传染性法氏囊病毒感染的生物标记与细胞SAGE文库的鉴定
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摘要
传染性法氏囊病病毒(Infectious bursal disease virus, IBDV)属于双RNA病毒科、禽双RNA病毒属,分为血清Ⅰ型和血清Ⅱ型。血清Ⅰ型对鸡有致病性,血清Ⅱ型对鸡无致病性。IBDV基因组由双股RNA组成：A节段和B节段,编码5种蛋白。B节段有一个ORF,编码RNA依赖的RNA聚合酶(VP1)。A节段有两个ORF,大ORF编码病毒的VP2、VP3和VP4蛋白；小ORF编码VP5蛋白。IBDV发病机理不清是该病控制的难点。为此,本研究在转录组和体液免疫水平上对IBDV与宿主的相互作用进行了分析。
首先,在转录组水平上,本研究分析了IBDV感染及IBDV编码基因转化对宿主Vero细胞基因转录的影响。以实验室已构建的5个Long-SAGE文库：正常Vero细胞文库、IBDV感染的Vero细胞文库、表达A节段编码蛋白的单克隆Vero细胞文库、表达VP243蛋白的单克隆Vero细胞文库、表达VP5蛋白的单克隆Vero细胞文库为基础,对文库中转录显著变化(P<0.05)的基因进行统计分析,发现217个可注释基因的转录发生显著变化。对差异转录基因进行聚类分析,发现蛋白质代谢、能量代谢、mRNA加工和细胞骨架相关基因转录变化最为显著。57个显著变化基因的荧光定量PCR验证结果显示,荧光定量PCR结果与文库数据的符合程度是60%-70%,确证了IBDV感染及IBDV编码基因Vero细胞SAGE文库大规模测序基因标签数据的正确性,这为IBDV的发病机制研究提供了大量的基因生物信息。
以IBDV攻毒后的法氏囊组织和感染的鸡胚成纤维细胞RNA为模板,以IBDV感染Vero细胞的基因转录变化为线索,选取IBDV感染Vero细胞Long-SAGE文库中显著变化的15个基因和17个促炎性因子基因,设计鸡源引物,借助荧光定量PCR方法,分析了IBDV感染的法氏囊组织和鸡胚成纤维细‘胞的转录本变化。在IBDV感染的鸡胚成纤维细胞中,HMGN2的显著下调,推测与病毒感染细胞的细胞凋亡相关；促炎性因子表达普遍上调,其中β干扰素(IFN-p)转录本上调500倍以上,其他促炎性因子如IFN-G、BAFF、IL-6、IL-1B、IL-8和iNOS也上调20倍以上。在感染IBDV的法氏囊组织中,氯离子通道蛋白4(CLIC4)显著表达上调,推测与病毒的破膜释放相关；促炎性因子IL-6的转录上调1746倍。表达上调20倍以上的基因还有TNFSF8、IFN-B、IFN-G、IL-8和iNOS。这些数据说明,IBDV感染导致不同类型宿主细胞基因变化的种类具有一致性,但变化的趋势具有不完全一致性,充分显示了不同类型细胞的易感差异性。
在IBDV感染的法氏囊组织中,我们发现Cofilin基因转录显著上调,为分析Cofilin基因表达与IBDV感染之间的关系,本试验利用抗Cofilin蛋白抗体对IBDV感染的鸡胚成纤维细胞和法氏囊组织进行分析,结果发现IBDV感染能够激活Cofilin基因的表达。
基于IBDV感染宿主细胞的转录本信息,为了了解宿主对IBDV非结构蛋白VP4的信号反馈,以重组表达的VP4、VP3蛋白为抗原,制备了抗VP4、VP3单克隆抗体和多克隆抗体,建立了VP4抗体检测ELISA。分析了IBDV强毒株(NB株,BC6/85株)、中等偏强毒力株(MB43株)、中等毒力株(B87株)、低毒力毒株(NB弱毒株)感染21日龄SPF Leghorn鸡的VP4蛋白的体液免疫应答,ELISA抗体监测结果显示：IBDV强毒感染鸡VP4抗体于感染后23天阳转,40天阳性率达70%以上,VP3抗体于感染后20天阳转,23天阳性率达到100%；IBDV弱毒疫苗一次免疫鸡难以检出VP4抗体,VP3抗体于20天阳转,23天阳性率达到88.9%。中等毒力IBDV感染后,VP4抗体阳性率为10%到40%,中等偏强毒力IBDV感染后VP4抗体阳性率为30%到60%。以IBDV VP3、VP4基因Vero细胞表达蛋白的IFA检测出现了类似的结果。对感染后血清VP4抗体效价测定发现,强毒感染血清VP4抗体效价与中等偏强毒力、中等毒力、弱毒相比较,具有显著差异。抗VP3、VP4单克隆抗体的免疫组织化学检测进一步显示：VP3、VP4抗原可以在IBDV强毒感染鸡法氏囊中检测到,而在弱毒疫苗感染鸡中只能检测到VP3抗原,难以检测到VP4抗原。这些数据证明VP4抗原及抗体是鉴别IBDV野毒与疫苗毒感染的生物标记。
Infectious bursal disease virus (IBDV), a member of the genus Avibirnavirus in the family Birnaviridae, has two serotype, serotype 1 and serotype 2. Serotype 1 viruses are pathogenic for chickens, whereas serotype 2 viruses are avirulent. Genome of IBDV consists of two segments (A and B) of double-stranded RNA (dsRNA) and codes 5 proteins. The large segment A, contains two open reading frames (ORFs). A small ORF encodes the nonstructural protein VP5, which overlaps the N terminal region of a precursor polyprotein from the large ORF. The precursor polyprotein is autoproteolytically cleaved to form mature viral proteins VP2-VP4. Segment B encodes VP1, which has RNA-dependent RNA polymerase (RDRP) activity. Infectious bursal disease (IBD) is an immunosuppression disease; IBD cause the death of infected chicken or vaccination failure of other vaccines for immunosuppression. Today, this disease still threatens poultry industry. The pathogenesis is still unclear. In the present study, the interactions of IBDV and hosts were deeply researched at transcriptomic and humoral levels.
Firstly, at transcriptome level, this study analyzed the impacts of IBDV infection or IBDV coding genes transformation on host transcriptome. Based on 5 Long-SAGE libraries:normal Vero cell, IBDV infected Vero cell, monoclonal Vero cell line expressing A-segment coding proteins, monoclonal Vero cell line expressing VP243 protein, monoclonal Vero cell line expressing VP5 protein, differentially expressed genes were analyzed (p<0.05),217 differentially expressed genes were identified. the influences of different viral genes on Vero cell transcriptome and metabolism were summarized. By classing the differential expression genes, we discovered that the protein metabolism, energy metabolism and cytoskeleton related genes were influenced. Real-time PCR analysis was performed to verify 57 genes expression which were differentially expressed in Long-SAGE library. The coincidence of the real-time PCR data and Long-SAGE results ranges from 60% to 70%, this verified the correctness of Long-SAGE data. These data provides much information to research the pathogenesis of IBDV.
Total RNA was isolated from IBDV infected/uninfected bursa of Fabricius and chicken embryo fibroblast, and 15 differentially expressed genes in Long-SAGE and 17 pro-inflammatory cytokines were selected to perform real-time PCR to analyze the transcript changes. In chicken embryo fibroblast, HMGN2 was down-regulated significantly, it is possible that this gene involved in apoptosis of infected cells; pro-inflammatory cytokines were all up-regulated, IFN-P was up-regulated more than 500 times. Other pro-inflammatory cytokines such as IFN-y, BAFF, IL-6, IL-1B, IL-8 and INOS were up-reguated more than 20 times. In IBDV infected bursa of Fabricius, CLIC4 was up-regulated significantly, it is presumed that CLIC4 involved in virus release; pro-inflammatory cytokines were all up-regulated, IL-6 was up-regulated 1746 times. Other pro-inflammatory cytokines TNFSF8, IFN-β, IFN-y, TNFRSF1A, IL-8 and iNOS were up-reguated more than 20 times. These data indicates that the category of differential transcript genes in different host cells infected with IBDV is coincident, but, the tendency of gene transcript changes is not coincident, thus presents the infection variability of different type cells to IBDV.
Coffilin was down-regulated significantly in IBDV and analyzed the relation of IBDV infection and cofilin expression using anti-cofilin polyclonal antibody, the result indicates that the cells expressing cofilin are susceptive to IBDV. Because that ADF and cofilin have 70% homology and similar function, for differentiating the function of this two genes in IBDV infection, we tried to prepare anti-ADF and anti-cofilin monoclonal antibody, only anti-ADF antibody was obtained. However, ADF is expressed in all cells.
Based on the information of Long-SAGE libraries and real-time PCR, for understanding host feeding back to IBDV VP4 protein, using recombinant expressed VP3 and VP4 proteins, the humoral immunity to VP4 protein was analyzed in 21-day old SPF Leghorn chicken infected respectively with virulent IBDV (NB strain, BC6/85), intermediate plus virulent IBDV (MB43), intermediate virulent IBDV (B87) and attenuated IBDV (attenuated NB strain). The serum antibody was detected by ELISA:in virulent IBDV infected chickens, VP4 antibody seroconverted at 23 days post infection (dpi), seropositivity ratio of VP4 antibody is more than 70% at 40 dpi; VP3-antibody seroconverts at 20 dpi, seropositivity ratio of VP3 antibody is 100% at 23 dpi. In intermediate plus virulent IBDV infected chickens, VP4 antibody seroconverted at 23 dpi, seropositivity ratio of VP4 antibody is less than 55% at 40 dpi; seropositivity ratio of VP3 antibody is 100% at 27 dpi. In intermediate virulent IBDV infected chickens, VP4 antibody seroconverted at 23 dpi, seropositivity ratio of VP4 antibody is less than 37% at 40 dpi; seropositivity ratio of VP3 antibody is 100% at 27 dpi. In attenuated IBDV infected chickens, VP4 antibody was scarcely detectable after first vaccination, seropositivity ratio of VP4 antibody is less than 22% after second vaccination; seropositivity ratio of VP3 antibody was nearly 100% after first vaccination. Similar results were obtained by IFA using Vero cell expressing IBDV VP3 or VP4 protein. Furthermore, immunohistochemistry assay using aniti-VP3/VP4 monoclonal antibodies showed:VP3 and VP4 proteins can detected in bursa of Fabricius infected with virulent IBDV, however, in bursa of Fabricius infected with attenuated IBDV, only VP3 protein can be detected, the VP4 protein of IBDV can scarcely be detected. These data proves that VP4 protein antigen or specific antibodis are biomarker discriminating wild IBDV or vaccine virus infection.
首先,在转录组水平上,本研究分析了IBDV感染及IBDV编码基因转化对宿主Vero细胞基因转录的影响。以实验室已构建的5个Long-SAGE文库：正常Vero细胞文库、IBDV感染的Vero细胞文库、表达A节段编码蛋白的单克隆Vero细胞文库、表达VP243蛋白的单克隆Vero细胞文库、表达VP5蛋白的单克隆Vero细胞文库为基础,对文库中转录显著变化(P<0.05)的基因进行统计分析,发现217个可注释基因的转录发生显著变化。对差异转录基因进行聚类分析,发现蛋白质代谢、能量代谢、mRNA加工和细胞骨架相关基因转录变化最为显著。57个显著变化基因的荧光定量PCR验证结果显示,荧光定量PCR结果与文库数据的符合程度是60%-70%,确证了IBDV感染及IBDV编码基因Vero细胞SAGE文库大规模测序基因标签数据的正确性,这为IBDV的发病机制研究提供了大量的基因生物信息。
以IBDV攻毒后的法氏囊组织和感染的鸡胚成纤维细胞RNA为模板,以IBDV感染Vero细胞的基因转录变化为线索,选取IBDV感染Vero细胞Long-SAGE文库中显著变化的15个基因和17个促炎性因子基因,设计鸡源引物,借助荧光定量PCR方法,分析了IBDV感染的法氏囊组织和鸡胚成纤维细‘胞的转录本变化。在IBDV感染的鸡胚成纤维细胞中,HMGN2的显著下调,推测与病毒感染细胞的细胞凋亡相关；促炎性因子表达普遍上调,其中β干扰素(IFN-p)转录本上调500倍以上,其他促炎性因子如IFN-G、BAFF、IL-6、IL-1B、IL-8和iNOS也上调20倍以上。在感染IBDV的法氏囊组织中,氯离子通道蛋白4(CLIC4)显著表达上调,推测与病毒的破膜释放相关；促炎性因子IL-6的转录上调1746倍。表达上调20倍以上的基因还有TNFSF8、IFN-B、IFN-G、IL-8和iNOS。这些数据说明,IBDV感染导致不同类型宿主细胞基因变化的种类具有一致性,但变化的趋势具有不完全一致性,充分显示了不同类型细胞的易感差异性。
在IBDV感染的法氏囊组织中,我们发现Cofilin基因转录显著上调,为分析Cofilin基因表达与IBDV感染之间的关系,本试验利用抗Cofilin蛋白抗体对IBDV感染的鸡胚成纤维细胞和法氏囊组织进行分析,结果发现IBDV感染能够激活Cofilin基因的表达。
基于IBDV感染宿主细胞的转录本信息,为了了解宿主对IBDV非结构蛋白VP4的信号反馈,以重组表达的VP4、VP3蛋白为抗原,制备了抗VP4、VP3单克隆抗体和多克隆抗体,建立了VP4抗体检测ELISA。分析了IBDV强毒株(NB株,BC6/85株)、中等偏强毒力株(MB43株)、中等毒力株(B87株)、低毒力毒株(NB弱毒株)感染21日龄SPF Leghorn鸡的VP4蛋白的体液免疫应答,ELISA抗体监测结果显示：IBDV强毒感染鸡VP4抗体于感染后23天阳转,40天阳性率达70%以上,VP3抗体于感染后20天阳转,23天阳性率达到100%；IBDV弱毒疫苗一次免疫鸡难以检出VP4抗体,VP3抗体于20天阳转,23天阳性率达到88.9%。中等毒力IBDV感染后,VP4抗体阳性率为10%到40%,中等偏强毒力IBDV感染后VP4抗体阳性率为30%到60%。以IBDV VP3、VP4基因Vero细胞表达蛋白的IFA检测出现了类似的结果。对感染后血清VP4抗体效价测定发现,强毒感染血清VP4抗体效价与中等偏强毒力、中等毒力、弱毒相比较,具有显著差异。抗VP3、VP4单克隆抗体的免疫组织化学检测进一步显示：VP3、VP4抗原可以在IBDV强毒感染鸡法氏囊中检测到,而在弱毒疫苗感染鸡中只能检测到VP3抗原,难以检测到VP4抗原。这些数据证明VP4抗原及抗体是鉴别IBDV野毒与疫苗毒感染的生物标记。
Infectious bursal disease virus (IBDV), a member of the genus Avibirnavirus in the family Birnaviridae, has two serotype, serotype 1 and serotype 2. Serotype 1 viruses are pathogenic for chickens, whereas serotype 2 viruses are avirulent. Genome of IBDV consists of two segments (A and B) of double-stranded RNA (dsRNA) and codes 5 proteins. The large segment A, contains two open reading frames (ORFs). A small ORF encodes the nonstructural protein VP5, which overlaps the N terminal region of a precursor polyprotein from the large ORF. The precursor polyprotein is autoproteolytically cleaved to form mature viral proteins VP2-VP4. Segment B encodes VP1, which has RNA-dependent RNA polymerase (RDRP) activity. Infectious bursal disease (IBD) is an immunosuppression disease; IBD cause the death of infected chicken or vaccination failure of other vaccines for immunosuppression. Today, this disease still threatens poultry industry. The pathogenesis is still unclear. In the present study, the interactions of IBDV and hosts were deeply researched at transcriptomic and humoral levels.
Firstly, at transcriptome level, this study analyzed the impacts of IBDV infection or IBDV coding genes transformation on host transcriptome. Based on 5 Long-SAGE libraries:normal Vero cell, IBDV infected Vero cell, monoclonal Vero cell line expressing A-segment coding proteins, monoclonal Vero cell line expressing VP243 protein, monoclonal Vero cell line expressing VP5 protein, differentially expressed genes were analyzed (p<0.05),217 differentially expressed genes were identified. the influences of different viral genes on Vero cell transcriptome and metabolism were summarized. By classing the differential expression genes, we discovered that the protein metabolism, energy metabolism and cytoskeleton related genes were influenced. Real-time PCR analysis was performed to verify 57 genes expression which were differentially expressed in Long-SAGE library. The coincidence of the real-time PCR data and Long-SAGE results ranges from 60% to 70%, this verified the correctness of Long-SAGE data. These data provides much information to research the pathogenesis of IBDV.
Total RNA was isolated from IBDV infected/uninfected bursa of Fabricius and chicken embryo fibroblast, and 15 differentially expressed genes in Long-SAGE and 17 pro-inflammatory cytokines were selected to perform real-time PCR to analyze the transcript changes. In chicken embryo fibroblast, HMGN2 was down-regulated significantly, it is possible that this gene involved in apoptosis of infected cells; pro-inflammatory cytokines were all up-regulated, IFN-P was up-regulated more than 500 times. Other pro-inflammatory cytokines such as IFN-y, BAFF, IL-6, IL-1B, IL-8 and INOS were up-reguated more than 20 times. In IBDV infected bursa of Fabricius, CLIC4 was up-regulated significantly, it is presumed that CLIC4 involved in virus release; pro-inflammatory cytokines were all up-regulated, IL-6 was up-regulated 1746 times. Other pro-inflammatory cytokines TNFSF8, IFN-β, IFN-y, TNFRSF1A, IL-8 and iNOS were up-reguated more than 20 times. These data indicates that the category of differential transcript genes in different host cells infected with IBDV is coincident, but, the tendency of gene transcript changes is not coincident, thus presents the infection variability of different type cells to IBDV.
Coffilin was down-regulated significantly in IBDV and analyzed the relation of IBDV infection and cofilin expression using anti-cofilin polyclonal antibody, the result indicates that the cells expressing cofilin are susceptive to IBDV. Because that ADF and cofilin have 70% homology and similar function, for differentiating the function of this two genes in IBDV infection, we tried to prepare anti-ADF and anti-cofilin monoclonal antibody, only anti-ADF antibody was obtained. However, ADF is expressed in all cells.
Based on the information of Long-SAGE libraries and real-time PCR, for understanding host feeding back to IBDV VP4 protein, using recombinant expressed VP3 and VP4 proteins, the humoral immunity to VP4 protein was analyzed in 21-day old SPF Leghorn chicken infected respectively with virulent IBDV (NB strain, BC6/85), intermediate plus virulent IBDV (MB43), intermediate virulent IBDV (B87) and attenuated IBDV (attenuated NB strain). The serum antibody was detected by ELISA:in virulent IBDV infected chickens, VP4 antibody seroconverted at 23 days post infection (dpi), seropositivity ratio of VP4 antibody is more than 70% at 40 dpi; VP3-antibody seroconverts at 20 dpi, seropositivity ratio of VP3 antibody is 100% at 23 dpi. In intermediate plus virulent IBDV infected chickens, VP4 antibody seroconverted at 23 dpi, seropositivity ratio of VP4 antibody is less than 55% at 40 dpi; seropositivity ratio of VP3 antibody is 100% at 27 dpi. In intermediate virulent IBDV infected chickens, VP4 antibody seroconverted at 23 dpi, seropositivity ratio of VP4 antibody is less than 37% at 40 dpi; seropositivity ratio of VP3 antibody is 100% at 27 dpi. In attenuated IBDV infected chickens, VP4 antibody was scarcely detectable after first vaccination, seropositivity ratio of VP4 antibody is less than 22% after second vaccination; seropositivity ratio of VP3 antibody was nearly 100% after first vaccination. Similar results were obtained by IFA using Vero cell expressing IBDV VP3 or VP4 protein. Furthermore, immunohistochemistry assay using aniti-VP3/VP4 monoclonal antibodies showed:VP3 and VP4 proteins can detected in bursa of Fabricius infected with virulent IBDV, however, in bursa of Fabricius infected with attenuated IBDV, only VP3 protein can be detected, the VP4 protein of IBDV can scarcely be detected. These data proves that VP4 protein antigen or specific antibodis are biomarker discriminating wild IBDV or vaccine virus infection.
引文
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