Balıklarda Sindirim Fizyolojisi

Özet

Bu bölüm, balıklarda sindirim sisteminin morfolojik ve fizyolojik çeşitliliğini kapsamlı bir şekilde incelemektedir. Balıkların beslenme alışkanlıklarına (etçil, otçul, hepçil) göre şekillenen sindirim kanalı yapıları; mideye sahip (gastrik) ve midesiz (agastrik) türler arasındaki temel farklar vurgulanarak ele alınmıştır. Özofagus, mide, intestinal bulbus, pilorik sekum ve bağırsak gibi organların yanı sıra; pankreas ve karaciğerin sindirimdeki kritik rolleri detaylandırılmıştır. Bölümde özellikle protein, lipit ve karbonhidratların enzimatik sindirimi ile bu besin maddelerinin enterositlerdeki moleküler emilim mekanizmaları (PepT1, SGLT1, CD36 gibi taşıyıcılar yardımıyla) güncel literatür ışığında açıklanmıştır. Ayrıca vitamin ve minerallerin emilim süreçleri ile çevresel sıcaklık ve rasyon içeriğinin sindirim verimliliği üzerindeki adaptif etkileri tartışılmaktadır. Bu çalışma, balık besleme stratejilerinin geliştirilmesi ve metabolik süreçlerin anlaşılması için temel bir fizyolojik perspektif sunmaktadır.

 

This chapter provides a comprehensive review of the morphological and physiological diversity of the digestive system in fish. It explores the structures of the alimentary canal shaped by various feeding habits (carnivorous, herbivorous, omnivorous), highlighting the fundamental distinctions between gastric and agastric species. The critical roles of organs such as the esophagus, stomach, intestinal bulb, pyloric caeca, and intestine, alongside accessory organs like the pancreas and liver, are detailed. The chapter specifically explains the enzymatic digestion of proteins, lipids, and carbohydrates, as well as the molecular absorption mechanisms of nutrients (e.g., transporters like PepT1, SGLT1, and CD36) in light of current scientific literature. Furthermore, the absorption processes of vitamins and minerals, and the adaptive impacts of environmental temperature and dietary composition on digestive efficiency are discussed. This study offers an in-depth perspective on the fundamental physiological processes essential for optimizing fish nutrition strategies and understanding metabolic pathways.

Referanslar

Le, H.T., et al., Intestinal function of the stomachless fish, ballan wrasse (Labrus bergylta). Frontiers in Marine Science, 2019. 6: p. 140.

LS, S., Digestive functions in teleost fishes. Fish nutrition, 1989: p. 331–421.

Shalaby, W., Comparative morphological and histological studies on the adaptation of esophagus and stomach to the feeding habits in some coral reef fishes at Hurghada, Red Sea, Egypt. Egyptian Journal of Aquatic Biology and Fisheries, 2020. 24(5): p. 289–306.

Palladino, A., et al., A morphological and ultrastructural study of the anterior digestive tract of adult nile tilapia oreochromis niloticus. Animals, 2023. 13(3): p. 420.

Breves, J.P., et al., Transcriptional regulation of esophageal, intestinal, and branchial solute transporters by salinity, growth hormone, and cortisol in Atlantic salmon. Journal of Experimental Zoology Part A: Ecological and Integrative Physiology, 2024. 341(1): p. 107–117.

Richardson, S., et al., Esophageal expression of Na+/H+ exchanger 2 and-3 (nhe2 and-3) gene transcripts in Atlantic salmon during smoltification and seawater acclimation. Physiology, 2023. 38(S1): p. 5731352.

Bakke, A.M., C. Glover, and Å. Krogdahl, Feeding, digestion and absorption of nutrients, in Fish physiology. 2010, Elsevier. p. 57–110.

Alves, L.C.M., et al., Morphological, Histological and Histochemical Analysis of the Digestive Tract in Panga Pangasius hypophthalmus (Teleostei: Siluriformes). Anatomia, Histologia, Embryologia, 2025. 54(2): p. e70026.

Mokhtar, D.M., M.M. Hussein, and R.K. Sayed, Novel identification and microscopy of the intestinal bulb of molly fish (Poecilia sphenops) with a focus on its role in immunity. Microscopy and Microanalysis, 2022. 28(5): p. 1827–1839.

Mokhtar, D.M., E.A. Abd-Elhafez, and A.H. Hassan, Microanalysis of the intestinal bulb of grass carp (Ctenopharyngodon idella): Histological, histochemical, immunohistochemical, and scanning electron microscopical studies. Microscopy and Microanalysis, 2021. 27(6): p. 1564–1572.

Purushothaman, K., et al., Morpho-histological characterisation of the alimentary canal of an important food fish, Asian seabass (Lates calcarifer). PeerJ, 2016. 4: p. e2377.

Buddington, R.K. and J.M. Diamond, Pyloric ceca of fish: a" new" absorptive organ. American Journal of Physiology-Gastrointestinal and Liver Physiology, 1987. 252(1): p. G65–G76.

Tlak Gajger, I., et al., Histochemical Analysis and Distribution of Digestive Enzymes in the Gastrointestinal System of the European Barracuda Sphyraena sphyraena (Linnaeus, 1758). Animals, 2024. 14(19): p. 2798.

Cho, J.-H., et al., Morphology, histology, and histochemistry of the digestive tract of the marbled flounder Pseudopleuronectes yokohamae. Animals, 2023. 13(5): p. 936.

Denstadli, V., et al., Lipid absorption in different segments of the gastrointestinal tract of Atlantic salmon (Salmo salar L.). Aquaculture, 2004. 240(1-4): p. 385–398.

Veillette, P.A., et al., Osmoregulatory physiology of pyloric ceca: regulated and adaptive changes in chinook salmon. Journal of Experimental Zoology Part A: Comparative Experimental Biology, 2005. 303(7): p. 608–613.

Ballesteros, N.A., et al., The pyloric caeca area is a major site for IgM+ and IgT+ B cell recruitment in response to oral vaccination in rainbow trout. PLoS One, 2013. 8(6): p. e66118.

Jiao, F., et al., A comparison of digestive strategies for fishes with different feeding habits: Digestive enzyme activities, intestinal morphology, and gut microbiota. Ecology and Evolution, 2023. 13(9): p. e10499.

Liu, Y., G. He, and H. Zhou, Substitution system of animal and plant protein source in aquatic feed and its relationship with feeding habits of cultured fish. Hebei Fisheries, 2014. 8: p. 54–57.

Bryan, P.G., Food habits, functional digestive morphology, and assimilation efficiency of the rabbitfish Siganus spinus (Pisces, Siganidae) on Guam. 1975.

Banan Khojasteh, S.M., The morphology of the post-gastric alimentary canal in teleost fishes: a brief review. Int. J. of Aquatic Science, 2012. 3(2): p. 71–88.

Day, R.D., I.R. Tibbetts, and S.M. Secor, Physiological responses to short-term fasting among herbivorous, omnivorous, and carnivorous fishes. Journal of Comparative Physiology B, 2014. 184(4): p. 497–512.

Genten, F., E. Trewinghe, and A. Danguy, Digestive system. Atlas of Fish Histology, Science Publishers, 2009: p. 75–91.

Guillaume, J., Nutrition and feeding of fish and crustaceans. 2001: Springer Science & Business Media.

Hardy, R.W. and S.J. Kaushik, Fish nutrition. 2021: Academic press.

Verdile, N., et al., A detailed study of rainbow trout (Onchorhynchus mykiss) intestine revealed that digestive and absorptive functions are not linearly distributed along its length. Animals, 2020. 10(4): p. 745.

Qi, Z., et al., The Structure of Digestive Tract Coordinating Digestion and Respiration in an Air-Breathing Weatherloach, Misgurnus anguillicaudatus. Biology, 2024. 13(6): p. 381.

Ray, A.K. and E. Ringø, The gastrointestinal tract of fish. Aquaculture nutrition: Gut health, probiotics and prebiotics, 2014: p. 1–13.

GHOSH, S.K. and P. CHAKRABARTI, Comparative studies on histology and histochemistry of pancreas between Labeo calbasu (Hamilton, 1822) and Mystus gulio (Hamilton, 1822). Iranian Journal of Ichthyology, 2016. 3(4): p. 251–265.

Geyer, H., M.M. Nel, and J. Swanepoel, Histology and ultrastructure of the hepatopancreas of the tigerfish, Hydrocynus forskahlii. Journal of Morphology, 1996. 227(1): p. 93–100.

Senoo, H., The Laboratory Rat-Handbook of Experimental Animals by George J. 2000, Krinke Academic Press.

Hussein, M.M., R.K. Sayed, and D.M. Mokhtar, Structural and immunohistochemical characterization of pancreas of Molly fish (Poecilia sphenops), with a special reference to its immune role. Microscopy Research and Technique, 2023. 86(12): p. 1667–1680.

Ichlasul Akmal, A.M., S.R. Laila, and T. Wresdiyati, HISTOMORPHOLOGY OF PANCREATIC TISSUE IN MEDAKA FISH (Oryzias javanicus): STUDY OF DIABETIC ANIMAL MODEL DEVELOPMENT. Indonesian Journal of Veterinary Science/Jurnal Kedokteran Hewan, 2023. 17(1).

Cheng, C.-H.C., P.A. Cziko, and C.W. Evans, Nonhepatic origin of notothenioid antifreeze reveals pancreatic synthesis as common mechanism in polar fish freezing avoidance. Proceedings of the National Academy of Sciences, 2006. 103(27): p. 10491–10496.

Rocha, E. and R.A. Monteiro, Histology and cytology of fish liver: a review. 1999.

Попова, О. and Л. Агафонова, Особенности метаболизма желчных кислот у рыб. Международный вестник ветеринарии, 2022(1): p. 61–65.

Hofer, R., The adaptation of digestive enzymes to temperature, season and diet in roach, Rutilus rutilus and rudd Scardinius erythrophthalmus; Proteases. Journal of Fish Biology, 1979. 15(4): p. 373–379.

Li, H., et al., Comparison of growth, digestive enzyme activity, immune response and intestinal microbiota in large yellow croaker (Larimichthys crocea) with different sizes. Aquaculture Reports, 2024. 36: p. 102059.

Bone, Q. and R. Moore, Biology of fishes. 2008: Taylor & Francis.

Karasov, W.H., Tests of the adaptive modulation hypothesis for dietary control of intestinal nutrient transport. American Journal of Physiology-Regulatory, Integrative and Comparative Physiology, 1992. 263(3): p. R496–R502.

Hani, Y.M.I., et al., Digestive enzymes and gut morphometric parameters of threespine stickleback (Gasterosteus aculeatus): Influence of body size and temperature. PLoS One, 2018. 13(4): p. e0194932.

Bowyer, J.N., et al., Temperature and dissolved oxygen influence growth and digestive enzyme activities of yellowtail kingfish S eriola lalandi (V alenciennes, 1833). Aquaculture Research, 2014. 45(12): p. 2010–2020.

Kofuji, P.Y.M., et al., Seasonal changes in proteolytic enzymes of yellowtail Seriola quinqueradiata (Temminck & Schlegel; Carangidae) fed extruded diets containing different protein and energy levels. Aquaculture Research, 2005. 36(7): p. 696–703.

Martinez-Llorens, S., et al., Digestive tract morphology and enzyme activities of juvenile diploid and triploid Atlantic salmon (Salmo salar) fed fishmeal-based diets with or without fish protein hydrolysates. PloS one, 2021. 16(1): p. e0245216.

Dıaz-López, M., et al., Characterization of fish acid proteases by substrate–gel electrophoresis. Comparative Biochemistry and Physiology Part B: Biochemistry and Molecular Biology, 1998. 121(4): p. 369–377.

Torrissen, K.R. and R. Male, Trypsin isozymes: Development, digestion, and structure. FOOD SCIENCE AND TECHNOLOGY-NEW YORK-MARCEL DEKKER-, 2000: p. 215–270.

Day, R.D., et al., Enzymatic digestion in stomachless fishes: how a simple gut accommodates both herbivory and carnivory. Journal of Comparative Physiology B, 2011. 181(5): p. 603–613.

García-Meilán, I., et al., Effects of dietary protein-to-lipid ratio on digestive and absorptive processes in sea bass fingerlings. Aquaculture, 2016. 463: p. 163–173.

Heu, M., H. Kim, and J. Pyeun, Comparison of trypsin and chymotrypsin from the viscera of anchovy, Engraulis japonica. Comparative Biochemistry and Physiology Part B: Biochemistry and Molecular Biology, 1995. 112(3): p. 557–567.

Natale, S., et al., First insight into temporal variation of digestive enzyme activities in flathead grey mullet (Mugil cephalus) during the ongrowing phase. Aquaculture Reports, 2025. 41: p. 102652.

Sheng, Z., et al., Functional properties of protein hydrolysates on growth, digestive enzyme activities, protein metabolism, and intestinal health of larval largemouth bass (Micropterus salmoides). Frontiers in Immunology, 2022. 13: p. 913024.

Debnath, S. and S.K. Saikia, Absorption of protein in teleosts: a review. Fish Physiology and Biochemistry, 2021. 47(2): p. 313–326.

Krogdahl, Å. and A.M. Bakke-McKellep, Fasting and refeeding cause rapid changes in intestinal tissue mass and digestive enzyme capacities of Atlantic salmon (Salmo salar L.). Comparative Biochemistry and Physiology Part A: Molecular & Integrative Physiology, 2005. 141(4): p. 450–460.

Verri, T., et al., Transport of di‐and tripeptides in teleost fish intestine. Aquaculture Research, 2010. 41(5): p. 641–653.

Storelli, C., et al., Brush-border amino acid transport mechanisms in carnivorous eel intestine. American Journal of Physiology-Regulatory, Integrative and Comparative Physiology, 1989. 257(3): p. R506–R510.

Kaushik, S.J. and I. Seiliez, Protein and amino acid nutrition and metabolism in fish: current knowledge and future needs. Aquaculture research, 2010. 41(3): p. 322–332.

Christensen, H.N., Role of amino acid transport and countertransport in nutrition and metabolism. Physiological reviews, 1990. 70(1): p. 43–77.

Orozco, Z.G.A., et al., Spatial mRNA expression and response to fasting and refeeding of neutral amino acid transporters slc6a18 and slc6a19a in the intestinal epithelium of Mozambique tilapia. Frontiers in physiology, 2018. 9: p. 212.

To, V.P., K. Masagounder, and M.E. Loewen, SLC transporters ASCT2, B0AT1‐like, y+ LAT1, and LAT4‐like associate with methionine electrogenic and radio‐isotope flux kinetics in rainbow trout intestine. Physiological Reports, 2019. 7(21): p. e14274.

Vacca, F., et al., The teleost fish PepT1-type peptide transporters and their relationships with neutral and charged substrates. Frontiers in Physiology, 2023. 14: p. 1186475.

Zhang, X., et al., Temperature regulated nutrient sensing and metabolism of amino acids in juvenile turbot (Scophthalmus maximus L.). Marine Life Science & Technology, 2025: p. 1–14.

Kanai, Y. and M.A. Hediger, The glutamate/neutral amino acid transporter family SLC1: molecular, physiological and pharmacological aspects. Pflügers Archiv, 2004. 447(5): p. 469–479.

Comesaña, S., et al., Amino acid carriers of the solute carrier families 7 (SLC7) and 38 (SLC38) are involved in leucine sensing in the brain of Atlantic salmon (Salmo salar). Frontiers in Marine Science, 2021. 8: p. 711508.

Verrey, F., System L: heteromeric exchangers of large, neutral amino acids involved in directional transport. Pflügers Archiv, 2003. 445(5): p. 529–533.

Broer, S., Amino acid transport across mammalian intestinal and renal epithelia. Physiological reviews, 2008. 88(1): p. 249–286.

Yang, J., et al., Cloning and molecular characterization of cationic amino acid transporter y+ LAT1 in grass carp (Ctenopharyngodon idellus). Fish physiology and biochemistry, 2014. 40(1): p. 93–104.

Hyde, R., P.M. Taylor, and H.S. Hundal, Amino acid transporters: roles in amino acid sensing and signalling in animal cells. Biochemical Journal, 2003. 373(1): p. 1–18.

Tocher, D.R., Metabolism and functions of lipids and fatty acids in teleost fish. Reviews in fisheries science, 2003. 11(2): p. 107–184.

Kurtovic, I., et al., Lipases from mammals and fishes. Reviews in fisheries science, 2009. 17(1): p. 18–40.

Nolasco-Soria, H., et al., Optimization of Classical Lipase Activity Assays for Fish Digestive Tract Samples. Fishes, 2024. 9(7): p. 261.

Bolsoni-Lopes, A. and M.I.C. Alonso-Vale, Lipolysis and lipases in white adipose tissue–An update. Archives of endocrinology and metabolism, 2015. 59: p. 335–342.

Lafontan, M. and D. Langin, Lipolysis and lipid mobilization in human adipose tissue. Progress in lipid research, 2009. 48(5): p. 275–297.

Sun, J., et al., Lipolytic enzymes involving lipolysis in Teleost: Synteny, structure, tissue distribution, and expression in grass carp (Ctenopharyngodon idella). Comparative Biochemistry and Physiology Part B: Biochemistry and Molecular Biology, 2016. 198: p. 110–118.

Morais, S., et al., Dietary neutral lipid level and source in marine fish larvae: effects on digestive physiology and food intake. Aquaculture, 2007. 268(1-4): p. 106–122.

Tocher, D.R., Fatty acid requirements in ontogeny of marine and freshwater fish. Aquaculture research, 2010. 41(5): p. 717–732.

Sheridan, M.A., Lipid dynamics in fish: aspects of absorption, transportation, deposition and mobilization. Comparative Biochemistry and Physiology Part B: Comparative Biochemistry, 1988. 90(4): p. 679–690.

Eslamloo, K., et al., Transcriptome profiling of antiviral immune and dietary fatty acid dependent responses of Atlantic salmon macrophage-like cells. BMC genomics, 2017. 18(1): p. 706.

Agulleiro, M.J., et al., High transcript level of fatty acid-binding protein 11 but not of very low-density lipoprotein receptor is correlated to ovarian follicle atresia in a teleost fish (Solea senegalensis). Biology of reproduction, 2007. 77(3): p. 504–516.

Wang, Y., et al., Effects of nutritional status and diet composition on fatty acid transporters expression in zebrafish (Danio rerio). Aquaculture Research, 2019. 50(3): p. 904–914.

Esteves, A., et al., Fatty acid binding proteins have the potential to channel dietary fatty acids into enterocyte nuclei [S]. Journal of lipid research, 2016. 57(2): p. 219–232.

Selvam, C., et al., Intracellular trafficking of fatty acids in the fish intestinal epithelial cell line RTgutGC. Frontiers in Marine Science, 2022. 9: p. 954773.

Oxley, A., et al., Enzyme activities of intestinal triacylglycerol and phosphatidylcholine biosynthesis in Atlantic salmon (Salmo salar L.). Comparative Biochemistry and Physiology Part B: Biochemistry and Molecular Biology, 2005. 141(1): p. 77–87.

Enes, P., et al., Nutritional regulation of hepatic glucose metabolism in fish. Fish physiology and biochemistry, 2009. 35(3): p. 519–539.

Krogdahl, Å., G.I. Hemre, and T. Mommsen, Carbohydrates in fish nutrition: digestion and absorption in postlarval stages. Aquaculture nutrition, 2005. 11(2): p. 103–122.

Spiro, R.G., Protein glycosylation: nature, distribution, enzymatic formation, and disease implications of glycopeptide bonds. Glycobiology, 2002. 12(4): p. 43R–56R.

Kuz’mina, V.V., Classical and modern concepts in fish digestion. Feeding and digestive functions of fishes, 2008: p. 85–154.

Goel, K., Carbohydrase activity in the digestive system of some teleost fishes. Acta Physiologica Academiae Scientiarum Hungaricae, 1975. 46(3): p. 191–196.

Agrawal, V., K. Sastry, and S. Kaushab, Digestive enzymes of three teleost fishes. Acta Physiologica Academiae Scientiarum Hungaricae, 1975. 46(2): p. 93–98.

Weinrauch, A.M., C.M. Schaefer, and G.G. Goss, Activity and post-prandial regulation of digestive enzyme activity along the Pacific hagfish (Eptatretus stoutii) alimentary canal. PLoS One, 2019. 14(4): p. e0215027.

Steinmann, K., et al., How They Eat: An Examination of Digestive Physiology of Sympatric Prickleback Fishes. Physiology, 2024. 39(S1): p. 2151.

Sari, D.N., et al., High carbohydrate increases amylase, plasma glucose, and gene expression related to glycolysis in giant gourami Osphronemus goramy. Fish Physiology and Biochemistry, 2022. 48(6): p. 1495–1505.

Kuz'Mina, V., Influence of age on digestive enzyme activity in some freshwater teleosts. Aquaculture, 1996. 148(1): p. 25–37.

Choct, M., Feed non-starch polysaccharides: chemical structures and nutritional significance. Feed milling international, 1997. 191(1): p. 13–26.

Stone, D.A., Dietary carbohydrate utilization by fish. Reviews in fisheries Science, 2003. 11(4): p. 337–369.

Sinha, A.K., et al., Non-starch polysaccharides and their role in fish nutrition–A review. Food Chemistry, 2011. 127(4): p. 1409–1426.

Maas, R.M., et al., Carbohydrate utilisation by tilapia: a meta‐analytical approach. Reviews in Aquaculture, 2020. 12(3): p. 1851–1866.

Das, K. and S. Tripathi, Studies on the digestive enzymes of grass carp, Ctenopharyngodon idella (Val.). Aquaculture, 1991. 92: p. 21–32.

German, D.P. and R.A. Bittong, Digestive enzyme activities and gastrointestinal fermentation in wood-eating catfishes. Journal of Comparative Physiology B, 2009. 179(8): p. 1025–1042.

Logothetis, E., M. Horn, and K. Dickson, Gut morphology and function in Atherinops affinis (Teleostei: Atherinopsidae), a stomachless omnivore feeding on macroalgae. Journal of Fish Biology, 2001. 59(5): p. 1298–1312.

Crossman, D.J., J.H. Choat, and K.D. Clements, Nutritional ecology of nominally herbivorous fishes on coral reefs. Marine Ecology Progress Series, 2005. 296: p. 129–142.

Horn, M., et al., Structure and function of the stomachless digestive system in three related species of New World silverside fishes (Atherinopsidae) representing herbivory, omnivory, and carnivory. Marine Biology, 2006. 149(5): p. 1237–1245.

German, D.P., Do herbivorous minnows have “plug-flow reactor” guts? Evidence from digestive enzyme activities, gastrointestinal fermentation, and luminal nutrient concentrations. Journal of Comparative Physiology B, 2009. 179(6): p. 759–771.

Manjakasy, J.M., et al., Functional morphology of digestion in the stomachless, piscivorous needlefishes Tylosurus gavialoides and Strongylura leiura ferox (Teleostei: Beloniformes). Journal of Morphology, 2009. 270(10): p. 1155–1165.

Solovyev, M., E. Kashinskaya, and E. Gisbert, A meta-analysis for assessing the contributions of trypsin and chymotrypsin as the two major endoproteases in protein hydrolysis in fish intestine. Comparative Biochemistry and Physiology Part A: Molecular & Integrative Physiology, 2023. 278: p. 111372.

Wilson, R.á., Utilization of dietary carbohydrate by fish. Aquaculture, 1994. 124(1-4): p. 67–80.

Kamalam, B.S., F. Medale, and S. Panserat, Utilisation of dietary carbohydrates in farmed fishes: new insights on influencing factors, biological limitations and future strategies. Aquaculture, 2017. 467: p. 3–27.

Polakof, S., et al., Glucose metabolism in fish: a review. Journal of Comparative Physiology B, 2012. 182(8): p. 1015–1045.

Zhang, Y., et al., A comparative genomics study of carbohydrate/glucose metabolic genes: from fish to mammals. BMC genomics, 2018. 19(1): p. 246.

Blanco, A.M., et al., Ghrelin facilitates GLUT2-, SGLT1-and SGLT2-mediated intestinal glucose transport in goldfish (Carassius auratus). Scientific Reports, 2017. 7(1): p. 45024.

Liang, H., et al., Molecular characterization and identification of facilitative glucose transporter 2 (GLUT2) and its expression and of the related glycometabolism enzymes in response to different starch levels in blunt snout bream (Megalobrama amblycephala). Fish physiology and biochemistry, 2018. 44(3): p. 869–883.

Karasov, W.H. and C. Martínez del Rio, Physiological ecology: how animals process energy, nutrients, and toxins. 2020.

Sitjà-Bobadilla, A., et al., Disruption of gut integrity and permeability contributes to enteritis in a fish-parasite model: a story told from serum metabolomics. Parasites & vectors, 2019. 12(1): p. 486.

Zhang, Y., et al., Effects of high carbohydrate diet-modulated microbiota on gut health in Chinese perch. Frontiers in Microbiology, 2020. 11: p. 575102.

Zhou, Y.-L., et al., High dietary starch impairs intestinal health and microbiota of largemouth bass, Micropterus salmoides. Aquaculture, 2021. 534: p. 736261.

Soengas, J.L., et al., Glucokinase and hexokinase expression and activities in rainbow trout tissues: changes with food deprivation and refeeding. American Journal of Physiology-Regulatory, Integrative and Comparative Physiology, 2006. 291(3): p. R810–R821.

Dholariya, S.J. and J.A. Orrick, Biochemistry, fructose metabolism. 2022.

Haskovic, M., et al., Nucleotide sugar profiles throughout development in wildtype and galt knockout zebrafish. Journal of Inherited Metabolic Disease, 2020. 43(5): p. 994–1001.

Peña-Llopis, S., M.D. Ferrando, and J.B. Peña, Fish tolerance to organophosphate-induced oxidative stress is dependent on the glutathione metabolism and enhanced by N-acetylcysteine. Aquatic Toxicology, 2003. 65(4): p. 337–360.

Panserat, S., E. Plagnes-Juan, and S. Kaushik, Nutritional regulation and tissue specificity of gene expression for proteins involved in hepatic glucose metabolism in rainbow trout (Oncorhynchus mykiss). Journal of Experimental Biology, 2001. 204(13): p. 2351–2360.

Panserat, S., et al., Hepatic glucokinase is induced by dietary carbohydrates in rainbow trout, gilthead seabream, and common carp. American Journal of Physiology-Regulatory, Integrative and Comparative Physiology, 2000. 278(5): p. R1164–R1170.

Council, N.R., et al., Nutrient requirements of fish and shrimp. 2011: National academies press.

Bjerkeng, B., Carotenoid pigmentation of salmonid fishes-recent progress1. Avances en Nutrición Acuicola, 2000.

During, A. and E.H. Harrison, Mechanisms of provitamin A (carotenoid) and vitamin A (retinol) transport into and out of intestinal Caco-2 cells. Journal of lipid research, 2007. 48(10): p. 2283–2294.

Ross, A.C., Vitamin A and retinoic acid in T cell–related immunity. The American journal of clinical nutrition, 2012. 96(5): p. 1166S–1172S.

Shastak, Y. and W. Pelletier, Vitamin A in fish well-being: integrating immune strength, antioxidant capacity and growth. Fishes, 2024. 9(8): p. 330.

Fernández, I., et al., Fat-soluble vitamins in fish: A transcriptional tissue-specific crosstalk that remains to be unveiled and characterized, in Emerging issues in fish larvae research. 2018, Springer. p. 159–208.

Buddington, R.K., et al., A high retinol dietary intake increases its apical absorption by the proximal small intestine of juvenile sunshine bass (Morone chrysops× M. saxatilis). The Journal of nutrition, 2002. 132(9): p. 2713–2716.

Ndayibagira, A. and P. Spear, Esterification and hydrolysis of vitamin A in the liver of brook trout (Salvelinus fontinalis) and the influence of a coplanar polychlorinated biphenyl. Comparative Biochemistry and Physiology Part C: Pharmacology, Toxicology and Endocrinology, 1999. 122(3): p. 317–325.

Reboul, E., et al., Vitamin D intestinal absorption is not a simple passive diffusion: evidences for involvement of cholesterol transporters. Molecular nutrition & food research, 2011. 55(5): p. 691–702.

Bouillon, R., et al., Skeletal and extraskeletal actions of vitamin D: current evidence and outstanding questions. Endocrine reviews, 2019. 40(4): p. 1109–1151.

Lock, E.J., et al., The significance of vitamin D for fish: a review. Aquaculture nutrition, 2010. 16(1): p. 100–116.

Lall, S.P. and L.M. Lewis-McCrea, Role of nutrients in skeletal metabolism and pathology in fish—An overview. Aquaculture, 2007. 267(1-4): p. 3–19.

Lie, Ø., A. Sandvin, and R. Waagbø, Transport of alpha-tocopherol in Atlantic salmon (Salmo salar) during vitellogenesis. Fish Physiology and Biochemistry, 1994. 13(3): p. 241–247.

Parazo, M.P., et al., Distribution of α‐and γ‐tocopherols in Atlantic salmon (Salmo salar) tissues. Lipids, 1998. 33(7): p. 697–704.

Hamre, K., Metabolism, interactions, requirements and functions of vitamin E in fish. Aquaculture nutrition, 2011. 17(1): p. 98–115.

Miller, G.W., et al., The α-tocopherol transfer protein is essential for vertebrate embryogenesis. 2012.

Krossøy, C., R. Waagbø, and R. Ørnsrud, Vitamin K in fish nutrition. Aquaculture Nutrition, 2011. 17(6): p. 585–594.

Udagawa, M., Physiological role of vitamin K in fish-review. 2000.

Park, J., et al., Vitamin K variations in conger eel (Conger myriaster) influenced by harvest time and size. Food Science and Preservation, 2023. 30(6): p. 929–943.

Udagawa, M. and T. Murai, Content of phylloquinone and menaquinone in the tissues of mummichog Fundulus heteroclitus fed diets containing different forms of vitamin K. Journal of nutritional science and vitaminology, 2001. 47(2): p. 91–95.

Beato, S., F.J. Toledo-Solís, and I. Fernández, Vitamin K in vertebrates’ reproduction: Further puzzling pieces of evidence from teleost fish species. Biomolecules, 2020. 10(9): p. 1303.

Abdelhamid, A.F., et al., Impacts of different levels of vitamin K on the growth performance, hematological parameters, and immunological response of juvenile Nile tilapia (Oreochromis niloticus). Aquaculture International, 2024. 32(1): p. 477–488.

Lin, M.F. and S.Y. Shiau, Requirements of vitamin C (l‐ascorbyl‐2‐sulphate and l‐ascorbyl‐2‐polyphosphate) and its effects on non‐specific immune responses of grouper, Epinephelus malabaricus. Aquaculture Nutrition, 2005. 11(3): p. 183–189.

Bae, J.-Y., et al., Re-evaluation of the optimum dietary vitamin C requirement in juvenile eel, Anguilla japonica by using L-ascorbyl-2-monophosphate. Asian-Australasian journal of animal sciences, 2012. 25(1): p. 98.

Cai, Q., et al., Dietary vitamin C affects growth, antioxidant status and serum immune parameter of juvenile hybrid grouper (Epinephelus fuscoguttatus♀× Epinephelus lanceolatus♂) fed low fishmeal diets. Aquaculture, 2022. 556: p. 738285.

Brown, S.B., et al., Implications of thiamine deficiency in Great Lakes salmonines. Journal of Aquatic Animal Health, 2005. 17(1): p. 113–124.

Lall, S.P. and S.J. Kaushik, Nutrition and metabolism of minerals in fish. Animals, 2021. 11(09): p. 2711.

Tacon, A.G., The nutrition and feeding of farmed fish and shrimp; a training manual. 1: The essential nutrients. 1987.

Bury, N.R., P.A. Walker, and C.N. Glover, Nutritive metal uptake in teleost fish. Journal of experimental biology, 2003. 206(1): p. 11–23.

Bury, N., et al., Intestinal iron uptake in the European flounder (Platichthys flesus). Journal of Experimental Biology, 2001. 204(21): p. 3779–3787.

Kwong, R.W. and S. Niyogi, The interactions of iron with other divalent metals in the intestinal tract of a freshwater teleost, rainbow trout (Oncorhynchus mykiss). Comparative Biochemistry and Physiology Part C: Toxicology & Pharmacology, 2009. 150(4): p. 442–449.

Jiang, Y., et al., Genome wide identification, phylogeny and expression of zinc transporter genes in common carp. PLoS One, 2014. 9(12): p. e116043.

Chen, G.-H., et al., Dietary zinc addition influenced zinc and lipid deposition in the fore-and mid-intestine of juvenile yellow catfish Pelteobagrus fulvidraco. British Journal of Nutrition, 2017. 118(8): p. 570–579.

Jeong, J. and D.J. Eide, The SLC39 family of zinc transporters. Molecular aspects of medicine, 2013. 34(2-3): p. 612–619.

Gelecek

19 Şubat 2026

Lisans

Lisans