CN101914500A - Gene-recombined H9N1 subtype avian influenza virus and application thereof - Google Patents
Gene-recombined H9N1 subtype avian influenza virus and application thereof Download PDFInfo
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- CN101914500A CN101914500A CN2009102722783A CN200910272278A CN101914500A CN 101914500 A CN101914500 A CN 101914500A CN 2009102722783 A CN2009102722783 A CN 2009102722783A CN 200910272278 A CN200910272278 A CN 200910272278A CN 101914500 A CN101914500 A CN 101914500A
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Abstract
The invention relates to a gene-recombined H9N1 subtype avian influenza virus and application thereof to preparing inactivated vaccines for preventing infection of an H9N2 subtype avian influenza virus. The virus is preserved in the China Center for Type Culture Collection (CCTCC) in August 3rd, 2009, and the preservation number is CCTCC V200913. The virus comprises an HA gene containing the H9N2 subtype avian influenza virus A/Chicken/Hubei/C1/2007 (H9N2) and 7 internal genes (PB2, PB1, PA, NP, NA, M and NS) containing the H1N1 influenza virus A/PR/8/34 (H1N1). The invention provides the application of the H9N1 subtype avian influenza virus to preparing the inactivated vaccines for preventing infection of the H9N2 subtype avian influenza virus, and the application of the H9N1 subtype avian influenza inactivated vaccines to distinguishing the immunity (H9N1) or field strain (H9N2) infection condition.
Description
Technical field
The present invention relates to a kind of H9N1 subtype avian influenza virus of gene recombination, and the application of the inactivated vaccine that should virus infects at preparation prevention H9N2 subtype avian influenza virus, this virus has been preserved in Chinese typical culture collection center on August 3rd, 2009, the address is a China. Wuhan. and Wuhan University, its called after influenza virus H9N1 hypotype A/Wuhan/Re-1/2009 (H9N1) that classifies, deposit number is CCTCC NO:V200913.
Background technology
(Avian Influenza is that (Avian Influenza Virus, the bird that AIV) causes infects and/or disease syndrome by avian influenza virus AI) in bird flu.AIV belongs in classification: viral boundary (Vira) ... orthomyxoviridae family (Orthomyxoviridae) ... Influenza Virus (Influenza Virus A and B) ... avian influenza virus (Avian InfluenzaVirus).The genome of AIV is made up of 8 sub-thread strand RNA fragments.According to surface protein antigenicity difference, AIV is divided into 16 hemagglutinin (HA) and 9 neuraminidases (NA) hypotype.HA is the main immunogen protein of AIV, it can induce body to produce antibody-mediated specific humoral immune response, thereby the antibody of anti-HA can combining or the viral infection that neutralizes of the fusion process of virus envelope and endocytosis body film by viral interference and sialic acid acceptor.H9N2 hypotype AIV occurred in the turkey in the U.S. first first in 1966, extensively distributed with popular in the wild fowl in the whole world and poultry at present.1998, the investigator was separated to the H9N2 avian influenza virus first from the human sample of China mainland, be separated to this subtype virus once more afterwards in patient's body of Hong Kong and China mainland; 2003 and 2007, the investigator repeatedly was separated to the H9N2 avian influenza virus in patient's body in Hong Kong.The incident of the frequent infected person of H9N2 avian influenza virus indicates that this subtype virus has the public health meaning.Along with the threat that the H9N2 avian influenza virus causes the mankind, people recognize, stop the valid approach of H9N2 avian influenza virus to people's group diffusion, are exactly the effective animal derived H9N2 avian influenza virus of control.Since nineteen ninety-six, China's aviculture is subjected to the puzzlement of H9N2 bird flu always, and especially 1998, this disease had caused catastrophic loss for China's aviculture.The H9N2 totivirus deactivation vaccine of chicken embryo production makes the control of this disease obtain effect preferably in the application of aviculture, but because the isolating street strain of terrain chicken embryo bad adaptability causes the vaccine of existing use to have shortcomings such as cost height, duration of immunity weak point; In addition, behind existing vaccine immunity animal, can't distinguish is the antibody that produces behind immunity back or the wild virus infection.
For solving an above-mentioned difficult problem, people have developed marker vaccine (marker vaccine) or have been referred to as DIVA vaccine (differentiating infected from vaccinated animals), be exactly after giving animal immune with a kind of vaccine, by the serology detection method that matches with it, immune animal and natural infected animal can be made a distinction.During 1999-2001 Italy breaks out the H7N1 bird flu, adopt the H7N3 strain isolated to be prepared into inactivated vaccine, connect breeder flock after, animal has produced the high antigenic antibody of anti-HA of tiring, and the attack that can resist the H7N1 avian influenza virus fully.Simultaneously, immune animal has produced the NA antibody at N3, can and infect poultry with immune poultry by means of test kit and make a distinction.Utilize this marker vaccine Italy eliminated effectively by the H7N1 avian influenza virus cause popular.When implementing the DIVA strategy, the generation of vaccine strain is a critical problem.At present, by the reverse genetics operative technique, can produce the purpose virus stain easily.
Summary of the invention
The H9N1 subtype avian influenza virus that the purpose of this invention is to provide a kind of gene recombination, called after A/Wuhan/Re-1/2009 (H9N1), this virus has height chicken embryo adaptability, but to chicken and chicken embryo no pathogenicity; With this virus as seed culture of viruses, after in the chicken embryo, breeding also deactivation, be prepared into oil emulsion inactivated vaccine, China's popular H9N2 subtype avian influenza virus had specific immune originality, in addition, by to the specific NA detection of antibodies of immune animal, can distinguish immunity (H9N1) or wild poison (H9N2) infection state.
The objective of the invention is to realize: the H9N1 subtype avian influenza virus of gene recombination by following technical scheme, it is characterized in that: its composition comprises HA gene with H9N2 subtype avian influenza virus A/Chicken/Hubei/C1/2007 (H9N2) and 7 internal gene PB2, PB1, PA, NP, NA, M and the NS of H1N1 influenza virus A/PR/8/34 (H1N1), and its preserving number is CCTCC-V200913.
This virus is received by China typical culture collection center on August 3rd, 2009, and is registered on the books.The viability of this culture has detected on August 31st, 2009 and has finished, and the result is survival.
Press such scheme, described HA gene has the nucleotide sequence shown in the sequence table SEQ ID NO.1.
Press such scheme, 7 internal gene PB2, PB1, PA, NP, NA, M and the NS of described H1N1 influenza virus A/PR/8/34 (H1N1) have the nucleotide sequence shown in sequence table SEQ ID NO.2～SEQ ID NO.8.
The H9N1 subtype avian influenza virus of above-mentioned gene recombination has specific immune originality to China's popular H9N2 subtype avian influenza virus, has height chicken embryo adaptability, but to chicken and chicken embryo no pathogenicity.
The H9N1 subtype avian influenza virus of above-mentioned gene recombination is in the application of the inactivated vaccine of preparation prevention H9N2 subtype avian influenza virus infection.
The application of above-mentioned H9N1 subtype avian influenza inactivated vaccine in distinguishing immunity (H9N1) or wild poison (H9N2) infection state.
The technology that the present invention adopts is that the viral RNA reverse transcription is become cDNA; obtain a kind of technology of RNA viruses by the plasmid that contains viral genome cDNA; by the reverse genetic manipulation technology; expression vector pHW2000 is gone in the HA gene clone of China popular H9N2 subtype avian influenza virus A/Chicken/Hubei/C1/2007 (H9N2); simultaneously; 7 gene (PB2 with influenza virus A/PR/8/34 (H1N1); PB1; PA; NP; NA; M and NS) also be cloned into expression vector pHW2000; 8 expression plasmids have been obtained; difference called after pHW-H9-HA, pHW-PR8-PB2; pHW-PR8-PB1; pHW-PR8-PA; pHW-PR8-NP; pHW-PR8-NA; pHW-PR8-M and pHW-PR8-NS.8 expression plasmid cotransfection 293T cells with above-mentioned have obtained the H9N1 subtype avian influenza virus of gene recombination.
With the H9N1 subtype avian influenza virus of above-mentioned gene recombination, after chicken embryo propagation and deactivation, be prepared into inactivated vaccine, produce the HI antibody of high titre behind the immune animal, can prevent the infection of H9N2 subtype avian influenza virus.
Behind the H9N1 subtype avian influenza virus inactivated vaccine immune animal with above-mentioned gene recombination, detection specificity NA antibody can be distinguished immunity (H9N1) or wild poison (H9N2) infection state.
Beneficial effect of the present invention:
1. the present invention utilizes the reverse genetics operative technique of influenza virus, has obtained H9N1 recombinant fowl influenza virus (A/Wuhan/Re-1/2009 (H9N1)).This virus has kept the distinctive antigenicity of China's H9 subtype avian influenza virus, as producing vaccine strain, will produce good special protection with this to the infection of H9N2 subtype avian influenza epidemic strain;
2. the avian influenza vaccine of existing use is the totivirus inactivated vaccine of chicken embryo breeding, and vaccine quality is directly related with virus antigen content.The H9N1 subtype avian influenza virus of gene recombination of the present invention contains 6 internal gene (PB2, PB1, PA, NP, M and NS) of chicken embryo adapted strain A/PR/8/34 (H1N1), the viral growth titre is compared with wild poison, can improve about 10 times, vaccine as vaccine strain production, can reduce the production of vaccine cost greatly, improve vaccine quality;
3. can distinguish vaccine immunity and wild virus infection, this also is the influence factor that avian influenza vaccine uses, the H9N1 subtype avian influenza virus of gene recombination of the present invention has kept the NA gene of A/PR/8/34 (H1N1), by to immune animal specificity NA detection of antibodies, can distinguish immunity and the wild virus infection situation of animal.
Embodiment
Further introduce the present invention below in conjunction with embodiment, but embodiment can not be construed as limiting the invention.
Embodiment 1:
The H9N1 subtype avian influenza virus of gene recombination of the present invention (A/Wuhan/Re-1/2009 (H9N1)), preserving number is CCTCC-V200913, its composition comprises HA gene with H9N2 subtype avian influenza virus A/Chicken/Hubei/C1/2007 (H9N2) and 7 internal gene PB2, PB1, PA, NP, NA, M and the NS of H1N1 influenza virus A/PR/8/34 (H1N1), and its preparation method comprises following step:
1. be template with China H9N2 subtype avian influenza virus A/Chicken/Hubei/C1/2007 (H9N2) strain RNA, with Auele Specific Primer RT-PCR amplification HA full-length cDNA, and insertion contains rna plymerase i, the BsmB I site of the RNA bidirectional transcription vector pHW2000 of rna plymerase ii.
2.RT-PCR 7 gene fragment PB2 of the high titre chicken of amplification influenza virus embryo adapted strain, PB1, PA, NP, NA, M and NS whole gene cDNA, insert among the pHW2000 as stated above respectively, 8 expression plasmids have been obtained, difference called after pHW-H9-HA, pHW-PR8-PB2, pHW-PR8-PB1, pHW-PR8-PA, pHW-PR8-NP, pHW-PR8-NA, pHW-PR8-M and pHW-PR8-NS.
3. utilize the liposome transfection method that the two-way expression plasmid of above-mentioned 8 genes is imported the 293T cell of individual layer simultaneously, 72 hours harvested cell supernatants are inoculated 10 age in days chick embryo allantoic cavities, collect allantoic fluid after 72 hours and detect blood clotting (HA) activity.The HA positive is the H9N1 subtype avian influenza virus of gene recombination.
Embodiment 2:
The growth characteristics of the H9N1 subtype avian influenza virus of gene recombination
With after 10 times of dilutions of H9N1 subtype avian influenza virus of gene recombination (10
-1-10
-10), be inoculated in instar chicken embryo on the 10th, every embryo 0.2ml, 5 pieces of chicken embryos of each dosage group.35 ℃ of cultivations are collected chick embryo allantoic liquid and are detected blood clotting (HA) activity after 72 hours, its HA tires and can reach 1024, and chicken embryo median infective dose (EID
50) can reach 10
10.3EID
50/ ml shows that these vaccine strain growth characteristics are good.
Embodiment 3:
The pathogenic analysis of the H9N1 subtype avian influenza virus of gene recombination
After 10 times of dilutions of H9N1 subtype avian influenza virus virus with gene recombination, inoculate instar chicken embryo on the 10th, every embryo 0.2ml.35 ℃ of cultivations were observed the chicken embryo every 6 hours, collected dead chicken embryo, until back 7 days of inoculation, and the mean time to death (MDT) of statistics chicken embryo.Experimental result shows, the MDT of recombinant virus＞168 hour show that this recombinant virus is to chicken embryo no pathogenicity.
In addition, the H9N1 subtype avian influenza virus of the gene recombination after 10 times of dilutions is inoculated 7 age in days chickens, observe postvaccinal clinical symptom, incidence and death condition, observed continuously 21 days.The result shows that behind the virus infection, experimental chicken does not have any clinical symptom, all survivals, shows that this virus is to chicken no pathogenicity, safety.
Embodiment 4:
The antigenicity analysis of the H9N1 subtype avian influenza virus of gene recombination
18 H9N2 avian influenza virus of China 2000-2007 popular Local Isolates repeatedly behind the immune rabbit, is prepared into hyper-immune serum.Carry out the HI experiment with the H9N1 subtype avian influenza virus of gene recombination and above-mentioned hyper-immune serum, measure the antigenicity of recombinant virus.The result shows that the H9N1 subtype avian influenza virus of gene recombination can cross reaction all take place with all hyper-immune serums, its HI tires 〉=and 640, show that this viral immunogenicity is good.
Embodiment 5:
Inductive antibody response behind the oil emulsion inactivated vaccine immune animal
With H9N1 subtype avian influenza virus inoculation instar chicken embryo on the 10th of gene recombination, the viral allantoic fluid (〉=10 of results after 72 hours
8EID
50/ ml), adding final volume and be 0.1% formaldehyde and carry out deactivation, the viral allantoic fluid that deactivation is up to the standards is 1: 1.5 oily adjuvant of adding in water and oil phase ratio, is prepared into oil emulsion inactivated vaccine.Female BALB/c mouse immunity in 4-6 age in week oil emulsion inactivated vaccine 0.1ml/ only, the immune oil emulsion inactivated vaccine of the chicken of different days (7 ages in days, 14 ages in days and 36 ages in week), be respectively 0.25ml/ only, 0.25ml/ only and 0.5ml/ only, determination of serum HI antibody titer is gathered in back 28 days of immunity.The result shows, behind back 14 days HI antibody titer 〉=256,28 of BALB/c mouse immunity day 〉=1024; Behind different days chicken (1 age in week, 2 ages in week and 36 ages in week) HI antibody titer 〉=256,28 of immunity after 14 days day 〉=1024.
Embodiment 6:
The protection of oil emulsion inactivated vaccine is renderd a service
Female BALB/c mouse immunity in 4-6 age in week oil emulsion inactivated vaccine 0.1ml/ only, the immune oil emulsion inactivated vaccine of the chicken of different days (7 ages in days, 14 ages in days and 36 ages in week) be respectively 0.25ml/ only, 0.25ml/ only and 0.5ml/ only, back 28 days of immunity is attacked with the H9N2 avian influenza virus.Observe clinical symptom, incidence and the death condition of infection animal, continuous monitoring 21 days.The result shows that the immune group BALB/c mouse is all survived (protection ratio 100%), and control group is all dead; The immune chicken of different days does not have any clinical symptom and all survivals, and tangible clinical symptom appears in the chicken of control group, and its mortality ratio can reach 25%.
Embodiment 7:
Serology differential diagnosis
With 20 of H9N1 gene recombination bird flu oil emulsion inactivated vaccine immunity 14 age in days chickens, 0.25ml/, serum was collected in immunity in back 28 days; Simultaneously, with 10
7EID
5020 of the H9N2 avian influenza 14 age in days chickens of dosage, serum was collected in immunity in back 28 days.With N2 is diagnostic antigen, and the standard NI test method that adopts WHO to recommend detects the serum of H9N1 gene recombination bird flu oil emulsion inactivated vaccine immunity and the serum of H9N2 avian influenza.The result shows that all chickens of immune oil emulsion inactivated vaccine are all negative, and the serum of H9N2 avian influenza all is positive, and this shows immunity and the infection of adopting the serology detection method can distinguish animal.
The A/Wuhan/Re-1/2009 of gene recombination (H9N1) virus gene sequence
SEQ?ID?No.1
AGCAAAAGCAGGGGAATTTCACAACCAGTCAAAATGGAAACAATATCACTAATAGCTATACTACTAGTAGTAACAGTAA
GCAATGCAGATAAAATCTGCATCGGCTACCAATCAACAAACTCCACAGAACCTGTGGATACGCTAACAGAAAACAATGT
CCCTGTGACACATGCTAAAGAATTGCTCCACACAGAGCACAATGGGATGCTGTGTGCAACAAATCTGGGACATCCTCTC
ATTCTAAACACCTGTACCATTGAAGGACTGATCTATGGCAACCCTTCTCGTGATCAGCTGTTGGGAGGAGGAAAATGGT
CCTACATCGTCGAAAGACCATCGGCCGTTAATGGAATGTGTTACCCTGGGAATGTAGAAAACCTAGAGGAACTAAGATC
ACTCTTTAGTTCTGCTAGTTCCTACCAAAGAATTCGGATCTTTCCAGACACGATCTGGAATGTGTCTTACAATGGAACA
AGCAAAGCATGTTCAGATTCATTCTACAGAAGCATGAGATGGCTGACTCAAAAGAACACCGCTTACCCTATTCAAGACG
CCCAATACACAAATAATAGAGGAAAGAGCATTCTTTTCATGTGGGGCATAAATCACCCACCCACCGATACTGTACAGAC
AAATTTGTACACAAGGACCGACACAACAACAAGTGTGACAACAGAGGATATAACTAGAACCTTCAAACCAATGATAGGG
CCAAGGCCCCTTGTCAATGGTCAGCAGGGGAGAATTGATTATTATTGGTCGGTATTAAAACCAGGTCAGACATTGCGAA
TAAGATCCAATGGGAATCTAATTGCTCCATGGTATGGACACATTCTTCCAGGAGAGAGCCACGGAAGAATCCTGAAGAC
TGATTTAAACAGTGGTAACTGTGTAGTGCAATGTCAGACTGAAAGAGGTGGCTTAAACACCACATTGCCGTTCCACAAT
GTCAGTAAGTATGCATTTGGGAACTGCCCAAAGTATGTTGGAGTAAAGAGTCTCAGACTGGCAGTTGGTCTAAGAAATG
TGCCTGCTAGATCAAGTAGAGGACTATTTGGGGCTATAGCTGGTTTCATAGAGGGAGGTTGGTCAGGGTTAGTCGCTGG
TTGGTATGGGTTCCAGCATTCAAATGATCAAGGGGTAGGTATGGCTGCAGATAGAGGGTCAACTCAAAGGGCAATTGAC
AAAATAACATCCAAAGTGAATAATATAGTCGATAAAATGAACAAGCAGTATGAAATTATTGATCATGAATTCAGCGAGG
TTGAAACTAGACTCAATATGATCAATAATAAGATTGATGATCAAATACAAGACATATGGGCTTATAATGCAGAATTGCT
AGTGCTGCTTGAAAATCAGAAAACACTCGATGAACATGATGCAAATGTGAACAATCTATATAACAAAGTGAAGAGGGCA
CTGGGTTCCAATGCCATGGAAGACGGGAAAGGATGTTTTGAGCTATACCATAAATGTGCTGATCAGTGCATGGAGACAA
TTCGGAACGGGACCTATAACAGGAGAAAGTATAAAGAGGAAGCAAAACTAGAAAGACAGAAAATAGAAGGGGTCAAGCT
GGAATCTGAAGGAACTTACAAAATCCTCACCATTTATTCGACTGTCGCCTCATCTCTTGTGATTGCAATGGGGTTTGCT
GCCTTCTTGTTCTGGGCCATGTCCAATGGATCTTGCAGATGCAACATTTGTATATAATTGGCAAAAACACCCTTGTTTC
TACT
SEQ?ID?No.2
AGCGAAAGCAGGTCAATTATATTCAATATGGAAAGAATAAAAGAACTAAGAAATCTAATGTCGCAGTCTCGCACCCGCG
AGATACTCACAAAAACCACCGTGGACCATATGGCCATAATCAAGAAGTACACATCAGGAAGACAGGAGAAGAACCCAGC
ACTTAGGATGAAATGGATGATGGCAATGAAATATCCAATTACAGCAGACAAGAGGATAACGGAAATGATTCCTGAGAGA
AATGAGCAAGGACAAACTTTATGGAGTAAAATGAATGATGCCGGATCAGACCGAATGATGGTATCACCTCTGGCTGTGA
CATGGTGGAATAGGAATGGACCAATGACAAATACAGTTCATTATCCAAAAATCTACAAAACTTATTTTGAAAGAGTCGA
AAGGCTAAAGCATGGAACCTTTGGCCCTGTCCATTTTAGAAACCAAGTCAAAATACGTCGGAGAGTTGACATAAATCCT
GGTCATGCAGATCTCAGTGCCAAGGAGGCACAGGATGTAATCATGGAAGTTGTTTTCCCTAACGAAGTGGGAGCCAGGA
TACTAACATCGGAATCGCAACTAACGATAACCAAAGAGAAGAAAGAAGAACTCCAGGATTGCAAAATTTCTCCTTTAAT
GGTTGCATACATGTTGGAGAGAGAACTGGTCCGCAAAACGAGATTCCTCCCAGTGGCTGGTGGAACAAGCAGTGTGTAC
ATTGAAGTGTTGCATTTGACTCAAGGAACATGCTGGGAACAGATGTATACTCCAGGAGGGGAAGTGAGGAATGATGATG
TTGATCAAAGCTTGATTATTGCTGCTAGGAACATAGTGAGAAGAGCTGCAGTATCAGCAGATCCACTAGCATCTTTATT
GGAGATGTGCCACAGCACACAGATTGGTGGAATTAGGATGGTAGACATCCTTAGGCAGAACCCAACAGAAGAGCAAGCC
GTGGATATATGCAAGGCTGCAATGGGACTGAGAATTAGCTCATCCTTCAGTTTTGGTGGATTCACATTTAAGAGAACAA
GCGGATCATCAGTCAAGAGAGAGGAAGAGGTGCTTACGGGCAATCTTCAAACATTGAAGATAAGAGTGCATGAGGGATA
TGAAGAGTTCACAATGGTTGGGAGAAGAGCAACAGCCATACTCAGAAAAGCAACCAGGAGATTGATTCAGCTGATAGTG
AGTGGGAGAGACGAACAGTCGATTGCCGAAGCAATAATTGTGGCCATGGTATTTTCACAGGAGGATTGTATGATAAAAG
CAGTTAGAGGTGATCTGAATTTCGTCAATAGGGCGAATCAGCGATTGAATCCTATGCATCAACTTTTAAGACATTTTCA
GAAGGATGCGAAAGTGCTTTTTCAAAATTGGGGAGTTGAACCTATCGACAATGTGATGGGAATGATTGGGATATTGCCC
GACATGACTCCAAGCATCGAGATGTCAATGAGAGGAGTGAGAATCAGCAAAATGGGTGTAGATGAGTACTCCAGCACGG
AGAGGGTAGTGGTGAGCATTGACCGGTTTTTGAGAATCCGGGACCAACGAGGAAATGTACTACTGTCTCCCGAGGAGGT
CAGTGAAACACAGGGAACAGAGAAACTGACAATAACTTACTCATCGTCAATGATGTGGGAGATTAATGGTCCTGAATCA
GTGTTGGTCAATACCTATCAATGGATCATCAGAAACTGGGAAACTGTTAAAATTCAGTGGTCCCAGAACCCTACAATGC
TATACAATAAAATGGAATTTGAACCATTTCAGTCTTTAGTACCTAAGGCCATTAGAGGCCAATACAGTGGGTTTGTAAG
AACTCTGTTCCAACAAATGAGGGATGTGCTTGGGACATTTGATACCGCACAGATAATAAAACTTCTTCCCTTCGCAGCC
GCTCCACCAAAGCAAAGTAGAATGCAGTTCTCCTCATTTACTGTGAATGTGAGGGGATCAGGAATGAGAATACTTGTAA
GGGGCAATTCTCCTGTATTCAACTACAACAAGGCCACGAAGAGACTCACAGTTCTCGGAAAGGATGCTGGCACTTTAAC
TGAAGACCCAGATGAAGGCACAGCTGGAGTGGAATCCGCTGTTCTGAGGGGATTCCTCATTCTGGGCAAAGAAGACAGG
AGATATGGGCCAGCATTAAGCATCAATGAACTGAGCAACCTTGCGAAAGGAGAGAAGGCTAATGTGCTAATTGGGCAAG
GAGACGTGGTGTTGGTAATGAAACGAAAACGGGACTCTAGCATACTTACTGACAGCCAGACAGCGACCAAAAGAATTCG
GATGGCCATCAATTAGTGTCGAATAGTTTAAAAACGACCTTGTTTCTACT
SEQ?ID?No.3
AGCGAAAGCAGGCAAACCATTTGAATGGATGTCAATCCGACCTTACTTTTCTTAAAAGTGCCAGCACAAAATGCTATAA
GCACAACTTTCCCTTATACCGGAGACCCTCCCTACAGCCATGGGACAGGAACAGGATACACCATGGATACTGTCAACAG
GACACATCAGTACTCAGAAAAGGGAAGATGGACAACAAACACCGAAACTGGAGCACCGCAACTCAACCCGATTGATGGG
CCACTGCCAGAAGACAATGAACCAAGTGGTTATGCCCAAACAGATTGTGTATTGGAAGCAATGGCTTTCCTTGAGGAAT
CCCATCCTGGTATTTTTGAAAACTCGTGTATTGAAACGATGGAGGTTGTTCAGCAAACACGAGTAGACAGGCTGACACA
AGGCCGACAGACCTATGACTGGACTCTAAATAGAAACCAGCCTGCTGCAACAGCATTGGCCAACACAATAGAAGTGTTC
AGATCAAATGGCCTCACGGCCAATGAGTCTGGAAGGCTCATAGACTTCCTTAAGGATGTAATGGAGTCAATGAACAAAG
AAGAAATGGGGATCACAACTCATTTTCAGAGAAAGAGACGGGTGAGAGACAATATGACTAAGAAAATGATAACACAGAG
AACAATAGGTAAAAGGAAGCAGAGATTGAACAAAAGGAGTTATCTAATTAGAGCATTGACCCTGAACACAATGACCAAA
GATGCTGAGAGAGGGAAGCTAAAACGGAGAGCAATTGCAACCCCAGGGATGCAAATAAGGGGGTTTGTATACTTTGTTG
AGACACTGGCAAGAAGTATATGTGAGAAACTTGAACAATCAGGGTTGCCAGTTGGAGGCAATGAGAAGAAAGCAAAGTT
GGCAAATGTTGTAAGGAAGATGATGACCAATTCTCAGGACACCGAACTTTCTTTCACCATCACTGGAGATAACACCAAA
TGGAACGAAAATCAGAATCCTCGGATGTTTTTGGCCATGATCACATATATGACCAGAAATCAGCCCGAATGGTTCAGAA
ATGTTCTAAGTATTGCTCCAATAATGTTCTCAAACAAAATGGCGAGACTGGGAAAAGGGTATATGTTTGAGAGCAAGAG
TATGAAACTTAGAACCCAAATACCTGCAGAAATGCTAGCAAGCATTGATTTGAAATATTTCAATGATTCAACAAGAAAG
AAGATTGAAAAAATCCGACCGCTCTTAATAGAGGGGACTGCATCATTGAGCCCTGGAATGATGATGGGCATGTTCAATA
TGTTAAGCACTGTATTAGGCGTCTCCATCCTGAATCTTGGACAAAAGAGATACACCAAGACTACTTACTGGTGGGATGG
TCTTCAATCCTCTGACGATTTTGCTCTGATTGTGAATGCACCCAATCATGAAGGGATTCAAGCCGGAGTCGACAGGTTT
TATCGAACCTGTAAGCTACTTGGAATCAATATGAGCAAGAAAAAGTCTTACATAAACAGAACAGGTACATTTGAATTCA
CAAGTTTTTTCTATCGTTATGGGTTTGTTGCCAATTTCAGCATGGAGCTTCCCAGTTTTGGGGTGTCTGGGATCAACGA
GTCAGCGGACATGAGTATTGGAGTTACTGTCATCAAAAACAATATGATAAACAATGATCTTGGTCCAGCAACAGCTCAA
ATGGCCCTTCAGTTGTTCATCAAAGATTACAGGTACACGTACCGATGCCATAGAGGTGACACACAAATACAAACCCGAA
GATCATTTGAAATAAAGAAACTGTGGGAGCAAACCCGTTCCAAAGCTGGACTGCTGGTCTCCGACGGAGGCCCAAATTT
ATACAACATTAGAAATCTCCACATTCCTGAAGTCTGCCTAAAATGGGAATTGATGGATGAGGATTACCAGGGGCGTTTA
TGCAACCCACTGAACCCATTTGTCAGCCATAAAGAAATTGAATCAATGAACAATGCAGTGATGATGCCAGCACATGGTC
CAGCCAAAAACATGGAGTATGATGCTGTTGCAACAACACACTCCTGGATCCCCAAAAGAAATCGATCCATCTTGAATAC
AAGTCAAAGAGGAGTACTTGAAGATGAACAAATGTACCAAAGGTGCTGCAATTTATTTGAAAAATTCTTCCCCAGCAGT
TCATACAGAAGACCAGTCGGGATATCCAGTATGGTGGAGGCTATGGTTTCCAGAGCCCGAATTGATGCACGGATTGATT
TCGAATCTGGAAGGATAAAGAAAGAAGAGTTCACTGAGATCATGAAGATCTGTTCCACCATTGAAGAGCTCAGACGGCA
AAAATAGTGAATTTAGCTTGTCCTTCATGAAAAAATGCCTTGTTTCTACT
SEQ?ID?No.4
AGCGAAAGCAGGTACTGATCCAAAATGGAAGATTTTGTGCGACAATGCTTCAATCCGATGATTGTCGAGCTTGCGGAAA
AAACAATGAAAGAGTATGGGGAGGACCTGAAAATCGAAACAAACAAATTTGCAGCAATATGCACTCACTTGGAAGTATG
CTTCATGTATTCAGATTTTCACTTCATCAATGAGCAAGGCGAGTCAATAATCGTAGAACTTGGTGATCCAAATGCACTT
TTGAAGCACAGATTTGAAATAATCGAGGGAAGAGATCGCACAATGGCCTGGACAGTAGTAAACAGTATTTGCAACACTA
CAGGGGCTGGGAAACCAAAGTTTCTACCAGATTTGTATGATTACAAGGAGAATAGATTCATCGAAATTGGAGTAACAAG
GAGAGAAGTTCACATATACTATCTGGAAAAGGCCAATAAAATTAAATCTGAGAAAACACACATCCACATTTTCTCGTTC
ACTGGGGAAGAAATGGCCACAAAGGCAGACTACACTCTCGATGAAGAAAGCAGGGCTAGGATCAAAACCAGACTATTCA
CCATAAGACAAGAAATGGCCAGCAGAGGCCTCTGGGATTCCTTTCGTCAGTCCGAGAGAGGAGAAGAGACAATTGAAGA
AAGGTTTGAAATCACAGGAACAATGCGCAAGCTTGCCGACCAAAGTCTCCCGCCGAACTTCTCCAGCCTTGAAAATTTT
AGAGCCTATGTGGATGGATTCGAACCGAACGGCTACATTGAGGGCAAGCTGTCCCAAATGTCCAAAGAAGTAAATGCTA
GAATCGAACCTTTTTTGAAAACAACACCACGACCACTTAGACTTCCGAATGGGCCTCCCTGTTCTCAGCGGTCCAAGTT
CCTGCTGATGGATGCCTTAAAATTAAGCATTGAGGACCCAAGTCATGAAGGAGAGGGAATACCGCTATATGATGCAATC
AAATGCATGAGAACATTCTTTGGATGGAAGGAACCCAATGTTGTTAAACCACACGAAAAGGGAATAAATCCAAATTATC
TTCTGTCATGGAAGCAAGTACTGGCAGAACTGCAGGACATTGAGAATGAGGAGAAAATTCCAAAGACTAAAAATATGAA
GAAAACAAGTCAGCTAAAGTGGGCACTTGGTGAGAACATGGCACCAGAAAAGGTAGACTTTGACGACTGTAAAGATGTA
GGTGATTTGAAGCAATATGATAGTGATGAACCAGAATTGAGGTCGCTTGCAAGTTGGATTCAGAATGAGTTTAACAAGG
CATGCGAACTGACAGATTCAAGCTGGATAGAGCTCGATGAGATTGGAGAAGATGTGGCTCCAATTGAACACATTGCAAG
CATGAGAAGGAATTATTTCACATCAGAGGTGTCTCACTGCAGAGCCACAGAATACATAATGAAGGGGGTGTACATCAAT
ACTGCCTTGCTTAATGCATCTTGTGCAGCAATGGATGATTTCCAATTAATTCCAATGATAAGCAAGTGTAGAACTAAGG
AGGGAAGGCGAAAGACCAACTTGTATGGTTTCATCATAAAAGGAAGATCCCACTTAAGGAATGACACCGACGTGGTAAA
CTTTGTGAGCATGGAGTTTTCTCTCACTGACCCAAGACTTGAACCACACAAATGGGAGAAGTACTGTGTTCTTGAGATA
GGAGATATGCTTCTAAGAAGTGCCATAGGCCAGGTTTCAAGGCCCATGTTCTTGTATGTGAGAACAAATGGAACCTCAA
AAATTAAAATGAAATGGGGAATGGAGATGAGGCGTTGCCTCCTCCAGTCACTTCAACAAATTGAGAGTATGATTGAAGC
TGAGTCCTCTGTCAAAGAGAAAGACATGACCAAAGAGTTCTTTGAGAACAAATCAGAAACATGGCCCATTGGAGAGTCC
CCCAAAGGAGTGGAGGAAAGTTCCATTGGGAAGGTCTGCAGGACTTTATTAGCAAAGTCGGTATTCAACAGCTTGTATG
CATCTCCACAACTAGAAGGATTTTCAGCTGAATCAAGAAAACTGCTTCTTATCGTTCAAGCTCTTAGGGACAACCTGGA
ACCTGGGACCTTTGATCTTGGGGGGCTATATGAAGCAATTGAGGAGTGCCTAATTAATGATCCCTGGGTTTTGCTTAAT
GCTTCTTGGTTCAACTCCTTCCTTACACATGCATTGAGTTAGTTGTGGCAGTGCTACTATTTGCTATCCATACTGTCCA
AAAAAGTACCTTGTTTCTACT
SEQ?ID?No.5
AGCAAAAGCAGGGTAGATAATCACTCACTGAGTGACATCAAAATCATGGCGTCCCAAGGCACCAAACGGTCTTACGAAC
AGATGGAGACTGATGGAGAACGCCAGAATGCCACTGAAATCAGAGCATCCGTCGGAAAAATGATTGGTGGAATTGGACG
ATTCTACATCCAAATGTGCACCGAACTTAAACTCAGTGATTATGAGGGACGGTTGATCCAAAACAGCTTAACAATAGAG
AGAATGGTGCTCTCTGCTTTTGACGAAAGGAGAAATAAATACCTGGAAGAACATCCCAGTGCGGGGAAAGATCCTAAGA
AAACTGGAGGACCTATATACAGGAGAGTAAACGGAAAGTGGATGAGAGAACTCATCCTTTATGACAAAGAAGAAATAAG
GCGAATCTGGCGCCAAGCTAATAATGGTGACGATGCAACGGCTGGTCTGACTCACATGATGATCTGGCATTCCAATTTG
AATGATGCAACTTATCAGAGGACAAGAGCTCTTGTTCGCACCGGAATGGATCCCAGGATGTGCTCTCTGATGCAAGGTT
CAACTCTCCCTAGGAGGTCTGGAGCCGCAGGTGCTGCAGTCAAAGGAGTTGGAACAATGGTGATGGAATTGGTCAGGAT
GATCAAACGTGGGATCAATGATCGGAACTTCTGGAGGGGTGAGAATGGACGAAAAACAAGAATTGCTTATGAAAGAATG
TGCAACATTCTCAAAGGGAAATTTCAAACTGCTGCACAAAAAGCAATGATGGATCAAGTGAGAGAGAGCCGGAACCCAG
GGAATGCTGAGTTCGAAGATCTCACTTTTCTAGCACGGTCTGCACTCATATTGAGAGGGTCGGTTGCTCACAAGTCCTG
CCTGCCTGCCTGTGTGTATGGACCTGCCGTAGCCAGTGGGTACGACTTTGAAAGAGAGGGATACTCTCTAGTCGGAATA
GACCCTTTCAGACTGCTTCAAAACAGCCAAGTGTACAGCCTAATCAGACCAAATGAGAATCCAGCACACAAGAGTCAAC
TGGTGTGGATGGCATGCCATTCTGCCGCATTTGAAGATCTAAGAGTATTAAGCTTCATCAAAGGGACGAAGGTGGTCCC
AAGAGGGAAGCTTTCCACTAGAGGAGTTCAAATTGCTTCCAATGAAAATATGGAGACTATGGAATCAAGTACACTTGAA
CTGAGAAGCAGGTACTGGGCCATAAGGACCAGAAGTGGAGGAAACACCAATCAACAGAGGGCATCTGCGGGCCAAATCA
GCATACAACCTACGTTCTCAGTACAGAGAAATCTCCCTTTTGACAGAACAACCATTATGGCAGCATTCACTGGGAATAC
AGAGGGGAGAACATCTGACATGAGGACCGAAATCATAAGGATGATGGAAAGTGCAAGACCAGAAGATGTGTCTTTCCAG
GGGCGGGGAGTCTTCGAGCTCTCGGACGAAAAGGCAGCGAGCCCGATCGTGCCTTCCTTTGACATGAGTAATGAAGGAT
CTTATTTCTTCGGAGACAATGCAGAGGAGTACGACAATTAAAGAAAAATACCCTTGTTTCTACT
SEQ?ID?No.6
AGCGAAAGCAGGGGTTTAAAATGAATCCAAATCAGAAAATAATAACCATTGGATCAATCTGTATGGTAGTCGGACTAAT
TAGCCTAATATTGCAAATAGGAAATATAACCTCAATATGGATTAGCCATTCAATTCAAACTGGAAGTCAAAACCATACT
GGAATATGCAACCAAAACATCATTACCTATAAAAATAGCATCTGGGTAAAGGACCCAACTTCAGTGATATTAACCGGCA
ATTCATCTCTTTGTCCCATCCGTGGGTGGGCTATATACAGCAAAGACAATAGCATAAGAATTGGTTCCAAAGGAGACGT
TTTTGTCATAAGAGAGCCCTTTATTTCATGTTCTCACTTGGAATGCAGGACCTTTTTTCTGACCCAAGGTGCCTTACTG
AATGACAGGCATTCAAATGGGACTGTTAAGGACAGAAGCCCTTATAGGGCCTTAATGAGCTGCCCTGTCGGTGAAGCTC
CGTCCCCGTACAATTCAAGATTTGAATCGGTTGCTTGGTCAGCAAGTGCATGTCATGATGGCATGGGCTGGCTAACAAT
CGGAATTTCAGGTCCAGATAATGGAGCAGTGGCTGTATTAAAATACAACGGCATAATAACTGAAACCATAAAAAGTTGG
AGGAAGAAAATATTGAGGACACAAGAGTCTGAATGTGCCTGTGTAAATGGTTCATGTTTTACTATAATGACTGATGGCC
CGAGTGATGGGCTGGCCTCGTACAAAATTTTCAAGATCGAAAAGGGGAAGGTTACTAAATCAATAGAGTTGAATGCACC
TAATTCTCACTATGAGGAATGTTCCTGTTACCCTGATACCGGCAAAGTGATGTGTGTGTGCAGAGACAATTGGCATGGT
TCGAACCGGCCGTGGGTGTCTTTCGATCAAAACCTGGATTATCAAATAGGATACATCTGCAGTGGGGTTTTCGGTGACA
ACCCGCGTCCCAAAGATGGAACAGGCAGCTGTGGTCCAGTGTATGTTGATGGAGCAAACGGAGTAAAGGGATTTTCATA
TAGGTATGGTAATGGTGTTTGGATAGGAAGGACCAAAAGTCACAGTTCCAGACATGGGTTTGAGATGATTTGGGATCCT
AATGGATGGACAGAGACTGATAGTAAGTTCTCTGTGAGGCAAGATGTTGTGGCAATGACTGATTGGTCAGGGTATAGCG
GAAGTTTCGTTCAACATCCTGAGCTAACAGGGCTAGACTGTATAAGGCCGTGCTTCTGGGTTGAATTAATCAGGGGACG
ACCTAAAGAAAAAACAATCTGGACTAGTGCGAGCAGCATTTCTTTTTGTGGCGTGAATAGTGATACTGTAGATTGGTCT
TGGCCAGACGGTGCTGAGTTGCCATTCACCATTGACAAGTAGTCTGTTCAAAAAACTCCTTGTTTCTACT
SEQ?ID?No.7
AGCAAAAGCAGGTAGATATTGAAAGATGAGTCTTCTAACCGAGGTCGAAACGTACGTTCTCTCTATCATCCCGTCAGGC
CCCCTCAAAGCCGAGATAGCACAGAGACTTGAAGATGTCTTTGCAGGGAAGAACACCGATCTTGAGGTTCTCATGGAAT
GGCTAAAGACAAGACCAATCCTGTCACCTCTGACTAAGGGGATTTTAGGATTTGTGTTCACGCTCACCGTGCCCAGTGA
GCGAGGACTGCAGCGTAGACGCTTTGTCCAAACTGCCCTTAATGGGAACGGGGATCCAAATAACATGGACAAAGCAGTT
AAACTGTATAGGAAGCTCAAGAGGGAGATAACATTCCATGGGGCCAAAGAAATCTCACTCAGTTATTCTGCTGGTGCAC
TTGCCAGTTGTATGGGCCTCATATACAACAGGATGGGGGCTGTGACCACTGAAGTGGCATTTGGCCTGGTATGTGCAAC
CTGTGAACAGATTGCTGACTCCCAGCATCGGTCTCATAGGCAAATGGTGACAACAACCAATCCACTAATCAGACATGAG
AACAGAATGGTTTTAGCCAGCACTACAGCTAAGGCTATGGAGCAAATGGCTGGATCGAGTGAGCAAGCAGCAGAGGCCA
TGGAGGTTGCTAGTCAGGCTAGGCAAATGGTGCAAGCGATGAGAACCATTGGGACTCATCCTAGCTCCAGTGCTGGTCT
GAAAAATGATCTTCTTGAAAATTTGCAGGCCTATCAGAAACGAATGGGGGTGCAGATGCAACGGTTCAAGTGATCCTCT
CGCTATTGCCGCAAATATCATTGGGATCTTGCACTTGACATTGTGGATTCTTGATCGTCTTTTTTTCAAATGCATTTAC
CGTCGCTTTAAATACGGACTGAAAGGAGGGCCTTCTACGGAAGGAGTGCCAAAGTCTATGAGGGAAGAATATCGAAAGG
AACAGCAGAGTGCTGTGGATGCTGACGATGGTCATTTTGTCAGCATAGAGCTGGAGTAAAAAACTACCTTGTTTCTACT
SEQ?ID?No.8
AGCAAAAGCAGGGTGACAAAAACATAATGGATTCAAACACTGTGTCAAGCTTTCAGGTAGATTGCTTTCTTTGGCATGT
CCGCAAGCGAGTTGCAGACCAAGAACTAGGTGATGCCCCATTCCTTGATCGGCTTCGCCGAGATCAGAAATCCCTAAGA
GGAAGGGGCAGCACTCTCGGTCTGGACATCGAGACAGCCACACGTGCTGGAAAGCAGATAGTGGAGCGGATTCTGAAAG
AAGAATCCGATGAGGCACTTAAAATGACCATGGCCTCTGTACCTGCGTCGCGTTACCTAACTGACATGACTCTTGAGGA
AATGTCAAGGGACTGGTCCATGCTCATACCCAAGCAGAAAGTGGCAGGCCCTCTTTGTATCAGAATGGACCAGGCGATC
ATGGATAAGAACCTCATACTGAAAGCGAACTTCAGTGTGATTTTTGACCGGCTGGAGACTCTAATATTGCTAAGGGCTT
TCACCGAAGAGGGAGCAATTGTTGGCGAAATTTCACCATTGCCTTCTCTTCCAGGACATACTGCTGAGGATGTCAAAAA
TGCAGTTGGAGTCCTCATCGGAGGGCTTGAATGGAATGATAACACAGTTCGAGTCTCTGAAACTCTACAGAGATTCGCT
TGGAGAAGCAGTAATGAGAATGGGAGACCTCCACTCACTCCAAAACAGAAACGAGAAATGGCGGGAACAATTAGGTCAG
AAGTTTGAAGAAATAAGATGGTTGATTGAAGAAGTGAGACACAAACTGAAGGTAACAGAGAATAGTTTTGAGCAAATAA
CATTTATGCAAGCCTTACATCTATTGCTTGAAGTGGAGCAAGAGATAAGAACTTTCTCATTTCAGCTTATTTAATAATA
AAAAACACCCTTGTTTCTACT
Claims (5)
1. the H9N1 subtype avian influenza virus of gene recombination, it is characterized in that: its composition comprises HA gene with H9N2 subtype avian influenza virus A/Chicken/Hubei/C1/2007 (H9N2) and 7 internal gene PB2, PB1, PA, NP, NA, M and the NS of H1N1 influenza virus A/PR/8/34 (H1N1), and its preserving number is CCTCC-V200913.
2. by the H9N1 subtype avian influenza virus of the described gene recombination of claim 1, it is characterized in that: described HA gene has the nucleotide sequence shown in the sequence table SEQ ID NO.1.
3. by the H9N1 subtype avian influenza virus of claim 1 or 2 described gene recombination, it is characterized in that: 7 internal gene PB2, PB1, PA, NP, NA, M and the NS of described H1N1 influenza virus A/PR/8/34 (H1N1) have the nucleotide sequence shown in sequence table SEQ ID NO.2～SEQ ID NO.8.
4. the H9N1 subtype avian influenza virus of the described gene recombination of claim 1 is in the application of the inactivated vaccine of preparation prevention H9N2 subtype avian influenza virus infection.
5. by the described application of claim 4, it is characterized in that the application of inactivated vaccine in distinguishing immunity (H9N1) or wild poison (H9N2) infection state that described prevention H9N2 subtype avian influenza virus infects.
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Cited By (3)
| Publication number | Priority date | Publication date | Assignee | Title |
|---|---|---|---|---|
| CN102586194A (en) * | 2011-12-01 | 2012-07-18 | 哈药集团生物疫苗有限公司 | H9 subtype avian influenza virus Vero cell adapted strain and application thereof |
| CN104312983A (en) * | 2014-10-20 | 2015-01-28 | 中国疾病预防控制中心病毒病预防控制所 | Recombinant influenza virus and application thereof |
| CN104372023A (en) * | 2014-11-10 | 2015-02-25 | 中国农业大学 | Method for rescuing influenza virus |
-
2009
- 2009-09-28 CN CN2009102722783A patent/CN101914500A/en active Pending
Cited By (4)
| Publication number | Priority date | Publication date | Assignee | Title |
|---|---|---|---|---|
| CN102586194A (en) * | 2011-12-01 | 2012-07-18 | 哈药集团生物疫苗有限公司 | H9 subtype avian influenza virus Vero cell adapted strain and application thereof |
| CN104312983A (en) * | 2014-10-20 | 2015-01-28 | 中国疾病预防控制中心病毒病预防控制所 | Recombinant influenza virus and application thereof |
| CN104372023A (en) * | 2014-11-10 | 2015-02-25 | 中国农业大学 | Method for rescuing influenza virus |
| CN104372023B (en) * | 2014-11-10 | 2017-02-08 | 中国农业大学 | Method for rescuing influenza virus |
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