Mechanical leverage in the middle ear of the American bullfrog, Rana catesbeiana

Textbooks lump the middle ears of 'submammalian Tetrapoda' as being 'one-ossicle ears'. Conventionally the anuran middle ear is depicted with a shaft-like skeletal unit connecting the tympanic membrane to the inner ear. This shaft comprises mediad a long bony columella and laterad a short cartilaginous extracolumella. But dissection of Rana catesbeiana ears showed: the extracolumella, as long as the columella, is proximally expanded in the vertical plane, forming dorsal and ventral heads. The medio-dorsal head is movably jointed to the columella, between these two there is an obtuse angle ventrad; the extracolumellar medio-ventral head is anchored by a ligament to the middle-ear cavity ceiling. When the tympanic membrane moves outwards, pulling the extracolumella, the medio-dorsal head of the extracolumella must be forced inwards, rotating on the ventral anchorage, pushing the columella towards the inner ear. The ossicular chain thus includes a mechanical lever, possessing the magnitude of the ratio length:width of the extracolumella; this is additional to the lever known from the columellar footplate, which rotates on its firm ventral attachment. These levers are confirmed physiologically, by the difference between the inner-ear sensitivity (shown by isopotential audiograms of microphonic potentials) when stimulated by a vibrator first at the tympanic membrane, then at the proximal stump of the amputated columella. Perusal of the primary literature showed that this morphology is widespread among anuran ears.