Contrast enhancement improves the representation of /epsilon/-like vowels in the hearing-impaired auditory nerve.

This study examines the neural representation of the vowel /epsilon/ in the auditory nerve of acoustically traumatized cats and asks whether spectral modifications of the vowel can restore a normal neural representation. Four variants of /epsilon/, which differed primarily in the frequency of the second formant (F2), were used as stimuli. Normally, the rate-place code provides a robust representation of F2 for these vowels, in the sense that rate changes encode changes in F2 frequency [Conley and Keilson, J. Acoust. Soc. Am. 98, 3223 (1995)]. This representation is lost after acoustic trauma [Miller et al., J. Acoust. Soc. Am. 105, 311 (1999)]. Here it is shown that an improved representation of the F2 frequency can be gained by a form of high-frequency emphasis that is determined by both the hearing-loss profile and the spectral envelope of the vowel. Essentially, the vowel was high-pass filtered so that the F2 and F3 peaks were amplified without amplifying frequencies in the trough between F1 and F2. This modification improved the quality of the rate and temporal tonotopic representations of the vowel and restored sensitivity to the F2 frequency. Although a completely normal representation was not restored, this method shows promise as an approach to hearing-aid signal processing.

Discriminability of vowel representations in cat auditory-nerve fibers after acoustic trauma.

This paper attempts to connect deficits seen in the neural representation of speech with perceptual deficits. Responses of auditory-nerve fibers were studied in cats exposed to acoustic trauma. Four synthetic steady-state vowels were used as test signals; these stimuli are identical, except that the second format (F2) resonator in the synthesizer was set to 1.4, 1.5, 1.7, or 2 kHz, producing four spectra that differ mainly in the vicinity of the F2 frequency. These stimuli were presented to a large population (523) of auditory-nerve fibers in four cats with sloping high-frequency threshold shifts that reached 50-70 dB at 2-4 kHz. In normal animals, May et al. [Auditory Neurosci 3, 135-162 (1996)] showed previously that the discharge rates of fibers with best frequencies near the F2 frequencies provide enough information to allow discrimination of these stimuli at the performance levels shown by cats in behavioral experiments. Here it is shown that, after acoustic trauma, there is essentially no rate information which would allow the vowels with different F2 frequencies to be discriminated. However, information that could allow discrimination remains in the temporal (phase-locked) aspects of the responses.

Effects of high sound levels on responses to the vowel "eh" in cat auditory nerve.

The vowel "eh" was used to study auditory-nerve responses at high sound levels (60-110 dB). By changing the playback sampling rate of the stimulus, the second formant (F2) frequency was set at best frequency (BF) for fibers with BFs between 1 and 3 kHz. For vowel stimuli, auditory-nerve fibers tend to phase-lock to the formant component nearest the fiber's BF. The responses of fibers with BFs near F2 are captured by the F2 component, meaning that fibers respond as if the stimulus consisted only of the F2 component. These narrowband responses are seen up to levels of 80-100 dB, above which a response to F1 emerges. The F1 response grows, at the expense of the F2 response, and is dominant at the highest levels. The level at which the F1 response appears is BF dependent and is higher at lower BFs. This effect appears to be suppression of the F2 response by F1. At levels near 100 dB, a component 1/component 2 transition is observed. All components of the vowel undergo the transition simultaneously, as judged by the 180 degrees phase inversion that occurs at the C2 transition. Above the C2 threshold, a broadband response to many components of the vowel is observed. These results demonstrate that the neural representation of speech in normal ears is degraded at high sound levels, such as those used in hearing aids.

Spectral envelope coding in cat primary auditory cortex: linear and non-linear effects of stimulus characteristics.

Electrophysiological studies in mammal primary auditory cortex have demonstrated neuronal tuning and cortical spatial organization based upon spectral and temporal qualities of the stimulus including: its frequency, intensity, amplitude modulation and frequency modulation. Although communication and other behaviourally relevant sounds are usually complex, most response characterizations have used tonal stimuli. To better understand the mechanisms necessary to process complex sounds, we investigated neuronal responses to a specific class of broadband stimuli, auditory gratings or ripple stimuli, and compared the responses with single tone responses. Ripple stimuli consisted of 150-200 frequency components with the intensity of each component adjusted such that the envelope of the frequency spectrum is sinusoidal. It has been demonstrated that neurons are tuned to specific characteristics of those ripple stimulus including the intensity, the spacing of the peaks, and the location of the peaks and valleys (C. E. Schreiner and B. M. Calhoun, Auditory Neurosci., 1994; 1: 39-61). Although previous results showed that neuronal response strength varied with the intensity and the fundamental frequency of the stimulus, it is shown here that the relative response to different ripple spacings remains essentially constant with changes in the intensity and the fundamental frequency. These findings support a close relationship between pure-tone receptive fields and ripple transfer functions. However, variations of other stimulus characteristics, such as spectral modulation depth, result in non-linear alterations in the ripple transformation. The processing between the basilar membrane and the primary auditory cortex of broadband stimuli appears generally to be non-linear, although specific stimulus qualities, including the phase of the spectral envelope, are processed in a nearly linear manner.

An identified model for human wrist movements.

We have performed tests to find the mechanical properties of the hand and muscles driving wrist flexion and extension, and have identified parameters of a model. The hand acts as a nearly pure inertial load over most of its range of motion. It can be approximated as a rigid body rotating about a single axis. Viscosity of the wrist joint is negligible. Passive elastic torques are also small, except at extreme wrist angles. We measured torque as a function of wrist angle for maximum voluntary contractions, and angular velocity as a function of load. The torque/velocity curves for shortening muscles are well approximated by a Hill equation. To measure the "series elasticity" of the muscle equivalents, we imposed step changes in torque. The series stiffness is a monotonically increasing function of the preload, or "active state", in the Hill sense. We discuss the relationship of the measured parameters to properties of isolated muscles. To see the implications of the model structure for the "inverse problem" of identifying motor control signals, we simulated four models of different complexities, and found best fits to movement data, assuming simple pulse-shaped inputs. Inferred inputs depend strongly on model complexity. Finally, we compared the best fit control signals to recorded electromyograms.