麦冬OjERF基因的克隆与功能研究
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摘要
麦冬(ophiopogpn japonicus)是我国重要的百合科多年生常绿草本植物,其植株类似草、开花,喜好温暖和湿润气候。在中国分布广泛,经常被用作可观赏的园林植物。它具有抗旱、耐盐、耐荫、抗寒、绿期长等优良特性。同时它具有很好的药用价值,长期作为滋阴类中药广泛应用于临床中。因此利用现代分子生物学技术,挖掘其优质基因并进行功能鉴定,对品种改良将有重要的意义。另一方面AP2/ERF类转录因子已经在许多物种中被克隆,它们在信号的传递、下游应答基因的表达调控和细胞发育过程中发挥重要的作用。它们可以与GCC-box、DRE/CRT等顺式作用元件结合,参与生物和非生物胁迫响应。通过转基因的方法单独导入AP2/ERF类转录因子的研究,已经成为物种改良的研究热点。
由于全球水资源紧缺和土壤盐渍化严重,干旱和盐胁迫已经成为限制植株生长和农作物产量的主要限制因素。植株的耐盐和抗旱性涉及到多个复杂代谢途径,研究人员对耐盐、抗旱基因工程的研究还处在初级阶段,其分子作用机制也尚不清楚。因此,克隆与抗旱、耐盐相关的转录因子基因,揭示其参与的信号转导途径网络,并通过转基因技术研究该基因在逆境胁迫中发挥的功能,为植物抗逆新品种的选育提供理论依据。
本研究以麦冬为实验材料,从中克隆了ERF类转录因子OjERF基因,并对其生物学特性进行分析,通过转化烟草,对其在植株抗旱和耐盐性方面的影响以及调控机理进行了深入的研究,具体的研究结果如下：
1.采用RT-PCR方法从麦冬中分离了ERF类转录因子OjERF基因,该基因开放阅读框长1047bp,编码348个氨基酸。氨基酸序列分析显示：它含有一个AP2/ERF典型结构域,推测它具有核定位信号和转录激活区,N-端保守的氨基酸MCGGAII元件,将它归属于ERF类转录因子第四类,是百合科转录因子的新基因。
2. RT-PCR分析结果表明OjERF基因在根茎叶中都表达,不具有组织特异表达性,表达水平为叶>茎>根。OjERF基因在麦冬中的表达受到干旱、高盐、低温、ABA和ET不同程度的诱导,变化规律各不相同,说明麦冬OjERF基因可能在逆境胁迫中有一定的作用,同时也可能通过ABA信号转导途径参与逆境胁迫响应。
3.通过PEG介导法将构建好的PEZR(K)-LC-OjERF质粒转化已经制备好的水稻原生质体,亚细胞定位结果显示：OjERF蛋白定位于细胞核中。
4. OjERF基因与GAL4DB融合后转化酵母细胞,研究表明：OjERF基因具有转录激活功能,可以激活报告基因表达。
5.将OjERF基因全长cDNA序列,克隆至原核表达载体pEASY-E1Expression,构建了重组质粒pEASY-E1-OjERF,经过诱导,表达产物SDA-PAGE电泳检测表明,目的蛋白在大肠杆菌中成功表达。
6.构建植物表达载体PBI121-OjERF,通过农杆菌侵染叶盘法转化烟草植株,并对TO代植株进行分子鉴定。对转基因烟草进行了耐盐和抗旱性分析,结果显示,在高盐胁迫下,转基因烟草的叶绿素含量是野生型对照的3倍；经过20d的干旱处理,转基因烟草和对照相比,降低了电解质渗透率和丙二醛含量的积累,提高了脯氨酸含量的积累和SOD、CAT酶活性,说明转基因烟草具有更强的抗旱和耐盐能力。另外,在正常生长条件下,转基因烟草NtERD10B、NtERD10C、NtERF5、NtSOD和NtCAT15种抗逆相关基因的表达水平要明显高于非转基因对照,由此推测,麦冬OjER1基因超表达后能够高效激活下游抗逆相关基因的表达。
7.实时荧光定量PCR结果显示：转基因烟草ABA合成相关基因NtSDR的表达量会高于非转基因对照。通过ELASA法测定体内ABA含量同样表明,转基因烟草体内有较高的ABA含量,这可能说明OjERF基因可以与多种顺式作用元件结合,通过激活下游抗逆相关基因和ABA合成基因的表达,使体内ABA含量积累,最终通过ABA依赖的信号转导途径提高植株对干早和高盐胁迫的抵抗力。上述结果表明,过量表达OjERF基因提高了烟草的抗旱耐盐能力。
Mondo Grass is an important Liliaceae perennial evergreen herb, like grass, flowering plants, preferring growing in warm and humid climate. It is distributed widely in China, often used as an ornamental garden plant. It has excellent characteristics of drought, salt, shade, cold and long green period. At the same time, it has good medicinal value.The Yin and traditional Chinese medicines are widely used in clinical for the long-term. Therefore, it will have important implications for improving varieties by cloning quality gene and functional identification with modern molecular biology techniques. On the other hand, the AP2/ERF transcription factors have been cloned in many species, they are play important roles in the transmission of the signal, regulation of downstream responses gene expression and cell development. They can be combined with the GCC-box, DRE/CRT cis-acting elements involved in biotic and abiotic stress response. The research on improving plant resistance through transformation separate AP2/ERF transcription factors has become hotspots.
Because of the serious global shortage of water resources and soil salinization.Drought and Salt stress have become a major limiting factor in plant growth and crop yields. Salt and drought-resistance involve a number of complex metabolic pathways and the molecular mechanism has not been known clearly. The research is still in the initial stage. Cloning drought and salt-related transcription factor and reveals its participation in signal transduction network, and study the function of the gene by transgenic technology in adversity stress, which will provide theoretical basis for breeding of new varieties of plants under environmental stresses.
In this study, we cloned the transcription factor OjERF gene from Mondo Grass and identified its biological characteristics, and through transformation tobacco plants to study the mechanism of regulation of plant drought and salt tolerance. The specific experimental results are as follows:
1. The ERF Class transcription factor OjERF gene was isolated by RT-PCR method from Mondo Grass.The open reading frame of the gene is1047bp encoding348amino acids. The predicted amino acid sequence analysis showed that:it contained a AP2/ERF typical domain, a putative nuclear localization signal and a transcription activation. It is a new gene in Liliaceae and N-teminal has a conserved amino MCGGAII motif, which belongs to the fourth class of the ERF transcription factor.
2. RT-PCR analysis results showed that the OjERF gene which had no tissue-specific expression characteristics was expressed in the roots, stems and leaves. The expression level was leaf> stem> root. OjERF gene expression were also affected by drought, high salt, low temperature, ABA and ET, but their variation degrees are different which illustrated the OjERF gene may have acertain role in abiotic stresses and also probably through the ABA signal transduction pathways involved in the Stress Response.
3. By PEG-mediated constructed PEZR (K)-LC-OjERF plasmid transformation into rice protoplast cells. Subcellular localization results showed:OjERF protein located in the nucleus.
4. OjERF gene was fused into GAL4DB and transformed into yeast cells. Studies have shown that: OjERF gene had transcriptional activation ability and also can activate the reporter gene expression.
5. The full-length cDNA sequence of OjERF gene, was cloned into the prokaryotic expression vector pEASY-E1Expression to construct the recombinant plasmid pEASY-E1-OjERF. After induced, expression product was detected by SDA-PAGE electrophoresis and showed that the target protein was successfully expressed in E. coli cell.
6. The OjERF gene was cloned into PBI121and then transferred into tobacco by Agrobacterium infection. TO generation plants were identified by molecular method. Salt-tolerance and drought-tolerance experiments showed that under high salt stress, the chlorophyll content of transgenic tobacco is3-fold greater than that in the wild-type control; After20d drought treatment, the transgenic tobacco had lower electrolyte leakage and malondialdehyde content and higher proline content and SOD, CAT activity compared to the control, which suggest that transgenic tobacco plants have stronger drought and salt tolerance. Under normal growth conditions, resistance related gene of NtERD10B, NtERD10C, NtERF5, NtSOD and NtCAT1expression level were significantly higher than that of non-transgenic control. Overexpression of the OjERF gene can efficiently activate downstream resistance related gene expression.
7. Quantitative real-time PCR results showed that:The expression level of ABA synthesis genes NtSDR was higher in the transgenic tobacco than that in the non-transgenic control. Measured by ELASA found that ABA content is in the transgenic tobacco plants were higher than wide-type,which may indicate the OjERF gene can be combined with multiple cis-acting elements and activiates the expression of stress responsive and ABA biosynthesis-related genes consequently causing ABA accumulation, and ultimately by the ABA-dependent signal transduction pathways to improve plant resistance to drought and high salt stress. These results indicated that overexpression OjERF can enhance tobacco tolerance under high salinity and drought.
由于全球水资源紧缺和土壤盐渍化严重,干旱和盐胁迫已经成为限制植株生长和农作物产量的主要限制因素。植株的耐盐和抗旱性涉及到多个复杂代谢途径,研究人员对耐盐、抗旱基因工程的研究还处在初级阶段,其分子作用机制也尚不清楚。因此,克隆与抗旱、耐盐相关的转录因子基因,揭示其参与的信号转导途径网络,并通过转基因技术研究该基因在逆境胁迫中发挥的功能,为植物抗逆新品种的选育提供理论依据。
本研究以麦冬为实验材料,从中克隆了ERF类转录因子OjERF基因,并对其生物学特性进行分析,通过转化烟草,对其在植株抗旱和耐盐性方面的影响以及调控机理进行了深入的研究,具体的研究结果如下：
1.采用RT-PCR方法从麦冬中分离了ERF类转录因子OjERF基因,该基因开放阅读框长1047bp,编码348个氨基酸。氨基酸序列分析显示：它含有一个AP2/ERF典型结构域,推测它具有核定位信号和转录激活区,N-端保守的氨基酸MCGGAII元件,将它归属于ERF类转录因子第四类,是百合科转录因子的新基因。
2. RT-PCR分析结果表明OjERF基因在根茎叶中都表达,不具有组织特异表达性,表达水平为叶>茎>根。OjERF基因在麦冬中的表达受到干旱、高盐、低温、ABA和ET不同程度的诱导,变化规律各不相同,说明麦冬OjERF基因可能在逆境胁迫中有一定的作用,同时也可能通过ABA信号转导途径参与逆境胁迫响应。
3.通过PEG介导法将构建好的PEZR(K)-LC-OjERF质粒转化已经制备好的水稻原生质体,亚细胞定位结果显示：OjERF蛋白定位于细胞核中。
4. OjERF基因与GAL4DB融合后转化酵母细胞,研究表明：OjERF基因具有转录激活功能,可以激活报告基因表达。
5.将OjERF基因全长cDNA序列,克隆至原核表达载体pEASY-E1Expression,构建了重组质粒pEASY-E1-OjERF,经过诱导,表达产物SDA-PAGE电泳检测表明,目的蛋白在大肠杆菌中成功表达。
6.构建植物表达载体PBI121-OjERF,通过农杆菌侵染叶盘法转化烟草植株,并对TO代植株进行分子鉴定。对转基因烟草进行了耐盐和抗旱性分析,结果显示,在高盐胁迫下,转基因烟草的叶绿素含量是野生型对照的3倍；经过20d的干旱处理,转基因烟草和对照相比,降低了电解质渗透率和丙二醛含量的积累,提高了脯氨酸含量的积累和SOD、CAT酶活性,说明转基因烟草具有更强的抗旱和耐盐能力。另外,在正常生长条件下,转基因烟草NtERD10B、NtERD10C、NtERF5、NtSOD和NtCAT15种抗逆相关基因的表达水平要明显高于非转基因对照,由此推测,麦冬OjER1基因超表达后能够高效激活下游抗逆相关基因的表达。
7.实时荧光定量PCR结果显示：转基因烟草ABA合成相关基因NtSDR的表达量会高于非转基因对照。通过ELASA法测定体内ABA含量同样表明,转基因烟草体内有较高的ABA含量,这可能说明OjERF基因可以与多种顺式作用元件结合,通过激活下游抗逆相关基因和ABA合成基因的表达,使体内ABA含量积累,最终通过ABA依赖的信号转导途径提高植株对干早和高盐胁迫的抵抗力。上述结果表明,过量表达OjERF基因提高了烟草的抗旱耐盐能力。
Mondo Grass is an important Liliaceae perennial evergreen herb, like grass, flowering plants, preferring growing in warm and humid climate. It is distributed widely in China, often used as an ornamental garden plant. It has excellent characteristics of drought, salt, shade, cold and long green period. At the same time, it has good medicinal value.The Yin and traditional Chinese medicines are widely used in clinical for the long-term. Therefore, it will have important implications for improving varieties by cloning quality gene and functional identification with modern molecular biology techniques. On the other hand, the AP2/ERF transcription factors have been cloned in many species, they are play important roles in the transmission of the signal, regulation of downstream responses gene expression and cell development. They can be combined with the GCC-box, DRE/CRT cis-acting elements involved in biotic and abiotic stress response. The research on improving plant resistance through transformation separate AP2/ERF transcription factors has become hotspots.
Because of the serious global shortage of water resources and soil salinization.Drought and Salt stress have become a major limiting factor in plant growth and crop yields. Salt and drought-resistance involve a number of complex metabolic pathways and the molecular mechanism has not been known clearly. The research is still in the initial stage. Cloning drought and salt-related transcription factor and reveals its participation in signal transduction network, and study the function of the gene by transgenic technology in adversity stress, which will provide theoretical basis for breeding of new varieties of plants under environmental stresses.
In this study, we cloned the transcription factor OjERF gene from Mondo Grass and identified its biological characteristics, and through transformation tobacco plants to study the mechanism of regulation of plant drought and salt tolerance. The specific experimental results are as follows:
1. The ERF Class transcription factor OjERF gene was isolated by RT-PCR method from Mondo Grass.The open reading frame of the gene is1047bp encoding348amino acids. The predicted amino acid sequence analysis showed that:it contained a AP2/ERF typical domain, a putative nuclear localization signal and a transcription activation. It is a new gene in Liliaceae and N-teminal has a conserved amino MCGGAII motif, which belongs to the fourth class of the ERF transcription factor.
2. RT-PCR analysis results showed that the OjERF gene which had no tissue-specific expression characteristics was expressed in the roots, stems and leaves. The expression level was leaf> stem> root. OjERF gene expression were also affected by drought, high salt, low temperature, ABA and ET, but their variation degrees are different which illustrated the OjERF gene may have acertain role in abiotic stresses and also probably through the ABA signal transduction pathways involved in the Stress Response.
3. By PEG-mediated constructed PEZR (K)-LC-OjERF plasmid transformation into rice protoplast cells. Subcellular localization results showed:OjERF protein located in the nucleus.
4. OjERF gene was fused into GAL4DB and transformed into yeast cells. Studies have shown that: OjERF gene had transcriptional activation ability and also can activate the reporter gene expression.
5. The full-length cDNA sequence of OjERF gene, was cloned into the prokaryotic expression vector pEASY-E1Expression to construct the recombinant plasmid pEASY-E1-OjERF. After induced, expression product was detected by SDA-PAGE electrophoresis and showed that the target protein was successfully expressed in E. coli cell.
6. The OjERF gene was cloned into PBI121and then transferred into tobacco by Agrobacterium infection. TO generation plants were identified by molecular method. Salt-tolerance and drought-tolerance experiments showed that under high salt stress, the chlorophyll content of transgenic tobacco is3-fold greater than that in the wild-type control; After20d drought treatment, the transgenic tobacco had lower electrolyte leakage and malondialdehyde content and higher proline content and SOD, CAT activity compared to the control, which suggest that transgenic tobacco plants have stronger drought and salt tolerance. Under normal growth conditions, resistance related gene of NtERD10B, NtERD10C, NtERF5, NtSOD and NtCAT1expression level were significantly higher than that of non-transgenic control. Overexpression of the OjERF gene can efficiently activate downstream resistance related gene expression.
7. Quantitative real-time PCR results showed that:The expression level of ABA synthesis genes NtSDR was higher in the transgenic tobacco than that in the non-transgenic control. Measured by ELASA found that ABA content is in the transgenic tobacco plants were higher than wide-type,which may indicate the OjERF gene can be combined with multiple cis-acting elements and activiates the expression of stress responsive and ABA biosynthesis-related genes consequently causing ABA accumulation, and ultimately by the ABA-dependent signal transduction pathways to improve plant resistance to drought and high salt stress. These results indicated that overexpression OjERF can enhance tobacco tolerance under high salinity and drought.
引文
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