Songs of birds comprise hierarchical sets of vocal gestures. In zebra
finches, songs Include notes and syllables (groups of notes) delivered
in fixed sequences. During singing, premotor neurons in the forebrain
nucleus HVc exhibited reliable changes In activity rates whose patter
ns were uniquely associated with syllable identity. Neurons in the for
ebrain nucleus robustus archistriatalis, which receives input from the
HVc, exhibited precisely rimed and structured bursts of activity that
were uniquely associated with note identity. Hence, units of vocal be
havior are represented hierarchically in the avian forebrain. The repr
esentation of temporal sequences at each level of the hierarchy may be
established by means of a decoding process involving Interactions of
higher level input with intrinsic local circuitry. Behavior is apparen
tly represented by precise temporal patterning of spike trains at lowe
r levels of the hierarchy.