About the Authors

Carole J. Proctor

c.j.proctor@ncl.ac.uk

Affiliation Center for Integrated Systems Biology of Ageing and Nutrition, Institute for Ageing and Health, Newcastle University, Newcastle upon Tyne, United Kingdom

Paul J. Tangeman

Affiliation Section of Ophthalmology and Neurosciences, Leeds Institute of Molecular Medicine, St James's University Hospital, Leeds, United Kingdom

Helen C. Ardley

Affiliation Section of Ophthalmology and Neurosciences, Leeds Institute of Molecular Medicine, St James's University Hospital, Leeds, United Kingdom

Competing Interests

The authors have declared that no competing interests exist.

Author Contributions

Conceived and designed the experiments: HA. Performed the experiments: PT HA. Analyzed the data: PT HA. Wrote the paper: CP HA. Built the mathematical model, ran simulations, analysed results: CP. Consulted with other authors regarding experiments: CP.