花斑裸鲤GeHSP70基因的克隆及其对重金属Cu~(2+)胁迫的应答
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摘要
【目的】克隆花斑裸鲤GeHSP70基因,并进行生物信息学分析;研究正常和Cu~(2+)胁迫条件下GeHSP70的表达模式。【方法】采用分子生物学技术克隆花斑裸鲤GeHSP70基因的全长cDNA,分析其生物信息学特征。采用Real-time PCR技术分析正常和0.01mg/L Cu~(2+)胁迫(胁迫0,12,24,48h)条件下GeHSP70mRNA在花斑裸鲤肝脏、肾脏、脑和鳃4个组织中的表达情况,并采用ELISA试剂盒检测肝脏、鳃组织中GeHSP70蛋白的表达水平。【结果】花斑裸鲤GeHSP70序列共1 387bp,由123bp的5′-UTR、592bp的3′-UTR和672bp的编码区构成,编码223个氨基酸;GeHSP70蛋白是疏水性蛋白,分子质量为11.15ku,等电点为5.01,包含2个HSP70家族的特征序列:IDLGTTYS(11-18位氨基酸)和IFDLGGGTFDVSIL(199-212位氨基酸)。GeHSP70基因核苷酸序列与鲫鱼的同源性最高(93%),在系统进化树中与同科或属中鱼种聚合成一个分支,显示出高度保守性。Real-time PCR结果表明,正常条件下花斑裸鲤GeHSP70在鳃组织中的表达量最高,其次是肝脏,在脑和鳃组织中表达量较低;随着0.01mg/L Cu~(2+)胁迫时间的延长,花斑裸鲤肝脏、肾脏和鳃组织的GeHSP70mRNA表达量先升后降,在24h表达量最高,而脑组织的表达量持续升高;GeHSP70在花斑裸鲤肝脏和鳃组织的蛋白水平表达量总体均表现为先升后降,表明Cu~(2+)能够诱导GeHSP70的表达。【结论】成功克隆了花斑裸鲤GeHSP70cDNA全长,在0.01mg/L Cu~(2+)胁迫下花斑裸鲤GeHSP70在mRNA水平和蛋白水平呈规律性应答,表明GeHSP70是潜在的反映水环境质量的生物标志物。
【Objective】This study cloned Gymnocypris eckloni GeHSP70 gene and analyzed its bioinformatics and expression pattern under normal condition and Cu2+stress.【Method】The full-length cDNA of Gymnocypris eckloni GeHSP70 gene was cloned by molecular biology technology,and its bioinformatics characteristics were analyzed.The GeHSP70 mRNA expressions in liver,kidney,brain and gill under normal and 0.01 mg/L Cu2+stress(0,12,24 and 48h)were analyzed by Real-time PCR technology.TheELISA kit was used to determine the protein level of GeHSP70 in liver and gill.【Result】Gymnocypris eckloni GeHSP70 sequence is 1 387 bp in length with 123 bp 5′-UTR,592 bp 3′-UTR and 672 bp encoding region,encoding 223 amino acids.GeHSP70 is a hydrophobic protein with molecular weight of 11.15 ku and an isoelectric point of 5.01.It includes 2 HSP70 family feature sequences of IDLGTTYS(11-18 amino acid)and IFDLGGGTFDVSIL(199-212 amino acid).The nucleotide sequence of GeHSP70 gene has the highest homology with Carassius auratus(93%).Real-time PCR showed that the expression of Gymnocypris eckloni GeHSP70 gill tissue under normal conditions is the highest,followed by liver,brain and gill tissues.With the extension of 0.01mg/L Cu2+exposure,GeHSP70 mRNA expressions of Gymnocypris eckloni in liver,kidney and gill tissues firstly increased then decreased with the peak at 24 h,while the expression in brain tissue increased all the time.The protein levels in Gymnocypris eckloni liver and gill tissues first increased and then decreased,indicating that Cu2+induced the expression of GeHSP70.【Conclusion】This study successfully cloned the full-length Gymnocypris eckloni cDNA of GeHSP70,and GeHSP70 showed regular response at mRNA and protein levels to 0.01mg/L Cu2+exposure,indicating that GeHSP70 is a potential biomarker to reflect the quality of water environment.
【Objective】This study cloned Gymnocypris eckloni GeHSP70 gene and analyzed its bioinformatics and expression pattern under normal condition and Cu2+stress.【Method】The full-length cDNA of Gymnocypris eckloni GeHSP70 gene was cloned by molecular biology technology,and its bioinformatics characteristics were analyzed.The GeHSP70 mRNA expressions in liver,kidney,brain and gill under normal and 0.01 mg/L Cu2+stress(0,12,24 and 48h)were analyzed by Real-time PCR technology.TheELISA kit was used to determine the protein level of GeHSP70 in liver and gill.【Result】Gymnocypris eckloni GeHSP70 sequence is 1 387 bp in length with 123 bp 5′-UTR,592 bp 3′-UTR and 672 bp encoding region,encoding 223 amino acids.GeHSP70 is a hydrophobic protein with molecular weight of 11.15 ku and an isoelectric point of 5.01.It includes 2 HSP70 family feature sequences of IDLGTTYS(11-18 amino acid)and IFDLGGGTFDVSIL(199-212 amino acid).The nucleotide sequence of GeHSP70 gene has the highest homology with Carassius auratus(93%).Real-time PCR showed that the expression of Gymnocypris eckloni GeHSP70 gill tissue under normal conditions is the highest,followed by liver,brain and gill tissues.With the extension of 0.01mg/L Cu2+exposure,GeHSP70 mRNA expressions of Gymnocypris eckloni in liver,kidney and gill tissues firstly increased then decreased with the peak at 24 h,while the expression in brain tissue increased all the time.The protein levels in Gymnocypris eckloni liver and gill tissues first increased and then decreased,indicating that Cu2+induced the expression of GeHSP70.【Conclusion】This study successfully cloned the full-length Gymnocypris eckloni cDNA of GeHSP70,and GeHSP70 showed regular response at mRNA and protein levels to 0.01mg/L Cu2+exposure,indicating that GeHSP70 is a potential biomarker to reflect the quality of water environment.
引文
[1]申志兴,王国杰,杨成,等.青海省花斑裸鲤现状及保护对策[J].青海畜牧业,2011(2):17-19.Shen Z X,Wang G J,Yang C,et al.The status and protection measures of the Gymnocypris eckloni in Qinghai province[J].Qinghai Animal Husbandry,2011(2):17-19.
[2] Craig E A.The heat shock response[J].CRC Crit Rev Biochem,1985,18(2):39-80.
[3] Frydman J.Folding of newly translated proteins in vivo:the role of molecular chaperones[J].Annu Rev Biochem,2001,70:603-649.
[4] Lindquist S,Craig E A.The heat-shock proteins[J].Annual Review of Genetics,2007,22:631-677.
[5] Cara J B,Aluru N,Moyano F J,et al.Food deprivation induces HSP70and HSP90protein expression in larval gilthead sea bream and rainbow trout[J].Comp Biochem Physiol B Biochem Mol Biol,2005,142(4):426-431.
[6]沈骅,王晓蓉,张景飞.应用应激蛋白HSP70作为生物标志物研究锌、铜及其联合毒性对鲫鱼肝脏的影响[J].环境科学学报,2004,24(5):895-899.Shen H,Wang X R,Zhang J F.The effects of the stress protein HSP70on the liver of Crucian carp were studied as biomarkers[J].Journal of Environmental Science,2004,24(5):895-899.
[7]沈骅,王晓蓉,张景飞,等.低浓度Zn对幼龄鲫鱼肝脏组织应激蛋白HSP70诱导的影响[J].农业环境科学学报,2004,23(3):441-443.Shen H,Wang X R,Zhang J F,et al.The influence of low concentration Zn on HSP70induced by liver tissue of young Crucian carp[J].Journal of Agricultural Environmental Science,2004,23(3):441-443.
[8]沈骅,王晓蓉,张景飞,等.低浓度Pb2+、Cd2+对鲫鱼肝脏组织中HSP70诱导的影响[J].环境污染与防治,2004,26(4):244-247.Shen H,Wang X R,Zhang J F,et al.The influence of low concentration Pb2+and Cd2+on HSP70induced in the liver of Crucian carp[J].Environment Pollution and Control,2004,26(4):244-247.
[9] Sanders B M.Stress-proteins in aquatic organisms:an environmental perspective[J].Crit Rev Toxicol,1993,23:49-75.
[10]张勇,徐钢春,杜富宽,等.美洲鲥hsp70的分子特征及其对运输应激的应答[J].动物学杂志,2016,51(2):268-280.Zhang Y,Xu G C,Du F K,et al.Molecular characteristics of the Alosa sapidissima hsp70and its response to transport stress[J].Chinese Journal of Zoology,2016,51(2):268-280.
[11]祝璟琳,王国良.鱼类HSP70的研究进展[J].宁波大学学报(理工版),2007,20(4):446-450.Zhu J L,Wang G L.Research progress on fish HSP70[J].Journal of Ningbo University(Polytechnic Edition),2007,20(4):446-450.
[12] Xie Y H.Molecular characterization of the HSP70and HSP90genes in Asian clam(Corbicula fluminea)and their expression analysis during heavy metal exposure[J].Gene,2017,16(1):146-150.
[13]刘迪秋,葛锋,陈朝银,等.重金属铜、镉对鲫肝脏基因表达的影响[J].水产科学,2010,17(6):227-230.Liu D Q,Ge F,Chen C Y,et al.Effects of heavy metal copper and cadmium on the gene expression of the Crucian carpliver[J].Fisheries Science,2010,17(6):227-230.
[14]刘福军,张饮江,王明学.铜对鱼类慢性毒性研究进展[J].水生生物学报,2003,27(3):302-307.Liu F J,Zhang Y J,Wang M X.The progress of copper on chronic toxicity of fish[J].Journal of Aquatic Biology,2003,27(3):302-307.
[15]凌明圣.分子伴侣HSP70研究进展[J].国外医学分子生物学分册,1997,15(5):227-230.Ling M S.Molecular partner HSP70research progress[J].Foreign Medical Molecular Biology Branch,1997,15(5):227-230.
[16] Grajg E A.Heat shock protein and molecular chaperones:mediate,ons of proteins stress 70and chaperones 60in diversed pecies[J].Cell,1994,34:253-258.
[17] Deane E E,Woo N Y S.Cloning and characterization of the hsp70multigene family from silver sea bream:modulated gene expression between warm and cold temperature acclimation[J].Biochem Biophys Res Commun,2005,330:776-783.
[18] Demand J,Lfiders J.The carboxy-terminal domain of HSC70provides binding sites for a distinct set of chaperone cofactorsⅢ[J].Molecular and Cellular Biology,1998,18(4):2023-2028.
[19] Vayssier M,Leguerhier F,Fabian J F,et al.Cloning and analysis of a Trichinella briotovi gene encoding a cytoplasmic heat shock protein of 72kD[J].Parasitology,1999,119:81-93.
[20]刘旭东.牙鲆hsp70与hsc70基因的克隆、表达及其SNPs与耐热性状的相关性分析[D].青岛:中国海洋大学,2011.Liu X D.The correlation analysis between Paralichthys olivaceus hsp70and hsc70gene cloning,expression and SNPs and heat resistance traits[D].Qingdao:China Ocean University,2011.
[21]明建华,谢骏,刘波,等.团头鲂hsp70cDNA的克隆、序列分析以及热应激对其mRNA表达的影响[J].中国水产科学,2009,16(5):635-648.Ming J H,Xie J,Liu B,et al.The cloning,sequence analysis and thermal stress of the Megalobrama amblycephala were affected on the expression of mRNA[J].Journal of Fishery Sciences of China,2009,16(5):635-648.
[22]何珊,梁旭方,李观贵,等.鲢鱼、草鱼和尼罗罗非鱼热休克蛋白70基因cDNA全序列的克隆与分析[J].环境科学学报,2009,29(11):2324-2330.He S,Liang X F,Li G G,et al.Cloning and analysis of the gene cDNA sequence in Hypophthalmichthys molitrix,Ctenopharyngodon idella,and Oreochromis niloticus heat shock protein 70gene[J].Journal of Environmental Sciences,2009,29(11):2324-2330.
[23]刘庆全.淞江鲈(Trachidermus fasciatus Heckel)hsp70全序列的克隆及其表达的初步研究[D].上海:复旦大学,2013.Liu Q Q.A preliminary study on the Trachidermus fasciatus Heckel cloning and expression of hsp70 sequences[D].Shanghai:Fudan University,2013.
[24] Zhang A,Zhou X,Wang X,et al.Characterization of two heat shock proteins(Hsp70/Hsc70)from grass carp(Ctenopharyngodon idella):evidence for their differential gene expression,protein synthesis and secretion in LPS challenged peripheral blood lymphocytes[J].Comp Biochem Physiol B Biochem Mol Biol,2011,159(2):109-114.
[25]张娟,张其中,张占会,等.黄颡鱼HSC70基因及其组织表达分析[J].水生生物学报,2009,33(3):426-433.Zhang J,Zhang Q Z,Zhang Z H,et al.Analysis of Pseudobagrus fulvidraco HSC70 gene and its tissue expression[J].Journal of Aquatic Biology,2009,33(3):426-433.
[26] Qu L Y,Xiang J H,Sun X Q,et al.Expression analysis of HSP70in various tissues of Chlamys farreri under thermal stress[J].High Technology Letters,2005,15(5):96-100.
[27] Jing J,Liu H,Chen H,et al.Acute effect of copper and cadmium exposure on the expression of heat shock protein 70in the Cyprinidae fish Tanichthys albonubes[J].Chemosphere,2013,1(14):1-10.
[28]周小飞.草鱼热休克蛋白HSC70和HSP70在免疫反应中不同基因表达、蛋白质合成和分泌的特征探讨[C].成都:电子科技大学,2011.Zhou X F.The characteristics of different gene expression,protein synthesis and secretion of Ctenopharyngodon idella HSC70and HSP70in immune response[D].Chengdu:University of Electronic Science and Technology,2011.
[29]王云彪,侯晓丽,徐镜波.温度与铜联合胁迫下鲤鱼HSP70的组织特异性表达[C]//中国环境科学学会.中国环境科学学会学术年会论文集.北京:中国环境出版社,2011:3714-3717.Wang Y B,Hou X L,Xu J B.Tissue specific expression of tissue specific table of Cyprinus carpio HSP70under combined temperature and copper stress[C]//China Society of Environmental Sciences.Paper on the academic annual meeting of the China Society of Environmental Science.Beijing:China Environmental Press,2011:3714-3717.
[30] Mohammad M N A,Mona S Z,Elsayed A K,et al.Use of fish as bio-indicator of the effects of heavy metals pollution[J].Aquac Res Development,2015,6(4):1-13.
[31] Kim S G,Jee J H,Kang J C.Cadmium accumulation and elimination in tissues of juvenile olive flounder,Paralichthys olivaceus after sub-chronic cadmium exposure[J].Environmental Pollution,2004,127:117-123.
[32] Zhang A Y,Guo Y F,Zhang S N,et al.Cytokine effects and cellular signaling pathways of grass carp HSP70in head kidney leukocytes[J].Fish&Shellfish Immunology,2015,46:550-556.
[33] Xing H,Li S,Wang X,et al.Effects of atrazine and chlorpyrifos on the mRNA levels of HSP70and HSC70in the liver,brain,kidney and gill of common carp(Cyprinus carpio)[J].Chemosphere,2013,90:910-916.
[34] Idriss A A,Ahmad A K.Heavy metal concentrations in fishes from Juru River,estimation of the health risk[J].Bull Environ Contam Toxicol,2015,94:204-208.
[35] Drevnick P E,Sandheinrich M B,Oris J T.Increased ovarian follicular apoptosis in fathead minnows(Pimephales promelas)exposed to dietary methylmercury[J].Aquat Toxicol,2006,79:49-54.
[36] Sfakianakis D G,Renieri E,Kentouri M,et al.Effect of heavy metals on fish larvae deformities:a review[J].Enviro Res,2015,137:246-255.
[37] Adeyemo O K,Adedeji O B,Offor C C.Blood lead level as biomarker of environmental lead pollution in feral and cultured African catfish(Clarias gariepinus)[J].Nigerian Vet,2010,31:139-147.
[38] Price B D,Calderwood S K.Heat-induced transcription from RNA polymeraseⅡandⅢand HSF binding activity are coordinately regulated by the products of the heat shock genes[J].Cell Physiol,1992,153:392-401.
[2] Craig E A.The heat shock response[J].CRC Crit Rev Biochem,1985,18(2):39-80.
[3] Frydman J.Folding of newly translated proteins in vivo:the role of molecular chaperones[J].Annu Rev Biochem,2001,70:603-649.
[4] Lindquist S,Craig E A.The heat-shock proteins[J].Annual Review of Genetics,2007,22:631-677.
[5] Cara J B,Aluru N,Moyano F J,et al.Food deprivation induces HSP70and HSP90protein expression in larval gilthead sea bream and rainbow trout[J].Comp Biochem Physiol B Biochem Mol Biol,2005,142(4):426-431.
[6]沈骅,王晓蓉,张景飞.应用应激蛋白HSP70作为生物标志物研究锌、铜及其联合毒性对鲫鱼肝脏的影响[J].环境科学学报,2004,24(5):895-899.Shen H,Wang X R,Zhang J F.The effects of the stress protein HSP70on the liver of Crucian carp were studied as biomarkers[J].Journal of Environmental Science,2004,24(5):895-899.
[7]沈骅,王晓蓉,张景飞,等.低浓度Zn对幼龄鲫鱼肝脏组织应激蛋白HSP70诱导的影响[J].农业环境科学学报,2004,23(3):441-443.Shen H,Wang X R,Zhang J F,et al.The influence of low concentration Zn on HSP70induced by liver tissue of young Crucian carp[J].Journal of Agricultural Environmental Science,2004,23(3):441-443.
[8]沈骅,王晓蓉,张景飞,等.低浓度Pb2+、Cd2+对鲫鱼肝脏组织中HSP70诱导的影响[J].环境污染与防治,2004,26(4):244-247.Shen H,Wang X R,Zhang J F,et al.The influence of low concentration Pb2+and Cd2+on HSP70induced in the liver of Crucian carp[J].Environment Pollution and Control,2004,26(4):244-247.
[9] Sanders B M.Stress-proteins in aquatic organisms:an environmental perspective[J].Crit Rev Toxicol,1993,23:49-75.
[10]张勇,徐钢春,杜富宽,等.美洲鲥hsp70的分子特征及其对运输应激的应答[J].动物学杂志,2016,51(2):268-280.Zhang Y,Xu G C,Du F K,et al.Molecular characteristics of the Alosa sapidissima hsp70and its response to transport stress[J].Chinese Journal of Zoology,2016,51(2):268-280.
[11]祝璟琳,王国良.鱼类HSP70的研究进展[J].宁波大学学报(理工版),2007,20(4):446-450.Zhu J L,Wang G L.Research progress on fish HSP70[J].Journal of Ningbo University(Polytechnic Edition),2007,20(4):446-450.
[12] Xie Y H.Molecular characterization of the HSP70and HSP90genes in Asian clam(Corbicula fluminea)and their expression analysis during heavy metal exposure[J].Gene,2017,16(1):146-150.
[13]刘迪秋,葛锋,陈朝银,等.重金属铜、镉对鲫肝脏基因表达的影响[J].水产科学,2010,17(6):227-230.Liu D Q,Ge F,Chen C Y,et al.Effects of heavy metal copper and cadmium on the gene expression of the Crucian carpliver[J].Fisheries Science,2010,17(6):227-230.
[14]刘福军,张饮江,王明学.铜对鱼类慢性毒性研究进展[J].水生生物学报,2003,27(3):302-307.Liu F J,Zhang Y J,Wang M X.The progress of copper on chronic toxicity of fish[J].Journal of Aquatic Biology,2003,27(3):302-307.
[15]凌明圣.分子伴侣HSP70研究进展[J].国外医学分子生物学分册,1997,15(5):227-230.Ling M S.Molecular partner HSP70research progress[J].Foreign Medical Molecular Biology Branch,1997,15(5):227-230.
[16] Grajg E A.Heat shock protein and molecular chaperones:mediate,ons of proteins stress 70and chaperones 60in diversed pecies[J].Cell,1994,34:253-258.
[17] Deane E E,Woo N Y S.Cloning and characterization of the hsp70multigene family from silver sea bream:modulated gene expression between warm and cold temperature acclimation[J].Biochem Biophys Res Commun,2005,330:776-783.
[18] Demand J,Lfiders J.The carboxy-terminal domain of HSC70provides binding sites for a distinct set of chaperone cofactorsⅢ[J].Molecular and Cellular Biology,1998,18(4):2023-2028.
[19] Vayssier M,Leguerhier F,Fabian J F,et al.Cloning and analysis of a Trichinella briotovi gene encoding a cytoplasmic heat shock protein of 72kD[J].Parasitology,1999,119:81-93.
[20]刘旭东.牙鲆hsp70与hsc70基因的克隆、表达及其SNPs与耐热性状的相关性分析[D].青岛:中国海洋大学,2011.Liu X D.The correlation analysis between Paralichthys olivaceus hsp70and hsc70gene cloning,expression and SNPs and heat resistance traits[D].Qingdao:China Ocean University,2011.
[21]明建华,谢骏,刘波,等.团头鲂hsp70cDNA的克隆、序列分析以及热应激对其mRNA表达的影响[J].中国水产科学,2009,16(5):635-648.Ming J H,Xie J,Liu B,et al.The cloning,sequence analysis and thermal stress of the Megalobrama amblycephala were affected on the expression of mRNA[J].Journal of Fishery Sciences of China,2009,16(5):635-648.
[22]何珊,梁旭方,李观贵,等.鲢鱼、草鱼和尼罗罗非鱼热休克蛋白70基因cDNA全序列的克隆与分析[J].环境科学学报,2009,29(11):2324-2330.He S,Liang X F,Li G G,et al.Cloning and analysis of the gene cDNA sequence in Hypophthalmichthys molitrix,Ctenopharyngodon idella,and Oreochromis niloticus heat shock protein 70gene[J].Journal of Environmental Sciences,2009,29(11):2324-2330.
[23]刘庆全.淞江鲈(Trachidermus fasciatus Heckel)hsp70全序列的克隆及其表达的初步研究[D].上海:复旦大学,2013.Liu Q Q.A preliminary study on the Trachidermus fasciatus Heckel cloning and expression of hsp70 sequences[D].Shanghai:Fudan University,2013.
[24] Zhang A,Zhou X,Wang X,et al.Characterization of two heat shock proteins(Hsp70/Hsc70)from grass carp(Ctenopharyngodon idella):evidence for their differential gene expression,protein synthesis and secretion in LPS challenged peripheral blood lymphocytes[J].Comp Biochem Physiol B Biochem Mol Biol,2011,159(2):109-114.
[25]张娟,张其中,张占会,等.黄颡鱼HSC70基因及其组织表达分析[J].水生生物学报,2009,33(3):426-433.Zhang J,Zhang Q Z,Zhang Z H,et al.Analysis of Pseudobagrus fulvidraco HSC70 gene and its tissue expression[J].Journal of Aquatic Biology,2009,33(3):426-433.
[26] Qu L Y,Xiang J H,Sun X Q,et al.Expression analysis of HSP70in various tissues of Chlamys farreri under thermal stress[J].High Technology Letters,2005,15(5):96-100.
[27] Jing J,Liu H,Chen H,et al.Acute effect of copper and cadmium exposure on the expression of heat shock protein 70in the Cyprinidae fish Tanichthys albonubes[J].Chemosphere,2013,1(14):1-10.
[28]周小飞.草鱼热休克蛋白HSC70和HSP70在免疫反应中不同基因表达、蛋白质合成和分泌的特征探讨[C].成都:电子科技大学,2011.Zhou X F.The characteristics of different gene expression,protein synthesis and secretion of Ctenopharyngodon idella HSC70and HSP70in immune response[D].Chengdu:University of Electronic Science and Technology,2011.
[29]王云彪,侯晓丽,徐镜波.温度与铜联合胁迫下鲤鱼HSP70的组织特异性表达[C]//中国环境科学学会.中国环境科学学会学术年会论文集.北京:中国环境出版社,2011:3714-3717.Wang Y B,Hou X L,Xu J B.Tissue specific expression of tissue specific table of Cyprinus carpio HSP70under combined temperature and copper stress[C]//China Society of Environmental Sciences.Paper on the academic annual meeting of the China Society of Environmental Science.Beijing:China Environmental Press,2011:3714-3717.
[30] Mohammad M N A,Mona S Z,Elsayed A K,et al.Use of fish as bio-indicator of the effects of heavy metals pollution[J].Aquac Res Development,2015,6(4):1-13.
[31] Kim S G,Jee J H,Kang J C.Cadmium accumulation and elimination in tissues of juvenile olive flounder,Paralichthys olivaceus after sub-chronic cadmium exposure[J].Environmental Pollution,2004,127:117-123.
[32] Zhang A Y,Guo Y F,Zhang S N,et al.Cytokine effects and cellular signaling pathways of grass carp HSP70in head kidney leukocytes[J].Fish&Shellfish Immunology,2015,46:550-556.
[33] Xing H,Li S,Wang X,et al.Effects of atrazine and chlorpyrifos on the mRNA levels of HSP70and HSC70in the liver,brain,kidney and gill of common carp(Cyprinus carpio)[J].Chemosphere,2013,90:910-916.
[34] Idriss A A,Ahmad A K.Heavy metal concentrations in fishes from Juru River,estimation of the health risk[J].Bull Environ Contam Toxicol,2015,94:204-208.
[35] Drevnick P E,Sandheinrich M B,Oris J T.Increased ovarian follicular apoptosis in fathead minnows(Pimephales promelas)exposed to dietary methylmercury[J].Aquat Toxicol,2006,79:49-54.
[36] Sfakianakis D G,Renieri E,Kentouri M,et al.Effect of heavy metals on fish larvae deformities:a review[J].Enviro Res,2015,137:246-255.
[37] Adeyemo O K,Adedeji O B,Offor C C.Blood lead level as biomarker of environmental lead pollution in feral and cultured African catfish(Clarias gariepinus)[J].Nigerian Vet,2010,31:139-147.
[38] Price B D,Calderwood S K.Heat-induced transcription from RNA polymeraseⅡandⅢand HSF binding activity are coordinately regulated by the products of the heat shock genes[J].Cell Physiol,1992,153:392-401.
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