[1] Univ Calif San Francisco, Sloan Swartz Ctr Theoret Neurobiol, Dept Physiol, San Francisco, CA 94143 USA
[2] Univ Calif San Francisco, Howard Hughes Med Inst, San Francisco, CA 94143 USA
[3] Univ Calif San Francisco, Dept Physiol, San Francisco, CA 94143 USA
[4] Princeton Univ, Dept Phys, Princeton, NJ 08544 USA