H1N1和H3N2亚型猪流感病毒的拯救及其致病基因的分析
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摘要
猪流感是由猪流感病毒引起的一种急性呼吸道传染病,该病以传染性强、发病率高和死亡率低等为特点。猪流感病毒为A型流感病毒,该病毒的基因组为分节段的单股负链RNA。这一特点使病毒能通过基因突变和基因重配不断进化。NSl和PB1-F2蛋白是流感病毒编码的两个重要的非结构蛋白。本研究开展了H1和H3亚型猪流感病毒流行情况的血清学调查,并分析了NSl和PBl-F2蛋白的突变及基因重配对古典猪流感病毒生物学特性的影响。
1.待宰肉猪H1和H3亚型猪流感病毒抗体血清学检测
为了了解待宰肉猪中H1和H3亚型猪流感病毒的流行情况,采用血凝抑制试验对采集于南京市某3个肉联厂的483份血清样品进行H1和H3亚型猪流感病毒的抗体检测。实验结果表明,H1亚型抗体阳性的血清样品为32份,阳性率为6.6%；H3亚型抗体阳性的血清样品为53份,阳性率为10.9%；H1和H3亚型均为阳性的血清样品2份,阳性率为0.4%。实验结果说明被调查的待宰肉猪受到H1或H3亚型猪流感病毒不同程度的感染。
2.猪流感病毒的拯救及体外生物学特性的分析
为了建立猪流感病毒拯救的反向遗传操作系统,用RT-PCR方法分别扩增出猪流感病毒A/Swine/Guangdong/1/2004(GD/04)和A/Swine/Shanghai/1/2005(SH/05)的8个基因片段,通过与双向转录表达载体pBD连接,构建成含流感病毒8个基因片段的转录表达重组质粒。将含流感病毒8个基因片段的重组质粒共转染293T细胞,拯救出GD/04和SH/05病毒。拯救出的病毒分别命名为rGD/04和rSH/05。对rGD/04和rSH/05病毒的生长曲线、病毒蚀斑形态、病毒NP蛋白在细胞内的分布和全基因组测序等进行分析后证实拯救病毒的体外生物学特性和基因组均与亲本病毒保持一致。实验结果说明成功拯救出GD/04和SH/05病毒。GD/04和SH/05病毒反向遗传操作系统的成功建立为后续的研究奠定了基础。
3.古典猪流感NS1蛋白突变病毒的构建及对小鼠的致病性分析
序列分析表明,古典猪H1N1流感病毒的NS1蛋白在过去80年问不断进化,最显著的特征为该蛋白C-端最末尾4个氨基酸残基序列的变化和该蛋白C-端末尾11个氨基酸残基的缺失。然而,发生在NSl蛋白C-端末尾的这些变化对古典猪H1N1流感病毒的毒力是否有影响至今仍不清楚。以反向遗传学技术为基础,我们以古典猪H1N1病毒A/Swine/Shanghai/1/2005为框架构建出3株NS1蛋白突变病毒(RSEV.GSEI和EPEV)和1株野生型病毒(PEQK),并以小鼠为动物模型分析这些病毒的毒力。实验结果表明,RSEV和PEQK病毒都不能在小鼠肺脏中复制；而GSEI和EPEV病毒却不需要在小鼠体内适应就能在小鼠肺脏中复制。攻毒后第4天,GSEI和EPEV病毒组小鼠肺脏中的病毒滴度分别为2.3×103pfu/g和0.7×103pfu/g.对攻毒后第6天的GSEI和EPEV病毒组小鼠的肺脏进行病理分析后发现这两株病毒均能在小鼠肺脏中引起轻度的病毒性肺炎(肺泡壁增厚)。上述实验结果说明,古典猪H1N1流感病毒NS1蛋白C-端最末尾4个氨基酸残基序列GSEI和EPEV与该病毒在小鼠上表现出的毒力相关且能使该病毒跨越种间屏障从猪传播给小鼠。
4.古典猪流感PB1-F2蛋白突变病毒的构建及对小鼠的致病性分析
PB1-F2蛋白是流感病毒PB1基因编码的一个小分子量非结构蛋白。研究表明,该蛋白能增强高致病性流感病毒和实验室小鼠适应流感病毒在小鼠模型上的毒力。但PB1-F2蛋白是否也能增强低致病性流感病毒在小鼠模型上的毒力还不清楚。通过反向遗传学技术,我们以低致病性的古典猪H1N1流感病毒(PEQK)为框架,构建出一株表达PBl-F2蛋白的PEQK-F2病毒。我们先前的实验表明,PEQK病毒不能在小鼠肺脏中复制,因此我们又以能在小鼠肺脏中复制的GSEI病毒(PEQK病毒的NS1蛋白突变病毒)为框架,构建出一株表达PB1-F2蛋白的GSEI-F2病毒。实验结果表明,PEQK-F2病毒不能在小鼠肺脏中复制,GSEI-F2病毒在小鼠肺脏中的清除速度比GSEI病毒的快(接毒后第6天仍能在小鼠肺脏中分离到GSEI病毒,而GSEI-F2病毒则在接毒后第4天就分离不到)。上述结果说明,PB1-F2蛋白不能促使PEQK-F2病毒在小鼠肺脏中复制,但PB1-F2蛋白能加快病毒从肺脏中清除。
5.古典猪流感基因重配病毒的构建及对豚鼠的致病性分析
本实验室于2004年和2005年分别分离到两株猪流感病毒A/Swine/Gongdong/1/2004(GD/04)和A/Swine/Shanghai/1/2005(SH/05).实验表明,GD/04病毒能在豚鼠肺脏中复制,而SH/05病毒却不能。为了分析与GD/04病毒感染豚鼠有关的基因,本实验以反向遗传学技术为基础,以SH/05病毒为框架,构建出5株含GD/04病毒基因的SH-PB2(GD)、SH-NS(GD)、SH-HA,NA(GD). SH-PB2,HA,NA(GD)和SH-HA,NA,NS(GD)基因重配病毒,并分析这些基因重配病毒对豚鼠的致病性。攻毒后第3天在豚鼠的气管和肺脏中均未检测到这5株基因重配病毒。血凝抑制试验发现,SH-PB2(GD)病毒能刺激豚鼠产生针对该病毒的中和抗体。这一结果说明,GD/04病毒的PB2、HA、NA、NS基因与该毒株是否能感染豚鼠并不直接相关,进一步说明流感病毒毒力相关基因的复杂性。
Swine influenza is a acute respiratory infectious diseases caused by influenza A virus, which is characterized by highly infectivity, high morbility, and low mortality. The genome of influenza A virus is segmented negative single-stranded RNA, which enables the dynamic evolution of virus by both antigenic drift and genetic shift. Both of the NS1and PB1-F2proteins have multiple accessory functions during viral infection. In the present study, serologic surveillance of swine H1and H3subtypes influenza A virus infections in finishing pigs was carried out, and the effects of mutations in NS1and PB1-F2proteins and gene reassortment on classical swine influenza A virus were also analyzed.
1. Detection of antibodies against H1and H3subtypes of swine influenza A viruses in finishing pigs
In order to investigate the prevalent state of swine influenza virus of H1and H3subtypes in finishing pigs.483swine blood samples were collected and investigated serologically using trace hemagglutination inhibition test. Thirty-two samples were characterized positive for H1subtype, of which the positive rate was6.6%. Fifty-three samples were characterized positive for H3subtype, of which the positive rate was10.9%. Two samples were characterized positive for H1and H3subtypes, of which the positive rate was0.4%. The results showed that the finshing pigs were infected in different degrees with swine influenza virus of H1or H3subtype.
2. Rescue of the swine influenza A viruses and characterization of the rescued viruses in vitro
Full-length cDNAs of swine influenza viruses (A/Swine/Guangdong/1/2004(GD/04) and A/Swine/Shanghai/1/2005(SH/05)) were amplified and cloned into the bidirectional pBD vector. The eight plasmids were co-transfected into293T cells to generate rGD/04and rSH/05viruses. The rescued viruses and the parental viruses shared similar biological properties:growth kinetics, plaque phenotype, and distribution of the NP protein in the virus-infected cells. Moreover,The gene sequence were identical between the rescued viruses and parental viruses. The successful rescue of the GD/04and SH/05viruse established a foundation for the further studies.
3. Recombination of NS1protein mutant classical swine influenza A viruses and determination of the pathogenicity of the viruses in mice
The NS1protein of classical swine H1N1influenza A virus evolved dynamically during the past80years, most notably changes happened in the four C-terminal sequences and the C-terminal truncation of11amino acids. However, the role of these changes on the virulence of classical swine H1N1influenza A virus remains unknown. Using reverse genetics, three NS1mutant viruses (RSEV, GSEI, and EPEV) and a wild-type virus (PEQK) were generated from A/Swine/Shanghai/1/2005virus and the pathogenicity of the viruses was determined in mice. The results showed that RSEV and PEQK viruses could not replicate in the lungs of mice. By contrast, GSEI and EPEV viruses could replicate in the lungs of mice without prior adaptation. The viral titers in lungs from GSEI and EPEV virus-infected mice were2.3×103pfu/g and0.7×101pfu/g at four day postinfection, respectively. Mild-to-moderate alveolitis was observed in the histopathological test of lungs from GSEI and EPEV virus-infected mice. The results indicated that C-terminal GSEI and EPEV motifs of NS1protein involved in viral virulence and facilitated the A/Swine/Shanghai/1/2005virus crossing the species barrier from swine to mice.
4. Recombination of PB1-F2protein mutant classical swine influenza A viruses and determination of the pathogenicity of the viruses in mice
The PB1-F2is a small protein encoded by an alternative reading frame of the PB1gene. Expression of the full-length PB1-F2protein has been showed to increase the pathogenicity of the highly pathogenic influenza A viruses and laboratory mouse-adapted influenza A viruses in mouse model. However, the effects of the full-length PB1-F2protein on the low pathogenic influenza A virus still remains unknown. Using reverse genetics, a PEQK-F2virus that expressed a90amino acids PB1-F2protein was generated from a low pathogenic influenza virus A/Swine/Shanghai/1/2005(PEQK). Our previous study showed that the PEQK virus did not replicate in the lungs of mice. Thus, a GSEI-F2virus was generated in the background of GSEI virus (NS1protein mutant virus of PEQK virus, which can replicate in the lungs of mice). The pathogenicity of the viruses was determined in mice. The results showed that the PB1-F2protein could not facilitate the replication of the PEQK-F2virus in the lung of mouse, and that the expression of PB1-F2protein accelerated the clearance of GSEI-F2virus from the lungs of mice. These findings suggested that while the PB1-F2protein did not facilitate the PEQK-F2virus crossing species barriers, it attenuates the virulence of GSEI-F2virus in the mice.
5. Recombination of gene reassortment classical swine influenza A viruses and determination of the pathogenicity of the viruses in guinea pigs
Two swine influenza A viruses, A/Swine/Gongdong/1/2004(GD/04) and A/Swine/Shanghai/1/2005(SH/05), were isolated in2004and2005, respectively. It was showed that the GD/04virus could replicate in the respiratory tract of guinea pigs without prior adaption, while the SH/05viurs could not. To analyze the gene(s) contribute the replication of the GD/04virus in the respiratory tract of guinea pigs. In the present study, five gene reassortment viruses (SH-PB2(GD)、SH-NS(GD)、SH-HA,NA(GD)、 SH-PB2,HA,NA(GD) and SH-HA,NA,NS(GD)) were generated using reverse genetics in the background of SH/05, and the pathogenicity of the viruses were determined in guinea pigs. All of the viruses did not be detected in both the upper and lower respiratory tract of guinea pigs. However, the SH-PB2(GD) virus induced antibody against the virus in the guinea pigs, which was detected by heamgglutinin inhibition. The results indicated that the PB2, HA, NA, and NS genes did not contribute to the replication of the GD/04virus in the guinea pigs, and that the pathogenicity of the influenza A virus is a polygenic trait.
1.待宰肉猪H1和H3亚型猪流感病毒抗体血清学检测
为了了解待宰肉猪中H1和H3亚型猪流感病毒的流行情况,采用血凝抑制试验对采集于南京市某3个肉联厂的483份血清样品进行H1和H3亚型猪流感病毒的抗体检测。实验结果表明,H1亚型抗体阳性的血清样品为32份,阳性率为6.6%；H3亚型抗体阳性的血清样品为53份,阳性率为10.9%；H1和H3亚型均为阳性的血清样品2份,阳性率为0.4%。实验结果说明被调查的待宰肉猪受到H1或H3亚型猪流感病毒不同程度的感染。
2.猪流感病毒的拯救及体外生物学特性的分析
为了建立猪流感病毒拯救的反向遗传操作系统,用RT-PCR方法分别扩增出猪流感病毒A/Swine/Guangdong/1/2004(GD/04)和A/Swine/Shanghai/1/2005(SH/05)的8个基因片段,通过与双向转录表达载体pBD连接,构建成含流感病毒8个基因片段的转录表达重组质粒。将含流感病毒8个基因片段的重组质粒共转染293T细胞,拯救出GD/04和SH/05病毒。拯救出的病毒分别命名为rGD/04和rSH/05。对rGD/04和rSH/05病毒的生长曲线、病毒蚀斑形态、病毒NP蛋白在细胞内的分布和全基因组测序等进行分析后证实拯救病毒的体外生物学特性和基因组均与亲本病毒保持一致。实验结果说明成功拯救出GD/04和SH/05病毒。GD/04和SH/05病毒反向遗传操作系统的成功建立为后续的研究奠定了基础。
3.古典猪流感NS1蛋白突变病毒的构建及对小鼠的致病性分析
序列分析表明,古典猪H1N1流感病毒的NS1蛋白在过去80年问不断进化,最显著的特征为该蛋白C-端最末尾4个氨基酸残基序列的变化和该蛋白C-端末尾11个氨基酸残基的缺失。然而,发生在NSl蛋白C-端末尾的这些变化对古典猪H1N1流感病毒的毒力是否有影响至今仍不清楚。以反向遗传学技术为基础,我们以古典猪H1N1病毒A/Swine/Shanghai/1/2005为框架构建出3株NS1蛋白突变病毒(RSEV.GSEI和EPEV)和1株野生型病毒(PEQK),并以小鼠为动物模型分析这些病毒的毒力。实验结果表明,RSEV和PEQK病毒都不能在小鼠肺脏中复制；而GSEI和EPEV病毒却不需要在小鼠体内适应就能在小鼠肺脏中复制。攻毒后第4天,GSEI和EPEV病毒组小鼠肺脏中的病毒滴度分别为2.3×103pfu/g和0.7×103pfu/g.对攻毒后第6天的GSEI和EPEV病毒组小鼠的肺脏进行病理分析后发现这两株病毒均能在小鼠肺脏中引起轻度的病毒性肺炎(肺泡壁增厚)。上述实验结果说明,古典猪H1N1流感病毒NS1蛋白C-端最末尾4个氨基酸残基序列GSEI和EPEV与该病毒在小鼠上表现出的毒力相关且能使该病毒跨越种间屏障从猪传播给小鼠。
4.古典猪流感PB1-F2蛋白突变病毒的构建及对小鼠的致病性分析
PB1-F2蛋白是流感病毒PB1基因编码的一个小分子量非结构蛋白。研究表明,该蛋白能增强高致病性流感病毒和实验室小鼠适应流感病毒在小鼠模型上的毒力。但PB1-F2蛋白是否也能增强低致病性流感病毒在小鼠模型上的毒力还不清楚。通过反向遗传学技术,我们以低致病性的古典猪H1N1流感病毒(PEQK)为框架,构建出一株表达PBl-F2蛋白的PEQK-F2病毒。我们先前的实验表明,PEQK病毒不能在小鼠肺脏中复制,因此我们又以能在小鼠肺脏中复制的GSEI病毒(PEQK病毒的NS1蛋白突变病毒)为框架,构建出一株表达PB1-F2蛋白的GSEI-F2病毒。实验结果表明,PEQK-F2病毒不能在小鼠肺脏中复制,GSEI-F2病毒在小鼠肺脏中的清除速度比GSEI病毒的快(接毒后第6天仍能在小鼠肺脏中分离到GSEI病毒,而GSEI-F2病毒则在接毒后第4天就分离不到)。上述结果说明,PB1-F2蛋白不能促使PEQK-F2病毒在小鼠肺脏中复制,但PB1-F2蛋白能加快病毒从肺脏中清除。
5.古典猪流感基因重配病毒的构建及对豚鼠的致病性分析
本实验室于2004年和2005年分别分离到两株猪流感病毒A/Swine/Gongdong/1/2004(GD/04)和A/Swine/Shanghai/1/2005(SH/05).实验表明,GD/04病毒能在豚鼠肺脏中复制,而SH/05病毒却不能。为了分析与GD/04病毒感染豚鼠有关的基因,本实验以反向遗传学技术为基础,以SH/05病毒为框架,构建出5株含GD/04病毒基因的SH-PB2(GD)、SH-NS(GD)、SH-HA,NA(GD). SH-PB2,HA,NA(GD)和SH-HA,NA,NS(GD)基因重配病毒,并分析这些基因重配病毒对豚鼠的致病性。攻毒后第3天在豚鼠的气管和肺脏中均未检测到这5株基因重配病毒。血凝抑制试验发现,SH-PB2(GD)病毒能刺激豚鼠产生针对该病毒的中和抗体。这一结果说明,GD/04病毒的PB2、HA、NA、NS基因与该毒株是否能感染豚鼠并不直接相关,进一步说明流感病毒毒力相关基因的复杂性。
Swine influenza is a acute respiratory infectious diseases caused by influenza A virus, which is characterized by highly infectivity, high morbility, and low mortality. The genome of influenza A virus is segmented negative single-stranded RNA, which enables the dynamic evolution of virus by both antigenic drift and genetic shift. Both of the NS1and PB1-F2proteins have multiple accessory functions during viral infection. In the present study, serologic surveillance of swine H1and H3subtypes influenza A virus infections in finishing pigs was carried out, and the effects of mutations in NS1and PB1-F2proteins and gene reassortment on classical swine influenza A virus were also analyzed.
1. Detection of antibodies against H1and H3subtypes of swine influenza A viruses in finishing pigs
In order to investigate the prevalent state of swine influenza virus of H1and H3subtypes in finishing pigs.483swine blood samples were collected and investigated serologically using trace hemagglutination inhibition test. Thirty-two samples were characterized positive for H1subtype, of which the positive rate was6.6%. Fifty-three samples were characterized positive for H3subtype, of which the positive rate was10.9%. Two samples were characterized positive for H1and H3subtypes, of which the positive rate was0.4%. The results showed that the finshing pigs were infected in different degrees with swine influenza virus of H1or H3subtype.
2. Rescue of the swine influenza A viruses and characterization of the rescued viruses in vitro
Full-length cDNAs of swine influenza viruses (A/Swine/Guangdong/1/2004(GD/04) and A/Swine/Shanghai/1/2005(SH/05)) were amplified and cloned into the bidirectional pBD vector. The eight plasmids were co-transfected into293T cells to generate rGD/04and rSH/05viruses. The rescued viruses and the parental viruses shared similar biological properties:growth kinetics, plaque phenotype, and distribution of the NP protein in the virus-infected cells. Moreover,The gene sequence were identical between the rescued viruses and parental viruses. The successful rescue of the GD/04and SH/05viruse established a foundation for the further studies.
3. Recombination of NS1protein mutant classical swine influenza A viruses and determination of the pathogenicity of the viruses in mice
The NS1protein of classical swine H1N1influenza A virus evolved dynamically during the past80years, most notably changes happened in the four C-terminal sequences and the C-terminal truncation of11amino acids. However, the role of these changes on the virulence of classical swine H1N1influenza A virus remains unknown. Using reverse genetics, three NS1mutant viruses (RSEV, GSEI, and EPEV) and a wild-type virus (PEQK) were generated from A/Swine/Shanghai/1/2005virus and the pathogenicity of the viruses was determined in mice. The results showed that RSEV and PEQK viruses could not replicate in the lungs of mice. By contrast, GSEI and EPEV viruses could replicate in the lungs of mice without prior adaptation. The viral titers in lungs from GSEI and EPEV virus-infected mice were2.3×103pfu/g and0.7×101pfu/g at four day postinfection, respectively. Mild-to-moderate alveolitis was observed in the histopathological test of lungs from GSEI and EPEV virus-infected mice. The results indicated that C-terminal GSEI and EPEV motifs of NS1protein involved in viral virulence and facilitated the A/Swine/Shanghai/1/2005virus crossing the species barrier from swine to mice.
4. Recombination of PB1-F2protein mutant classical swine influenza A viruses and determination of the pathogenicity of the viruses in mice
The PB1-F2is a small protein encoded by an alternative reading frame of the PB1gene. Expression of the full-length PB1-F2protein has been showed to increase the pathogenicity of the highly pathogenic influenza A viruses and laboratory mouse-adapted influenza A viruses in mouse model. However, the effects of the full-length PB1-F2protein on the low pathogenic influenza A virus still remains unknown. Using reverse genetics, a PEQK-F2virus that expressed a90amino acids PB1-F2protein was generated from a low pathogenic influenza virus A/Swine/Shanghai/1/2005(PEQK). Our previous study showed that the PEQK virus did not replicate in the lungs of mice. Thus, a GSEI-F2virus was generated in the background of GSEI virus (NS1protein mutant virus of PEQK virus, which can replicate in the lungs of mice). The pathogenicity of the viruses was determined in mice. The results showed that the PB1-F2protein could not facilitate the replication of the PEQK-F2virus in the lung of mouse, and that the expression of PB1-F2protein accelerated the clearance of GSEI-F2virus from the lungs of mice. These findings suggested that while the PB1-F2protein did not facilitate the PEQK-F2virus crossing species barriers, it attenuates the virulence of GSEI-F2virus in the mice.
5. Recombination of gene reassortment classical swine influenza A viruses and determination of the pathogenicity of the viruses in guinea pigs
Two swine influenza A viruses, A/Swine/Gongdong/1/2004(GD/04) and A/Swine/Shanghai/1/2005(SH/05), were isolated in2004and2005, respectively. It was showed that the GD/04virus could replicate in the respiratory tract of guinea pigs without prior adaption, while the SH/05viurs could not. To analyze the gene(s) contribute the replication of the GD/04virus in the respiratory tract of guinea pigs. In the present study, five gene reassortment viruses (SH-PB2(GD)、SH-NS(GD)、SH-HA,NA(GD)、 SH-PB2,HA,NA(GD) and SH-HA,NA,NS(GD)) were generated using reverse genetics in the background of SH/05, and the pathogenicity of the viruses were determined in guinea pigs. All of the viruses did not be detected in both the upper and lower respiratory tract of guinea pigs. However, the SH-PB2(GD) virus induced antibody against the virus in the guinea pigs, which was detected by heamgglutinin inhibition. The results indicated that the PB2, HA, NA, and NS genes did not contribute to the replication of the GD/04virus in the guinea pigs, and that the pathogenicity of the influenza A virus is a polygenic trait.
引文
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