Among anuran amphibians, only two species, Odorrana tormota and Huia cavitympanum, are known to possess recessed tympanic membranes. Odorrana tormota is the first non-mammalian vertebrate demonstrated to communicate with ultrasonic frequencies (above 20 kHz), and the frogs' sunken tympana are hypothesized to play a key role in their high-frequency hearing sensitivity. Here we present the first data on the vocalizations of H. cavitympanum. We found that this species emits extraordinarily high-frequency calls, a portion of which are comprised entirely of ultrasound. This represents the first documentation of an anuran species producing purely ultrasonic signals. In addition, the vocal repertoire of H. cavitympanum is highly variable in frequency modulation pattern and spectral composition. The frogs' use of vocal signals with a wide range of dominant frequencies may be a strategy to maximize acoustic energy transmission to both nearby and distant receivers. The convergence of these species' call characteristics should stimulate additional, phylogenetically based studies of other lower vertebrates to provide new insight into the mechanistic and evolutionary foundations of high-frequency hearing in all vertebrate forms.
Odorrana tormota; Huia cavitympanum; ultrasonic communication; ultrasound; convergence
Direct acoustic stimulation of the cochlea by the sound-pressure difference between the oval and round windows (called the “acoustic route”) has been thought to contribute to hearing in some pathological conditions, along with the normally dominant “ossicular route.” To determine the efficacy of this acoustic route and its constituent mechanisms in human ears, sound pressures were measured at three locations in cadaveric temporal bones [with intact and perforated tympanic membranes (TMs)]: (1) in the external ear canal lateral to the TM, PTM; (2) in the tympanic cavity lateral to the oval window, POW; and (3) near the round window, PRW. Sound transmission via the acoustic route is described by two concatenated processes: (1) coupling of sound pressure from ear canal to middle-ear cavity, HPCAV≡PCAV/PTM, where PCAV represents the middle-ear cavity pressure, and (2) sound-pressure difference between the windows, HWPD≡(POW−PRW)/PCAV. Results show that: HPCAV depends on perforation size but not perforation location; HWPD depends on neither perforation size nor location. The results (1) provide a description of the window pressures based on measurements, (2) refute the common otological view that TM perforation location affects the “relative phase of the pressures at the oval and round windows,” and (3) show with an intact ossicular chain that acoustic-route transmission is substantially below ossicular-route transmission except for low frequencies with large perforations. Thus, hearing loss from TM perforations results primarily from reduction in sound coupling via the ossicular route. Some features of the frequency dependence of HPCAV and HWPD can be interpreted in terms of a structure-based lumped-element acoustic model of the perforation and middle-ear cavities.
Lungfishes are the closest living relatives of the tetrapods, and the ear of recent lungfishes resembles the tetrapod ear more than the ear of ray-finned fishes and is therefore of interest for understanding the evolution of hearing in the early tetrapods. The water-to-land transition resulted in major changes in the tetrapod ear associated with the detection of air-borne sound pressure, as evidenced by the late and independent origins of tympanic ears in all of the major tetrapod groups. To investigate lungfish pressure and vibration detection, we measured the sensitivity and frequency responses of five West African lungfish (Protopterus annectens) using brainstem potentials evoked by calibrated sound and vibration stimuli in air and water. We find that the lungfish ear has good low-frequency vibration sensitivity, like recent amphibians, but poor sensitivity to air-borne sound. The skull shows measurable vibrations above 100 Hz when stimulated by air-borne sound, but the ear is apparently insensitive at these frequencies, suggesting that the lungfish ear is neither adapted nor pre-adapted for aerial hearing. Thus, if the lungfish ear is a model of the ear of early tetrapods, their auditory sensitivity was limited to very low frequencies on land, mostly mediated by substrate-borne vibrations.
lungfish; hearing; vibration; tetrapod; sound; evolution
Insights into the onset of evolutionary novelties are key to the understanding of amniote origins and diversification. The possession of an impedance-matching tympanic middle ear is characteristic of all terrestrial vertebrates with a sophisticated hearing sense and an adaptively important feature of many modern terrestrial vertebrates. Whereas tympanic ears seem to have evolved multiple times within tetrapods, especially among crown-group members such as frogs, mammals, squamates, turtles, crocodiles, and birds, the presence of true tympanic ears has never been recorded in a Paleozoic amniote, suggesting they evolved fairly recently in amniote history.
In the present study, we performed a morphological examination and a phylogenetic analysis of poorly known parareptiles from the Middle Permian of the Mezen River Basin in Russia. We recovered a well-supported clade that is characterized by a unique cheek morphology indicative of a tympanum stretching across large parts of the temporal region to an extent not seen in other amniotes, fossil or extant, and a braincase specialized in showing modifications clearly related to an increase in auditory function, unlike the braincase of any other Paleozoic tetrapod. In addition, we estimated the ratio of the tympanum area relative to the stapedial footplate for the basalmost taxon of the clade, which, at 23∶1, is in close correspondence to that of modern amniotes capable of efficient impedance-matching hearing.
Using modern amniotes as analogues, the possession of an impedance-matching middle ear in these parareptiles suggests unique ecological adaptations potentially related to living in dim-light environments. More importantly, our results demonstrate that already at an early stage of amniote diversification, and prior to the Permo-Triassic extinction event, the complexity of terrestrial vertebrate ecosystems had reached a level that proved advanced sensory perception to be of notable adaptive significance.
The most common consequences of acute acoustic trauma (AAT) are hearing loss at frequencies above 3 kHz and tinnitus. In this study, we have used functional Magnetic Resonance Imaging (fMRI) to visualize neuronal activation patterns in military adults with AAT and various tinnitus sequelae during an auditory “oddball” attention task. AAT subjects displayed overactivities principally during reflex of target sound detection, in sensorimotor areas and in emotion-related areas such as the insula, anterior cingulate and prefrontal cortex, in premotor area, in cross-modal sensory associative areas, and, interestingly, in a region of the Rolandic operculum that has recently been shown to be involved in tympanic movements due to air pressure. We propose further investigations of this brain area and fine middle ear investigations, because our results might suggest a model in which AAT tinnitus may arise as a proprioceptive illusion caused by abnormal excitability of middle-ear muscle spindles possibly link with the acoustic reflex and associated with emotional and sensorimotor disturbances.
Acoustic trauma; fMRI; Middle ear; Proprioception; Tinnitus
The hypothesis is tested that an open-canal hearing device, with a microphone in the ear canal, can be designed to provide amplification over a wide bandwidth and without acoustic feedback. In the design under consideration, a transducer consisting of a thin silicone platform with an embedded magnet is placed directly on the tympanic membrane. Sound picked up by a microphone in the ear canal, including sound-localization cues thought to be useful for speech perception in noisy environments, is processed and amplified, and then used to drive a coil near the tympanic-membrane transducer. The perception of sound results from the vibration of the transducer in response the electromagnetic field produced by the coil. Sixteen subjects (ranging from normal-hearing to moderately hearing-impaired) wore this transducer for up to a ten-month period, and were monitored for any adverse reactions. Three key functional characteristics were measured: 1) the maximum equivalent pressure output (MEPO) of the transducer; 2) the feedback gain margin (GM), which describes the maximum allowable gain before feedback occurs; and 3) the tympanic-membrane damping effect (DTM), which describes the change in hearing level due to placement of the transducer on the eardrum. Results indicate that the tympanic-membrane transducer remains in place and is well tolerated. The system can produce sufficient output to reach threshold for those with as much as 60 dBHL of hearing impairment for up to 8 kHz in 86% of the study population, and up to 11.2 kHz in 50% of the population. The feedback gain margin is on average 30 dB except at the ear canal resonance frequencies of 3 and 9 kHz, where the average was reduced to 12 dB and 23 dB respectively. The average value of DTM is close to 0 dB everywhere except in the 2–4 kHz range, where it peaks at 8 dB. A new alternative system that uses photonic energy to transmit both the signal and power to a photodiode and micro-actuator on an EarLens platform is also described.
The hearing threshold in atrophic tympanic membrane is assessed in 35 individuals. Assessment of hearing threshold in patients having atrophic tympanic membrane. Prospective clinical study. Tertiary referral centre. Thirty-five patients who had atrophic tympanic membrane in one ear and normal tympanic membrane of the other ear which was used as control, were selected Hearing threshold of patients having atrophic tympanic membrane. Twenty-nine patients with atrophic tympanic membrane had absolutely normal PTA of the ear and the opposite ear with normal tympanic membrane had similar normal PTA. Majority of the patients with atrophic tympanic membrane have normal hearing.
Atrophic tympanic membrane; Pure tone audiogram
A salient characteristic of most auditory systems is their capacity to analyse the frequency of sound. Little is known about how such analysis is performed across the diversity of auditory systems found in animals, and especially in insects. In locusts, frequency analysis is primarily mechanical, based on vibrational waves travelling across the tympanal membrane. Different acoustic frequencies generate travelling waves that direct vibrations to distinct tympanal locations, where distinct groups of correspondingly tuned mechanosensory neurons attach. Measuring the mechanical tympanal response, for the first time, to acoustic impulses in the time domain, nanometre-range vibrational waves are characterized with high spatial and temporal resolutions. Conventional Fourier analysis is also used to characterize the response in the frequency domain. Altogether these results show that travelling waves originate from a particular tympanal location and travel across the membrane to generate oscillations in the exact region where mechanosensory neurons attach. Notably, travelling waves are unidirectional; no strong back reflection or wave resonance could be observed across the membrane. These results constitute a key step in understanding tympanal mechanics in general, and in insects in particular, but also in our knowledge of the vibrational behaviour of anisotropic media.
tympanal membrane; frequency; place principle; biomechanics; time-resolved laser vibrometry
Conductive hearing loss (CHL) is known to produce hearing deficits, including deficits in sound localization ability. The differences in sound intensities and timing experienced between the two tympanic membranes are important cues to sound localization (ILD and ITD, respectively). Although much is known about the effect of CHL on hearing levels, little investigation has been conducted into the actual impact of CHL on sound location cues. This study investigated effects of CHL induced by earplugs on cochlear microphonic (CM) amplitude and timing and their corresponding effect on the ILD and ITD location cues. Acoustic and CM measurements were made in 5 chinchillas before and after earplug insertion, and again after earplug removal using pure tones (500 Hz to 24 kHz). ILDs in the unoccluded condition demonstrated position and frequency dependence where peak far-lateral ILDs approached 30 dB for high frequencies. Unoccluded ear ITD cues demonstrated positional and frequency dependence with increased ITD cue for both decreasing frequency (± 420 µs at 500 Hz, ± 310 µs for 1–4 kHz ) and increasingly lateral sound source locations. Occlusion of the ear canal with foam plugs resulted in a mild, frequency-dependent conductive hearing loss of 10–38 dB (mean 31 ± 3.9 dB) leading to a concomitant frequency dependent increase in ILDs at all source locations. The effective ITDs increased in a frequency dependent manner with ear occlusion as a direct result of the acoustic properties of the plugging material, the latter confirmed via acoustical measurements using a model ear canal with varying volumes of acoustic foam. Upon ear plugging with acoustic foam, a mild CHL is induced. Furthermore, the CHL induced by acoustic foam results in substantial changes in the magnitudes of both the ITD and ILD cues to sound location.
Conductive hearing loss; Interaural level differences (ILD); Interaural timing differences (ITD); Otitis media with effusion
Background noise poses a significant obstacle for auditory perception, especially among individuals with hearing loss. To better understand the physiological basis of this perceptual impediment, the present study evaluated the effects of background noise on the auditory nerve representation of head-related transfer functions (HRTFs). These complex spectral shapes describe the directional filtering effects of the head and torso. When a broadband sound passes through the outer ear en route to the tympanic membrane, the HRTF alters its spectrum in a manner that establishes the perceived location of the sound source. HRTF-shaped noise shares many of the acoustic features of human speech, while communicating biologically relevant localization cues that are generalized across mammalian species. Previous studies have used parametric manipulations of random spectral shapes to elucidate HRTF coding principles at various stages of the cat’s auditory system. This study extended that body of work by examining the effects of sound level and background noise on the quality of spectral coding in the auditory nerve. When fibers were classified by their spontaneous rates, the coding properties of the more numerous low-threshold, high-spontaneous rate fibers were found to degrade at high presentation levels and in low signal-to-noise ratios. Because cats are known to maintain accurate directional hearing under these challenging listening conditions, behavioral performance may be disproportionally based on the enhanced dynamic range of the less common high-threshold, low-spontaneous rate fibers.
spectral integration; auditory nerve; rate representation; sound localization; background noise
Turtles, like other amphibious animals, face a trade-off between terrestrial and aquatic hearing. We used laser vibrometry and auditory brainstem responses to measure their sensitivity to vibration stimuli and to airborne versus underwater sound. Turtles are most sensitive to sound underwater, and their sensitivity depends on the large middle ear, which has a compliant tympanic disc attached to the columella. Behind the disc, the middle ear is a large air-filled cavity with a volume of approximately 0.5 ml and a resonance frequency of approximately 500 Hz underwater. Laser vibrometry measurements underwater showed peak vibrations at 500–600 Hz with a maximum of 300 µm s−1 Pa−1, approximately 100 times more than the surrounding water. In air, the auditory brainstem response audiogram showed a best sensitivity to sound of 300–500 Hz. Audiograms before and after removing the skin covering reveal that the cartilaginous tympanic disc shows unchanged sensitivity, indicating that the tympanic disc, and not the overlying skin, is the key sound receiver. If air and water thresholds are compared in terms of sound intensity, thresholds in water are approximately 20–30 dB lower than in air. Therefore, this tympanic ear is specialized for underwater hearing, most probably because sound-induced pulsations of the air in the middle ear cavity drive the tympanic disc.
underwater sound; evolution; cochlea; auditory brainstem response
The position of testudines in vertebrate phylogeny is being re-evaluated. At present, testudine morphological and molecular data conflict when reconstructing phylogenetic relationships. Complicating matters, the ecological niche of stem testudines is ambiguous. To understand how turtles have evolved to hear in different environments, we examined middle ear morphology and scaling in most extant families, as well as some extinct species, using 3-dimensional reconstructions from micro magnetic resonance (MR) and submillimeter computed tomography (CT) scans. All families of testudines exhibited a similar shape of the bony structure of the middle ear cavity, with the tympanic disk located on the rostrolateral edge of the cavity. Sea Turtles have additional soft tissue that fills the middle ear cavity to varying degrees. When the middle ear cavity is modeled as an air-filled sphere of the same volume resonating in an underwater sound field, the calculated resonances for the volumes of the middle ear cavities largely fell within testudine hearing ranges. Although there were some differences in morphology, there were no statistically significant differences in the scaling of the volume of the bony middle ear cavity with head size among groups when categorized by phylogeny and ecology. Because the cavity is predicted to resonate underwater within the testudine hearing range, the data support the hypothesis of an aquatic origin for testudines, and function of the middle ear cavity in underwater sound detection.
Lizard ears are clear examples of two-input pressure-difference receivers, with up to 40-dB differences in eardrum vibration amplitude in response to ipsi- and contralateral stimulus directions. The directionality is created by acoustical coupling of the eardrums and interaction of the direct and indirect sound components on the eardrum. The ensuing pressure-difference characteristics generate the highest directionality of any similar-sized terrestrial vertebrate ear. The aim of the present study was to measure the gain of the direct and indirect sound components in three lizard species: Anolis sagrei and Basiliscus vittatus (iguanids) and Hemidactylus frenatus (gekkonid) by laser vibrometry, using either free-field sound or a headphone and coupler for stimulation. The directivity of the ear of these lizards is pronounced in the frequency range from 2 to 5 kHz. The directivity is ovoidal, asymmetrical across the midline, but largely symmetrical across the interaural axis (i.e., front–back). Occlusion of the contralateral ear abolishes the directionality. We stimulated the two eardrums with a coupler close to the eardrum to measure the gain of the sound pathways. Within the frequency range of maximal directionality, the interaural transmission gain (compared to sound arriving directly) is close to or even exceeds unity, indicating a pronounced acoustical transparency of the lizard head and resonances in the interaural cavities. Our results show that the directionality of the lizard ear is caused by the acoustic interaction of the two eardrums. The results can be largely explained by a simple acoustical model based on an electrical analog circuit.
lizard; tympanum; vibrometry; directional; hearing; reptile
Vibratory measurements of the structures of the ear are key to understanding much of the pathology in mouse models of hearing loss. Unfortunately the high-speed sampling required to interrogate the high end of the mouse hearing spectrum is beyond the reach of most optical coherence tomography (OCT) systems. To address this issue, we have developed an algorithm that enables phase-sensitive OCT measurements over the full range of the mouse hearing spectrum (4–90 kHz). The algorithm phase-locks the line-trigger to the acoustic stimulation and then uses interleaved sampling to reconstruct the signal with higher temporal sampling. The algorithm was evaluated by measuring the vibratory response of mouse tympanic membrane to a pure tone stimulus.
The hearing of tetrapods including humans is enhanced by an active process that amplifies the mechanical inputs associated with sound, sharpens frequency selectivity, and compresses the range of responsiveness. The most striking manifestation of the active process is spontaneous otoacoustic emission, the unprovoked emergence of sound from an ear. Hair cells, the sensory receptors of the inner ear, are known to provide the energy for such emissions; it is unclear, though, how ensembles of such cells collude to power observable emissions.
Methodology and Principal Findings
We have measured and modeled spontaneous otoacoustic emissions from the ear of the tokay gecko, a convenient experimental subject that produces robust emissions. Using a van der Pol formulation to represent each cluster of hair cells within a tonotopic array, we have examined the factors that influence the cooperative interaction between oscillators.
Conclusions and Significance
A model that includes viscous interactions between adjacent hair cells fails to produce emissions similar to those observed experimentally. In contrast, elastic coupling yields realistic results, especially if the oscillators near the ends of the array are weakened so as to minimize boundary effects. Introducing stochastic irregularity in the strength of oscillators stabilizes peaks in the spectrum of modeled emissions, further increasing the similarity to the responses of actual ears. Finally, and again in agreement with experimental findings, the inclusion of a pure-tone external stimulus repels the spectral peaks of spontaneous emissions. Our results suggest that elastic coupling between oscillators of slightly differing strength explains several properties of the spontaneous otoacoustic emissions in the gecko.
Fishes have evolved a diversity of sound-generating organs and acoustic signals of various temporal and spectral content. Additionally, representatives of many teleost families such as otophysines, anabantoids, mormyrids and holocentrids possess accessory structures that enhance hearing abilities by acoustically coupling air-filled cavities to the inner ear. Contrary to the accessory hearing structures such as Weberian ossicles in otophysines and suprabranchial chambers in anabantoids, sonic organs do not occur in all members of these taxa. Comparison of audiograms among nine representatives of seven otophysan families from four orders revealed major differences in auditory sensitivity, especially at higher frequencies (> 1 kHz) where thresholds differed by up to 50 dB. These differences showed no apparent correspondence to the ability to produce sounds (vocal versus non-vocal species) or to the spectral content of species-specific sounds. In anabantoids, the lowest auditory thresholds were found in the blue gourami Trichogaster trichopterus, a species not thought to be vocal. Dominant frequencies of sounds corresponded with optimal hearing bandwidth in two out of three vocalizing species. Based on these results, it is concluded that the selective pressures involved in the evolution of accessory hearing structures and in the design of vocal signals were other than those serving to optimize acoustic communication.
Time-averaged holograms describing the sound-induced motion of the tympanic membrane (TM) in cadaveric preparations from three mammalian species and one live ear were measured using opto-electronic holography. This technique allows rapid measurements of the magnitude of motion of the tympanic membrane surface at frequencies as high as 25 kHz. The holograms measured in response to low and middle-frequency sound stimuli are similar to previously reported time-averaged holograms. However, at higher frequencies (f > 4 kHz), our holograms reveal unique TM surface displacement patterns that consist of highly-ordered arrangements of multiple local displacement magnitude maxima, each of which is surrounded by nodal areas of low displacement magnitude. These patterns are similar to modal patterns (two-dimensional standing waves) produced by either the interaction of surface waves traveling in multiple directions or the uniform stimulation of modes of motion that are determined by the structural properties and boundary conditions of the TM. From the ratio of the displacement magnitude peaks to nodal valleys in these apparent surface waves, we estimate a Standing Wave Ratio of at least 4 that is consistent with energy reflection coefficients at the TM boundaries of at least 0.35. It is also consistent with small losses within the uniformly stimulated modal surface waves. We also estimate possible TM surface wave speeds that vary with frequency and species from 20 to 65 m/s, consistent with other estimates in the literature. The presence of standing wave or modal phenomena has previously been intuited from measurements of TM function, but is ignored in some models of tympanic membrane function. Whether these standing waves result either from the interactions of multiple surface waves that travel along the membrane, or by uniformly excited modal displacement patterns of the entire TM surface is still to be determined.
We report the results of anatomical and vibrometric studies of the middle ear of the African clawed frog, Xenopus laevis. The cartilaginous tympanic disk of Xenopus shows pronounced sexual dimorphism, that of male frogs being much larger than that of females, relative to body size. The stapes footplate, however, is not enlarged in males. The cucullaris muscle was found to insert on the stapes in frogs of both sexes. Using laser interferometry to examine the response of middle ear structures to airborne sound, the stapes footplate was found to vibrate close to 180° out-of-phase with the tympanic disk across a range of frequencies, this resembling the relationship between tympanic membrane and footplate movement previously described in ranid frogs. By contrast, whereas there is a pronounced difference in vibration velocity between tympanic membrane and footplate in ranids, the footplate vibration velocity in Xenopus was found to be similar to that of the tympanic disk. This may be interpreted as an adaptation to improve the detection of sound underwater.
Sound pressure was mapped in the bony ear canal of gerbils during closed-field sound stimulation at frequencies from 0.1 to 80 kHz. A 1.27-mm-diam probe-tube microphone or a 0.17-mm-diam fiber-optic miniature microphone was positioned along approximately longitudinal trajectories within the 2.3-mm-diam ear canal. Substantial spatial variations in sound pressure, sharp minima in magnitude, and half-cycle phase changes occurred at frequencies >30 kHz. The sound frequencies of these transitions increased with decreasing distance from the tympanic membrane (TM). Sound pressure measured orthogonally across the surface of the TM showed only small variations at frequencies below 60 kHz. Hence, the ear canal sound field can be described fairly well as a one-dimensional standing wave pattern. Ear-canal power reflectance estimated from longitudinal spatial variations was roughly constant at 0.2–0.5 at frequencies between 30 and 45 kHz. In contrast, reflectance increased at higher frequencies to at least 0.8 above 60 kHz. Sound pressure was also mapped in a microphone-terminated uniform tube—an “artificial ear.” Comparison with ear canal sound fields suggests that an artificial ear or “artificial cavity calibration” technique may underestimate the in situ sound pressure by 5–15 dB between 40 and 60 kHz.
This study compares measurements of ear-canal reflectance (ECR) to other objective measurements of middle-ear function including, audiometry, umbo velocity (VU), and tympanometry in a population of strictly defined normal hearing ears.
Data were prospectively gathered from 58 ears of 29 normal hearing subjects, 16 female and 13 male, aged 22–64 years. Subjects met all of the following criteria to be considered as having normal hearing. (1) No history of significant middle-ear disease. (2) No history of otologic surgery. (3) Normal tympanic membrane (TM) on otoscopy. (4) Pure-tone audiometric thresholds of 20 dB HL or better for 0.25 – 8 kHz. (5) Air-bone gaps no greater than 15 dB at 0.25 kHz and 10 dB for 0.5 – 4 kHz. (6) Normal, type-A peaked tympanograms. (7) All subjects had two “normal” ears (as defined by these criteria). Measurements included pure-tone audiometry for 0.25 – 8 kHz, standard 226 Hz tympanometry, Ear canal reflectance(ECR) for 0.2 – 6 kHz at 60 dB SPL using the Mimosa Acoustics HearID system, and Umbo Velocity (VU ) for 0.3 – 6 kHz at 70–90 dB SPL using the HLV-1000 laser Doppler vibrometer (Polytec Inc).
Mean power reflectance (|ECR|2) was near 1.0 at 0.2– 0.3 kHz, decreased to a broad minimum of 0.3 to 0.4 between 1 and 4 kHz, and then sharply increased to almost 0.8 by 6 kHz. The mean pressure reflectance phase angle (∠ECR) plotted on a linear frequency scale showed a group delay of approximately 0.1 ms for 0.2 – 6 kHz. Small significant differences were observed in |ECR|2 at the lowest frequencies between right and left ears, and between males and females at 4 kHz. |ECR|2 decreased with age, but reached significance only at 1 kHz. Our ECR measurements were generally similar to previous published reports. Highly significant negative correlations were found between |ECR|2 and VU for frequencies below 1 kHz. Significant correlations were also found between the tympanometrically determined peak total compliance and |ECR|2 and The results suggest that middle-ear compliance VU at frequencies below 1 kHz. contributes significantly to the measured power reflectance and umbo velocity at frequencies below 1 kHz, but not at higher frequencies.
This study has established a database of objective measurements of middle ear function (ear-canal reflectance, umbo velocity, tympanometry) in a population of strictly defined normal hearing ears. The data will promote our understanding of normal middle ear function, and will serve as a control for comparison to similar measurements made in pathological ears.
Although tympanic membrane perforations are common, there have been few systematic studies of the structural features determining the magnitude of the resulting conductive hearing loss. Our recent experimental and modeling studies predicted that the conductive hearing loss will increase with increasing perforation size, be independent of perforation location (contrary to popular otologic belief), and increase with decreasing size of the middle-ear and mastoid air space (an idea new to otology).
To test our predictions regarding determinants of conductive hearing loss in tympanic membrane perforations against clinical data gathered from patients.
Prospective clinical study.
Tertiary referral center.
Patients with tympanic membrane perforations without other middle-ear disease.
Main Outcome Measures
Size and location of perforation; air-bone gap at 250, 500, 1,000, 2,000, and 4,000 Hz; and tympanometric estimate of volume of the middle-ear air spaces.
Isolated tympanic membrane perforations in 62 ears from 56 patients met inclusion criteria. Air-bone gaps were largest at the lower frequencies and decreased as frequency increased. Air-bone gaps increased with perforation size at each frequency. Ears with small middle-ear volumes, ≤4.3 ml (n = 23), had significantly larger air-bone gaps than ears with large middle-ear volumes, >4.3 ml (n = 39), except at 2,000 Hz. The mean air-bone gaps in ears with small volumes were 10 to 20 dB larger than in ears with large volumes. Perforations in anterior versus posterior quadrants showed no significant differences in air-bone gaps at any frequency, although anterior perforations had, on average, air-bone gaps that were smaller by 1 to 8 dB at lower frequencies.
The conductive hearing loss resulting from a tympanic membrane perforation is frequency-dependent, with the largest losses occurring at the lowest sound frequencies; increases as size of the perforation increases; varies inversely with volume of the middle-ear and mastoid air space (losses are larger in ears with small volumes); and does not vary appreciably with location of the perforation. Effects of location, if any, are small.
Audiometry; Conductive hearing loss; Perforation; Tympanic membrane; Tympanometry
Recent clinical reports found a high incidence of recurrent otitis media in children suffering hyperacusis, a marked intolerance to an otherwise ordinary environmental sound. However, it is unclear whether the conductive hearing loss caused by otitis media in early age will affect sound tolerance later in life. Thus, we have tested the effects of tympanic membrane (TM) damage at an early age on sound perception development in rats. Two weeks after the TM perforation, more than 80% of the rats showed audiogenic seizure (AGS) when exposed to loud sound (120 dB SPL white noise, < 1 minute). The susceptibility of AGS lasted at least sixteen weeks after the TM damage, even the hearing loss recovered. The TM damaged rats also showed significantly enhanced acoustic startle responses compared to the rats without TM damage. These results suggest that early age conductive hearing loss may cause an impaired sound tolerance during development. In addition, the AGS can be suppressed by the treatment of vigabatrin, acute injections (250 mg/kg) or oral intakes (60 mg/kg/day for 7 days), an antiepileptic drug that inhibits the catabolism of GABA. c-Fos staining showed a strong staining in the inferior colliculus (IC) in the TM damaged rats, not in the control rats, after exposed to loud sound, indicating a hyper-excitability in the IC during AGS. These results indicate that early age conductive hearing loss can impair sound tolerance by reducing GABA inhibition in the IC, which may be related to hyperacusis seen in children with otitis media.
Otitis media; Audiogenic seizure; Acoustic startle reflex; Hyperacusis; Inferior colliculus; GABA
The conductive hearing of 18 patients with juvenile chronic arthritis was studied. Tympanometry was therefore undertaken for the assessment of functional state of the tympanic membrane, the ossicular chain with its ligaments and muscles, and the air cushion of the tympanic cavity. Acoustic reflex was present in all patients but an abnormal tympanometric pattern (type As) was found bilaterally or unilaterally in 10 (mostly severe cases) patients. These findings indicate that juvenile chronic arthritis changes middle ear function, probably as a result of inflammation of the synovial ossicular joints, which is followed by stiffness of the tympanic membrane and ossicular chain.
Central nervous system (CNS) involvement by chronic lymphocytic leukemia (CLL) can present with dramatic neurologic findings or can be quite subtle, discovered only at the time of autopsy. We describe a case of CLL in a patient who presented initially with hearing loss and was ultimately found to have involvement of the tympanic membrane. She noted improvement of her hearing after induction therapy but was not aware at the time of the involvement of her CNS with CLL. Upon worsening of hearing at the time of relapse, she was evaluated by imaging and CSF analysis as well as biopsy of the tympanic membrane, and involvement of the CNS was confirmed. She has received CNS-directed therapy with intrathecal liposomal cytarabine and intravenous CNS-directed therapy and has noted improved hearing and resolution of her imaging and CSF findings. This is the first reported case of tympanic membrane involvement with CLL and describes potentially effective methods for managing this challenging complication.
Based on comparisons of ear canal and scala vestibuli pressures the gerbil middle ear transmits sound with a gain of ~25 dB that is almost flat from 2 to 40 kHz, and with a delay-like phase corresponding to a 25 – 30 μs delay. How the middle ear is able to transmit sound with such high temporal and amplitude fidelity is not known, and is particularly mysterious given the complex motion the ossicles and tympanic membrane (TM) are known to undergo. To explore this question, we looked at the velocities of the manubrium and along a line on the TM. The TM motion was complex, and could be approximated as the combination of a wave-like motion and an in-and-out piston-like motion. The manubrium underwent bending at some stimulus frequencies and therefore its motion was not like a rigid body. It had a complex motion with frequency fine structure that seemed likely to be derived from resonances on the drum-like TM.
motion; manubrium; tympanic membrane; gerbil; middle ear