Two-tone distortion at different longitudinal locations on the basilar membrane

When listening to two tones at frequency f1 and f2 (f2>f1), one can hear pitches not only at f1 and f2 but also at distortion frequencies f2-f1, (n+1)f1-nf2, and (n+1)f2-nf1 (n=1,2,3...). Such two-tone distortion products (DPs) also can be measured in the ear canal using a sensitive microphone. These ear-generated sounds are called otoacoustic emissions (OAEs). In spite of the common applications of OAEs, the mechanisms by which these emissions travel out of the cochlea remain unclear. In a recent study, the basilar membrane (BM) vibration at 2f1-f2 was measured as a function of the longitudinal location, using a scanning laser interferometer. The data indicated a forward traveling wave and no measurable backward wave. However, this study had a relatively high noise floor and high stimulus intensity. In the current study, the noise floor of the BM measurement was significantly decreased by using reflective beads on the BM, and the vibration was measured at relatively low intensities at more than one longitudinal location. The results show that the DP phase at a basal location leads the phase at an apical location. The data indicate that the emission travels along the BM from base to apex as a forward traveling wave, and no backward traveling wave was detected under the current experimental conditions.     

Cochlear model with three-dimensional fluid, inner sulcus and feed-forward mechanism

A three-dimensional model of the guinea pig cochlea using the phase-integral method is presented. This model incorporates the viscous fluid effects in the cochlea, dimensional and material property variation along the cochlear duct and the active feed-forward mechanism of the outer hair cells. Two degrees of freedom of the basilar membrane are considered, which results in two traveling waves propagating along the duct for a given frequency. Basilar membrane response with the active feed-forward mechanism compares favorably with published experimental measurements.     

A comparison of OAEs arising from different generation mechanisms in guinea pig

Otoacoustic emissions provide unambiguous evidence that the cochlea supports energy propagation both towards, and away from, the stapes. The standard wave model for energy transport and cochlear mechanical amplification provides for compressional and inertial waves to transport this energy, the compressional wave through the fluids and the inertial wave along the basilar membrane via fluid coupling. It is generally accepted that energy propagation away from the stapes is dominated by a traveling wave mechanism along the basilar membrane. The mechanism by which energy is predominantly transported back to the stapes remains controversial. Here, we compared signal onset delay measurements and rise/steady-state/fall times for SFOAEs and 2f1-f2 OAEs (f2/f1=1.2) obtained using a pulsed-tone paradigm in guinea pig. Comparison of 2f1-f2 OAE signal onset delay for the OAE arising from the f2 region with SFOAE signal onset delay (matched to the f2 stimulus frequency) based on signal onset occurring at 10% of the peak signal amplitude was suggestive of a bi-directional traveling wave mechanism. However, significant variability in signal onset delay and signal rise, steady-state duration, and fall times for both the 2f1-f2 OAE and SFOAE was found, qualifying this interpretation. Such variability requires explanation, awaiting further studies.     

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 delays measured with amplitude-modulated tone-burst-evoked OAEs

Delay times in the mammalian cochlea, whether from measurement of basilar membrane (BM) vibration or otoacoustic emissions (OAEs) have, to date, been largely based on phase-gradient estimates from steady-state responses. Here we report cochlear delays measured directly in the time domain from OAEs evoked by amplitude-modulated tone-burst (AMTB) stimuli. Measurement using OAEs provides a non-invasive estimate of cochlear delay but is confounded by the complexity of generation of such OAEs. At low to moderate stimulus levels, and provided that the stimulus frequency range does not include a region of the cochlea where there is a large change in effective reflectance, AMTB stimuli evoke an OAE with an envelope shape that is similar to the stimulus and allow a direct calculation of cochlear group delay. Such delays are commensurate with BM estimates of delay, estimates of cochlear delay inferred from neural recordings, and previous OAE measures of delay in the guinea pig. However, a nonlinear distortion mechanism, variation in effective reflectance, and intermodulation distortion products generated by the nonlinear interaction in the cochlea of the carrier and sidebands of the AMTB stimulus, may all contribute to OAEs arising with envelope shapes that are not a scaled representation of the stimulus, confounding the estimation of cochlear group delay.     

Recent developments in cochlear physiology

Recent findings in cochlear physiology have caused many of our long held ideas about how sound is analyzed by the ear to be reevaluated. This article describes changes which have occurred in three classical ideas of cochlear transduction: (1) There is a gradient of frequency representation along the cochlea with high frequencies being represented at the base and lower frequencies represented progressively toward the apex. It is now known that the specific frequency which is represented at a given location along the cochlea is not invariant but changes systematically during the normal development of hearing. (2) The place code and frequency tuning along the cochlea are due to the conventional traveling wave of von Békésy and basilar membrane mechanics. Experiments in nonmammalian vertebrates which lack a traveling wave have shown that other mechanisms, including the mechanical resonance of hair cell stereocilia, may contribute to tonotopic organization and frequency tuning. It is possible that hair cell stereocilia also contribute to frequency representation and tuning in the mammalian cochlea. (3) The vibration of the basilar membrane to sound is determined by its passive mechanical properties. It is now known that the response of the basilar membrane, and that of the cochlear partition as a whole, is influenced by physiological processes which utilize metabolic energy. The active processes are likely expressed through the motile activity of outer hair cells.     

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.     

Spatial representation of basilar membrane movement with 3D computer graphics

A mathematical model of the cochlea was implemented on a computer. The basilar membrane motion was computed for single, two, and multi-tone stimuli as well as for musical sounds and vowels. The pattern of the travelling waves were presented in three-dimensional color computer graphics. The high performance 3D graphics system performs local hidden surface removal, 3D geometric transformations and supports local lighting models to generate truly realistic shading for complex 3D objects. An addressable 1280 by 1024 pixel matrix assures crisp, precise resolution of the finest detail in the graphic images. The spatial pattern of basilar membrane motion conveys an impression of the image of acoustic stimuli on the basilar membrane. Firstly, the motion pattern of the travelling wave to a single tone is presented. The superposition of several tones (two-tone, multi-tone) causes a superposition of the travelling waves along the basilar membrane whereby the place principle in the cochlear partition becomes more clearly recognizable. Sounds (flute and violin) and vowels (German "u" and "i") evoke a complex motion pattern on the basilar membrane. The realization of the chronological order of movements on the basilar membrane can be made by computer animation. This enables the analysis of the space-time patterns of complex acoustic stimuli.     

Otoacoustic Emissions from Residual Oscillations of the Cochlear Basilar Membrane in a Human Ear Model

Sounds originating from within the inner ear, known as otoacoustic emissions (OAEs), are widely exploited in clinical practice but the mechanisms underlying their generation are not entirely clear. Here we present simulation results and theoretical considerations based on a hydrodynamic model of the human inner ear. Simulations show that, if the cochlear amplifier (CA) gain is a smooth function of position within the active cochlea, filtering performed by a middle ear with an irregular, i.e., nonsmooth, forward transfer function suffices to produce irregular and long-lasting residual oscillations of cochlear basilar membrane (BM) at selected frequencies. Feeding back to the middle ear through hydrodynamic coupling afforded by the cochlear fluid, these oscillations are detected as transient evoked OAEs in the ear canal. If, in addition, the CA gain profile is affected by irregularities, residual BM oscillations are even more irregular and tend to evolve towards self-sustaining oscillations at the loci of gain irregularities. Correspondingly, the spectrum of transient evoked OAEs exhibits sharp peaks. If both the CA gain and the middle-ear forward transfer function are smooth, residual BM oscillations have regular waveforms and extinguish rapidly. In this case no emissions are produced. Finally, and paradoxically albeit consistent with observations, simulating localized damage to the CA results in self-sustaining BM oscillations at the characteristic frequencies (CFs) of the sites adjacent to the damage region, accompanied by generation of spontaneous OAEs. Under these conditions, stimulus-frequency OAEs, with typical modulation patterns, are also observed for inputs near hearing threshold. This approach can be exploited to provide novel diagnostic tools and a better understanding of key phenomena relevant for hearing science.     

Experimental look at cochlear mechanics.

The mechanisms by which the organ of Corti is stimulated by acoustic stimuli are discussed on the basis of experimental observations. This discussion refers to the resonance theory as well as to the traveling wave (TW) theory. The measurement of the basilar membrane displacements, of the cochlear microphonic (CM) responses to pure tones and impulses, and the recording of the intracochlear acoustic pressure seem to indicate that, at least in the basal part of the cochlea and for frequencies up to the characteristic frequency of a given location, the cochlear responses do not exhibit large phase lags and long delays which characterize the one-dimensional long-wave models (in which a TW transports the energy along the cochlear partition). These experimental observations suggest that the cochlear partition is excited simultaneously as a whole, more or less like a bank of resonators, as proposed a long time ago by Helmholtz.     

Effects of a perilymphatic fistula on the passive vibration response of the basilar membrane

In this study, a three-dimensional finite-element model of the passive human cochlea was created. Dynamic behavior of the basilar membrane caused by the vibration of the stapes footplate was analyzed considering a fluid-structure interaction with the cochlear fluid. Next, the effects of a perilymphatic fistula (PLF) on the vibration of the cochlea were examined by making a small hole on the wall of the cochlea model. Even if a PLF existed in the scala vestibuli, a traveling wave was generated on the basilar membrane. When a PLF existed at the basal end of the cochlea, the shape of the traveling wave envelope showed no remarkable change, but the maximum amplitude became smaller at the entire frequency range from 0.5 to 5kHz and decreased with decreasing frequency. In contrast, when a PLF existed at the second turn of the cochlea, the traveling wave envelope showed a notch at the position of the PLF and the maximum amplitude also became smaller. This model assists in elucidating the mechanisms of hearing loss due to a PLF from the view of dynamics.     

How does the inner ear generate distortion product otoacoustic emissions?. Results from a realistic model of the human cochlea

There is general agreement that distortion product (DP) otoacoustic emissions elicited by stimuli up to 80-90 dB SPL originate from the saturating nonlinearity of the cochlear amplifier at the basilar membrane site, S, where the responses to the two primary tones overlap. There are, however, different interpretations of how the inner ear transmits the effects of this process to the stapes. The supporters of transmission line models assert that the phenomenon depends upon two main mechanisms: (1) the generation of forward and backward traveling waves (TWs) by DP oscillations at S; (2) the backward propagation of wave components reflected by 'micromechanical impedance perturbations' at the sites where the DP TWs peak. However, quantitative predictions based on this view are still lacking. In contrast, here we show, using a nonlinear hydrodynamic model, that the emissions are propagated almost instantaneously through the fluid.     

Some current concepts of cochlear mechanics

The development of the current concepts is reviewed in historical perspective. Helmholtz's hypothesis of basilar-membrane resonance was partially confirmed and partially defeated by Békésy's experiments on models and postmortem cochlear preparations. He discovered that sound was propagated in the cochlea in the form of traveling waves which reached a flat maximum at a frequency-dependent location. Mathematical theory explained this type of sound propagation as a special case of surface waves. Johnstone and his coworkers discovered that the maximum of cochlear vibration in living animals was much sharper than postmortem, and more recently Khanna and Johnstone independently determined the maximum to be nearly as sharp as the tuning curves of the inner hair cells and the auditory-nerve fibers. These findings, together with the work at the Massachusetts Institute of Technology on alligator lizards, have led to new concepts of cochlear mechanics which include hypothesized micromechanical processes in the organ of Corti. These concepts deal not only with the sharpness of basilar-membrane tuning but also with the details of the basilar-membrane amplitude and phase characteristics, as well as with the hair cell and neural tuning curves and response phases. They suggest that some sharpening of the tuning curves occurs between the basilar membrane and hair-cell responses. Such sharpening has been demonstrated in lizards, but in the mammalian ear, the relation is less clear.     

Estimation of Round-Trip Outer-Middle Ear Gain Using DPOAEs

The reported research introduces a noninvasive approach to estimate round-trip outer-middle ear pressure gain using distortion product otoacoustic emissions (DPOAEs). Our ability to hear depends primarily on sound waves traveling through the outer and middle ear toward the inner ear. The role of the outer and middle ear in sound transmission is particularly important for otoacoustic emissions (OAEs), which are sound signals generated in a healthy cochlea and recorded by a sensitive microphone placed in the ear canal. OAEs are used to evaluate the health and function of the cochlea; however, they are also affected by outer and middle ear characteristics. To better assess cochlear health using OAEs, it is critical to quantify the effect of the outer and middle ear on sound transmission. DPOAEs were obtained in two conditions: (i) two-tone and (ii) three-tone. In the two-tone condition, DPOAEs were generated by presenting two primary tones in the ear canal. In the three-tone condition, DPOAEs at the same frequencies (as in the two-tone condition) were generated by the interaction of the lower frequency primary tone in the two-tone condition with a distortion product generated by the interaction of two other external tones. Considering how the primary tones and DPOAEs of the aforementioned conditions were affected by the forward and reverse outer-middle ear transmission, an estimate of the round-trip outer-middle ear pressure gain was obtained. The round-trip outer-middle ear gain estimates ranged from ?39 to ?17 dB between 1 and 3.3 kHz.     

Spectral Ripples in Round-Window Cochlear Microphonics: Evidence for Multiple Generation Mechanisms

The cochlear microphonic (CM) results from the vector sum of outer hair cell transduction currents excited by a stimulus. The classical theory of CM generation—that the response measured at the round window is dominated by cellular sources located within the tail region of the basilar membrane (BM) excitation pattern—predicts that CM amplitude and phase vary little with stimulus frequency. Contrary to expectations, CM amplitude and phase-gradient delay measured in response to low-level tones in chinchillas demonstrate a striking, quasiperiodic pattern of spectral ripples, even at frequencies >?5 kHz, where interference with neurophonic potentials is unlikely. The spectral ripples were reduced in the presence of a moderate-level saturating tone at a nearby frequency. When converted to the time domain, only the delayed CM energy was diminished in the presence of the saturator. We hypothesize that the ripples represent an interference pattern produced by CM components with different phase gradients: an early-latency component originating within the tail region of the BM excitation and two delayed components that depend on active cochlear processing near the peak region of the traveling wave. Using time windowing, we show that the early, middle, and late components have delays corresponding to estimated middle-ear transmission, cochlear forward delays, and cochlear round-trip delays, respectively. By extending the classical model of CM generation to include mechanical and electrical irregularities, we propose that middle components are generated through a mechanism of “coherent summation” analogous to the production of reflection-source otoacoustic emissions (OAEs), while the late components arise through a process of internal cochlear reflection related to the generation of stimulus-frequency OAEs. Although early-latency components from the passive tail region typically dominate the round-window CM, at low stimulus levels, substantial contributions from components shaped by active cochlear processing provide a new avenue for improving CM measurements as assays of cochlear health.     

Predicting the effect of post-implant cochlear fibrosis on residual hearing

Intracochlear scarring is a well-described sequela of cochlear implantation. We developed a mathematical model of passive cochlear mechanics to predict the impact that this might have upon residual acoustical hearing after implantation. The cochlea was modeled using lumped impedance terms for scala vestibuli (SV), scala tympani (ST), and the cochlear partition (CP). The damping of ST and CP was increased in the basal one half of the cochlea to simulate the effect of scar tissue. We found that increasing the damping of the ST predominantly reduced basilar membrane vibrations in the apex of the cochlea while increasing the damping of the CP predominantly reduced basilar membrane vibrations in the base of the cochlea. As long as intracochlear scarring continues to occur with cochlear implantation, there will be limitations on hearing preservation. Newer surgical techniques and electrode technologies that do not result in as much scar tissue formation will permit improved hearing preservation.     

Measurement of basilar membrane vibrations and evaluation of the cochlear condition

The experimental procedure for measuring basilar membrane responses to acoustic signals is described. The surgical procedure developed for opening the cochlea with minimal trauma is presented. Each experiment included sound pressure level measurements to define the input signal, cochlear microphonic (CM) measurements to monitor the cochlear condition, interferometric measurements and histological evaluation of the cochleas. The characteristic frequency (CF) and the sensitivity at CF for the basilar membrane response is correlated with the change of CM response observed in six animals. It is demonstrated that both tuning characteristics are extremely sensitive to cochlear trauma as evidenced by changes of CM.     

Olivocochlear efferents: anatomy, physiology, function, and the measurement of efferent effects in humans

This review covers the basic anatomy and physiology of the olivocochlear reflexes and the use of otoacoustic emissions (OAEs) in humans to monitor the effects of one group, the medial olivocochlear (MOC) efferents. MOC fibers synapse on outer hair cells (OHCs), and activation of these fibers inhibits basilar membrane responses to low-level sounds. This MOC-induced decrease in the gain of the cochlear amplifier is reflected in changes in OAEs. Any OAE can be used to monitor MOC effects on the cochlear amplifier. Each OAE type has its own advantages and disadvantages. The most straightforward technique for monitoring MOC effects is to elicit MOC activity with an elicitor sound contralateral to the OAE test ear. MOC effects can also be monitored using an ipsilateral elicitor of MOC activity, but the ipsilateral elicitor brings additional problems caused by suppression and cochlear slow intrinsic effects. To measure MOC effects accurately, one must ensure that there are no middle-ear-muscle contractions. Although standard clinical middle-ear-muscle tests are not adequate for this, adequate tests can usually be done with OAE-measuring instruments. An additional complication is that most probe sounds also elicit MOC activity, although this does not prevent the probe from showing MOC effects elicited by contralateral sound. A variety of data indicate that MOC efferents help to reduce acoustic trauma and lessen the masking of transients by background noise; for instance, they aid in speech comprehension in noise. However, much remains to be learned about the role of efferents in auditory function. Monitoring MOC effects in humans using OAEs should continue to provide valuable insights into the role of MOC efferents and may also provide clinical benefits.     

Human whole-nerve response to clicks of various frequency.

The averaged VIIIth nerve response to third-octave clicks at 500, 2000 and 8000 Hz was recorded from the promontory of 4 normal-hearing young adults. As click frequency is lowered, the N, latency increases in a manner consistent with the changes in velocity of the cochlear traveling wave. This finding suggests that clicks of different spectral content stimulate different regions of the basilar membrane. N amplitude shows a general increase with frequency; this observation appears related to the increased synchrony of neural firing that results from the higher velocity of the traveling wave in the more basal portions of the cochlea.