Active tympanic tuning facilitates sound localization in animals with internally coupled ears

Earlier studies have reported that numerous vertebrate taxa have skeletal muscle(s) attaching directly, or indirectly, onto the tympanic membrane. The present study links these prior studies by quantitatively modeling the influence of skeletal muscle contraction on tympanic tension, tympanic dampening, and, ultimately, the fundamental frequency. In this way, the efficacy of these tympanic muscles to dynamically alter the sensory response of the vertebrate ear is quantified. Changing the tension modifies the eardrum's fundamental frequency, a key notion in understanding hearing through internally coupled ears (ICE) as used by the majority of terrestrial vertebrates. Tympanic tension can also be modulated by altering the pressure acting on the deep (medial) surface of the tympanum. Herein we use the monitor lizard Varanus as an example to demonstrate how active modulation of the pharyngeal volume permits tuning of an ICE auditory system. The present contribution offers a behaviorally and biologically realistic perspective on the ICE system, by demonstrating how an organism can dynamically alter its morphology to tune the auditory response. Through quantification of the relationships between tympanic surface tension, damping, membrane fundamental frequency, and auditory cavity volume, it can be shown that an ICE system affords a biologically relevant range of tuning.