高致病性猪繁殖与呼吸综合征病毒人工感染模型的建立及其部分生物学特性研究
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摘要
猪繁殖与呼吸综合征(PRRS)是以妊娠母猪晚期流产、早产、死产和产弱仔数量增加,以及新生仔猪与生长猪严重呼吸道疾病为主要特征。目前,该病在全球大部分地区流行,是猪主要的传染病之一。2006年5月以来,最初称为“高热综合症”的猪病在我国南方部分省区首先暴发。随后,该病在我国广泛传播,给我国养猪业造成巨大的经济损失。业已证明,引发该病的主要病原是北美洲型的高致病性猪繁殖与呼吸综合征病毒(HP-PRRSV)变异株。虽然“高热综合症”可引起各种年龄的猪发病和死亡,但HP-PRRSV分离株对不同年龄猪的毒力和体内致病性、致病特征是否与临床发病相似,疾病的致病机理、免疫应答,以及预防高致病性蓝耳病(HP-PRRS)的疫苗免疫效果等尚未清楚。弄清这些问题最终都离不开动物感染实验。
为了建立一个可靠的研究HP-PRRS的普通猪人工感染模型,本研究用含3 104TCID50的HuN4病毒液,通过滴鼻和肌注方式分别人工感染30d、60d和180d龄的普通猪。结果表明,PRRSV变异株HuN4可引起上述三种年龄猪发生典型的PRRS,与生产中猪临床发病特征相似。其中,30d、60d仔猪严重发病,发病率均为100%,死亡率分别为100%和60%。病猪出现持续高热(41.0～42.0°C)、呼吸困难等临床症状和肺脏实变、淋巴结肿大等解剖病变,以及以间质性肺炎、淋巴样组织炎症和脑炎为主的组织病理变化。而180d龄感染猪在上述各方面都比其他两个年龄组猪发病轻。结果证明,30d和60d龄的普通仔猪适合作为HP-PRRSV变异株HuN4的感染模型。
为了进一步研究PRRSV变异株代表性毒株HuN4的体内致病性和免疫应答,以及了解病毒感染剂量对疾病的影响,用已建立的感染模型,以3 103～3 105 TCID50三个感染剂量的HuN4第5代细胞毒,对60日龄的普通仔猪进行人工感染试验,评价感染仔猪临床发病特征、体液免疫应答、病毒血症和组织病毒含量,以及血清中细胞因子IFN-γ和IL-10的蛋白含量。结果表明,PRRSV变异株HuN4三个不同感染剂量的病毒都可引起感染仔猪严重的临床疾病,但最小感染剂量组猪的临床症状比其他病毒感染组相对轻一些。三个病毒感染组表现明显和一致的临床特征包括持续高热(41.0～41.9°C)、高发病率和高死亡率(60～100%)、明显的PRRS临床症状和多系统器官的严重病理学损伤;可诱导迅速和强烈的体液免疫应答,感染后(PI)7d大部分感染仔猪针对PRRSV特异抗体的血清阳转;病毒血症检测结果表明,该病毒可在猪体内迅速和大量复制,从大部分病猪感染后3d的猪血清分离到病毒,并且持续感染至少21d;血清和主要器官组织病毒定量检测结果表明,血清病毒平均滴度在感染后7d达到高峰,最大感染剂量组病毒核酸含量高达9.7logs copies/ml。组织病毒含量为4.66～9.05 logs copies/g不等。试验还发现病毒感染猪血清中细胞因子IFN-γ和IL-10浓度升高以及这两种细胞因子表达比例失调。总的实验结果表明,HP-PRRSV变异株对仔猪具有高度致病性,适合用作评价预防HP-PRRS疫苗的强毒参考株。此外,根据实验数据,我们初步制定了HP-PRRSV变异株HuN4对60d龄普通仔猪的强毒标准和发病标准。
为了建立一种可检测HP-PRRSV变异株及其细胞传代致弱株和国内经典毒株的荧光定量RT-PCR方法,研究设计针对PRRSV ORF7基因的特异性引物和TaqMan荧光探针,建立了一种检测PRRSV的快速、敏感、特异、重复性好的TaqMan荧光定量RT-PCR方法。该方法在检测101拷贝/μL～107拷贝/μL模板范围内具有良好的线性关系。比常规RT-PCR更敏感,可分别检测最少为10拷贝的阳性标准品和1TCID50病毒。应用建立的方法检测了HP-PRRSV HuN4第5代和第65代细胞毒人工感染仔猪的血清组织样品的病毒含量。结果证明该方法适合用于病毒的早期感染检测、研究病毒在体内的复制和持续性感染,以及HP-PRRSV弱毒苗的研制及其免疫效果评价等。
重组猪IFN-γ可抑制PRRSV在培养细胞的复制。而且可从感染PRRSV的猪淋巴结、肺脏和外周血单核细胞(PBMC)检测到猪IFN-γ。本研究旨在建立快速、敏感和特异的猪IFN-γmRNA TaqMan荧光定量RT-PCR检测方法,为研究猪IFN-γ等细胞因子在猪体内表达变化等提供检测技术。针对猪IFN-γ基因和管家基因cyclophilin A (CyPA)核苷酸序列分别设计特异性引物和TaqMan荧光探针,以重组质粒pMD18-T-IFN-γ和pMD18-T-CyPA为标准品,进行实时荧光定量RT-PCR检测,并构建猪IFN-γmRNA和CyPA的荧光定量RT-PCR标准曲线。结果表明,建立的方法在检测101拷贝/μL～107拷贝/μL模板范围内具有良好的线性关系,相关系数R2均高达0.999,扩增效率均高于99.0%;可检测最少为100拷贝的阳性标准品。应用建立的方法检测HP-PRRSV变异株HuN4人工感染猪后21d的肺、脾和淋巴结组织样品,发现IFN-γmRNA在上述组织表达量很低。结果证明,该方法快速、敏感性高、特异性强和重复性好,可直接应用于临床样品的检测。
Porcine reproductive and respiratory syndrome (PRRS) is characterized by late-term abortions and delivery of increase number of stillbirths or premature and weak piglets in sows and severe respiratory disease in neonatal and growing pigs. Now the disease is endemic in most areas around the world and recognized as one of most important diseases of swine. In May 2006, a severe disease, initially called‘pig high fever syndrome’initially occurred in some provinces in south of China and then spread widely in China, resulting in huge economic losses in pig industry of China. Now a highly pathogenic porcine reproductive and respiratory syndrome virus (HP-PRRSV) variant has been confirmed to be the main pathogen of the severe outbreaks. Pigs of all age were clinically sick and dead in the severe infection. However, whether the virulence and in vivo pathogenicity and clinical features of the new isolates of PRRSV to pigs of all age were similar to those observed in the field, the pathogenesis and immune response of the virus and the immune efficacy of new vaccines for protection against the highly pathogenic porcine reproductive and respiratory syndrome (HP-PRRS) are still unclear. Animal experimentally infection needs to be done to answer the questions.
To develop a reliable conventional pig model to study the HP-PRRS, three groups of 30-, 60- and 180-day-old pigs were infected with 3×104 50% tissue culture infective dose (TCID50) per pig of a representative HP-PRRSV strain, 5th-passage HuN4 (HuN4-F5) intramuscularly and intranasally. The results showed that the HP-PRRSV HuN4 could induce the typical PRRS in all the conventional pigs and the clinical features were similar to those observed in the field. Among them, 30-day-old and 60-day-old pigs exhibited severe clinical signs with 100% morbidity and 100% and 60% mortality, respectively. Persistently high fever (41.0～42.0°C), dyspnoea, gross lesions of consolidation of lungs,enlargement and/or haemorrhage of lymph nodes and histopathological lesions of interstitial pneumonia, the inflammation of the lymphoid tissue and encephalitis were observed in the two exposed groups. However, the clinical signs, gross lesions and microscopic lesions of the infected 180-day-old pigs were slight compared with other infected groups. So the results demonstrated that the 30 and 60-day-old conventional pigs are suitable to be the infection model for the HP-PRRSV HuN4 strain.
To further investigate the in vivo pathogenicity and virulence and immune responses of the newly emerging PRRSV, 3 groups of 60-day-old conventional piglets were inoculated intranasally with a representative strain of the HP-PRRSV variant, HuN4 with 3 different infection doses(3 103～3 105 TCID50). The clinical features, humoral immune responses, viremia, the viral load in different tissues and levels of IFN-γand IL-10 cytokine production in serum in this study were evaluated. The results revealed that 3 different infection doses of virus could induce severe disease in piglets and the infected pigs in the least infection dose group exhibited a little mild clinical signs than those of other infection dose groups. The significantly and consistently clinical characteristics of the PRRSV-exposed groups consisted of persistently high fever (41.0～41.9℃) and high morbidity and mortality (60～100%), the marked clinical signs of PRRS and severe histopathologic damages in multiple organs. It induced rapid and intense humoral immune responses and seroconversion was detected in most of infected pigs at 7 days post-infection (DPI).The virus rapidly and vigorously replicated in vivo. The virus was isolated from the serum of most of infected pigs at 3 DPI and the viremia was detected in all the infected pigs by 21 DPI days. The highest average titer of viremia peaked near 9.7logs copies/ml serum at 7 DPI. The viral load in different tissue reached 4.66～9.05 logs copies/g. Elevated levels of IFN-γand IL-10 cytokine production in serum and an unbalanced profile of Th1 (IFN-γ) /Th2 (IL-10) responses in this study were also observed. Taken together, our results demonstrated that the HP-PRRSV variant HuN4 strain is a highly pathogenic for piglets and suitable to be a reference strain of highly virulent PRRSV for evaluating the efficacy of the new vaccines to prevent the HP-PRRS. Additionally, according to the experiment data, we initially made a standard of high virulence of the HuN4 for the 60-day-old conventional pigs and a clinical standard of HP-PRRS.
To establish a fluorescent quantitative reverse transcription-polymerase chain reaction (RT-PCR) assay for the detection of HP-PRRSV and its attenuated virus and the classical NA-type PRRSV in China,the specific primers and fluorogenetic probe based on the ORF7 gene of PRRSV were designed and a quantitative TaqMan RT-PCR for detecting PRRSV had been developed. The established technique was time-saving, highly sensitive and specific and had reliable reproducibility. The assay for the quantitation of PRRSV template was found to be in the 101～107copies/μL linear dynamic range with excellent linearity. It is more sensitive than conventional RT-PCR and could detect the least template of 10 copies and 1 TCID50 virus. Using the technique, serum and tissue samples from pigs experimentally infected with HP-PRRSV HuN4-F5, and the sixty-fifth passage attenuated virus (HuN4-F65) were collected to detect viral load. The results demonstrated the established technique is suitable for detection PRRSV in early infection, replication and persistent infection in vivo, development and evaluation the efficacy of HP-PRRSV attenuated virus vaccines.
The recombinant porcine interferon-γ(IFN-γ) can inhibit the replication of PRRSV in vitro and the IFN-γcould be detected from the lymph node, lung and peripheral blood monocytes (PBMC) of PRRSV infected pigs. To establish a fluorescent quantitative RT-PCR assay for the detection of porcine IFN-γ, the specific primers and fluorogenetic probes based on the gene of porcine IFN-γand housekeeping gene cyclophilin A (CyPA) were designed, by using the recombinant plasmids pMD18-T-IFN-γand pMD18-T-CyPA as standard products, a real-time quantitative TaqMan reverse transcription-polymerase chain reaction (RT-PCR) was performed to construct the standard curves of porcine IFN-γand CyPA. The results indicated that the assay for the quantitation of porcine IFN-γtemplate could work well in the range of 101～107copies/μL with excellent linearity, the coefficient correlation R squared values reached 0.999 and amplification efficiency was higher than 99.0%. The sensitivity was sufficient to detect the least template of 100 copies. IFN-γmRNA in tissue samples from the spleen, lung and lymph node in the euthanatized pigs infected with the HuN4-F5 virus at 21DPI were collected and quantitatively detected by the established assay. The results showed that the expression of IFN-γmRNA in infected pigs is very low with a range of 6.44×102～1.04×104 copies/μL cDNA. Therefore, The currently established porcine IFN-γspecific real time RT-PCR was time-saving, highly sensitive and specific, and had reliable reproducibility and suitable to detect the clinical specimens.
为了建立一个可靠的研究HP-PRRS的普通猪人工感染模型,本研究用含3 104TCID50的HuN4病毒液,通过滴鼻和肌注方式分别人工感染30d、60d和180d龄的普通猪。结果表明,PRRSV变异株HuN4可引起上述三种年龄猪发生典型的PRRS,与生产中猪临床发病特征相似。其中,30d、60d仔猪严重发病,发病率均为100%,死亡率分别为100%和60%。病猪出现持续高热(41.0～42.0°C)、呼吸困难等临床症状和肺脏实变、淋巴结肿大等解剖病变,以及以间质性肺炎、淋巴样组织炎症和脑炎为主的组织病理变化。而180d龄感染猪在上述各方面都比其他两个年龄组猪发病轻。结果证明,30d和60d龄的普通仔猪适合作为HP-PRRSV变异株HuN4的感染模型。
为了进一步研究PRRSV变异株代表性毒株HuN4的体内致病性和免疫应答,以及了解病毒感染剂量对疾病的影响,用已建立的感染模型,以3 103～3 105 TCID50三个感染剂量的HuN4第5代细胞毒,对60日龄的普通仔猪进行人工感染试验,评价感染仔猪临床发病特征、体液免疫应答、病毒血症和组织病毒含量,以及血清中细胞因子IFN-γ和IL-10的蛋白含量。结果表明,PRRSV变异株HuN4三个不同感染剂量的病毒都可引起感染仔猪严重的临床疾病,但最小感染剂量组猪的临床症状比其他病毒感染组相对轻一些。三个病毒感染组表现明显和一致的临床特征包括持续高热(41.0～41.9°C)、高发病率和高死亡率(60～100%)、明显的PRRS临床症状和多系统器官的严重病理学损伤;可诱导迅速和强烈的体液免疫应答,感染后(PI)7d大部分感染仔猪针对PRRSV特异抗体的血清阳转;病毒血症检测结果表明,该病毒可在猪体内迅速和大量复制,从大部分病猪感染后3d的猪血清分离到病毒,并且持续感染至少21d;血清和主要器官组织病毒定量检测结果表明,血清病毒平均滴度在感染后7d达到高峰,最大感染剂量组病毒核酸含量高达9.7logs copies/ml。组织病毒含量为4.66～9.05 logs copies/g不等。试验还发现病毒感染猪血清中细胞因子IFN-γ和IL-10浓度升高以及这两种细胞因子表达比例失调。总的实验结果表明,HP-PRRSV变异株对仔猪具有高度致病性,适合用作评价预防HP-PRRS疫苗的强毒参考株。此外,根据实验数据,我们初步制定了HP-PRRSV变异株HuN4对60d龄普通仔猪的强毒标准和发病标准。
为了建立一种可检测HP-PRRSV变异株及其细胞传代致弱株和国内经典毒株的荧光定量RT-PCR方法,研究设计针对PRRSV ORF7基因的特异性引物和TaqMan荧光探针,建立了一种检测PRRSV的快速、敏感、特异、重复性好的TaqMan荧光定量RT-PCR方法。该方法在检测101拷贝/μL～107拷贝/μL模板范围内具有良好的线性关系。比常规RT-PCR更敏感,可分别检测最少为10拷贝的阳性标准品和1TCID50病毒。应用建立的方法检测了HP-PRRSV HuN4第5代和第65代细胞毒人工感染仔猪的血清组织样品的病毒含量。结果证明该方法适合用于病毒的早期感染检测、研究病毒在体内的复制和持续性感染,以及HP-PRRSV弱毒苗的研制及其免疫效果评价等。
重组猪IFN-γ可抑制PRRSV在培养细胞的复制。而且可从感染PRRSV的猪淋巴结、肺脏和外周血单核细胞(PBMC)检测到猪IFN-γ。本研究旨在建立快速、敏感和特异的猪IFN-γmRNA TaqMan荧光定量RT-PCR检测方法,为研究猪IFN-γ等细胞因子在猪体内表达变化等提供检测技术。针对猪IFN-γ基因和管家基因cyclophilin A (CyPA)核苷酸序列分别设计特异性引物和TaqMan荧光探针,以重组质粒pMD18-T-IFN-γ和pMD18-T-CyPA为标准品,进行实时荧光定量RT-PCR检测,并构建猪IFN-γmRNA和CyPA的荧光定量RT-PCR标准曲线。结果表明,建立的方法在检测101拷贝/μL～107拷贝/μL模板范围内具有良好的线性关系,相关系数R2均高达0.999,扩增效率均高于99.0%;可检测最少为100拷贝的阳性标准品。应用建立的方法检测HP-PRRSV变异株HuN4人工感染猪后21d的肺、脾和淋巴结组织样品,发现IFN-γmRNA在上述组织表达量很低。结果证明,该方法快速、敏感性高、特异性强和重复性好,可直接应用于临床样品的检测。
Porcine reproductive and respiratory syndrome (PRRS) is characterized by late-term abortions and delivery of increase number of stillbirths or premature and weak piglets in sows and severe respiratory disease in neonatal and growing pigs. Now the disease is endemic in most areas around the world and recognized as one of most important diseases of swine. In May 2006, a severe disease, initially called‘pig high fever syndrome’initially occurred in some provinces in south of China and then spread widely in China, resulting in huge economic losses in pig industry of China. Now a highly pathogenic porcine reproductive and respiratory syndrome virus (HP-PRRSV) variant has been confirmed to be the main pathogen of the severe outbreaks. Pigs of all age were clinically sick and dead in the severe infection. However, whether the virulence and in vivo pathogenicity and clinical features of the new isolates of PRRSV to pigs of all age were similar to those observed in the field, the pathogenesis and immune response of the virus and the immune efficacy of new vaccines for protection against the highly pathogenic porcine reproductive and respiratory syndrome (HP-PRRS) are still unclear. Animal experimentally infection needs to be done to answer the questions.
To develop a reliable conventional pig model to study the HP-PRRS, three groups of 30-, 60- and 180-day-old pigs were infected with 3×104 50% tissue culture infective dose (TCID50) per pig of a representative HP-PRRSV strain, 5th-passage HuN4 (HuN4-F5) intramuscularly and intranasally. The results showed that the HP-PRRSV HuN4 could induce the typical PRRS in all the conventional pigs and the clinical features were similar to those observed in the field. Among them, 30-day-old and 60-day-old pigs exhibited severe clinical signs with 100% morbidity and 100% and 60% mortality, respectively. Persistently high fever (41.0～42.0°C), dyspnoea, gross lesions of consolidation of lungs,enlargement and/or haemorrhage of lymph nodes and histopathological lesions of interstitial pneumonia, the inflammation of the lymphoid tissue and encephalitis were observed in the two exposed groups. However, the clinical signs, gross lesions and microscopic lesions of the infected 180-day-old pigs were slight compared with other infected groups. So the results demonstrated that the 30 and 60-day-old conventional pigs are suitable to be the infection model for the HP-PRRSV HuN4 strain.
To further investigate the in vivo pathogenicity and virulence and immune responses of the newly emerging PRRSV, 3 groups of 60-day-old conventional piglets were inoculated intranasally with a representative strain of the HP-PRRSV variant, HuN4 with 3 different infection doses(3 103～3 105 TCID50). The clinical features, humoral immune responses, viremia, the viral load in different tissues and levels of IFN-γand IL-10 cytokine production in serum in this study were evaluated. The results revealed that 3 different infection doses of virus could induce severe disease in piglets and the infected pigs in the least infection dose group exhibited a little mild clinical signs than those of other infection dose groups. The significantly and consistently clinical characteristics of the PRRSV-exposed groups consisted of persistently high fever (41.0～41.9℃) and high morbidity and mortality (60～100%), the marked clinical signs of PRRS and severe histopathologic damages in multiple organs. It induced rapid and intense humoral immune responses and seroconversion was detected in most of infected pigs at 7 days post-infection (DPI).The virus rapidly and vigorously replicated in vivo. The virus was isolated from the serum of most of infected pigs at 3 DPI and the viremia was detected in all the infected pigs by 21 DPI days. The highest average titer of viremia peaked near 9.7logs copies/ml serum at 7 DPI. The viral load in different tissue reached 4.66～9.05 logs copies/g. Elevated levels of IFN-γand IL-10 cytokine production in serum and an unbalanced profile of Th1 (IFN-γ) /Th2 (IL-10) responses in this study were also observed. Taken together, our results demonstrated that the HP-PRRSV variant HuN4 strain is a highly pathogenic for piglets and suitable to be a reference strain of highly virulent PRRSV for evaluating the efficacy of the new vaccines to prevent the HP-PRRS. Additionally, according to the experiment data, we initially made a standard of high virulence of the HuN4 for the 60-day-old conventional pigs and a clinical standard of HP-PRRS.
To establish a fluorescent quantitative reverse transcription-polymerase chain reaction (RT-PCR) assay for the detection of HP-PRRSV and its attenuated virus and the classical NA-type PRRSV in China,the specific primers and fluorogenetic probe based on the ORF7 gene of PRRSV were designed and a quantitative TaqMan RT-PCR for detecting PRRSV had been developed. The established technique was time-saving, highly sensitive and specific and had reliable reproducibility. The assay for the quantitation of PRRSV template was found to be in the 101～107copies/μL linear dynamic range with excellent linearity. It is more sensitive than conventional RT-PCR and could detect the least template of 10 copies and 1 TCID50 virus. Using the technique, serum and tissue samples from pigs experimentally infected with HP-PRRSV HuN4-F5, and the sixty-fifth passage attenuated virus (HuN4-F65) were collected to detect viral load. The results demonstrated the established technique is suitable for detection PRRSV in early infection, replication and persistent infection in vivo, development and evaluation the efficacy of HP-PRRSV attenuated virus vaccines.
The recombinant porcine interferon-γ(IFN-γ) can inhibit the replication of PRRSV in vitro and the IFN-γcould be detected from the lymph node, lung and peripheral blood monocytes (PBMC) of PRRSV infected pigs. To establish a fluorescent quantitative RT-PCR assay for the detection of porcine IFN-γ, the specific primers and fluorogenetic probes based on the gene of porcine IFN-γand housekeeping gene cyclophilin A (CyPA) were designed, by using the recombinant plasmids pMD18-T-IFN-γand pMD18-T-CyPA as standard products, a real-time quantitative TaqMan reverse transcription-polymerase chain reaction (RT-PCR) was performed to construct the standard curves of porcine IFN-γand CyPA. The results indicated that the assay for the quantitation of porcine IFN-γtemplate could work well in the range of 101～107copies/μL with excellent linearity, the coefficient correlation R squared values reached 0.999 and amplification efficiency was higher than 99.0%. The sensitivity was sufficient to detect the least template of 100 copies. IFN-γmRNA in tissue samples from the spleen, lung and lymph node in the euthanatized pigs infected with the HuN4-F5 virus at 21DPI were collected and quantitatively detected by the established assay. The results showed that the expression of IFN-γmRNA in infected pigs is very low with a range of 6.44×102～1.04×104 copies/μL cDNA. Therefore, The currently established porcine IFN-γspecific real time RT-PCR was time-saving, highly sensitive and specific, and had reliable reproducibility and suitable to detect the clinical specimens.
引文
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