黄姑鱼锰超氧化物歧化酶基因的克隆及氨氮和亚硝态氮胁迫对其表达的影响
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摘要
锰超氧化物歧化酶(MnSOD)是一种含金属辅基的抗氧化酶,广泛存在于各种需氧生物中,能将氧自由基快速歧化为分子氧(O_2)和过氧化氢(H_2O_2)。本研究首次获得了黄姑鱼MnSOD基因的cDNA序列,其全长958 bp,包括47 bp的5′端非编码区(untranslated region,UTR)、233 bp的3′UTR和678 bp的开放阅读框(open reading frame,ORF),编码225个氨基酸残基(aa)。氨基酸序列分析显示,MnSOD含有一条信号肽序列(1～27 aa),4个Mn结合位点(His 53、101、190和Asp 186)和一条保守的锰/铁SOD特征序列(186～193 aa)。系统进化树分析显示,黄姑鱼MnSOD在进化上与大黄鱼最近,并与其他鱼类(斜带石斑鱼、暗纹东方鲀、牙鲆、斑马鱼和日本鳗鲡)聚为一支。荧光定量PCR检测显示,MnSOD基因在所检测的11个黄姑鱼组织/器官中均有表达,其中心脏中表达量最高,其次为脑、肝脏、鳃、中肾、肠、胃、头肾、肌肉和鳔,在脾脏中表达量最低。氨氮和亚硝态氮对黄姑鱼的急性毒性实验显示,黄姑鱼对氨氮胁迫更为敏感,其氨氮和亚硝态氮的96 h半致死浓度(LC50)分别为20.23 mg/L (换算成非离子氨0.57 mg/L)和99.08 mg/L,安全浓度分别为2.02 mg/L (换算成非离子氨0.06 mg/L)和9.91 mg/L。此外,黄姑鱼经氨氮和亚硝态氮急性攻毒后,其肝脏、鳃和头肾中MnSOD基因的表达水平均不同程度上调,推测MnSOD的上调是为了及时清除由氨氮和亚硝态氮刺激产生的氧自由基,或可用作水体污染检测的早期生物标志物。
Manganese superoxide dismutase(MnSOD), a transition-metal-containing antioxidant enzyme that dismutates the superoxide radicals to either ordinary molecular oxygen(O_2) or hydrogen peroxide(H_2O_2), widely existed in many aerobic organisms. In this study, the cDNA of Mn SOD, identified from Nibea albiflora, was 958 bp in length including 5′-untranslated region(UTR) of 47 bp, 3′-UTR of 233 bp and an open reading frame(ORF) of678 bp encoding a polypeptide of 225 amino acids. The deduced amino acid sequence analysis showed that MnSOD contains a putative signal peptide in the N-terminus(1-27 aa), four Mn binding sites(His 53, 101, 190 and Asp 186) and a Mn/Fe SOD signature sequence(186-193 aa). Phylogenetic tree analysis indicated that N. albiflora has the closest relationship with Larimichthys crocea, and was classified into the same cluster with other teleosts(Epinephelus coioides, Takifugu fasciatus, Paralichthys olivaceus, Danio rerio and Anguilla japonica). Quantitative real-time qRT-PCR analysis showed that the mRNA transcripts of MnSOD were detected in all examined tissues and the predominant distribution was in heart, followed by brain, liver, gill, kidney, intestine, stomach,head-kidney, muscle and swim bladder, and the minimum level was displayed in spleen. Moreover, during the acute toxic experiment of ammonia nitrogen or nitrite nitrogen, N. albiflora showed more sensitive to ammonia nitrogen, and the 96 h median lethal concentration(LC_(50)) value was found to be 20.23 mg/L for ammonia nitrogen(or 0.57 mg/L for non-ionic ammonia) with safe concentration(SC) values of 2.02 mg/L(or 0.06 mg/L for nonionic ammonia) and 99.08 mg/L with SC of 9.91 mg/L for nitrite nitrogen, successively. Furthermore, the temporal expression of MnSOD was significantly up-regulated in the liver, gill and head-kidney of N. albiflora. The expression features of the MnSOD suggested its important role in scavenging oxygen free radicals caused by ammonia nitrogen/nitrite nitrogen and could be used as an early biomarker for detection of environmental pollution.
Manganese superoxide dismutase(MnSOD), a transition-metal-containing antioxidant enzyme that dismutates the superoxide radicals to either ordinary molecular oxygen(O_2) or hydrogen peroxide(H_2O_2), widely existed in many aerobic organisms. In this study, the cDNA of Mn SOD, identified from Nibea albiflora, was 958 bp in length including 5′-untranslated region(UTR) of 47 bp, 3′-UTR of 233 bp and an open reading frame(ORF) of678 bp encoding a polypeptide of 225 amino acids. The deduced amino acid sequence analysis showed that MnSOD contains a putative signal peptide in the N-terminus(1-27 aa), four Mn binding sites(His 53, 101, 190 and Asp 186) and a Mn/Fe SOD signature sequence(186-193 aa). Phylogenetic tree analysis indicated that N. albiflora has the closest relationship with Larimichthys crocea, and was classified into the same cluster with other teleosts(Epinephelus coioides, Takifugu fasciatus, Paralichthys olivaceus, Danio rerio and Anguilla japonica). Quantitative real-time qRT-PCR analysis showed that the mRNA transcripts of MnSOD were detected in all examined tissues and the predominant distribution was in heart, followed by brain, liver, gill, kidney, intestine, stomach,head-kidney, muscle and swim bladder, and the minimum level was displayed in spleen. Moreover, during the acute toxic experiment of ammonia nitrogen or nitrite nitrogen, N. albiflora showed more sensitive to ammonia nitrogen, and the 96 h median lethal concentration(LC_(50)) value was found to be 20.23 mg/L for ammonia nitrogen(or 0.57 mg/L for non-ionic ammonia) with safe concentration(SC) values of 2.02 mg/L(or 0.06 mg/L for nonionic ammonia) and 99.08 mg/L with SC of 9.91 mg/L for nitrite nitrogen, successively. Furthermore, the temporal expression of MnSOD was significantly up-regulated in the liver, gill and head-kidney of N. albiflora. The expression features of the MnSOD suggested its important role in scavenging oxygen free radicals caused by ammonia nitrogen/nitrite nitrogen and could be used as an early biomarker for detection of environmental pollution.
引文
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[3]韩春艳,郑清梅,陈桂丹,等.氨氮胁迫对奥尼罗非鱼非特异性免疫的影响[J].南方水产科学,2014,10(3):47-52.Han C Y,Zheng Q M,Chen G D,et al.Effect of ammonia-N stress on non-specific immunity of tilapia(Oreochromis niloticus×O.areus)[J].South China Fisheries Science,2014,10(3):47-52(in Chinese).
[4]Benli A?K,K?ksal G,?zkul A.Sublethal ammonia exposure of Nile tilapia(Oreochromis niloticus L.):effects on gill,liver and kidney histology[J].Chemosphere,2008,72(9):1355-1358.
[5]Sinha A K,AbdElgawad H,Giblen T,et al.Antioxidative defences are modulated differentially in three freshwater teleosts in response to ammonia-induced oxidative stress[J].PLoS One,2014,9(4):e95319.
[6]Jia R,Han C,Lei J L,et al.Effects of nitrite exposure on haematological parameters,oxidative stress and apoptosis in juvenile turbot(Scophthalmus maximus)[J].Aquatic Toxicology,2015,169:1-9.
[7]Tucker C S,Francis-Floyd R,Beleau M H.Nitriteinduced anemia in channel catfish,Ictalurus punctatus rafinesque[J].Bulletin of Environmental Contamination and Toxicology,1989,43(2):295-301.
[8]Moraes G,Avilez I M,Hori T S.Comparison between biochemical responses of the teleost pacu and its hybrid tambacu(Piaractus mesopotamicus×Colossoma macro- pomum)to short-term nitrite exposure[J].Brazilian Journal of Biology,2006,66(4):1103-1108.
[9]Devasagayam T P A,Tilak J C,Boloor K K,et al.Free radicals and antioxidants in human health:current status and future prospects[J].Journal of the Association of Physicians of India,2004,52:794-804.
[10]Hwang Y S,Jeong M,Park J S,et al.Interleukin-1βstimulates IL-8 expression through MAP kinase and ROS signaling in human gastric carcinoma cells[J].Oncogene,2004,23(39):6603-6611.
[11]Winston G W.Oxidants and antioxidants in aquatic animals[J].Comparative Biochemistry and PhysiologyPart C:Comparative Pharmacology,1991,100(1-2):173-176.
[12]李建喜,杨志强,王学智.活性氧自由基在动物机体内的生物学作用[J].动物医学进展,2006,27(10):33-36.Li J X,Yang Z Q,Wang X Z.Biological function of reactive oxygen free radicals in animals[J].Progress in Veterinary Medicine,2006,27(10):33-36(in Chinese).
[13]Jacob R A.The integrated antioxidant system[J].Nutrition Research,1995,15(5):755-766.
[14]张克烽,张子平,陈芸,等.动物抗氧化系统中主要抗氧化酶基因的研究进展[J].动物学杂志,2007,42(2):153-160.Zhang K F,Zhang Z P,Chen Y,et al.Antioxidant defense system in animals[J].Chinese Journal of Zoology,2007,42(2):153-160(in Chinese).
[15]Yost F J Jr,Fridovich I.An iron-containing superoxide dismutase from Escherichia coli[J].The Journal of Biological Chemistry,1973,248(14):4905-4908.
[16]Wuerges J,Lee J W,Yim Y I,et al.Crystal structure of nickel-containing superoxide dismutase reveals another type of active site[J].Proceedings of the National Academy of Sciences of the United States of America,2004,101(23):8569-8574.
[17]Gómez-Anduro G A,Barillas-Mury C V,PeregrinoUriarte A B,et al.The cytosolic manganese superoxide dismutase from the shrimp Litopenaeus vannamei:molecular cloning and expression[J].Developmental&Comparative Immunology,2006,30(10):893-900.
[18]Dhar S K,St Clair D K.Manganese superoxide dismutase regulation and cancer[J].Free Radical Biology and Medicine,2012,52(11-12):2209-2222.
[19]Giulivi C,Boveris A,Cadenas E.The steady-state concentrations of oxygen radicals in mitochondria[M]//Gilbert D L,Colton C A.Reactive Oxygen Species in Biological Systems.Boston,MA:Springer,2002:77-102.
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[21]Li S J,Yan T,Yang J Q,et al.The role of cellular glutathione peroxidase redox regulation in the suppression of tumor cell growth by manganese superoxide dismutase[J].Cancer Research,2000,60(14):3927-3939.
[22]Wang M,Kirk J S,Venkataraman S,et al.Manganese superoxide dismutase suppresses hypoxic induction of hypoxia-inducible factor-1αand vascular endothelial growth factor[J].Oncogene,2005,24(55):8154-8166.
[23]Miriyala S,Spasojevic I,Tovmasyan A,et al.Manganese superoxide dismutase,MnSOD and its mimics[J].Biochimica et Biophysica Acta(BBA)-Molecular Basis of Disease,2012,1822(5):794-814.
[24]Pérez-Sánchez J,Borrel M,Bermejo-Nogales A,et al.Dietary oils mediate cortisol kinetics and the hepatic mRNA expression profile of stress-responsive genes in gilthead sea bream(Sparus aurata)exposed to crowding stress.Implications on energy homeostasis and stress susceptibility[J].Comparative Biochemistry and Physiology Part D:Genomics and Proteomics,2013,8(2):123-130.
[25]Arockiaraj J,Palanisamy R,Bhatt P,et al.A novel murrel Channa striatus mitochondrial manganese superoxide dismutase:gene silencing,SOD activity,superoxide anion production and expression[J].Fish Physiology and Biochemistry,2014,40(6):1937-1955.
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