家蚕核型多角体病毒（BmNPV）Bm61、orf74的基因分析
摘要
杆状病毒是节肢动物特别是鳞翅目昆虫重要的病原微生物。到目前为止,已有47种杆状病毒完成了全基因组测序。序列分析发现,仅有30个基因在所有已经测序的杆状病毒中都存在,称为核心基因。在鳞翅目昆虫杆状病毒中,有62个基因是保守的。
本研究以家蚕核型多角体病毒(Bombyx mori nucleopolyhedrovirus,BmNPV)核心基因Bm61和非核心基因orf74为研究对象,从基因的转录、表达、亚细胞定位、病毒结构定位、基因缺失等方面,研究基因的基本特性及其功能,从而为丰富杆状病毒分子生物学提供理论基础。论文主要结论如下:
1.BmNPV的orf61(Bm61)位于基因组61,188—61,892 nt,全长399bp,编码一个含133个氨基酸残基的蛋白,预测分子量约为15.5 kDa,Bm61是AcMNPV orf75的同源序列。在Bm61基因ATG上游存在杆状病毒晚期转录基序ATAAG,在终止密码子TAA下游发现转录终止信号AATAAA。
2.RT-PCR分析发现,Bm61在病毒感染宿主细胞12h开始转录,直到72h。利用大肠杆菌表达系统表达了BM61融合蛋白并制备了多克隆抗体。利用抗体进行Western blot分析,发现Bm61的表达产物在病毒感染宿主细胞后12 h被检测出来,一直持续72 h都有表达,大小为15.5kDa,与预测的大小一致,说明Bm61是一个晚期表达基因,而且没有转录后修饰。利用抗体检测发现BM61定位在细胞核膜和细胞核内周。
3.利用phageλRed重组酶和Bac-to-Bac系统,在家蚕核型多角体病毒Bacmid中成功敲除Bm61基因。缺失Bm61的Bacmid转染细胞后,只有少量的细胞荧光,说明缺失Bm61导致病毒在细胞间的传染出现异常。PCR显示转染细胞的上清中没有BV产生,而Bm61的拯救病毒表现出和野生病毒相同的特性。证明Bm61是BV产生的必需基因。进一步分析Bm61缺失不影响病毒基因组复制和gp64的转录。
4.orf74位于BmNPV基因组的69,987-70,449 nt之间,全长462 bp,编码154个氨基酸残基,预测分子量大约为17.3 kDa。在orf74基因ATG上游9nt处有一个杆状病毒晚期转录基序TTAAG。orf74是AcMNPVorf91的同源序列,
5.orf74的转录分析表明,orf74从病毒感染后12h开始转录,一直持续到96h,可能是一个晚期表达基因。利用ORF74与GFP融合对其亚细胞定位进行了观察,发现ORF74的表达集中在细胞核。
6.敲除orf74基因后构建了缺失orf74的病毒vBm-ko。vBm-ko在BmN上的复制结果表明:orf74的敲除不影响病毒在BmN细胞上增殖和基因组复制。幼虫分析表明orf74的敲除可以延长病毒杀死幼虫的时间,但是病毒的产量没有明显差异。这些结果表明orf74可能是病毒复制的非必须基因,可能与病毒的毒力有关。
The family Baculoviridae is a highiy selective pathogen in arhropods, mainly in insects of the order Lepidoptera.So far,the genomes of 47 baculoviruses have been sequeneced.And it is revealed that 30 genes are baculovirus core genes and 62 genes are conserved in Lepidoptera baculoviruses after analyzing the whole genomes.
Bombyx mori nueleopolyhedrovirus is one of most studied baeuloviruses;some genes have not been characterized.In this study,a non-conserved genes(orf74) and a conserved genes(Bm61) in Lepidoptera baculoviruses are characterized.These two genes were characterized from the transcription,expression,cellular location and knockout.The object of this study is to enhance our understanding of baculovirus molecular biology.The results are as following:
1.orf61(Bm61) is located at nt 61,188—61,892 of BmNPV genome, containing 399 bp and coding 133 aa with a predicted molecular weight of 15.5 kDa.Bm61 is the homology sequence of AcMNPV的orf75.The late transcription motif of baculoviridae,ATAAG was located at the upstream of ATG.And the transcription terminal signal,AATAAA was found at the downstream of the terminal codon TAA.
2.The RT-PCR results showed that Bm61 genes was transcribed in the infected cells from 12h to 72h.Bm61 fusion protein was expressed using the E.coli expression system and polyclonal antibodies were harvested. The western blot analysis showed that Bm61 was detected in the infected cells from 12 to 72 h,which further suggesting that Bm61 is a late gene. The detected Bm61 had a molecular weight of 15.5 kDa,same as the predicted one,which suggesting that Bm61 have no post-transcription modification.The sub-cellular localization of the Bm61 proteins was investigated by immunofluorescence.It was found that the proteins were located on the nuclear membrane and the nuclear ring zone.
3.Bm61 deletion Bacmid was generated by Red-recombination in E. coli DH10B.A fragment of Bm61 gene was knocked out from the BmNPV Bacmid and the Cm gene was inserted into the same site.The gfp and polh genes were transposed into the recombinant Bacmid which was used to transfect BmN cells.Cells transfected with Bm61-knockout Bacmid showed a few fluorescent spots,which indicated that there was no spread of the virus beyond the cells transfected with Bm61 deletion Bacmid.The results of PCR showed that there was no BV in the supernatant of transfected cells,which indicated that it reduced the production of BV induced of the deletion of Bm61.However,Bm61-recovered Bacmid showed the same feature of wild type Bacmid.The deletion of Bm61 did not affect the replication of the virus genome and the transcription of gp64.Therefore,it was concluded that the Bm61 gene was an essential gene for BV production.
4.orf74 is located at nt 69,987-70,449 of BmNPV genome(nt), contains 462 bp and codes 154 aa,with a predicted molecular weight of 17.3 kDa.The late transcription motif of baculoviridae,TTAAG was found at 9 nts.orf74 is the homology sequence of AcMNPVorf91.
5.The results of RT-PCR showed thal orf74 genes was transcribed in the infected cells from 12h to 96h,which indicated that orf74 was probably a late gene.ORF74 fused with GFP was used to investigate its subcellular location.ORF74 was found to be located in the nucleus.
6.orf74 deletion Bacmid,vBm-ko was generated by Red-recombination in E.coli DH10B.The analysis of orf74-knockout virus replication in BmN cells showed that the deletion of orf74 did not affect both the reproduction and the replication of the virus genome in BmN cells. The bioassay indicated that the deletion of off74 could extend the survival time of the infected larvae,but the yield of virus had no changes.These results showed that orf74 was a non-essential gene for virus replication,but related to the virulence.
本研究以家蚕核型多角体病毒(Bombyx mori nucleopolyhedrovirus,BmNPV)核心基因Bm61和非核心基因orf74为研究对象,从基因的转录、表达、亚细胞定位、病毒结构定位、基因缺失等方面,研究基因的基本特性及其功能,从而为丰富杆状病毒分子生物学提供理论基础。论文主要结论如下:
1.BmNPV的orf61(Bm61)位于基因组61,188—61,892 nt,全长399bp,编码一个含133个氨基酸残基的蛋白,预测分子量约为15.5 kDa,Bm61是AcMNPV orf75的同源序列。在Bm61基因ATG上游存在杆状病毒晚期转录基序ATAAG,在终止密码子TAA下游发现转录终止信号AATAAA。
2.RT-PCR分析发现,Bm61在病毒感染宿主细胞12h开始转录,直到72h。利用大肠杆菌表达系统表达了BM61融合蛋白并制备了多克隆抗体。利用抗体进行Western blot分析,发现Bm61的表达产物在病毒感染宿主细胞后12 h被检测出来,一直持续72 h都有表达,大小为15.5kDa,与预测的大小一致,说明Bm61是一个晚期表达基因,而且没有转录后修饰。利用抗体检测发现BM61定位在细胞核膜和细胞核内周。
3.利用phageλRed重组酶和Bac-to-Bac系统,在家蚕核型多角体病毒Bacmid中成功敲除Bm61基因。缺失Bm61的Bacmid转染细胞后,只有少量的细胞荧光,说明缺失Bm61导致病毒在细胞间的传染出现异常。PCR显示转染细胞的上清中没有BV产生,而Bm61的拯救病毒表现出和野生病毒相同的特性。证明Bm61是BV产生的必需基因。进一步分析Bm61缺失不影响病毒基因组复制和gp64的转录。
4.orf74位于BmNPV基因组的69,987-70,449 nt之间,全长462 bp,编码154个氨基酸残基,预测分子量大约为17.3 kDa。在orf74基因ATG上游9nt处有一个杆状病毒晚期转录基序TTAAG。orf74是AcMNPVorf91的同源序列,
5.orf74的转录分析表明,orf74从病毒感染后12h开始转录,一直持续到96h,可能是一个晚期表达基因。利用ORF74与GFP融合对其亚细胞定位进行了观察,发现ORF74的表达集中在细胞核。
6.敲除orf74基因后构建了缺失orf74的病毒vBm-ko。vBm-ko在BmN上的复制结果表明:orf74的敲除不影响病毒在BmN细胞上增殖和基因组复制。幼虫分析表明orf74的敲除可以延长病毒杀死幼虫的时间,但是病毒的产量没有明显差异。这些结果表明orf74可能是病毒复制的非必须基因,可能与病毒的毒力有关。
The family Baculoviridae is a highiy selective pathogen in arhropods, mainly in insects of the order Lepidoptera.So far,the genomes of 47 baculoviruses have been sequeneced.And it is revealed that 30 genes are baculovirus core genes and 62 genes are conserved in Lepidoptera baculoviruses after analyzing the whole genomes.
Bombyx mori nueleopolyhedrovirus is one of most studied baeuloviruses;some genes have not been characterized.In this study,a non-conserved genes(orf74) and a conserved genes(Bm61) in Lepidoptera baculoviruses are characterized.These two genes were characterized from the transcription,expression,cellular location and knockout.The object of this study is to enhance our understanding of baculovirus molecular biology.The results are as following:
1.orf61(Bm61) is located at nt 61,188—61,892 of BmNPV genome, containing 399 bp and coding 133 aa with a predicted molecular weight of 15.5 kDa.Bm61 is the homology sequence of AcMNPV的orf75.The late transcription motif of baculoviridae,ATAAG was located at the upstream of ATG.And the transcription terminal signal,AATAAA was found at the downstream of the terminal codon TAA.
2.The RT-PCR results showed that Bm61 genes was transcribed in the infected cells from 12h to 72h.Bm61 fusion protein was expressed using the E.coli expression system and polyclonal antibodies were harvested. The western blot analysis showed that Bm61 was detected in the infected cells from 12 to 72 h,which further suggesting that Bm61 is a late gene. The detected Bm61 had a molecular weight of 15.5 kDa,same as the predicted one,which suggesting that Bm61 have no post-transcription modification.The sub-cellular localization of the Bm61 proteins was investigated by immunofluorescence.It was found that the proteins were located on the nuclear membrane and the nuclear ring zone.
3.Bm61 deletion Bacmid was generated by Red-recombination in E. coli DH10B.A fragment of Bm61 gene was knocked out from the BmNPV Bacmid and the Cm gene was inserted into the same site.The gfp and polh genes were transposed into the recombinant Bacmid which was used to transfect BmN cells.Cells transfected with Bm61-knockout Bacmid showed a few fluorescent spots,which indicated that there was no spread of the virus beyond the cells transfected with Bm61 deletion Bacmid.The results of PCR showed that there was no BV in the supernatant of transfected cells,which indicated that it reduced the production of BV induced of the deletion of Bm61.However,Bm61-recovered Bacmid showed the same feature of wild type Bacmid.The deletion of Bm61 did not affect the replication of the virus genome and the transcription of gp64.Therefore,it was concluded that the Bm61 gene was an essential gene for BV production.
4.orf74 is located at nt 69,987-70,449 of BmNPV genome(nt), contains 462 bp and codes 154 aa,with a predicted molecular weight of 17.3 kDa.The late transcription motif of baculoviridae,TTAAG was found at 9 nts.orf74 is the homology sequence of AcMNPVorf91.
5.The results of RT-PCR showed thal orf74 genes was transcribed in the infected cells from 12h to 96h,which indicated that orf74 was probably a late gene.ORF74 fused with GFP was used to investigate its subcellular location.ORF74 was found to be located in the nucleus.
6.orf74 deletion Bacmid,vBm-ko was generated by Red-recombination in E.coli DH10B.The analysis of orf74-knockout virus replication in BmN cells showed that the deletion of orf74 did not affect both the reproduction and the replication of the virus genome in BmN cells. The bioassay indicated that the deletion of off74 could extend the survival time of the infected larvae,but the yield of virus had no changes.These results showed that orf74 was a non-essential gene for virus replication,but related to the virulence.
引文
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