拟南芥Rad23家族基因在胁迫应答中的功能分析
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摘要
紫外辐射敏感蛋白RAD23，是一种核苷酸切除修复因子，参与了DNA损伤核酸修复，细胞应激反应和生长发育等。真核生物体内的RAD23蛋白参与DNA损伤修复功能已有广泛研究，但RAD23蛋白在植物体内对细胞光损伤修复作用尚不清楚，因此，本文通过对Rad23家族基因生物信息学的分析以及大量突变体植株系统的研究，得到了如下研究结果：
(1)分析了Rad23-4基因的时空表达特性及其蛋白定位，并系统研究了Rad23-4基因在紫外胁迫应答过程中的作用。结果显示：紫外胁迫处理后，rad23-4与野生型相比其花粉颗粒活性明显降低，种子败育率增加以及体内花青素含量明显降低。对一系列与NER修复相关基因进行分析表明，突变体Rad23-4T-DNA插入突变体内rad4基因mRNA表达量急剧降低，突变体内rad1、rad7、rad16基因表达和野生型表达中变化不大。其体内rad-4基因表达量和花青素含量降低，表明Rad23-4基因在植物的紫外胁迫应答中起重要作用。
(2)实验成功构建了pCold-RAD23重组载体，pCold-RAD23重组载体经IPTG诱导后获得了表达，利用纯化后的重组蛋白制备了多克隆抗体。通过ELISA和WB方法检测制备的抗体具有较高的效价和特异性。同时，WB实验检测该抗体能识别植物体内的RAD23蛋白，免疫组化实验发现RAD23蛋白在花粉粒中表达较多。为进一步研究RAD23重组蛋白在体外是否具有紫外损伤修复功能，检测了重组蛋白处理后细胞的表型，实验发现构建的重组蛋白pCold-RAD23对紫外线损伤Hela细胞具有保护作用。
(3)通过在ABA、NaCl、蔗糖和甘露醇等胁迫条件处理下，分析了拟南芥4种Rad23突变体植株的表型，实验发现Rad23-1、Rad23-3T-DNA插入突变体对ABA、NaCl、蔗糖和甘露醇都不敏感，而Rad23-2、Rad23-4T-DNA插入突变体对ABA和NaCl敏感，但对蔗糖和甘露醇不敏感。用ABA和NaCl处理后，与野生型相比，Rad23-2、Rad23-4T-DNA插入突变体萌发率明显降低，根伸长也受到抑制。同时，实验分析了Rad23-2、Rad23-4突变体和野生型体内胁迫诱导相关基因表达情况，结果表明：与野生型相比，Rad23-2、Rad23-4T-DNA突变体内胁迫应答基因RA29A/B，DREB1A/2A的表达降低，将35S::Rad23-2，35S::Rad23-4过表达到缺失突变体中，表型均得到回复，证明Rad23-2，Rad23-4能正调控相关胁迫应答基因。
(4)运用GUS染色方法检测了Rad23-2基因的GUS活性，结果发现Rad23-2基因在幼苗以及成年植株的花粉粒中GUS活性较强，而花瓣、萼片和雌蕊等组织中的GUS活性较弱；rad23-2突变体与野生型相比，rad23-2突变体的花粉粒发育不良，萌发率下降，种子败育率增加，将35S::Rad23-2过表达到rad23-2缺失突变体中，表型得到回复。对一系列与花粉和绒毡层发育相关的基因检测分析发现，EMS1和TPD1在突变体rad23-2和野生型表达中变化不大，而MYB33/MYB65和AMS在rad23-2缺失突变体中的表达量明显低于野生型，而将35S::Rad23-2过表达到rad23-2缺失突变体中，能回复MYB33/MYB65和AMS基因的表达，证明Rad23-2基因通过正调控MYB33/MYB65和AMS基因来影响花粉粒发育。
Ultraviolet radiation sensitive protein RAD23, is a nucleotide excision repair factor, mainly involved in repairing the DNA damage, such as repairing the DNA damage by UV light, cellular stress response and growth. The function of RAD23protein involved in DNA damage repair has been studied extensively in Eukaryotic, but it is not clear the detailed function of RAD23proteins in plant.In this study, the bioinformatics and functions of Rad23family genes were analyzed.The detailed informaitons were listed as follows:
(1)The expression level and protein location of Rad23-4was analyzed in different organs of Arabidopsis thaliana. The detailing functions of Rad23-4were analyzed after UV-B–treated. Compared with the wild type, the Rad23-4mutants showed increased numbers of collapstion pollen grain, lower levels of pollen grain germination, and higher seed abortion rates after UV-B–treated. Furthermore, the Rad23-4mutants decreased the anthocyanin production.Several NER genes were analyzed in Rad23-4mutants and wild-type.The expression level of rad1, rad7and rad16were not obviously different between Rad23-4mutants and wild-type, but the rad-4gene and anthocyanin production were obviously decreased. All results supported that the Rad23-4gene plays important roles in UV-B stress response.
(2)The expression vector of pCold-RAD23recombinant protein was induced by IPTG. Then the recombinant protein was used as an antigen to prepare polyclonal antiserum in mice. The results showed that the polyclonal antiserum could specific reaction with the RAD23protein of Arabidopsis by WB and ELISA assays. Furthermore, the immunolocalization results showed that the expression level of RAD23protein was found to be high in the pollen grains. To further investigate the photo-protective mechanisms of RAD23protein from UV light irradiation, The Hela cells were irradiated by UV light and incubated with the fusion protein of pCold-RAD23, the results showed that the recombinant RAD23protein could protect the Hela cells from UV irradiation.
(3) The phenotype of Rad23family gene was anlyzed after treating with ABA, NaCl, mannitol and sucrose. The results showed that the Rad23-1and Rad23-3mutant were not sensitive to ABA, NaCl, mannitol and sucrose,while the Rad23-2and Rad23-4mutant were sensitive to ABA, NaCl, but not sensitive to mannitol and sucrose. Compared with the wild type, the Rad23-2and Rad23-4mutants showed germination decreased and seedling root shorten after treating with ABA and salt. Several stress response genes were analyzed in Rad23-2, Rad23-4mutants and wild-type.Compared with the wild type, the expression level of RA29A/B and DREB1A/2A were obviously decreased. The35S::rad23-2and35S::rad23-4transgene were sufficient to restore the expression of RA29A/B and DREB1A/2A. These results indicated that Rad23-2and Rad23-4gene might positive regulate the expression of stress response genes
(4) The GUS activity of Rad23-2gene was analyzed in different organs of Arabidopsis thaliana by GUS stain assay. From the results, the expression level of Rad23-2gene was found to be high in seedlings and pollen grains while low in petals, sepals, and pistil tissues. In addition, the phenotypes of rad23-2mutant, wild-type,35S::rad23-2were observed, and the rad23-2mutants showed increased numbers of abnormal pollen grain, lower levels of pollen grain germination, and higher seed abortion rates than wild-type. This defects can be rescued by35S::rad23-2, a transgene that restores the rad23-2expression of rad23-2mutant. Several genes underlying microspore and tapetum development were analyzed. These results showed that the expression levels of EMS1, TPD1and DYT1were not significantly different between the rad23-2mutants and wild-type. However,the levels of MYB33/MYB65and AMS gene expression in the Rad23-2mutant were significantly lower than those in the wild-type and the35S::rad23-2transgene was sufficient to restore the expression of MYB33/MYB65and AMS.These results indicated that Rad23-2might regulate pollen grains development via regulating the expression of MYB33/MYB65and AMS.
(1)分析了Rad23-4基因的时空表达特性及其蛋白定位，并系统研究了Rad23-4基因在紫外胁迫应答过程中的作用。结果显示：紫外胁迫处理后，rad23-4与野生型相比其花粉颗粒活性明显降低，种子败育率增加以及体内花青素含量明显降低。对一系列与NER修复相关基因进行分析表明，突变体Rad23-4T-DNA插入突变体内rad4基因mRNA表达量急剧降低，突变体内rad1、rad7、rad16基因表达和野生型表达中变化不大。其体内rad-4基因表达量和花青素含量降低，表明Rad23-4基因在植物的紫外胁迫应答中起重要作用。
(2)实验成功构建了pCold-RAD23重组载体，pCold-RAD23重组载体经IPTG诱导后获得了表达，利用纯化后的重组蛋白制备了多克隆抗体。通过ELISA和WB方法检测制备的抗体具有较高的效价和特异性。同时，WB实验检测该抗体能识别植物体内的RAD23蛋白，免疫组化实验发现RAD23蛋白在花粉粒中表达较多。为进一步研究RAD23重组蛋白在体外是否具有紫外损伤修复功能，检测了重组蛋白处理后细胞的表型，实验发现构建的重组蛋白pCold-RAD23对紫外线损伤Hela细胞具有保护作用。
(3)通过在ABA、NaCl、蔗糖和甘露醇等胁迫条件处理下，分析了拟南芥4种Rad23突变体植株的表型，实验发现Rad23-1、Rad23-3T-DNA插入突变体对ABA、NaCl、蔗糖和甘露醇都不敏感，而Rad23-2、Rad23-4T-DNA插入突变体对ABA和NaCl敏感，但对蔗糖和甘露醇不敏感。用ABA和NaCl处理后，与野生型相比，Rad23-2、Rad23-4T-DNA插入突变体萌发率明显降低，根伸长也受到抑制。同时，实验分析了Rad23-2、Rad23-4突变体和野生型体内胁迫诱导相关基因表达情况，结果表明：与野生型相比，Rad23-2、Rad23-4T-DNA突变体内胁迫应答基因RA29A/B，DREB1A/2A的表达降低，将35S::Rad23-2，35S::Rad23-4过表达到缺失突变体中，表型均得到回复，证明Rad23-2，Rad23-4能正调控相关胁迫应答基因。
(4)运用GUS染色方法检测了Rad23-2基因的GUS活性，结果发现Rad23-2基因在幼苗以及成年植株的花粉粒中GUS活性较强，而花瓣、萼片和雌蕊等组织中的GUS活性较弱；rad23-2突变体与野生型相比，rad23-2突变体的花粉粒发育不良，萌发率下降，种子败育率增加，将35S::Rad23-2过表达到rad23-2缺失突变体中，表型得到回复。对一系列与花粉和绒毡层发育相关的基因检测分析发现，EMS1和TPD1在突变体rad23-2和野生型表达中变化不大，而MYB33/MYB65和AMS在rad23-2缺失突变体中的表达量明显低于野生型，而将35S::Rad23-2过表达到rad23-2缺失突变体中，能回复MYB33/MYB65和AMS基因的表达，证明Rad23-2基因通过正调控MYB33/MYB65和AMS基因来影响花粉粒发育。
Ultraviolet radiation sensitive protein RAD23, is a nucleotide excision repair factor, mainly involved in repairing the DNA damage, such as repairing the DNA damage by UV light, cellular stress response and growth. The function of RAD23protein involved in DNA damage repair has been studied extensively in Eukaryotic, but it is not clear the detailed function of RAD23proteins in plant.In this study, the bioinformatics and functions of Rad23family genes were analyzed.The detailed informaitons were listed as follows:
(1)The expression level and protein location of Rad23-4was analyzed in different organs of Arabidopsis thaliana. The detailing functions of Rad23-4were analyzed after UV-B–treated. Compared with the wild type, the Rad23-4mutants showed increased numbers of collapstion pollen grain, lower levels of pollen grain germination, and higher seed abortion rates after UV-B–treated. Furthermore, the Rad23-4mutants decreased the anthocyanin production.Several NER genes were analyzed in Rad23-4mutants and wild-type.The expression level of rad1, rad7and rad16were not obviously different between Rad23-4mutants and wild-type, but the rad-4gene and anthocyanin production were obviously decreased. All results supported that the Rad23-4gene plays important roles in UV-B stress response.
(2)The expression vector of pCold-RAD23recombinant protein was induced by IPTG. Then the recombinant protein was used as an antigen to prepare polyclonal antiserum in mice. The results showed that the polyclonal antiserum could specific reaction with the RAD23protein of Arabidopsis by WB and ELISA assays. Furthermore, the immunolocalization results showed that the expression level of RAD23protein was found to be high in the pollen grains. To further investigate the photo-protective mechanisms of RAD23protein from UV light irradiation, The Hela cells were irradiated by UV light and incubated with the fusion protein of pCold-RAD23, the results showed that the recombinant RAD23protein could protect the Hela cells from UV irradiation.
(3) The phenotype of Rad23family gene was anlyzed after treating with ABA, NaCl, mannitol and sucrose. The results showed that the Rad23-1and Rad23-3mutant were not sensitive to ABA, NaCl, mannitol and sucrose,while the Rad23-2and Rad23-4mutant were sensitive to ABA, NaCl, but not sensitive to mannitol and sucrose. Compared with the wild type, the Rad23-2and Rad23-4mutants showed germination decreased and seedling root shorten after treating with ABA and salt. Several stress response genes were analyzed in Rad23-2, Rad23-4mutants and wild-type.Compared with the wild type, the expression level of RA29A/B and DREB1A/2A were obviously decreased. The35S::rad23-2and35S::rad23-4transgene were sufficient to restore the expression of RA29A/B and DREB1A/2A. These results indicated that Rad23-2and Rad23-4gene might positive regulate the expression of stress response genes
(4) The GUS activity of Rad23-2gene was analyzed in different organs of Arabidopsis thaliana by GUS stain assay. From the results, the expression level of Rad23-2gene was found to be high in seedlings and pollen grains while low in petals, sepals, and pistil tissues. In addition, the phenotypes of rad23-2mutant, wild-type,35S::rad23-2were observed, and the rad23-2mutants showed increased numbers of abnormal pollen grain, lower levels of pollen grain germination, and higher seed abortion rates than wild-type. This defects can be rescued by35S::rad23-2, a transgene that restores the rad23-2expression of rad23-2mutant. Several genes underlying microspore and tapetum development were analyzed. These results showed that the expression levels of EMS1, TPD1and DYT1were not significantly different between the rad23-2mutants and wild-type. However,the levels of MYB33/MYB65and AMS gene expression in the Rad23-2mutant were significantly lower than those in the wild-type and the35S::rad23-2transgene was sufficient to restore the expression of MYB33/MYB65and AMS.These results indicated that Rad23-2might regulate pollen grains development via regulating the expression of MYB33/MYB65and AMS.
引文
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