Tuning Curves Of Auditory Nerve Fibers Research Articles

Estimation of Cochlear Frequency Selectivity Using a Convolution Model of Forward-Masked Compound Action Potentials.

Frequency selectivity is a fundamental property of the peripheral auditory system; however, the invasiveness of auditory nerve (AN) experiments limits its study in the human ear. Compound action potentials (CAPs) associated with forward masking have been suggested as an alternative to assess cochlear frequency selectivity. Previous methods relied on an empirical comparison of AN and CAP tuning curves in animal models, arguably not taking full advantage of the information contained in forward-masked CAP waveforms. To improve the estimation of cochlear frequency selectivity based on the CAP, we introduce a convolution model to fit forward-masked CAP waveforms. The model generates masking patterns that, when convolved with a unitary response, can predict the masking of the CAP waveform induced by Gaussian noise maskers. Model parameters, including those characterizing frequency selectivity, are fine-tuned by minimizing waveform prediction errors across numerous masking conditions, yielding robust estimates. The method was applied to click-evoked CAPs at the round window of anesthetized chinchillas using notched-noise maskers with various notch widths and attenuations. The estimated quality factor Q10 as a function of center frequency is shown to closely match the average quality factor obtained from AN fiber tuning curves, without the need for an empirical correction factor. This study establishes a moderately invasive method for estimating cochlear frequency selectivity with potential applicability to other animal species or humans. Beyond the estimation of frequency selectivity, the proposed model proved to be remarkably accurate in fitting forward-masked CAP responses and could be extended to study more complex aspects of cochlear signal processing (e.g., compressive nonlinearities).

Timing of cochlear responses inferred from frequency-threshold tuning curves of auditory-nerve fibers

Links between frequency tuning and timing were explored in the responses to sound of auditory-nerve fibers. Synthetic transfer functions were constructed by combining filter functions, derived via minimum-phase computations from average frequency-threshold tuning curves of chinchilla auditory-nerve fibers with high spontaneous activity (Temchin et al., 2008), and signal-front delays specified by the latencies of basilar-membrane and auditory-nerve fiber responses to intense clicks (Temchin et al., 2005). The transfer functions predict several features of the phase-frequency curves of cochlear responses to tones, including their shape transitions in the regions with characteristic frequencies of 1 kHz and 3–4 kHz (Temchin and Ruggero, 2010). The transfer functions also predict the shapes of cochlear impulse responses, including the polarities of their frequency sweeps and their transition at characteristic frequencies around 1 kHz. Predictions are especially accurate for characteristic frequencies <1 kHz.

A ratchet mechanism for amplification in low-frequency mammalian hearing

The sensitivity and frequency selectivity of hearing result from tuned amplification by an active process in the mechanoreceptive hair cells. In most vertebrates, the active process stems from the active motility of hair bundles. The mammalian cochlea exhibits an additional form of mechanical activity termed electromotility: its outer hair cells (OHCs) change length upon electrical stimulation. The relative contributions of these two mechanisms to the active process in the mammalian inner ear is the subject of intense current debate. Here, we show that active hair-bundle motility and electromotility can together implement an efficient mechanism for amplification that functions like a ratchet: Sound-evoked forces, acting on the basilar membrane, are transmitted to the hair bundles, whereas electromotility decouples active hair-bundle forces from the basilar membrane. This unidirectional coupling can extend the hearing range well below the resonant frequency of the basilar membrane. It thereby provides a concept for low-frequency hearing that accounts for a variety of unexplained experimental observations from the cochlear apex, including the shape and phase behavior of apical tuning curves, their lack of significant nonlinearities, and the shape changes of threshold tuning curves of auditory-nerve fibers along the cochlea. The ratchet mechanism constitutes a general design principle for implementing mechanical amplification in engineering applications.

Two modes of motion of the alligator lizard cochlea: Measurements and model predictions

Measurements of motion of an in vitro preparation of the alligator lizard basilar papilla in response to sound demonstrate elliptical trajectories. These trajectories are consistent with the presence of both a translational and rotational mode of motion. The translational mode is independent of frequency, and the rotational mode has a displacement peak near 5 kHz. These measurements can be explained by a simple mechanical system in which the basilar papilla is supported asymmetrically on the basilar membrane. In a quantitative model, the translational admittance is compliant while the rotational admittance is second order. Best-fit model parameters are consistent with estimates based on anatomy and predict that fluid flow across hair bundles is a primary source of viscous damping. The model predicts that the rotational mode contributes to the high-frequency slopes of auditory nerve fiber tuning curves, providing a physical explanation for a low-pass filter required in models of this cochlea. The combination of modes makes the sensitivity of hair bundles more uniform with radial position than that which would result from pure rotation. A mechanical analogy with the organ of Corti suggests that these two modes of motion may also be present in the mammalian cochlea.

Shapes of cat auditory nerve fiber tuning curves

Tuning curves of auditory nerve fibers in normal-hearing cats were fitted by a computational model comprising four processes. One process accounts for sensitivity in tuning curve tails and consists of an approximation to bandpass filtering by extracochlear structures. The second and third processes describe passive and active components of basilar membrane (BM) mechanics, respectively. The former consists of a lowpass filter function, which provides baseline threshold sensitivity and filtering above characteristic frequency (CF), and the latter consists of a Gaussian that accounts for sharp tuning and high sensitivity around CF. A fourth process, modeled as a high-pass filter, was needed in many fits to account for breaks and plateaus in threshold sensitivity at frequencies above CF. The latter three processes operated on cochlear spatial coordinates rather than stimulus frequency. The four-process description closely accounted for shapes of most tuning curves. Tuning curve tails possessed minima at 40–80 dB SPL, and minima increased with fiber CF. High-frequency cutoffs of tail filters tended to increase with CF, but low-frequency cutoffs were generally constant across CF. Functions describing tails varied from ear to ear but behaved in a similar manner for fibers from a single ear. Passive components of BM resonances possessed baselines with sensitivities that decreased with CF and cutoff slopes that increased with CF. The magnitude of the active component increased smoothly with CF over an 80 + dB range, and its spatial extent was essentially constant at 1.5 mm or 6% of cochlear length regardless of gain magnitude, fiber CF, or threshold sensitivity. Tuning curves from fibers with high and medium spontaneous rates (SRs) and similar CFs had nearly identical shapes, with the sole difference being essentially constant differences in sensitivity across the entire excitatory frequency range. Tuning curve shapes from fibers with low SRs were more variable. These could either resemble those obtained from similarly-tuned fibers with higher SRs, or they could exhibit lower tip-to-tail ratios and reduced active component magnitudes. The latter were typically associated with low maximum discharge rates.

Comparative analysis of spectro-temporal receptive fields, reverse correlation functions, and frequency tuning curves of auditory-nerve fibers.

The tuning properties of single auditory-nerve fibers (ANFs) are characterized with spectro-temporal receptive fields (STRFs), reverse correlation functions (revcors), and frequency tuning curves (FTCs). Measures of tuning and latency from the STRFs and revcors are largely comparable to the traditional measures of tuning from FTCs and measures of latency from peristimulus time histograms (PSTHs), but several important differences are found. As is well known, revcors can only characterize low (< 6 kHz) best frequency (BF) units, whereas STRFs are able to characterize all units studied (BFs ranging from 0.26-23 kHz), except for a few very low-BF examples. Whereas tuning bandwidth derived from revcor exceeds that measured from FTCs at all BFs and increases with sound level. STRF bandwidth is comparable to FTC bandwidth, except at low BFs, and is stable with sound level. The STRF may reflect nonlinear properties of auditory-nerve fibers such as refractoriness and two-tone suppression that are absent in the FTC and revcor characterizations. The principal drawback of the STRF is its narrow dynamic range.

Automatic gain control in the bat's sonar receiver and the neuroethology of echolocation

The sensitivity of the echolocating bat, Eptesicus fuscus, to sonar echoes at different time delays after sonar emissions was measured in a two-choice echo detection experiment. Since echo delay is perceptually equivalent to target range, the experiment effectively measured sensitivity to targets at different ranges. The bat's threshold for detecting sonar echoes at a short delay of only 1.0 msec after emissions (corresponding to a range of 17 cm) was 36 dB SPL (peak to peak), but the threshold decreased to 8 dB SPL at a longer delay of 6.4 msec (a range of 1.1 m). Prior research has shown that, at even longer delays (corresponding to ranges of 3 to 5 m), the bat's threshold is in the region of 0 dB SPL. Contractions of the bat's middle ear muscles synchronized with the production of echolocation sounds cause a transient loss in hearing sensitivity which appears to account for the observed echo detection threshold shifts. The bat's echo detection thresholds increase by approximately 11 dB for each reduction in target range by a factor of 2 over the span from 17 cm to 1.1 m. As range shortens, the amplitude of echoes from small targets also increases, by 12 dB for each 2-fold reduction in range. Thus, when approaching a target, the bat compensates for changes in echo strength as target range shortens by changing its hearing threshold. Since this compensation appears to occur in the middle ear, the bat regulates echoes reaching the cochlea to a stable amplitude during its approach to a target such as a flying insect. In addition to this automatic gain control linked to target range, the bat aims its head to track a target's position during approach, thus stabilizing echo amplitude even if the target's direction changes. We hypothesize that the bat's directional emissions, directional hearing, middle ear muscle contractions, and head aim response collectively create a three-dimensional spatial tracking filter which the bat locks onto targets to stabilize echo amplitudes during interception of prey. We further hypothesize that this regulation, which cancels echo amplitude changes caused by the target's changing spatial position, leaves the bat free to observe echo amplitude changes caused by the target's own actions, such as insect wing beats. Elimination of spatially dependent echo amplitude changes removes the cause of potentially troublesome changes in neural response latency and keeps stimulation from echoes in the "tip" region of auditory nerve fiber tuning curves.(ABSTRACT TRUNCATED AT 400 WORDS)

Effects of continuous noise backgrounds on rate response of auditory nerve fibers in cat.

This study describes the effects of broadband background noise on the average discharge rate to best-frequency (BF) tones of auditory nerve fibers in the cat. The effects of exposure to long-term continuous noise are compared to the effects of noise gated on and off simultaneously with test tones. Addition of background noise causes a shift of the dynamic portion of tone-evoked rate versus level functions to higher tone intensities. The shift occurs at a mean rate of 0.61 dB of shift for each 1-dB increment in noise level. The rate of shift is independent of best frequency and spontaneous discharge rate. The noise level at which the shift begins is frequency dependent and is consistent with the frequency-dependent bandwidths of auditory nerve fiber tuning curves. The adjustment of the dynamic range shows many similarities to two-tone suppression. Therefore, it is most likely that it is caused by suppression of the response to the BF test tone by energy present in the noise at surrounding frequencies. At high noise levels, the ability of auditory nerve fibers to respond to test tones is limited by the rate response to the noise. As noise level increases, the discharge rate it evokes approaches a fiber's saturation rate and ultimately eliminates the fiber's ability to respond to test tones. Low spontaneous rate fibers, which have been shown to have higher thresholds and wider dynamic range (17,29), are significantly more resistant to saturation by high noise levels. Exposure to broadband noise prior to onset of test tones produces an overall decrement in response rate. This phenomenon is similar to the effects of short-term adaptation (32) and seems to develop independently of the shift of dynamic range. At high noise levels, previous exposure to the noise produces a small dynamic range shift. This effect is similar to that produced by suppression but is smaller. The effect is occluded in continuous noise backgrounds by the adjustment of sensitivity produced by suppression.

The effects of furosemide on the endocochlear potential and auditory-nerve fiber tuning curves in cats

Furosemide was injected intravenously in anesthetized cats while the endocochlear potential (EP) and tuning curves from fibers of the auditory nerve were simultaneously monitored in the same ear. The characteristic frequencies (CF's) of the fibers studied ranged from 0.25 to 28 kHz. Furosemide administration produced reversible, dose-related decreases in the EP which were accompanied by threshold elevations and alterations in the tuning curves of auditory nerve fibers. There was approximately l dB of threshold elevation for every millivolt decrease in the EP. For fibers with CF's above 3 kHz, threshold elevation at CF (in the tip of the tuning curve) was three times that in the tail of the tuning curve. Threshold elevation in the tip was always accompanied by threshold elevation in the tuning curve tail. Threshold elevation was dependent upon the CF of the fiber, with higher CF fibers showing larger threshold shifts than lower CF fibers. Threshold elevation was accompanied by a systematic shift in the CF of the fiber. Fibers with CF above 1 kHz exhibited downward shifts in CF, while those with CF less than 1 kHz generally exhibited upward shifts in CF.

Eighth-nerve action-potential tuning curves in cats before and after inducement of an acute noise trauma.

Potentials and limitations of eighth-nerve action-potential tuning curves (APTC) in providing us with a measure of frequency selectivity are studied in cats. APTCs are measured, using a forward-masking technique. They are essentially wider than auditory-nerve fiber tuning curves. Width of APTCs is dependent upon test-tone sound pressure level (SPL) and masking criterion. This can be explained by assuming that APs are formed by discharges of fiber with different CFs. Acoustic traumata are induced halfway into the experiments in order to measure APTCs before and during the trauma in the same animal. This broad-band noise exposure resulted in a long-lasting temporary threshold shift which remained fairly steady during the experiment. In the traumatized ear APTCs are wider than in the normal ear. This widening, also observed in auditory-nerve fibers, is not simply an SPL effect.

Physiological masking functions in normal and acoustically traumatized cats

In cats that have undergone acoustic traumatization, it has been observed that several characteristics in the shape of single auditory nerve fiber tuning curves correspond to histological changes in the hair cells of the cochlea [N. Y. S. Kiang, M. C. Liberman, and R. A. Levine, Ann. Otol. Rhinol. Laryngol. 85, 752–768 (1976)]. Dallos and Cheatham (J. Acoust. Soc. Am. 59, 591–597 (1976)] using the whole‐nerve action potential as a response have demonstrated masking curves in normal animals similar in shape to single‐fiber tuning curves. We have measured whole‐nerve action potential tuning curves (AP tuning curve) in both normal and acoustically traumatized cats to investigate the sensitivity of such a measure in reflecting changes in single‐fiber tuning curves. A tonal masker in a forward‐masking paradigm was used to elicit the measured responses. AP tuning curves were generated at several signal frequencies in order to measure the physiological changes at different places along the cochlea. A preliminary review of the data indicate that the AP tuning curves in acoustically traumatized animals can show a broadening of the tip region relative to normals.

Tails of tuning curves of auditory-nerve fibers.

Békésy's interest in improving telephone communication led him eventually to make observations on the mechanical properties of the basilar membrane. For apical parts of the membrane he obtained resonance curves with steep high-frequency slopes and gradual low-frequency slopes. Tuning curves for auditory-nerve fibers in anesthetized cats also show significant low-frequency tails. Even fibers with high characteristic frequencies (CF) respond to tones throughout a wide range of low and middle frequencies. For low-frequency tones at moderate and high stimulus levels the responses of high-CF fibers tend to be time-locked either to the same phase of the stimulus or to a phase that differs by a half cycle. For complex stimuli such as speech, which contains many low-frequency components, these fibers behave as broadly tuned elements, discharging with time patterns that are principally determined by the temporal characteristics of the acoustic waveform. Results of experiments using low- and high-frequency masking stimuli reinforce the view that responses of high-CF fibers to low-frequency stimuli should not be ignored in theories that seek to describe the role of the auditory nerve in speech communication.

Stria vascularis as source of endocochlear potential.

Stria vascularis as source of endocochlear potential.