玉米ZmPP2C6-01基因的克隆及表达分析
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摘要
为了研究玉米蛋白磷酸酶2C (PP2C)基因对非生物胁迫的响应及生理功能,以耐旱玉米自交系豫882为试验材料,对玉米20%PEG6000胁迫处理转录组进行分析,筛选受干旱诱导强烈表达的PP2C基因,然后对其进行克隆、生物信息学分析及表达模式分析。结果表明,在玉米20%PEG6000胁迫处理转录组中筛选到一个干旱胁迫下显著上调表达的PP2C蛋白基因,命名为ZmPP2C6-01。ZmPP2C6-01基因的开放阅读框为1 242 bp,编码413个氨基酸,编码蛋白质的分子质量为43.8 ku,等电点为6.6,是一种亲水性蛋白。ZmPP2C6-01蛋白与高粱的PP2C蛋白亲缘关系最近,而与冬小麦、粗山羊草等的PP2C蛋白亲缘关系相对较远。ZmPP2C6-01蛋白定位于细胞核中。利用实时荧光定量PCR分析发现,ZmPP2C6-01基因在玉米根中的表达量最高,其次是叶,茎中最低;PEG胁迫下,ZmPP2C6-01基因在根中的表达量大部分时间点均高于对照,同时在叶中呈上调表达模式;在ABA、NaCl、高温胁迫下,ZmPP2C6-01基因在根中大部分时间点的表达量均低于对照,整体呈下降趋势,而在叶中的表达量均高于对照,呈上调表达模式。综上表明,ZmPP2C6-01基因响应干旱等逆境胁迫,但表达模式不一致。
In order to study the response of maize serine/threonine protein phosphatase PP2 C gene to abiotic stress and its physiological function, the transcriptome of maize inbred line Yu 882 was analyzed under 20% PEG6000 stress. The PP2 C gene strongly expressed under drought stress was screened and cloned, while the bioinformatics analysis and expression pattern analysis were carried out. A PP2 C protein gene that was up-regulated significantly by drought stress was found in the 20% PEG6000 stress-treated transcriptome of maize, and was named as ZmPP2 C6-01. The open reading frame of ZmPP2 C6-01 gene was 1 242 bp, encoding 413 amino acids. The encoded protein had a molecular mass of 43.8 ku and an isoelectric point of 6.6. It was a hydrophilic protein. The ZmPP2 C6-01 protein was closely related to the PP2 C protein of sorghum, and also had the close relationship with the PP2 C protein of winter wheat and Aegilops tauschii. The ZmPP2 C6-01 protein was localized in the nucleus. Real-time quantitative PCR analysis showed that ZmPP2 C6-01 gene had the highest expression in maize roots, followed by leaves and stems. Under the condition of PEG stress, the expression level of ZmPP2 C6-01 gene in roots was higher than that of the control in most of the time, and it was up-regulated in leaves. However, under the stresses of ABA, NaCl and high temperature, the expression level of ZmPP2 C6-01 gene in roots was lower than that of the control, and the overall expression showed a downward trend, while the expression level in leaves was higher than that of the control, showing an up-regulated expression pattern. In summary, the ZmPP2 C6-01 gene responded to drought and other stresses, but the expression patterns were inconsistent.
In order to study the response of maize serine/threonine protein phosphatase PP2 C gene to abiotic stress and its physiological function, the transcriptome of maize inbred line Yu 882 was analyzed under 20% PEG6000 stress. The PP2 C gene strongly expressed under drought stress was screened and cloned, while the bioinformatics analysis and expression pattern analysis were carried out. A PP2 C protein gene that was up-regulated significantly by drought stress was found in the 20% PEG6000 stress-treated transcriptome of maize, and was named as ZmPP2 C6-01. The open reading frame of ZmPP2 C6-01 gene was 1 242 bp, encoding 413 amino acids. The encoded protein had a molecular mass of 43.8 ku and an isoelectric point of 6.6. It was a hydrophilic protein. The ZmPP2 C6-01 protein was closely related to the PP2 C protein of sorghum, and also had the close relationship with the PP2 C protein of winter wheat and Aegilops tauschii. The ZmPP2 C6-01 protein was localized in the nucleus. Real-time quantitative PCR analysis showed that ZmPP2 C6-01 gene had the highest expression in maize roots, followed by leaves and stems. Under the condition of PEG stress, the expression level of ZmPP2 C6-01 gene in roots was higher than that of the control in most of the time, and it was up-regulated in leaves. However, under the stresses of ABA, NaCl and high temperature, the expression level of ZmPP2 C6-01 gene in roots was lower than that of the control, and the overall expression showed a downward trend, while the expression level in leaves was higher than that of the control, showing an up-regulated expression pattern. In summary, the ZmPP2 C6-01 gene responded to drought and other stresses, but the expression patterns were inconsistent.
引文
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[2] Mizoi J,Shinozaki K,Yamaguchi-Shinozaki K.AP2/ERF family transcription factors in plant abiotic stress responses[J].Biochim Biophys Acta,2012,1819(2):86-96.doi:10.1016/j.bbagrm.2011.08.004.
[3] 陈金焕,夏新莉,尹伟伦.植物2C类蛋白磷酸酶及其在逆境信号转导中的作用[J].北京林业大学学报,2010,32(5):168-171.doi:10.13332/j.1000-1522.2010.05.004.Chen J H,Xia X L,Yin W L.Plant 2C protein phosphatase and its role in stress signal transduction[J].Journal of Beijing Forestry University,2010,32(5):168-171.
[4] Schweighofer A,Hirt H,Meskiene L.Plant PP2C phosphatases:emerging functions in stress signaling[J].Trendsin Plant Science,2004,9(5):236-243.doi:10.1016/j.tplants.2004.03.007.
[5] Meskiene I,Baudouin E,Schweighofer A,Liwosz A,Jonak C,Rodriguez P L,Jelinek H,Hirt H.Stress-induced protein phos-phatase 2C is a negative regulator of a mitogen-activated protein kinase[J].J Biol Chem,2003,278(21):18945-18952.doi:10.1074/jbc.M300878200.
[6] Shi Y.Serine/threonine phosphatases:Mechanism through structure[J].Cell,2009,139(3):468-84.doi:10.1016/j.cell.2009.10.006.
[7] Miyazaki S,Koga R,Bohnert H J.Tissue and environmental response-specific expression of 10 PP2C transcripts in Mesembryanthemum crystallinum [J].Mol Gen Genet,1999,261(2):307-316.doi:10.1007/s004380050971.
[8] Vranov E,Langebartels C,Van Montagu M,Inze D,Van Camp W.Oxidative stress,heat shock and drought differentially affect expression of a tobacco protein phosphatase 2C[J].J Exp Bot,2000,51(351):1763-1764.doi:10.1093/jexbot/51.351.1763.
[9] 倪飞.鸢尾蛋白磷酸酶2C(PP2C)基因的克隆和功能分析及拟南芥PP2C基因家族的鉴定和基因组学分析[D].泰安:山东农业大学,2009.Ni F.Cloning and functional analysis of iris protein phosphatase 2C(PP2C) gene and identification and genomics analysis of Arabidopsis PP2C gene family[D].Taian:Shandong Agricultural University,2009.
[10] Yang Q,Liu K,Niu X,Wang Q,Wan Y,Yang F,Li G,Wang Y,Wang R.Genome-wide identification of PP2C genes and their expression profiling in response to drought and cold stresses in Medicago truncatula[J].Scientific Reports,2018,8(1):12841.doi:10.1038/s41598-018-29627-9.
[11] Zhang F,Wei Q H,Shi J C,Jin X,He Y,Zhang Y,Luo Q C,Wang Y S,Chang J L,Yang G X,He G Y.Brachypodium distachyon BdPP2CA6 interacts with BdPYLs and BdSnRK2 and positively regulates salt tolerance in transgenic Arabidopsis[J].Frontiers Plant Science,2017,28(8):264.doi:10.3389/fpls.2017.00264.
[12] Singh A,Jha S K.Jha S K,Bagri J,Pandey G K.ABA inducible rice protein phosphatase 2C confers ABA insensitivity and abiotic stress tolerance in Arabidopsis[J].PLoS One,2015,10(4):e0125168.doi:10.1371/journal.pone.0125168.
[13] Guo P,Shi W C,Li L L,Bao Y J.GMsPP2C,a protein phosphatase 2C gene of alfalfa,confers enhanced salt tolerance in Arabidopsis[J].International Journal of Agriculture and Biology,2018,20(1):62-70.doi:10.17957/IJAB/15.0362.
[14] 闵东红,薛飞洋,马亚男,陈明,徐兆师,李连城,刁现民,贾冠清,马有志.谷子PP2C基因家族的特性[J].作物学报,2013,39 (12):2135-2144.doi:10.3724/SP.J.1006.2013.02135.Min D H,Xue F Y,Ma Y N,Chen M,Xu Z S,Li L C,Diao X M,Jia G Q,Ma Y Z.Characteristics of millet PP2C gene family[J].Acta Agronomica Sinica,2013,39 (12):2135-2144.
[15] Reyes D,Rodriguez D,Gonzalez-Garcia M P,Lorenzo O,Nicolas G,Garcia-Martinez J L,Nicolas C.Overexpression of a protein phosphatase 2C from beech seeds in Arabidopsis shows phenotypes related to abscisic acid responses and gibberellin biosynthesis[J].Plant Physiol,2006,141(4):1414-1424.doi:10.1104/pp.106.084681.
[16] Saavedra X,Modrego A,Rodriguez D,Gonzalez-Garcia MP,Sanz L,Nicolas G,Lorenzo O.The nuclear interactor PYL8/RCAR3 of Fagus sylvatica FsPP2C1 is a positive regulator of abscisic acid signaling in seeds and stress[J].Plant Physiol,2010,152(1):133-150.dio10.1104/pp.109.146381.
[17] Liu L X,Hu X L,Song J A,Zong X J,Li D P,Li D Q.Over-expression of a Zea mays L.protein phosphatase 2C gene (ZmPP2C) in Arabidopsis thaliana decreases tolerance to salt and drought[J].J Plant Physiol,2009,166(5):531-542.doi:10.1016/j.jplph.2008.07.008.
[18] Santiago J,Rodrigues A,Saez A,Rubio S,Antoni R,Dupeux F,Park S Y,M rquez J A,Cutler S R,Rodriguez P L.Modulation of drought resistance by the abscisic acid-receptor PYL5 through inhibition of clade A PP2Cs[J].Plant Journal,2009,60(4):575-588.doi:10.1111/j.1365-313X.2009.03981x.
[19] Jia H F,Lu D,Sun J H,Li C L,Xing Y,Qin L,Shen Y Y.Type 2C protein phosphatase ABI1 is a negative regulator of strawberry fruit ripening[J].J Exp Bot,2013,64(6):1677-1687.doi:10.1093/jxb/ert028.
[20] Li Y S,Sun H,Wang Z F,Duan M,Huang S D,Yang J,Zhang H S.A novel nuclear proteinphosphatase 2C negatively regulated by ABL1 is involvedin abiotic stress and panicle development in rice[J].Mol Biotechnol,2013,54(2):703-710.doi:10.1007/s12033-012-9614-8.
[21] Xue T T,Wang D,Zhang S Z,Ehlting J,Ni F,Jakab S,Zheng C,Zhong Y.Genome-wide and ex-pression analysis of protein phosphatase 2C in rice and Arabidopsis[J].BMC Genomics,2008,20(9):550.doi:10.1186/1471-2164-9-550.
[22] 刘兆明,刘宗旨,白庆武,方荣祥.Agroinfiltration 在植物分子生物学研究中的应用[J].生物工程学报2002,18(4):411-414.doi:10.3321/j.issn:1000-3061.2002.04.003.Liu Z M,Liu Z Z,Bai Q W,Fang R X.Application of agroinfiltration in plant molecular biology research[J].Chinese Journal of Biotechnology,2002,18(4):411-414.
[23] Livak K J,Schmittgen T D.Analysis of relative gene expression data using real-time quantitative PCR and the 2-ΔΔCt method[J].Method,2001,25(4):402-408.doi:10.1006/meth.2001.1262.
[24] 邓克勤,郭新红,汪启明,刘选明.拟南芥磷酸酶基因亚细胞定位与组织表达[J].西北植物学报,2009,29(2):234-239.doi:1000-4025(2009)02-0234-06.Deng K Q,Guo X H,Wang Q M,Liu X M.Subcellular localization and tissue expression of Arabidopsis phosphatase gene[J].Acta Botanica Boreali-Occidentalia Sinica,2009,29(2):234-239.
[25] 官晓敏.百脉根LjPP2C蛋白磷酸酶负调控MPK6信号转导途径[D].武汉:华中农业大学,2015.Guan X M.The negative regulation of MPK6 signal transduction pathway by LjPP2C protein phosphatase[D].Wuhan:Central China Agricultural University,2015.
[26] 郭鹏,张士刚,邢鑫,姜健.欧美杨PdPP2C基因的克隆与功能分析[J].北京林业大学学报.2015,37(2):100-106.doi:10.13332/j.cnki.jbfu.2015.02.018.Guo P,Zhang S G,Xing X,Jiang J.Cloning and functional analysis of PdPP2C gene from Populu[J].Journal of Beijing Forestry University,2015,37 (2):100-106.
[27] 胡晓丽.玉米根系蛋白磷酸酶ZmPP2C2基因的分离、功能鉴定及转基因烟草对逆境的响应[D].泰安:山东农业大学,2008.Hu X L.Isolation,functional identification of maize root protein phosphatase ZmPP2C2 gene and response of transgenic tobacco to stress[D].Taian:Shandong Agricultural University,2008.
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