Neutrophils Are Needed for an Effective Immune Response Against Pulmonary Rat Coronavirus 1 Infection, but also Contribute to Pathology 2 3 Running Title: Neutrophils in rat coronavirus infection 4 Contents Category: Animal Viruses- Positive-Strand RNA 5 6 Anoria K. Haick1,2, Joanna P. Rzepka1,3, Elizabeth Brandon1, Onesmo B. Balemba1, and Tanya 7 A. Miura1* 8 1Department of Biological Sciences, University of Idaho, 875 Perimeter Dr., MS 3051, Moscow, 9 ID 83844-3051, USA 10 2Current Address: Department of Obstetrics and Gynecology, University of Washington School 11 of Medicine, 1959 Northeast Pacific Street, Seattle, WA 98195, USA 12 3Current Address: Veterinary Medical Research and Development Incorporated, 425 Northwest 13 Albion Drive, Pullman, WA 99163, USA 14 *Corresponding Author. Email: