水稻OsPCS分子进化及耐Cd突变体cadH-5 ASC-GSH代谢
摘要
植物螯合肽(PCs)在植物累积和抵抗重金属胁迫中起着至关重要的作用,PCs是由PC合酶(PCS)催化谷胱甘肽(GSH)通过谷氨酰半胱氨酸转肽反应得到的。1999年3个独立的实验室各自从拟南芥(Arabidopsis thaliana L.)中分离到了产生PCs的关键基因PCS。到目前为止,在高等植物中已报道了将近20个PCS基因,但在水稻(Oryzasaliva L.)中仅克隆到一个基因,且其功能尚未研究。随着模式植物拟南芥和水稻基因组序列测序的完成,通过对染色体的注释,不断有功能不明确的基因被发现,我们通过生物信息学相关手段分析NCBI、TIGR和DDBJ数据库相关水稻的序列信息,发现在水稻中存在14个PCS基因,分属两个基因亚族,即功能结构域为PHYTOCHELATIN(含133个氨基酸残基)的PCS基因亚族和功能结构域为PHYTOCHEL SYNTH(含167个氨基酸残基)的类PCS基因亚族。其中3个基因属于PCS基因亚族,11个基因属于类PCS基因亚族。RT-PCR验证基因存在性的结果表明:有三个基因(OsPCS3,OsPCS8,and OsPCS11)在根、根茎转换区、节、节间、叶柄、叶及穗中均未发现有表达,由此推测可能为假基因或基因时序表达,而其它11个基因在叶及穗中均有表达。代表PCS基因亚族的OsPCS9可以为Cd~(2+)和Zn~(2+)诱导,代表类PCS基因亚族的OsPCS7可以为Hg~(2+)和pb~(2+)诱导。由此表明:PCS基因亚族和类PCS基因亚族在植物生存与重金属脱毒中起重要作用。
前人的研究表明:GSH和PCs在Cd脱毒中起关键作用,拟南芥突变体cad2-1和cad1-3分别为GSH的PCs缺失突变体,表现出对Cd的敏感性。植物对Cd的抗性或钝感依赖于植物的抗氧化保护系统,在这个抗氧化保护系统中抗坏血酸-谷胱甘肽(ASC-GSH)循环起着至关重要的作用。尽管Cd不能激发费通型反应(Fenton-typereactions),但它可以间接地激活质膜NADPH氧化酶,从而减少受Cd胁迫组织活性氧(ROS)的产生。我们用0.75mM CdCl_2在水稻T-DNA/Ds突变体库筛选到一个Cd耐受突变体cadH-5。用0.5mM CdCl_2持续处理2d、4d、6d、8d及12d,结果表明:在根、茎、叶中突变体与野生型相比均积累较多的Cd,因此突变体是一个相对于野生型是Cd高积累的植物;亚细胞分布显示,细胞壁、细胞液及膜系统中突变体与野生型相比均积累较多的Cd,但细胞器中的累积则比较复杂:同时发现,不管是哪个处理点,突变体中PCs含量均高于野生型,而过氧化氢则相反,野生型高于突变体;GSH/GSSG、ASC/DHA和NADPH/NADP~+的比率,突变体高于野生型;APX、GR、DHAR和MDHAR的活性也是突变体高于野生型。我们的结果表明,持续用0.5mMCdCl_2处理,还原态的水平突变体高于野生型,参与的相关酶活性也是突变体高于野生型。因此,突变体对Cd耐受的原因,是因为其有更好的抗氧化保护系统。
Phytochelatin synthases (PCS) are cytosol proteins that may play vital roles in heavy metal detoxicity in bacteria, yeasts, plants, or worms. More than 10 PCS-encoded genes from different species have been reported during the last decade. But, there is little known in rice. We identified 3 putative PCS and 11 putative PCS-like genes in rice genome by the bioinformatic analysis of recently completed rice genome data, classified them into 2 subfamilies on the basis of domain sequence similarity by constructing phylogenetic trees, and localized them in rice chromosomes. RT-PCR was performed to demonstrate that 3 putative PCS sequences (OsPCS3, OsPCS8, and OsPCS11) could not be detected in the roots, stem base, nodes, internodes or petioles, but all the remaining 11 putative rice PCS genes were found to be expressed in the leaves and panicles, OsPCS7 was induced by Hg~(2+) and Pb~(2+), while OsPCS9 was induced by Cd~(2+) and Zn~(2+) in roots, suggesting that PCS or PCS-like genes might play crucial roles in survival or in different heavy metal detoxification.
Screening a Cd tolerance and accumulation high yield cereal is a promising method for phytoremidiation, with this purpose, rice plants were mutagenized by Agrobacterium tumefaciens- based gene derived system and hydroponically screend a Cd tolerance mutant cadH-5. Exposure with 0.75mM CdCl_2 for 15d, significant phenotypic differences between the wild-type (WT) and the mutant could be easily distinguished. Successive exposure with 0.5mM CdCl2 for 0d (as control), 2d, 4d, 6d, 8d, or 12d, results showed that (1) Cd accumulated to higher leves in roots, stems and leaves. Subcellular distribution of Cd indicated that cell wall, sytosol and membrane accumulated more Cd in mutant cadH-5 than WT, unfortunately, in cell wall, Cd accumulation was varied, (2) hydrogen peroxide (H_2O_2) accumulation was higher in WT, while cell wall polysaccharide and phytochelations (PCs) were lower compared to the mutant cadH-5, (3) GSH/GSSG, ASC/DHA and NADPH/NADP~+ ratios were lower in the WT than in the mutant cadH-5, and (4) ascorbate peroxidase (APX, EC 1.11.1.11), glutathione peroxidase (GR, EC 1.6.4.2), dehydroascorbate reductase (DHAR, EC 1.8.5.1) and monodehydroascorbate reductase (MDHAR, EC 1.6.5.4) activities were lower in WT compared to the mutant cadH-5. Our results suggest that under successive Cd stress, the ASC-GSH cycle was inhibited more in the WT than in the cadH-5 plants, rendering the WT less able to scavenge reactive oxygen species (ROS). The study provides a possible physiological explanation for the mutant cadH-5 is higher accumulation and tolerance to Cd toxicity.
前人的研究表明:GSH和PCs在Cd脱毒中起关键作用,拟南芥突变体cad2-1和cad1-3分别为GSH的PCs缺失突变体,表现出对Cd的敏感性。植物对Cd的抗性或钝感依赖于植物的抗氧化保护系统,在这个抗氧化保护系统中抗坏血酸-谷胱甘肽(ASC-GSH)循环起着至关重要的作用。尽管Cd不能激发费通型反应(Fenton-typereactions),但它可以间接地激活质膜NADPH氧化酶,从而减少受Cd胁迫组织活性氧(ROS)的产生。我们用0.75mM CdCl_2在水稻T-DNA/Ds突变体库筛选到一个Cd耐受突变体cadH-5。用0.5mM CdCl_2持续处理2d、4d、6d、8d及12d,结果表明:在根、茎、叶中突变体与野生型相比均积累较多的Cd,因此突变体是一个相对于野生型是Cd高积累的植物;亚细胞分布显示,细胞壁、细胞液及膜系统中突变体与野生型相比均积累较多的Cd,但细胞器中的累积则比较复杂:同时发现,不管是哪个处理点,突变体中PCs含量均高于野生型,而过氧化氢则相反,野生型高于突变体;GSH/GSSG、ASC/DHA和NADPH/NADP~+的比率,突变体高于野生型;APX、GR、DHAR和MDHAR的活性也是突变体高于野生型。我们的结果表明,持续用0.5mMCdCl_2处理,还原态的水平突变体高于野生型,参与的相关酶活性也是突变体高于野生型。因此,突变体对Cd耐受的原因,是因为其有更好的抗氧化保护系统。
Phytochelatin synthases (PCS) are cytosol proteins that may play vital roles in heavy metal detoxicity in bacteria, yeasts, plants, or worms. More than 10 PCS-encoded genes from different species have been reported during the last decade. But, there is little known in rice. We identified 3 putative PCS and 11 putative PCS-like genes in rice genome by the bioinformatic analysis of recently completed rice genome data, classified them into 2 subfamilies on the basis of domain sequence similarity by constructing phylogenetic trees, and localized them in rice chromosomes. RT-PCR was performed to demonstrate that 3 putative PCS sequences (OsPCS3, OsPCS8, and OsPCS11) could not be detected in the roots, stem base, nodes, internodes or petioles, but all the remaining 11 putative rice PCS genes were found to be expressed in the leaves and panicles, OsPCS7 was induced by Hg~(2+) and Pb~(2+), while OsPCS9 was induced by Cd~(2+) and Zn~(2+) in roots, suggesting that PCS or PCS-like genes might play crucial roles in survival or in different heavy metal detoxification.
Screening a Cd tolerance and accumulation high yield cereal is a promising method for phytoremidiation, with this purpose, rice plants were mutagenized by Agrobacterium tumefaciens- based gene derived system and hydroponically screend a Cd tolerance mutant cadH-5. Exposure with 0.75mM CdCl_2 for 15d, significant phenotypic differences between the wild-type (WT) and the mutant could be easily distinguished. Successive exposure with 0.5mM CdCl2 for 0d (as control), 2d, 4d, 6d, 8d, or 12d, results showed that (1) Cd accumulated to higher leves in roots, stems and leaves. Subcellular distribution of Cd indicated that cell wall, sytosol and membrane accumulated more Cd in mutant cadH-5 than WT, unfortunately, in cell wall, Cd accumulation was varied, (2) hydrogen peroxide (H_2O_2) accumulation was higher in WT, while cell wall polysaccharide and phytochelations (PCs) were lower compared to the mutant cadH-5, (3) GSH/GSSG, ASC/DHA and NADPH/NADP~+ ratios were lower in the WT than in the mutant cadH-5, and (4) ascorbate peroxidase (APX, EC 1.11.1.11), glutathione peroxidase (GR, EC 1.6.4.2), dehydroascorbate reductase (DHAR, EC 1.8.5.1) and monodehydroascorbate reductase (MDHAR, EC 1.6.5.4) activities were lower in WT compared to the mutant cadH-5. Our results suggest that under successive Cd stress, the ASC-GSH cycle was inhibited more in the WT than in the cadH-5 plants, rendering the WT less able to scavenge reactive oxygen species (ROS). The study provides a possible physiological explanation for the mutant cadH-5 is higher accumulation and tolerance to Cd toxicity.
引文
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