William S. Rhode

Observations of the Vibration of the Basilar Membrane in Squirrel Monkeys using the Mössbauer Technique

The amplitude and the phase of vibration of the basilar membrane and the bony limbus of the cochlea were measured in living squirrel monkeys using the Mossbauer technique. In the middle ear, the vibration of the malleus (and occasionally the incus) was measured. The Mossbauer technique makes possible the measurement of very small velocities, e.g., 0.2 mm/sec. This sensitivity permits measurement of the motion of the malleus at sound‐pressure levels (SPLs) of 90 to 110 dB and measurement of the motion of the basilar membrane at 70 to 120 dB SPL, depending on the frequency. The basilar membrane vibrates nonlinearly for frequencies which produce the largest deflections at the spot on the basilar membrane under observation. The ratio of the displacement of the basilar membrane to that of the malleus was observed to have the following characteristics: (1) As the frequency is increased from a low value, its amplitude increases at 6 dB/oct until just below the maximum ratio where the slope increases to about 24 ...

Some observations on cochlear mechanics.

A set of experiments was conducted using the Mössbauer effect to determine the vibratory characteristics of the basilar membrane, Reissners membrane, the malleus, incus, and oval window in squirrel monkey. A few measurements were also made in guinea pig in the basal cochlear region. The nonlinear vibration properties of the basilar membrane are described in detail for the midfrequency region in the squirrel monkey. Only in this region have nonlinear effects been observed. A comparison of mechanical and neural data indicates good qualitative agreement.

Evidence from Mössbauer experiments for nonlinear vibration in the cochlea

The Mossbauer technique has been applied to the measurement of vibration of the basilar membrane in the squirrel monkey cochlea. Both steady‐state and transient responses have been recorded in the 7–8‐kHz locus of the cochlea. The steady‐state response indicates that the basilar membrane vibrates nonlinearly for frequencies of stimulation near or greater than the characteristic frequency. The nonlinearity can be observed at the lowest levels of stimulation, 70–80 dB SPL, for which measurements could be made. The nonlinearity extends to lower frequencies and the basilar membrane transfer function tends to broaden as SPL is increased. Rapid postmortem changes occur in the cochlea: (1) the amplitude of the transfer ratio (basilar membrane/malleus) decreases 10–15 dB over a period of several hours with a downward shift of 1.5–3 kHz in the characteristic frequency of the basilar membrane at a given location; (2) the low‐frequency slope of the transfer ratio settles to 6 dB/octave by 6 h after death; (3) the sl...

Basilar membrane mechanics in the hook region of cat and guinea-pig cochleae: sharp tuning and nonlinearity in the absence of baseline position shifts.

A heterodyne laser interferometer was used to observe the movements of small (approximately 20 microns) stainless-steel beads placed on the basilar membrane in the hook region of cat and guinea-pig cochleae. In several preparations, the displacement patterns observed exhibited sharp nonlinear tuning; in one cat this tuning was comparable to that commonly observed in single auditory-nerve fibers. The most sensitive frequencies of the preparations ranged from 31-40 kHz in the cat, and 28-32 kHz in the guinea-pig. The sharp tuning and nonlinearity of the basilar membrane responses was not apparent in surgically or acoustically traumatized preparations. The response nonlinearities were susceptible to temporary threshold shifts and disappeared within a few minutes post-mortem. Stimulus-related shifts in the baseline position of the basilar membrane were not apparent at low stimulus levels. Such shifts were occasionally observed at higher stimulus levels (e.g., > 90 dB SPL), but never approached the fundamental (oscillatory) component of basilar membrane vibration in magnitude. These findings are discussed in relation to previous observations by other workers.

Nonlinear mechanics at the apex of the guinea-pig cochlea

A heterodyne laser interferometer was used to observe the sound-evoked displacement patterns of Reissners membrane and various other structures in the apical turn of the guinea-pig cochlea. Most structures (including the basilar membrane) were similarly tuned, and had best frequencies in the 200-350Hz range. A distinct notch was usually observed approximately 0.7 octaves above the best frequency, and amplitude- and phase-plateaus were observed at higher frequencies. In most other respects, however, the mechanical tuning resembled the frequency-threshold curves of low frequency cochlear nerve fibers. In five reasonably intact, in vivo preparations, the frequency of the mechanical sensitivity notch was intensity-dependent: Compressive nonlinearities were observed above approximately 80 dB SPL on the low-frequency side of the notch, with antagonistically expansive nonlinearities on the high-frequency side. Two-tone suppression was observed in one of these preparations. Stimulus-related baseline position shifts were observed in another in vivo preparation. No such nonlinearities were observed in structurally damaged and/or > 1 hour post-mortem preparations. However, more robust nonlinearities were observed in all preparations at higher levels of stimulation (e.g. > 100-110 dB SPL). These high-level nonlinearities diminished only slowly after death, and gave rise to various effects, including time-dependent (i.e. adapting) and severely distorted (e.g. peak-split and/or dc-shifted) responses.

Characteristics of tone-pip response patterns in relationship to spontaneous rate in cat auditory nerve fibers

The responses of single auditory nerve (AN) fibers in the cat were recorded in response to 25 ms tone pips. Peristimulus time histograms (PSTH) of discharge patterns recorded from fibers with high spontaneous rates (high SRs), show that the discharge rate rapidly adapts to a much lower steady-state level over a 15 ms period with shorter times for units with best frequencies (CFs) greater than 5 kHz. The PSTHs of auditory nerve fibers with low SRs do not show this pattern of rapid adaptation. Differences between the high and low SR populations include higher thresholds, better tuning, and longer latency in the low SR population. The peak-to-steady-state discharge ratio is an increasing function of SR and CF; it varies from 1.0 for fibers with SR = 0 to over 8 for fibers with high SRs and CFs near 10 kHz. This ratio increases with increasing stimulus intensity and stimulus recovery time. The high SR population shows a number of responses to transients which are weak or absent in the low SR population. Increasing the recovery time shortened the latency of both high and low SR AN fibers by as much as 1 ms. A number of other response properties of AN fibers are also reported that are important when interpreting the responses of cochlear nucleus neurons to tone pips.

Study of mechanical motions in the basal region of the chinchilla cochlea

Measurements from the 1-4-mm basal region of the chinchilla cochlea indicate the basilar membrane in the hook region (12-18 kHz) vibrates essentially as it does more apically, in the 5-9-kHz region. That is, a compressive nonlinearity in the region of the characteristic frequency, amplitude-dependent phase changes, and a gain relative to stapes motion that can attain nearly 10,000 at low levels. The displacement at threshold for auditory-nerve fibers in this region (20 dB SPL) was approximately 2 nm. Measurements were made at several locations in individual animals in the longitudinal and radial directions. The results indicate that there is little variability in the phase of motion radially and no indication of higher-order modes of vibration. The data from the longitudinal studies indicate that there is a shift in the location of the maximum with increasing stimulus levels toward the base. The cochlear amplifier extends over a 2-3-mm region around the location of the characteristic frequency.

Transient response of the basilar membrane measured in squirrel monkeys using the Mössbauer effect.

Measurements of the transient response of the basilar membrane were conducted using the Mossbauer effect on 33 squirrel monkeys using an experimental preparation identical to that of Rhode (1971). The stimuli were acoustic clicks 150 μsec in duration repeated 100 000–400 000 times. The amplitude of the click was varied and the responses of the malleus and of the basilar membrane at a point in the basal turn were measured. The basilar membrane’s click response is oscillatory, with a period near that of the characteristic frequency. The first few response peaks behave almost linearly with stimulus intensity, while the later peaks exhibit a pronounced nonlinearity. This behavior is shown to be consistent with the nonlinearity reported using steady‐state measurement methods (Rhode, 1971). The transient response observed in some of the preparations was very lightly damped; however, a wide range in the damping of the responses was found in the different animals. A progressive increase in the rate of decay of th...

Two-tone suppression and distortion production on the basilar membrane in the hook region of cat and guinea pig cochleae

Two-tone suppression and two-tone distortion were investigated at the level of the basilar membrane in the hook region of cat and guinea pig cochleae using a displacement-sensitive laser interferometric measurement system. The system allowed measurements to be performed at physiological stimulus levels in the cochlear region tuned to 30-35 kHz in cat and 29 kHz in guinea pig. The amplitude of vibration of the basilar membrane due to a probe tone at the characteristic frequency (CF) was attenuated during the presentation of a simultaneous suppressor tone either above or below CF. The amount of suppression depended on the intensities of both probe and suppressor, and the relationship of the suppressor frequency to the CF. Suppressors at frequencies more than an octave below the CF attenuated the responses to the CF probe at a rate of up to 1 dB/dB, with little variation based on suppressor frequency. As the suppressor frequency was increased above CF the rate of suppression decreased rapidly. The lowest suppressor intensity at which attenuation of the probe response was observed did not vary in direct proportion to the probe intensity. This suppression threshold often varied only a few dB SPL when the probe was varied over a 20 dB SPL range. In a few instances the rate of attenuation was as much as a factor of two greater at the lowest probe intensities than at higher intensities. It is noteworthy that suppression was found when the frequency of the suppressor was either above or below CF in the same preparation. Low frequency suppressor tones suppress basilar membrane motion at the CF when the basilar membrane undergoes displacement toward either scala. The maximum suppression occurs around 100 microseconds after the peak excursions caused by the low frequency biasing tone. Two-tone distortion products were often observed even at stimulus levels below those causing two-tone suppression at the site studied. The cubic difference tone (CDT) was the most prominent of the distortion products. The level of the CDT component varied nonmonotonically with the level of either of the primary tones. Responses at the difference frequency between the two primaries were usually below the noise floor of the recording system. The existence of both two-tone distortion and two-tone suppression was dependent on the presence of a cochlear nonlinearity.

Basilar membrane mechanics in the 6–9kHz region of sensitive chinchilla cochleae

The vibration of the basilar membrane in the 6-9 kHz region in the chinchilla cochlea has been studied using a displacement sensitive interferometer. Displacements of 0.7-1.4 nm at 0 dB sound pressure level have been obtained. At the characteristic frequency (CF), rate-of-growth (ROG) functions computed as the slope of input-output (IO) functions can be as low as 0.1 dB/dB. IO functions for frequencies > CF have ROGs near 0 dB/dB and can have notches characterized by both negative slopes and expansive ROGs, i.e., > 1 dB/dB. For frequencies < 0.6*CF, ROGs > 1.2 dB/dB were found. Cochlear gain is shown to be greater than 60 dB in sensitive preparations with a single cochlea having nearly 80 dB gain. The compressive nature of the cochlea remains at all levels though it is masked at frequencies > CF when the amplitude of a compression wave exceeds that of the traveling wave. The compression wave produces the plateau region of the mechanical response at high intensities and has a nearly constant phase versus frequency function implying a high velocity. The summation of the traveling and compression waves explains the occurrence of the notches in both the IO and iso-intensity functions. Vibration of the osseous spiral limbus may alter the drive to inner hair cells.