About the Authors

Timothy Z Vitalis

Affiliations The Michael Smith Laboratories, University of British Columbia, Vancouver, British Columbia, Canada , Biomedical Research Centre, University of British Columbia, Vancouver, British Columbia, Canada , The Michael Smith Laboratories, University of British Columbia, Vancouver, British Columbia, Canada

Qian-Jin Zhang

Address Current address: Cellular Biology and Anatomy, LSU Health Sciences Center, Shreveport, Louisiana, United States of America

Affiliations The Michael Smith Laboratories, University of British Columbia, Vancouver, British Columbia, Canada , Biomedical Research Centre, University of British Columbia, Vancouver, British Columbia, Canada , The Michael Smith Laboratories, University of British Columbia, Vancouver, British Columbia, Canada

Judie Alimonti

Address Current address: Manitoba Institute of Cell Biology, Winnipeg, Manitoba, Canada

Affiliations The Michael Smith Laboratories, University of British Columbia, Vancouver, British Columbia, Canada , Biomedical Research Centre, University of British Columbia, Vancouver, British Columbia, Canada , Department of Microbiology and Immunology, University of British Columbia, Vancouver, British Columbia, Canada , The Michael Smith Laboratories, University of British Columbia, Vancouver, British Columbia, Canada

Susan S Chen

Affiliations The Michael Smith Laboratories, University of British Columbia, Vancouver, British Columbia, Canada , Biomedical Research Centre, University of British Columbia, Vancouver, British Columbia, Canada , Department of Zoology, University of British Columbia, Vancouver, British Columbia, Canada , The Michael Smith Laboratories, University of British Columbia, Vancouver, British Columbia, Canada

Genc Basha

Affiliations The Michael Smith Laboratories, University of British Columbia, Vancouver, British Columbia, Canada , Biomedical Research Centre, University of British Columbia, Vancouver, British Columbia, Canada , The Michael Smith Laboratories, University of British Columbia, Vancouver, British Columbia, Canada

Alex Moise

Address Current address: Case Western Reserve University, Cleveland, Ohio, United States of America

Affiliations The Michael Smith Laboratories, University of British Columbia, Vancouver, British Columbia, Canada , Biomedical Research Centre, University of British Columbia, Vancouver, British Columbia, Canada , Department of Zoology, University of British Columbia, Vancouver, British Columbia, Canada , The Michael Smith Laboratories, University of British Columbia, Vancouver, British Columbia, Canada

Jacqueline Tiong

Affiliations The Michael Smith Laboratories, University of British Columbia, Vancouver, British Columbia, Canada , Biomedical Research Centre, University of British Columbia, Vancouver, British Columbia, Canada , Department of Microbiology and Immunology, University of British Columbia, Vancouver, British Columbia, Canada , The Michael Smith Laboratories, University of British Columbia, Vancouver, British Columbia, Canada

Mei Mei Tian

Affiliations The Michael Smith Laboratories, University of British Columbia, Vancouver, British Columbia, Canada , Biomedical Research Centre, University of British Columbia, Vancouver, British Columbia, Canada , Department of Microbiology and Immunology, University of British Columbia, Vancouver, British Columbia, Canada , The Michael Smith Laboratories, University of British Columbia, Vancouver, British Columbia, Canada

Kyung Bok Choi

Affiliations The Michael Smith Laboratories, University of British Columbia, Vancouver, British Columbia, Canada , Biomedical Research Centre, University of British Columbia, Vancouver, British Columbia, Canada , The Michael Smith Laboratories, University of British Columbia, Vancouver, British Columbia, Canada

Douglas Waterfield

Affiliations Department of Oral Biology, Faculty of Dentistry, University of British Columbia, Vancouver, British Columbia, Canada , The Michael Smith Laboratories, University of British Columbia, Vancouver, British Columbia, Canada

Andy Jeffries

Affiliations The Michael Smith Laboratories, University of British Columbia, Vancouver, British Columbia, Canada , Biomedical Research Centre, University of British Columbia, Vancouver, British Columbia, Canada , The Michael Smith Laboratories, University of British Columbia, Vancouver, British Columbia, Canada

Wilfred A Jefferies

To whom correspondence should be addressed. E-mail: wilf@brc.ubc.ca

Affiliations The Michael Smith Laboratories, University of British Columbia, Vancouver, British Columbia, Canada , Biomedical Research Centre, University of British Columbia, Vancouver, British Columbia, Canada , Department of Microbiology and Immunology, University of British Columbia, Vancouver, British Columbia, Canada , Department of Zoology, University of British Columbia, Vancouver, British Columbia, Canada , Department of Medical Genetics, University of British Columbia, Vancouver, British Columbia, Canada , The Michael Smith Laboratories, University of British Columbia, Vancouver, British Columbia, Canada

Competing Interests

The University of British Columbia and its scientists (TZV, QZ, JA, and WAJ) received equity in GeneMax as part of a technology license agreement between the University of British Columbia and GeneMax. WAJ is a director of GeneMax. GeneMax provided partial financial support for this study.

Author Contributions

WAJ conceived and designed the experiments. TZV, QZ, JA, SSC, GB, AM, JT, MMT, KBC, DW, and AJ performed the experiments. TZV and WAJ analyzed the data and wrote the paper.