The relationship between magnetic and electrophysiological responses to complex tactile stimuli

Background  

Magnetoencephalography (MEG) has become an increasingly popular technique for non-invasively characterizing neuromagnetic field changes in the brain at a high temporal resolution. To examine the reliability of the MEG signal, we compared magnetic and electrophysiological responses to complex natural stimuli from the same animals. We examined changes in neuromagnetic fields, local field potentials (LFP) and multi-unit activity (MUA) in macaque monkey primary somatosensory cortex that were induced by varying the rate of mechanical stimulation. Stimuli were applied to the fingertips with three inter-stimulus intervals (ISIs): 0.33s, 1s and 2s.     

Representation of harmonic complex stimuli in the ventral cochlear nucleus of the chinchilla.

The representation of Schroeder-phase harmonic complex sounds in the ventral cochlear nucleus (VCN) of the anesthetized chinchilla was studied. Stimuli consisted of a series of harmonically related sinusoids, multiples of a fundamental frequency (f0), summed in either negative (-SCHR) or positive (+SCHR) Schroeder phase. Psychoacoustic experiments performed in humans by other investigators have revealed that masking effects of -SCHR stimuli are larger than those found using +SCHR stimuli as maskers. In our laboratory, basilar membrane measurements at the base of the chinchilla cochlea show that responses to -SCHR stimuli are less "peaked," or modulated, than responses to +SCHR stimuli. We also found that suppression of a characteristic-frequency (CF) tone by -SCHR stimuli is larger than that evoked by +SCHR stimuli. Rate-intensity functions display higher firing rates in responses to -SCHR stimuli than in those produced by +SCHR stimuli. Firing rates evoked by either -SCHR or +SCHR stimuli saturate at lower values than those obtained in responses to CF tones. Rate and synchrony suppressions by -SCHR stimuli were larger than those evoked by +SCHR stimuli. Auditory nerve fiber responses to Schroeder complex stimuli share most of the properties of VCN responses, indicating little additional processing by the VCN.     

Perceptual interactions in the loudness of combined auditory and vibrotactile stimuli

The loudness of auditory (A), tactile (T), and auditory-tactile (A+T) stimuli was measured at supra-threshold levels. Auditory stimuli were pure tones presented binaurally through headphones; tactile stimuli were sinusoids delivered through a single-channel vibrator to the left middle fingertip. All stimuli were presented together with a broadband auditory noise. The A and T stimuli were presented at levels that were matched in loudness to that of the 200-Hz auditory tone at 25 dB sensation level. The 200-Hz auditory tone was then matched in loudness to various combinations of auditory and tactile stimuli (A+T), and purely auditory stimuli (A+A). The results indicate that the matched intensity of the 200-Hz auditory tone is less when the A+T and A+A stimuli are close together in frequency than when they are separated by an octave or more. This suggests that A+T integration may operate in a manner similar to that found in auditory critical band studies, further supporting a strong frequency relationship between the auditory and somatosensory systems.     

Dependence of loudness growth on skirts of excitation patterns

Over a range of 50 dB, the loudness of a 100-Hz tone was measured in the presence of a broadband noise with a low-frequency cutoff at 200 Hz. The noise was varied in intensity along along with the tone so that the signal-to-noise ratio remained constant at either 0 or--10 dB. Listeners judged the loudness of the tone by loudness matching, magnitude estimation, and magnitude production. The noise markedly decreased the tone's rate of loudness growth but not the range over which loudness grows. The overall decrease in steepness of the 100-Hz loudness function was greater than that previously reported at higher frequencies. It is hypothesized that the decrease was greater because the spread of excitation at 100 Hz was more effectively contained than at higher frequencies. Support for this hypothesis is given by measures of intensity discrimination at 100 Hz.     

Further evaluation of a model of loudness perception applied to cochlear hearing loss.

This paper describes further tests of a model for loudness perception in people with cochlear hearing loss. It is assumed that the hearing loss (the elevation in absolute threshold) at each audiometric frequency can be partitioned into a loss due to damage to outer hair cells (OHCs) and a loss due to damage to inner hair cells (IHCs) and/or neurons. The former affects primarily the active mechanism that amplifies the basilar membrane (BM) response to weak sounds. It is modeled by increasing the excitation level required for threshold, which results in a steeper growth of specific loudness with increasing excitation level. Loss of frequency selectivity, which results in broader excitation patterns, is also assumed to be directly related to the OHC loss. IHC damage is modeled by an attenuation of the calculated excitation level at each frequency. The model also allows for the possibility of complete loss of IHCs or functional neurons at certain places within the cochlea ("dead" regions). The parameters of the model (OHC loss at each audiometric frequency, plus frequency limits of the dead regions) were determined for three subjects with unilateral cochlear hearing loss, using data on loudness matches between sinusoids presented alternately to their two ears. Further experiments used bands of noise that were either 1-equivalent rectangular bandwidth (ERB) wide or 6-ERBs wide, centered at 1 kHz. Subjects made loudness matches for these bands of noise both within ears and across ears. The model was reasonably accurate in predicting the results of these matches without any further adjustment of the parameters.     

On the relations of intensity jnd's to loudness and neural noise

It is shown experimentally that, in contradiction of the fundamental concept of Fechner's law, the intensity jnd for auditory sinusoidal signals follows loudness, rather than its derivative with respect to sound intensity. The evidence is obtained by comparing the jnd's of a population with normal hearing to those of a population with hearing loss accompanied by loudness recruitment. Although the recruitment increases the slope of the loudness function, the jnd's of both populations were found to be practically equal when the loudness were equal. The phenomenon is accounted for mathematically by assuming that psychophysically relevant neural noise depends not only on the magnitude of loudness, but also on its derivative with respect to sound intensity. A related derivation accounts for the near miss to Weber's law.     

Effects of phase duration and electrode separation on loudness growth in cochlear implant listeners

Loudness estimates were obtained in a group of four adult subjects implanted with the Nucleus-22 multielectrode cochlear implant device, for a range of pulse amplitudes and different fixed phase durations and electrode separations. The stimulus was a 200-ms long train of biphasic pulses presented at 500 pulses/s. Subjects estimated loudness as a number from 0 ("don't hear it") to 100 ("uncomfortably loud"). Loudness was found to grow exponentially with pulse amplitude, at a rate that was dependent upon the phase duration as well as the electrode separation. An equation of the form L = e(lambda + gamma M)(D theta)I, where L is the estimated loudness, M is the separation between electrodes of a stimulating pair, D is the phase duration, I is current amplitude, and lambda, gamma, and theta are constants, appears to describe the observed data adequately. The findings are remarkably consistent across subjects.     

Relation of loudness to loudness just noticeable difference of tone

1IntroductionAnimportantprobleminpsychoacousticsandaudiologyishowtointerprettherelationofloudnesstoloudnessjustnoticeabledifference(JND).ThefirstpostulateabouttherelationwajsproposedbyFechner,whosuggestedthatloudnessofatonewascoulltofloudnessJNDfromhearingthreshold.AccordingtoFechner'spostulateandWeber'slaw[2],theloudnessLofatonewithintensityIisL=Inl.(1)Butagreatnumberofexperimentsdemonstratedthatloudnessisapowerfunctionofintensity3[')3f4]L=I",(2)wheretheexponentorvariesslightlywithinten…     

Auditory steady-state responses to chirp stimuli based on cochlear traveling wave delay

This study investigates the use of chirp stimuli to compensate for the cochlear traveling wave delay. The temporal dispersion in the cochlea is given by the traveling time, which in this study is estimated from latency-frequency functions obtained from (1) a cochlear model, (2) tone-burst auditory brain stem response (ABR) latencies, (3) and narrow-band ABR latencies. These latency-frequency functions are assumed to reflect the group delay of a linear system that modifies the phase spectrum of the applied stimulus. On the basis of this assumption, three chirps are constructed and evaluated in 49 normal-hearing subjects. The auditory steady-state responses to these chirps and to a click stimulus are compared at two levels of stimulation (30 and 50 dB nHL) and a rate of 90s. The chirps give shorter detection time and higher signal-to-noise ratio than the click. The shorter detection time obtained by the chirps is equivalent to an increase in stimulus level of 20 dB or more. The results indicate that a chirp is a more efficient stimulus than a click for the recording of early auditory evoked responses in normal-hearing adults using transient sounds at a high rate of stimulation.     

Temporal integration of loudness in listeners with hearing losses of primarily cochlear origin.

To investigate how hearing loss of primarily cochlear origin affects the loudness of brief tones, loudness matches between 5- and 200-ms tones were obtained as a function of level for 15 listeners with cochlear impairments and for seven age-matched controls. Three frequencies, usually 0.5, 1, and 4 kHz, were tested in each listener using a two-interval, two--alternative forced--choice (2I, 2AFC) paradigm with a roving-level, up-down adaptive procedure. Results for the normal listeners generally were consistent with published data [e.g., Florentine et al., J. Acoust Soc. Am. 99, 1633-1644 (1996)]. The amount of temporal integration--defined as the level difference between equally loud short and long tones--varied nonmonotonically with level and was largest at moderate levels. No consistent effect of frequency was apparent. The impaired listeners varied widely, but most showed a clear effect of level on the amount of temporal integration. Overall, their results appear consistent with expectations based on knowledge of the general properties of their loudness-growth functions and the equal-loudness-ratio hypothesis, which states that the loudness ratio between equal-SPL long and brief tones is the same at all SPLs. The impaired listeners' amounts of temporal integration at high SPLs often were larger than normal, although it was reduced near threshold. When evaluated at equal SLs, the amount of temporal integration well above threshold usually was in the low end of the normal range. Two listeners with abrupt high-frequency hearing losses (slopes > 50 dB/octave) showed larger-than-normal maximal amounts of temporal integration (40 to 50 dB). This finding is consistent with the shallow loudness functions predicted by our excitation-pattern model for impaired listeners [Florentine et al., in Modeling Sensorineural Hearing Loss, edited by W. Jesteadt (Erlbaum, Mahwah, NJ, 1997), pp. 187-198]. Loudness functions derived from impaired listeners' temporal-integration functions indicate that restoration of loudness in listeners with cochlear hearing loss usually will require the same gain whether the sound is short or long.     

Asymmetry of masking between complex tones and noise: partial loudness

This experiment examined the partial masking of periodic complex tones by a background of noise, and vice versa. The tones had a fundamental frequency (F0) of 62.5 or 250 Hz, and components were added in either cosine phase (CPH) or random phase (RPH). The tones and the noise were bandpass filtered into the same frequency region, from the tenth harmonic up to 5 kHz. The target alone was alternated with the target and the background; for the mixture, the background and target were either gated together, or the background was turned on 400 ms before, and off 200 ms after, the target. Subjects had to adjust the level of either the target alone or the target in the background so as to match the loudness of the target in the two intervals. The overall level of the background was 50 dB SPL, and loudness matches were obtained for several fixed levels of the target alone or in the background. The resulting loudness-matching functions showed clear asymmetry of partial masking. For a given target-to-background ratio, the partial loudness of a complex tone in a noise background was lower than the partial loudness of a noise in a complex tone background. Expressed as the target-to-background ratio required to achieve a given loudness, the asymmetry typically amounted to 12-16 dB. When the F0 of the complex tone was 62.5 Hz, the asymmetry of partial masking was greater for CPH than for RPH. When the F0 was 250 Hz, the asymmetry was greater for RPH than for CPH. Masked thresholds showed the same pattern as for partial masking for both F0's. Onset asynchrony had some effect on the loudness matching data when the target was just above its masked threshold, but did not significantly affect the level at which the target in the background reached its unmasked loudness. The results are interpreted in terms of the temporal structure of the stimuli.