A Review:Toxicity of Ammonia-N to Fish and Detoxification Strategy of Fish
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摘要
In natural water, the content of ammonia nitrogen is always kept at a low level. However, the ammonia nitrogen content in water will be elevated or even excessive when the density of aquaculture is too large or exogenous ammonia are discharged into the water, causing a series of ad-verse effects on fish survival. Within a certain range, fishes reduce ammonia production in vivo by converting ammonia into glutamine and urea, and increase NH3 volatilization and NH4+emissions to ease ammonia toxicity, but the ability to alleviate ammonia poisoning vary with fish species and size. The ammonia nitrogen sources, existence form, factors affecting toxicity, toxic mechanism and detoxification strategy of fish are reviewed, in or-der to provide reference for improving the ability of fish to alleviate ammonia nitrogen toxicity and healthy farming.
In natural water, the content of ammonia nitrogen is always kept at a low level. However, the ammonia nitrogen content in water will be elevated or even excessive when the density of aquaculture is too large or exogenous ammonia are discharged into the water, causing a series of ad-verse effects on fish survival. Within a certain range, fishes reduce ammonia production in vivo by converting ammonia into glutamine and urea, and increase NH3 volatilization and NH4+emissions to ease ammonia toxicity, but the ability to alleviate ammonia poisoning vary with fish species and size. The ammonia nitrogen sources, existence form, factors affecting toxicity, toxic mechanism and detoxification strategy of fish are reviewed, in or-der to provide reference for improving the ability of fish to alleviate ammonia nitrogen toxicity and healthy farming.
In natural water, the content of ammonia nitrogen is always kept at a low level. However, the ammonia nitrogen content in water will be elevated or even excessive when the density of aquaculture is too large or exogenous ammonia are discharged into the water, causing a series of ad-verse effects on fish survival. Within a certain range, fishes reduce ammonia production in vivo by converting ammonia into glutamine and urea, and increase NH3 volatilization and NH4+emissions to ease ammonia toxicity, but the ability to alleviate ammonia poisoning vary with fish species and size. The ammonia nitrogen sources, existence form, factors affecting toxicity, toxic mechanism and detoxification strategy of fish are reviewed, in or-der to provide reference for improving the ability of fish to alleviate ammonia nitrogen toxicity and healthy farming.
引文
[1]LI B,FAN QX,YANG K,et al.Effects of chronic ammonia stress on foraging,growth,and haematological parameters of Yellow Catfish(Pelteobagrus fulvidraco)Juveniles[J].Chinese Journal of Applied and Environmental Biology,2011,17(6):824-828.(in Chinese).
[2]XIAO W,LI DY,XU Y,et al.Effects of chronic external ammonia stress on growth,immunity and metabolism of juvenile GIFT tilapia(Oreochromis niloticus)[J].South China Fisheries Science,2015,11(4):81-87.(in Chinese).
[3]HUANG HJ.The effects of ammonia and ration size on growth and toxicological response of mullet,Lisa haematocheila[D].Shanghai:Shanghai Ocean University,2012.(in Chinese).
[4]LEMARIE G,DOSDAT A,COVES D,et al.Effect of chronic ammonia exposure on growth of European seabass(Dicentrarchus labrax)juveniles[J].Aquaculture,2004,229(1):479-491.
[5]FOSS A,SIIKAVUOPIO SI,SATHER BS,et al.Effect of chronic ammonia exposure on growth in juvenile Atlantic cod[J].Aquaculture,2004,237(1-4):179-189.
[6]KOLAREVIC J,SELSET R,FELIP O,et al.Influence of long term ammonia exposure on Atlantic salinon(Salmo salar.L)parr growth and welfare[J].Aquaculture Research,2013,44(1):1649-1664.
[7]WANG X,WANG L,YAO C,et al.Alternation of immune parameters and cellular energy allocation of Chlamys farrreri under ammonia-Nexposure and Vibrio auguillarum challenge[J].Fish&Shellfish Immunology,2012,32(5):741-749.
[8]WANG X,LI R,LIU DC,et al.Effects of Ammonia on growth and energy budgets of Redfin Puffer Taki fugu rubripes[J].Fisheries Science,2014,33(7):433-437.(in Chinese).
[9]CHEN JC,LEE Y.Effects of nitrite exposure on acid-base balance,respiratory protein,and ion concentrations of giant freshwater prawn Mocrohrochium rosenhergii at low pH[J].Archives of Environmental Contamination&Toxicology,1997,33(3):290-297.
[10]LI Q,GONG SY,LI M.Chronic ammonia toxicity induces glutamine accumulation,oxidative damage and immunosuppression of juvenile yellow catfish Pelteobagrus fulvidraco[J].Journal of Fisheries of China,2015,39(5):728-734.(in Chinese).
[11]ZHAO HT.Effects of Ammonia on haematological Immune parameters of juvenile Southern catfish(Silurus meridionalis chen)[D].Chongqing:Southwestern University,2006.(in Chinese).
[12]ZHOU X,DONG YW,WANG F,et al.The effect of high ammonia concentration on gill structure alternation and expression of sod and hsp90 genes in grass carp[J].Acta Hydrobiologica Sinica,2013,37(2):321-328.(in Chinese).
[13]ZHANG WX,SUN SM,GE XP,et al.Acute effects of ammonia exposure on histopathology of gill,liver and kidney in juvenile Megalobrama amblycephala and the post-exposure recovery[J].Journal of Fisheries of China,2015,39(2):233-244.(in Chinese).
[14]EMERSONK,RUSSO RC,LUND RE,et al.Aqueous ammonia equilibrium calculations:effect of pH and temperature[J].Journal De L’office Des Recherches Sur Les Pe cheries Du Canada,2011,32(12):2379-2383.
[15]LUO J,DU T,LIU CW,et al.Toxic effects of ammonia to Hemifusus tuba Juveniles at different ph and salinity[J].Chinese Journal of Zoology,2010,45(3):102-109.(in Chinese).
[16]WEIRICH CR,TOMASSO JR,SMITH TIJ.Toxicity of ammonia and nitrite to sunshine bass in selected environmenu[J].Journal of Aquatic Animal Health,1993,5(1):64-72.
[17]KTR M,KUMLU M,EROLDOGAN OT.Effects of temperature on acute toxicity of ammonia to Penoems.sernisulcotms,juveniles[J}.Aquaculture,2004,241(1-4):479-489.
[18]MUMMERT AK,NEVES RJ,NEWOOMB TJ,et al.Sensitivity of juvenile freshwater mussels(Lampsilis fasciola,Villosa iris)to total and un-ionized ammonia[J].Environmental Toxicology&Chemistry,2003,22(11):2545-2553.
[19]THURSTON RV,RUSSO RC,VINOGRADOVGA.Ammonia toxicity to fishes:Effect of p Hon the toxicity of the unionized ammonia species[J].Environmental Science&Technology,1981,15(7):837-840.
[20]LI B,FAN QX,ZHANG L,et al.Acute toxic effects of ammonia and nitrite on yellow catfish(Pelteobagrus fulvidraco)at different dissolved oxygen levels[J].Freshwater Fisheries,2009,39(3):31-35.(in Chinese).
[21]TANG XM,SUI Z,TIAN JB,et al.Effects of salinity on metabolic rate of juvenile turbot(Scophamus maximus)[J].South China Fisheries Science,2006,2(4):54-58.(in Chinese).
[22]CHEN JC,NAN FH.Changes of oxygen consumption and ammonia-N excretion by Peuaeus chinensis Osbeck at different temperature and salinity levels[J].Journal of Qustacean Biology,1993,13(4):706-712.
[23]CHAI XJ,HU ZH,XU JZ,et al.Effect of salinity and p H on oxygen consumption rate and ammonia excretion rate in Juvenile Nibea japonica[J].Journal of Zhejiang Ocean University(Natural Science Edition),2009,28(2):146-150.(in Chinese).
[24]JIANG M,HUANG ZQ,PENG ZR,et al.Research on ammonia excretion and oxygen consumption in Allogynogenetic sliver crucian carp[J].Journal of Shanghai Fisheries University,2007,16(1):28-32.(in Chinese).
[25]HU XD,SUN CB,CAI HX,et al.Toxicity of Ammonia-N to Penaeus monodon under the Different Salinities[J].Guangxi Sciences,2009,16(2):206-209.
[26]LV F,HUANG JT,LV LL,et al.Study on the toxicities of ammonia-n to Perinereis aibuhitensis under different salinities[J].Journal of Anhui Agricultural Sciences,2009,37(7):2986-2987.(in Chinese).
[27]RANDALL DJ,TSUI TK.Ammonia toxicity in fish[J].Marine Pollution Bulletin,2002,45(1-12):17-23.
[28]LIU Y.Effects of environmental factors on toxicity of ammonia nitrogen and calibration of aquatic biological benchmarks for ammonia nitrogen[D].Nanjing:Nanjing University,2014.(in Chinese).
[29]EPA-822-D-09-001,Draft 2009 update of aquatic life ambient water quality criteria for ammonia:Freshwater[S].Washington DC,2009.
[30]GB11607-89,Water quality standard for fisheries[S].Beijing:China Standards Press,1990.(in Chinese).
[31]IP YK,CHEW SF,RANDALL DJ.Ammonia toxicity,tolerance,and excretion[J].Fish Physiology,2001,20(1):148.
[32]BENLI ACK,KOKSAL G,OZKUL A.Sublethal ammonia exposure of Nile tilapia(Oreochromis niloticus L):Effects on gill,liver and kidney histology[J].Chemosphere,2008,72(9):1355-1358.
[33]WILKIE MP,PAMENTER ME,DUQUETTES,et al.The relationship between NMDA receptor function and the high ammonia tolerance of anoxia-tolerant goldfish[J].Journal of Experimental Biology,2011,214(24):4107-4120.
[34]LI B,FAN QX,YANG K,et al.Effects of chronic ammonia stress on foraging,growth,and haematological parameters of yellow catfish(Pelteobagrus fulvidraco)Juveniles[J].Chinese Journal of Applied and Environmental Biology,2011(6):824-828.(in Chinese).
[35]QIAO SF,LI HS.Causes and control techniques of acute ammonia poisoning in water[J].Hebei Fisheries,2005(2):27-29.(in Chinese).
[36]WEE NLJ,TNG YYM,CHENG HT,et al.Ammonia toxicity and tolerance in the brain of the African sharptooth catfish,Clarias gariepinus[J].Aquatic Toxicology,2007,82(3):204-213.
[37]JAYAKUMAR AR,NORENBERG MD.The Na-K-Cl co-transporter in astrocyte swelling[J].Metabolic Brain Disease,2010,25(1):31-38.
[38]VEAUVY CM,MCDONALD MD,VAN AU-DEKERKE J,et al.Ammonia affects brain nitrogen metabolism but not hydration status in the gulf toadfish(Opsanus beta)[J].Aquatic Toxicology,2005,74(1):32-46.
[39]RAO VLR.Nitric oxide in hepatic encephalopathy and hyperammonemia[J].Neurochemistry International,2002,41(2-3):161-170.
[40]RODRIGO R,ERCEG S,FELIPO V.Neurons exposed to ammonia reproduce the differential alteration in nitric oxide modulation of guanylate cyclase in the cerebellum and cortex of patients with liver cirrhosis[J].Neurobiology of Disease,2005,19(1-2):150-161.
[41]RODRIGO R,CAULI O,BOIX J,et al.Role of NMDA receptors in acute liver failure and ammonia toxicity:therapeutical implications[J].Neurochemistry International,2009,55(1-3):113-118.
[42]BRAISSANT O,MCLIN VA,CLRDALBU C.Ammonia toxicity to the brain[J].Journal of Inherited Metabolic Disease,2012,36(4):595-612.
[43]KLEJMANA,WEGRZYNOWICZ M,SZAT-MARI EM,et al.Mechanisms of ammoniainduced cell death in rat cortical neurons:Roles of NMDA receptors and glutathione[J].Neurochemistry International,2005,47(1-2):51-57.
[44]BRUSILOW SW.Hyperammonemic encephalopathy[J].Medicine,2002,81(3):240-249.
[45]TSUI TKN,RANDALL DJ,HANSON L,et al.Dogmas and controversies in the handling of nitrogenous wastes:ammonia tolerance in the oriental weatherloach Misgurnus anguillicoudotus[J].Journal of Experimental Biology,2004,207(12):1977-1983.
[46]IP YK,LEONG MWF,SIM MY,et al.Chronic and acute ammonia toxicity in mudskippers,Periophthalmodon schlosseri and Boleophtholmus hoddoerti:brain ammonia and glutamine contents,and effects of methionine sulfoximine and MK801[J].Journal of Experimental Biology,2005,208(10):1993-2004.
[47]SINHA AK,GIBLENT,ABDELGAWAD H,et al.Regulation of amino acid metabolism as a defensive strategy in the brain of three freshwater teleosts in response to high environmental ammonia exposure[J].Aquatic Toxicology,2013,130-131(2):86-96.
[48]PART P,WRIGHT PA,WOOD CM.Urea and water permeability in dogfish(Squalus acanthias)gills[J].Compartitive Biochemistry&Physiology Part A Molecular&Integrative Physiology,1998,119(1):117-123.
[49]WILSON JM,IWATA K,IWAMA GK,et al.Inhibition of ammonia excretion and production in rainbow trout during severe alkaline exposure[J].Comparative Biochemistry&Physiology B Comparative Biochemistry,1998,121(1):99-109.
[50]LIM CB,CHEW SF,ANDERSON PM,et al.Reduction in the rates of protein and amino acid catabolism to slow down the accumulation of endogenous ammonia:a strategy potentially adopted by mudskippers(Periophtholmodon schlosseri and Boleophtholmus boddoerti)during aerial exposure in constant darkness[J].Journal of Experimental Biology,2001,204(9):1605-1614.
[51]TSUI TK,RANDALL DJ,CHEW SF,et al.Accumulation of ammonia in the body and NH3volatilization from alkaline regions of the body surface during ammonia loading and exposure to air in the weather loach Misgurnus anguilicaudatus[J].Journal of Experimental Biology,2002,205(5):651-659.
[52]CHEW SF,JIN Y,IP Y.The loach Misgurnus anguilicaudatus reduces amino acid catabolism and accumulates alanine and glutamine during aerial exposure[J].Physiol Biochem Zool,2001,74(2):226-237.
[53]WILSON RW,TAYLOR EW.Transbranchial ammonia gradiems and acid-base responses to high external ammonia concentration in rainbow trout(Oncorhyuchus mykiss)acclimated to different salinities[J].Journal of Experimental Biology,1992,166(1):95-112.
[54]RANDALL DJ,WILSON JM,PENG KW,et al.The mudskipper,Periophtholmodon schloesseri,actively transports NHq’against a concentration gradient[J].American Journal of Physiology,2000,277(6):1562-1567.
[55]WILSON JM,RANDALL DJ,DONOWITZM,et al.Immunolocalization of ion-transport proteins to branchial epithelium mitochondriarich cells in the mudskipper(Periophtholmodon schlosseri)[J].Journal of Experimental Biology,2000,203(15):2297-2310.
[56]LIU H,XIE S,ZHU X,et al.Effects of dietary ascorbic acid supplementation on the growth performance,immune and stress response in juvenile Leiocassis longirostris,Gunther exposed to ammonia[J].Aquaculture Research,2008,39(15):1628-1638.
[57]CHEN XM.Effects of V-aminoJ-acid on growth,immunity and resistance to ammonia nitrogen stress of Jian Cyprinus carpio[D].Changchun:Jilin Agricultural University,2015.(in Chinese).
[58]ZHANG YL.Effects of vitamin e on growth,immunity and stress resistance of Juvenile moire Grouper[D].Shanghai:Shanghai Ocean University,2015.(in Chinese).
[59]HU Y,HUANG Y,WEN H,et al.Effect of vitamin C on growth,immunity and anti-ammonia-nitrite stress ability in juvenile black carp(Mylopharyngodon piceus)[J].Journal of Fisheries of China,2013,37(4)565-573.(in Chinese).
[60]LUO N,DING ZL,ZHANG YX,et al.Effects of dietary linolenic acid content on growth,antioxidant capacity,non-specific immunity and anti-ammonia-nitrite stress ability of oriental river prawn(Macrobrachium nipponense)[J].Acta Zoonutrimenta Sinica,2017,29(1):134-146.(in Chinese).
[61]SONG C,NIU CJ,CHEN XR.The effect of chitosan on some non-specific immune indices of the juvenile soft-shelled turtle Pelodiscus sinensis[J].Acta Zoologica Sinica,2007,53(2):270-277.(in Chinese).
[2]XIAO W,LI DY,XU Y,et al.Effects of chronic external ammonia stress on growth,immunity and metabolism of juvenile GIFT tilapia(Oreochromis niloticus)[J].South China Fisheries Science,2015,11(4):81-87.(in Chinese).
[3]HUANG HJ.The effects of ammonia and ration size on growth and toxicological response of mullet,Lisa haematocheila[D].Shanghai:Shanghai Ocean University,2012.(in Chinese).
[4]LEMARIE G,DOSDAT A,COVES D,et al.Effect of chronic ammonia exposure on growth of European seabass(Dicentrarchus labrax)juveniles[J].Aquaculture,2004,229(1):479-491.
[5]FOSS A,SIIKAVUOPIO SI,SATHER BS,et al.Effect of chronic ammonia exposure on growth in juvenile Atlantic cod[J].Aquaculture,2004,237(1-4):179-189.
[6]KOLAREVIC J,SELSET R,FELIP O,et al.Influence of long term ammonia exposure on Atlantic salinon(Salmo salar.L)parr growth and welfare[J].Aquaculture Research,2013,44(1):1649-1664.
[7]WANG X,WANG L,YAO C,et al.Alternation of immune parameters and cellular energy allocation of Chlamys farrreri under ammonia-Nexposure and Vibrio auguillarum challenge[J].Fish&Shellfish Immunology,2012,32(5):741-749.
[8]WANG X,LI R,LIU DC,et al.Effects of Ammonia on growth and energy budgets of Redfin Puffer Taki fugu rubripes[J].Fisheries Science,2014,33(7):433-437.(in Chinese).
[9]CHEN JC,LEE Y.Effects of nitrite exposure on acid-base balance,respiratory protein,and ion concentrations of giant freshwater prawn Mocrohrochium rosenhergii at low pH[J].Archives of Environmental Contamination&Toxicology,1997,33(3):290-297.
[10]LI Q,GONG SY,LI M.Chronic ammonia toxicity induces glutamine accumulation,oxidative damage and immunosuppression of juvenile yellow catfish Pelteobagrus fulvidraco[J].Journal of Fisheries of China,2015,39(5):728-734.(in Chinese).
[11]ZHAO HT.Effects of Ammonia on haematological Immune parameters of juvenile Southern catfish(Silurus meridionalis chen)[D].Chongqing:Southwestern University,2006.(in Chinese).
[12]ZHOU X,DONG YW,WANG F,et al.The effect of high ammonia concentration on gill structure alternation and expression of sod and hsp90 genes in grass carp[J].Acta Hydrobiologica Sinica,2013,37(2):321-328.(in Chinese).
[13]ZHANG WX,SUN SM,GE XP,et al.Acute effects of ammonia exposure on histopathology of gill,liver and kidney in juvenile Megalobrama amblycephala and the post-exposure recovery[J].Journal of Fisheries of China,2015,39(2):233-244.(in Chinese).
[14]EMERSONK,RUSSO RC,LUND RE,et al.Aqueous ammonia equilibrium calculations:effect of pH and temperature[J].Journal De L’office Des Recherches Sur Les Pe cheries Du Canada,2011,32(12):2379-2383.
[15]LUO J,DU T,LIU CW,et al.Toxic effects of ammonia to Hemifusus tuba Juveniles at different ph and salinity[J].Chinese Journal of Zoology,2010,45(3):102-109.(in Chinese).
[16]WEIRICH CR,TOMASSO JR,SMITH TIJ.Toxicity of ammonia and nitrite to sunshine bass in selected environmenu[J].Journal of Aquatic Animal Health,1993,5(1):64-72.
[17]KTR M,KUMLU M,EROLDOGAN OT.Effects of temperature on acute toxicity of ammonia to Penoems.sernisulcotms,juveniles[J}.Aquaculture,2004,241(1-4):479-489.
[18]MUMMERT AK,NEVES RJ,NEWOOMB TJ,et al.Sensitivity of juvenile freshwater mussels(Lampsilis fasciola,Villosa iris)to total and un-ionized ammonia[J].Environmental Toxicology&Chemistry,2003,22(11):2545-2553.
[19]THURSTON RV,RUSSO RC,VINOGRADOVGA.Ammonia toxicity to fishes:Effect of p Hon the toxicity of the unionized ammonia species[J].Environmental Science&Technology,1981,15(7):837-840.
[20]LI B,FAN QX,ZHANG L,et al.Acute toxic effects of ammonia and nitrite on yellow catfish(Pelteobagrus fulvidraco)at different dissolved oxygen levels[J].Freshwater Fisheries,2009,39(3):31-35.(in Chinese).
[21]TANG XM,SUI Z,TIAN JB,et al.Effects of salinity on metabolic rate of juvenile turbot(Scophamus maximus)[J].South China Fisheries Science,2006,2(4):54-58.(in Chinese).
[22]CHEN JC,NAN FH.Changes of oxygen consumption and ammonia-N excretion by Peuaeus chinensis Osbeck at different temperature and salinity levels[J].Journal of Qustacean Biology,1993,13(4):706-712.
[23]CHAI XJ,HU ZH,XU JZ,et al.Effect of salinity and p H on oxygen consumption rate and ammonia excretion rate in Juvenile Nibea japonica[J].Journal of Zhejiang Ocean University(Natural Science Edition),2009,28(2):146-150.(in Chinese).
[24]JIANG M,HUANG ZQ,PENG ZR,et al.Research on ammonia excretion and oxygen consumption in Allogynogenetic sliver crucian carp[J].Journal of Shanghai Fisheries University,2007,16(1):28-32.(in Chinese).
[25]HU XD,SUN CB,CAI HX,et al.Toxicity of Ammonia-N to Penaeus monodon under the Different Salinities[J].Guangxi Sciences,2009,16(2):206-209.
[26]LV F,HUANG JT,LV LL,et al.Study on the toxicities of ammonia-n to Perinereis aibuhitensis under different salinities[J].Journal of Anhui Agricultural Sciences,2009,37(7):2986-2987.(in Chinese).
[27]RANDALL DJ,TSUI TK.Ammonia toxicity in fish[J].Marine Pollution Bulletin,2002,45(1-12):17-23.
[28]LIU Y.Effects of environmental factors on toxicity of ammonia nitrogen and calibration of aquatic biological benchmarks for ammonia nitrogen[D].Nanjing:Nanjing University,2014.(in Chinese).
[29]EPA-822-D-09-001,Draft 2009 update of aquatic life ambient water quality criteria for ammonia:Freshwater[S].Washington DC,2009.
[30]GB11607-89,Water quality standard for fisheries[S].Beijing:China Standards Press,1990.(in Chinese).
[31]IP YK,CHEW SF,RANDALL DJ.Ammonia toxicity,tolerance,and excretion[J].Fish Physiology,2001,20(1):148.
[32]BENLI ACK,KOKSAL G,OZKUL A.Sublethal ammonia exposure of Nile tilapia(Oreochromis niloticus L):Effects on gill,liver and kidney histology[J].Chemosphere,2008,72(9):1355-1358.
[33]WILKIE MP,PAMENTER ME,DUQUETTES,et al.The relationship between NMDA receptor function and the high ammonia tolerance of anoxia-tolerant goldfish[J].Journal of Experimental Biology,2011,214(24):4107-4120.
[34]LI B,FAN QX,YANG K,et al.Effects of chronic ammonia stress on foraging,growth,and haematological parameters of yellow catfish(Pelteobagrus fulvidraco)Juveniles[J].Chinese Journal of Applied and Environmental Biology,2011(6):824-828.(in Chinese).
[35]QIAO SF,LI HS.Causes and control techniques of acute ammonia poisoning in water[J].Hebei Fisheries,2005(2):27-29.(in Chinese).
[36]WEE NLJ,TNG YYM,CHENG HT,et al.Ammonia toxicity and tolerance in the brain of the African sharptooth catfish,Clarias gariepinus[J].Aquatic Toxicology,2007,82(3):204-213.
[37]JAYAKUMAR AR,NORENBERG MD.The Na-K-Cl co-transporter in astrocyte swelling[J].Metabolic Brain Disease,2010,25(1):31-38.
[38]VEAUVY CM,MCDONALD MD,VAN AU-DEKERKE J,et al.Ammonia affects brain nitrogen metabolism but not hydration status in the gulf toadfish(Opsanus beta)[J].Aquatic Toxicology,2005,74(1):32-46.
[39]RAO VLR.Nitric oxide in hepatic encephalopathy and hyperammonemia[J].Neurochemistry International,2002,41(2-3):161-170.
[40]RODRIGO R,ERCEG S,FELIPO V.Neurons exposed to ammonia reproduce the differential alteration in nitric oxide modulation of guanylate cyclase in the cerebellum and cortex of patients with liver cirrhosis[J].Neurobiology of Disease,2005,19(1-2):150-161.
[41]RODRIGO R,CAULI O,BOIX J,et al.Role of NMDA receptors in acute liver failure and ammonia toxicity:therapeutical implications[J].Neurochemistry International,2009,55(1-3):113-118.
[42]BRAISSANT O,MCLIN VA,CLRDALBU C.Ammonia toxicity to the brain[J].Journal of Inherited Metabolic Disease,2012,36(4):595-612.
[43]KLEJMANA,WEGRZYNOWICZ M,SZAT-MARI EM,et al.Mechanisms of ammoniainduced cell death in rat cortical neurons:Roles of NMDA receptors and glutathione[J].Neurochemistry International,2005,47(1-2):51-57.
[44]BRUSILOW SW.Hyperammonemic encephalopathy[J].Medicine,2002,81(3):240-249.
[45]TSUI TKN,RANDALL DJ,HANSON L,et al.Dogmas and controversies in the handling of nitrogenous wastes:ammonia tolerance in the oriental weatherloach Misgurnus anguillicoudotus[J].Journal of Experimental Biology,2004,207(12):1977-1983.
[46]IP YK,LEONG MWF,SIM MY,et al.Chronic and acute ammonia toxicity in mudskippers,Periophthalmodon schlosseri and Boleophtholmus hoddoerti:brain ammonia and glutamine contents,and effects of methionine sulfoximine and MK801[J].Journal of Experimental Biology,2005,208(10):1993-2004.
[47]SINHA AK,GIBLENT,ABDELGAWAD H,et al.Regulation of amino acid metabolism as a defensive strategy in the brain of three freshwater teleosts in response to high environmental ammonia exposure[J].Aquatic Toxicology,2013,130-131(2):86-96.
[48]PART P,WRIGHT PA,WOOD CM.Urea and water permeability in dogfish(Squalus acanthias)gills[J].Compartitive Biochemistry&Physiology Part A Molecular&Integrative Physiology,1998,119(1):117-123.
[49]WILSON JM,IWATA K,IWAMA GK,et al.Inhibition of ammonia excretion and production in rainbow trout during severe alkaline exposure[J].Comparative Biochemistry&Physiology B Comparative Biochemistry,1998,121(1):99-109.
[50]LIM CB,CHEW SF,ANDERSON PM,et al.Reduction in the rates of protein and amino acid catabolism to slow down the accumulation of endogenous ammonia:a strategy potentially adopted by mudskippers(Periophtholmodon schlosseri and Boleophtholmus boddoerti)during aerial exposure in constant darkness[J].Journal of Experimental Biology,2001,204(9):1605-1614.
[51]TSUI TK,RANDALL DJ,CHEW SF,et al.Accumulation of ammonia in the body and NH3volatilization from alkaline regions of the body surface during ammonia loading and exposure to air in the weather loach Misgurnus anguilicaudatus[J].Journal of Experimental Biology,2002,205(5):651-659.
[52]CHEW SF,JIN Y,IP Y.The loach Misgurnus anguilicaudatus reduces amino acid catabolism and accumulates alanine and glutamine during aerial exposure[J].Physiol Biochem Zool,2001,74(2):226-237.
[53]WILSON RW,TAYLOR EW.Transbranchial ammonia gradiems and acid-base responses to high external ammonia concentration in rainbow trout(Oncorhyuchus mykiss)acclimated to different salinities[J].Journal of Experimental Biology,1992,166(1):95-112.
[54]RANDALL DJ,WILSON JM,PENG KW,et al.The mudskipper,Periophtholmodon schloesseri,actively transports NHq’against a concentration gradient[J].American Journal of Physiology,2000,277(6):1562-1567.
[55]WILSON JM,RANDALL DJ,DONOWITZM,et al.Immunolocalization of ion-transport proteins to branchial epithelium mitochondriarich cells in the mudskipper(Periophtholmodon schlosseri)[J].Journal of Experimental Biology,2000,203(15):2297-2310.
[56]LIU H,XIE S,ZHU X,et al.Effects of dietary ascorbic acid supplementation on the growth performance,immune and stress response in juvenile Leiocassis longirostris,Gunther exposed to ammonia[J].Aquaculture Research,2008,39(15):1628-1638.
[57]CHEN XM.Effects of V-aminoJ-acid on growth,immunity and resistance to ammonia nitrogen stress of Jian Cyprinus carpio[D].Changchun:Jilin Agricultural University,2015.(in Chinese).
[58]ZHANG YL.Effects of vitamin e on growth,immunity and stress resistance of Juvenile moire Grouper[D].Shanghai:Shanghai Ocean University,2015.(in Chinese).
[59]HU Y,HUANG Y,WEN H,et al.Effect of vitamin C on growth,immunity and anti-ammonia-nitrite stress ability in juvenile black carp(Mylopharyngodon piceus)[J].Journal of Fisheries of China,2013,37(4)565-573.(in Chinese).
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