatty acids, inflammation, and inflammatory diseases. Am J Clin Nutr 83, 6 Suppl., 1505S–1519S.
27.Yaqoob, P & Calder, PC (2007) Fatty acids and immune function: new insights into mechanisms. Br J Nutr 98, S41–S45.
28.Montero, D & Izquierdo, M (2010) Welfare and health of fish fed vegetable oils as alternative lipid sources to fish oil. In Fish Oil Replacement and Alternative Lipid Sources in Aquaculture Feeds, pp. 439–485 [Turchini, GM, Ng, WK and Tocher, DR, editors]. Boca Raton, FL: CRC Press.
29.Calder, PC (1996) Immunomodulatory and anti-inflammatory effects of n-3 polyunsaturated fatty acids. Proc Nutr Soc 55, 737–774.
30.Calder, PC (2002) Dietary modification of inflammation with lipids. Proc Nutr Soc 61, 345–358.
31.Yaqoob, P (2004) Fatty acids and the immune system: from basic science to clinical applications. Proc Nutr Soc 63, 89–104.
32.Secombes, CJ, Hardie, LJ & Daniels, G (1996) Cytokines in fish: an update. Fish Shellfish Immun 6, 291–304.
33.Calder, PC (2005) Polyunsaturated fatty acids and inflammation. Biochem Soc Trans 33, 423–427.
34.Izquierdo, MS & Koven, W (2011) Lipids. In Larval Fish Nutrition, pp. 47–81 [Holt, JG, editor]. Hoboken, NJ: Wiley-Blackwell.
35.Serhan, CN (2006) Novel chemical mediators in the resolution of inflammation: resolvins and protectins. Anesthesiol Clin North Am 24, 341–364.
36.Marcheselli, VL, Hong, S, Lukiw, WJ, et al. (2003) Novel docosanoids inhibit brain ischemia-reperfusion-mediated leukocyte infiltration and pro-inflammatory gene expression. J Biol Chem 278, 43807–43817.
37.Montero, D, Terova, G, Rimoldi, S, et al. (2015) Modulation of adrenocorticotrophin hormone (ACTH)-induced expression of stress-related genes by PUFA in inter-renal cells from European sea bass (Dicentrarchus labrax). J Nutr Sci 4, e16.
38.Farooqui, AA, Horrocks, LA & Farooqui, T (2007) Modulation of inflammation in brain: a matter of fat. J Neurochem 101, 577–599.
39.Torrecillas, S, Román, L, Rivero-Ramírez, F, et al. (2017) Supplementation of arachidonic acid rich oil in European sea bass juveniles (Dicentrarchus labrax) diets: effects on leucocytes and plasma fatty acid profiles, selected immune parameters and circulating prostaglandins levels. Fish Shellfish Immun 64, 437–445.
40.Hwang, SW, Cho, H, Kwak, J, et al. (2000) Direct activation of capsaicin receptors by products of lipoxygenases: endogenous capsaicin-like substances. Proc Natl Acad Sci U S A 97, 6155–6160.
41.Liu, J, Caballero, MJ, Izquierdo, M, et al. (2002) Necessity of dietary lecithin and eicosapentaenoic acid for growth, survival, stress resistance and lipoprotein formation in gilthead sea bream, Sparus aurata. Fish Sci 68, 1165–1172.
42.Montero, D, Tort, L, Izquierdo, MS, et al. (1998) Depletion of serum alternative complement pathway activity in gilthead seabream caused by α-tocopherol and n-3 HUFA dietary deficiencies. Fish Physiol Biochem 18, 399–407.
43.Ganga, R, Tort, L, Acerete, L, et al. (2006) Modulation of ACTH-induced cortisol release by polyunsaturated fatty acids in interrenal cells from gilthead seabream, Sparus aurata. J Endocrinol 190, 39–45.
44.Benedito-Palos, L, Ballester-Lozano, GF, Simó, P, et al. (2016) Lasting effects of butyrate and low FM/FO diets on growth performance, blood haematology/biochemistry and molecular growth-related markers in gilthead sea bream (Sparus aurata). Aquaculture 454, 8–18.
45.Fernández-Palacios, H, Norberg, B, Izquierdo, M, et al. (2011) Effects of broodstock diet on eggs and larvae. In Larval Fish Nutrition, pp. 151–181 [Holt, JG, editor]. Oxford: Wiley-Blackwell.
46.Ibeas, C, Izquierdo, MS & Lorenzo, A (1994) Effect of different levels of n-3 highly unsaturated fatty acids on growth and fatty acid composition of juvenile gilthead seabream (Sparus aurata). Aquaculture 127, 177–188.
47.AOAC (1995) Official Methods of Analysis of AOAC International, 15th ed. Arlington, VA: AOAC International.
48.Folch, J, Lees, M & Sloane-Stanley, G (1957) A simple method for the isolation and purification of total lipides from animal tissues. J Biol Chem 226, 497–509.
49.Christie, WW (1982) Lipid Analysis. Oxford: Pergamon Press.
50.Izquierdo, MS, Watanabe, T, Takeuchi, T, et al. (1990) Optimum EFA levels in Artemia to meet the EFA requirements of red sea bream (Pagrus major). In The Current Status of Fish Nutrition in Aquaculture: The Proceedings of the Third International Symposium on Feeding and Nutrition in Fish, Toba, Japan, pp. 221–232.
51.Sepulcre, MP, López-Castejón, G, Meseguer, J, et al. (2007) The activation of gilthead seabream professional phagocytes by different PAMPs underlines the behavioural diversity of the main innate immune cells of bony fish. Mol Immunol 44, 2009–2016.
52.García-Castillo, J, Pelegrín, P, Mulero, V, et al. (2002) Molecular cloning and expression analysis of tumor necrosis factor α from a marine fish reveal its constitutive expression and ubiquitous nature. Immunogenetics 54, 200–207.
53.Pelegrín, P, García-Castillo, J, Mulero, V, et al. (2001) Interleukin-1β isolated from a marine fish reveals up-regulated expression in macrophages following activation with lipopolysaccharide and lymphokines. Cytokine 16, 67–72.
54.Cuesta, A, Ángeles Esteban, M & Meseguer, J (2006) Cloning, distribution and up-regulation of the teleost fish MHC class II alpha suggests a role for granulocytes as antigen-presenting cells. Mol Immunol 43, 1275–1285.
55.Acerete, L, Balasch, JC, Castellana, B, et al. (2007) Cloning of the glucocorticoid receptor (GR) in gilthead seabream (Sparus aurata): differential expression of GR and immune genes in gilthead seabream after an immune challenge. Comp Biochem Physiol Part B: Biochem Mol Biol 148, 32–43.
56.Avella, MA, Gioacchini, G, Decamp, O, et al. (2010) Application of multi-species of Bacillus in sea bream larviculture. Aquaculture 305, 12–19.
57.Santos, CRA, Power, DM, Kille, P, et al. (1997) Cloning and sequencing of a full-length sea bream (Sparus aurata) β-actin cDNA. Comp Biochem Physiol Part B: Biochem Mol Biol 117, 185–189.
58.Laizé, V, Pombinho, AR & Cancela, ML (2005) Characterization of Sparus aurata osteonectin cDNA and in silico analysis of protein conserved features: evidence for more than one osteonectin in Salmonidae. Biochimie 87, 411–420.
59.Livak, KJ & Schmittgen, TD (2001) Analysis of relative gene expression data using real-time quantitative PCR and the 2-ΔΔCT method. Methods 25.
60.Schmittgen, TD & Livak, KJ (2008) Analyzing real-time PCR data by the comparative CT method. Nat Protocol 3, 1101–1108.
61.Sokal, RR & Rohlf, FJ (1969) The Principles and Practice of Statistics in Biological Research. San Francisco, CA: WH Freeman and Company
62.Gluckman, PD, Hanson, MA, Spencer, HG, et al. (2005) Environmental influences during development and their later consequences for health and disease: implications for the interpretation of empirical studies. Proc Royal Soc B: Biol Sci 272, 671–677.
63.Mourente, G & Odriozola, JM (1990) Effect of broodstock diets on total lipids and fatty acid composition of larvae of gilthead sea bream (Sparus aurata L.) during yolk sac stage. Fish Physiol Biochem 8, 103–110.
64.Adam, A-C, Skjærven, KH, Whatmore, P, et al. (2018) Parental high dietary arachidonic acid levels modulated the hepatic transcriptome of adult zebrafish (Danio rerio) progeny. PLOS ONE 13, e0201278.
65.Morais, S, Mendes, AC, Castanheira, MF, et al. (2014) New formulated diets for Solea senegalensis broodstock: effects of parental nutrition on biosynthesis of long-chain polyunsaturated fatty acids and performance of early larval stages and juvenile fish. Aquaculture 432, 374–382.
66.Hastings, N, Agaba, M, Tocher, DR, et al. (2001) A vertebrate fatty acid desaturase with Δ5 and Δ6 activities. Proc Natl Acad Sci U S A 98, 14304–14309.
67.Bell, JG, Tocher, DR, Farndale, BM, et al. (1997) The effect of dietary lipid on polyunsaturated fatty acid metabolism in Atlantic salmon (Salmo salar) undergoing parr-smolt transformation. Lipids 32, 515–525.
68.Zheng, X, Seiliez, I, Hastings, N, et al. (2004) Characterization and comparison of fatty acyl Δ6 desaturase cDNAs from freshwater and marine teleost fish species. Comp Biochem Physiol B Biochem Mol Biol 139, 269–279.
69.Francis, DS, Peters, DJ & Turchini, GM (2009) Apparent in vivo Δ-6 desaturase activity, efficiency, and affinity are affected by total dietary C18 PUFA in the freshwater fish Murray cod. J Agric Food Chem 57, 4381–4390.
70.Thanuthong, T, Francis, DS, Senadheera, SPSD, et al. (2011) LC-PUFA biosynthesis in rainbow trout is substrate limited: use of the whole body fatty acid balance method and different 18 : 3n-3/18 : 2n-6 ratios. Lipids 46, 1111–1127.
71.Fernández-Palacios, H, Izquierdo, MS, Robaina, L, et al. (1995) Effect of n-3 HUFA level in broodstock diets on egg quality of gilthead sea bream (Sparus aurata L.). Aquaculture 132, 325–337.
72.Izquierdo, MS, Fernandez-Palacios, H & Tacon, AGJ (2001) Effect of broodstock nutrition on reproductive performance of fish. Aquaculture 197, 25–42.
73.Tocher, DR (2003) Metabolism and functions of lipids and fatty acids in teleost fish. Rev Fish Sci 11, 107–184.
74.Betancor, MB, Sprague, M, Sayanova, O, et al. (2016) Nutritional evaluation of an EPA-DHA oil from transgenic Camelina sativa in feeds for post-smolt Atlantic salmon (Salmo salar L.). PLOS ONE 11, e0159934.
75.Betancor, MB, Sprague, M, Usher, S, et al. (2015) A nutritionally-enhanced oil from transgenic Camelina sativa effectively replaces fish oil as a source of eicosapentaenoic acid for fish. Sci Rep 5, 8104.
76.Thomassen, MS, Rein, D, Berge, GM, et al. (2012) High dietary EPA does not inhibit Δ5 and Δ6 desaturases in Atlantic salmon (Salmo salar L.) fed rapeseed oil diets. Aquaculture 360–361, 78–85.
77.Rowley, AF, Knight, J, Lloyd-Evans, P, et al. (1995) Eicosanoids and their role in immune modulation in fish – a brief overview. Fish Shellfish Immunol 5, 549–567.
78.Song, C, Li, X, Leonard, BE, et al. (2003) Effects of dietary n-3 or n-6 fatty acids on interleukin-1β-induced anxiety, stress, and inflammatory responses in rats. J Lipid Res 44, 1984–1991.
79.Ganga, R (2010) Effect of feeding gilthead seabream (Sparus aurata) with different levels of n-3 and n-6 lipids vegetable oils on fish health and resistance to stress, Universidad de Las Palmas de Gran Canaria. http://hdl.handle.net/10553/4781 (accessed September 2018).
80.Petursdottir, DH & Hardardottir, I (2007) Dietary fish oil increases the number of splenic macrophages secreting TNF-alpha and IL-10 but decreases the secretion of these cytokines by splenic T cells from mice. J Nutr 137, 665–670.
81.Kumar, KA, Lalitha, A, Pavithra, D, et al. (2013) Maternal dietary folate and/or vitamin B12 restrictions alter body composition (adiposity) and lipid metabolism in Wistar rat offspring. J Nutr Biochem 24, 25–31.
82.Bell, JG, McVicar, AH, Park, MT, et al. (1991) High dietary linoleic acid affects the fatty acid compositions of individual phospholipids from tissues of Atlantic salmon (Salmo salar): association with stress susceptibility and cardiac lesion. J Nutr 121, 1163–1172.
83.Basu, S & Srivastava, P (2003) Heat shock proteins in immune response. In Progress in Inflammation Research, pp. 33–42 [van Eden, W, editor]. Basel, Germany: Birkhäuser.