马铃薯Y病毒CP基因不同区段对发夹RNA和人工的miRNA介导的病毒抗性的影响
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摘要
基因沉默是生物体长期以来形成的一种防御机制,阻止外源基因、转座因子、病毒等外源核酸的侵入,保护生物基因组的完整性。小干扰RNA(small interfering RNA, siRNA)和微小RNA(microRNA, miRNA)是两种序列特异性地转录后基因表达的调节因子,是沉默RNA的最主要组成部分。RNA介导的病毒抗性(RNA mediated virus resistance, RMVR)是RNA沉默的一种表现形式,侵入植物细胞的病毒基因组与转基因转录产物具有部分同源性,因此在转录后基因沉默所导致的RNA降解过程中,RNA降解系统识别侵入细胞内的病毒基因组并将其与转基因RNA一起降解。RMVR具有抗病程度高(近乎免疫)、抗性持久、生物安全性高等优点。在基因沉默的研究中已发现“位置效应”的存在,siRNA的有效性可能受到靶序列所处的位置影响。在RNA介导的植物病毒抗性研究中,目前尚未见有关“位置效应”的系统报道。本研究以马铃薯Y病毒外壳蛋白为靶基因分别以hpRNA和人工的miRNA两种方式构建RNA干扰的植物表达载体,系统性的比较PVY CP基因不同区段对RNA介导的病毒抗性的影响,为寻找引发基因沉默的有效片段及更好利用RMVR培育抗病毒转基因植物提供了依据。研究结果和主要结论如下:
(1)马铃薯Y病毒CP基因不同区段对发夹RNA介导的病毒抗性的影响将PVY CP基因人工的划分为16段各50bp的区段。通过PCR扩增,得到相应的不同区段的正向和反向的片段,通过酶切连入植物表达载体pROKⅡ中;热激法转化大肠杆菌DH5α,筛选并获得hpRNA结构的植物表达载体pROK CPs(pROK CP9~pROK CP16)。
用成功构建的16个植物表达载体(另外8个为吴斌构建)瞬时侵染本生烟验证其有效性,Northern Blot结果显示,16个植物表达载体均能在植物体内成功表达产生siRNA,且瞬时表达的siRNA能有效地下调靶基因PVY CP的表达。
将构建的8个植物表达载体利用冻融法导入农杆菌LBA4404。叶盘法转化烟草NC89,经卡那霉素抗性筛选和PCR检测确定转基因植株。T0代转基因植株自交,种子置于含卡那霉素的筛选培养基上进行筛选,共得到的pROK CP9~pROK CP16各50株、62株、56株、55株、45株、63株、72株和55株T1代转基因植株。抗病性分析发现除了转基因株系pROK CP1、pROK CP2、pROK CP3、pROK CP4未获得抗性植株外,其余12个转基因株系中抗性植株的比例分别为27.15%、6.27%、67.65%、70.83%、66.01%、61.34%、66.04%、23.59%、11.11%、55.56%、77.78%和67.25%。结果表明在利用hpRNA介导的抗PVY病毒的研究中,以PVY CP基因上的50bp序列为茎足以介导宿主植株的基因沉默的产生,但由于hpRNA靶位置的差异产生不同的抗病效率,靶向于3′端nt701~nt750区段表现最高水平的抗性;且以CP基因3′端为靶序列的hpRNA转基因植株比以5′端为靶序列的植株更能有效地介导对PVY的抗性。转基因植株的Northern杂交分析表明,转基因都在RNA水平上得到了表达,抗病植株中RNA的积累量明显低于同类型的感病植株,抗性与RNA积累量呈负相关,证实病毒抗性是由RNA介导的;对转基因植株的siRNA表达量的分析,表明植株对病毒的敏感程度与siRNA积累量之间没有明显的相关性。
对T2代的转基因植株的抗病性分析结果发现,几乎所有的抗性转基因植株的后代均表现很强的抗病毒侵染能力,说明由hpRNA介导的PVY抗性在T2代植株中能够稳定遗传。
(2)马铃薯Y病毒CP基因不同区段对人工的miRNA介导的病毒抗性的影响以PVY CP基因的全长cDNA序列为靶基因,根据miRNA的特征选取8个不同的位置,M1(nt96~nt115)、M2(nt140~nt159)、M3(nt322~nt341)、M4(nt380~nt399)、M5(nt475~nt494)、M6(nt567~nt586)、M7(nt679~nt698)和M8(nt735~nt754)为amiRNA的靶区。选用拟南芥miR319a前体作为amiRNA表达的骨架,通过PCR扩增的方式替代掉原有的pre miR319a中具有生物学功能的miRNA和miRNA*,得到含有amiRNA和amiRNA*的DNA片段。将扩增产物连入植物表达载体pROKⅡ中。热激法转化大肠杆菌DH5α,筛选并获得重组表达载体pROK amiRcp1~pROK amiRcp8。
用构建的amiRNA表达载体瞬时侵染本生烟,Northern Blot验证其有效性,结果显示,8个amiRNA植物表达载体均能在植物体内成功表达amiRNA,且瞬时表达的amiRNA能有效地下调靶PVY CP基因的表达。
将成功构建的amiRNA表达载体利用冻融法导入农杆菌LBA4404。叶盘法转化烟草NC89。再生植株经卡那霉素抗性筛选和PCR检测确定为转基因植株。T0代自交后,经卡那霉素筛选和PCR鉴定,分别获得pROK amiRcp1~pROK amiRcp8的T1代转基因植株98株、96株、114株、116株、117株、111株、110株和116株。抗病性结果显示,表达靶向于马铃薯Y病毒CP基因的amiRNA转基因植株能介导对马铃薯Y病毒的抗性,抗性比例分别为37.68%、50.29%、53.76%、37.75%、29.86%、17.05%、26.27%、64.69%。针对马铃薯Y病毒CP基因不同区段设计的amiRcps其转基因植株所介导的抗病性效果之间存在差异,靶向于3′端的amiRcp 8(nt735~nt754)能表现更高水平的抗性。转基因植株的Northern杂交分析表明,转基因都在RNA水平上得到了表达,且植株的抗性与转基因的表达量成负相关,表明amiRNA介导的病毒抗性是由RNA介导的。对转基因植株的amiRNA的Northern Blot分析,结果显示,所有抗病转基因植株中都有amiRNA存在,且植株的抗性与amiRNA的表达量成正相关。
对T2代的转基因植株的抗病性分析结果发现,所有的抗性转基因植株的后代都表现高的抗病率,表明由amiRNA介导的PVY抗性在T2代植株中能够稳定遗传。
RNA silencing is a host defence against invasive nucleic acid, such as viruses or transposable retro elements, preserving the integrality of biology. Small interfering and microRNA which is the most important parts of RNA silencing system are the inducing factor of post transcript gene silencing. RNA mediated virus resistance (RMVR) is regarded as an effective strategy in plant resistance against viruses. The expression of virus specific dsRNA which has homology to the invasive virus will active the post transcriptional gene silencing response of the host plant. The RNA silencing system will recognize complementary invasive viral RNAs and triggers degradation of both the transgene RNA and the corresponding viral RNA. RMVR has been widely used in plant antiviral research for its advantage, such as nearly immunity; sustaining resistance; no biosafety problem. In the application of gene silencing, the depending of the postion of target sequence has been found. There is no systematically study in the RNA mediated virus resistance. In our studies, we designed hairpin RNA constructs and artificial miRNA consructs based on different regions of the CP gene of PVY and generated transgenic plants. We aim at disserting if these kinds of tansgene construction could make transgenic plant exhibit different resistant ratio. The result will contribute to the selection of the target sequence and successfully apply the strategy of RMVR, which has an important theory and application value. The main results and conclusions presented in this thesis are as follows:
(1) Different origins of PVY CP gene influence the resistance of hairpin expressing plants against PVY
The PVY CP gene was divided into 16 regions. For every region, two PCR fragments were amplified from the whole length of the CP gene. Two PCR fragments were inverted inserted into binary vector pROKⅡ. The ligated products were transferred into E.coli DH5αby heat shock. The plant expression vectors obtained were dubbed pROK CPs (pROK CP9~pROK CP16).
The 16 hpRNA constructs (the other 8 hpRNA expression vectors were constructed by Wu Bin) were transferred into N. benthamiana through Agrobacterium tumfaciens mediated transient infection. Northern blot analysis revealed siRNAs were detected in all pROK CPs in?ltrated leaves in transient assay. And all 16 biologically active siRNAs that could effectively down regulate the expression of target mRNA.
The 16 hpRNA constructs were introduced into NC89 mediated by LBA4404. T1 50, 62, 56, 55, 45, 63, 72, 55 transgenic lines exhibiting kanamycin resistance and positive results in PCR tests for each vector were regenerated after T0 selfing. Virus resistance assay revealed that the 16 transgenic tobacco groups exhibited varying degrees of virus resistance in preventing PVY infection. All the transgenic plants of pROK CP1, pROK CP2, pROK CP3, and pROK CP4 are susceptive to the PVY infection; The resistance ratios of pROK CP5~ pROK CP16 are 27.15%, 6.27%, 67.65%, 70.83%, 66.01%, 61.34%, 66.04%, 23.59%, 11.11%, 55.56%, 77.78% and 67.25%. We demonstrate that 50bp length hpRNA construct is effective for RMVR; however, the hairpin constructs harboring different cDNA regions of the CP gene engendered different silencing efficiencies, the hpRNA targeting the 3′end (nt701~nt750) region exhibited the most highly virus resistance. The plants which harboring the hpRNAi constructs targeting 3′of PVY CP can induced higher virus resistance than those targeting 5′of PVY CP.
Northern blot analysis revealed that the trangene have been expressed in level of RNA, and the transcripts accumulation level of resistant plants was lower than that of the susceptible transgenic plants. There was an inverse correlation between the resistance and the amount of RNA accumulation in the transgenic plants. The results proved this resistance was RNA mediated. siRNAs were present in all selected resistant and susceptible transgenic plants; No obvious correlation was observed between the expression level of sequence speciflc siRNAs and virus resistance.
Virus assary in the T2 generation showed that most of the progeny transgenic plants of resistant plant still exhibited high resistance. The results indicated that hpRNA mediated virus resistance can be stably inherited in T2 generation.
(2) Different origins of PVY CP gene influence the resistance of artificial miRNA expressing plants against PVY We designed eight amiRNAs M1 (nt96~nt115), M2 (nt140~nt159), M3 (nt322~nt341), M4 (nt380~nt399), M5 (nt475~nt494), M6 (nt567~nt586), M7 (nt679~nt698) and M8 (nt735~nt754) derived from PVY Coat Protein RNA sequences. Arabidopsis thaliana miR319a precursor was used as backbone. Through oligonucleotide directed mutagenesis, the natural miR319 and miR319* sequences were replaced with synthetic sequences, each corresponding to a designed amiRNA that targets one region of the PVY CP gene. The generated fragment was digested and inserted into plant binary expression vector pROKⅡto generate pROK amiRcps.
The 8 amiRNA constructs were transferred into N. benthamiana. In transient assay, Northern blot analysis revealed amiRNAs were detected in all pROK amiRcps in?ltrated leaves. All plants with amiRNA expression exhibited a significantly decreased CP accumulation
The eight amiRcp expression constructs were introduced into NC89 mediated by LBA4404. After the kanamycin resistance and PCR test, T1 plants 98, 96, 114, 116, 117, 111, 110, 116 were screened. The resistance ratios of amiRcps expressing plants were 37.68%, 50.29%, 53.76%, 37.75%, 29.86%, 17.05%, 26.27%, 64.69%. Virus resistance assay revealed that expression of amiRNA can effective in preventing PVY infection; but not all amiRcps targeting viral CP sequence are equally effective. The amiRcp 8 targeting the 3′end (nt735~nt754) region exhibited high virus resistance.
Northern blots showed that the trangene have been expressed in level of RNA. There was an inverse correlation between the resistance and the amount of RNA accumulation in the transgenic plants. The results proved this resistance was RNA mediated. AmiRNAs were present in all selected resistant transgenic plants and there is a positive correlation between virus resistance and expression level of amiRNA.
Virus resistance assay in the T2 generation revealed that most of the progeny transgenic plants of resistant plant still exhibited high resistance. It indicated that amiRNA mediated virus resistance can be stably inherited in T2 generation.
(1)马铃薯Y病毒CP基因不同区段对发夹RNA介导的病毒抗性的影响将PVY CP基因人工的划分为16段各50bp的区段。通过PCR扩增,得到相应的不同区段的正向和反向的片段,通过酶切连入植物表达载体pROKⅡ中;热激法转化大肠杆菌DH5α,筛选并获得hpRNA结构的植物表达载体pROK CPs(pROK CP9~pROK CP16)。
用成功构建的16个植物表达载体(另外8个为吴斌构建)瞬时侵染本生烟验证其有效性,Northern Blot结果显示,16个植物表达载体均能在植物体内成功表达产生siRNA,且瞬时表达的siRNA能有效地下调靶基因PVY CP的表达。
将构建的8个植物表达载体利用冻融法导入农杆菌LBA4404。叶盘法转化烟草NC89,经卡那霉素抗性筛选和PCR检测确定转基因植株。T0代转基因植株自交,种子置于含卡那霉素的筛选培养基上进行筛选,共得到的pROK CP9~pROK CP16各50株、62株、56株、55株、45株、63株、72株和55株T1代转基因植株。抗病性分析发现除了转基因株系pROK CP1、pROK CP2、pROK CP3、pROK CP4未获得抗性植株外,其余12个转基因株系中抗性植株的比例分别为27.15%、6.27%、67.65%、70.83%、66.01%、61.34%、66.04%、23.59%、11.11%、55.56%、77.78%和67.25%。结果表明在利用hpRNA介导的抗PVY病毒的研究中,以PVY CP基因上的50bp序列为茎足以介导宿主植株的基因沉默的产生,但由于hpRNA靶位置的差异产生不同的抗病效率,靶向于3′端nt701~nt750区段表现最高水平的抗性;且以CP基因3′端为靶序列的hpRNA转基因植株比以5′端为靶序列的植株更能有效地介导对PVY的抗性。转基因植株的Northern杂交分析表明,转基因都在RNA水平上得到了表达,抗病植株中RNA的积累量明显低于同类型的感病植株,抗性与RNA积累量呈负相关,证实病毒抗性是由RNA介导的;对转基因植株的siRNA表达量的分析,表明植株对病毒的敏感程度与siRNA积累量之间没有明显的相关性。
对T2代的转基因植株的抗病性分析结果发现,几乎所有的抗性转基因植株的后代均表现很强的抗病毒侵染能力,说明由hpRNA介导的PVY抗性在T2代植株中能够稳定遗传。
(2)马铃薯Y病毒CP基因不同区段对人工的miRNA介导的病毒抗性的影响以PVY CP基因的全长cDNA序列为靶基因,根据miRNA的特征选取8个不同的位置,M1(nt96~nt115)、M2(nt140~nt159)、M3(nt322~nt341)、M4(nt380~nt399)、M5(nt475~nt494)、M6(nt567~nt586)、M7(nt679~nt698)和M8(nt735~nt754)为amiRNA的靶区。选用拟南芥miR319a前体作为amiRNA表达的骨架,通过PCR扩增的方式替代掉原有的pre miR319a中具有生物学功能的miRNA和miRNA*,得到含有amiRNA和amiRNA*的DNA片段。将扩增产物连入植物表达载体pROKⅡ中。热激法转化大肠杆菌DH5α,筛选并获得重组表达载体pROK amiRcp1~pROK amiRcp8。
用构建的amiRNA表达载体瞬时侵染本生烟,Northern Blot验证其有效性,结果显示,8个amiRNA植物表达载体均能在植物体内成功表达amiRNA,且瞬时表达的amiRNA能有效地下调靶PVY CP基因的表达。
将成功构建的amiRNA表达载体利用冻融法导入农杆菌LBA4404。叶盘法转化烟草NC89。再生植株经卡那霉素抗性筛选和PCR检测确定为转基因植株。T0代自交后,经卡那霉素筛选和PCR鉴定,分别获得pROK amiRcp1~pROK amiRcp8的T1代转基因植株98株、96株、114株、116株、117株、111株、110株和116株。抗病性结果显示,表达靶向于马铃薯Y病毒CP基因的amiRNA转基因植株能介导对马铃薯Y病毒的抗性,抗性比例分别为37.68%、50.29%、53.76%、37.75%、29.86%、17.05%、26.27%、64.69%。针对马铃薯Y病毒CP基因不同区段设计的amiRcps其转基因植株所介导的抗病性效果之间存在差异,靶向于3′端的amiRcp 8(nt735~nt754)能表现更高水平的抗性。转基因植株的Northern杂交分析表明,转基因都在RNA水平上得到了表达,且植株的抗性与转基因的表达量成负相关,表明amiRNA介导的病毒抗性是由RNA介导的。对转基因植株的amiRNA的Northern Blot分析,结果显示,所有抗病转基因植株中都有amiRNA存在,且植株的抗性与amiRNA的表达量成正相关。
对T2代的转基因植株的抗病性分析结果发现,所有的抗性转基因植株的后代都表现高的抗病率,表明由amiRNA介导的PVY抗性在T2代植株中能够稳定遗传。
RNA silencing is a host defence against invasive nucleic acid, such as viruses or transposable retro elements, preserving the integrality of biology. Small interfering and microRNA which is the most important parts of RNA silencing system are the inducing factor of post transcript gene silencing. RNA mediated virus resistance (RMVR) is regarded as an effective strategy in plant resistance against viruses. The expression of virus specific dsRNA which has homology to the invasive virus will active the post transcriptional gene silencing response of the host plant. The RNA silencing system will recognize complementary invasive viral RNAs and triggers degradation of both the transgene RNA and the corresponding viral RNA. RMVR has been widely used in plant antiviral research for its advantage, such as nearly immunity; sustaining resistance; no biosafety problem. In the application of gene silencing, the depending of the postion of target sequence has been found. There is no systematically study in the RNA mediated virus resistance. In our studies, we designed hairpin RNA constructs and artificial miRNA consructs based on different regions of the CP gene of PVY and generated transgenic plants. We aim at disserting if these kinds of tansgene construction could make transgenic plant exhibit different resistant ratio. The result will contribute to the selection of the target sequence and successfully apply the strategy of RMVR, which has an important theory and application value. The main results and conclusions presented in this thesis are as follows:
(1) Different origins of PVY CP gene influence the resistance of hairpin expressing plants against PVY
The PVY CP gene was divided into 16 regions. For every region, two PCR fragments were amplified from the whole length of the CP gene. Two PCR fragments were inverted inserted into binary vector pROKⅡ. The ligated products were transferred into E.coli DH5αby heat shock. The plant expression vectors obtained were dubbed pROK CPs (pROK CP9~pROK CP16).
The 16 hpRNA constructs (the other 8 hpRNA expression vectors were constructed by Wu Bin) were transferred into N. benthamiana through Agrobacterium tumfaciens mediated transient infection. Northern blot analysis revealed siRNAs were detected in all pROK CPs in?ltrated leaves in transient assay. And all 16 biologically active siRNAs that could effectively down regulate the expression of target mRNA.
The 16 hpRNA constructs were introduced into NC89 mediated by LBA4404. T1 50, 62, 56, 55, 45, 63, 72, 55 transgenic lines exhibiting kanamycin resistance and positive results in PCR tests for each vector were regenerated after T0 selfing. Virus resistance assay revealed that the 16 transgenic tobacco groups exhibited varying degrees of virus resistance in preventing PVY infection. All the transgenic plants of pROK CP1, pROK CP2, pROK CP3, and pROK CP4 are susceptive to the PVY infection; The resistance ratios of pROK CP5~ pROK CP16 are 27.15%, 6.27%, 67.65%, 70.83%, 66.01%, 61.34%, 66.04%, 23.59%, 11.11%, 55.56%, 77.78% and 67.25%. We demonstrate that 50bp length hpRNA construct is effective for RMVR; however, the hairpin constructs harboring different cDNA regions of the CP gene engendered different silencing efficiencies, the hpRNA targeting the 3′end (nt701~nt750) region exhibited the most highly virus resistance. The plants which harboring the hpRNAi constructs targeting 3′of PVY CP can induced higher virus resistance than those targeting 5′of PVY CP.
Northern blot analysis revealed that the trangene have been expressed in level of RNA, and the transcripts accumulation level of resistant plants was lower than that of the susceptible transgenic plants. There was an inverse correlation between the resistance and the amount of RNA accumulation in the transgenic plants. The results proved this resistance was RNA mediated. siRNAs were present in all selected resistant and susceptible transgenic plants; No obvious correlation was observed between the expression level of sequence speciflc siRNAs and virus resistance.
Virus assary in the T2 generation showed that most of the progeny transgenic plants of resistant plant still exhibited high resistance. The results indicated that hpRNA mediated virus resistance can be stably inherited in T2 generation.
(2) Different origins of PVY CP gene influence the resistance of artificial miRNA expressing plants against PVY We designed eight amiRNAs M1 (nt96~nt115), M2 (nt140~nt159), M3 (nt322~nt341), M4 (nt380~nt399), M5 (nt475~nt494), M6 (nt567~nt586), M7 (nt679~nt698) and M8 (nt735~nt754) derived from PVY Coat Protein RNA sequences. Arabidopsis thaliana miR319a precursor was used as backbone. Through oligonucleotide directed mutagenesis, the natural miR319 and miR319* sequences were replaced with synthetic sequences, each corresponding to a designed amiRNA that targets one region of the PVY CP gene. The generated fragment was digested and inserted into plant binary expression vector pROKⅡto generate pROK amiRcps.
The 8 amiRNA constructs were transferred into N. benthamiana. In transient assay, Northern blot analysis revealed amiRNAs were detected in all pROK amiRcps in?ltrated leaves. All plants with amiRNA expression exhibited a significantly decreased CP accumulation
The eight amiRcp expression constructs were introduced into NC89 mediated by LBA4404. After the kanamycin resistance and PCR test, T1 plants 98, 96, 114, 116, 117, 111, 110, 116 were screened. The resistance ratios of amiRcps expressing plants were 37.68%, 50.29%, 53.76%, 37.75%, 29.86%, 17.05%, 26.27%, 64.69%. Virus resistance assay revealed that expression of amiRNA can effective in preventing PVY infection; but not all amiRcps targeting viral CP sequence are equally effective. The amiRcp 8 targeting the 3′end (nt735~nt754) region exhibited high virus resistance.
Northern blots showed that the trangene have been expressed in level of RNA. There was an inverse correlation between the resistance and the amount of RNA accumulation in the transgenic plants. The results proved this resistance was RNA mediated. AmiRNAs were present in all selected resistant transgenic plants and there is a positive correlation between virus resistance and expression level of amiRNA.
Virus resistance assay in the T2 generation revealed that most of the progeny transgenic plants of resistant plant still exhibited high resistance. It indicated that amiRNA mediated virus resistance can be stably inherited in T2 generation.
引文
白庆荣,朱俊华,刘晓玲,朱常香,宋云枝,温孚江。利用RNA介导的抗病性获得抗2种病毒的转基因烟草。植物病理学报,2005,35:148~154
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