Contributors and authorship Yi Jiang designed the study and conducted all the experiments, analysis and interpretation of the data, and wrote the manuscript. Sheng Zhou helped perform the recombinant IBV construction. Xu Cheng, Xiumei Zhao, Yan Yu, and Mingyan Gao participated in the whole animal experiment in different groups. Xu Cheng conducted the experiments involving viral shedding. All authors read and approved the final manuscript.