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However, the role and mechanism of IBV and H9N2 AIV co-infection remain largely unknown. Specific-pathogen-free (SPF) chickens were inoculated with IBV 2 days before H9N2 virus inoculation (IBV\/H9N2); with IBV and H9N2 virus simultaneously (IBV+H9N2); with H9N2 virus 2 days before IBV inoculation (H9N2\/IBV); or with either IBV or H9N2 virus alone. Severe respiratory signs, pathological damage, and higher morbidity and mortality were observed in the co-infection groups compared with the IBV and H9N2 groups. In general, a higher virus load and a more intense inflammatory response were observed in the three co-infection groups, especially in the IBV\/H9N2 group. The same results were observed in the transcriptome analysis of the trachea of the SPF chickens. Therefore, IBV might play a major role in the development of respiratory disease in chickens, and secondary infection with H9N2 virus further enhances the pathogenicity by inducing a severe inflammatory response. These findings may provide a reference for the prevention and control of IBV and H9N2 AIV in the poultry industry and provide insight into the molecular mechanisms of IBV and H9N2 AIV co-infection in chickens.<\/jats:p>","DOI":"10.3389\/fvets.2021.824179","type":"journal-article","created":{"date-parts":[[2022,2,8]],"date-time":"2022-02-08T00:16:31Z","timestamp":1644279391000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":27,"title":["Infectious Bronchitis Virus Infection Increases Pathogenicity of H9N2 Avian Influenza Virus by Inducing Severe Inflammatory Response"],"prefix":"10.3389","volume":"8","author":[{"given":"Lingchen","family":"Kong","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Renrong","family":"You","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Dianchen","family":"Zhang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Qingli","family":"Yuan","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Bin","family":"Xiang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jianpeng","family":"Liang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Qiuyan","family":"Lin","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Chan","family":"Ding","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ming","family":"Liao","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Libin","family":"Chen","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Tao","family":"Ren","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1965","published-online":{"date-parts":[[2022,2,8]]},"reference":[{"key":"B1","doi-asserted-by":"publisher","first-page":"3995","DOI":"10.1128\/JVI.06540-11","article-title":"Discovery of seven novel Mammalian and avian coronaviruses in the genus deltacoronavirus supports bat coronaviruses as the gene source of alphacoronavirus and betacoronavirus and avian coronaviruses as the gene source of gammacoronavirus and deltacoronavirus","volume":"86","author":"Woo","year":"2012","journal-title":"J Virol."},{"key":"B2","doi-asserted-by":"publisher","first-page":"1517","DOI":"10.1007\/s00705-020-04628-0","article-title":"The emergence of SARS, MERS and novel SARS-2 coronaviruses in the 21st century","volume":"165","author":"da Costa","year":"2020","journal-title":"Arch Virol."},{"key":"B3","doi-asserted-by":"publisher","first-page":"1324","DOI":"10.3201\/eid2606.200299","article-title":"Co-infection with SARS-CoV-2 and influenza A virus in patient with pneumonia, China","volume":"26","author":"Wu","year":"2020","journal-title":"Emerg Infect Dis."},{"key":"B4","doi-asserted-by":"publisher","first-page":"e128","DOI":"10.1016\/j.jinf.2020.05.041","article-title":"Co-infection of SARS-CoV-2 and influenza virus in early stage of the COVID-19 epidemic in Wuhan, China","volume":"81","author":"Zheng","year":"2020","journal-title":"J Infect."},{"key":"B5","doi-asserted-by":"publisher","first-page":"1008","DOI":"10.1002\/jmv.26364","article-title":"High prevalence of SARS-CoV-2 and influenza A virus (H1N1) coinfection in dead patients in Northeastern Iran","volume":"93","author":"Hashemi","year":"2020","journal-title":"J Med Virol."},{"key":"B6","doi-asserted-by":"publisher","first-page":"57","DOI":"10.1099\/0022-1317-68-1-57","article-title":"Completion of the sequence of the genome of the coronavirus avian infectious bronchitis virus","volume":"68","author":"Boursnell","year":"1987","journal-title":"J Gen Virol"},{"key":"B7","doi-asserted-by":"publisher","first-page":"100","DOI":"10.1016\/j.vetmic.2016.08.017","article-title":"Pathogenicity of virulent infectious bronchitis virus isolate YN on hen ovary and oviduct","volume":"193","author":"Zhong","year":"2016","journal-title":"Vet Microbiol."},{"key":"B8","doi-asserted-by":"publisher","first-page":"2030","DOI":"10.3390\/ijms18102030","article-title":"Infectious bronchitis virus variants: molecular analysis and pathogenicity investigation. 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