Long-Chain Polyunsaturated Fatty Acids in Fish: Recent Advances on Desaturases and Elongases Involved in Their Biosynthesis

Authors

  • Óscar Monroig Instituto de Acuicultura Torre de la Sal, IATS-CSIC
  • Juan C. Navarro Instituto de Acuicultura Torre de la Sal, IATS-CSIC
  • Douglas R. Tocher Institute of Aquaculture, School of Natural Sciences, University of Stirling

Keywords:

biosynthesis, desaturase, elongase, essential fatty acid, fish nutrition, long-chain polyunsaturated fatty acids

Abstract

The pathways of long-chain polyunsaturated fatty acid (LC-PUFA) biosynthesis have been extensively investigated in farmed fish both for determining the specific dietary essential fatty acids that guarantee optimal growth and development in captivity, and also for maintaining high levels of the healthy n-3 LC-PUFA for human consumers. It is now well established that the capacity for LC-PUFA biosynthesis of a particular species depends upon the complement of the key enzymes required, namely the fatty acyl desaturases (Fad) and elongases of very long-chain fatty acids (Elovl). This paper reviews the recent progress made on the molecular aspects of Fad- and Elovl-encoding genes isolated so far from fish species. More specifically, we discuss the potential implications that Fad and Elovl functions and expression patterns have on the ability of a particular fish species to thrive on sustainable vegetable-based aquafeeds.

Downloads

Download data is not yet available.

References

Agaba, M., Tocher, D.R., Dickson, C., Dick, J.R., Teale, A.J. (2004) Zebrafish cDNA encoding multifunctional fatty acid elongase involved in production of eicosapentaenoic (20:5n-3) and docosahexaenoic (22:6n-3) acids. Mar. Biotechnol. 6, 251-261.

Agaba, M.K., Tocher, D.R., Dickson, C.A., Zheng, X., Dick, J.R., Teale, A.J. (2005) Cloning and functional characterisation of polyunsaturated fatty acid elongases from marine and freshwater teleost fish. Comp. Biochem. Physiol. 142B, 342–352.

Alimuddin., Yoshizaki, G., Kiron, V., Satoh, S., Takeuchi, T. (2005) Enhancement of EPA and DHA biosynthesis by over-expression of masu salmon Δ6-desaturase-like gene in zebrafish. Transgenic Res. 14, 159–165.

Alimuddin., Yoshizaki, G., Kiron, V., Satoh, S., Takeuchi, T. (2007) Expression of masu salmon Δ5-desaturase-like gene elevated EPA and DHA biosynthesis in zebrafish. Mar. Biotechnol. 9, 92– 100

Alimuddin, Kiron, V., Satoh, S., Takeuchi, T., Yoshizaki, G. (2008) Cloning and over-expression of a masu salmon (Oncorhynchus masou) fatty acid elongase-like gene in zebrafish. Aquaculture 282, 13–18

Aveldaño, M.I. (1987) A novel group of very long chain polyenoic fatty acids in dipolyunsaturated phosphatidylcholines from vertebrate retina. J. Biol. Chem. 262, 1172–1179.

Aveldaño, M.I. (1988) Phospholipid species containing long and very long polyenoic fatty acids remain with rhodopsin after hexane extraction of photoreceptor membranes. Biochemistry 27, 1229-1239.

Aveldaño, M.I., Robinson, B.S., Johnson, D., Poulos, A. (1993) Long and very long chain polyunsaturated fatty acids of the n-6 series in rat seminiferous tubules. active desaturation of 24:4n-6 to 24:5n-6 and concomitant formation of odd and even chain tetraenoic and pentaenoic fatty acids up to C32. J. Biol. Chem. 268, 11663–11669.

Bell, J.G., Waagbø, R. (2008) Safe and nutritious aquaculture produce: benefits and risks of alternative sustainable aquafeeds. In: Holmer, M., Black, K. D., Duarte, C. M., Marba, N., Karakassis, I. (Eds.), Aquaculture in the ecosystem. Berlin, Springer Verlag BV, pp.185-225.

Bell, M.V., Tocher D.R. (2009a) Biosynthesis of fatty acids; general principles and new directions. Pp. 211–236 in Lipids in Aquatic Ecosystems, M. T. Arts, M. Brett, and M. Kainz, eds. New York, NY: Springer-Verlag.

Bell, J.G., Tocher, D.R. (2009b) Farmed Fish: The impact of diet on fatty acid compositions. In: Oils and Fats Handbook volume 4; Fish Oils, (Rossell, B., Ed), pp. 171-184. Leatherhead Food International, Leatherhead.

Brouwer, I.A., Geelen, A., Katan, M.B. (2006) n-3 Fatty acids, cardiac arrhythmia and fatal coronary heart disease. Prog. Lipid Res. 4, 357-367.

Buzzi, M., Henderson, R.J., Sargent, J.R. (1996) 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 1299, 235-244.

Buzzi, M., Henderson, R.J., Sargent, J.R. (1997) Biosynthesis of docosahexaenoic acid in trout hepatocytes proceeds via 24-carbon intermediates. Comp Biochem Physiol 116B, 263–267.

Calder, P.C., Yaqoob, P. (2009) Understanding omega-3 polyunsaturated fatty acids. Postgrad. Med. 121, 148-57.

Cameron, D.J., Tong, Z., Yang, Z., Kaminoh, J., Kamiyah, S., Chen, H., Zeng, J., Chen, Y., Lou, L., Zhang, K. (2007) Essential role of Elovl4 in very long chain fatty acid synthesis, skin permeability barrier function, and neonatal survival. Int. J. Biol. Sci. 3, 111-119.

Carmona-Antoñanzas, G., Monroig, Ó., Dick, J.R., Davie, A., Tocher D.R. 2011. Biosynthesis of very long-chain fatty acids (C > 24) in Atlantic salmon: Cloning, functional characterisation, and tissue distribution of an Elovl4 elongase. Comp. Biochem. Physiol. 159B, 122-129.

Cherian, G., Sim, J.S. (2001) Maternal dietary -linolenic acid (18:3n-3) alters n-3 polyunsaturated metabolism and liver enzyme activity inhatched chicks. Poult. Sci. 80, 901–905.

Cook, H.W., McMaster, R.C.R. (2004) in Biochemistry of Lipids, Lipoproteins and Membranes, eds Vance DE, Vance JE (Elsevier, Amsterdam). pp 181-204.

Damude, H.G., Zhang, H., Farrall, L., Ripp, K.G., Tomb, J-F., Hollerbach, D., Yadav, N.S. (2006) Identification of bifunctional �Δ12/3 fatty acid desaturases for improving the ratio of 3 to 6 fatty acids in microbes and plants. Proc. Natl. Acad. Sci. USA 103, 9446–9451.

D‟Andrea, S., Guillou, H., Jan, S., Catheline, D., Thibault, J.N., Bouriel, M., Rioux, V., Legrand, P. (2002) The same rat Δ6-desaturase not only acts on 18- but also on 24-carbon fatty acids in very-long-chain polyunsaturated fatty acid biosynthesis. Biochem. J. 364, 49–55.

De Antueno, R.J., Knickle, L.C., Smith, H., Elliot, M.L., Allen, S.J., Nwaka, S., Winther, M.D. (2001) Activity of human Δ5 and Δ6 desaturases on multiple n-3 and n-6 polyunsaturated fatty acids. FEBS Lett. 509, 77–80.

Eilander, A., Hundscheid, D.C., Osendarp, S.J., Trander, C., Zock, P.L. (2007) 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 76, 189-203.

Food and Agricultural Organization of the United Nations (FAO) (2009) The State of World Fisheries and Aquaculture 2008 (SOFIA), FAO Fisheries and Aquaculture Department, Rome.

Funk, C.D. (2001) Prostaglandins and leukotrienes: Advances in eicosanoid biology. Science 294, 1871-1875.

Furland, N.E., Maldonado, E.N., Aveldaño, M.I. (2003) Very long chain PUFA in murine testicular triglycerides and cholesterol esters. Lipids 38, 73-80.

Furland, N.E., Oresti, G.M., Antollini, S.S., Venturino, A., Maldonado, E.N., Aveldaño, M.I. (2007a) Very long-chain polyunsaturated fatty acids are the major acyl groups of sphingomyelins and ceramides in the head of mammalian spermatozoa. J. Biol. Chem. 282, 18151-18161.

Furland, N.E., Maldonado, E.N., Ayuza-Aresti, P., Aveldaño, M.I. (2007b) Changes in lipids containing long- and very long-chain polyunsaturated fatty acids in cryptorchid rat testes. Biol. Reprod. 77, 181-188.

González-Rovira, A., Mourente, G., Zheng, X., Tocher, D.R., Pendón, C. (2009) Molecular and functional characterization and expression analysis of a Δ6 fatty acyl desaturase cDNA of European sea bass (Dicentrarchus labrax L.). Aquaculture 298, 90–100.

Gregory, M., See, V.H.L., Gibson, R.A., Shuller, K.A. (2010) Cloning and functional characterisation of a fatty acyl elongase from southern bluefin tuna (Thunnus maccoyii). Comp. Biochem. Physiol. 155B, 178-185.

Guillou, H., Zadravec, D., Martin, P.G.P., Jacobsson, A. (2010) The key roles of elongases and desaturases in mammalian fatty acid metabolism: Insights from transgenic mice. Prog. Lipid Res. 49, 186-199.

Hastings, N., Agaba, M., Tocher, D.R., Leaver, M.J., Dick, J.R., Sargent, J.R., Teale A.J. (2001) A vertebrate fatty acid desaturase with Δ5 and Δ6 activities. Proc. Natl. Acad. Sci. USA 98, 14304-14309.

Hastings, N., Agaba, M.K., Tocher, D.R., Zheng, X., Dickson, C.A., Dick, J.R., Teale, A.J. (2005) Molecular cloning and functional characterization of fatty acyl desaturase and elongase cDNAs involved in the production of eicosapentaenoic and docosahexaenoic acids from α-linolenic acid in Atlantic salmon (Salmo salar). Mar. Biotechnol. 6, 463-474.

Ishak, S.D., Tan, S-H., Khong, H-K., Jaya-Ram, A., Enyu, Y-L., Kuah, M-K., Shu-Chien, A.C. (2008) Upregulated mRNA expression of desaturase and elongase, two enzymes involved in highly unsaturated fatty acids biosynthesis pathways during follicle maturation in zebrafish. Reprod. Biol. Endocrinol. 6, 56-65.

Izquierdo, M.S., Socorro, J., Arantzamendi, L., Hernández-Cruz, C.M. (2000) Recent advances in lipid nutrition in fish larvae. Fish Physiol. Biochem. 22, 97-107.

Izquierdo, M.S., Fernández-Palacios, H., Tacon, A.G.J. (2001) Effect of broodstock nutrition on reproductive performance of fish. Aquaculture 197, 25-42.

Izquierdo, M.S., Robaina, L., Juárez-Carrillo, E., Oliva, V., Hernández-Cruz, C.M., Afonso, J.M. (2008) Regulation of growth, fatty acid composition and delta 6 desaturase expression by dietary lipids in gilthead seabream larvae (Sparus aurata). Fish Physiol. Biochem. 34, 117–127.

Jakobsson, A., Westerberg, R., Jacobsson, A. (2006) Fatty acid elongases in mammals: Their regulation and roles in metabolism. Prog. Lipid Res. 45, 237-249.

Jaya-Ram, A., Ishak, S.D., Enyu, Y-L., Kuah, M-K., Wong, K-L., Shu-Chien, A.C. (2011) Molecular cloning and ontogenic mRNA expression of fatty acid desaturase in the carnivorous striped snakehead fish (Channa striata). Comp. Biochem. Physiol. 158A, 415–422

Leaver, M.J., Bautista, J.M., Björnsson, T., Jönsson, E., Krey, G., Tocher, D.R., Torstensen, B.E. (2008a) Towards fish lipid nutrigenomics: current state and prospects for finfish aquaculture. Rev. Fisheries Sci. 16, 71-92.

Leaver, M.J., Villeneuve, L.A., Obach, A., Jensen, L., Bron, J.E., Tocher, D.R., Taggart, J.B. (2008b) 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 Genomics 9, 299.

Li, W., Chen, Y., Cameron, D.J., Wang, C., Karan, G., Yang, Z., Zhao, Y., Pearson, E., Chen, H., Deng, C., Howes, K., Zhang, K. (2007a) Elovl4 haploinsufficiency does not induce early onset retinal degeneration in mice. Vision Res. 47, 714-722.

Li, W., Sandhoff, R., Kono, M., Zerfas, P., Hoffman, V., Ding, B.C-H., Proia, R.L., Deng, C-X. (2007b) Depletion of ceramides with very long chain fatty acids causes defective skin permeability barrier function, and neonatal lethality in ELOVL4 deficient mice. Int. J. Biol. Sci. 3, 120-128.

Li, Y., Monroig, Ó., Zhang, L., Wang, S., Zheng, X., Dick, J.R., You, C., Tocher, D.R. (2010) Vertebrate fatty acyl desaturase with Δ4 activity. Proc. Natl. Acad. Sci. USA 107, 16840-16845.

Marquardt, A., Stöhr, H., White, K., Weber, B.H.F. (2000) cDNA cloning, genomic structure, and chromosomal localization of three members of the human fatty acid desaturase family. Genomics 66, 175-183.

Mazorra, C., Bruce, M., Bell, J.G., Davie, A., Alorend, E., Jordan, N., Rees, J., Papanikos, N., Porter, M., Bromage, N. (2003) Dietary lipid enhancement of broodstock reproductive performance and egg and larval quality in Atlantic halibut (Hippoglossus hippoglossus). Aquaculture 227, 21-33.

McMahon, A., Jackson, S.N., Woods, A.S., Kedzierski, W. (2007) A Stargardt disease-3 mutation in the mouse Elovl4 gene causes retinal deficiency of C32-C36 acyl phosphatidylcholines. FEBS Lett. 581, 5459-5463.

Mohd-Yusof, N.Y., Monroig, Ó., Mohd-Adnan, A., Wan, K.-L., Tocher, D.R. (2010) Investigation of highly unsaturated fatty acid metabolism in the Asian sea bass, Lates calcarifer. Fish Physiol. Biochem. 36, 827-843.

Monroig, Ó., Rotllant, J., Sánchez, E., Cerdá-Reverter, J.M., Tocher, D.R. (2009) Expression of long-chain polyunsaturated fatty acid (LC-PUFA) biosynthesis genes during zebrafish Danio rerio early embryogenesis. Biochim. Biophys. Acta 1791, 1093–1101.

Monroig, Ó., Zheng, X., Morais, S., Leaver, M.J., Taggart, J.B., Tocher, D.R. (2010a) Multiple genes for functional Δ6 fatty acyl desaturases (Fad) in Atlantic salmon (Salmo salar L.): Gene and cDNA characterization, functional expression, tissue distribution and nutritional regulation. Biochim. Biophys. Acta 1801, 1072–1081.

Monroig, Ó., Rotllant, J., Cerdá-Reverter, J.M., Dick, J.R., Figueras, A., Tocher, D.R. (2010b) Expression and role of Elovl4 elongases in biosynthesis of very long-chain fatty acids during zebrafish Danio

rerio early embryonic development. Biochim. Biophys. Acta 1801, 1145-1154.

Monroig, Ó, Webb, K., Ibarra-Castro, L., Holt, G.J., Tocher, D.R. (2011a) Biosynthesis of long-chain polyunsaturated fatty acids in marine fish: Characterization of an Elovl4-like elongase from cobia Rachycentron canadum and activation of thepathway during early life stages. Aquaculture 312, 145–153.

Monroig, Ó., Li, Y., Tocher, D.R. (2011b) Delta-8 desaturation activity varies among fatty acyl desaturases of teleost fish: high activity in delta-6 desaturases of marine species. Comp. Biochem. Physiol. 159B, 206-213.

Morais, S., Monroig, Ó., Zheng, X., Leaver, M.J., Tocher, D.R. (2009) Highly unsaturated fatty acid synthesis in Atlantic salmon: characterization of Elovl5- and Elovl2-like elongases. Mar. Biotechnol. 11, 627–639.

Morais, S., Mourente, G., Ortega, A., Tocher, J.A. and Tocher, D.R. (2011) Expression of fatty acyl desaturase and elongase genes, and evolution of DHA:EPA ratio during development of Atlantic bluefin tuna (Thunnus thynnus L.). Aquaculture 313, 129-139.

Navarro, J.C., McEvoy, L.A., Bell, M.V., Amat, F., Hontoria, F., Sargent, J.R. (1997) Effect of different dietary levels of docosahexaenoic acid (DHA, 22:6-3) on the DHA composition of lipid classes in sea bass larvae eyes. Aquacult. Internat. 5, 509-516.

Naylor, R.L., Hardy, R.W., Bureau, D.P., Chiu, A., Elliot, M., Farrell, A.P., Forster, I., Gatlin, D.M., Goldburg, R.J., Hua, K., Nichols, P.D. (2009) Feeding aquaculture in an era of finite resources. Proc. Natl. Acad. Sci. USA 106, 15103-15110.

Noble R.C., Shand J.H. (1985) Unsaturated fatty acid compositional changes and desaturation during the embryonic development of the chicken (Gallus domesticus). Lipids 20, 278–282.

Nugteren, D.H. (1965) The enzymatic chain elongation of fatty acids by rat-liver microsomes. Biochim. Biophys. Acta 106, 280-290.

Oxley, A., Jolly, C., Eide, T., Jordal, A-E.O., Svardal, A., Olsen R-E. (2010) The combined impact of plant-derived dietary ingredients and acute stress on the intestinal arachidonic acid cascade in Atlantic salmon (Salmo salar). Br. J. Nutr. 103, 851-861.

Park, W.J., Kothapalli, K.S., Lawrence, P., Tyburczy, C., Brenna, J.T. (2009) An alternate pathway to long-chain polyunsaturates: The FADS2 gene product Delta8-desaturates 20:2n-6 and 20:3n-3. J. Lipid Res. 50, 1195-1202.

Pickova, J., Dutta, P.C., Larsson, P.-O., Kiessling, A. (1997) Early embryonic cleavage pattern, hatching success, and egg-lipid fatty acid composition: comparison between two cod (Gadus morhua) stocks. Can. J. Fish. Aquat. Sci. 54, 2410-2416.

Poulos, A. (1995) Very long chain fatty acids in higher animals – A review. Lipids 30, 1-14.

Pratoomyot, J., Bendiksen, E.Å., Bell, J.G., Tocher, D.R. (2008) Comparison of effects of vegetable oils blended with southern hemisphere fish oil and decontaminated northern hemisphere fish oil on

growth performance, composition and gene expression in Atlantic salmon (Salmo salar L.). Aquaculture 280, 170–178.

Robinson, B.S., Johnson, D.W., Poulos, A. (1990) Unique molecular species of phosphatidylcholine containing very-long-chain (C24-C38) polyenoic fatty acids in rat brain. Biochem. J. 265, 763-7.

Rodríguez, C., Cejas, J.R., Martín, M.V., Badía, P., Samper, M., Lorenzo, A. (1998) Influence of n-3 highly unsaturated fatty acid deficiency on the lipid composition of broodstock gilthead seabream (Sparus aurata L.) and on egg quality. Fish Physiol. Biochem. 18, 177-187.

Ruyter, B., Røsjø, C., Grisdale-Helland, B., Rosenlund, G., Obach, A., Thomassen, M.S. (2003) Influence of temperature and high dietary linoleic acid content on esterification, elongation, and desaturation of PUFA in Atlantic salmon hepatocytes. Lipids 38, 833-840.

Salem Jr, N., Litman, B., Kim, H-Y., Gawrisch K. (2001) Mechanisms of action of docosahexaenoic acid in the nervous system. Lipids 36, 945-949.

Santigosa, E., Geay, F., Tonon, T., Le Delliou, H., Kuhl, H., Reinhardt, R., Corcos, L., Cahu, C., Zambonino-Infante, J.L., Mazurais, D., (2011) Cloning, tissue expression analysis, and functional characterization of two Δ6-desaturase variants of sea bass (Dicentrarchus labrax L.). Mar. Biothechnol. 13, 22-31.

Sargent, J.R., McEvoy, L.A., Bell, J.G. (1997) Requirements, presentation and sources of polyunsaturated fatty acids in marine fish larval feeds. Aquaculture 155, 117–127.

Sargent, J. ., cEvoy, .A., Est vez, A., Bell, G., Bell, M., Henderson, J., Tocher, D.R. (1999) Lipid nutrition of marine fish during early development: current status and future directions. Aquaculture 179, 217–229.

Sargent, J.R., Tocher, D.R., Bell, J.G. (2002) The Lipids, in: Halver, J.E., Hardy, R.W., (Eds.), Fish Nutrition, third ed., Academic Press, San Diego, 2002, pp. 181-257.

Sarker, M.A-A., Yamamoto, Y., Haga, Y., Sarker, M.S.A., Miwa, M., Yoshizaki, G., Satoh, S. (2011) Influences of low salinity and dietary fatty acids on fatty acid composition and fatty acid desaturase and elongase expression in red sea bream Pagrus major. Fish. Sci. 77, 385–396.

Sayanova, O., Haslam, R., Guschina, I., Lloyd, D., Christie, W.W., Harwood, J.L., Napier, J.A. (2006) A bifunctional �Δ12,Δ15-desaturase from Acanthamoeba castellanii directs the synthesis of highly unusual n-1 series unsaturated fatty acids. J. Biol. Chem. 281, 36533–36541.

Seiliez, I., Panserat, S., Kaushik, S., Bergot, P. (2001) Cloning, tissue distribution and nutritional regulation of a Δ6-desaturase-like enzyme in rainbow trout. Comp. Biochem. Physiol. 130B, 83–93.

Seiliez, I., Panserat, S., Corraze, G., Kaushik, S., Bergot, P. (2003) Cloning and nutritional regulation of a Δ6-desaturase-like enzyme in the marine teleost gilthead sea bream (Sparus aurata). Comp. Biochem. Physiol. 135B, 449–460.

Serra, M., Gauthier, L.T., Fabrias, G., Buist, P.H. (2006) Δ11 desaturases of Trichoplusia ni and Spodoptera littoralis exhibit dual catalytic behaviour. Insect Biochem. Mol. Biol. 36, 822–825.

Sprecher, H. (2000) Metabolism of highly unsaturated n-3 and n-6 fatty acids. Biochim. Biophys. Acta. 1486, 219-231.

Suh, M., Wierzbicki, A.A., Clandini, M.T. (2002) Dietary n-3 FA modulate long and very long chain FA content, rhodopsin content, and rhodopsin phosphorylation in rat rod outer segment after light exposure. Lipids 37, 253-60.

Tacon, A.G.J., Metian, M. (2008) Global overview on the use of fish meal and fish oil in industrially compounded aquafeeds: Trends and future prospects. Aquaculture 285, 146-158.

Taggart, J.B., Bron, J.E., Martin, S.A.M., Seear, P.J., Høyheim, B., Talbot, R., Carmichael, S.N., Villeneuve, L.A.N., Sweeney, G.E., Houlihan, D.F., Secombes, C.J., Tocher, D.R., Teale, A.J. (2008) A description of the origins, designs and performance of the TRAITS-SGP Atlantic salmon Salmo salar L. cDNA microarray. J. Fish Biol. 72, 2071-2094.

Tocher, D.R., Dick, J.R. (1999) Polyunsaturated fatty acid metabolism in a cell culture model of essential fatty acid deficiency in a freshwater fish, carp (Cyprinus carpio). Fish Physiol. Biochem. 21, 257-267.

Tocher, D.R., Dick, J.R. (2000) Essential fatty acid deficiency in freshwater fish: the effects of linoleic, α- linolenic, γ-linolenic and stearidonic acids on the metabolism of [1-14C]18:3n-3 in a carp cell culture model. Fish Physiol. Biochem. 22, 67-75.

Tocher, D.R., Bell, J.G., Dick, J.R., Sargent, J.R. (1997) Fatty acid desaturation in isolated hepatocytes from Atlantic salmon (Salmo salar): Stimulation by dietary borage oil containing γ-linolenic acid. Lipids 32, 1237-1247.

Tocher, D.R., Bell, J.G., MacGlaughlin, P., McGhee, F., Dick, J.R. (2001) Hepatocyte fatty acid desaturation and polyunsaturated fatty acid composition of liver in salmonids: Effects of dietary vegetable oil. Comp. Biochem Physiol. 130B, 257-270.

Tocher, D.R., Agaba, M., Hastings, N., Teale, A.J. (2003) Biochemical and molecular studies of the fatty acid desaturation pathway in fish. In: Browman, H.I., Skiftesvik, A.B. (Eds.), The Big Fish Bang – Proceedings of the 26th Annual Larval Fish Conference, pp. 211-227. Institute of Marine Nutrition, Bergen.

Tocher, D.R., Zheng, X., Schlechtriem, C., Hastings, N., Dick, J.R., Teale, A.J. (2006) Highly unsaturated fatty acid synthesis in marine fish: cloning, functional characterization and nutritional regulation of fatty acid Δ6 desaturase of Atlantic cod (Gadus morhua L.). Lipids 41, 1003-1016.

Tocher, D.R. (2010) Fatty acid requirements in ontogeny of marine and freshwater fish. Aquaculture Res. 41, 717-732.

Torrejon, C., Jung, U.J., Deckelbaum, R.J. (2007) n-3 Fatty acids and cardiovascular disease: Actions and molecular mechanisms. Prostaglandins Leukot. Essent. Fatty Acids 77, 319–326.

Turchini, G.M., Ng, W.-K., Tocher, D.R. (Eds.) (2010) Fish Oil Replacement and Alternative Lipid Sources in Aquaculture Feeds. Taylor & Francis, CRC Press, Boca Raton.

Worm, B., Barbier, E.B., Beaumont, N., Duffy, J.E., Folke, C., Halpern, B.S., Jackson, J.B.C., Lotze, H.K., Micheli, F., Palumbi, S.R., Sala, E., Selkoe, K.A., Stachowicz, J.J., Watson, R. (2006) Impacts of biodiversity loss on ocean ecosystem services. Science 314, 787–790.

Yamamoto, Y., Kabeya, N., Takeuchi, Y., Alimuddin, Haga, Y., Satoh, S., Takeuchi, T., Yoshizaki, G. (2010) Cloning and nutritional regulation of polyunsaturated fatty acid desaturase and elongase of a marine teleost, the nibe croaker Nibea mitsukurii. Fish. Sci. 76, 463–472.

Zhang, S., Sakuradani, E., Ito, K., Shimizu, S. (2007) Identification of a novel bifunctional Δ12/Δ15 fatty acid desaturase from a basidiomycete, Coprinus cinereus TD#822-2. FEBS Lett. 581, 315-319.

Zheng, X., Seiliez, I., Hastings, N., Tocher, D.R., Panserat, S., Dickson, C.A., Bergot, P., Teale A.J. (2004a) Characterization and comparison of fatty acyl Δ6 desaturase cDNAs from freshwater and marine teleost fish specie. Comp. Biochem. Physiol. 139B, 269-279.

Zheng, X., Tocher, D.R., Dickson, C.A., Bell, J.G., Teale, A.J. (2004b) Effects of diets containing vegetable oil on expression of genes involved in highly unsaturated fatty acid biosynthesis in liver of Atlantic salmon (Salmo salar). Aquaculture 236, 467-483.

Zheng, X., Tocher, D.R., Dickson, C.A., Dick, J.R., Bell, J.G., Teale A.J. (2005a) Highly unsaturated fatty acid synthesis in vertebrates: new insights with the cloning and characterisation of a Δ6 desaturase of Atlantic salmon. Lipids 40 13-24.

Zheng, X., Torstensen, B.E., Tocher, D.R., Dick, J.R., Henderson, R.J., Bell, J.G. (2005b) Environmental and dietary influences on highly unsaturated fatty acid biosynthesis and expression of fatty acyl desaturases and elongase genes in liver of Atlantic salmon (Salmo salar). Biochim. Biophys. Acta 1734, 13–24.

Zheng, X., Ding, Z., Xu, Y., Monroig, O., Morais, S., Tocher, D.R. (2009a) Physiological roles of fatty acyl desaturase and elongase in marine fish: Characterisation of cDNAs of fatty acyl Δ6 desaturase and Elovl5 elongase of cobia (Rachycentron canadum). Aquaculture 290, 122-131.

Zheng, X., Leaver, M.J., Tocher, D.R. (2009b) Regulation of fatty acyl desaturase (FAD) gene transcription: isolation and characterisation of Δ6 FAD gene promoters of Atlantic salmon (Salmo salar L.) and Atlantic cod (Gadus morhua L.). Comp. Biochem. Physiol. 154B, 255-263.

Downloads

How to Cite

Monroig, Óscar, Navarro, J. C., & R. Tocher, D. (2011). Long-Chain Polyunsaturated Fatty Acids in Fish: Recent Advances on Desaturases and Elongases Involved in Their Biosynthesis. Avances En Nutrición Acuicola. Retrieved from https://nutricionacuicola.uanl.mx/index.php/acu/article/view/93

Similar Articles

<< < 5 6 7 8 9 10 11 12 13 14 > >> 

You may also start an advanced similarity search for this article.