稻瘟菌诱导性水稻OsFd和OsFNR基因的cDNA克隆与诱导表达
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摘要
对受病原物诱导的植物基因的研究有助于我们进一步了解植物—病原物互作机理,并可以将这些防卫相关基因应用到植物抗病基因工程中。本研究开展了受稻瘟菌侵染诱导表达的水稻OsFd和OsFNR的cDNA克隆与诱导表达工作;并进行了OsFd和OsFNR的水稻转化和原核表达。
植物Fd(Ferredoxin,铁氧还蛋白)和FNR(Ferredoxin- NADP+ Reductase,铁氧还蛋白-NADP+还原酶) 都是多基因家族,在植物体中可分为光合作用与非光合作用两个类型。光合型的位于植物叶绿体内,其作用是将来自于光合系统I的电子传递给NADP+,生成NADPH。非光合Fd和FNR,主要分布在根和成熟果实等非光合器官中,可能参与亚硝酸还原酶,亚硫酸盐还原酶,谷氨酸合成酶等的还原作用。
本文以从非亲和性稻瘟菌小种131侵染的、水稻品种爱知旭的稻叶中获得的、2个cDNA片段20A03和21C01作探针,筛选非亲和性稻瘟菌侵染的水稻稻叶cDNA文库,获得了相应的全长cDNA克隆OsFd、OsFNR。
OsFd是新发现的水稻Fd基因,其cDNA全长为848bp,完整的ORF为459bp,推定的蛋白产物有153个氨基酸,分子量16KD。该基因的cDNA序列及其推定的氨基酸序列与玉米FdIII基因同源性最高,分别为83%和81%。
Northern杂交分析表明:非亲和性稻瘟菌小种131能在6小时内强烈诱导OsFd转录,亲和性稻瘟菌小种在48小时内开始诱导OsFd转录;在SA(水杨酸)喷洒的稻叶中,OsFd基因转录物在处理后6个小时内明显地诱导积累,而且在处理后的6个小时至72个小时范围内,OsFd转录物的积累量逐渐增加。与水稻基因组序列的同源性分析表明:OsFd在水稻基因组的第三条染色体上,只有一个拷贝,有一个内含子。
OsFNR基因全长1495bp,开放阅读框为1137bp(103-1238bp)。该基因编码378个氨基酸、分子量为41.6KD的蛋白。 BLAST序列同源性分析表明,该基因cDNA序列与水稻胚中的FNR同源,后者仅有电子登记,但没有进一步的研究报道。
Northern杂交分析表明,在亲和性稻瘟菌007小种处理中,在接种后的60小时内
OsFNR被诱导转录;在131接种的处理中,接种6小时内OsFNR就大量转录积累,一直到接种后的72小时OsFNR转录物积累的水平持续不减弱;在SA处理下,6个小时内已经有转录物积累,而且在取样的72个小时范围内,OsFNR基因转录物的积累量逐渐增加。水稻基因组序列分析结果表明OsFNR在水稻基因组的第七条染色体上,只有一个拷贝,有6个内含子。
通过与发表的植物Fd、FNR的同源性分析,推断OsFd和OsFNR属于非光合类型。根据OsFd、OsFNR受稻瘟菌诱导表达动态特征:受亲和性稻瘟菌诱导转录在48小时后,时间晚、表达量低;受非亲和性稻瘟菌浸染6小时内就高水平转录积累,直到72小时水平不降低,另外受SA诱导在6小时内就高水平转录积累,直到72小时水平持续升高,SA被证明是许多防御基因表达和植物表现系统获得抗性所必需的。综合以上分析,推断OsFd和OsFNR都与水稻抗稻瘟病的防卫反应密切相关;它们是催化同一氧化还原反应的相邻的两个电子供体,这个氧化还原反应可能是水稻抗稻瘟病的防卫反应的关键组成部分。
为证明OsFd和OsFNR的生化特性,构建了它们的重组原核表达载体pGEX-4T-3/OsFd和pGEX-4T-3/OsFNR,并分别诱导其在大肠杆菌中表达,均诱导出大小与期望相符的特异蛋白,初步证明OsFd和OsFNR的推定蛋白产物是正确的。
为进一步证明OsFd和OsFNR在水稻抗稻瘟病中的作用,构建了它们的sense和hairpin表达载体:pCAMBIA1301/OsFd-sense、pCAMBIA1301/OsFd-hairpin、 pCAMBIA1301/OsFNR-sense 、pCAMBIA1301/OsFNR-hairpin,并通过农杆菌介导法将这四种表达载体分别转入水稻品种爱知旭中,分别获得83、110、43和121株转基因水稻。为抗病性检测等进一步的研究打下基础。
此外,克隆四个水稻乙醛酸循环酶基因:OsICL(异柠檬酸裂解酶)、OsCS(柠檬酸合成酶)、OsMS(苹果酸合成酶)和OsMDH(苹果酸脱氢酶)。通过分析水稻乙醛酸循环的标志酶OsICL在稻瘟菌侵染条件下的转录积累动态,认为乙醛酸循环受亲和性稻瘟菌侵染的诱导。
The functional identification and expressing characterization of plant genes that are induced during interaction with pathogen will be a great help to reveal the mechanism of plant-pathogen interaction; Isolation of these genes may provide a good choice for engineering plant disease resistance.
In this study, two differential displayed fragments, 21A03 and 21C01, were isolated from rice leaves (cv. Aichiasahi) infected by an incompatible race 131 of Magnaporthe grisea and used as probes to screen the cDNA library, which was constructed using mRNA from the p131 pathogen infected leaves. By this screening, two kinds of full length cDNA clones were obtained, of which, one is a Fd (ferredoxin) gene named as OsFd and the other is FNR (ferredoxin NADP+ oxidoreductase) gene named as OsFNR.
The cDNA sequence of OsFd is 848bp and contains one intact open reading frame. The cDNA sequence shares 83% identity with that of maize FdIII from maize. It encodes a protein of 153 amino acid residues. The deduced amino acid sequence shares 81% identity with that of maize FdIII. OsFd transcript in rice leaves can be induced by compatible and incompatible blast fungus infection and SA. Genomic Rice genome contained one copy sequence homologous to OsFd gene.
The size of nucleotide sequence of OsFNR is 1495bp, contained one open reading frame encoding 378 amino acid residues. Sequence analysis indicated that the cDNA sequence of OsFNR is identical to that of FNR from rice embryo. Northern blotting showed that OsFNR transcript in rice leaves can be induced by incompatible blast fungus infection and SA, suggesting that the OsFNR induction be involved in the defense of rice to M. grisea.
Homologous analysis to Rice Genome Database suggests that OsFNR exists as one copy in rice genome.
In order to further analyze the biochemical characteristics of OsFd and OsFNR the prokaryotic expressing recombination vectors pGEX-4T-3/OsFd and pGEX-4T-3/OsFNR
were constructed and transformed into BL21 (DE3), respectively. The optimum expressing conditions for the both were approached, SDS-PAGE analysis showed that the products of recombinants OsFd and OsFNR were both specific.
To clarify if OsFd and OsFNR are required for resistance in rice to blast, two types of rice expressing vectors (sense and hairpin) of OsFd and OsFNR were constructed and cloned into binary pCAMBIA 1301, and then transformed into rice by Agrobacterium tumefaciens-mediated transformation (cv. Aichiasahi), respectively. T0 transgenic rice lines of the four rice expressing vectors were obtained. There are 83, 110 ,43, 121 lines in transformation of sense and hairpin expressing vectors of OsFd and OsFNR , respectively.
In conclusion, OsFd and OsFNR participate in the process of rice resistance to blast.
On the other hand, we cloned the four genes from the glyoxylate cycle in rice. They are ICL( isocitrate lyase), CS(citrate synthase),MDH(malate dehydrogenase), MS(malate synthase) genes and they are named as OsICL,. OsCS ,OsMDH and OsMS, respectively. OsICL transcript in rice leaves can be induced by comapatible blast fungi infection.
植物Fd(Ferredoxin,铁氧还蛋白)和FNR(Ferredoxin- NADP+ Reductase,铁氧还蛋白-NADP+还原酶) 都是多基因家族,在植物体中可分为光合作用与非光合作用两个类型。光合型的位于植物叶绿体内,其作用是将来自于光合系统I的电子传递给NADP+,生成NADPH。非光合Fd和FNR,主要分布在根和成熟果实等非光合器官中,可能参与亚硝酸还原酶,亚硫酸盐还原酶,谷氨酸合成酶等的还原作用。
本文以从非亲和性稻瘟菌小种131侵染的、水稻品种爱知旭的稻叶中获得的、2个cDNA片段20A03和21C01作探针,筛选非亲和性稻瘟菌侵染的水稻稻叶cDNA文库,获得了相应的全长cDNA克隆OsFd、OsFNR。
OsFd是新发现的水稻Fd基因,其cDNA全长为848bp,完整的ORF为459bp,推定的蛋白产物有153个氨基酸,分子量16KD。该基因的cDNA序列及其推定的氨基酸序列与玉米FdIII基因同源性最高,分别为83%和81%。
Northern杂交分析表明:非亲和性稻瘟菌小种131能在6小时内强烈诱导OsFd转录,亲和性稻瘟菌小种在48小时内开始诱导OsFd转录;在SA(水杨酸)喷洒的稻叶中,OsFd基因转录物在处理后6个小时内明显地诱导积累,而且在处理后的6个小时至72个小时范围内,OsFd转录物的积累量逐渐增加。与水稻基因组序列的同源性分析表明:OsFd在水稻基因组的第三条染色体上,只有一个拷贝,有一个内含子。
OsFNR基因全长1495bp,开放阅读框为1137bp(103-1238bp)。该基因编码378个氨基酸、分子量为41.6KD的蛋白。 BLAST序列同源性分析表明,该基因cDNA序列与水稻胚中的FNR同源,后者仅有电子登记,但没有进一步的研究报道。
Northern杂交分析表明,在亲和性稻瘟菌007小种处理中,在接种后的60小时内
OsFNR被诱导转录;在131接种的处理中,接种6小时内OsFNR就大量转录积累,一直到接种后的72小时OsFNR转录物积累的水平持续不减弱;在SA处理下,6个小时内已经有转录物积累,而且在取样的72个小时范围内,OsFNR基因转录物的积累量逐渐增加。水稻基因组序列分析结果表明OsFNR在水稻基因组的第七条染色体上,只有一个拷贝,有6个内含子。
通过与发表的植物Fd、FNR的同源性分析,推断OsFd和OsFNR属于非光合类型。根据OsFd、OsFNR受稻瘟菌诱导表达动态特征:受亲和性稻瘟菌诱导转录在48小时后,时间晚、表达量低;受非亲和性稻瘟菌浸染6小时内就高水平转录积累,直到72小时水平不降低,另外受SA诱导在6小时内就高水平转录积累,直到72小时水平持续升高,SA被证明是许多防御基因表达和植物表现系统获得抗性所必需的。综合以上分析,推断OsFd和OsFNR都与水稻抗稻瘟病的防卫反应密切相关;它们是催化同一氧化还原反应的相邻的两个电子供体,这个氧化还原反应可能是水稻抗稻瘟病的防卫反应的关键组成部分。
为证明OsFd和OsFNR的生化特性,构建了它们的重组原核表达载体pGEX-4T-3/OsFd和pGEX-4T-3/OsFNR,并分别诱导其在大肠杆菌中表达,均诱导出大小与期望相符的特异蛋白,初步证明OsFd和OsFNR的推定蛋白产物是正确的。
为进一步证明OsFd和OsFNR在水稻抗稻瘟病中的作用,构建了它们的sense和hairpin表达载体:pCAMBIA1301/OsFd-sense、pCAMBIA1301/OsFd-hairpin、 pCAMBIA1301/OsFNR-sense 、pCAMBIA1301/OsFNR-hairpin,并通过农杆菌介导法将这四种表达载体分别转入水稻品种爱知旭中,分别获得83、110、43和121株转基因水稻。为抗病性检测等进一步的研究打下基础。
此外,克隆四个水稻乙醛酸循环酶基因:OsICL(异柠檬酸裂解酶)、OsCS(柠檬酸合成酶)、OsMS(苹果酸合成酶)和OsMDH(苹果酸脱氢酶)。通过分析水稻乙醛酸循环的标志酶OsICL在稻瘟菌侵染条件下的转录积累动态,认为乙醛酸循环受亲和性稻瘟菌侵染的诱导。
The functional identification and expressing characterization of plant genes that are induced during interaction with pathogen will be a great help to reveal the mechanism of plant-pathogen interaction; Isolation of these genes may provide a good choice for engineering plant disease resistance.
In this study, two differential displayed fragments, 21A03 and 21C01, were isolated from rice leaves (cv. Aichiasahi) infected by an incompatible race 131 of Magnaporthe grisea and used as probes to screen the cDNA library, which was constructed using mRNA from the p131 pathogen infected leaves. By this screening, two kinds of full length cDNA clones were obtained, of which, one is a Fd (ferredoxin) gene named as OsFd and the other is FNR (ferredoxin NADP+ oxidoreductase) gene named as OsFNR.
The cDNA sequence of OsFd is 848bp and contains one intact open reading frame. The cDNA sequence shares 83% identity with that of maize FdIII from maize. It encodes a protein of 153 amino acid residues. The deduced amino acid sequence shares 81% identity with that of maize FdIII. OsFd transcript in rice leaves can be induced by compatible and incompatible blast fungus infection and SA. Genomic Rice genome contained one copy sequence homologous to OsFd gene.
The size of nucleotide sequence of OsFNR is 1495bp, contained one open reading frame encoding 378 amino acid residues. Sequence analysis indicated that the cDNA sequence of OsFNR is identical to that of FNR from rice embryo. Northern blotting showed that OsFNR transcript in rice leaves can be induced by incompatible blast fungus infection and SA, suggesting that the OsFNR induction be involved in the defense of rice to M. grisea.
Homologous analysis to Rice Genome Database suggests that OsFNR exists as one copy in rice genome.
In order to further analyze the biochemical characteristics of OsFd and OsFNR the prokaryotic expressing recombination vectors pGEX-4T-3/OsFd and pGEX-4T-3/OsFNR
were constructed and transformed into BL21 (DE3), respectively. The optimum expressing conditions for the both were approached, SDS-PAGE analysis showed that the products of recombinants OsFd and OsFNR were both specific.
To clarify if OsFd and OsFNR are required for resistance in rice to blast, two types of rice expressing vectors (sense and hairpin) of OsFd and OsFNR were constructed and cloned into binary pCAMBIA 1301, and then transformed into rice by Agrobacterium tumefaciens-mediated transformation (cv. Aichiasahi), respectively. T0 transgenic rice lines of the four rice expressing vectors were obtained. There are 83, 110 ,43, 121 lines in transformation of sense and hairpin expressing vectors of OsFd and OsFNR , respectively.
In conclusion, OsFd and OsFNR participate in the process of rice resistance to blast.
On the other hand, we cloned the four genes from the glyoxylate cycle in rice. They are ICL( isocitrate lyase), CS(citrate synthase),MDH(malate dehydrogenase), MS(malate synthase) genes and they are named as OsICL,. OsCS ,OsMDH and OsMS, respectively. OsICL transcript in rice leaves can be induced by comapatible blast fungi infection.
引文
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