摘要
促分裂原活化蛋白激酶（MAPK）级联途径是真核生物中普遍存在的高度保守的信号转导模式。MAPK级联途径由三个蛋白激酶组成：MAPKKK、MAPKK和MAPK，这三个信号转导组分通过逐级磷酸化作用将外界信号级联放大并向下传递。大量研究表明，MAPK级联途径在植物调控生物与非生物胁迫，生长发育以及植物激素信号转导过程中起着重要作用。系统进化树分析发现，MAPK家族成员主要分为A，B，C，D四族，有关A，B和C族MAPK功能的研究比较深入，但关于D族MAPK功能研究的相关报道主要集中在水稻中。本研究从玉米品种郑单958叶片中克隆到一个D组MAPK基因，命名为ZmMPK17，对其序列特征，表达模式，过表达转基因烟草的抗逆生理功能，以及其上游的组分做了初步研究。主要结果如下：
（1）从玉米中分离到一个促分裂原活化蛋白激酶基因：ZmMPK17，该基因的开放阅读框长1476bp，编码一个491个氨基酸的蛋白，对其序列分析发现，该蛋白具有MAPK进化上保守的11个蛋白激酶催化结构域。在第七和第八结构域之间的激活环中有一个MAPKK的磷酸化位点TDY基序。进化树分析表明，ZmMPK17属于D族的MAPK。
（2）将ZmMPK17与GFP荧光蛋白融合后，瞬时转化洋葱表皮细胞，发现ZmMPK17主要定位于细胞核，并且ZmMPK17蛋白的C-末端延长区对其核定位是必须的。
（3）通过Northern blot及qRT-PCR分析发现，ZmMPK17在玉米根、茎、叶中均有表达，在根中表达量最高。4℃、ABA、SA、MeJA和ETH处理均能诱导ZmMPK17的表达，而PEG、H2O2、NaCl和高温则抑制ZmMPK17的表达。H2O2和Ca2+介导PEG和4℃对ZmMPK17表达的调控。
（4）对ZmMPK17基因的启动子分析发现，启动子中含有多个响应生物与非生物胁迫的顺式作用元件。对ProZmMPK17::GUS转基因烟草GUS染色分析表明，低温（8℃）、ABA、SA和JA处理不同程度地诱导了GUS基因的表达，而PEG处理则抑制GUS的表达。这些结果与表达分析的结果基本一致。
（5）将ZmMPK17及其失活突变基因ZmMPK17-AF构建pBI121表达载体，成功转化烟草，Northern blot和Western blot结果表明，ZmMPK17在转基因植株中正常表达。
（6）在渗透胁迫下，过表达ZmMPK17的转基因烟草积累了较少的ROS，具有较高的抗氧化酶活性（APX和CAT），MV处理后表现出较强的抗氧化能力，这表明过表达ZmMPK17通过ROS的清除，增强对渗透胁迫的抗性。
（7）在低温胁迫下，过表达ZmMPK17及失活型ZmMPK17-AF的转基因烟草种子萌发率明显高于WT，并且积累了较多的渗透调节物质（包括可溶性糖和脯氨酸）。这表明过表达ZmMPK17增强了转基因烟草株系的低温抗性，并且ZmMPK17激活环中TDY基序的磷酸化对其低温抗性并不是必须的。
（8）在渗透和低温胁迫下，过表达ZmMPK17增强了转基因烟草株系中胁迫相关基因的表达。
（9）用病原菌CMV和PVY侵染后，过表达ZmMPK17的转基因烟草株系表现出较轻发病症状，CMV和PVY的积累受到明显抑制，而且抗病相关基因的表达要高于WT和AF株系。
（10）利用酵母双杂交实验发现，ZmMPK17并没有与CaMs和ZmMKKs发生互作，并且从cDNA文库中也没有筛选到与ZmMPK17互作的蛋白。ZmMPK17与其它蛋白的相互作用，还需要通过其它方法做进一步的验证。
Mitogen-activated protein kinase (MAPK) cascades are universal modules of signaltransduction, which are highly conserved in eukaryotes. The MAPK cascades are composedof MAPK, MAPKK (MAPK kinase) and MAPKKK (MAPKK kinase). The intracellularsignals were relayed and amplified through sequential phosphorylations ofMAPKKK→MAPKK→MAPK. In plants，numerous studies show that MAPKs play animportant role in the regulation of biotic and abiotic stresses, development and growth andphytohormones cues. Phylogenetic trees reveal that plant MAPKs can be divided into at leastfour groups (A–D). The most extensively studied MAPKs are in groups A, B and C, reportsabout the function of group D MAPKs is emerging. Currently, the study of group D MAPKhas been concentrated mainly in monocot modle plant rice. In this study, we isolated andcharacterized a novel group D MAPK gene, ZmMPK17, from maize (Zea mays L.Zhengdan958). Sequence characteristic, expression analysis, functional studies on theover-expressing ZmMPK17transgenic plants, and the upstream kinases of ZmMPK17wereanalyzed in this work. The main results are as follows:
(1) We isolated a MAPK gene, designated ZmMPK17, from Zea mays. The ZmMPK17genehas an open reading frame of1,476bp encoding a protein of491amino acid residues.Alignment of the deduced protein sequence with other MAPKs from plants indicated thatZmMPK17contains all11conserved subdomains that are characteristic of MAP kinases. Aspecific dual phosphorylation activation motif TDY (aa175-177) between subdomains VIIand VIII was found. Phylogenetic tree analysis showed that ZmMPK17belongs to group D.
(2) The fusion protein ZmMPK17-GFP was expressed transiently in onion epidermis. Theresults clearly indicate that ZmMKK4is localized in the nucleus, and the C-terminal domainextension (CD) of ZmMPK17is essential for its nuclear localization.
(3) ZmMPK17was ubiquitously detected in roots, stems and leaves. The transcription level inthe roots was higher than that in the stems and leaves.4℃, ABA, SA, MeJA and ETHtreatments led to an increase of ZmMPK17expression, whereas PEG, H2O2, NaCl and45℃ resulted in a decline in ZmMPK17transcript levels. H2O2and Ca2+mediate PEG and4℃-induced the regulation of ZmMPK17at transcription level.
(4) Many potential cis-elements related to biotic and abiotic stress responsiveness were foundin the promoter region of ZmMPK17. The response of ProZmMPK17::GUS transgenic plantsto different treatments was investigated. Upon8℃, ABA, SA and MeJA application, theexpression of GUS was induced, but PEG treatment caused a decrease in transcription level ofGUS. The GUS staining analyses were consistent with the expression pattern of ZmMPK17.
(5) The full coding regions of ZmMPK17and constitutively inactive version ZmMPK17-AFligated into the binary pBI121expression vector under the control of the CaMV35S promoter,and transgenic tobacco was obtained. Northern blot and Western blot analysis revealed thatZmMPK17and ZmMPK17-AF were expressed normally in engineered lines.
(6) ZmMPK17-overexpressing tobacco plants accumulated less reactive oxygen species (ROS)and presented higher activities of antioxidant enzymes (APX and CAT) under osmotic stress.After methyl-niologen (MV) treatment, ZmMPK17-overexpressing lines showed increasedtolerance to oxidative stress. These observations demonstrate that overexpression ofZmMPK17in transgenic tobacco enhanced osmotic stress tolerance by affecting ROSscavenging system.
(7) Overexpression of ZmMPK17and ZmMPK17-AF improved transgenic lines tolerance tolow temperature stress. Compared to WT plants, transgenic lines showed a significantlyhigher germination rate, and accumulated more proline and soluble sugars. These resultsindicate that overexpression of ZmMPK17increased transgenic tobacco tolerance to coldstress, and the phosphorylation of TDY motif was not essential for full function in plants.
(8) Under osmotic and low temperature stress, ZmMPK17-overexpressing transgenic tobaccoplants increased the expression levels of stress-responsive marker genes.
(9) ZmMPK17-overexpressing transgenic tobacco plants showed slight disease symptomsafter inoculated with cucumber mosaic virus (CMV) and potato virus Y (PVY), and theaccumulation of CMV and PVY was suppressed. The expression of PR genes was elevated inZmMPK17-overexpressing plants in compare with WT and ZmMPK17-AF-overexpressingplants.
(10) Using the Yeast Two-Hybrid System, we found that ZmMPK17does not interact with CaMs and ZmMKKs, we also did not obtain any ZmMPK17-interacting proteins from aproteome-wide Y2H screen of the maize leaf cDNA library. Interacting protein screen forZmMPK17need further research.
引文
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