文章信息
- 麻浩, 王爽, 周亚丽
- MA Hao, WANG Shuang, ZHOU Yali
- 植物中钙依赖蛋白激酶的研究进展
- Research progress of calcium-dependent protein kinases in plants
- 南京农业大学学报, 2017, 40(4): 565-572
- Journal of Nanjing Agricultural University, 2017, 40(4): 565-572.
- http://dx.doi.org/10.7685/janu.201701036
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文章历史
- 收稿日期: 2017-01-19
钙是植物必需的大量元素之一, 可以稳定细胞膜结构, 维持细胞壁结构, 对植物代谢中产生的酸进行中和消除过量有机酸对细胞的毒害, 提高植物的抗病害能力。同时, 钙离子(Ca2+)是植物细胞信号转导中的第二信使。真核生物中发现的蛋白激酶很多, 与逆境信号传递关系最密切的主要有:分裂原激活蛋白激酶(mitogen-activated protein kinase, MAPK)、钙依赖蛋白激酶(calcium-dependent protein kinase, CDPK)、受体蛋白激酶(receptor protein kinase, RPK)、核糖体蛋白激酶(ribosomal protein kinase)、转录调控蛋白激酶(transcription regulation protein kinase)。其中, 钙依赖蛋白激酶由Hetherington等[1]于1982年在豌豆(Pisum sativum L.)中首先报道, 目前是植物中研究较多的一类丝氨酸/苏氨酸蛋白激酶, 分布于植物和一些原生生物中[2], 在细菌、真菌、酵母、线虫和动物中尚未发现[3-5]。
目前, 植物中许多CDPK的基因已被克隆, 如:在拟南芥(Arabidopsis thaliana)基因组中发现了34个CDPK基因[3, 6]; 水稻(Oryza sativa)基因组中存在31个CDPK基因[7-8]; 玉米(Zea mays L.)基因组中克隆出40个CDPK基因[9]; 小麦(Triticum aestivum L.)基因组中鉴定出20个CDPK基因[10]; 毛果杨(Populus trichocarpa)中有30个CDPK基因[11]; 番茄(Lycopersicon esculentum Mill.)中发现了29个CDPK基因[12]。
1 钙依赖蛋白激酶在植物体内的分布和结构 1.1 钙依赖蛋白激酶的分布CDPK在植物体内分布广泛, 包括根、茎、叶、花、花粉、果实和种子等[13-18]; 同时在胚细胞、花粉细胞、分生细胞、保卫细胞和木质部细胞中也发现有CDPK存在[17, 19]。植物CDPK以可溶性和膜结合两种形式存在, 分布几乎涉及到细胞核、细胞骨架和所有的细胞器(如:线粒体、叶绿体、液泡、内质网、过氧化物酶体和油体等)[13, 15, 20-33]。
1.2 钙依赖蛋白激酶的结构植物中CDPK以单肽链形式存在, 从蛋白质的N端到C端存在4个结构域:可变区、催化区、连接区和调控区[2, 4-5, 34], 典型的CDPK结构如图 1[35]所示。在分子进化角度上, 早期植物种属古老的CDPK基因可能来自于编码Ca2+/CaM蛋白激酶催化区和自抑制区的基因和编码CaM基因的融合, 这可能是CDPK结构特征的根源所在。
可变区:植物CDPK的N末端由20~200个氨基酸残基组成的序列称为可变区。不同种属的CDPK蛋白该区域在氨基酸水平上残基数变化很大, 同源性很低[36-37]。
催化区:催化区具有典型的丝氨酸/苏氨酸蛋白激酶的催化保守序列, 由300多个氨基酸残基组成。不同种属的CDPK蛋白该区域同源性可达80%以上, 此区域的活性部位同源性达到100%[34, 37-38]。
连接区:连接区是CDPK结构域中最为保守的区域, 由20~30个氨基酸组成[34, 37]。当无Ca2+或Ca2+低于某一浓度时, 该区域与催化区结合, 抑制激酶活性; 当Ca2+高于某一浓度时, 该区域解除对蛋白激酶结构域的抑制, 使其恢复激酶活性[4], 因此又称为自抑制区[39]。
调控区:调控区是Ca2+结合的结构域。该区域保守性很低, 不同的CDPK调控区有很大的差异[37, 40], 大多数CDPK含有4个与Ca2+结合的保守的EF手型结构(EF hand structure), 通过该手型结构使CDPK在不依赖于CaM的条件下与Ca2+高度亲和[2, 39, 41]。但有些CDPK却只有3个EF手型结构, 如:拟南芥的AtCPK7、AtCPK8、AtCPK10和AtCPK14等[35], AtCPK13仅有2个EF手型结构, 而AtCPK25只有1个EF手型结构[4, 35]。
2 钙依赖蛋白激酶的功能Ca2植物细胞中的第二信使, 通过其下游CDPK的感受和转导进而引起细胞内的生物化学反应, 从而调控植物对多种非生物逆境信号的响应(图 2)。
2.1 钙依赖蛋白激酶参与调节激素反应CDPK可作为调节激素反应的钙效应器, 脱落酸(ABA)、生长素(IAA)、赤霉素(GA)、油菜素内酯(BR)、细胞分裂素(CK)和茉莉酸(JA)等均可诱导CDPK的表达量发生改变[42-43]。
CDPK/Ca2+介导的ABA信号通道中, AtCPK 10参与干旱条件下拟南芥气孔的调节[44]; AtCPK 4、AtCPK11和AtCPK12是拟南芥的种子萌发过程中的调控因子[45-47]; 在水稻中, OsCPK 4的表达受到ABA的诱导[31]; OsDi 19-4作为OsCDPK14的下游基因, 正向调节水稻ABA应答基因的表达[32]; OsCPK 21在ABA信号通道中起正调控作用[48]; 玉米ZmCPK 11参与了ABA诱导的抗氧化防御反应过程, 并且ZmCPK 11位于ZmMPK5的上游, 同时参与组织伤害应答反应[30, 49]。在IAA胁迫下, 绿豆(Vigna radiata)CDPK的转录明显增加[50]; 生长素还可以使胚发生期的苜蓿(Medicago sativa L.)CDPK转录增加[51]。Zhang等[52]发现GA处理可诱导烟草NtCPK 4转录水平提高; GA处理还可以诱导水稻幼苗OsCDPK 13转录水平的提高, 而ABA和BR却抑制该蛋白的活性[43, 53]。
2.2 钙依赖蛋白激酶参与调节植物对环境胁迫的应答逆境胁迫是诱导CDPK表达的因素之一, 研究表明CDPKs参与调节了多种植物对环境胁迫的应答反应, 干旱、高温、冷害、盐害、光照等多种环境因子都能引起CDPK基因的差异表达和其mRNA的特异性积累[53-56]。
在拟南芥中, 干旱和高盐胁迫下AtCDPK 1和AtCDPK2对应的mRNA迅速表达[54], AtCPK 6参与盐害和干旱相关的保卫细胞的甲基茉莉酸信号的正调控[57], AtCPK8与CATALASE3(CAT3) 互作并通过磷酸化CAT3第261位丝氨酸残基来调节CAT3的活性, 从而调节植物体内活性氧的平衡, 提高植物的抗旱性[58-59]; AtCPK 23在干旱和盐害胁迫中起负调控因子作用[60]; AtCPK 27的表达受到NaCl的诱导并有可能正向调节盐胁迫信号转导过程[33]。在水稻中, OsCPK 4的表达受到高盐和干旱的诱导[31]; OsCDPK 7 基因的超量表达参与在干旱、冷害和盐胁迫下的信号转导[61]; OsCPK 9可通过促进气孔关闭而提高植物的抗旱性[62]; OsCPK 12通过调节OsAPx2、OsAPx8或OsrbohI的转录水平来提高植物的耐盐能力[63]; OsCPK 17改变水稻对冷胁迫的适应性, 但是不影响关键冷胁迫诱导基因的表达[64]; 过表达OsCPK 21基因可以提高水稻的耐盐性[48]; OsCDPK 25受热激诱导上调表达[18]。在小麦中, TaCDPK 1和TaCDPK5参与小麦对低磷的响应[65]; 在聚乙二醇、NaCl、4 ℃和H2O2处理下, TaCPK 7的表达量明显提高[66]。有研究发现, 在玉米中, ZmCPK 1在低温胁迫下上调表达, 而ZmCPK 25则下调表达[67]; 用ABA和NaCl处理玉米后ZmCPK 4下调表达, ZmCPK4过表达的拟南芥对ABA的敏感程度和抗旱性均有所提高[29]; 机械损伤3和6 h以后, 在玉米受伤叶片和其邻近叶片中ZmCPK 11积累表达[68]。在烟草(Nicotiana tabacum L.)中, NtCDPK 1在机械损伤处理后2 h转录开始, 11 h时转录水平下降, 28 h后转录停止[43, 55, 69]; Romeis等[70]发现干旱可以诱导NtCDPK 2和NtCDPK3的表达; NtCPK4在NaCl处理30 min后达到最高水平, 在处理2 h后表达量恢复到初始水平[52]; 干旱和盐害可以影响NtCDPK 12的表达[71]。在番茄中, LeCDPK 1的表达水平在受伤的叶片中4 h时达到最高, 在临近叶片8~12 h达到最高, 而在远端的叶片需18 h才可达到最大值, 该基因表达量的提高与番茄中的可溶性CDPK的含量和活性的增加有关[72-73]; 在高温胁迫下, 番茄LeCPK 2能使番茄免受高温胁迫的伤害[74]。在大麦(Hordeum vulgare L.)中, 营养生长阶段的干旱胁迫条件下, HvCPK 7、HvCPK8和HvCPK2相对表达显著增加[75]。
这些研究说明, CDPK在植物适应逆境过程中起到至关重要的作用, 然而其参与植物产生防御反应和防御信号转导的机制还不清楚。
2.3 钙依赖蛋白激酶参与植株生长和发育的调控植物生长发育的过程绝大多数都要受到Ca2+的调节[76]。CDPK的组织特异性表达说明CDPK参与了植物的早期发育阶段, 如胚发生、种子发育和萌发[69]。
CDPK在植物体中并不决定器官的形成, 但可能具有调控植物器官正常发育的作用。有研究表明胡萝卜(Daucus carota L. var. sativa Hoffm.)中的Ca2+能够提高胚形成的频率, 而缺失钙离子会阻碍体细胞胚的形成。非生物压力例如氧化和低渗压力能够导致细胞质Ca2+的浓度增加, 并且可以引起细胞周期进展的延迟[77]。
CDPK在种子形成和萌发过程中也具有重要的调节作用[69]。SPK是在未成熟的水稻种子胚乳中特异表达的一种CDPK, 对水稻储藏物的生物合成非常重要, 将SPK沉默后, 水稻积累存储产物(如淀粉)的能力下降; 另外, 由于不能利用蔗糖, 种子变得极为松软。这一结果表明, SPK是一种蔗糖合酶激酶, 为存储产物的生物合成提供底物, 参与了存储产物的生物合成途径[78]。在蓖麻(Ricinus communis L.)种子早期发育过程中, RcCDPK 2的表达量先上升后下降[79]。
CDPK也参与块茎的发育。在马铃薯(Solanum tuberosum L.)中, StCDPK 1和StCDPK3在块茎发育的不同阶段存在着时空差异性表达情况:StCDPK3仅在早期伸长的块茎中表达, 而StCDPK1则在块茎顶端的膨胀处表达[80]。
3 钙依赖蛋白激酶的底物越来越多的CDPK底物通过体外磷酸化反应[20]、蛋白互作[22]和保守序列分析[81]被鉴定出来(表 1)[4, 82-83], 如:转录因子、热激蛋白、蛋白磷酸酶和离子通道蛋白等, 这些底物通过与CDPK的互作将Ca2+信号级联放大并向下游传导, 进而对相关基因表达、酶代谢、细胞骨架动态变化以及离子和水分的跨膜运输等进行调节, 使植物在生长发育、应对抵抗非生物胁迫和生物胁迫等方面产生相应的变化[3], 如:Milla等[84]和Curran等[85]先后发现AtCPK4、AtCPK10、AtCPK11和AtCPK16拥有共同底物AtDi19, 同时AtCPK10还可以与HSP1互作[44]; 而AtCPK4、AtCPK11、AtCPK12和AtCPK32都可以与ABF4互作[45-47]; AtCPK8的直接作用底物是拟南芥过氧化氢酶CAT3[58]; AtCPK12以蛋白磷酸酶ABI2作为底物[47]; 阴离子通道蛋白SLAC1是AtCPK21和AtCPK23共同的下游底物[25, 27]。
许多研究已经鉴定出一些与CDPK的作用底物相应的潜在磷酸化基序, 目前已知的基序主要分为:1) 经典磷酸化基序:Φ-5-X-R-3-X-X-[ST0]-。其中:Φ为疏水性氨基酸(下同); X为任何氨基酸(下同); R为碱性氨基酸(下同)。2) ACA2磷酸化基序:[R-9-R-8-X-R-6]-Φ-5-X-X-X-X-S0-X-R+2-。3) ACS磷酸化基序:Φ-3-R-2-Φ-1-S0-Φ+1-x-K+3-R+4[4, 85-86]。
4 大豆钙依赖蛋白激酶研究进展Harmon等[86]于1987年在大豆中第1次分离、纯化和鉴定出CDPK蛋白。随后, Harper等[34]对该蛋白进行蛋白酶水解纯化并进行测序, 根据蛋白测序结果设计简并引物, 从大豆基因组中扩增出长度为151 bp的DNA片段, 进而通过杂交探针从大豆cDNA文库中筛选出第1个编码大豆CDPK蛋白的基因——GmCDPKSK 5。随后, Lee等[87]从大豆中克隆出2个CDPK同系物GmCDPKβ和GmCDPKγ基因。Tang等[88]克隆了3个大豆种子GmCDPKSeeda、GmCDPKSeedb和GmCDPKSeedc基因。迄今, 大豆中共报道了GmCDPKSeeda、GmCDPKSeedb、GmCDPKSeedc、GmCDPKSK 5、GmCDPKβ和GmCDPKγ等6个CDPK基因。Lee等[89]在对不同CDPK同系物的底物特性的研究中发现, 不同的同系物受Ca2+活化所需的Ca2+浓度也不同, 大豆SK5、β、γ磷酸化syntide-2所需Ca2+的K0.5(米氏常数)分别为0.06、0.4、1.0 μmol · L-1[17]。赵弘巍等[90]研究发现大豆叶片质膜上相对分子质量为57×103的钙依赖蛋白激酶具有较强的体外自磷酸化活性, 且人工诱导衰老处理可明显促进其体外自磷酸化水平, 同时外源6-BA预处理可有效抑制其体外自磷酸化水平, 说明该激酶可能参与外源细胞分裂素对大豆叶片衰老的调控过程。Liu等[91]发现在大豆体内GmCDPKSK5和GmCDPKγ可以通过调节GmSerat2;1的磷酸化参与抵御氧化胁迫。Wang等[92]和王爽等[93]研究发现, 在高温高湿胁迫下, 种子田间劣变抗性品种‘湘豆3号’和种子田间劣变不抗品种‘宁镇1号’在种子活力形成时期的种子中钙依赖蛋白激酶CDPK受高温高湿胁迫诱导在品种间呈显著差异积累。进一步研究发现, GmCDPKSK 5在发育种子中呈组织特异性高表达, 而且GmCDPKSK5在高种子活力品种活力形成时期响应高温高湿胁迫呈显著上调表达, 而在低种子活力品种中呈下调表达, 说明其与高温高湿胁迫下春大豆种子活力的形成相关[93]。通过构建高温高湿胁迫下春大豆种质cDNA膜蛋白酵母双杂交文库, 以GmCDPKSK 5 cDNA克隆为诱饵对文库进行筛选和回转验证, 获得了6个与之互作的蛋白:1个LEA蛋白、1个翻译控制肿瘤蛋白、1个种子成熟蛋白、1个微粒体油酸脱氢酶以及2个未知功能蛋白[94]。
5 展望CDPK在参与植物激素应答和响应逆境胁迫机制方面取得了很大进展, 但仍有很多方面需要进行深入的研究。植物体内存在一套非常完善的抵御胁迫的应答机制, 同一刺激能引发不同信号的传递途径, 同时各途径之间存在相互作用, 所以研究CDPK参与不同信号转导以及蛋白激酶之间的相互关系, 对于揭示植物逆境信息传递的机制有重要意义。
今后, 本实验室将在前期研究的基础上, 以GmCDPKSK 5为对象, 在大豆种子活力形成关键时期(R6~R7期)进一步对高温高湿胁迫下与GmCDPKSK5互作的关键蛋白及其互作方式, 高、低种子活力春大豆种质种子钙信号特征及其对GmCDPKSK 5表达影响, GmCDPKSK5与互作蛋白受钙信号调控的方式及GmCDPKSK 5参与种子活力形成的机制等方面进行深入的研究, 以期揭示GmCDPKSK5参与田间高温高湿胁迫下春大豆种子活力形成的机制。
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