1. Ecole Polytechnique Fédérale de Lausanne, Lausanne CH-1015, Switzerland
2. School of Informatics, University of Edinburgh, Edinburgh EH8 9AB, U.K.
3. Computer Science, Neuroscience, and Biomedical Engineering, University of Southern California, Los Angeles, CA 90089, U.S.A.
4. Computer Science, Neuroscience, and Biomedical Engineering, University of Southern California, Los Angeles, CA 90089, U.S.A.; Max-Planck-Institute for Intelligent Systems, Tübingen 72076, Germany; and ATR Computational Neuroscience Laboratories, Kyoto 619-0288, Japan