Topography of vibration frequency responses on the bony tympano-periotic complex of the pilot whale Globicephala macrorhynchus.

In modern Cetacea, the ear bone complex comprises the tympanic and periotic bones forming the tympano-periotic complex (TPC), differing from temporal bone complexes of other mammals in form, construction, position, and possibly function. To elucidate its functioning in sound transmission, we studied the vibration response of 32 pairs of formaldehyde-glutaraldehyde-fixed TPCs of Globicephala macrorhynchus, the short-finned pilot whale (legally obtained in Taiji, Japan). A piezoelectric-crystal-based vibrator was surgically attached to a location on the cochlea near the exit of the acoustic nerve. The crystal delivered vibrational pulses through continuous sweeps from 5 to 50 kHz. The vibration response was measured as a function of frequency by Laser Doppler Vibrometry at five points on the TPC. The aim of the experiment was to clarify how the vibration amplitudes produced by different frequencies are distributed on the TPC. At the lowest frequencies (<12 kHz), no clear differential pattern emerged. At higher frequencies the anterolateral lip of the TP responded most sensitively with the highest displacement amplitudes, and response amplitudes decreased in orderly fashion towards the posterior part of the TPC. We propose that this works as a lever: high-frequency sounds are most sensitively received and cause the largest vibration amplitudes at the anterior part of the TP, driving movements with lower amplitude but greater force near the posteriorly located contact to the ossicular chain, which transmits the movements into the inner ear. Although force (pressure) amplification is not needed for impedance matching in water, it may be useful for driving the stiffly connected ossicles at the high frequencies used in echolocation.

Relative size variation of the otoliths, swim bladder, and Weberian apparatus structures in piranhas and pacus (Characiformes: Serrasalmidae) with different ecologies and its implications for the detection of sound stimuli.

The Weberian apparatus of otophysan fishes confers acute hearing that is hypothesized to allow these fishes to assess the environment and to find food resources. The otophysan family Serrasalmidae (piranhas and pacus) includes species known to feed on falling fruits and seeds (frugivore/granivores) that splash in rivers, herbivorous species associated with torrents and rapids (rheophiles), and carnivores that feed aggressively within shoals. Relevant sound stimuli may vary among these ecological groups and hearing may be tuned to different cues among species. In this context, we examined size variation of the Weberian ossicles, swim bladder chambers, and otoliths of 20 serrasalmid species from three broad feeding ecologies: frugivore/granivores, rheophiles, and carnivores. We performed 3D-reconstructions of high resolution tomographic data (µCT) from 54 museum specimens to estimate the size of these elements. We then tested for an ecology effect on covariation of auditory structure size and body size and accounted for phylogeny with phylogenetic generalized least squares analyses. Among ecological groups, we observed differences in relative sizes of otoliths associated with sound pressure and particle motion detection, and variation in Weberian ossicle size that may impact sound transmission. Rheophiles, which live in noisy environments, possess the strongest modifications of these structures.

Human ossicular-joint flexibility transforms the peak amplitude and width of impulsive acoustic stimuli.

The role of the ossicular joints in the mammalian middle ear is still debated. This work tests the hypothesis that the two synovial joints filter potentially damaging impulsive stimuli by transforming both the peak amplitude and width of these impulses before they reach the cochlea. The three-dimensional (3D) velocity along the ossicular chain in unaltered cadaveric human temporal bones (N = 9), stimulated with acoustic impulses, is measured in the time domain using a Polytec (Waldbronn, Germany) CLV-3D laser Doppler vibrometer. The measurements are repeated after fusing one or both of the ossicular joints with dental cement. Sound transmission is characterized by measuring the amplitude, width, and delay of the impulsive velocity profile as it travels from the eardrum to the cochlea. On average, fusing both ossicular joints causes the stapes velocity amplitude and width to change by a factor of 1.77 (p = 0.0057) and 0.78 (p = 0.011), respectively. Fusing just the incudomalleolar joint has a larger effect on amplitude (a factor of 2.37), while fusing just the incudostapedial joint decreases the stapes velocity on average. The 3D motion of the ossicles is altered by fusing the joints. Finally, the ability of current computational models to predict this behavior is also evaluated.

Sheep as a large animal ear model: Middle-ear ossicular velocities and intracochlear sound pressure.

Animals are frequently used for the development and testing of new hearing devices. Dimensions of the middle ear and cochlea differ significantly between humans and commonly used animals, such as rodents or cats. The sheep cochlea is anatomically more like the human cochlea in size and number of turns. This study investigated the middle-ear ossicular velocities and intracochlear sound pressure (ICSP) in sheep temporal bones, with the aim of characterizing the sheep as an experimental model for implantable hearing devices. Measurements were made on fresh sheep temporal bones. Velocity responses of the middle ear ossicles at the umbo, long process of the incus and stapes footplate were measured in the frequency range of 0.25-8 kHz using a laser Doppler vibrometer system. Results were normalized by the corresponding sound pressure level in the external ear canal (PEC). Sequentially, ICSPs at the scala vestibuli and tympani were then recorded with custom MEMS-based hydrophones, while presenting identical acoustic stimuli. The sheep middle ear transmitted most effectively around 4.8 kHz, with a maximum stapes velocity of 0.2 mm/s/Pa. At the same frequency, the ICSP measurements in the scala vestibuli and tympani showed the maximum gain relative to the PEC (24 dB and 5 dB, respectively). The greatest pressure difference across the cochlear partition occurred between 4 and 6 kHz. A comparison between the results of this study and human reference data showed middle-ear resonance and best cochlear sensitivity at higher frequencies in sheep. In summary, sheep can be an appropriate large animal model for research and development of implantable hearing devices.

Simultaneous 3D imaging of sound-induced motions of the tympanic membrane and middle ear ossicles.

Efficient transfer of sound by the middle ear ossicles is essential for hearing. Various pathologies can impede the transmission of sound and thereby cause conductive hearing loss. Differential diagnosis of ossicular disorders can be challenging since the ossicles are normally hidden behind the tympanic membrane (TM). Here we describe the use of a technique termed optical coherence tomography (OCT) vibrography to view the sound-induced motion of the TM and ossicles simultaneously. With this method, we were able to capture three-dimensional motion of the intact TM and ossicles of the chinchilla ear with nanometer-scale sensitivity at sound frequencies from 0.5 to 5 kHz. The vibration patterns of the TM were complex and highly frequency dependent with mean amplitudes of 70-120 nm at 100 dB sound pressure level. The TM motion was only marginally sensitive to stapes fixation and incus-stapes joint interruption; however, when additional information derived from the simultaneous measurement of ossicular motion was added, it was possible to clearly distinguish these different simulated pathologies. The technique may be applicable to clinical diagnosis in Otology and to basic research in audition and acoustics.

Transitional mammalian middle ear from a new Cretaceous Jehol eutriconodont.

The transference of post-dentary jaw elements to the cranium of mammals as auditory ossicles is one of the central topics in evolutionary biology of vertebrates. Homologies of these bones among jawed vertebrates have long been demonstrated by developmental studies; but fossils illuminating this critical transference are sparse and often ambiguous. Here we report the first unambiguous ectotympanic (angular), malleus (articular and prearticular) and incus (quadrate) of an Early Cretaceous eutriconodont mammal from the Jehol Biota, Liaoning, China. The ectotympanic and malleus have lost their direct contact with the dentary bone but still connect the ossified Meckel's cartilage (OMC); we hypothesize that the OMC serves as a stabilizing mechanism bridging the dentary and the detached ossicles during mammalian evolution. This transitional mammalian middle ear narrows the morphological gap between the mandibular middle ear in basal mammaliaforms and the definitive mammalian middle ear (DMME) of extant mammals; it reveals complex changes contributing to the detachment of ear ossicles during mammalian evolution.

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.