Effects of different dietary lipid sources on fatty acid composition and gene expression in common carp

https://doi.org/10.17221/248/2019-CJASCitation:Ren H., Zhang G., Huang Y., Gao X. (2020): Effects of different dietary lipid sources on fatty acid composition and gene expression in common carp. Czech J. Anim. Sci., 65: 51-57.
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The effects of fatty acid composition in artificial feed on the change in the fatty acid composition of carp muscles and the relationship between Δ6-Fad and Elovl5 genes participating in the regulation of fatty acid synthesis were studied. Juveniles were fed three semi-purified diets (D1–D3) for 6 weeks with different lipid sources: D1, fish oil with high highly unsaturated fatty acids (HUFA); D2, corn oil with high linoleic acid (18:2n-6, LA), D3, linseed oil with high α-linolenic acid (18:3n-3, LNA); then, samples were taken to explore the molecular mechanism and the factors which affect the synthesis of carp HUFA. The content of LA and arachidonic acid (20:4n-6, AA) in common carp fed Diet 2 was higher than in carp receiving D3 (P < 0.05), but the contents of eicosapentaenoic acid (20:5n-3, EPA) and docosahexaenoic acid (22:6n-3, DHA) were lower than in carp fed D1 and D2 (P < 0.05). The liver transcript abundance of Δ6-Fad and Elovl5 in fish fed D2 and D3 at the end of 6 weeks was generally higher than the abundance in the initial stage and in the fish fed D1 (P < 0.05). The results suggest that the common carp can biosynthesise HUFA, and the type and content of fatty acids in feed affected not only the composition and content of fatty acids in common carp muscles, but also the Δ6-Fad and Elovl5 gene expression involved in the biosynthesis of HUFA. Feeding high levels of n-3 HUFA diet can increase the body content of EPA and DHA in common carp. The results of this research may provide a theoretical basis for choosing an appropriate source of lipid for common carp feeds.

 

References:
Ackman RG. Freshwater fish lipids – an overlooked source of beneficial long-chain n-3 fatty acids. Eur. J. Lipid Sci. Tech. 2002;104:253-254. https://doi.org/10.1002/1438-9312(200205)104:5<253::AID-EJLT253>3.0.CO;2-B
 
Alimuddin KV, Satoh S, Takeuchi T, Toshio T, Goro Y. Cloning and overexpression of a masu salmon (Oncorhynchus masou) fatty acid elongase-like gene in zebrafish. Aquaculture. 2008;282:13-18. https://doi.org/10.1016/j.aquaculture.2008.06.033
 
Bell JG, Dick JR, Vicar MC, Sargent JR, Thompson KD. Dietary sunflower, linseed and fish oils affect phospholipid fatty acid composition, development of cardiac lesions, phospholipase activity and eicosanoid production in Atlantic salmon (Salmo salar). Prostaglandins Leukot. Essent. Fatty Acids. 1993;49:665-673. https://doi.org/10.1016/0952-3278(93)90075-8
 
Buzzi M, Henderson RJ, Sargent JR. The desaturation and elongation of linolenic acid and eicosapentaenoic acid by hepatocytes and liver microsomes from rainbow trout (Oncorhynchus mykiss) fed diets containing fish oil or olive oil. Biochim. Biophys. Acta. 1996;1299:235-244.
 
Cook HW. Fatty acid desaturation and chain elongation in eukaryote. In: Vance DE, Vance JE, editors. Biochemistry of Lipids, Lipoproteins and Membranes. Amsterdam, the Netherlands: Elsevier; 1996. pp. 129–152.
 
Donmez M. Determination of fatty acid compositions and cholesterol levels of some freshwater fish living in Porsuk Dam, Turkey. Chem. Nat. Compd. 2009;45:4-17. https://doi.org/10.1007/s10600-009-9219-z
 
Eilander A, Hundscheid DC, Osendarp SJ, Transler C, Zock PL. Effects of n-3 long chain polyunsaturated fatty acid supplementation on visual and cognitive development throughout childhood: A review of human studies. Prostaglandins Leukot. Essent. Fatty Acids. 2007;76:189-203.  https://doi.org/10.1016/j.plefa.2007.01.003
 
Folch J, Lees M, Sloane Stanley GH. A simple method for the isolation and purification of total lipides from animal tissues. J. Biol. Chem. 1957;226:497-509.
 
Gregory MK, See VH, Gibson RA, Schuller KA. Cloning and functional characterisation of a fatty acyl elongase from southern bluefin tuna (Thunnus maccoyii). Comp. Biochem. Physiol. B Biochem. Mol. Biol. 2010;155:178-185.
 
Guler GO, Kiztanir B, Aktumsek A, Citil OB, Ozparlak H. Determination of the seasonal changes on total fatty acid composition and ω3/ω6 ratios of carp (Cyprinus carpio L.) muscle lipids in Beysehir Lake (Turkey). Food Chem. 2008;108:689-694. https://doi.org/10.1016/j.foodchem.2007.10.080
 
Izquierdo MS, Robaina L, Juarez-Carrillo E, Oliva V, Hernandez-Cruz CM, Afonso JM. Regulation of growth, fatty acid composition and delta-6 desaturase expression by dietary lipids in gilthead seabream larvae (Sparus aurata). Fish Physiol. Biochem. 2008;34:117-127.
 
Jakobsson A, Westerberg R, Jacobsson A. Fatty acid elongases in mammals: their regulation and roles in metabolism. Prog. Lipid Res. 2006;45:237-249. https://doi.org/10.1016/j.plipres.2006.01.004
 
Kim S, Lim B, Kim K, Do K. QTL fine mapping for intramuscular fat and fatty acid composition using high-density SNP chip array on SSC12 in Korean native pig × Yorkshire F2 population. Czech J. Anim. Sci. 2019;64:180-188.
 
Li YY, Hu CB, Zheng YJ, Xia XA, Xu WJ, Wang SQ, Chen WZ, Sun ZW, Huang JH. The effects of dietary fatty acids on liver fatty acid composition and delta-6 desaturase expression differ with ambient salinities in Siganus canaliculatus. Comp. Biochem. Physiol. B Biochem. Mol. Biol. 2008;151:183-190.
 
Liu YF. Analysis of fatty acid composition of five freshwater fishes in China. J. Fish. China. 1991;15:169-171.
 
Luo YK. Fatty acid composition of muscle and guts in seven freshwater fishes. J. China Agri. Univ. 2001;4:108-111.
 
Monroig O,·Kabeya N. Desaturases and elongases involved in polyunsaturated fatty acid biosynthesis in aquatic invertebrates: A comprehensive review. Fisheries Sci. 2018;84:911-928. https://doi.org/10.1007/s12562-018-1254-x
 
Mourente G, Tocher DR. In vivo metabolism of [1-14C] linolenic acid (18:3n−3) and [1-14C] eicosapentaenoic acid (20:5n-3) in marine fish: Time-course of the desaturation/elongation pathway. Biochim. Biophys. Acta. 1994;1212:109-118.
 
Prchal M, Vandeputte M, Gela D, Dolezal M, Buchtova H, Rodina M, Flajshans M, Kocour M. Estimation of genetic parameters of fatty acids composition in flesh of market size common carp (Cyprinus carpio L.) and their relation to performance traits revealed that selective breeding can indirectly affect flesh quality. Czech J. Anim. Sci. 2018;63:280-291.
 
Ruyter B, Rosjo C, Masoval K, Einen O, Thomassen MS. Influence of dietary n-3 fatty acids on the desaturation and elongation of [1-14C] 18:2n-6 and [1-14C] 18:3n-3 in Atlantic salmon hepatocytes. Fish Physiol. Biochem. 2000;23:151-158.
 
Sargent JR, Tocher DR, Bell JG. The lipids. In: Halver JE, Hardy RW, edtors. Fish Nutrition. San Diego, USA: Academic Press; 2002. pp 181-257.
 
Shaliutina-Kolesova A, Xian M, Nian R. Antioxidant defence system in protein fractions of common carp (Cyprinus carpio) seminal plasma. Czech J. Anim. Sci. 2019;64:265-271.
 
Tocher DR. Fatty acid requirements in ontogeny of marine and freshwater fish. Aquac. Res. 2010;41:717-732. https://doi.org/10.1111/j.1365-2109.2008.02150.x
 
Tocher DR, Bell JG, McGhee F, Dick JR, Fonseca-Madrigal J. Effects of dietary lipid level and vegetable oil on fatty acid metabolism in Atlantic salmon (Salmo salar) over the entire production cycle. Fish Physiol. Biochem. 2003;29:193-209.
 
Tocher DR, Zheng X, Schlechtriem C, Hastings N, Dick JR, Teale AJ. Highly unsaturated fatty acid synthesis in marine fish: Cloning, functional characterization, and nutritional regulation of fatty acyl Δ6-desaturase of Atlantic cod (Gadus morhua L.). Lipids. 2006;41:1003-1016. https://doi.org/10.1007/s11745-006-5051-4
 
Ulmann L, Bouzianne M, Mimouni V, Belleville J, Poisson JP. Relationship between rat liver microsomal Δ6- and Δ5-desaturase activities and fatty acid composition: Comparative effects of coconut and salmon oils during protein restriction. J. Nutr. Biochem. 1992;3:188-193.
 
Zhao Y, Wang LK, Gao J. Effects of dietary highly unsaturated fatty acid levels on growth, fatty acid profiles, antioxidant activities, mucus immune responses and hepatic lipid metabolism related gene expressions in loach (Misgurnus anguillicaudatus) juveniles. Aquac. Res. 2019;50:2486-2495.
 
Zheng X, Tocher DR, Dickson CA, Bell JG, Teale AJ. Effects of diets containing vegetable oil on expression of genes involved in polyunsaturated fatty acid biosynthesis in liver of Atlantic salmon (Salmo salar). Aquaculture. 2004;236:467-483. https://doi.org/10.1016/j.aquaculture.2004.02.003
 
Zheng X, Tocher DR, Dickson CA, Bell JG, Teale AJ. Highly unsaturated fatty acid synthesis in vertebrates: New insights with the cloning and characterization of a Δ6-desaturase of Atlantic salmon. Lipids. 2005a;40:13-24. https://doi.org/10.1007/s11745-005-1355-7
 
Zheng X, Torstensen BE, Tocher DR, Dick JR, Henderson RJ, Bell JG. Environmental and dietary influences on highly unsaturated fatty acid biosynthesis and expression of fatty acyl desaturase and elongase genes in liver of Atlantic salmon (Salmo salar). Bioch. Biophys. Acta. 2005b;1734:13-24.  https://doi.org/10.1016/j.bbalip.2005.01.006
 
Zheng X, King Z, Xu Y, Monroig O, Morais S, Tocher DR. Physiological roles of fatty acyl desaturases and elongases in marine fish: Characterisation of cDNAs of fatty acyl Δ6-desaturase and Elovl5 elongase of cobia (Rachycentron canadum). Aquaculture. 2009;290:122-131. https://doi.org/10.1016/j.aquaculture.2009.02.010
 
Ackman RG. Freshwater fish lipids – an overlooked source of beneficial long-chain n-3 fatty acids. Eur. J. Lipid Sci. Tech. 2002;104:253-254. https://doi.org/10.1002/1438-9312(200205)104:5<253::AID-EJLT253>3.0.CO;2-B
 
Alimuddin KV, Satoh S, Takeuchi T, Toshio T, Goro Y. Cloning and overexpression of a masu salmon (Oncorhynchus masou) fatty acid elongase-like gene in zebrafish. Aquaculture. 2008;282:13-18. https://doi.org/10.1016/j.aquaculture.2008.06.033
 
Bell JG, Dick JR, Vicar MC, Sargent JR, Thompson KD. Dietary sunflower, linseed and fish oils affect phospholipid fatty acid composition, development of cardiac lesions, phospholipase activity and eicosanoid production in Atlantic salmon (Salmo salar). Prostaglandins Leukot. Essent. Fatty Acids. 1993;49:665-673. https://doi.org/10.1016/0952-3278(93)90075-8
 
Buzzi M, Henderson RJ, Sargent JR. The desaturation and elongation of linolenic acid and eicosapentaenoic acid by hepatocytes and liver microsomes from rainbow trout (Oncorhynchus mykiss) fed diets containing fish oil or olive oil. Biochim. Biophys. Acta. 1996;1299:235-244.
 
Cook HW. Fatty acid desaturation and chain elongation in eukaryote. In: Vance DE, Vance JE, editors. Biochemistry of Lipids, Lipoproteins and Membranes. Amsterdam, the Netherlands: Elsevier; 1996. pp. 129–152.
 
Donmez M. Determination of fatty acid compositions and cholesterol levels of some freshwater fish living in Porsuk Dam, Turkey. Chem. Nat. Compd. 2009;45:4-17. https://doi.org/10.1007/s10600-009-9219-z
 
Eilander A, Hundscheid DC, Osendarp SJ, Transler C, Zock PL. Effects of n-3 long chain polyunsaturated fatty acid supplementation on visual and cognitive development throughout childhood: A review of human studies. Prostaglandins Leukot. Essent. Fatty Acids. 2007;76:189-203.  https://doi.org/10.1016/j.plefa.2007.01.003
 
Folch J, Lees M, Sloane Stanley GH. A simple method for the isolation and purification of total lipides from animal tissues. J. Biol. Chem. 1957;226:497-509.
 
Gregory MK, See VH, Gibson RA, Schuller KA. Cloning and functional characterisation of a fatty acyl elongase from southern bluefin tuna (Thunnus maccoyii). Comp. Biochem. Physiol. B Biochem. Mol. Biol. 2010;155:178-185.
 
Guler GO, Kiztanir B, Aktumsek A, Citil OB, Ozparlak H. Determination of the seasonal changes on total fatty acid composition and ω3/ω6 ratios of carp (Cyprinus carpio L.) muscle lipids in Beysehir Lake (Turkey). Food Chem. 2008;108:689-694. https://doi.org/10.1016/j.foodchem.2007.10.080
 
Izquierdo MS, Robaina L, Juarez-Carrillo E, Oliva V, Hernandez-Cruz CM, Afonso JM. Regulation of growth, fatty acid composition and delta-6 desaturase expression by dietary lipids in gilthead seabream larvae (Sparus aurata). Fish Physiol. Biochem. 2008;34:117-127.
 
Jakobsson A, Westerberg R, Jacobsson A. Fatty acid elongases in mammals: their regulation and roles in metabolism. Prog. Lipid Res. 2006;45:237-249. https://doi.org/10.1016/j.plipres.2006.01.004
 
Kim S, Lim B, Kim K, Do K. QTL fine mapping for intramuscular fat and fatty acid composition using high-density SNP chip array on SSC12 in Korean native pig × Yorkshire F2 population. Czech J. Anim. Sci. 2019;64:180-188.
 
Li YY, Hu CB, Zheng YJ, Xia XA, Xu WJ, Wang SQ, Chen WZ, Sun ZW, Huang JH. The effects of dietary fatty acids on liver fatty acid composition and delta-6 desaturase expression differ with ambient salinities in Siganus canaliculatus. Comp. Biochem. Physiol. B Biochem. Mol. Biol. 2008;151:183-190.
 
Liu YF. Analysis of fatty acid composition of five freshwater fishes in China. J. Fish. China. 1991;15:169-171.
 
Luo YK. Fatty acid composition of muscle and guts in seven freshwater fishes. J. China Agri. Univ. 2001;4:108-111.
 
Monroig O,·Kabeya N. Desaturases and elongases involved in polyunsaturated fatty acid biosynthesis in aquatic invertebrates: A comprehensive review. Fisheries Sci. 2018;84:911-928. https://doi.org/10.1007/s12562-018-1254-x
 
Mourente G, Tocher DR. In vivo metabolism of [1-14C] linolenic acid (18:3n−3) and [1-14C] eicosapentaenoic acid (20:5n-3) in marine fish: Time-course of the desaturation/elongation pathway. Biochim. Biophys. Acta. 1994;1212:109-118.
 
Prchal M, Vandeputte M, Gela D, Dolezal M, Buchtova H, Rodina M, Flajshans M, Kocour M. Estimation of genetic parameters of fatty acids composition in flesh of market size common carp (Cyprinus carpio L.) and their relation to performance traits revealed that selective breeding can indirectly affect flesh quality. Czech J. Anim. Sci. 2018;63:280-291.
 
Ruyter B, Rosjo C, Masoval K, Einen O, Thomassen MS. Influence of dietary n-3 fatty acids on the desaturation and elongation of [1-14C] 18:2n-6 and [1-14C] 18:3n-3 in Atlantic salmon hepatocytes. Fish Physiol. Biochem. 2000;23:151-158.
 
Sargent JR, Tocher DR, Bell JG. The lipids. In: Halver JE, Hardy RW, edtors. Fish Nutrition. San Diego, USA: Academic Press; 2002. pp 181-257.
 
Shaliutina-Kolesova A, Xian M, Nian R. Antioxidant defence system in protein fractions of common carp (Cyprinus carpio) seminal plasma. Czech J. Anim. Sci. 2019;64:265-271.
 
Tocher DR. Fatty acid requirements in ontogeny of marine and freshwater fish. Aquac. Res. 2010;41:717-732. https://doi.org/10.1111/j.1365-2109.2008.02150.x
 
Tocher DR, Bell JG, McGhee F, Dick JR, Fonseca-Madrigal J. Effects of dietary lipid level and vegetable oil on fatty acid metabolism in Atlantic salmon (Salmo salar) over the entire production cycle. Fish Physiol. Biochem. 2003;29:193-209.
 
Tocher DR, Zheng X, Schlechtriem C, Hastings N, Dick JR, Teale AJ. Highly unsaturated fatty acid synthesis in marine fish: Cloning, functional characterization, and nutritional regulation of fatty acyl Δ6-desaturase of Atlantic cod (Gadus morhua L.). Lipids. 2006;41:1003-1016. https://doi.org/10.1007/s11745-006-5051-4
 
Ulmann L, Bouzianne M, Mimouni V, Belleville J, Poisson JP. Relationship between rat liver microsomal Δ6- and Δ5-desaturase activities and fatty acid composition: Comparative effects of coconut and salmon oils during protein restriction. J. Nutr. Biochem. 1992;3:188-193.
 
Zhao Y, Wang LK, Gao J. Effects of dietary highly unsaturated fatty acid levels on growth, fatty acid profiles, antioxidant activities, mucus immune responses and hepatic lipid metabolism related gene expressions in loach (Misgurnus anguillicaudatus) juveniles. Aquac. Res. 2019;50:2486-2495.
 
Zheng X, Tocher DR, Dickson CA, Bell JG, Teale AJ. Effects of diets containing vegetable oil on expression of genes involved in polyunsaturated fatty acid biosynthesis in liver of Atlantic salmon (Salmo salar). Aquaculture. 2004;236:467-483. https://doi.org/10.1016/j.aquaculture.2004.02.003
 
Zheng X, Tocher DR, Dickson CA, Bell JG, Teale AJ. Highly unsaturated fatty acid synthesis in vertebrates: New insights with the cloning and characterization of a Δ6-desaturase of Atlantic salmon. Lipids. 2005a;40:13-24. https://doi.org/10.1007/s11745-005-1355-7
 
Zheng X, Torstensen BE, Tocher DR, Dick JR, Henderson RJ, Bell JG. Environmental and dietary influences on highly unsaturated fatty acid biosynthesis and expression of fatty acyl desaturase and elongase genes in liver of Atlantic salmon (Salmo salar). Bioch. Biophys. Acta. 2005b;1734:13-24.  https://doi.org/10.1016/j.bbalip.2005.01.006
 
Zheng X, King Z, Xu Y, Monroig O, Morais S, Tocher DR. Physiological roles of fatty acyl desaturases and elongases in marine fish: Characterisation of cDNAs of fatty acyl Δ6-desaturase and Elovl5 elongase of cobia (Rachycentron canadum). Aquaculture. 2009;290:122-131. https://doi.org/10.1016/j.aquaculture.2009.02.010
 
Ackman RG. Freshwater fish lipids – an overlooked source of beneficial long-chain n-3 fatty acids. Eur. J. Lipid Sci. Tech. 2002;104:253-254. https://doi.org/10.1002/1438-9312(200205)104:5<253::AID-EJLT253>3.0.CO;2-B
 
Alimuddin KV, Satoh S, Takeuchi T, Toshio T, Goro Y. Cloning and overexpression of a masu salmon (Oncorhynchus masou) fatty acid elongase-like gene in zebrafish. Aquaculture. 2008;282:13-18. https://doi.org/10.1016/j.aquaculture.2008.06.033
 
Bell JG, Dick JR, Vicar MC, Sargent JR, Thompson KD. Dietary sunflower, linseed and fish oils affect phospholipid fatty acid composition, development of cardiac lesions, phospholipase activity and eicosanoid production in Atlantic salmon (Salmo salar). Prostaglandins Leukot. Essent. Fatty Acids. 1993;49:665-673. https://doi.org/10.1016/0952-3278(93)90075-8
 
Buzzi M, Henderson RJ, Sargent JR. The desaturation and elongation of linolenic acid and eicosapentaenoic acid by hepatocytes and liver microsomes from rainbow trout (Oncorhynchus mykiss) fed diets containing fish oil or olive oil. Biochim. Biophys. Acta. 1996;1299:235-244.
 
Cook HW. Fatty acid desaturation and chain elongation in eukaryote. In: Vance DE, Vance JE, editors. Biochemistry of Lipids, Lipoproteins and Membranes. Amsterdam, the Netherlands: Elsevier; 1996. pp. 129–152.
 
Donmez M. Determination of fatty acid compositions and cholesterol levels of some freshwater fish living in Porsuk Dam, Turkey. Chem. Nat. Compd. 2009;45:4-17. https://doi.org/10.1007/s10600-009-9219-z
 
Eilander A, Hundscheid DC, Osendarp SJ, Transler C, Zock PL. Effects of n-3 long chain polyunsaturated fatty acid supplementation on visual and cognitive development throughout childhood: A review of human studies. Prostaglandins Leukot. Essent. Fatty Acids. 2007;76:189-203.  https://doi.org/10.1016/j.plefa.2007.01.003
 
Folch J, Lees M, Sloane Stanley GH. A simple method for the isolation and purification of total lipides from animal tissues. J. Biol. Chem. 1957;226:497-509.
 
Gregory MK, See VH, Gibson RA, Schuller KA. Cloning and functional characterisation of a fatty acyl elongase from southern bluefin tuna (Thunnus maccoyii). Comp. Biochem. Physiol. B Biochem. Mol. Biol. 2010;155:178-185.
 
Guler GO, Kiztanir B, Aktumsek A, Citil OB, Ozparlak H. Determination of the seasonal changes on total fatty acid composition and ω3/ω6 ratios of carp (Cyprinus carpio L.) muscle lipids in Beysehir Lake (Turkey). Food Chem. 2008;108:689-694. https://doi.org/10.1016/j.foodchem.2007.10.080
 
Izquierdo MS, Robaina L, Juarez-Carrillo E, Oliva V, Hernandez-Cruz CM, Afonso JM. Regulation of growth, fatty acid composition and delta-6 desaturase expression by dietary lipids in gilthead seabream larvae (Sparus aurata). Fish Physiol. Biochem. 2008;34:117-127.
 
Jakobsson A, Westerberg R, Jacobsson A. Fatty acid elongases in mammals: their regulation and roles in metabolism. Prog. Lipid Res. 2006;45:237-249. https://doi.org/10.1016/j.plipres.2006.01.004
 
Kim S, Lim B, Kim K, Do K. QTL fine mapping for intramuscular fat and fatty acid composition using high-density SNP chip array on SSC12 in Korean native pig × Yorkshire F2 population. Czech J. Anim. Sci. 2019;64:180-188.
 
Li YY, Hu CB, Zheng YJ, Xia XA, Xu WJ, Wang SQ, Chen WZ, Sun ZW, Huang JH. The effects of dietary fatty acids on liver fatty acid composition and delta-6 desaturase expression differ with ambient salinities in Siganus canaliculatus. Comp. Biochem. Physiol. B Biochem. Mol. Biol. 2008;151:183-190.
 
Liu YF. Analysis of fatty acid composition of five freshwater fishes in China. J. Fish. China. 1991;15:169-171.
 
Luo YK. Fatty acid composition of muscle and guts in seven freshwater fishes. J. China Agri. Univ. 2001;4:108-111.
 
Monroig O,·Kabeya N. Desaturases and elongases involved in polyunsaturated fatty acid biosynthesis in aquatic invertebrates: A comprehensive review. Fisheries Sci. 2018;84:911-928. https://doi.org/10.1007/s12562-018-1254-x
 
Mourente G, Tocher DR. In vivo metabolism of [1-14C] linolenic acid (18:3n−3) and [1-14C] eicosapentaenoic acid (20:5n-3) in marine fish: Time-course of the desaturation/elongation pathway. Biochim. Biophys. Acta. 1994;1212:109-118.
 
Prchal M, Vandeputte M, Gela D, Dolezal M, Buchtova H, Rodina M, Flajshans M, Kocour M. Estimation of genetic parameters of fatty acids composition in flesh of market size common carp (Cyprinus carpio L.) and their relation to performance traits revealed that selective breeding can indirectly affect flesh quality. Czech J. Anim. Sci. 2018;63:280-291.
 
Ruyter B, Rosjo C, Masoval K, Einen O, Thomassen MS. Influence of dietary n-3 fatty acids on the desaturation and elongation of [1-14C] 18:2n-6 and [1-14C] 18:3n-3 in Atlantic salmon hepatocytes. Fish Physiol. Biochem. 2000;23:151-158.
 
Sargent JR, Tocher DR, Bell JG. The lipids. In: Halver JE, Hardy RW, edtors. Fish Nutrition. San Diego, USA: Academic Press; 2002. pp 181-257.
 
Shaliutina-Kolesova A, Xian M, Nian R. Antioxidant defence system in protein fractions of common carp (Cyprinus carpio) seminal plasma. Czech J. Anim. Sci. 2019;64:265-271.
 
Tocher DR. Fatty acid requirements in ontogeny of marine and freshwater fish. Aquac. Res. 2010;41:717-732. https://doi.org/10.1111/j.1365-2109.2008.02150.x
 
Tocher DR, Bell JG, McGhee F, Dick JR, Fonseca-Madrigal J. Effects of dietary lipid level and vegetable oil on fatty acid metabolism in Atlantic salmon (Salmo salar) over the entire production cycle. Fish Physiol. Biochem. 2003;29:193-209.
 
Tocher DR, Zheng X, Schlechtriem C, Hastings N, Dick JR, Teale AJ. Highly unsaturated fatty acid synthesis in marine fish: Cloning, functional characterization, and nutritional regulation of fatty acyl Δ6-desaturase of Atlantic cod (Gadus morhua L.). Lipids. 2006;41:1003-1016. https://doi.org/10.1007/s11745-006-5051-4
 
Ulmann L, Bouzianne M, Mimouni V, Belleville J, Poisson JP. Relationship between rat liver microsomal Δ6- and Δ5-desaturase activities and fatty acid composition: Comparative effects of coconut and salmon oils during protein restriction. J. Nutr. Biochem. 1992;3:188-193.
 
Zhao Y, Wang LK, Gao J. Effects of dietary highly unsaturated fatty acid levels on growth, fatty acid profiles, antioxidant activities, mucus immune responses and hepatic lipid metabolism related gene expressions in loach (Misgurnus anguillicaudatus) juveniles. Aquac. Res. 2019;50:2486-2495.
 
Zheng X, Tocher DR, Dickson CA, Bell JG, Teale AJ. Effects of diets containing vegetable oil on expression of genes involved in polyunsaturated fatty acid biosynthesis in liver of Atlantic salmon (Salmo salar). Aquaculture. 2004;236:467-483. https://doi.org/10.1016/j.aquaculture.2004.02.003
 
Zheng X, Tocher DR, Dickson CA, Bell JG, Teale AJ. Highly unsaturated fatty acid synthesis in vertebrates: New insights with the cloning and characterization of a Δ6-desaturase of Atlantic salmon. Lipids. 2005a;40:13-24. https://doi.org/10.1007/s11745-005-1355-7
 
Zheng X, Torstensen BE, Tocher DR, Dick JR, Henderson RJ, Bell JG. Environmental and dietary influences on highly unsaturated fatty acid biosynthesis and expression of fatty acyl desaturase and elongase genes in liver of Atlantic salmon (Salmo salar). Bioch. Biophys. Acta. 2005b;1734:13-24.  https://doi.org/10.1016/j.bbalip.2005.01.006
 
Zheng X, King Z, Xu Y, Monroig O, Morais S, Tocher DR. Physiological roles of fatty acyl desaturases and elongases in marine fish: Characterisation of cDNAs of fatty acyl Δ6-desaturase and Elovl5 elongase of cobia (Rachycentron canadum). Aquaculture. 2009;290:122-131. https://doi.org/10.1016/j.aquaculture.2009.02.010
 
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