菜心SOD基因的克隆及其在低温条件下的表达分析
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摘要
采用同源克隆法从菜心中获得3个SOD基因,并进行生物信息学分析,采用qRT-PCR分析3个基因在不同组织器官和低温胁迫下的表达模式。结果表明:(1)获得Cu/Zn-SOD、Fe-SOD、Mn-SOD基因的ORF,分别命名为BclCZSD、BclFSD、BclMSD,序列长分别为459、639、696bp,分别编码152、212、231个氨基酸。(2)生物信息学分析显示,3种蛋白均为稳定的亲水性蛋白,均不存在跨膜结构和信号肽,BclCZSD的二级结构以无规则卷曲为主,含有2个Cu/Zn-SOD结构域,BclFSD和BclMSD的二级结构以α-螺旋为主,含有一个相同的Mn/Fe-SOD结构域;进化分析显示BclCZSD和BclMSD与油菜最先聚在一个分支,BclFSD与甘蓝、萝卜、油菜、芜菁聚在一个分支。(3)qRT-PCR结果显示,BclCZSD、BclFSD和BclMSD基因在菜心根、茎、叶和叶柄中均有表达,且在根、茎、叶和叶柄中的表达模式不完全相同;低温条件下,3个基因的表达量随着胁迫时间的延长均呈先升高后降低的变化趋势。研究结果为进一步探讨菜心SOD基因在低温胁迫下的响应机制奠定了基础。
In this study,three SOD genes were cloned from Brassica campestris L.ssp.chinensis(L.)var.utilis Tsen et Lee by homologous cloning and bioinformatics were analyzed.The expression patterns of three SODgenes in different tissues and under low temperature stress were analyzed by qRT-PCR.The results showed that:(1)the ORF of Cu/Zn-SOD,Fe-SODand Mn-SOD genes were obtained and named BclCZSD,BclFSDand BclMSD.Their sequence length were 459,639 and 696 bp,encoding 152,212 and 231 amino acids,respectively.(2)Bioinformatics analysis showed that all three proteins were stable hydrophilic proteins without transmembrane structure and signal peptide.The secondary structure of BclCZSD protein was mainly random coil and had two Cu/Zn-SOD domains,BclFSD and BclMSD proteins were mainly alpha helix and had one Mn/Fe-SOD domain.Evolutionary analysis showed that BclCZSD and BclMSD proteins were first clustered in one branch with SOD protein in Brassica napus,and BclFSD protein was clustered in one branch with Brassica oleracea,Raphanus sativus,Brassica napus and Brassica rapa.(3)The results of qRT-PCR showed that BclCZSD,BclFSDand BclMSD genes were expressed with different expression profile in the roots,stems,leaves and petioles.The expression level of the 3 genes increased first and then decreased with the extension of time under low temperature.These results provide a foundation for the low temperature response mechanism study of SODgenes in Brassica campestris L.ssp.chinensis(L.)var. utilis Tsen et Lee.
In this study,three SOD genes were cloned from Brassica campestris L.ssp.chinensis(L.)var.utilis Tsen et Lee by homologous cloning and bioinformatics were analyzed.The expression patterns of three SODgenes in different tissues and under low temperature stress were analyzed by qRT-PCR.The results showed that:(1)the ORF of Cu/Zn-SOD,Fe-SODand Mn-SOD genes were obtained and named BclCZSD,BclFSDand BclMSD.Their sequence length were 459,639 and 696 bp,encoding 152,212 and 231 amino acids,respectively.(2)Bioinformatics analysis showed that all three proteins were stable hydrophilic proteins without transmembrane structure and signal peptide.The secondary structure of BclCZSD protein was mainly random coil and had two Cu/Zn-SOD domains,BclFSD and BclMSD proteins were mainly alpha helix and had one Mn/Fe-SOD domain.Evolutionary analysis showed that BclCZSD and BclMSD proteins were first clustered in one branch with SOD protein in Brassica napus,and BclFSD protein was clustered in one branch with Brassica oleracea,Raphanus sativus,Brassica napus and Brassica rapa.(3)The results of qRT-PCR showed that BclCZSD,BclFSDand BclMSD genes were expressed with different expression profile in the roots,stems,leaves and petioles.The expression level of the 3 genes increased first and then decreased with the extension of time under low temperature.These results provide a foundation for the low temperature response mechanism study of SODgenes in Brassica campestris L.ssp.chinensis(L.)var. utilis Tsen et Lee.
引文
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[2]张淑杰,杨再强,陈艳秋,等.低温、弱光、高湿胁迫对日光温室番茄花期生理生化指标的影响[J].生态学杂志,2014,33(11):2 995-3 001.ZHANG S J,YANG Z Q,CHEN Y Q,et al.Effects of low temperature,weak light and high humidity stresses on the physiological and biochemical indicators of greenhouse tomato during flowering period[J].Chinese Journal of Ecology,2014,33(11):2 995-3 001.
[3]邵怡若,许建新,薛立,等.低温胁迫时间对4种幼苗生理生化及光合特性的影响[J].生态学报,2013,33(14):4 237-4 247.SHAO Y R,XU J X,XUE L,et al.Effects of low temperature stress on physiological-biochemical indexes and photosynthetic characteristics of seedlings of four plant species[J].Acta Ecologica Sinica,2013,33(14):4 237-4 247.
[4]朱恒光,顾万荣,王泳超,等.DCPTA对低温下玉米叶片生理生化特征的影响[J].核农学报,2015,29(3):549-556.ZHU H G,GU W R,WANG Y C,et al.Effects of DCPTAon the physiological and biochemical characteristics in maize seedling leaves under low temperature stress[J].Journal of Nuclear Agricultural Sciences,2015,29(3):549-556.
[5]徐龙,陈火英,蒋明敏,等.茄子MnSOD基因的克隆及表达分析[J].植物生理学报,2016,52(10):1 537-1 545.XU L,CHEN H Y,JIANG M M,et al.Cloning and expression analysis of MnSODfrom Solanum melongena[J].Plant Physiology Journal,2016,52(10):1 537-1 545.
[6]HROUART D,VAN MONTAGU M,INZD.Redox-activated expression of the cytosolic copper/zinc superoxide dismutase gene in Nicotiana[J].Proceedings of the National A-cademy of Sciences of the United States of America,1993,90(7):3 108-3 112.
[7]高俊杰,秦爱国,于贤昌.低温胁迫下嫁接对黄瓜叶片SOD和CAT基因表达与活性变化的影响[J].应用生态学报,2009,20(1):213-217.GAO J J,QIN A G,YU X C.Effects of grafting on cucumber leaf SOD and CAT gene expression and activities under low temperature stress[J].Chinese Journal of Applied Ecology,2009,20(1):213-217.
[8]曾秀存,孙万仓,孙佳,等.白菜型冬油菜铁超氧化物歧化酶(Fe-SOD)基因的克隆及表达分析[J].中国农业科学,2013,46(21):4 603-4 611.ZENG X C,SUN W C,SUN J,et al.Cloning and expression analysis of Fe superoxide dismutase(Fe-SOD)gene from winter turnip rape(Brassica rapa L.)[J].Scientia Agricultura Sinica,2013,46(21):4 603-4 611.
[9]曾秀存,刘自刚,史鹏辉,等.白菜型冬油菜铜锌超氧化物歧化酶(Cu/Zn-SOD)基因的克隆及其在低温条件下的表达[J].作物学报,2014,40(4):636-643.ZENG X C,LIU Z G,SHI P H,et al.Cloning and expression analysis of copper and zinc superoxide dismutase(Cu/ZnSOD)gene fromBrassica campestris L.[J].Acta Agronomica Sinica,2014,40(4):636-643.
[10]李钰,郑文寅,冯春,等.非生物逆境胁迫下普通小麦烟农19幼苗FeSOD基因表达分析[J].作物杂志,2016,(4):75-79.LI Y,ZHENG W Y,FENG C,et al.Analysis of FeSODgene expression in normal wheat yannong 19seedlings under abiotic stress[J].Crops,2016,(4):75-79.
[11]朱磊,韩苹,张鹤,等.芥菜锰超氧化物歧化酶基因BjMnSOD1的克隆和分析[J].植物生理学报,2017,53(4):625-631.ZHU L,HAN P,ZHANG H,et al.Cloning and analysis of superoxide dismutase gene BjMnSOD1from Brassica juncea[J].Plant Physiology Journal,2017,53(4):625-631.
[12]余海霞,罗聪,徐趁,等.杧果Mn-MiSOD基因的克隆与表达模式分析[J].植物生理学报,2015,51(12):2 214-2 222.YU H X,LUO C,XU C,et al.Molecular cloning and expression analysis of Mn-MiSOD gene from Mango[J].Plant Physiology Journal,2015,51(12):2 214-2 222.
[13]刘家林,欧阳林娟,曾嘉丽,等.水稻SOD基因家族的全基因组分析及逆境胁迫下表达研究[J].分子植物育种,2018,16(9):2 753-2 760.LIU J L,OUYANG L J,ZENG J L,et al.Genome-wide analysis of rice SOD gene family and expression research under stress[J].Molecular Plant Breeding,2018,16(9):2 753-2 760.
[14]赵静,梁建生,吴雪玲,等.高盐低温胁迫下水稻叶细胞ROS清除系统的相关基因表达[J].西北植物学报,2015,35(5):872-883.ZHAO J,LIANG J S,WU X L,et al.Expression profiling of rice ROS scavenging system related genes under salt or low temperature stress[J].Acta Botanica Boreali-Occidentalia Sinica,2015,35(5):872-883.
[15]VAN CAMP W,CAPIAU K,VAN MONTAGU M,et al.Enhancement of oxidative stress tolerance in transgenic tobacco plants overproducing Fe-superoxide dismutase in chloroplasts[J].Plant Physiology,1996,112(4):1 703-1 714..
[16]MCKERSIE B D,MURNAGHAN J,JONES K S,et al.I-ron-superoxide dismutase expression in transgenic alfalfa increases winter survival without a detectable increase in photosynthetic oxidative stress tolerance[J].Plant Physiology,2000,122(4):1 427-1 437.
[17]KLIEBENSTEIN D J,MONDE R A,LAST R L.Superoxide dismutase in Arabidopsis:an eclectic enzyme family with disparate regulation and protein localization[J].Plant Physiology,1998,118(2):637-650.
[18]LIN Y L,LAI Z X.Superoxide dismutase multigene family in longan somatic embryos:a comparison of Cu/Zn-SOD,FeSOD,and Mn-SOD gene structure,splicing,phylogeny,and expression[J].Molecular Breeding,2013,32(3):595-615.
[19]MOLINA-RUEDA J J,TSAI C J,KIRBY E G.The Populus superoxide dismutase gene family and its responses to drought stress in transgenic poplar overexpressing apine cytosolic glutamine synthetase(GS1a)[J].PLoS One,2013,8(2):e56 421.
[20]SCANDALIOS J G.Oxygen stress and superoxide dismutases[J].Plant Physiology,1993,101(1):7-12.
[21]李玉坤,王学敏,高洪文,等.东方山羊豆Cu/ZnSOD基因的克隆及表达分析[J].遗传,2012,34(1):95-101.LI Y K,WANG X M,GAO H W,et al.Cloning and expression analysis of Cu/ZnSOD gene from Galega orientalis L.[J].Hereditas,2012,34(1):95-101.
[22]王鹏,段文静,王玉昆,等.小麦光温敏雄性不育系BS366铜锌超氧化物歧化酶基因的克隆及表达分析[J].植物遗传资源学报,2017,18(5):939-951.WANG P,DUAN W J,WANG Y K,BAI J F,et al.Cloning and expression analysis of a cupro-zinc superoxide dismutase gene from wheat photo-thermosensitive genic male sterile line BS366[J].Journal of Plant Genetic Resources,2017,18(5):939-951.
[23]李琳玲,程华,许锋,等.银杏铁型超氧化物歧化酶基因(GbFeSOD)的克隆与表达[J].果树学报,2009,26(6):840-846.LI L L,CHENG H,XU F,et al.Molecular cloning,characterization and expression of iron superoxide dismutase gene fromGinkgo biloba[J].Journal of Fruit Science,2009,26(6):840-846.
[24]韩立敏,化文平,王喆之.菘蓝铁型超氧化物歧化酶基因IiFeSOD的克隆与胁迫表达分析[J].广西植物,2013,33(5):663-668.HAN L M,HUA W P,WANG Z Z.Clone and stress expression analysis of iron superoxide dismutase fromIsatis indigotica[J].Guihaia,2013,33(5):663-668.
[25]程华,李琳玲,许锋,等.银杏锰型超氧化物歧化酶GbMnSOD基因的克隆与表达[J].园艺学报,2009,36(9):1 283-1 290.CHENG H,LI L L,XU F,et al.Molecular cloning,characterization and expressionof a manganese superoxide dismutase gene fromGinkgo biloba[J].Acta Horticulturae Sinica,2009,36(9):1 283-1 290.
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