Endemicity of H9N2 and H5N1 avian influenza viruses in poultry in China poses a serious threat to poultry industry and public health

Frontiers of Agricultural Science and Engineering

Volumn 3, Issue 1, 2016, Pages 11-24

  Hu, Jiao ^a   Liu, Xiufan ^a  

^a YANGZHOU UNIVERSITY   (China)

Author keywords

Avian influenza virus; H5N1; H9N2; Novel viruses; Public health

Indexed keywords

EID: 85006292502     PISSN: 20957505     EISSN: 2095977X     Source Type: Journal    
DOI: 10.15302/J-FASE-2016092     Document Type: Review

References (179)

1. Stallknecht D E, Shane S M. Host range of avian influenza virus in free-living birds. Veterinary Research Communications, 1988, 12 (2-3): 125-141
2. Webster R G, Bean W J, Gorman O T, Chambers T M, Kawaoka Y. Evolution and ecology of influenza A viruses. Microbiological Reviews, 1992, 56(1): 152-179
3. Krauss S, Walker D, Pryor S P, Niles L, Chenghong L, Hinshaw V S, Webster R G. Influenza A viruses of migrating wild aquatic birds in North America. Vector Borne and Zoonotic Diseases, 2004, 4 (3): 177-189
4. Olsen B, Munster V J, Wallensten A, Waldenstrom J, Osterhaus A D, Fouchier R A. Global patterns of influenza a virus in wild birds. Science, 2006, 312(5772): 384-388
5. Spackman E. The ecology of avian influenza virus in wild birds: what does this mean for poultry? Poultry Science, 2009, 88(4): 847-850
6. Krauss S, Stallknecht D E, Negovetich N J, Niles L J, Webby R J, Webster R G. Coincident ruddy turnstone migration and horseshoe crab spawning creates an ecological 'hot spot' for influenza viruses. Proceedings of the Royal Society B: Biological Sciences, 1699, 2010(277): 3373-3379
7. Horimoto T, Kawaoka Y. Influenza: lessons from past pandemics, warnings from current incidents. Nature Reviews Microbiology, 2005, 3(8): 591-600
8. Kuiken T, Rimmelzwaan G, van Riel D, van Amerongen G, Baars M, Fouchier R, Osterhaus A. Avian H5N1 influenza in cats. Science, 2004, 306(5694): 241
9. Quirk M. Zoo tigers succumb to avian influenza. Lancet Infectious Diseases, 2004, 4(12): 716
10. Songserm T, Amonsin A, Jam-on R, Sae-Heng N, Pariyothorn N, Payungporn S, Theamboonlers A, Chutinimitkul S, Thanawongnuwech R, Poovorawan Y. Fatal avian influenza A H5N1 in a dog. Emerging Infectious Diseases, 2006, 12(11): 1744-1747
11. Thanawongnuwech R, Amonsin A, Tantilertcharoen R, Damrongwatanapokin S, Theamboonlers A, Payungporn S, Nanthapornphiphat K, Ratanamungklanon S, Tunak E, Songserm T, Vivatthanavanich V, Lekdumrongsak T, Kesdangsakonwut S, Tunhikorn S, Poovorawan Y. Probable tiger-to-tiger transmission of avian influenza H5N1. Emerging Infectious Diseases, 2005, 11 (5): 699-701
12. Palese P. The genes of influenza virus. Cell, 1977, 10(1): 1-10
13. Lamb R A, Lai C J. Sequence of interrupted and uninterrupted mRNAs and cloned DNA coding for the two overlapping nonstructural proteins of influenza virus. Cell, 1980, 21(2): 475-485
14. Lamb R A, Lai C J, Choppin P W. Sequences of mRNAs derived from genome RNA segment 7 of influenza virus: colinear and interrupted mRNAs code for overlapping proteins. Proceedings of the National Academy of Sciences of the United States of America, 1981, 78(7): 4170-4174
15. Wise H M, Foeglein A, Sun J, Dalton R M, Patel S, Howard W, Anderson E C, Barclay W S, Digard P. A complicated message: identification of a novel PB1-related protein translated from influenza A virus segment 2 mRNA. Journal of Virology, 2009, 83 (16): 8021-8031
16. Chen W, Calvo P A, Malide D, Gibbs J, Schubert U, Bacik I, Basta S, O'Neill R, Schickli J, Palese P, Henklein P, Bennink J R, Yewdell J W. A novel influenza A virus mitochondrial protein that induces cell death. Nature Medicine, 2001, 7(12): 1306-1312
17. Jagger BW, Wise HM, Kash J C, Walters K A, Wills NM, Xiao Y L, Dunfee R L, Schwartzman L M, Ozinsky A, Bell G L, Dalton R M, Lo A, Efstathiou S, Atkins J F, Firth A E, Taubenberger J K, Digard P. An overlapping protein-coding region in influenza A virus segment 3 modulates the host response. Science, 2012, 337 (6091): 199-204
18. Wise H M, Hutchinson E C, Jagger B W, Stuart A D, Kang Z H, Robb N, Schwartzman LM, Kash J C, Fodor E, Firth A E, Gog J R, Taubenberger J K, Digard P. Identification of a novel splice variant form of the influenza A virus M2 ion channel with an antigenically distinct ectodomain. PLoS Pathogens, 2012, 8(11): e1002998
19. Selman M, Dankar S K, Forbes N E, Jia J J, Brown E G. Adaptive mutation in influenza A virus non-structural gene is linked to host switching and induces a novel protein by alternative splicing. Emerging Microbes & Infections, 2012, 1(11): e42
20. Muramoto Y, Noda T, Kawakami E, Akkina R, Kawaoka Y. Identification of novel influenza A virus proteins translated from PA mRNA. Journal of Virology, 2013, 87(5): 2455-2462
21. Vasin A V, Temkina O A, Egorov V V, Klotchenko S A, Plotnikova M A, Kiselev O I. Molecular mechanisms enhancing the proteome of influenza A viruses: an overview of recently discovered proteins. Virus Research, 2014, 185: 53-63
22. Yamayoshi S, Watanabe M, Goto H, Kawaoka Y. Identification of a novel viral protein expressed from the PB2 segment of influenza A virus. Journal of Virology, 2016, 90(1): 444-456
23. Chen B L, Zhang Z J, Chen W B. Isolation and preliminary serological characterization of type A influenza viruses from chickens. Journal of Veterinary Medicine, 1994, 22(2): 3-5
24. Xu X, Subbarao K, Cox N J, Guo Y. Genetic characterization of the pathogenic influenza A/Goose/Guangdong/1/96 (H5N1) virus: similarity of its hemagglutinin gene to those of H5N1 viruses from the 1997 outbreaks in Hong Kong. Virology, 1999, 261(1): 15-19
25. Zhou J, Wang D, Gao R, Zhao B, Song J, Qi X, Zhang Y, Shi Y, Yang L, Zhu W, Bai T, Qin K, Lan Y, Zou S, Guo J, Dong J, Dong L, Wei H, Li X, Lu J, Liu L, Zhao X, Huang W, Wen L, Bo H, Xin L, Chen Y, Xu C, Pei Y, Yang Y, Zhang X, Wang S, Feng Z, Han J, Yang W, Gao G F, Wu G, Li D, Wang Y, Shu Y. Biological features of novel avian influenza A (H7N9) virus. Nature, 2013, 499(7459): 500-503
26. Tan K X, Jacob S A, Chan K G, Lee L H. An overview of the characteristics of the novel avian influenza A H7N9 virus in humans. Frontiers in Microbiology, 2015, 6: 140
27. Bi Y, Zhang Z, Liu W, Yin Y, Hong J, Li X, Wang H, Wong G, Chen J, Li Y, Ru W, Gao R, Liu D, Liu Y, Zhou B, Gao G F, Shi W, Lei F. Highly Pathogenic Avian Influenza A(H5N1) Virus Struck Migratory Birds in China in 2015. Scientific Reports, 2015, 5: 12986
28. Xu L, Bao L, Yuan J, Li F, Lv Q, Deng W, Xu Y, Yao Y, Yu P, Chen H, Yuen K Y, Qin C. Antigenicity and transmissibility of a novel clade 2.3.2.1 avian influenza H5N1 virus. Journal of General Virology, 2013, 94(12): 2616-2626
29. Garten R J, Davis C T, Russell C A, Shu B, Lindstrom S, Balish A, Sessions W M, Xu X, Skepner E, Deyde V, Okomo-Adhiambo M, Gubareva L, Barnes J, Smith C B, Emery S L, Hillman M J, Rivailler P, Smagala J, de Graaf M, Burke D F, Fouchier R A, Pappas C, Alpuche-Aranda C M, Lopez-Gatell H, Olivera H, Lopez I, Myers C A, Faix D, Blair P J, Yu C, Keene KM, Dotson P D Jr, Boxrud D, Sambol A R, Abid S H, St George K, Bannerman T, Moore A L, Stringer D J, Blevins P, Demmler-Harrison G J, Ginsberg M, Kriner P, Waterman S, Smole S, Guevara H F, Belongia E A, Clark P A, Beatrice S T, Donis R, Katz J, Finelli L, Bridges C B, Shaw M, Jernigan D B, Uyeki T M, Smith D J, Klimov A I, Cox N J. Antigenic and genetic characteristics of swine-origin 2009 A (H1N1) influenza viruses circulating in humans. Science, 2009, 325(5937): 197-201
30. Lee A J, Das S R, Wang W, Fitzgerald T, Pickett B E, Aevermann B D, Topham D J, Falsey A R, Scheuermann R H. Diversifying selection analysis predicts antigenic evolution of 2009 pandemic H1N1 influenza A virus in humans. Journal of Virology, 2015, 89 (10): 5427-5440
31. Cui L, Liu D, Shi W, Pan J, Qi X, Li X, Guo X, Zhou M, Li W, Li J, Haywood J, Xiao H, Yu X, Pu X, Wu Y, Yu H, Zhao K, Zhu Y, Wu B, Jin T, Shi Z, Tang F, Zhu F, Sun Q, Wu L, Yang R, Yan J, Lei F, Zhu B, Liu W, Ma J, Wang H, Gao G F. Dynamic reassortments and genetic heterogeneity of the human-infecting influenza A (H7N9) virus. Nature Communications, 2014, 5: 3142
32. Su S, Bi Y H, Wong G, Gray G C, Gao G F, Li S. Epidemiology, evolution, and recent outbreaks of avian influenza virus in China. Journal of Virology, 2015, 89(17): 8671-8676
33. Shi W, Li W, Li X, Haywood J, Ma J, Gao G F, Liu D. Phylogenetics of varied subtypes of avian influenza viruses in China: potential threat to humans. Protein & Cell, 2014, 5(4): 253-257
34. Guo Y J, Krauss S, Senne D A, Mo I P, Lo K S, Xiong X P, Norwood M, Shortridge K F, Webster R G, Guan Y. Characterization of the pathogenicity of members of the newly established H9N2 influenza virus lineages in Asia. Virology, 2000, 267(2): 279-288
35. Li K S, Xu K M, Peiris J S, Poon L L, Yu K Z, Yuen K Y, Shortridge K F, Webster R G, Guan Y. Characterization of H9 subtype influenza viruses from the ducks of southern China: a candidate for the next influenza pandemic in humans? Journal of Virology, 2003, 77(12): 6988-6994
36. Liu H, Liu X, Cheng J, Peng D, Jia L, Huang Y. Phylogenetic analysis of the hemagglutinin genes of twenty-six avian influenza viruses of subtype H9N2 isolated from chickens in China during 1996-2001. Avian Diseases, 2003, 47(1): 116-127
37. Xu K M, Li K S, Smith G J, Li J W, Tai H, Zhang J X, Webster R G, Peiris J S, Chen H, Guan Y. Evolution and molecular epidemiology of H9N2 influenza A viruses from quail in southern China, 2000 to 2005. Journal of Virology, 2007, 81(6): 2635-2645
38. Lu J H, Liu X F, Shao W X, Liu Y L, Wei D P, Liu H Q. Phylogenetic analysis of eight genes of H9N2 subtype influenza virus: a mainland China strain possessing early isolates' genes that have been circulating. Virus Genes, 2005, 31(2): 163-169
39. Zhang P, Tang Y, Liu X, Peng D, Liu W, Liu H, Lu S, Liu X. Characterization of H9N2 influenza viruses isolated from vaccinated flocks in an integrated broiler chicken operation in eastern China during a 5 year period (1998-2002). Journal of General Virology, 2008, 89(12): 3102-3112
40. Gu M, Chen H, Li Q, Huang J, Zhao M, Gu X, Jiang K, Wang X, Peng D, Liu X. Enzootic genotype S of H9N2 avian influenza viruses donates internal genes to emerging zoonotic influenza viruses in China. Veterinary Microbiology, 2014, 174(3-4): 309-315
41. Wang Q, Ju L, Liu P, Zhou J, Lv X, Li L, Shen H, Su H, Jiang L, Jiang Q. Serological and virological surveillance of avian influenza A virus H9N2 subtype in humans and poultry in Shanghai, China, between 2008 and 2010. Zoonoses and Public Health, 2015, 62(2): 131-140
42. Wang N, Ruan Z, Wan Y, Wang B, Zhang S H, Ge X Y. Identification of a novel strain of influenza A (H9N2) virus in chicken. Virologica Sinica, 2015, 30(4): 309-312
43. Bi Y, Lu L, Li J, Yin Y, Zhang Y, Gao H, Qin Z, Zeshan B, Liu J, Sun L, Liu W. Novel genetic reassortants in H9N2 influenza A viruses and their diverse pathogenicity to mice. Virology Journal, 2011, 8(1): 505
44. Pu J, Wang S, Yin Y, Zhang G, Carter R A, Wang J, Xu G, Sun H, Wang M, Wen C, Wei Y, Wang D, Zhu B, Lemmon G, Jiao Y, Duan S, Wang Q, Du Q, Sun M, Bao J, Sun Y, Zhao J, Zhang H, Wu G, Liu J, Webster R G. Evolution of the H9N2 influenza genotype that facilitated the genesis of the novel H7N9 virus. Proceedings of the National Academy of Sciences of the United States of America, 2015, 112(2): 548-553
45. Li Q, Zhou L, Zhou M, Chen Z, Li F, Wu H, Xiang N, Chen E, Tang F, Wang D, Meng L, Hong Z, Tu W, Cao Y, Li L, Ding F, Liu B, Wang M, Xie R, Gao R, Li X, Bai T, Zou S, He J, Hu J, Xu Y, Chai C, Wang S, Gao Y, Jin L, Zhang Y, Luo H, Yu H, He J, Li Q, Wang X, Gao L, Pang X, Liu G, Yan Y, Yuan H, Shu Y, Yang W, Wang Y, Wu F, Uyeki T M, Feng Z. Epidemiology of human infections with avian influenza A(H7N9) virus in China. New England Journal of Medicine, 2014, 370(6): 520-532
46. Wu A, Su C, Wang D, Peng Y, Liu M, Hua S, Li T, Gao G F, Tang H, Chen J, Liu X, Shu Y, Peng D, Jiang T. Sequential reassortments underlie diverse influenza H7N9 genotypes in China. Cell Host & Microbe, 2013, 14(4): 446-452
47. Liu M, Li X, Yuan H, Zhou J, Wu J, Bo H, Xia W, Xiong Y, Yang L, Gao R, Guo J, Huang W, Zhang Y, Zhao X, Zou X, Chen T, Wang D, Li Q, Wang SW, Chen S, Hu M, Ni X, Gong T, Shi Y, Li J, Zhou J, Cai J, Xiao Z, Zhang W, Sun J, Li D, Wu G, Feng Z, Wang Y, Chen H, Shu Y. Genetic diversity of avian influenza A (H10N8) virus in live poultry markets and its association with human infections in China. Scientific Reports, 2015, 5: 7632
48. Alexander D J, Brown I H. History of highly pathogenic avian influenza. Revue Scientifique et Technique, 2009, 28(1): 19-38
49. Guo Y, Xu X, Wan X. Genetic characterization of an avian influenza A (H5N1) virus isolated from a sick goose in China. Chinese Journal of Experimental and Clinical Virology, 1998, 12 (4): 322-325 (in Chinese)
50. Chin P S, Hoffmann E, Webby R, Webster R G, Guan Y, Peiris M, Shortridge K F. Molecular evolution of H6 influenza viruses from poultry in Southeastern China: prevalence of H6N1 influenza viruses possessing seven A/Hong Kong/156/97 (H5N1)-like genes in poultry. Journal of Virology, 2002, 76(2): 507-516
51. Hu J, Hu Z, Song Q, Gu M, Liu X, Wang X, Hu S, Chen C, Liu H, Liu W, Chen S, Peng D, Liu X. The PA-gene-mediated lethal dissemination and excessive innate immune response contribute to the high virulence of H5N1 avian influenza virus in mice. Journal of Virology, 2013, 87(5): 2660-2672
52. Li C, Yu K, Tian G, Yu D, Liu L, Jing B, Ping J, Chen H. Evolution of H9N2 influenza viruses from domestic poultry in Mainland China. Virology, 2005, 340(1): 70-83
53. Smith G J D, Donis R O. Nomenclature updates resulting from the evolution of avian influenza A(H5) virus clades 2.1.3.2a, 2.2.1, and 2.3.4 during 2013-2014. Influenza and Other Respiratory Viruses, 2015, 9(5): 271-276
54. Gu M, Zhao G, Zhao K, Zhong L, Huang J, Wan H, Wang X, Liu W, Liu H, Peng D, Liu X. Novel variants of clade 2.3.4 highly pathogenic avian influenza A(H5N1) viruses, China. Emerging Infectious Diseases, 2013, 19(12): 2021-2024
55. Guan Y, Smith G J. The emergence and diversification of panzootic H5N1 influenza viruses. Virus Research, 2013, 178(1): 35-43
56. Chen J, Fang F, Yang Z, Liu X, Zhang H, Zhang Z, Zhang X, Chen Z. Characterization of highly pathogenic H5N1 avian influenza viruses isolated from poultry markets in central China. Virus Research, 2009, 146(1-2): 19-28
57. Guan Y, Peiris J S, Lipatov A S, Ellis T M, Dyrting K C, Krauss S, Zhang L J, Webster R G, Shortridge K F. Emergence of multiple genotypes of H5N1 avian influenza viruses in Hong Kong SAR. Proceedings of the National Academy of Sciences of the United States of America, 2002, 99(13): 8950-8955
58. Li K S, Guan Y, Wang J, Smith G J, Xu K M, Duan L, Rahardjo A P, Puthavathana P, Buranathai C, Nguyen T D, Estoepangestie A T, Chaisingh A, Auewarakul P, Long H T, Hanh N T, Webby R J, Poon L L, Chen H, Shortridge K F, Yuen K Y, Webster R G, Peiris J S. Genesis of a highly pathogenic and potentially pandemic H5N1 influenza virus in eastern Asia. Nature, 2004, 430(6996): 209-213
59. Chen H, Smith G J, Li K S, Wang J, Fan X H, Rayner J M, Vijaykrishna D, Zhang J X, Zhang L J, Guo C T, Cheung C L, Xu K M, Duan L, Huang K, Qin K, Leung Y H, Wu W L, Lu H R, Chen Y, Xia N S, Naipospos T S, Yuen K Y, Hassan S S, Bahri S, Nguyen T D, Webster R G, Peiris J S, Guan Y. Establishment of multiple sublineages of H5N1 influenza virus in Asia: implications for pandemic control. Proceedings of the National Academy of Sciences of the United States of America, 2006, 103(8): 2845-2850
60. Sonnberg S, Webby R J, Webster R G. Natural history of highly pathogenic avian influenza H5N1. Virus Research, 2013, 178(1): 63-77
61. Neumann G, Green M A, Macken C A. Evolution of highly pathogenic avian H5N1 influenza viruses and the emergence of dominant variants. Journal of General Virology, 2010, 91(8): 1984-1995
62. Gu M, Liu W, Cao Y, Peng D, Wang X, Wan H, Zhao G, Xu Q, Zhang W, Song Q, Li Y, Liu X. Novel reassortant highly pathogenic avian influenza (H5N5) viruses in domestic ducks, China. Emerging Infectious Diseases, 2011, 17(6): 1060-1063
63. Zhao G, Gu X, Lu X, Pan J, Duan Z, Zhao K, Gu M, Liu Q, He L, Chen J, Ge S, Wang Y, Chen S, Wang X, Peng D, Wan H, Liu X. Novel reassortant highly pathogenic H5N2 avian influenza viruses in poultry in China. PLoS ONE, 2012, 7(9): e46183
64. Hu J, Zhao K, Liu X, Wang X, Chen Z, Liu X. Two highly pathogenic avian influenza H5N1 viruses of clade 2.3.2.1 with similar genetic background but with different pathogenicity in mice and ducks. Transboundary and Emerging Diseases, 2013, 60 (2): 127-139
65. de Vries E, Guo H, Dai M, Rottier P J, van Kuppeveld F J, de Haan C A. Rapid emergence of highly pathogenic avian influenza subtypes from a subtype H5N1 hemagglutinin variant. Emerging Infectious Diseases, 2015, 21(5): 842-846
66. Zhang P, Tang Y, Liu X, Liu W, Zhang X, Liu H, Peng D, Gao S, Wu Y, Zhang L, Lu S, Liu X. A novel genotype H9N2 influenza virus possessing human H5N1 internal genomes has been circulating in poultry in eastern China since 1998. Journal of Virology, 2009, 83(17): 8428-8438
67. Gu M, Huang J, Chen Y, Chen J, Wang X, Liu X, Liu X. Genome sequence of a natural reassortant H5N2 avian influenza virus from domestic mallard ducks in eastern China. Journal of Virology, 2012, 86(22): 12463-12464
68. Fan S, Zhou L, Wu D, Gao X, Pei E, Wang T, Gao Y, Xia X. A novel highly pathogenic H5N8 avian influenza virus isolated from a wild duck in China. Influenza and Other Respiratory Viruses, 2014, 8(6): 646-653
69. Wu H, Peng X, Xu L, Jin C, Cheng L, Lu X, Xie T, Yao H, Wu N. Novel reassortant influenza A (H5N8) viruses in domestic ducks, eastern China. Emerging Infectious Diseases, 2014, 20(8): 1315-1318
70. Bi Y, Mei K, Shi W, Liu D, Yu X, Gao Z, Zhao L, Gao G F, Chen J, Chen Q. Two novel reassortants of avian influenza A (H5N6) virus in China. Journal of General Virology, 2015, 96(5): 975-981
71. Homme P J, Easterday B C. Avian influenza virus infections. I. Characteristics of influenza A-turkey-Wisconsin-1966 virus. Avian Diseases, 1970, 14(1): 66-74
72. Kawaoka Y, Chambers T M, Sladen W L, Gwebster R. Is the gene pool of influenza viruses in shorebirds and gulls different from that in wild ducks? Virology, 1988, 163(1): 247-250
73. Alexander D J. A review of avian influenza in different bird species. Veterinary Microbiology, 2000, 74(1-2): 3-13
74. Arafa A S, Hagag NM, Yehia N, Zanaty A M, Naguib MM, Nasef S A. Effect of cocirculation of highly pathogenic avian influenza H5N1 subtype with low pathogenic H9N2 subtype on the spread of infections. Avian Diseases, 2012, 56(4 s1): 849-857
75. Butt K M, Smith G J, Chen H, Zhang L J, Leung Y H, Xu K M, Lim W, Webster R G, Yuen K Y, Peiris J S, Guan Y. Human infection with an avian H9N2 influenza A virus in Hong Kong in 2003. Journal of Clinical Microbiology, 2005, 43(11): 5760-5767
76. Cong Y L, Pu J, Liu Q F, Wang S, Zhang G Z, Zhang X L, Fan W X, Brown E G, Liu J H. Antigenic and genetic characterization of H9N2 swine influenza viruses in China. Journal of General Virology, 2007, 88(7): 2035-2041
77. Cong Y L, Wang C F, Yan C M, Peng J S, Jiang Z L, Liu J H. Swine infection with H9N2 influenza viruses in China in 2004. Virus Genes, 2008, 36(3): 461-469
78. Lin Y P, Shaw M, Gregory V, Cameron K, Lim W, Klimov A, Subbarao K, Guan Y, Krauss S, Shortridge K, Webster R, Cox N, Hay A. Avian-to-human transmission of H9N2 subtype influenza A viruses: Relationship between H9N2 and H5N1 human isolates. Proceedings of the National Academy of Sciences of the United States of America, 2000, 97(17): 9654-9658
79. Peiris J S M, Guan Y, Markwell D, Ghose P, Webster R G, Shortridge K F. Cocirculation of avian H9N2 and contemporary 'human' H3N2 influenza A viruses in pigs in southeastern China: potential for genetic reassortment? Journal of Virology, 2001, 75 (20): 9679-9686
80. Peiris M, Yuen K Y, Leung C W, Chan K H, Ip P L S, Lai RW M, Orr W K, Shortridge K F. Human infection with influenza H9N2. Lancet, 1999, 354(9182): 916-917
81. Huang Y W, Li X D, Zhang H, Chen B Z, Jiang Y L, Yang L, Zhu W F, Hu S X, Zhou S Y, Tang Y L, Xiang X Y, Li F C, Li W C, Gao L D. Human infection with an avian influenza A (H9N2) virus in the middle region of China. Journal of Medical Virology, 2015, 87(10): 1641-1648
82. Saito T, Lim W, Suzuki T, Suzuki Y, Kida H, Nishimura S I, Tashiro M. Characterization of a human H9N2 influenza virus isolated in Hong Kong. Vaccine, 2001, 20(1-2): 125-133
83. ShiWF, GibbsMJ, Zhang Y Z, Zhang Z, Zhao XM, Jin X, Zhu C D, Yang M F, Yang N N, Cui Y J, Ji L. Genetic analysis of four porcine avian influenza viruses isolated from Shandong, China. Archives of Virology, 2008, 153(1): 211-217
84. Xu C T, FanWX, Wei R, Zhao H K. Isolation and identification of swine influenza recombinant A/Swine/Shandong/1/2003(H9N2) virus. Microbes and Infection, 2004, 6(10): 919-925
85. Yu H, Zhou Y J, Li G X, Ma J H, Yan L P, Wang B, Yang F R, Huang M, Tong G Z. Genetic diversity of H9N2 influenza viruses from pigs in China: a potential threat to human health? Veterinary Microbiology, 2011, 149(1-2): 254-261
86. Cheng V C, Chan J F, Wen X, WuWL, Que T L, Chen H, Chan K H, Yuen K Y. Infection of immunocompromised patients by avian H9N2 influenza A virus. Journal of Infection, 2011, 62(5): 394-399
87. Kim J A, Cho S H, Kim H S, Seo S H. H9N2 influenza viruses isolated from poultry in Korean live bird markets continuously evolve and cause the severe clinical signs in layers. Veterinary Microbiology, 2006, 118(3-4): 169-176
88. Sun Y, Pu J, Jiang Z, Guan T, Xia Y, Xu Q, Liu L, Ma B, Tian F, Brown E G, Liu J. Genotypic evolution and antigenic drift of H9N2 influenza viruses in China from 1994 to 2008. Veterinary Microbiology, 2010, 146(3-4): 215-225
89. Liu J H, Okazaki K, Shi W M, Wu Q M, Mweene A S, Kida H. Phylogenetic analysis of neuraminidase gene of H9N2 influenza viruses prevalent in chickens in China during 1995-2002. Virus Genes, 2003, 27(2): 197-202
90. Aamir U B, Wernery U, Ilyushina N, Webster R G. Characterization of avian H9N2 influenza viruses from United Arab Emirates 2000 to 2003. Virology, 2007, 361(1): 45-55
91. Wu R, Sui Z W, Zhang H B, Chen Q J, Liang W W, Yang K L, Xiong Z L, Liu Z W, Chen Z, Xu D P. Characterization of a pathogenic H9N2 influenza A virus isolated from central China in 2007. Archives of Virology, 2008, 153(8): 1549-1555
92. Deng G, Bi J, Kong F, Li X, Xu Q, Dong J, Zhang M, Zhao L, Luan Z, Lv N, Qiao J. Acute respiratory distress syndrome induced by H9N2 virus in mice. Archives of Virology, 2010, 155(2): 187-195
93. Liu L, Zi L, Zhou J, Zhu Y, Dong J, Zhao X, Guo J, Shu Y. Pathogenesis and immunogenicity of an avian H9N2 influenza virus isolated from human. Biomedical and Environmental Sciences, 2011, 24(5): 530-536
94. Li X, Qi W, He J, Ning Z, Hu Y, Tian J, Jiao P, Xu C, Chen J, Richt J, Ma W, Liao M. Molecular basis of efficient replication and pathogenicity of H9N2 avian influenza viruses in mice. PLoS ONE, 2012, 7(6): e40118
95. Lin Z Q, Xu C T, Liu B, Ji Y H, Fu Y G, Guo J H, Zhu Q Y. Analysis of the phylogeny of Chinese H9N2 avian influenza viruses and their pathogenicity in mice. Archives of Virology, 2014, 159(10): 2575-2586
96. Li Y, Shan Y F, Chi Y, Wen T, Han X D. Acute lung injury induced by H9N2 virus in mice. Chinese Medical Journal, 2014, 127(20): 3576-3580
97. Mok C K, Yen H L, YuMY, Yuen KM, Sia S F, ChanMC, Qin G, Tu W W, Peiris J S. Amino acid residues 253 and 591 of the PB2 protein of avian influenza virus A H9N2 contribute to mammalian pathogenesis. Journal of Virology, 2011, 85(18): 9641-9645
98. Wang J, Sun Y, Xu Q, Tan Y, Pu J, Yang H, Brown E G, Liu J. Mouse-adapted H9N2 influenza A virus PB2 protein M147L and E627K mutations are critical for high virulence. PLoS ONE, 2012, 7(7): e40752
99. Liu Q T, Huang J Q, Chen Y X, Chen H Z, Li Q H, He L, Hao X L, Liu J J, Gu M, Hu J, Wang X Q, Hu S L, Liu X W, Liu X F. Virulence Determinants in the PB2 Gene of a Mouse-Adapted H9N2 Virus. Journal of Virology, 2015, 89(1): 877-882
100. Sun Y, Tan Y, Wei K, Sun H, Shi Y, Pu J, Yang H, Gao G F, Yin Y, Feng W, Perez D R, Liu J. Amino acid 316 of hemagglutinin and the neuraminidase stalk length influence virulence of H9N2 influenza virus in chickens and mice. Journal of Virology, 2013, 87(5): 2963-2968
101. Zhang K, Zhang Z, Yu Z, Li L, Cheng K, Wang T, Huang G, Yang S, Zhao Y, Feng N, Fu J, Qin C, Gao Y, Xia X. Domestic cats and dogs are susceptible to H9N2 avian influenza virus. Virus Research, 2013, 175(1): 52-57
102. Zhou P, Wang L, Huang S, Fu C, He H, Hong M, Su S, Li S. Beagle dogs have low susceptibility to BJ94-like H9N2 avian influenza virus. Infection, Genetics and Evolution, 2015, 31: 216-220
103. Wan H, Sorrell E M, Song H, Hossain M J, Ramirez-Nieto G, Monne I, Stevens J, Cattoli G, Capua I, Chen L M, Donis R O, Busch J, Paulson J C, Brockwell C, Webby R, Blanco J, Al-Natour M Q, Perez D R. Replication and transmission of H9N2 influenza viruses in ferrets: evaluation of pandemic potential. PLoS ONE, 2008, 3(8): e2923
104. Gao R, Bai T, Li X, Xiong Y, Huang Y, Pan M, Zhang Y, Bo H, Zou S, Shu Y. The comparison of pathology in ferrets infected by H9N2 avian influenza viruses with different genomic features. Virology, 2016, 488: 149-155
105. Sun Y, Bi Y, Pu J, Hu Y, Wang J, Gao H, Liu L, Xu Q, Tan Y, Liu M, Guo X, Yang H, Liu J. Guinea pig model for evaluating the potential public health risk of swine and avian influenza viruses. PLoS ONE, 2010, 5(11): e15537
106. Zhang K, Xu W, Zhang Z, Wang T, Sang X, Cheng K, Yu Z, Zheng X, Wang H, Zhao Y, Huang G, Yang S, Qin C, Gao Y, Xia X. Experimental infection of non-human primates with avian influenza virus (H9N2). Archives of Virology, 2013, 158(10): 2127-2134
107. Bonfante F, Patrono L V, Aiello R, BeatoMS, Terregino C, Capua I. Susceptibility and intra-species transmission of the H9N2 G1 prototype lineage virus in Japanese quail and turkeys. Veterinary Microbiology, 2013, 165(1-2): 177-183
108. Nili H, Mohammadi A, Habibi H, Firouzi S. Pathogenesis of H9N2 virus in Chukar partridges. Avian Pathology, 2013, 42(3): 230-234
109. Iqbal M, Yaqub T, Mukhtar N, Shabbir M Z, McCauley J W. Infectivity and transmissibility of H9N2 avian influenza virus in chickens and wild terrestrial birds. Veterinary Research, 2013, 44 (1): 100
110. Zhang C, Xuan Y, Shan H, Yang H, Wang J, Wang K, Li G, Qiao J. Avian influenza virus H9N2 infections in farmed minks. Virology Journal, 2015, 12(1): 180
111. Li P, Chen C, Han K, Zhang F, Zhu Y, Ling Z, Zhang X, Jiang S, Xie Z. Molecular characterization of H9N2 influenza virus isolated from mink and its pathogenesis in mink. Veterinary Microbiology, 2015, 176(1-2): 88-96
112. Perkins L E, Swayne D E. Pathobiology of A/chicken/Hong Kong/220/97 (H5N1) avian influenza virus in seven gallinaceous species. Veterinary Pathology, 2001, 38(2): 149-164
113. Alexander D J, Allan W H, Parsons D G, Parsons G. The pathogenicity of four avian influenza viruses for fowls, turkeys and ducks. Research in Veterinary Science, 1978, 24(2): 242-247
114. Alexander D J, Parsons G, Manvell R J. Experimental assessment of the pathogenicity of eight avian influenza A viruses of H5 subtype for chickens, turkeys, ducks and quail. Avian Pathology, 1986, 15(4): 647-662
115. Wood G W, Parsons G, Alexander D J. Replication of influenza A viruses of high and low pathogenicity for chickens at different sites in chickens and ducks following intranasal inoculation. Avian Pathology, 1995, 24(3): 545-551
116. Ellis T M, Barry Bousfield R, Bissett L A, Dyrting K C, Luk G S M, Tsim S T, Sturm-ramirez K, Webster R G, Guan Y, Peiris J S M. Investigation of outbreaks of highly pathogenic H5N1 avian influenza in waterfowl and wild birds in Hong Kong in late 2002. Avian Pathology, 2004, 33(5): 492-505
117. Sturm-Ramirez K M, Ellis T, Bousfield B, Bissett L, Dyrting K, Rehg J E, Poon L, Guan Y, Peiris M, Webster R G. Reemerging H5N1 influenza viruses in Hong Kong in 2002 are highly pathogenic to ducks. Journal of Virology, 2004, 78(9): 4892-4901
118. Swayne D E. Understanding the complex pathobiology of high pathogenicity avian influenza viruses in birds. Avian Diseases, 2007, 51(s1): 242-249
119. Liu J, Xiao H, Lei F, Zhu Q, Qin K, Zhang XW, Zhang X L, Zhao D, Wang G, Feng Y, Ma J, Liu W, Wang J, Gao G F. Highly pathogenic H5N1 influenza virus infection in migratory birds. Science, 2005, 309(5738): 1206
120. Hu J, Hu Z, Mo Y, Wu Q, Cui Z, Duan Z, Huang J, Chen H, Chen Y, Gu M, Wang X, Hu S, Liu H, Liu W, Liu X, Liu X. The PA and HA gene-mediated high viral load and intense innate immune response in the brain contribute to the high pathogenicity of H5N1 avian influenza virus in mallard ducks. Journal of Virology, 2013, 87(20): 11063-11075
121. Kajihara M, Sakoda Y, Soda K, Minari K, Okamatsu M, Takada A, Kida H. The PB2, PA, HA, NP, and NS genes of a highly pathogenic avian influenza virus A/whooper swan/Mongolia/3/2005 (H5N1) are responsible for pathogenicity in ducks. Virology Journal, 2013, 10(1): 45
122. Song J, Feng H, Xu J, Zhao D, Shi J, Li Y, Deng G, Jiang Y, Li X, Zhu P, Guan Y, Bu Z, Kawaoka Y, Chen H. The PA protein directly contributes to the virulence of H5N1 avian influenza viruses in domestic ducks. Journal of Virology, 2011, 85(5): 2180-2188
123. Hulse-Post D J, Franks J, Boyd K, Salomon R, Hoffmann E, Yen H L, Webby R J, Walker D, Nguyen T D, Webster R G. Molecular changes in the polymerase genes (PA and PB1) associated with high pathogenicity of H5N1 influenza virus in mallard ducks. Journal of Virology, 2007, 81(16): 8515-8524
124. Bröjer C van Amerongen G, van de Bildt M, van Run P, Osterhaus A, Gavier-Widen D, Kuiken T. Pathogenicity and tissue tropism of currently circulating highly pathogenic avian influenza A virus (H5N1; clade 2.3.2) in tufted ducks (Aythya fuligula). Veterinary Microbiology, 2015, 180(3-4): 273-280
125. Leigh Perkins L E, Swayne D E. Pathogenicity of a Hong Kongorigin H5N1 highly pathogenic avian influenza virus for emus, geese, ducks, and pigeons. Avian Diseases, 2002, 46(1): 53-63
126. Brown J D, Stallknecht D E, Beck J R, Suarez D L, Swayne D E. Susceptibility of North American ducks and gulls to H5N1 highly pathogenic avian influenza viruses. Emerging Infectious Diseases, 2006, 12(11): 1663-1670
127. Brown J D, Stallknecht D E, Swayne D E. Experimental infection of swans and geese with highly pathogenic avian influenza virus (H5N1) of Asian lineage. Emerging Infectious Diseases, 2008, 14 (1): 136-142
128. Pasick J, Berhane Y, Embury-Hyatt C, Copps J, Kehler H, Handel K, Babiuk S, Hooper-McGrevy K, Li Y, Le Q M, Phuong S L. Susceptibility of Canada Geese (Branta canadensis) to highly pathogenic avian influenza virus (H5N1). Emerging Infectious Diseases, 2007, 13(12): 1821-1827
129. Pantin-Jackwood M J, Swayne D E. Pathobiology of Asian highly pathogenic avian influenza H5N1 virus infections in ducks. Avian Diseases, 2007, 51(s1): 250-259
130. Keawcharoen J, van Riel D, van Amerongen G, Bestebroer T, BeyerWE, van Lavieren R, Osterhaus A D M E, Fouchier R A M, Kuiken T. Wild ducks as long-distance vectors of highly pathogenic avian influenza virus (H5NI). Emerging Infectious Diseases, 2008, 14(4): 600-607
131. Zhao K, Gu M, Zhong L, Duan Z, Zhang Y, Zhu Y, Zhao G, Zhao M, Chen Z, Hu S, Liu W, Liu X, Peng D, Liu X. Characterization of three H5N5 and one H5N8 highly pathogenic avian influenza viruses in China. Veterinary Microbiology, 2013, 163(3-4): 351-357
132. Yu Z, Gao X, Wang T, Li Y, Li Y, Xu Y, Chu D, Sun H, Wu C, Li S, Wang H, Li Y, Xia Z, Lin W, Qian J, Chen H, Xia X, Gao Y. Fatal H5N6 Avian Influenza Virus Infection in a Domestic Cat and Wild Birds in China. Scientific Reports, 2015, 5: 10704
133. Wu H, Lu R, Peng X, Xu L, Cheng L, Lu X, Jin C, Xie T, Yao H, Wu N. Novel reassortant highly pathogenic H5N6 avian influenza viruses in poultry in China. Infection, Genetics and Evolution, 2015, 31: 64-67
134. Shen H, Wu B, Chen Y, Bi Y, Xie Q. Influenza A (H5N6) virus reassortant, Southern China, 2014. Emerging Infectious Diseases, 2015, 21(7): 1261-1262
135. Li J, Gu M, Liu D, Liu B, Jiang K, Zhong L, Liu K, Sun W, Hu J, Wang X, Hu S, Liu X. Phylogenetic and biological characterization of three K1203 (H5N8)-like avian influenza A virus reassortants in China in 2014. Archives of Virology, 2016, 161(2):289-302
136. Li Y, Shi J, Zhong G, Deng G, Tian G, Ge J, Zeng X, Song J, Zhao D, Liu L, Jiang Y, Guan Y, Bu Z, Chen H. Continued evolution of H5N1 influenza viruses in wild birds, domestic poultry, and humans in China from 2004 to 2009. Journal of Virology, 2010, 84 (17): 8389-8397
137. Pantin-Jackwood M, Swayne D E, Smith D, Shepherd E. Effect of species, breed and route of virus inoculation on the pathogenicity of H5N1 highly pathogenic influenza (HPAI) viruses in domestic ducks. Veterinary Research, 2013, 44: 62
138. Kwon Y K, Thomas C, Swayne D E. Variability in pathobiology of South Korean H5N1 high-pathogenicity avian influenza virus infection for 5 species of migratory waterfowl. Veterinary Pathology, 2010, 47(3): 495-506
139. Uno Y, Usui T, Soda K, Fujimoto Y, Takeuchi T, Ito H, Ito T, Yamaguchi T. The pathogenicity and host immune response associated with H5N1 highly pathogenic avian influenza virus in quail. Journal of Veterinary Medical Science, 2013, 75(4): 451-457
140. Isoda N, Sakoda Y, Kishida N, Bai G R, Matsuda K, Umemura T, Kida H. Pathogenicity of a highly pathogenic avian influenza virus, A/chicken/Yamaguchi/7/04 (H5N1) in different species of birds and mammals. Archives of Virology, 2006, 151(7): 1267-1279
141. Smietanka K, Minta Z, Reichert M, Olszewska M, Wyrostek K, Jozwiak M, van den Berg T. Experimental infection of juvenile domestic and Canada geese with two different clades of H5N1 high pathogenicity avian influenza virus. Veterinary Microbiology, 2013, 163(3-4): 235-241
142. Songserm T, Amonsin A, Jam-On R, Sae-Heng N, Meemak N, Pariyothorn N, Payungporn S, Theamboonlers A, Poovorawan Y. Avian influenza H5N1 in naturally infected domestic cat. Emerging Infectious Diseases, 2006, 12(4): 681-683
143. Kwon Y K, Lipatov A S, Swayne D E. Bronchointerstitial pneumonia in guinea pigs following inoculation with H5N1 high pathogenicity avian influenza virus. Veterinary Pathology, 2009, 46(1): 138-141
144. Li X, Jin M, Yu Z, Zhang A, Chen H, Qian P. Pathogenicity of a goose isolate of highly pathogenic H5N1 influenza a virus for chickens, mice, and pigs. Acta Virologica. English Ed., 2008, 52 (1): 41-46
145. Smietanka K, Minta Z, Wyrostek K, Jozwiak M, Olszewska M, Domanska-Blicharz K, Reichert M, Pikula A, Habyarimana A, van den Berg T. Susceptibility of pigeons to glade 1 and 2.2 high pathogenicity avian influenza H5N1 virus. Avian Diseases, 2011, 55(1): 106-112
146. Wu H, Peng X, Xu L, Jin C, Cheng L, Lu X, Xie T, Yao H, Wu N. Characterization of a novel highly pathogenic H5N2 avian influenza virus isolated from a duck in eastern China. Archives of Virology, 2014, 159(12): 3377-3383
147. Feng H X, Liu Y Y, Song Q Q, Ling Z S, Zhang F X, Zhu Y L, Jiang S J, Xie Z J. Interspecies transmission of canine influenza virus H5N2 to cats and chickens by close contact with experimentally infected dogs. Veterinary Microbiology, 2014, 170(3-4): 414-417
148. Pulit-Penaloza J A, Sun X, Creager H M, Zeng H, Belser J A, Maines T R, Tumpey T M. Pathogenesis and Transmission of Novel Highly Pathogenic Avian Influenza H5N2 and H5N8 Viruses in Ferrets and Mice. Journal of Virology, 2015, 89(20): 10286-10293
149. Song Q Q, Zhang F X, Liu J J, Ling Z S, Zhu Y L, Jiang S J, Xie Z J. Dog to dog transmission of a novel influenza virus (H5N2) isolated from a canine. Veterinary Microbiology, 2013, 161(3-4): 331-333
150. Wang D, Tang G, Huang Y, Yu C, Li S, Zhuang L, Fu L, Wang S, Li N, Li X, Yang L, Lan Y, Bai T, Shu Y. A returning migrant worker with avian influenza A (H7N9) virus infection in Guizhou, China: a case report. Journal of Medical Case Reports, 2015, 9(1): 109
151. Li X, Fu Y, Yang J, Guo J, He J, Weng S, Jia Y, Liu B, Zhu Q, Chen H. Genetic and biological characterization of two novel reassortant H5N6 swine influenza viruses in mice and chickens. Infection, Genetics and Evolution, 2015, 36: 462-466
152. Pan M, Gao R, Lv Q, Huang S, Zhou Z, Yang L, Li X, Zhao X, Zou X, Tong W, Mao S, Zou S, Bo H, Zhu X, Liu L, Yuan H, Zhang M, Wang D, Li Z, Zhao W, Ma M, Li Y, Li T, Yang H, Xu J, Zhou L, Zhou X, Tang W, Song Y, Chen T, Bai T, Zhou J, Wu G, Li D, Feng Z, Gao G F, Wang Y, He S, Shu Y. Human infection with a novel highly pathogenic avian influenza A (H5N6) virus: Virological and clinical findings. Journal of Infection, 2016, 72 (1):52-59
153. Bae Y J, Lee S B, Min K C, Mo J S, Jeon E O, Koo B S, Kwon H I, Choi Y K, Kim J J, Kim J N, Mo I P. Pathological evaluation of natural cases of a highly pathogenic avian influenza virus, subtype H5N8, in broiler breeders and commercial layers in South Korea. Avian Diseases, 2015, 59(1): 175-182
154. Kang H M, Lee E K, Song B M, Jeong J, Choi J G, Jeong J, Moon O K, Yoon H, Cho Y, Kang Y M, Lee H S, Lee Y J. Novel reassortant influenza A (H5N8) viruses among inoculated domestic and wild ducks, South Korea, 2014. Emerging Infectious Diseases, 2015, 21(2): 298-304
155. Carnaccini S, Crossley B, Breitmeyer R, Charlton B R, Bland M, Fowler K, De La Torre F, Torchetti M K, Wong S S, Wilson D, Jones A, Senties-Cue C G. Diagnosis and control of a LPAI H5N8 outbreak in a Japanese quail (Coturnix coturnix japonica) commercial flock in the central valley of California. Avian Diseases, 2015, 59(2): 344-348
156. Richard M, Herfst S, van den Brand J M A, Lexmond P, Bestebroer T M, Rimmelzwaan G F, Koopmans M, Kuiken T, Fouchier R A M. Low virulence and lack of airborne transmission of the Dutch highly pathogenic avian influenza virus H5N8 in Ferrets. PLoS ONE, 2015, 10(6): e0129827
157. Kim Y I, Pascua P N, Kwon H I, Lim G J, Kim E H, Yoon S W, Park S J, Kim SM, Choi E J, Si Y J, Lee O J, ShimWS, Kim SW, Mo I P, Bae Y, Lim Y T, Sung M H, Kim C J, Webby R J, Webster R G, Choi Y K. Pathobiological features of a novel, highly pathogenic avian influenza A (H5N8) virus. Emerging Microbes Infect, 2014, 3(10): e75
158. Song BM, Kang HM, Lee E K, Jung J, Kang Y, Lee H S, Lee Y J. Pathogenicity of H5N8 virus in chickens from Korea in 2014. Journal of Veterinary Science, 2015, 16(2): 237-240
159. Shi H, Ashraf S, Gao S, Lu J, Liu X. Evaluation of transmission route and replication efficiency of H9N2 avian influenza virus. Avian Diseases, 2010, 54(1): 22-27
160. Zhong L, Wang X, Li Q, Liu D, Chen H, Zhao M, Gu X, He L, Liu X, Gu M, Peng D, Liu X. Molecular mechanism of the airborne transmissibility of H9N2 avian influenza A viruses in chickens. Journal of Virology, 2014, 88(17): 9568-9578
161. Li X, Shi J, Guo J, Deng G, Zhang Q, Wang J, He X, Wang K, Chen J, Li Y, Fan J, Kong H, Gu C, Guan Y, Suzuki Y, Kawaoka Y, Liu L, Jiang Y, Tian G, Bu Z, Chen H. Genetics, receptor binding property, and transmissibility in mammals of naturally isolated H9N2 Avian Influenza viruses. PLoS Pathogens, 2014, 10 (11): e1004508
162. Wu R, Sui Z, Liu Z, Liang W, Yang K, Xiong Z, Xu D. Transmission of avian H9N2 influenza viruses in a murine model. Veterinary Microbiology, 2010, 142(3-4): 211-216
163. Lv J, Wei B, Yang Y, YaoM, Cai Y, Gao Y, Xia X, Zhao X, Liu Z, Li X, Wang H, Yang H, Roesler U, Miao Z, Chai T. Experimental transmission in guinea pigs of H9N2 avian influenza viruses from indoor air of chicken houses. Virus Research, 2012, 170(1-2): 102-108
164. Fusaro A, Monne I, Salviato A, Valastro V, Schivo A, Amarin N M, Gonzalez C, Ismail M M, Al-Ankari A R, Al-Blowi MH, Khan O A, Maken Ali A S, Hedayati A, Garcia Garcia J, Ziay G M, Shoushtari A, Al Qahtani K N, Capua I, Holmes E C, Cattoli G. Phylogeography and evolutionary history of reassortant H9N2 viruses with potential human health implications. Journal of Virology, 2011, 85(16): 8413-8421
165. Guan Y, Shortridge K F, Krauss S, Webster R G. Molecular characterization of H9N2 influenza viruses: Were they the donors of the 'internal' genes of H5N1 viruses in Hong Kong? Proceedings of the National Academy of Sciences of the United States of America, 1999, 96(16): 9363-9367
166. Kimble J B, Sorrell E, Shao H, Martin P L, Perez D R. Compatibility of H9N2 avian influenza surface genes and 2009 pandemic H1N1 internal genes for transmission in the ferret model. Proceedings of the National Academy of Sciences of the United States of America, 2011, 108(29): 12084-12088
167. Gao Y, Zhang Y, Shinya K, Deng G, Jiang Y, Li Z, Guan Y, Tian G, Li Y, Shi J, Liu L, Zeng X, Bu Z, Xia X, Kawaoka Y, Chen H. Identification of amino acids in HA and PB2 critical for the transmission of H5N1 avian influenza viruses in a mammalian host. PLoS Pathogens, 2009, 5(12): e1000709
168. Zhang Y, Zhang Q, Kong H, Jiang Y, Gao Y, Deng G, Shi J, Tian G, Liu L, Liu J, Guan Y, Bu Z, Chen H. H5N1 hybrid viruses bearing 2009/H1N1 virus genes transmit in guinea pigs by respiratory droplet. Science, 2013, 340(6139): 1459-1463
169. Herfst S, Schrauwen E J A, Linster M, Chutinimitkul S, de Wit E, Munster V J, Sorrell E M, Bestebroer T M, Burke D F, Smith D J, Rimmelzwaan G F, Osterhaus A D M E, Fouchier R A M. Airborne transmission of influenza A/H5N1 virus between ferrets. Science, 2012, 336(6088): 1534-1541
170. Imai M, Watanabe T, Hatta M, Das S C, Ozawa M, Shinya K, Zhong G X, Hanson A, Katsura H, Watanabe S, Li C J, Kawakami E, Yamada S, Kiso M, Suzuki Y, Maher E A, Neumann G, Kawaoka Y. Experimental adaptation of an influenza H5 HA confers respiratory droplet transmission to a reassortant H5 HA/H1N1 virus in ferrets. Nature, 2012, 486(7403): 420
171. Li Q, Wang X, Gu M, Zhu J, Hao X, Gao Z, Sun Z, Hu J, Hu S, Liu X. Novel H5 clade 2.3.4.6 viruses with both α-2, 3 and α-2, 6 receptor binding properties may pose a pandemic threat. Veterinary Research, 2014, 45(1): 127
172. Yu M, Qi W, Huang Z, Zhang K, Ye J, Liu R, Wang H, Ma Y, Liao M, Ning Z. Expression profile and histological distribution of IFITM1 and IFITM3 during H9N2 avian influenza virus infection in BALB/c mice. Medical Microbiology and Immunology, 2015, 204(4): 505-514
173. Bi J, Deng G, Dong J, Kong F, Li X, Xu Q, Zhang M, Zhao L, Qiao J. Phylogenetic and molecular characterization of H9N2 influenza isolates from chickens in Northern China from 2007-2009. PLoS ONE, 2010, 5(9): e13063
174. Chen F, Yan Z Q, Liu J, Ji J, Chang S, Liu D, Qin J P, Ma J Y, Bi Y Z, Xie Q M. Phylogenetic analysis of hemagglutinin genes of 40 H9N2 subtype avian influenza viruses isolated from poultry in China from 2010 to 2011. Virus Genes, 2012, 45(1): 69-75
175. Sun Y, Pu J, Fan L, Sun H, Wang J, Zhang Y, Liu L, Liu J. Evaluation of the protective efficacy of a commercial vaccine against different antigenic groups of H9N2 influenza viruses in chickens. Veterinary Microbiology, 2012, 156(1-2): 193-199
176. Swayne D E. Impact of vaccines and vaccination on global control of avian influenza. Avian Diseases, 2012, 56(4 s1): 818-828
177. Smith G J, Fan X H, Wang J, Li K S, Qin K, Zhang J X, Vijaykrishna D, Cheung C L, Huang K, Rayner J M, Peiris J S, Chen H, Webster R G, Guan Y. Emergence and predominance of an H5N1 influenza variant in China. Proceedings of the National Academy of Sciences of the United States of America, 2006, 103 (45): 16936-16941
178. Hu X, Liu D, Wang M, Yang L, Zhu Q, Li L, Gao G F. Clade 2.3.2 avian influenza virus (H5N1), Qinghai Lake region, China, 2009-2010. Emerging Infectious Diseases, 2011, 17(3): 560-562
179. Qin T, Yin Y, Yu Q, Huang L, Wang X, Lin J, Yang Q. CpG oligodeoxynucleotides facilitate delivery of whole inactivated H9N2 influenza virus via transepithelial dendrites of dendritic cells in nasal mucosa. Journal of Virology, 2015, 89(11): 5904-5918