水稻RAD21同源基因RIX4的分子特性研究
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摘要
本论文前期工作中,通过mRNA差别显示获得了受白叶枯菌(Xanthomonas oryzae pv.oyzae)诱导表达的差异cDNA片段。国际互连网Blastn查询表明该基因为新基因。本课题旨在对这一与白叶枯菌抗性相关的基因进行进一步研究。
应用RT-PCR技术获得了该基因完整ORF序列,该序列全长2,187bp,编码一个728氨基酸的预测蛋白,分子量约为80.35KD,等电点为4.65。经序列拼接获得该基因的约5.3kb的基因组DNA序列,将该基因组DNA序列与eDNA序列进行比对分析,发现RIX4基因含11个外显子和10个内含子,所有10个内含子都符合保守的/GT-AG/特征。Blastp分析结果表明,RIX4蛋白在N末端含Pfam04825,在C末端含Pfam04824保守结构域,这两个保守结构域在RAD21/REC8类蛋白中很保守,表明RIX4蛋白属于RAD21/REC8类粘合蛋白。其蛋白的二级结构由21.98% a-螺旋,7.01%延伸和71.02%随意卷曲三种结构模块组成。
在不育系和保持系水稻花粉母细胞减数分裂期幼穗中得到了RIX4基因另一转录本RIX4-5片段。对RIX4基因的5个转录本进行了序列和剪接模式的分析,结果表明第6和第7外显子序列的选择性剪接模式属于常见的选择外显子上不同的5'或3'剪接位点进行选择性剪接的模式,而第6和9外显子上序列的选择性剪接模式不符合常见的7种剪接模式,是一种新的剪接模式。稻瘟病菌的侵染有利于RIX4基因前体mRNA剪接形成RIX4-1转录本,白叶枯病菌的侵染有利于RIX4基因前体mRNA剪接形成RIX4-4转录本。
运用大肠杆菌原核表达系统表达出RIX4多肽,并通过免疫新西兰大白兔制备了多克隆抗体,以此抗体检测了植物体内RIX4蛋白的含量。在受病原菌CR3诱导的水稻IR26愈伤组织中,在诱导0～60min的水稻愈伤组织中均能检测到RIX4基因编码的全长蛋白,而且随着诱导时间的延长,检测到的蛋白量增多。在处于花粉母细胞减数分裂旺盛期的不育系和保持系的幼穗中,保持系中RIX4蛋白含量均高于不育系。在受白叶枯菌CR3侵染不同时期的IR26愈伤组织和不育系及保持系幼穗中仅能检测到RIX4基因的全长蛋白,而检测不到其它转录本所编码的蛋白。在受稻瘟菌生理小种ZB15(2001-046)诱导的近等基因系H_7R和H_7S的叶片及生理小种01-14诱导的嘉兴02-03和中早27的叶片中,RIX4蛋白含量会随着诱导时间的变化而变化,在不同的水稻品系中变化趋势不同,但在受稻瘟菌诱导的水稻叶片中均能检测到全长蛋白和RIX4-4转录本所编码的蛋白,在蛋白水平上证明了RIX4基因存在不同的转录本。
构建了RIX4基因RNA干涉表达载体,通过基因枪法对水稻粳稻品种中花11进行转化,获得9株转pCAMBIA-HANNIBAL-RIX4(461)质粒的转基因植株和12株转
The differential expressed cDNA fragments were obtained by DD-PCR from IR26 callus induced by Xanthomonas oryzae pv. oyzae;(Xoo) in previous work.. Blastn searching showed it was a novel gene then named RIX4. This work was designed to study further the splicing pattern of this gene, the relationship between them, expression characteristics and the function.The full-length ORF sequence of RIX4 gene was 2,187 bp long, encoding a 728 amino acid residues presumed protein, with MW 80.35KD, pI 4.65. The genomic DNA of RIX4 gene has about 5.3 kb. Comparison analysis of the sequences between the genomic DNA and cDNA showed that this gene has 11 exons and 10 introns, and all of the 10 introns have the /GT-AG/ consensus characteristics.The RIX4 protein has an entire N-terminal domain Pfam04825 and a C-terminal domain Pfam04824 which are conserved in previously identified RAD21/REC8 proteins. It was evidenced that RIX4 protein belongs to RAD21/REC8 cohesion protein. The secondary structure prediction of RIX4 was conducted on websites. The result showed that RIX4 consisted of 21.98% alpha helix, 7.01% extended strand and 71.02% random coil. The random coil.as the most abundant structure, penetrated through the RIX4 protein, while the alpha helix and extended strands relatively gathered to form three obvious helix/strand groups.Another novel transcript RIX4-5 was obtained from young panicles in both sterile lines and maintainer lines when the distance between the pulvinus of the flag leaf and the pulvinus of the second leaf was zero. Analysis of the sequences and splicing patterns of all the 5 RIX4 transcripts was conducted. The results showed that the alternative splicing patterns in exon 6 and 7 are identical with the typical pattern, namely, alternative splicing on the different 5'or 3' site. But the alternative splicing patterns in exon 6 and 9 are novel. The study showed that RIX4-1 transcript was produced in leaves inoculation with Magneporthe grisea(M. grisea) and RJX4-4 transcript was predominant in callus inoculation with Xoo.The fragment encoding the polypeptide of RIX4 protein N termius was cloned into pET-28a and expressed in the strain BL21 of E.coli. After induction by IPTG at 37℃,the expressed protein was purified by Ni~2+-NTA agrose column. The antiserum was raised by immunizing New Zealand rabbit and used to analysize the content of RIX4 protein in plants. The full length protein can be detected in the IR26 callus induced by pathogen Xoo. during 0~60 min, and the content increased with induction time. The RIX4 protein can also be detected in the young
panicles of both sterile lines and maintainer lines when the distance between the pulvinus of the flag leaf and the pulvinus of the second leaf was zero, and the content in maintainer lines was higher than in the former.But protein encoded by other transcripts was not detected in all the situations above. In the leaves of rice near isogenic line (NIL) H7R and H7S induced by M. grisea isolate ZB15 (2001-046) and the leaves of varieties Jaxing 02-03 and Zhongzao 27 by isolate 01-14, the content of RIX4 full-length protein varied along with the induction time, but the variation was not identical. Furthermore, both full length protein and protein encoded by RIX4-4 transcript can be detected in all leaves induced by M. grisea. Thus, our result evidenced that the RIX4 gene had different transcripts in the protein level.RNAi expression vectors were constructed and transformed into Zhonghua 11 through shotgun. 9 (pCAMBIA-HANNIBAL-iUX^l)) and 12 (pCAMBIA-HANNIBAL-iUX4(397)) transgenic plants were obtained, respectively. Identification of resistance of sense and antisense transgenic plants to Xoo showed that the RIX4 gene conferred rice more effective resistance to the pathogen.Peiai 64S is a photoperiod- and thermo- sensitive genie male sterile (PTSGMS) line of rice, which is male sterile under long day/high temperature and partial fertile under short day/low temperature. A cDNA array representing 3,328 unique rice genes was used to profile the gene expression patterns in the young panicles of Peiai 64S under these two conditions. The statistical data showed that up to 14.60% of genes exhibited up- or down-regulated expressions in the plants under long day/high temperature compared with plants under short day/low temperature, but the expression abudance of RIX4 gene was not differential. Only four genes were up-regulated while 482 genes down-regulated. Real-time PCR with all the up-regulated and 9 randomly selected down-regulated genes confirmed the differential expressions detected by the array, indicating that the constructed cDNA array is reliable. These differently expressed genes participated in almost all cell biological responses. Analysis of up- and down-regulated genes revealed distinctive changes between the mRNA abundances of MMK1 and MMK.2, both of which are analogs of MAPK, and significant down-regulation of several transcription factors. It is proposed that changes occurred in the MAPK signal transduction pathway might disturb the transcription control necessary for morphogenesis of pollens and corresponding physiological functions.
应用RT-PCR技术获得了该基因完整ORF序列,该序列全长2,187bp,编码一个728氨基酸的预测蛋白,分子量约为80.35KD,等电点为4.65。经序列拼接获得该基因的约5.3kb的基因组DNA序列,将该基因组DNA序列与eDNA序列进行比对分析,发现RIX4基因含11个外显子和10个内含子,所有10个内含子都符合保守的/GT-AG/特征。Blastp分析结果表明,RIX4蛋白在N末端含Pfam04825,在C末端含Pfam04824保守结构域,这两个保守结构域在RAD21/REC8类蛋白中很保守,表明RIX4蛋白属于RAD21/REC8类粘合蛋白。其蛋白的二级结构由21.98% a-螺旋,7.01%延伸和71.02%随意卷曲三种结构模块组成。
在不育系和保持系水稻花粉母细胞减数分裂期幼穗中得到了RIX4基因另一转录本RIX4-5片段。对RIX4基因的5个转录本进行了序列和剪接模式的分析,结果表明第6和第7外显子序列的选择性剪接模式属于常见的选择外显子上不同的5'或3'剪接位点进行选择性剪接的模式,而第6和9外显子上序列的选择性剪接模式不符合常见的7种剪接模式,是一种新的剪接模式。稻瘟病菌的侵染有利于RIX4基因前体mRNA剪接形成RIX4-1转录本,白叶枯病菌的侵染有利于RIX4基因前体mRNA剪接形成RIX4-4转录本。
运用大肠杆菌原核表达系统表达出RIX4多肽,并通过免疫新西兰大白兔制备了多克隆抗体,以此抗体检测了植物体内RIX4蛋白的含量。在受病原菌CR3诱导的水稻IR26愈伤组织中,在诱导0～60min的水稻愈伤组织中均能检测到RIX4基因编码的全长蛋白,而且随着诱导时间的延长,检测到的蛋白量增多。在处于花粉母细胞减数分裂旺盛期的不育系和保持系的幼穗中,保持系中RIX4蛋白含量均高于不育系。在受白叶枯菌CR3侵染不同时期的IR26愈伤组织和不育系及保持系幼穗中仅能检测到RIX4基因的全长蛋白,而检测不到其它转录本所编码的蛋白。在受稻瘟菌生理小种ZB15(2001-046)诱导的近等基因系H_7R和H_7S的叶片及生理小种01-14诱导的嘉兴02-03和中早27的叶片中,RIX4蛋白含量会随着诱导时间的变化而变化,在不同的水稻品系中变化趋势不同,但在受稻瘟菌诱导的水稻叶片中均能检测到全长蛋白和RIX4-4转录本所编码的蛋白,在蛋白水平上证明了RIX4基因存在不同的转录本。
构建了RIX4基因RNA干涉表达载体,通过基因枪法对水稻粳稻品种中花11进行转化,获得9株转pCAMBIA-HANNIBAL-RIX4(461)质粒的转基因植株和12株转
The differential expressed cDNA fragments were obtained by DD-PCR from IR26 callus induced by Xanthomonas oryzae pv. oyzae;(Xoo) in previous work.. Blastn searching showed it was a novel gene then named RIX4. This work was designed to study further the splicing pattern of this gene, the relationship between them, expression characteristics and the function.The full-length ORF sequence of RIX4 gene was 2,187 bp long, encoding a 728 amino acid residues presumed protein, with MW 80.35KD, pI 4.65. The genomic DNA of RIX4 gene has about 5.3 kb. Comparison analysis of the sequences between the genomic DNA and cDNA showed that this gene has 11 exons and 10 introns, and all of the 10 introns have the /GT-AG/ consensus characteristics.The RIX4 protein has an entire N-terminal domain Pfam04825 and a C-terminal domain Pfam04824 which are conserved in previously identified RAD21/REC8 proteins. It was evidenced that RIX4 protein belongs to RAD21/REC8 cohesion protein. The secondary structure prediction of RIX4 was conducted on websites. The result showed that RIX4 consisted of 21.98% alpha helix, 7.01% extended strand and 71.02% random coil. The random coil.as the most abundant structure, penetrated through the RIX4 protein, while the alpha helix and extended strands relatively gathered to form three obvious helix/strand groups.Another novel transcript RIX4-5 was obtained from young panicles in both sterile lines and maintainer lines when the distance between the pulvinus of the flag leaf and the pulvinus of the second leaf was zero. Analysis of the sequences and splicing patterns of all the 5 RIX4 transcripts was conducted. The results showed that the alternative splicing patterns in exon 6 and 7 are identical with the typical pattern, namely, alternative splicing on the different 5'or 3' site. But the alternative splicing patterns in exon 6 and 9 are novel. The study showed that RIX4-1 transcript was produced in leaves inoculation with Magneporthe grisea(M. grisea) and RJX4-4 transcript was predominant in callus inoculation with Xoo.The fragment encoding the polypeptide of RIX4 protein N termius was cloned into pET-28a and expressed in the strain BL21 of E.coli. After induction by IPTG at 37℃,the expressed protein was purified by Ni~2+-NTA agrose column. The antiserum was raised by immunizing New Zealand rabbit and used to analysize the content of RIX4 protein in plants. The full length protein can be detected in the IR26 callus induced by pathogen Xoo. during 0~60 min, and the content increased with induction time. The RIX4 protein can also be detected in the young
panicles of both sterile lines and maintainer lines when the distance between the pulvinus of the flag leaf and the pulvinus of the second leaf was zero, and the content in maintainer lines was higher than in the former.But protein encoded by other transcripts was not detected in all the situations above. In the leaves of rice near isogenic line (NIL) H7R and H7S induced by M. grisea isolate ZB15 (2001-046) and the leaves of varieties Jaxing 02-03 and Zhongzao 27 by isolate 01-14, the content of RIX4 full-length protein varied along with the induction time, but the variation was not identical. Furthermore, both full length protein and protein encoded by RIX4-4 transcript can be detected in all leaves induced by M. grisea. Thus, our result evidenced that the RIX4 gene had different transcripts in the protein level.RNAi expression vectors were constructed and transformed into Zhonghua 11 through shotgun. 9 (pCAMBIA-HANNIBAL-iUX^l)) and 12 (pCAMBIA-HANNIBAL-iUX4(397)) transgenic plants were obtained, respectively. Identification of resistance of sense and antisense transgenic plants to Xoo showed that the RIX4 gene conferred rice more effective resistance to the pathogen.Peiai 64S is a photoperiod- and thermo- sensitive genie male sterile (PTSGMS) line of rice, which is male sterile under long day/high temperature and partial fertile under short day/low temperature. A cDNA array representing 3,328 unique rice genes was used to profile the gene expression patterns in the young panicles of Peiai 64S under these two conditions. The statistical data showed that up to 14.60% of genes exhibited up- or down-regulated expressions in the plants under long day/high temperature compared with plants under short day/low temperature, but the expression abudance of RIX4 gene was not differential. Only four genes were up-regulated while 482 genes down-regulated. Real-time PCR with all the up-regulated and 9 randomly selected down-regulated genes confirmed the differential expressions detected by the array, indicating that the constructed cDNA array is reliable. These differently expressed genes participated in almost all cell biological responses. Analysis of up- and down-regulated genes revealed distinctive changes between the mRNA abundances of MMK1 and MMK.2, both of which are analogs of MAPK, and significant down-regulation of several transcription factors. It is proposed that changes occurred in the MAPK signal transduction pathway might disturb the transcription control necessary for morphogenesis of pollens and corresponding physiological functions.
引文
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