Vertebrate pressure-gradient receivers.

The eardrums of all terrestrial vertebrates (tetrapods) are connected through Eustachian tubes or interaural canals. In some of the animals, these connections create pressure-gradient directionality, an enhanced directionality by interaction of sound arriving at both sides of the eardrum and strongly dependent on interaural transmission attenuation. Even though the tympanic middle ear has originated independently in the major tetrapod groups, in each group the ancestral condition probably was that the two middle ears were exposed in the mouth cavity with relatively high interaural transmission. Recent vertebrates form a continuum from perfect interaural transmission (0 dB in a certain frequency band) and pronounced eardrum directionality (30-40 dB) in the lizards, over somewhat attenuated transmission and limited directionality in birds and frogs, to the strongly attenuated interaural transmission and functionally isolated pressure receiver ears in the mammals. Since some of the binaural interaction already takes place at the eardrum in animals with strongly coupled ears, producing enhanced interaural time and level differences, the subsequent neural processing may be simpler. In robotic simulations of lizards, simple binaural subtraction (EI cells, found in brainstem nuclei of both frogs and lizards) produces strongly lateralized responses that are sufficient for steering the animal robustly to sound sources.