1. Max Planck Institute for Biological Cybernetics, Spemannstrasse 38, 72076 Tübingen, Germany, University of Southern California, 3641 Watt Way, Los Angeles, CA 90089, USA
2. University of Southern California, 3641 Watt Way, Los Angeles, CA 90089, USA, ATR Computational Neuroscience Laboratory, 2-2-2 Hikaridai, Seika-cho, Soraku-gun, Kyoto 619-0288, Japan