Hearing without a tympanic ear.

The ability to sense and localize sound is so advantageous for survival that it is difficult to understand the almost 100 million year gap separating the appearance of early tetrapods and the emergence of an impedance-matching tympanic middle ear - which we normally regard as a prerequisite for sensitive hearing on land - in their descendants. Recent studies of hearing in extant atympanate vertebrates have provided significant insights into the ancestral state(s) and the early evolution of the terrestrial tetrapod auditory system. These reveal a mechanism for sound pressure detection and directional hearing in 'earless' atympanate vertebrates that may be generalizable to all tetrapods, including the earliest terrestrial species. Here, we review the structure and function of vertebrate tympanic middle ears and highlight the multiple acquisition and loss events that characterize the complex evolutionary history of this important sensory structure. We describe extratympanic pathways for sound transmission to the inner ear and synthesize findings from recent studies to propose a general mechanism for hearing in 'earless' atympanate vertebrates. Finally, we integrate these studies with research on tympanate species that may also rely on extratympanic mechanisms for acoustic reception of infrasound (<20 Hz) and with studies on human bone conduction mechanisms of hearing.