蛋白磷酸化修饰在保卫细胞响应非生物胁迫中的作用机制
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摘要
蛋白质的磷酸化是一种重要的蛋白质翻译后修饰方式。蛋白质的磷酸化与去磷酸化在植物气孔运动过程中起着关键的调节作用。目前,保卫细胞磷酸化蛋白质组学的主要研究内容包括鉴定磷酸化蛋白、定位磷酸化位点、定量磷酸化水平,进而揭示磷酸化和去磷酸化在植物气孔运动过程中所起的生物学功能。Open Stomata 1 (OST1)/SnRK2.6是蔗糖非酵解型蛋白激酶SnRK2(sucrose non-fermenting receptor kinase)家族的成员,具有典型的丝氨酸/苏氨酸蛋白激酶保守域,并主要在保卫细胞中表达。当植物处于正常生长环境时,ABA的受体PYR/PYL/RCAR不能作用于蛋白磷酸酶2C (PP2Cs,protein phosphatase 2Cs),PP2Cs通过与OST1互作抑制OST1的活性;在逆境胁迫下,PP2Cs解除对OST1蛋白激酶的抑制,随后OST1蛋白激酶启动对下游信号组分的调控作用并引起气孔运动。通过综述磷酸化修饰的定性和定量分析方法,以及蛋白质磷酸化修饰在保卫细胞应对非生物胁迫中的作用机制,提出了保卫细胞磷酸化蛋白组学领域目前存在的挑战和研究前景,旨在为深入了解保卫细胞应答非生物胁迫的气孔运动机制提供参考和新方向。
Protein phosphorylation modification is recognized as the most important post-translational modification. Protein phosphorylation and dephosphorylation in guard cells under abiotic stress play essential roles in stomatal movement under stress conditions. Recently, the main research contents of phosphorylation proteomics in guard cells include identification of phosphorylated proteins, localization of phosphorylation sites, quantification of phosphorylation levels, and revealing the biological functions of phosphorylation and dephosphorylation in stomata movement. Open Stomata 1(OST1)/SnRK2.6, a major SnRK2-type protein kinase, has a typical conserved domain of Ser/Thr protein kinase, and it mainly expresses in guard cells. ABA receptor(PYR/PYL/RCAR) can't interact with Protein Phosphatase 2Cs(PP2Cs) under normal condition, and inhibit the activity of SnRK2.6(OST1) through interact with it. Under stress conditions, protein phosphatase type 2C relieves the inhibition of protein kinase OST1, which activates the regulation of the downstream components and induces stomatal movement. This review summarized the qualitative and quantitative analysis of phosphorylation modification, and mechanism of protein phosphorylation modification in protecting cells in response to abiotic stress. In addition, we also proposed provided some proposals for further research on guard cell phosphoprotomics. Our study aims are to provide a reference and indicate new directions for the mechanism study of stomata movement in response to abiotic stress.
Protein phosphorylation modification is recognized as the most important post-translational modification. Protein phosphorylation and dephosphorylation in guard cells under abiotic stress play essential roles in stomatal movement under stress conditions. Recently, the main research contents of phosphorylation proteomics in guard cells include identification of phosphorylated proteins, localization of phosphorylation sites, quantification of phosphorylation levels, and revealing the biological functions of phosphorylation and dephosphorylation in stomata movement. Open Stomata 1(OST1)/SnRK2.6, a major SnRK2-type protein kinase, has a typical conserved domain of Ser/Thr protein kinase, and it mainly expresses in guard cells. ABA receptor(PYR/PYL/RCAR) can't interact with Protein Phosphatase 2Cs(PP2Cs) under normal condition, and inhibit the activity of SnRK2.6(OST1) through interact with it. Under stress conditions, protein phosphatase type 2C relieves the inhibition of protein kinase OST1, which activates the regulation of the downstream components and induces stomatal movement. This review summarized the qualitative and quantitative analysis of phosphorylation modification, and mechanism of protein phosphorylation modification in protecting cells in response to abiotic stress. In addition, we also proposed provided some proposals for further research on guard cell phosphoprotomics. Our study aims are to provide a reference and indicate new directions for the mechanism study of stomata movement in response to abiotic stress.
引文
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[9] GRENNAN A K.Protein S-nitrosylation:potential targets and roles in signal transduction[J].Plant Physiology,2007,144(3):1237-1239.
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[19] LAOHAVISIT A,BROWN A T,CICUTA P,et al.Annexins:components of the calcium and reactive oxygen signaling network[J].Plant Physiology,2010,152(4):1824-1829.
[20] SHI K,LI X,ZHANG H,et al.Guard cell hydrogen peroxide and nitric oxide mediate elevated CO2-induced stomatal movement in tomato[J].New Phytologist,2015,208(2):342-353.
[21] LI X,SUN Z,SHAO S,et al.Tomato–Pseudomonas syringae interactions under elevated CO2concentration:the role of stomata[J].Journal of Experimental Botany,2014,66(1):307-316.
[22] KUDLA J,BATISTIC O,HASHIMOTO K.Calcium signals:the lead currency of plant information processing[J].The Plant Cell,2010,22(3):541-563.
[23] DING Y,LI H,ZHANG X,et al.OST1 kinase modulates freezing tolerance by enhancing ICE1 stability in Arabidopsis[J].Developmental Cell,2015,32(3):278-289.
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[25] MUSTILLI A C,MERLOT S,VAVASSEUR A,et al.Arabidopsis OST1 protein kinase mediates the regulation of stomatal aperture by abscisic acid and acts upstream of reactive oxygen species production[J].The Plant Cell,2002,14(12):3089-3099.
[26] NAKASHIMA K,YAMAGUCHI-SHINOZAKI K.ABA signaling in stress-response and seed development[J].Plant Cell Reports,2013,32(7):959-970.
[27] ZHANG T,CHEN S,HARMON A C.Protein phosphorylation in stomatal movement[J].Plant Signaling&Behavior,2014,9(11):e972845.
[28] LEE S C,LIM C W,LAN W,et al.ABA signaling in guard cells entails a dynamic protein-protein interaction relay from the PYL-RCAR family receptors to ion channels[J].Molecular Plant,2013,6(2):528-538.
[29] MUNEMASA S,HAUSER F,PARK J,et al.Mechanisms of abscisic acid-mediated control of stomatal aperture[J].Current Opinion in Plant Biology,2015,28:154-162.
[30] MULLER H M,SCHAFER N,BAUER H,et al.The desert plant Phoenix dactylifera closes stomata via nitrate-regulated SLAC1anion channel[J].New Phytologist,2017,216(1):150-162.
[31] VAHISALU T,KOLLIST H,WANG Y F,et al.SLAC1 is required for plant guard cell S-type anion channel function in stomatal signaling[J].Nature,2008,452(7186):487.
[32] GEIGER D,SCHERZER S,MUMM P,et al.Activity of guard cell anion channel SLAC1 is controlled by drought-stress signaling kinase-phosphatase pair[J].Proceedings of the National Academy of Sciences,2009,106(50):21425-21430.
[33] ACHARYA B R,JEON B W,ZHANG W,et al.Open Stomata 1(OST1)is limiting in abscisic acid responses of Arabidopsis guard cells[J].New Phytologist,2013,200(4):1049-1063.
[34] BRANDT B,BRODSKY D E,XUE S,et al.Reconstitution of abscisic acid activation of SLAC1 anion channel by CPK6 and OST1 kinases and branched ABI1 PP2C phosphatase action[J].Proceedings of the National Academy of Sciences,2012,109(26):10593-10598.
[35] VAHISALU T,PUZORJOVA I,BROSCHE M,et al.Ozone-triggered rapid stomatal response involves the production of reactive oxygen species,and is controlled by SLAC1 and OST1[J].The Plant Journal,2010,62(3):442-453.
[36] GRONDIN A,RODRIGUES O,VERDOUCQ L,et al.Aquaporins contribute to ABA-triggered stomatal closure through OST1-mediated phosphorylation[J].The Plant Cell,2015,27(7):1945.
[37] HUBBARD K E,SIEGEL R S,VALERIO G,et al.Abscisic acid and CO2signalling via calcium sensitivity priming in guard cells,new CDPK mutant phenotypes and a method for improved resolution of stomatal stimulus–response analyses[J].Annals of Botany,2011,109(1):5-17.
[38] LIANG S,LU K,WU Z,et al.A link between magnesium-chelatase H subunit and sucrose nonfermenting 1(SNF1)-related protein kinase SnRK2.6/OST1 in Arabidopsis guard cell signalling in response to abscisic acid[J].Journal of Experimental Botany,2015,66(20):6355-6369.
[39] MEYER S,MUMM P,IMES D,et al.AtALMT12 represents an R-type anion channel required for stomatal movement in Arabidopsis guard cells[J].The Plant Journal,2010,63(6):1054-1062.
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