About the Authors

Laura B Goodman

Affiliation Department of Microbiology and Immunology, Cornell University, Ithaca, New York, United States of America

Arianna Loregian

Affiliation Department of Histology, Microbiology and Medical Biotechnologies, University of Padua, Padua, Italy

Gillian A Perkins

Affiliation Department of Clinical Sciences, Cornell University, Ithaca, New York, United States of America

Josie Nugent

Affiliation Centre for Preventive Medicine, Animal Health Trust, Newmarket, United Kingdom

Elizabeth L Buckles

Affiliation Department of Biomedical Sciences, Cornell University, Ithaca, New York, United States of America

Beatrice Mercorelli

Affiliation Department of Histology, Microbiology and Medical Biotechnologies, University of Padua, Padua, Italy

Julia H Kydd

Current address: School of Veterinary Medicine and Science, University of Nottingham, Sutton Bonington, United Kingdom

Affiliation Centre for Preventive Medicine, Animal Health Trust, Newmarket, United Kingdom

Giorgio Palù

Affiliation Department of Histology, Microbiology and Medical Biotechnologies, University of Padua, Padua, Italy

Ken C Smith

Current address: Royal Veterinary College, Department of Pathology and Infectious Diseases, London, United Kingdom

Affiliation Centre for Preventive Medicine, Animal Health Trust, Newmarket, United Kingdom

Nikolaus Osterrieder

To whom correspondence should be addressed. E-mail: no34@cornell.edu

Affiliation Department of Microbiology and Immunology, Cornell University, Ithaca, New York, United States of America

Nicholas Davis-Poynter

Current address: Herpesvirus Molecular Pathogenesis Unit, Sir Albert Sakzewski Virus Research Centre, Royal Children's Hospital Clinical Medical Virology Centre, University of Queensland, Queensland, Australia

Affiliation Centre for Preventive Medicine, Animal Health Trust, Newmarket, United Kingdom

Competing Interests

The authors have declared that no competing interests exist.

Author Contributions

LBG performed the majority of the experiments, analyzed the data, and wrote the manuscript. AL, BM, and GP designed and performed the polymerase activity assays. ELB, KCS, and GAP performed postmortem examinations of animals. GAP, JN, and JHK participated in the animal studies, and GAP assisted with qPCR analyses. LBG, AL, GAP, JN, ELB, BM, JHK, GP, KCS, NO, and NDP revised the manuscript. NO and NDP share senior authorship.