Integrative physiology of fundamental frequency control in birds

One major feature of the remarkable vocal repertoires of birds is the range of fundamental frequencies across species, but also within individual species. This review discusses four variables that determine the oscillation frequency of the vibrating structures within a bird's syrinx. These are (1) viscoelastic properties of the oscillating tissue, (2) air sac pressure, (3) neuromuscular control of movements and (4) source-filter interactions. Our current understanding of morphology, biomechanics and neural control suggests that a complex interplay of these parameters can lead to multiple combinations for generating a particular fundamental frequency. An increase in the complexity of syringeal morphology from non-passeriform birds to oscines also led to a different interplay for regulating oscillation frequency by enabling control of tension that is partially independent of regulation of airflow. In addition to reviewing the available data for all different contributing variables, we point out open questions and possible approaches.