2009-2011年国内传染性支气管炎病毒分子流行病学研究及以马立克病毒为载体的基因工程疫苗研制
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摘要
禽传染性支气管炎病毒(avian infectious bronchitis virus,IBV)属于冠状病毒科冠状病毒属中的γ-冠状病毒,由该病毒感染引起的疾病称为禽传染性支气管炎(avian infectious bronchitis,IB),是一种对鸡危害非常严重的急性、高度传染性的病毒性呼吸道疾病,可导致鸡的增重和饲料报酬降低,也能引起蛋鸡产蛋量和蛋的品质下降等,因此具有重要的经济意义,免疫预防是目前控制该病的最为有效的手段。IBV的重要特点之一就是血清型众多,不同血清型之间交叉保护性较低甚至完全不能保护,因此在生产中必须选用与流行毒株血清型一致的疫苗才能取得良好的免疫保护效果。不同血清型的IBV具有典型的地域性流行特征,而且由于病毒基因组的突变和毒株间的重组使得新的血清型不断出现,使免疫预防日益复杂化。本研究的目的旨在通过系统的流行病学监测了解近年来不同基因型IBV在我国的流行状况,通过生物信息学分析阐明不同IBV基因型间遗传演化的规律和发展趋势,并针对流行的优势基因型毒株研制新型疫苗。
1、2009-2011年间国内IBV流行病学监测与流行株的遗传进化分析
2009-2011年间,共从国内主要养鸡省份分离到IBV流行毒株54株,其中肉鸡源毒株52株,蛋鸡源毒株2株。在肉鸡源毒株中我们发现存在较高比例的IBV与H9亚型AIV的混合感染(25/52),发生混合感染的鸡群往往发病更为严重,且混合感染样品对接种的鸡胚表现出更强的致病性,该结果提示这两种病毒之间可能存在一定程度的致病协同作用。对所有分离毒株S1基因进行克隆和基因测序,共发现4种不同的S1基因长度,分别为1611bp、1617bp、1620bp和1626bp,说明S1基因中碱基的插入和缺失是一种非常普遍的现象。将54个分离株S1基因序列和近年来在国内及周边国家和地区流行的13个基因型共57个参考毒株的S1基因序列用Mega4.1软件进行遗传进化分析,结果显示54个分离株可以分为QX型(42株)、HN-08型(5株)、LSC/991型(2株)、TW-I型(2株)、Mass型(1株)、793/B型(1株)和CHⅢ型(1株)等7个基因型。在所有分离株中,QX型分离株数量最多,占总数的77.8%(42/54)。从进化树中还可以看出,QX型分离株又可以进一步细分为两个不同的基因亚群-Cluster Ⅰ(21株)和Cluster Ⅱ(21株),经典的QX型参考毒株QXIBV和LX4株均属于Cluster Ⅰ亚群,而Cluster Ⅱ亚群中的毒株均在2010年才开始出现。为了更加系统地描述同期内国内不同基因型IBV的分布情况,我们将已经在GenBank发表全长S1基因序列的所有同时期分离株共296株合并进行遗传进化分析,结果显示,350个同时期分离株可分为19个基因型,其中11个基因型均能根据现有文献报道找到同源的参考毒株,根据各基因型中包含的毒株数量多少依次为：QX型(219株)、HN-08型(26株)、LSC/99I型(22株)、LDT3/03型(12株)、TW-I型(12株)、CH Ⅲ型(11株)、Mass型(10株)、LDL/971型(3株)、CH VI型(3株)、TW-Ⅱ型(2株)和793/B型(1株),此外还有8个基因型未能归入已被正式命名的已知基因型,因此暂定名为未定型Ⅰ～未定型Ⅷ。从分型结果可以看出,我国流行的IBV基因型具有典型的地域性分布特征,大多数在周边国家和地区广泛流行的基因型在我国并未发现。
为阐明国内不同基因型毒株之间的遗传演化关系,根据遗传进化和基因分型的结果,从不同基因型中选择代表性毒株,用RDP3.31软件对S1基因进行重组分析。结果显示当前国内流行的毒株中存在广泛的重组现象,不同基因型间的重组事件不仅导致同一基因型内部发生分化,出现了不同基因亚群,而且经过多次的同源重组后可能会导致新的基因型毒株的出现。综合不同基因型间发生的系列重组事件,可以推断：QX型、LSC/991型、793/B型、LDL/971型、TW-Ⅰ型和TW-Ⅱ型是目前在我国流行的所有基因型毒株的原型毒株,除此以外的新出现的基因型均是在这6个基因型的基础上经过一次或多次的同源重组而产生的变异株。
2、Mass型H120活疫苗对当前流行的主要基因型代表毒株的免疫保护效力试验
根据遗传进化分析的结果,从当前主要流行的三个IBV基因型中挑选出4个代表毒株：AH/2009/1(QX型Cluster Ⅰ亚群)、JS/2010/12(QX型Cluster Ⅱ亚群)、JS/2009/5(LSC/99I型)和JS/2010/6(HN08型),分别用Mass型IBV活疫苗(H120株)进行免疫保护效力试验,以评价现有疫苗对流行毒株的免疫保护状况。结果显示H120疫苗株对不同基因型的IBV毒株免疫保护效力存在较大差异,对同属于Mass基因型的M41强毒株能够提供完全保护，且能显著降低攻毒后气管和肾脏的排毒率；对于QX型毒株AH/2009/1和JS/2010/12,基本达到临床保护标准,但试验鸡气管和肾脏带毒的比例远远高于M41攻毒组,说明局部免疫保护效果不是非常理想；对于HN08型的JS/2010/6株和LSC/991型的JS/2009/5株保护效果均不理想,攻毒后鸡群表现出较为严重的临床症状和相当高的气管和肾脏排毒率。该结果提示我们,现有疫苗已经不能对当前流行的主要基因型毒株提供良好的免疫保护,急需研究开发与流行毒株基因型一致的新疫苗用于该病的防控。
3、以马立克病毒为载体研制QX型IBV基因工程疫苗
本研究以MDV CV1988/Rispens疫苗毒株为载体,以其IRS-US间隔区作为插入位点,选择网状内皮组织增生症病毒(REV)基因组长末端重复序列(LTR)作为启动子,构建含QX型IBV分离株CK/CH/JS/06Ⅱ S1基因及报告基因EGFP的转移载体pUP-LTR-EGFP-S1-DOWN。将MDV基因组DNA与转移载体共转染鸡胚成纤维细胞(CEF),经同源重组获得重组病毒,通过多轮荧光筛选和纯化,最终获得的重组病毒命名为rMDV-EGFP-S1。进一步通过Cre重组酶介导的重组反应敲除EGFP报告基因,获得仅带有LTR启动子和S1基因的重组病毒,命名为rMDV-S1。通过PCR和间接免疫荧光试验(IFA)对重组病毒进行鉴定,结果表明S1基因完全按照预定方式正确插入到MDV基因组中并能够表达S1蛋白。将重组病毒在CEF上进行连续30代传代培养,经PCR和IFA试验鉴定,该重组病毒具有良好的遗传稳定性。
重组病毒rMDV-S1。对QX型IBV强毒株CK/CH/JS/06Ⅱ的免疫攻毒保护效力试验结果显示,重组病毒按5×103PFU/羽的剂量免疫1日龄SPF鸡,免疫后4w攻毒,免疫组试验鸡发病率和死亡率分别为30%和5%,对照组则分别为100%和30%,经χ2检验差异显著(p<0.05)。排毒检测结果显示,免疫组气管排毒率在攻毒后部分时间点与对照组相比差异显著,攻毒后14d扑杀时肾脏排毒率差异极显著(p<0.01)。免疫组试验鸡攻毒后的增重未受明显影响,与对照组相比差异显著(p<0.05)。病理组织学检查结果显示与对照组相比免疫组试验鸡肾脏的病变程度相对较轻。所有结果均表明重组病毒对QX型强毒株的攻击能够提供良好的免疫保护。
重组病毒对MDV强毒株RB1B的免疫攻毒保护试验结果显示,与亲本病毒CV1988/Rispens相比,对MDV本身的免疫保护效率未受明显影响,说明该重组病毒可以作为二联疫苗同时用于IB和MD两种疾病的免疫预防。
Avian infectious bronchitis virus (IBV), a member of the genus Gammacoronavirus of the family Coronaviridae, causes avian infectious bronchitis (IB), an acute, highly contagious respiratory disease in chickens. IB is of major economic importance because it is a cause of poor weight gain and feed efficiency and reduced egg production and egg quality. Live or inactivated vaccines containing strains of IBV from multiple serotypes are routinely used to protect chickens against IB infection in commercial settings. In nature, IBVs can evolve rapidly to give rise to new antigenic variants, which have complicated and increased the cost of attempts to prevent the disease by immunization. The aims of this study were to elucidate the predominant genotypes circulating in China in recent years and define their phylogenetic relationships, as well as to develop a new recombinant viral vaccine effective against the current epidemic genotypes using Marek's disease virus (MDV) as a vector.
1. Epidemiological surveillance and phylogenetic analysis of IBVs circulating in China between2009and2011
A total of54IBV field strains were isolated from chicken flocks suspected of infection with IB in8provinces of China between2009and2011. Of the54strains collected,52were isolated from broiler chickens and2were isolated from egg-laying chickens. Frequent high mortality (25/52flocks) caused by coinfection with IBV and H9subtype avian influenza viruses (AIV) have been observed in broiler chicken flocks in this study. The multiple infection samples also exhibited more severe pathogenicity in inoculated embryos. This phenomenon suggested that there is potential synergism between IBV and H9subtype AIV.
The S1genes of all isolates were cloned and sequenced, revealing4different S1genes, including1611-,1617-,1620-and1626-nucleotide isoforms. This demonstrated that nucleotide insertion or deletion is common in the S1gene. The sequences of S1genes from all isolates and57reference strains representing different genotypes recently circulating in China and surrounding countries or areas were aligned, and a phylogenetic tree was constructed. The results revealed that all of the isolates could be grouped into7genotypes:QX (42isolates). HN-08(5isolates), LSC7991(2isolates). TW-I (2isolates), Mass (1isolate),793/B (1isolate) and CH Ⅲ (1isolate). The QX type has been the predominant genotype recently circulating in China, and strains of this type could be further subdivided into Cluster Ⅰ (21isolates) and Cluster Ⅱ (21isolates). The classical QX-type reference strain QXIBV is in Cluster Ⅰ, and all Cluster Ⅱ strains were found in samples obtained prior tosince2010.
To more clearly describe the distribution of the different genotypes in China,296isolates between2009and2011with complete S1genes published in GenBank were combined for further phylogenetic analysis. These results revealed that all350isolates could be grouped into19genotypes, including11known genotypes and8novel genotypes. The11known genotypes were QX (219isolates), HN-08(26isolates), LSC/991(22isolates), LDT3/03(12isolates), TW-I (12isolates), CH III (11isolates), Mass (10isolates), LDL/971(3isolates), CH VI (3isolates), TW-II (2isolates) and793/B (1isolate). The8novel genotypes were tentatively named undetermined genotypes I-VIII. These results strongly suggested that IBV epidemiology in China is very complex and shows distinct regional distribution characteristics; interestingly, most of the genotypes circulating in surrounding countries and areas are not found in China.
To clarify the phylogenetic relationships between the different IBV genotypes circulating in China, the S1gene sequences of some strains in different genotypes were chosen for recombination analysis with RDP3.31software. The results revealed that recombination between S1genes is a common phenomenon. A series of recombination incidents not only accelerate genotype differentiation but also lead to the emergence of new genotypes. According to the results of recombinant analysis, we were able to conclude that all current epidemic genotypes in China originated from6parental genotypes:QX, LSC/99I,793/B, LDL/97I, TW-Ⅰ and TW-Ⅱ. We further concluded that all newly emerged genotypes were variants generated by one or more homologous recombination events between those6prototype strains.
2. Protective efficacy of Mass type H120live vaccine against some genotypes of IBV
Four IBV strains representing three predominant genotypes were selected to evaluate the protective efficacy offered by Mass type H120live vaccine according to the results of our phylogenetic analysis. The corresponding genotypes were QX type Cluster1(AH/2009/I), QX type Cluster Ⅱ (JS/2010/12). LSC/99I type (JS/2009/5) and HN08type (JS/2010/6), respectively. In this experiment,3-day-old young SPF chickens were vaccinated with HI20live vaccine and challenged21d post-vaccination with4field isolates and another virulent Mass type IBV reference strain M41. The morbidity, mortality and virus shedding from the trachea and kidney of challenged chickens were recorded for each group. The results showed that the protective efficacy was different against every virus. The vaccine conferred complete protection against the homologous Mass type M41strain, and the chickens showed low shedding rates post-challenge. Although the vaccination also conferred clinical protection against the QX-like strains AH/2009/1and JS/2010/12. high rates of virus shedding were detected in the tracheae and kidneys of challenged chickens. The vaccination did not provide good protection against strains JS/2009/5and JS/2010/6, and the challenged chickens showed serious clinical signs of infection and high rates of virus shedding. All of these results suggested that the Mass type vaccine cannot provide complete protection against new epidemic genotypes of IBVs isolated in China, and it is therefore urgent to develop new vaccines to control this disease.
3. Construction of a recombinant Marek's disease virus expressing the QX-type IBV S1protein
In this study, we constructed a recombinant CVI988/Rispens, a strain of type Ⅰ MDV, that expresses the S1protein of the QX type IBV strain CK/CH/JS/06Ⅱ. The IRS-US intergenic region of the MDV genome was used as the gene insertion site in which the LTR promoter from the reticuloendotheliosis virus (REV) and S1gene were incorporated into the MDV genome. The transfer vector pUP-LTR-EGFP-S1-DOWN and MDV genomic DNA were co-transfected into chicken embryo fibroblast cells (CEF) to generate the recombinant virus rMDV-EGFP-S1by homologous recombination. The fluorescent plaques formed by recombinant virus were picked and purified by repeated passages in CEFs. Then, the EGFP gene was knocked out by Cre-mediated recombination, and the rendered recombinant virus containing the LTR promoter and the S1gene was designated rMDV-S1. The recombinant virus was identified through PCR and an indirect immunofluorescence antibody assay (IFA), and the results showed that the S1gene was correctly inserted into the MDV genome and was stably expressed. The rMDV-S1virus was genetically stable after30sequential passages in CEFs.
In the experiment evaluating the protective efficacy of rMDV vaccination against IBV,1-day-old SPF chickens were immunized with5×103PFU rMDV-S1per bird and challenged with QX type virulent IBV strain CK/CH/JS/06Ⅱ. The morbidity and mortality were30%and5%, respectively, compared to100%and30%in the control group, a significant difference (p <0.05), as determined by chi-squared(χ2) test, in the virus shedding test, there was a significant difference in the number of tracheal swabs that tested positive for the virus between the rMDV-S1-vaccinated group and the control group (p<0.05). The virus detection rate from the kidneys of the immunized chickens was lower than that in the control group. There was a highly significant difference (p<0.01) in the number of kidneys that tested positive for the virus between the vaccinated and control groups. All of these results demonstrated that rMDV-S1is an effective and promising recombinant vaccine for the prevention of QX-like IBV infection.
In an experiment evaluating the protective efficacy of rMDV vaccination against MDV, the rMDV-Sl showed similar protective efficacy against virulent MDV RB1B and its parent virus CVI988/Rispens. This result showed that the rMDV-Sl can serve as a vaccine for the concomitant immunoprophylaxis of IB and MD.
1、2009-2011年间国内IBV流行病学监测与流行株的遗传进化分析
2009-2011年间,共从国内主要养鸡省份分离到IBV流行毒株54株,其中肉鸡源毒株52株,蛋鸡源毒株2株。在肉鸡源毒株中我们发现存在较高比例的IBV与H9亚型AIV的混合感染(25/52),发生混合感染的鸡群往往发病更为严重,且混合感染样品对接种的鸡胚表现出更强的致病性,该结果提示这两种病毒之间可能存在一定程度的致病协同作用。对所有分离毒株S1基因进行克隆和基因测序,共发现4种不同的S1基因长度,分别为1611bp、1617bp、1620bp和1626bp,说明S1基因中碱基的插入和缺失是一种非常普遍的现象。将54个分离株S1基因序列和近年来在国内及周边国家和地区流行的13个基因型共57个参考毒株的S1基因序列用Mega4.1软件进行遗传进化分析,结果显示54个分离株可以分为QX型(42株)、HN-08型(5株)、LSC/991型(2株)、TW-I型(2株)、Mass型(1株)、793/B型(1株)和CHⅢ型(1株)等7个基因型。在所有分离株中,QX型分离株数量最多,占总数的77.8%(42/54)。从进化树中还可以看出,QX型分离株又可以进一步细分为两个不同的基因亚群-Cluster Ⅰ(21株)和Cluster Ⅱ(21株),经典的QX型参考毒株QXIBV和LX4株均属于Cluster Ⅰ亚群,而Cluster Ⅱ亚群中的毒株均在2010年才开始出现。为了更加系统地描述同期内国内不同基因型IBV的分布情况,我们将已经在GenBank发表全长S1基因序列的所有同时期分离株共296株合并进行遗传进化分析,结果显示,350个同时期分离株可分为19个基因型,其中11个基因型均能根据现有文献报道找到同源的参考毒株,根据各基因型中包含的毒株数量多少依次为：QX型(219株)、HN-08型(26株)、LSC/99I型(22株)、LDT3/03型(12株)、TW-I型(12株)、CH Ⅲ型(11株)、Mass型(10株)、LDL/971型(3株)、CH VI型(3株)、TW-Ⅱ型(2株)和793/B型(1株),此外还有8个基因型未能归入已被正式命名的已知基因型,因此暂定名为未定型Ⅰ～未定型Ⅷ。从分型结果可以看出,我国流行的IBV基因型具有典型的地域性分布特征,大多数在周边国家和地区广泛流行的基因型在我国并未发现。
为阐明国内不同基因型毒株之间的遗传演化关系,根据遗传进化和基因分型的结果,从不同基因型中选择代表性毒株,用RDP3.31软件对S1基因进行重组分析。结果显示当前国内流行的毒株中存在广泛的重组现象,不同基因型间的重组事件不仅导致同一基因型内部发生分化,出现了不同基因亚群,而且经过多次的同源重组后可能会导致新的基因型毒株的出现。综合不同基因型间发生的系列重组事件,可以推断：QX型、LSC/991型、793/B型、LDL/971型、TW-Ⅰ型和TW-Ⅱ型是目前在我国流行的所有基因型毒株的原型毒株,除此以外的新出现的基因型均是在这6个基因型的基础上经过一次或多次的同源重组而产生的变异株。
2、Mass型H120活疫苗对当前流行的主要基因型代表毒株的免疫保护效力试验
根据遗传进化分析的结果,从当前主要流行的三个IBV基因型中挑选出4个代表毒株：AH/2009/1(QX型Cluster Ⅰ亚群)、JS/2010/12(QX型Cluster Ⅱ亚群)、JS/2009/5(LSC/99I型)和JS/2010/6(HN08型),分别用Mass型IBV活疫苗(H120株)进行免疫保护效力试验,以评价现有疫苗对流行毒株的免疫保护状况。结果显示H120疫苗株对不同基因型的IBV毒株免疫保护效力存在较大差异,对同属于Mass基因型的M41强毒株能够提供完全保护，且能显著降低攻毒后气管和肾脏的排毒率；对于QX型毒株AH/2009/1和JS/2010/12,基本达到临床保护标准,但试验鸡气管和肾脏带毒的比例远远高于M41攻毒组,说明局部免疫保护效果不是非常理想；对于HN08型的JS/2010/6株和LSC/991型的JS/2009/5株保护效果均不理想,攻毒后鸡群表现出较为严重的临床症状和相当高的气管和肾脏排毒率。该结果提示我们,现有疫苗已经不能对当前流行的主要基因型毒株提供良好的免疫保护,急需研究开发与流行毒株基因型一致的新疫苗用于该病的防控。
3、以马立克病毒为载体研制QX型IBV基因工程疫苗
本研究以MDV CV1988/Rispens疫苗毒株为载体,以其IRS-US间隔区作为插入位点,选择网状内皮组织增生症病毒(REV)基因组长末端重复序列(LTR)作为启动子,构建含QX型IBV分离株CK/CH/JS/06Ⅱ S1基因及报告基因EGFP的转移载体pUP-LTR-EGFP-S1-DOWN。将MDV基因组DNA与转移载体共转染鸡胚成纤维细胞(CEF),经同源重组获得重组病毒,通过多轮荧光筛选和纯化,最终获得的重组病毒命名为rMDV-EGFP-S1。进一步通过Cre重组酶介导的重组反应敲除EGFP报告基因,获得仅带有LTR启动子和S1基因的重组病毒,命名为rMDV-S1。通过PCR和间接免疫荧光试验(IFA)对重组病毒进行鉴定,结果表明S1基因完全按照预定方式正确插入到MDV基因组中并能够表达S1蛋白。将重组病毒在CEF上进行连续30代传代培养,经PCR和IFA试验鉴定,该重组病毒具有良好的遗传稳定性。
重组病毒rMDV-S1。对QX型IBV强毒株CK/CH/JS/06Ⅱ的免疫攻毒保护效力试验结果显示,重组病毒按5×103PFU/羽的剂量免疫1日龄SPF鸡,免疫后4w攻毒,免疫组试验鸡发病率和死亡率分别为30%和5%,对照组则分别为100%和30%,经χ2检验差异显著(p<0.05)。排毒检测结果显示,免疫组气管排毒率在攻毒后部分时间点与对照组相比差异显著,攻毒后14d扑杀时肾脏排毒率差异极显著(p<0.01)。免疫组试验鸡攻毒后的增重未受明显影响,与对照组相比差异显著(p<0.05)。病理组织学检查结果显示与对照组相比免疫组试验鸡肾脏的病变程度相对较轻。所有结果均表明重组病毒对QX型强毒株的攻击能够提供良好的免疫保护。
重组病毒对MDV强毒株RB1B的免疫攻毒保护试验结果显示,与亲本病毒CV1988/Rispens相比,对MDV本身的免疫保护效率未受明显影响,说明该重组病毒可以作为二联疫苗同时用于IB和MD两种疾病的免疫预防。
Avian infectious bronchitis virus (IBV), a member of the genus Gammacoronavirus of the family Coronaviridae, causes avian infectious bronchitis (IB), an acute, highly contagious respiratory disease in chickens. IB is of major economic importance because it is a cause of poor weight gain and feed efficiency and reduced egg production and egg quality. Live or inactivated vaccines containing strains of IBV from multiple serotypes are routinely used to protect chickens against IB infection in commercial settings. In nature, IBVs can evolve rapidly to give rise to new antigenic variants, which have complicated and increased the cost of attempts to prevent the disease by immunization. The aims of this study were to elucidate the predominant genotypes circulating in China in recent years and define their phylogenetic relationships, as well as to develop a new recombinant viral vaccine effective against the current epidemic genotypes using Marek's disease virus (MDV) as a vector.
1. Epidemiological surveillance and phylogenetic analysis of IBVs circulating in China between2009and2011
A total of54IBV field strains were isolated from chicken flocks suspected of infection with IB in8provinces of China between2009and2011. Of the54strains collected,52were isolated from broiler chickens and2were isolated from egg-laying chickens. Frequent high mortality (25/52flocks) caused by coinfection with IBV and H9subtype avian influenza viruses (AIV) have been observed in broiler chicken flocks in this study. The multiple infection samples also exhibited more severe pathogenicity in inoculated embryos. This phenomenon suggested that there is potential synergism between IBV and H9subtype AIV.
The S1genes of all isolates were cloned and sequenced, revealing4different S1genes, including1611-,1617-,1620-and1626-nucleotide isoforms. This demonstrated that nucleotide insertion or deletion is common in the S1gene. The sequences of S1genes from all isolates and57reference strains representing different genotypes recently circulating in China and surrounding countries or areas were aligned, and a phylogenetic tree was constructed. The results revealed that all of the isolates could be grouped into7genotypes:QX (42isolates). HN-08(5isolates), LSC7991(2isolates). TW-I (2isolates), Mass (1isolate),793/B (1isolate) and CH Ⅲ (1isolate). The QX type has been the predominant genotype recently circulating in China, and strains of this type could be further subdivided into Cluster Ⅰ (21isolates) and Cluster Ⅱ (21isolates). The classical QX-type reference strain QXIBV is in Cluster Ⅰ, and all Cluster Ⅱ strains were found in samples obtained prior tosince2010.
To more clearly describe the distribution of the different genotypes in China,296isolates between2009and2011with complete S1genes published in GenBank were combined for further phylogenetic analysis. These results revealed that all350isolates could be grouped into19genotypes, including11known genotypes and8novel genotypes. The11known genotypes were QX (219isolates), HN-08(26isolates), LSC/991(22isolates), LDT3/03(12isolates), TW-I (12isolates), CH III (11isolates), Mass (10isolates), LDL/971(3isolates), CH VI (3isolates), TW-II (2isolates) and793/B (1isolate). The8novel genotypes were tentatively named undetermined genotypes I-VIII. These results strongly suggested that IBV epidemiology in China is very complex and shows distinct regional distribution characteristics; interestingly, most of the genotypes circulating in surrounding countries and areas are not found in China.
To clarify the phylogenetic relationships between the different IBV genotypes circulating in China, the S1gene sequences of some strains in different genotypes were chosen for recombination analysis with RDP3.31software. The results revealed that recombination between S1genes is a common phenomenon. A series of recombination incidents not only accelerate genotype differentiation but also lead to the emergence of new genotypes. According to the results of recombinant analysis, we were able to conclude that all current epidemic genotypes in China originated from6parental genotypes:QX, LSC/99I,793/B, LDL/97I, TW-Ⅰ and TW-Ⅱ. We further concluded that all newly emerged genotypes were variants generated by one or more homologous recombination events between those6prototype strains.
2. Protective efficacy of Mass type H120live vaccine against some genotypes of IBV
Four IBV strains representing three predominant genotypes were selected to evaluate the protective efficacy offered by Mass type H120live vaccine according to the results of our phylogenetic analysis. The corresponding genotypes were QX type Cluster1(AH/2009/I), QX type Cluster Ⅱ (JS/2010/12). LSC/99I type (JS/2009/5) and HN08type (JS/2010/6), respectively. In this experiment,3-day-old young SPF chickens were vaccinated with HI20live vaccine and challenged21d post-vaccination with4field isolates and another virulent Mass type IBV reference strain M41. The morbidity, mortality and virus shedding from the trachea and kidney of challenged chickens were recorded for each group. The results showed that the protective efficacy was different against every virus. The vaccine conferred complete protection against the homologous Mass type M41strain, and the chickens showed low shedding rates post-challenge. Although the vaccination also conferred clinical protection against the QX-like strains AH/2009/1and JS/2010/12. high rates of virus shedding were detected in the tracheae and kidneys of challenged chickens. The vaccination did not provide good protection against strains JS/2009/5and JS/2010/6, and the challenged chickens showed serious clinical signs of infection and high rates of virus shedding. All of these results suggested that the Mass type vaccine cannot provide complete protection against new epidemic genotypes of IBVs isolated in China, and it is therefore urgent to develop new vaccines to control this disease.
3. Construction of a recombinant Marek's disease virus expressing the QX-type IBV S1protein
In this study, we constructed a recombinant CVI988/Rispens, a strain of type Ⅰ MDV, that expresses the S1protein of the QX type IBV strain CK/CH/JS/06Ⅱ. The IRS-US intergenic region of the MDV genome was used as the gene insertion site in which the LTR promoter from the reticuloendotheliosis virus (REV) and S1gene were incorporated into the MDV genome. The transfer vector pUP-LTR-EGFP-S1-DOWN and MDV genomic DNA were co-transfected into chicken embryo fibroblast cells (CEF) to generate the recombinant virus rMDV-EGFP-S1by homologous recombination. The fluorescent plaques formed by recombinant virus were picked and purified by repeated passages in CEFs. Then, the EGFP gene was knocked out by Cre-mediated recombination, and the rendered recombinant virus containing the LTR promoter and the S1gene was designated rMDV-S1. The recombinant virus was identified through PCR and an indirect immunofluorescence antibody assay (IFA), and the results showed that the S1gene was correctly inserted into the MDV genome and was stably expressed. The rMDV-S1virus was genetically stable after30sequential passages in CEFs.
In the experiment evaluating the protective efficacy of rMDV vaccination against IBV,1-day-old SPF chickens were immunized with5×103PFU rMDV-S1per bird and challenged with QX type virulent IBV strain CK/CH/JS/06Ⅱ. The morbidity and mortality were30%and5%, respectively, compared to100%and30%in the control group, a significant difference (p <0.05), as determined by chi-squared(χ2) test, in the virus shedding test, there was a significant difference in the number of tracheal swabs that tested positive for the virus between the rMDV-S1-vaccinated group and the control group (p<0.05). The virus detection rate from the kidneys of the immunized chickens was lower than that in the control group. There was a highly significant difference (p<0.01) in the number of kidneys that tested positive for the virus between the vaccinated and control groups. All of these results demonstrated that rMDV-S1is an effective and promising recombinant vaccine for the prevention of QX-like IBV infection.
In an experiment evaluating the protective efficacy of rMDV vaccination against MDV, the rMDV-Sl showed similar protective efficacy against virulent MDV RB1B and its parent virus CVI988/Rispens. This result showed that the rMDV-Sl can serve as a vaccine for the concomitant immunoprophylaxis of IB and MD.
引文
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