水稻C2H2型锌指蛋白ZFP182和ZFP36在ABA诱导的抗氧化防护中的功能分析
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摘要
植物特别是作物对非生物逆境(干旱、冷害、高盐环境等)的适应性对于作物的产量和品质有着非常重要的影响,干旱是植物逆境最普遍的形式,在许多地区是农业发展的瓶颈。植物响应干旱胁迫的一种重要反应是积累植物激素脱落酸(abscisic acid,ABA)。水分胁迫下所积累的ABA能增加植物体内ROS的产生和上调抗氧化防护系统的活性,增强植物对水分胁迫的耐性。而在这一信号网络中,一些转录因子如MYB、WRKY、热激因子和多种类型锌指蛋白的含量增加,它们共同参与维持ROS的稳定含量。因而认为这些转录因子家族在植物对逆境的应答过程中起着非常重要的调控作用。但锌指蛋白在ABA诱导的抗氧化防护中的作用机理,尤其与H2O2、MAPK、 NADPH氧化酶等ABA信号途径中的主要组分的关系并不清楚。
本研究从水稻中分离参与ABA、H2O2应答反应的锌指蛋白转录因子着手,通过原生质体瞬时表达和转基因技术,使目标转录因子基因过表达或者沉默,通过对比转基因与野生型植株抗氧化防护酶、NADPH氧化酶以及MAPK的基因表达及酶活,深入探讨其在水稻耐逆性中的作用机理,对提高水稻抗逆性的遗传育种改良有重要的理论指导意义。主要研究结果如下：
利用生物信息学和RT-PCR的方法从水稻幼苗的叶片中分离出两个锌指蛋白基因ZFP182和ZFP36,分别与拟南芥ZAT12和ZAT10有较高的同源性。其编码产物都含有两个典型的C2H2型锌指结构,ZFP182编码产物为170个氨基酸残基,分子量为18.215kDa,而ZFP36编码产物为220个氨基酸,分子量为22.804kDa。通过亚细胞定位实验发现,这两个锌指蛋白基因均定位于细胞核中。荧光定量RT-PCR分析表明：ABA和H2O2处理后,水稻叶片中ZFP182和ZFP36基因转录水平与对照组相比,出现双阶段快速、瞬时的上调。H2O2清除剂(DMTU)的预处理能有效抑制ABA诱导的ZFP182和ZFP36基因的转录。NO也能上调ZFP182和ZFP36基因的表达；这些结果提示,ABA、H2O2和NO均能诱导ZFP182和ZFP36基因的转录,而且ABA诱导水稻锌指蛋白基因表达增强是通过ABA诱导产生的内源H2O2来起作用的。
为了进一步研究ZFP182和ZFP36的功能,本研究分别构建了ZFP182和ZFP36基因的过量表达(sense)、反义抑制(antisense)和RNA干涉(sense/antisense, dsRNA)的植物表达载体,通过农杆菌介导法转化水稻愈伤组织,成功获得ZFP182的RNA干扰植株；ZFP36的过量表达和反义抑制以及RNA干扰的植株。分别以转基因水稻F1代和野生型水稻为材料,外源ABA处理可以提高细胞内SOD和APX的活性,与野生型相比,RNA干扰及反义抑制的转基因植株的SOD和APX的活性显著降低。通过原生质体瞬时表达体系,我们也发现过表达ZFP182和ZFP36基因可以明显提高SOD和APX酶的活性,而干扰这两个锌指蛋白基因,SOD和APX的活性显著下降。经ABA处理后,SOD和APX的活性均略有上升,实验结果与以转基因植株为材料的结果完全吻合,充分说明了ZFP182和ZFP36基因确实参与了ABA诱导的抗氧化防护反应,而且在其中扮演了重要的角色,对植物增强抗氧化防护,提高胁迫耐性具有重要意义。
真核生物中的MAPK级联系统在外源刺激信号转入胞内应答受体／感应器的下游过程中起枢纽作用。植物ABA信号途径中也有MAPK级联系统的参与。本研究以水稻幼苗叶片为材料,采用药理学方法和荧光定量RT-PCR技术研究了MAPK和ZFPl82和ZFP36基因转录在ABA信号途径之间的关系。MAPKK抑制剂(PD98059和U0126)的预处理能有效抑制ABA诱导的ZFP182和ZFP36基因的转录。在水稻原生质体中,ABA处理能够上调ZFP182和ZFP36的表达。利用dsRNA技术,分别将OsMPK1和OsMPK5基因(分别与AtMPK6和AtMPK3同源)沉默,ABA对ZFP182和ZFP36基因转录的诱导被阻断。而利用dsRNA技术或者RNAi突变体分别使ZFP182和ZFP36基因沉默,OsMPK1和OsMPK5基因的表达不受影响；说明在ABA信号途径中,OsMPK1/OsMPK5和ZFP36/ZFP182呈现出一种直线关系,即OsMPK1和OsMPK5作为ZFP182和ZFP36的上游因子发挥作用。
本实验室已有的研究结果显示ABA及H202均可以诱导OsDM3基因表达,提示这一激酶可能参与干旱与氧化胁迫的信号转导。然而,这一激酶在植物对胁迫反应中涉及的下游元件还不清楚。OsDM3与锌指蛋白转录因子的关系更是未见报道。本研究发现,钙通道阻断剂EGTA、LaCl3, CaM拮抗剂TFP、W7和CCaMK抑制剂KN-93的预处理能有效抑制ABA诱导的OsZF182和OsZF36基因的转录。通过原生质体瞬时表达及dsRNA技术,干扰OsDMI3基因的表达,能有效的阻断ABA对锌指蛋白基因ZFP182和ZFP36的诱导,暗示在ABA信号途径中,OsDMI3位于ZFP182和ZFP36的上游,调控这两个锌指蛋白基因的表达。有意思的是,利用原生质体瞬时沉默技术和转基因技术,分别干扰ZFP182和ZFP36基因的表达,可以显著降低ABA诱导的OsDMI3基因的转录和活性,暗示ZFP182和ZFP36基因又可以调控OsDM3基因的表达。上述研究结果表明,在ABA信号通路中,OsDM3和ZFP182/ZFP36之间可能存在一个交叉对话的机制：OsDMI3作为上游因子调控ZFP182和ZFP36基因的表达,而锌指蛋白ZFP182和ZFP36作为转录因子,可能通过直接作用于OsDMI3的启动子区域或者通过作用于其他的一些激酶或者因子间接的对OsDMI3起调控作用。具体机制还有待于进一步深入的研究。
本实验室以往以玉米叶片作为材料的实验阐明,在ABA信号途经中,NADPH、H202和MAPK之间存在一个正反馈调节机制：即ABA诱导ZmrbohA-D基因的转录,增加RBOH的活性,诱导质外体H202产生,活化MAPK,继而通过激活相关转录因子从而再次激活Rboh基因的表达,增加RBOH的活性,导至H202的进一步产生,形成一个RBOH介导的ROS放大过程。但是,对于ZFP182和ZFP36是否参与这个正反馈调节过程并不清楚。本研究用NADPH的抑制剂(DPI)预处理能有效抑制ABA诱导的ZFP182和ZFP36基因的转录,说明在ABA信号系统中,Osrbohs基因位于ZFP182和ZFP36基因的上游。利用水稻原生质体瞬时表达和dsRNA技术干扰ZFP182或ZFP36基因,对ABA信号相关的OsrbohB和Osrbohl基因转录并无影响。DAB染色结果显示,ABA诱导4h之内,野生型和RNAi的转基因植株中,H202的积累并无明显差异。上述几点暗示,ZFP182和ZFP36基因可能并不参与ABA信号途径中H202信号放大的正反馈调节机制,但并不能排除有其它的C2H2型锌指蛋白参与ABA信号途径中H202信号放大的正反馈调节机制。
The adaptability of plants especially crops to abiotic stress such as drought, cold and high salinity has a very significant impact for the production and quality of crops. Drought is the most common form of plant stresses and is critical bottleneck to the development of agriculture in many countries. The accumulation of the plant hormone abscisic acid (ABA) plays important roles in the responses to water stress. ABA induced by water stress can cause an enhancement in the generation of ROS and antioxidant defences to enhance plant tolerance to water stress. In this signal network, a number of transcription factors such as MYB, WRKY, heat shock factor and multiple types of zinc finger protein content increased, they participate in maintaining stable levels of ROS. So that, these transcription factors play important regulation in plant responses to abiotic stresses. However, it is not clear whether these members of C2H2-type ZFPs are involved in the ABA-induced antioxidant defense and if so, what the relationship between the C2H2-type ZFPs, H2O2, MAPK and NADPH oxidase in the ABA signaling is.
In this study, two ABA-and H2O2-responsive C2H2-type ZFP genes, ZFP36and ZFP182were identified in rice. By protoplast transient expression system and transgenic technology, we made the targen gene over-expression or silence. The enzyme activity and gene expression of antioxidant enzymes, NADPH oxidase and MAPK were investigated in transgenic rice and wild type rice to explore the mechanism of ZFP36and ZFP182in rice stress tolerance. It has an important theoretical significance to improve rice resistance. The results as follows:
In this study, two zinc finger protein genes ZFP182and ZFP36was identified from rice (Oryza sativa Japonica) by bioinformatics method and RT-PCR approach which are clustered with Zat12and Zat10respectively. Sequence analysis showed that ZFP182and ZFP36both contained two typical C2H2zinc finger domains. ZFP182encodes a18.215kDa polypeptide with170amino acids and ZFP36encodes a22.804kDa polypeptide with 220amino acids. It was found by subcellular localization, the two zinc finger protein genes are located in the nucleus. Real-time quantitative RT-PCR analysis showed that:Treatments with ABA and H2O2, a biphasic response in the expression of ZFP182and ZFP36was observed. Pretreatment with reactive oxygen species scavengers (DMTU) substantially arrested ABA-induced gene expression. NO can also up-regulate the expression of ZFP182and ZFP36. These results suggest that ABA, H2O2and NO can increase ZFP182and ZFP36gene expression and H2O2is required for the ABA-induced up-regulation expressions of ZFP182and ZFP36in rice.
For further study of the function of ZFP182and ZFP36, overexpression (sense) vector, antisense inhibition (anti sense) expression vector, and RNA interference (sense/antisense, dsRNA) expression vector of ZFP182and ZFP36gene were constructed, and successfully transferred to rice callus, using agrobacterium-mediated transformation approach. We have get the RNA interference plants of ZFP182; overexpression and antisense suppression and RNA interference plants of ZFP36. We used F1generation of transgenic rice and the wild type rice. Respectively as materials, exogenous ABA could increase intracellular SOD and APX activity. Compare with the wild type, SOD and APX activity of the transgenic plants which were RNA interfered or antisense inhibited decreased significantly. By protoplast transient expression system, we found that overexpression of ZFP182and ZFP36gene could increase the SOD and APX activity, while interfering the two zinc finger protein, SOD and APX activity decrease significantly. After ABA treatment, SOD and APX activity were increased slightly. These results suggest that ZFP182and ZFP36not only participate but also play an important role in ABA-induced antioxidant protection. It is significant to enhance plant antioxidant protection and increase stress tolerance.
MAPK cascade in the eukaryotic system play a pivotal role in the downstream processes that exogenous stimuli into intracellular responses receptors/sensors. MAPK cascade also involve ABA signaling pathway in plants. In this study, we use pharmacological approaches and quantitative RT-PCR technique to discuss the relationship of MAPK and ZFP182/ZFP36gene transcription between the ABA signaling pathway in leaves of rice seedlings. MAPKK inhibitor (PD98059and U0126) pretreatment can effectively inhibit the ABA-induced gene transcription of ZFP182and ZFP36. In rice protoplasts, ABA treatment induced the expression of ZFP36and ZFP182, and the ABA-induced up-regulation in the expression of ZFP36and ZFP182was almost fully blocked in the protoplasts transfected with dsRNAs against OsMPK1and OsMPK5, which show the highest similarity with AtMPK6and AtMPK3respectively. By contrast, the reduction in the expression of ZFP36and ZFP182in the protoplasts silencing transiently ZFP36and ZFP182and in the RNAi mutants of ZFP36and ZFP182did not affect the ABA-induced up-regulation in the expression OsMPK1and OsMPK5. These data indicate that there are a linear relationship between OsMPK1/OsMPK5and ZFP36/ZFP182in ABA signaling, and OsMPKl and OsMPK5function upstream of ZFP182and ZFP36in the ABA signaling.
Previous studies showed that ABA and H2O2induced the expression of OsDMI3, which suggested that OsDIM3may be involved in drought and oxidative stress signal transduction. However, the downstream components involved in the responses of this kinase to stress are unclear. The relationship between OsDIM3and ZFPs is not reported. Our studies showed pretreatment with Calcium channel blockers (EGTA and LaC13), CaM inhibitor (W7and TFP) or CaMK inhibitor KN-93substantially arrested ABA-induced OsZF182and OsZF36gene expression. Using rice protoplast transient expression and dsRNA technology to have OsDMI3gene silencing, ABA-induced ZFP182and ZFP36gene expression was significantly decreased, it indicates that OsDMI3gene locate upstream of the two zinc finger proteins gene ZFP182and ZFP36and control the expression of transcription factors in the ABA signaling system. Interestingly, while interference ZFP182and ZFP36genes respectively, ABA-induced OsDMI3gene expression and enzyme activity were significantly decreased, it indicate that ZFP182and ZFP36genes regulate OsDMI3gene expression. These results clearly suggest that there is a cross talk between OsDMJ3and OsZF182IOsZF36in the ABA signaling:OsDMJ3as a upstream factor regulates the expression of ZFP182and ZFP36, while ZFP182and ZFP36as transcription factors maybe act directly on the promoter of OsDM3, or regulate OsDIM3indirectly through acting on other kinases or factors. Specific mechanism needs further study.
Previous studies showed that there is a positive feedback loop involving NADPH oxidase, H2O2, and MAPK in ABA signaling in maize leaves. ABA-induced H2O2production activates MAPKs, which can amplify H2O2signal by regulating the activity of NADPH oxidase in ABA signaling. However, it is unknown whether ZFP36and ZFP182are involved in the positive feedback loop in the ABA signaling. Our studies showed pretreatment with NADPH inhibitor DPI can effectively inhibit the ABA-induced gene transcription of ZFP182and ZFP36. it indicates that Osrbohs locate upstream of the two zinc finger proteins gene ZFP182and ZFP36in the ABA signaling system. The RNAi silencing of ZFP36and ZFP182in rice protoplasts did not affect the ABA-induced up-regulation in the expression of the NADPH oxidase genes, OsrbohB and Osrbohl. The results of DAB staining showed ABA treatment caused the similar increases in the production of H2O2in the leaves of the wild-type plants and the RNAi mutants of ZFP36and ZFP182during the4h of treatment. The above points suggest that ZFP36and ZFP182maybe not mediate the feedback regulation of H2O2production in the ABA signaling. However, we cannot exclude the possible involvement of other C2H2-type ZFPs in the positive feedback loop in ABA signaling.
本研究从水稻中分离参与ABA、H2O2应答反应的锌指蛋白转录因子着手,通过原生质体瞬时表达和转基因技术,使目标转录因子基因过表达或者沉默,通过对比转基因与野生型植株抗氧化防护酶、NADPH氧化酶以及MAPK的基因表达及酶活,深入探讨其在水稻耐逆性中的作用机理,对提高水稻抗逆性的遗传育种改良有重要的理论指导意义。主要研究结果如下：
利用生物信息学和RT-PCR的方法从水稻幼苗的叶片中分离出两个锌指蛋白基因ZFP182和ZFP36,分别与拟南芥ZAT12和ZAT10有较高的同源性。其编码产物都含有两个典型的C2H2型锌指结构,ZFP182编码产物为170个氨基酸残基,分子量为18.215kDa,而ZFP36编码产物为220个氨基酸,分子量为22.804kDa。通过亚细胞定位实验发现,这两个锌指蛋白基因均定位于细胞核中。荧光定量RT-PCR分析表明：ABA和H2O2处理后,水稻叶片中ZFP182和ZFP36基因转录水平与对照组相比,出现双阶段快速、瞬时的上调。H2O2清除剂(DMTU)的预处理能有效抑制ABA诱导的ZFP182和ZFP36基因的转录。NO也能上调ZFP182和ZFP36基因的表达；这些结果提示,ABA、H2O2和NO均能诱导ZFP182和ZFP36基因的转录,而且ABA诱导水稻锌指蛋白基因表达增强是通过ABA诱导产生的内源H2O2来起作用的。
为了进一步研究ZFP182和ZFP36的功能,本研究分别构建了ZFP182和ZFP36基因的过量表达(sense)、反义抑制(antisense)和RNA干涉(sense/antisense, dsRNA)的植物表达载体,通过农杆菌介导法转化水稻愈伤组织,成功获得ZFP182的RNA干扰植株；ZFP36的过量表达和反义抑制以及RNA干扰的植株。分别以转基因水稻F1代和野生型水稻为材料,外源ABA处理可以提高细胞内SOD和APX的活性,与野生型相比,RNA干扰及反义抑制的转基因植株的SOD和APX的活性显著降低。通过原生质体瞬时表达体系,我们也发现过表达ZFP182和ZFP36基因可以明显提高SOD和APX酶的活性,而干扰这两个锌指蛋白基因,SOD和APX的活性显著下降。经ABA处理后,SOD和APX的活性均略有上升,实验结果与以转基因植株为材料的结果完全吻合,充分说明了ZFP182和ZFP36基因确实参与了ABA诱导的抗氧化防护反应,而且在其中扮演了重要的角色,对植物增强抗氧化防护,提高胁迫耐性具有重要意义。
真核生物中的MAPK级联系统在外源刺激信号转入胞内应答受体／感应器的下游过程中起枢纽作用。植物ABA信号途径中也有MAPK级联系统的参与。本研究以水稻幼苗叶片为材料,采用药理学方法和荧光定量RT-PCR技术研究了MAPK和ZFPl82和ZFP36基因转录在ABA信号途径之间的关系。MAPKK抑制剂(PD98059和U0126)的预处理能有效抑制ABA诱导的ZFP182和ZFP36基因的转录。在水稻原生质体中,ABA处理能够上调ZFP182和ZFP36的表达。利用dsRNA技术,分别将OsMPK1和OsMPK5基因(分别与AtMPK6和AtMPK3同源)沉默,ABA对ZFP182和ZFP36基因转录的诱导被阻断。而利用dsRNA技术或者RNAi突变体分别使ZFP182和ZFP36基因沉默,OsMPK1和OsMPK5基因的表达不受影响；说明在ABA信号途径中,OsMPK1/OsMPK5和ZFP36/ZFP182呈现出一种直线关系,即OsMPK1和OsMPK5作为ZFP182和ZFP36的上游因子发挥作用。
本实验室已有的研究结果显示ABA及H202均可以诱导OsDM3基因表达,提示这一激酶可能参与干旱与氧化胁迫的信号转导。然而,这一激酶在植物对胁迫反应中涉及的下游元件还不清楚。OsDM3与锌指蛋白转录因子的关系更是未见报道。本研究发现,钙通道阻断剂EGTA、LaCl3, CaM拮抗剂TFP、W7和CCaMK抑制剂KN-93的预处理能有效抑制ABA诱导的OsZF182和OsZF36基因的转录。通过原生质体瞬时表达及dsRNA技术,干扰OsDMI3基因的表达,能有效的阻断ABA对锌指蛋白基因ZFP182和ZFP36的诱导,暗示在ABA信号途径中,OsDMI3位于ZFP182和ZFP36的上游,调控这两个锌指蛋白基因的表达。有意思的是,利用原生质体瞬时沉默技术和转基因技术,分别干扰ZFP182和ZFP36基因的表达,可以显著降低ABA诱导的OsDMI3基因的转录和活性,暗示ZFP182和ZFP36基因又可以调控OsDM3基因的表达。上述研究结果表明,在ABA信号通路中,OsDM3和ZFP182/ZFP36之间可能存在一个交叉对话的机制：OsDMI3作为上游因子调控ZFP182和ZFP36基因的表达,而锌指蛋白ZFP182和ZFP36作为转录因子,可能通过直接作用于OsDMI3的启动子区域或者通过作用于其他的一些激酶或者因子间接的对OsDMI3起调控作用。具体机制还有待于进一步深入的研究。
本实验室以往以玉米叶片作为材料的实验阐明,在ABA信号途经中,NADPH、H202和MAPK之间存在一个正反馈调节机制：即ABA诱导ZmrbohA-D基因的转录,增加RBOH的活性,诱导质外体H202产生,活化MAPK,继而通过激活相关转录因子从而再次激活Rboh基因的表达,增加RBOH的活性,导至H202的进一步产生,形成一个RBOH介导的ROS放大过程。但是,对于ZFP182和ZFP36是否参与这个正反馈调节过程并不清楚。本研究用NADPH的抑制剂(DPI)预处理能有效抑制ABA诱导的ZFP182和ZFP36基因的转录,说明在ABA信号系统中,Osrbohs基因位于ZFP182和ZFP36基因的上游。利用水稻原生质体瞬时表达和dsRNA技术干扰ZFP182或ZFP36基因,对ABA信号相关的OsrbohB和Osrbohl基因转录并无影响。DAB染色结果显示,ABA诱导4h之内,野生型和RNAi的转基因植株中,H202的积累并无明显差异。上述几点暗示,ZFP182和ZFP36基因可能并不参与ABA信号途径中H202信号放大的正反馈调节机制,但并不能排除有其它的C2H2型锌指蛋白参与ABA信号途径中H202信号放大的正反馈调节机制。
The adaptability of plants especially crops to abiotic stress such as drought, cold and high salinity has a very significant impact for the production and quality of crops. Drought is the most common form of plant stresses and is critical bottleneck to the development of agriculture in many countries. The accumulation of the plant hormone abscisic acid (ABA) plays important roles in the responses to water stress. ABA induced by water stress can cause an enhancement in the generation of ROS and antioxidant defences to enhance plant tolerance to water stress. In this signal network, a number of transcription factors such as MYB, WRKY, heat shock factor and multiple types of zinc finger protein content increased, they participate in maintaining stable levels of ROS. So that, these transcription factors play important regulation in plant responses to abiotic stresses. However, it is not clear whether these members of C2H2-type ZFPs are involved in the ABA-induced antioxidant defense and if so, what the relationship between the C2H2-type ZFPs, H2O2, MAPK and NADPH oxidase in the ABA signaling is.
In this study, two ABA-and H2O2-responsive C2H2-type ZFP genes, ZFP36and ZFP182were identified in rice. By protoplast transient expression system and transgenic technology, we made the targen gene over-expression or silence. The enzyme activity and gene expression of antioxidant enzymes, NADPH oxidase and MAPK were investigated in transgenic rice and wild type rice to explore the mechanism of ZFP36and ZFP182in rice stress tolerance. It has an important theoretical significance to improve rice resistance. The results as follows:
In this study, two zinc finger protein genes ZFP182and ZFP36was identified from rice (Oryza sativa Japonica) by bioinformatics method and RT-PCR approach which are clustered with Zat12and Zat10respectively. Sequence analysis showed that ZFP182and ZFP36both contained two typical C2H2zinc finger domains. ZFP182encodes a18.215kDa polypeptide with170amino acids and ZFP36encodes a22.804kDa polypeptide with 220amino acids. It was found by subcellular localization, the two zinc finger protein genes are located in the nucleus. Real-time quantitative RT-PCR analysis showed that:Treatments with ABA and H2O2, a biphasic response in the expression of ZFP182and ZFP36was observed. Pretreatment with reactive oxygen species scavengers (DMTU) substantially arrested ABA-induced gene expression. NO can also up-regulate the expression of ZFP182and ZFP36. These results suggest that ABA, H2O2and NO can increase ZFP182and ZFP36gene expression and H2O2is required for the ABA-induced up-regulation expressions of ZFP182and ZFP36in rice.
For further study of the function of ZFP182and ZFP36, overexpression (sense) vector, antisense inhibition (anti sense) expression vector, and RNA interference (sense/antisense, dsRNA) expression vector of ZFP182and ZFP36gene were constructed, and successfully transferred to rice callus, using agrobacterium-mediated transformation approach. We have get the RNA interference plants of ZFP182; overexpression and antisense suppression and RNA interference plants of ZFP36. We used F1generation of transgenic rice and the wild type rice. Respectively as materials, exogenous ABA could increase intracellular SOD and APX activity. Compare with the wild type, SOD and APX activity of the transgenic plants which were RNA interfered or antisense inhibited decreased significantly. By protoplast transient expression system, we found that overexpression of ZFP182and ZFP36gene could increase the SOD and APX activity, while interfering the two zinc finger protein, SOD and APX activity decrease significantly. After ABA treatment, SOD and APX activity were increased slightly. These results suggest that ZFP182and ZFP36not only participate but also play an important role in ABA-induced antioxidant protection. It is significant to enhance plant antioxidant protection and increase stress tolerance.
MAPK cascade in the eukaryotic system play a pivotal role in the downstream processes that exogenous stimuli into intracellular responses receptors/sensors. MAPK cascade also involve ABA signaling pathway in plants. In this study, we use pharmacological approaches and quantitative RT-PCR technique to discuss the relationship of MAPK and ZFP182/ZFP36gene transcription between the ABA signaling pathway in leaves of rice seedlings. MAPKK inhibitor (PD98059and U0126) pretreatment can effectively inhibit the ABA-induced gene transcription of ZFP182and ZFP36. In rice protoplasts, ABA treatment induced the expression of ZFP36and ZFP182, and the ABA-induced up-regulation in the expression of ZFP36and ZFP182was almost fully blocked in the protoplasts transfected with dsRNAs against OsMPK1and OsMPK5, which show the highest similarity with AtMPK6and AtMPK3respectively. By contrast, the reduction in the expression of ZFP36and ZFP182in the protoplasts silencing transiently ZFP36and ZFP182and in the RNAi mutants of ZFP36and ZFP182did not affect the ABA-induced up-regulation in the expression OsMPK1and OsMPK5. These data indicate that there are a linear relationship between OsMPK1/OsMPK5and ZFP36/ZFP182in ABA signaling, and OsMPKl and OsMPK5function upstream of ZFP182and ZFP36in the ABA signaling.
Previous studies showed that ABA and H2O2induced the expression of OsDMI3, which suggested that OsDIM3may be involved in drought and oxidative stress signal transduction. However, the downstream components involved in the responses of this kinase to stress are unclear. The relationship between OsDIM3and ZFPs is not reported. Our studies showed pretreatment with Calcium channel blockers (EGTA and LaC13), CaM inhibitor (W7and TFP) or CaMK inhibitor KN-93substantially arrested ABA-induced OsZF182and OsZF36gene expression. Using rice protoplast transient expression and dsRNA technology to have OsDMI3gene silencing, ABA-induced ZFP182and ZFP36gene expression was significantly decreased, it indicates that OsDMI3gene locate upstream of the two zinc finger proteins gene ZFP182and ZFP36and control the expression of transcription factors in the ABA signaling system. Interestingly, while interference ZFP182and ZFP36genes respectively, ABA-induced OsDMI3gene expression and enzyme activity were significantly decreased, it indicate that ZFP182and ZFP36genes regulate OsDMI3gene expression. These results clearly suggest that there is a cross talk between OsDMJ3and OsZF182IOsZF36in the ABA signaling:OsDMJ3as a upstream factor regulates the expression of ZFP182and ZFP36, while ZFP182and ZFP36as transcription factors maybe act directly on the promoter of OsDM3, or regulate OsDIM3indirectly through acting on other kinases or factors. Specific mechanism needs further study.
Previous studies showed that there is a positive feedback loop involving NADPH oxidase, H2O2, and MAPK in ABA signaling in maize leaves. ABA-induced H2O2production activates MAPKs, which can amplify H2O2signal by regulating the activity of NADPH oxidase in ABA signaling. However, it is unknown whether ZFP36and ZFP182are involved in the positive feedback loop in the ABA signaling. Our studies showed pretreatment with NADPH inhibitor DPI can effectively inhibit the ABA-induced gene transcription of ZFP182and ZFP36. it indicates that Osrbohs locate upstream of the two zinc finger proteins gene ZFP182and ZFP36in the ABA signaling system. The RNAi silencing of ZFP36and ZFP182in rice protoplasts did not affect the ABA-induced up-regulation in the expression of the NADPH oxidase genes, OsrbohB and Osrbohl. The results of DAB staining showed ABA treatment caused the similar increases in the production of H2O2in the leaves of the wild-type plants and the RNAi mutants of ZFP36and ZFP182during the4h of treatment. The above points suggest that ZFP36and ZFP182maybe not mediate the feedback regulation of H2O2production in the ABA signaling. However, we cannot exclude the possible involvement of other C2H2-type ZFPs in the positive feedback loop in ABA signaling.
引文
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