野鸟禽流感监测及哺乳动物跨种传播能力评估
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摘要
禽流感(Avian Influenza, A1)是由甲型流感病毒所引起的各种家禽及野生禽类感染疾病综合征。根据致病性不同可划分为高致病性禽流感和低致病性禽流感。近儿年来,屡有禽流感病毒特别是低致病性禽流感突破种间屏障作用,直接感染人类或其它哺乳动物,甚至致人死亡事件的情况发生。野鸟作为储存宿主在禽流感病毒传播中的作用一直受到高度关注：H5N1亚型禽流感随野鸟迁徙在不到两年的时间里传播到14个国家40多个地区；感染人的H7N9亚型禽流感病毒致少有2个基因节片源于野鸟携带病毒的变异；导致江西感染病例的人H10N8亚型禽流感病毒来自于野鸟。这些事件说明禽流感特别是高致病性禽流感已经严重危及到人类的健康与生命安全,并有进一步突破种间屏障引发人间流感大流行的潜在可能。在野生禽流感监测中,方面,发达国家己将野鸟禽流感监测列入疫病监测中的常态化监测工作。中国林业局自2008年起开展了主动监测工作。因野鸟采样极为困难,国家投入相对较少,我国对野鸟禽流感的监测工作还相对落后,如2013年在发生H7N9人感染事件前,周边的日韩等国均已在野鸭中发生该病毒的前体病毒,而我国缺少相关毒株的监测,无法有效溯源。另一方面,H5、H6、H7、H9、H10等多亚型禽源病毒向人类跨种传播的现象提示我们可能有更多亚型的禽流感病毒会对人类健康构成威胁,因此,深入了解作为储存宿主的野鸟中禽流感病毒的分布情况和野鸟源病毒对哺乳动物的跨种传播作用,对于加深我们对流感病毒的认知,做好禽流感的防控工作有着重要的意义。
本研究采集了14省市48个地区的15152份野鸟样品,进行了流感病毒的分离鉴定,分离得到57株病毒,进行一步分析表明野鸟流感病毒携带率约为0.38%,秋季样品分离率高于春季。拭子样品分离率约为0.42%,粪便(环境)样品分离率为0.40%,组织样品分离率为0.18%。分离的57株病毒包括8种HA和7种NA亚型,其中H3、H4亚型为优势流行毒株,但野鸟携带高致病性H5亚型禽流感在病毒的比例较高(17.5%),说明防控形势依然严峻。为进一步了解这些毒株的起源及与国内流行株(包括人、猪流行株)之间的关系,本研究对30株代表病毒进行了全基因测序。HA基因进化分析表明,除H3亚型具有宿主特异性外,其余亚型均存在禽、猪或禽、人或三者基因重组现象。NA基因进化关系更加复杂,但与HA进化关系无协同或一致性。相比于外部基因,内部基因(PB2、PB、PA、NP、M、NS)进化关系更是错综复杂,均存在禽、猪、人宿主间基因节片交换现象。以上结果表明野鸟是流感病毒天然的贮存库和基因库,为不同亚型或不同宿主的流感病毒提供基因片段。此外,首次在国内野鸟中分离到H13N6和H5N8病毒,提示进一步加强候鸟监测工作将有助于提前了解可能进入家禽中的病毒。
为了解这些病毒对哺乳动物的致病性,本研究以小鼠为模型对28株病毒进行感染与致病能力研究。所用用的28株病毒均能在小鼠的肺脏、鼻甲骨中复制,但病毒滴度差别较大,以H5N8最高,可达105.7EID50/ml。 H5N1、H5N8、H4N6、H3N2、H7、H9N2业型毒株对小鼠均有不同程度的致死性,其中H5N8的致死率达100%,其余H3和H6亚型的病毒虽然能使小鼠感染,但并无特别的临床特征。本结果提示我们上述源于野鸟的病毒已不需要适应就可能直接感染哺乳动物。
为进一步评估毒株在哺乳动物间水平传播的能力,分别进行了受体结合特性分析和豚鼠间水平传播能力评估。受体结合特性测定结果表明H3业型病毒均具有SAα-2,6Gal(人样流感病毒受体)结合特性,H7、H9亚型也具有一定的SAα-2,6Gal结合特性。本研究选取H3和H9亚型毒株在豚鼠间进行了接触传播研究,结果表明S89(H3N2)、ZH47(H3N3)和JL-2(H9N2)均具有一定的豚鼠间传播能力,其中以S89(H3N2)传播能力最高(2/3)。利用反向遗传学技术分析了决定S89(H3N2)与SAα-2,6Gal结合的关键位点,发现L274I影响病毒受体结合特性,表明除已发现的系列位点外,受体结合区域的其它位点同样会影响病毒受体识别能力。
为进一步分析野鸟、野生动物乃致一些家养动物携带病原体的背景,我们建立了病毒性病原体宏基因组学分析方法,结果表明,该方法无样品类型特殊性差别,对环境、组织样品均普遍适用。应用该方法首次对野鸟体内病毒种类及其丰度进行了研究,检测到多种病毒(鹅圆环病毒、雉疱疹病毒、鸭乙型肝炎病毒、鹤乙型肝炎病毒、禽流感病毒等)。在对突发大规模死亡的家养猪样品分析中发现猪流行性腹泻病毒和新型猪嵴病毒。吉林省鼠出血热监测表明,宿主动物携带的汉坦病毒仍以汉城病毒为主,其遗传进化呈现多样性。这些研究表明该方法对新型或未知病毒性病原体分析具有重要应用价值。
Avian Influenza is a variety infection disease syndrome of poultry and wild birds caused by influenza A virus. Avian influenza virus can be divided into highly pathogenic and low pathogenic avian influenza virus according to its pathogenic in avian. In recent years, avian influenza virus especially low pathogenic avian influenza virus frequent crossed the species barrier, directly infection of humans or other mammals, even causing death event in human. The role of wild birds as reservoir host in the spread of the virus has been highly gained concerns. H5N1subtype avian influenza has spread across14countries and more than40areas along with migratory wild birds in less than two years. At least two genes of human infection H7N9subtype avian influenza virus come from wild bird virus variation. The genes of caused human infection H10N8avian influenza virus in Jiangxi are also originated from wild birds. Avian influenza especially highly pathogenic avian influenza has seriously endangered to human beings, and has the potential to break through the species barrier and caused next influenza pandemic. On the one hand, the developed countries have the wild bird influenza monitoring included in the normal monitoring disease surveillance. Chinese forestry bureau have carried out active surveillance since2008. However, the monitoring work of avian influenza in wild birds in China is relatively backward because of the sampling is extremely difficult from wild bird; especially the national investment is relatively limited. For example, before the event H7N9infection in2013, precursor virus have been detected in the wild duck by our neighboring countries such as Japan and South Korea, but our traceability was not effective because of lack monitoring related strains. On the other hand, the phenomenon of H5, H6, H7, H9, H10subtype avian viruses infection human suggested that more subtype of avian influenza virus would pose a threat to human health. Therefore, it is necessary to in-depth understanding of the influenza virus distribution among wild birds and mammalian cross species transmission, not only for deepen our understanding of the influenza virus, but has an important significance to avian influenza prevention and control.
In this study,15152samples were collected from48regions of14provinces, and then isolation and identification of influenza virus were performed.57strains of avian influenza virus were obtained from wild birds in this survey, the influenza virus positive rate is about0.38%, and the rate of autumn sample is higher than its in spring. Swab sample isolation rate is about0.42%, feces (environment) sample isolation rate is0.40%, and tissue sample isolation rate is0.18%.57strains of virus containing8HA and7NA subtypes, H3, H4subtype is the predominant strains, but highly pathogenic avian influenza H5subtype is also distributed to a wide range in wild birds, accounting for17.5%of the overall isolation of virus, the situation of prevention and control of this disease still grim. In order to understand the relationship between the origin of these strains and the domestic epidemic strains (including human, porcine epidemic strain),30strains was selected for study by whole genome sequencing. Phylogenetic analysis of HA gene showed that, except for the H3subtype with host specificity in avian; the other subtypes were originated from bird, pig or poultry, or all three gene recombination. NA gene evolution was more complex, but no synergistic or consistency with HA in relation to the evolution. Compared to the external gene, the evolutionary relationship of genes (PB2, PB1, PA, NP, M, and NS) was more complex, the phenomenon of gene segments recombination of influenza virus from bird, pig, human host was also observed. The results indicated that wild bird is a reservoir host and natural genes bank of influenza virus genes, providing gene segments for different subtypes and different hosts. In addition, there is need to further strengthen the monitoring of migratory birds because of first isolation H13N6and H5N8virus in wild birds.
In order to understand the pathogenicity of these viruses to mammals, mice were selected as model to evaluate the characterization of infection and pathogenic of28representative virus strains isolated from wild birds. The selected viruses can be replicated in mouse lungs, nasal turbinates, but the virus titer different, the H5N8subtype is the highest, up to105.7EID50/ml. The H3N2, H5N1, H5N8, H4N6, H7, H9N2subtype strains can cause mice death, the mortality of H5N8reached up to100%, the rest of the H3and H6subtypes of the virus can make mice infected without clinical special features. These results showed that the sourced wild bird virus may directly infect humans without adaption. The receptor binding property and horizontal transmission between guinea pigs were analyzed to evaluate transmission in mammals. Receptor binding assay showed that the H3subtype virus has SA a-2,6Gal (human-like influenza virus receptor) binding property, H7, H9subtypes has SA α-2,6Gal binding property in some degree. H3and H9subtypes strains were selected for contacted transmission in guinea pigs, the S89(H3N2), ZH47(H3N3) and JL-2(H9N2) can transmit in guinea pigs, in which S89(H3N2) was the highest (2/3). The recombinant rS89strains were constructed by reverse genetics technology and found that rS89-HA-L2741can influence the S89virus SA α-2,6Gal receptor binding characteristics. These results showed that except the series of sites have been found; other sites receptor binding may also affect the recognition ability of virus receptor.
We established viral metagenomic to analyze the pathogen of wild birds, wild domestic mammalian. The method is universal for different type samples, such as environment, tissue, or swab samples. This method was first used to study viruses and distribution of wild bird, and detect a variety of virus (goose circovirus, pheasant herpes virus, duck hepatitis B virus, crane, hepatitis B virus, avian influenza virus etc.). Porcine epidemic diarrhea virus and porcine novel kobuvirus were found in outbreak of pig death. Most of Hantaviruses carried by animal hosts in Jilin Province were Seoul viruses, of which the genetics showed a diversity. These results showed that the method is important to new or unknown viral pathogens.
本研究采集了14省市48个地区的15152份野鸟样品,进行了流感病毒的分离鉴定,分离得到57株病毒,进行一步分析表明野鸟流感病毒携带率约为0.38%,秋季样品分离率高于春季。拭子样品分离率约为0.42%,粪便(环境)样品分离率为0.40%,组织样品分离率为0.18%。分离的57株病毒包括8种HA和7种NA亚型,其中H3、H4亚型为优势流行毒株,但野鸟携带高致病性H5亚型禽流感在病毒的比例较高(17.5%),说明防控形势依然严峻。为进一步了解这些毒株的起源及与国内流行株(包括人、猪流行株)之间的关系,本研究对30株代表病毒进行了全基因测序。HA基因进化分析表明,除H3亚型具有宿主特异性外,其余亚型均存在禽、猪或禽、人或三者基因重组现象。NA基因进化关系更加复杂,但与HA进化关系无协同或一致性。相比于外部基因,内部基因(PB2、PB、PA、NP、M、NS)进化关系更是错综复杂,均存在禽、猪、人宿主间基因节片交换现象。以上结果表明野鸟是流感病毒天然的贮存库和基因库,为不同亚型或不同宿主的流感病毒提供基因片段。此外,首次在国内野鸟中分离到H13N6和H5N8病毒,提示进一步加强候鸟监测工作将有助于提前了解可能进入家禽中的病毒。
为了解这些病毒对哺乳动物的致病性,本研究以小鼠为模型对28株病毒进行感染与致病能力研究。所用用的28株病毒均能在小鼠的肺脏、鼻甲骨中复制,但病毒滴度差别较大,以H5N8最高,可达105.7EID50/ml。 H5N1、H5N8、H4N6、H3N2、H7、H9N2业型毒株对小鼠均有不同程度的致死性,其中H5N8的致死率达100%,其余H3和H6亚型的病毒虽然能使小鼠感染,但并无特别的临床特征。本结果提示我们上述源于野鸟的病毒已不需要适应就可能直接感染哺乳动物。
为进一步评估毒株在哺乳动物间水平传播的能力,分别进行了受体结合特性分析和豚鼠间水平传播能力评估。受体结合特性测定结果表明H3业型病毒均具有SAα-2,6Gal(人样流感病毒受体)结合特性,H7、H9亚型也具有一定的SAα-2,6Gal结合特性。本研究选取H3和H9亚型毒株在豚鼠间进行了接触传播研究,结果表明S89(H3N2)、ZH47(H3N3)和JL-2(H9N2)均具有一定的豚鼠间传播能力,其中以S89(H3N2)传播能力最高(2/3)。利用反向遗传学技术分析了决定S89(H3N2)与SAα-2,6Gal结合的关键位点,发现L274I影响病毒受体结合特性,表明除已发现的系列位点外,受体结合区域的其它位点同样会影响病毒受体识别能力。
为进一步分析野鸟、野生动物乃致一些家养动物携带病原体的背景,我们建立了病毒性病原体宏基因组学分析方法,结果表明,该方法无样品类型特殊性差别,对环境、组织样品均普遍适用。应用该方法首次对野鸟体内病毒种类及其丰度进行了研究,检测到多种病毒(鹅圆环病毒、雉疱疹病毒、鸭乙型肝炎病毒、鹤乙型肝炎病毒、禽流感病毒等)。在对突发大规模死亡的家养猪样品分析中发现猪流行性腹泻病毒和新型猪嵴病毒。吉林省鼠出血热监测表明,宿主动物携带的汉坦病毒仍以汉城病毒为主,其遗传进化呈现多样性。这些研究表明该方法对新型或未知病毒性病原体分析具有重要应用价值。
Avian Influenza is a variety infection disease syndrome of poultry and wild birds caused by influenza A virus. Avian influenza virus can be divided into highly pathogenic and low pathogenic avian influenza virus according to its pathogenic in avian. In recent years, avian influenza virus especially low pathogenic avian influenza virus frequent crossed the species barrier, directly infection of humans or other mammals, even causing death event in human. The role of wild birds as reservoir host in the spread of the virus has been highly gained concerns. H5N1subtype avian influenza has spread across14countries and more than40areas along with migratory wild birds in less than two years. At least two genes of human infection H7N9subtype avian influenza virus come from wild bird virus variation. The genes of caused human infection H10N8avian influenza virus in Jiangxi are also originated from wild birds. Avian influenza especially highly pathogenic avian influenza has seriously endangered to human beings, and has the potential to break through the species barrier and caused next influenza pandemic. On the one hand, the developed countries have the wild bird influenza monitoring included in the normal monitoring disease surveillance. Chinese forestry bureau have carried out active surveillance since2008. However, the monitoring work of avian influenza in wild birds in China is relatively backward because of the sampling is extremely difficult from wild bird; especially the national investment is relatively limited. For example, before the event H7N9infection in2013, precursor virus have been detected in the wild duck by our neighboring countries such as Japan and South Korea, but our traceability was not effective because of lack monitoring related strains. On the other hand, the phenomenon of H5, H6, H7, H9, H10subtype avian viruses infection human suggested that more subtype of avian influenza virus would pose a threat to human health. Therefore, it is necessary to in-depth understanding of the influenza virus distribution among wild birds and mammalian cross species transmission, not only for deepen our understanding of the influenza virus, but has an important significance to avian influenza prevention and control.
In this study,15152samples were collected from48regions of14provinces, and then isolation and identification of influenza virus were performed.57strains of avian influenza virus were obtained from wild birds in this survey, the influenza virus positive rate is about0.38%, and the rate of autumn sample is higher than its in spring. Swab sample isolation rate is about0.42%, feces (environment) sample isolation rate is0.40%, and tissue sample isolation rate is0.18%.57strains of virus containing8HA and7NA subtypes, H3, H4subtype is the predominant strains, but highly pathogenic avian influenza H5subtype is also distributed to a wide range in wild birds, accounting for17.5%of the overall isolation of virus, the situation of prevention and control of this disease still grim. In order to understand the relationship between the origin of these strains and the domestic epidemic strains (including human, porcine epidemic strain),30strains was selected for study by whole genome sequencing. Phylogenetic analysis of HA gene showed that, except for the H3subtype with host specificity in avian; the other subtypes were originated from bird, pig or poultry, or all three gene recombination. NA gene evolution was more complex, but no synergistic or consistency with HA in relation to the evolution. Compared to the external gene, the evolutionary relationship of genes (PB2, PB1, PA, NP, M, and NS) was more complex, the phenomenon of gene segments recombination of influenza virus from bird, pig, human host was also observed. The results indicated that wild bird is a reservoir host and natural genes bank of influenza virus genes, providing gene segments for different subtypes and different hosts. In addition, there is need to further strengthen the monitoring of migratory birds because of first isolation H13N6and H5N8virus in wild birds.
In order to understand the pathogenicity of these viruses to mammals, mice were selected as model to evaluate the characterization of infection and pathogenic of28representative virus strains isolated from wild birds. The selected viruses can be replicated in mouse lungs, nasal turbinates, but the virus titer different, the H5N8subtype is the highest, up to105.7EID50/ml. The H3N2, H5N1, H5N8, H4N6, H7, H9N2subtype strains can cause mice death, the mortality of H5N8reached up to100%, the rest of the H3and H6subtypes of the virus can make mice infected without clinical special features. These results showed that the sourced wild bird virus may directly infect humans without adaption. The receptor binding property and horizontal transmission between guinea pigs were analyzed to evaluate transmission in mammals. Receptor binding assay showed that the H3subtype virus has SA a-2,6Gal (human-like influenza virus receptor) binding property, H7, H9subtypes has SA α-2,6Gal binding property in some degree. H3and H9subtypes strains were selected for contacted transmission in guinea pigs, the S89(H3N2), ZH47(H3N3) and JL-2(H9N2) can transmit in guinea pigs, in which S89(H3N2) was the highest (2/3). The recombinant rS89strains were constructed by reverse genetics technology and found that rS89-HA-L2741can influence the S89virus SA α-2,6Gal receptor binding characteristics. These results showed that except the series of sites have been found; other sites receptor binding may also affect the recognition ability of virus receptor.
We established viral metagenomic to analyze the pathogen of wild birds, wild domestic mammalian. The method is universal for different type samples, such as environment, tissue, or swab samples. This method was first used to study viruses and distribution of wild bird, and detect a variety of virus (goose circovirus, pheasant herpes virus, duck hepatitis B virus, crane, hepatitis B virus, avian influenza virus etc.). Porcine epidemic diarrhea virus and porcine novel kobuvirus were found in outbreak of pig death. Most of Hantaviruses carried by animal hosts in Jilin Province were Seoul viruses, of which the genetics showed a diversity. These results showed that the method is important to new or unknown viral pathogens.
引文
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