1. Department of Physiology and Biophysics, Center for Neuroscience, and Computational Bioscience Program, University of Colorado School of Medicine, Aurora, Colorado 80045
2. Department of Applied Mathematics, University of Colorado, Boulder, Colorado 80309
3. Learning in Machines and Brains Program, Canadian Institute for Advanced Research, Toronto, Ontario M5G 1Z8, Canada
4. Department of Neurosciences, Case Western Reserve University School of Medicine, Cleveland, Ohio 44106;
5. Department of Physiology and Biophysics, Case Western Reserve University School of Medicine, Cleveland, Ohio 44106