Hebbian and neuromodulatory mechanisms interact to trigger associative memory formation

aLaboratory for Neural Circuitry of Memory, RIKEN Brain Science Institute, Wako, Saitama 351-0198, Japan;

bDepartment of Life Sciences, Graduate School of Arts and Sciences, University of Tokyo, Tokyo 153-0198, Japan;

cCenter for Neural Science, New York University, New York, NY 10003;

cCenter for Neural Science, New York University, New York, NY 10003;

aLaboratory for Neural Circuitry of Memory, RIKEN Brain Science Institute, Wako, Saitama 351-0198, Japan;

aLaboratory for Neural Circuitry of Memory, RIKEN Brain Science Institute, Wako, Saitama 351-0198, Japan;

aLaboratory for Neural Circuitry of Memory, RIKEN Brain Science Institute, Wako, Saitama 351-0198, Japan;

aLaboratory for Neural Circuitry of Memory, RIKEN Brain Science Institute, Wako, Saitama 351-0198, Japan;

aLaboratory for Neural Circuitry of Memory, RIKEN Brain Science Institute, Wako, Saitama 351-0198, Japan;

cCenter for Neural Science, New York University, New York, NY 10003;

dDepartment of Bioengineering, Department of Psychiatry and Behavioral Sciences, and

eHoward Hughes Medical Institute, Stanford University, Stanford, CA 94305;

fMcGovern Institute for Brain Research, Department of Brain and Cognitive Sciences, Massachusetts Institute of Technology, Cambridge, MA 02139; and

cCenter for Neural Science, New York University, New York, NY 10003;

gThe Emotional Brain Institute, Nathan Kline Institute for Psychiatric Research, Orangeburg, NY 10962