ABA诱导的OsDMI3基因的表达分析与亚细胞定位
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摘要
脱落酸(abscisic acid, ABA)是一种重要的植物激素,不仅在植物的生长发育中而且还在植物对各种环境胁迫反应中起重要作用。已有研究表明ABA的作用模式与植物细胞中氧化胁迫有关,ABA能够诱导NADPH氧化酶引起活性氧产生及抗氧化基因的表达,从而提高植物抗氧化防护能力。Ca2+/CaM依赖性蛋白激酶(CBK)是一种Ser/Thr类蛋白激酶,在动物中,Ca2+/CaM依赖性蛋白激酶(CaMK)已经被很好的研究,迄今人们已在多种植物中分离出大量CBK家族成员,但大多数植物CBK的功能没有被阐明,已有的研究提示这一激酶可能参与盐、低温、干旱与氧化胁迫的信号转导。因此,鉴定ABA诱导的氧化胁迫信号途径与植物CBK基因的关系,对于揭示ABA信号的细胞和分子机理具有重要的意义。
本实验以水稻为研究对象,利用生物信息学和分子生物学相结合的方法,以半定量RT-PCR为主要研究手段对水稻中一条OsCBK基因OsDMI3与ABA和H202的关系进行了研究,并通过构建原生质体瞬时表达载体对其进行亚细胞定位,主要结果如下：
1.用100μmol·L-1 ABA处理4h的时间进程结果显示,OsDMI3基因在水稻叶片和根中的转录水平和对照相比,分别表现为30 min和60 min产生上调表达高峰之后逐渐降低至接近对照水平。用ABA合成抑制剂fluridon(FLU)预处理水稻,研究结果表明：干旱胁迫下,水稻叶片和根中ABA合成受抑制的(FLU处理)与不受抑制的相比,OsDMI3基因的表达均受到了明显的抑制,外源施加ABA恢复后,基本上清除了这种抑制作用。这些结果初步说明ABA可以上调OsDMI3的表达。
2.10mmol·L-1 H2O2处理后,水稻叶片和根中OsDMI3基因的转录水平和对照相比,在4h处理时间均产生两个上调表达高峰；为了确定H202是否参与了ABA诱导的OsDMI3基因在水稻中的转录,本研究应用H202清除剂DMTU和NADPH氧化酶抑制剂DPI浓度分别为100μmol·L-1和10mmol·L-1预处理后的样品较对照样品,结果显示,当ABA处理60min时清除剂和抑制剂预处理对OsDMI3基因的表达并没有明显的抑制。而当ABA处理至60min至90min时,OsDMI3基因在水稻叶片和根中的转录均受到明显抑制。证明了ABA诱导水稻OsDMI3基因表达增强是通过ABA诱导产生的内源H202来起作用的。这些结果初步阐明了在ABA信号途径中ABA、H2O2与植物CBK三者之间的关系,从而为ABA诱导植物细胞抗氧化防护系统的信号转导途径提供了新的证据。
3.通过设计含有酶切位点的引物和PCR方法,从本实验室已保存的水稻OsDMI3基因的模板上扩增完整的基因编码区,并利用DNA重组技术将OsDMI3基因连到含有35S启动子的植物瞬间表达载体pXZP008上,构建了全长基因瞬时表达载体35S-pXZOsDMI3。将表达载体用PEG方法转化玉米原生质体,以黄色荧光蛋白YFP为报告基因,通过激光共聚焦显微镜观察。结果表明OsDMI3主要定位在玉米的细胞质中。为进一步研究ABA诱导的OsDMI3基因表达与ABA诱导的抗氧化防护基因如：Sod4, Cat1及Apx1等表达的关系做准备,为深入了解和丰富ABA信号转导途径网络关系图奠定基础。
Abscisic acid (ABA), one of the most important phytohormones, play an important roles not only to regulate many important aspects of plant-growth and development, but also to regulate plant adaptive responses to various adverse environmental conditions. Previous studies showed that one model of ABA action are related to plant oxidative stress signaling. ABA could cause the generation of hydrogen peroxide (H2O2) and enhance the capacity of antioxidant defense systems. Ca2+/CaM-dependent protein kinase(CBK) is Ser/Thr protein kinase. Ca2+/CaM- dependent protein kinases(CaMK) have been well researched in animals. Until now, many Ca2+/CaM-dependent protein kinases have been isolated from different plants. But the functions of most of CBKs in plants have not been clarified.Previous studies showed that CBK is likely to take part in the signal transduction of salt, cold, drought and Oxidative stress. Therefore, it is important to identify the relationship between ABA-induced oxidative stress signaling pathways and CBK gene to reveal the cellular and molecular mechanism of ABA signal transduction.
In this study, we did something about that by using rice as model material and bioinformatics methods combine with molecular biology methods. An effort was made to elucidate the relationship between CBK genes(OsDMI3) and ABA, H2O2 in semi-quantitative PCR. Then we use the constructs (35S-pXZOsDMI3) and transient expression system to determine potential subcellular locations of OsDMI3. The results are as follows:
1.100μmol·L-1 ABA treatment in the time-course of 4h showed that the transcript level of OsDMI3 in rice leves and roots reached maximum values in 30 min and 60min, then renduced to the control level. Through the treatment of ABA systhesis inhibitor fluridon(FLU), the results are that the transcript level of OsDMI3 in rice leves and roots was inhibited more significantly in the rice which were treated by ABA systhesis inhibitor fluridon(FLU) compared to the other rice which were not treated. However, the treatment of ABA largely removed this inhibition. These suggested that ABA can make the transcript level of OsDMI3 increasing.
2.10 mmol·L-1 H2O2 treatment resulted in an analogical enhancement in the transcript level of OsDMI3 gene analyzed by RT-PCR. The time-course analysis of OsDMI3 gene expressions showed that the transcript level reached two maximum values in 4h of H2O2 treatment. To make sure that H2O2 took part in ABA signal transduction, pretreatments with H2O2 scavenger and inhibitor DMTU and DPI did not suppressed the increases of gene expression within 60min of ABA treatment. But the increases of gene expression within 90min of ABA treatment was significantly suppressed. These suggested that H2O2 is required for the ABA-induced up-regulation expressions of OsDMI3 gene in rice. A new evidence was provided to ABA-induced signal transduction in plant cell antioxidant protection system.
3. To determine the location of OsDMI3 and through designing the primers which include restriction enzyme cutting site, the open reading frames were amplified. The gene were inserted into the pXZP008, respectively, under control of the CaMV35S promoter that allows the transient expression of protein in C-terminal fusion with YFP. The constructs (35S-pXZOsDMI3) were delivered into maize protoplast by PEG method. Images of OsDMI3-YFP fusion were captured with a confocal laser scanning microscope. OsDMI3-YFP were located to cytoplasm. This research prepares for further study about the relationship between ABA-induced OsDMI3 gene expression and protective antioxidant genes such as:Sod4, Cat1 and Apx1. The reorganized protoplast transient expression vector laid the foundation for understanding and expounding the network of ABA signal transduction pathway.
本实验以水稻为研究对象,利用生物信息学和分子生物学相结合的方法,以半定量RT-PCR为主要研究手段对水稻中一条OsCBK基因OsDMI3与ABA和H202的关系进行了研究,并通过构建原生质体瞬时表达载体对其进行亚细胞定位,主要结果如下：
1.用100μmol·L-1 ABA处理4h的时间进程结果显示,OsDMI3基因在水稻叶片和根中的转录水平和对照相比,分别表现为30 min和60 min产生上调表达高峰之后逐渐降低至接近对照水平。用ABA合成抑制剂fluridon(FLU)预处理水稻,研究结果表明：干旱胁迫下,水稻叶片和根中ABA合成受抑制的(FLU处理)与不受抑制的相比,OsDMI3基因的表达均受到了明显的抑制,外源施加ABA恢复后,基本上清除了这种抑制作用。这些结果初步说明ABA可以上调OsDMI3的表达。
2.10mmol·L-1 H2O2处理后,水稻叶片和根中OsDMI3基因的转录水平和对照相比,在4h处理时间均产生两个上调表达高峰；为了确定H202是否参与了ABA诱导的OsDMI3基因在水稻中的转录,本研究应用H202清除剂DMTU和NADPH氧化酶抑制剂DPI浓度分别为100μmol·L-1和10mmol·L-1预处理后的样品较对照样品,结果显示,当ABA处理60min时清除剂和抑制剂预处理对OsDMI3基因的表达并没有明显的抑制。而当ABA处理至60min至90min时,OsDMI3基因在水稻叶片和根中的转录均受到明显抑制。证明了ABA诱导水稻OsDMI3基因表达增强是通过ABA诱导产生的内源H202来起作用的。这些结果初步阐明了在ABA信号途径中ABA、H2O2与植物CBK三者之间的关系,从而为ABA诱导植物细胞抗氧化防护系统的信号转导途径提供了新的证据。
3.通过设计含有酶切位点的引物和PCR方法,从本实验室已保存的水稻OsDMI3基因的模板上扩增完整的基因编码区,并利用DNA重组技术将OsDMI3基因连到含有35S启动子的植物瞬间表达载体pXZP008上,构建了全长基因瞬时表达载体35S-pXZOsDMI3。将表达载体用PEG方法转化玉米原生质体,以黄色荧光蛋白YFP为报告基因,通过激光共聚焦显微镜观察。结果表明OsDMI3主要定位在玉米的细胞质中。为进一步研究ABA诱导的OsDMI3基因表达与ABA诱导的抗氧化防护基因如：Sod4, Cat1及Apx1等表达的关系做准备,为深入了解和丰富ABA信号转导途径网络关系图奠定基础。
Abscisic acid (ABA), one of the most important phytohormones, play an important roles not only to regulate many important aspects of plant-growth and development, but also to regulate plant adaptive responses to various adverse environmental conditions. Previous studies showed that one model of ABA action are related to plant oxidative stress signaling. ABA could cause the generation of hydrogen peroxide (H2O2) and enhance the capacity of antioxidant defense systems. Ca2+/CaM-dependent protein kinase(CBK) is Ser/Thr protein kinase. Ca2+/CaM- dependent protein kinases(CaMK) have been well researched in animals. Until now, many Ca2+/CaM-dependent protein kinases have been isolated from different plants. But the functions of most of CBKs in plants have not been clarified.Previous studies showed that CBK is likely to take part in the signal transduction of salt, cold, drought and Oxidative stress. Therefore, it is important to identify the relationship between ABA-induced oxidative stress signaling pathways and CBK gene to reveal the cellular and molecular mechanism of ABA signal transduction.
In this study, we did something about that by using rice as model material and bioinformatics methods combine with molecular biology methods. An effort was made to elucidate the relationship between CBK genes(OsDMI3) and ABA, H2O2 in semi-quantitative PCR. Then we use the constructs (35S-pXZOsDMI3) and transient expression system to determine potential subcellular locations of OsDMI3. The results are as follows:
1.100μmol·L-1 ABA treatment in the time-course of 4h showed that the transcript level of OsDMI3 in rice leves and roots reached maximum values in 30 min and 60min, then renduced to the control level. Through the treatment of ABA systhesis inhibitor fluridon(FLU), the results are that the transcript level of OsDMI3 in rice leves and roots was inhibited more significantly in the rice which were treated by ABA systhesis inhibitor fluridon(FLU) compared to the other rice which were not treated. However, the treatment of ABA largely removed this inhibition. These suggested that ABA can make the transcript level of OsDMI3 increasing.
2.10 mmol·L-1 H2O2 treatment resulted in an analogical enhancement in the transcript level of OsDMI3 gene analyzed by RT-PCR. The time-course analysis of OsDMI3 gene expressions showed that the transcript level reached two maximum values in 4h of H2O2 treatment. To make sure that H2O2 took part in ABA signal transduction, pretreatments with H2O2 scavenger and inhibitor DMTU and DPI did not suppressed the increases of gene expression within 60min of ABA treatment. But the increases of gene expression within 90min of ABA treatment was significantly suppressed. These suggested that H2O2 is required for the ABA-induced up-regulation expressions of OsDMI3 gene in rice. A new evidence was provided to ABA-induced signal transduction in plant cell antioxidant protection system.
3. To determine the location of OsDMI3 and through designing the primers which include restriction enzyme cutting site, the open reading frames were amplified. The gene were inserted into the pXZP008, respectively, under control of the CaMV35S promoter that allows the transient expression of protein in C-terminal fusion with YFP. The constructs (35S-pXZOsDMI3) were delivered into maize protoplast by PEG method. Images of OsDMI3-YFP fusion were captured with a confocal laser scanning microscope. OsDMI3-YFP were located to cytoplasm. This research prepares for further study about the relationship between ABA-induced OsDMI3 gene expression and protective antioxidant genes such as:Sod4, Cat1 and Apx1. The reorganized protoplast transient expression vector laid the foundation for understanding and expounding the network of ABA signal transduction pathway.
引文
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