Acoustic analysis of primate air sacs and their effect on vocalization.

This paper presents an analysis of the acoustic impedance of primate air sacs and their interaction with the vocal tract. A lumped element model is derived and it is found that the inertance of the neck and the volume of the air sac are relevant, as well as the mass and stiffness of the walls (depending on the tissue). It is also shown that at low frequencies, radiation from the air sac can be non-negligible, even if the mouth is open. It is furthermore shown that an air sac can add one or two low resonances to the resonances of the oral tract, and that it shifts up the oral tract's resonances below approximately 2000 Hz, and shifts them closer together. The theory was verified by acoustic measurements and applied to the red howler monkey (Alouatta seniculus) and the siamang (Symphalangus syndactylus). The theory describes the physical models and the siamang calls correctly, but appears incomplete for the howler monkey vocalizations. The relation between air sacs and the evolution of speech is discussed briefly, and it is proposed that an air sac would reduce the ability to produce distinctive speech, but would enhance the impression of size of the vocalizer.