玉米抗逆相关基因ZmRAV1和ZmTCX8.1的功能研究
摘要
作物研究中抗逆相关基因的功能研究对了解作物在逆境胁迫下的抗逆机制、提高作物的抗逆性非常重要。本文从affymatrix数据中挑取了两个受干旱、高盐和ABA等逆境胁迫处理后诱导表达上调的转录本/表达序列标签,他们在玉米中对应的全长基因编码两个转录因子蛋白。
其中,RAV1蛋白作为植物中的一类转录因子在植物生长发育的多个方面起着至关重要的作用,目前在拟南芥、辣椒和水稻等多种植物中已有相关的功能研究,但是在玉米中还没有关于该类蛋白的报道。本文从玉米自交系Han21中克隆得到一个新的RAV1蛋白编码基因ZmRAV1。ZmRAV1预测编码了一个转录因子,整个结构中包含两个与DNA直接结合的结构域AP2和B3,这两个结构域在部分RAV类转录因子家族的成员中保守存在。在洋葱表皮细胞中对35S::YFP-ZmRAV1融合蛋白进行瞬时表达分析的结果显示ZmRAV1蛋白定位于细胞核中。在玉米中,ZmRAV1的表达受到脱水,高盐和ABA等胁迫的诱导。与野生型植株相比,ZmRAV1过表达转基因株系提高了对盐胁迫和渗透胁迫的抗性。Illumina测序的结果显示,相对于野生型植株,ZmRAV1过表达转基因株系中一些盐胁迫应答基因,主要是一些和ROS清除相关的基因的表达出现上调。野生型植株和35S::ZmRAV1转基因株系中的抗过氧化物酶酶活测定的结果表明,转基因株系的耐盐性和其体内维持的高水平的POD含量有关。这些结果表明,ZmRAV1作为一个正向转录调节因子参与了植物对盐胁迫和渗透胁迫的抗性应答信号传导途径。
另外,从玉米中分离克隆得到了一个TSO1类似的基因,将其命名为ZmTCX8.1。该基因预测编码一个转录因子,将ZmTCX8.1和其在玉米及水稻中的同源基因进行氨基酸序列分析显示其含有两个保守的CXC结构域。对ZmTCX8.1的融合蛋白进行瞬时表达分析结果显示,ZmTCX8.1蛋白定位于细胞核中。用Real-Time定量PCR的方法对玉米中ZmTCX8.1的表达分析结果显示,其表达受到脱水和ABA胁迫的诱导。对拟南芥过表达转基因株系的表型分析显示,ZmTCX8.1的过表达显著提高了植株对ABA介导的包括种子萌发、幼苗生长和气孔关闭等多种效应的敏感程度。通过检测野生型株系和转基因株系中ABA信号通路相关基因ABIl,ABI3,ABI4和ABI5的表达量显示这些基因的表达在转基因株系中都出现明显上调,说明ZmTCX8.1的过表达在一定程度上恢复了ostl突变体对ABA介导的气孔关闭不敏感的表型。ZmTCX8.1在ABA介导的ROS合成中起着重要的作用,并且可能通过影响植物中ROS的合成从而参与ABA信号传导通路的调控。对内源ABA含量测定的结果表明,相对于野生型植株,转基因株系中内源ABA的含量明显下降,并且ABA1,ABA3,NCED3和AA03等ABA合成相关基因的表达量也明显下调。以上结果表明,ZmTCX8.1作为正向调控因子参与了ABA的信号传导,还对内源ABA的合成起了负向的调控作用。
本文中对这两个抗性相关基因的功能研究,有利于进一步了解玉米在逆境胁迫下的抗逆机制,为下一步通过分子操作技术提高作物的抗逆性奠定了一定的实验基础。
The functional analysis of the stress tolerance related genes is very important for the mechanism of stress tolerance under stress, and for the improvement of the stress tolerance in crops. Two transcripts which were up-regulated under drought, high salnity and ABA treatments were chosen from the affymatrix data, and the full length genes of the two transcript encode two transcription factor proteins.
RAV1proteins as plant physiology and transcription factors play crucial roles in several aspects of development, has been widely studied in plants Arabidopsis, pepper and rice. However, the functions of RAVs in maize have not ever been descripted. A novel RAV1protein gene, ZmRAV1, was amplified from Zea maize inbred line Han21. It encodes a transcription factor with two distinct DNA-binding domains AP2and B3which appear simultaneously uniquely in some members of RAVs family. Transient expression assays of35S::YFP-ZmRAV1fusion construct in onion epidermis cells revealed the ZmRAV1protein to be localized in the nucleus. The expression of ZmRAV1was up-regulated in maize by dehydration, Salt and ABA stresses. Overexpression of ZmRAV1in the transgenic Arabidopsis plants enhanced the salt and osmotic tolerance compared with the wild type. Illumina sequencing shown that a number of salt responsive genes, primary the ROS scavenge related genes were up-regulated in ZmRAV1transgenic line compared with wild type plants. The detection of the activity of antioxidant enzyme in WT and35S::ZmRAV1plant under salt stress shown that, high maintance of POD contributes to the salt tolerance of Arabidopsis transgenic lines. These data suggest that ZmRAV1functions as a transcriptional activator may involved in the salt and osmotic resistance signaling pathways in plants.
We have isolated and cloned a TSO1-like gene from maize, and we named it ZmTCX8.1. It is speculated to encode a transcription factor, and amino acid sequence analysis showed that it contains two conserved CXC domains. Constructing the ZmTCX8.1-GFP confusion vector and transforming it to the epidermis cells showed that the ZmTCX8.1localized to the nucleus. The expression pattern analysis by real-time quantitative PCR indicated that ZmTCX8.1is induced under exogenous ABA and dehydration Stresses. The phenotype analysis of the transgenic plants showed that overexpression of the ZmTCX8.1dramatically enhanced the sensitivity of the Arabidopsis in all the major ABA response, including ABA-induced inhibition of seed germination and post-germination growth arrest, and ABA-induced stomatal closure. We have detected the expression of some ABA signaling pathway genes such as AB11, AB13, AB14and AB15in transgenic and wild-type plants, and found that the expression levels of these genes were significantly improved in transgenic Arabidopsis lines. Overexpression of ZmTCX8.1in ostl mutant effectively rescued the defective ABA-sensitivity phenotype in stomatal closure. ZmTC×8.1was shown to play import role in ROS production in response to ABA, indicating that ZmTC×8.1involved in ABA signal transduction pathway at least partly through affecting ROS production in plant cell. We further measured the endogenous ABA level, and found that endogenous ABA level was dramatically lower in the ZmTCX8.1transgenic lines when compared with wild-type. Hence, we detected the expression of several ABA biosynthesis genes such as ABA1, ABA3, NCED3and AAO3. Consistence to the measurement results of endogenous ABA level, the expression levels of these genes were significantly reduced in transgenic Arabidopsis lines compared with wild-type. Based on the results above, we speculated that ZmTCX8.1might be involved in the regulation of ABA signaling and inhibition of endogenous ABA biosynthesis though feedback regulation.
The function analysis of the two stress tolerance genes in this paper make the benefit to the further acknowledge of the mechanism of stress tolerance in maize, and provide the experimental basis for the improvement of stress tolerance by the molecular technology in crops.
其中,RAV1蛋白作为植物中的一类转录因子在植物生长发育的多个方面起着至关重要的作用,目前在拟南芥、辣椒和水稻等多种植物中已有相关的功能研究,但是在玉米中还没有关于该类蛋白的报道。本文从玉米自交系Han21中克隆得到一个新的RAV1蛋白编码基因ZmRAV1。ZmRAV1预测编码了一个转录因子,整个结构中包含两个与DNA直接结合的结构域AP2和B3,这两个结构域在部分RAV类转录因子家族的成员中保守存在。在洋葱表皮细胞中对35S::YFP-ZmRAV1融合蛋白进行瞬时表达分析的结果显示ZmRAV1蛋白定位于细胞核中。在玉米中,ZmRAV1的表达受到脱水,高盐和ABA等胁迫的诱导。与野生型植株相比,ZmRAV1过表达转基因株系提高了对盐胁迫和渗透胁迫的抗性。Illumina测序的结果显示,相对于野生型植株,ZmRAV1过表达转基因株系中一些盐胁迫应答基因,主要是一些和ROS清除相关的基因的表达出现上调。野生型植株和35S::ZmRAV1转基因株系中的抗过氧化物酶酶活测定的结果表明,转基因株系的耐盐性和其体内维持的高水平的POD含量有关。这些结果表明,ZmRAV1作为一个正向转录调节因子参与了植物对盐胁迫和渗透胁迫的抗性应答信号传导途径。
另外,从玉米中分离克隆得到了一个TSO1类似的基因,将其命名为ZmTCX8.1。该基因预测编码一个转录因子,将ZmTCX8.1和其在玉米及水稻中的同源基因进行氨基酸序列分析显示其含有两个保守的CXC结构域。对ZmTCX8.1的融合蛋白进行瞬时表达分析结果显示,ZmTCX8.1蛋白定位于细胞核中。用Real-Time定量PCR的方法对玉米中ZmTCX8.1的表达分析结果显示,其表达受到脱水和ABA胁迫的诱导。对拟南芥过表达转基因株系的表型分析显示,ZmTCX8.1的过表达显著提高了植株对ABA介导的包括种子萌发、幼苗生长和气孔关闭等多种效应的敏感程度。通过检测野生型株系和转基因株系中ABA信号通路相关基因ABIl,ABI3,ABI4和ABI5的表达量显示这些基因的表达在转基因株系中都出现明显上调,说明ZmTCX8.1的过表达在一定程度上恢复了ostl突变体对ABA介导的气孔关闭不敏感的表型。ZmTCX8.1在ABA介导的ROS合成中起着重要的作用,并且可能通过影响植物中ROS的合成从而参与ABA信号传导通路的调控。对内源ABA含量测定的结果表明,相对于野生型植株,转基因株系中内源ABA的含量明显下降,并且ABA1,ABA3,NCED3和AA03等ABA合成相关基因的表达量也明显下调。以上结果表明,ZmTCX8.1作为正向调控因子参与了ABA的信号传导,还对内源ABA的合成起了负向的调控作用。
本文中对这两个抗性相关基因的功能研究,有利于进一步了解玉米在逆境胁迫下的抗逆机制,为下一步通过分子操作技术提高作物的抗逆性奠定了一定的实验基础。
The functional analysis of the stress tolerance related genes is very important for the mechanism of stress tolerance under stress, and for the improvement of the stress tolerance in crops. Two transcripts which were up-regulated under drought, high salnity and ABA treatments were chosen from the affymatrix data, and the full length genes of the two transcript encode two transcription factor proteins.
RAV1proteins as plant physiology and transcription factors play crucial roles in several aspects of development, has been widely studied in plants Arabidopsis, pepper and rice. However, the functions of RAVs in maize have not ever been descripted. A novel RAV1protein gene, ZmRAV1, was amplified from Zea maize inbred line Han21. It encodes a transcription factor with two distinct DNA-binding domains AP2and B3which appear simultaneously uniquely in some members of RAVs family. Transient expression assays of35S::YFP-ZmRAV1fusion construct in onion epidermis cells revealed the ZmRAV1protein to be localized in the nucleus. The expression of ZmRAV1was up-regulated in maize by dehydration, Salt and ABA stresses. Overexpression of ZmRAV1in the transgenic Arabidopsis plants enhanced the salt and osmotic tolerance compared with the wild type. Illumina sequencing shown that a number of salt responsive genes, primary the ROS scavenge related genes were up-regulated in ZmRAV1transgenic line compared with wild type plants. The detection of the activity of antioxidant enzyme in WT and35S::ZmRAV1plant under salt stress shown that, high maintance of POD contributes to the salt tolerance of Arabidopsis transgenic lines. These data suggest that ZmRAV1functions as a transcriptional activator may involved in the salt and osmotic resistance signaling pathways in plants.
We have isolated and cloned a TSO1-like gene from maize, and we named it ZmTCX8.1. It is speculated to encode a transcription factor, and amino acid sequence analysis showed that it contains two conserved CXC domains. Constructing the ZmTCX8.1-GFP confusion vector and transforming it to the epidermis cells showed that the ZmTCX8.1localized to the nucleus. The expression pattern analysis by real-time quantitative PCR indicated that ZmTCX8.1is induced under exogenous ABA and dehydration Stresses. The phenotype analysis of the transgenic plants showed that overexpression of the ZmTCX8.1dramatically enhanced the sensitivity of the Arabidopsis in all the major ABA response, including ABA-induced inhibition of seed germination and post-germination growth arrest, and ABA-induced stomatal closure. We have detected the expression of some ABA signaling pathway genes such as AB11, AB13, AB14and AB15in transgenic and wild-type plants, and found that the expression levels of these genes were significantly improved in transgenic Arabidopsis lines. Overexpression of ZmTCX8.1in ostl mutant effectively rescued the defective ABA-sensitivity phenotype in stomatal closure. ZmTC×8.1was shown to play import role in ROS production in response to ABA, indicating that ZmTC×8.1involved in ABA signal transduction pathway at least partly through affecting ROS production in plant cell. We further measured the endogenous ABA level, and found that endogenous ABA level was dramatically lower in the ZmTCX8.1transgenic lines when compared with wild-type. Hence, we detected the expression of several ABA biosynthesis genes such as ABA1, ABA3, NCED3and AAO3. Consistence to the measurement results of endogenous ABA level, the expression levels of these genes were significantly reduced in transgenic Arabidopsis lines compared with wild-type. Based on the results above, we speculated that ZmTCX8.1might be involved in the regulation of ABA signaling and inhibition of endogenous ABA biosynthesis though feedback regulation.
The function analysis of the two stress tolerance genes in this paper make the benefit to the further acknowledge of the mechanism of stress tolerance in maize, and provide the experimental basis for the improvement of stress tolerance by the molecular technology in crops.
引文
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