Auditory nerve responses to imposed displacements of the turtle basilar membrane

Impulse activity of single auditory nerve fibres was recorded in the isolated half-head of the turtle in response to displacements of a piezoelectric probe placed on the basilar membrane. The temporal pattern of firing in response to sinusoidal displacements of amplitude 0.1-1.0 nm r.m.s. at a fibre's characteristic frequency could be matched to that generated by low-level tonal stimuli delivered to the tympanum. Frequency-threshold curves for acoustic and mechanical stimuli had similar shapes and differed only at frequencies above 500 Hz where the middle ear should filter acoustic but not direct mechanical stimuli. Step displacements of the basilar membrane gave a transient periodic discharge which resembled the responses to acoustic clicks. Most fibres initially increased their firing rate for rarefaction clicks and displacements towards the scala tympani.     

Energy Flux in the Cochlea: Evidence Against Power Amplification of the Traveling Wave

Traveling waves in the inner ear exhibit an amplitude peak that shifts with frequency. The peaking is commonly believed to rely on motile processes that amplify the wave by inserting energy. We recorded the vibrations at adjacent positions on the basilar membrane in sensitive gerbil cochleae and tested the putative power amplification in two ways. First, we determined the energy flux of the traveling wave at its peak and compared it to the acoustic power entering the ear, thereby obtaining the net cochlear power gain. For soft sounds, the energy flux at the peak was 1 ± 0.6 dB less than the middle ear input power. For more intense sounds, increasingly smaller fractions of the acoustic power actually reached the peak region. Thus, we found no net power amplification of soft sounds and a strong net attenuation of intense sounds. Second, we analyzed local wave propagation on the basilar membrane. We found that the waves slowed down abruptly when approaching their peak, causing an energy densification that quantitatively matched the amplitude peaking, similar to the growth of sea waves approaching the beach. Thus, we found no local power amplification of soft sounds and strong local attenuation of intense sounds. The most parsimonious interpretation of these findings is that cochlear sensitivity is not realized by amplifying acoustic energy, but by spatially focusing it, and that dynamic compression is realized by adjusting the amount of dissipation to sound intensity.     

A mechanism for simultaneous suppression of tone burst-evoked otoacoustic emissions

Tone burst-evoked otoacoustic emission (TBOAE) components in response to a 1 kHz tone burst are suppressed by the simultaneous presence of tone bursts at higher frequencies. To date, the underlying cause of this "simultaneous suppression" of TBOAEs is unclear. This paper describes a potential mechanism based on local nonlinear interactions between basilar membrane (BM) travelling waves, and tests the extent to which it is able to account for this specific suppression phenomenon. A simple mathematical model based on local nonlinear interactions was developed, and its predictions for a range of tone burst pairs were compared to corresponding TBOAE suppression data recorded from fourteen normally hearing human ears at a level of 60 dB p.e. SPL. Model predictions and mean TBOAE suppression data showed close agreement for all pairs of tone bursts. These results suggest that simultaneous suppression of TBOAEs can be explained solely in terms of the local nonlinear interaction-based mechanism. However, the involvement of other mechanisms, involving components generated at places basal to their characteristic place along the BM, cannot be excluded.     

Wever and Lawrence revisited: effects of nulling basilar membrane movement on concomitant whole-nerve action potential

It has been assumed for decades that mechanically stimulating hair cells, both inner and outer (IHC, OHC), leads to CM and subsequent neural activity. A test of that assumption was attempted in this experiment. Tone-pips of 300 msec duration at 4 or 5 kc/s with fast rise times were simultaneously presented to the cochleae of 10 chinchillas, through the external meatus and a hole drilled into the scala tympani. A round-window electrode allowed the recording of CM and computer-averaged whole-nerve action potentials (CAP). Stimulus levels and relative phase could be adjusted to yield CAPs of similar amplitude and shape to either stimulus alone. When the two stimuli were combined, the vectorial CM could be changed by about 30 db between maximum and minimum levels when delta phi was changed by 180 degrees. However, the combined CAP was relatively insensitive to delta phi. If basilar membrane motion was minimized at CM minimum, the data mean that some other principle than basilar membrane motion must underlie or generate neural activity. These data are not consistent with the traditional view that basilar membrane motion underlies sensitivity and frequency discrimination, and are congruent with theories of sensitivity of hair cells or their stereocilia to direct acoustic or electric stimulation, with basilar membrane mechanical stimulation assigned some secondary role. The author offers an electromodel comprising one system of basilar membrane motion of supramolecular dimensions leading to mechanical stimulation of OHCs and their large CM, and a second parallel system excited by the same stapes displacements but of submolecular dimensions leading to a propagated acoustic wave through the cochlear partition and to acoustic----electric transduction by the tectorial membrane; the output of that membrane is picked up in the fluids of the subtectorial space by the electro-sensitive IHCs and analyzed by them in some unknown manner for frequency. These IHCs are then the sole direct precursors of neural activity. A seeming anomaly was found in that at delta phi = CM minimum, when the traditional model would predict reduced basilar membrane movement, a reduced CM, consequent reduction in neural activity, and an increase in the latency of the N1 component of the CAP, latency was in fact slightly but uniformly decreased. It was suggested that in this phase condition the larger CM may have been correlated with the suppressive action of the OHCs upon the IHCs.     

Travelling wave motion along the pigeon basilar membrane

The basilar membrane (BM) motion in the pigeon was measured using the M?ssbauer technique. Tonotopic frequency mapping and travelling wave motion were observed over the basal 35% of the BM. The sensitivity and sharpness of the BM tuning depended on the physiological condition of the cochlea. The observed amplitude responses did not match the frequency threshold tuning curves of single primary auditory fibers.     

Effect of modulation of basilar membrane position on the cochlear microphonic

Certain characteristics of the extracellular cochlear microphonic (CM) recorded by intracochlear electrodes change in a bimodal manner as a function of prior acoustic exposure, intensity of stimulation, or stimulus frequency. In the present study, it as shown that blasing the basilar membrane position toward scala tympani serves to enhance the CM amplitude when the cochlea is unfatigued, when low-intensity stimuli are used, or when frequencies below the best frequency of a differential electrode pair are used. Conversely, after acoustic fatigue, or during high-intensity or high-frequency stimulation, the microphonic potential is enhanced by a movement of the basilar membrane toward the scala vestibuli. The two populations of hair cells, whose responses are enhanced and diminished on opposing positions of the basilar membrane, are probably outer and inner hair cells.     

Processing of acoustic stimuli in the inner ear--a review of recent research results

This review presents recent findings on the micromechanics of the basilar membrane. Active processes are essential for basilar membrane motion. It may be that contractile proteins within the outer hair cells play an important role for this amplification. The coding of acoustic information within the auditory nerve depends on spectral analysis (place information) as well as on the time structure of the stimulus. This latter time analysis seems to play a major role.     

Functional role of the olivo-cochlear bundle: a motor unit control system in the mammalian cochlea

A fiber optic lever is applied to the measurement of the motion of the basilar membrane motion in guinea pigs. In response to intense tones from either ear, the motion includes a substantial summating shift in the mean position in addition to a travelling wave originally described by von Békésy. His stroboscopic technique and most techniques used since have been concentrated upon measuring vibrations of the basilar membrane synchronous with the stimulus and have been insensitive to variations in the baseline position such as a summating component of motion analogous to the extracellular summating potential. In addition to the role of the outer hair cells in providing normal hearing sensitivity, they evidently play a role in regulating the mean position of the basilar membrane. For a fixed frequency, the polarity of the mean position varies systematically with sound level and place and summates with time since onset. Since these cells are the target cells for the olivocochlear bundle, homeostasis in the cochlea would appear to be linked efferent function and involve cochlear mechanics. The negative damping hypothesis asserts that hair cell activity is necessary for low thresholds. The results presented here demonstrate that OHC activity exists independent of neural thresholds. The discussion develops the concept that threshold losses are due to a mismatch of opposing tonic forces which normally maintain the mean position of the basilar membrane. Structure is examined in relation to function and the group of outer hair cells innervated by a single medial efferent neuron is identified as a motor unit. Implications of central control of individual motor units include peripheral involvement in selective attention tasks.     

Development of the mouse cochlea database (MCD)

The mouse cochlea database (MCD) provides an interactive, image database of the mouse cochlea for learning its anatomy and data mining of its resources. The MCD website is hosted on a centrally maintained, high-speed server at the following URL: http://mousecochlea.umn.edu. The MCD contains two types of image resources, serial 2D image stacks and 3D reconstructions of cochlear structures. Complete image stacks of the cochlea from two different mouse strains were obtained using orthogonal plane fluorescence optical microscopy (OPFOS). 2D images of the cochlea are presented on the MCD website as: viewable images within a stack, 2D atlas of the cochlea, orthogonal sections, and direct volume renderings combined with isosurface reconstructions. In order to assess cochlear structures quantitatively, "true" cross-sections of the scala media along the length of the basilar membrane were generated by virtual resectioning of a cochlea orthogonal to a cochlear structure, such as the centroid of the basilar membrane or the scala media. 3D images are presented on the MCD website as: direct volume renderings, movies, interactive QuickTime VRs, flythrough, and isosurface 3D reconstructions of different cochlear structures. 3D computer models can also be used for solid model fabrication by rapid prototyping and models from different cochleas can be combined to produce an average 3D model. The MCD is the first comprehensive image resource on the mouse cochlea and is a new paradigm for understanding the anatomy of the cochlea, and establishing morphometric parameters of cochlear structures in normal and mutant mice.     

New efficient stimuli for evoking frequency-specific auditory steady-state responses

ASSR is a promising tool for the objective frequency-specific assessment of hearing thresholds in children. The stimulus generally used for ASSR recording (single amplitude-modulated carrier) only activates a small area on the basilar membrane. Therefore, the response amplitude is low. A stimulus with a broader frequency spectrum can be composed by adding several cosines whose frequency intervals comply with the desired stimulus repetition rate. Compensation of the travelling wave delay on the basilar membrane is possible with a stimulus of this type. Through this, a better synchronization of the neural response can be obtained and, as a result, higher response amplitudes can be expected, particularly for low-frequency stimuli. The additional introduction of a frequency offset enables the use of a q-sample test for the response detection, especially important at 500 Hz. The results of investigations carried out on a large group of normally hearing test subjects have confirmed the efficiency of this stimulus design. The new stimuli lead to significantly improved ASSRs with higher SNRs and thus higher detection rates and shorter detection times.     

Backward Propagation of Otoacoustic Emissions

IntroductionsSounds impinging the eardrum are transmitted viamiddle ear ossicles to the oval window. Stapes vibra-tion creates pressure difference between the scala tym-pani and the scala vestibuli. This pressure differencecauses a movement of cochlear partition and adjacentcochlear fluids. Basilar membrane vibrations result indeflection of hair cell stereocilia, which gate ion chan-nels on their tips. This mechanical-to-electrical trans-duction process converts mechanical vibrations intoelec…     

Basilar membrane tuning properties in the specialised cochlea of the CF-bat, Rhinolophus ferrumequinum

The greater horseshoe bat has greatly expanded frequency mapping, and morphological specialisations, in the first half turn of its cochlea and a sudden transition to normal mapping. Amplitude and phase of vibration have been measured on various structures in the expanded and normal regions and have not revealed any sharply tuned responses. Amplitudes are much lower than those found in other species and frequently show a deep notch in the 77–84 kHz region. The high-frequency cut-off frequencies are tonotopically organised but deviate from the Bruns map, so that hair-cell tuning appears to occur at a frequency at which basilar membrane vibration is small. In the basal region, phase differences were frequently found between the inner and outer parts of the basilar membrane. These appear to be due to interaction between two components of motion and are probably not indicative of a further filtering mechanism. There is no evidence for reflection of the travelling wave at the transition.     

Simultaneous suppression of tone burst-evoked otoacoustic emissions--effect of level and presentation paradigm

There is conflict in the literature over whether individual frequency components of a transient-evoked otoacoustic emission (TEOAE) are generated within relatively independent "channels" along the basilar membrane (BM), or whether each component may be generated by widespread areas of the BM. Two previous studies on TEOAE suppression are consistent with generation within largely independent channels, but with a degree of interaction between nearby channels. However, both these studies reported significant suppression only at high stimulus levels, at which the "nonlinear" presentation paradigm was used. The present study clarifies the separate influences of stimulus level and presentation paradigm on this type of suppression. TEOAEs were recorded using stimulus tone bursts at 1, 2 and 3 kHz and a complex stimulus consisting of a digital addition of the three tone bursts, over a range of stimulus levels and both "linear" and "nonlinear" presentation paradigms. Responses to the individual tone bursts were combined offline and compared with responses to the complex stimuli. Results clearly demonstrate that TEOAE suppression under these conditions is dependent upon stimulus level, and not upon presentation paradigm. It is further argued that the data support the "local" rather than "widespread" model of TEOAE generation, subject to nonlinear interactions between nearby generation channels.     

Mechanics of the basilar membrane in Caiman crocodilus

Vibration measurements were made at a number of positions near the proximal (basal) end of the basilar membrane, and on the columella footplate, of Caiman crocodilus using a capacitive probe. The measurements established the existence of a mechanical travelling wave in this species. They showed no significant change of mechanical tuning with temperature, and were highly significantly different from previous reports of neural temperature sensitivity (Smolders, J. and Klinke, R. (1984): J. Comp. Physiol. 155, 19-30). Thus the neural sensitivity to temperature change appears not to depend upon basilar membrane mechanics. One interpretation of this is that the basilar membrane passively precedes an active temperature-sensitive filter. It was also found that the limbus supporting the basilar membrane had a measurable, but unturned, vibration and that the effect of draining scala tympani for the measurements was to increase the basilar membrane tuning frequency by a factor of about 1.5.     

Acoustic characteristics of different target vowels during the laryngeal telescopy

Objective

The aim of this study was to investigate the acoustic characteristics of target vowels phonated in normal voice persons while performing laryngeal telescopy. The acoustic characteristics are compared to show the extent of possible difference to speculate their impact on phonation function.

Methods

Thirty-four male subjects aged 20–39 years with normal voice were included in this study. The target vowels were /i/ and /?/. Recording of voice samples was done under natural phonation and during laryngeal telescopy. The acoustic analysis included the parameters of fundamental frequency, jitter, shimmer and noise-to-harmonic ratio.

Results

The sound of a target vowel /?/ was perceived identical in more than 90% of the subjects by the examiner and speech language pathologist during the telescopy. Both /i/ and /?/ sounds showed significant difference when compared with the results under natural phonation. There was no significant difference between /i/ and /?/ during the telescopy.

Conclusion

The present study showed that change in target vowels during laryngeal telescopy makes no significant difference in the acoustic characteristics. The results may lead to the speculation that the phonation mechanism was not affected significantly by different vowels during the telescopy. This study may suggest that in the principle of comfortable phonation, introduction of the target vowels /i/ and /?/ is practical.     

Synchronized responses of primary auditory fibre-populations in Caiman crocodilus (L.) to single tones and clicks

Measurements of the responses to tones and clicks were made from single primary auditory fibres of the caiman. The distribution of the amplitude and phase of the fundamental component of the response rate modulation over the best frequencies of the fibres is comparable to that reported in the cat, despite the fact that the basilar membrane in caiman is only 4.5 mm long. However, much higher intensities are needed in the caiman (75-85 dB SPL) than reported in the cat (20 dB SPL) to obtain systematic distributions of the phase of the responses, probably due to the larger scatter of the phase responses in the caiman. The slopes of the phase distributions are very similar to those in cat. Single unit phase responses as a function of stimulus frequency at 85 dB SPL can be approximated by one, or in fibres with low best frequency, two straight lines. At lower intensities the deviation of the phase-frequency responses from a straight line increases as the group delay at the best frequency becomes larger. The shortest latencies of click responses are obtained with rarefaction clicks. Group delay estimates obtained from the responses to clicks and from the straight line approximations of the phase-frequency responses are related in a way expected for linear filter systems and accurately predict the measured distributions of the phase of the responses over the neural best frequency. The obtained group delays and click latencies in the caiman are very similar to those reported by other workers in the cat, the squirrel monkey and the treefrog, despite large morphological and probably functional differences of their inner ears. The click latencies are also very similar to those in the pigeon. The results are consistent with the existence of a mechanical travelling wave reported previously on the basilar membrane of the caiman, but at the same stimulus level the phase characteristic of the present single unit responses is steeper and the wave length estimates from the neural population phase distributions are shorter than those observed directly in the motion of the basilar membrane. Since the neural responses are an indirect estimate of the basilar membrane motion it cannot be decided whether the difference between neural and mechanical data is due to deterioration of the basilar membrane responses during the direct measurements or whether the basilar membrane response is sharpened by additional tuning mechanisms.     

Influence of encoding and acoustic similarity on the ear advantage and lag effect in dichotic listening.

Experiments with simultaneous and time lag dichotic listening conditions were used to test two hypotheses concerning the right ear advantage and lag effect in dichotic listening. One hypothesis is based on the similarity of acoustic spectra, and the other is based on a categorization of speech sounds as being either encoded or not encoded. Natural vowels and consonant-vowel syllables were used to obtain seven different types of speech stimuli: stop vowel syllables, fricative vowel syllables, stop burst noise, fricative noise, stop vowel transitions, fricative vowel transitions and steady state vowels. The presentation conditions were monaural, simultaneous dichotic, and dichotic with interaural time delays of 15, 30, 60, and 90 msec. With monaural presentations, all stimuli were identifiable above chance levels. For the simultaneous dichotic condition, significant right ear advantages occurred for stop vowel syllables, fricative vowel syllables, stop burst noise, and steady state vowels. For the time lag conditions, stop vowel syllables, stop bursts, and fricative noise produced consistent lag effects, but steady state vowels produced consistent lead effects. In general, the results gave stronger support to the hypothesis of acoustic similarity than to the encoding hypothesis in that stop burst noise produced both a right ear advantage and a lag effect whereas consonant-vowel transitions produced neither a right ear advantage nor a lag effect.     

Identification of synthetic vowels by patients using the Symbion multichannel cochlear implant

In this report we describe the vowel identification ability of eight patients who scored above 70% correct on a test of spondee word identification. The stimuli were 12 synthetic vowels in "bVt" format. The vowels differed in the frequency of F1, F2, and F3. The mean identification accuracy was 60% correct. Front vowels and dipthongs, (3), and (u) were relatively well identified. The vowels in "but," "bought," and "bout," which were characterized by high F1s and low F2s were not well identified. The results are consistent with a model of recognition in which F1 is specified, with relatively good accuracy, by a rate code and in which extreme values of F2 are specified by a rate/place code.     

Implications from cochlear implant insertion for cochlear mechanics

It is usually thought that the displacements of the two inner ear windows induced by sound stimuli lead to pressure differences across the basilar membrane and to a passive mechanical traveling wave progressing along the membrane. However, opening a hole in the sealed inner ear wall in experimental animals is surprisingly not accompanied by auditory threshold elevations. It has also been shown that even in patients undergoing cochlear implantation, elevation of threshold to low-frequency acoustic stimulation is often not seen accompanying the making of a hole in the wall of the cochlea for insertion of the implant. Such threshold elevations would be expected to result from opening the cochlea, reducing cochlear impedance, altering hydrodynamics. These considerations can be taken as additional evidence that it may not be the passive basilar membrane traveling wave which elicits hearing at low sound intensities, but rather factors connected with cochlear fluid pressures and fluid mechanics.     

Cochlear mechanisms at low frequencies in the guinea pig

Summary The study of the cochlear microphonic and of the intracochlear sound pressure in guinea pigs shows that the behavior of the cochlea at very low frequencies is controlled by three discrete elements: (a) the compliance of the whole basilar membrane; (b) the acoustic resistance of the helicotrema; (c) the compliance of the round window. The part of each of these elements has been established. The compliance of the whole basilar membrane produces constant amplitudes at frequencies lower than the minimum frequency at which a travelling wave is present (130 Hz). In fact, this constant amplitude range is limited by connection of the two cochlear scalae through the helicotrema resistance. This protecting mechanism produces an attenuation slope for frequencies lower than 80 Hz. The compliance of the round window does not modify the slope of the cochlear microphonic, but it induces a constant sound pressure in scala tympani up to 200 Hz. Decreasing of the sound pressure in the scala vestibuli is, therefore, limited for frequencies less than 30 Hz by this constant value of the sound pressure in scala tympani.Presented at the 18th Workshop on Inner Ear Biology in Montpellier/La Grande Motte, September 14–16, 1981