Russian Journal of Biological Physics and Chemisrty Russian Journal of Biological Physics and Chemisrty АКТУАЛЬНЫЕ ВОПРОСЫ БИОЛОГИЧЕСКОЙ ФИЗИКИ И ХИМИИ 2499-9962 54563 ЭКОЛОГИЧЕСКАЯ БИОФИЗИКА ECOLOGICAL BIOPHYSICS ЭКОЛОГИЧЕСКАЯ БИОФИЗИКА Skeletal muscles of marine fish and molecular oxygen utilization systems (brief overview) Скелетные мышцы морских рыб и молекулярные системы утилизации кислорода (краткий обзор) Солдатов А А Soldatov A A alekssoldatov@yandex.ru Парфенова И А Parfenova I A ФИЦ "Институт биологии южных морей им. А.О. Ковалевского РАН"; Севастопольский государственный университет ru A.O. Kovalevsky Institute of Biology of Southern Seas, Russian Academy of Sciences; Sevastopol State University ru Крымский федеральный университет им. В.И. Вернадского ru V.I. Vernadsky Crimean Federal University ru 25 09 2020 25 09 2020 5 3 524 535 20 09 2020 20 09 2020 https://rusjbpc.ru/en/nauka/article/54563/view

Обобщена информация об особенностях клеточного состава и молекулярных системах утилизации кислорода в скелетных мышцах костистых рыб. Рассмотрено их состояние в условиях нормоксии и гипоксии. Отмечено, что белые мышечные волокна имеют нескомпенсированный тип организации дыхательной цепи митохондрий с явным преобладанием содержания цитохромов группы аа3 . При этом мышечная ткань имеет эффективные ферментативные системы, способные нейтрализовать образующийся при острой гипоксии и аноксии токсический лактат, переводя его в этанол, аланин, СО2 , глюкозу, жиры. Показано, что функционирование цикла Кребса в условиях анаэробиоза, обеспечивает дополнительный ресинтез АТР на основе углеводных и белковых субстратов, ключевым элементом которого является высоко активный малик-фермент. Делается вывод о том, что скелетные мышцы и ряд других тканей у рыб изначально ориентированы на функционирование в условиях острого дефицита кислорода, а гипоксические состояния для них скорее являются нормой, чем исключением.

The information on the characteristics of the cellular composition and molecular oxygen utilization systems in skeletal muscles of bony fish is summarized. Their state under conditions of normoxia and hypoxia is considered. It has been noted that white muscle fibers possess an uncompensated type of organization of the respiratory chain of mitochondria with a pronounced predominance in the content of cytochromes of the aa3 group. At the same time, muscle tissue has effective enzymatic systems that can neutralize toxic lactate formed during acute hypoxia and anoxia, converting it to ethanol, alanine, CO2 , glucose and lipids. It has been shown that the functioning of the Krebs cycle under conditions of anaerobiosis provides additional ATP resynthesis based on carbohydrate and protein substrates, with a highly active malik-enzyme as the key element. It is concluded that the skeletal muscles and a number of other tissues in fish have been initially oriented towards functioning under conditions of acute oxygen deficiency, and for these organisms hypoxic conditions seem to be normal and not extraordinary ones.

 скелетные мышцы костистые рыбы метаболизм дыхательная цепь митохондрий гипоксия skeletal muscles marine fish metabolism the respiratory chain of mitochondria hypoxia Работа выполнена в рамках госзадания (№ гос. регистрации АААА-А18-118021490093-4) и при частичной поддержке проекта РФФИ (№ 20-44-920001).

Nilsson G.E., Renshaw G.M.C. Hypoxic survival strategies in two fishes: extreme anoxia tolerance in the North European crucian carp and natural hypoxic preconditioning in a coral-reef shark. J. Exp. Biol., 2004, vol. 207, pp. 3131-3139. 04. Nilsson G.E., Renshaw G.M.C. Hypoxic survival strategies in two fishes: extreme anoxia tolerance in the North European crucian carp and natural hypoxic preconditioning in a coral-reef shark. J. Exp. Biol., 2004, vol. 207, pp. 3131-3139. 04. Joyce S. The dead zones: oxygen-starved coastal waters. Environ. Health Persp., 2000, vol. 108, no. 3, pp. A120-A125. Joyce S. The dead zones: oxygen-starved coastal waters. Environ. Health Persp., 2000, vol. 108, no. 3, pp. A120-A125. El Albani A., Bengtson S., Canfield D.E., Bekker A., Macchiarelli R., Mazurier A., Meunier A. Large colonial organisms with coordinated growth in oxygenated environments 2.1 Gyr ago. Nature, 2010, vol. 466, no. 7302, pp. 100-104. El Albani A., Bengtson S., Canfield D.E., Bekker A., Macchiarelli R., Mazurier A., Meunier A. Large colonial organisms with coordinated growth in oxygenated environments 2.1 Gyr ago. Nature, 2010, vol. 466, no. 7302, pp. 100-104. Danovaro R., Dell'Anno A., Pusceddu A., Gambi C., Heiner I., Kristensen R.M. The first metazoa living in permanently anoxic conditions. BMC Biol., 2010, vol. 8, no. 1, p. 30. Danovaro R., Dell'Anno A., Pusceddu A., Gambi C., Heiner I., Kristensen R.M. The first metazoa living in permanently anoxic conditions. BMC Biol., 2010, vol. 8, no. 1, p. 30. Soldatov A.A., Andreenko T.I., Sysoeva I.V., Sysoev A.A. Tissue specificity of metabolism in the bivalve mollusc Anadara inaequivalvis Br. under conditions of experimental anoxia. J. Evolutionary Biochem. Physiol., 2009, vol. 45, no. 3, pp. 349-355. Soldatov A.A., Andreenko T.I., Sysoeva I.V., Sysoev A.A. Tissue specificity of metabolism in the bivalve mollusc Anadara inaequivalvis Br. under conditions of experimental anoxia. J. Evolutionary Biochem. Physiol., 2009, vol. 45, no. 3, pp. 349-355. Soldatov A.A., Andreeva A.Yu., Novitskaya V.N., Parfenova I.A. Coupling of membrane and metabolic function in nucleated erythrocytes of Scorpaena porcus L. under hypoxia in vivo and in vitro. J. Evolutionary Biochem. Physiol., 2014, vol. 50, no. 5, pp. 409-415. Soldatov A.A., Andreeva A.Yu., Novitskaya V.N., Parfenova I.A. Coupling of membrane and metabolic function in nucleated erythrocytes of Scorpaena porcus L. under hypoxia in vivo and in vitro. J. Evolutionary Biochem. Physiol., 2014, vol. 50, no. 5, pp. 409-415. Савина М.В. Механизмы адаптации тканевого дыхания в эволюции позвоночных. С.-Петербург: Наука, 1992, 200 с. Savina M.V. Mehanizmy adaptacii tkanevogo dyhaniya v evolyucii pozvonochnyh. S.-Peterburg: Nauka, 1992, 200 s. Soldatov A.A. Cytochrome System and Oxygen Tension in Muscle Tissue of Salt-Water Fish with Different Natural Activity. J. Evolutionary Biochem. Physiol., 1996, vol. 32, no. 2, pp. 112-115. Soldatov A.A. Cytochrome System and Oxygen Tension in Muscle Tissue of Salt-Water Fish with Different Natural Activity. J. Evolutionary Biochem. Physiol., 1996, vol. 32, no. 2, pp. 112-115. Солдатов А.А., Парфенова И.А. Стехиометрия цитохромов и напряжение кислорода в скелетных мышцах морских рыб. Укр. биох. журн., 2014, т. 96, № 2, с. 60-67. Soldatov A.A., Parfenova I.A. Stehiometriya citohromov i napryazhenie kisloroda v skeletnyh myshcah morskih ryb. Ukr. bioh. zhurn., 2014, t. 96, № 2, s. 60-67. Hughes G.M., Johnston I.A. Some responses of the electric ray (Torpedo marmorata) to low ambient oxygen tensions. J. Exp. Biol., 1978, vol. 73, pp. 107-117. Hughes G.M., Johnston I.A. Some responses of the electric ray (Torpedo marmorata) to low ambient oxygen tensions. J. Exp. Biol., 1978, vol. 73, pp. 107-117. Waarde A. Biochemistry of non-protein nitrogenous compounds in fish including the use of amino acids for anaerobic energy production. Comp. Biochem. Physiol., 1988, vol. 91 B, no. 2, pp. 207-228. Waarde A. Biochemistry of non-protein nitrogenous compounds in fish including the use of amino acids for anaerobic energy production. Comp. Biochem. Physiol., 1988, vol. 91 B, no. 2, pp. 207-228. Waarde A., Thillart G., Kesbeke F. Anaerobic energy metabolism of the European eel, Anguilla anguilla L. J. Comp. Physiol., 1983, vol. 149 B, no. 4, pp. 469-475. Waarde A., Thillart G., Kesbeke F. Anaerobic energy metabolism of the European eel, Anguilla anguilla L. J. Comp. Physiol., 1983, vol. 149 B, no. 4, pp. 469-475. Страйер Л. Биохимия. М.: Мир, 1984, т. 1, 232 с. Strayer L. Biohimiya. M.: Mir, 1984, t. 1, 232 s. Johnston I.A. Implications of Muscle Growth Patterns for the Colour and Texture of Fish. Farmed fish quality kestin. Osney Mead Oxford: Blackwell Science Ltd, 2001, pp. 13-30. Johnston I.A. Implications of Muscle Growth Patterns for the Colour and Texture of Fish. Farmed fish quality kestin. Osney Mead Oxford: Blackwell Science Ltd, 2001, pp. 13-30. Love R.M. The chemical biology of fish. London: Academic Press, 1980, vol. 2, 943 p. Love R.M. The chemical biology of fish. London: Academic Press, 1980, vol. 2, 943 p. Drummond G. Muscle metabolism. Fortschritte der Zoollogie, 1967, vol. 18, pp. 359-429. Drummond G. Muscle metabolism. Fortschritte der Zoollogie, 1967, vol. 18, pp. 359-429. Wittenberger C. Metabolic interaction between isolated white and red carp muscle. Rev. Roum. Biol. Ser. Zool., 1973, vol. 18, pp. 71-76. Wittenberger C. Metabolic interaction between isolated white and red carp muscle. Rev. Roum. Biol. Ser. Zool., 1973, vol. 18, pp. 71-76. Солдатов А.А., Парфенова И.А. Напряжение кислорода в крови, скелетных мышцах и особенности тканевого метаболизма кефали-сингиля в условиях экспериментальной гипотермии. Пробл. криобиологии, 2009, т. 19, № 3, с. 290-300. Soldatov A.A., Parfenova I.A. Napryazhenie kisloroda v krovi, skeletnyh myshcah i osobennosti tkanevogo metabolizma kefali-singilya v usloviyah eksperimental'noy gipotermii. Probl. kriobiologii, 2009, t. 19, № 3, s. 290-300. Солдатов А.А. Напряжение кислорода в крови, скелетных мышцах и особенности тканевого метаболизма у кефали-сингиля (Liza aurata Risso, 1810) в условиях острой экспериментальной гипоксии. Актуальные вопросы биологической физики и химии. 2018, т. 3, № 4, с. 724-729. Soldatov A.A. Napryazhenie kisloroda v krovi, skeletnyh myshcah i osobennosti tkanevogo metabolizma u kefali-singilya (Liza aurata Risso, 1810) v usloviyah ostroy eksperimental'noy gipoksii. Aktual'nye voprosy biologicheskoy fiziki i himii. 2018, t. 3, № 4, s. 724-729. Knoerr S., Bohl M., Braunbeck T. Development of red and white muscle in juvenile Danube salmon (Hucho hucho) under the influence of different water currents - a contribution to optimized conditions for reintroduction into natural habitats. Verh. Ges. Ichthyol., 1998, vol. 1, pp. 109-127. Knoerr S., Bohl M., Braunbeck T. Development of red and white muscle in juvenile Danube salmon (Hucho hucho) under the influence of different water currents - a contribution to optimized conditions for reintroduction into natural habitats. Verh. Ges. Ichthyol., 1998, vol. 1, pp. 109-127. Saenger A.M., Kim Z.S., Adam H. The fine structure of muscle fibres of roach, Rutilus rutilus (L.), and chub, Leuciscus cephalus (L.), Cyprinidae, Teleostei: Interspecific differences and effects of habitat and season. J. Fish Biol., 1990, vol. 36, no. 2, pp. 205-213. Saenger A.M., Kim Z.S., Adam H. The fine structure of muscle fibres of roach, Rutilus rutilus (L.), and chub, Leuciscus cephalus (L.), Cyprinidae, Teleostei: Interspecific differences and effects of habitat and season. J. Fish Biol., 1990, vol. 36, no. 2, pp. 205-213. Dickson K.A., Johnson N.M., Donley J.M., Hoskinson J.A., Hansen M.W., Tessier J.D. Ontogenetic changes in characteristics required for endothermy in juvenile black skipjack tuna (Euthynnus lineatus). J. Exp. Biol., 2000, vol. 203, no. 20, pp. 3077-3087. Dickson K.A., Johnson N.M., Donley J.M., Hoskinson J.A., Hansen M.W., Tessier J.D. Ontogenetic changes in characteristics required for endothermy in juvenile black skipjack tuna (Euthynnus lineatus). J. Exp. Biol., 2000, vol. 203, no. 20, pp. 3077-3087. Leonard J.B.K. Regional variation in muscle metabolic enzymes in individual American shad (Alosa sapidissima). Can. J. Zool., 1999, vol. 77, no. 8, pp. 1322-1326. Leonard J.B.K. Regional variation in muscle metabolic enzymes in individual American shad (Alosa sapidissima). Can. J. Zool., 1999, vol. 77, no. 8, pp. 1322-1326. Leonard J.B.K., McCormick S.D. Effects of migration distance on whole-body and tissue-specific energy use in American shad (Alosa sapidissima). Can. J. Fish. Aquat. Sci., 1999, vol. 56, no. 7, pp. 1159-1171. Leonard J.B.K., McCormick S.D. Effects of migration distance on whole-body and tissue-specific energy use in American shad (Alosa sapidissima). Can. J. Fish. Aquat. Sci., 1999, vol. 56, no. 7, pp. 1159-1171. Leonard J.B.K., McCormick S.D. The effect of migration distance and timing on metabolic enzyme activity in an anadromous clupeid, the American shad (Alosa sapidissima). Fish. Physiol. Biochem., 1999, vol. 20, no. 2, pp. 163-179. Leonard J.B.K., McCormick S.D. The effect of migration distance and timing on metabolic enzyme activity in an anadromous clupeid, the American shad (Alosa sapidissima). Fish. Physiol. Biochem., 1999, vol. 20, no. 2, pp. 163-179. Hemre G.I., Kahrs F. 14C-glucose injection in Atlantic cod, Gadus morhua, metabolic responses and excretion via the gill membrane. Aquacult. Nutr., 1997, vol. 3, no. 1, pp. 3-8. Hemre G.I., Kahrs F. 14C-glucose injection in Atlantic cod, Gadus morhua, metabolic responses and excretion via the gill membrane. Aquacult. Nutr., 1997, vol. 3, no. 1, pp. 3-8. Hyvaerinen H., Holopainen I.J., Piironen J. Anaerobic wintering of crucian carp (Carassius carassius L.). 1. Annual dynamics of glycogen reserves in nature. Comp. Biochem. Physiol., 1985, vol. 82 A, no. 4, pp. 797-803. Hyvaerinen H., Holopainen I.J., Piironen J. Anaerobic wintering of crucian carp (Carassius carassius L.). 1. Annual dynamics of glycogen reserves in nature. Comp. Biochem. Physiol., 1985, vol. 82 A, no. 4, pp. 797-803. West T.G., Schulte P.M., Hochachka P.W. Implications of hyperglycemia for post-exercise resynthesis of glycogen in trout skeletal muscle. J. Exp. Biol., 1994, vol. 189, pp. 69-84. West T.G., Schulte P.M., Hochachka P.W. Implications of hyperglycemia for post-exercise resynthesis of glycogen in trout skeletal muscle. J. Exp. Biol., 1994, vol. 189, pp. 69-84. Modigh M., Tota B. Mitochondrial respiration in the ventricular myocardium and in the white and deep red myotomal muscles of juvenile tuna fish (Thunnus thynnus L.). Acta Physiol. Scand., 1975, vol. 93, no. 3, pp. 289-294. Modigh M., Tota B. Mitochondrial respiration in the ventricular myocardium and in the white and deep red myotomal muscles of juvenile tuna fish (Thunnus thynnus L.). Acta Physiol. Scand., 1975, vol. 93, no. 3, pp. 289-294. Jabarsyah A., Tsuchimoto M., Yada O., Kozuru Ya., Miyake T., Misima T., Wang Q., Tachibana K. Comparison of biochemical and physiological characteristics among white, pink, and red muscle fibers in carp (cultured). Fish Sci., 2000, vol. 66, no. 3, pp. 586-593. Jabarsyah A., Tsuchimoto M., Yada O., Kozuru Ya., Miyake T., Misima T., Wang Q., Tachibana K. Comparison of biochemical and physiological characteristics among white, pink, and red muscle fibers in carp (cultured). Fish Sci., 2000, vol. 66, no. 3, pp. 586-593. Yada O., Tsuchimoto M., Wang Q., Apablaza P.A.G., Jabarsyah A., Tachibana K. Differences of muscle fiber type and temporal change of K-value among parts toward depth of dorsal muscle in carp (cultured). Fish Sci., 2000, vol. 66, no. 1, pp. 147-152. Yada O., Tsuchimoto M., Wang Q., Apablaza P.A.G., Jabarsyah A., Tachibana K. Differences of muscle fiber type and temporal change of K-value among parts toward depth of dorsal muscle in carp (cultured). Fish Sci., 2000, vol. 66, no. 1, pp. 147-152. Froyland L., Lie O., Berge R. Mitochondrial and peroxisomal β-oxidation capacities in various tissues from Atlantic salmon Salmo salar. Aquacult. Nutr., 2000, vol. 6, no. 2, pp. 85-89. Froyland L., Lie O., Berge R. Mitochondrial and peroxisomal β-oxidation capacities in various tissues from Atlantic salmon Salmo salar. Aquacult. Nutr., 2000, vol. 6, no. 2, pp. 85-89. Carpene E., Martin B., Libera L.D. Biochemical differences in lateral muscle of wild and farmed gilthead sea bream (Sparus aurata L.). Fish. Physiol. Biochem., 1998, vol. 19, no. 2, pp. 229-238. Carpene E., Martin B., Libera L.D. Biochemical differences in lateral muscle of wild and farmed gilthead sea bream (Sparus aurata L.). Fish. Physiol. Biochem., 1998, vol. 19, no. 2, pp. 229-238. De’Oliveira E.G., Urbinati E.C., Souza V.L., Roviero D.P. Glycogen levels in different tissues of pacu (Piaractus mesopotamicus, Holmberg, 1887). Bol. Inst. Pesca-Sao-Paulo, 1997, vol. 24, pp. 89-95. De’Oliveira E.G., Urbinati E.C., Souza V.L., Roviero D.P. Glycogen levels in different tissues of pacu (Piaractus mesopotamicus, Holmberg, 1887). Bol. Inst. Pesca-Sao-Paulo, 1997, vol. 24, pp. 89-95. El-Casfi M. Effects of water low salinity on tissular lipids in white and red muscle of Liza aurata. Ichtyophysiol. Acta., 1998, no. 21, pp. 15-25. El-Casfi M. Effects of water low salinity on tissular lipids in white and red muscle of Liza aurata. Ichtyophysiol. Acta., 1998, no. 21, pp. 15-25. El-Casfi M., Romdhane M.S., Chanussot F., Cherif A. Variation of the lipid composition of the muscle of mullet Liza aurata living in two sites of different salinity. Proc. Symp. Brest., 19-20, November 1998. Plouzane France Ifremer, 2000, no. 27, pp. 130-139. El-Casfi M., Romdhane M.S., Chanussot F., Cherif A. Variation of the lipid composition of the muscle of mullet Liza aurata living in two sites of different salinity. Proc. Symp. Brest., 19-20, November 1998. Plouzane France Ifremer, 2000, no. 27, pp. 130-139. Kiessling A., Aasgaard T., Storebakken T., Johansson L., Kiessling K.H. Changes in the structure and function of the epaxial muscle of rainbow trout (Oncorhynchus mykiss) in relation to ration and age. 3. Chemical composition. Aquaculture, 1991, vol. 93, no. 4, pp. 373-387. Kiessling A., Aasgaard T., Storebakken T., Johansson L., Kiessling K.H. Changes in the structure and function of the epaxial muscle of rainbow trout (Oncorhynchus mykiss) in relation to ration and age. 3. Chemical composition. Aquaculture, 1991, vol. 93, no. 4, pp. 373-387. De’Oliveira E.G., Urbinati E.C., Souza V.L., Roviero D.P. Visceral lipsomatic index (VLSI) and total lipid levels in different body tissues of pacu (Piaractus mesopotamicus, Holmberg, 1887). Bol. Inst. Pesca-Sao-Paulo, 1997, vol. 24, pp. 97-103. De’Oliveira E.G., Urbinati E.C., Souza V.L., Roviero D.P. Visceral lipsomatic index (VLSI) and total lipid levels in different body tissues of pacu (Piaractus mesopotamicus, Holmberg, 1887). Bol. Inst. Pesca-Sao-Paulo, 1997, vol. 24, pp. 97-103. George J.C., Stevens E.D. Fine structure and metabolic adaptation of red and white muscles in tuna. Environ. Biol. Fish, 1978, vol. 3, no. 2, pp. 185-191. George J.C., Stevens E.D. Fine structure and metabolic adaptation of red and white muscles in tuna. Environ. Biol. Fish, 1978, vol. 3, no. 2, pp. 185-191. Driedzic W.R., Hochachka P.W. The unansweres question of high anaerobic capabilities of carp white muscle. Can. J. Zool., 1975, vol. 53, pp. 706-712. Driedzic W.R., Hochachka P.W. The unansweres question of high anaerobic capabilities of carp white muscle. Can. J. Zool., 1975, vol. 53, pp. 706-712. Johnston I.A., Davison W., Goldspink G. Energy metabolism of carp swimming muscles. J. Comp. Physiol., 1977, vol. 114, no. 2, pp. 203-216. Johnston I.A., Davison W., Goldspink G. Energy metabolism of carp swimming muscles. J. Comp. Physiol., 1977, vol. 114, no. 2, pp. 203-216. Martinez M.L., Landry C., Boehm R., Manning S., Cheek A.O., Rees B.B. Effects of long-term hypoxia on enzymes of carbohydrate metabolism in the Gulf killifish, Fundulus grandis. J. Exp. Biol., 2006, vol. 209, no. 19, pp. 3851-3861. Martinez M.L., Landry C., Boehm R., Manning S., Cheek A.O., Rees B.B. Effects of long-term hypoxia on enzymes of carbohydrate metabolism in the Gulf killifish, Fundulus grandis. J. Exp. Biol., 2006, vol. 209, no. 19, pp. 3851-3861. Матюхин В.А., Нешумова Т.В., Дементьев Я.В. Изменения температуры красных и белых мышц байкальского хариуса Thymallus arcticus baicalensis Dyb. при различных скоростях плавания. Вопр. ихтиол., 1975, т. 15, № 5, c. 884-889. Matyuhin V.A., Neshumova T.V., Dement'ev Ya.V. Izmeneniya temperatury krasnyh i belyh myshc baykal'skogo hariusa Thymallus arcticus baicalensis Dyb. pri razlichnyh skorostyah plavaniya. Vopr. ihtiol., 1975, t. 15, № 5, c. 884-889. Coughlin D.J., Rome L.C. Muscle activity in steady swimming scup, Stenotomus chrysops, varies with fiber type and body position. Bull. Mar. Biol. (Lab. Woods Hole), 1999, vol. 196, no 2, pp. 145-152. Coughlin D.J., Rome L.C. Muscle activity in steady swimming scup, Stenotomus chrysops, varies with fiber type and body position. Bull. Mar. Biol. (Lab. Woods Hole), 1999, vol. 196, no 2, pp. 145-152. Кондратьева Т.П., Астахова Л.П. Морфологические и биохимические особенности белых и красных мышц при различных физиологических состояниях. Тр. Карадагского филиала ИнБЮМ НАН Украины, Севастополь, 1997, c. 111-120. Kondrat'eva T.P., Astahova L.P. Morfologicheskie i biohimicheskie osobennosti belyh i krasnyh myshc pri razlichnyh fiziologicheskih sostoyaniyah. Tr. Karadagskogo filiala InBYuM NAN Ukrainy, Sevastopol', 1997, c. 111-120. Chance B., Oshino N., Sugano T., Mayevsky A. Basic principles of tissue oxygen determination from mitochondrial signals. Adv. Exp. Med. Biol., 1973, vol. 37, pp. 277-292. Chance B., Oshino N., Sugano T., Mayevsky A. Basic principles of tissue oxygen determination from mitochondrial signals. Adv. Exp. Med. Biol., 1973, vol. 37, pp. 277-292. Wilson A., Owen Cn., Erecinska M. Quantitative dependence of mitochondrial oxidative phosphorylation on oxygen concentration: A mathematical model. Arch. Biochem. Biophys., 1979, vol. 195, no. 2, pp. 494-504. Wilson A., Owen Cn., Erecinska M. Quantitative dependence of mitochondrial oxidative phosphorylation on oxygen concentration: A mathematical model. Arch. Biochem. Biophys., 1979, vol. 195, no. 2, pp. 494-504. Lubbers D., Kessler M. Oxygen supply and the rate of tissue respiration. Oxygen transport in blood and tissue. Stuttgart: Tieme, 1968, pp. 90-99. Lubbers D., Kessler M. Oxygen supply and the rate of tissue respiration. Oxygen transport in blood and tissue. Stuttgart: Tieme, 1968, pp. 90-99. Савина М.В., Маслова Г.М., Демин В.И., Бакланова С.М. Исследование цитохромов в соматической и сердечной мышцах миноги Lampetra fluviatilis L. Ж. эволюц. биох. физиол., 1981, т. 17, № 3, c. 246-253. Savina M.V., Maslova G.M., Demin V.I., Baklanova S.M. Issledovanie citohromov v somaticheskoy i serdechnoy myshcah minogi Lampetra fluviatilis L. Zh. evolyuc. bioh. fiziol., 1981, t. 17, № 3, c. 246-253. Richardson T., Tappel A.L., Smith L.M., Houle C.R. Polyunsaturated fatty acids in mitochondria. J. Lipid Res., 1962, vol. 3, pp. 344-350. Richardson T., Tappel A.L., Smith L.M., Houle C.R. Polyunsaturated fatty acids in mitochondria. J. Lipid Res., 1962, vol. 3, pp. 344-350. Wodtke E. Temperature adaptation of biological membranes. Compensation of the molar activity of cytochrome C oxidase in the mitochondrial enegy-transducing membrane during thermal acclimation of the carp (Cyprinus carpio L.). Biochim. biophys. acta., 1981, vol. 640, no. 3, pp. 710-720. Wodtke E. Temperature adaptation of biological membranes. Compensation of the molar activity of cytochrome C oxidase in the mitochondrial enegy-transducing membrane during thermal acclimation of the carp (Cyprinus carpio L.). Biochim. biophys. acta., 1981, vol. 640, no. 3, pp. 710-720. Солдатов А.А., Парфенова И.А. Цитохромная система и уровень миоглобина в скелетных мышцах кефали-сингиля (Liza aurata Risso) в условиях экспериментальной гипотермии. Труды ИБВВ РАН, 2017, № 80 (83), с. 69-75. Soldatov A.A., Parfenova I.A. Citohromnaya sistema i uroven' mioglobina v skeletnyh myshcah kefali-singilya (Liza aurata Risso) v usloviyah eksperimental'noy gipotermii. Trudy IBVV RAN, 2017, № 80 (83), s. 69-75. Wilson M.T., Bonaventura J., Brunory M. Mitochondrial cytochrome content and cytochrome oxidase activity of some amazonian fish. Comp. Biochem Physiol., 1979, vol. 62 A, pp. 245-249. Wilson M.T., Bonaventura J., Brunory M. Mitochondrial cytochrome content and cytochrome oxidase activity of some amazonian fish. Comp. Biochem Physiol., 1979, vol. 62 A, pp. 245-249. Martinez M., Dutil J.D., Guderley H. Longitudinal and allometric variation in indicators of muscle metabolic capacities in Atlantic cod (Gadus morhua). J. Exp. Zool., 2000, vol. 287, no. 1, pp. 38-45. Martinez M., Dutil J.D., Guderley H. Longitudinal and allometric variation in indicators of muscle metabolic capacities in Atlantic cod (Gadus morhua). J. Exp. Zool., 2000, vol. 287, no. 1, pp. 38-45. Rodnick K.J., Williams S.R. Effects of body size on biochemical characteristics of trabecular cardiac muscle and plasma of rainbow trout (Oncorhynchus mykiss). Comp. Biochem. Physiol., 1999, vol. 122 A, no. 4, pp. 407-413. Rodnick K.J., Williams S.R. Effects of body size on biochemical characteristics of trabecular cardiac muscle and plasma of rainbow trout (Oncorhynchus mykiss). Comp. Biochem. Physiol., 1999, vol. 122 A, no. 4, pp. 407-413. Garenc C., Couture P., Laflamme M., Guderley H. Metabolic correlates of burst swimming capacity of juvenile and adult threespine stickleback (Gasterosteus aculeatus). J. Comp. Physiol., 1999, vol. 169, no. 2, pp. 113-122. Garenc C., Couture P., Laflamme M., Guderley H. Metabolic correlates of burst swimming capacity of juvenile and adult threespine stickleback (Gasterosteus aculeatus). J. Comp. Physiol., 1999, vol. 169, no. 2, pp. 113-122. Phillips M.C.L., Moyes C.D., Tufts B.L. The effects of cell ageing on metabolism in rainbow trout (Oncorhynchus mykiss) red blood cells. J. Exp. Biol., 2000, vol. 203, no. 6, pp. 1039-1045. Phillips M.C.L., Moyes C.D., Tufts B.L. The effects of cell ageing on metabolism in rainbow trout (Oncorhynchus mykiss) red blood cells. J. Exp. Biol., 2000, vol. 203, no. 6, pp. 1039-1045. Soldatov A.A., Savina M.V. Effect of hypoxia on the content and stoichiometry of cytochromes in muscle of the gray mullet Liza aurata. J. Evolutionary Biochem. Physiol., 2008, vol. 44, no. 5, pp. 599-604. Soldatov A.A., Savina M.V. Effect of hypoxia on the content and stoichiometry of cytochromes in muscle of the gray mullet Liza aurata. J. Evolutionary Biochem. Physiol., 2008, vol. 44, no. 5, pp. 599-604. Sidell B.D. Turnover of cytochrome c in skeletal muscle of green sunfish (Lepomis cyanellus R.) during thermal acclimation. J. Exp. Zool., 1977, vol. 199, no. 2, pp. 233-250. Sidell B.D. Turnover of cytochrome c in skeletal muscle of green sunfish (Lepomis cyanellus R.) during thermal acclimation. J. Exp. Zool., 1977, vol. 199, no. 2, pp. 233-250. Blier P.U., Lemieux H. The impact of the thermal sensitivity of cytochrome c oxidase on the respiration rate of Arctic charr red muscle mitochondria. J. Comp. Physiol., 2001, vol. 171, no. 3, pp. 247-253. Blier P.U., Lemieux H. The impact of the thermal sensitivity of cytochrome c oxidase on the respiration rate of Arctic charr red muscle mitochondria. J. Comp. Physiol., 2001, vol. 171, no. 3, pp. 247-253. Wilson F.R., Somero G., Prosser C.L. Temperature-metabolism relations of two species of Sebastes from different thermal environments. Comp. Biochem. Physiol., 1974, vol. 47, no. 2 B, pp. 485-491. Wilson F.R., Somero G., Prosser C.L. Temperature-metabolism relations of two species of Sebastes from different thermal environments. Comp. Biochem. Physiol., 1974, vol. 47, no. 2 B, pp. 485-491. Sebert P., Simon B., Barthelemy L. Hydrostatic pressure induces a state resembling histotoxic hypoxia in Anguilla Anguilla. Comp. Biochem. Physiol., 1993, vol. 105 A, no. 2, pp. 255-258. Sebert P., Simon B., Barthelemy L. Hydrostatic pressure induces a state resembling histotoxic hypoxia in Anguilla Anguilla. Comp. Biochem. Physiol., 1993, vol. 105 A, no. 2, pp. 255-258. Cohen A., Nugegoda D., Gagnon M.M. Metabolic responses of fish following exposure to two different oil spill remediation techniques. Ecotoxicol. Environ. Saf., 2001, vol. 48, no. 3, pp. 306-310. Cohen A., Nugegoda D., Gagnon M.M. Metabolic responses of fish following exposure to two different oil spill remediation techniques. Ecotoxicol. Environ. Saf., 2001, vol. 48, no. 3, pp. 306-310. Reddy S.J., Kalarani V., Tharakanadha B., Reddy D.C., Ramamurthi R. Changes in energy metabolism of the fish, Labeo rohita in relation to prolonged lead exposure and recovery. J. Ecotoxicol. Environ. Monitoring, 1998, vol. 8, no. 1, pp. 43-53. Reddy S.J., Kalarani V., Tharakanadha B., Reddy D.C., Ramamurthi R. Changes in energy metabolism of the fish, Labeo rohita in relation to prolonged lead exposure and recovery. J. Ecotoxicol. Environ. Monitoring, 1998, vol. 8, no. 1, pp. 43-53. Вержбинская Н.А. Цитохромная система мозга в филогенезе позвоночных животных. Физиол. ж. СССР, 1953, т. 39, no. 1, с. 17-26. Verzhbinskaya N.A. Citohromnaya sistema mozga v filogeneze pozvonochnyh zhivotnyh. Fiziol. zh. SSSR, 1953, t. 39, no. 1, s. 17-26. Buchner T., Abele D., Poertner H.O. Oxyconformity in the intertidal worm Sipunculus nudus: the mitochondrial background and energetic consequences. Comp. Biochem. Physiol., 2001, vol. 129, no. 1, pp. 109-120. Buchner T., Abele D., Poertner H.O. Oxyconformity in the intertidal worm Sipunculus nudus: the mitochondrial background and energetic consequences. Comp. Biochem. Physiol., 2001, vol. 129, no. 1, pp. 109-120. Tschischka K., Abele D., Poertner H.O. Mitochondrial oxyconformity and cold adaptation in the polychaete Nereis pelagica and the bivalve Arctica islandica from the Baltic and White Seas. J. Exp. Biol., 2000, vol. 203, no. 21, pp. 3355-3368. Tschischka K., Abele D., Poertner H.O. Mitochondrial oxyconformity and cold adaptation in the polychaete Nereis pelagica and the bivalve Arctica islandica from the Baltic and White Seas. J. Exp. Biol., 2000, vol. 203, no. 21, pp. 3355-3368. Matschak T.W., Tyler D.D., Stickland N.C. Metabolic enzyme activities in Atlantic salmon (Salmo salar L.) embryos respond more to chronic changes in oxygen availability than to environmental temperature. Fish Physiol. Biochem., 1998, vol. 18, no. 2, pp. 115-123. Matschak T.W., Tyler D.D., Stickland N.C. Metabolic enzyme activities in Atlantic salmon (Salmo salar L.) embryos respond more to chronic changes in oxygen availability than to environmental temperature. Fish Physiol. Biochem., 1998, vol. 18, no. 2, pp. 115-123. Bickler P.E., Buck L.T.Hypoxia tolerance in reptiles, amphibians, and fishes: life with variable oxygen availability. Annu. Rev. Physiol., 2007, vol. 69, pp. 145-170. Bickler P.E., Buck L.T.Hypoxia tolerance in reptiles, amphibians, and fishes: life with variable oxygen availability. Annu. Rev. Physiol., 2007, vol. 69, pp. 145-170. Vornanen M., Paajanen V. Seasonality of dihydropyridine receptor binding in the heart of an anoxia-tolerant vertebrate, the crucian carp (Carassius carassius L.). Am. J. Physiol. Regul. Integr. Comp. Physiol., 2004, vol. 287, no. 5, pp. R1263-R1269. Vornanen M., Paajanen V. Seasonality of dihydropyridine receptor binding in the heart of an anoxia-tolerant vertebrate, the crucian carp (Carassius carassius L.). Am. J. Physiol. Regul. Integr. Comp. Physiol., 2004, vol. 287, no. 5, pp. R1263-R1269. Hochachka P. Defence strategies against hypoxia and hypothermia. Science, 1986, vol. 231, pp. 324-241. Hochachka P. Defence strategies against hypoxia and hypothermia. Science, 1986, vol. 231, pp. 324-241. Zhou B.S., Wu R.S., Randall D.J., Lam P.K., Ip Y.K., Chew S.F. Metabolic adjustments in the common carp during prolonged hypoxia. J. Fish Biol., 2000, vol. 57, no. 5, pp. 1160-1171. Zhou B.S., Wu R.S., Randall D.J., Lam P.K., Ip Y.K., Chew S.F. Metabolic adjustments in the common carp during prolonged hypoxia. J. Fish Biol., 2000, vol. 57, no. 5, pp. 1160-1171. Chew S.F., Gan J., Ip Y.K. Nitrogen metabolism and excretion in the swamp eel, Monopterus albus, during 6 or 40 days of estivation in mud. Physiol. Biochem. Zool., 2005, vol. 78, no. 4, pp. 620-629. Chew S.F., Gan J., Ip Y.K. Nitrogen metabolism and excretion in the swamp eel, Monopterus albus, during 6 or 40 days of estivation in mud. Physiol. Biochem. Zool., 2005, vol. 78, no. 4, pp. 620-629. Dorigatti M., Krumschnabel G., Schwarzbaum P.J., Wieser W. Effects of hypoxia on energy metabolism in goldfish hepatocytes. Comp. Biochem. Physiol., 1997, vol. 117 B, no. 1, pp. 151-158. Dorigatti M., Krumschnabel G., Schwarzbaum P.J., Wieser W. Effects of hypoxia on energy metabolism in goldfish hepatocytes. Comp. Biochem. Physiol., 1997, vol. 117 B, no. 1, pp. 151-158. Herbert N.A., Wells R.M. The aerobic physiology of the air-breathing blue gourami, Trichogaster trichopterus, necessitates behavioural regulation of breath-hold limits during hypoxic stress and predatory challenge. J. Comp. Physiol. B., 2001, vol. 171, no. 7, pp. 603-612. Herbert N.A., Wells R.M. The aerobic physiology of the air-breathing blue gourami, Trichogaster trichopterus, necessitates behavioural regulation of breath-hold limits during hypoxic stress and predatory challenge. J. Comp. Physiol. B., 2001, vol. 171, no. 7, pp. 603-612. Ip Y.K., Kuah S.S., Chew S.F. Strategies adopted by the mudskipper Boleophthalmus boddaerti to survive sulfide exposure in normoxia or hypoxia. Physiol. Biochem. Zool., 2004, vol. 77, no. 5, pp. 824-837. Ip Y.K., Kuah S.S., Chew S.F. Strategies adopted by the mudskipper Boleophthalmus boddaerti to survive sulfide exposure in normoxia or hypoxia. Physiol. Biochem. Zool., 2004, vol. 77, no. 5, pp. 824-837. MacCormack T.J., Lewis J.M., Almeida-Val V.M., Val A.L., Driedzic W.R. Carbohydrate management, anaerobic metabolism, and adenosine levels in the armoured catfish, Liposarcus pardalis (castelnau), during hypoxia. J. Exp. Zoolog. A. Comp. Exp. Biol., 2006, vol. 305, no. 4, pp. 363-375. MacCormack T.J., Lewis J.M., Almeida-Val V.M., Val A.L., Driedzic W.R. Carbohydrate management, anaerobic metabolism, and adenosine levels in the armoured catfish, Liposarcus pardalis (castelnau), during hypoxia. J. Exp. Zoolog. A. Comp. Exp. Biol., 2006, vol. 305, no. 4, pp. 363-375. Moraes G., Chippari A.R., Guerra C.D.R., Gomes L.C., Souza R.H.S. Immediate changes on metabolic parameters of the freshwater teleost fish Piaractus mesopotamicus (PACU) under severe hypoxia. Bol. Tec. Cepta., 1997, vol. 10, pp. 45-52. Moraes G., Chippari A.R., Guerra C.D.R., Gomes L.C., Souza R.H.S. Immediate changes on metabolic parameters of the freshwater teleost fish Piaractus mesopotamicus (PACU) under severe hypoxia. Bol. Tec. Cepta., 1997, vol. 10, pp. 45-52. Moraes G., Choudhuri J.V., Souza R.H.S. Metabolic strategies of Hypostomus regani (Cascudo), a freshwater teleost fish under extreme environmental hypoxia. Bol. Tec. Cepta., 1997, vol. 10, pp. 35-44. Moraes G., Choudhuri J.V., Souza R.H.S. Metabolic strategies of Hypostomus regani (Cascudo), a freshwater teleost fish under extreme environmental hypoxia. Bol. Tec. Cepta., 1997, vol. 10, pp. 35-44. Pincetich C.A., Viant M.R., Hinton D.E., Tjeerdema R.S. Metabolic changes in Japanese medaka (Oryzias latipes) during embryogenesis and hypoxia as determined by in vivo 31P NMR. Comp. Biochem. Physiol. C. Toxicol. Pharmacol, 2005, vol. 140, no. 1, pp. 103-13. Pincetich C.A., Viant M.R., Hinton D.E., Tjeerdema R.S. Metabolic changes in Japanese medaka (Oryzias latipes) during embryogenesis and hypoxia as determined by in vivo 31P NMR. Comp. Biochem. Physiol. C. Toxicol. Pharmacol, 2005, vol. 140, no. 1, pp. 103-13. Van Ginneken V.J.T., Van den Thillart G.E.E.J., Muller H.J., Van Deursen S., Onderwater M., Visee J., Hopmans V., Van Vliet G., Nicolay K. Phospholylation state of red and white muscles in tilapia during graded hypoxia: an in vivo 31P-NMR study. Am. J. Physiol. Regul. Integrat. Comp. Physiol., 1999, vol. 277, pp. 1501-1512. Van Ginneken V.J.T., Van den Thillart G.E.E.J., Muller H.J., Van Deursen S., Onderwater M., Visee J., Hopmans V., Van Vliet G., Nicolay K. Phospholylation state of red and white muscles in tilapia during graded hypoxia: an in vivo 31P-NMR study. Am. J. Physiol. Regul. Integrat. Comp. Physiol., 1999, vol. 277, pp. 1501-1512. Rees B.B., Bowman J.A.L., Schulte P.M. Structure and sequence conservation of a putative hypoxia response element in the lactate dehydrogenase-B gene of Fundulus. Biol. Bull., 2001, vol. 200, pp. 247-251. Rees B.B., Bowman J.A.L., Schulte P.M. Structure and sequence conservation of a putative hypoxia response element in the lactate dehydrogenase-B gene of Fundulus. Biol. Bull., 2001, vol. 200, pp. 247-251. Bosworth C.A., Chou C.W., Cole R.B., Rees B.B. Protein expression patterns in zebrafish skeletal muscle: initial characterization and the effects of hypoxic exposure. Proteomics, 2005, vol. 5, no. 5, pp. 1362-1371. Bosworth C.A., Chou C.W., Cole R.B., Rees B.B. Protein expression patterns in zebrafish skeletal muscle: initial characterization and the effects of hypoxic exposure. Proteomics, 2005, vol. 5, no. 5, pp. 1362-1371. Ju Z., Wells M.C., Heater S.J., Walter R.B. Multiple tissue gene expression analyses in Japanese medaka (Oryzias latipes) exposed to hypoxia. Comp. Biochem. Physiol. C., 2007, vol. 145, no. 1, pp. 134-144. Ju Z., Wells M.C., Heater S.J., Walter R.B. Multiple tissue gene expression analyses in Japanese medaka (Oryzias latipes) exposed to hypoxia. Comp. Biochem. Physiol. C., 2007, vol. 145, no. 1, pp. 134-144. Vleugel M.M., Bos R., Buerger H., van der Groep P., Saramaki O.R., Visakorpi T., van der Wall E., van Diest P.J. No amplifications of hypoxia-inducible factor-1alpha gene in invasive breast cancer: a tissue microarray study. Cell. Oncol., 2004, vol. 26, no. 5-6, pp. 347-351. Vleugel M.M., Bos R., Buerger H., van der Groep P., Saramaki O.R., Visakorpi T., van der Wall E., van Diest P.J. No amplifications of hypoxia-inducible factor-1alpha gene in invasive breast cancer: a tissue microarray study. Cell. Oncol., 2004, vol. 26, no. 5-6, pp. 347-351. Jensen M.A., Gesser H. Influence of inorganic phosphate and energy state on force in skinned cardiac muscle from freshwater turtle and rainbow trout. J. Comp. Physiol. B., 1999, vol. 169, no. 6, pp. 439-444. Jensen M.A., Gesser H. Influence of inorganic phosphate and energy state on force in skinned cardiac muscle from freshwater turtle and rainbow trout. J. Comp. Physiol. B., 1999, vol. 169, no. 6, pp. 439-444. Overgaard J., Gessre H. Force development, energy state and ATP production of cardiac muscle from turtles and trout during normoxia and severe hypoxia. J. Exp. Biol., 2004, vol. 207, pp. 1915-1924. Overgaard J., Gessre H. Force development, energy state and ATP production of cardiac muscle from turtles and trout during normoxia and severe hypoxia. J. Exp. Biol., 2004, vol. 207, pp. 1915-1924. Smith R.W., Houlihan D.F., Nilsson H.E., Alexandre J. Tissue-specific changes in RNA synthesis in vivo during anoxia in crucian carp. Am. J. Physiol. Regul. Integrat. Comp. Physiol., 1999, vol. 277, pp. 690-697. Smith R.W., Houlihan D.F., Nilsson H.E., Alexandre J. Tissue-specific changes in RNA synthesis in vivo during anoxia in crucian carp. Am. J. Physiol. Regul. Integrat. Comp. Physiol., 1999, vol. 277, pp. 690-697. MacCormack T.J., Driedzic W.R. The impact of hypoxia on in vivo glucose uptake in a hypoglycemic fish, Myoxocephalus scorpius. Am. J. Physiol. Regul. Integr. Comp. Physiol., 2007, vol. 292, no. 2, pp. R1033-R1042. MacCormack T.J., Driedzic W.R. The impact of hypoxia on in vivo glucose uptake in a hypoglycemic fish, Myoxocephalus scorpius. Am. J. Physiol. Regul. Integr. Comp. Physiol., 2007, vol. 292, no. 2, pp. R1033-R1042. Plante S., Chabot D., Dutil J. Hypoxia tolerance in Atlantic cod. J. Fish Biol., 1998, vol. 53, no. 6, pp. 1342-1356. Plante S., Chabot D., Dutil J. Hypoxia tolerance in Atlantic cod. J. Fish Biol., 1998, vol. 53, no. 6, pp. 1342-1356. Raaij M.T.M., Pit D.S.S., Balm P.H.M., Steffens A.B., Thillart G.E.E.J.M. Behavioral strategy and the physiological stress response in rainbow trout exposed to severe hypoxia. Horm. Behav., 1996, vol. 30, no. 1, pp. 85-92. Raaij M.T.M., Pit D.S.S., Balm P.H.M., Steffens A.B., Thillart G.E.E.J.M. Behavioral strategy and the physiological stress response in rainbow trout exposed to severe hypoxia. Horm. Behav., 1996, vol. 30, no. 1, pp. 85-92. Raaij M.T.M., Thillart G.E.E.J.M., Vianen G.J., Pit D.S.S., Balm P.H.M., Steffens A.B. Substrate mobilization and hormonal changes in rainbow trout (Oncorhynchus mykiss L.) and common carp (Cyprinus carpio L.) during deep hypoxia and subsequent recovery. J. Comp. Physiol., 1996, vol. 166, no. 7, pp. 443-452. Raaij M.T.M., Thillart G.E.E.J.M., Vianen G.J., Pit D.S.S., Balm P.H.M., Steffens A.B. Substrate mobilization and hormonal changes in rainbow trout (Oncorhynchus mykiss L.) and common carp (Cyprinus carpio L.) during deep hypoxia and subsequent recovery. J. Comp. Physiol., 1996, vol. 166, no. 7, pp. 443-452. Wells R.M., Baldwin J. Plasma lactate and glucose flushes following burst swimming in silver trevally (Pseudocaranx dentex: Carangidae) support the "releaser" hypothesis. Comp. Biochem. Physiol. A. Mol. Integr. Physiol., 2006, vol. 143, no. 3, pp. 347-352. Wells R.M., Baldwin J. Plasma lactate and glucose flushes following burst swimming in silver trevally (Pseudocaranx dentex: Carangidae) support the "releaser" hypothesis. Comp. Biochem. Physiol. A. Mol. Integr. Physiol., 2006, vol. 143, no. 3, pp. 347-352. Jackson D.C. Acid-base balance during hypoxic hypometabolism: selected vertebrate strategies (review). Respir. Physiol. Neurobiol., 2004, vol. 141, no. 3, pp. 273-283. Jackson D.C. Acid-base balance during hypoxic hypometabolism: selected vertebrate strategies (review). Respir. Physiol. Neurobiol., 2004, vol. 141, no. 3, pp. 273-283. Zhang Z., Wu R.S.S., Mok H.O.L., Wang Y., Poon W.W.L., Cheng S.H., Kong R.Y.C. Isolation, characterization and expression analysis a hypoxia-responsive glucose transporter gene from the grass carp, Ctenopharyngodon idellus. Eur. J. Biochem., 2003, vol. 270, pp. 3010-3017. Zhang Z., Wu R.S.S., Mok H.O.L., Wang Y., Poon W.W.L., Cheng S.H., Kong R.Y.C. Isolation, characterization and expression analysis a hypoxia-responsive glucose transporter gene from the grass carp, Ctenopharyngodon idellus. Eur. J. Biochem., 2003, vol. 270, pp. 3010-3017. Chippari-Gomes A.R., Gomes L.C., Lopes N.P., Val A.L., Almeida-Val V.M. Metabolic adjustments in two Amazonian cichlids exposed to hypoxia and anoxia. Comp. Biochem. Physiol. B. Biochem. Mol. Biol., 2005, vol. 141, no. 3, pp. 347-355. Chippari-Gomes A.R., Gomes L.C., Lopes N.P., Val A.L., Almeida-Val V.M. Metabolic adjustments in two Amazonian cichlids exposed to hypoxia and anoxia. Comp. Biochem. Physiol. B. Biochem. Mol. Biol., 2005, vol. 141, no. 3, pp. 347-355. Bidinotto P.M., Moraes G., Souza R.H.S. Hepatic glycogen and glucose in eight tropical freshwater teleost fish: A procedure for field determinations of micro samples. Bol. Tec. Cepta., 1997, vol. 10, pp. 53-60. Bidinotto P.M., Moraes G., Souza R.H.S. Hepatic glycogen and glucose in eight tropical freshwater teleost fish: A procedure for field determinations of micro samples. Bol. Tec. Cepta., 1997, vol. 10, pp. 53-60. Johnston I.A., Bernard L.M. Utilization of the ethanol pathway in carp following exposure to anoxia. J. Exp. Biol., 1983, vol. 104, pp. 73-78. Johnston I.A., Bernard L.M. Utilization of the ethanol pathway in carp following exposure to anoxia. J. Exp. Biol., 1983, vol. 104, pp. 73-78. Jurss K. Athanol - ein Endprodukt des anaeroben Stoffwechsels von Fischen. Biol. Rdsch., 1982, vol. 20, no. 3, pp. 178. Jurss K. Athanol - ein Endprodukt des anaeroben Stoffwechsels von Fischen. Biol. Rdsch., 1982, vol. 20, no. 3, pp. 178. Lutz P.L., Nilsson G.E. Contrasting strategies for anoxic brain survival - glycolysis up or down. J. Exp. Biol., 1997, vol. 200, pp. 411-419. Lutz P.L., Nilsson G.E. Contrasting strategies for anoxic brain survival - glycolysis up or down. J. Exp. Biol., 1997, vol. 200, pp. 411-419. Waversveld J., Addink A.D.F., Thillart G. The anaerobic energy metabolism of goldfish determined by simultaneous direct and indirect calorimetry during anoxia and hypoxia. J. Comp. Physiol., 1989, vol. 159 B, no. 3, pp. 263-268. Waversveld J., Addink A.D.F., Thillart G. The anaerobic energy metabolism of goldfish determined by simultaneous direct and indirect calorimetry during anoxia and hypoxia. J. Comp. Physiol., 1989, vol. 159 B, no. 3, pp. 263-268. Thillart G., Waarde A. Teleosts in hypoxia: aspects of anaerobic metabolism. Mol. Physiol. 1985, vol. 8, no. 3, pp. 393-409. Thillart G., Waarde A. Teleosts in hypoxia: aspects of anaerobic metabolism. Mol. Physiol. 1985, vol. 8, no. 3, pp. 393-409. Waarde A. Aerobic and anaerobic ammonia production by fish. Comp. Biochem. Physiol., 1983, vol. 74, no. 4, pp. 675-684. Waarde A. Aerobic and anaerobic ammonia production by fish. Comp. Biochem. Physiol., 1983, vol. 74, no. 4, pp. 675-684. Jorgensen J.B., Mustafa T. The effect of hypoxia on carbohydrate metabolism in flounder (Platichthys flesus L.) II. High energy phosphate compounds and the role of glycolytic and gluconeogenetic enzymes. Comp. Biochem. Physiol., 1980, vol. 67 B, pp. 249-256. Jorgensen J.B., Mustafa T. The effect of hypoxia on carbohydrate metabolism in flounder (Platichthys flesus L.) II. High energy phosphate compounds and the role of glycolytic and gluconeogenetic enzymes. Comp. Biochem. Physiol., 1980, vol. 67 B, pp. 249-256. Waarde A., De Wilde H.M. Nitrogen metabolism in goldfish, Carassius auratus (L.) pathway of aerobic and muscle mitochondria. Comp. Biochem. Physiol., 1982, vol. 72 B, no. 1, pp. 133-136. Waarde A., De Wilde H.M. Nitrogen metabolism in goldfish, Carassius auratus (L.) pathway of aerobic and muscle mitochondria. Comp. Biochem. Physiol., 1982, vol. 72 B, no. 1, pp. 133-136. Lushchak V.I., Smirnova Y.D., Storey K.B. AMP-deaminase from sea scorpion white muscle: properties and redistribution under hypoxia. Comp. Biochem. Physiol., 1998, vol. 119 B, no. 3, pp. 611-618. Lushchak V.I., Smirnova Y.D., Storey K.B. AMP-deaminase from sea scorpion white muscle: properties and redistribution under hypoxia. Comp. Biochem. Physiol., 1998, vol. 119 B, no. 3, pp. 611-618. Светличный Л.С., Юнева Т.В., Шульман Г.Е., Хаусман Дж. А. Использование белка в энергетическом обмене ветвистоусого рачка Moina macrura при различном содержании кислорода в воде. Докл. РАН, 1994, т. 337, № 3, c. 428-430. Svetlichnyy L.S., Yuneva T.V., Shul'man G.E., Hausman Dzh. A. Ispol'zovanie belka v energeticheskom obmene vetvistousogo rachka Moina macrura pri razlichnom soderzhanii kisloroda v vode. Dokl. RAN, 1994, t. 337, № 3, c. 428-430. Столбов А.Я., Ставицкая Е.Н., Шульман Г.Е. Потребление кислорода и экскреция азота у черноморских рыб различной экологической специализации при гипоксических режимах. Гидробиол. ж., 1995, т. 31, № 1, c. 73-78. Stolbov A.Ya., Stavickaya E.N., Shul'man G.E. Potreblenie kisloroda i ekskreciya azota u chernomorskih ryb razlichnoy ekologicheskoy specializacii pri gipoksicheskih rezhimah. Gidrobiol. zh., 1995, t. 31, № 1, c. 73-78. Шульман Г.Е., Аболмасова Г.И., Столбов А.Я. Использование белка в энергетическом обмене гидробионтов. Усп. совр. биол., 1993, т. 113, № 5, c. 576-586. Shul'man G.E., Abolmasova G.I., Stolbov A.Ya. Ispol'zovanie belka v energeticheskom obmene gidrobiontov. Usp. sovr. biol., 1993, t. 113, № 5, c. 576-586. Романенко В.Д., Арсан О.М., Соломатина В.Д. Механизмы температурной акклимации рыб. К.: Наук. думка, 1991, 192 с. Romanenko V.D., Arsan O.M., Solomatina V.D. Mehanizmy temperaturnoy akklimacii ryb. K.: Nauk. dumka, 1991, 192 s. Owen T.G., Hochachka P.W. Purification and properties of dolphin muscle aspartate and alanine transaminases and their possible roles in the energy metabolism of diving mammals. Biochem. J., 1974, vol. 143, pp. 541-553. Owen T.G., Hochachka P.W. Purification and properties of dolphin muscle aspartate and alanine transaminases and their possible roles in the energy metabolism of diving mammals. Biochem. J., 1974, vol. 143, pp. 541-553. Mommsen Th.P., French C.J., Hochachka P.W. Sites and patterns of protein and amino acid utilization during spawning migration of salmon. Can. J. Zool., 1980, vol. 58, pp. 1785-1799. Mommsen Th.P., French C.J., Hochachka P.W. Sites and patterns of protein and amino acid utilization during spawning migration of salmon. Can. J. Zool., 1980, vol. 58, pp. 1785-1799. Almeida-Val V.M., Val A.L., Duncan W.P., Souza F.C., Paula-Silva M.N., Land S. Scaling effects on hypoxia tolerance in the Amazon fish Astronotus ocellatus (Perciformes: Cichlidae): contribution of tissue enzyme levels. Comp. Biochem. Physiol. B. Biochem. Mol. Biol., 2000, vol. 125, no. 2, pp. 219-226. Almeida-Val V.M., Val A.L., Duncan W.P., Souza F.C., Paula-Silva M.N., Land S. Scaling effects on hypoxia tolerance in the Amazon fish Astronotus ocellatus (Perciformes: Cichlidae): contribution of tissue enzyme levels. Comp. Biochem. Physiol. B. Biochem. Mol. Biol., 2000, vol. 125, no. 2, pp. 219-226. Panepucci L., Fernandes M.N., Sanches J.R., Rantin F.T. Changes in lactate dehydrogenase and malate dehydrogenase activities during hypoxia and after temperature acclimation in the armored fish, Rhinelepis strigosa (Siluriformes, Loricariidae). Rev. Bras. Biol., 2000, vol. 60, no. 2, pp. 353-360. Panepucci L., Fernandes M.N., Sanches J.R., Rantin F.T. Changes in lactate dehydrogenase and malate dehydrogenase activities during hypoxia and after temperature acclimation in the armored fish, Rhinelepis strigosa (Siluriformes, Loricariidae). Rev. Bras. Biol., 2000, vol. 60, no. 2, pp. 353-360. Skorkowski E.F. Mitochondrial malic enzyme from crustacean and fish muscle. Comp. Biochem. Physiol., 1988, vol. 90 B, pp. 19-24. Skorkowski E.F. Mitochondrial malic enzyme from crustacean and fish muscle. Comp. Biochem. Physiol., 1988, vol. 90 B, pp. 19-24. Брагин Е.О., Дергунов А.Д. и др. Роль фосфолипазы А2 в аноксическом повреждении энергозависимых функций митохондрий. Вопр. мед. Химии, 1977, т. 23, № 5, с. 673-677. Bragin E.O., Dergunov A.D. i dr. Rol' fosfolipazy A2 v anoksicheskom povrezhdenii energozavisimyh funkciy mitohondriy. Vopr. med. Himii, 1977, t. 23, № 5, s. 673-677. Abele D., Puntarulo S. Formation of reactive species and induction of antioxidant defence systems in polar and temperate marine invertebrates and fish. Comp. Biochem. Physiol. A. Mol. Integr. Physiol., 2004, vol. 138, no. 4, pp. 405-415. Abele D., Puntarulo S. Formation of reactive species and induction of antioxidant defence systems in polar and temperate marine invertebrates and fish. Comp. Biochem. Physiol. A. Mol. Integr. Physiol., 2004, vol. 138, no. 4, pp. 405-415. Hermes-Lima M., Zenteno-Savin T. Animal response to drastic changes in oxygen availability and physiological oxidative stress (review). Comp. Biochem. Physiol. C. Toxicol. Pharmacol, 2002, vol. 133, no. 4, pp. 537-556. Hermes-Lima M., Zenteno-Savin T. Animal response to drastic changes in oxygen availability and physiological oxidative stress (review). Comp. Biochem. Physiol. C. Toxicol. Pharmacol, 2002, vol. 133, no. 4, pp. 537-556. Hattink J., De Boeck G., Blust R. Toxicity, accumulation, and retention of zinc by carp under normoxic and hypoxic conditions. Environ. Toxicol. Chem., 2006, vol. 25, no. 1, pp. 87-96. Hattink J., De Boeck G., Blust R. Toxicity, accumulation, and retention of zinc by carp under normoxic and hypoxic conditions. Environ. Toxicol. Chem., 2006, vol. 25, no. 1, pp. 87-96.