Physiological Insights Into the Social-Context-Dependent Changes in the Rhythm of the Song Motor Program

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Abstract
Precisely timed behaviors are central to the survival of almost all organisms. Song is an example of a learned behavior under exquisite temporal control. Song tempo in zebra finches ( Taeniopygia guttata) is systematically modified depending on social context. When male zebra finches sing to females (directed), it is produced with a faster motor pattern compared with when they sing in isolation (undirected). We measured heart rate and air sac pressure during directed and undirected singing to quantify motivation levels and respiratory timing. Heart rate was significantly higher when male birds sang to females and was negatively correlated with song duration. The change in song tempo between directed and undirected song was accounted for by varying the duration of vocal expiratory events, whereas the duration of silent inspirations was unchanged. Song duration increased with repeated singing during directed song bouts, which was caused by a uniform increase in the duration of both expirations and inspirations. These results illustrate the importance of motivational state in regulating song tempo and demonstrate that multiple timing oscillators are necessary to control the rhythm of song. At least two different neural oscillators are required to control context-dependent changes in song tempo. One oscillator controlling expiratory duration varies as function of social context and another controlling inspiratory duration is fixed. In contrast, the song tempo change affecting expiratory and inspiratory duration within a directed bout of song could be achieved by slowing the output of a single oscillator.