ThLEA3基因过表达提高黄檗抗渗透胁迫能力的机理研究
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摘要
晚期胚胎富集蛋白(Late embryogenesis abundant protein, LEA)是一类在植物种子成熟期间高富集的与渗透调节有关的多功能的逆境蛋白。研究结果表明LEA基因可以在干旱和ABA处理条件下诱导表达,并且LEA蛋白的表达与ABA变化相一致；很多LEA基因的过表达能提高植物的抗旱能力,但其参与植物抗逆胁迫的分子机制还未深入研究,对其结构和功能的了解尚不完善,且存在很多争议。本研究主要以转柽柳LEA3基因的转基因黄檗(Phellodendron amurense Rupr.)为研究对象,通过对转基因黄檗在渗透胁迫下的生理指标变化、气孔关闭程度、荧光分子探针标记、非损伤微测技术和转录组测序等研究NO、H2O2和ABA在诱导气孔关闭过程中的变化规律,初步研究柽柳LEA3基因在渗透胁迫下快速诱导气孔关闭的调控机理,探索在渗透胁迫条件下的分子保护机制。其主要的实验结果如下：
(1)经7%PEG6000处理10小时后,对照的黄檗叶片有明显的萎蔫现象,叶片严重失水；而转ThMT3转基因黄檗的叶片保持正常,说明ThLEA3基因的过表达提高了转基因黄檗的抗渗透胁迫能力。各种生理指标的比较结果表明,在渗透胁迫下,与对照相比,转基因黄檗的SOD、CAT的活性没有明显的差异,而脯氨酸含量、可溶性蛋白含量、丙二醛(MDA)含量均高于非转基因对照。
(2)扫描电镜观察结果表明,7%PEG6000处理5小时后,转基因株系的气孔关闭,而非转基因对照的气孔是开着的,同时保卫细胞发生变形,说明ThLEA3转基因黄檗组培苗通过气孔的快速关闭来抵抗渗透胁迫。
(3)荧光分子探针检测结果表明,转基因黄檗气孔中的NO荧光强度高于对照,而转基因黄檗气孔中的H202荧光强度低于对照。ABA、H2O2、SNP及PEG6000处理能引起转基因株系的气孔中Ca2+的荧光强度,并引起气孔关闭；而不能引起对照的气孔关闭。非损伤微测技术检测结果表明PEG6000处理引起转基因植株气孔中Ca2+内流,H+、K+外流,最终导致气孔的关闭,而对照气孔中Ca2+、H+、K+全部外流,导致气孔不能关闭和保卫细胞的破坏。
(4)内源ABA含量检测结果表明,在渗透处理之前非转基因对照(NT)叶片中ABA含量高于ThLEA3转基因植株。而7%PEG6000处理之后,两者ABA的含量提高幅度都很大,但是胁迫后的还是对照叶片中ABA含量高于转基因叶片。
(5)为进一步在基因水平上验证相关基因的表达,我们利用Illumina高通量测序技术测定了转基因及对照组培苗叶片的转录组。测序分析结果表明,ThLEA3转基因引起大量基因表达量的变化,即有30,494个差异表达的基因,其中有12263个基因是上调表达的,有18231个基因是下调表达的。进一步分析表明,与对照相比,ABA合成代谢途径的各类基因在转基因黄檗中呈现出从上游基因上调到下游基因下调逐渐过渡的趋势,而且ABA的生物合成途径的最后一个关键基因abscisic aldehyde oxidase (AAO)的同源基因全部下调表达。另外,调控NO合成的NOS和NR基因都上调表达。
Late embryogenesis abundant proteins (LEA proteins) is a group of multifunctional adversity protein, which accumulating late in plant seeds embryogenesis. The results showed that LEA can be induced by drought and ABA stresses, and the expression level is positively relative to ABA content. Many studies indiated that LEA overexpression could improve the drought resisitant ability. However, up to now, less information on LEA functional moleculat mechanism and not perfect understanding of structure and function were known, also lots of controversy exsited. In this study, ThLEA3transgenic Phellodendron amurense Rupr. was used as research object, by the methods of physiological indexes change, stomatal closure degree, fluorescence molecular probe, SIET technology and Illumina high-throughout sequencing, change of NO, H2O2and ABA in stomatal closure process were investigated. We firstly researched the mechanism of rapidly stomatal closure under osmotic stress, and speculated the pobable molecular protective mechanism under this condition. The maining results are listed as following:
(1) After7%PEG6000treated for10h, the leaves of non-transgenic plant apparently wilting with servious water loss, while ThLEA3transgenic plant still grown normally, indicatingThLEA3transgenic plant confer to osmotic tolerance. Different physiological indexes results showed that there is no differece of SOD, CAT activity between non-transgenic and transgenic plant under osmotic stress, while proline, soluble protein and MDA content in transgenic plant were higher than that in the control one.
(2) SEM observation result indicate that stoma of transgenic plant closed, while stoma of non-transgenic plant was opened, even deformed under7%PEG6000, suggesting ThLEA3improved osmotic tolerance by rapidly stomatal closure.
(3) Fluorescence probe detection results showed that, NO in transgenic P. amurense stoma was stronger than that in non-transgenic plant. But H2O2fluorescence result was opposite to NO. Meanwhile,Ca2+fluorescence in transgenic plant stoma was higher under ABA, H2O2, NO donror SNP and PEG6000treatment, also induced stomatal closure. SIET result showed that Ca2+was influxed, K+and H+were effluxed under PEG6000stress, finally, induced stomatal closure. But Ca2+、H+、K+in stoma of non-transgenic plant effluxed, resulting in gard cell damaged and not closed.
(4) Content of endogenious ABA in non-transgenic plant leaves was higher than that in transgeni plant leaves with non-treatment. After7%PEG6000treatment, ABA in both ones greatly increased, however, ABA content in non-transgenic plant leaves was still higher than this in transgenic plant.
(5) To further verify the expression of relative genes at trascrition level, transcriptome of non-and transgenic plant leaves were analyzed by Illumina high-throughout sequencing. The sequencing results indicated that ThLEA3overexpression changed expression level of30,494genes, among which,12263were up-regulated, and18231were down-regulated. Further analysis results showed that genes involve in ABA synthesis pathway presents the transition from up-rugulaion to down-regulation trend in transgenic plant compared with non-transgenic plant. In addition, the homologous genes of abscisic aldehyde oxidase, the last key step, involve in ABA synthesis were all down-regulated. Interestingly, all the gene relative to NO synthesis, such as NOS and NR were up-rugulated.
(1)经7%PEG6000处理10小时后,对照的黄檗叶片有明显的萎蔫现象,叶片严重失水；而转ThMT3转基因黄檗的叶片保持正常,说明ThLEA3基因的过表达提高了转基因黄檗的抗渗透胁迫能力。各种生理指标的比较结果表明,在渗透胁迫下,与对照相比,转基因黄檗的SOD、CAT的活性没有明显的差异,而脯氨酸含量、可溶性蛋白含量、丙二醛(MDA)含量均高于非转基因对照。
(2)扫描电镜观察结果表明,7%PEG6000处理5小时后,转基因株系的气孔关闭,而非转基因对照的气孔是开着的,同时保卫细胞发生变形,说明ThLEA3转基因黄檗组培苗通过气孔的快速关闭来抵抗渗透胁迫。
(3)荧光分子探针检测结果表明,转基因黄檗气孔中的NO荧光强度高于对照,而转基因黄檗气孔中的H202荧光强度低于对照。ABA、H2O2、SNP及PEG6000处理能引起转基因株系的气孔中Ca2+的荧光强度,并引起气孔关闭；而不能引起对照的气孔关闭。非损伤微测技术检测结果表明PEG6000处理引起转基因植株气孔中Ca2+内流,H+、K+外流,最终导致气孔的关闭,而对照气孔中Ca2+、H+、K+全部外流,导致气孔不能关闭和保卫细胞的破坏。
(4)内源ABA含量检测结果表明,在渗透处理之前非转基因对照(NT)叶片中ABA含量高于ThLEA3转基因植株。而7%PEG6000处理之后,两者ABA的含量提高幅度都很大,但是胁迫后的还是对照叶片中ABA含量高于转基因叶片。
(5)为进一步在基因水平上验证相关基因的表达,我们利用Illumina高通量测序技术测定了转基因及对照组培苗叶片的转录组。测序分析结果表明,ThLEA3转基因引起大量基因表达量的变化,即有30,494个差异表达的基因,其中有12263个基因是上调表达的,有18231个基因是下调表达的。进一步分析表明,与对照相比,ABA合成代谢途径的各类基因在转基因黄檗中呈现出从上游基因上调到下游基因下调逐渐过渡的趋势,而且ABA的生物合成途径的最后一个关键基因abscisic aldehyde oxidase (AAO)的同源基因全部下调表达。另外,调控NO合成的NOS和NR基因都上调表达。
Late embryogenesis abundant proteins (LEA proteins) is a group of multifunctional adversity protein, which accumulating late in plant seeds embryogenesis. The results showed that LEA can be induced by drought and ABA stresses, and the expression level is positively relative to ABA content. Many studies indiated that LEA overexpression could improve the drought resisitant ability. However, up to now, less information on LEA functional moleculat mechanism and not perfect understanding of structure and function were known, also lots of controversy exsited. In this study, ThLEA3transgenic Phellodendron amurense Rupr. was used as research object, by the methods of physiological indexes change, stomatal closure degree, fluorescence molecular probe, SIET technology and Illumina high-throughout sequencing, change of NO, H2O2and ABA in stomatal closure process were investigated. We firstly researched the mechanism of rapidly stomatal closure under osmotic stress, and speculated the pobable molecular protective mechanism under this condition. The maining results are listed as following:
(1) After7%PEG6000treated for10h, the leaves of non-transgenic plant apparently wilting with servious water loss, while ThLEA3transgenic plant still grown normally, indicatingThLEA3transgenic plant confer to osmotic tolerance. Different physiological indexes results showed that there is no differece of SOD, CAT activity between non-transgenic and transgenic plant under osmotic stress, while proline, soluble protein and MDA content in transgenic plant were higher than that in the control one.
(2) SEM observation result indicate that stoma of transgenic plant closed, while stoma of non-transgenic plant was opened, even deformed under7%PEG6000, suggesting ThLEA3improved osmotic tolerance by rapidly stomatal closure.
(3) Fluorescence probe detection results showed that, NO in transgenic P. amurense stoma was stronger than that in non-transgenic plant. But H2O2fluorescence result was opposite to NO. Meanwhile,Ca2+fluorescence in transgenic plant stoma was higher under ABA, H2O2, NO donror SNP and PEG6000treatment, also induced stomatal closure. SIET result showed that Ca2+was influxed, K+and H+were effluxed under PEG6000stress, finally, induced stomatal closure. But Ca2+、H+、K+in stoma of non-transgenic plant effluxed, resulting in gard cell damaged and not closed.
(4) Content of endogenious ABA in non-transgenic plant leaves was higher than that in transgeni plant leaves with non-treatment. After7%PEG6000treatment, ABA in both ones greatly increased, however, ABA content in non-transgenic plant leaves was still higher than this in transgenic plant.
(5) To further verify the expression of relative genes at trascrition level, transcriptome of non-and transgenic plant leaves were analyzed by Illumina high-throughout sequencing. The sequencing results indicated that ThLEA3overexpression changed expression level of30,494genes, among which,12263were up-regulated, and18231were down-regulated. Further analysis results showed that genes involve in ABA synthesis pathway presents the transition from up-rugulaion to down-regulation trend in transgenic plant compared with non-transgenic plant. In addition, the homologous genes of abscisic aldehyde oxidase, the last key step, involve in ABA synthesis were all down-regulated. Interestingly, all the gene relative to NO synthesis, such as NOS and NR were up-rugulated.
引文
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