[1] Univ Calif San Francisco, Nina Ireland Lab Dev Neurobiol, Ctr Neurobiol & Psychiat, Dept Psychiat, San Francisco, CA 94143 USA
[2] Univ Calif San Francisco, Program Neurosci, San Francisco, CA 94143 USA
[3] Univ Calif San Francisco, Program Dev Biol & Biomed Sci, San Francisco, CA 94143 USA
[4] Columbia Univ Coll Phys & Surg, Dept Biochem & Mol Biophys, New York, NY 10032 USA
[5] Columbia Univ Coll Phys & Surg, Howard Hughes Med Inst, New York, NY 10032 USA