Songbirds tune their vocal tract to the fundamental frequency of their song

In human speech, the sound generated by the larynx is modified by articulatory movements of the upper vocal tract, which acts as a variable resonant filter concentrating energy near particular frequencies, or formants, essential in speech recognition. Despite its potential importance in vocal communication, little is known about the presence of tunable vocal tract filters in other vertebrates. The tonal quality of much birdsong, in which upper harmonics have relatively little energy, depends on filtering of the vocal source, but the nature of this filter is controversial. Current hypotheses treat the songbird vocal tract as a rigid tube with a resonance that is modulated by the end-correction of a variable beak opening. Through x-ray cinematography of singing birds, we show that birdsong is accompanied by cyclical movements of the hyoid skeleton and changes in the diameter of the cranial end of the esophagus that maintain an inverse relationship between the volume of the oropharyngeal cavity and esophagus and the song's fundamental frequency. A computational acoustic model indicates that this song-related motor pattern tunes the major resonance of the oropharyngeal-esophageal cavity to actively track the song's fundamental frequency.