家蚕核型多角体病毒分子流行病学调查及宿主特异性研究
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摘要
家蚕核型多角体病毒(Bombyx mori nucleopolyhedrovirus, BmNPV)是家蚕血液型脓病病原,对家蚕幼虫产生致死性的感染,该病是养蚕业常见且危害较大的一种病毒病。广西地处亚热带地区,具有明显的气候优势,过去几年中广西蚕桑业得到迅猛发展,然而,由于养殖密度大,生产期较长,往往容易造成蚕病特别是病毒性传染病的流行。广西几乎每年都会出现BmNPV的流行,而且这些年来由于养殖规模不断扩大以及病原体在养蚕区不断积累,BmNPV流行显示出逐渐加剧的趋势,给广西养蚕业带来了严重的经济损失。为了广西养蚕业的可持续发展,对广西各地BmNPV流行情况进行调查,了解病毒传播特点,将有助于制定相应措施遏制BmNPV在养蚕区扩大化流行。
本研究从广西各养蚕区分别收集了45个野生BmNPV分离毒株,对这些分离株的唯一两个极晚期表达基因多角体蛋白polh基因和微管相关蛋白p10基因进行克隆测序,构建系统进化树。序列分析结果显示,polh基因高度保守,而p10基因变异较大,频繁出现核苷酸点突变,表现出密码子使用多样化,并没有明显的密码子偏好。根据p10基因系统进化树分析,广西BmNPV分离株主要分为三个群clades I、Ⅱ和Ⅲ。广西BmNPV分离株的地理分布显示,clade I(?)口Ⅱ分群中的分离株来源地域较为集中而且毗邻,提示在这些区域内出现局部的流行与传播。cladeⅢ的分离株呈不规律分布于广西各地,提示有可能发生远距离的病毒传播。
广西桑蚕资源丰富,蚕虫或者蚕蛹可以作为一种理想的生物反应器,利用杆状病毒表达系统在其中大量生产重组蛋白,不仅省时省力而且经济节约。狂犬病病毒糖蛋白(Rabies virus glycoprotein, RVG)是主要保护性抗原,能诱导机体产生中和性抗体。本研究使用了一个经过基因工程改良的杆状病毒表达系统,即与BmNPV基因组同源的bacmid缺失了几丁质酶和半胱氨酸酶基因,构建了携带RVG基因的重组bacmid,同时引入在哺乳动物细胞内具有转录活性的巨细胞瘤病毒启动子／增强子CMV-IE序列。将重组bacmid与脂质体混合注射家蚕幼虫血腔,进行体内转染获得重组杆状病毒。鉴定结果显示,重组RVG在蚕虫机体组织中获得大量表达,重组表达的RVG最终定位于细胞质膜,重组RVG分子量略小于在哺乳动物细胞中表达的天然RVG。但是,多个抗RVG单克隆抗体皆与之发生反应,且反应的程度与天然RVG的基本一致,提示重组RVG的结构和功能是完好的。
针对BmNPV宿主特异性研究表明,无论是体内还是体外BmNPV的宿主域相当狭窄。草地夜蛾(Spodoptera frugiperda, Sf)细胞系Sf9一直被认为是BmNPV的非受纳细胞系,BmNPV不能在其中进行有效的复制增殖。本研究中,两个携带有外源报告基因的重组BmNPV,即携带狂犬病病毒RVG基因及巨细胞瘤病毒启动子CMV-IE序列的rBm-PCMV-RVG和携带绿色荧光蛋白GFP基因的rBm-GFP,被构建并表现出对Sf9细胞具有感染性。该重组BmNPV可以在Sf9细胞中复制增殖,被感染的Sf9细胞无明显细胞病变,显示一种无症状温和型的感染状态,并且在被感染的Sf9细胞上可以检测到由低pH值引发的膜融合反应。以重组rBm-GFP接种Sf9细胞,GFP荧光灶形成和子代病毒滴度增加证实了病毒可以从Sf9细胞中出芽。然而,只有在接种病毒量较大时,重组BmNPV才能在Sf9细胞中复制增殖,且增殖过程与其在受纳家蚕细胞系BmN中增殖过程不同；而当接种病毒量较小时,重组BmNPV在Sf9细胞中产生顿挫感染。另外,重组rBm-GFP在Sf9细胞中经过系列传代后显示出比传代前的原始病毒复制能力更强。
根据GenBank发表的家蚕抗BmNPV病毒脂肪酶基因Bmlipase-1和丝氨酸蛋白酶基因BmSP-2序列,分别设计相应引物,以广西优良家蚕品系“两广二号”为实验材料,成功克隆了该品系家蚕两个抗病毒基因的完整开放阅读框(open reading frame, ORF),测序并与不同蚕来源的同源基因序列进行比较。实验结果显示：“两广二号”家蚕Bmlipase-1基因ORF长度是885bp,编码294个氨基酸,BmSP-2基因ORF长度是855bp,编码284个氨基酸；同源性比较表明二者与其他品系家蚕甚至野蚕和蓖麻蚕的同源序列高度保守,核苷酸和推导氨基酸序列的同源性皆在92%以上,其中Bmlipase-1更为保守,同源性达到99%以上；序列分析发现各种蚕来源的Bmlipase-1基因脂肪酶活化部位(GXSXG,X为任一氨基酸)的氨基酸序列完全一致,BmSP-2基因酶催化三联体(His95, Asp142,Ser236)位点及附近氨基酸序列也完全相同。以上结果说明这两个抗病毒基因在桑蚕遗传进化过程中保持高度一致,提示其可能在机体消化或者免疫防御方面起着重要生理作用。将这两个抗病毒基因连接到原核表达载体pET32a(+),在大肠杆菌BL21中融合表达目的蛋白,SDS-PAGE和Western blot结果显示：这两个抗病毒基因都在大肠杆菌中表达成功,融合表达的Bmlipase-1蛋白分子量约为47kD, BmSP-2的分子量约为42kD,与预期大小一致。
Bombyx mori nucleopolyhedrovirus (BmNPV) is a severe pathogen that lethally infects the Bombyx mori silkworm. BmNPV disease is a common but quite destructive viral disease in sericulture. Guangxi is located in southern China, where the subtropical climate is perfect for mulberry cultivation and silkworm husbandry, and the sericulture industry in Guangxi has been developed extensively during the past few years. However, due to the high raising density and long production period, the prevalence of silkworm diseases especially viral epidemics are likely to happen. The BmNPV epidemic takes place almost every year in Guangxi, and recently becomes more serious resulting from the large-scale development of sericulture and the accumulation of the pathogen in the silkworm-raising regions in Guangxi. This intractable disease has been caused considerable economic losses in Guangxi sericulture industry. In order to keep sustainable development of sericulture industry in Guangxi, it is essential to perform an investigation on the epidemic condition of BmNPV in Guangxi and make a better understanding of the BmNPV transmission, which will help to devise strategies for preventing this epidemic disease to spread widely in the silkworm-raising regions.
In this study,45wild BmNPV isolates were collected from different silkworm-raising regions in China's Guangxi Zhuang Autonomous Region. Two highly expressed very late genes from each isolate, polh and p10, were sequenced and subjected to phylogenetic analysis. The polh gene was found to be highly conserved, while the p10gene was more variable frequently harboring point mutations and displaying variations in codon use without obvious codon bias. The BmNPV isolates from Guangxi were separated into three main clades, Ⅰ, Ⅱ and Ⅲ,according to the p10gene phylogenetic tree. The geographical distribution of clade Ⅰ isolates in Guangxi showed a concentrated pattern and that of clade Ⅱ isolates showed a connected pattern. Local transmission of this pathogen clearly occurred in the silkworm-raising regions in Guangxi. And clade Ⅲ isolates were irregularly scattered throughout Guangxi, which suggested long-distance transmission may happen. This study may provide some data on BmNPV transmission in the silkworm-raising regions and be helpful in devising strategies for the prevention and control of BmNPV disease.
There is quite a rich resource of silkworm production in Guangxi. Silkworm serves as an ideal bioreactor for abundantly producing recombinant protein by using a baculovirus expression system. Rabies virus glycoprotein (RVG) is the major protective antigen which can induce neutralizing antibody against lethal rabies virus infection. In this study, the complete open reading frame (ORF) of RVG gene was cloned and inserted into a modified bacmid/BmNPV deleting both chitinase and cysteine-protease genes. A mammalian cell-activate promoter cytomegalovirus immediate-early promoter-enhancer (CMV-IE) was also constructed into the bacmid. The recombinant baculovirus was generated by syringe injection of the silkworm larvae hemocoel with the recombinant bacmid and Cellfectin Reagent mixture. Results showed that recombinant RVG was highly expressed in the tissues of recombinant baculovirus infected silkworm larvae. The recombinant RVG was displayed on the membrane surface and its molecular weight was slightly smaller than that of the natural RVG expressed in mammalian cells. But the reacting patterns of anti-RVG monoclonal antibodies with the recombinant RVG were almost the same as those with the natural RVG, suggested that the structure and function of the recombinant RVG were intact.
Studies on BmNPV host specificity have shown that this virus possess relatively narrow host ranges both in vivo and in vitro. Spodoptera frugiperda cell line Sf9was classically considered to be nonpermissive for BmNPV proliferation. In this study, two recombinant BmNPV carrying exogenous reporter gene, the recombinant rBm-PCMV-RVG carrying RVG gene and CMV-IE promoter sequence and the recombinant rBm-GFP carrying green fluorescent protein (GFP) gene, were constructed and showed infectivity in Sf9cells. There was no obvious cytopathic effect in the Sf9cells infected with recombinant BmNPV, showing an inapparent and mild pattern of infection. Low-pH-triggered envelope fusion was detected in the infected Sf9cells. When inoculating Sf9cells with rBm-GFP, virus budding was confirmed by the formation of GFP fluorescent focus and the increase of budded virion yield in Sf9cells. However, the viral replication only occured with high inoculation dose and showed a pattern different from that in permissive BmN cells. To the contrary, low inoculation dose led to abortive infection. The rBm-GFP after serial passages in Sf9cells showed more efficient replication than the original virus without serial passage.
According to the sequences of Bombyx mori silkworm antiviral protein genes Bmlipase-1(lipase) and BmSP-2(serine protease) published in Genbank, specific primers were designed to clone the complete ORF of the two genes from Guangxi excellent silkworm race "Liangguang NO.2". The cloned products were sequenced and analyzed with the homologs derived from other silkworm species. The results show that Bmlipase-1is composed of885bp nucleotides coding294amino acids and BmSP-2is composed of855bp nucleotides coding284amino acids. Sequence analysis reveal that the two genes from "Liangguang NO.2" shared more than92%homology with those from different Bombyx mori races even Bombyx mandarina and Samia cynthia ricini, especially for Bmlipase-1which reached more than99%homology. Amino acid residues for the lipase active site of Bmlipase-1and serine protease catalytic triad of BmSP-2are identical among these gene sequences. The data above suggested that these two antiviral genes are highly conserved during silkworm species genetic evolution and may play important roles in immune defence as well as food digestion. Then the cloned genes were ligated into protokaryon expression vector pET32a(+) and expressed in E.coli BL21. Fused proteins were detected by SDS-PAGE and Western blot. According to the result, the protein mass of expressed Bmlipase-1is about47kD and BmSP-2is about42kD, which consisted with the anticipation.
本研究从广西各养蚕区分别收集了45个野生BmNPV分离毒株,对这些分离株的唯一两个极晚期表达基因多角体蛋白polh基因和微管相关蛋白p10基因进行克隆测序,构建系统进化树。序列分析结果显示,polh基因高度保守,而p10基因变异较大,频繁出现核苷酸点突变,表现出密码子使用多样化,并没有明显的密码子偏好。根据p10基因系统进化树分析,广西BmNPV分离株主要分为三个群clades I、Ⅱ和Ⅲ。广西BmNPV分离株的地理分布显示,clade I(?)口Ⅱ分群中的分离株来源地域较为集中而且毗邻,提示在这些区域内出现局部的流行与传播。cladeⅢ的分离株呈不规律分布于广西各地,提示有可能发生远距离的病毒传播。
广西桑蚕资源丰富,蚕虫或者蚕蛹可以作为一种理想的生物反应器,利用杆状病毒表达系统在其中大量生产重组蛋白,不仅省时省力而且经济节约。狂犬病病毒糖蛋白(Rabies virus glycoprotein, RVG)是主要保护性抗原,能诱导机体产生中和性抗体。本研究使用了一个经过基因工程改良的杆状病毒表达系统,即与BmNPV基因组同源的bacmid缺失了几丁质酶和半胱氨酸酶基因,构建了携带RVG基因的重组bacmid,同时引入在哺乳动物细胞内具有转录活性的巨细胞瘤病毒启动子／增强子CMV-IE序列。将重组bacmid与脂质体混合注射家蚕幼虫血腔,进行体内转染获得重组杆状病毒。鉴定结果显示,重组RVG在蚕虫机体组织中获得大量表达,重组表达的RVG最终定位于细胞质膜,重组RVG分子量略小于在哺乳动物细胞中表达的天然RVG。但是,多个抗RVG单克隆抗体皆与之发生反应,且反应的程度与天然RVG的基本一致,提示重组RVG的结构和功能是完好的。
针对BmNPV宿主特异性研究表明,无论是体内还是体外BmNPV的宿主域相当狭窄。草地夜蛾(Spodoptera frugiperda, Sf)细胞系Sf9一直被认为是BmNPV的非受纳细胞系,BmNPV不能在其中进行有效的复制增殖。本研究中,两个携带有外源报告基因的重组BmNPV,即携带狂犬病病毒RVG基因及巨细胞瘤病毒启动子CMV-IE序列的rBm-PCMV-RVG和携带绿色荧光蛋白GFP基因的rBm-GFP,被构建并表现出对Sf9细胞具有感染性。该重组BmNPV可以在Sf9细胞中复制增殖,被感染的Sf9细胞无明显细胞病变,显示一种无症状温和型的感染状态,并且在被感染的Sf9细胞上可以检测到由低pH值引发的膜融合反应。以重组rBm-GFP接种Sf9细胞,GFP荧光灶形成和子代病毒滴度增加证实了病毒可以从Sf9细胞中出芽。然而,只有在接种病毒量较大时,重组BmNPV才能在Sf9细胞中复制增殖,且增殖过程与其在受纳家蚕细胞系BmN中增殖过程不同；而当接种病毒量较小时,重组BmNPV在Sf9细胞中产生顿挫感染。另外,重组rBm-GFP在Sf9细胞中经过系列传代后显示出比传代前的原始病毒复制能力更强。
根据GenBank发表的家蚕抗BmNPV病毒脂肪酶基因Bmlipase-1和丝氨酸蛋白酶基因BmSP-2序列,分别设计相应引物,以广西优良家蚕品系“两广二号”为实验材料,成功克隆了该品系家蚕两个抗病毒基因的完整开放阅读框(open reading frame, ORF),测序并与不同蚕来源的同源基因序列进行比较。实验结果显示：“两广二号”家蚕Bmlipase-1基因ORF长度是885bp,编码294个氨基酸,BmSP-2基因ORF长度是855bp,编码284个氨基酸；同源性比较表明二者与其他品系家蚕甚至野蚕和蓖麻蚕的同源序列高度保守,核苷酸和推导氨基酸序列的同源性皆在92%以上,其中Bmlipase-1更为保守,同源性达到99%以上；序列分析发现各种蚕来源的Bmlipase-1基因脂肪酶活化部位(GXSXG,X为任一氨基酸)的氨基酸序列完全一致,BmSP-2基因酶催化三联体(His95, Asp142,Ser236)位点及附近氨基酸序列也完全相同。以上结果说明这两个抗病毒基因在桑蚕遗传进化过程中保持高度一致,提示其可能在机体消化或者免疫防御方面起着重要生理作用。将这两个抗病毒基因连接到原核表达载体pET32a(+),在大肠杆菌BL21中融合表达目的蛋白,SDS-PAGE和Western blot结果显示：这两个抗病毒基因都在大肠杆菌中表达成功,融合表达的Bmlipase-1蛋白分子量约为47kD, BmSP-2的分子量约为42kD,与预期大小一致。
Bombyx mori nucleopolyhedrovirus (BmNPV) is a severe pathogen that lethally infects the Bombyx mori silkworm. BmNPV disease is a common but quite destructive viral disease in sericulture. Guangxi is located in southern China, where the subtropical climate is perfect for mulberry cultivation and silkworm husbandry, and the sericulture industry in Guangxi has been developed extensively during the past few years. However, due to the high raising density and long production period, the prevalence of silkworm diseases especially viral epidemics are likely to happen. The BmNPV epidemic takes place almost every year in Guangxi, and recently becomes more serious resulting from the large-scale development of sericulture and the accumulation of the pathogen in the silkworm-raising regions in Guangxi. This intractable disease has been caused considerable economic losses in Guangxi sericulture industry. In order to keep sustainable development of sericulture industry in Guangxi, it is essential to perform an investigation on the epidemic condition of BmNPV in Guangxi and make a better understanding of the BmNPV transmission, which will help to devise strategies for preventing this epidemic disease to spread widely in the silkworm-raising regions.
In this study,45wild BmNPV isolates were collected from different silkworm-raising regions in China's Guangxi Zhuang Autonomous Region. Two highly expressed very late genes from each isolate, polh and p10, were sequenced and subjected to phylogenetic analysis. The polh gene was found to be highly conserved, while the p10gene was more variable frequently harboring point mutations and displaying variations in codon use without obvious codon bias. The BmNPV isolates from Guangxi were separated into three main clades, Ⅰ, Ⅱ and Ⅲ,according to the p10gene phylogenetic tree. The geographical distribution of clade Ⅰ isolates in Guangxi showed a concentrated pattern and that of clade Ⅱ isolates showed a connected pattern. Local transmission of this pathogen clearly occurred in the silkworm-raising regions in Guangxi. And clade Ⅲ isolates were irregularly scattered throughout Guangxi, which suggested long-distance transmission may happen. This study may provide some data on BmNPV transmission in the silkworm-raising regions and be helpful in devising strategies for the prevention and control of BmNPV disease.
There is quite a rich resource of silkworm production in Guangxi. Silkworm serves as an ideal bioreactor for abundantly producing recombinant protein by using a baculovirus expression system. Rabies virus glycoprotein (RVG) is the major protective antigen which can induce neutralizing antibody against lethal rabies virus infection. In this study, the complete open reading frame (ORF) of RVG gene was cloned and inserted into a modified bacmid/BmNPV deleting both chitinase and cysteine-protease genes. A mammalian cell-activate promoter cytomegalovirus immediate-early promoter-enhancer (CMV-IE) was also constructed into the bacmid. The recombinant baculovirus was generated by syringe injection of the silkworm larvae hemocoel with the recombinant bacmid and Cellfectin Reagent mixture. Results showed that recombinant RVG was highly expressed in the tissues of recombinant baculovirus infected silkworm larvae. The recombinant RVG was displayed on the membrane surface and its molecular weight was slightly smaller than that of the natural RVG expressed in mammalian cells. But the reacting patterns of anti-RVG monoclonal antibodies with the recombinant RVG were almost the same as those with the natural RVG, suggested that the structure and function of the recombinant RVG were intact.
Studies on BmNPV host specificity have shown that this virus possess relatively narrow host ranges both in vivo and in vitro. Spodoptera frugiperda cell line Sf9was classically considered to be nonpermissive for BmNPV proliferation. In this study, two recombinant BmNPV carrying exogenous reporter gene, the recombinant rBm-PCMV-RVG carrying RVG gene and CMV-IE promoter sequence and the recombinant rBm-GFP carrying green fluorescent protein (GFP) gene, were constructed and showed infectivity in Sf9cells. There was no obvious cytopathic effect in the Sf9cells infected with recombinant BmNPV, showing an inapparent and mild pattern of infection. Low-pH-triggered envelope fusion was detected in the infected Sf9cells. When inoculating Sf9cells with rBm-GFP, virus budding was confirmed by the formation of GFP fluorescent focus and the increase of budded virion yield in Sf9cells. However, the viral replication only occured with high inoculation dose and showed a pattern different from that in permissive BmN cells. To the contrary, low inoculation dose led to abortive infection. The rBm-GFP after serial passages in Sf9cells showed more efficient replication than the original virus without serial passage.
According to the sequences of Bombyx mori silkworm antiviral protein genes Bmlipase-1(lipase) and BmSP-2(serine protease) published in Genbank, specific primers were designed to clone the complete ORF of the two genes from Guangxi excellent silkworm race "Liangguang NO.2". The cloned products were sequenced and analyzed with the homologs derived from other silkworm species. The results show that Bmlipase-1is composed of885bp nucleotides coding294amino acids and BmSP-2is composed of855bp nucleotides coding284amino acids. Sequence analysis reveal that the two genes from "Liangguang NO.2" shared more than92%homology with those from different Bombyx mori races even Bombyx mandarina and Samia cynthia ricini, especially for Bmlipase-1which reached more than99%homology. Amino acid residues for the lipase active site of Bmlipase-1and serine protease catalytic triad of BmSP-2are identical among these gene sequences. The data above suggested that these two antiviral genes are highly conserved during silkworm species genetic evolution and may play important roles in immune defence as well as food digestion. Then the cloned genes were ligated into protokaryon expression vector pET32a(+) and expressed in E.coli BL21. Fused proteins were detected by SDS-PAGE and Western blot. According to the result, the protein mass of expressed Bmlipase-1is about47kD and BmSP-2is about42kD, which consisted with the anticipation.
引文
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